CONTENTS.

PART I.—SEptemser, 1875.

I. Onthe Morphology of the Skull in the Woodpeckers (Picidee) and Wrynecks (Yungide).

By Prof. W. K. Parker, “B.S. (Communicated by H. T. Stainton, Lsq., Sec.

MIE Te SOG!) (Elecegyk. Vin)et Yen at ee yn EOE BY Space ay

IT. On some Atlantic Crustacea from the ‘ Challenger’ Expedition. By Dr. R. v.

Witiemoes-Sunm, Naturalist to the Expedition. (Communicated by Professor

Wayyiin DHOMSON,, 27-12... G8.) . (Plates WRX TDD) ee 28

III. On the Structure and Systematic Position of Stephanoscyphus mirabilis, the Type of

a new Order of Hydrozoa. By Professor Atuman, ID., LL.D., F.RS., §c.,

NEGCS ETAT SOGNe a (lab eNO cate A get eS eee oo Wo) we og op Wl wa MO!

PART IJ.—Drcremper, 1875.

IV. On the external Anatomy of Tanais vittatus, occurring with Limnoria and Chelura

‘terebrans im excavated Pier-wood. By Joun Denis Macponatp, JLD., F.B.S.,

Staff-Surgeon R.N., Assistant Professor of Naval Hygiene, Netley Medical School.

(Communicated by W. T. Tatseuton Dyur, IA., F.L.S.) (Plate XV.). . 67

V. On Valencinia Armandi, a new Nemertean. By W.C. M‘Intosu, ILD., F. RSF,

iS CPAATONON Taliare ret ie nen (ah tert te Met Me cn Me a ae ar at mete eee he

L ty J

PART Mii =juny, 1876:

VI. On thirty-one Species of Marine Planarians, collected partly by the late Dr. Kelaart,

F.L.S., at Trincomalee, and partly by Dr. Collingwood, F.L.S., in the Eastern

Seas. By Dr. Cutupert Cottinewoop. (Plates XVII.—XIX.) . . page 83

VII. On the Structure and Development of the Bird’s Skull. By W. K. ParKxsr,

HBS Holes 166) (bart Us) ((Rlates XGXeSXeXevail)| 2). 7 et

PART [V.—Janvary, 1877.

VIII. A Monograph of the Lepidopterous Genus Castnia and some allied Groups. By

Prof. J. O. Wxstwoop, U/.4., F.L.S., fe. (Plates XXVITI-XXXIII.). . 155

IX. On a new Genus of Trematoda, and some new or little-known Parasitic Hirudinei.

By Joann Denis Macponaup, ID., F.R.S., Deputy Inspector-General R.N.,

Professor of Naval Hygiene Army Medical School. (Communicated by G. EB.

DOBSON, Al MB: EES.) 1( Plate eXeXox Vi) see

X. On the Osteology and Pterylosis of the Spoon-billed Sandpiper (Kurynorhynchus

pygmeeus, Linn.). By Joun AnDERSON, ID., PRS, LS. ZS. S§¢.,

Curator of Indian Museum and Professor of Comparative Anatomy at Medical

College, Calcutta. (Plate XXXV. and 2 Woodeuts.). ..... =. . 213

PART V.—Ocroper, 1877.

XI. On Steere’s Sponge, a new Genus of the Hexactinellid Group of the Spongida. By

James Moris, ILD., LS, .GS., ae (Plates XXXVI. & XXXVII. and 2

WVOOd CUES Ait Meier aeon hae ce. ear wie eh tee Sy oo nae eae en

XII. Notes upon the Oxystomatous Crustacea. By Epwarv J. Mizrs, £.L.S., #.Z.S8.,

Assistant in the Zoological Department, British Museum. (Plates XXXVIII—

2.4) eee ae Ce rn Cee rein MC NG ee Vg ge BRS

XIII. On the Domestic Pig of Prehistoric times in Britain, and on the mutual Relations

of this Variety of Pig and Sus scrofa ferus, Sus cristatus, Sus andamanensis, and

Sus barbatus. By GrorGEe Roxiirston, I.D., F.R.S., F.L.S., Linacre Professor

of Anatomy ond Physiology, Oxford. (Plates XLIXLIII.) . .. . . 251

beaer J

XIV. On two new Forms of Deep-sea Ascidians, obtained during the Voyage of H.ILS.

‘Challenger. By H. N. Moseuey, .A., F.RS., Fellow of Exeter College,

Oxford. (Communicated by Dr. J. Muniz, F.Z.8.) (Plate XLIV.) . page 287

XV. On new Forms of Actiniaria dredged in the Deep Sea; with a Description of certain

Pelagic Surface-swimming Species. By H. N. Mosetey, W_4., F.R.S., Fellow of

Exeter College, Oxford, late Naturalist on board H.U.S. ‘ Challenger. (Com-

municated by Dr. J. Murie, F.L.S.) (Plate XLV.) . . . . . . . . 295

_ PART VI.—Novemser, 1877.

XVI. On the Birds collected by Professor J. B. Steere in the Philippine Archipelago.

By R. Bowpier Suarer, /.L.S., 2.Z.S., e., Senior Assistant Zoological Depart-

ment, British Museum. (Plates XLVI-LIV.‘and 2 Woodcuts.) . . . . 3807

PART VII.—Avetst, 1878.

XVII. On the Male Genital Armature in the European Rhopalocera. By ¥. BUCHANAN

Wisin I De hei Ss, so) (Plates UV-Vis a se DT

XVIII. Worphology of the Mammalian Ossicula auditis. By ALBAN H. G. Doran,

F.R.C.S. (Communicated by Professor Firowrr, F.R.S., FDS. &c. (Plates

AV AINTATESOXCIE ES Jarek Se Sameer tala he es PR oem Tia bee eat See eit

XIX. On the Annelida obtained during the cruise of H.U.S. ‘Valorous’ to Davis Strait

im 1875. By W. Carmicuart M‘Intosu, ID., LL.D., P.R.SS. L. & #., FLS.,

Sex (Rlateniaxevy and: 2 Woodeuts:)i 28 5 Yeah e eae a 2 ws & 499

XX. On the Sacral Plexus and Sacral Vertebre of Lizards and other Vertebrata. By

St. GzorcE Mrvart, Sec. Linn. Soc., and the Rev. Ropert Cxuarxe, F.L.S.

(Ristes avy, &UxeVE andi 9: Woodeuts.)) 2 5 9 20. 2 a «o/s 518

PART VIII.—Junz, 1879.

XXI. The Butterflies of Malacca. By ARTHUR GARDINER ButiER, /.LS., F.Z.S8.,

Senior Assistant in the Zoological Department, British Museum. (Plates LXVIIT.

£5, Tike TIN) SONS) AE CoC ch Me ee an ord ta eae ON a ee a A Re ee RE

brevet

XXII. On certain Organs of the Cidaride. By Cuares Stewart, Hsq., F.L.S.,

U.R.C.S., Lecturer on Comparative Anatomy, St. Thomas’s Hospital. (Plate

TX.) Retest) OR Til! she) oo SOE ee Menge Sena ao

XXIII. On Hypsiprimnodon, Ramsay, a Genus indicative of a distinct Family (Pleopo-

didze), in the Diprotodont Section of the Marsupialia. By Professor Owry, C.B.,

F.R.S., F.L.S., &c., Superintendent of the Natural-History Departments in the

British Museum. (Plates LXXI. & LXXII. with 8 Woodcuts.) , . . . 578

XXIV. Descriptions of some minute Hymenopterous Insects. By Prof. J. O. WEstwoop,

MAR Hh itaS.. ces, \(Rlate XX.) oe egeeeiediae os): (cee

Divers ee ei eee ke RS ies gE a ele

ERRATA.

Page 59, bottom line, for loupe read pocket-lens

», 240, line 15 from top, for WV. plicata read NV. sinuata

» 066, ,, 5 from top, for Albisara read Abisara

» 068, ,, 18 from bottom, for Tagisdes read Tagiades

THE

TRANSACTIONS

THE LINNEAN SOCIETY

LONDON.

SECOND SERIES.—ZOOLOGY.

VOLUME I.

PART. THE FIRST.

LON DION:

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET:

SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON-HOUSE ;

AND BY LONGMANS, GREEN, READER, AND DYER, PATHERNOSTER=-ROW.

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CONTENTS.

PART I.—1875.

I. On the Morphology of the Skull in the Woodpeckers (Picide) and Wrynecks

(Yungide). By W. K. Parker, Hsg., B.S. (Communicated by H. 'T. Statyton,

BE SG MISCO USS e AT NT a Pha adc ckeeaunes one) ess) a Neal wah Oe pace!

II. On some Atlantic Crustacea from the ‘ Challenger’ Expedition. By Dr. R. v.

Witiemors-Sunm, Naturalist to the Expedition. (Communicated by Prof. WYvILLE

PET ORESONE Die Hrs scale Se) ead iin oe Mi eA a Rl Un Oe Gee aeb

Ill. On the Structure and Systematic Position of Stephanoscyphus mirabilis, the Type

of anew Order of Hydrozoa. By Prof. Attman, U.D., LL.D., F.RS., &e., Pres.

RET ES OOM tiheaate Iie oye dined careoniet sb you ist heet pexh OW Le deck Paeces Ol

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PROTA ALE! ThA valootwaaiee) 4 eh ea RD ea

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+ See Melos AL VG hivale KANO) | : oats says a i oh Sather LY Aa

Se ae ae he amaty, eu 4 eae . }

SANS BAS Silitetid, cud QoeGnnlgale \w eae Sinaateatee di are nngil w

Wa aye CaN ASA OO 4 nt Le ORE ea con \ oh

TRANSACTIONS

THE LINNEAN SOCIETY.

I. On the Morphology of the Skull in the Woodpeckers (Picidee) and Wrynecks

(Yungide). By W. K. Parxsr, Esq., F.R.S. (Communicated by H. T. Statnton,

Esq., Sec. L.S8.)

(Plates I.-V.) ,

Read April 2nd, 1874.

INTRODUCTION.

MopERN zoologists, when at fault in their taxonomy, not knowing where to place

some perplexing specimen or type, begin to look to the working morphologist for

counsel and help in their difficulties.

These workers are merely separated for convenience’ sake: a thorough zoologist finds

no time for embryology ; an embryologist is most grateful to the skilled and accom-

plished classifier who arranges the various members of each faunal group ready to

his hand.

Hach kind of labourer has the greatest need of the results brought out by the other:

the patient dissector waits for the treasures supplied to him by the more mercurial

taxonomist ; whilst he, in turn, profits by the work of one to whom a single type may

serve for the labour of a year or more. Yet both are learning to look beneath the

surface of things, a growing knowledge of the types showing both that close kinship is

often marked by great difference in outward form, and that it is easy to be beguiled by

the external likeness of forms—isomorphic, indeed, but far apart zoologically. Never-

theless, on the whole, the keen eye of the zoological expert seldom errs in the grouping

of forms, even by their outward characters alone: but there are types that will baffle

all that skill; and then other counsel has to be called in.

Embryology, however, does not exist as a handmaid to zoology; its aims are higher

by far than that; and as for the zoologists proper, they exist for the morphologists, and

not the morphologists for them. ‘The familiar term “ zygodactyie” for birds with a

certain form of foot, has been very useful; and yet how much ignorance it may be made

to hide! It seems to be something when one knows that a certain bird belongs to that

- group; and yet a Cuckoo, a Parrot, and a Woodpecker come none the nearer each other

zoologically by the possession of that kind of foot.

SECOND SERIES.—ZOOLOGY, VOL. I. B

2 MR. W. K. PARKER ON THE MORPHOLOGY OF

The Woodpeckers, like the Parrots, form a well-defined group—too well defined for

the morphologist, who is pained for the paleontological losses the bird-fauna has sus-

tained, judging from analogy that the mixed forms are lost which would have yoked

the most divergent groups together. In the late geological period we have merely

the twigs of a few branches of the great genealogical bird-tree; the main branches and

the trunk are buried in the strata. Yet not the less does the morphologist seek in the

early stages of the young and embryo bird for the lineaments of the old feathered

fathers of the existent types. Hidden from the eye of the mere taxonomist, they yet

may be revealed to him, in some degree, by embryology; but birds cannot be inter-

preted morphologically, irrelatively to other vertebrata; judged merely by themselves,

and compared among themselves only, they present enigmas at every turn.

We all know now how peculiar is that type of ornithic face which is found in Passe-

rine birds, and is called by Professor Huxley the “Aigithognathous”’ palate. Nitzsch de-

scribed it; and I have long been familiar with its more marked peculiarities; but its

morphological importance dawned upon me when I saw that the parts of that complex

face, so conjugated and so metamorphosed, were really built up of elements which had

their true counterparts or “symmorphs”’ in the snake. But the snake does but repeat

these parts from the Amphibia; and the Amphibia borrow them from the cartilaginous

Fishes, amongst the lowest of which, namely the Lamprey, may be found the fullest

development, both morphologically and functionally, of cartilages that form the sub-

stratum of the most peculiar part of a Sparrow’s face.

Considered ornithologically, the ‘“ Picide”’ are full of interest; but looked at from a

higher stand-point they are almost of greater importance to the morphologist than the

Passerinz, having for the most part the same curious repetition of parts borrowed from

below, but also retaining more evident marks of their ascent from the quasi-pupal and

quasi-larval forms of the groups beneath.

The Woodpeckers have been designated “‘Celeomorphe” by Professor Huxley (Proce.

Zool. Soc. April 1867); but although the “ Picidee”’ and “ Yungide”’ are thus named as

to their general form, yet that author has not given a term for their peculiar facial

structure. With clearest insight, however, they are spoken of as “ not Desmognathous,

the palate of these birds exhibiting rather a degradation and simplification of the

Agithognathous structure” (p. 468). The fact is they are like early embryos of the

Passerine, in their palatal region arrested at a most simple and Lacertian stage,

whilst in other respects they are metamorphosed and specialized beyond any other

kind of birds. As far as their upper face is concerned, their arrested ‘“ maxillo-

palatines,”’ symmetrical “ vomers,” ‘“ septo-maxillaries,” and feebly developed turbinal

scrolls, entitle them to a name which shall be a memorial of their Lacertian facial

morphology. I therefore propose to call them the ‘‘ Saurognathe.”

On the Structure and Development of the Face in the “ Picidee (Celeomorphe,” H..,

*“ Saurognathe,” P.).

The necessity for a morphological study of birds becomes manifest when it is re-

membered that Coracias was for a very long time classed with the “ Corvide,” and that,

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 3

until the appearance of Professor Huxley’s paper, the best ornithologists classed birds of

the Cuckoo-form, together with the Woodpeckers and Wrynecks, as “ Picarie.’ So

much as the author of that paper made out of the palate of the Pici has been of great

value to me. My own study of the structure of these birds, both of hard and soft parts,

dates back to the early part of the year 1843; the unpublished results of that labour, in

the form of minutely careful drawings, are still of considerable value for reference.

More recently, before my friend’s researches were made, I had from time to time taken

up my preparations of the British species for the sake of comparing them with the skulls

of other ornithic types, and for their own elucidation. This was never fully satisfactory ;

and even with the help of Professor Huxley’s work, until very lately they continued to

be enigmatic.

In addition to my older material, including some nestling Wrynecks in spirits, the

opportunity to dissect three young green Woodpeckers, and to inspect and figure the

skulls of three southern species *, has resulted in the figures and descriptions which

I now offer. In the first species I shall describe the whole face, but in the others merely

the preeoral structures. This is in consonance with other researches of the same kind in

Avian families.

Morphologically considered, the palate is composed of just so much of the vertebrate

face as exists in front of the oral (double) cleft, namely :—the trabecule cranii, and the

parts developed around them in front and above; and below and behind those foremost

structures, so much of the mandibular arch as develops quasi-independently in the

maxillo-palatine lobe the pedate process of the mandibular arch.

These proper palatine structures form a secondary arch in front of the mouth, distinct

enough in the higher types; but in cartilaginous fishes, amphibia, &c. it shows its true

nature, in the former as a pedate spur of cartilage, growing forwards from the primary

apex of the visceral rod, and in the latter either as a fore-growing spur, as in the

** Urodela,” or as a conjugational band tying together the trabecula and the mandible, as

in the “ Anura.” ;

Thus in the large series of types which I have determined to compare together, it has

seemed fit to me to take a very small territory; yet that territory contains parts that

have undergone the greatest amount of metamorphosis of any in the whole body of a

high and noble vertebrate, and moreover being, in the bird, the skeletal framework of the

whole upper face, these parts are, as it were, an index of the amount of speciali-

zation undergone by any particular type—the ruling determining structures that lead to

all, and really demand all, the changes that take place in the rest of the organism. This

is especially explained for the benefit of those who will accuse me, and have already

accused Professor Huxley, of taking a narrow view of the Bird-types—touching with

the point of a needle some little tract, but unacquainted with and not able to appre-

ciate the Bird as a whole. This we calmly but indignantly deny; and the allegation

is the more ungrateful in that we do not come as those who would have dominion

over the faith of the mere zoologists, but as helpers of their work, letting in light from

a new chink.

* My helpers in this matter are my kind friends Messrs. Murie, Bartlett, Dines, and Salvin.

4 MR. W. K. PARKER ON THE MORPHOLOGY OF

On the Structure and Development of the Skull in Picus (Gecinus) viridis.

The extraordinary complexity of the palatine structures in the Green Woodpecker can

only be understood by reference to the unfused elements in the fledgling (Plate I, fig. 2,

for the palate as a whole, and Plate I. figs. 3-8, and Plate IT. figs. 5-7, showing details

and sections). In the palatal view of the young bird the nasal structures are purposely

left out for the better display of the facial bones—that part of the lateral ethmoid which

belongs to the 1st preoral, the ‘“‘ os uncinatum,’ being coloured, as it is an essentially

upper palatine element.

Beginning at the apical region of the trabeculz, we see rudimentary basipterygoids

set on the side of the basifacial region, where it begins to form the rounded “ rostrum”

(Plate I. fig. 2, d.pg). This parasphenoidal under-balk sharpens in front, and ends

behind the free fore edge of the mesoethmoid. The premaxillaries have together a high

triangular outline, and the outer bar (dentary portion) ends behind by lying for some dis-

tance on the dentary region of the maxillary ; so that whilst many birds are isomorphic

of the typical “‘ Teleostei’’ in having an internally placed “os mystaceum,” this bird

exposes as large a maaillary margin as the salmon. ‘The palatine processes of the pree-

maxillaries equal the solid part in length, and in breadth they equal the outer or

dentary bar. Their position also is somewhat analogous; for they pass on the iside of

the “ preepalatine,” whereas in many Passerines they pass on the outside. This reversed

situation of processes is seen again in certain delicate Australian Passerines, namely

Acanthorhynchus and Ptilotis. 'The nasal processes of the preemaxillaries are broad,

flat, and fibrous, and they show a median suture late in life. The “os uncinatum” (0. w)

is part of the “ ectoethmoid,’ and becomes as large as its distinct symmorph in those

Parrots which have an imperfect suborbital bar.

The symmetrical ‘“vomers” are not at first easy to find at this stage, being very

small; but by removing the palatine they can be seen as small styles, attached to the tips

of the long mesopterygoid processes (Plate III. figs 5 & 6, v, m.pg). The symmetrical

“ septo-maxillaries ” (s.mx) are also to be found, by careful search, on the inside of the

palatine bars; one was seen on the right, and two on the left side. The pterygoids (pg)

are of moderate length, and they have a long styloid “ mesopterygoid process ”’ ( Plate ITT.

figs, 5 & 6, pg, m.pg); on their upper surface the pterygoids have a free spur, pointing in

the same direction as the mesopterygoid: this is very long in the old bird (Plate III.

fig..7, pg); it is peculiar to the “ Picide.” The “ epipterygoid”’ hook is aborted; and

the mesopterygoid not being differentiated, there is then an evident arrest of metamor-

phosis in this proximal pier segment of the pterygopalatine arcade. The prepalatine

bar is normally long and slender; it is closely wedged in between the maxillary style

and the retral palatine process of the preemaxillary. At the middle this narrow part

ceases ; the broad posterior half is gently suleate along its middle; and the inner portion

sends forward a styloid interpalatine spur (Plate I. fig. 2, pa, i.pa). The “ interpalatine

edges”’ of the two bones are almost parallel from the spur to the end of the base. The

broad part begins to narrow at its outer edge midway ; and forming an obtuse angle there,

the transpalatine is aborted. The upper edges meet at the mid line; and thus the nasal

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 5

canal is three fourths closed. The commissure above is peculiar: each bone has a car-

tilaginous selvedge; these meet together, coalesce, and in front endostosis takes place,

so that a dagger-shaped wedge of bone divides the sides of this arch. This is the “ medio-

palatine” (Plate I. fig. 2, and Plate IIT. figs.5 & 6, m.pa). The “ ethmo-palatine”

region is premorse, having no long spur as in most birds. The secondary parts of the

palatine arch equal in interest those formed directly in the primary bar: these are the

maxillary, palato-maxillary, and jugal. The maxillary (Plate I. fig. 2, and Plate III:

fig. 6, mx) is most developed in this species; but the maxillo-palatine plate is merely a

small suboval outgrowth, joined to the main bar by a wide isthmus; it is convex above

and concave below.

This process is far apart from its fellow, and grows out of the narrow maxillary bar

nearer to the posterior than the anterior end; both of these regions are long narrow

styles; and the bone grows outwards most where it gives off the inner plate or “ maxillo-

palatine.” The essential nature of the maxillary is well seen here as a long bony

bar applied to the outside of the second preeoral arch—long, because of the very pro-

gnathous face of the bird; in this case its palatine portion is scarcely more developed

than in the Teleostean fish. The posterior splint, the jugal, is of the normal size, and, as

in many of the arboreal birds, has no separate quadrato-jugal attached to it. Besides the

« maxillo-palatine ” process, there is, as in certain of the “‘ Coracomorphe,” Cardinals,

Buntings, &c., a “ palato-maxillary ” bone. This, however, gives a peculiar asymmetry

to these birds; for it only appears on the /ef¢ side (Plate I. fig. 2, p.mx): in the young

bird it is a small wedge attached to the inner side of the main maxillary, some distance

in front of the “ maxillo-palatine process.” Thus we see that the single mammalian

maxillary may be represented by ¢hree bones in the bird: the “ postmaxillary,” which

I first observed in the young Emu, is about as common as the distinct palato-maxillary ;

the “ septo-maxillary’’ belongs to the first preeoral arch *.

The newly fledged Green Woodpecker presents an excellent subject for determining

the morphological meaning of the parts composing the nasal labyrinth. These have

always to be worked out with the preoral arches, with which they are even more inti-

mately blended, by metamorphosis, than the postoral with the ear-sac. A vertical section

of the head, made a little to the left of the axial cartilage, is very instructive (Plate I.

fig. 3). This section takes in the whole length of the trabecular arch from its apex,

now involved in the basisphenoidal mass (6.s), to its azygous extremity, the shrunken

preenasal (p.m). The “ parasphenoid”’ has its “rostrum”” still distinct ; but the broad

hinder portion has been ingrafted on the apices of the trabecule and the squared end

of the “investing mass,” to form that composite cranio-facial bone, the “ basisphenoid.”’

The bony matter has reached to the ‘“ optic commissure” and the sides of the deeply

cupped “ sella turcica.”” A line drawn obliquely through the middle of the circular

optic foramen (2) and the oval interorbital space (é.0. s) would exactly divide the facial

from the cranial elements, so soon, by metamorphosis, wrought into a common mags,

* In the “ Fowl” paper {note at bottom of p. 773) I have spoken of the “ septo-maxillary” as “distinct in no

bird.” Only within the last few years have I found the true centre; and it is very commonly present, not, however,

in the subject of that paper, the Fowl.

6 MR. W. K. PARKER ON THE MORPHOLOGY OF

and only partly redivided by this same interorbital window. Here turns up the ex-

planation as to why the anterior sphenoid (p.s) should be perched up high above the

ethmoidal and basisphenoidal base-line. This part, with its arrested “ orbito-sphenoidal ”

lips, is at present unossified ; it has, afterwards at least, three centres—one for the body,

and a large pair for the wings; and it ends above in the peaked extremity of the frontal

region of the mesoethmoid (m.eth). This latter region is very large; but its lower half

is borrowed territory, belonging to the subjacent portion of the long trabecular com-

missure with its ascending crest; four fifths of this composite plate is already ossified,

about to become the largest mesoethmoid (relatively) in the vertebrate series: I speak

of that of Birds generally. In the adult the double nasal sac is separated for the most

part by the merest film of membrane (Plate IT. fig. 5, c. fm) : this fenestrate space has com-

menced as a “‘ notch” by the absorption of part of the trabecular tract (Plate I. fig. 3

c.f.n), both base and crest. This is the beginning of the notch which at once divides

the first facial arch into a proximal and a distal portion, and also separates the middle

wall of the nasal labyrinth into a vestibular and a true sensory region. The septum nasi

{s.n) is a thinnish continuous plate of cartilage, terminated by the stunted preenasal

above, and by the thick projecting trabecular cornua below (fig. 3, p.n, c.tv). An outer

side view of the same part of the face (fig. 4) shows that the great median ethmoidal

ossification (“ pars perpendicularis”’) is growing outwards into the alee, or upper part of

the “ectoethmoid,’ whilst the lower part (“‘ pars plana’’) has its own ossification

(p-e; p-p)-

The angle below the centre for the pars plana is the substratum that should form the

“os uncinatum ;”’ the mass in front of this part, and below the superior turbinal (w. <b),

is the nasal gland (m.g); the “ aliseptal”’ (a/.s), or root of the inferior turbinal, is seen

above; in front of this the “ alinasal”’ fold (al.2) ; below this the nasal floor (x. f), and

altogether in front the pedate trabecular cornu (c. ¢7). All these parts are brought into

daylight by transverse sections (Plate I. figs. 5-8). In the foremost of these (fig. 5) the

recurrent trabecular horns (c. ¢) are shown as two thick lobular masses of cartilage, with

a binding isthmus formed between the ends of the “ cornua.” This is very similar to

what is seen in an ordinary Mammal, e.g. the Pig. Hedging in the tuberous mass we

see the nasal processes of the preemaxillaries (nv. px), the dentary part (d. px), the fore

shaft of the maxillary (mx), the palatine spur of the preemaxillary (p. px), and outside

this the preepalatine bar (p.pa). In the next section (fig. 6) the same bony sections are

shown, with the exception of the palatine portion of the premaxillary. The “ nasal

nerves’ (7.7%) come into view here below the fore part of the septum (s. ), and at the top

this wall has grown into a pair of roof-plates (aliseptals), which develop into a large

thick half-scroll, the “ alinasal turbinal” (al. t+). Outside these valvular growths

there is an infero-lateral development of cartilage, which at first is a wall and then a floor

(n. w,.f). In the succeeding section (fig. 7) new parts come into view: the two arms

of the nasal bone (m); the interpalatine spur (i.pa); and the fore part of the “inferior

turbinal ” (7. ¢b). The section of this arrested scroll is hammer-shaped, an outgrowth

being developed inside as well as outside the root; the outer coil is the rudiment of the

scroll, which in Rhea and Tinamus is coiled upon itself three times (“Ostrich skull,”

THE SKULL IN THE WOODPECKERS AND WRYNECKS. i

' plate x. figs. 14 & 15, and plate xv. fig. 7. ¢b); in the Fowl (“ Fowl’s skull,” plate Ixxxvi.

figs. 4—6, 7. tb) it has two coils. In the same figure (7) the alinasal turbinal (a/. ¢0) is

seen to form one nearly perfect coil. It is somewhat more like the inferior turbinal of

ordinary birds. In these things the Woodpecker is but little more specialized than a

Lizard or Sea-Turtle.

Behind, the inferior turbinal (fig. 8) is continuous above with the aliethmoid (a/.e),

which infolds itself, is pinched in and crenate, and becomes a rudimentary superior tur-

binal. This will be better seen in the thoroughly ossified parts of the adult (Plate V.

fig. 2, w. tb): where the inferior turbinal ends, running into the front of the antorbital

plate, there a rudimentary “middle turbinal”’ is seen (ibid. m. ¢b). In the young bird

this section (Plate I. fig. 8) shows the termination of the nasal floor (n.f), curving

upwards to the partially ossified “ perpendicular plate” (p.e) ; the nasal nerves (n. 7),

the palatine (pa), the nasals, frontals, and nasal processes of the preemaxillaries (n, f, n.p2)

are also cut through, and a good view is obtained of the extent and relations of the

“‘ maxillo-palatine processes” (mwx.p). The condition of the nasal floor in this section

is preparatory to a remarkable metamorphosis, to be described hereafter. The adult skull

and face (Plate II.) can now be interpreted, notwithstanding the mixture there is in it

of arrest and of an unusual degree of metamorphosis.

_ The palate (figs. 1 & 2) is a remarkable piece of basketwork; and as in this species

ossification is intense, the nasal labyrinth brings in its many bony parts to add to the

complexity. The preoral facial bones are thin and splintery, but are, notwithstanding,

very strong. The strongest part is the solid fore end of the preemaxillary; yet it is

quite hollow, forming scarcely any diploé except short side props at the junction of the

lateral with the lower plate. The head is the hammer, and the bony beak the helve, to

this curious combination of axe and chisel, the dense horny sheath forming an apt “suc-

cedaneum ” for steel. The bindings on and interlacements of the facial plates and bars

have their meaning in this, that they carry off the jar from the brain of this plumy artisan.

Taking the parts in detail, we find that the “ basipterygoid processes” (b.pg) are arrested,

and yet the pterygoids cleave close to the converging cranio-facial base.

The cleanly cut groove formed by the pterygo-palatine bars glides smoothly under the

rounded “ parasphenoidal rostrum,” (7. 6.8); this is a synovial joint : the rostrum ends in

a free style behind the oblique fore edge of the ethmoid, which spreads into two small

wings (figs. 1, 2, 3, 7.0.s, p.e). The “notch” from being small and kidney-shaped

(Plate I. fig. 3, ¢.f. 7), has grown into a very large nearly closed fenestra, which runs

forward to near the end of the septum. Thus the septum nasi (Plate IT. fig. 3, s.z) is

reduced to a sharp bony keel running down a short way below the double sac; and the

trabecular remnant, with its stunted przenasal beak (p.2), has an equally low crest.

One third of the trabecular bar is absorbed entirely behind; and scarcely half the

remainder is ossified by its centre; the rest is a continued growth of bony matter from

the “ septum” and the “ trabecular cornua” (fig. 1, s.”,¢.¢r). The “notch” forming

the cranio-facial hinge does not run through; it is arrested earlier than in the Gal-

linaceous birds (“ Fowl’s Skull,” plate Ixxxvi. fig. 14); and being anchylosed to the

ethmoid, the face sits more stiffly on the head than in most birds. The palatine process of

8 MR. W. K. PARKER ON THE MORPHOLOGY OF

the preemaxillary (figs. 1, 2,3, p.pa) comes, besides, under the fore part of the septum with

its recurrent wings; it has anchylosed with the inner edge of the long lathy pre-

palatine bars (p.pa). There is also a slender ‘“‘ median palatine process” to the pre-

maxillaries (m.p. px); in the specimens before me it is distinct from the nasal septum ;

but in another of my preparations, now in the College of Surgeons, the two parts are

anchylosed. The dentary region of the premaxillary overlaps, and is completely fused

with the maxillary; the nasal processes (fig. 5, 2.px) are well anchylosed to their nasal

and frontal surroundings, with a small part of their own median suture evident.

It is in the intermediate splints, however, rather than in the parasphenoid and pre-

maxillaries, that the peculiarity of the Woodpecker’s palate is most remarkable; these

are the “ vomers”’ and “ septo-maxillaries.”

It is evident that Professor Huxley had not appreciated the true nature of the latter

bones in the Reptilian skull in his ‘Elements ;’ for both in the Lizard and the Snake

(fig. 90, p. 225) they are described as ossifications in the internasal cartilage. Such

ossification I have never seen; and the bones figured are what I at first called in the

Reptile “ praevomers,”’ and now call “ septo-maxillaries ” *.

Seen from below (Plate IT. figs. 1 & 2, v.s.mz), the vomerine bones are attached loosely,

behind, to the premorse “ethmo-palatine”’ (e.pa), but more strongly to the inner

edge of the “ maxillo-palatine” plate (ma.p) and of the alinasal and inferior turbinals

(al.tb, 7. tb). Their position is vertical, their convex face within towards the septum

nasi, and their concave face towards the turbinal scrolls.

The inner view (fig. 3) shows them edgewise; but they are also shown as dissected

away, with the inner edge of the “ maxillo-palatine ” in figs. 4 & 5: here the piece of

the maxillary is thrown back to display the other pieces. The palatine bones and

maxillo-palatine plates lie directly beneath the vomerine series ; in figs. 2 & 6 the palatine

is cut away to display them better. The main septo-maxillary (s.ma%) is a conchoidal

bone, equal in size to the broadest, inturned part of the maxillary just adjacent. It has

a strong keel on its septal or convex surface ; and its hinder margin is broadly transverse,

ragged at its angles, and has its edge thickened on the inner surface, like the edge of a

bivalve shell. The lower edge of the bone is attached to the maxillo-palatine; and its

narrowed anterior dentate margin carries a ragged wedge of bone, the “ anterior septo-

maxillary” (s.ma’). Between the dorsal keel and the uncinate posterior angle there is a

rough wedge of bone one fourth its size: this is the main “vomer”’ (v’); it is sharp in

front and notched behind. In the ligament which attaches the main vomerine piece to

stunted “‘ ethmo-palatine” edge (e.pa), there is another, much smaller, irregularly oblong

bony centre. ‘his is the posterior vomer (v”). The large conchoidal “ septo-maxillary ”

is a compound bone, as I shall soon show. These four pairs of bones do not, however,

exhaust the vomerine series; for the tract of ligamentous tissue hedging in the great

*‘notch’’ below, from the mesoethmoid to the septum nasi, has in it two “ median septo-

* My first term is perfectly correct if the facial arches are looked upon as a series of rib-like hoops growing down-

ward from the head. These bones appear to have been confused with the Mammalian “ cornets inférieurs ” by most

authors. Professor Owen (‘ Report on Archetype,’ 1846, p. 293) speaks of them in Reptiles as “‘ The small and in-

constant ossifications in the capsule of the organ of smell.”

THE SKULL IN THE WOODPECKERS AND: WRYNECKS. 9

maxillaries,” small ovate-oblong patches of bone (figs. 1, 2, 5, m. s.mx). A single bone in

this region is not uncommon in Birds, as I have already shown. These apparently

azygous bones will be explained by a younger specimen.

In this species (the Green Woodpecker) the main septo-maxillary piece rivals its

symmorph in many of the Lizard kind; and when all the details of this instructive but

perplexing skull are mastered, I am confident that the term “saurognathous” for this

kind of palate will not be thought inappropriate.

The nasal labyrinth is so intimately connected with the trabecular arch that its

description may well come in here. I have already shown its structure in the July bird,

and also the relation of the median part to the first arch.

In this species the turbinals and al nasi are endosteally ossified; the alee project

strongly in front, and in infolding themselves form a remarkable boss on each side,

which is strongly attached to the fore part of the septum nasi by continuous ossification

(figs. 1, 2, 5, al.n, c.tr). These bosses are the trabecular cornua. The “alinasal

turbinal,” coiled once upon itself (Plate V. figs. 6 & 7, al. tb), is of enormous size; the

“inferior turbinal,”’ equally simple (Plate V. figs. 7 & 8, 2. ¢), is one fourth less. In a

front view of the antorbitals and perpendicular ethmoid (Plate V. fig. 2, p. e) we see the

“rostrum ” of the “ parasphenoid,” the fore end of the median ethmoid, flat-faced below,

and growing into the aliethmoid above the elegant ear-shaped “superior turbinals”’

(wu. tb), the rudiments of the “middle” (m. tb), and scars where the inferior turbinals

(«. tb) have been broken away. The parts are intensely anchylosed ; but we can discover

the passages for the first and nasal branch of the fifth nerve (1, 5), the great size of the

“ ectoethmoid,”’ mounting to the top above, and sending backward a large “ uncinate”

process below. The frontal bones (/), strongly roofing in the forehead, are thrown into

a solid ridge by the tips of the ‘ cornua majora” (4r. 1), which turn to the right side,

and lie in an obliquely directed smooth groove.

The second, or pterygo-palatine arch (figs. 1, 2, 4) has a short posterior segment, or

pier, the pterygoid (pg). This bone is broad, inwardly bent, and bifurcate (Plate

III. fig. 7, pg); for it sends upwards and forwards a process, or muscular attachment,

behind the true mesopterygoid process (m.pg): the latter does not become segmented

off; and hence this bespeaks less metamorphosis than what obtains in a Passerine bird.

There is no “epipterygoid’’ process. The long, flat, elastic palatines (pa), are anchy-

losed to the dentary and palatine portions of the preemaxillary, but not to any part of the

maxilla or their own splints. The “transpalatine” angle is aborted and obtuse; the

inferior surface of the broad part of the bone is gently sulcate along its middle. This

broad part is of great extent, and ends in the slender “interpalatine” spine (¢.pa),

which looks towards the free retral end of the palatine plate of the premaxillary, but does

not in this instance help it to enclose an oval space; it may do so in very old birds.

The postpalatine region is narrow, the bones attenuating suddenly ; and the “ ethmo-

palatine” lamine (e.pa) are of very small extent, and have no spine, or only a rudiment

for attachment to the vomerine ligament ; the plate is fenestrate on the right side. The

commissure is finished by the endosteal mediopalatine (fig. 2, m.pa), a lozenge-shaped

bone, dagger-like behind. The palatine arch is seen to be fenced on its outer side by a

SECOND SERIES.—ZOOLUGY, VOL. I. Cc

10 MR. W. K. PARKER ON THE MORPHOLOGY OF

bony lath, which runs from the body of the preemaxillary to the cup in the outer face of

the quadrate (fig. 4, q). The main part of this outer bar is the maxillary; the terminal

style is the jugal (see also Plate I. fig. 2, mx, j). The ear-shaped “maxillo-palatine”’ is

scooped above, where the ‘‘inturned nasal wall” (Plate I. fig. 8) and alinasal turbinal

(al. tb) rest upon it. Opposite the palato-pramaxillary hook we see a smallish, oval,

bony plate, on the left side of the palate—the palato-maxillary (Plate II. figs. 1, 2,

p.m).

In this species I cannot pass by the postoral arches without some notice, notwith-

standing the greater interest of the preorals. The structure and development of the

mandible (Plate V. fig. 1) corresponds very exactly with that of the Crow-form; the

internal angular process is strong and recurved, and the posterior process short and

broad. The five pairs of splints, and one pair of endosteal bones (‘‘ articulo-Meckelian ”’),

form but one V-shaped bone in the adult, with but little trace of sutures, and with no

fenestra. The ossified “symphysis” is shortish; and that end of the bone is formed on

the model of the upper beak.

The stapedial apex of the largely aborted second postoral arch has some peculiarities of

importance. The ¢rue stapedial, or periotic portion (Plate I. fig. 11), is rather large and

roughly oval, the side towards the “‘ opisthotic” bar separating the fenestra ovalis from

the fenestra rotunda being straightest. .

The capitular portion of the arch, continuously ossified with the base, is the flattish

*““mediostapedial’’ (m.s¢); a bony rod from this bar runs down the anterior “ infra-

stapedial”” bar (z.s¢), bringing to mind the small bone in the “‘stapedius” muscle of

the mammal; it is its symmorph. 'This double ‘“infrastapedial” is new to me; it ends

below in a spatulate stylohyal (st.h), which has a proximal ossicle just below the infra-

stapedial fenestra. The extrastapedial (e.s¢) is faleate and broad-backed ; the “‘supra-

stapedial”’ (s.st) is small; and from it and from the proximal end of the extrastapedial

a fibrous fan arises, which supplements the small “tubercular” head of this facial rod.

The proximal portion of the large “ stapedial fenestra” (f) is hidden in this view ; but it

scoops the faleate ‘‘ extrastapedial”’ beneath its thick outer or back part.

The rest of the arch, save its merest extremity, is membranous; and the terminal

rudiments (“‘cornua minora”’) have coalesced. These (Plate I. figs. 9,10, c.h) form,

even in my youngest specimen, a spearhead-shaped piece, notched deeply above, and

less below; for there the coalescence of the two ossified rods is greatest. The upper sur-

face is flat, the lower subcarinate; the whole piece is covered with a barbed horny sheath

(Plate V. fig. 1, ¢.h). The rest of the hyoidean skeleton is a most exquisitely elastic

structure of thread-like bones, ossified, at first, endosteally (Plate I. figs. 10, 11, 6.07).

The basihyal (Plate V. fig. 1, 6.67) is 13 inch in length, the proximal part of the next arch

(1st branchial) (67. 1) 1 inch 2 lines, and the distal portion is about 43 inches in length;

each ramus of the 3rd postoral is therefore about 6 inches long. There isno “ urohyal.”’

In this the Woodpeckers agree with the Gannet, Baleniceps, Rhea, and some other

“‘ Microglosse.”

Besides the smooth, shallowish, double groove on the upper part of the head, there are

to be noticed the feather-pits with which this skull is dinted. The whole skull is very

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 1a

strong, and the orbits well marked; the interorbital space is small; the lachrymal is a

little bone attached to the nasal. The exoccipital and basitemporal together, with the

help of about three tympanics, that soon coalesce, form the most curious tympanum seen

in the class; it is like a cowrie shell: the Duck comes nearest to the Pici in this

respect. Even with materials at hand, I cannot go further into the skull, as the fore

Jace is at present my proper field.

Since the above description of the skull of the fledglings and the adult Green Wood-

pecker were written, Samuel Whitbread, Esq., F.R.S., has sent* me a first-autwmmn spe-

cimen of this kind; and this stage happily bridges over the great distance between the

fledgling and the old bird. Its description may well come in after that of the adult

skull; for it serves as a most perfect key to the mysteriously complex basketwork of the

old bird’s palate, and also sheds a clear light upon the embryo-passerine characters seen in

these birds.

The progress of anchylosis through the five additional months has been intense, and

many new parts have appeared. This is in conformity with what was observed in the

Fowl, many of the metamorphic changes taking place during the first year, although so

much had been gone through in the egg. |

The anterior trabecular splints, the preemaxillaries (Plate IT. fig. 6, px), have coalesced

with the palatines and maxillaries; and the appearance of free ends to their palatine

processes, well seen in Picus minor (Plate IV. fig. 1, p.sp.x), does not appear to be the

ease here, the retral snags being due in the Green Woodpecker to additional ossicles.

All the three membrane bones that lie near the septum, on each side, belong. to the

vomerine series (vomers and “ septo-maxillaries”’); and those which had appeared on the

fledgling (Plate I. fig. 2, v, s.mx) have now attained their fullest free development.

Yet I cannot find the “posterior vomer” of the old bird here (Plate IT. figs. 2-5, x”).

The series of “ mediotrabecular splints” forms a long chain, reaching from the anterior

end of the przenasal style (p.m) to the “ ethmo-palatine laminz.”

‘Beginning at the hinder extremity of the series, we have the true symmorphs of the

vomerine crura of the Passerine bird (the “shoulders” of the compound bone I shall

describe presently); and only on the right side do I see any attempt at cutting off the

“posterior vomer”’ (Plate IT. figs. 6 & 8 v”).

These “ free vomerine crura”’ are long, thin, half-coiled bones, convex outside, concave

within, thick below, sharp and serrate above. They do not reach their proper fastening-

point (the ethmo-palatines) behind, having become dislocated by growth (see Plate IIT.

figs. 5 & 6, v, for their primordial “ Rhynchosaurian ” position) f.

Attached to the fore end of the vomer by fibrous tissue, is a somewhat slenderer

needle-shaped bone (Plate IT. figs. 6 & 8, sma’). This is the hindermost of the “ septo-

maxillary chain ;’”’ it undergirds the “inturned alinasal lamine ” (¢. a. 1), to which it is

strongly tied by fibrous tissue. A similar bone, but still slenderer, is attached to the

* This bird was sent from Southill (Biggleswade) on the 1st of November, 1872; it was a young of the first

autumn, and therefore half a year old.

+ Compare those figures of the youngest Green Woodpecker’s vomer with Dr. Giinther’s figures and descriptions

of Hatteria (Phil. Trans. 1867, part ii. plate i. fig. 2, p. 5).

c 2

12 MR. W. K. PARKER ON THE MORPHOLOGY OF

front of this (s.ma*); this reaches to, and is bound under, the trabecular cornu (c. ¢r)

The retral growths of the alinasal folds have also two short irregularly oval patches of

fibrous bone, the 3rd and 2nd “septo-maxillaries”’ (s.ma’*, s.ma’); and these have

running in front of them the first of the septo-maxillary series (s.ma'); it is a frail,

slightly inbent style, running as far forward as the preenasal (fig. 3, pz). The ossicles

at the mid line have a character which was not suspected in the examination of the

adult; and they were not discernible in the fledglings; the section of these (Plate L.

figs. 7 & 8) showed no submesial cartilage inside the inturned alinasal lamina (¢. a/, 1.w).

Here, however late and abortively developed, we have the “labial cartilages” ossified,

though independently of the other vomerine bones (vomers and “ septo-maxillaries ”),

and very unsymmetrical in shape. The /eft bone (J. c’) is hindermost; it is tuberculate

behind, broad and grooved below in the middle, and styloid in front; this styloid part is

strongly bound beneath the right piece (J. c'). This latter is a sessile leaf of bone, on a

long fore-and-aft style, the foremost style being tied beneath the true septal bone or

trabecular ossicle (s.”), In the embryo of the Linnet (to be described in another paper)

are to be seen the labial cartilages of the Passerine bird, which so soon coalesce, and then

send backward exogenous “crura;’’ here, in the Woodpecker, they are abortively developed,

come nearer and nearer to the mid line (see in the old bird, Plate IT. figs. 1, 2, 3, s. 2, s.mza),

and appear like mere patches of fibrous bone. Almost all the elements of the complex

highly metamorphosed Passerine vomer keep apart in the Woodpecker, but not al/; the

5th septo-maxillary coalesces with the ossified inturned lamina (Plate II. figs. 2 & 5,

s.mx,%. al); and that part of the nasal capsule, contrary to all experience, detaches

itself from the rest, and becomes the scooped part of the complex 5th septo-maxillary of

the old bird. In the Passerines this part of the nasal capsule coalesces with, and often

becomes largely ossified by, the contiguous vomerine bones. In the old bird (Plate IT.

figs. 1-5) the Ist, 2nd, & 3rd septo-maxillaries coalesce with the contiguous parts of

the preemaxillary and infero-mesial parts of the nasal capsule. In the subadult bird

(Plate IL.) the huge, simple “alinasal turbinals” (a/. 70) are ossifying below; the “in-

ferior turbinal” (7. ¢b) is almost entirely soft. The inferior part of the labyrinth, in

front, has the very constant endosteal patch of bone; it has already ossified the long

“preenasal”’ style (which afterwards becomes blunt, Plate II. fig. 1, p.m); laterally

(fig. 7) it is affecting the “ trabecular cornua” (ce. ¢7); and behind it runs along the fore -

half of the septum below ; the rest of the septum below, or rather its trabecular addition,

has the usual infero-posterior osseous centre (¢7, s. 2), which is attached to the right ossified

labial cartilage. Binding the “inturned lamina,” below is the sessile, leafy ‘‘ maxillo-pala-

tine” plate; and undergirding this is the contiguous part of the palatine (pa), whose

interpalatine process (ipa), far forwards in position, ties itself to the outer and under

surface of the 4th and 5th septo-maxillaries. The rest of the parts of this most re-

markably complex facial basketwork have been described already; and the reader will

now be able to follow me whilst I give the details of other species of Woodpeckers.

The first of these exhibits the “ vomerine cartilages,” but unossified, and is only to be

understood by references to the condition of these parts in various Passerine birds. These

references will be given in due time and place.

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 13

On the Palatal Structures of Hemilophus fulvus.

This Celebesian species, from Mr. Salvin’s collection, is about the size of our native

Gecinus viridis; it presents some modifications of great interest. In many respects the

description of the last species would serve for this; but the practised eye soon detects a

number of modifications. The rudimentary basipterygoids (Plate III. fig. 1, b.pg) are

sharper and more divergent; the part answering to the “os uncinatum” (0.2) has less

suborbital development, but is more distinct in outline. The anterior border of the

“«meso-ethmoid ” is somewhat sharper than in G'ecinus; the nasal septum and the rest of

the labyrinth are ossified to nearly the same degree. The three palatal bands of the pre-

maxillary (fig. 1 pa) are all fused together by bony deposit; and this excess of ossifica-

tion has filled in the angle between the side of the palatine and the end of the pre-

maxillary palatine plate. Opposite the sharp interpalatine spurs (¢.pa) there is a

mass of cartilage, not seen, in this form at least, in any dissection of mine, in the other

“ Picide.” This is evidently two flat plates that have coalesced at the mid line (fig. 2,

l. €), and then, becoming carinate below, send forwards a blunt, and backwards a sharp

spur. The septo-maxillary bones are strongly attached to, but are not grafted upon, this

cartilage. Here we have the vomerine cartilages of the Wren, developed only on their

front portion, not metamorphosed into part of the vomerine bar, but distinct as in the

Rhea. As to the morphological meaning of this double cartilage, I am fairly satisfied

that it answers to a pair of upper labials. In front of the shoulders of this cartilage

there is, on each side. a falciform septo-maxillary; it reaches to where the premax-

illary style has coalesced with the inner edge of the palatine. Behind the broad end of

the right-hand style there are four bony grains; on the /eft side there is one larger and

one smaller grain. All these are strongly tied to the interpalatine style, to the labial

cartilage, and to the rounded fore end of each true “vomer.” These latter bones (v)

reach from the septo-maxillary grains to the short snags of the “ ethmo-palatine” (e.pa).

They are bowed inwards, as in the embryo of the Passerines, have a thick lower and a

crested upper edge; they are free in front, but anchylosed behind.

Between these, behind, I find one “median septo-maxillary’’ (m. s.mx) attached to

the base of the free edge of the ethmoid. It is probable that the larger posterior septo-

maxillary on the left side was once three grains of bones, and that each vomer ossified

from-two points, as in Gecinus. Anyhow this is as curious a piece of facial morphology

as any to be seen in the whole range of the vertebrata, and makes the observer very

cautious of predicting what he shall or shall not find. The palatine arch is, on the

whole, so closely like that of the green species, that it requires no fresh description; the

small “ palato-maxillary ” (p.mz) on the /ef¢ side is very similar to that of Gecinus.

On the Palatal Structures of Picus analis.

Mr. Salvin’s rich Neotropical collection has supplied me with a Javan Woodpecker

intermediate in size between our native P. minor and P. major. In Picus analis the

fore edge of the “‘basitemporal”’ plate (Plate ITI. fig. 3, 0.¢) is elegantly trilobate, the

outer enlargements being partly due to the presence of rudimentary basipterygoids.

14 MR. W. K. PARKER ON THE MORPHOLOGY OF

The “rostrum” of the “ parasphenoid”’ (pa.s) is rather short, and glides in the pterygo-

palatine groove. Its pointed end comes short of the anterior sharp edge of the ‘‘ meso-

ethmoid.”

The nasal septum (s. 7), ‘‘ anterior and inferior turbinals” (al.td, 7. tb) are well ossified,

as in Hemilophus ; these are only faintly indicated in the figure, to give prominence to

the facial arches.

The median palatine part of the preemaxillary (px) lies above the lateral palatine

plates (p.pz), so as to form a rounded sulcus; the latter have their retral tips anchy-

losed to the vomerine bar. The dentary regions of the preemaxillary (d. px) run con-

tinuously into the maxillary bar (mz).

The palatine process of the preemaxillary is connected with the stunted feeble ethmo-

palatine plate (e.pa) by a delicate, sinuous, rugose band of bone, which gradually broadens

backwards (figs. 3 & 4, p.px, s.mx,v). Here, whatever number of ossicles existed in the

growing bird, in the adult are all undistinguishable; the only distinction between septo-

maxillary and vomer is in certain irregularities in this frail bony bar. Even here a

“median septo-maxillary”’ (m. s.mx) bounds the lower outline of the “hinge:” it is

fixed at the septal end.

The pterygoids (pq), each with its large muscle-snag and scythe-shaped “‘ mesoptery-

goid”’ process, are quite normal, as also are the palatines, which, however, have deve-

loped a gnawed transpalatine rudiment (¢.pa). The “interpalatine” spurs have the

same irregular eroded appearance; the ‘“‘mediopalatine” lozenge (m.pa) is imbedded

in more cartilage than in Hemilophus.

The “maxillo-palatine” processes (ma.p) form merely a small subtriangular lip to the

inner edge of the main maxillary bar (mx), and look broader than they are, because of the

outbowing of the bone on the dentary edge, where the angle of the mouth is. Small as

the ingrowth is, yet it is scooped below, to form an air-passage to the body of the bone.

All trace of suture in the jugal bar is gone, as in the other species. Here we come

athwart the most developed specimen of the odd “palato-maxillary ” bone (p.mz) ; it is

a wedge-shaped plate, strongly attached to the overlying “alinasal wall,” with its

enclosed turbinal. The “os uncinatum”’ (fig. 3, 0. uw) has a semidistinct appearance,

being well marked out at the angle of the large well-ossified “pars plana.” The la-

crymal is, as in Gecinus and Hemilophus, a smallish semioval plate, which becomes

anchylosed to the hinder edge of the descending crus of the nasal, as in Thrushes. tp

On the Palate of Picus major.

The description of P. analis may serve also for this for the most part; yet this is a

stouter kind, and the palatal bars are straighter and stronger (Plate I. fig. 1). The

specimen from which the drawing was taken, the gift of Dr. Murie, was rather young,

and it had lost the “ palato-maxillary” and the “mediopalatine” by maceration ; its

less-developed condition gives a softer state of the ossifying nasal labyrinth. Yet

already the vomerine series had become fused together and to the preemaxillary and

palatine bones (s.ma, v, p.pa, e.pa). This species has a free “interpalatine” spur,

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 15

and a more decided “transpalatine” angle; and the “os uncinatum’’ is much more

distinct than in P. analis; their “ maxillo-palatine” lobes are about equal. i

The main body of the palatine is more deeply grooved than in Gecinus; in this it

agrees with P. analis; the “ethmo-palatine” hook shows some distinctness from the

end of the vomer. The absence of a “median septo-maxillary” is probably due to

maceration.

On the Palate of Picus minor vel striolatus.

Some years ago Mr. Bartlett put into my hands two ripe nestlings of this small

species; these were immediately dissected, and have been greatly valued because of the

distinctness of their sutures.

A palatal view of one of these skulls (Plate IV. fig. 1) well serves to illustrate the

structure in Picus proper (Plate IV. figs. 1 & 2). In front of the very wide “ basi-

temporal” plate, with its median beak, we see a pair of very arrested “ basipterygoids”’

(d.pg) on the wide-rooted, short, sharp-pointed “parasphenoid” (pas) which underlies

the massive “‘meso-ethmoid.” In front of the “hinge” the nasal septum (s. 2) is still

cartilaginous to a great extent, although a bony tract from the trabecular cornu has run

backwards beneath it. The fore part of the nasal labyrinth is still cartilaginous ; but the

“pars plana” behind has coalesced with the “meso-ethmoid.” The premaxillaries are

shorter here, and their palatine processes (p. px) curve inwards at their end, thus free-

ing themselves from the inner edge of the palatine. Immediately above the styloid

“interpalatine” processes*(7.pa) there are seen three “septo-maxillaries” (figs. 1 & 2,

s.mx), one on the right, larger, and two on the left, lesser ossicles. Their clavate ends

are attached to the clubbed fore end of the subarcuate vomers, which are suddenly fili-

form, and then expand into a twisted crest, where they join behind to the broad,

eroded, inturned “ethmo-palatines” (e.pa,v). The palatines are very broad and thin,

and have an obtuse “transpalatine” angle (¢.pa). Their upper plates are uniting by

synchondrosis ; and a small “ medio-palatine ” centre has appeared on the left side, in front

(m.pa). The terminal part of the palatine is sharp and notched, and is overlain by

the large falciform “‘ mesopterygoid process.

The pterygoids (pg) are more slender than usual; and the muscle-snag above is short

at present. The “os uncinatum” (0. w) is represented as yet by the unossified condition

of the “ecto-ethmoidal” angle. A small cuneiform “ palato-maxillary” (p.pr) has

appeared in the angle between the palatine and maxillary ; this latter bone sends inwards

a small subtriangular “maxillo-palatine” lip (mx.p), scarcely larger than, and not so

distinct as, the smallest process of the kind seen in the Bird Class, namely, in Wenura

superba. I may mention the large “orbito-sphenoids” seen on each side in these

specimens, as they indicate a more general character of the bird than we have in the

Crow form, in the higher types of which, at least, these do not appear.

The lacrymal is, at present, a very small, short thread of bone behind the nasal.

The jugals (7) are quite distinct from the zygomatic process of the maxillary.

16 MR. W. K. PARKER ON THE MORPHOLOGY OF

On the Palate of Picumnus minutus.

This Woodpecker, from Bahia, Brazil, is about the size of our native Golden-crested

Wren; it leads on to the lesser group, or the Wrynecks, having soft tail-quills. In

several respects it is of extreme interest, especially as relating to the Passerines of its

own geographical area, namely those low forms of the Western Notogea the “ Cotin-

gide” and “ Formicariide.” J have had to work out the parts of this bird’s palate

from the fractured skull in a dry skin, kindly given me by Osbert Salvin, Esq. Labour of

this kind was never more richly rewarded.

The nasal septum (Plate V. figs. 3 & 6, s.) is semiosseous; the rest of the labyrinth

is principally persistent cartilage. The upper anterior and posterior regions of the

septum are all occupied with one continuous bony tract, which also runs into the ale

of the lower and trabecular region.

Between and behind these wings there are three centres of bone, the last of them

being the familiar postero-inferior bone of the septum, a subterminal trabecular centre.

The cranio-facial “ notch ” is not perfect, and the ossified septum and meso-ethmoid (p.e)

are articulated together.

A distinct ‘“ septo-maxillary ’”’ (s.mx) could be seen on the left side, and in front of it

a thickening of the same nature. The right bone had coalesced with the inner edge of

the palatine, that part which receives increment from the palatal process of the pre-

maxillary.

The vomers (figs. 4, 5, v) retain their embryonic (“ Rhynchosaurian’’) condition, as in

the young of Gecinus viridis (Plate ITI. figs. 5, 6, v), being arcuate “horns” attached

to the mesopterygoid processes of the pterygoid (pg). These latter processes are normal,

but shorter than in our native species; and the rest of the bone is similar to that of

Picus minor, but stouter.

The palatines, as seen from below (fig. 3, pa, pt.pa), struck me at once as being sin-

cular, the postpalatine flanges not being bevelled off, as in the other kinds, but largely

developed as in the ‘“‘ Formicariide”’ and “ Cotingide”’ amongst the Passerines of South

America*, and notably also in the Australian Lyre-bird (lenura superba). In these birds,

however, Thamnophilus, Pachyramphus, &c. these side walls of the posterior narial tunnel

are continuous with the rest of the palatal ossification; but in Picwmnus (Plate V. fig. 5)

they are distinct laminee of bone, each with a down-growing ala.

These “postpalatine” bones end in front behind the “interpalatine’’ spurs, and

below the “ ethmo-palatine” laminee, which here, as in the Green Woodpecker, are feebly

developed. ‘The interpalatine spurs are well developed, and are more normally placed,

further backwards, than in the “ Picide” generally. The rest of the palatine is quite

like that of its congeners. The maxillaries (mx) have a very rudimentary maxillo-

palatine process (ma.p); and the separate “ palato-maxillary ” (p.mzx) on the eft side is

represented by an enlargement of the inner edge of the bone, which was once, most pro-

bably, quite separate.

* T cannot give illustrations of these Passerine structures here ; they are freely given in the first part of my paper

on the “ Aigithognathe,” about to appear in the ‘ Zoological Transactions.’

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 17

_ The jugals (7) are slender, save at the quadrate end, where they are very solid and

inhooked.

Altogether this small far-western type is extremely instructive, and helps to lead the

imagination down to extinct types in which the characters of the Hemipod, the Low

Passerine, and the Woodpecker were existent in one generalized form—a form and a type

only a step or two above the raft-breasted Ostrich tribe,

On the Development of the Face in the Subfamily Yungide: example

Yunx torquilla.

We now come to the feeblest members of the “ Celeomorphe ;” yet the Wrynecks

are stout little birds, with the same restless energetic character as the larger types.

The possession of several nestlings of this form, the gift of Mr. Dines, has enabled me

to work at a still earlier condition than that of the youngest specimen of Gecinus

viridis.

The palate (Plate IV. fig. 3) shows that these birds are true Woodpeckers, and have

no special affinity to the Crow and its allies.

Here we have the same form of occipital condyle (0. ¢), the same massive broad “ basi-

temporal”’ plate (4.2), the rudimentary “ basipterygoids”’ (b.pg), and, what is very instruc-

tive, an explanation of the very remarkable cowrie-shaped tympanum seen in those types.

The “ basitemporal”’ forms a large retral hook where it joins the “exoccipital” (e.0),

and is then roundly scooped at its edge. Here the lobular growths of the adult are seen

to be due to the presence, of a ‘‘ tympanic chain;” three of these bones are now to be

counted on each side (Plate IT. fig. 1, as compared with Plate IV. fig. 3, ty).

In Yunex the “ parasphenoidal rostrum” (pa.s) is much longer than in Pécus, and the

face much shorter, especially the preemaxillary (px). This is most remarkable when

compared with Gecinus and Hemilophus: something is due, however, to age; for these

specimens are very young.

The solid part of the premaxillary is very short indeed; but the three main bars

are long and extremely slender (d.px, p.px); they have the same relations as in Picus.

The nasal septum, with its thick trabecular base, is still unossified, and indeed scarcely

hardens at all in this type, nor do the turbinals. Here the preenasal cartilage (fig. 3, p.2)

is still large.

There are no septo-maxillaries at this stage; nor am I aware of their existence in the

adult*. The “ vomers,” however, are very noticeable. At first I expected to find them

partly preformed in hyaline cartilage, as in the ‘‘ Coracomorphe,”’ especially as Hemi-

lophus retains in its adult state its large or prevomerine labial plate. It is not so; they

arise, as in Gecinus viridis, in a band of soft connective web on each side of a remarkable

recess in the mucous membrane of the nose, the azygous counterpart of those paired.

recesses called in Man the “ sphenoidal sinuses” (Plate IV. fig. 11, v, m.sph).

Below the vomers is the palatal mucous membrane (Plate IV. figs. 10, 11, v). Seen

from below (fig. 4, v) they are remarkably like those of a young Passerine, being pointed

es The adult skull may be seen in the macerated condition in the College of Surgeons ; it was prepared by me with

the other bones, and drawn thirty-two years ago.

SECOND SERIES.— ZOOLOGY, VOL. I. D

18 MR. W. K. PARKER ON THE MORPHOLOGY OF

behind, broad and obliquely premorse in front. Outside they are overlapped by the long

ethmo-palatal style (e.pa), whilst the filiform very elongated mesopterygoid processes

(m.pg) nearly reach them behind. Here the “ parasphenoidal rostrum” (pa.s) ends,

emarginate and toothed, whilst in front the vomerine pieces are tied to the nasal wall by

a band of membrane continuous with that which forms a commissure to the two bones.

In front of the vomers this membrane thickens at the mid line, being scooped submesially

for nervesand vessels. This thick alate tract of fibrous tissue, however, is the symmorph

of the cartilaginous mass in Hemilophus (Plate III. fig. 2, 7. c); and the fibrous bands in

which the vomers are developed are the degraded counterparts of the free labial cartilages

of the Rhea, the Hemipod, and the “Coracomorph.” The whole “ectoethmoid” (p.p)

is still soft; but the “ trabeculo-palatine”’ conjugational lobe is marked out (0.w).

The inbent clinging pterygoids are thoroughly Picine, with their exceedingly large and

long mesopterygoid (m.pg), and having no “ epipterygoid”’ spur. They are underbound

by the filiform out-turned ends of the palatines (figs. 3 & 4, pa), which are slightly

grooved in their gradually broadening wider part. This region forms a very obtuse

angle, the Passerine transpalate being quite gone. They differ in this from Picus.

The interpalatine and ethmo-palatine styles are equally long and slender (i.pa, e.pa) ;

and the prepalatine bar is filiform and of great length. I have not been able to find a

*‘ mediopalatine.” The maxillaries (mx) have an extremely small palatine lip (mz.p)

opposite the thick angular process; the jugal process is very long, and is separate

from the jugal (j). The palatine edge of the maxillary is more abortively developed on

the left than on the right side; but the deficiency is supplemented by the palato

maxillary ossicle (p.mz).

A series of sectional views will illustrate the relations of the nasal sacs to the rest of the

face. The first of these (Plate IV. fig. 5) is through the body and fore part of the palatine

process of the preemaxillary and the preenasal cartilage (pa, p.px, pn). The next (fig. 6)

cuts through the fore part of the alinasal cartilage (al.7), the alinasal turbinal (al.¢0), the

fore part of the septum with its trabecular base (s. 2), and the related parts of the facial

bars (.px, pa, mx, p.px). The next section (fig. 7) shows that the alinasal turbinal

(al.tb) is coiled upon itself once and a half, whereas it is scarcely coiled once in

Gecinus (Plate I. fig. 7, al.tb). In this section the upper and lateral parts of the pre-

maxillary, the maxillaries, the two crura of the nasals, and the preepalatine bar are cut

through: here the septum (s. 7) is at its greatest depth. Further back (fig. 8), behind

the maxillary angle, the inferior turbinals have come into view (i. td). They do not finish

one coil, and are scarcely more developed than in Gecinus (Plate II. figs. 7 & 8,7. 6). The

“ alinasal turbinal”’ (ad. tb) is there also behind its root; it has lost most of its coil, but

has developed an inner outgrowth. This section is through the notched part of the

cranio-facial axis. Still further back, behind the alinasal turbinal, and behind the notch,

we come to the thick-topped shelving “ mesoethmoid” (p.e), with its bony matter; the

alee above are the “ ali-ethmoids” (a/.e). Here on each side of the lower nasal nerves

there is a large flap of the nasal fioor, which reappears behind. The “ali-ethmoids’’ grow

down and form a subvertical wall, from the lower part of which the posterior extremity

of the inferior turbinal is growing. Above this there is a new scroll. All these scrolls

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 19

begin above; the one in front of this is the inturned and flattened fold which represents

the upper turbinal (w. ¢b). Here the inferior turbinal (i. 42) sends out an inner out-

growth like the one in front of it.

The middle turbinal is a mere ridge in the front of the pars plana, where the inferior

turbinal ends (see in Gecinus, Plate V. fig. 2, m.¢b). When the razor has cut through

the thickest part of the rapidly ossifying “‘ mesoethmoid,” we get a view of the orbital face

of each “ pars plana” (Plate IV. fig. 10, pe, pp) with its conjugational angle corresponding

to the ‘os uncinatum.”’ It is in this section that we get a view of the ‘‘ median sphenoidal

sinus” (m.sph) and the two vomers; (see enlarged view, fig. 11) in this region; both

endostosis and ectostosis are at work, hardening the “ perpendicular plate,’’ which grows

outward into the “ ali-ethmoids,” which, infolding themselves and becoming pinched

at the fold, give us ultimately the ear-shaped wavy crenate upper turbinal (compare fig. 9,

with Plate V. fig. 2, w. tb).

Already I find a small styloid lacrymal behind the nasal crus, and a distinct centre in

the orbito-sphenoidal region.

Summary of characters of the ‘‘Saurognathous”’ type of Palate.

A. Trabecular Arch.

1. Retention and ossification of trabecular cornua, and of basitrabecular (przenasal)

rostrum.

2. Addition thereto, in certain types, of a pair of “ labial” cartilages, which do not

form the foundation of an oxhead-shaped compound vomer as in the Passerine.

3. Detachment, in Gecinus, at least, of part of the nasal wall, after the ingrafting on

it of a septo-maxillary.

4.. Great number and bilateral independence of the vomerine series of bones, some

of which are azygous.

B. Palatine Arch.

1. Absence of a distinct mesopterygoid, the process answering thereto being very

long, and in the embyo reaching the vomer of that side, as in the New-

Zealand Lizard Hatteria.

2. A dagger-shaped basipalatine between the right and left bones.

3. Absence of a distinct “ transpalatine.”

4, Presence of a postpalatine bone in the most generalized form (namely, Picumnus).

5. Absence of epipterygoid hook, and presence of a large spur for muscular attach-

ment on the pterygoid.

6. Abortive development of maxillo-palatine plates, and presence of a distinct palato-

maxillary bone on the left side only.

C. Hyoidean Arch.

1. Abortive development and early fusion of the ceratohyals.

2. Double “ infrastapedial ” and ossified stylohyal.

20 MR. W. K. PARKER ON THE MORPHOLOGY OF

D. Third postoral arch.

Length and tenuity of basal piece and upper and lower pieces of the cornua caynes

hyals, or 1st cerato- and epibranchials).

E. Large size, but great simplicity, of nasal turbinal folds.

F. Cowrie-shaped tympanic cavity,

Concluding Remarks.

The “ Celeomorphe ” of Huxley form a most natural and well-defined group—a group

equal, zoologically, to the Pigeons or the Parrots. Evidently this differentiation has

taken place through the gradual extinction, during long secular periods, of conjugational

types more generalized than those now extant. Iam not without evidence as to the

truth of this view; but it must be discussed elsewhere ; yet proofs will be given in proper

time and place.

Those who are familiar with the Lacertian type of skull will agree with me as to the

propriety of such a morphological term for the Woodpeckers and Wrynecks as ‘“ Sau-

rognathe.” It will exactly fill up the lacuna left by my friend in the masterly paper

already referred to. The view there expressed that these birds have a Passerine foun-

dation, but that they are somewhat abortively developed, arrested one way and wonder-

fully specialized in another, will be seen to be the exact truth of the matter.

Corresponding with what I am showing in papers for the ‘ Zoological Transactions,’

with regard to the “ Aigithognathe (Passerinz,’ “ Coracomorphe”’), the most embry-

onic and least specialized Woodpecker is a South-American form, namely Picwmnus.

We have seen how numerous the bony centres are in this group; and it is not easy to

account for this peculiarity. They are very numerous in all birds, but least so in the

Thrushes (“Turdidz’’); whilst in the “ Coracomorphz” generally there is an average

development as to numbers. There is some reason for these differences ; but it is hidden

amongst the extinct forms, both warm- and cold-blooded.

Teleologically, the “ Picidze’”’ cannot possibly benefit by the numerous small bones

that havea temporarily separate existence in their palate, many of these being mere grains,

and others most feeble splints.

The general agreement of the palate of Picuwmnus with that of the last of the Rhyn-

chosaurian Lizards (namely Hatteria) is very suggestive. In the dow South-American

Passerinz, the “‘ Formicariide”’ and the “Cotingide,” the essentially Reptilian face

shows itself most clearly. .

But that which characterizes the whole of the ‘‘Celeomorphze”’ is the want of fusion

of the parts of the palate at the mid line. This is evidently an ancient reptilian

“atavism.” This must be admitted in a metaphorical sense, if not in an actual: it is a

remarkable verisimilitude, if not a real historic truth.

Fig.

THE SKULL IN THE WOODPECKERS AND WRYNECKS.

DESCRIPTION OF THE PLATES.

Puate I.

1. Palate of Picus major, adult. x 4 diameters.

2. Palate of Gecinus viridis, oldest fledgling. x 24 diameters.

3. Vertical section of skull of G. viridis, younger fledgling. x 24 diameters.

4. Side view of face of ditto. x 24 diameters.

Ist section of ditto. x 5 diameters.

. 2nd section of ditto. x 5 diameters.

. 8rd section of ditto: x 5 diameters.

. 4th section of ditto. x 5 diameters.

. Part of tongue of youngest Gecinus (lower view). x 10 diameters.

10. Part of ditto (upper view). x 10 diameters.

11. Stapes of adult Gecinus. x 74 diameters.

mn Oo BB & OD

Puate II.

Gecinus viridis (continued).

. Palate of adult. x 24 diameters.

. Part of same. x 5 diameters.

Septum nasi. x 4 diameters.

. Vomerine bones (outer view). x 4 diameters.

. Vomerine bones (inner view) x 4 diameters.

Palate of specimen six months old. x 4 diameters.

. Part of nasal capsule of ditto. x 4 diameters.

. Part of fig. 6. x 8 diameters.

Prats II.

. Palate of Hemilophus fulvus (adult). x 24 diameters.

. Part of same. x 5 diameters.

. Palate of Picus analis (adult). x 5 diameters.

. Part of same. x 10 diameters.

. Part of palate of Gecinus viridis (older fledgling) from above. x 43 diameters.

. Ditto (younger fledgling). x 44 diameters.

. Pterygoid and part of palate of adult Gecinus. x 24 diameters.

Puate IV.

. Palate of Picus minor (fledgling). x 6 diameters.

. Part of same. x 12 diameters.

. Palate of Yuna torquilla (nestling). x 54 diameters.

. Part of same. x 11 diameters.

ON THE SKULL IN THE WOODPECKERS AND WRYNECKS.

5. Ist of a series of vertically transverse sections of face of Yunx torquilla. x 8 diameters.

6. 2nd section of ditto. x 8 diameters.

3rd section of ditto. x 8 diameters.

. 4th section of ditto. x 8 diameters.

. 5th section of ditto. x 8 diameters.

. 10. 6th section of ditto. x 8 diameters.

.11. Part of same. x 24 diameters.

PuatTe V.

. Side view of skull of Gecinus viridis. x 2 diameters.

. Fore view of ethmoid of ditto. x 2 diameters.

. Palate of Picumnus minutus (from below). x 10 diameters.

. Part of ditto (from above). x 10 diameters.

. Part of dito (side view). x 10 diameters.

. Side view of septum nasi of Picumnus minutus. x 10 diameters.

. Diagram of visceral arches and sense-capsules in the embryo of an aérial bird at the middle of

incubation :—o/, nasal sac; e, eye; au, auditory sac; tr, trabecula; p.n, prenasal; c. tr, tra-

becular cornu; pa, palatal; pg, pterygoid; g, quadratum; mk, Meckel’s cartilage ; st, stipes ;

5, 5th nerve; s.st, suprastapedial; e.st, extrastapedial ; i.st, infrastapedial ; st.h, stylohyal ;

c.hy, ceratohyal; bh, basihyal; uh, urohyal; br. 1, 1st branchial.

Trans.Linw Soc.,SER.2.ZooL.Vor.1 Tas. |.

WK. P. ddl.ad nat; GWeet lth,

WWest &Co. imp.

a Traws.Liny.Soc.,Ser.2. Zoot Von.1. Tas. II.

\, WE Padel.ad nat; GWest lth.

j ff . 2 . «

7 Gecinus viridis.

W.West &Co. imp.

‘ Trans.Linw. Soc. Ser. 2. Zoou.Vou..Tas. 3.

WK. P. del. ad nat. C.West ith.

' ines Ae &. Hemilophus fulvus. Figs 3,4. Picus analis, Figs 5-7. Gecinus viridis.

W. West & C? imp!

Tears) Lin Soe. SER 2.Zoor. Vow.|. Tas 4

a Cana

SST

RAE LINSS

WEP. del. ad nat.G.West lith

j Figs 1,4. Picus mmor. Fig? 3—11. Yumx torquilla. WiWerhe: C2 eue

Trans. Linw.Soc.Ser.2.Zoor Vor. Tas. 5.

WK PR. del ad nat G. West lith

: - Oe ER : f W. West & C° imp

Fige1,2a@- Gecinus viridis. Figs 3—6. Picumnus minutus.

[ 23 |

II. On some Atlantic Crustacea from the ‘ Challenger’ Expedition. By Dr. R. v.

WILLEMOES-SuuM, Naturalist to the Expedition. Communicated by Prof. WYVILLE

THomson, F.L.S. & L.S.

(Plates VI-XIII.)

Read May 7th, 1874.

AMONG the great quantity of deep-sea Crustacea which have been. brought up during

our cruise in the Atlantic, either by the dredge or the trawl, I have selected the most

interesting ones for immediate description. Besides, readers will find in this paper the

description of both sexes of an interesting Nebalia from the shallow water of the

Bermudas, some remarks on the structure of Cystisoma, and a note on the develop-

ment of a Land-Crab from the Cape-Verds. Some of the deep-sea forms, the Wille-

moesie (Deidamie), Astacus, Zaleucus, and Gnathophausia, have been already figured in

‘Nature,’ in the “* Notes from the ‘ Challenger’”’ published by Prof. Wyville Thomson ;

and a short description, taken from my first notes on the subject, has been added. In

this paper it is now intended to give a more detailed account of their structure and their

systematic position. The paper is divided into the following seven parts :—

I. On a blind deep-sea Tanaid.

II. On Cystisoma Neptunus (Thawmops pellucida).

III. On a Nebalia from the Bermudas.

IV. On some genera of Schizopods with a free dorsal shield.

V. On the development of a Land-Crab.

VI. On a blind deep-sea Astacus.

VII. On Willemoesia (Grote), a deep-sea Decapod allied to Eryon.

I have only worked out such species as, from their structure and zoological affinities,

present a greater interest or belong to groups about which we had a satisfactory lite-

rature on board. Should, nevertheless, a want of literary information be remarked in

some point or other, the reader is asked kindly to remember that this paper was worked

out on board a ship cruising in the Atlantic.

H.M.S. ‘ Challenger,’

Cape of Good Hope, November 1873.

I. ON A BLIND DEEP-SEA TANAID. (PI. XII. figs. 1-9.)

During our cruise in the Atlantic, members of the Anisopoda tribe were found twice.

First, on the 2nd of May a real Tanais was got from a depth of 1700 fathoms off the

North-American coast; I cannot determine whether this animal belongs to a known

species or not. Later, when cruising among the Azorean islands San Miguel and Santa

24 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

Maria, we got from a depth of 1000 fathoms another Anisopod, which belongs to the

interesting genus Apseudes, according to Mr. C. Spence Bate* the only Isopod in which

the antennal scale, so common in the Macrura, is present on the lower antenne, and, as

we may add, one of the few in which there are two flagella in the anterior antenna.

In the species of Apseudes hitherto known the eyes are pedunculated: in this deep-sea

animal there are no eyes at all; hence we might call it 4. ceca. It has a length of

6 millims., the females having the same length as the males.

The cephalothorax, to which is attached the first pair of gnathopods, has a spiny

rostrum and two prominent spines on each side. The first antenne (fig. 1, a’, and fig. 2)

show a very strong funiculus and two flagella, one of which consists of twenty-four, the

other of eight joints. The longer is double the length of the smaller one.

- The exterior or second antennee (fig. 1, a’, and fig. 3) have a smaller flagellum, con-

sisting of eight joints and an antennal scale articulating with the first joint of the

funiculus (fig. 3, s). The mandibul have a palpus consisting of several joints.

The maxilliped (fig. 9) has strong tufts of hairs on the last three joints. The two

enathopoda are very different in form: the first one (fig. 4) is short, and terminates

in a strong recurved chela; while the second is much longer (fig. 5), and has enlarged

and flattened joints with strong spines, as may also be seen in the other species of

Apseudes. The pereiopoda are strong but not very much enlarged appendages. We

have drawn the first and second as well as the top of the fourth; the third is terminated

by a bifid claw, as is the sixth in A. falpa, Mont., which we have not drawn, as only

once could we clearly get sight of it. When afterwards it was tried again to get a view

of the two claws, it was impossible to separate them from each other.

The first five segments of the pleon are short, and possess small biramous pleopoda.

The last segment is elongated, as in 4. talpa, and shows at the top two pairs of caudal

sete.

In the females there are breeding-lamelle at the base of the second gnathopoda and

the first two pereiopoda, which are wanting in the males. -This is the only difference

which I could find out between the two sexes.

Most of the specimens which we got were washed out from the mud which the. dredge

had brought up. We did not meet with this Crustacean again.

_ This is not the first instance of blind Isopods being described, as some subterranean +

and parasitical { forms are well known to have no eyes, or to lose them, as Anceus does,

when getting old. There is also a whole family, the Munopside, which is blind, though

most of its members live in moderate depths. A very large member of this family

has also been got by us in the deep sea; it cannot, however, be described now, as the

necessary literature is not at hand.

II. ON Ovsrisoua Neprowvs (Taavmors petrrvcrpa). (Pl. XI. figs. 4-8.)

In a postscript to my paper on Thawmops pellucida (Phil. Trans. of the Royal So-

ciety, 1873) I have described the male of this remarkable Amphipod, and given some

* Facts and Arguments for Darwin, by Fritz Miller, translated by Dallas (London), 1869, p. 132.

+ Pitanethes albus and Lyphloniscus Steinir. ; + Bopyride and the female of Anceus.

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 25

details on the structure of both sexes, which could only be discovered when it was

possible to dissect one of the specimens. I now have to publish some figures which

could not be given in my first paper, and add a few words to them. But before doing

this, I must declare that the name of Thaumops pellucida must be withdrawn, as this

animal has been already described under the name of Cystisoma Neptunus by Guérin-

Méneville*, from a single specimen caught in the Indian ocean. I see this from Spence

Bate’s ‘Catalogue of Amphipodous Crustacea’ +, which I have just received, and which

contains a copy of Guérin-Méneville’s rather bad figure and a short description of a

specimen, which probably is a male.

Guérin-Méneville (and Spence Bate?) regards Cystisoma as belonging to the Hyperide

family, and takes the antenne (which are in front of the head) as representing the

second pair. Why they should be homologous with these I do not know; nor do I think

they are; for in Phronimids, Oxycephalids, and Hyperids the second antenne have a

funiculus consisting of several (more than two) joints. Now in Cystisoma the antennze

have got two joints, which seem to represent the one a funiculus joint, and the other the

flagellum of the first antennze of Phronima.

Against a union of Cystisoma with the Hyperids may be advanced, besides the form

of the head (which is more Typhid-like) and the absence of the second antennz in both

sexes, the absence of a palpus on its mandible (Pl. XI. fig. 6). The palpus is always

present, according to Claus, in Hyperids, but is wanting in Phronimids. Besides, there

are so many peculiarities distinguishing Cystisoma, which I have enumerated in my

above-mentioned paper, that I do not think it possible to keep it in one of the hitherto

received families, but that now, when we know its structure better, we have to establish

for it a special family called Cystisomide.

The first maxilla has been described, and is now figured in Pl. XI. fig. 7. The second

maxilla could not be found, and is probably represented by an organ which might have

arisen from the union of the second maxille and the under lip (Pl. XI. fig. 8). This

organ (7) is attached to the third joint of the reduced maxillipedes, and surely acts phy-

siologically as an under lip. The maxillipedes are reduced and fused together so as to

form one organ consisting of two basal joints and two claws (fig. 8, ™), between which is

attached the organ which we suppose to represent the maxillz and the under lip.

The male differs by the absence of glands at the top of nearly all the appendages,

especially in the last pair of pereiopoda, which, according to this, have not the same

clumsy appearance as in the female. The two testes begin just behind the stomach

(fig. 5,7), and send vasa deferentia to the last segment of the pereion, where two simple

genital openings are to be seen between the last pair of legs (fig. 5, a g).

Probably (as in Phronima) the full-grown male is somewhat smaller than the female,

it seems that Cystisoma Neptunus can attain a very considerable size; for the last

and largest male which we got in the trawl has a length of 108 millims. It seems

that these oceanic amphipods have a very wide geographical distribution. Cystisoma,

as we have seen, was first described from the Indian Ocean; and Oxycephalus piscator is

* Guérin-Méneyille, in Revue Zoologique, July 1842, p, 214. 7 Page 311, pl. 1. fig. 7.

SECOND SERIES.—ZOOLOGY, VOL. I. £

26 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

mentioned as occurring in the same. Now we have found in the Atlantic not only four

specimens of Cystisoma, but also numerous specimens of two species of Oxycephalus, one

of which seems to me to be identical with O. piscator.

III. On a Nezzazrs FROM THE BERMUDAS. (Plate VI.)

During our stay in the Bermudas Mr. Murray collected one day some small Crustacea.

in Harrington Sound, a bay which only communicates with the sea through a narrow

passage. Among them I was very glad to find a Nebalia, and went then myself to the

bay, where this interesting little Crustacean was not rare under stones and corals. I

collected, perhaps, twenty females and two males, and was able to make some observa-

tions on them which, added to the embryological researches of Mecznikoff and the

anatomical remarks of Claus* on both sexes of the Mediterranean N. G'eoffroy?, will, I

think, enlarge somewhat our knowledge of these Crustacea, which for such a long time

have been considered to be Phyllopods, but, according to the above-named papers, are

more rightly placed among the Schizopods.

The female of this Nedalia, which I am going to call N. longipes, has a length of

6 millims., and is, when living, of a milk-white colour. The shell, posteriorly and above

somewhat flattened, has an oval form; it terminates anteriorly in a sharp and slender

rostrum, which covers the pedunculated eyes; the upper surfaces of these eyes are entirely

covered with small spines, so that the perception of the light is only possible through

the facets of the under surface. The shell does not quite extend so far backwards as in

N. Geoffroyi, where it reaches the fifth abdominal segment.

The anterior antenne (figs. 1, 2, a, and fig. 5) consist of four joints, the last of which is

a denticulated claw (fig. 5, /), From the base of this claw start two flagella (fig. 5, a and

b), the shorter of which is transformed into a large squamiform appendage, fringed with

hairs; the other one is more slender and densely covered with olfactory hairs.

The posterior antenne (fig. 7) consist of three joints only, the last of which is densely

coyered on its protuberances with tufts of hairs, especially at the point of insertion of the

flagellum, which itself is divided into nine imperfect joints marked by tufts of hair.

A comparison of the mandibula and the two mazille (figs. 8-10) with the corresponding

parts in V. Geoffroy: shows that there are only very slight differences between them. I

saw the palpus of the first maxilla (“ Putzfuss,” Claus) in the same position in the male

as in the female, z.e. bent backwards.

More important are the pectoral feet; for on their account especially Nebalia was

formerly supposed to be a true Phyllopod. If this Bermudan species had been the first

Nebalia which had been noticed, I do not think that this would have occurred ; for in the

feet of this one the phyllopodal characters are very little perceptible. In fig. 3 I have

figured a pectoral foot of NV. Jongipes, and in fig. 4, for comparison, one of NV. Geoffroyi

(taken from Claus’s paper); so that the corresponding parts will be easily found out.

The two-eared branchial appendage of the basal joint (figs. 3 & 4, K A) is very large in WV.

Geoffroyi, and in N. longipes only rudimentary. The appendage of the second joint has

* Von Siebold und Kolliker’s Zeitschrift fiir wissenschaftliche Zoologie, vol. xxii. p, 323.

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 27

entirely lost its flattened form, and is a sort of leg, which, however, like the main branch,

is not jointed, but is densely covered with hairs on one side. When comparing such a foot

with the pereiopod of a true Schizopod, as, for example, Lophogaster (Sars, Beskrivelse

over Lophogaster typicus, tab. il. fig. 35), we see in both on the basal joint the bran-

chial appendages, which in Nebalia longipes are very simple, and in Lophogaster divided

and arborescent. From the second joint starts the palpus, which in our case has a

different shape from that of Lophogaster, but which morphologically is the same thing.

In the foot itself we find in Lophogaster well-defined joints and a terminating claw, in

N. Geoffroyt we still find the joints, but in our case we find no joints; in all three, how-

ever, it is not difficult to find out the corresponding parts.

The first four pairs of abdominal feet or pleopods (fig. 11) have a large basal joint den-

ticulated exteriorly, to which two rami are attached, the interior of which is longer and

has at its base a little appendage, which is by means of a denticulation at the top united

to a similar one of the other pleopod. It has been called by Claus, who found it in W.

Geoffroy, the retinaculum. Such appendages have been shown by Claus also in Stoma-

topods, and figured by Milne-Edwards from the Decapod genus Afya. The fifth and

sixth pair of pleopods resemble very much those in WV. Geoffroyi.

The two joints of the fwrca are covered at the top with tufts of long and dense hairs.

Among twenty to twenty-five females I discovered two or three males, easily recogni-

zable by their minute size and the enlarged flagellum of the anterior antenne, which here

are transformed into real prehensile organs. The males are only 8 millims. long, and

differ from the females also by being more slender, by their shorter pleopods and their

antennee, which are on the whole somewhat shorter and bigger. The pectoral feet are

exactly the same as in the females. In the males of WV. Geoffroyi the second pair of

antenne have an enormously long flagellum, something like the Cuma male; while in our

Nebalia the second antenna differs very little from that of the female, nor are the olfac-

tory hairs to be seen in much greater quantity, unless perhaps at the top and the outer

edge of the third joint. The first antenna (fig.6), however, is very different in the male ;

for its squamiform flagellum is greatly swollen up, and the joints are only to be discerned

as small protuberances with some hairs on them on one side. This conformation is

rather a phyllopodal one—so far as I can make out here, something like that found in

Estheria.

I found, like Claus, a ductus ejaculatorius directed towards the base of the eighth

pereiopod, but could not exactly see its opening.

This Nebalia differs from the two species hitherto known, by the spines on the upper

surface of its eyes, by the tuft of hairs at the top of the furca, by the form of its legs, in

which the phyllopodal character has been nearly entirely lost, and the schizopodal cha-

racter has been more approached than in any other species of this genus. in the male,

however, characters have been found which are wanting in the male of NV. Geoffroyi. We

see a modification of the first antenna reminding us of Astheria, but no prehensile organs

on any of the feet. This species therefore approaches, so far as the shape of its legs is

concerned, the higher Crustacea more than NV. Geoffroyi; but the prehensile organs of

its male are characters which show, again, that there remains a good deal of phyllopodal

28 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

element in Nebalia. We have now, however, discovered another Schizopod, in which the

legs are entirely normal and the shield is phyllopodal (Gnathophausia), but evidently

not to be derived from the Hstheria-like shell of Nebalia, but from the carapace of Apus.

And in this same Gnathophausia there is an abdominal segment which shows that, now

being essentially a Schizopod, it is in genealogical connexion with forms which have had

more segments in the abdomen than the Schizopods have.

These two forms, though both Schizopods, show characters which connect the lower

Crustacea more and more with the higher ones. Perhaps the fossil forms might give some

further evidence. We may also hope to get more forms from the deep sea which will

enable us more and more to understand the curious genera which are known under the

names of Ceratiocaris, Dithyrocaris, &c.

IV. ON sOME GENERA OF SCHIZOPODS WITH A FREE DoRSAL SHIBLD.

1. On GNATHOPHAUSIA, a new Genus of Lophogastride. (Plates IX. & X. figs. 2-4.)

During our cruise from the Bermudas to the Azores, Madeira, Cape-Verds, and from

there to the Brazilian coast, three species of this new genus of Schizopods came up in

the trawl or in the dredge. One of these species (now probably figured in Professor

Thomson’s “‘ Notes from the ‘Challenger,’” in ‘Nature’) is very large, probably the largest

Schizopod which we know. Of this one, which I have called Gn. gigas, only one male

was caught; of another, smaller species, Gn. zoéa, two males and two females; and of the

third, Gn. gracilis, only one male. Of these I have dissected one female of Gn. zoéa. I

have also made drawings of the legs of Gu. gigus, which I could not dissect ; but finding

that they have a close resemblance to those of the former one, I shall not publish them.

A sketch of the oral apparatus, however, and of a maxilliped, of the larger species are

among my figures.

| GNATHOPHAUSIA *, n. gen.

1.. GNATHOPHAUSIA GIGAS, 0. Sp.

The specimen, a male, came up from a depth of 2200 fathoms in lat. 38° 22’ N. and

long. 37° 21' W., between the Bermudas and the Azores.

Ti has a length of 142 millims., and much resembles at first sight a large Caridid

shrimp ; but the abdomen is straight and not bent down by the enlargement of the third

pleon-segment, as is the case in the Caridide.

'' The Carapace.—Its anterior part extends as a triangular rostrum (85 millims. in

length), starting in the form of a high ridge from the midst of the hepatic region. From

its base to the top of the rostrum the dorsal shield has a length of 64 millims.; it is a

flexible parchment-like duplicature of the skin, covering entirely the lateral branchiz,

and touching with two lateral and backward-directed spines the middle of the second

abdominal ring. Its upper posterior edges, however, do not quite cover the last segment

of the pereion. In Lophogaster it leaves two segments entirely free, and its borders are

not united with the pereion. Here the carapace starts from the cephalic and the first

pereion segments as in Apus and Nebalia. It is in no connexion with the five posterior

* Tya0os and davotos.

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 29

segments of the pereion. On its upper surface there are five longitudinal ridges and two

transverse sulci, which can best be understood from the figures (Pl. X. figs. 2 & 2 a),

These ridges divide the carapace into five regions, which we might call the regiones

cephalica, hepatica, cardiaca, branchialis, and marginalis. I think it unnecessary to give

for each the exact dimensions.

In front there is a cleft for the pedunculated eyes, above which there is a sharp spine ;

laterally, above the cleft, which is occupied especially by the second maxilla, there is

another spine. The lateral edges are more or less flattened, and form two “rounded

plates,” as Sars calls them in his paper on Lophogaster *, between which there is the

just-mentioned cleft. This can best be explained by Pl. X. fig. 8, where I have figured

the parts of the mouth and the upper lateral edges of the dorsal shield, seen from below.

There the upper plate (2), to which are attached the muscles of the oral apparatus, covers

to some extent the mandible; the lower plate (y) protects the palpus and the accessory

eye of the second maxilla from behind.

In Lophogaster there is also to be seen a rudiment of a rostrum and the longitudinal

ridge (Sars, fig. 1). The lateral spines are very much like those of Guathophausia; the

lateral edges of the shield, however, are not prolonged to such an extent backwards as in

our genus, and the posterior margin leaves, as has been mentioned already, two segments

entirely free.

The eyes issue from between the rostral portion of the shield and the base of the inner

antennz ; they have a length of 8 millims., and show two joints in their peduncle.

The «ner antenne consist of a short and stout funiculus and two flagella. The funi-

culus is four-jointed, and attains a third of the length of the squamiform appendage of

the outer antenna. It is twice the diameter of the eyes in width, and is densely covered

with hairs on its inner side. The exterior flagellum is very long (130 millims.), and flat-

tened at its base (2 millims.), but the interior flagellum rounded and much shorter (length

40 millims.). Underneath and behind the inner antennz is the large base of the squami-

form appendage, showing outside a strong spine. The appendage itself entirely covers the

funiculus, and has a length of 19 millims.: its inner border is even, but densely covered

with hairs; the outer one has at the base a rounded prominence, and further on four

sharp spines, in addition to the top spine. A longitudinal ridge (a trace of which may

also be seen in the cordiform appendage of Lophogaster—Sars, fig. 13 e) divides the

appendage into a larger and a smaller portion. Underneath this appendage we see a

three-jointed long funiculus of the exterior antenna (length 5 millims.), the flagellum of

which has the length of the shorter one of the inner antenna (40 millims.).

Oral apparatus. (Plate X. fig. 3.)

The upper lip is a large cordiform plate (63 millims. in length by 5 millims. in width),

the top of which reaches upwards to the first joint of the antenne interioris funiculus.

A short groove descends from its centre towards the mouth ; this organ is nearly of the

same shape here as in Lophagaster (Sars, fig. 17). Very Lophogaster-like are also the

strong mandibles (fig. 8, md), to which are attached two three-jointed palpi, the inner

* «Beskrivelse over Lophogaster typicus’ (Christiania, 1862), pl. i. figs. 3 & 4, and p. 2.

30 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

side of which is densely covered with hairs (fig. 3, mp). The palpi are situated on both

sides of the upper lip, and reach higher up than the funiculus of the exterior antenna.

The first pair of maxillz could not be inspected, as they are entirely covered by the second

pair, which, like the whole oral apparatus, have a strong resemblance to those of Lopho-

gaster (Sars, fig. 28, cd). All the parts mentioned there are also to be found in our case

(fig. 8, ma) :—on the inner side the two digitiform manducatory processes covered with

stiff hairs; close to them on both sides a flagellum (fig. 3, /), which is jointed in Gna-

thophausia; and on the outsides the palpus (fig. 3, p) in the form of a large leaf-like

appendage fringed with hairs and occupying, like a large cover, the whole space between

mandibles and maxillipeds. From the basal joint of these maxille start a pair of small

black pigmented accessory eyes, the peduncles of which have a length of 2 millims.

Their position and direction are such that they just peep out between the inferior mar-

gin of the dorsal shield and the gnathopods. A dissection of these accessory eyes is

of course impossible in our case; but I think one is right in supposing that they consist,

like the accessory eyes of the Euphausiide, of a lens, nervous rods, and a pigmented sub-

stance. They have been found in the Euphausiide on the basal joint of the pereiopods,

as well as of the pleopods, and in Thysanopoda norvegica even up to the number of eight

pairs. In Zophogaster there are none, nor have accessory eyes ever been observed on the

maxille of any animal.

There is only one pair of maxillipeds, as there is in Lophogaster; and in both the

enathopods are used for walking. On the large transverse basal joint of the maxillipeds

we have a long flattened palpus (Pl. IX. fig. 16, p), and somewhat further on a small

flagellum (fig. 16, f). The second and third joints of these maxillipeds are very short ;

the fourth is longer, the fifth very short again; and the sixth is the terminating claw.

Gnathopods, Pereiopods, and Branchie.

We have remarked already that the gnathopods do not essentially differ from the perei-

opods, and are used for walking. At the base of all of them, with the exception of the

last pair of pereiopods, are the arborescent gills, the branches of which are partly situated

underneath the lateral parts of the dorsal shield, partly projecting into the water. There

are, as far as I can see without dissecting or spoiling our specimen, three branches, one

of them on the sides, one between the legs, and a larger one leaning on the pectoral shield,

which is entirely covered by two rows of branchiz. In the whole the arrangement seems

to me to be exactly the same as in Lophogaster; and the pictures given by Sars (figs. 16,

35, 37, and 45) give also a very fair idea of the branchial appendages of Gnathophausia.

The gnathopods and pereiopods (Pl. IX. figs. 8-14 may be compared here, though these

figures are drawn from Gz. zoéa) consist each of seven joints, if one counts that basal one

to which the branchize are attached. To the second joint the palpi are attached, which

differ somewhat from those of Lophogaster; they are shorter, and in Gnathophausia

the ramus palpi rapidly decreases in width, and has a great resemblance to the ramus of

one of the pleopods. The palpus of the first pereiopod has a length of 13 millims. The

third and fourth joints are in all the legs very short, and do not offer any thing parti-

cular; but the fifth and sixth are very different from each other in different legs. In

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. al

the first gnathopod they are even, very strong, and somewhat curved inwards; there are

only hairs at the inner side, not on the lateral parts of the joints, which in the second

gnathopod have got three lines of hairs, and in the following ones five and six. In the

last three pereiopods there are not only lines of hairs on the lateral parts of these joints,

but their point of attachment is strongly marked in the chitinous substance, so that

these joints look as if they were divided into five and six subjoints. And the same thing

is the case throughout with the fifth jomt. Such subjointed pereiopods are also found

in Yysis, but not in any other genus of Schizopods.

In the gnathopods as well as in the pereiopods the last joint is a short claw—even in

_ the first pair of the former, where it is wanting in Lophogaster.

The Pleon and its Appendages.

The abdomen has a length of 78 millims. It is square-shaped; for its segments are

nearly as large as they are high. The second segment, for example, is 11 millims. in

height by 10 millims. in width. The edges of the segments are even; only laterally

there are squamiform processes, which cover each other, and each of which has a rounded

anterior and a sharpened posterior portion. The shortest segment is the fifth; the sixth

is much larger, and is divided by a deep sulcus into two parts, which look perfectly like

. two distinct segments. One is the more inclined at first to think so, as the lateral squa-

miform processes, which terminate all the preceding segments, are attached to the first

of these two subdivisions. In the second one there are only two short spines on each

side. Sars has described something very much like this in Lophoguster ; and as we now

know that Medalia, in which there are nine abdominal segments, has essentially schizo-

podal characters, while Gnathophausia in its unattached dorsal shield has decidedly a

phyllopodal character, this pseudo-amplification of the abdominal segments seems to me

to be exceedingly interesting, as it shows that Gnathophausia must be in genealogical

connexion with forms (or a descendant of forms) in which there are more than seven

segments in the pleon.

The telson has a length of 30 millims.; on its surface there is a longitudinal groove

bordered by two ridges; the lateral borders of the telson are denticulated; at its end

there are two spines, between which there is a semicircular cleft. The caudal appendages

are both jointed (Pl. X. fig. 2); the interior are somewhat shorter than the exterior ones ;

and both are shorter than the telson ; at their basal articulation there is a spine on both

sides.

The five pairs of pleopods consist of a pear-shaped basal joint, at the posterior side ot

which there is a protuberance; at their inner side they are covered with a fringe of hairs.

‘The two annulated rami do not present any thing particular. Appendages such as have

been found here in the Huphausiidz do not exist in Guathophausia.

Before fixing the generic and specific characters of this large species, [ shall now give

a shorter description of the smaller one (of which four specimens were got), and point out

especially the characters in which it differs from the larger one.

32, DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

2. GNATHOPHAUSIA ZOBA, 0. sp.

In its general appearance this species differs from Gn. gigas at once by the presence

of a long posterior spine at the dorsal shield.

The specimens, two males and two females, were got in very different places. (1)

June 30,a male was obtained off the Azores in a depth of 1000 fathoms; it has a length

of 49 millims., the rostrum being 18 millims. long. (2) August 25, in lat. 1° 47’ N., long.

24° 26' W.,a male came up from a depth of 1650 fathoms: length 58 millims., with a

rostrum of 17 millims. (8) August 26, a female from 1500 fathoms. in lat. 1° 22'N., long.

26° 36’ W.: length 81 millims., with a rostrum of 14 millims. (4) September 12, off Rio

San Francisco on the coast of Brazil, another female came up from 750 fathoms: length

313 (rostrum 7) millims. According to this the size of the males varies between 49 and

58 millims., and that of the females between 313 and 81.

The substance of the dorsal shield is flexible, like strong parchment ; it is as in Gn.

gigas a duplicature of the skin, adhering to the body only in the first segments, and in

no connexion with the five posterior segments of the pereion. A ridge running over the

whole length of the shield is continued into a long triangular rostrum, which, in the

males, is much longer (3 of the length of the whole animal) than in the females (+ of the

length). The posterior spine is a continuation of this ridge, and has in both sexes a length

of 6-8 millims. Both rostrum and posterior spine are triangular and armed with small

secondary spines, which in the former are to be seen on all sides, in the latter only on two

sides. The posterior spine reaches backward to the third abdominal segment; and all the

segments of the pereion are covered by the dorsal shield, with the exception of the lateral

parts of the last one.

The posterior angles of the shield are rounded. In front there is a sharp spine above

each eye, and another somewhat smaller one below it, just inside, at the place where we see

the round plate, to which are attached the muscles of the oral apparatus. A transverse

sulcus, a transverse ridge, and two longitudinal ridges (besides the larger central one)

divide the shield, as in the former species, into cephalic, hepatic, cardiac, branchial, and

marginal regions (Pl. X. fig. 4).

The inner or first antenne hardly differ from those of the preceding species. Their

funiculus only reaches to the middle of the squamiform appendage. The outer or second

antenne differ from those in Gn. gigas by the shape of their scale, in which only one

spine is visible (Pl. IX. fig. 2, a1), while in the larger species there are five spines.

The parts of the mouth in this species could be dissected and submitted to micro-

scopical examination, which showed that they are very much like those in Lophogaster.

The very strong and big mandibule (Pl. IX. fig. 3) have a palpus the second joint

of which is somewhat recurved. The third joint shows two kinds of hairs—very short

ones, which also Sars has figured in his genus, and longer ones at the top. ‘The wpper

lip has the same cordiform shape as in the preceding species; and the wader lip (Pl. IX.

fig. 4) is more clumsy than that of Lophogaster, and does not show so deep an incision,

but has the same small hairs at its anterior border. The first maxilla is distinguished

by very strong bristles in its inferior portion, and has a palpus which, among Schizopods,

has hitherto only been found in Nedalia (fig. 5, p); at the top of this palpus we find very

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 30

long hairs which by far surpass it in length. In the larger species I could not examine

the first maxilla; but it probably has got a palpus too. Lophogaster has not got it; and

it is very interesting to find it in Gnathophausia, as it is one more connecting link between

this genus and Nebalia. The second maville are large organs (fig. 6), covering the former

ones and exhibiting a flagellum, a two-jointed palpus, and an accessory eye. I tried

to make out the structure of this eye by hardening it and making it transparent, but did

not succeed ; all I saw was the cornea and a dark pigment inside. For him, however,

who has seen the same organ in the larger species, there is not the slightest doubt that

these sense-organs on the second maxille are the same things which have been called

accessory eyes in the Huphausiidee. They are present in all the specimens of the genus

Gnathophausia in the same place.

In the mawillipeds (fig. 7) of this species I could not find the palpus, which in the

larger is so clearly visible (fig. 16). Possibly it is wanting in the female; for I do not

think that it escaped my attention as I looked for it with the microscope.

The two gnathopoda (figs. 8 & 9) have got at their base breeding-lamelle and three

pairs of gills, one of which is situated on the ventral side, while two are laterally covered

by the shield. No subjoints can be seen in these first two legs, traces of them are to be

found in the first and second pereiopods (figs. 10 & 11); but they are only very clearly

visible on the third and fourth (figs. 12 & 13). In the last (in which I have drawn the

breeding-lamella, fig. 14) they are wanting, as are also the gills at its base. Among

Mysidee we have the same thing which we find now in Lophogastride, for these sub-

joints which are wanting in Siriella (as they here are in Lophogaster) are to be met with

in Mysis.

The abdomen of Gn. zoéa differs somewhat from that of the larger species. We do not

find here those squamiform prolongations, but have simply a few small spines at the

posterior and inferior angle of each segment. In the sixth we have the same peculiar

arrangement of its being divided by a deep ridge into two pseudo-segments which do not

articulate with each other. There is a spine bent backwards at the end of the first one,

and several small spines at the end of the second.

The ¢elson (8 millims. in length) terminates in a half-moon; the caudal appendages

are both jointed; the pleopoda are very slender (fig. 15), and show a small tubercle at

‘the inner posterior side.

The male differs from the female in this species by the length of the rostrum, which

in the former has one third of the length of the whole body, and in the latter only one

fifth. In the size of the inner antennz I see no great difference. Whether the male or

the female attains a larger size I cannot say; the largest specimen of G. zoéa which we

got was a female; besides, it is very easy to tell a male from a female by the want of

‘breeding-lamelle in the former.

3. GNATHOPHAUSIA GRACILIS, n. sp. (Pl. IX. fig. 1).

Only one male of this smallest Gnathophausia-species has been hitherto got; it came up

on the 26th of August, in lat. 1° 22’ N., long. 26° 36’ W., 170 miles east of St. Paul’s rocks;

from a depth of 1500 fathoms. It was on this same day that one of the specimens of Gz.

SECOND SERIES.—ZOOLOGY, VOL. I. F :

34 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

zoéa and the male of Petalophthalmus were got. The specimen has a length of 41 millims.,

with a carapace of 24 and a rostrum of 14 millims. The length of the rostrum (one third

of the whole length) and the absence of the breeding-lamelle show that it is a male.

From the two preceding species this little Lophogastrid differs by the presence of two

spines at the anterior and posterior lateral angles of the carapace and by the presence of

spines at the top of the first, second, and fifth abdominal rmg. From Gn. gigas it is

easily distinguished by the presence of the middle posterior spine of the carapace and —

the absence of the five spines on the lamellar appendage. There are also several spines

on the line which divides the carapace in its full length, which are absent in the other

species.

As in other respects it shows no characters which have not already been found in

the two preceding species, I refer to the figure (fig. 1), and shall not describe in detail

all its parts.

Probably the genus G'nathophausia is represented at the bottom of the deep sea by

many species; but I do not think that these species are very numerous in individuals ;

for the trawl always brings up many more Peneids and Caridids than Schizopods.

Colour, and Mode of Life.

All the Gnathophausias, when they came up, were bright red, a colour which is com-

mon to all deep-sea Crustacea, but which they very soon lose in alcohol.

Like Lophogaster, which was found by Sars at the bottom of the sea in 50-60 fathoms,

these Schizopods live probably near the bottom without ever coming to the surface

(except perhaps in the larval stage). They evidently live together with a rich fauna,

of which the trawl has given us some idea. As to the animals which live together with

the larger species, I am obliged to cite here those which were got the day before and the

day after the capture of Gi. gigas; for on that same day when it came up the trawl con-

tained nothing else. In the first of these stations (June 24, lat. 38° 3’ N., long. 39° 19!

W.) we found a depth of 2175 fathoms; and in the other one, two days afterwards (lat.

38° 25’ N., long. 35° 30’ W.), when we already approached the Azores, there was a depth

of 1675 fathoms. We got there several sponges, a Palythoa, a Pennatulid, a small hydroid

polyp, the Sea-urchin Salenia, several Asterids, some Bryozoa, one Annelid (Hermione), a

Pyenogonid * (Zetes, sp.?), and some blind Isopods, one of which belongs to the Munop-

side family.

' The smaller species were got in different localities and depths (750-1850 fathoms)—

near the Azores, not far from St. Paul’s rocks, and near the coast of Brazil. Together

with them were found many inhabitants of the deep sea, some of which I shall now men-

tion. We got there the sponges Hyalonema, Tisiphonia, and Halichondria, the celebrated

Uinbellularia grenlandica, Fungia symmetrica, and a Flabellum. Among the Hchino-

derms (which were very numerous) may be mentioned Chirodota and Synapta, a Bathy-

crinus, Astropecten, Astrogonium, Asteracanthion, and the Urchins Hehinocyamus and

* From several reasons I supposed, when beginning our operations, that we should get Pycnogonids in great

abundance and variety; but just the contrary was the case, as hitherto, after ten months’ deep-sea dredging, we have

only got from two to three Pycnogonids, belonging, as far as I can make out here, to Kroyer’s genus Zetes.

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 35

Salenia. Among the worms, which, unfortunately, usually come up in single and, in most

cases, spoiled specimens, we found some Annelids, a long Nemertean, and a very large

Balanoglossus*. Crustacea there are many—Peneids, Caridids, a Pagurus, and the genera

Chalaraspis and Petalophthalmus (which are described in this paper). Only one Brachi-

opod and several genera of Gastropods (Dentaliwm, Mitra, Pleuronectia, Newra, Limopsis,

and Cardium) belong to this fauna. There were also several fishes, one of which (the

Lophioid Melanocetus) certainly lives at the bottom in great depths.

The Systematic Position

of our genus is easily fixed after what we have seen of its resemblances to Lophogaster ;

for, after all that has been said about it, nobody, I am sure, will doubt that Guathophausia

is a member of the family Lophogastride. Now fortunately Sars has so well discussed

the affinities of Lophogaster, that the systematic position of our genus is easily under-

stood when we say that it is a Lophogaster with a free dorsal shield (like an Apus), with

a palpus on its first maxilla (like Nebalia), with accessory eyes on its second maxilla

and with subjoints in its perelopods:(like Mysis). Gnathophausia, it is true, has many

peculiarities which are not found in the genus Lophogaster; but I must nevertheless

maintain that it is closely allied to that form. The gnathopoda and pereiopoda all having

the shape of legs, the three branchiz, two of which are laterally covered by the carapace

and one not, and the presence of breeding-lamellee on all the legs, are fundamental cha-

racters which, according to our present systematical arrangement of the Schizopods,

assign it its place among the Lophogastride. The subdivision of the sixth abdominal

segment, the shape of all the appendages and of the telson, are nearly exactly the same

in the two genera; but there are other characters which Lophogaster has not, characters

which one might call atavistic ones ; and these are (1) the looseness of the dorsal shield,

(2) the presence of a palpus maxille prime, (3) the subjoimts in the legs, and (4) the

accessory eyes on the second maxille.

The subjoimts in the pereiopods (3) have been hitherto only known as a peculiarity of

the genus Mysis, and the accessory eyes (4) as occurring in some genera of the Euphau-

slide family. We therefore find characters united in Gnathophausia which were only

known separate in the two other families of Schizopods. Hereafter, however, we shall

find in a Mysid a character which until now was supposed to characterize only the

Lophogastride, and shall see that these deep-sea forms of Schizopods combine in many

ways the characters of the shallow-water forms, the types of the hitherto acknowledged

families. But we shall aiso see that they can nevertheless be regarded as belonging to

those families (with the exception of one genus), as they have not a mere mixture of

characters, but are Mysids or Lophogastrids which possess some characters of the other

families.

* Two fragments of this interesting worm were got August 19, in lat. 5° 48’ N., long. 14° 20’ W.., off the west

coast of Africa, from a depth of 2500 fathoms. The whole animal had probably a length of 4 inches; the head and

a piece of the body, however, were the only things we got. These fragments were very remarkable for the vivid

colours which they displayed: the head was yellow, the collar bright red, and the body of a yellowish red. Of this,

the Neapolitan species and the one which I discovered in the sand near Copenhagen, in shallow water, show nothing,

as they are uniformly white; a priori one would have been inclined to suppose just the contrary.

30 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

The palpus of the first maxilla (2) and the free dorsal shield (1) are both characters

which, among Schizopods, we only find in Nedalia, a genus which has hitherto had a very

isolated position, and was considered a connecting link between these and the Phyllopods.

In Plate VI. fig. 9, we have figured the first maxilla with its long palpus in our Nebalia

longipes. ‘The palpus there is a mere continuation of the maxilla; here, however (where

it has very little resemblance to the Nedalia palpus), it seems to be in articulation

with the maxilla. This is a character which I have hitherto only found in G'nathophausia,

and confirms my opinion that this genus is, next to Mebalia, the most aberrant one among

the Schizopods.

In Lophogaster as well as in Guathophausia the dorsal shield is of a parchment-like

consistency. In the former its posterior and lateral edges are free, but the rest of the

shield forms the integument of the pereion-segments. A free dorsal shield has hitherto

never been found in a true Schizopod, but is also peculiar to Nebalia. Now Nebalia is

considered to be an aberrant Schizopod with nine abdominal segments ; and it seems to

me to be very interesting for the explanation of the semidivision of the last pleon-segment

in Gnathophausia, that this genus has two other phyllopodal characters, in the configura-

tion of its dorsal shield and the presence of a palpus on the first maxilla; but in all the

main points Gnathophausia is much more a typical Schizopod than Nebalia, and, I think,

not even closely allied to it. Probably both derive their origin from a series of animals

which were neither Schizopods nor Phyllopods, and to which possibly belonged the fossil

forms Dithyrocaris, Ceratiocaris, &c. I do not know how far these forms are really allied

to Gnathophausia, as I have not the literature on them on board; but I strongly suspect

that also on them some light may be thrown by the discovery of this interesting genus.

The Characters of the Lophogastride,

therefore, ought to be modified in several points; the following characters ought to be

added to the family-description :—connexion of the shield with the segments of the pereion

complete or incomplete ; with or without accessory eyes on the second maxilla; with or

without a palpus on the first maxilla; and with or without subjoints in the pereiopods.

- Besides, all the family characters given by Sars are applicable to both genera.

Characters of the Genus and Species.

GNATHOPHAUSIA™, n. gen.

Carapace in no connexion with the five posterior segments of the pereion; a long

rostrum ; first antenne with a short and stout pedunculus and two flagella, a long and

a shorter Inner one; second antennee with an oblong squamiform appendage ; stout

mandibles, with a three-jointed palpus; a palpus on the first maxille; accessory eyes,

palpus, and flagellum on the second maxillze; maxilliped showing nothing particular.

First gnathopods terminated by a claw; all the gnathopods and pereiopods leg-like, the

latter showing subjoints on their penultimate joint. Six pairs of gills at the base of the

gnathopods and the four anterior pereipods ; two branches of the arborescent gills laterally

* From yvalos and gavavos.

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 37

covered by the carapace, one branch turned inside between the legs and in free commu-

nication with the water. Well developed pleopods. Caudal appendages jointed.

1. GN. GIGAS, 0. sp.

Scale of the second antennz with five spines, not jointed; short spines at the lateral

angles of the dorsal shield, no spine in the midst of the upper posterior margin.

_ Length 142 millims. Colour red.

2. GN. ZOBA, 0. sp.

Scale of the second antennz with only one spine, jointed ; dorsal shield with a sharp

spine in the midst of its upper posterior margin, lateral posterior angles rounded.

Length 59 millims. Colour red.

3. GN. GRACILIS, n. sp.

Scale of the second antennz with one spine; dorsal shield with a spine in the middle

of its upper posterior margin, and two spines in its anterior as well as in its posterior

lateral angles.

Length 41 millims. Colour red.

2. On CHALARASPIS, the Type of a new Schizopod Family. (Plate VIII.)

This little Crustacean is the commonest Schizopod of the deep-sea fauna, and seems to

enjoy a very wide bathymetrical and geographical distribution ; for we got it from depths

of 350-2500 fathoms, off the west coast of Africa, as well as off the east coast of South

America, down to the 35th degree of southern latitude. Whenever in the mid-Atlantic

true deep-sea animals came up in the dredge or the trawl, I was sure to get at least a

fragment of this Crustacean. Unfortunately most of the specimens were spoiled ; only a

few males and females were in a good state of preservation. The animal is so soft that it

does not stand the long passage of the trawl through the water; and even if it is not

broken to pieces, its carapace is usually overturned. On two occasions, however, I got

specimens which are quite intact; and these, as well as the many fragments, furnished

ample opportunity of studying this genus in all its details.

The females of this shrimp have a length of 35 millims., the carapace having from

above a length of 8 millims. and laterally of 12 millims. The males are 37 millims. in

length, but somewhat more slender than the females; their dorsal shield has the same

length from above as that of the females, but is laterally somewhat shorter (114 millims.).

The carapace is very soft, especially in the males, and in connexion with only the first

segments of the pereion. It has a very short, rounded rostrum (fig. 1 a), and on the

frontal border a pair of short spines underneath the eyes. The lateral anterior border is

rounded in the female, and somewhat angular in the male. The posterior border of the

carapace shows a deep excision, especially in the male, so that its rounded lateral borders

cover a great deal more of the pereion-segments than the upper margin (fig. 1). The

carapace is divided by a longitudinal ridge and two transverse sulci into several regions,

which in old females are very well marked; in the males, however, they are not always

plainly visible.

38 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

Underneath the short rounded rostrum, between the eyes, there is a small tubercle

(fig. 1 a, 7) which in the males is somewhat more prominent than in the females.

The eyes have the same position as in Gnathophausia, and are 2 millims. in length.

They cover somewhat the first joint of the funiculus of the first antenne, which consists

of three joints, the last of which has a lateral prolongation (fig. 2). Between the third

joint and the insertion of the flagella a small subjoint is to be seen. The external fla-

gellum is much bigger than the internal. Hairs are in both sexes at the inner side of

the funiculus, the size of which is also very much the same in both. The scale of the

second antenna is jointed and ovoid. The slender funiculus has three joints and a

flagellum, which exceeds in length the whole animal.

The wpper lip (fig. 11) is cordiform, with the point directed upwards. The mandibule

(fig. 4) are not particularly strong, and have a three-jointed palpus, the second joint of

which is somewhat bent backwards, and the third rounded at the top.

The excision between the two pieces of the under lip (fig. 5) is not so deep as in Gna-

thophausia. On its anterior border we find the same hairs which may be seen in the

Lophogastride.

The first mazilla (fig. 6) has two manducatory processes: the upper one has many

strong teeth ; while the under one has only a few hairs, no denticulation.

The second maxille (fig. 7) have a very elongated flagellum (/1) and a small two-jointed

palpus (7).

The maxillipedes (fig. 8) have a very large palpus and a flagellum on their enlarged

basal joint. They are terminated by a small denticulated claw.

The gnathopoda and the first pereiopod in this genus (which hitherto has not shown so

very great differences from Lophogaster) are not leg-like, but have all the shortened mawil-

lipedal form and function (figs. 1,9, & 10). This is the same in both sexes. The branchize

and (in the female) the breeding-lamelle begin at the base of the first gnathopod. The

form of this appendage (fig. 9) can best be seen from the figure. The terminating claw is

not denticulated ; the second is somewhat longer than the first, but shows no peculiarities.

It is very astonishing that in both sexes of this genus the first pereiopod has the same

maxillipedal form and function as the gnathopoda. This is the case in no other Schizopod

family. Also the form of its various joints (fig. 10) is very much like that of the gnatho-

pod-joints.

Among the four remaining pereiopods we have again two very different forms: three

of them are enormously elongated (figs. 1 & 12; they have a length of 27 millims.) and

terminated by a strong and recurved claw, which has many spines at its inner side,

whereas the last pereiopod (fig. 1, p*, & fig. 18) does not end by a claw, but by a simple

rounded joint, is very much shorter than the three preceding leg-like pereiopods, and all

over covered with hairs.

There are arborescent branchie at the base of all the gnathopoda and pereiopoda with

the exception of the last one. ach pair of branchiz consists of three branches, two of

which are laterally covered by the carapace; the third branch is bent inside between the

appendages, and is accordingly in free communication with the water. In this respect

there is no difference between this genus and the Lophogastride.

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 39

There are seven pairs of breeding-lamelle at the base of the gnathopoda and pereiopoda,

increasing in size towards the end of the pereion. This also is quite the same thing as

has been seen in the Lophogastride.

The abdomen of Chalaraspis differs from that of the family just mentioned especially

by the absence of the pseudo-segment. The last segment of the pleon is elongated and

perfectly smooth.

The pleopods are very perfect; they have a strong basal joint and two rami. They

offer nothing particular either in the male or the female.

The posterior borders of the quadrangular ¢e/son show a slight denticulation (fig. 1,0) ;

the outer appendages of the caudal fin are jointed.

The males differ from the females only by being somewhat more slender and by the

absence of the breeding-lamelle. The small tubercle underneath the rostrum seems to

be more prominent in the males than in the females.

Systematic Position of the Genus.

Among the three families of Schizopods (Myside, Huphausiide, and Lophogastridz)

hitherto known, this new form no doubt approaches most the latter, with which it

has in common the position and shape,of the branchiz, the breeding-lamelle, and the

pleopods; but the presence of only four leg-like appendages, and the fact that four

pairs of appendages act here as maxillipedes, do not allow us to class it together with

them, as here both genera (Lophogaster and Gnathophausia) have got seven pairs of leg-

like appendages. It depends, of course, on what one considers the main point of view

from which a classification of the Schizopods ought to be made, whether from the form

and position of the branchize, or from the number of appendages which are transformed

to maxillipedes. I think the best plan is to carry on the hitherto accepted principle—to

establish different families for those groups in which there is a different number of leg-

like appendages. One should only deviate from this principle in case an animal should

be discovered which in every other respect is so closely allied to a certain group that the

conformation of the legs appears only as a very secondary acquisition ; but from what

we have hitherto seen of the steadiness in which the same number of mazxillipedal

appendages is kept in forms which differ so much from each other as Guathophausia

does from Lophogaster, or Petalophthalmus from Mysis, I doubt very much whether such

forms will ever be found.

For the genus Chalaraspis one ought therefore to establish a separate family, Chala-

raspidze, holding its place between the Euphausiidz and Lophogastride, and characterized

(until further discoveries show whether it has got several living members differing from

each other) by the

Characters of the Genus.

CHALARASPIS*, n. g.

Dorsal shield or carapace in no connexion with the posterior segments of the pereion ;

rostrum short, rounded ; eyes present; no accessory eyes ; first antennze having a stout -

* XaXapos and doris.

40 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

funiculus and two flagella; scale of the second antenne jointed; upper lip triangular ;

mandibule with a three-jointed palpus; no deep excision in the under lip. Second

process of the first maxilla without denticulation ;. second maxilla with an elongated

flagellum and a small palpus; maxillipedes terminated by a denticulated claw, with large

palpus and flagellum. Gnathopoda and first pereiopod used as maxillipedes, shortened

and recurved; second, third, and fourth pereiopod having nearly the length of the body,

terminated by a long denticulated claw ; last pair of pereiopods shorter, terminated by a

rounded joint, very hairy, and without branchie. Three branches of branchiz on the

base of the gnathopoda and first four pereiopoda, two of which are covered by the cara-

pace. Breeding-lamellze on all the gnathopoda and pereiopoda. No pseudo-segmentation

on the sixth segment of the pleon. External caudal appendages jointed.

CHALARASPIS UNGUICULATA, 0. sp.

Length 35-37 millims.

Mode of Life, and Colour.

We have already seen at the beginning of this description that this is the commonest

deep-sea Schizopod with as wide a geographical as bathymetrical distribution. It lives

together with the same fauna as Gnathophausia zoéa, which several times came up

accompanied by Chalaraspis. Its colour is a bright red.

3. On PETALOPHTHALMUS, a Mysidiform Schizopod. (Plate VII.)

In Prof. Wyville Thomson’s ‘Depths of the Sea,’ p. 176, there is a quotation from a

preliminary note of the Rev. A. Merle-Norman, according to which specimens of Ethusa

granulata, a brachyurous Decapod, taken in 110-370 fathoms, are apparently blind, but

have two remarkable spiny eye-stalks, with a smooth rounded termination, where the eye

itself is ordinarily situated. In other specimens of the same crab the rostrum disappears

entirely, and the two eye-stalks, approaching each other, assume its functions. In

accordance with the altered conditions of life, the eyes and their stalks seem to undergo

in Hpethusa the most extraordinary modifications.

The same thing which now-a-days happens to a crab living in different conditions of

life has evidently also happened (but in the course of ages probably) to a Schizopod, in

which we find at the top of the eye-stalks, instead of organs of vision, two flat spherical

terminations in which no trace of an eye (which one would expect to find in this place)

has been left. But this peculiarity in the Schizopod seems not to be a modification of

individuals exposed to certain conditions of life; for we got specimens from very different

depths (1590-2500 fathoms) and latitudes, from the mid-Atlantic near St. Paul’s Rocks

down to the southern regions of Tristan d’Acunha, in both sexes of which the conforma-

tion of the eye is exactly the same. Though in the whole these Schizopods approach the

eenus Mysis, we shall find that they have several peculiarities which do not allow us to

suppose that this is a Jfysis which, like the above- mentioned Lyethusa, has changed

when exposed to certain conditions of life, and of which specimens might exist now-

a-days in full possession of their organs of vision. Once, of course, this process must

have taken place, but probably at a time when such Schizopods as Gnathophausia and

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION. 41

Chalaraspis were the only representatives of the order, from which only in later periods

of the earth the well-known shallow-water members of the Schizopod families have

developed.

T have called the animal which I shall now describe Petalophthalmus, and the species

P. armiger. The first specimen, a female, was caught by the trawl in lat. 2° 25’ N.,

long. 20° 1’ W., about midway between Cape Palmas and St. Paul’s Rocks, from a depth

of 2500 fathoms. We afterwards got a male in lat. 1° 22' N., long. 26° 36’ W., about

170 miles east of St. Paul’s Rocks, from 1500 fathoms, and, finally, a female in lat.

35° 41' S., long. 20°55' W., from 100 fathoms, about 400 miles west of Inaccessible Island

(Tristan d’Acunha group). The male has a length of 37 millims., the largest female of

44 millims., and the smaller one of 30 millims. .

The carapace, soft and parchment-like, is in no connexion with five posterior segments

of the body. In the female, where it has not quite one half of the total length of the

animal (it is 18 millims. in length in a specimen of 44 millims.), it covers in its ordinary

position nearly all the segments of the pereion. It has a small rostrum (fig. 1), which

is very little prominent, a small spine at its anterior border underneath the eye, and

sharpened anterior angles. The posterior and inferior angles are rounded; and the sur-

face of the whole carapace is perfectly smooth. A slight transverse sulcus divides it into

an anterior and a posterior portion, the latter of which is by far the largest.

Underneath the rostrum are the eye-stalks, terminated by concave spherical plates

having a diameter of 3 millims.; they are simply a duplicature of chitinous matter. On

examining them under the microscope I could not find a trace of any eye-like structure.

The first antenne have a three-jointed funiculus (I consider the small joint in fig. 3,

which is not to be seen in the first antenna of the male, to be not a fourth, but an ex-

ternal pseudo-joint) and two flagella, the external of which is enlarged at its base and

covered with hairs at the inner side.

On the second antenna (fig. 4) there is a lamellar appendage (fig. 5) which has no

spines.

The labrum is subcordiform (fig. 6). The mandibule (fig. 7) have a very strong

manducatory portion, with many denticulations and stiff hairs. The second joint of their

palpus is enlarged; the third has a rounded point and many strong hairs at its inner side.

The under lip is somewhat like that of Chalaraspis, and bordered anteriorly with small

hairs. I could not draw it, as I saw only a part of it, the organ being broken when I

tried to take it out.

The first maxille have two processes, showing many hairs and a strong denticulation

(fig. 8); they have no palpus. The second mazille (fig. 9) have both palpus (p) and fla-

gellum (fi). The mawilliped (fig. 10) has a much more perfect palpus, quite close to its

flagellum, than Grathophausia has; besides, there is at the inner side of the third joint

one of those flat lamellar appendages which are known in the maxilliped and first ena-

thopod of Mysis and Siriella, and which in the male of our genus are indeed present

on both appendages, in the female, however, only on the maxilliped.

The first gnathopod (fig.11) has still maxillipedal functions, though it is rather elon-

gated; its penultimate joint, however, is enlarged and recurved, and terminated by a

SECOND SERIES.—ZOOLOGY, VOL. I. G

42, . DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC ~

rounded claw, which fits into an excision at its top. The second gnathopod (fig. 12) is

perfectly leg-like. Its tarsus, like that of the following pereiopoda, is not subjointed, as

in Wysis and Gnathophausia, though there are hairs at intervals on it as in those Crus-

taceans.

The five pereiopoda are all very much alike; only the fifth is somewhat shorter than

the rest. I have only drawn the third (fig. 18).

At the base of ad2 the gnathopoda and pereiopoda we find breeding-lamelle (fig. 1, and

figs. 12 & 18), the posterior ones nearly as long as the legs. This is a character not

found in any other Mysids, and, among all the Schizopods, hitherto only known in the

Lophogastridze.

The rings of the pleon are perfectly smooth, and do not present any thing particular.

The sixth segment is longer than the rest. The five pairs of pleopoda are only rudi-

mentary and exceedingly small (fig. 14); they consist of a small basal joint and only one

ramus. The last ones, especially the fifth, are longer than the first.

_ The ¢ail (fig. 1) is formed by the telson, which shows a little excision and two appen-

dages on each side, the exterior one of each pair jointed.

Branchie are entirely wanting.

The male differs very much from the female. Its soft and free carapace is much shorter,

not much more than + of the total length; for in a specimen of 37 millims. it has a

length of 10 millims. Thus the last two segments of the pereion are entirely uncovered ;

while several of the preceding ones are laterally uncovered, the upper part of the cara-

pace being much longer than its lateral parts.

Very extraordinary are the prehensile organs of the male—the first antenna, the pal-

pus mandibularis, the maxilliped, and the first gnathopod—altogether forming a for-

midable prehensile apparatus (fig. 2).

The funiculus of the first antenna is enlarged, and all the joints are elongated; it

has a length of 10 millims. Its flagella, however, are of ordinary size, the exterior

one being not enlarged at the base. The second antenne (fig. 2, a’) do not differ

from those of the female. The principal prehensile organ is the palpus mandibulze

(fig. 2, pm), the second and third joints of which are enormously enlarged, and the third

armed with strong sete. The maxilliped (fig. 15) and the first gnathopod (fig. 16) have

enlarged joints, and are terminated by strong claws. At their inner side is the lamellar

appendage (figs. 15 & 16, da), which we found in the female only in the maxilliped.

Unfortunately the only male which we caught has lost the second gnathopod and

the first pereiopod. The second pereiopod is terminated by a tuft of hairs instead of a

claw (fig. 2, prp?). The three last ones do not differ from those of the female, except by

being somewhat more slender.

In the male the pleopods are more perfect (fig. 17); for, besides being stronger, they

have two rami, one of which is broad and bordered at its rounded top with hairs, the other

pointed and nearly filiform. The last pleopoda are longer than the preceding ones.

The telson of the male (fig. 2@) does not show such a deep excision as that of the

female.

No traces of branchiz are to be seen.

CRUSTACEA FROM THE * CHALLENGER’ EXPEDITION. 43

Surely these characters of the Podophthalmus males ave quite exceptional among Schi-

zopods, and much more like what we find in certain ge in than what we should

expect to find among the higher Crustaceans. i

Systematic Position of the Genus.

-The six pairs of leg-like appendages, the rudimentary condition of the abdominal feet,

and the absence of the branchiz assign to this Schizopod its place among the Mysidee. It

differs from them only by the free dorsal shield and the presence of breeding-lamellz on

seven appendages, which it has in common with the Lophogastridee. It is also sur-

prising to find that a member of this latter family has a Mysid character (the subjoints

of the legs in Gnathophausia), which, however, is only a peculiarity of one of its generic

groups (genus Mysis and subgenera). It accordingly stands in the same relations to

Myside as Gnathophausia stands to Lophogastridee—there being in both a character

which shows some distant relationship to the other group, and both having a free cara-

pace, which is not to be found in the hitherto known Schizopods.

Among Myside there are two generic groups :—Wysis and the genera connected with

it, characterized by joints in the tarsal parts of the legs and the absence of branchial

appendages on the pleopoda; and S%riel/a, in which the legs are simple and terminated

by a claw, as in our case, but in which there are branchial appendages on the pleopoda,

which Petalophthalmus has not got. This genus therefore belongs to none of these

generic groups, and can only be united with the Myside if the family characters are so

far enlarged as to allow the entrance within it of an animal which has a free dorsal

shield and breeding-lamelle on seven appendages.

The eyes having disappeared, and the eye-stalks having assumed a spherical termi-

nation, are of course entirely secondary characters, and probably not even of generic

value.

Characters of the Genus.

PETALOPHTHALMUS*, nov. gen. 4. .

Carapace in no connexion with the five posterior segments of the pereion; rostrum

very short; three joints in the funiculus of the first antenne; scale of the second

antenna without spines; labrum subcordiform ; mandibule having a large manducatory

process and a three-jointed palpus. Very strong teeth and setze on both processes of the

first maxilla; second maxilla consisting of a manducatory portion (two elongated pro-

cesses and a larger lamellar one), a ciliated flagellum, and a palpus. Maxilliped in

both sexes with a lamellar appendage on one of the basal joints ; first gnathopod having

the same only in the male; in the female it is elongated, and its last two joints are

clumsy and recurved; second gnathopod and the five pereiopoda leg-like; pleopoda in

the female quite rudimentary, with only one ramus; in the male somewhat more per-

fect, with two rami; telson showing a slight excision at the top; outer appendages

jointed.

’ No branchiz on the pereion, nor any branchial appendages on the pleopoda of the

male. Seven pairs of breeding-lamelle at the base of the pectoral appendages.

* Tléradoy and d¢Oadpos.

G 2

4A, DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

The male differs from the female by the rudimentary condition of its carapace and the

metamorphosis of the first antennz, the palpus mandibularis, the maxilliped, and first

gnathopod into prehensile organs.

PETALOPHTHALMUS ARMIGER, N. Sp.

Eyes wanting. Eyestalks with spherical concave terminations at the place where one

would expect to find the eyes. Length 37-44 millims.

Habits of Life, and Colour.

Petalophthalmus is a member of that deep-sea fauna which we sketched when de-

scribing Gnathophausia zoéa, in which, however, it seems not to be common. Its colour

is a bright red.

We have now described (1) a lophogastriform genus (Guathophausia), with three

species, (2) a genus allied to the former, but forming a new family (Chalaraspis), and (3)

a Mysidiform genus (Petalophthalmus)—all of which have a free carapace and certain

peculiarities which show that their existence dates from a period in which the family

characters of Schizopods were not so sharply defined as they are now. They are pre-

cursors of the Myside and Lophogastride, and representatives of another family, having

characters which, among the Schizopods, are only shared by Nebalia, but in all other

respects entirely different from this genus. Now the idea presents itself whether it would

be advisable to bring all these Schizopods together into one group, the chief and only

universal character of which would be that they all have a free dorsal shield. We could

best describe this in the following synoptical way :—

(9 abdominal segments, palpus maxille prime ..................-. NEBALIA.

( Tlegs, subjoints, palpus maxille prime, supple-

Group I. | mentary eyes, Lophogastriform ............ GNATHOPHAUSIA.

Dorsal shield free. < :

(aleilomuael Seg 1) g legs, no subjoints, 7 breeding-lamelle, Mysi-

BEEN: COOH cconcconoas0oucc08 Pacgansodmeooas PETALOPHTHALMUS.

\. 4 legs, no subjoints, branchiz as in Lophogaster.. CHALARASPIS.

C (7 legs, no subjoints, no palpus maxille prime,

Group II. no supplementary eyes...............----- LopHogastER.

ove elite 228 7 abdominal seg- } 6 legs, with or without subjoints, 2 breeding-

tened to the )

ments. lena samemiaaie s ucian digit eee ee ore ooo Sole Mysis and Srrierra.

pereion.

8 legs (the last wanting or rudimentary), branchiz

| free, supplementary eyes...........-.....-. KUPHAUSTA.

This arrangement of the Schizopods would, no doubt, have a certain advantage, as it

prevents the destruction of the family characters of our old and well-known forms in

favour of a few deep-sea genera, all of which have a certain peculiarity which they share

only with Nebalia. But it is not a natural genealogical arrangement; for, from what

has been said, there is no doubt that Gnathophausia is, if 1 may use the expression, an

ancient Lophogaster, and Petalophthalmus an ancient Mysis. I therefore propose to

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION. 45

unite these with the families of the shallow-water forms, and would suggest for the

Schizopods the following systematic arrangement :—

9 abdominal segments, 8 legs ...........-.- 02 -eeeer creer eens I, NeparupZ.

Ghlogs cee opersnctsboieyceslsversats ckovche etevathu ay before IL. Mysinz.

: Sep Sk eectausiegsscvsrssveiai ness ler Syaueuaveos'cseiako eleyais Ill. EvpHavsiipz.

7 abdominal segments, A NEBR hte sisheccyoreiae forse celeaies sare wate ate IV. CaanarasPipz.

MUG RS rviareys ae cools eee Sera lan betel wae VY. LopHogasrripx.

In order to expose the genealogical connexion between the genera of these families,

it would be necessary to prepare a large table; but I think that their relations will be

well understood from what I have said about the subject in the preceding pages, as well

as from a comparison of my plates with the figures given on the other genera by Sars,

Claus, and other authors.

Two questions will arise, of course, from a comparison of the deep-sea forms with the

shallow-water forms:—(1) Are there no Schizopods in’ the depths which show the

ordinary conformation of a shield fastened to the pereion? and (2) As the Mysidee and

Lophogastride have relations of very extraordinary characters living in the deep sea,

are there no forms which represent the third of the hitherto acknowledged families—the

Euphausiide? These questions are partly answered by the discovery of a very large

member of this family, which came up in the trawl in lat. 35° 41’ S., long. 20° 55’ W..,

from a depth of 1900 fathoms. This animal is a male, has a length of 84 millims., and

belongs to Dana’s genus Euphausia; for its dorsal shield is perfectly fastened to the

pereion, and of the eight pairs of legs only six are developed, the last two being only

represented by the branchial appendages and the palpi. But, unlike all other species

of Huphausia hitherto known, it has no accessory eyes. There are very long olfac-

tory hairs on the enlarged first antennze, and the peculiar appendages to the pleo-

poda which characterize the males of this genus. I shall figure and describe this

species, which I intend to call Huphausia simplex, n. sp., when we come home, and

when it will be possible to compare it with the other species of the genus (which have

been described by Dana and Claus). From the discovery of this Huphausia, it appears

that not all the deep-sea Schizopods offer the said peculiarity, and that, among very

peculiar genera such as have been described above, a form is also to be found which

differs from the surface-Huphausie only by the absence of accessory eyes. ‘These, I

think, are to be considered a secondary adaptation to pelagic life; and therefore 2. sim-

plex is probably a more ancient form than #. Wiilleri, splendens, and superba. Probably,

however, this will not be the last discovery we make in this interesting group; and I

think it is very likely that a form will come up which is in the same relations to the

Huphausiide as Gnathophausia is to the Lophogastride, and Petalophthalmus to the

Mysidez. Then we should have made one step forward in the knowledge of Schizopodal

genealogy, a step which will probably lead to further discoveries concerning the rela-

tions of the Phyllopods to the Schizopods—of the lower to the higher crustaceans.

Perhaps it will now also be possible to understand certain fossil forms which seem to

be allied to Nebalia and to Gunathophausia.

46 . DR. R. V: WILLEMOES-SUHM ON SOME ATLANTIC,

V. On-THE DEVELOPMENT OF A LAND-CRAB.

(Plate XI. figs. = 3:

In most books on natural history we find concerning the land-crabs a remark that,

according to Mr. Westwood’s observations, they migrate to the sea when the time for de-

positing their eggs has come, and that the young ones, when coming out, have the same

form as their parents. The same thing isto be observed in Astacus fluviatilis; and the

knowledge of these two facts has been much in the way of the general acceptance of

Mr. Vaughan Thompson’s first observations on the Zoéa-stages of crabs. Also his state-

that he found Zoéa-brood in the eggs of a Gecarcinus has been doubted ; for, according to

Fritz Miller *, Mr. Bell considered himself justified in eliminating Thompson’s observa-

tion, because he could only have examined ovigerous females preserved in alcohol. The

following lines, however, will show that there is no reason to doubt Mr. Thompson’s

statement. Fritz Miller himself, though we owe him many observations on land-crabs,

says only that there is Zoéa-brood in Ocypoda and Gelasimus, but had no occasion to

observe the Gecarcinoids.

Watching the habits of these crabs is always attended with some difficulties, as De

of them are nocturnal. In St. Thomas I only succeeded in getting a few Gelasimi.

Only in the Bermudas did we get Gecarcinoids:—the large Cardisoma guanhumi,

which was caught by torch-light in the interior of the island, but all the specimens we

got were males ; besides, Mr. Moseley caught Gecarcinus lateralis and Ocypoda rhombea,

and I myself caught several crabs allied to Boscia. 1 watched there also for a long time

the lively Grapsus cruentatus, which lives in great quantities in Hungry Bay, among the

mangrove trees. It has entirely taken to terrestrial habits, makes deep holes and runs up

and down the mangrove trees. ‘he holes, however, it always makes so near the shore,

that it is sure to find water at a depth of 2 or 3 feet. What its mode of development

is I could not find out, as it was evidently not their breeding-season (June). Arriving

in the beginning of August at the Cape-Verd Islands, I found in St. Vincent many holes

on shore inhabited by Ocypoda hippeus, a crab running with marvellous velocity over the

sand, and very valuable to me, as it gave me ample opportunity of studying the interest-

ing entrance to the branchial cavity described by Fritz Miller. But also in these I could

discover no trace of eggs or of young. Holes: made by the same crab we found

again in the island of San Jago, where they are to be seen in abundance near the

houses of the little village which surrounds the old cathedral of Ribiera Grande. But

besides Ocypoda hippeus, we got in San Jago another large land-crab, caught by Mr.

Moseley’s seining-party at night in the bay of Porto Praya, a true Gecarcinoid, be-

longing to the same genus (Car disoma) to which the Bermuda ones belonged. Unfor-

tunately I cannot make out its specific name, as it is not mentioned in Milne-Edwards’s

‘ Histoire Naturelle des Crustacés’ +; but I suppose it is an animal well known to Euro-

pean carcinologists as inhabiting the Cape-Verd Islands, and, very likely, also the coast

* «Facts and Arguments for Darwin,’ by Fritz Miller (translated by W. 8. Dallas. London: 1869), p. 48.

+ Probably described in the same author’s ‘ Crustacés des fles du Cap Vert.’

CRUSTACEA FROM THE ‘ CHALLENGER” EXPEDITION. AT

of the African continent. This Cardisoma inhabits large holes all along the shore,

especially in the palm-groves on both sides of the town. But when I went there the next

day to catch more of them, the difficulties in getting them were too great; for the holes

were more than 8 feet deep. The one, however, which Mr. Moseley had caught, a

female, proved to be very interesting; for after having killed it in spirits, I found the

next-day that its abdominal feet were covered with strings of eggs, that most of these

eggs were empty, some of them containing an embryo, and that among the empty eggs

there was a good number of young animals newly come out. These young ones, how-

ever, were not like their mother, but Zoéas. The greater number of them had evidently

left the mother already; some were still to-be found among the empty capsules; and

others had not yet emerged. In fig. 3 Mr. Wild has drawn me sucha string of eggs

(each 0°42 millim. in diam., having the form of a grape), in which the berry-like

peduneulated eggs surround a common axis.

The Zoéa of Cardisoma (fig..1) leaves the eggs in a somewhat more advanced state

than the Zoéa of Carcinas menas. If we compare it with the figures given by Mr. Spence

Bate (‘Philosophical Transactions,’ 1858, pl. xl.), we find that it represents a middle

stage between the larva of Carcinus menas which has just left the egg and the one

after its first moult (figs. A and B). With the former it has in common the want of the

frontal spine, with the latter the presence of setze on most of the appendages, and the

more developed caudal fin.

The carapace is very globular, and its dorsal spine is only small. The frontal spine,

entirely wanting now, is probably only developed after the next moult. The eyes are

very large. I could not see the pedunculus; but as I had only dead animals before me,

which had been for some time in spirit, I do not deny that a very short pedunculus

might exist. The first antenna has a few more hairs than in the second stage of

Carcinus menas; but the second ones are very much like each other in both cases (a'

and @’, fig. 1). The parts of the mouth (m) had equally already got their hair. I had

some difficulty in getting a side view of them, and could have only isolated them with

very great difficulty, which was not worth while, as there was no chance of getting

later stages of this larva. The two gnathopoda have four long hairs in the last joint,

which in our case (gp' and gp’) are not so long, and not feathered, as they are in the

second Zoéa-stage of Carcinus menas. I have not seen any buds of the pereiopods

behind those two legs, which are present in the young Carcinus menas.

Each segment of the pleon presents some black pigment spots, some of which were

also-observed in the carapace. There are no appendages on the pleon. The last segment

is bifurcate; and at the inner side of the two terminal pieces six feathered setae may be

observed, very much like those in fig. 21” of Spence Bate’s larva.

I think there is no doubt that these Zoéas, as they partly had done already, leave the

mother and lead a pelagic life until they have undergone all their metamorphosis ; and

IT hope that in other parts of the world which we are about to visit, I shall be abie to

complete these observations on the propagation of the land-crabs, and to make out in

which genera there is a metamorphosis, and in which not.

I may add here an observation which I made some years ago in Italy, and which,

48 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

if not quite directly concerning our subject, at least touches it; for the animal I am

to speak about is the freshwater crab of Southern Europe, Zelphusa fluviatilis. In the

mountains above Spezzia I caught some specimens at a spring coming out from the Bocca

Lupara; and one of these being a female, carried with it nearly 150 young ones under

the abdomen, the smallest of which had a length of 3 millims. I think this shows clearly

that also Telphusa is, with Gecarcinus and Astacus, one of the few Decapods which

develop directly without metamorphosis.

VI. ON A BLIND DEEP-SEA ASTACUS.

(Plate X. fig. 1.)

In a very successful haul on the 15th of March near Sombrero Island, W. I., we got,

from a depth of 450 fathoms, a blind Astacus, one of the claws of which is developed to

an extraordinary extent. In the notes which I gave about it to Professor Thomson, I

have called this animal Astacus zaleucus ; for, strange as it may seem at first sight, there

is no reason for separating this animal from the genus Astacus, except perhaps the want

of the eye andits pedunculus. The male of Astacus zaleucus has a length of 110 millims.,

the cephalothorax having a length of 50, the abdomen of 60 millims.

The carapace is laterally very much compressed, and divided by a deep transverse

sulcus into two portions or regions. At the top of the anterior one is the rostrum,

having a length of 9 millims. Its borders have six small teeth. Before the rostrum there

are two large triangular patches, which are densely covered with short hairs; and these

continue from both sides towards the rostrum. On both sides of the anterior portion

there are besides several small spines, and two sulci, which establish again a small mar-

ginal and triangular subdivision. The posterior portion of the carapace can also be

subdivided, into a quadrangular upper portion and two lateral regions. The last segment

of the pereion is in articulation with the carapace, quite in the same way in which it is

in all Astaci,

On both sides of the rostrum there are two qaean places, where in other Species the

pedunculus of the eye is fastened. Here there is not a trace of it.

The four-jointed funiculus of the antenna interior bears two very hairy flagella, of the

length of 40 millims, The base of the antenna exterior terminates on one side in a sharp

spine. The squamiform appendage is somewhat curved outwards, reaches the middle of

the second funiculus-joint, which is rather long, and has at the inner side eight sharp

spines, The flagellum of the antenna exterior has a length of 130 millims. Underneath

its base the large opening of the ‘‘ green gland” is clearly to be seen.

The margin of the upper lip is somewhat tilted upwards, and beset with 6 small

spines. About the mandible I have nothing particular to say. The chewing-plates of

the first maxillee are very strong, and the palpi of the maxillipedes very large. They look

nearly like a ramus of one of the swimmerets. The palpi of the two gnathopods,

however, have the ordinary form. The second gnathopod has thrice the length of the

first one.

The pereiopods are all very hairy, especially their first two joints, which, with the

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION. . 49

exception of the right claw, are all densely covered with hairs. To three pairs there are

chele; the fourth has a small recurved claw at the end; and in the fifth the sixth joint

is nothing but a very small stump. Very remarkable are the first pair of pereiopods; the

chela of the right one has a length of 100 millims., and that of the left one of only

48. Between the two chels, however, there is not only the difference in length, but

also in width, as the carpus in the right one is very much expanded, spiny, and

hairless; in the left one it is even and covered with hairs and spines. The two digiti

chele have large teeth at their inner side, 57 on the digitus mobilis, and 62 on the oppo-

site one; so that the claws of the larger pereiopod look like the jaws of some ferocious

reptile.

The genital openings are here at the base of the third pereiopod, where we also found

them in the male of Deidamia leptodactyla.

Between the last pairs of pereiopods there is a small triangular pectoral plate.

The pleon is very flat; and the sharp angles formed by a longitudinal line on both

sides give it nearly a square shape. Its lateral pieces do not terminate in sharp spines,

nor are they very large; the edges of these, as well as the whole surface of the third

and fourth segments, are covered by dense and short hairs. The telson is a square, as

it is in many Astaci; its lateral appendages are very large, and extend, like wings, to

both sides.

The pleopoda of the first, very short segment are styliform appendages; the other

pleopods have a very solid unjointed base and two broad rami, the interior of which is

longer than the exterior (fig.1q@). All these, as well as the caudal appendages, are

densely fringed with hairs.

, Characters of the Species.

ASTACUS ZALEUCUS, 0. Sp.

Hyes and eyestalks wanting entirely. Chelee of the first pereiopod unequally developed,

the right one having more than double the length of the left one. Hight spines at the

squamiform appendage of the outer antenna. Genital opening of the male at the base of

the third pair of pereiopods.

Those who consider the eyestalk an appendage of a separate segment of its own

would probably be inclined to make another genus of this form. But I think this could

be the only reason for separating it generically from Astacus, and for me not a sufficient

one. We find in Deidamia that the genital opening of the male is, in one species, at the

third pereiopod, and in another, undoubtedly belonging to the same genus, at the fifth.

I therefore think that also this cannot be a reason for separating it generically from

Astacus, in which genus this species will be the first one made known as an inhabitant of

the sea.

To the Astacus of the Mammoth-cave this crayfish has only a physiological resem-

blanee; for even in those species in which the eyes are abortive the stalks at least are

present. But we have another form, which we got in deep water near the Bermudas,

in which the eyes have the same hidden position as in A. pellucidus, to which it has in

all other respects an extreme likeness; only in the marine species the squamiform ap-

SECOND SERIES.—ZOOLOGY, VOL. I. H

50 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

pendage of the outer antenna is entirely wanting. The same thing is the case in a

crustacean recently described by Mr. Wood-Mason from deep water near the Andaman

Islands, which he calls Nephropsis Stewarti*. Iam inclined to think that the crusta-

cean which we got near the Bermudas belongs to the same genus; but as I have not

seen the plate which Mr. Mason has added to his original paper, I shall defer its descrip-

tion until we come back.

Colour, and Mode of Life.

A. zaleucus came up on the swabs of the dredge, together with the large chele of

another, smaller specimen, the body of which was lost. The one we got had the red

colour of all deep-sea Crustacea. It lived on a bottom of globigerine ooze, evidently

frequented by a great many animals. Several sponges and seven echinoderms (Ophio-

musium, Luidia, Archaster, Astrogoniun, Oidaris, and Hchinus), a Planularia, Mopsea,

and an Jsis, several Annelids and a Sipunculus, a Galathea, a Peneid, an Arcturus, and

the fine crustacean to which was given the name of Willemoesia crucifera—four Bryozoa,

a Dentalium, and many small shells—finally, a fish probably belonging to the genus

Chauliodus, were got in the same place with this extraordinary Astacus. Unfortunately

our stay in the West Indies was only a very short one. The few hauls, however, which

we had near Sombrero Island and St. Thomas showed us that a great quantity of new

and probably also interesting animals live there in moderate depths (800-400 fathoms).

VIL. ON Witrmorsra, A DEEP-SEA DECAPOD ALLIED TO Hrvon.

(Plate XII. fig. 10, and Plate XIII.)

In the “ Notes from the Challenger,” sent by Professor Wyville Thomson to ‘ Nature,’

two blind crustacea have been described, both of which were found on our cruise from

Teneriffe to St. Thomas. The larger one of these remarkable animals was called by me

Deidamia leptodactyla, obtained on the 4th of March, by the dredge, in a most perfect state,

from a depth of 1900 fathoms, in lat. 21° 38’ N., long. 44° 39° W. Another specimen

was caught in the trawl in lat. 35° 41’ S., and long. 20° 55’ W., about 400 miles W. of

Inaccessible Island (‘Tristan d’Acunha), from the same depth at which the first was got.

The second, smaller species, which I proposed to call D. crucifera, was got on the 15th of

March, close to Sombrero Island, West Indies, from a depth of 450 fathoms, and equally

well preserved. I shall now give a description of both of them, but especially of the

larger one, which may be considered the type of the genus, and of the smaller one as far

as it offers any differences of importance.

The generic name Deidamia, however, must be dropped, as (according to Mr. Grote,

who has done me the honour of proposing for it the name Willemoesia) it was given

by Dr. Clemens in 1859 to a genus of North-American Sphingidee.

1. WILLEMOESIA LEPTODACTYLA.

The body of this crayfish approaches in form closer to the well-known fossil qa

than any other crustacean known to me, the body being flattened above, and there being

* “On Nephropsis Stewarti, a new Genus and Species of Macrurous Crustaceans,” by James Wood-Mason, ‘ Annals

and Magazine of Natural History,’ No. 67, July 1878.

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION. 51

no lateral compression as in Astacus or in Palinurus. The cephalothorax exceeds in

width half of its length; and the abdomen too is very flat. The whole animal, being a

male, has a length of 120 millims.* (Plate XIII. fig. 1.)

The cephalothoraz has a length of 60 millims. by 33 in width at its base. Its upper

lateral edges are very sharp, and formed by a series of lateral spines, those of the under-

side, however, rounded. The whole cephalothorax, owing to this configuration of the

carapace, is square, the upper surface being wider than the under one, and somewhat

convex, the side parts being not rounded as in Astacus but perfectly flattened ; also the

upper surface forms a square, the angles of which are terminated anteriorly by two

spines, and are posteriorly rounded. A line of spines runs longitudinally through the

midst of the carapace, and divides it into two portions, being terminated anteriorly by

a sharp spine. The lateral spiny edges are divided by small but clearly perceptible

fissures into four portions, the first of them containing 8, the second 5, the third 5,

and the last 6 larger spines. A deep transverse sulcus divides the carapace into an

anterior and a posterior portion, which by secondary elevated lines are again divided

into several regions, which we might call the stomachal, hepatic, lateral, cardiac, and

branchial regions, each of which will be easily seen in the drawing (fig. 1). The inner

borders of the carapace of the under surface are perfectly smooth; and the rounded exte-

rior edges of the same surface are covered by a fringe of long and beautiful hairs. The

whole carapace is covered by very small spines, all directed forward, giving it a certain

likeness in appearance to the well-known chagrin of the sharks.

The eyes are entirely wanting ; nor is there, as in Astacus zaleucus, any place left open

where you might expect to find them.

Immediately underneath the frontal edge of the carapace we find in a prominent place

the wing-like expansions of the joints, funiculi antennarum anteriorwm. They are (espe-

cially at the inner side) densely covered with hairs, and very closely approach each

other. The funiculus consists of three joints, and has a length of 10 millims. Its in-

ternal flagellum, consisting of very many rings, has a length of 65; while the external

is very short, being only 10 millims. in length. On the same line with the inner antennee

and close to them, are the outer ones, the funiculus of which is a little more elongated,

and consists equally of three joints (12 millims. in length). The squamiform appendage

is very small (8 millims. in length by 2 in width), lancet-shaped, and inserted at the base

of the first joint. The flagellum is 60 millims. in length.

The opening of the mouth is angular (fig. 4); and you find above its rejected border a

small tuft of hairs. I could not find any labrum. The mandibule, covering nearly the

whole opening with their strong manducatory processes, have behind these a strong and

hairy palpus (fig. 5, pm), and at their base the two slender organs (fig. 5, Jab) which repre-

sent the labiwm or under lip. The first maxilla (fig. 6 a) shows avery small rudiment of a

flagellum (1), which in the second mawilla (fig. 6 b) attains so considerable dimensions.

The maztluped has two manducatory processes (fig. 7), a palpus (fig. 7, p), and a

flagellum, which is divided into two portions, the upper one (jl’) being attached to

the palpus, and the under one to the base of the maxilliped (/?).

* This description refers to the larger of the two males procured.

52 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

The first guathopod (fig. 8), as well as the second (fig. 9), are terminated by a denticulated

claw, and covered by hairs at their inner side. It is quite certain that the gnathopods

(or the second and third maxillipeds) have no palpi.

The pereiopods (fig. 1) are all terminated by chelee, but of very different sizes. The length

of the first pair, enormously developed, is 155 millims., thus exceeding by 35 millims. the

length of the whole body. The basal joint (the coxa) is very large and flat, the second,

however, very thin at its articulation, bent somewhat backwards, and increasing in size

only at its top, where it articulates with the broad base of the third joint. This is the

longest one, being covered on its upper surface by a series of small spines, and terminated

at its top articulation by two recurved spines. The fourth joint is again very slender;

and so is also the delicate chela, the immovable finger of which is armed with a large

transverse spine. This first pair of pereiopods are perfectly smooth ; all the others, however,

are covered with hairs up to the top. They are much smaller and clumsier, the second

pair having only a length of 39 millims. At the coxal joint of the third pereiopod

are the genital openings.

As to the underpart of the cephalothorax, it may still be mentioned that there is no

distinct opening for the so-called green glands, that there is a strong spine between the two

maxillipeds above the entrance of the mouth, and that an elevated line, covered with

a fringe of hairs, runs over the sternal shield between the first pair of pereiopods.

The abdomen (pleon) has a length of 60 mm., and consists of six segments and the

telson. The top of the second to fifth segments is marked by a strong spine, which is

bent forward, and which in the first and sixth segments is only represented by a small ele-

vation. There are also in the first segment no squamiform lateral appendages, which in

the subsequent ones are very large, but decrease in size from the second to the sixth. All

of them are fringed with long hairs; and so are also the telson and the caudal appendages,

the form of which can best be seen from Mr. Wild’s excellent drawing.

The first pair of pleopods is transformed to styliform appendages (fig. 2), our specimens

being both males. The four subsequent ones have a length of 33 millims., and are evi-

dently very powerful organs of locomotion. They consist (fig. 3) of a very strong basal

joint, to which the external palpus is attached, of smaller second joint, which goes

laterally over into the second interior palpus, and of a very small and styliform third

joint (fig. 3, a)—quite the same arrangement which we find in W. erucifera and in an un-

described Palinurid from the deep sea near the Bermudas (fig. 12). All the four pleopods

are alike; and the posterior ones are very little smaller than the anterior ones.

Of the internal organization I can only say that the carapace is wonderfully trans-

parent for the size of the animal, and permits the heart to be seen, which is situated

just between the hepatic and stomachal regions.

2. WILLEMOESIA CRUCIFERA, n.sp. (Plate XII. fig. 10, and Plate XIII. figs 10 and 11.)

The species we have hitherto considered is very delicate, and has very long fringes

of hairs on the legs, the carapace, and the pleon. All this is less the case in the

smaller species, which is much more solid, not transparent at all, and in which the

fringes are not so large and beautiful. Nevertheless also this one is a very elegant

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION. 53

creature, which on the whole has a strong family likeness to W. leptodactyla. It has

a length of 42 millims. x

The form of the carapace is nearly the same as in the former species. It has a length

of 19 millims., by 18 of greatest width; at its base, however, it is only 10 millims. wide.

The denticulation of the edges is a much stronger one than in the preceding species;

there is a line of spines divided by two incisions into three divisions, the first of which

contains 7, the second 5, and the third 17 spines on each side, all bent forward and

somewhat upward. Two very prominent lines, one longitudinal, ending at the front with

a spine, and one transverse, form the figure of a cross; and following the lines of

caruncles, you might trace the same regions which we have described in the former

species, but which in this case are not so clearly visible. At the frontal border there are

two spines on the right side and four spines on the left side of the median spine. The

whole carapace is finely covered by very small spines, which look like very fine chagrin.

The interior antennz have several spines on the inner side of their three funiculus-

joints, but not such a wing-like expansion as we have observed in the former species.

The exterior flagellum has a length of 17, the interior of 7 millims. The exterior

antenna does not offer any thing particular.

The parts of the mouth have exactly the same form as in the preceding species ; and

the absence of palpi at the base of the gnathopods can also be stated in this case.

Of the five pairs of pereiopods only four are terminated by chele ; and this is the chief

difference from W. leptodactyla. The first pair also are proportionally much shorter,

(only 34 millims. long), and have no spine on the slender fingers of the chela. In the

third and fourth pairs of pereiopods the chele are much more slender than in the first.

The last one is terminated by a simple recurved claw. The last four pairs of pereiopods

are covered with hairs, but not so densely as in WV. leptodactyla. The genital opening

is, in this species, not on the coxal joint of the third pair of pereiopods, but, as is usually

the case, on that of the fifth.

The abdomen has a length of 23 millims. Its first segment bears one spine; and all

the following ones have two spines on a little prominence. Telsen, caudal appendages,

and lateral processes of the segment are fringed with hairs.

The first pair of pleopods is transformed into two styliform and flattened appendages.

In the second pair (fig. 10) we observe the peculiarity that the third styliform joint is

covered by another, smaller joint, the morphological value of which I do not understand.

In the other three pairs (fig. 11) this supernumerary joint is not attached to the third one ;

so that these are very much like the pleopods of W. leptodactyla.

Colour, and- Habits of Life.

Especially the larger of these two species was very beautiful when it came up; for it

was of a fine red, while the hairs bordering it were yellowish. The smaller one also was

red. This colour, however, rapidly disappeared when the specimens had been for a

certain time in spirit.

As was also the case with Gnathophausia, the larger species of the two was caught im

very deep, the smaller one in shallower water. W. leptodactyla was found on a plateau

54 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

which probably extends southwards from Greenland, and on which in the north are

situated the Azores, and in the south Tristan d’Acunha. Before getting to it from

Teneriffe we had the greatest depth of that section of the ocean, 3150 fathoms ; and after-

wards again we gradually came down to a depth of 3000 fathoms before getting into the

shallower water of the West Indies. On that plateau we got with the first specimen a

sponge, some Bryozoa, a Brachiopod, a Lamellibranchiate, a Gastropod, and a small

Shrimp, besides this magnificent Decapod, which was entangled in the swabs of the

dredge. With the second specimen a great many animals came up, among which I may

mention a Palythoa, a Fungia, and a Pennatulid. A Holothuria, an Archaster, and an

Urchin represented the Echinoderms. Crustacea were present in large numbers—espe-

cially Peneid and Caridid Shrimps and the Schizopods which I have called Petalophthat-

mus and Chalaraspis, together with a large species of Huphausia. The smaller species

was got near Sombrero Island, W.I., ina depth of 450 fathoms, together with Astacus

zaleucus ; and in describing this species I have already given an idea of the fauna which

lives in the same place with these crustacea.

Characters of the Genus, and Systematic Position.

WILLEMOESIA, Grote *.

Carapace large, flattened, and quadrangular. Anterior margin and lateral borders

denticulated, the latter divided by two incisions into three denticulated portions. A

longitudinal line divides the carapace into a right anda left portion. Hyes and eye-stalks

entirely wanting. Funiculus of the first antennze rather large, its inner flagellum longer

than the outer one. Small lamellar appendage at the base of the second attenne, the

flagellum of which has the same length as the longer one of the inner antennz. Man-

ducatory portion of the mandibula very strong. A large flagellum attached to the second

maxilla. The palpus and flagellum of the maxillipedes reach the base of the first

antennee. No palpi at the base of the gnathopoda (second and third maxillipeds). First

pair of pereiopoda very elongated, terminated by a slender pair of chele. Subsequent

pereiopoda much shorter.

Small styliform appendages at the inner side of the ramus interior of the pleopoda.

Abdomen flattened, divided by a longitudinal line of spines into aright anda left portion.

Telson rapidly decreasing in size, longer than the shield-like caudal appendages.

1. W. LEPTODACTYLA, 0. sp.

Lateral borders of the carapace not very much expanded. Denticulation not very deep.

First pair of pereiopoda longer than the body.. All the pereiopoda terminated by chele,

Length 120 millims.

2. W. CRUCIFERA, 0. sp.

Carapace with wing-like expanded lateral borders. Deep incisions and strong den-

ticulation at its lateral borders. First pair of pereiopoda shorter than the body. Only

four pereiopoda terminated by chele. Length 42 millims.

* Tn ‘Nature,’ October, 1873.

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION. 55

Among the living Decapoda Macrura there is hardly a group with which Willemoesia

could be said to be very closely allied. Nearest to it are undoubtedly the Scyllarine ; but

these, like all the genera of the family Palinuride, differ from it in the absence of the

lamellar appendage of the second antenne, and in the presence of palpi at the base of

the gnathopoda, which, as we have seen, are wanting in this new genus. Nor can it, for

this latter reason, be referred to the Astacidee, with which it has in common the presence

of the antennal scale. ©

It is very astonishing, indeed, that, among all crustaceans known to us, Willemoesia

approaches most closely the fossil Eryontide. If we compare, for example, our figure

of W. crucifera (Plate XII. fig. 10) with a figure of Hryon arctiformis, and the description

of the “Tribu des Eryons,” given by Milne-Edwards* (and probably taken especially

from Desmarest’s ‘ Crustacés Fossiles’), we find most striking resemblances between the

two forms. In W. crucifera as well as in Eryon the carapace has nearly half the length of

the whole body; and in both forms its lateral borders are wing-like expansions which

are divided by two deep incisions into three portions. The anterior border of the carapace

is nearly straight in both forms.

Eryon was probably not blind ; for the eye-stalks have been found in several specimens.

Its antennz seem to be somewhat more reduced than in Willemoesia; but the second

pair of them has, according to Desmarest, “ une écaille assez large, ovoide et forte-

ment échancrée.” This is the chief difference between Hryon and the Palinuride, and

the same in which also Willemoesia differs from that group.

Milne-Edwards says nothing on the parts of the mouth; but according to Quenstedt

they had a very large mandibula, one of the teeth of which was preeminently strong.

This is very much like what we find in Willemoesia; but in the fossil genus palpi were

present at the base of the first and second gnathopods, which are wanting in the living

genus. ‘The first pair of pereiopoda is in both forms longer than the following ones, and

terminated by a pair of long and slender chelze. In Hryon three pairs of pereiopoda, in

W. leptodactyla five, and in W. erucifera four are terminated by chelee. The form of the

last pereiopod in Z#. arctiformis is exactly the same as in WV. crucifera; and the abdomen

of these two forms is, as the above-mentioned figures show, so very much alike in the

two forms, that, if the last pair of pereiopoda and the pleon of Hryon were presented to me

without my knowing to what they belonged, I should undoubtedly declare them to be

parts of the genus Willemoesia. There are the same line of spines at the top of the

rings, the same winglike expansions on both sides, and that characteristic “ nageoire

caudale, dont la lame médiane est pointue et les quatre lames latérales moins longues

que la médiane et hastiformes.”” Also the fine fringe of hairs which distinguishes the

caudal fin of Willemoesia is to be seen in the fossil crustacean.

Eryon differs from the living genus chiefly by the presence of eye-stalks and of palpi at

the base of the gnathopoda. According to Quenstedt the latter were observed only with

some difficulty ; and their presence seems not to be beyond all doubt. I shall only on my

return be able to look myself over the original specimens and papers, and then, I hope,

be able to give a more detailed account on the relations of Willemoesia to Eryon.

* Histoire Naturelle des Crustacés, tome ii. (Paris, 1837) p. 278.

56 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

That they must be very close appears already from what we have said; and among the

Eryontidz this new deep-sea genus must take its place. The Eryontide, now consisting

of two genera, must either form a new family intermediate between the Astacide and

Palinuridee, or be united with the latter.

During our cruise in the Atlantic we got deep-sea animals (7. e. from more than 350

fathoms; stations where only Foraminifera-shells have been got are not included) from

67 stations. After every haul a list of these animals, as much specialized as our know-

ledge of the thing allowed us to do, has been put down by me in the so-called Station-

book. As it might be of some interest to know how often the different groups of animals

have been got, I have made the necessary additions, and shall terminate this paper by

giving the results as far as Crustacea are concerned. According to this,

Cimipediathavesbeenscot momma ilem na enn WUIGIES:

Ostracoda hen cn t.. Mecy tat ee ie mace oem RCL

STO MVOC. one | Gilkaniol eich. Surin womb candy 9 C HMNES.

Wsopoday, | ei oa) eu 26 FR el oe en

Cumiaced OU OA OD COT OREO Te

Stomatopoda. . ........ . J time.

Schizopoda, |. . . «°% 5 =. « =.» -12\times.

Decapoda Macrura - 4) so. 5 2 ss . 22-5

IDecapock lke, 5 8! 556 6 os eo Wily

EXPLANATION OF PLATES.

Prats, VI.

All the figures, with the exception of fig. 4, refer to Nebalia longipes.

Fig. 1. A female. Low power. a, antenna of the first pair; 6, antenna of the second pair; Oc, eye;

R, rostrum; M, stomach; D, intestine; F, joint of the furca.

Fig. 2. A male. Letters the same as in fig 1. d, palpus of the first maxilla (‘‘ Putzfuss” Claus) ;

G, ductus eiaculatorius.

_ Fig. 3, Leg of a female: a, basal joint with the branchial appendage (K, A) ; 8, second joint with lateral

appendage (NN, A); y, main branch.

Fig. 4. Phyllopodal foot of the female of N. Geoffroyi, copied from Claus. Letters the same as in fig. 3.

y—-v, Joints of the main branch.

5. First antenna of a female. &. fourth joint, a claw. a, External, 6, ternal flagellum.

6. First antenna of a male. Letters the same as in fig. 5.

Vig. 7. Second antenna of a female.

8. Mandibula; p, palpus

9. First maxilla; p, palpus of a female.

Tig. 10. Second maxilla

Fig. 11. One of the anterior pleopoda: a, basal joimt; @, ramus exterior ; y, ramus interior; 6, retina-

culum corresponding to the same on the other side.

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION.

Fig. 12. Fifth

Fig. 18. Sixth } pleopod of a female.

Prats VII.

All the figures refer to Petalophthalmus armiger. All drawn with very low power,

except figs. 7-9.

Fig. 1. A female, x 2 nat. size. a’, first antenna; ap, appendix lamellaris; a, second antenna;

pm, palpus mandibule ; mp, maxilliped; gn, gnathopoda; prp, pereiopoda; Ja, breeding-

lamelle.

Fig. 1 a. Caudal fin of the same.

Fig. 2. A male, x 2 nat. size. a’, first antenna; ap, appendix lamellaris ; a*, second antenna; pm,

enlarged palpus mandibule; mp, maxilliped; gn’, first gnathopod; prp?, second pereiopod ;

prp’, fifth pereiopod.

Fig. 2 a. Caudal fin of the same.

Fig. 3. First antenna of the female.

Fig. 4. Second antenna of the same.

Fig. 5. Scale of the second antenna.

Fig. 6. Upper lip.

Fig. 7. Mandibula of the female: pm, palpus mandibule.

Fig. 8. First maxilla of the same.

Fig. 9. Second maxilla of the same: p, palpus; fl, flagellum. ;

Fig. 10. Maxilliped of the same: a/, appendix lamellaris ; p, palpus; fl, flagellum.

Fig. 11. First gnathopod of the same.

Fig. 12. Second gnathopod of the same.

Fig. 13. Third pereiopod of the same.

Fig. 14. Pleopoda 1-5 of the same.

Vig. 15. Maxilliped of the male: /a, lamellar appendage.

Fig. 16. First gnathopod of the same: Ja, lamellar appendage.

Fig. 17. First, fourth, and fifth pleopoda of the same.

Prave VIL.

All the figures refer to Chalaraspis ungwiculata.

Fig. 1. A male, x2 nat. size. a’, first antenna; a, second antenna; gu, gnathopoda; p, pereiopoda.

Fig. la. Head of the male, seen from above: sc, small tubercle beneath the rostrum and between the

eyes.

Fig. 16. Caudal fin of the same.

Fig. 2. First antenna of a female.

Fig. 3. Second autenna, with lamellar appendage of the same.

Fig. 4. Mandibula of the same.

Fig. 5. Under lip of the same.

Fig. 6. First maxilla of the same.

Fig. 7. Second maxilla of the same: p, palpus ; fl, flacellum.

Fig. 8. Maxilliped ofthe same: p, palpus; ji, flagellum.

Fig. 9. First gnathopod

5 ; : |of the same.

Fig. 10. First pereiopod

Fig. 11. Upper lip of the same.

SECOND SERIES.—ZOOLOGY, VOL. I.

heed

Soe Ss

g@ do Go do OO

fras|

Fig

2 Oo, Hirst

DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

- 12. Fourth pereiopod i

. 13. Fifth pereiopod fof the same.

. 14, Anterior pleopod

Puate IX.

1. Gnathophausia gracilis, male, x 4 nat. size.

2. Second antenna of Gn. zoéa: al, appendix lamellaris ; fw, funiculus; jl, flagellum.

3. Mandibula of the same ; p, palpus.

. 4. Labium of the same.

. 5, Hirst maxilla of the same, higher power: p, palpus maxille.

ig. 6. Second maxilla of the same: fl, flagellum ; p, palpus; 0, accessory eye.

- Maxilhped of the same: jl, flagellum.

g. 15. One of the anterior pleopoda of the same.

g. 16. Maxilliped of Gn. gigas: fl, flagellum ; p, palpus.

g. 17. Outlines of the eye of the same.

PLATE X.

. 1. Astacus zaleucus, male, nat. size

g. la. One of its right pleopoda.

Nig. 2. Gnathophausia gigas, male, nat. size.

. 2a, The same, side view (the carapace is lifted somewhat to show that it is not connected with the

pereion), nat. size.

Hig. 3. Oral apparatus of the same: /, labrum; md, mandibula; mp, palpus mandibule ; p, palpus of the

second maxilla; jl, its flagellum; ac, accessory eyes on the second maxilla; basal joint of

the maxilliped with its palpus (the other joints of m have not been drawn, in order not to

confuse the figure) ; 2 and y, plates formed by the flattened inferior borders of the carapace.

4. Gnathophausia zoéa, male, x 4 nat. size.

Prats XI.

‘ig. 1. Zoéa of Cardisoma, sp.? from San Jago (Cape-Verd Islands). High power. a’, first, a?, second

antenna; m, parts of the mouth; gp, the gnathopoda.

». 2. Caudal fin of the same, seen from above. High power.

e. 3. A string of eggs and egg-capsules, partly containing zoéas, partly left by them. ‘Taken from

the abdominal feet of the mother. Low power.

lig. . Cystisoma neptunus (Thaumops pellucida), male. Nat. size.

ig. 5. Genital organs of a smaller male: A, segments of the percion; B, first segment of the pleon ;

t, testes; vd, vasa deferentia; ag, aperture genitales.

. 6, Mandibula of the same. High power.

7. Maxilla of the same.

8. Maxillipedes (m) and second maxilla, with (/) labium (?) of the same.

Pruate XII.

Vigs, 1-9 refer to Apseudes ceca.

Vig.

1. A female, x 4 nat. size: a, antenna; gn, gnathopoda.

Fig

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Fig.

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION.

. 2, First antenna.

. 8. Second antenna: s, scale; f, flagellum.

. 4. First gnathopod.

. 5. Second gnathopod.

s. 6, 7. First and second pereiopods.

. 8. Last joints of the fourth pereiopod.

. 9. Maxilliped.

. 10. Willemoesia crucifera, x 4 nat. size.

Pruate XIII.

. Willemoesia leptodactyla, male, nat. size.

. First styliform pleopod.

. Upper border of the opening of the mouth.

. Mandibula, with the palpus (ym) and the left labium (Jad).

- 6a. First maxilla: fl, rudiment of a flagellum.

oF WOH

g. 66. Second maxilla: fi, flagellum.

g. 7. (First) maxilliped: p, palpus; fl, upper half of the flagellum ; fi”, under half.

g. 8. First gnathopod (second maxilliped).

9. Second gnathopod (third maxilliped).

10. Second left pleopod of W. crucifera, with two styliform appendages.

11. Third right pleopod of the same, with one styliform appendage.

- Second pleopod of the right side: a@, styliform appendage to the inner ramus.

12. Second right pleopod of an undescribed Palinurid from deep water near-the Bermudas.

Wherever the words “high power” are not added, a low power of a Hartnack’s microscope, or, in

case it was not possible to dissect the animal, a loupe has been used.

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[ 61 ]

III. On the Structure and Systematic Position of Stephanoscyphus mirabilis, the Type

of a new Order of Hydrozoa. By Prof. Attman, W.D., DL.D., FR.S., &c., Pres.

Linn. Soe.

(Plate XIV.)

Read November 19th, 1874.

WHILST engaged last May in zoological researches on the coast of Antibes, I was

struck by some small patches of one of the horny sponges which had attached them-

selves to stones at a slight distance below the surface of the sea, and which had arrested

my attention by what appeared to be unusually distinct and well-defined oscula. After

transferring some specimens to a phial of sea-water, I was astonished to find that from

each of the apparent oscula a beautiful crown of delicate tentacles soon began to develop

itself. The tentacles were quite like those of certain hydroids; and the appearance was

such as might easily have led to the belief that the anticipated form had at last been

found which would decide the zoological position of the sponges, and remove all doubt of

the hydrozoal structure recently assigned to them*. A closer examination, however,

showed that the orifices on the surface were not really oscula, and that the tentacles

proceeded, not from the sponge itself, but from a totally different organism, which lay

imbedded in its substance, and which sent tubular prolongations to the surface, where

they opened in the manner of genuine oscula.

It was further evident that the organism thus associated with the sponge was in the

highest degree interesting ; and a careful study of its structure made it plain that, while

in all essential points it possessed a true hydrozoal organization, it differed from all the

recognized orders of Hydrozoa so widely as to necessitate our assuming it as the type of

an entirely new one.

To this remarkable animal I have given the generic name of Stephanoscyphus. When

freed from the surrounding sponge, it is seen to be composed of an assemblage of zooids,

which are included in chitinous tubes, and associated into a composite colony by a common

basal plexus of tubes, the aspect of the whole being very much that of an ordinary hydroid

trophosome (figs. 2,3). In all the specimens examined the common plexus lay at the

base of the sponge, and the tubes given off from this penetrated the sponge-tissue, and,

rapidly increasing in width, ultimately reached the surface, on which they opened. The

tubes really extended beyond the general surface; but each carried up the sponge-sub-

stance with it, so that the orifices appeared elevated on the summit of conical papillee—

a condition very common in the oscula of sponges (fig. 1).

The distal extremity of the animal can extend itself beyond the safes of the chitinous

tubes, and can again withdraw itself far into the interior. When extended (fig. 3, a), it

displays a coronal of tentacles, which surround a wide terminal orifice. The tentacles

spring from the very margin of this orifice ; for there is nothing like a hypostome or pro-

* Haeckel, ‘ Kalkschwimme.’

62, PROF. ALLMAN ON STEPHANOSCYPHUS MIRABILIS.

boscis, such as in the true hydroids we find intervening between the tentacles and the

mouth of the hydranth ; and it was the great width of the terminal orifice, with its mar-

ginal circlet of tentacles, so different from the narrow mouth of a hydroid elevated on

the summit of a hypostome, that first led me to doubt the justice of referring Stephano-

scyphus to the true hydroids. The appearance of the tentacular crown, indeed, was not

altogether unlike that of certain zoanthal Actinozoa; but all grounds for assigning to

Stephanoscyphus such actinozoal relations were removed by dissection.

The tentacles form a single circlet, which is composed of two closely approximated

alternating series of about eighteen tentacles in each, the outer series being slightly

shorter than the inner, and usually depressed in extension, while the inner series is held

more elevated. Not unfrequently the whole circlet may be seen bent back upon the

outer side of the chitinous tube (fig. 3, ¢). The tentacles are very contractile: when

fully extended they are long and filiform; but in certain states of contraction they

assume a very distinctly clavate form (fig. 8, 6). Their structure is entirely that of the

typical hydroid tentacle: they present the septate condition so characteristic of such

tentacles ; and they carry along their length, just as in most hydroids, somewhat spirally

arranged slightly elevated clusters of thread-cells (fig. 4). In complete retraction the

tentacles are not only greatly shortened, but are thrown inwards and entirely with-

drawn within the terminal orifice of the animal, which is then nearly or altogether

closed over them (fig. 3, d).

In the completely extended state of the animal, four longitudinal canals may be traced

from the base of the tentacular crown for some distance backwards in the walls of the

body ; but the general want of transparency, and the way in which the greater part of

the animal is concealed by the surrounding sponge, renders it impossible to obtain,

without dissection, a satisfactory view of the deeper-lying parts. Sections, however, were

easily made of specimens which had been preserved in spirits; and some further im-

portant points of structure were thus determined.

When a longitudinal section (fig. 5) of such specimens is made through the axis of

the body, it is seen that the terminal orifice opens into a wide thin-walled cavity, which

becomes continuous posteriorly with the canal which occupies the axis of the stem.

The contracted tentacles are now also brought into view with their extremities thrown

back, and with the whole tentacular crown entirely withdrawn into the cavity.

By the same section another very important piece of structure becomes apparent; for

‘it may be now seen that the knife has divided a circular canal which runs in the walls of

the body round the terminal orifice. This canal is very obvious; it is remarkable for its

width, and easily admits the passage into it of a needle. It supports the tentacular

circlet; and when a transverse section of the body is made along its course, it is seen to

be quite continuous and destitute of septa. It possesses a very distinct endodermal

lining.

With the circular canal the distal extremities of the longitndinal canals are in con-

nexion, and they probably open into it. ‘These canals can now be traced far back in the

walis of the body. They lie quite close to the inner surface of the walls ; and their sides

project into the cavity as four slightly elevated ridges. In the deeper parts of this

PROF. ALLMAN ON STEPHANOSCYPHUS MIRABILIS. 63

cavity they present a somewhat lobed or pouched appearance; but beyond this they

become lost in the thickened walls of the body, so that their actual termination could

not be made out. Their walls are thick; and their lumen, which is very narrow, is

doubtless lined by an endoderm; but this could not be demonstrated with the same

distinctness as in the circular canal.

The walls of the cavity which occupies the axis of the narrow portion, or stem, are

much thicker than those of the wide distal chamber, and posteriorly encroach so much

upon the axile cavity as to reduce this to a very narrow canal.

Thread-cells exist throughout in the ectodermal layer; but these are everywhere

exceedingly minute. Those examined from the body-walls in spirit specimens were

nearly spherical, and under a 3‘; immersion-lens were seen to include a spirally coiled

filament ; they were about the gso9 of an inch in diameter.

The more posterior or proximal parts of the animal are fixed in the external chitinous

perisarcal tube, and possess no obvious contractility; but the more anterior or distal

portion is very contractile, and is quite free from the tube, so that it can move up and

down in it during extension and retraction, quite like a campanularian hydranth in its

hydrotheca.

The surrounding chitinous tube, indeed, increases so rapidly in width towards its

distal end as to resemble here the hydrotheca of a campanularian (fig. 3). It is marked

by annular ruge for some distance from its free extremity, and usually by fine longi-

tudinal striz ; but towards its attached end the marking disappears.

At rather irregular distances along its length its inner surface sends off remarkable

processes, which project far into the interior (fig. 5, a, a, and fig. 3, e). At four equi-

distant points, situated on the same level, a thin chitinous lamina separates from the

inner surface of the tube, and forms a hollow pyramidal process. The four processes

thus formed radiate towards the axis, pushing before them the soft parts, and forming a

stelliform constriction of the body (fig. 6). Between each of these four radiating pro-

cesses a much smaller one is sometimes present, thus raising to eight the number of pro-

cesses in a set, and indicating an increase in number by consecutive intercalations of

younger ones between those previously formed. These imperfect septa, notwithstanding

their striking appearance, do not seem to possess any important significance in the orga-

nization. They will, however, suggest a comparison with the tabule of the tabulate

corals, and may be regarded as offering some support to Agassiz’s view of the hydrozoal

nature of these animals. j

In none of the specimens examined could generative elements be found. It is almost

certain, however, that the absence of obvious generative elements is the result of the

immature condition of the reproductive organs. We should expect to find them in the

walls of the longitudinal canals; and I believe that the pouched condition observed in

these canals in the deeper part of their course is only an early state of the reproductive

system.

64 PROF. ALLMAN ON STEPHANOSCYPHUS MIRABILIS.

General Remarks—Homology.

From the account now given of Stephanoscyphus mirabilis, it is evident that we have in

this remarkable animal a tue hydrozoal structure ; but while its apparent form and habit

are those of a hydroid trophosome, its organization is, in its essential features, that of a

hydroid planoblast or medusa. If we imagine a campanularian hydroid in which the

hydranths have been replaced by peculiarly modified medusze, capable, as in the hydranth,

of exsertion beyond the hydrothec, and of retraction within them, we shall form very

nearly a true conception of Stephanoscyphus. 'The wide thin-walled chamber into which

the terminal orifice opens would thus correspond to the umbrella of a medusa; and the

terminal orifice itself, though neither lithocysts nor ocelli appear to be developed

on its margin, would be homologous, not with the mouth of a hydranth, but with the

codonostome or umbrella-aperture of a medusa; while the tentacular crown would admit

of a direct comparison, not with the tentacular crown of a hydranth, but with the mar-

ginal tentacles of a medusa. The true mouth would thus have to be sought for at the

point where the wide distal cavity passes into the narrow proximal cavity which occupies

the axis of the stem. Then there are the four longitudinal canals running symmetrically

in the umbrella-walls, and the circular canal surrounding the codonostome, all so many

obvious features of medusal organization. It is true that no manubrium appears to be

developed, and the bottom of the umbrella-cavity seems to pass uninterruptedly into the

true gastric cavity (which is continued through the axis of the stem), neither is there a

velum developed at the codonostome, nor do the walls possess the gelatinous consistence

and transparency of the typical medusze; but these special modifications in no degree

invalidate the points of homology here insisted on. In the absence of a manubrium we

are reminded of the swimming-bells or nectocalices of the Siphonophora, which, though

closely resembling in all other respects a true hydroid medusa, and undoubtedly homo-

logous with it, are destitute of a manubrium.

If the views now advocated be correct, the wide cavity into which the terminal orifice

opens ought to have no endodermal lining: and this I believe to be really the fact; for a

very careful examination has failed to demonstrate more than a single layer in its walls.

On the contrary, the endoderm of the circular canal is very obvious; and though a

proper lining membrane cannot be distinctly recognized in the longitudinal canals, this

ought not, in an observation rendered difficult by the narrowness of these canals and by

the want of transparency of the tissues, to be urged as an argument against its presence.

On the other hand, the axile cavity of the stem appears to possess a well-developed endo-

derm, and the proper gastric cavity must be regarded as confined to this part.

The peculiar position assumed by the tentacles in extreme retraction is very different

from the condition of these parts in the retracted state of the hydranth in a hydroid

trophosome, and quite corresponds with that which they present in the greater number

of young medusiform planoblasts, in which before the planoblast becomes free the

marginal tentacles are thrown back, and lie entirely within the cavity of the umbrella.

1 have elsewhere * endeavoured to show that the radiating canals of a medusa are —

strictly homologous with the tentacles of a hydranth, and that, if the latter were con-

* Monograph of Gymnoblastic Hydroids.

PROF. ALLMAN ON STEPHANOSCYPHUS MIRABILIS, 65

nected with one another by an extension of the ectoderm, we should have the tentacular

crown of the hydranth converted in nearly all essential points into the umbrella with the

radiating canals of a medusa.

Now in certain species of Hydroida (Laomedea flexuosa for example) the tentacles are

actually connected at their bases by an ectodermal web. Here, then, there is little

needed to render the parallelism complete but the presence of a circular canal. Ste-

phanoscyphus, however, supplies the missing element, and, by the addition of a well-

developed circular canal, beautifully confirms the view here taken of the homological

relations between hydranth and medusa.

Whether the relation between Stephanoscyphus mirabilis and the sponge with which

it is found associated is one of parasitism, or whether the association is merely acci-

dental, it is at present impossible to say. At all events, in no instance did I meet with

the Stephanoscyphus free from the accompanying sponge.

From the account now given it is obvious that Stephanoscyphus must form the type

of a distinct order of Hydrozoa. With the order Hydroida it possesses very decided

affinities, but it is nevertheless removed from the Hydroida by a distance at least as great

as that which separates these from the Siphonophora. For the new order I propose the

name of THECOMEDUS£; and if we bring together the characters given above, so as to

enable us to assign to Stephanoscyphus its true systematic position, we shall obtain

the following diagnostic category :—

Subkingdom CHLENTERATA.

Class HyDROZzOA.

Order THECOMEDUS&.

THECOMEDUS2.

Animal composite, zooids medusiform, with circular and radiating canals, and included

in a chitinous rooted perisare, which forms thecze, within which they are retractile.

Genus Stephanoscyphus.

Name from orépavoc, a crown, and sxioc, a cup, in allusion to the cup-like body with

its crown of marginal tentacles.

Gen. char.—Terminal orifice surrounded by a single circlet of simple filiform contractile

tentacula; four longitudinal canals; velum and manubrium undeveloped.

Generative elements formed in the walls of the longitudinal canals ?

Stephanoscyphus mirabilis.

Perisarcal tubes springing from a basal tubular plexus, and rising to a height of about

one third of an inch, simple, rapidly enlarging upwards, annulated for some distance

from their distal extremities ; tentacles about 36 in number.

Hatitat.—Immersed in the substance of certain corneous sponges.

Locality —Antibes, Mediterranean coast of France.—G. J. A.

SECOND SERIES.—ZOOLOGY, VOL. I. K

66 PROF. ALLMAN ON STEPHANOSCYPHUS MIRABILIS.

DESCRIPTION OF PLATE.

PuaTe XIV.

Fig. 1. Stephanoscyphus mirabilis, with its investing sponge : nat. size.

2. Perisarcal tubes of the same freed from the sponge: nat. size.

3. A group of zooids magnified. a, the animal fully exserted and the tentacles extended; 5, the

animal partially retracted ; c, the animal with the tentacles thrown back over the sides of the

tube; d, the animal completely retracted; e, an empty perisarcal tube: the internal chitinous

processes are seen through its walls.

4. Distal portion of a tentacle seen im optical longitudinal section : more highly magnified.

5. Longitudinal section through a perisarcal tube and the retracted animal, from a specimen preserved

in spirit: a, a,imternal chitious processes.

6. Transverse section of same through one of the verticils of chitinous processes.

7. Thread-cell from body.

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Trans. Linn. Soc. Ser 2. Zoou No.l Tas.

W. West & C° amp.

mirabilis.

Steph anos cyphus

GJ Allman del.

WH Wesley lth.

THE .

TRANSACTIONS

THE LINNEAN SOCIETY

LONDON.

SECOND SERIES.—ZOOLOGY.

VOLUME I.

PART THE SECOND.

PAO) NEDEOAN:

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET:

SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON-HOUSE;

. AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER-ROW.

M.DCCC.LXXV.

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[ 67 ]

IV. On the external Anatomy of Tanais vittatus, occurring with Limnoria and Chelura

terebrans in excavated Pier-wood. By JoHN Dents Macponatp, I.D., F.R.S.,

Staff-Surgeon R.N., Assistant Professor of Naval Hygiene, Netley Medical School.

(Communicated by W.'T. Tuisetton Dyer, WA, F.L.S.)

(Plate XV.)

Read Feb. 18th, 1875.

WHILE searching for Limnoria terebrans in excavated wood from N etley Hospital

Pier my attention was diverted to another little Crustacean, rather smaller in size, but

wearing the outward appearance of a chelate Decapod so completely that it was only by

counting its legs and more minute inspection of the leading features of its structure I

was enabled to determine it to be a veritable Isopod, though of an aberrant type.

It proved to be conformable with the characters of Tanais Cavolini (Edw.) = Crossurus

vittatus (Rathke), which was originally found in the Gulf of Naples, the habitat assigned

to it by Milne-Edwards; but it has since been shown to have a wider range. Its occur-

rence under the above-mentioned circumstances is worth recording ; for I have also found

it in perforated and decayed wood from the pier at Alum Bay, Isle of Wight; and I have

no doubt that it would be captured in other localities if it were sought for in the burrows

of Limnoria.

Quite exceptionally, in the Tanaidee the head and first thoracic segment coalesce above

to form a perfect little carapace or cephalothorax, which, in the little creature under

consideration, is somewhat triangular in form, with an obtuse but distinct rostrum in

front, from beneath which the two pairs of antenne project forwards; and immediately

to the right and left of these are the orbital notches for the eyes.

These latter organs are very prominent, in articulation with the carapace, and all but

peduneculated. They are conical in form, with the apex directed forwards, and rather

more convex or rounded on the outer than on the inner side. Round lenticular cornes,

not unlike those of Limnoria, but much smaller, are recognized by their brilliancy, and

the relief of a central mass of black pigment radiating amongst the little nervous cups in

connexion with them.

Having thus commenced with the eyes, I shall describe the members of the succeeding

segments seriatim, which will render their homological relations more intelligible.

The two pairs of antenne (figs. 1, 2, & 3) are nearly of equal length, rather shorter

than the cephalothorax, and disposed in two lines, one above another. The superior

antennee (a, figs. 2 & 3), which are much stouter and somewhat longer than the inferior,

are each composed of three joints, successively diminishing in their proportions, and

setaceous at their distalends. Several bristles of a ligulate form (fig. 2, e), with transverse

strize near their base, occur amongst the simple ones in the tuft at the tip of the terminal

segment. A labyrinth-like excavation at the upper and inner side of the base of the first

SECOND SERIES.—ZOOLOGY, VOL. I. L

68 DR. J. D. MACDONALD ON TANAIS VITTATUS.

articulation (fig. 2,¢) represents the so-called olfactory organ of the Decapod. The second

or inferior pair of antenneze are narrower than the superior, and composed of five joints,

of which the first, third, and fifth are shorter than the second and fourth; and the ulti-

mate joint is tipped with a brush of long and simple hairs, springing from what would

appear to be an aborted multiarticulate filament.

The oral organs consist of a spatulate upper lip, fringed with short hairs (fig. 6, a), a

pair of two-branched mandibles without palps (0), and articulated laterally with the buccal

frame, two pairs of maxille (¢ & d), and a pair of five-jointed jaw-feet (/f), inclosing, at

their base, two membraneous plates, forming a lower lip (¢).

The internal maxille (¢) are in the form of rounded membraneous lamelle, fringed

with fine hairs, and presenting a Single-jointed palpiform process near their summit

externally.

The external maxillee(d), on the contrary, are strong and sabre-shaped, with a spiny

and setaceous extremity. ‘Two oval leaflets (d’), connected with these internally, form a

tongue-like organ in the mouth; and at their base externally, but concealed by the cara-

pace, is a conical appendage (d”) or flagellum, with long diverging hairs at the tip. In

the possession of this appendage, as Fritz Miller has pointed out, the Tanaidee make a

still further approach to the Decapoda. Thus, in his ‘ Facts and Arguments for Darwin’*,

he remarks, “ Whilst in all other Oniscoida the abdominal feet serve for respiration,

these in our cheliferous Isopod (Zanais dubiws(?) Kr.), are solely motory organs, into

which no blood-corpuscle ever enters ; and the chief seat of respiration is, as in the Zoez,

in the lateral parts of the carapace, which are abundantly traversed by currents of blood,

and beneath which a constant stream of water passes, maintained, as in the Zoez and the

adult Decapod, by an appendage of the second pair of maxille, which is wanting in all

other Edriophthalmata.” In passing, I may mention that the figure of Tanais dubws, so

called, loc. cit., is rather more conformable to the genus Leptochelia. In this connexion

Bate and Westwood’s beautiful work on the British sessile-eyed Crustacea may be con-

sulted with advantage. But to return to the subject, the joints of the jaw-fect (f) are

furnished with rigid comb-like bristles, arising from the inner border of the third and

fourth, and the free extremity of the fifth. The lower lip (e) is also fringed with fine

short hairs.

Following the jaw-feet just noticed are the seven pairs of legs common to all the Kdri

ophthalmata, the first two and last five respectively corresponding with the two external

‘pairs of jaw-feet, and the five pairs of legs appertaining to the thorax of the Decapod.

The two first limbs (fig. 7) are strongly chelate, and many times thicker than the

ambulatory legs succeeding them. The fixed claw presents a trenchant upper border,

more prominent in the middle, with which, at least in the female, the movable claw cor-

responds more or less closely. In the male, however, (fig. 15) both claws are more

arcuate. As in the beak of the Crossbill, the opposable tips of the claws are curved

with sufficient obliquity to cross over one another, the index taking the outer side. They

are of a rich reddish amber tint, similar to that of the dental processes of the mandibles.

* Translated by W. 8. Dallas, F.L.S8.

DR. J. D. MACDONALD ON TANAIS VITTATUS. 69

The first or basal portions of these limbs form two oval projections beneath the thorax ;

the second is a small triangular piece, with which the third is so connected as to present

a rounded projection posteriorly, while it extends forwards to articulate with the manus.

In the female (figs. 1 & 7) eight or nine stout bristles spring in a row at the outer side

of the base of the cutting-border of the manus, while in the male they are interrupted in

the middle. Other bristles occur near the articulation, 7. e. one on the inner and several

on the outer side.

As already mentioned, the first thoracic ring having coalesced with the head, the six

following pairs of legs arise from the six apparent thoracic segments. ‘The first visible

segment, distinct from the carapace, must, therefore, carry the second pair of legs. These

have long curved thigh-pieces, with which the succeeding joints of the limbs are so arti-

culated as to permit them to play their legitimate part as external jaw-feet, curving

inwards and forwards round the basal prominence of the first pair. Their claws (fig. 8)

are long, simpie, styliform, and but slightly curved; they are themselves rather longer

than the other legs.

The claws of the third and fourth pairs (figs. 9 & 10) are simply uncinate, while those

of the last three pairs (figs. 11, 12, & 13), which are curved forwards, are also combed

on either side, though this is not quite so apparent on the inner side in the last pair.

Instead of it, however, they present a close row of stiff serrated hairs at the extremity of

the penultimate joint, which is moreover roughened with rasping tubercles (13 a), quite

like those which beset the whole surface of the body in Limnoria. In all the legs, with

the exception of first and second pairs, the third segment presents a row of stout spines on

either side of its articulation with the fourth.

The abdomen consists of five segments, the first three of which bear the respiratory

lamelle and swimmerets together. The fourth is much smaller than those preceding it,

and without appendage. The fifth is rather more than twice the length of the fourth,

broadly shield-shaped, with four rigid hairs projecting backwards from the middle of

its posterior border. Besides the swimmerets there is a fourth pair of appendages

(figs. 1, 4, 5, & 14), which are rather conspicuous on account of their projection beyond

the terminal segment. They have a simple folded lamella on the inner side, and much

resemble a pair of three-jointed antennz, with stout diverging hairs at the distal end of

each joint.

The inner ovate and acute leaflets of the abdominal appendages (fig. 4, 6) lie flat, and

are apparently motionless ; while the outer ones, which are rather semicircular (a), cross

over them superficially towards opposite sides. Both borders of the deep leaflets and the

outer border of the superficial ones, as well as that of the broad propodia, are beautifully

fringed with long flat acute and feathery setze (d). Similar feathery hairs forma trans-

verse linear band or fascicle upon the dorsum of the first and second segments.

The generative pores lie side by side near the middle of the sternal plate, between the

two last thoracic limbs.

The female (fig. 1) carries her eggs in a membraneous expansion or saccule under the

thorax ; and, from having observed the chele and other anatomical characters of the

adult in the young soon after leaving the egg, I think it may be concluded that there is

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V. On Valencinia Armandi*, a new Nemertean.

By W.C. M‘Intosu, U.D., F.R.S.H., BLS.

(Plate XVI.)

Read June 17th, 1875.

WHILE searching for a rare Annelid (Mea mirabilis) that oceurs near low water in

the extensive waste of sand at Southport, a slender thread, which glistened in the October

sun like gossamer, attracted notice in the upturned sand. This was the proboscis of the

ruptured Nemertean, which exists in considerable abundance in the sand amongst the

tubes of Lanice conchilega, associated with Amphiporus lactifloreus, A. hastatus, Micrura

Jusca, and other forms usually characteristic of the laminarian and coralline regions of

the north. It appears to live in a burrow in the sand, and readily takes to a case or tube

in confinement. A

The body is 7 or 8 inches in length, and not much thicker than a stout thread. It is

rounded in front and in the middle, but becomes extremely flattened towards the tail,

which terminates in a somewhat lanceolate point. The anterior part of the body is

whitish, the posterior pale buff, while the tail is translucent. The head is rounded in

front, flattened, and slightly enlarged behind the tip. There is no trace of eyes, fissure,

or furrow. On the whole the Nemertean very much resembled Carinella linearis, from

Lochmaddy; and it was placed in spirit under this impression. More careful scrutiny,

however, reveals that it properly comes near the genus Valencinia, De Quatrefages,

char. emend., the only British example of which (Valencinia lineiformis) was dredged in

numbers amongst the shell-gravel of Bressay Sound, in Shetland. The Zetlandic species

is larger, and has an evident row of eyes on each side of the head, besides other points of

distinction, which will by-and-by be alluded to.

In the structure of the cutaneous system Valencinia Armandi agrees with its allies.

The entire surface is clothed with cilia; and the cutis has the usual cellulo-granular

appearance (fig. 9, @), with numerous vertical streaks. The latter are not conspicuous in

the Zetlandic species (fig. 10, a), which has a minutely cellulo-granular cutis, very easily

removed in the preparations. In the new form the cutis is bounded externally by a some-

what definite edge, which probably represents a rudimentary cuticle. In the next layer

(4, figs. 9 & 10) a considerable difference exists between the two species. In Valencinia

Armandi the basis-layer is very thin, while in /. lineiformis it is dense. In the snout

of the former the cutis is proportionally thicker, but the basis-layer is not increased—

another contrast; for the latter is especially developed on the snout in the Zetlandic

species. In VY. Armandi there is an increase of this layer opposite the nerve-cords. In

connexion with the cutaneous system, it may be mentioned that, in a large Canadian

* Named after M. Armand de Quatrefages, who long ago did such good work in the group.

74. W. C. MIINTOSH ON VALENCINIA ARMANDI.

Carmella, fibres from the elaborate muscular system of the snout pass from the trans-

verse bands into the cutaneous layer; so that, with the arrangement of the basis-layer, the

tissue has a characteristic arborescent appearance.

Underneath the basis-layer is a well-marked belt of circular muscular fibres. Towards

the tip of the snout these fibres are not developed as an external coat, probably because

the complex system of internal muscles, hereafter to be described, renders their presence

less necessary. Immediately behind the anterior border of the snout (fig. 1) a differen-

tiation of the cutaneous stroma occurs. At first, in front of the proboscidian aperture,

a single distinct transverse band appears, with numerous vascular meshes beneath it,

such probably representing the area of communication between the lateral trunks. Two

transverse muscular bands, however, as soon as the proboscidian canal is reached (fig. 2,

e & e’), pass from side to side, leaving a considerable space between them; and this is

occupied by vertical bundles, which enclose various meshes, the chief being in the centre

for the proboscidian canal, while several smaller vascular spaces exist on each side.

Very shortly, however, as we proceed backward, greater differentiation occurs; for the

transverse bands increase in strength, and numerous longitudinal muscular fibres appear

above and beneath their outer borders. A transverse section, therefore, in this region

(fig. 2) shows beneath the cutis (a) the pale and comparatively thin basis-layer (0). The

two great transverse muscular bands (e & e’) pass right across, forming a kind of ellipse,

each spreading out so as almost to meet its fellow at the circumference on each side, and

perforated here and there by the vertical bands. Numerous vertical bands extend

from the superior and inferior borders of the ellipse to the circumference, often in an

arborescent manner, and become lost at the basis-layer. The spaces thus left between

the outer edge of the ellipse and the latter are filled with powerful bundles of longitudinal

muscular fibres (7d). The area of the ellipse, again, presents the canal for the proboscis

(ao) in the centre, with its pale glandular lining internally, and externally its strong coat

of longitudinal muscular fibres. This canal is supported on each side by vertical mus-

cular bands, whose fibres, spreading out at each transverse muscle, pass to the circum-

ference above and beneath. Two bundles of fibres are especially conspicuous on each

side. Then come a great vascular space and a small one between it and the circumfer-

ence. Between these spaces and at the outer edge similar vertical bands of fibres occur,

with numerous longitudinal fasciculi. Along the inferior border of the superior trans-

verse muscle and the superior border of the inferior are many longitudinal bundles, form-

‘ing a somewhat regularly diminishing series towards the central line. Proceeding back-

ward, it is found that the chief changes in the great transverse bands are their diminu-

tion in length, increase in bulk, and more evident divergence at the extremities, so that

their lateral fibres almost decussate. The superior and inferior vertical bands are also

more boldly developed. There are still two vascular tubes on each side of the probosci-

dian canal, a larger and a smaller. By-and-by there is a tendency to further differentia-

tion in the proboscidian and in the vascular area, as shown by the greater development

of a lateral vertical band, composed internally of longitudinal, and externally of vertical

fibres. This becomes marked as soon as the anterior border of the ganglia and the turn-

ing over of the proboscidian sheath are reached, while only a single vascular channel is"

W. C. M‘SINTOSH ON VALENCINIA ARMANDI. 75

present on each side. A thin ring of circular muscular fibres can now be traced all

round the united area of the proboscis and the vascular canals, the great transverse

bands being inflected at each side where the circular fibres pass. Between the united

area and the circumference of the body we have now the ever-increasing ganglia, which

rest inferiorly on the outer division of the inferior transverse band, and are bounded

superiorly by the outer fibres of the upper transverse band. Externally they apparently

abut on the basis-layer. About this region a curious maculation of the part within the

basis-layer is observed, especially superiorly—a feature due to the presence of two or three

narrow longitudinal separate layers of muscular fibres, which form a complete investment.

Just in front of the mouth they form a rather thick compound layer, best marked supe-

riorly and inferiorly; but the advent of the oral aperture separates the latter into two

great masses. Some circular fibres also appear to be present at the basis-layer, but they

are much less conspicuous than the longitudinal. A section made through the ganglia,

almost in the plane of both commissures (fig. 3), shows three rows of these interrupted

muscles (d’) superiorly and inferiorly; but the lateral regions are less distinct. The

ganglia (/) surround the united area of the proboscidian and vascular systems; and, besides,

their other investments have increased in complexity. The external circular and internal

longitudinal fibres of the area still form a complete ring; but there is also an inferior

appendix in the shape of a canal (s) on each side, separated from the great vascular tunnel

by a powerful transverse band (m). From the centre of the latter superiorly springs a

process ( p), which separates the great vascular tubes, and terminates above in the circular

coat of the proboscidian chamber(ao). Then, shortly afterwards, a series of longitudinal

fibres appear in the isthmus between the vascular channels, and run round the inner

circular coat, which further presents on its inner surface a complete lining of longitu-

dinal fibres. The two vascular canals appended to the inferior border of the transverse

band are soon separated by a broad transverse belt, and by-and-by are obliterated by the

mouth. The broad transverse belt just mentioned spreads out on each side underneath

the diminishing ganglia; and some of the inferior fibres join the external longitudinal

series of separate muscles. The only representative of the great longitudinal muscular

layer afterwards so highly developed is a narrow arch (fig. 4, d), extending from the upper

end of each great vascular channel, and meeting at the raphe in the dorsal median line.

This, however, soon increases in thickness, and presents two channels (fig. 4, vs) between

its inner surface and the proboscidian sheath; while behind the mouth it quite encircles

the body. The circular muscular coat (c) likewise completes the circuit of the body

immediately after the mouth. The separated layer of longitudinal fibres (d’) outside

the circular layer soon breaks up into small fasciculi, which, in transverse section, form

a dotted belt outside the nerve-cords.

There is thus a very characteristic difference between this form and Valencinia linei-

formis or Carinella. In V. linetformis the snout, though very much longer in front of

the mouth, is less complex, there is only one vascular channel (of large size) on each

side of the proboscidian canal, and the ganglia have no longitudinal fibres between them

and the basis-layer. From without inwards there are the cutis, basis-layer, and circular

_ muscular fibres, while the rest of the area, with the exception of the three channels just

SECOND SERIES.—ZOOLOGY, VOL. I. M

76 W.C. M‘INTOSH ON VALENCINIA ARMANDI.

mentioned, is filled up by the great longitudinal muscular layer. The nerves lie in the

substance of the latter at once, whereas in the new form the nerves are placed quite out-

side the great longitudinal coat, and the whole region is much more complex. Ina

large Canadian example of Carinella*, again, the nerves lie outside all the muscular

layers, but the complexity of the muscular system of the snout is equally wonderful.

The tip is formed of an interwoven series of longitudinal, vertical, and transverse

fibres—which lie within the circular coat, and interdigitate with certain decussating

fibres of the latter at its inner margin. A very symmetrical arrangement of this

kind occurs in the middle line at the tip in front of the proboscidian aperture. At first

the snout is quite filled up with these elaborate muscles, with the exception of the pro-

boscidian canal; then a series of vascular spaces form above the latter, and range

themselves on each side, while a great longitudinal muscular mass occurs inferiorly.

Strong vertical bands on each side of the proboscidian canal, and transverse muscles

below it, map out the region very effectively. Thus at the ganglia a transverse section

shows within the cutaneous layer the circular muscular coat, with it decussating fibres

dorsally and ventrally. The great longitudinal coat forms a complete circuit, pierced

here and there by the vertical and other fibres. There are apparently two great vascular

channels above the great transverse muscular band on each side of the proboscidian canal ;

but the septa may only indicate temporary division of a single channel. Another large

canal lies in the centre below the transverse band; but the occurrence of the mouth soon

obliterates the latter. It is probable that all the canals communicate (as in the Anopla),

and the circulating medium is represented in the preparations by a minutely granular

substance.

The occurrence of the special layers of separated longitudinal fibres in the new form is

therefore a feature of interest; and since they are confined to the head and cesophageal

region of the body, it is probable they aid the animal in its progress amongst the sand.

Within the great longitudinal muscular layer of Valencinia Armandi is superiorly the

cireular muscular coat of the proboscidian sheath, which divides opposite the upper part

of the nerve-trunks (fig. 5, os), and sends a strong band along the ventral surface of the

cesophageal region of the alimentary canal. Between the latter band and the chief longi-

tudinal layer is the great vascular channel (v) on each side.

The external circular muscular layer (¢) shows, some distance behind the mouth, a

tendency to form superiorly two distinct inner bands (fig. 5, ce), extending from one

nerve-cord to the other; and a little further backward the differentiation is observed all

round the body. The circular coat also increases in thickness; but little further change

ensues in this layer in its course backward, except the alteration of its relations to tke

nerve-trunks, which pass from its outer border into the great longitudinal layer.

The great longitudinal muscular layer forms a continuous investment, indented by the

two small vascular canals (vs) superiorly, and the great vascular channels (v) inferiorly

on its inner side, and by the great nerve-trunks externally. As we proceed backward a

tendency to enlargement is noticed at the regions above and below the latter, the coat

* Dredged by Mr. Whiteaves at various points in the Gulf of St. Lawrence.

W. C. M‘INTOSH ON VALENCINIA ARMANDI. vi

generally gets thicker, the median raphe superiorly and inferiorly more distinct, and the

fasciculi more regularly arranged. Then the nerve presses more and more on the circular

coat, penetrates it, indents the longitudinal layer deeply, and finally is included in its

outline (fig. 6, 7), though not at first surrounded by its fasciculi. The next step (which

oecurs a little further backward) is the increase in the relative thickness of this coat

on the disappearance of the great internal circular layer.. The dorsal region dimi-

nishes (in transverse section) to a blunt point at the raphe (fig. 7, 7), forms a consider-

able enlargement at the side of the proboscidian sheath, becomes broad and prominent

internally near the lateral nerve-trunk (which it now encloses), contracts a little again

below the nerve, and increases largely towards the inner edge of the vascular canal (this

being its thickest part), and, lastly, diminishes rapidly towards the ventral raphe. The

arrangement of the fasciculi has now become very regular, viz. in the form of long

rows running, chiefly in pairs, from without inward (fig. 9, d), and which fairly indicate

a stage in the penniform condition of the muscles, observed, for instance (first by M.

Marion), in Amphiporus spectabilis, De Quatref. (Drepanophorus, Hubrecht).

In Valencinia lineiformis the arrangement of the muscular fasciculi in the same region

(from the great vascular trunk downward) takes the form of long narrow rows

(fig. 10, d), quite different in appearance from those in the present form. Dorsally the

rows in VY. Armand: are for the most part directed downward and outward on each side

(fig. 7), then downward and inward, nearly horizontal above the nerve-cord, irregularly

arranged round the latter, and with the chief mass external to the trunk, then upward

and inward below the nerve, and upward and outward from near the vascular canal to the

ventral raphe. The whole field, therefore, has a striking appearance; and the arrange-

ment continues, with little change, to the posterior region of the body. At the tail,

again, the layer becomes somewhat thinner, and the diminution of the proboscidian

sheath causes a greater proportional increase near the dorsal median line, while the nerve-

trunks have the fasciculi only on their inner side.

To comprehend the next muscular layer (the inner circular iayer, fig. 6, 0s), it is

necessary to examine the commencement of the proboscidian sheath. Just in front of

the mouth the sheath (fig. 4, a0) is complete—that is, has externally a strong layer of

circular fibres, and internally a coat of more slenderly developed longitudinal fibres. The

inferior part of the sheath rests (over the mouth) on a series of longitudinal muscular

fibres (z). Behind the mouth a very noteworthy change rapidly takes place, viz. the

gradual absoption of the contiguous portion of the proboscidian sheath into the upper

wall of the digestive chamber, and the envelopment of both by the strong inner circular

layer. The latter is first formed by a splitting of the outer (circular) fibres of the pro-

boscidian sheath (fig. 5, os); and while the coat remaining between the sheath and the

digestive canal gradually diminishes, this new circular belt soon envelops the united

area in a dense muscular mass. Indeed so little of the thin partition between the

sheath and the digestive wall is visible, that at first sight it seems as if both chambers .

floated in a common cavity. After a considerable course this special inner circular layer

diminishes, the proboscidian sheath gets thicker, and its proper layers more evident, and

finally the former disappears, while the latter (fig. 7, a0) continues to the posterior end

M 2

78 W. C. M‘INTOSH ON VALENCINIA ARMANDI.

of the worm. The greatly developed layer of circular muscular fibres must exercise a

special influence on this region of the alimentary canal, probably as an cesophageal con-

strictor. The arrangement is one common in the Carinellide.

The proboscidian sheath (a 0) commences, as usual, at the incurvation in front of the

ganglia, and, as formerly described, becomes more or less atrophied behind the mouth,

while the great inner circular layer developed from its outer fibres acts as the chief

constrictor for this chamber as well as the alimentary canal. At first (figs. 3 & 4, ao) it

clearly possesses two muscular layers, an external circular and internal longitudinal ; but

after the change behind the mouth is completely established, all that can be noticed

inferiorly is a circular band in the tissue of the upper layer of the digestive chamber. A

reversion to the primary condition ensues on the diminution of the inner circular

muscular layer, the proboscidian sheath striking off from the latter and the alimentary

wall as a special tube, which is continued to the posterior end of the worm, where,

however, it forms an extremely small chamber (fig. 8,a0). In Carinella annulata it is

not continued to the posterior region of the body, and, as formerly observed, absence

or diminution of the proboscidian chamber (é.e. the cavity within the sheath) in the

Lineidz and Carinellide is accompanied by greatly enlarged lateral vessels.

The proboscis (figs. 4 & 7, A) agrees with that of the Carinellide ; but its very minute

size prevented an accurate discrimination of the external region into two layers. It had

the aspect of an elastic coat with external circular fibres. A reexamination of more

favourable examples of the proboscides of the Carinellidee has enabled me to correct an

error into which I fell on a former occasion, in stating that the external coat consisted

of elastic, and the next of circular muscular fibres. The latter are external, and the

structureless elastic internal. The next layer in Valencinia Armandi consists of a

powerful longitudinal muscular coat ; then comes a considerable circular muscular layer,

upon which the glandular lining of the organ rests. The glandular papille, as usual,

contain many rod-like bodies. The proboscis terminates posteriorly in a ribbon of lon-

gitudinal fibres.

Digestive System.—The mouth (in transverse section, fig. 3, w) forms a small lon-

gitudinal slit a little behind the ganglia, and is much less prominent than in /. linei-

formis, as well as more anterior in position. It leads into the glandular cesophageal

region, which soon becomes clasped by the powerful internal circular muscular layer

formerly alluded to (figs. 5 & 6,7). The walls of this region are thinner, but of firmer

consistence than the succeeding portion, which commences near the termination of the

special circular muscular layer. In transverse section (fig. 7, 7’) the densest masses of

the refracting (fatty ?) granules, which form the chief portion of the wall, lie on each side

above the nerve-trunks, while a layer beneath the proboscidian sheath joins them supe-

riorly, and a V-shaped bend ventrally completes the circuit. Towards the posterior end

of the body (fig. 8, 7’) the canal assumes the form of an ellipse in transverse section, the

- walls are lax, translucent, and more finely granular, and the cilia appear to be longer.

It is held in position in this region by circular fibres and vertical bands.

Nervous System.—There is nothing special in the structure of the ganglia, which, near

their commissures, are seen in fig.3, h; but the presence of a special longitudinal series

W. C. M‘INTOSH ON VALENCINIA ARMANDI. 79

of separated muscular fasciculi (d’) between them and the basis-layer is a new feature in

the Carinellide. In regard to the nerve-trunks on each side, it is an important fact that

while (in common with the ganglia) they lie clearly outside the proper circular and lon-

gitudinal muscular layers of the body-wall anteriorly, they gradually perforate both in

their progress backward to take up a position in the middle of the longitudinal layer.

Towards the tail the nerve-trunk again gets to the outer border of the latter, and almost

touches the circular coat (fig. 8, 2); but it does not seem to get quite clear of the lon-

gitudinal fibres, some of which form a thin line externally.

In regard to the vascular system, the general features are those of the Carinellid.

There are two great longitudinal trunks (v, v) ; placed within the longitudinal muscular

layer of the body. In the csophageal region (figs. 5 & 6) the trunks agree with

the arrangement in the common Carimella—that is, are situated outside the special

inner circular muscular layer. In the large Canadian Carimella, again, they lie within

this special layer, between it and the wall of the cesophageal region of the alimentary

tract. Anteriorly the trunks terminate in the new species in the vascular channels on

each side of the proboscidian canal, those of opposite sides: communicating in front. The

arrangement, indeed, forms a kind of vascular mesh-work towards the tip of the snout;

and the contents probably exercise an important influence on the mobile tissues of the

region when driven forward by the contraction of the great lateral trunks. At the

mouth (fig. 4) the latter lie nearly opposite the nerve-trunks on each side internally ; in

the anterior region of the body each vessel descends to a lower plane (fig. 6), while pos-

teriorly (fig. 8) the distance between the nerve and vessel greatly increases, the latter

having advanced inward almost to the wall of the centrally situated alimentary canal.

Two other accessory systems yet remain to be described. The first of these consists of

a vessel with highly contractile walls (va, fig. 5), which lies anteriorly within the pro-

boscidian sheath, just above the point where the wall of the digestive canal meets it on

each side. It begins immediately after the mouth, and extends backward only a short

distance ; but its connexions anteriorly and posteriorly have not yet been ascertained.

There can be little doubt about the nature of this structure, which is quite different

from the collections of proboscidian corpuscles and coagulated fluid often seen in similar

positions. The wall of each vessel is definitely formed of a hyaline contractile substance,

in which numerous granules are imbedded. The physiological importance of this arrange-

ment is apparent, and it is especially interesting since the discovery of a regular series of

canals into the cavity of the proboscidian sheath of Amphiporus spectabilis (Drepano-

phorus, Hubrecht). The second system consists in the establishment of a channel on

each side underneath the great longitudinal muscular layer (vs, figs. 4, 5, & 6), as soon

as the upper arch of the proboscidian sheath is formed. These channels assume con-

siderable dimensions, but do not appear to be very contractile, since they have only

a hyaline film all round, and thus stand in marked contrast to the great vascular

channels inferiorly, or even the intra-proboscidian trunks above mentioned. ‘They have

a lengthened course, but disappear towards the posterior end of the body.

None’ of the examples contained reproductive organs; so that this department in

the group is still in need of further investigation. In the large Carinella from Canada.

80 W.C. M‘SINTOSH ON VALENCINIA ARMANDI.

the ovaries are filled with large ova, and stretch from a little below the nerve-trunks on

each side upward to the dorsal region of the great longitudinal muscular coat, and

encroach considerably on the alimentary chamber behind the proboscidian sheath,

EXPLANATION OF PLATE.

Prats XVI.

The following letters are used to designate the respective parts throughout the series :—

a. Cutis.

b. Basis-layer of the cutaneous system.

c. Cireular muscular coat of the body.

d. Longitudinal muscular layer of the body.

ao. Proboscidian sheath.

d’. Special layer of separated longitudinal muscles anteriorly.

Ah. Ganglia.

j. Wsophageal region of the digestive system.

j.. Alimentary canal proper.

mn. lateral nerves.

r. Median raphe.

v. Great longitudinal vascular trunks.

w. Mouth.

A. Proboscis.

Fig. 1. Transverse (vertical) section near the tip of the snout of Valencinia Armandi: v, v, vascular

meshes by which the great trunks unite. x 210 diam.

Fig. 2. Transverse section of the snout behind the former. The two horizontal bands of fibres (e and e’)

have now attained considerable development ; and numerous vertical fibres pass from their sur-

faces to the circumference: ao, canal for the proboscis in the snout. Numerous longitudinal

muscular bundles (d) occur along the inner faces of the horizontal bands and in the forks of the

vertical series. x 210 diam. i

ig. 3. Transverse (and somewhat flattened) section just in front of the mouth, through the ganglia (A).

The separated series of longitudinal muscles (d’) are very conspicuous in this region, though a

process of the mouth (w) interrupts the circuit inferiorly. The proboscidian sheath (ao) is some-

what flattened out. The two accessory vascular meshes, s, s, are probably homologous with

those found in a similar situation in the typical Anopla; m, powerful transverse muscular band,

separating the chief from the accessory vascular channels ; m’, ring of circular and longitudinal

fibres surrounding the united area of the proboscidian and the vascular systems. In this pre-

paration the proboscidian sheath is differentiated from the general investment of the united area

les]

very distinctly. x 90 diam.

Fig. 4. Similar section through the mouth. The separated series of longitudinal muscles (d’) are still

very evident. They form aconsiderable mass on each side of the mouth inferiorly. The basis-

layer (b) of the cutaneous system is greatly developed im this region. The proboscidian sheath

is wholly independent of the digestive system, a special band of longitudinal fibres (z) besides

the sheath proper and other tissues intervening. The commencement of the great longitudinal

W. C. M‘IINTOSH ON VALENCINIA ARMANDI. 81

layer of the body-wall is seen at d; vs, vs, the two dorsal accessory vascular channels between the

proboscidian sheath and the longitudinal muscular coat. x 90 diam.

.5. Section through the anterior esophageal region, showing the intraproboscidian vessel, va. The

proboscis, A, has been cut obliquely. The dorsal accessory vascular channels, vs, are very

distinct. The cutaneous basis-layer, 6; shows traces of the separated series of longitudinal fibres,

but they are insignificant. At os is a slight inflection of the inner circular coat, indicating the

point of separation of the thin layer still representing the proboscidian sheath. x 90 diam.

Fig. 6. Section through the internal circular muscular layer, os, in its complete condition. x 90 diam.

Fig. 7. Transverse section behind the esophageal region, and after the disappearance of the special inner

circular muscular layer (os, fig. 6). The great development of the longitudinal muscular layer of

the body is apparent, as well as the regularity of its arrangement; o/, inner (longitudinal) fibres

of the proboscidian sheath ; oc, outer (circular) of the same. x 90 diam.

Fig. 8. Section about one fifth of an inch from the tip of the tail, somewhat flattened by compression. The

proboscidian sheath (ao) is now very minute. The great longitudinal muscular layer (d@)

presents no regular arrangement of its fasciculi; and the lateral nerve-cord has almost reached

the circular coat (ce) exteriorly. The area on each side of the now attenuated alimentary

So]

canal is filled up by a translucent fibrous stroma, which in this preparation has been much

stretched, especially on the left side. x90 diam.

Fig. 9. Portion of the body-wall towards the middle of V. Armandi, from the great lateral vessel outwards,

showing the arrangement of the muscular fasciculi of the longitudimal layer. x 350diam.

Fig. 10. Section of a similar portion in Valencinia lineiformis. The great thickness of the basis-layer is

conspicuous. The arrangement of the longitudinal muscular fasciculi also contrasts with the

former. x 350 diam.

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THE

TRANSACTIONS

THE LINNEAN SOCTETY

LONDON.

SECOND SERIES.—ZOOLOGY.

VOLUME I.

PART THE THIRD.

1 OMNGDE OR INE:

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET

SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON-HOUSE;

AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER-ROW.

M.DCCC.LXXVI.

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CONTENTS.

PART III.—Jouty, 1876.

VI. On thirty-one Species of Marine Planarians, collected partly by the late Dr. KELAART,

F.L.S., at Trincomalee, and partly by Dr. Couuinewood, F.L.S., m the Hastern

Sco o Gera COULINGWOOD) |). eey ia eles Sereda oss ey) ePAgserss

VII. On the Structure and Development of the Bird’s Skull. By W. K. ParKer,

ERAS LS Gio (UR ALbe DI) is oe Pe a ee Me eg SOD

[ 83 ]

VI. On thirty-one Species of Marine Planarians, collected partly by the late Dr. Kevaart,

F.L.S., at Trincomalee, and partly by Dr. Couinewoop, E.LLS., in the Eastern

Seas. By Dr. CoLLINGwoop.

(Plates XVII. to XIX.)

Read March 18th, 1875.

Convenrs.

I. Prefatory and introductory. IV. Description of twelve new species of marine Pla-

II. Characters of two new genera of Kuryleptide. narians collected by Dr. Collingwood, and deli-

III. Synopsis of the genera and species described and neated in the accompanying figures.

figured in the present paper. Y. Dr. Kelaart’s descriptions of his figures of marine

Planarians collected in Ceylon.

THE species of Planarians which I am about to lay before the Society were, as far

as I was concerned, the fruit of many searches upon rocky shores of the coast of China,

Borneo, Singapore, &c., and represent a considerable amount of labour near low-

water mark in turning over stones, pieces of coral, &c. in such situations. The little

animals in question, although extremely interesting, are all more or less minute; and it

requires not only a sharp but an educated eye to discover them; for so delicate are they,

and often so thin in texture, that out of the water, and adhering to a more or less muddy

or weed-grown stone, it is by no means easy to detect them. It must not be under-

stood, however, that these searches were unrewarded except by the comparatively small

number of specimens or new species here recorded. Many other animals of interest

divided the attention, as, for instance, nudibranchiate Mollusca, of which a number

of new species await the notice of the Society, and other no less interesting creatures.

The Planarians in many respects rival the Nudibranchs in gracefulness of form and

beauty of colouring; but they are even more delicate than the latter. When a small

spot of colour upon the under surface of a stone betrays the presence of one of these

little animals, a camel’s-hair brush is necessary to dislodge it safely ; and a bottle of salt

water must be at hand, into which to transfer the prize.

It may be thought that a dozen species is but a small number to reward one’s labour

on coasts which are supposed to abound with unknown forms of animal life. But

a good deal of experience in shore-hunting on our own coasts prepared me for the

fact, which I found amply corroborated on foreign shores, that we must not look for

a large number of species at any one locality, unless the same locality is examined many

times, and at different seasons of the year—conditions not always, or, indeed, often

practicable. These little organisms appear to be sporadically distributed over wide

areas. ‘They are seldom met with gregarious; and three or four species captured upon

any given occasion might be reckoned a prize more than ordinarily satisfactory.

SECOND SERIES.—ZOOLOGY, VOL. I. N

84: DR. COLLINGWOOD ON THIRTY-ONE

Although, however, I never lost an opportunity of examining a shore when I could do

so towards low tide, the opportunities for such rambles were necessarily limited.

And yet, although they appeared to be so sparsely distributed, a fact would come to

light every now and then which seemed to point out that they were very abundant in

the aggregate. For instance, a single visit to a certain spot would perhaps result in the

discovery of some minute organism identical with one found on another occasion at a

point separated perhaps by a hundred miles from the first-mentioned locality, or, as was

the case in some instances, by the whole extent of the China Sea. How was it that

I should stumble upon such an inconspicuous organism a second time at a spot so

distant from the first, unless they were abundantly distributed through the inter-

mediate spaces? Doubtless an explorer at the same places would still find many new

forms, and leave yet many more for a successor.

With regard to the Planarians collected by the late Dr. Kelaart, and included in this

collection, it is necessary that I should say a few words concerning how they came into

my possession. The late Mr. Albany Hancock, of Newecastle-on-Tyne, hearing from me,

in the course of correspondence, that I had collected some marine Planarians which

appeared to be new, informed me that he had in his possession a number of drawings

of similar animals made by Dr. Kelaart (formerly a Fellow of this Society) from speci-

mens collected by himself in Ceylon, and principally at Trmcomalee. These drawings

were briefly characterized by him in a paper “ On some new and little-known Ceylon

Invertebrata,’ published in the Journal of the Ceylon branch of the Royal Asiatic

Society. Part of this paper was printed in the ‘ Annals of Natural History’ for 1859;

but the portion relating to the Planarians was not so published; nor, need I say, were

the drawings published at all. Dr. Kelaart issued a ‘ Prodromus Faune Zeylanice,’ a

portion of which is in the Society’s Library, and which contains the description of the

Planarians here figured ; but I have nowhere met in the literature of the Planarians

with any reference to either of these publications, which appear to be unknown.

Mr. Hancock therefore suggested that if I could publish Kelaart’s species along with my

own, they would, as he expressed it, ‘‘ make a very interesting monograph on the

subject—would, in fact, make a valuable contribution towards the history of these little-

known animals of the eastern seas.”’ He therefore sent me the drawings, which, re-

garding them as a testament jointly from Dr. Kelaart and Mr. Hancock, both deceased,

I have accordingly submitted to a careful examination, and done my best to distribute

them according to the genera and species of the best classification. I may say that,

with Mr. Hancock, I consider these drawings are sufficient for identification; but they

are unaccompanied by details, and the descriptions of them are brief. This is to be

regretted, since the value of such drawings depends no less upon the accuracy of the

verbal descriptions taken from the living animals than upon the correctness and minute-

ness with which they are depicted. Unfortunately the animals are so extremely

perishable and delicate, that no means yet devised can possibly exhibit them after

death in any thing approaching to their natural appearance or beauty of form and

brilliancy of colouring. Good drawings and accurate descriptions, therefore, are of the

utmost value and importance; and either one without the other is consequently of

SPECIES OF MARINE PLANARIANS. 85

impaired value. Of my own drawings I can, of course, vouch for the faithfulness. They

were always taken from the living animal while in health and activity ; and any thing

striking and remarkable could not fail to be noticed and recorded.

The term Planaria, which was at first applied to these animals as a generic name for

them all, has been long since restricted to a small division of them which is not marine.

Linnzeus, Pallas, Lamarck, and Cuvier placed them among parenchymatous Entozoa ;

but the investigations of Dugés, von Baer, Quatrefages, and others have by degrees raised

them to their proper and natural position. This is now assigned to them under the

Turbellarian Annelids. The genera described by Ehrenberg and De Quatrefages out of

the old genus Planaria have in turn for the most part been split up or given place to

more recent classification. Ihave endeavoured to assign positions to the present species

under the arrangement of Diesing (‘ Systema Helminthum’), as revised by W. Stimp-

son in his ‘‘ Prodromus descriptionis animalium evertebratorum” of the South-Pacific

Federal Expedition, under Capt. Jno. Rodgers (Proc. Acad. Nat. Scien. Philad. 1857,

p- 19 &c.). Stimpson enumerates thirty-eight genera of Turbellaria Dendroccela (Max

Schultze, Wiegm. Arch. 1849, p. 292), including several new genera resulting from that

expedition. Of these thirty-eight genera, eight are fluviatile, four terricolous, one inhabits

brackish water, and oneis pelagic. The remaining twenty-four consist of ordinary marine

genera, under which group the present species would be all included. I have ventured

to constitute a new genus of two of the Ceylon species, from a character which does not

appear to be shared with any of those previously described. Inasmuch, however, as all

the Ceylon species have already been named by their collector and delineator, I shall

of course adopt his specific terms, only inserting them in what appears to me to be their

proper generic group.

The marine Turbellaria Dendroceela are for the most part flat, thin, soft, and delicate

animals, remarkable for their beautiful colours and often graceful forms and movements.

They have usually a pair of tentacles, these occasionally being mere folds or ear-like projec-

tions of the anterior margin (Huryleptidze)—sometimes true tentacles, situated occipitally

(Stylochidz), or even dorsally, in which case they are retractile (Planoceridz). In some

families, however, tentacles are entirely absent (Typhloleptidee and Leptoplanidee). The

margin is usually ample, and much folded or puckered, which is very noticeable when

the animal is at rest; but when in motion, this ample pallium is gracefully waved

in a vertical direction, and aids greatly such species as are natatory in swimming,

which they effect with ease and rapidity. They possess either two genital apertures (sub-

tribe Digonopora), as is the case with all the present species, or only a single one (sub-

tribe Monogonopora), as is the case with the true Planariade. The mouth is variously

situated on the median line of the under surface, and is in turn antecentral, subcentral,

or postcentral; and the dendriform gastrovascular ramifications are often more or less

visible upon the under surface.

One of their most peculiar characteristics consists of the eye-spots, small black specks

of varying size and position, sometimes clustered into congeries of a symmetrical form,

and sometimes scattered over the anterior and marginal portions of the upper surface, or

on the tentacles. These eye-spots, in some cases at least, consist of a highly refracting

86 DR. COLLINGWOOD ON THIRTY-ONE

spherical body (crystalline), surrounded with pigment; and they are always situated in a

part which is free from the usual pigmentary matter of the animal, and where the skin

is reduced to a minimum of thickness (Baer). In any case they can but very imperfectly

perform the function of vision; and in one family (‘Typhloleptide) they appear to be

altogether absent *. Nevertheless these animals all seem to avoid the light, by concealing

themselves under stones and in situations where no light can reach them.

Their ova are deposited in thin flakes upon sea-weeds and stones, and in appearance are

not unlike those of the Nudibranchs, for which at first sight they might be mistaken.

They resemble these mollusks also in their habit of floating, back downwards, upon the

surface of the water. Nor do the resemblances end here ; for some genera of the Hury-

leptidee are provided with numerous papillze, more or less covering the whole upper sur-

face, after the manner of an Holis, which character earned for them the generic name

of Holidiceros from De Quatrefages.

It has often been said, and is copied from one book into another, that these animals

have the power of spontaneous fission, by means of which two individuals are produced

from one. I can only say that, of the numerous specimens I have met with, kept alive,

and carefully watched, I never saw any thing like this process take place in any instance.

Still I cannot assert that it does not happen.

It is worthy of remark that among the 19 species of Dr. Kelaart, no less than 11

appear to be referable to the genus Hurylepta (Hemp. and Ehrenb.), while of my own 12

species only one is of that genus. Mot one of Kelaart’s claims rank under Quatrefages’

old genus Proceros, in which one fourth of my own species oceur. In each collection is

found a new genus of Euryleptidz ; and the whole of the 31 species are included under

one subtribe—Digonopora, or monecious Turbellaria.

If.

(Characters of two new genera of Burylepide.)

TURBELLARIA DENDROCGLA.

Genus ACANTHOZOON, Coll.

Caput subdiscretum, tentaculis parvis approximatis. Corpus supra spinulis brevibus nigris ubique

imstructum.

Thysanozoon,

ACGANTHOZOON, Proceros,

Hurylepta.

Genus SPHYNGICEPS, Coll.

Corpus leve, caput discretum, tentaculis magnis subdistantibus ; ocelli occipitales et capitales.

Proceros,

SPHYNGICEPS,

Eurylepta.

* With regard to the eye-spots, structurally and as supposed organs of vision, consult H. N. Moseley’s remarks

thereon, ‘‘ Anat. and Histol. Land-Planarians of Ceylon,” Proe. Roy. Soc. 1873, p. 172 &e.

SPECIES OF MARINE PLANARIANS. 87

(Synopsis of the Genera and Species described and figured in the present Paper.)

[Note—Diesing’s main divisions were founded upon the presence or absence of tentacles ; but Stimp-

son pertinently remarks that such a division separates allied forms, such as Stylochus and Leptoplana,

while widely separated ones, such as Hurylepta and Planaria, were approximated. In view of this I

therefore follow Stimpson in foundmg the subtribes upon the presence or absence of a second genital

aperture, instead of the mere presence or absence of tentacular appendages.—C. C.]

TURBELLARIA DENDROCGLA.

(Tubus alimentarius dendritice ramosus.)

Subtribus I. Digonovora. Two genital apertures.

(All the species are included in this subtribe.)

Family EURYLEPTID&.

Genus THYSANOZOON, Grube.

1. Thysanozoon Alderi, Coll. Labuan.

2. Thysanozoon Allmanni, Col/. Singapore.

3. Thysanozoon auropunctatum, Kel. Aripo.

Genus AcANTHOZOON, Col'., n.g.

4, Acanthozoon armatum, Kel. Ceylon.

5. Acanthozoon papilio, Kel. Ceylon.

Genus SpHyNneicers, Coll., n. g.

6. Sphyngiceps lacteus, Coll. Singapore.

Genus Proceros, Quatref.

. Proceros concinnus, Coll. Labuan.

. Proceros Hancockanus, Coll. Singapore.

. Proceros Buskiu, Coll. Singapore.

<> OO

Genus Hunyuepra, Hemp. & Ehren.

10. Eurylepta fusca, Kel. Ceylon.

11. Eurylepta atraviridis, Kel. Ceylon.

12. Hurylepta undulata, Kel. Ceylon.

13. Hurylepta violacea, Kel. Ceylon.

14. Eurylepta dulcis, Kel.

15. Eurylepta purpurea, Kel. Ceylon.

16. Eurylepta viridis, Kel. Ceylon.

17. Eurylepta affinis, Kel. Ceylon.

18. Eurylepta cerebralis, Kel. Ceylon.

19. Eurylepta striata, Kel. Ceylon.

20. Eurylepta zeylanica, Kel. Ceylon.

21. Eurylepta Kelaartu, Coll. Simgapore.

DR. COLLINGWOOD ON THIRTY-ONE

Family TY PHLOLEPTID &.

Genus TYPHLOLEPTA, Oerst.

. Typhlolepta Byerleyana, Coll. Pulo Barundum, west coast of Borneo.

Family LEPTOPLANIDA.

Genus CenTRostomum, Diesing.

. Centrostomum ocellatum, Kel. Ceylon.

. Centrostomum punctatum, Kel. Ceylon.

Genus EiasmopEs, Le Conte.

. Elasmodes obtusus, Coll. Singapore.

Genus LepropLana, Hemp. et Ehr.

. Leptoplana patellensis, Co//. Simon’s Bay.

. Leptoplana aurantiaca, Coll, Singapore.

Family STYLOCHID &.

Genus STYLOCHOPLANA, Stimpson.

. Stylochoplana elegans, Kel. Ceylon.

. Stylochoplana meleagrina, Kel. Ceylon.

Genus StyLocuopsis, Stimpson.

. Stylochopsis malayensis, Coll. Pulo Barundum, west coast of Borneo.

Family PLANOCERIDA.

Genus PLanocera, Blainv.

. Planocera thesea, Kel. Ceylon.

IV.

(Descriptions of the Marine Planarians collected in the Eastern Seas by Dr. Colling-

wood, F.L.S., and delineated in the accompanying figures.)

1. Thysanozoon Alder.

Length 2+ inches; breadth 14 inch.

Body thin, with very irregular margin, amply folded and puckered.

Upper surface of a general light brownish colour, with a narrow, pale external margin,

within which is a broad, black border, somewhat shaded and marbled. Down the

median line for about three quarters of its extent runs an irregular, black marbling ;

a faint marbled pattern of pale brown is diffused over the general surface; and a

lens discloses also a fine ramification of a darker tinge throughout.

The whole upper surface is studded with small papillee of a conical form, the foot-

stalks of which are pale, and the distal extremities orange. Many of these papille

SPECIES OF MARINE PLANARIANS. 89

arise from an elevated white spot or tubercle, such tubercles producing only one

papilla each; and other papille exist upon the black margin, as well as on the

general surface.

Under surface whitish, edged with black, the part answering to the black marbled line

on the dorsum being here opaque white.

‘Head blackish, angular, raised somewhat above the general plane of the body, flexible,

and having two projecting angles or folded tentacles.

This was a very beautiful and striking animal, looking as if studded with golden beads.

it swam freely, with a graceful vertical vermicular movement of the ample sides.

One was found under stones about two feet under water at low tide, upon a reef of

the island of Labuan, coast of Borneo, on August 22. I have great gratification in

naming it after the late Mr. Joshua Alder, of Newcastle, a gentleman with whom I was

long in correspondence, and from whom, though we never met, I always received

uniform kindness and urbanity.

Fig. 1: a, general appearance of upper surface; 6, head and tentacles; c, portion of upper surface,

enlarged, showing papille; d, a papilla; e, a papilla-bearing tubercle.

2. Thysanozoon Allmani, Coll.

Length 23/5 inches ; breadth ? inch.

Body translucent, papillose.

Upper surface light brown, becoming darker towards the margin, and with an irregular

edging of opaque white all round, excepting the head. An elevated ridge runs

along the median line of the dorsum.

‘The whole upper surface is covered with clavate and pointed papille, of a deep

brown colour, and varying in size, the smallest being the lightest-coloured and most

numerously clustered, and occurring along the median ridge.

Under surface grey, darkening to deep brown at the sides, and edged with opaque

white.

Head with two long tentacles, often thrown back, and presenting the appearance of

hare’s ears. Tentacles dark brown, tipped with white. Twominute white tentacles

are situated in front of the head, beneath the hare-like ones.

Hye-spots situated in a light-coloured spot immediately. posterior to the head, in a double

cluster, consisting of two small crescentic patches of minute black spots.

A splendid species, the edges much corrugated, unless the animal is in full movement,

when the folds almost disappear. It swims freely, with a graceful serpentine movement,

and crawls with moderate rapidity and with an undulating movement. When at. rest

the animal assumes a nearly circular form, with numerous marginal folds.

Two specimens found at Singapore, west of the harbour, under pieces of dead coral,

on the beach between tide-marks, Nov. 22nd.

I have much pleasure in connecting this species with the name of the President of this

Society, so well known for his researches among the Invertebrate division of animais.

Fig. 2: a, animal in movement (dorsal surface) ; 6, papille, large and small, magnified; c, eye-spots

(magnified).

90 DR. COLLINGWOOD ON THIRTY-ONE

3. Sphyngiceps lacteus, Coll.

Length 4% inch ; breadth { inch.

Body graceful in form, semitransparent.

Upper surface cream-coloured, irregularly spotted with sparse and minute black dots,

and having faint marbling on either side of the median ridge, which is very con-:

spicuous. Margin irregularly blotched with red, and the whole body edged with a

narrow black line.

Under surface whitish, edged with red blotches and a black streak as on the upper side,

but somewhat less distinct. A broad streak of white occupies the anterior third

of the median line.

Head small and narrow, but rendered very conspicuous at times by its being raised up,

so that the median ridge is thus very much elevated in front. Head furnished

with two folded tentacles.

Eye-spots round, immediately posterior to the head, which has in front of it a pair of

larger single spots.

This is a remarkably graceful little animal, crawls with great rapidity, and with a

gliding movement, then suddenly throws its head back, and, leaving the side of the vessel,

swims with a graceful serpentine movement, sometimes very fast ; floats also on its back

on the surface of the water. It has the habit of throwing back its head very far, and

remaining in that position (rampant, as it were), at which time the head and anterior

portion of the body become narrow and tubular, giving the appearance of a neck. When

at rest it expands laterally, becoming in shape as broad as long.

One specimen found under a coral block, west of Singapore Harbour, at low water, on

Noy. 22nd.

Fig. 3: a, animal in motion; 0, head, enlarged, showing eye-spots.

4. Proceros concinnus, Coll.

Length 2 inch.

Body narrow, entire, without marginal foldings.

Upper surface cream-colour, approaching to yellow, with an edging of blue all round,

composed of small and larger spots running into one another. A similar blue streak

runs along the median line from a little behind the head to some distance from the

posterior extremity, through about three quarters the length.

Under surface cream-colour, similarly edged with blue, a brownish streak running

through the whole median line, the arbusculiform alimentary tube of a straw-

colour, occupying the anterior third. .

Head with two folded tentacles.

Eye-specks conglomerated in a small round spot midway between the head and the blue

median line.

A very pretty and lively species, particularly fond of floating on its back or crawling

rapidly along. I obtained one specimen on a reef at Labuan, near low-water mark, and

SPECIES OF MARINE PLANARIANS. 91

a second at Pulo Daak, a small island between Labuan and the mainland of Borneo,

August 25th.

Fig. 4: a, animal extended, upper surface ; 4, animal extended, under surface; c, eye-spots.

5. Proceros Hancockanus, Coll.

Length 1,45 inch; breadth ;8; inch.

Body velvety, opaque.

Upper surface of a deep velvety brown, edged with a double margin of equal widths, the

inner deep orange, the outer opaque white. Along the centre of the back was a

slightly elevated ridge.

Under surface grey, darkening towards the sides.

Head small, tentacles simply folded, long and graceful, the orange margin disappearing,

and the white alone being prseent.

Hye-specks in an oval elongated cluster, immediately posterior to the head.

A very beautiful species, no less from its rich colour than from its active movements.

Swimming by an undulating movement, it also floated on its back on the surface of the

water, after the manner of the Nudibranchiata. It crawled slowly, and with an undu-

lating motion. In texture it appeared somewhat translucent, and when suddenly dis-

turbed became much paler, as though by a sudden contraction of the tissues. In one

specimen a temporary slit appeared, occupying the middle third of the dorsal ridge,

rounded anteriorly, wedge-shaped posteriorly, through which slit the internal organiza-

tion was clearly visible. The slit remained open one day, and afterwards closed. The

intestines appeared sometimes as though protruding through this slit, which was not

exhibited by a second specimen. On placing one of these animals in glycerine, it imme-

diately discharged a quantity of a brownish-coloured fluid, and contracted considerably

in bulk.

Two specimens were obtained on succeeding days from among stones and old coral

blocks, between tide-marks, west of Singapore Harbour, Nov. 21st.

This species I have much pleasure in connecting with the name of the late Mr. Albany

Hancock, the coadjutor of Alder, from whom, although we never met, I received many

| kindly letters up to near the time of his lamented decease.

Fig. 5 : a, the animal in motion, exhibiting the open dorsal fissure; 4, the cluster of eye-specks.

6. Proceros Buskii, Coll.

Length 4 inch; breadth } inch.

Body opaque, flat, smooth.

Upper surface rich velvety olive-green, edged with pale yellow. :

Under surface dark grey, the dendritic marking whitish, and occupying the anterior half

of the median line.

Head with two folded earlike antennal projections.

Eye-spots in a circular cluster, difficult to detect owing to the dark colour of the animal,

and situated upon a ridge formed by the elevation of the antennal head.

SECOND SERIES.—ZOOLOGY, VOL. I. fc)

92 DR. COLLINGWOOD ON THIRTY-ONE

A very handsome active species, crawls rapidly, at the same time throwing back its

head, and floats, back downwards, on the surface of the water. One specimen only, found

under a stone, and upon a small grey incrusting sponge, between tide-marks in Singa-

pore Harbour, west of the town, December 3rd.

I have named this species after my friend Mr. Busk, than whom no one has been

more assiduous or successful in his investigations among the lower forms of Inver-

tebrata.

Fig. 6: a, animal at rest, dorsal surface; 6, head.

7. Eurylepta Kelaartu, Coll.

Length ;45 inch; breadth + inch.

Body small, smooth, thin.

Upper surface mottled dark purple, the mesial line presenting a slightly elevated ridge

of a darker colour.

Under surface the same colour as the upper, but paler.

Head small, with minute ear-like processes.

Hye-spots minute, roundish.

An actively crawling creature, occasionally throws its head back. Two specimens

were found in Singapore Harbour, west of the town, under stones, and apparently feeding

upon a small incrusting sponge. One of these specimens was of a lighter lake colour.

December 3rd.

As out of the 19 species delineated by Dr. Kelaart I have referred no less than 11 to

the genus Hurylepta, while of my own 12 this is the sole one of that genus, I have

dedicated it to him—a compliment to his memory which his industry in this as well as

in other branches of zoology well deserves.

Fig. 7: a, animal in motion; 0, eye-spot, magnified.

8. Typhlolepta Byerleyana, Coll.

Length ? inch ; breadth 2 inch.

Body smooth, thin, and the lateral parts very ample and puckered.

Upper surface beautifully marbled with light-brown rings, including roundish spaces of

a whitish colour, smaller rings being between the interstices of the larger; most

crowded and darkest in colour along the median line, paler and more delicate

towards the sides.

Under surface of a pale grey, the dendritic marking in the centre of an opaque white.

This very beautiful species I obtained from under a piece of coral on Pulo Barundum,

off the west coast of Borneo. Not having a brush at hand, I had great difficulty in dis-

lodging it from the crevices. Its movements were very contorted, and it did not exhibit

much activity. One specimen only was procured, October 6th.

I have named this elegant species after my friend Mr. J. Byerley, F.L.S., of Seacombe,

Cheshire, a gentleman who in the hardly-spared intervals of active practice has ex-

SPECIES OF MARINE PLANARIANS. 93

hibited the greatest interest in zoological science, and with great industry and labour

compiled the ‘ Fauna of Liverpool.’

Fig. 8. General appearance of upper surface.

9. Hlasmodes obtusus, Coll.

Length -§;; breadth + inch.

Body thin, delicate, smooth, semitransparent, dendritic.

Upper surface pale brown, with a dark, shaded brown streak along the middle half of the

median line, from which radiate delicate pale brown markings, which fade as they

reach the margin.

Under surface pale, and exhibiting the radiating markings, though fainter.

Head indistinct.

Eye-spots two, somewhat crescentic, situated upon a round, white, transparent space

immediately in front of the median dark brown streak.

A very lively, active, and graceful animal. When in motion and fully extended it has

a broad or truncated appearance anteriorly. It crawls rapidly, floats feet uppermost,

and swims gracefully and rapidly by a vertical movement of the sides of the body, which

is performed by quick and sudden jerks.

One specimen was obtained in Singapore Harbour, west of the town, from beneath a

stone between tide-marks, December 3rd.

Fig. 9. Upper surface of the animal.

10. Leptoplana patellensis, Coll.

Length $ an inch.

Body entire, opaque.

Upper surface cream-colour, smooth, beautifully mottled with rich light brown. A

ridge runs along the median dorsal line, irregularly marked with a darker brown,

from which the general mottling radiates to the margin, where it is palest.

Under surface of an opaque whitish colour, the dendritic marking occupying the middle

third, of an opaque white, and surrounded with dots of the same.

Eye-spots irregular and indistinct, consisting of an oval ring at the anterior part of the

median line, and on either side an irregular patch, that on the left roundish, and on

the right crescentic. ‘These spots when magnified appear roundish, but not circular,

and do not present any regular figure.

This animal moves with a leech-like motion, fixing itself by its anterior and posterior

ends alternately. It showed no inclination to swim or float like most of the Planarians,

nor even to leave the bottom of the vessel of water in which it was contained.

Two specimens were obtained from under the mantle of a large limpet (Patella oculus),

which I knocked off with my foot from the granite boulders of Simon’s Bay, Cape of

Good Hope, May 23rd.

Fig. 10: a, upper surface, animal in motion ; 0, head, enlarged, showing arrangement of eye-spots.

0 2

94 DR. COLLINGWOOD ON THIRTY-ONE

11. Leptoplana aurantiaca, Coll.

Length ;%; inch; breadth ¢ inch.

Body slender, semitransparent, entire.

Upper surface, general colour orange-chrome, with a median ridge of a pinkish colour.

From this ridge radiate a number of minute dendritic processes of a bright chrome-

colour, which approach the margin, where the body becomes perfectly translucent.

Sparse white spots are scattered irregularly over the general surface.

Under surface similar to the upper, only paler, as though from the colour being seen

through the semitransparent body.

Head. The anterior portion of the body is without a distinct head or tentacles, but

notched, and apparently folded; on the left side of the notch appeared a tentaculi-

form process tipped with a black spot, and having also two or three small black

specks in its neighbourhood.

Hiye-specks in a hippocrepiform congeries immediately anterior to the pik median line;

the spots few, and larger than usual compared with the smallness of the animal.

This species moves somewhat slowly, crawling like a slug, and also swims or floats

upon its back, like the nudibranchs, upon the surface of the water.

I obtained several specimens between tide-marks under stone westward of Singapore

Harbour in November. |

_ Fig. 11. Upper surface of the animal.

12. Stylochopsis malayensis, Coll.

Length 13 inch; breadth 3 inch.

Body smooth, folds ample; general colour a rich, velvety, deep brownish black, with a

narrow border of deep chrome, external to which is a second narrow edge of dull

white.

Under surface nearly as dark as the upper, with a central irregular line of rose-colour.

Tentacles large, separate, anterior, supporting ocelli.

One specimen found under a coral block on Pulo Barundum, west coast of Borneo,

between tide-marks, Oct. 6.

Fig. 12. Upper surface of the animal.

Vie

(Dr. Kelaart’s description of his figures of Marine Planarians taken at Ceylon, all of

which, except two, he included under the genus Planaria. They are therefore, in

this list, referred to their proper genera.)

Genus THysANOozoon, Grube.

18. Thysanozoon auropunctatum, Kel.

A large species.

Upper surface, a vich violet brown, darker in the centre, and edged all round with a

‘SPECIES OF MARINE PLANARIANS. 95

border of pure white. Thickly studded with papille, small and conical, the bases of

which are black, the apices golden yellow, and the intermediate band white.

Under surface pale purple, very dark towards the margin all round, but having the

narrow white border as above. :

Head furnished with two small rudimentary tentacles. Mouth situated between the

middle and anterior third.

Found at Aripo, 28th February.

Fic. 13: a, upper surface of animal; 0, papill, magnified.

Genus ACANTHOZOON, Coll., n. g. (see p. 86).

14. Acanthozoon armatum, Kel.

Length 14 inch; breadth 14 inch.

Upper surface of a dark purple colour, covered with short black spines.

Under surface pale purple, smooth.

Tentacles folded, but somewhat distinctly formed.

Fig. 14. Upper surface. j

15. Acanthozoon papilio, Kel.

Length about 1 inch.

Upper surface yellow, covered with small black spines; margin whitish.

Under surface pale yellow.

Tentacles as in last species, black, tipped with white.

Looks very like a butterfly moving in the water.

Fig. 15. Upper surface.

Genus EuryLerta, Hemp. et Ehr.

16. Eurylepta fusca, Kel.

Length 13 inch.

Upper surface dusky brown.

Under surface paler brown.

Tentacles inconspicuous and approximated.

Fig. 16. Upper surface.

17. Eurylepta atraviridis, Kel.

Upper surface dark mottled green, with a darker broad streak through the whole median

line. An edging of pale green runs all round.

Tentacles like the last.

Fig. 17. Upper surface.

18. Hurylepta undulata, Kel.

Length 2 inches.

Upper surface pale yellow, with undulating lines and spots of purplish brown, producing

a marbled appearance. Margin and median line purplish.

Tentacles rudimentary.

Vig. 18. Upper surface.

96 DR. COLLINGWOOD ON THIRTY-ONE

19. Hurylepta violacea, Kel.

Length 1} inch; breadth ? inch.

Upper surface violet purple, edged with bright yellow; median line yellowish.

Under surface rose-coloured.

Tentacles rudimentary.

Ova yellowish.

This beautiful species, in a quiescent state, resembles some variety of Pansy (Kel.). It

appears to be nearly allied to Planaria zebra, Leuck. See Riippell’s Atlas, t. 3. f. 1a, 6.

Fig. 19: a, upper surface; 4, under surface.

20. Eurylepta dulcis, Kel.

Length. 1 inch.

Upper surface light green, minutely spotted with reddish brown; margin white; median

line brown.

Tentacles rudimentary.

Fig. 20. Upper surface.

21. Hurylepta purpurea, Kel.

Length about 14 inch.

Upper surface of a beautiful purple colour.

Under surface paler purple, darkening towards the margin.

Tentacles very small and rudimentary.

Fig. 21. Upper surface.

22. Hurylepta viridis, Kel.

Length about 1} inch.

Upper surface green, spotted with brown; margin darkly grizzled brown.

Under surface paler green. ;

Tentacles rudimentary, small, brown.

This species appears to be nearly allied to Planaria limbata, Leuck. See Riippell’s

Atlas, t. 3. f. 4.—C. C.

Fig. 22. Upper surface.

23. Hurylepta affinis, Kel.

Upper surface purple, with yellow border.

Tentacles very small and rudimentary.

Fig. 23. Upper surface.

24, Hurylepta cerebralis, Kel.

Length 53 inches; breadth 3 inches..

Upper surface of a yellowish brown colour, and minutely streaked with fine wavy brown

lines; border ample, edged with black and streaked with white.

Under surface of a delicate salmon-colour, with a narrow blackish border.

Head with rudimentary tentacles, formed by two folds of the margin; mouth large,

placed on the anterior third of the lower surface; lips white.

Ova greenish white.

SPECIES OF MARINE PLANARIANS. 97

This was the largest specimen observed. Its colour and the ample foldings of the

margin call to mind the appearance of convoluted brain-substance.

Fig. 24. Upper surface.

25. Eurylepta striata, Kel.

Length 24 inches.

Upper surface brownish purple, streaked with brown; marginal folds ample, and edged

with a narrow border of dark brown.

Under surface pale orange-brown, darker towards the margin, and edged with a narrow

border of brown.

Fig. 25. Upper surface.

26. Eurylepta zeylanica, Kel.

Length 23 inches; breadth 13 inch.

Upper surface dark purplish chocolate-brown ; margin crenated, white, with an inner

border of orange, and another thin one of black.

Under surface paler.

Ova white.

Apparently allied to Hurylepta interrupta, Stimp. (Proc. Ac. Se. Phil. 1857, p. 26).—

Cre:

Fig. 26. Upper surface.

Genus CENTROSTOMUM, Dies.

27. Centrostomum ocellatum, Kel.

Length 2 inches.

Animal gelatinous.

Upper surface pale yellowish brown, with dark brown ocellated spots.

Under surface pale buff.

Tentacles none.

Mouth near the centre.

Hye-spots occipital.

Ova white.

Of this and the next species Dr. Kelaart made a new genus, which he called Penula,

the principal character of which was the absence of any form of tentacles. But this

character is not that of a genus, but of a subtribe, called by Diesing Aceridec ; and the

families Typhloleptide, Leptoplanide, &c. are therein included.—C. C.

Fig. 27. Upper surface.

28. Centrostomum punctatum, Kel.

Length 12 inch.

Upper surface white, shaded and minutely punctated with reddish brown.

Under surface very delicate white, clouded with faint reddish brown.

Fig. 28. Upper surface.

These two species (particularly the first) seem to bear a great resemblance to the Pla-

naria gigas, from the Red Sea, figured in Riippell’s Atlas, t. 3. f. 5.—C. C.

98 , DR. COLLINGWOOD ON MARINE PLANARIANS.

Genus STYLOCHOPLANA, Stim. -

29. Stylochoplana elegans, Kel.

Length 1 inch.

Upper surface pale yellow, shaded with greenish brown, and dotted with black; margin

black, lined with orange.

Under surface whitish, with brownish margin.

Tentacles two, small, reddish, tipped with red, occipital.

Vig. 29. Upper surface.

30. Stylochoplana meleagrina, Kel.

Length 12 inch.

Upper surface sixiped with broad streaks, the inner pure white, the others light purplish

and whitish ; the median line brownish red, edged with black ; the margin waved and

edged narrowly with black.

Tentacles small, oval, occipital. There are also two linear appendages on the occipital

region above the eye-spots.

Fig. 30: a, upper surface ; b, under surface.

Genus PLANOCERA (Blainville).

31. Planocera thesea, Kel.

Length 1} inch.

Upper surface chocolate-brown, edged with yellow.

Under surface pale purple, darker towards the margin.

Tentacles white, tipped with red, rising from depressions or cups placed near the middle

third of the body.

Mouth central.

Fig. 31. Upper surface.

| EprrortaAL ADDENDUM, with EXPLANATION of Puates XVII. to XIX.

The figures im the three Plates are consecutively numbered, and correspond with the numbers and

references appended at the close of each description of the species in the text. The original drawings of

figs. 1 to 12 (Pl. XVII. and part of Pl. XVIII.) were designed by Dr. Cuthbert Collingwood ; those of figs.

13 to 81 (part of Pl. XVIII. and the whole of Pl. XIX.) are by Dr. Kelaart. Owing to the protracted

absence of Dr. Collingwood abroad, the proof sheets have not had the advantage of his revision. This

has unfortunately led to a discrepancy in the lettering in the plates. It was originally intended that the

author’s own newly described species should be distinguished by Arabic, and Dr. Kelaart’s by Roman

numbers; but the artist has not carried out this minor matter of detail. The lettering im the text has

not been altered to correspond with the artistic error in the plates, which were printed aff before the

correction of proof sheets was undertaken.

Trans. Linn. Soc, Ser. 2. Zoon Vou.l'Tas. 17.

C Collingwood delt G. French Angas lith W. West&C° imp

New Species of Planaria.

Trans Linn Soc Szr.2 Zoor Vou | Tas 18.

¢. Collingwood. delt

Geo French Angos lth

New species of Planaria.

W.West & C2 ump

}

Trans. Low.Soc.Ser 2.Zoon Vou. 1. Tas 19.

Dr Feloort deleod not W West & Co ump

Geo French Angas ith.

New species of Planaria.

[22 all

VII. On the Structure and Development of the Bird's Skull. By W. K. PARKER,

THES, LM beSe, Cfo (Ceehen 18)

(Plates XX.-X XVII.)

Read December 16th, 1875.

INTRODUCTORY REMARKS.

IN a paper of mine on the Development of the Bird’s Skull (Phil. Trans. 1869, pp. 755-

_ 807), a single species, the Common Fowl, was used as the subject; this was worked out

to as great an extent as was possible to me at the time.

Year by year since then steady work at other types of the Vertebrata has, by letting

in fresh light, shown me some serious deficiencies in that paper, and my mind has not

been able to rest without an attempt to superadd something of value to the older piece

of research.

I may at once remark that, criticising this my own work, I find that the eye and the

hand had done their work better than the md; the plates are clearer and truer to

nature than the descriptions. :

The Memoir which followed that on the Skull of the Fowl treated of this part in the

Common Frog, a type so diverse from that of the Vertebrata generally as to mislead me

somewhat in my comparisons, and to make me doubt the correctness of my earlier

descriptions. Further researches, however, have set my mind at ease on several points,

notably those on the Salmon’s Skull (Phil. Trans. 1878, pp. 95-145, plates i—viii.).

There was another reason, however, why it was desirable to supplement the paper on

the Skull of the common Gallinaceous bird, namely, that it is itself a very simple semi-

struthious type, and conveys but an imperfect idea of the remarkable metamorphoses the

Reptilian type of skull can undergo when subjected to the intense life-energy of a high-

class perching or climbing bird.

A third reason has acted as a propelling force in this matter: this was a strong desire

to clothe with flesh and form the masterly skeleton paper on the “ Classification of

Birds” given by Professor Huxley to the Zoological Society a few years ago (Proc. Zool.

Soe. 1867, pp. 415-472).

In that paper the palatal structures are chosen as the ground of classification; and

although, to a hasty observer, these might seem to be a mere fragment of the whole bird,

yet they are, indeed, the parts to which all others follow suit—the upper jaw and palate,

the first and part of the second facial arch. To these parts every thing else in the Bird is

correlated ; they rule, as it were, the whole economy of the Bird.

Renewed researches, which have been zealously followed up by me for some years, on

the structure and development of the “ trabeculee cranii,” “ pterygo-palatine arcades,”

and nasal labyrinth, have yielded results that seem to me to be of great value.

SECOND SERIES.—ZOOLOGY, VOL. I. P

100 MR. W. K. PARKER ON THE STRUCTURE

In this newer work the forms of the Fish and the Reptile have been before my eyes

night and day. These have been to me as the larval and pupal stages through which

may be traced the changing materials out of which that vertebrate imago, the Bird,

is constructed, its hot blood infusing new life and producing unlooked-for trans-

formations. ;

Now that cranial morphology is beginning to embody itself in a clear and intel-

ligible shape, it may be well to explain the meaning of certain terms that will of neces-

sity be in frequent use.

The term metamorphosis I use in exactly the same manner and for the same purposes

as the entomologist, namely, as describing the whole series of changes undergone by the

skull or skeleton throughout the life-history of the individual.

The borrowed term-isomorphic is used, in a zoological sense, for very similar types in

quite distinct groups: the Sun-birds (‘‘ Nectariniidee””) may be said to be the isomorphs

of the Humming-birds (“ Trochilidee ”’).

The term symmorphic may be used for any morphological part or parts in one type

which agree in their structure and development with those of some other types; thus

the “ Desmognathous” palate of the Parrot and that of the Goose are symmorphic; such

agreeing structures are also said to be homologous or representative.

Professor Huxley, in his paper (op. cit. p. 454), places the Goatsuckers and Humming-

birds with the Passerine types, supposing them to have the true “ Mgithognathous ”

palate. This is an error: they are Insessorial Schizognaths.

The Woodpeckers (“ Picidee,”’ p. 448) had this service done to them, namely, their vomer

was shown to be double, and their maxillaries to have a very rudimentary palatine plate.

Thus they could not be treated of as belonging to the Cuckoo tribe (“‘ Coceygomorphee”’).

The palatal structures (trabecule, pterygo-palatines, and nasal capsules) of the Wood-

peckers show that these birds can be classified with none of the groups established by

Professor Huxley, as their face is intensely Lacertian and even Ophidian. I have there-

fore called this group of birds the “ Saurognathe” (Trans. Linn. Soe., Zool. pt. 1,

1875, pp. 1-22, Plates I.—V.).

That peculiarity of the Bird’s face which has been termed “ Aigithognathous ” will be

explained. It exists in all the “ Coracomorphe”’ and, besides these, in the Swifts, or

“‘ Cypselomorphe ”’ (op. cit. p. 468).

The importance of the fact that a low, superstruthious bird like the Hemipod should

offer itself as a stock (“phylum”) to the half myriad of known Passerines cannot be

easily overrated *.

My principal reason for describing the types of skull presented here is not zoological

but morphological. I wish, in some degree, to fill up that which remains over of cranial

* On the distribution of “ Alectoromorphe,” see Huxley, Proc. Zool. Soc. 1868, pp. 294-319. ‘The Tinamous

stand with one foot in that group and with the other over the struthious border, and the larger “ Cracide” (Craa

globicera) and “* Megapodide ” (Megacephalon maleo) are “ Desmognathous” (see Huxley “‘ On Classification,” p. 423,

for a figure of the former). Thus the Fowl tribe sweeps over a huge zoological space, immediately overlying the

stratum occupied by the “ Ratite.” Amongst the ancient extinct forms of this group we must look in imagination

for the forefathers of the Birds of Prey, the Climbers, and the Perchers.

AND DEVELOPMENT OF THE BIRD’S SKULL. 101

morphology after that of the Fowl has been exhausted, so that the student’s conception

of the Bird’s skull may be richer and more in accordance with the fulness of Nature.

I shall take the present opportunity to give figures of skulls that contain elements

frequently spoken of in other papers by me, but not illustrated, such as the “ inter-

palatines’’ and “ mesopterygoids.” I shall also have to add other pieces, with their

names, namely, the “ medio-palatine”’ and the “ palato-maxillary.”’

Processes that grow in a secondary manner out of the simple facial bars, for union of

bar with bar, serially, are called here “‘ conjugational processes ;” the union of the right

and left bars of one arch at the mid line is called a “‘ commissure.”

Further Researches on the Chick's Skull.

My first stage (Phil. Trans. 1869, plate Ixxxi. figs. 1 & 2) is rather wnderdated as the

fourth day ; those given are of the fifth, and those soon to be described run nearly to the

sixth day. The second stage (figs. 3-13) should be put as dating from the sixth and

seventh days of incubation.

One of the first points to be determined in the morphology of the skull is the structure

of the “ investing mass” of the notochord and the “ rafters of the cranium” (“ tra-

beculze cranii”’), the first pair of facial bars.

The figures given in my former paper show the distinctness of these parts much more

definitely than I supposed at the time. These I now proceed to explam. At page 758

I speak of the ends of the investing mass as forming the cartilaginous “ lingul sphenoi-

dales,” thus confusing the free truncated end of the investing mass (plate Ixxxi.

fig. 2, 1.g), which looks forwards and outwards, with the free tops of the trabecular bars,

which are better seen in fig. 8, and in plate Ixxxii. figs. 1 & 3, l.g.

The explanation of these parts was made by the exquisitely lucid structures of the

embryo Salmon ; and now, if these figures of the Fowl be compared with those, it will be

seen that the trabecular apices are at first embraced by the squared out-turned ends of

the investing mass, and that in the next and third stages the curved trabecular tips,

where they surround the pituitary body, open out like the blades of a pair of callipers,

and growing more distinct and solid, form, in the third stage (plate Ixxxii. figs. 1,3, & 4,

lg), the free tongue-like cartilages, the meaning of which I for a long while failed

to see.

The first figure in the present paper (Plate XX. fig. 1, é, pm) ciears up another very

different point, namely, the formation of the “ prenasal rostrum,” or azygous con-

tinuation of the distal part of the trabecular arch.

In my first paper, on the “ Skull in the Ostrich tribe” (Phil. Trans. 1866, plate vii. p.x,

p- 122), the prenasal cartilage is described as being formed by confluence of the “ tra-

beculz ;” but in the second paper (p. 759) [have corrected this statement, but not without

falling into another error, namely, that of supposing the “ale nasi’’ to be formed directly

out of the “cornua trabecule.’” The anomalous position of the parts is caused by the

“© mesocephalic flexure,” the trabecule being positively hooked backwards.

The ‘alinasal, aliseptal, and the aliethmoidal cartilages are regions of membrane

which chondrify later than, but not separately from, the trabecule, the common

102 MR. W. K. PARKER ON THE STRUCTURE

descending nasal septum becoming solid at its sides continuously with the crest that

grows upwards from each trabecula. The broadest part of the “trabecular commissure ”

(Fowl’s Skull, plate Ixxxi. figs. 1 & 2, tr) develops into two important structures,

namely, the subnasal floor (plate Ixxxii. fig. 11, s.v./) and the conjugational process,

which corresponds with the “ ethmo-palatine,”’ or orbital plate of the palatine. The sub-

nasal or “ supervomerine”’ lamine are of small extent and temporary in the Fowl

(plate Ixxxii. fig. 11, s.v./); and the conjugational process does not develop into a

distant structure, but remains as part of the antorbital wall, its infero-external angle

(plate Ixxxiil. fig. 5, p.p). In many birds, however, it hecomes separately ossified as the

“os uncinatum” of Magnus, and takes its place as a kind of transverse bone in the

“« Musophagide ” and other “‘ Coccygomorphee ”*.

In the first of my new illustrations of the Chick’s Skull (Plate XX. fig. 1, tr, pn),

the down-bent skull of a chick, intermediate between the first and second stages of my

former paper, has been sliced off in an obliquely transverse manner, cutting away the

fore part of the projecting optic lobes (C’) and thalamencephalon (C’’), and removing

most of the rudimentary hemispheres (C”). Underpropping the primary vesicle (C’’)

are two thick cartilages, which are severed at their commissure ; these are the trabeculez

at their bend backwards. From this position they send downwards an azygous sharp

keel. This is the nasal cartilage; it looks backwards.

In the next stage (“ Fowl’s Skull,” plate Ixxxi. fig. 3, p.m) it is still retral, but has

become thick and rounded.

In another section (fig. 2), taken at a different angle to the axis of the head and seen

from its front face and not from behind, as in the last figure, the trabecular knuckle

has been cut off, and with it the prenasal cartilage. Here the under-placed nasal sacs

(ns) are exposed, and the hooked trabeculze are cut through so as to expose them in

section, both in the meso-ethmoidal region above, and under the outer nostrils below.

In this section the fore part of the eyeballs (e) and the hemispheres (C”) are cut

through ; the floor of the nose shows cartilage in it only where the cornua trabeculz are

cut through; the prenasal has been pared off. In this rudimentary nasal labyrinth

rudiments of the superior and inferior turbinals (wf, i¢b) are seen, and the membrane

infolding them is the early condition of the aliethmoidal and aliseptal cartilages. The

eyeball is seen to rest on a very neatly formed ledge; this is the jugo-maxillary eleva-

tion; and on the side of this is a lesser ridge, which contains the second preoral bar,

the palato-pterygoid.

This palatine bar (Pa) is composed at present of indifferent tissue; it is triangular in

section, and the ascending crest is its ethmo-palatine region ; the conjugational process

is correlated to that of the trabeculz.

Tn a somewhat more advanced skull, sixth day (figs. 3 & 4), the eyeballs have been cut

through at their middle and the hemispheres at their hindermost part. This belongs toa

* Sce Dr. Reinhardt’s paper, “Om en hidtil ukjendt Knogle i Hovedskallen hos Turakoerne.” Copenhagen, 1871.

In my paper on the “ Ostrich’s Skull” (p. 158), I referred the “ os uncinatum” to the ethmoidal category ; but no

explanation of its true meaning was had until I saw the secondary bud growing from the outside of each trabecular

bar in the Salmon. :

AND DEVELOPMENT OF THE BIRD’S SKULL. 103

head that had begun to straighten, and nearly comes up to the earliest part of my second

stage (“ Fowl’s Skull,” plate lxxxi. fig. 3).

The interorbital septum (pe) has here been cut through in the anterior sphenoidal

region, and, as in the Mammal, the sphenoidal sinuses (sps) are exposed ; they have not,

however, any retral continuation of the nasal wall in a cartilaginous state, and they are

very narrow. Below, thestrongly compressed trabeculee (¢r) have coalesced, and are now

composed of hyaline cartilage; although the commissure is finished below, the crests of

the trabeculee are distinct. These solidify more tardily ; and their conversion into true

cartilage taking place at the same time as the cranial floor with its double descending

keel, there is at present no line of demarcation between these regions.

Afterward (see “ Fowl’s Skull,” plate Ixxxii. figs. 4, 5, & 11, p.s, 0.s) the mem-

branous interorbital space (é.0.s) redivides these dissimilar tracts, leaving for some

extent a descending presphenoidal keel and an ascending trabecular crest.

The orbito-sphenoids (figs. 3 & 4, os) are small, and lessen rather than increase in size.

The palatines are here narrowing towards the pterygoid region.

The next section (fig. 5) is through the optic lobes (C’), the pituitary body (py), and

the back part of the eyeball.

The manner in which the face is tucked under the head at this stage makes a most

remarkable display of parts possible in one section: these are the apices of the tra-

becular (¢7) and pterygo-palatine arches ( pg), and at the same time the distal extremities

of the first and second postorals (mk, ch, b.hy). The internal carotid arteries (éc) are

seen piercing the fibrous pituitary floor mesiad of the ends of the ‘trabeculz, the

actual apices of which look downwards in the figure. On each side of the high and

narrow mouth-roof (m) are seen nearly the whole of the pterygoids (pg); their “ trabecular

process,” or epipterygoid apex, looks a little inwards below, and is seen enclosed in the

rudiment of a “ soft palate.”

The end of each “ sphenoidal sinus” (sps) is here seen to reach to the side of the sella

turcica. The floor of the mouth (m) contains laterally the Meckelian rods (mk), supra-

mesially the tips of the small distal cerato-hyals (ch), and below the basi-hyal (b.hy).

Behind the optic lobes and eyeballs the head has become very narrow. The next section

(fig. 6), which has been made through the fore part of the auditory capsules, shows

this.

The notochord (ve) lies between the two halves of the investing mass (iv), which has

completely coalesced with the periotic cartilage. This section shows the cochlear cavities

(cl) and those of the anterior semicircular canals (asc) with their ampulle. Below the

cochlee are seen the internal carotids (ic), and towards the mid line the submucous

tissue is of great thickness; it forms the nidus in which those remarkable ornithic

detachments of the parasphenoid, the basitemporals (0¢), are developed. All but the

fore ends of the mandibular rods are here shown, and also their piers, the quadrates ;

mesiad of these are seen the basi- and cerato-hyals (bhy, chy).

With regard to the parasphenoid, although it is developed as an azygous bone ante-

riorly, yet the thick bed of stroma which forms its mother-substance is a symmetrical

104 MR. W. K. PARKER ON THE STRUCTURE

growth from the two sides (see fig. 4), when these masses are approximating below the

coalesced trabeculz.

Before dismissing the chick’s skull I have to speak of the posterior conjugational pro-

cesses connecting the trabeculze with the next facial bar, the pterygo-palatine.

These parts, the basipterygoids with their articular plate and the corresponding

structures in the pterygoids, are thoroughly worked out in my paper on the “ Fowl’s

Skull” (plate Ixxxiii. p. 781); but the general morphology, their symmorphism with the

anterior conjugationals of the same arch, was not clearly seen.

Before closing this supplemental description of the fowl’s skull, I may remark that

the fenestra which severs the perpendicular ethmoid from the nasal septum (‘“ Fowl’s

Skull,” plate Ixxxiii. fig. 4, ¢fic) is the redifferentiation of the trabecule from their

surroundings.

In many birds another fenestra, besides the “hinge” and the interorbital space, is

found in the nasal septum; and in the Goose tribe a fourth appears between the upper

turbinals; to this is added, in Pelecanoides urinatriz, or Petrel, a fifth large fenestra,

close behind the antorbitals.

In the adult Swift (Cypselus apus) there are fowr membranous fenestre in the inter-

orbital space, instead of two as in Pelecanoides, and one as in Birds generally. This

gradual fretting-away of growth-tracts which at one stage of metamorphosis obscure

essentially distinct parts is of extreme interest; the highest and, as it were, the #mago

forms show this most, notwithstanding that they use up and mix together in many ways

parts that remain thoroughly distinct in types that may be compared to the pupe and

larve. As the two preoral arches are intimately interblended, not only with each other

but also with the nasal labyrinth and the cranium, it may be well to set down fairly what

splints or secondary bones become applied to or grafted on these arches.

To the hinder pair, the “ palato-pterygoids,”’ secondary developments of the mandible

or first postoral, there belong the maxillaries, palato-maxillaries, interpalatines, jugals,

and quadrato-jugals.

The trabecular arch draws to itself, as subcutaneous or submucous bones, the follow-

ing—namely, the premaxillaries, septo-maxillaries, vomer or vomers, and the para-

sphenoid ; the “ os uncinatum”’ is an endosteal part.

The splints of the mandibular arch were described in my former paper. The tympanic

chain of the Bird is a masked splint-series to the segmented and metamorphosed apex of

the hyoid arch.

On the Structure and Development of the Facial Arches in the Passerine types,

“‘ Coracomorphee ’’ (Huxley).

In the paper already referred to, Professor Huxley has sought to group the flying

birds in accordance with the spirit of Humboldt’s plant-groups in his ‘ Views of Nature.’

The group under notice, then, is his “ Coracomorphe,” or Crow form (p. 469); this is

so large that (as my friend Mr. Osbert Salvin informs me) whilst the birds already known

and described are about ten thousand, half, or well nigh half, of them belong to this one

group. Now the type the skull of which I have already worked out, namely that of

AND DEVELOPMENT OF THE BIRD’S SKULL. 105

the Fowl, stands as the head of another group, the “ Alectoromorphe;” and this

represents also a broad and goodly family of forms. In another paper, ‘“‘ On the Classi-

fication and Distribution of the Alectoromorphze”’ (Proc. Zocl. Soc. 1868, pp. 294-319),

our author has broken this fine family into fragments, separating (see pp. 302-304)

not only the Pigeons (‘ Peristeromorphe”’), which are “ Altrices’’ and have tender

young, but the Sand-Grouse also, as the ‘“‘ Pteroclomorphe,”’ and the Hemipods as “ Tur-

nicimorphe.” As they stand at present, no cutting and contriving will make the

lesser groups, such as the ‘“ Charadriomorphe,”’ ‘‘ Pteroclomorphe,”’ and the like,

correspond to or in anywise be the equivalents of the great army comprised under the

Crow form. Much as I value my friend’s paper, I do but consider it in the light of a

sign-post to guide others who are footing it on the same road, and not as a finished work

to rest in.

In one of the most outlying of the Crow forms, the House-Martin (Chelidon urbica),

I have been able to demonstrate the second morphological stage of the second postoral

arch. I did not succeed in this point in the Fowl; but now that it is done it will do

duty for the whole of the “ Sauropsida.”

The metamorphosis of this, as of the other arches, has been fully worked out in the

“ Tchthyopsida,’” in my researches on the skull of the Frog and the Salmon; but

neither of these throw any light upon the long-stalked, trifoliate stapes of the Reptile

and the Bird. Professor Huxley, in his paper ‘‘ On the Representatives of the Malleus

and Incus” (Proc. Zool. Soc. 1869, pp. 391-407), says (p. 398), “the suprastapedial

cartilage”’ [he is describing the parts in Sphenodon] “turns out to be nothing more

than the proximal end of the hyoidean arch, while the stapes and its appendages are

exclusively related to this arch, and have nothing whatever to do with the mandibular

arch.”

True! and yet he does not say how much of the stapes belongs to the auditory capsule.

In the Urodela the stapes is cut out of that capsule, like a dung, and in Batrachia the

head of the upper segment of the hyoid arch articulates with the auditory stapes, its

uppermost part being segmented off as a “ pars orbicularis.” In the Mammalia the

Batrachia are closely followed (Phil. Trans. 1874, plates xxvi—xxxvii.). In my former

paper on the Fowl’s head (plate Ixxxi. figs. 5, 9, & 10, st) I showed, in the second stage,

a thick somewhat flattened club of cartilage, the knobbed end of which fitted into the

fenestra ovalis, whilst the other or distal incurved end was trifoliate, the upper process

being the “ suprastapedial ”’ rudiment, the lower the “infrastapedial,’”’ and the longer,

bowed leaf the “‘ extrastapedial.”

In a young Have-Swallow, at about the same astel, the upper end was seen to be

continuous with the periotic capsule im front of its fusion with the exoccipital cartilage,

and above the outer end of the rudimentary cochlea. This stapedial bar (Plate XX.

fig. 7, mst) has the characteristic curve of the summit of the facial arches, and at this

stage has coalesced with the auditory capsule as is the wont of these bars (see the

quadrate in continuity with the periotic cartilage, as figured in Huxley’s ‘ Elements,’

p- 138, fig. 57, F* Qu). This is undoubtedly the second or coalescing stage of the

second postoral; it has, however, only coalesced by the strong posterior head; the

106 MR. W. K. PARKER ON THE STRUCTURE

aborted anterior head, comparable to the “ tubercle” of a rib, and answering to the

short crus of the incus, has been left free. The anterior head, or “‘ suprastapedial” (s.s¢),

is, however, continuous permanently with the auditory capsule in Sphenodon (Huxley,

«On Malleus and Incus,” p. 397, fig. 4, S.S¢), and also, as I find, in the Bull-frog (Rana

pipiens). Notwithstanding that in Sphenodon the posterior head is free, and the anterior

is fixed, contrary to what is here shown in the Chelidon, yet in these two, the adult

Lizard and the embryo Bird, the “ extrastapedial”’ (Plate XX. fig. 7, est, and Huxley

op. cit. fig. 4, H.St) has a similar shape; even the foramen seen in Sphenodon (a) soon

appears in the Bird. The “ infrastapedial” (Plate XX. fig. 7, ist) of this young bird

is longer than I found it in the Fowl, even after the oval tract of ear-wall had come

away with its “ mediostapedial”’ stalk. This is not all; for its pointed apex is con-

tinuous with a band of cells of indifferent tissue which runs downwards towards the

entering carotid artery at the skull-base.

In Sphenodon (op. cit. fig. 4, Sth) the scooped “ extrastapedial ” passes directly down-

wards into a bar which is labelled Sth (stylo-hyal); but in the Crocodile (op. cit.

figs. 1 & 2, Sth) the stylo-hyal is a short club of cartilage, flattish, with its broad end

upwards, looking backwards, and having a clear membranous space between it and the

“ extrastapedial.”

The stylo-hyal of Chelidon answers to this description exactly ; and if the reader will

turn to Professor Huxley’s figure in this part of the Fowl (op. cié. fig.5, A, B, I.St) he

will see that it is there, but that it has become confluent with the infrastapedial. In

figs. 5,9, & 10, st, of my former paper, this part is not shown; but in a chick one day

older (fig. 14, i.s¢) it is certainly figured, but not distinct. Not having seen it in the

earlier stage, I could not choose but find it in the next, as it had then coalesced with

the “ infrastapedial.’”” In certain birds this part ossifies; and this fact led me to call it

the “stylo-hyal” *. The distal part only of the rest of the arch is chondrified, namely,

the small cerato-hyal (chy), which, with its fellow, lies in the substance of the tongue.

The basal element, “ basi-urohyal” (bhy, 667), is a stout wedge, which narrows, and

then is flat and emarginate behind. The so-called “thyro-hyals”” (= 1st branchials, br’)

are very large, the upper or proximal piece embracing the occiput. In this illustra-

tion the lingual skeleton is drawn as flattened out in the minute preparation, so as to

display all the parts at once.

I take this opportunity further to illustrate Professor Huxley’s account of these parts

in his invaluable paper (op. cit. p. 398), and first give the condition of these parts in a

fledgling Crow (Corvus corone). The membrana tympani (fig. 8, mfy) is shown from

the inner side, with the stapedial and mandibular structures; the articular end of the

lower jaw is shown slightly dislocated, and the ‘“ siphonium”’ is cut through.

This tube, the siphoniwm of Nitzsch, conveys air, not from the quadrate to the lower

jaw (see Huxley, op. cit. p. 391), as in the Crocodile, but it directly opens into the

tympanic cavity in the Birdt.

* Full often have I argued with the author of the paper just quoted that this dilated, decurved, sometimes bony

extremity of the bird’s “ infrastapedial” was the true “ stylo-hyal;” but he always turned his deaf ear towards

me. Jam silent now; but “ he will rise up at the voice of the bird,” so sharp and clear.

+ The dissection here figured was made to dispel a doubt of Professor Huxley’s as to the correctness of Nitzsch’s

AND DEVELOPMENT OF THE BIRD’S SKULL. 107

In the adult of this species, the Carrion-Crow, the siphonium is enringed by the

lowest and largest of the seven bones that form the curious tympanic chain. Figures of

these have been given in my paper on the “ Coracomorphe ”’ in the ‘ Transactions’ of the

Zoological Society.

The stapedial plate (st), turned a little forward for display, is continuous, as bone,

with the mediostapedial (mst), and then come the outspread leaflets of cartilage. The

* extrastapedial” (est) is sickle-shaped and broad-backed, its thick convex back being

strongly connected with the inner face of the membrani tympani (mty); and the fibres

of the perichondrial coat are interwoven with those of the drum-skin itself, and also meet,

from without, those of the tendon of the “ tensor tympani” muscle (stm).

The tongue-shaped rudiment of the suprastapedial (sst) lies in the base of a two-

winged membrane, the fibrous supplement of this part.

The cartilaginous strap running downwards from the extrastapedial to the entering

carotid (ic) is the infrastapedial (ist) and stylo-hyal (sth) in one band. The fore end of

the mandible (ms, d) shows that, as in the Fish, the mandibular rods do not meet to

form a “mentum” in these prognathous types; they keep distinct, and each receives

its own dentary.

The form and disposition of these parts in the adult are shown in fig. 9. They are

those of the Daw (Corvus monedula); the references are the same as in fig. 8.

In the Crows of the southern world there are many important differences of structure

from what is seen in the northern species: notably, the condition of the stylo-hyal is

instructive. In the Piping Crow (Gymnorhina tibicen) the general structure corresponds

very exactly with that of Corvus; but the stylo-hyal (figs. 10 & 11, sth) ends below in a

cheese-knife-shaped plate, which lies upon the outer edge of the basitemporal, and is

well ossified. Slender as in the mediostapedial shaft (mst), it is pneumatic and com-

municates with the tympanic cavity by both proximal and distal passages.

Being desirous that this, together with the paper on the Fow/’s Skull, should enable

the student to form an accurate conception of the skull of the Carimate Bird, as

distinguished not only from that of other classes, but also from that of the “ Ratite,’” or

Struthious order, I now proceed to show some of the most important modifications of the

ornithic face. Happily for me, I have at hand a strong and sharply expressed outline of

this subject ; and now it will be easy to make a more extended study and comparison of the

facial morphology of the higher birds (see Huxley’s “ Classification of Birds,’’ Proc. Zool.

Soc. 1867, pp. 415-472).

My former papers illustrate the “ Deomaeeinttous” face as seen in the raft-breasted

Ostrich (Huxley, op. cit. p. 425) and the keel-breasted Tinamou (Phil. Trans. 1866, plates

ix.—xv. pp. 118-183) ; the “ Schizognathous”’ face (Huxley, op. cit. p. 426), as seen in the

Common Fowl (Phil. Trans. 1869, plates Ixxxi—Ixxxvii. pp. 755-837); and the “ Aigi-

thognathee ” have been illustrated in the Zoological Transactions for 1875 *.

T return now to the Sparrow-faced birds, “ Mgithognathee”’ (see Huxley, op. cit. p. 450).

description, as that made the Bird differ from the Crocodile in the disposition of the pneumatic duct. It does differ,

as I have shown above.

* See Part I., in vol. ix., plates 654-62. Part II. will soon appear.

SECOND SERIES.—ZOOLOGY, VOL. I. Q

108 MR. W. K. PARKER ON THE STRUCTURE

The development of the preoral arches and nasal capsule in the “ Fringillide ” (which

in the person of the Sparrow gives name to the group) will show the peculiarities of this

type, a type which takes in all Professor Huxley’s “ Coracomorphz” (op. cit. p. 469),

the Swifts amongst his “ Cypselomorphe ” (p. 468), and the Hemipods also, which lie

down lowly, in an ornithic stratwm immediately over the “ Tinamide”’ *.

In embryos of the Brown Linnet (Linota cannabina), at about the end of the fifth

day of incubation, I find the facial arches and nasal capsules (Plate XXI. figs. 1 & 2)

in a very instructive condition. Nearly the whole of the long “ trabecular commissure ”

is shown, the bar being severed in the prepituitary region, where the ethmoid and basi-

sphenoid meet; the prenasal rostrum (pz) is in full size, as the continuation of the

great cartilaginous balk.

This balk is wasting between the ethmoidal and septal regions, preparatory to its

peculiar ornithic segmentation.

The open or naturally cloven state of the palate is here shown; for the folds of the

nasal labyrinth are well seen below, and the “ median nares” (mm) are large and

patulous. Neither the true olfactory nor the inferior turbinal regions are displayed in

this figure, only the immensely developed alinasal structures, so very secondary in

importance and size in the Mammal. The trabecule being bowed and folded over in

front send backwards, on each side, a flap of cartilage; this may be called the “ recur-

rent fold” (rec). These two flaps form a commissure below the fore end of the

septum nasi; and this becoming ossified from the alee, sometimes appears as an anterior

nasal bridge, a desmognathic band close behind the fore beak (e. g. in Dicholophus).

The posterior of the three folds, the inturned lamina (éa/), seen in the preparation, is

the swelling alinasal wall passing into the floor of the nasal vestibule. Here it is far in

front of the maxillo-palatine hook (map); but after a short while it will lie on this bony

bar, and become intimately connected with it by the binding fibres belonging to each

structure. Between the plaits already described is seen a middle cartilage; this is an

outgrowth of the wall or posterior plate. It is the alinasal turbinal, and ends in a

double free edge.

Although the premaxillaries (px) and maxillaries (mx) are already well developed, yet

one does not see here the broad forked vomer of this type. Looking, however, carefully

between the wasting middle bar and the inturned alinasal wall (¢a/), there is to be seen

at this stage a small oval cartilage; as this appears to me to be as yet undescribed, I

propose to call it the ‘‘ vomerine cartilage” (vc). It ossifies endosteally, and has even

now a small core of osteoblasts with the finest deposit of calcareous matter. One of these

cartilages is more highly magnified in fig. 2, and also with it part of the nasal wall, ali-

nasal turbinal, and maxillo-palatine.

This cartilage is small here in the Linnet, but very large, as I shall soon show, in

other “ Hgithognathe ;” and as it forms the mother patch for each vomerine moiety, it

* In the paper under notice (p. 459) Professor Huxley placed the Hemipods among the Fowls, ‘“ Alectoro-

morphe ;” but in another equally important contribution to Ornithology, namely, his “ Classification and Distribution

of the Alectoromorphe ” (Proc. Zool. Soc. 1868, pp. 294-319), he places the Hemipods apart, haying gained a clearer

insight into their peculiar characters (see p. 804). They are now his “ Turnicimorphe.”

AND DEVELOPMENT OF THE BIRD’S SKULL. 109

is of great interest, and isnew to me. If any morphologist knows what it answers to in

the lower vertebrate types it were well that he should explain it *.

As the metamorphosis of these parts is very rapid, it is necessary to examine embryos

not more than a day or so in advance of or behind each other. Inno longer a period, what

is displayed in fig. 1 is exchanged for what I have shown in fig.3. Here the small osseo-

cartilaginous nuclei have developed into the large ox-faced vomer, with its broad anterior

part grafted on each side upon the alinasal wall, which turns directly inwards, and

spreads in a pedate form to apply itself to the vomer. On one side of the V-shaped

emargination the vomerine piece is fenestrate; the bony matter here becomes detached,

and an additional bone formed. ‘This additional bone is seen in the fledgling (see it in

the young Sparrow, fig. 4, smx). Not only does the vomer grow into the nasal wall, but

the alinasal turbinal (fig. 3, atb) also; hence the huge size of the upper vomerine groove,

which is so striking in the Crow tribe (“ Corvide” proper); for the thick common end

of the wall and the turbinal approach the septum nasi, and, rising high on each side,

embrace the base of the septum. Another noteworthy characteristic of the Passerine

face is the elbowed form of the palatines near their hinder third; this out-bent part does

not ossify ‘‘ by first intention,” but lingers until it is solid hyaline cartilage, and then

receives its own endosteal centre. This new bone is an arrested “ transverse’ element,

my “ transpalatine”’ (¢pa).

That part of the Passerine palatine which is arrested from forming a bony palatal floor

generally exists as a free fore-looking spine. This is well seen in the Sparrow (fig. 4,

ipa); it is the “ interpalatine process.” In Ducks and Swans this is a separate knuckle

of bone; and in looking over a large number of Starlings’ skulls, I find it in certain of

them (see Plate XX. fig. 12, ipa).

Other parts of the Passerine face must be noticed.; but I turn now to the most striking

condition of the vomer I have hitherto found in the group: it is seen in the native

Wren (Troglodytes vulgaris). When the whole palate of the bird is viewed (Plate X XI.

fig. 5, v), the vomer is seen to be as large, relatively, as in the “ Ratite;”’ but its

anterior third is composed of true cartilage. In a spirit-specimen, by careful dissection,

I have been able to make out what is shown in fig. 6 on a larger scale. Here one of the

palatines (pa) and both of the maxillo-palatine hooks are removed; they are, however,

indicated by a dotted outline.

* Finding the development of the Passerine “vomer” so totally unlike what I always supposed, namely, that it is

an endosteal ossification of cartilage, my mind cast about for some symmorph in the cold-blooded types and in the

Reptilian birds (Ratite). I had not far to seek; for lying unexplained in my memory were certain structures in the

Snakes and also in the struthious Rhea, which I had figured years ago (Phil. Trans. 1866, plate x. fig. 14, the alate

cartilages above v), but which had hitherto resisted all efforts at explanation. On reexamination of these parts in the

Rhea chick, 1 find that they are free cartilages in the mazillo-palatine region, and attached to those processes and to

the fore forks of the double vomer by fibrous tissue. In the Snake the nasal glands and their bony capsules, the

vomers and septo-mawillaries (P), are flanked on their outside by an arcuato-spatulate flap of cartilage, continuous

by a narrow stalk with the recurrent lamine, or separate from that process, both states occurring in the same skull.

As in the furculum of the same bird (Linota, see “‘ Shoulder-girdle and Sternum,” plate xv. figs. 12-15), the Linnet’s

““yomer ” is a morphological compound, and the counterparts of its component elements are persistently separate in

the cold-blooded types.

110 MR. W. K. PARKER ON THE STRUCTURE

Part of the dialate septal base (¢r) and part of the nasal labyrinth of one side is here

shown. We here see that the trabeculze have only partially lost their flatness, and form

a, partial subnasal floor, and that the alinasal wall (mv), after giving off the “ alinasal”

turbinal (at), ends externally in a blunt process, and internally in an incurved bifurcate

rod (ial).

The forks of this rod are continuous with the corresponding vomerine half ; the halves

of the vomer are joined by a long commissure into a flat bone, notched at both ends; the

long posterior ends articulate with the ethmo-palatines, and the shorter anterior forks pass

into a cartilage shaped like a breast-plate. ‘This plate, being gently emarginate behind,

forms a fenestra with the help of the bony vomer; it narrows gently forwards, and ends

in a pair of straight styles.

This plate is nothing more or less than a forward continuation of the vomerine

cartilages (vc), the unossified parts being bridged over by a commissure. For relative

size, these vomerine cartilages * have their rivals only in the Snake, the Rhea, the

Hemipod, and in certain low-typed Passerines of the southern world (Notogea), namely,

Aneretes parulus, Pipra auricapilla, and in that remarkable type Pachyrhamphus.

Other characters in the Wren’s face are of importance, as illustrating what is com-

monly found in the numerous related forms.

The lacrymal (fig. 7, 7), often absent, is here very small; the lateral ethmoid (e¢h, pe),

on the other hand, is very large, and shows itself on the upper view ; its inferior angle is

separately ossified as an “‘ os uncinatum”’ (oz), or palato-trabecular conjugational bone.

Between and below the eyes the trabecular crested bar (¢) has been freely differentiated

from the ethmo-presphenoidal bar (pe), and the long overlapping process of the pterygoid

has become segmented into a distinct “‘ mesopterygoid ” (ms.pg), soon, however, to

ankylose with the palatine. In this rather young bird the transpalatine (fig. 5, tpa) was

scarcely confluent with the broad part of the long and slender palatine (pa). The deli-

cate pterygoids (fig. 5, pg) here are more like the epipterygoid of the Lizard than its ptery-

goid ; but the terminal part (apex of the bar) is alone upturned. This upturned part is

very long in some “ Coracomorphe ”—e. g. Coccothraustes, one of the most specialized of

the conirostral division of the group. In certain subfamilies of the “ Coracomorphe,”

although the maxillo-palatine processes do not solder the upper jaws together, yet the

palate is made more solid by means of an additional bony wedge, the “ palato-maxillary ;”

this grows in between the prepalatine bar and the body of the maxillary. In the “Car-

dinalide”’ (Plate XX. fig. 18, pa.mx) it is a large and thick bone; it is somewhat

smaller in the Buntings (Linberiza, Plectrophanes, Phrygilus), and also somewhat smaller

still in the “ Ieteridze,” “ Sylvicolidee”” (Mniotilta, Dendreca, Trichas, Chlorophanes),

and in the “ Tanagridee”’ (Zanagra, Prionocheilus, Stephanophorus).

That curious bone, the ‘‘ os uncinatum,” the matrix of which was first developed as a

conjugational bud from the trabecula, is very large in the Grosbeak ( Coccothraustes), as

in its climbing isomorphs the Parrots. The “posterior conjugational processes,” “ basi-

pterygoids,” and their counterparts on the pterygoid bones are early arrested in the

* Tf the Wren had possessed a “ recurrent alinasal,” like the long forked one of Vireosylvia olivacea, the two

subseptal floors would have reached each other.

AND DEVELOPMENT OF THE BIRD’S SKULL. aal

*« Coracomorphe ;” spiny rudiments, however, are not seldom seen, especially in southern

forms, as, for instance, Gymnorhina, Homorus, Dendrocolaptes, but most clearly of all

in a form. allied to the Sun-birds, namely Anthreptes malaccensis.

I have purposely mentioned these modifications of the face in the very numerous types

of the Crow form, and if it had been possible I should have sought to make the “ Agi-

thognathe”’ run parallel with the ‘“ Coracomorphe ;” but Nature herself forbids this.

Yet it is some gain to have been able, as Professor Huxley has done, to take in a lump

nearly half of the myriad of known birds, to show that all these are ‘ Aigithognathous,”

and to callthem by the name of the commonest and yet one of the highest of the group,

namely, the Crow. Here the zoological group cannot be made to hold all the mor-

phologically agreeing types, the ““symmorphs.”’ This will be felt still more strongly in

the next group to be considered, the “‘ Coccygomorphe”’ (Huxley, op. cit. p. 466),

which, with an extension of its boundaries to be proposed here, will contain “ Aigi-

thognathe,” “ Schizognathee,”’ and “ Desmognathee.”’

To make the matter a little clearer, it is necessary to show that there are serena

varieties or degrees in the morphological types of the ornithic palate, as designated by

Professor Huxley. In the “ Agithognathous” palate I find three varieties, namely :—

a. Incomplete.

6. Complete: var. 1.

¢. Complete: var. 2.

d. Compound.

a. Incomplete “ Megithognathism” occurs in the “Turnicimorphs,” Hemipodius or

Turniz. Here the vomerine cartilages are very large and incompletely ossified, and the

broad double vomer has a septo-maxillary at each angle; but these bones are only

strongly tied to the “ alinasal” cartilage, and do not graft themselves upon it.

b. Complete: var. 1. This occurs in some of the lowest harsh-voiced ‘ Coraco-

morphe.’ The vomers are developed in large “ vomerine cartilages,” which they often

only partially ossify ; but these osseous tracts are distinct from those of the often bony

alinasal walls and turbinals: a small septo-maxillary on each side generally appears,

limpet-like, on the angle of the alinasal inturned cartilage, but does not run into it.

This is well seen in Pachyrhamphus, Pipra, and Thamnophilus.

c. Complete: var. 2. This occurs in an-immense group, comprising most of the

“Coracomorphe”’ and also the Swifts (“ Cypselidee’’); here the vomerine bones are

grafted upon the nasal wall, and thus the bird loses its primary schizognathism,

d. Compound. 'This type occurs where, in a perfectly egithognathous face, desmo-

gnathism is produced by ankylosis of the inner edge of the maxillaries with a highly

ossified alinasal wall and nasal septum.—EHxamples, Gymnorhina tibicen, Paradisea

papuana, Artamus leucorhinus. Of this type a feebler form is produced when the

maxillaries only coalesce with the ossified alinasal wall, as in Dendrocolaptes albicollis,

Thamnophilus doliatus, and Phytotoma rara.

For Professor Huxley’s account of the Desmognathous palate, I refer to his paper

(p. 485). Ihave one remark to make, namely, that the vomer is very broad antero-

inferiorly (as in the Chelonia) in the most perfectly desmognathous Raptorial palate,

112 MR. W. K. PARKER ON THE STRUCTURE

that of the “ Falconide,’’ also that the vomer is not small in the “ Ardeide”’ and

Spoonbill, but very large; it is primarily double in them, but mostly single in the “Coc-

cygomorphe ”’ (when present) and in the “ Chenomorphe.”’ In the gallinaceous “ Schizo-

enathee’’ the vomer is azygous ; in the charadrians it is double.

The varieties of the Desmognathous palate are four, namely :—

a. Direct, as in Falcons and Geese, when the maxillo-palatines meet below at the

mid line, as in the mammal. Two subvarieties of this form occur—in the Falcon, where

the nasal septum is ankylosed to the hard palate, and in the Goose, where it remains

free:

b. Indirect. This is very common, and is best seen in Eagles, Vultures, and Owls;

the maxillo-palatines are ankylosed to the nasal septum by their inner margin, but are

separated from each other by a chink. This is well seen also in the fledgling of the

Falcon, which is ixdirectly desmognathous at that early stage.

c. Imperfectly direct. This is where the maxillo-palatine plates are united by har-

mony-sutures, and not by coalescence. Example, Dicholophus cristatus.

d. Imperfectly indirect. Here the maxillo-palatine plates are closely articulated with

and separated by the median septo-maxillary, but there is no ankylosis. Hxample,

Megalema asiatica.

e. Double desmognathism. <A fifth variety may be added in such a case as Podargus,

where the palatines as well as the maxillaries largely coalesce below; to aless extent this

is seen in the gigantic species of the Hornbills, e. g. Buceros birostris (see Huxley, op.

cit. p. 446, fig. 28, PZ).

Having thus analyzed their morphological characters, we shall be better able to deter-

mine their zoological value. Although my proper business in this paper is the deter-

mination of purely morphological meanings, yet I venture to offer these to the zoologist

embarrassed with the dazzling variety of types.

On the Structure and Development of the Face im Cuculine Birds.

After the many hundreds of the “ Coracomorphe”’ have been set aside, there yet

remains a large residuum of high-class “ Arboreal birds.”

Some of these are easy to cull from the rest, as they “pass under the hands of him

that telleth them,” without any confusion. Such are the Pigeons; such also are the

Parrots; and the two “ Families” of the Woodpeckers stand apart. A legion, however,

of other forms follow; these, for convenience’ sake, may take their group-name from the

Cuckoo, e. g. “ Coccygomorphe.”

Professor Huxley’s ‘‘ Cypselomorphe”’ (op. cit. p. 468) is his worst group; and he

has evidently tied its subdivisions together under some degree of misconception of their

palatal morphology. rochilus has not “a true Passerine vomer” (op. cit. p. 454), and

neither this bird nor the Goatsucker is Mgithognathous : Cypselus is, however ; and yet,

lying close by the true Swallows (“Hirundinidz”’), it differs in many most important

particulars from them, having no singing-muscles, no czca coli, and is possessed of

organs of flight very similar to those of the Humming-bird.

In fundamental cranio-facial morphology the Swifts are most Coracomorphous, and

AND DEVELOPMENT OF THE BIRD’S SKULL. 113

the Goatsuckers least: the Goatsucker, agreeing in very important characters with

Podargus and Steatornis on the one hand, and with the Trogons on the other, lies alto-

gether on a lower and more embryonic level than the “ Coracomorphe,”’ especially those

to which the Swifts are allied. These other ‘“‘ Families” may take their place for the

present with the other equally discordant “‘ Coccygomorphe,” and the term “ Cypse-

lomorphz’’ be allowed to drop. The Swift, so far as its cranio-facial morphology is

concerned, is an ‘“‘ Aigithognath,” and I have treated of it in a paper which has been

read at the Zoological Society. The Goatsucker and Humming-bird come under the

plan of the present memoir.

On the Structure of the Face in Caprimulgus europzus.

The moth-shaped mouth of the Fern-Owl is roofed in in a manner peculiar to itself;

but its gaping face is developed by a process in nowise different from that which operates

in. its nearer or more distant relations.

In the basal figure (Plate XXI. fig. 8) the trabecular structures are largely hidden

by the palatines ; and to understand this most frog-like of all the skulls of the “‘ Carinatze,”’

the student must examine the actual object.

Bony matter is here spun to its uttermost degree of fineness, and the resulting sub-

stance is something more like feather than bone. The whole shape is flattened and

outspread, and, as in the Batrachia, the originally flat broad trabecule have retained

much of their primary form. In front, however (Plate XXI. fig. 8, and Plate XXII.

fig. 1, sv), they are narrowed, and quite absorbed anteriorly, as in all other birds; at

least, the azygous prenasal outgrowth is early absorbed. Behind, the bony trumpets

that ‘grow from the ossifying apices of the trabecule spread far apart, forming the

“anterior tympanic recesses” (atr), and enclosing the V-shaped double Hustachian

tube (ez).

The trabeculz, as they surround and converge in front of the pituitary body, are con-

verted into a broad and winged mass of bone by the parasphenoid (pas), and at each

side of the common Eustachian opening there are two “ conjugational processes ”’

(fig. 8, bpg). ‘The lower of these is the “‘ basipterygoid ” proper, and it carries the free

cartilaginous plate for pterygoidean hingement; this flat broad process looks forward.

Above this is a retral hook, which merely holds a ligament.

As in the Frog, the trabecular and parasphenoidal structures project but little below

the lateral ethmoids (Plate X XI. fig. 8, pp), so that each pars plana seems to grow out

from the median, beam-like, large-winged outgrowths: they have carried the “ palatine

conjugational process” (ow) with them. This does not ossify into an “ os uncinatum,” as

is common in the “ Cuculinz,” but remains persistently soft as hyaline cartilage.

In front, between these ecto-ethmoidal wings, the broad parasphenoid and the trabecular

base of the meso-ethmoid each separately taper to a point; and here the first facial arch

has part of its long commissure absorbed to form the cranio-facial hinge.

In the pin-feathered nestling the trabecular base of the septum nasi (Plate XXII.

fig. 1, sv) is rather solid, but the azygous rostrum has been already absorbed. In the

adult (Plate XXTI. fig. 8, sz) the “ posterior septal bone” is rather solid than spongy, and

114 MR. W. K. PARKER ON THE STRUCTURE ©

its edges can be seen in an upper view (Plate X XI. fig. 10) projecting beyond the nasal

processes of the premaxillaries and upper spur of the nasals, and in the lower view

(fig. 8) above and in front of the vomer. But the rest of the trabecular part of the

septum nasi is absorbed, and only its nasal part is retained; it is bony, and the bony

matter passes a short distance into the paired “ale” (figs. 8 & 10). As to the secondary

or splint-bones of the trabecular arch, I have already spoken of the parasphenoid: the

others are, in this case, the premaxillaries and the vomers. The former (fig. 8, px)

appear very small on the lower surface, but above (fig. 10) each half is seen to be equal to

the corresponding maxillary, and to overlap it, as in birds generally and Teleostean Fishes.

The dentary margin, however, does not reach nearly so far back as in birds generally, and

this double bone is ornithically very small.

The other double bone isthe vomer. This is a long bar of spongy bone, rather bluntly

pointed at the fore end and sharp behind (figs. 8 & 9, v); it is quite distinct from the

overlying ethmo-palatal bars in newly fledged birds (fig. 9); but in the adult (fig. 8) it

becomes a derived commissural band to the palatine arch, as in a large number of birds,

notably the “ Coracomorphe.”

Flat above, where the palatal bars lie upon it (fig. 9, v, epa), the vomer is carinate below

throughout its entire length (fig. 8). In the younger (pin-feathered) fledglings I found

the vomers distinct throughout their whole length (Plate XXII. fig. 1, v). They were

long styles, approximating very closely in front, and gently diverging behind. They

were seen to have been formed by ossification of a fibrous and not a cartilaginous tract.

In more mature fledglings the fore ends had united (Plate XXI. fig. 11, v), and the

hinder ends were still more divergent. These again approximate, ankylosis takes place

thoroughly fore and aft, and the inferior surface ossifying, the interpalatine “ raphe ”

acquires its long keel. The fore end of the vomer in the adult (fig. 8) becomes rounded,

losing the beaked form it had in the newly fledged bird (fig. 9), and at this part below

the maxillo-palatine hooks (map) are notched on their inner margin to articulate with

this extremity of the vomer. A character like this gives the palate a very ‘“ Aigitho-

enathous”’ appearance, as this curious articulation, which bridges over the chink on each

side, is often seen in southern “ Coracomorphe,” both those of low and also of a high

type, as in Grallaria, Thamnophilus, Dicrurus, Dendrocolaptes, Homorus, and G'ymno-

rhina. But I have already said that these vomers are membrane bones; and I here show,

in three stages, the longish ligament which binds the vomer to the internal alinasal

wall with its long worm-like turbinal (Plate XXII. fig. 1, and Plate XXT. figs. 9 & 11,

aln,v). In the second of these three stages (fig. 11), the inturned alinasal wall is seen

to lie on the broad maxillo-palatine ; but it stops far short of the vomer, which also fails

to spread its shoulders towards the cartilage.

The palatine arch of the Fern-Owl equally departs from the Coracomorphous type, and

cleaves to what is signally Cwculine—I mean, in the broad sense of that word.

It has been already shown that the distinctively Passerine or Corvine character of the

palatine is the metamorphosis of the primary bar into fowr pairs of true morphological

endoskeletal elements, to say nothing of the secondary bones. But in Caprimulgus, my

youngest specimen (Plate XXIT. fig. 1) only shows a trace of true cartilage on the edge

AND DEVELOPMENT OF THE BIRD’S SKULL. 115

of the overgrown “ transverse”’ region; and the first and only palatine centre runs riot

over both morphological territories. Also in the hinder division of the arch-moiety the

pterygoids correspond to those of the young nestling, but not to those of the fledgling Pas-_

serine; for each pterygoid sends a spike-shaped process over the palatine; but this does

not become a “‘mesopterygoid ” bone (Plate XXI. fig. 11, and Plate XXII. fig. 1, pg).

The lower fore-looking spines of the palatines do not increase much in size—compare

the “ interpalatine”’ of the young (Plate XXII. fig. 1, ipa) with those of the adult

(Plate XXI. fig. 8). But the upper or orbital processes, the ethmo-palatine bars, grow

exceedingly, and meet by suture at the mid line (compare Plate XXTI. figs. 8, 9, & 11,

epa). This apposition is a step towards a true facial “‘ commissure.” The palatines are

elegantly bowed outwards from each other behind this tract, and then form a complete

commissure, like that of the trabecule (Plate XXI. figs. 8 £9). By the time this bird

is fully fledged the posterior palatal region is joined together at the mid line by a

cartilage common to both sides, and this cartilage becomes ossified by two endosteal

centres ; these I propose to call the anterior and posterior “ medio-palatines ” (Plate X XI.

figs. 8 & 9, mpa). Now the morphologist, at first ‘rounding about in darkness,” feels

himself safely within the great Cuculine territory, knowing that there and amongst the

** Celeomorphous ”’ tribes (Picidz) this structure is alone to be found *.

With regard to the articulation of the pterygoids with the basipterygoid processes

(bpg), Caprimulgus is countenanced by many types within the great Cuculine circle.

I see them everywhere in smaller or larger rudiments; but they are perfect in a very

near relation, the Oil-bird (Steatornis), and also in the Trogons and Humming-birds.

There are but two pairs of palatine splints, the maxillaries and jugo-quadrato-jugals

(me, 7); the latter are mere curved needles of bone; the former are elegantly ornithic pieces,

with, however, characters that are peculiar to this bird. In the young (Plate XXI.

fig. 11, and Plate XXIT. fig. 1, mx) the maxillary differs but little from that of Struthio

camelus (** Ostrich Skull,” plate viii. fig. 2, and Huxley, “Classification of Birds,”

p. 420, fig. 1, xp); for the palatal plate is rough, large, and 4-sided, and the cuneiform

body of the bone is very similar with its zygomatic process. In the adult, however

(Plate XXI. fig. 8), the change is as though the most delicate chiselling had been

fining it down into a very Passerine element ; the maxillo-palatine hooks especially might

belong to a southern Crow. The bone has its own caprimulgine diagnostic in an

f-shaped fore-turned spur, which is applied, for defence, to the nasal wall on each side

(Plate XXI. fig. 10, mp.mx). The figures show how thoroughly non-passerine is the

nasal labyrinth (Plate XXI. figs. 10 & 11, and Plate XXII. fig. 1, aln, als, atb, itb);

the upper view (fig. 11) shows, through the transparent tubular nostril-walls, the ali-

nasal turbinal (a/b), which is exposed in the inferior view of the younger specimen.” The

inferior turbinal (2¢b) is similarly displayed in the two figures, its line of insertion being

seen from above, and its anterior half is shown in the lower view (Plate XXII. fig. 1, 7¢6).

I have already described the relation of the nasal wall to the vomer, and have only to

* T am well aware that in the larger Gralle, Mycterta, Baleniceps, Ciconia, &c., and the “ Totipalmatz,” the

palatines form a bony “ commissure ;” but there is no key-stone in their palate. In the Pelicans this common bar

sends upwards a crest which is the symmorph of the trabecular crest.

SECOND SERIES.— ZOOLOGY, VOL. I. R

116 MR. W. K. PARKER ON THE STRUCTURE

add that the upper and middle turbinals are not more developed than in the common

Fowl. The small nasal septum (sz), diminishing forwards, is the only part ossified in

front of the hinge, the bone running a little into the alee ; and also in the true olfactory

region the root of the simple aliethmoid alcoil is alone ossified ; and this growth comes

directly from the perpendicular wall, or meso-ethmoid. The lachrymal (fig. 8, /) is a large

spongy bone, broad in the roof of the skull, pedate where it binds upon the zygoma,

and it entirely excludes the lateral ethmoid from the upper and outer regions of the face.

This is a Cuculine character.

On the Structure and Development of the Face in the Humming-birds

(** Trochilide’’).

Professor Huxley (op. cit. p. 454) says “ Trochilus has the true Passerine yomer, with

its broad and truncated anterior and deeply cleft posterior end. I have not yet been

able to obtain a perfectly satisfactory view of the structure and arrangement of the

palatine bones in the Humming birds.” That same paper contains several figures drawn

from his friend’s preparations ; and I wish those which could have been supplied to him

of the Humming-bird’s skull had been used by him also; they would have prevented the

error contained in the above quotation. I have been long familiar with these parts in

the old and nestling bird, in specimens from Barbadoes (? Lampornis); and now I am

able to add the structure of the largest known species, from the Andes of Peru (Patagona

gigas) *.

It might have been supposed that the Humming-bird would have been especially

related to its “ Tenuirostral’? isomorphs the Sun-birds (“ Nectariniide”’) and their

allies; but I find nothing in these, except extreme prognathism, which indicates any

special relationship. Having carefully dissected and drawn several scores of Passerine

types, to say nothing of the Cuculines over the border, it seems to me not altogether a

difficult task to weigh and measure the relationships of these bright “ minims” of

vertebrate life. No isomorphism apparent in types that grow from the uppermost twigs

of the life-tree should satisfy the ornithologist in matters of affinity; down by the

root must morphological observation be commenced. I had hoped to find the Trochiline

congeners in southern-world forms, such as Nectarophilus and Anthreptes, from Celebes,

or in the Australian genera Acanthorhynchus and Ptilotis; but such is not the case.

Morphologically—I do not say zoologically—I am nearer my starting-point down among

the superstruthious Hemipods, and the Passerines that have sprung up from that low

level, such as Pipra, Pachyrhamphus, and Thamnophilus.

All this is said in relation to Passerine modification, as such; not as seeking to lower

the idea of the specialization of the Trochiline type, which is without rival. These

zoological ideas force themselves forwards; but my work is with the morphology of the

region of the face.

* The old and young specimens came to me, the former in spirits, the latter dried in a letter, from my friend the

Rev. Jno. Philp, for several years missionary in that island. The gigantic form, also a spirit specimen, belongs to my

excellent co-worker in this field, Professor Alfred Newton, F.R.S.

AND DEVELOPMENT OF THE BIRD’S SKULL. 117

The forth-drawing (prognathism) of the facial bars has been carried to its uttermost

extent in the Humming-bird; but the secondary parts of the trabecular arch occupy two

thirds of the length anteriorly, their endoskeletal axis or prenasal rostrum early under-

going absorption. Behind, the apices of this arch are buried and lost in the outworks

of the tympanic cavity; and below, just in front of this part, the great splint, the para-

sphenoid, marks the bowed converging bars (Plate XXII. figs. 2,3, 4, 8, pas). Here, just

outside the common Hustachian meatus (ew), the trabeculae have sent out conjuga-

tional spurs, with their obliquely fore-looking facets for the pterygoids.

This high form gives us the extreme of a series commenced in the Struthionidz, where

the hinder end of the pterygoid is wedged in in the Emu between the quadrate and

basipterygoid process. Hven in some “ Ratitee” (see “ Ostrich Skull,” plates vii—xiv.)

the articulation is further forwards, whilst the series is obtained by passing to the

Tinamou (plate xv.), the Syrrhaptes (Trans. Zool. Soc. 1863, vol. v. pl. xxxvi. fig. 1);

then come the Plovers, Pigeons, Owls, &c., where it is in the middle; the “ Alectoro-

morphze”’ and ‘‘ Chenomorphe,’’ where it is at the anterior third; and, lastly, here in the

* Trochilide”’ it is quite at the fore end. Of necessity, this relative motion, as it were>

of the palatine on the trabecular arch is attended with shortening of the basipterygoid

processes as we pass along the series from the Emu to the Humming-bird. The tra-

becular base of the meso-ethmoid (Plate XXII. fig. 3, pas) is short, broad, pointed, and

keeled in front; and the hinge between this part and the septum nasi is finished by

uncleft cartilage above (Plate XXII. fig. 4, eth, sn). The remainder of the trabecular bar

is one with a most small ¢rwe nasal septum; there is a larger posterior and a smaller

anterior osseous centre in this small cartilaginous keel (fig. 4, sv). There are several

secondary bones applied to the trabecular arch besides the hinder plate, or parasphenoid.

In front the premaxillaries (pr) take the lead; they are here at their highest develop-

ment; they reign in this type, especially, and all things else in the organism are in con-

formity with them.

In the old bird (fig. 2) the premaxillaries are three fourths the length of the face; in

the nestling (fig. 8) they are only two thirds; so that it is evident that this excessive

prognathism takes place by the working of a general morphological law.

The boundaries of this double bone are best seen in the young (fig. 8); the dentary

processes (dpa) nearly reach the maxillo-palatines (map) behind, and the palatine pro-

cesses (figs. 8 & 9, ppx) are the twentieth of an inch shorter; as usual, the nasal pro-

cesses (fig. 5, wpa) are strongly grafted on and ankylosed to the forehead. Laterally

the upper and lower portions of the premaxillaries are thinned out extremely, and present.

ragged edges towards each other: this can be seen in the lower view (fig. 2). The halves

of the vomer are thoroughly fused together in my young specimen (figs. 8 & 9); they show

no sign of being formed on any thing else than a tract of fibrous tissue, and their likeness

and dissimilarity to the true Passerine vomer is well shown.

The long anterior spike in which the coalesced pieces terminate in front would not of

itself exclude them from the “ Aigithognathe ;” in the typical Chough (Pregilus graculus),

and in the aberrant Lyre-bird (Wenura superba), the double vomer is blunt-pointed in

front ; but in them, supero-laterally, the scars are seen in the dry skull where the alinasal

R 2

118 MR. W. K. PARKER ON THE STRUCTURE

cartilage has been macerated away. Besides the absence of the vomerine cartilages, we

have, then, in the “ Trochilide,” a lax relation of the vomer to the alinasal wall (see

figs. 2, 3; 8, 9, v, ial), the two being connected by a fibrous ligament. But the bone is

well shouldered, and above the broadish part the additional bones are seen, namely, the

‘* septo-maxillaries”’ (figs. 2-4, sma), so that the morphological elements are all here

except the pair of vomerine cartilages ; yet the metamorphosis of the parts is arrested

and the elements remain distinct. Besides the paired septo-maxillaries (smz), there is an

odd one, lying on the vomer, below the septum nasi, and in front of the small pair of centres

(fig. 4, jma’). These three,bones coexist in certain Aigithognathee, as Gymnorhina, where

they all coalesce with the vomer, and Homorus, where the median piece remains distinct:

This azygous piece I often find when the vomer itself is azygous, as in Anser palustris,

Psophia crepitans, Circus cyaneus, Haliastur indus, &c.; it is related, morphologically,

to the base of the cranio-facial notch, and is distinct from the centre which ossifies the

postero-inferior part of the septum nasi. Thus the unspecialized submucous plates seen

in the vomerine region of the Sturgeon reappear in their old profusion in these nobler

types, but are specialized to the highest degree.

The next arch, the pterygo-palatine, has as much and as little of the Passerine type as

the one described. ‘The palatines themselves are of great length (figs. 8 & 9, pa), and

are extremely slender, especially the distal part; they thicken gradually to the widest

part, and then gently taper off to their posterior end. On the inside of the widest part

each bone sends forward a long interpalatine style (¢pa) ; and on the outside a wart-like

protuberance represents the transpalatine of the “ Algithognathe.” This is only seen

in the adult, and isa mere periosteal outgrowth (figs. 3 & 4, ¢pa).

In some low-type Passerines, such as Pipra and Pachyrhamphus, the transp2latine is not

much larger; in the young Humming-bird the palatines are as much deficient here as

those of the Hemipods. The ethmo-palatine lamina (figs. 9 & 10, epa) is very much

stunted, the reverse of what obtains in the Swift and Goatsucker, where it is of extreme

length; and in them the vomerine crura articulate and then coalesce with these lamine,

whereas they remain distinct in the “ Trochilide.” The short stoutish pterygoids (pq)

are scooped in front, where they rest upon the end of the palatines ; in the young (fig. 10,

pg, mspg) the mesopterygoid piece is nearly distinct from the body of the bone; it

becomes quite separate, and then coalesces with the palatine. Both palatines and

pterygoids are faceted with cartilage to glide upon the short, stout, flat-faced basi-

pterygoids (bpg). Hach pterygoid is sharp-edged both above and below, and above the

shallow cup for the pterygoid there is scarcely any epipterygoid process (epg). In this

these birds are contrary to the “‘ Aigithognathe.” In the young (figs. 8 & 9, mx) the

maxillaries are clearly traceable; the broad flat part is about equal in length to the

long terminal spike, which is equal to that of the palatine; already, the posterior style

(zygomatic process) has coalesced witb the long jugal (j). The dentary edge is quite

overlapped by the premaxillary, opposite the end of which we see the characteristic

ornithie process, the “ maxillo-palatine.”

These outgrowths from the maxillaries are of great interest to the morphologist ; here,

in the Humming-bird, they show relationship as by the most delicate test. In the young

AND DEVELOPMENT OF THE BIRD’S SKULL. 119

(figs. 8 & 9, map) they agree in general with those of young Coracomorphe and Hemipods,

but in particular with both young and adult of that family of birds which has most

similarity to them in general bodily structure, namely Swifts (‘‘ Cypselide”). The

process itself is at first a mere hook, moderately thick, rather long, curving at first

inwards, and then directly backwards. This perfectly retrorsal rod runs directly along

the inside of the palatine; in the Swift it clears itself of the palatine, and is altogether

more flattened and sickle-shaped.

The maxillo-palatine of the ‘“Trochilidz’’ becomes more and more underlaid by the

palatine (figs. 2 & 3), so that a lower view shows little of them, and the specimen has to

be carefully studied in oblique aspects before its form and relations can be understood

(fig. 4, pa, mxp). Then it is seen that the little rod has become a flat thin plate, with a

terminal hook, and with its lower edge entirely ankylosed tothe palatine. This is similar

to the condition of things found in Cranes and Plovers, and totally unlike any thing seen

in the “ Agithognathe.” As far as my memory serves me, I have never seen any

ankylosis of these parts in the ‘‘ Coracomorphe,” not even in such highly ossified faces

as those of Phytotoma, Coccothraustes, Gymnorhina, or Artamus; the palatines, even

in these, keep scrupulously apart from the overlying maxillo-palatines.

The huge nasal labyrinth is much more in conformity with its counterparts in the Pas-

serine than the facial bars. The alee nasi (alm), like those of the Hemipods, are large

mussel-like valves, and have this special character, namely, that the widest part of the

nostril is behind; the teleological meaning of this. structure “lies at the door,” like the

other specializations of this type.

Unlike the generality of the small and delicate species of the “‘Coracomorphe,” such

as Acanthorhynchus, Nectarophilus, Phylloscopus, Troglodytes, Chelidon, &c., the nasal

outworks and walls are extensively ossified. The posterior septal bone is present, and

has a smaller ossicle in front of it (fig. 4, sv), and the anterior alar region (figs. 2-6, aln)

is ossified right and left in the recurrent lamina (7cc) and in the free valves; in the

latter there is a margin of cartilage left, and this has a selvedge of bone formed by a

distinct centre on each side. ‘This is the sickle-shape counterpart of the annular alinasal

ossicle of certain Parrots (Melopsittacus undulatus &c.). The large alinasal turbinals

(ath) are ossified to a considerable extent, the inferior turbinals (ib) scarcely at all.

The anterior nasal floor is but little developed; yet it sends inwards its inturned lamina

(ial), a broad unossified flap, to be tied to the vomer by a broad ligament (éal, v). The

true olfactory region (ecto- and meso-ethmoids) is of great size, and is densely ossified :

thoroughly Passerine, it has completely aborted the lacrymals, and appears on each side

of the frontals above (figs. 5 & 6, eeth), whilst below it forms most extensive antorbital

wings (pp), the suborbital part of which, reaching two thirds of the way towards the

quadrate, has no separate ossification of the ‘‘ os uncinatum.”

The skull is depressed: grooved above symmetrically for the long hyoid cornua

(8rd arch), it has its ethmo-presphenoidal plate (ye) solid, as in the Carp, and non-

fenestrate. The orbito-sphenoidal lips are narrow, and the orbital plates of the frontal

very little developed, so that the tilted floor of the cranium is largely membranous. The

120 MR. W. K. PARKER ON THE STRUCTURE

pier (quadrate) and proximal part of the mandible is much like that of the Swift. The

symphysis is long, and the distal part a mere copy of the upper jaw.

The 2nd and 3rd postoral arches are merely outdrawn modifications of the normally

ornithic hyoid bone, much less specialized than in the Woodpeckers. The true cerato-

hyals (fig. 7, chy) only extend for one fourth up the interior of the double horny sheath ; but

another fourth is filled, first with fibrous and then with fatty tissue. A short tract of

each bar is ossified, and all but the small bilobate end of the middle, or basi-urohyal bar

(bhy, bbr). The proximal part of the 3rd postoral (67) is only about one fourth the

length of the flat distal piece, which reaches, when the tongue is retracted, as far as to

the premaxillary apices: both are fully ossified.

Any one who will repeat these observations will find that the “ Trochilide”’ have no

right within the Passerine territory, and that they have at length diverged very widely

from the most striking tsomorphs within that range. The ‘‘ Coccygomorphe,” if they

be considered broadly, will be seen to be their truest relations; they are not, however, a

normal branch, but should be looked upon as forming a very distinct secondary leader,

arising close to the root of this widely ramifying group. It is well worthy of remark that

that Caprimulgine outlier of the “ Aigithognathe,” the Swift, comes very much nearer

the Humming-bird than the “ Nectariniide,” birds so mimetic and isomorphic of these

gorgeous “‘ vertebrated insects.”

On the Structure of the Face in Scythrops novee-hollandiz.

It would be difficult to find a more central form than Scythrops for the whole of the

* Coccygomorphe.” It appears to be equally allied to the Hornbills (“ Buceride’’),

the Toucans (“ Rhamphastide”’), the Plantain-eaters (“ Musophagidze”’), and the true

Cuckoos (“ Cuculidee’’). This paper, however, is given, not so much for the elucidation

of a difficult taxonomy, as for the purpose of exhibiting the more striking modifications

of the face and nasal organs in highly specialized birds. Ankylosis is in this case carried

to its uttermost limits as far as is consistent with the well-being of the type; yet the

regions can easily be determined. Rudiments of basipterygoid processes (Plate XXIII.

fig. 1, bpg) remain on each side of the subcarinate posterior part of the basifacial bar;

the bar becomes rounded and synovial where it glides along the common palatine groove

(fig. 2, pe, pa). In front of the hinge the bony septum nasi becomes lost in the delicate

diploé that, sponge-like, fills the. large upper jaw. Besides the huge premaxillaries,

which have melted into the common mass of the upper face, there is a well-marked

vomer (figs. 1 & 2, v), which appears as a thickish subcarinate plate of bone between the

maxillo-palatine wedge and the approximated interpalatine lips. This lesser bony wedge

thickens again above, and then thins out to articulate by fibrous tissue with a supero-

posterior ossicle. The vomer of Scythrops is an azygous bone, of the Chelonian type,

such as is also seen in Raptorial Birds, and in Mammalia of the Cat tribe ; but the hinder

part has been detached, and has coalesced with the palatines (ptpa, v).

There is, however, another element of the trabecular arch in this bird, which makes it

one of great interest to the morphologist; this is the os uncinatum. Notwithstanding

the normally large size of the lacrymal (fig. 3, 7), the ecto-ethmoid, or pars plana (pp),

AND DEVELOPMENT OF THE BIRD’S SKULL. 121

is a great squarish plate of bone; as in Caprimulgus, the infero-external angle of this

plate has its ossification stopped short, leaving that tract which primarily grew out of

the trabecule to conjugate itself with its counterpart process on the palatine bar. This

yoke-band is proximally broad and triangular (figs. 1 & 3, ow), and the apex of the

triangle, which alone is ossified, grows into two long and slender lashes of cartilage, one

of which runs directly across to its endoskeletal symmorph, tying itself to the very apex

of the ethmo-palatine crest (fig. 3, epa, ow), whilst the other runs forwards and down-

wards, and is attached to the upper surface of the palatine, near its outer edge and

opposite the fore margin of the maxillo-palatine floor (fig. 1, ow, mxp). The strong

palatine arch has, notwithstanding its intense specialization, attained the very form

which the foremost (trabecular) arch assumes as soon as it can be seen in a vertebrate

embryo as clouded bands of granular bioplasm. Here we have symmetrical sigmoid

bands, which behind (at their apex) are strongly hooked upwards and inwards; these

bands approximate, coalesce, and then diverge to their free flat anterior ends. Whilst

undergoing this junction the trabeculz form not only a commissure but a crest ; here, in

the palatine arch of Scythrops (figs. 1-3, pa, epa, pg, mspg), each bar grows upwards at

the commissure, which forms a semicylindrical trough above (fig. 2), on each side of which

the ethmo-palatine processes (trabeculo-palatine conjugational outgrowths) project.

Each pterygoid (pg) expands in a very ichthyic manner on the fore edge of the

quadrate, the expansion being the posterior margin of the subterminal part of the bar ;

the real apex is the epipterygoid hook (epg), which is as well developed here as in the

Finches. Each pterygoid is a strong bone, sharply keeled both above and below; leafy

above and hooked below, where it articulates with the palatine. It has evidently yielded

a mesopterygoid element (mspg) to the upper cartilage of the palatine behind.

The postpalatine region is vertical, the prepalatine broadly horizontal; the middle

part is mixed. The upper edges have united by ankylosis, not, however, without a

key-stone, a detached postvomerine piece, and perhaps also a medio-palatine (figs.

1 & 2, v, pa).

The deep thick-edged interpalatine lips, where they embrace the exposed vomerine

plate, nearly meet (fig. 1, ipa), so that altogether the lower or hard palate is of great

extent and complexity. By far the greater part of the hard palate is due to the larger

growth of the median and lateral palatine portions of the overgrown premaxillaries

(ppzx); but there is no little of the maxillary element besides. The proximal or lateral

portion of the maxillo-palatine plates (figs. 2 & 3, mxp) is spongy and of great width ;

it rises (fig. 3) up in front of the ethmo-palatine (fig. 3, epa); but the heart-shaped

median portion, which belongs to both sides, fills in half the space between the median

palatine process of the premaxillary and the gap formed by the diverging palatines. My

figure (fig. 1) does not show more than the inferior surface of this common median

maxillo-palatine mass ; but above, in front of the vertical vomer, it swells out into a very

elegant bulb of spongy bone, which is exactly heart-shaped, the base looking upwards.

The ankylosed ‘“‘jugal” bar (j) is strong, and greatly inhooked behind. The small

ossified alee nasi with their bony rudiment of a turbinal (fig. 3, alm, atb), form a nostril

that looks downwards and backwards. The whole labyrinth is huddled into a small

122 MR. W. K. PARKER ON THE STRUCTURE

space between the nostril and antorbital, and the short inferior turbinal is ossified. The

ecto-ethmoid (fig. 3, pp) is entirely excluded from the frontal region.

On the Structure of the Skull in Megaleema asiatica.

Having described a good representative Cuculine face, I take as my next instance that

of a bird which has a doubtful and mixed morphology. The physiognomy of the living

bird corresponds with this; for the most trained eye, seeing it for the first time, is

arrested by a sort of play and interchange of features, now looking like a relative of the

Cuckoo, and anon seeming to belong to the Southern Shrikes or Crows (e.g. Vanga,

Coronica, Gymnorhina).

Seen from below (Plate X XIII. fig. 4), the imperfectly “ desmognathous” and sezmi-

passerine characters are seen at one view. ‘The bony structures have the same Ouwl-like

spongy character as is general in the Cuculine birds; in the ‘“ Coracomorphe ” they are

more fibrous and stronger; they are more cleanly built.

In Megalema the basipterygoids are quite aborted, and the basifacial beam is

rounded to its end, the parasphenoidal rostrum ending below the hinge as a blunt spur.

In front of the hinge there is a solid postseptal bone, which ossifies that part of the

trabecular bar; for the rest, the septum nasi is aborted to a great degree, yet the recur-

rent alinasal folds send backward a style of bone which reaches almost to the vomerine

forks. The nasal labyrinth has its scrolls stunted ; those of the “ale” are partly ossified,

and the alinasal wall has become thick bone externally (ain); in front this bar divides

the external space into an anterior ¢rwe and a posterior false narial opening, as in many

Cuculines.

The spongy premaxillaries are remarkable in having their thick, hollow, rod-like

palatine processes free posteriorly (fig. 4, ppx). The dentary termination is equally

thick, but its lower edge is sharp and dentate.

The os uncinatum is not distinct from the pars plana (figs. 4 & 5, ow, pp), but it has

retained its primary embryonic shape as a somewhat hooked triangular flap. The

vomerine series of trabecular splints is extremely interesting. There is no evidence of a

cartilaginous origin for the evidently double vomer (v); but its explanation must be

sought for in aseries of forms outside the Cuculine boundary, as well as near the border,

namely, in the Humming-bird. As in Gymnorhina, Homorus, and Trochilus there are

jiwe vomerine bones, two vomerine moieties, forming a most elegant Y-shaped bone, a

pair of septo-maxillaries, and a median piece (fig. 4, v, sma, sma’). When the septo-

maxillaries are well developed, as in the ‘‘ Ophidia,” they stretch themselves beyond the

vomers, fore and aft, as is well seen in the Cobra (Naja tripudians); but when they are

feebly developed, a bony rudiment may appear in any part of the tract occupied by them

in their larger growth.

Thus in the Sparrow (Plate X XI. fig. 4, smx) they appear outside the broad shoulder

of the vomer ; in the Humming-bird in front of the broadest part; and here, in MWegalema,

behind, answering to that terminal spike of the bone in the Cobra which articulates

with the ethmo-palatine crest. I am satisfied that this is the true interpretation of the

posterior pair of vomerine bones in Megalema; for the bone does not form a palatine com-

AND DEVELOPMENT OF THE BIRD’S SKULL. 123

nmussure, and the medio-palatine, which occurs there, is azygous. The median septo-

maxillary turns up again in the Woodpecker; it has been already described in the

Humming-bird (Plate XXII. figs. 3 & 4, sma’); it exists free in the Coracomorphous

genus Homorus, and ankylosed to the vomer in Gymnorhina. For a vomer like the

one figured here (fig. 4, 7) we must look to some Celebesian Passerines. I have it in

Nectarophilus Grayi and in Anthreptes malaccensis; especially characteristic is that of

the former *.

But the vomer in these Passerines is confluent by its forks with the inturned alinasal

wall; in Megalema the arms of the Y form a neat joint, apparently without a cavity

(Plate XXIII. fig. 4, v, map), with the corresponding maxillo-palatine plates. The nearest

relation of this type known to me at present is the Roller (Coracias garrula) : this is one

of Cuvier’s “‘ Passerine ;” but it is far less passerine than Megalema, and has a chelonian

azygous vomer. ‘The narrowish, parallel, neat palatines are very much like those of the

Roller, but they are narrower, have a less distinct inner keel, and have distinct interpalatine

processes (pa); the rudimentary transpalatine angle (¢pa) is nearly equal in the two

kinds. The large mesopterygoid (fig.5, mspg) is semidistinct, and has not united with the

palatines in this specimen. In Coracias these parts are quite passerine, the mesopterygoid

having united with the palatine on one side, the left, whilst on the other all these are

fused together. In Coracias the epipterygoid process is abortively developed; in

Megalema (fig. 4, epg) it is an elegant rounded lobe. In Coracias the maxillo-palatines

are large, and have a complete ankylosis with each other; in Megalema (fig. 4, mxp)

they are smaller and do not meet, being kept apart by the “ median septo-maxillary ”’

(smz’).

In Coracias the lacrymal has its largest development and the ecto-ethmoid the least ;

in Megalema these things are reversed, and the lacrymal (fig. 5, 7) is a mere point,

whilst the ecto-ethmoids (eeth, pp) are like those cf the Passerines and Hemipods, being

largely exposed in the frontal region. In the structure of the nostrils the two types

agree; but conformably with its more intensely cuculine character, the Roller has a

very massive, spongy, osseous nasal septum ; it also has very large but non-functional basi-

pterygoids.

The characteristic modifications of Coracias are exaggerated in Hurystomus, save that

its basipterygoids are aborted. In no Owls that I have seen does the bone become so

frail a sponge as in the enormous lacrymals of that type; the ecto-ethmoids are in it

very minute flaps, the outer part of which is not ossified, as in Caprimulgus. Hurystomus

is a Roller, making no inconsiderable approaches to the Owl-like nocturnal Cuculines

next to be noticed.

The first of these is the Guacharo, or “ Oil-bird” (Steatornis caripensis, Humb. ;

Podargus, Cuv., Temm.).

The high arch both of beak and skull makes this bird’s head most completely iso-

morphic of that of the Owl; its basipterygoids are well developed (of enormous breadth,

the articular facet being oblong rather than oval), thus agreeing with the Ow/ and Fern-

'* The figures and descriptions of these forms, and of many others referred to in this paper, will soon appear in

the ‘ Zoological Transactions.’

SECOND SERIES.—ZOOLOGY, VOL. I. s

124 MR. W. K. PARKER ON THE STRUCTURE

Owl at once, and departing from Podargus ; Steatornis also agrees with Caprimulgus in

having the proximal part of the large flat antorbital (ecto-ethmoid) ossified. I mention

these things here to distinguish this type from the next to be described; a full account

of it must be reserved for another time and place.

On the Structure of the Face in ine Giant Goatsucker (Podargas).

Having been led up to Podargus by aseries of marginal and of central Cuculine forms,

J am anxious in this paper to give the most intensely modified form of face that has as

yet come under my notice. A face and skull are present here which, like that of the

equally specialized skull of the Crocodile, give us a prolepsis of that of the Mammal,

with its hard palate, the last degree of desmognathism having been attained.

This apparently ancient type, perhaps an early tertiary or even secondary waif, is also,

like the Oil-bird, an isomorph of the Owl, but not to the same degree. The huge upper

face, mimetic of the succulent leaf of an Aloe or a Mesembryanthemum, is mainly com-

posed of the premaxillaries and maxillaries (Plate XXIII. fig. 6). The figures already

given of the small and somewhat Passerine Fern-Owl (Plate XXI. figs. 8-11 and

Plate XXII. fig. 1) will help the student to understand the development of this thick,

arched, scooped, and although apparently solid, yet in reality spongy face. From the

terminal hook of the neb to the frontal region above, and to the truncated palatines below,

there is no trace of a suture. To make the division below the fore face on the one hand,

and the hind face and skull on the other, more complete, the Jugal process of the maxil-

lary is absorbed, and the jugo-quadrato-jugal bar articulates with the maxillary imside

its dentary angle. Here, then, the only movement possible is at the cranio-facial

hinge; for the pterygoids overlap and are tightly strapped to the palatines. The great

main hinge is behind, namely, that of the quadrate upon the auditory wall (fis. 7, q).

The motion here is almost as limited as in the claw-hinge of a Lobster.

A rudiment of the basipterygoid is seen as a ridge, somewhat pointed in its middle,

which runs from the optic foramen below, and is soon indistinct. Between these ridges

the basifacial bar is subcarinate, and then flattens out, running into short bony wings,

which are the proximal parts of the ecto-ethmoids; these are squarish flat cartilages

(Plate XXIII. fig. 7, pp). The septum nasi (fig. 7) is flat, bony, and notched behind,

below the groove for the nasal nerve; the hinge between the septum and the meso-

ethmoid (fig. 7, pe) is perfect. ‘The fore part of the median ethmoid is slightly keeled

between the antorbitals; its line is forwards and upwards equally. The ale nasi and

all the turbinals are unossified, and are extremely small and feeble; for the descending

crus of each nasal, and the face-plate of the maxillaries, straighten the nasal cavities into

two widish tubes. The posterior trabecular splint, the parasphenoid, does not project in

front, and is very short; the great anterior splints (premaxillaries) have few rivals in

relative size. With the nasal labyrinth so aborted the vomerine bones were likely to be

feeble. A flat band of fibrous tissue runs froma very soft bar of bone in front (v) to the

ethmo-palatine hooks behind (fig. 9, epa). Below, in front, we see this spongy wedge ;

it is higher at its smooth lower face than the great maxillo-palatine plate (map) to which

it is ankylosed. Above, in the end of the membranous vomerine band, there is a small

AND DEVELOPMENT OF THE BIRD’S SKULL. 125

ossicle, which lies between the ethmo-palatine hooks (fig. 9, v*, epa); this is a minute

posterior vomer, this element being double, as in Scythrops and other “ Cuculine.”

Caprimulgus is very passerine in having a huge double (symmetrical) vomer (Plate

XXII. fig. 1, v); but Steatornis comes much nearer Podargus, and has an azygous vomer,

of a clubbed shape in front, articulating with the palatine commissure. But the flat

anterior foot of the Chelonian vomer is again, as in Podargus, represented by a small

bony wedge in the cleft behind the maxillo-palatine synostosis.

The elegant leafy palatine arch is as fraught with interest as any other part of this

bird. The long prepalatine bar is aborted (compare Plate XXI. fig. 8, prpa, with

Plate XXIII. fig. 6, prpa); the broad truncated ends of these, the shortest of ornithic

palatines, are joined to the great fore jaw by a half-ankylosed dentate suture. For

short distance they are distinct, but they soon converge; these interpalatine plates are

ankylosed together, and form a bony raphe below to this superaddition to the bird’s

hard palate. This line of union is half the length of the palatines; a rounded notch

separates the extreme ends of the bars. The transpalatine process is triangular, with a

rounded point; it is not one third the relative size of this part in Caprimulgus; in

Steatornis it is entirely gone, as in many Cuculines. In that intermediate Goat-

sucker the interpalatine processes are rounded lobes, that grow near to each other in a

horizontal manner, but do not meet; whilst in Caprimulgus (Plate XXI. fig. 8, ipa) the

interpalatine ridges run parallel a long way, a short distance apart, and end in a delicate

spur. In Steatornis the ethmo-palatal commissure shows no medio-palatal keystone as

in Caprimulgus (Plate XXI. fig. 9, mpa); but in Podargus, although this tract is very

short, only half the extent of the lower hard floor, yet here, again, we have the two

azygous medio-palatines, the undoubted symmorphs of those I have described in the

Fern-Owl (Plate XXI. fig. 9, mpa, and Plate XXIII. fig. 8, mpa). In Steatornis the

palatine commissure is longer than in either the Giant or the Common Goatsucker, more

like that of Scythrops (Plate XXIII. fig. 1, ipa); the posterior notch is similarly small.

In all these thin types the apex of the palatine arch, or epipterygoid, is aborted, and in

Podargus and Caprimulgus this bone is very spongy (Plate XXIII. figs. 6 & 7, pg,

and Plate XXI. fig. 8, pg); in Steatornis it is less spongy and more crested. In this

latter bird the facet for the basipterygoid has an overlapping process, by which it lies

over as well as against the basal facet; in Caprimulgus there is a slight rudiment

of this structure; in Podargus there is no joint at this part. All these three types

agree in being embryonic, as regards the mesopterygoid. This spur does not segment

off in either; in the old Fern-Owl it has become aborted; the young agrees with

Steatornis and Podargus (Plate XXI. fig. 11, and Plate XXIII. fig. 7, pg); it lies ina

depression and is strongly bound to the supero-median surface of the palatine. I find

no defined os uncinatum in either the Fern-Owl or Podargus; but in Steatornis it is as

well developed as in Scythrops (Plate XXIII. fig. 1, ow), and has two cartilaginous tapes

that tie it in front and behind.

The lacrymal is large in the Fern-Owl (Plate XXI. fig. 8, 7); in Podargus it seems to

be absent; but a small ossicle may have coalesced with the supero-external angle of the

nasals, in front of the narrow frontals, as is the case with a larger piece in Steatornis.

126 MR. W. K. PARKER ON THE STRUCTURE

The face of Steatornis is much like that of the Eared Owl (Asio otws), but the ale nasi are

much more ossified; they are quite soft in the other two, and ouly the fore edge is left

cartilaginous ; its septum nasi is also bony.

This agreement and disagreement, alternately, of these three Goatsuckers suggests the

probability that they each represent what were once large “ subfamilies;” but their

characters, by their intense inosculation, show that they belong to one highly natural

group.

The stapes of Podargus presents a very curious and instructive example of this hyootic

structure. The otic portion, or basal plug (figs. 10 & 11, st), and the medio-stapedial bar

(mst) are quite normal. The “suprastapedial”’ (ss¢) is large and unossified, and so

also is the infrastapedial (ifst), which ends below, continuously, in a spatulate recurved

stylo-hyal. The thing, however, to be noticed is the huge roughly spoon-shaped extra-

stapedial (est), all bony, convex outside, concavo-carinate within, and pneumatic; the

upper margin is evenly sinuous, the lower and the end sublobate; there is a fenestra

between it and the falcate suprastapedial (fig. 11). A ligament is seen growing from

the top of the extrastapedial spoon; and another larger ligament ties the supra-

stapedial to the prootic wall. The cartilage of the supra- and infrastapedial is con-

tinuous; the line of the original arch or bar is from the mediostapedial to the infra-

stapedial, whilst the suprastapedial becomes a second head; the extrastapedial is a

conjugational outgrowth. The bifurcation of the rod above produces processes which

are the isomorphs of the capitular and tubercular processes of a true rib; the supra-

stapedial resembles a tuberculum, and the mediostapedial a capitulum; the shaft, as we

saw in the Martin, chondrifies in patches, the backwardly bent stylo-hyal reuniting

with the upper piece by a later chondrification of the infrastapedial tract.

On the Structure of the Face in some other “‘ Coccygomorphe.”

In the Hornbills, the secondary growths of the trabecular and palatine arches, more

especially the former, acquire the uttermost degree of prognathic development consistent

with the mechanism of a bird’s skeleton as such. The result, taking in the correlated

modifications throughout the rest of the skeleton of these birds, is a form which makes the

conception of the living Péerodactyle an easy effort of the fancy. Pneumatic overgrowths,

of such a size as we see in the larger species, intensify the grotesqueness of the vreature ;

for the aneurismal bursting of the bony walls by the imprisoned breath has resulted in

their huge air-chambers, walled in merely by the horny epiderm and its filmy quick.

For a figure of the palate of Buceros bicornis the reader is referred to Professor Huxley’s

paper (op. cit. p. 446, fig. 28). Here the bony bar between the apertures marked a is the

foot of the Chelonian vomer, separated from its posterior portion by a membranous

space, and ankylosed to its surroundings, namely, the maxillo-palatines and the coalesced

interpalatines, behind; for these larger species have a palatinal hard palate. The styloid

posterior part is ankylosed with the palatines at their upper or ¢7we commissure. I see a

similar structure in B. ruficollis, Temm.; but the interpalatines are imperfect, and the

form of the vomerine foot is not so easily traceable. In this species the notch which

forms the cranio-facial hinge is reconverted into a fenestra by a tuberous growth of bone,

AND DEVELOPMENT OF THE BIRD’S SKULL. 127

a median septo-maxillary, which runs from the parasphenoidal beak to the bony septum

nasi, above the posterior vomer. The os uncinatum is not distinct in these Hornbills,

but the ecto-ethmoid terminates in a large pedate process which binds upon the zygoma ;

their lacrymal is aborted, and the large lateral ethmoid comes to the outside and top of

the face, as in Megalema and the Passerinee. With the complete ankylosis of the pala-

tines at their commissure, nothing can be learned from the adult as to the existence of a

medio-palatine keystone.

In the most exquisite of all vertebrate skulls, that of the Toucan (Ramphastos toco),

the structure largely agrees with the “ Buceride.”” The ecto-ethmoid projects as in them

and the Passerine, and the os uncinatum is represented by the pedate angle of that bony

plate. I see a small lozenge-shaped subcarinate vomer below the level of, but acting as

a secondary keystone to, the palatine commissure; this, however, is only the posterior

vomer. The anterior piece is a broad bifid ragged wedge of bone, with its narrow angular

end dovetailed in between the palatines, immediately above the small spiny semidistinct

interpalatines; its position is transversely vertical, and its size ten times that of the

hinder piece. The notch is complete that forms the wide hinge between this marvellous

face and skull; it is 14 inch across, and the beak of the parasphenoid is but little behind

a descending plate of bone, the sharply bifid middle portion of which is the posterior

angle of the nasal septum. The anterior vomer reaches upwards to these three bony

spikes by its ragged split upper end. The nostrils are nearly half an inch apart on the

upper surface, close in front of the hinge, and overshadowed by the rounded posterior

margin of the elevated beak. The thoroughly ossified ale nasi enclose a transversely

oval nostril; the other parts of the nasal labyrinth are all ossified, and the inferior

turbinals are exquisite little scrolls of scabrous bone, almost pisiform, and vertically

placed, the end of the scroll being seen from delow instead of in front as in other birds.

Between the splintery prepalatines and the maxillary synostosis there is a rather large

oval space; behind, the palatines are fused together.

The transpalatine region, with its retral spike, is something between what we see in

Megalema (Plate XXIII. fig. 4) and in Dacelo (Huxley, op. cit. p. 447, fig. 29, Pl). The

basipterygoids are small rudiments; the pterygoids are spongy, and have a small epi-

pterygoid lobe. This short ¢itial description may help the student to decipher the

morphology of this culmination of the “ Coceygomorphx.” TI have just referred to Pro-

fessor Huxley’s good woodcut ofthe palate of Dacelo ; in my specimens of the “ Alcedinze,”’

namely, Alcedo ispida, Upupa epops, Todus viridis, I see no vomer. These skulls are

thoroughly ‘‘ Desmognathous,” and are especially interesting as to their palatines, in

which the transpalatine sends backwards a spike, equal, in Alcedo, to the prepalatine

style in front ; in the young Hoopoe the transpalatine is obliquely set on to the palatine

as a distinct bone.

Inthe “ Musophagidee ” the os uncinatum attains its fullest development as an anterior

transverse bone (see Dr. Reinhardt’s paper, already quoted). In a species of Corythata

(2? buffoni) there are two medio-palatines and a posterior vomer; these lie behind the

short sutural commissure, in fibrous tissue ; for, posteriorly, the rounded palatines are

‘wide apart, exposing the parasphenoid.

128 MR. W. K. PARKER ON THE STRUCTURE

The foremost medio-palatine is a minute style, having the equally minute styliform

vomer beneath it; the second medio-palatine is a mere grain of bone *.

The nasal labyrinth is well developed in these birds, and is thoroughly ossified ; so that

the bony skull of no other bird is so useful to the student who would master this elegant

sense-organ.

On the Structure of the Skull in Birds of Prey (« Aetomorphe,” Huxley).

Both for this and for the Goose tribe I have sought to extend the lines that bound in

their “lot,” confident that in the Southern World we have extant, and, toa much greater

degree, in the buried Tertiary world there are extinct, many a type that cannot and could

not wear the ordinary ornithological harness. ‘The characters of an Hagle, a Falcon, or

an Owl may easily be set down; but when the scientific formalist comes athwart a bird

in which all these are shaken up together, like the lots in the helmet of Agamemnon,

then every thing breaks down, and his wire-drawn distinctions lose their value.

Yet these distinctions are not the less valuable, being applicable to the newest and most

specialized forms; but they melt away in certain types, just as the professional distinctions

of civilized life are seen to melt away when we contemplate the life and polity of

savage races.

I make these remarks in defence of the position in which it is here sought to place a

bird, the Cariama, the relationships of which seem to me to be self-evident, and which

has been looked at by most ornithologists through the coloured and refracting glasses of

their own arbitrary systems.

Having no misgivings as to the essentially Raptorial nature of the Cariama (Dicholo-

phus), I do the more boldly place it for comparison side by side with the two most

intensely specialized types in the whole group, viz. the Falcon and the Owl. Such a

collocation, however painful to the eyes of full-grown systematists, will, if its natural-

ness be proved, be of real value to the tentative worker, serving at least as a hint that

the strength of Nature will burst the new ropes we seek to bind her with, those bonds

being, indeed, as weak “as the tow when it smelleth the fire.”

On the Structure of the Skull and Face in Dicholophus cristatus.

The form of the whole head in this bird is Vulturine (Plate XXIV. fig. 2), as may be

seen at once if it be compared with that of Gyps fulvus (fig. 1) t-

* It is not always possible to be absolutely certain as to whether any given grain of bone belongs to the hinder

part of the yomerine region or to the palatine commissure.

+ How the eyes of naturalists have been holden against seeing the truth here I cannot understand ; the legs of the

bird, so thin and long, have robbed it of its aquiline glory. A few years since, showing two of these birds in the

gardens of the Zoological Society to my brother, a man of like tastes with myself, he at once recognized the Cariama

as a kind of Hawk, whilst it crouched upon the perch. Another individual was standing in the enclosure in

front of the aviary: “ What bird is that ?” I immediately asked. ‘I cannot say,” was the answer.

Capt. Burton (‘ Exploration of the Highlands of the Brazil, London, 1869, vol. i. p. 57) compares this bird with

the Secretary (Gypogeranus), and says that it has the same serpent-eating tastes (see also vol. ii. p. 18).

The keeper of the ayiaries where the Cariamas are kept informed me that they eat mice and sparrows, beating

them about before they kill them. On July 31st, 1872, I watched both D. cristatus and D. burmeistert dust them-

AND DEVELOPMENT OF THE BIRD’S SKULL. 129

The first and most notable difference between the skull of the Cariama and that of

Gyps fulous is that it is generally feebler (Plate XXIV. figs. 1& 2: fig. 1, Gyps; fig. 2,

Dicholophus), and its ossification is arrested; but it has the elongated vulturine face ;

whilst Gypogeranus comes close to the Harriers, Kites, and Buzzards.

The first and leading characteristic of the skull in this type is its palate, which cor-

responds to that of a young Falcon (Plate XXIV. figs. 3 & 4), and gives me that modi-

fication of the Desmognathous skull which I have termed (supra, p. 112) “ imperfectly

direct,”” the maxillo-palatine plates (mxp) being united by harmony-suture and not by

coalescence.

How this skull has been taken for one of the “ Schizognathous”’ type, and put with

that of Cranes and Bustards, I cannot understand. In respect of its feebly ossified nasal

labyrinth, this form approaches and agrees with other South-American types, namely

e “ Cathartide ;” and the skull of Sarcoramphus has greatly assisted me in working

this kind out; but there is no kind of Raptorial skull that it is not, more or less, in

harmony with, in one point or another; and the whole group of the Aetomorphe may

in some long past epoch have lain in the loins of a bird but little different from the

Cariama.

There are three main varieties in the general bony characters of the Raptores : in the

Cariama the skeleton has a fibrous lightness and delicacy, in the Owls a soft spongy

condition for the most part, the long bones being dense as to their walls; but these are

of extreme tenuity. In the Falcons the bones, everywhere, are almost like ivory in their

density, reminding one of those of Serpents.

The skeleton of the Cariama has very little in common with that of the Crane tribe ;

it is as delicate, relatively, as that of that lonely waif of a subfamily of the “ Gruine,”

the Sun-Bittern (Hurypyga helias), and still more like that of another subextinct type

of ancient and generalized Cranes, namely the Kagu (Rhinochetus). But the Crane

with which one would naturally compare it is the Trumpeter (Psophia); nothing, how-

ever, can be more fatal to the view of its being a Gruine bird than such a comparison :

I would rather make room for it amongst the Gallinaceous birds than see it so mis-

placed. In the genealogical life-tree of the families of Birds, some grow near to each

other, and have, as it were, their branches forming an acute angle with their leader, as

in the Lombardy poplar; whilst others, as in the black poplar, grow out at all angles,

most of which are wide and divergent.

If we take the ‘“‘ Coracomorphe,” they do so cling to each other, diverging very

gently; but the “ Aetomorphe” delight in wide and wild divergence; and although

manifestly ‘“‘ of one blood,” they yet have acquired the greatest variety in their speciali-

zations; but the Falcon is “ brother to the Owl.”

Notwithstanding the enfeebled length of the beak of the Cariama, the bones are

thoroughly fused together. The broad basitemporals (fig. 3, 0¢) end in front, as in the

selves by rolling exactly like the Common Fowl: this is in harmony with the fact that they impinge very closely

on the gallinaceous circle. They are Raptores with a very gallinaceous aspect; whilst the vulterine Guinea-hen and

the Brush-Turkey are Fowls with a very vulturine aspect.

130 MR. W. K. PARKER ON THE STRUCTURE

Owl (fig. 5), in two pouting lips; they underlie the common Eustachian opening (ew);

outside and in front of this emargination the basipterygoids (pg) are seen projecting

from those wings of the parasphenoid that form the “ anterior tympanic recess.” The

narrowed part of the parasphenoid (pas) is relatively slender, and the pterygo-palatine

bones undergird it for most of its extent. The great anterior palatine interspace is large,

but it is lessened in the dry skull by an ossification which has affected the retral part of

the trabeculee and al nasi (fig. 3, alz). A similar bony bridge is formed in Sarco-

ramphus papa by the maxillo-palatines where they join the nasal septum.

Most of the nasal capsule, as in pluvialine and gallinaceous birds, is unossified. The

vomer (fig. 3, v) is of a type rare in birds; it is thoroughly Chelonian, more so than in

any other bird save the Falcon (fig. 4, 7), although some approach is made to this in the

Sacred [bis and the Tree-Duck (Dendrocygna).

As in the other Raptores, it is truly azygous, so also it is in the Gallinacez; but in the

Cranes and all pluvialine birds it is composed of long lathy pieces, that converge and

ankylose, only keeping apart behind.

In the Chelonia the azygous knife-shaped vomer spreads out behind and above to

articulate with the orbital plates of the palatines, whilst in front it descends and helps

the maxillaries to form an anterior hard palate. Just so does this bone behave in the

Falcon and Cariama. In the Green Turtle (Chelone midas), however, the palatines have

a small palatine lamina joining this fore foot of the vomer; this part is arrested in the

Cariama and the Falcon (figs. 3 & 4, prpa’). In a side view the vomer of the Cariama

(fig. 3%, v) is seen to be very high in the middle, low and flat in front, and gradually to

lessen behind; the middle part is deficient of bone for a large space. This vomerine

fenestra is repeated in the large Falcons (Plate XXV. fig. 12). The articulation of

the vomer with the ascending or orbital plates of the palatines is through the medium

of the keystone-piece of that arch, the medio-palatine (mpa). This bone is of great

interest; hitherto I have only found it in Birds, and amongst them most developed in

Woodpeckers (Trans. Linn. Soc. ser. 2, Plate II. p.9). But it has just been shown to

exist very commonly in Cuculine types.

There is no abrupt line of demarkation between this ossicle and the paired meso-

pterygoids; and all the gradations are seen in the Raptores, where these intercalary

pieces are very variously developed. In the Cariama it is a roughish wedge of bone,

rounded behind, pointed in front, and wedged in between the right and left ethmo-

palatines laterally ; vertically is jammed in between the parasphenoid and vomer. There

is one ossicle here in the young of Falco tinnunculus (fig. 4, mspg); but this is the

apex of a right mesopterygoid, the left being suppressed. In Sarcoramphus (fig. 19)

it is more truly azygous; in Neophron (Plate XXV. figs. 14.&15) it is a notable right

mesopterygoid; but in that bird the mesopterygoid appears at both ends of its region

(fig. 16), a little grain of bone occurring at the junction of the pterygoid on the

same side.

In Helotarsus ecaudatus there is one small bone (a right mesopterygoid) in front

(Plate XXV. fig. 9, mspg); but certain Owls (Strix stridula, Plate XXV. fig. 20, and

AND DEVELOPMENT OF THE BIRD’S SKULL. 131

Ketupa ceylonensis, Plate XXV. fig. 11) come the nearest to Dicholophus in respect of

this bone and its relation to the vomer; but in these birds the vomer has no fore foot *.

The prepalatine bars (prpa) are long and slender; the transpalatine is rounded off

(¢pa), and passes into a short postpalatine (ptpa), on the inner margin of which the sub-

mesial muscular ridge ends ; the whole form and manner of these parts is Falconine, but

drawn out. The upper part, or ethmo-palatine (epa), meets its fellow process for some

extent in the middle, the fore end diverging for the medio-palatine keystone {mpa), and

the hinder for articulation with the diverging pterygoids (pg). Threads of bone represent

the interpalatine (¢pa), which, if developed, would have given a desmognathous palate,

bound together in the same manner, and composed of the same parts, as that of the

Green Turtle, which has the anterior vomerine foot inwedged between rudimentary

palatine plates of the maxillaries and palatines. This Reptile stands halfway between

the Lizard, with its highly schizognathous face, and the Crocodile, in which desmognathism

is carried to its uttermost, as in the Mammalian genus Wyrmecophaga. Walf the

charm of this research would vanish if we could possibly forget that the bird groups are

luxuriant outgrowths of the reptilian understock, and that their fiery nature once lay hid

in the stone-cold Reptile.

Some of the Raptorial characters of the Cariama are exaggerated. The shortness of

the pterygoids (pg) is greater than in the Owl and Falcon; they answer, however,

exactly in character to those of the latter, having no articular junction with the basi-

pterygoid processes (dpg), these being arrested. Their epipterygoid process (epg) is a

hooked flange. The splints of the face are confluent, the marginal line being continuous

from the neb to the quadrate hinge; the hinder jugal bar is slender. About two fifths

of the premaxillary mass is continuous in the fore palate; the palatine processes of the

premaxillaries are long, slender, and lie outside the prepalatine laths. At the middle

these large bones are again united by a commissure, namely, the alinasal bridge (adn).

But the most remarkable part of the face is the maxillo-palatine floor, which is

relatively larger than that of almost any bird; the processes forming this hard palate

are flat ear-shaped lobes, which unite at the mid line by harmony. They are pointed

behind to let in the foot of the vomer, and broad and notched in front, behind the large

anterior palatine space. These plates have the thin fibrous character of the main part of

the skeleton, instead of being dense as in the Falcons, or spongy as in the Owls. The

skulls of those types when young (figs. 4 & 5) are exactly correspondent in their essential

morphology to that of the Cariama, being of the “ imperfectly direct’ desmognathous

type.

The general form of the skull seen from above, with its alate lacrymals, is evidently

an enfeeblement of the vulturine type (fig. 2) as compared with that of Gyps fulvus

(fig. 1); and what arrest and feebleness there is belongs not to the Crane, with its dense

heavy skull and square-browed lacrymals, but is something intermediate between that

* T have the more carefully gone over these details because of the instructive fact that the structure of the Cariama

is best explained by that of the various Rapiorial subfamilies, and is not illustrated by members of the family to

which it has been supposed to belong. My notes on the soft parts of this bird, and the observations and comparisons

I have made of the rest of its skeleton, all tell the same tale.

SECOND SERIES.— ZOOLOGY, VOL. I. T

1382 MR. W. K. PARKER ON THE STRUCTURE

of the Fowl and the common Vulture. In the Crane, a truly pluvialine bird, the maxillo-

palatines project but little mesiad of the prepalatine bar, the huge almost struthious

double vomer being wholly displayed in the upturned palate.

Structurally, then, the Cariama is seen to have the remotest relation to the Pluvialine

type; and if the details just given be weighed and measured, they will show that its

Raptorial characters vastly preponderate ; that it is evidently a branch of the same stock

as the true Gallinacez, especially the Megapods and Curassows, and that it does not

belong to any extant type of Raptorial bird; but as the Palamedea is a Chenomorph, so

is the Cariama an Aetomorph, not holding out the flag of any one party of that common-

wealth, but being, as it were, a representative of the old fathers of the race, whose sons

have acquired such sharp characters of dissimilarity. It contains in itself not merely

the attributes of a single subfamily, as Hawk, Falcon, Owl, Vulture, but “all these in

their pregnant causes mixed.”

There is a bone on each side in the face of the Cariama which is not seen well in the

palatine view; this is the os uncinatum (Plate XXIV. fig. 2a, ow); it is well seen

in the side view.

As I have shown in the former part of this paper, this is common in the Cuculines ; in

Dicholophus it is similar to but larger than its counterpart in the Gulls (Plate XX VII.

fig. 11, ow). It has a like development in the Alabatross, and is abortively seen in Uria

troile and Alcatorda. I find it inno other Raptorial bird, nor in any of the Grallee that

lie on the same morphological level as the Cariama, and although not belonging to its

group have yet some affinity with it.

Here the bone is a notable style perched upon the zygoma, and underpropping the

lacrymal (/); it is, however, connected with the feebly developed pars plana (pp), the

free tongue-shaped part of which is unossified as in the pluvialine types.

This is the ornithic representative of the cartilaginous bar which yokes the trabecula

to the palatine in the Frog, and therefore of great interest morphologically.

On the Structure of the Skull and Face in the Falcon (Falco tinnunculus).

Having treasured up from time to time every obtainable stage of young birds, I am

able to compare the adult Cariama’s skuli with the immature skull (that of fledglings)

of the Falcon, the Hawk, and the Owl, the proper morphological counterparts of the

skull of such a root-form as the one just described *.

I trust to the trained eye of the ornithologist to see at once the essential agreement of

this Falconine palate (fig. 4) with that of the Cariama (fig. 3). Nature, anti-Procrustean

in all her works, has developed one skull into a form which is feline in its shortness,

massiveness, and strength, whilst the other is outdrawn, and, as it were, of a vulpine

shape ; yet the “habit” of both types is manifestly one. The difference depicted is not

the full measure of what would have been seen if an adult Kestrel had been selected for

comparison.

* The same method with regard to the relation of the Plovers to the Gulls has appeared to be fruitful to my

accomplished fellow workers, Professors Huxley and Newton; and their coinciding judgment has been to me in no

little degree determinative of renewed labour in this field.

AND DEVELOPMENT OF THE BIRD’S SKULL. 133

The transverse basitemporals (fig. 4, b¢) have joined their wings beneath the common

Eustachian opening (ew). Above this point the parasphenoid (pas) is seen with small

basipterygoid knobs (dpg), and above these and the basitemporals expands on each side

into a large trumpet of bone, whose mouth opens into the labyrinthic drum-cavity.

These trumpets form the “anterior tympanic recesses,” and are lipped by one or more

additional (tympanic) ossifications.

The fore part of the parasphenoid has the shape of a planter’s dibble, and ends, blunt-

pointed, above the palatine commissure, carrying the hinge mesethmoid on its grooved

upper face. The premaxillaries form a triangle whose sides are but little longer than

their (ideal) base from side to side, and after forming themselves separately on the

spatulate prenasal cartilage, combine at the mid line, not only above but also below,

aborting and taking the place of their model, growing even there much more solid than

their displaced prototype.

This is the most Chelonian of all the birds’ skulls examined by me; and knowing full

well that the bird’s skull, after it has fused its trabecule and acquired a cartilaginous

consistency, takes on immediately the most unmistakable Rhynchosaurian characters, I

am in nowise surprised to see the Faleon (which shoots upward so high above the

Chelonian) in its ascent passing close to and retaining much of the likeness of that

less specialized and much less metamorphosed type. ‘That the study of the modifications

produced by metamorphosis of the primordial face is of the utmost importance to

taxonomists is evident; and the vomer alone is, in this respect, worth more than all

Mr. Garrod’s muscles put together. In the Reptilian group the Chelonia are alone in

possessing an azygous vomer; and only a minority of the bird class have the same

character. Here, in the Falcon, this bone is not shaped like the proper ploughshare,

but like that less common implement the breast-plough, for sod-turning. The blade of

this bone is roundly three-toothed, its shaft flat vertically, down-bent in front and lifted

behind, and the handle is thick and forked. The blade is wedged behind in the maxillo-

palatine harmony, and between the ear-shaped maxillo-palatine lobes. The shaft divides

the median nares, right and left; the two-toothed handle bites the palatine commissural

plates ; it is fastened there by a wedge of bone, which has been driven in, as it were, on

the right side. This wedge is the right mesopterygoid (mspg). In this type I have

found no other vomerine bones.

The pterygoids (fig. 4, pg) are even less specialized at this stage than those of the

Cariama; they are larger relatively, have a less distinct auricular process (epg), and

their abortively developed apex only appears on the right side as the mesopterygoid wedge

(mspq). |

The only difference between the palatines and those of Dicholophus is their greater

shortness; also whilst the inner ridges end in the Falcon as a short interpalatine spur

(ipa), those of the other bird are ragged and formless. In both cases the transpalatine

region (tp) is evenly rounded off, but the outer lobe is scooped below for the palatine

muscles. In the jugum of the Cariama the elements once combined ; but I suppose these

were then the jugal process of the maxillary, the jugal itself, and the quadrato-jugal.

These two latter are not separate, as far as I can see, in the Falcon, which thus

T 2

134 MR. W. K. PARKER ON THE STRUCTURE

agrees with a large number of the “ Aves aérize,”” whereas in Hawks and Owls they are

both present. But the most striking part of their palatine outworks is the huge

maxillary (ma); it is, relatively to the premaxillary, as extensively developed as in the

Chelonia, although the dentary process of the latter bone overlaps it as in osseous Fishes,

and, indeed, in birds generally.

These parts are builded together in a manner which would please the eye of an

architect.

But the whole meaning of these parts can only be seen by the study of sections made

in young skulls. In a nestling younger than the one from which the palatal view was

taken, I have been able to decipher all those parts of the face in which the nasal capsules

have their furniture disposed amongst and within the foremost facial bars.

In this type the nasal labyrinth enfolds itself and encloses its passages with solid car-

tilage to a very unusual extent. Here (Plate X XV. fig. 1, em) the external nasal opening

is not a long slit, but a neat round opening; the rim is wrought, like the rim of a

cup, and is almost closed by a finger-shaped projection from the alinasal turbinal (a7).

This process looks outwards and backwards, and only leaves around it an oval chink.

The whole alinasal region (figs. 1-8, alm) is formed into a pair of oblong oval pouches,

like two carpels of an apocarpous fruit dehiscing; seen from below, there seem to he

two cells left out of four; and the still persistent lanceolate prenasal cartilage (pz)

running from the base of the septum (sz) carries out still further the similitude, it being

in the likeness of the fruit-axis. Behind, at this most bulbous part, these nasal pouches

are complete tubes, the floor having coalesced with the middle wall (fig. 3, adn, sm); this

is shown in section (fig. 5), also the apposition of those parts further forwards (fig. 6).

When the septum is removed, to show the inner face of the turbinals, it is seen that

the alinasal turbinal (fig. 2, a¢b) is a large leaf of cartilage, hollow outside and bulging

within (see also sections, figs. 5 & 6). This leaf is attached to the outer wall by its

upper edge, and is also confluent behind with the bulging floor, where that part rests on

the nasal nerves (x). Over the hinder third the true “inferior turbinal” arises; this

is at first a two-winged crest (fig. 5), but further backwards acquires the true type of the

organ, as seen in the “Carinate” generally; it has nearly three coils (see fig. 2, ib).

The middle turbinal is merely represented by an irregular ridge on the fore face of the

antorbital plate, and the upper turbinal by the coiling, on itself, of the aliethmoidal

plates, exactly as in the Common Fowl (Phil. Trans.).

- The septum nasi is connected by an uncleft isthmus of cartilage (fig. 4, sv) with the

perpendicular ethmoid; it is unusually deep and large for a bird (figs. 4-6). There is

one septal fenestra (sf) in addition to the “notch” between the septum and the ethmoid ;

this window is situate close behind the insertion of the alinasal bag. The hinder inser-

tion of that bag is much thickened, and at this part the nasal branch of the orbito-nasal

nerve (fig. 4, nm) bores its way through the cartilage, piercing the alinasal, and bur-

rowing in the septal cartilages; it soon gets below the septum (see figs. 2, 4, 5). Now

if these sections be compared with the palatal view (Plate XXIV. fig. 4), and these by the

reader with an adult skull before him, it will be easy to comprehend the compact and

complex desmognathism of this type, a type having the most highly ossified and, in many

AND DEVELOPMENT OF THE BIRD’S SKULL. 135

respects, the most specialized skull to be seen amongst the Raptores. The broad foot of

the vomer (Plate XXV. fig. 5, v) fits like a keystone between the large maxillo-palatine

slabs (map); and these are bound, externally, by the dentary plate and angle of the pre-

maxillaries (dpx). In front of the vomer (Plate XXIV. fig. 4) the maxillo-palatine plates

meet, and afterwards coalesce. Ultimately, when all these structures, without and

within, have coalesced by extended ossification, all but the vomerine undersetter, then

the lateral halves of the face are indissolubly bound together, and form a structure most

compact and strong.

On the Structure of the Skull in the Sparrow-Hawk (Accipiter nisus)*.

In greater feebleness of general structure the skull of this type is intermediate between

that of the Falcon and the Cariama; but the leanings of the Hawk are not so much

towards the latter as in the direction of that gigantic Vultwrine Harrier the Secretary

_ Bird (Gypogeranus). The skull of a partially fledged nestling yielded the results given

in Plate XXV. figs. 7&8. Here we see at once things that agree better with the skull

of the Cariama, and also things that are less in conformity with it than those just

described in the Falcon: whilst the Cariama agrees with the Falcons, Hagles, and the

more Aquiline types of Vultures in having the basipterygoids early arrested, the

Sparrow-Hawk shows them as strong short spurs in the young (Plate XXV. fig. 7, bpg),

and retains them as mammillary elevations in the adult. They are still more developéd

in the Harrier (Circus), more still in the Brahmany Kite (Haliastur indus), and perfectly

functional in the Secretary. In Accipiter the parasphenoid (pas) is stout, and bevelled

on each side by the under-gliding pterygo-palatines. The next median splint, the vomer

(v), is truly azygous, and is a long knife-blade at present; in the young it is neither bi-

dentate behind nor pedate in front; it alters but little afterwards, and agrees in this

respect with the whole Accipitrine division of the group.

The desmognathism is perfect, although, in the young bird, it is “ imperfectly direct,”

as in the adult Cariama. Already the septum nasi is fast fusing with the solid fore end

of the premaxillaries; and, behind, its nerve-bridges and connexion with the ale nasi

make it a fit wedge to fill up the space in front between the large lunate maxillo-

palatines (mxp), which, lying back to back, of necessity leave an open space in front for

the septum, and behind for the vomer. The premaxillaries form a less solid mass than

in the Falcon ; but this type is much more orthognathous than the Cariama. Whilst the

Hawk agrees with the gallinaceous types in having a truly azygous vomer, it differs from

them, and is in harmony with the huge groups of the Passerines and Pluvialines, in having

a large mesopterygoid wedge differentiated from the fore end of the proper pterygoid (figs.

7 & 8, mspg, pg). As usual, the transpalatine (¢pa) is rounded off, more developed than

in the Pluvialine types, but less than in the Passerines. The ethmo-palatine plates (epa)

are thin, curved, pointed laminz, embracing the head of the vomer; the interpalatine

ridge (ipa) has no free spur. ‘he prepalatine bars (prpa) reach as far forwards, or

nearly, as the trabecular axis of the premaxillaries ; they are thin laths of bone.

' * For further details on this type of skull, see M. Micr. Journ. 1873, plates v. & vi. pp. 45-50. In these plates.

for ep read op.

136 MR. W. K. PARKER ON THE STRUCTURE

Behind we see that the large mesopterygoids (mpg) appear at the mid line, between

the postpalatine processes (ptya). Their general shape, as sharp wedges, is seen in the

side view (fig. 8). The pterygoids (pg) differ but little from those of the Cariama, being

short, flattened out on the inside in front; whilst behind they send up a very rudi-

mentary epipterygoid plate (epg). The extent of the median part of the maxillaries, the

maxillo-palatine plates (map), is very large, equal indeed, fore and aft, to what is seen in

an ordinary mammal. They are spongy, have a somewhat lunate or, rather, ear-shaped

form, and their hinder part forms a large free lobe, curving outwards.

In addition to the jugal process of the maxillary, there are two slender styles on each

side, a jugal (j) and a quadrato-jugal (qj), and not merely one bone, as in the Falcon.

On the Structure of the Face in Buzzards, Kites, and Harriers.

Several subfamilies of the lesser rapacious birds range themselves under the Accipitrine

type as a distinct form, and inferior to the Falconine. Their cranio-facial structures

agree very closely with what is seen in the Eagles and the Aquiline Vultures.

The condition of the facial parts in a recently fledged Common Buzzard (Buteo vul-

garis) is very instructive, and is representative of what may be found in a large number

of Old-World Rapaces, excluding, of course, the culminating forms, the Falcons.

A large interorbital fenestra (Plate XXIV. fig. 9, tof) separates the ethmo-presphe-

noidal bar above from the ethmo-basisphenoidal bar below (pe, bs), and the flat roof of

the great ethmoid, covered by the frontals and nasals (f,7), is rapidly becoming bony

from the huge median centre (pe). This roof sends backwards the normally ornithic

spike over the olfactory nerve (1), which grooves the bony plate on each side. The

coiled aliethmoid (ale), the representative of the upper turbinal mass of the mammal,

is thus partly ossified.

The flat antorbital plate, or pars plana (pp), which represents the middle turbinal,

is already ossified, and has coalesced with the perpendicular plate or meso-ethmoid (pe) ;

it is roughly pentagonal in form.

But this orbito-nasal partition is a downward continuation of the ala growing from

the roof, the aliethmoid‘(ale); the two are connected by a narrowish band of carti-

lage: this band is separately ossified, forming an “upper ectethmoid” bone, as in the

Passerinze (Oorvus). It lies under the roof, however, and does not stand out as a

prefrontal mass, as in the Crow and its congeners.

- The nasal nerve runs into the nose from the orbit, inside this bar, skirting the upper

turbinal, and then sends most of its substance forwards and downwards (see dotted

line in the figure) to the base of the septum nasi.

The notch which divides the true olfactory region from the septum nasi (sz) is very

large, runs obliquely forwards and upwards, and ends a little distance from the top;

hence this cranio-facial cleft is imperfect. The septum itself, only partially ossified as

yet, is differentiated into a nasal or trabecular, and an upper or true alinasal region,

by a large, oval, internasal fenestra (inf). Below and in front of this space the septum

is ossified as one large bone (/so), whilst the upper part is soft, except at the roof,

behind; there is at that part an upper septo-nasal bone (uso). The large lower bone

AND DEVELOPMENT OF THE BIRD’S SKULL. 137

is burrowed by the nasal nerve; and above and in front of this burrowing there is a

transverse septal knob, where the inturned alinasal fold joins on to the septum. On

each side the large, spongy, pneumatic maxillo-palatines (map) flank the septum, and

have the fore end of the knife-shaped vomer (v) between them. The palatines (pa), the

mesopterygoids (mspg), and the pterygoids (pg) conform very closely to those of the

Sparrow-Hawk.

In the Harrier (Circus cyaneus) and in the Brahmany Kite (Halastur indus) I find

a separate, thin, “median septo-maxillary,” attached, vertically, to the upper edge of

the vomer. It is very possible that this state of things may be merely the appearance

of a second vomer; I am not certain on this point; parts primarily symmetrical do at

times become azygous or median by the overgrowth of one and the arrest of the other,

the stronger bone gaining the mid line, and dragging the feebler bone after it.

These feebler types have, as a rule, a flat superorbital on the free end of the long

superorbital process of the lacrymal. In the Falcon the ossification is continuous

along their frowning brow.

The reader is referred to Prof. Huxley’s “Classification of Birds” (Proc. Zool. Soc.

1867, p. 442, fig. 24) for the palatal view of the Secretary’s skull: that figure is defi-

cient ; it should show a small knife-shaped vomer, like that of a Fowl. The whole skull

of that bird comes very near to that of the Brahmany Kite (Haliastur mdus) and to

that of the Egyptian Kite (Zlanus ceruleus). Its general skeleton is hugely modified

on account of its stilted Crane-like stature; legs of any indifferency would have asked

for none of those specializations of shoulder and hip-girdle which we see in that type.

On the Palatine Bones in Eagles and Vultures.

I shall merely note a few things to be seen in these large groups of the gigantic

Raptores.

In the Bateleur Eagle (Helotarsus ecaudatus) there is a small median palatine (Plate

XXV. fig. 9, mspg) between the ethmo-palatine lamine.

In Neophron percnopterus I find a delicate knife-shaped vomer (Plate X XV. figs. 14,

15, v), the right. fork of which is evidently aborted by a small mesopterygoid (mspg);

and besides this, on the same side, there is another granule of bone, belonging to the

same region (fig. 16, mspg). At the end of vomer, thinned out, however, in front, there

is a small medio-septomaxillary (msmz).

In Gyps fulvus there is an unlooked-for amount of differentiation of the palatine arch ;

for even in old birds a sharp sutural line runs obliquely two thirds across, almost sepa-

rating the transpalatine (Plate X XV. figs. 17, 18, ¢pa) from the rest of the bar. Here also

I find a strong wedge-shaped mesopterygoid (mspg) permanently distinct.

For the skull of Cathartes aura the reader is referred to Prof. Huxley’s paper (op.

cit. p. 440, fig. 22); it is an instance of “indirect desmognathism,” the maxillo-pala-

tines not being developed more mesiad than in Plovers and Pigeons; but the bond

being formed of an ossified tract of the septum nasi and inturned alinasal fold, this

forms a bony bridge similar to that seen in the Cariama; but there it is the anterior

138 MR. W. K. PARKER ON THE STRUCTURE

part of the alinasal fold. These South-American types agree together in the very

limited ossification of the nasal sacs, so different from the Old-World Rapaces.

In the King Vulture (Sarcorhamphus papa) the palate agrees with that of its con-

gener Oathartes (Plate XXYV. fig. 19); but I find a small medio-palatine (mpa), and

also much larger and more functional basipterygoids (pg). In this bird the Eustachian

passages (ev) open out into a very patulous, triangular fossa; and all the basicranial

structures are very strongly soldered together. The basitemporals (6¢) have also a

large mammillate knob on each side for muscular attachment.

On the Structure of the Skull in the Owls (Strigidee).

The Owls are the most distinct group of the Rapaces, and in respect of their basis

cranii come very naturally after the Cathartide. They agree with the Cariama in their

digestive organs, and, what is also quite unaccountable, with the Pigeons, in having a

curious deficiency in the occipital arch, a hole, like an artificial pin-hole, being left unos-

sified, as it was formerly non-chondrified, over the great cranio-spinal opening. ‘The study

of the skull of the Owls in nestlings and young fledglings is of great interest, these ob-

servations being made before the skull has become one light, spongy, undivided mass.

In a fledgling of the Barn-Owl (Aluco flammeus) the details can be well made out

(Plate XXIV. fig. 5), and the likeness and unlikeness of the type to that of the highest,

the Falcon (fig. 4), and the lowest, the Cariama (fig. 3), is clearly seen.

The relation of the pterygoids to the basipterygoid processes (pg, dpg) is quite like what

is seen in the King Vulture; but the parasphenoid (pas) is very swollen and is early fused

with a very thick and spongy interorbital septum. Not only are the outworks of the

ear of larger extent, but the tympanic cavity has large “anterior recesses” and great

upper chambers, corresponding to the open gallery that connects the right and left

chamber in the Crocodiles. The Barn-Owl has a chain of small tympanic ossicles, six

on each side, running along the inferior surface of the junction of the “lower otic pro-

cess”’ of the quadrate and the wing of the prootic, to which it is articulated.

This lower otic process is as large relatively, and as divergent inwards from the

“upper otic process,” as in the Frog; it turns, however, more backwards than in the

Batrachian.

The orbital process, or blunt-pointed free pedicle of the mandibular suspensorium (qua-

drate), is far severed from the trabecular base, and turns forward as well as inwards. ‘The

rest of the mandibular arch is feeble; its united halves form an acute angle in Aluco

flammeus, and a much wider angle in Asio and Striv. Soitis with the palate, this part not

being nearly so outspread in Aluco flammeus as in the other types. Still in this species

the pterygoids (pg) diverge very much, and articulate with basipterygoid processes

(bpg) very far apart, on the massive parasphenoid (pas). The pterygoids are slender,

sigmoid, and cultrate; they are toothed in front, and have a small epipterygoid crest (epg)

behind. In this species the palatines (pa) are almost parallel, and they are very contrary

to most of the bones, being thin and lathy. In this species the palatines are as simple

as in the Common Fowl, the transpalatine process being suppressed, and the muscular

AND DEVELOPMENT OF THE BIRD’S SKULL. 139

grooves almost invisible. In older birds a small interpalatine spur can be seen, but

in the young it is not evident. The ethmo-palatines (fig. 7, epa) are very short, and

backwardly placed; they are high, looking obliquely forwards and upwards, and become

spongy; in front they send forwards a flat half-coiled horn of bone; this is lost as a

separate process in the adult, for it coalesces with the much enlarged vomer, each

process adding to the depth of its shelving sides, and leaving a large fossa above, which

opens into most exquisite air-cells.

The palatines are kept apart by a triple wedge, the vomer (v) below, and the early-

coalesced mesopterygoids (mspg) above. This structure can only be seen in very young

birds before they leave the nest. Now the vomer is a smallish spindle; afterwards it

is a large dagger-shaped bone, with a deeply sulcate upper edge.

The thin fibrous prepalatine bars (fig. 5, prpa) form rests for the huge, ear-shaped,

swollen maxillo-palatines (map); their tip articulates with the imside of the splintery

palatine process of the premaxillary (ppx). The dentary part of that bone (dpz) is not

strong, and the beak part is hollowed below. ‘The fore end of the ossified septum nasi

coalesces with the premaxillary, and thus an extensive bridge is formed over the anterior

palatine space and under the external nostrils, the same as, but much more extensive

than, that seen in the Cariama (fig. 3, al), where the ossification is arrested.

- The hollow part of the beak in front of the nostrils has a bony septum between its

air-cavities, formed by the premaxillaries themselves (px), and lying in front of and

below the true septum nasi.

A considerable chink, 35 of an inch wide, separates the maxillo-palatines (map) ;

so that, notwithstanding their great size, these birds are only “ indirectly desmognathous.”’

The septum nasi is thoroughly ossified, but not the outer ale. It is of great antero-

posterior extent.

The orbital septum is a thick cushion of soft bone, very much bevelled away below,

but running forwards at its upper part; this arises from the direction taken by the great

cranio-facial cleft, which runs forward as well as upward.

The weak diverging jugal bars are composed of three elements, as in Accipiter (mz,

J; 9j); both the premaxillary and the maxillary have an angular process.

But these three elements are not all that is to be seen in the jugal bar; for that addi-

tional bone which I have called the postmaxiilary is here present (Plate XXIV. fig. 8,

ptmx); it is a smallish oval grain of bone, and can be seen in young birds not long

escaped from the nest. In this bird also can be seen an additional bone behind the orbit,

the counterpart of the ichthyic postfrontal or ‘“ sphenotic.” The lacrymal (partly

shown in fig. 8, 7) is very large and spongy, and does not send out a superorbital process

as in the “ Diurne.”

In the Long-horned Owl (Aso ofus) the palatines are supplied with a large vomerine

keystone (Plate X XV. fig. 10, v). Here the inner part of the palatine arch may have had

a single medio-palatine, or a pair of mesopterygoids; but they are not distinct, although

the specimen was a bird of the first summer; these sutures are early lost in the Barn-

Owl; and the medio-palatine of the adult is often formed of two mesopterygoids in the

young. I suspect it is so in this case.

SECOND SERIES.—ZOOLOGY, VOL. I. U

140 ‘MR. W. K. PARKER ON THE STRUCTURE

The vomer (v) wedges in between the small ethmo-palatine laminze (epa); and above it

is a similar bone, the median septo-maxillary (msm): both the bones are similar in

shape; but the vomer lies behind the bone above it. In the large Ketwpa ceylonensis

there are two similar bones (Plate XXV. fig. 11, v, mpa); but they are much smaller,

and the vomer is in front of the other, which is a medio-palatine.

In Athene noctua these parts are all coalesced together. The palatine keystone, made

carinate by asmall vomer, just shows traces of its sutural connexion with the ethmo-pala-

tines. This bird has large superorbital processes to its frontals, not to its lacrymals.

In the Hooting-Owl (Strix stridula) these parts remain separate (Plate XXV. figs. 20,

21). There is a small vomer (v), and over it a similar bone, as in Asio otus; both of these

arrested bones are in front of the short ethmo-palatines (epa); but behind these there

is a veritable medio-palatine (fig. 20, mspg, fig. 21, mpa)*; and the antero-superior

ossicle (msma) has the same position as the small median bone which I have called

the “median septo-maxillary ””’ in the Harrier (Circus cyaneus).

This is the bone which may possibly be a rudiment of the vomer of the other side

(right or left); for in the “ Picidze,” certainly, I have seen submesial bones join the mid

line; and the stronger push the weaker out of place.

In none of the Rapaces have I seen any remains of the paired “ septo-maxillaries ’’ so

common in the Passerinz, where they are small (although so large in Ophidia and Lacer-

tilia). Asa correlate of this state of things, the labial cartilages appear to be wholly

suppressed in these birds.

In Stri2 stridula (Plate XXYV. fig. 20) the transpalatine region (tpa) is as well developed

as in the Diurnal types; altogether it has a most instructive skull, and one quite unri-

valled for its swollen, spongy lightness of osseous tissue.

We have, even in this short survey of so small a tract of their structure, seen that the

Rapacious birds have much in common, and that the least specialized, as the Cariama,

marvellously wedges itself in among the roots of the most specialized of all, even the

Falcons and the Owls.

On the Structure of the Skull in the Laughing- Gull (Gavia ridibunda).

Several years ago a friend supplied me with a goodly collection of the young of this

species, and I took an early opportunity of working them out.

One of the first things which struck me was the Pluvialine character of these pulli, and

this at once struck me as being completely in harmony with resemblances I had noted

in these birds to the common Lapwing (Vanellus cristatus).

I am not sure which side of the boundary between the Plovers and Gulls one elegant

type—the Pratincole (Glareola)—should be placed; nor whether any man knows the

exact proportions of the Plover and of the Tern it contains in its organization. The fact

that Gulls are neither “ Altrices,” proper, nor true “ Preecoces ”’ makes their relation to

the Pluvialine types the more instructive; for the amount of super-pluvialine metamor-

phosis is not great, although real and measurable.

* This bone has been lettered in two ways, by mistake ; both mpa and mspg are correct, as a sengle mesopterygoid

becomes a medio-palatine.

AND DEVELOPMENT OF THE BIRD’S SKULL. 141

My views as to these relationships were fairly pledged several years since (see “ Gal-

linaceous Birds and 'Tinamous,” Trans. Zool. Soc. vol. v. p. 150); and I was soon led to

see how many large Families of Birds were constructed according to the pattern of the

Plover*.

But this was only one among several nascent generalizations germinating in my head

at the time ; so that when Prof. Huxley’s new ideas were given out, they fell into a recep-

tive soil, so far as I was concerned.

In this, as in other branches of zoology and morphology, there has been for many

years a remarkable harmony in our views; and nothing more struck and delighted

me, in that coincidence, than my friend’s commendation of one of the large groups

that seemed to me to gather towards one point: I refer to the double “ Gallo-Anserine”

family.

That the embryology of both the Goose and Fowl] was marvellously similar, I had then

seen; but I was not aware how close the Palamedeas on one hand come to the

Curassows and mound-making Megapods on the other.

But that which concerns us most at present is the huge number of birds with open

palates (‘‘Schizognathe”’) that have either a filial relationship to the Plover, or are

cousins-german to it.

It thus comes to pass that when once the structure of a Plover’s skull is thoroughly

understood, it serves as a norma to which the skulls of whole groups of birds can be

brought for comparison.

This is true, not merely of the related Schizognathous birds, but also of those Grallee

which are desmognathous; for the likeness and the difference between the split skull

of a fowl and the downd skull of a Palamedea is very similar to the likeness and the

unlikeness of the skull of the Crane on the one hand, and of the Heron and Stork on

the other.

There is one thing which is apt to mislead the zoologist in the consideration of the

great group of “ Carinate” birds; they, with the “‘ Ratitz,” are unconsciously compared

with the whole of the Mammalia. At first view it might be thought that the Ratitze

equalled, as a group, the “non-placental”” mammals, and the Carinatz the placental ;

but the groups are not properly comparable; the Carinate birds, notwithstanding their

number and variety, could not be equalled with more than two or three at most of the

Mammalian Orders, and not with the whole herd of the Placentalia.

Those morphological modifications which lie at the root of the diverse specializations

of the Carinate birds take place, as we pass from type to type, in the gentlest manner; all

the types of skull may be seen to spring from a fundamental form, rich in Rhyncho-

saurian, and even in Batrachian characters. Such a skull the Ratitee all possess.

After analyzing a skull of this kind, and finding how it is built up, as it were, accord-

ing to old styles of skull-architecture, familiar to him who knows the lower or cold-

blooded types, the morphologist is brought to a stand; for here the lines of life diverge.

* This. was handled excellently by Prof. Huxley in his “ Classification of Birds” (Proc. Zool. Soc. 1867, pp. 426—

431 and 456-458).

142 MR. W. K. PARKER ON THE STRUCTURE |

Standing in the ways to see and to judge which is the right path for each Carinate

type, the worker finds it no “ far call” from the order-joining Tinamous to the Plovers on

one hand, and the Fowls on the other. j

The sucker which grows into the Passerine tree is not so clearly discernible in the

Tinamine stock—in such Tinamous as now exist; but just above where the fork is, the

Hemipods are unmistakable in their relation to the harsh-voiced Passerines of the southern

hemisphere.

Above, when the Tinamine branch forks amongst types of no great elevation, and in

which the Dromzeognathous skull has passed into the Schizognathous, we see the nobler

forms (“ Aves aérize”’) just superimposed upon and, as it were, growing from the lower

forms—the “‘ Preecoces.”

From above the Sand-Plover arose the Pigeon, in its older and now extinct forms;

and from above old types of the Gallinaceze the fundamental or stock-form of the Rapa-

cious bird sprung. The genus Dicholophus is but little changed from such a primitive

plunderer.

Skull of Gavia ridibunda. 1st stage.

There were five stages of the young of the Laughing-Gull amongst those examined by

me; to these are added the adult; the largest young were just ready for flight, the

smallest only about a week old.

After these had been worked out, I obtained half-ripe young of the Silvery Gull

(Larus argentatus) from the “ Lesser Orme’s Head,” near Llandudno ; but these, although

studied in various ways, have not been wholly worked out by me; their condition at that

early stage differs but little from embryos of the Common Fowl (see Phil. Trans. 1869,

plate Ixxxii., 3rd stage).

In the first stage the primordial skull (chondrocranium) is thoroughly formed; it has

also most of the osseous centres commenced, and some of them ankylosed to their neigh-

bours. The occipital region is very instructive ; the basiocciptal (Plate XX VI. fig. 2, bo)

is a long oval piece, enclosing the starved notochord, and separated by large cartilaginous ~

tracts from the lateral pieces, the exoccipitals (eo). These bones are very irregular

pentagons; they are creeping over the hinder part of the auditory capsule, the canals of

which are seen shining through the clear cartilage (figs. 1-3, hse, psc). The ex-

occipitals already half enclose the foramen magnum (fm) ; but a good space exists between

them and the double keystone piece, the superoccipital (so). This latter bone is com-

posed of two halves, which are only half-soldered together ; above they are quite distinct.

They now form a very elegant bony plate with many sides, some having a scooped and

others a rounded outline.

The archway of the foramen magnum is finished by the concave margin of the

coalesced part; above, behind, and below the parietals (p), the margin is a bifoil.

Between these margins the bone has an indrawn waist; on each side, at this part, there

is a non-cartilaginous space—a “fenestra” or “ fontanelle,” the lateral occipital fon-

tanelle so familiar to osteologists in Pluvialine and Anserine types (see in Vanellus

cristatus, Trans. Zool. Soc. vol. v. plate xxxvii. figs. 1, 2, & 4, J. 0. f).

AND DEVELOPMENT OF THE BIRD’S SKULL. 143

Outside these spaces the bone is ear-shaped, the substance of it spreading towards

the ear-capsule.

These capsules are very large, and being impacted into or, at least, grafted upon the

large occipital arch and wall, appear very considerably in the back of the head.

Ossification has appeared in them, and the largest bony piece, the prootic (pro), is seen

behind the quadrate (q) in the lateral view (fig. 1); the other centres will be described in

more advanced stages, but the opisthotic is already well developed, although much

smaller than the prootic; the epiotic has not yet appeared.

The common sphenoidal region is at this stage a part full of interest to the mor- |

phologist. The three divisions of the great winged parasphenoid, although ankylosed, are

very evident. They are the rostral portion in front (figs. 1 & 2, pas), and the basi-

temporal plates (d¢) behind. These three bones are setting up ossification in the

overlying chondrocranium, although developed at first as parostoses and quite inde-

pendent of it.

As in the Ichthyopsida, these primordial splints are very large in relation to the rest

of the skull. That this should be so, shows how little the Bird is related to the ordinary

Lacertian type of Reptile, where the only remnant (not always present) of the para-

sphenoid is the anterior part. On each side of the scooped (Eustachian) part of the

median bone there is a cartilaginous bud; this is the basipterygoid process (pg), a

typically Pluvialine structure.

_ Behind these knobs the bone spreads into large wings that enclose the “ anterior

tympanic recesses.” Beneath these imperfect trumpets lie the basitemporal bones (6¢) ;

they have nearly melted into each other at the mid line, and above they are being

soldered to the tympanic wings of the parasphenoid.

But the overlying cartilage around the pituitary body and beneath the optic nerve (2)

has become osseous directly from the bone beneath; this is the true basisphenoid (ds) ;

the square inturned large alisphenoids (qa/s) are also ossified.

A large bony leaf is seen in front of the orbital septum ; it is trifoliate behind, and does

not nearly reach the membranous window between the eyes (iof).

In the figure (fig. 1) the septal portion of the nasal labyrinth has been cut away (see

this part from below in fig. 7); this is at the cranio-facial “ notch,” and the part in front

is only supplied by the orbito-nasal branch of the fifth nerve, and not by the olfactory.

This bony plate is the perpendicular or mesethmoid (pe) ; its huge size has relation to

the abortive development of the anterior sphenoid, and its bony matter will not cease to

grow until it has reached that of the posterior sphenoid (4s) beneath the optic nerve (2).

Nothing but development could possibly throw the least light upon this anomaly ;

there are no orbito-sphenoidal alee, and the presphenoid is merely that bar of cartilage,

afterwards bone, which is seen bounding the posterior half of the interorbital fenestra

(tof) behind; it reaches neither the top nor the bottom of the interorbital chondro-

cranium.

The top of the chondrocranium finishes behind in a median projection, behind and

between the upper turbinal scrolls (aliethmoids); this is a rudiment of that cartilaginous

144 MR. W. K. PARKER’ ON THE STRUCTURE

cloth which is thrown over the brain-sac in the Salmon and Polypterus (see my memoir

on the “Salmon’s Skull,’ Phil. Trans. 1873, plate v. ¢.cv), the tegmen craniv.

The reason for the membranous space between the eyes is not primarily teleological.

Nature is not husbanding her cartilage as being poor in that substance; it is a secondary

cleft, tending to divide the overlying ethmoidal structures from the crest developed on

the coalesced trabecule.

The roofing bones are seen to be well developed, the frontals, parietals, and squamosals

(fp, sq); the latter are elegant subfalcate bones, running far along the postorbital part

of the frontals, and somewhat trilobate below, where they clamp the ear-capsule. The

columella (s¢) is ossified in its medio-stapedial region, and the quadrate also, not its pedicle

or orbital process; its otic facets and its double lower condyles are all soft as yet.

Tn this stage I have only brought into view the hinder facial parts ; the pterygoid (pg)

has now a long splint-like process which reaches to the corresponding crus of the vomer,

not here shown, but seen in fig. 12, v*.

Afterwards the absorbents fret a suture across the base of this spur, and convert it into

a separate mesopterygoid, which, however, is only distinct for a few weeks, for it soon

coalesces with the postero-superior edge of the palatine.

The relation of the jugal to the quadrato-jugal, and of the latter to the quadrate, ito

which its hooked end is inserted, is here shown (fig. 1, /, ¢/, q).

The long dentary, which has coalesced with its fellow at the “mentum,” has been

removed, but the proximal part of the free mandible is shown (fig. 4, outside; fig. 5,

inside).

Meckel’s cartilage (mz) is still large, and runs far forwards; its articular end, with its

short Pluvialine posterior and internal angular processes (pag, iag), is ossifying ; there is

an endosteal patch and a long ectosteal plate (ar).

I see no separate coronoid, a bone often indistinct in birds; but the splenial, angular,

and surangular (sp, ag, sag) are large and still free.

In these long-faced birds the glossal portions of the hyoid are more than their hypo-

hyals; they deserve the name here of cerato-hyals (fig. 6, chy); they are unossified and

confluent.

There is a common cartilaginous basibranchial element (40r), pointed and delicate

behind, and only the proximal part of the first branchial arch (thyro-hyals, brs) is

ossified ; the dorsal element is long, slender, and soft.

-The nasal cartilages have been cut away and turned on their dorsal face to show their

laminar outgrowths (fig. 7); the section was made close behind the hinge or notch (see

fig. 1), and a line in front of the meso-ethmoidal bony centre (pe).

Unlike what we see in Mammals, the turbinal that figures most largely here is that

belonging to the ale nasi. This alinasal turbinal (at)) is here a simple foliaceous

outgrowth (see it in section, fig. 8). Behind this, and rising above it, is the inferior tur-

binal (itd). This is a simple plate at its commencement; further back it is coiled upon

itself. In most birds this scroll has two coils.

* This is a primordial relation of the pterygoid and the vomer; it is found in the lowest Urodela, to say nothing

of the Rhynchosauria. See Huxley, Proc. Zool. Soc. 1874, pl. xxix., and Ginther, Phil. Trans. 1867, pl. i. fig. 2.

AND DEVELOPMENT OF THE BIRD’S SKULL. 145

Behind the inferior turbinal, and also above it, is the coil of the upper turbinal (wd),

which is merely the aliethmoidal region of the nasal roof turned in upon itself.

This is represented as cut through the middle in the inverted figure (fig. 7, wtb); this

and the front wall of the pars plana (see fig. 15), which has no definite middle turbinal on

it, form the proper olfactory territory.

Skull of Gavia ridibunda. 2nd stage.

A somewhat more advanced stage (Plate X XVI. fig. 11) is shown in section as far as

to the cranio-facial isthmus (cfc); this figure supplies the deficiency of the last in not

showing a sectional view ; and it also shows some advance in the osseous deposits.

This section has been madea little to the near side of the mid line, and shows the para-

sphenoid and basipterygoid and the whole interorbital septum uninjured. The back-

tilted large auditory capsule is seen to lie both on and im the occipital arch; for the bony

matter creeping over the junction of the anterior and posterior semicircular canals is not,

in the bird, a large separate epictic bone as in reptiles; a rudiment appears afterwards

in this bird as in some others; but at present the super- and exoccipital is largely walling

in the otic mass.

The main periotic centre, the prootic ( pro), is now a large oblong bone, and separated

by the unossified auditory crest from the alisphenoid (als), and by synchondrosis below

from the basisphenoid (0s) and the basioccipital (do). The auditory nerve, or portio

mollis, pierces it, as also the portio dura, or facial nerve. Outside these foramina the

bone is scooped for the superincumbent brain, and becomes a mere crest; the edge

itself is still soft.

A clear wedge of bone is seen between the prootic and the exocciptal (eo) ; this is the

opisthotic (op); in the specimen it was becoming ankylosed to the prootic, but I have

diagrammatically given it a sharp suture from observation of the first stage.

The large passage (8) between the exoccipital and the opisthotic is for the glosso-

pharyngeal and vagus nerves ; the anterior and posterior condyloid foramina (9) are seen

on the edge of the exoccipital.

A large unossified space still exists on the crown of the arch of the anterior semi-

circular canal, and also in the recess for the cerebellar flosculus below it. If the bony

matter from the superoccipital had been somewhat less, then this space would have cor-

responded exactly to the region occupied by the epiotic bone in the Reptiles; its rudi-

ment has not yet appeared in the Gull.

The lozenge-shaped basioccipital is nearing the side piece (40, eo); but there is a large

spheno-occipital synchondrosis, underfloored by the basitemporal plate (07).

From the great submucous bones, the parasphenoid proper, and the basitemporals

(pas, bt), an upgrowth of bone has taken place, metamorphosing the overlying chondro-

cranium.

This bone, the basisphenoid (4s), underprops the optic nerves at their exit, and

embraces the dipping pituitary body, which passes down, as in osseous Fishes, to lie on

the submucous bony tract ; in the chondrocranium the pituitary space is open.

Into this pouched space the internal carotid (éc) enters; behind this space the cerebral

146 MR. W. K. PARKER ON THE STRUCTURE

mass lies on a shelving wall, the postpituitary wall, a scooped and slanting tract of bone,

split at the mid line, from which the notochord has retreated. This fissure is the

remains of the posterior basicranial fontanelle of the Reptile.

The prepituitary region has the anterior clinoid wall reaching the bony alisphenoid

(als), and a pinched front portion, which forms the postero-inferior angle of the inter-

orbital plate, and which afterwards coalesces with the huge perpendicular ethmoid below

the presphenoid.

This great mesethmoidal plate has increased in size, and is growing into the frontal

region. The frontals themselves at present fail to wallin the space caused by the

abortion of the orbito-sphenoids in the chondrocranium; there is, therefore, a post-

orbital fontanelle (po).

The subfalcate squamosal helps the alisphenoid (sq, als) to finish the side walls; it

clamps the prootic, overlies the alisphenoid, reaches the frontal, and supplements the

parietal (p) *.

Skull of Gavia ridibunda. 3rd stage.

This stage, in pulli a few days older than the last, has been worked out principally to

display the exquisite architecture of the great skull-floor, including the occipital, tem-

poral, and posterior sphenoidal regions.

At this stage we see best this most curious piece of morphological mosaic, not quite

unlike the wall-work of the test of an Hehinus, but of a more complex type by far, and

yet having fewer parts as far as mere number goes.

Indeed two very distinct strata are condensed and calcified to produce these results ;

and this difference is indicated in the figures by the colouring of the inner stratum,

formed by ossification of the chondrocranium, and the outer, which is the inner stratum

of the submucous connective web, is left uncoloured.

The first thing that strikes the eye in the under view is the long bony dagger and the

deltoid bone which lies beneath and behind it. These divisions of the ichthyic para-

sphenoid are the rostrum and the basitemporals (Plate XX VII. figs. 1-3, pas, bt); on each

side the rostrum widens between the basipterygoid cartilages (dpg), which are lessening ;

at the mid line it is scooped correlative to the meeting together of the right and left

Eustachian tubes (ew), and behind it simulates the basitemporal wings, widening and

spreading.

_ These hinder wings form the “ anterior tympanic recess ”’ (atr, ppg) : they are long,

faleate, notched, and grooved plates of bone, and with the help of the underlying

deltoid plate, the coalesced basitemporals (02), they form a pair of very elegant bony

trumpets leading into the drum of the ear, and are a specialization of the first visceral

* One of the older antomists (Owen), not seeing the squamosal in the inside of the struthious skull, took one of

his remarkable intellectual leaps, and supposed that it was always absent from that part, a hard thing to find in any

of the ‘ Carinatee.” ,

This great theorist wanted the bone on the outside and further down for transcendental uses; it, however, refuses,

to leave its own supratemporal region, for it must be ready to graft itself on the auditory mass in all the hot-blooded

Vertebrata.

AND DEVELOPMENT OF THE BIRD’S SKULL. 147

cleft. The hinder angles of the basitemporal plate are greatly produced, forming horns

that underlie the exoccipitals, whilst the posterior concave edge has lying on it the fore

end of the basioccipital (40).

This latter bone has on each side, between it and the paired exoccipitals, a reentering

angle of soft cartilage in front; but behind the three meet in the very substance of

the transversely oval occipital condyle; here, in meeting, they form a suture like a

leech-bite.

For the most part the occipital arch is well ossified (Plate XXVII. figs. 1-4); but the

whole of the outer edge of this transversely crested structure is still unossified, the car-

tilage not merely existing on the tympanic wings of the exoccipitals, but also on the

lower half of the superoccipital ridge (¢eo, so). The original suture between the two

superoccipitals is still visible above; but the “ lateral occipital fontanelles ”’ are nearly

filled in by bone on each side the narrow waist of the median part of the keystone.

We note here already the partial obliteration of the two very important Pluvialine

characters in these Sea-Mew chicks, namely, the basipterygoids and these occipital

windows. Here specialization has taken place in two ways :—first, by arrest and absorp-

tion; and secondly, by the secondary covering in of a region not filled in by the

chondrocranium.

The next important point to be noticed in these chicks of the third stage is the recol-

lection, as it were, of the epiotics left out in the first growth of bony territories; they

are, however, very small, and broken up into two or even three subcentres (Plate XX VII.

fies. 2, 5, & 6, ep).

This is seen many a time in ornithic morphology ; the bones which get the first start

in growth are large, whilst those which lose it are small, feeble, and overshadowed.

The upper view of the skull-base (fig. 2) shows well the round, deep, pituitary cup,

pierced below by the internal carotids (ic); also the slit below the hinder pituitary

wall, which is the old gap caused by the retirement of the notochord. The elegant basi-

occipital lozenge somewhat wedges itself into the end of the large, long, compound

basisphenoid.

But perhaps the most notable bones of all are the prootics; immense are they if com-

pared with those of the mammal. They are scooped laterally for the brain, and have

a lunate notch in front, which by a similar notch in the alisphenoid (a/s) becomes the

foramen ovale (5).

The prootics tilt themselves back to such a degree that the great anterior semicircular

canal is thrown, at its junction with the posterior canal, into the fore edge of the super-

occipital (figs. 4 & 5, asc, so).

If this figure (2) be compared with the rest, it will be seen how neat is the carpentry

by which the great ethmo-trabecular plate rests in the grooved upper surface of the

parasphenoidal rostrum (pas).

There it stands as a fixture; but this plate, the great meso-ethmoid, is almost sawn

through from below in front of the rostrum, and its under beam is the light and loosely

braced yomer.

SECOND SERIES.—ZOOLOGY, VOL. I. x

148 MR. W. K. PARKER ON THE STRUCTURE

Skull of Gavia ridibunda. 4th stage.

A little older than the last, the young of this stage yielded me the perfect longitu-

dinally vertical section (Plate X XVI. fig. 12) now to be described: it runs close after

and further illustrates the third stage.

The first sight of this object, and indeed of its figure, suggests feathery lightness ; and

the whole sum of its specialization, as compared with what is seen in the lower classes of

Vertebrata, may be taken as typical of this group (the Carinatze), in which the Reptile

so marvellously culminates.

The actual brain-cavity only occupies one third of the entire length of the head; the

nasal region proper, where the olfactory nerves are distributed, is exactly in the middle;

the ear-capsules run into the hinder arch; and the eyes are fairly halfway between the

other sense-capsules.

The great mesethmoid (pe) has now reached the interorbital fenestra (tof) above,

and behind it the cartilage ends in a retral spike, and in a groove below this lies the

olfactory crus.

Then in front of that bony plate we see how the cranio-facial axis has been gnawed

away, and a main and a lesser notch are seen leaving still a thick cartilaginous isthmus

connecting the small septum nasi (sz) with the main ethmoid.

In front of the former plate there is a large open gap before we reach the short, solid,

hooked end of the premaxillary (pz).

The feeble maxillary (ma) is left im sitw, and its maxillo-palatine plate (map) is seen

rising obliquely and leaf-like, on each side of the cranio-facial notch; inside of it is the

forked and notched double vomer (v); the rest of the face has been removed.

Behind, we see what has been already described in the second stage (compare figs. 11

& 12), but with this difference, namely, that the bony territories are not hedged in with

cartilaginous balks, but have their margins edge to edge.

The prootic is still distinct; but the epiotic fragments are now lost in the coadapted

faces of the prootic and superoccipital ; also the opisthotic has coalesced with the pro-

otic and exoccipital.

The base of the cranial cavity is an ascending floor, concave below and behind, and

convex as it begins to close in in front: the brain sits on the rounded upper edge of the

presphenoid in front (Plate XXVI. figs. 16 & 17, ps).

Here the floor is the original membrano-cranium, the cartilage being stunted in this

part ; and any orbito-sphenoidal bones are merely such osseous centres as appear in the

fontanelle. Inthe Fowl there are two on each side, supplemented by the orbital plate of

the frontal. I see no evidence of any distinct centre here in this species; in the old

bird the bony matter merely runs a little way into the fontanelle from the presphenoid.

Skull of Gavia ridibunda. 5th stage.

In these ripe chicks the sutures are becoming obliterated fast, especially those of the

ossified chondrocranium; but there are still many clear divisional lines in the investing

or outer part of the skull.

AND DEVELOPMENT OF THE BIRD’S SKULL. 149

The occipito-otic region is now one large shell of bone (Plate XX VII. figs. 7-10); in

the hind region (fig. 10) the “ lateral occipital fontanelles ” (dof) are still open.

The basitemporal plate (fig. 9, df) has now obtained its perfect form; it is subtriangular,

but the sides are convex, and the base is emarginate, and it has ear-like processes at

its angles; these help to floor in the tympanic cavity. From the parasphenoidal rostrum

(pas) the basipterygoids have disappeared, so that the Pluvialine stage has been passed

in this part of the skull.

All the elements of the face are very slender and elastic, both those of the outer and

also those of the inner series. The great perpendicular ethmoid (pe) has reached the

“notch” in front and above, and hs nearly met the basisphenoid behind and below.

There are still large postorbital and a large interorbital fenestra ( pof, tof).

The nasal cartilages (the fore part of which had been removed in the specimen figured)

are all cartilaginous, with the exception of the back wall or pars plana (pp), which

is made bony by a centre of its own.

Outside this there is an anvil-shaped lacrymal, Pluvialine in form, but answering to

that of the larger types; its lower crus rests upon the outstretched antorbital plate

(Plate XXIV. fig. 7, 7, pp). The upper turbinal coil (aliethmoid) divides the groove for

the olfactory nerve from that for the trabecular or naso-orbital.

The nasal processes of the premaxillaries were laid upon ‘and wedged in between the

upper plate of the nasals (xpx,n). The upper plate of the nasals lies upon and is wedged

in between the frontals (7, f). The frontals are sinuously applied, behind, to the fore

margin of the parietals (f, p); this harmony-suture is the “coronal” of human anatomy.

A “sagittal suture” runs along the whole skull and face, from the fore end of the nasal

fossa, where the premaxillaries are soldered together, up to the notch in the upper surface

of the superoccipital (fig. 10, so), the remains of the divisional line of its two primary

halves.

The falcate squamosal (sq) elegantly binds the whole cranial cavity together on each

side; it runs upwards and downwards obliquely, behind, lying like a tile over the slant-

ing occipital edge.

The nasals, as is the wont of the Pluvialine types, are sharply split into two crura

(figs. 7 & 8, 7). These run forwards, tying down the upper and lower branches of the

premaxillaries (7, npx, dpa).

The pterygoids (pg) are long, slender, and inbent. They are pedate behind, having

an epipterygoid process, with an oval hollow above; and the proper end of the bone has

an acetabular facet ; both these embrace similar convexities on the fore margin of the

quadrate (q).

In front the pterygoid, having lost its mesopterygoid spike, which is now part of the

palatine, fits, by a short tooth, into a shallow socket in the end of the palatine (pa). In

front, the palatine is a long, slender needle of bone, which is now tied to, and afterwards

anchyloses with, similar needles, namely, the fore part of the maxillary (ma) and the

palatal process of the premaxillary (ppz).

The outer edge of the two-keeled hinder part of the palatine has no more outgrowth

as a proper transpalatine portion than the Plovers. (See my “ Gallinaceous Birds and

x 2

150 MR. W. K. PARKER ON THE STRUCTURE

Tinamous,” Trans. Zool. Soc. vol. v. plate xxxvii. figs. 1-5, for views of the skull of

Vanellus cristatus, useful for the whole comparison between the Larine and Pluvialine

skulls.)

The solid part of the premaxillaries is short. The maxillary widens at its middle, to

form the elegant spoon-shaped maxillo-palatine process (map), which is obliquely placed,

and has its “bowl” outside. The rest of that bone and the two pairs of jugals (7, ¢/)

are very slender spicules of bone.

The vomer (Plate XXVILI. figs. 9 & 9a, v) has the form so common in Pluvialine, Gruine,

Ralline, Alcine, and Ardeine types; it is formed of two lanceolate centres that unite for

their anterior two thirds by a sharpish angle. They become carinate below, and the

carina is, in the young bird (fig. 9a), divided into two tracts. These afterwards are joined

by fresh bone into one keel, which forks where the bone forks to form the crura that

unite with the under surface of that part of the palatine which was the distinct meso-

pterygoid bone.

The very pluvialine mandible (fig. 7, d, ar) has still a suture where the anterior and

posterior parts unite ; there is also an oval fenestra in front of the hinge.

Skull of Gavia ridibunda. 6th stage.

In the skull of the oldest specimen I have examined of this species the occipital

fenestree and the basipterygoids are entirely gone. The bone is light and thin, but

rather dense; scarcely a trace of suture is to be seen—for instance, only above the

cranio-facial hinge. The only bones free are the vomer, the quadrates, and the mandible.

The latter bone has its sutures between the dentary and the hinder part persistent.

The nasal capsules are only ossified in the olfactory region (aliethmoid and pars plana) ;

the vestibular cartilages remain soft, viz. the aliseptal with its inferior turbinal, and the

alinasal with its turbinal.

But there is a structure of intense interest attached to the outturned end of the pars

plana, and not differentiated from it in the cartilaginous state ; this is the “os uncinatum”’

(Plate XX VII. fig. 11, ow). This is a small triangular wedge of bone which rests upon

the zygoma. It is well shown in the Albatross (Diomedea). I have lately described it in

Dicholophus ; traces of it occur in Alcea torda and Uria troile; but its highest development

is found in certain arboreal birds—Plantain-eaters, Parrots, &c.

{his bone is the ethmo-palatine, or joining-piece between the trabecula and palatine

in front, and belongs to the same category as the basipterygoid.

On the roof of the skull large and elegant fosse exist, on which lie the long, tongue-

shaped supraorbital or nasal glands. In old birds the two long ceratohyals are only

soft at their apex or ventral extremity ; they ossify to a great extent, ankylose in the fore

part, then are somewhat bowed out apart, and then come close together in front of

the basihyal. This latter bone sends osseous matter into more than the front half of

the urohyal. The lower thyrohyals are ossified, The upper and more slender pieces

are half soft below and at the upper ends.

But little difference would be found between this bird in its development and the

AND DEVELOPMENT OF THE BIRD’S SKULL. 151

growing young of a species of Podiceps, Colymbus, Alca, or Uria; they all specialize

by passing just a stage or so beyond the simple Pluvialine pattern, but in different

degrees. The Petrels are close congeners of the Gulls; many of the smaller kinds

retain their basipterygoids. A large number of water-birds keep their “lateral occi-

pital fontanelles” open throughout life. (See, on the Pluvialine types, Huxley, Proc.

Zool. Soc. 1867, pp. 426-431.)

EO ot B® 09

DESCRIPTION OF THE PLATES.

PLATE XX.

. 1. Vertical section through head of embryo Fowl (Gallus domesticus), about the beginning of the

sixth day of incubation. x 7 diameters.

. The same, taken further back through the eyeballs and nasal sacs. x 7 diameters.

. The same, behind the nasal walls. x 7 diameters.

Part of same section. x 27 diameters.

. A section further back, through the eyes, pituitary space, and lower jaw. x 7 diameters.

. Section through ear-sacs of the same embryo. x 7 diameters.

. A longitudinally vertical section of the hind face of an embryo of the House-Martin (Chelidon

urbica). x 15 diameters.

ig. 8. Inner view of auditory structure and mandible in a fledgling Carrion-Crow (Corvus corone).

x 3 diameters.

. 9. Os quadratum and stapes of adult Jackdaw (Corvus monedula). x 6 diameters.

. 10. Stapes of Pipmg-Crow (Gymnorhina tibicen), inner view. x 10 diameters.

. 11. The same, edge view. x 10 diameters.

g. 12. Palatine view of skull of Starling (Sturnus vulgaris). x 4 diameters.

g. 13. A species of Cardinal (Cardinalis, ?sp.). x 54 diameters.

Prats X XI.

.1. Palate of embryo of Brown Linnet (Linota cannabina) of the sixth day of incubation.

x 15 diameters.

.2. Part of same. x 150 diameters.

. 3. Part of embryo of same bird, seventh day of incubation. » 20 diameters.

Fig.

4, Vomer and part of palatines of a fledgling Sparrow (Passer domesticus), seen from above.

x 44 diameters.

152 MR. W. K. PARKER ON THE STRUCTURE

Fig. 5. Palate of common Wren (Troglodytes vulgaris). x 51 diameters.

Fig. 6. Part of same. x 11 diameters.

Fig. 7. Side view of face of the Wren. x 54 diameters.

Fig. 8. Palate of adult Goatsucker (Caprimulgus europeus). x 24 diameters.

Fig. 9. Part of same, seen from above. x 44 diameters.

Fig. 10. Part of face of same bird, upper view. x 24 diameters.

Fig. 11. Part of palate of young Goatsucker, seen from above. x 4 diameters.

Puate XXII.

Fig. 1. Palate of a still younger Goatsucker (Caprimulgus europeus). x 4 diameters.

Fig. 2. Palate of Humming-bird (Patagona gigas). x 34 diameters.

Fig. 3. Part of same view. x 6% diameters.

Fig. 4. The same, seen obliquely. x 62 diameters.

Fig. 5. Cranium of same bird, seen from above. x 34 diameters.

Fig. 6. Part of same, side view. x 34 diameters.

Fig. 7. Hyoid bones of same bird. x 63 diameters.

Fig. 8. Palate of a nestling Humming-bird (? Lampornis) from Barbadoes. x 63 diameters.

Fig. 9. Part of same. x 10 diameters.

Fig. 10. Part of same, seen from above. x 10 diameters.

Prats XXIII.

Fig.

Fig. 10. Stapes of same, outer view. x 74 diameters.

. Palate of same, with lower plate removed, seen from below. x 24 diameters.

Fig. 1. Palate of Scythrops nove-hollandie. x 24 diameters.

Fig. 2. Section of palatine bones of the same bird. x 21 diameters.

Fig. 3. Side view of face of same bird. x 21 diameters.

Fig. 4. Palate of Megalema asiatica. x 3 diameters.

Fig. 5. Side view of face of same. x 3 diameters.

Fig. 6. Palate of Podargus. x 1z diameter.

Fig. 7. Side view of skull of same. x 17 diameter.

. Side view of palate of same. x 21 diameters.

Fig. 11. The same, inner view. x 7% diameters.

PuatTe XXIV.

Fig. 1. Skull of Gyps fulvus, from above. + nat. size.

Fig. 2. Skull of Dicholophus cristatus, from above. + nat. size.

Fig. 2a. Part of side view of the same skull. x 2 diameters.

Fig. 3. Palate of same. x 14 diameter.

Fig. 8a. Vomer of same. x 3 diameters.

Fig. 4. Palate of Falco tinnunculus (nestling). x 2 diameters.

Fig. 5. Palate of Aluco flammeus (fledgling). x 2 diameters.

Fig. 6. Part of same, from below. x 4 diameters.

. Part of same skull, side view. x 4 diameters.

. Skull of Buteo vulgaris (recently fledged), part of side view. x 13 diameter.

Fig.

Fig. 7. The same, from above. x 4 diameters.

Fig. 9

AND DEVELOPMENT OF THE BIRD’S SKULL. 153

Piatt XXV.

Fig. 1. Upper beak and nasal opening of Falco tinnunculus (nestling). x 8 diameters.

Fig. 2. The same in section, seen from the inside, and half the inferior turbinal showing. x 3 diameters.

Fig. 3. Nasal capsule of same, with bones removed, from below. x 3 diameters.

Fig. 4. The same in longitudinal section. x 3 diameters.

Fig. 5. Transverse section of same, through the beginning of the inferior turbinal. x 6 diameters.

Fig. 6. Transverse section of same, through alz nasi. x 6 diameters.

Fig. 7. Palate of Accipiter nisus (fledgling). x 4 diameters.

Fig. 8. Part of same, side view. x 4 diameters.

Fig. 9. Part of palate of Helotarsus ecaudatus. Nat. size.

Fig. 10. Part of palate of Asio otus, lower view. x 3 diameters.

Fig. 11. Similar figure of Ketupa ceylonensis. x 2 diameters.

Fig. 12. Vomer of Falco peregrinus (?), side view. x 3 diameters.

Fig. 13. The same, under view. x 38 diameters.

Fig. 14. Vomer of Neophron percnopterus, from below. x 38 diameters.

Fig. 15. The same, side view. x 3 diameters.

Fig. 16. Pterygoid of same bird. x 38 diameters.

Fig. 17. Part of palatine of Gyps fulvus, upper view. Nat. size.

Fig. 18. The same, from below. Nat. size.

Fig. 19. Palate (part) of Sarcorhamphus papa, from below. Nat. size.

Fig: 20. The same in Sfria stridula. x 2 diameters.

Fig. 21. Part of same, oblique view. x 4 diameters.

Note.—The letters mspg in fig. 20 and mpa in fig. 21 refer to the same bone; fig. 21 is inverted (by mistake).

Prate X XVI.

Fig. 1. Skull of Gavia ridibunda (first nestling), side view, partial. x 3 diameters.

Fig. 2. Same, from below. x 3 diameters.

Fig. 3. End view of same. x 3 diameters.

Fig. 4. Hinder part of mandible of same, outer view. x 3 diameters.

Fig. 5. Same, from within. x 3 diameters.

Fig. 6. Hyoid of same specimen. x 3 diameters.

Fig. 7. Nasal labyrimth of same. x 5 diameters.

Fig. 8. Transverse section of nasal vestibule of same. x 20 diameters.

Fig. 9. A like section, further back. x 20 diameters.

Fig. 10. Another section still further back. x 20 diameters.

Fig. 11. A longitudinal view of the inside of the skull of second nestling (partial). x 3 diameters.

Fig. 12. A similar (perfect) section of the skull of fourth nestling.

Fig. 13. Section of alee nasi of fourth nestling. x 20 diameters.

Fig. 14. Section through ethmoid of fifth nestling. x 5 diameters.

Fig. 15. Section near “ notch” of ethmoid of fourth nestling. x 5 diameters.

Fig. 16. Another section of same, through interorbital septum. x 5 diameters.

Fig. 17. A like section, behind the fenestra. x 5 diameters.

154 MR. W. K. PARKER ON THE STRUCTURE OF THE BIRD’S SKULL.

Puate XXVILI.

Fig. 1. Basis cranii of third nestling, from below. x 3 diameters.

Fig. la. Alisphenoid of third nestling. x 3 diameters.

Fig. 2. The same basis cranii, from above. x 3 diameters.

Fig. 8. Side view of same. x 3 diameters.

Fig. 4. End view of same. x 8 diameters.

Fig. 5. Inner view of auditory region of the same. x 6 diameters.

Fig. 6. Inner view of the other side. x 6 diameters.

7. Side view of skull of fifth nestlmg. x 1}? diameter.

. Upper view of same. x 14 diameter.

Fig.

Vig.

Fig. 9. Lower view of same. x 1? diameter.

Fig. 9a. Vomer of same, side view. x 4 diameters.

Fig. 10. End view of same skull. x 12 diameter.

Fig. 11. Part of side view of skull of the adult bird. x 6 diameters.

Trans. Linn. Soc. Ser 2. Zoont.Vou.l Tas 20.

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THE

TRANSACTIONS

THE LINNEAN SOCIETY

LONDON.

SECOND SERIES.—ZOOLOGY.

VOLUME I.

PART THE FOURTH.

tO Ne DeOrN:

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET :

SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON-HOUSE;

AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER- ROW.

- M.DCCC.LXXVII.

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CONTENTS.

PART IV.—Janvuary, 1877.

VIII. A Monograph of the Lepidopterous Genus Castnia and some allied Groups. By

JRO WESEWOODS MOAN, Pens aw SMe 5) 2 om ee pare dlns

IX. On a New Genus of Trematoda, and some new or little-known Parasitic Hirudinei.

By Joan Dents Macponatp, W.D., F.R.S., Deputy Inspector-General R.N.,

Professor of Naval Hygiene Army Medical School. (Communicated by G. EH.

DOBSON UMCA. SIE Beet Sa caustiod Tron Wa, Pees series hen, 209

X. On the Osteology and Pterylosis of the Spoon-billed Sandpiper (Eurynorhynchus

pygmeus, Linn.). By Jonn ANDERSON, ID., F.R.S.H., FLS., P.ZS., §¢.,

Curator of Indian Museum, and Prof. Comp. Anatomy at Medical College,

(Cian, sh ccm “ese We cpa: gh ble Rete Reis iyi ghey tala decal ci ae NT

PP etnat eo

Tain! a Ho See

NT I meee cc Cam

ah BAMo'VvO Waits Anion, Saw pinden’) wavy ayoradqobiqars oh Xp sage

ach aseg DRONE 2) 50! cq 1c po Ae I A Rs gs Joona

: F \ i ‘

| ianibnit toners winorkattil 0 osm amos baw hone Yami awl

* GAB Any d -eolosaeeh ysge 2 AL ML ,ctenoagell ava caOk Wi,

A yh GelnaioonmeteO ) ~ Joos Londo gare, QnSi YEE Ioan, Yo wweadhor a

} GOk, ms ¥ Pier ag. i fe) se a A ae aie 5 te (Gh i ohooh weoenokh,

- ado tomrstik) setabane holid-coog’ oY Ap eine Id huey yooleaied say

Lah BSN 2AM AT AG woonseck aah yh” CW atrotarg yay

AYN Ihibskl Io ymoinak .qudy Apvk Gan see svibal Wyo solo

Mec ae on) ne i erage acs, Se

[ 155 ]

VIII. A Monograph of the Lepidopterous Genus Castnia and some allied Groups.

By J. O. Westwoop, JZA., P.LS., Sc.

(Plates XXVIII.-XXXIIL.)

Read June 17th, 1875.

NOTWITHSTANDING the vast additions which have been made during the last

quarter of a century to the lists of species of Lepidopterous insects, especially by the

entomologists of our own country, the observation of Latreille, in his most valuable and

classical work the ‘Genera Crustaceorum’ &c., vol. iv. p. 186 ( Lepidopterorum ordo

entomologorum scopulus: horum insectorum etenim instrumenta cibaria simplicia; an-

tennee pro sexu diverse; metamorphoses permultorum nobis ignotz; idcirco nepotes

nostri methodum optimam soli conficient”’), is almost as true now as it was when

written, seventy years ago. In fact, the circumstance of such great numbers of species,

for the most part of exotic origin, being added to the already numerous tribes of Butter-

flies and Moths, without, in the greater number of cases, any actual knowledge of their

transformations or special structure being furnished by the captors or describers, has only

tended to render that “still darker which was dark enough before.” If, at the same

time, we turn to the generic distribution of these insects, we are bewildered by the

infinite number of new groups which have, as it appears to me, most unnecessarily been

introduced into the science, and the never-ending change in the employment of old

generic names. ‘This has in a great degree resulted from an insufficient appreciation of

the relative value of the characters which constitute a species and the illogical raising

of such characters to a generic rank. Thus no sooner had I published a monograph of

the Australian species of sac-bearing Moths, for which I adopted the generic name of

Oiketicus, proposed by the Rev. Lansdown Guilding, in the ‘Transactions of the Lin-

nean Society,’ in which paper I described the varying structure of the antennz, wing-

veins, and cases formed by the larve, than my specific details were seized upon, and each

species erected into a genus, characterized from my figures. Again, we have seen in a

recent American publication some of the Swallow-tailed Butterflies, which had uni-

versally been considered a species of the old genus Papilio, separated off from the

rest, and formed into different genera; whilst the old generic name Papilio has been

entirely removed from the family, and applied to one of the genera, or rather to one of

the species of the genus Vanessa—each of the other species of the latter group being also

raised to generic rank, whilst almost every species of North-American Hesperiide has

been formed into a separate genus. If, again, we look at the general classification of the

Order in the hope of supplying the desideratum of a ‘“ methodus optima,’ to which

Latreille especially alluded in the paragraph quoted above, I fear we are but little further

advaniced towards the attainment of that end than our forefathers. By the employment

SECOND SERIES. —ZOOLOGY, VOL, I. : ¥

156 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

of the transformations, and especially by the structure of the veins of the wings, feet,

antenne, palpi, tongue, and other characters of the imago, we have arrived at certain

results as to the characters of certain groups which had formerly been comparatively only

partially examined ; yet the want of a general revision of all the groups of the Order in

the same philosophical manner, has still left us in doubt as to the proper clue to the clas-

sification and relations of the families and still higher primary divisions of the Order.

I do not hesitate to say that the attempts to characterize and systematize the families

and subfamilies of Nocturnal Lepidoptera hitherto made are total failures.

Another difficulty, unknown to the older writers upon this order of insects, has arisen

from the exploration of numerous adjacent localities, which, whilst it has added greatly

to our knowledge of new and quite distinct species, has also shown that the wide geo-

eraphical range of a species is often attended with the development of slightly modified

races, which have by some writers been indifferently regarded as distinct species, or have

been sunk to the rank of local varieties. Thus, of the gigantic types of the diurnal

Lepidoptera which, from their size, have been well named Ornithopterus, and which are

natives of the East, we find the single species, Papilio Priamus, in Mr. Kirby’s Cata-

logue, made to comprise not fewer than seventeen of these local forms. These have been

specifically named, and regarded by others as distinct species, namely :—1. P. Pri-

amus, Linn.; 2. Panthous, Linn.; 3. Richmondia, G. R. Gray; 4. Cassandra, Scott;

5. Euphorion, G. R. Gray ; 6. Pronomus, G. R. Gray; 7. Poseidon, Doubleday ; 8. Cronius,

Felder; 9. Boisdwalii, Montrouzier; 10. Oceanus, Felder; 11. Arruana, Felder ;

12. Urvilliana, Guérin; 13. Triton, Felder; 14. Pegasus, Felder; 15. Archideus, Felder ;

16. Lydias, Felder; 17. Craesus, Wallace.

Another series of Hastern Butterflies, of which Papilio Paris is the type, distinguished

by having their wings powdered with golden-green atoms, exhibits a number of permanent

variations of no higher specific rank than those of the Priamus group in their respective

localities ; yet we find each of them, in Mr. Kirby’s Catalogue, given as a distinct

species.

The solution of this difficulty can only be hoped for by the very careful investigation

of such Protean species in their native haunts, with reference to their appearance in the

adjacent and intervening localities, so as, if possible, to determine how far the specific

type becomes modified (at least, so far as the markings of the wings are concerned) by

change of situation and other local circumstances. And here it is to be observed that

possibly too much weight has been eiven to the mere markings of the wings, without

sufficient attention to the question whether they are accompanied by structural differ-

ences. It was, for instance, with considerable doubt that I at first regarded several of

the Heliconiiform species of Oastnia described below as specifically distinct; the inves-

tigation of their wing-veins, however, satisfactorily proved that they were structurally

different from each other. But paradoxical as it may at first appear, the publication

of descriptions of Butterflies and Moths (unless belonging to well-defined and previously

characterized genera) has probably become the greatest obstacle in the way to our know-

ledge of the real “ methodus optima” desiderated by Latreille. Ido not hesitate to

affirm that the publication of the hundreds, nay, even thousands of careless and insuffi-

GENUS CASTNIA AND SOME ALLIED GROUPS. 157

cient descriptions, especially of Moths, in this country during the last twenty years, will

embarrass and disgust the student to such an extent as to retard the inquiry into the

general distribution of the Order, as well as the relations of the various families inter se,

for years to come. Without going to the extreme length which Mr. Scudder has done

in extending the characters of every genus over four or five closely printed pages (em-

bodying every point of the structure of the type in every stage of its existence, thereby

converting specific characters into generic ones), I yet hold that the description of a

Lepidopterous insect, without a thorough examination of its structural characters, is not

only detrimental to the science, but disgraceful to the author*.

In like manner the entire alteration in the old and well-established systems of celassi-

fication, both of the Rhopalocerous and Heterocerous Lepidoptera, and the substitution in

their stead of other systems which have only the charm of novelty and the love of change

to support them, is, as it appears to me, detrimental to the real advance of science. I

see, for instance, no reason or even advantage in removing the six-legged Papilionidee

from the head of the order, and substituting in their stead the Nymphalide, with their

imperfect fore feet, advocated by the German writers, and servilely adopted by their

English followers ; neither can I approve of the displacement of the gigantic Bombycide

(Saturnia, &c.) from the head of the Nocturnal Lepidoptera.

We thus perceive that, with respect to the discrimination of many species, or the limita-

tions of genera and families, or the general distribution of the primary groups of the

Order, we are scarcely further advanced than we were in the days of Latreille. The

genus Castnia of Fabricius affords an illustration of the various difficulties to which I

have above alluded. By Linnzeus and all the writers of the last century the species of

this interesting genus with which they were acquainted were regarded as Butterflies,

being arranged under the genus Papilio, on account of the clubbed structure of the

antennze. On the dismemberment of that great group, at the commencement of the

present century, by Fabricius and Latreille, the genus Castnia was established; but it

was still retained by the former as the 7th genus of the Butterflies, between the genera

Cethosia and Huplea, characterized only by the structure of the palpi and antenne + ;

whilst Latreille, in all his works, placed it (with Agarista &c.) at the head of the

Crepuscularia, between the Hesperiidee (with which the Uranie were united) and the

Sphingidee.

The connexion of these insects with the Diurnal Lepidoptera rests only on the structure

of their antenne, and on their evidently diurnal habits evinced by the brilliancy of their

colours. But when the antenne are carefully examined they do not bear out this con-

* It would. in this place, indeed be unjust not to refer to the admirable manner in which Mr. Stainton has worked

out the economy and structure of the small and difficult groups of the Linnean genus Tinea, in his different works on

the Microlepidoptera, whereby he has laid down the groundwork of a classification of these beautiful tribes, which

had previously been enveloped in doubt and difficulty. His figures also of the veining of the wings of these small

and difficult insects, as well as those given by the late Dr. Herrich-Schiffer in his ‘Systematische Bearbeitung der

Sehmetterlinge von Europa,’ will greatly assist the student in his investigation of the relations of these insects.

t System. Glossat. (Hd. Lliger, Mag. Ins. yi. 277, 1822). In Mr. Children’s abstract of this system, given in Taylor’s

Phil, Mag. for 1830, the generic name Castnia is misprinied Castina.

Y 2

158 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

nexion, differing materially from the real Rhopalocerous structure; whilst their con-

nexion with the Hesperiide is more remote than with any other group of the Diurna,

in consequence of the simple arrangement of the veins of the wings in the Hesperiidz.

The complicated arrangement of the wing-veins in Castnia, and the metallic gloss of the

very large scales (which are much larger than in any other Lepidopterous genus) with

which most of them are clothed, and the rich coloration of nearly all the species, in like

manner separate them from all the other groups of Heterocerous Lepidoptera.

The large and robust structure of the Castnia, together with their very ample wings,

separate them from all the groups to which they are assimilated by their transforma-

tion, except the gigantic Hepiali and Cossi, from which they are at once distinguished

by the veining of the wings and their clavate antenne.

In the complicated arrangement of the wing-veins, and especially in the existence of

the minute supplemental postcostal cell, we find an approximation to the Geometride ;

the details of so many species of which are illustrated by Herrich-Schaffer. But the weak

structure of these moths and their geometric larvee, destitute of several pairs of the

ventral pro-legs, will not allow any approximation between the Castnie and Geometre.

Another curious circumstance connected with the group, and which is not without its

significance in respect to their general relation, is the analogical resemblance exhibited

by some of the species to that strange family of Butterflies, the Heliconiide ; to these latter

we find similar resemblances exhibited by species, not only of other Heterocera, as

among the Arctiidee, but also amongst the most typical of all the butterflies forming the

family Papilionidee, as well as amongst the Pierides and Hrycinides.

Hence with regard to the actual relations of the Castnie, in the uncertainty which

still exists as to the preponderance to be given to the characters derived either from the

immature or perfect state of the articulated animals, we can arrive at no other con-

clusion than that which Dr. Boisduval has given in his last work, ‘ Hist. Nat. des Ins.

Lepidopt. hétérocéres :’'—‘‘ La famille des Castnides appelées Hesperi-sphinx par lillustre

Latreille, quoiqu’elles n’aient pas plus de rapports avec les Hesperies qu’avec les

Sphinx. Ce sont des insectes d’une création 4 part, qui sous leurs premiers états, ont a

la fois les habitudes des Sésiides et des Zeuzérides, mais auxquels il est difficile d’assigner

une place” (Pref. p. iv). .

The veining of the wings in this genus offers several modifications of much interest with

reference to the typical arrangement of these organs in the order Lepidoptera. In their

simplest form in the fore wings they may be thus described :—Ist. A strong simple branch-

less vein, running parallel with and close to the front margin of the fore wing (the costal

vein); 2nd. A vein arising close behind the base of the costal vein, but dividing into six

branches before reaching the extremity of the wing (the subcostal or postcostal vein and

its branches); 3rd. The strong median vein, separating into four branches; 4th. The

subanal vein, simple and branchless, but sometimes accompanied by one or two rudimental

longitudinal vein-like folds. In most instances, in order to give support to the middle of

the fore wing, there is a transverse veinlet connecting the 6th branch of the postcostal

(6 5*) and the 4th branch (¢ 8*) of the median veins, forming a closed cell (the discoidal

cell). But in all the families of Butterflies (except the Papilionide) the median vein has

GENUS CASTNIA AND SOME ALLIED GROUPS. 159

apparently only three branches, and the postcostal vein only five branches; the two

intermediate branches, namely the 6th branch of the postcostal (6 5*) and the 4th

branch (¢ 3*) of the median veins, have been assumed to be distinct from those veins,

and to constitute the remains of a vein intermediate between the postcostal and the

median, of which the basal portion has been aborted or seen only in a fold running

towards the base of the wing between those two veins. To these two detached branches

Mr. Doubleday applied the names of the 1st or upper discoidal nervule, and the 2nd

or lower discoidal nervule; the little transverse veinlet connecting the 5th branch of

the postcostal vein with the upper discoidal nervule he termed the upper disco-cel-

lular nervule; that connecting the upper and lower discoidal nervules together he

termed the middle disco-cellular nervule; and that connecting the fourth median (or

lower discoidal) nervule with the third branch of the median vein he termed the lower

disco-cellular nervule (these three disco-cellular nervules closing the discoidal cell); the

longitudinal fold or obsolete vein between the bases of the postcostal and median veins

within the discoidal cell, he termed the discoidal nervule.

From the very great number of specimens of Lepidopterous insects of all families,

both diurnal and nocturnal, which I have examined, I prefer to treat the first or upper

discoidal nervule as part of the system of the postcostal vein and the second or lower

discoidal nervule as portion of the median system. But in order to mark the distinction

between these two aberrant nervules or branches and the true branches, or veinlets, or

nervules of the postcostal and median veins, I have, in the Plates of details of structure

illustrating this memoir, distinguished the upper or first discoidal nervule with the mark

65*, and the lower or second discoidal nervule as ¢ 3*.

In thus differing from the system of venation proposed by the late Mr. Edward

Doubleday in his paper on Argynnis, published in the ‘ Linnean Transactions,’ and in

the introduction to the ‘ Genera of Diurnal Lepidoptera,’ I cannot but admit the propriety

of the adoption of the stems of the veins as of primary importance—contrasted with the

plan of Herrich-Schaffer and other German writers, who seem to regard the branches as

of primary value, counting them (quite independently of their origin from the stems)

numerically upwards from the anal angle, from 1 to 12:—No. 1 being the anal vein

(my d); Nos. 2,3, 4 the three branches of the median vein; 5 and 6 the two disco-

cellular veins (or my ¢3* and 0 5*); 7, 8, 9,10, 11 the five branches of the postcostal

vein, and 12 the costal vein+.

Many years ago I had a long controversy with the late M. Alex. Lefebvre on this

subject, in consequence of his adoption of the principle that the fold indicating the vein

in the discoidal cell of the fore wing, and running thence towards the outer margin,

formed the true le of demarcation between the branches of the postcostal and median —

veins. The fold in question is, of course, the representative of the discoidal vein; and an

examination of the figures accompanying this memoir, or those of Herrich-Schiaffer’s

+ Inhis elementary plate, H.-Schiffer omitted one of the branches of the postcostal vein, and so gave only 11 ribs

(as he terms the branches). Assmann (Schmett. Schles. tab. A. f. 1) counts the “‘ Adern” of the fore wing from 1 to 14,

adding the two supplemental anal veins, which are sometimes, although very rarely, to be observed.

160 MR. J. 0. WESTWOOD ON THE LEPIDOPTEROUS

work, will show the complete irregularity in the position of this discoidal vein and its

-divisions.

The large moths of the genus Gastropacha are excellent instances in which the

arrangement which I have endeavoured to establish is shown. In G. repanda (H.-Schaffer,

Syst. Bearb. Schm. Eur. pl. xviii. fig. 3), the median vein is seen to have four branches,

the upper one of which (¢ 3*) arises from the third branch (¢ 3), at the lower extremity

of the discoidal cell, whilst the other, upper, accessory branch, my 0 5*, arises from the

preceding, 0 5, far beyond the discoidal cell, forming as clearly a portion of the post-

costal system as c¢8* does of the median, there being no trace of a discoidal vein within

the discoidal cell.

Aglaope infausta (H.-Schaffer, Syst. Bearb. Schm. Eur. pl. xiii. fig. 32) presents us with

an interesting example, in which the arrangement of Mr. H. Doubleday is excellently

shown. Here the last branch of the postcostal vein (65) extends to the tip of the fore

wings, and the discoidal cell is divided into two nearly equal parts by the discoidal vein,

which, extending nearly from the base of the wing, branches into two equal branches

(my 6 5* and ¢ 3*), leaving the three branches of the median vein at equal distances

apart.

In Leucophasia sinapis (see H.-Sch. op. cit. pl. v. fig. 11) the arrangement of the

branches is very instructive. Here, in consequence of the very small size of the discoidal ~

cell, all the branches of the postcostal vein arise quite regularly on the anterior side of the

vein beyond the cell, the fifth (6 5) extending to the tip of the wing; but the two accessory

and the three branches of the median vein are all equally independent at their origin at

the end of the short oval discoidal cell, and appear to be parts of one system of veins

distinct from the postcostal.

In many of the Geometridee, as may be seen especially in the 9th plate of Herrich-

Schaffer’s work on European Lepidoptera, vol. vi., not only is the small postdiscoidal

cell, as I have called it (cellula accessoria of H.-Sch.), found in the Castnie present,

but it is even divided into two cellulets by a more or less oblique cross vein.

In the Hesperiide, on the other hand, the discoidal cell of the fore wing is elongate-

ovate, and all the branches of the postcostal and median veins, as well as the two

accessory branches (0 5* and ¢ 8*), arise from the extremity of the discoidal cell at

nearly equal distances apart, without any trace of the discoidal vein within the cell; so

that it is impossible, except from analogy, to determine whither each of the several

systems of veins extends.

The same arrangement is also seen amongst the He erbeeroes Lepidoptera, as in the

fore wing of Scardia mediella (H1.-Sch. op. cit. vi. pl. Microl. ii. fig. 1), and in Nemotois

scabriosella (ib. pl. iv. f. 32).

In some species of Hesperiidee, however, as in Hudamus (Megathymus) yucce, the

postcostal and median systems are easily distinguished, although their two outer

branches, 0 5* and ¢ 3*, are united by an extremely slender and angulated veinlet

imperfectly closing the discoidal cell.

From the numerous figures given in Pls. XXVIII. and XXIX., it will be seen that

the branches both of the postcostal and median veins exhibit much irregularity in their

GENUS CASTNI4 AND SOME ALLIED GROUPS. 161

arrangement as compared with the simple system seen in so many other groups of

Lepidoptera. The number of branches is normally present; but the 2nd, 3rd, 4th,

and 5th branches of the postcostal (d 2, 63, 64, and 65) are generally arranged so that

they form a small oblong premedian or postcostal cell. Looking at Pl. XXVIII. fig. 6

(C. cacica), we see that this is effected by the branch 63 throwing off a very short

transverse branchlet connecting it with 6 4; or, rather, we may say that 6 4: commences

transversely, and then suddenly bends at a right angle, and runs longitudinally, so that

the little transverse branchlet must be regarded as its basal portion. In 45 we have

the anomaly that the 5th branch arises at a short distance beyond the 1st branch, and

forms the hinder margin of the postcostal cell, throwing off a little oblique branchlet,

by which itis united to the 4th branch, and which in fact forms part of the extremity

of the postcostal cell. In like manner the median vein (c) has its third branch (c 3)

strongly angulated at a right angle at its base, whilst the branch ¢ 3* (lower discoidal

nervule of Doubleday) is carried backwards by a fold (z), the discoidal nervure of

Doubleday, and almost to the base of the wing f. Moreover the veinlet connecting 6 5*

and ¢ 3* is angulated below its middle, throwing off backwards a fold which unites

with 3, and which indicates the normal position of the discoidal nervure of Doubleday.

The object of this complicated arrangement, which has not hitherto been described,

appears to be to form a strong network of corneous veins for strengthening the wing,

so as to enable it to perform the strong flight which these muscles doubtless enable

the insect to perform.

Modifications of the arrangement described above exist in many of the species of

Castnia. :

In the splendid new species C. veraguana (fore wing, Pl. XXVIII. fig. 9) the distribution

is nearly similar to that of C. cacica; but there is no little transverse branchlet connecting

63 and 04, and consequently there is no closed postcostal cell, and the branch ¢ 3* arises

from ¢ 3 in the ordinary manner at a little distance in front of the transverse base of

e 3, making an acute angle instead of being pushed backwards a little distance, and

making an obtuse angle as in C. cacica. The curious species C. cochrus (Pl. XXVIII.

fig. 11) agrees with C. veraguana in this respect.

In C. Botsduvallii and other closely allied species (Pl. XX VIII. fig. 8) the small narrow

posteostal cell exists; but it only emits one branch at its extremity, which branches off

into 63, 64, and 05, and the angulated base of 05* arises from the posterior margin

of the postcostal cell near its extremity.

In C. icarus (Pl. XXVIII. fig. 13) the small postcostal cell exists, emitting 6 2 from its

front margin a little beyond the middle, whilst from its extremity one branch is emitted

which forks into 6 3 and 6 4, whilst 0 5 arises from the extremity of the postcostal cell,

close to the base of preceding branch, and 0 5* from 0 5 at a little distance from its base.

In C. Hubneri (Pl. XXVITI. fig. 14) there is no closed small postecostal cell, the little

transverse veinlet between 03 and 04 being wanting. In Gray’s figure, pl. 18. f. 2,

the vein is misrepresented as closing the cell.

In C. Latreiilii (Pl. XXVITI. fig. 15) the small postcostal cell exists, terminating in an

+ C. linus scarcely differs from C. cacica in the arrangement of the veins of its fore wings (Pl. XXIX. fig. 9).

162 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

acute point from which a single branch is emitted, which forks into two branches, 63

and 04, 65and 0 5* arising together from the hind margin of the cell at three fourths of

its length from the base. C.mygdon (Pl. XXVIII. fig.7) and C. inca agree with C. La-

treillu in this respect.

Among the Helicoinoid species with elongated fore wings, forming the genus Gazera,

Boisduval, C. cononia (Pl. XXIX. fig. 8) has a small postcostal cell, the 2nd branch of

the postcostal vein arising shortly before its extremity ; the 3rd and 4th branches arise

from its extremity a little distance apart, the short transverse branchlet connecting

65 with 6 5* springs close to the emission of 65 from the postcostal cell ; the 5th branch

arises from the postcostal cell nearly opposite the base of the 2nd branch, and appears

to extend backwards to the origin of the 1st branch of the postcostal. The branches

b 5* and ¢ 3* extend backwards, forming an acute angle within the discoidal cell, and

emit a branch-like fold (discoidal nervure) towards the base of the wing; ¢3 likewise

is extended backwards, uniting with this discoidal nervure, forming with it a rudimental

(postdiscoidal) cell.

In C. equadoria (Pl. XXVIII. fig. 10) the arrangement of the postcostal branches

agrees with C. cononia, except that the little transverse vein uniting branches 65 and

b 5* is rather nearer the base of the wing.

In ©. mimica (Pl. XXIX. fig. 11) and C. linus (Pl. XXIX. fig. 9) the postdiscoidal cell

is also present ; but it only emits one branch at its acute extremity, which forks at some

little distance beyond the cell into the two branches 03 and 0 4.

In C. truvilla (Pl. XXVIII. fig. 12), on the other hand, the postdiscoidal cell is not

completed, there being no connexion between the branches 63 and 64; the relative

position of the postcostal branches is otherwise identical with that of C. linus.

In C. Marcel Serresi (Pl. XXIX. fig. 2), forming the type of Boisduval’s genus

Ceretes, the small postdiscoidal cell is present, emitting the branches 0 2, 63, 64, and

65 from its extremity at smail distances apart, and the short transverse veinlet con-

necting 0 5 and 65* (and closing the discoidal cell dc) from its hinder margin beyond

the middle. The discoidal cell is traversed by a longitudital vein-like fold, the hinder

portion of the cell (d c*) forming a distinct cell by the addition of a veinlet extending

backward from the basal junction of the median branches ¢3* and ¢ 3—thus leaving

a narrow tract between it and the basal portion of the median vein c, which may be

called the premedian cell, p.m. c.

In C. chremes (P1. XXIX. fig. 4) the arrangement is nearly similar; but the vein-like

fold in the discoidal cell is less strongly defined, and the little transverse vein which

closes the discoidal cell does not arise from the hind margin of the postcostal cell, but

is pushed forwards and arises from the 5th postcostal branch close to its base.

On comparing the figures of the wings of C. cacica (Pl. XXVIII. figs. 6 and 6 a) with

those of C. Marcel Serresi (Pl. XXIX. figs. 2 and 3), it will be seen that the general

arrangement both of the veins and cells is identical.

The arrangement of the veins of the hind wings has been comparatively neglected in

the Lepidoptera, although Dr. Boisduval has partially, but not satisfactorily, used this

character in the generic distribution of his Castniaires. In the vast majority of the

GENUS CASTNIA AND SOME ALLIED GROUPS. 163

species of Lepidoptera the number of branches of the veins in the hind wings is smaller

than in the fore wings; but it will suffice to examine the wings of Hepialus humuli

(Pl. XXIX. figs. 27 & 28) to be convinced that the same elements normally exist in

the hind as in the fore wings. It will be seen from the lettering of the branches, that

the costal, subcostal with its six branches, median with its four branches, and anal veins

are present; and even the discoidal cell is cut up into the three discoidal, subdiscoidal,

and premedian divisions. This will enable us to determine satisfactorily the normal

condition of the veins and branches when some of the latter are not present in the hind

wings.

In C. cacica (Pl. XXVIII. fig. 6a) the subcostal vein has only two branches, the

second of which is united with the base of ¢ 3* by a very thin curved fold, whilst the

veinlet connecting the base of ¢3 with that of ¢3* extends backwards, and is continued

into a discoidal nervure extending nearly to the base of the wing.

In C. juturna (Pl. XXIX. fig. 1) the same arrangement exists, except that the thin

fold arising a little beyond the base of 6 2 and ¢ 3* is strongly angulated, and a very

slight longitudinal fold extends from the angle towards the base.

In C. cochrus (Pl. XXVIII. fig. 11a) the fold connecting the extreme base of ¢ 2

and ¢38* is almost obsolete, but is seen to spring from the discoidal vein at some distance

before the origin of ¢ 3*, thus leaving the discoidal cell nearly entirely open.

In C. Hubneri (Pl. XXVIII. fig. 14a) the discoidal is fairly divided into three portions,

namely, the discoidal, subdiscoidal, and premedian portions.

In C. paradoxa the vein or fold connecting the branches 62 and ¢ 3* is entirely

wanting, and the true discoidal cell is wanting; but its premedian portion is present,

in consequence of the branch c 3* being extended backwards, nearly to the base of the

wing. i

C. acreoides (Pl. XXIX. fig. 14) agrees with C. paradoxa, except that ¢2 and c3 are

united together at the base beyond the cell.

In C. linus (Pl. XXIX. fig. 10) the almost indistinct fold connecting the base of the

subcostal vein with the discoidal nervure is pushed still further back, and thus differs

materially in its relation to the branches of the median and subcostal veins.

In C. mimica the arrangement is the same as in C. linus, except that the transverse

fold closing the cell is obsolete, being only indicated by two minute tubercles at the

place of its origin (Pl. X XIX. fig. 12).

In both sexes of C. Marcel Serresi (Pl. X XIX. fig. 3) the postcostal vein and its two

branches are quite free; but c3* extends backwards beyond the base of ¢3, forking

into two branches, which curve backwards to the base of the wing, thus forming a sepa-

rate subdiscoidal cell, the anterior or discoidal part being open, and a very narrow pre-

median cell.

In the female of C. chremes (Pl. XXIX. fig. 5) the same arrangement takes place, but

the discoidal cell itself is partially closed by a thin fold; and in the male the branches

e3 and ¢ 3* arise close together at the extremity of the premedian cell.

Lhave thought it advisable to trace these veins and their branches to a normal state

in order, if possible, to ascertain their significance, either as sexual, specific, subgeneric,

SECOND SERIES.—ZOOLOGY, VOL. I. a

164 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

or generic characters. It appears to me that we have now sufficient materials before us

to assume that they possess no higher than specific importance.

The palpi in C. dedalus are closely applied to the face, extending upwards to three

fourths of the height of the eyes, with their inner margins close together; the apical

joint very minute, but distinct and pointed, but not at all projecting. In C. atymnus

they do not reach higher than half the height of the eyes, and the terminal joint is less

prominent.

In C. eudesmia the palpi are longer, and extend as much as one fourth of their length

above the top of the eyes; they are rather obliquely porrected, standing off from the

face, and are more slender. In C. decussata they are parallel, and reach to the top of

the eyes. In C. Hubneri they only reach to half the height of the eyes, and are much

more rugosely squamose. In C. Marcel Serresi they are small, not reaching so high as

half the height of the eyes, slender, oblique, pointed, and finely scaled. In C. cochrus

they are small, not reaching higher than the epistome, and are thickly squamose, but not

rugose. Such is also their structure in the Heliconiiform species, as well as in C. linus

and acreoides.

The geographical range of the whole of the species of Castnia and its immediate allies

extends only to Southern and Central America.

The transformations of the species of Castnia are of great interest, in connexion with

the natural relations of the group and its position with regard to the Rhopalocerous

division of the Order in which it was placed by the early systematists.

Madame Merian, in the 36th plate of her work on the insects of Surinam, gave a set

of figures as representing the transformations of Castnia licas. The caterpillar is eyi-

dently that of one of the larger Morphideous Butterflies; and the pupa is a chrysalis

fastened by its tail to the plant, with its head downwards, resting on the leaf, but with-

out any indication of a central girth of silk round the body. Dr. Klug was fortunate,

however, to obtain a pupa of C. therapow from the bulb of a Catasetwm, sent from

Central America to the Royal Gardens at Potsdam. ‘This pupa, figured in the Appendix

to his Memoir on the genus Synemon, agrees with those of Sesia, Cossus, Zeuzera, and

Hepialus, in its elongated general form and in the transverse rows of reflexed spines

with which the segments of the abdomen are furnished. It is, however, to Dr. Philippi,

of Santiago, Chile, that we are indebted for a knowledge of the larva of the genus Castnia.

In the ‘Stettiner entomologische Zeitung’ for 1863, p. 337, and plate iil., he has de-

seribed and figured the transformations of C. ewdesmia, the larva of which he found in

the stem of Pourretia coarctata, one of the Bromeliacez. It is 44 inches long, and

closely resembles the larva of a Prionus or Cossus, with a large prothoracic and short

meso- and metathoracic segments, with six thoracic, eight ventral, and two anal feet. It

is white and fleshy, with a fulvous-brown-coloured head and extremity of the body.

When full-grown it forms a large cocoon of bits of leaves, twigs, and other vegetable

matter, fastened together with silken threads. The pupa is dark chestnut-coloured, with

the abdominal segments paler.

In the Hopeian Collection at Oxford is preserved one of these cocoons (Pl. XXVIII.

fig. 4) from which I extracted a pupa of C. eudesmia, nearly arrived at the imago state,

GENUS CASTNIA AND SOME ALLIED GROUPS. 165

so that the scaly covering of the fore wings could be easily removed or raised, showing

the markings of the wings of the future moth of a diminished size (Pl. XXVIII. fig. 3).

The classification of the species of the genus Castnia has not hitherto been successful,

owing to the imperfect examination of the minute specific details of structure of many

of the species; nor can I hope to have been more fortunate than my predecessors.

Dalman, who described seventeen species of the genus in his memoir in the Swedish

Academy’s Transactions, remarked in his introduction that C. linus differed from the rest

in its more elongated and partially hyaline wings and abdomen, longer than the wings,

adding,—“ An proprium etsi propinquum genus he discrepantiz indicant?” In his

list of the species at the end of his paper he places C. cronus in a separate second section,

and C. linus in a third, without any characters; and his description of the former of

these two species is confined to its colour and markings.

Mr. G. R. Gray, m his memoir in the Entomological Society’s ‘Transactions,’ proposed

to divide the species into five sections :—

1. Those with the palpi very long, closely applied to the head, and covered with short

scales. The fore wings are subtriangular, thickly covered with scales, with the exterior

margin straight or slightly rounded ; the lower wings beneath with one guide (or bristle

working in the socket at the base of the fore wing). C. cyparissias, licus, evalthe,

Fonscolombet, &e. (pl. xiii. fig. 1).

2. Those with the fore wings elongate, triangular. The hind wings are expanded,

with the posterior margin somewhat truncated: C. ardalus, palatinus.

3. Those with the palpi short, reaching halfway towards the base of the antenn, and

covered with long scales; the fore wings covered with small scales, with the exterior

margin rounded. The outer margin of all prominently fringed with long scales: C.

Hiibneri (pl. xiii. fig. 2).

4, Those with the wings covered with minute scales, partly diaphanous; the fore pair

with the exterior margin rounded ; similar also in the hind pair, with three wing-guides :

C. cochrus, linus, and acreoides.

5. Those with the antennz similar to the others, but the hook at the apex is formed

of large scales. ‘The head of the male is very broad, and the eyes are much larger than

in the female. All the wings have the outer margin rounded, with a narrow fringe of

small scales, and with four wing-guides. ‘The sexes differ much in colour: C. nicon

(pl. xii. fig. 8), thats.

The plate accompanying the memoir of Gray gives the veining of the upper and lower

wings, the clava of the antennze, and the palpi of the 1st, 3rd, and 5th of these groups.

Mr. F. Walker, in his Catalogue of the Lepidoptera Heterocera, Part I., neglected the

various excellent characters pointed out by Mr. G. R. Gray, and divided the species

simply in accordance with the form of the wings, thus :—

Div. 1. Body thick.

Subdiy. 1. Fore wings slightly angular at the tips; hind wings rounded.

Sect. 1. Wings very broad. C. dedalus, Latreillii, atymnius, cochrus, &c. (16 species).

Sect. 2. Wings rather narrower; fore wings longer. C. satrapes, therapon, ardalus (Brecourtii) ,

invaria (.carus, var.), inca, palatinus, subvaria (phalaris).

Z 2

166 MR. J.O. WESTWOOD ON THE LEPIDOPTEROUS

Subdiv. 2. Fore wings more distinctly angular at the tips: hind wings slightly angular. 5 species—

C. Dalmanni, hegemon, orestes, Boisduvalii, and amycus.

Div. 2. Body more slender.

Subdiv. 1. Hind wings slightly angular. C. mygdon and phalaris.

Subdiv. 2. Hind wings quite rounded.

Sect 1. Fore wings rounded at the tips, moderately long. C. actinophous (acreoides), decussata,

cronis, strigata (Godarti), Hubnerz.

Sect. 2. Fore wings slightly truncated at the tips. C. chremes, thais.

Sect. 3. Fore wings rounded at the tips, very long; resembling the Heliconii in structure. C. linus

and acreoides.

Div. ?. C. Kirstenit.

Dr. Boisduval, in his ‘Species général des Lépidoptéres hétérocéres,’ published in

February 1875 (although dated 1874), has divided the Castnie (Tribu des Castniaires)

into the following genera :—

1. Castnra (including 53 species, without any attempt at subdivision). Characterized by

the “Palpes divergents de trois articles écailleux, montant 4 peine au niveau du

front, le dernier article petit et conique” (as shown in Gray’s figures 10, 1¢, 1d).

The “antennes en massue, cylindriques, terminées par une trés-petite houppe

soyeuse’”’ (as in Gray’s figure 1 a), the ‘‘ pattes—avec les jambes munies de deux

ergots.”’ The other characters cannot be regarded as generic.

2. CERETES (n. g., including C. chremes, thais, and Fabric (Marcel Serresi) = Gray’s

5th group and Walker’s Div. 2, Subdiv. 2, Sect. 2, characterized by the “ téte petite”

(overlooking the difference of the size of the head in the two sexes). Antennze with

a “longue massue fusiforme se terminant en pointe”’ (contrary to Gray’s descrip-

tion), “ palpes trés-velus appliqués fortement sur le front, non divergents, & articles

indistincts.” Fore wings with the “sommet un peu pointu,” and hind wings with

the “deux nervures médianes trés-rapprochées et coudées en sens inverse, mais ne

se joignant pas pour former une cellule fermée.” This last character is evidently

taken from Gray’s fig. 3, and answers to the hind wing of C. Marcel Serresi, but

not to that of C. chremes, in which the cellule is pro tanto ‘ fermée” by the fold, as

shown in my figure. The fact that Dr. Boisduval excludes C. diva, Butler (tricolor,

Felder) from this genus, and unites it (p. 531) with Castnia, notwithstanding its

extremely close affinity with C. chremes, is a sufficient proof of the unsatisfactory

character of the genus.

3. ORTHIA, n.g., having the head small, the antenne “en massue fusiforme, terminée par

une petite pointe, palpes tres-velus, trés-courts, a articles indistincts. Ailes allongées,

assez étroites, les inférieures ayant prés de la base une petite cellule elliptique

fermée.”

This genus is divided into two uncharacterized sections :—

+1. C. therapon, Koll. (paradoxa, Bdv.).

2. C. acreoides, Bdy.

GENUS CASTNIA AND SOME ALLIED GROUPS. 167

t13. C. pelasgus, Fab.

A. C. unifasciata, Felder (Voy. Novara).

5. O. augias, Bdy. (Herr.-Sch.).

6. O. nexa, nu. sp., regarded as the probable female of O. augias.

The genus Orthia, as constructed by Dr. Boisduval, is quite untenable. C. therapon

and acreoides have long fore wings, with a small front costal cell, the discoidal cell

closed as above described; whilst the hind wings have the discoidal cell open; that is,

there is no junction between the branches 6 2 and ¢3*, the small cell near the base of

the wing noticed by Boisduval being the cell which I have termed premedian. This’

cell, indeed, exists in C. pelasgus, the first of Boisduval’s second section of Orthia; but

the fore wings have no postcostal cell, the second and third postcostal branches arising

from a single base; and the 4th, 5th, and 6th branches also arising from a single basal

portion of the vein; the discoidal cell itself is quite open, there being no connexion

between the branches .65* and ¢3*. The very broadly triangular wings of C. pelasgus

will also remove it from the two preceding, and indeed from O. augias, which possesses

a small postcostal cell, a closed discoidal cell, and a closed discoidal cell in the hind

wings. I do not hesitate therefore to reject the genus Orthia.

4, GAZERA, n.&., characterized by the “‘ Antennes en massue allongée, terminées par une

5) oO” £

petite houppe soyeuse a peine visible. Palpes courts arrivant & peine au milieu du

front, de trois articles, le troisiéme trés-distinct, nu et pointu. Téte assez petite.

> P

Ailes allongées Héliconiformes, a écailles moins grossiéres que dans les Castnia pro-

prement dites; les inférieures ayant la cellule discoidale étroite fermée en angle

aigu. Abdomen un peu plus long que les ailes inférieures.”” Containing

1. C: limus, F. 4. C. columbina, Bdv.

2. C. heliconioides, Bdv. 5. C. zagrea, Feld.

3. C. personata, Web. 6. C. simulans, Bdv.

The elongate ovate form of the fore wings of the species of this group (to which, how-

ever, C. therapon and acre@oides must be added) is scarcely sufficient to separate them

generically (seeing the variable form of the wings in the other species) from the rest.

The other characters proposed by Boisduval are too variable in the different species to

possess generic value.

Finding it impracticable to carry out a distribution of the species of Castnia founded on

the form of the wings, as partially proposed by Mr. Walker, and rejecting Boisduval’s genera

for the reasons given above, I prefer to regard the species as constituting a single genus,

and shall content myself by dividing them into two sections :—1st. Those with the fore

wings more or less broad and triangular. 2nd. Those with the fore wings more or less

elongate-ovate, the latter nearly corresponding with Boisduval’s genus Gazera.

Genus I. CastTni..

Sect. I. Species with the fore wings more or less broad and triangular.

1, CASTNIA D&aDALUS. C. alis fuscis metallico nitentibus, anticis fascia media angusta,

obliqua lateribus parallelis fere ad angulum internum extensa alteraque subapicali

168 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

magis irregulari e lunulis connexis, albis; alis posticis fuscis extus nigricantibus,

fasciis duabus macularum parvarum albidarum (margine externo parallelis), interna

interdum minus distincta. Long. corp. unc. 23. Expans. alar. ant. unc. 6-73.

Papilio dedalus, Cram. i. pl. i. f.1, A, B; Dalm. Monogr. Castn. p. 7, 1 (Casin. ded.) ; Act. Holm.

1824, p. 898; Walker, List Lep. Het. B.M.i. p.16; Boisduval, H. n. Lép. Hét. p. 499.

P. cyparissias, Fabricius, Gen. Ins. Mant. 257; Ent. Syst. i. 1. p. 80; Herbst, pl. 118. f. 1,2; Shaw’s

Nat. Miscell. pl. 574, p. 142 (Castnia, c.) ; Godart, Enc. Méth. ix. 797, Ins. pl. 12. f. 1 (Castnia

cyp.); Gray, Trans. Hnt. Soc. u.. 142.

Hab. Cayenne, Amazonia. In Mus. Hopeiano Oxonie, et Boisduval.

2. CASTNIA GERON. C. alis fusco-griseis, certo situ violaceo viridique nitentibus, supe-

rioribus supra fasciis duabus albis, subtus medio pilis longis barbatis; inferioribus

supra basi discoque barbatis, margine postico flavido maculatis, subtus dense squa-

mosis, duplici serie e maculis albis obsoletis. Exp. alar. antic. unc. 5 lin. 4

(14 centim.).

Castnia geron, Kollar, Lep. Bras. Ann. Wien, i. 217, pl. 13. f. 1; Walker, List Lep. Het. B. M. i. p. 20;

Boisduv. H. n. Lép. Hét. p. 496.

Hab. Brasilia. In Mus. Vindobon.

3. CASTNIA VERAGUANA (Pl. XXX. fig. 1). OC. thorace alisque anticis castaneo-brunneis

his basi czeruleo viridique nitentibus, in medio fascia lata valde angulata, antice

albida, postice brunnescente, punctisque nonnullis ovalibus versus apicem albidis ;

alis posticis chalybeo-nigris, fascia pone medium (e maculis ovalibus composita)

margineque postico aurantiaco-rufis, abdomine nigro. Long. corp. unc. 24. Exp.

alar. antic. unc. 6 lin. 7.

Hab. Veragua (Salvin). In Mus. Hopeiano Oxoniz (olim Saunders).

This very fine new species is allied to C. cacica, Bdv., H.-Sch., but has the fore wings

rather less triangular, and they do not possess the small oblong cell seen in other species,

formed by the connexion of the second and third branches of the subcostal vein by a

small transverse veinlet (see Pl. XXVIII. fig. 9).

4. CASTNIA SCHREIBERSII. C. alis anticis olivaceo-fuscis, medio multo pallidioribus et

luteo-albidis, fascia obliqua intus attenuata e costa, ante medium, ad medium disci

-extensa, ubi attingit maculam magnam rotundam brunneam parum distinctam et

ad apicem ale extensam, angulo apicali interno in maculam alteram maximam

intus dilatatam brunneam; alis posticis nigris, vitta longitudinali pone medium

disci antice dilatata, guttisque nonnullis (8-5) submarginalibus pallide ceruleis.

Long. corp. une. 2-24. Expans. alar. antic. unc. 54 (14°15 centim.).

Castnia Schreibersui, Mikan, Delect. Flore et Faun. Brasiliensis (1820) ; Boisduval, H. n. Lép. Hétér.

p. 500.

C. Latreillti, Godart, Enc. Méth. ix. 797; G. R. Gray, Trans. Ent. Soc. L. ii. p. 142; Walker, List

Lep. Het. B. M. pt. i. p. 17; Perty, Del. An. Art. Brasil. pl. 30. f. 7; Schaufuss, ‘Nunq.

Otiosus,’ p. 8.

GENUS CASTNIA AND SOME ALLIED GROUPS. 169

C. actor, Dalman, Monogr. p. 8, pl. 1. f.1; Act. Holm. 1824, p. 398, pl. 1. fig. 1.

Eupalamnides ctesiphon, Hubner, Exot. Schm., ii.

Obs. Variat presertim mas vitta, vel guttis ceruleis alarum posticarum interdum

obsoletis, ut in figura Dalmanni. ‘Talia vidi in Mus. Parisiensi cum nomine C. Godarti

inscripta (nec C. Godartii, Ménétriés= C. strigata).

Hab. Brasilia, Rio Janeiro, Minas, Pernambuco. In Mus. Britann., Hopeiano

Oxonize, et Boisduval.

5. Castnra cactca. C. alis supra nigro-fuscis, ciliis albis; anticis striga tenui albida

fere recta margini postico fere parallela, punctoque parvo in medio disci interdum

obsoleto: alis posticis fascia denticulata paullo pone medium, ad angulum analem

dilatata, maculisque quinque rotundis submarginalibus aurantiacis (3) vel rufis (2).

Variat interdum guttis duabus parvis pallidis inter fasciam et marginem externum

alarum anticarum. Long. corp. unc. 2. Expans. alar. ant. unc. 53.

Castnia cacica (Boisduval, MS.), H.-Schaffer, Aussereur. Schm., i. f. 143; Boisduval, H. n. Lép. Hétér.

p- 502; Walker, Cat. Lep. Het. B. M. p. 1580.

Mas? C. procera, Boisduval.

Obs. An excellent figure of this fine species is contained in Jones’s unpublished

‘Icones,’ Dandi, pl. 63, from the collection of Mr. Francillon, but without indication of

locality or name.

Hab. Ad ripas fluv. Magdalenze, Colombia, Polochic Valley (Salvin), Bogota; Nica-

ragua (Belt). In Mus. Hopeiano Oxonie, et Boisduval.

6. CasTNIA PROCERA. ‘“ D’un noir brun 4a reflet verdatre. Les ailes supérieures, dont

la frange est blanche, sont traversées, sur leurs tiers postérieur, par une raie blanche

droite, réguliére, ne touchant pas tout-a-fait la céte. Ailes inférieures offrent un

peu au-dela du milieu une bande transverse, assez étroite, rouge, un peu crénelée, et

dilatée 4 langle anal; entre cette bande et l’extrémité il y a, en outre, une série de

quatre gros points de la méme couleur. Différe de la cacica par ses ailes supérieures

un peu plus pointues au sommet, par l’absence de la petite tache blanche dis-

coidale, par ses ailes inférieures dont la bande transversale est moitié plus étroite, et

enfin par le dessous des ailes, qui est entiérement d’un brun uniforme, ainsi que le

corps. Le méme port et la méme taille que la cacica.” (Exp. 13 centim.)

Castnia procera, Boisduval, H. n. Lép. Hétér. p. 5038.

Hab. Guatemala. In Mus. Boisduval.

7. CASTNIA PYLADES. C- alis nigro-fuscis metallico nitentibus, anticis fascia lata fusco-

lutea media oblique ad angulum internum apicalem extensa, marginibus irregulari-

bus fuscoque irrorata alteraque abbreviata ejusdem coloris subapicali irregular,

strigaque nigra undulata apicali; alis posticis fuscis, fascia subapicali utrinque

dentata rufo-fulva, nigro maculata maculis versus angulum analem plus minusve

confluentibus. Femina differt alis anticis inter basin et medium fascia indistincta

subalbida. Long. corp. unc. 12-2. Exp. alar. antic. unc. 43-53-64.

170 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

Papilio pylades, Cram. pl. 397. f. A,B; Herbst, Icon. pl. 185. f. 1,2; Shaw, Nat. Miscell. pl. 895 5

Godart, Enc. Méth. ix. 797 (Castnia p.); Dalm. Ins. pl. 12. f. 2; Monogr. p.9; Act. Holm. 1824,

p. 899; Gray, Trans. Ent. Soc. L. ii. p. 142 ; Walker, List Lep. Het. B. M.i. p.17; Boisduval, H.

n. Lép. Hétér. p. 501.

Hab. Cayenne, Para. In Mus. Britann., Hopeiano Oxonice, et Boisduval.

8. CASTNIA ZERYNTHIA. C. alis anticis olivaceo-fuscis medio pallidioribus lutescentibus

basi et apice magis nigricantibus, macula magna cuneata obliqua nigricante e costa

ad medium ale extensa albo marginata vittaque nigricante e medio fere ad basin

marginis interni, maculaque magna trigona costali inter medium et apicem albo bi-

euttata; alis posticis, basi fuscis, medio cum abdomine albis, limbo lato nigro, fulvo

guttato. Long. corp. unc. 24. Expans. alar. ant. unc. 5$ (14 centim.).

Casinia zerynthia, G. R. Gray, Trans. Ent. Soc. Lond. i. p. 144; Walker, List Lep. Het. B.M. 1. p. 18;

Boisduval, H. n. Lép. Hét. p. 506.

C. Langsdorfii, Ménétriés, Enum. Corp. An. Mus. Petrop. pl. xi. f. 1.

Hab. Brasilia, Bolivia. In Mus. Britann., Hopeiano Oxonie, Boisduval.

9. CasTNIA IcARuUS. C. alis anticis fuscis margine interno late griseo, fasciis duabus

parallelis albis obliquis e costa fere ad angulum internum extensis, punctisque

nonnullis interdum fasciam parvam simulantibus versus apicem; alis posticis

leete rubris basi griseis, strigis duabus undulatis pone medium margineque externo

nigris maculisque nonnullis prope medium costz albis et nigris. Long. corp. maris

une. 14, foem. une. 1g. Expans. alar. antic. maris unc. 34, foem. une. 43-5

(10-12 centim.).

Papilio icarus, Cram. i. pl. 18. f. A, B; Shaw’s Mise. pl. 672; Godart. Enc. Méth. ix. p.798 ; Ins. pl. 12.

f. 3 (Castnia ic.) ; Dalman, Monogr. p. 10; Act. Holm. 1824, p. 899; Gray, Trans. Ent. Soe. L. ii.

p- 144; Boisduval, Consp. Lep. Guatemala, p. 75; H.n. Lép. Hétéroc. p. 503; Walker, List Lep.

Het. B. M. i. p. 19; Hiibner, Samml. exot. Schmett. 1. (Urbanus celebris ic.).

Hab. Cayenne, Brasilia, Guatemala, Maranham. In Mus. Britann., Hopeiano Oxoniz,

et Boisduval.

Var. (C. invaria) alis posticis maculis discoidalibus fere obliteratis. “ Fusca, corpus

subtus et abdomen testacea: ale cyaneo viridique nitentes; antice fasciis tribus

hyalinis [albis]; posticee rufse basi albidze, fascia antica [sc. versus costam] maculari

hyalina [alba], fascia breviore margineque postico nigris, hoc maculis 2 vel 3 rufis.”

Expans. alar. ant. unc. 47.

Castnia invaria, Walker, List Lep. Het. B. M. i. p. 23; Boisduval, H. n. Lép. Hét. p. 530.

Hab. Brasilia. In Mus. Britann.

10. CastNIA PAPAGAYA (Pl. XXX. fig. 6). Corpore crasso fulvo-fuscescente, abdominis

lateribus magis rufis; alis anticis ovali-trigonis, in medio saturate fulvescentibus,

basi et margine apicali obscurioribus maculaque media costali fusca bipartita,

medio pallidiore, fasciaque obliqua irregulari fusca inter medium ale et apicem,

ad costam et marginem internum dilatata, in medio angustata et ibi subobliterata,

maculam parvam albam prope costam includente; alis postic’s rotundatis chalybeo-

GENUS CASTNIA AND SOME ALLIED GROUPS. 171

nigris, fasciis tribus transversis e maculis ovalibus compositis, rubris, fimbriaque lute-

scente; antennis rufescentibus; alis subtus rufis, anticis nigro variis. Long. corp.

une. 13. Expans. alar. antic. unc, 33.

Hab. Papagaya (Rogers). In Mus. Hopeiano Oxonie (olim Saunders).

This species is remarkable for its large robust body and short strong wings.

11. Castyra Datmannit (Pl. XXX. fig. 5). C. alis ferrugineis, maris fasciis duabus

obsoletis, foemine fascia alba angusta et punctis hyalinis duobus; posticis rubris,

basi strigis maculisque juxta marginem nigris, maris disco flavo et foeminz albo.

Expans. alar. antic. maris unc. 34, foem. une. 4.

Castnia Dalmannii, Gray, Trans. Ent. Soc. L. ii. 145; Walker, List Lep. Het..B. M. 1. p. 26; nec

Boisduval, H. n. Lép. Hét. p. 512, which = C. hegemon.

Fom. C. Grayi, Boisduval, H. n. Lép. Hét. p. 498.

Hab. Brasilia. In Mus. Britann., Boisduval.

Dr. Boisduval’s description of the wings of the insect to which he has applied the name

of C. Grayi, will be seen from my figure (Tab. XXX. f. 5) and Gray’s description to agree

entirely with that of the female of C. Dalmannii, whilst the description of his C. Dal-

mannit as completely agrees with C. hegemon. His description of C. Grayi is as follows :—

* Ailes supérieures d'un brun un peu ferrugineux, A reflet verdatre; elles ont entre le

milieu et le sommet une raie oblique d’un blanc pur, allant de la céte au bord interne,

précédée en dehors d’un petit point de sa couleur: ailes inférieures d’un brun olivatre

a la base, marquées au milieu d’une large tache blanche transversale, atteignant le bord

abdominal; ensuite le contour est rouge, divisé en arriére par un cordon de taches

noires assez grosses et bien marquées. Un peu plus grande que |’ Japyz.”

Hab. Brasilia. In Mus. Boisduval.

12. Castnta Paunasta. C.- alis anticis olivaceo-fuscis, fascia valde sinuata et irregulari,

grisea, e costa (ultra medium), tune dilatata subbifida et inde versus basin marginis

interni extensa, maculaque magna fere rotundata ejusdem coloris versus angulum

posticum internum, liturisque duabus arcuatis subapicalibus albis; alis posticis

nigris fascia lata fere media alba seu flavescente, striga rubra maculari pone medium

(in limbo late nigro) guttisque submarginalibus albidis. Long. corp. maris unc. 1%;

foeem.une. 13. Expans. alar. antic. maris une. 31, foem. une. 4 (9-12 centim.).

Var. a alis anticis magis nigricantibus, posticis fascia alba venis nigris gracilibus divisa.

Castnia Pallasia, Eschscholtz, in Kotzebue, Reise, iii. p. 217, pl. 6. f. 27; Thon, Archiv, ii. pl. 1. f. 5;

Boisduval, H. n. Lép. Hét. p. 507. :

C. ardalus, Dalman, Monogr. p. 17; Act. Holm. 1824, p. 403; Gray, Trans. Ent. Soc. L. ii. p. 147 ;

Walker, List Lep. Het. B. M. i. p. 24.

C. Brecourti, Godart, Ene. Méth. ix. p. 798; Gray, Trans. Ent. Soc. L. u. 147.

Hab. Brasilia. In Mus. Britann., Hopeiano Oxonie (Burchell, 29 Dec. 1825, Catombi,

Brazil), Boisduval.

Var. 8. Mas e Papagaya fasciis variis alarum minus dilatatis.

SECOND SERIES.—ZOOLOGY, VOL. I. 2 A

172 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

13. CASTNIA GALINTHIAS. C. alis anticis acuminatis, fuscis, nitidis, lineolis longi-

tudinalibus albis et nigris alternis (in interstitiis yvenarum); alis posticis nigris,

fascia media (venis divisa) seriebus duabus macularum plus minusve distinctarum

albis, margine postico obscure fulvo. Long. corp. fem. une. 13. Expans. alar.

antic. unc. 33.

Castnia galinthias, Hopfter, Neue Schmett. tab. iv. f. 4; Boisduval, H. n. Lép. Het. p. 412.

Hab. Brasilia, Novo Friborgo. In Mus. Hopeiano Oxonie et Berol.

14. CasTNIA HARMODIUS. C. alis fusco-nigricantibus metallico tinctis, anticis fimbria

alba, fascia media obliqua postice attenuata e costa ad angulum internum extensa,

altera sinuata, subapicali minore e maculis quinque formata; alis posticis fascia

intermarginali maculari rufa, ad angulum analem dilatata et striga obliqua nigra ~

divisa. Long. corp. unc. 13. EHxpans. alar. antic. une. 3g—-44.

Obs. Striga parva obliqua fasciam rubram submarginalem alz posticee in duas partes

dividens interdum obliterata est, inde fascia integra evadit.

Papilio harmodius, Cramer, ii. pl. 223. f. C, D; Dalman, Monogr. Act. Holm. 1824, p. 399 (Castnia h.) ;

Gray, Trans. Ent. Soc. L. ii. p. 143; Walker, Cat. Lep. Het. B. M.i. p. 21; Boisduval, H. n. Lép.

Hét. p. 525.

P. syphax, Fabricius, Syst. Ent. p.48; Ent. Syst. ii. 1,p.45; Herbst, Ins. vi. p. 73, pl. 134. f. 1, 2; Jones,

Tcones, 11. pl. 74. f. 1.

P. pirrha, Oliv. Enc. Méth. Ins. pl. 16. f. 2.

Castnia licas, var., Godart, Hnc. Méth. x. p. 797.

Hab. Cayenne, Maranham, Amazonia, Surinam, Para, Demerara. In Mus. Britann.,

Hopeiano Oxonie, Boisduval.

15. CASTNIA ATYMNIUS. OC. alis anticis nigro-fuscis metallico nitidis, fascia media albida

obliqua fere recta ad angulum analem extensa alteraque subapicali valde sinuata e

maculis 5 vel 6 formata (interdum obliteratis) ; alis posticis fuscis extus plus minusve

brunneis, fascia magna alba obliqua media versus costam attenuata, et ad angulum

analem valde dilatata, fimbria partim alba; antennis apice fulvis. Long. corp. une.

13-13. Expans. alar. antic. unc. 33-33.

Var. a maculis subapicalibus albis in alis anticis valde distinctis. (Colombia, Nicaragua,

Bogota.)

Var. 8 fascia media alarum anticarum alba distincta, fascia maculari subapicali fere

obliterata squamis nonnullis pallidioribus tantum distinguenda, vel omnino obliterata

(ut in figura Pertii), alis posticis brunneis albo fasciatis. (Colombia, St. Hstavan,

Venezuela. )

Var. y fascia media et fascia subapicali alarum anticarum fere obsoletis. (Nicaragua,

Santa Martha, Guatemala, Mexico.) .

Castnia atymnius, Dalman, Monogr. p. 12; Act. Holm. 1824, p. 400; Gray, Trans. Ent. Sec. L. un.

142; Walker, List Lep. Het. B. M.i. p. 17 (C. atymnus) ; Boisduval, Cons. Lepid. p. 75; H. n. Lép.

Hét. p. 528.

C. spixii, Perty, Del. An. Art. Bras. pl. 31. f. 3.

GENUS CASTNIA AND SOME ALLIED GROUPS. 173

Obs. C. futilis, Walker (List Lep. Brit. M. pt. vii. 1856, p. 1581), from Nicaragua, is

described from two old faded and rubbed specimens of this species in the British Museum.

Hab. Brasilia, Guatemala, Nicaragua, Honduras, Venezuela, Columbia, Bogota. In

Mus. Britann., Hopeiano Oxoniz, Boisduval.

16. Castnia Licus. C-.alis fusco-nigris metallico nitidis, anticis fascia media recta postice

attenuata ad angulum internum extensa, alteraque subapicali smuata maculari;

alis posticis fascia magna alba obliqua, versus costam attenuata, ad angulum

analem valde dilatata, serieque marginali macularum rubrarum. Long. corp. une.

14-14. Expans. alar. unc. 33-4 (7-11 centim.).

Var. a fascia maculari subapicali alarum anticarum fere obliterata.

Var. 6 alis anticis lutescentibus, basi, apice, marginibus fascixe medie nigris, maculis

alarum posticarum pallide fulvis.

Papilio licus, Fabricius, Ent. Syst. ii. 1, p. 47; Herbst, Ins. pl. 134. f. 38, 4; Drury, Ins. i. pl. 16. £.1, 2;

Merian, Ins. Surimam. pl. 86; Oliv. Enc. Méth. Ins. pl. 12. f. 4; Jones, Icones, iu. pl. 73. f£. 3, 4;

Godart, Enc. Méth. ix. p. 797 (Castnia licus) ; Cramer, iii. pl. 223. f. A, B; Dalman, Monogr.

p. 13; Act. Holm. 1824, (Castnia licus); Gray, Trans. Ent. Soc. L. 1. p. 143; Walker, List

Lep. Het. B. M.i. p. 18; Cuvier, Régne An. ed. Crochard, Ins. pl. 145. f. 1, 2; Boisduval, H. n.

Lép. Hét. p. 526; Chenu, Ency. Pap. 235. f. 402.

Obs. I am inclined to believe that this supposed species is formed of individuals of

C. atymnius, the chief difference consisting in the submarginal rows of red spots on the

hind wings, which, indeed, vary both in number and size in the specimens in the Oxford

Museum. Several faded specimens in the Hopeian Collection, marked C. futilis by Mr.

Walker, only differ from similar faded specimens of C. atymnius (also labelled C. futilis

by Mr. Walker) in having a few slight orange marks near the angle of the hind wings.

Var. a alis posticis fascia brevi alba e disco ad angulum internum extensa margine

aurantiaco (Walker, op. cit. vii. p. 1582).

Hab. America meridionalis, Ecuador, Trinidad, Cayenne, Surinam, Demerara, Para,

Brazil, Ega (Amazonia); mouth of river Una, near Para, 19 Jan. 1830 (Burchell);

Upper Orinoco (larva in Orchidearwm radicibus). In Mus. Britann., Hopeiano Oxonie,

Boisduval.

17. Castyta Botsptvattr. Mas C. alis anticis purpureo-fuscis metallico nitidis, basi,

macula trigona media costali, altera magna trigona inter medium cost et apicem

(guttas tres pellucidas gerente), his maculis intus cum altera magna trigona (e

medio marginis interni ad angulum externum extensa) connexis; alis posticis basi

nigris, in medio macula magna aurantiaco-rufa marginis antici, in strigam tenuem

ad marginem analem extensa, limbo late nigro, rufo plus minusve maculato, maculis

duplici serie congestis. Long. corp. unc. 14. Exp. alar. antic. unc. 23—4.

Foemina differt abdomine albo, macula postica alarum anticarum extus griseo-suffusa,

alisque posticis fascia media lata ad marginem analem extensa maculisque mar-

ginalibus omnibus flavidis.

Castnia Boisduvalii, Becker, MS.; Walker, List Lep. Het. B. M. i. p. 27; Schaufuss, Nung. Otiosus,

p- 9; Boisduval, H. n. Lép. Hét. p. 510.

2A 2

174 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

3. C. Besckei, Ménétriés, Enum. Anim. Mus. Petrop. u. pl. xi. f. 3.

9. C. Herrickii, Boisduval, MS.; Herr.-Schaffer, Samml. aussereur. Schmett. fig. 144; Walker, List Lep.

Het. B. M. p. 1581.

Hab. Brasilia, Bahia, Pernambuco, Papagaya, Parana, Bolivia. In Mus. Britann.,

Hopeiano Oxoniz, Boisduval.

18. CastN1a oRESTES. C. alis anticis plus minusve pallide luteo-fuscis, basi, fascia e

medio coste fere ad angulum internum extensa (ramum curvatum intus pone

medium disci emittente), macula magna irregulari trigona subapicali (guttas tres

indistinctas hyalinas gerente) margineque apicali obscurioribus ; alis posticis basi

eriseis, medio macula magna triangulari, e margine anali fere ad costam extensa,

alba, limbo lato nigro fasciis duabus irregularibus e maculis aurantiacis plus minusve

distinctis compositis notato. Long. corp. unc. 1-1}. Expans. alar. 25-32.

Castnia orestes, Boisd. H. n. Lép. Hét. p. 509.

C. umbrata, Ménétriés, Enum, An. Mus. Petrop. pl. xi. f. 2; Walker, List Lep. Het. B. M.1. p. 26.

Hab. Brasilia, Rio Janeiro, Venezuela. In Mus. Britann., Hopeiano Oxonie,

Boisduval.

19. CastNIA IncA. C. alis anticis fusco, luteo et brunneo variegatis, maculis duabus

pallidioribus costalibus, una media subtrigona, altera semirotundata inter primam et

euttas tres hyalinas subapicales, hujus macule margo posticus niger obliquus, et cum

striga tenui obliqua prope basin marginis interni retro extensa continuus; alis

posticis basi fuscis; medio fulvo-rufis, ultra medium nigris, fascia maculari albida

submarginali. Long. corp. unc. 14. Expans. alar. antic. unc. 383-43 (9-10 centim.).

Var. a alis posticis serie macularum parvarum rotundarum parum distincta, inter fasciam

fulvam et maculas submarginales.

Castnia inca (Boisduval, MS.), Herr.-Schaffer, Samml. aussereur. Schm. f. 488, 489; Walker, List

Lep. Het. B.M. i. p. 24; Hopffer, Neue Schmett. Heft ii. pl. 4. £2; Boisduval, H. n. Léep. Hét.

p- 524.

Hab. Mexico prope Oajaca, Honduras, Venezuela. In Mus. Britann., Hopeiano

Oxoniz, Boisduval.

20, CASTNIA HECATE. (. alis fuscis olivaceo nitidis, fascia ocellari (margini postico fere

parallela) postice dilatata strigaque obliqua alba e medio coste ad medium disci

extensa, ubi fasciz ocellari se conjungit ; alis posticis rufo-fulvis, fascia pone medium

limboque tenui marginali nigris. Long. corp. unc. 17-13. Expans. alar. antic.

unc. 3¢-4¢ (8 centim. ).

Castnia hecate (Boisduval, MS.), Herrich-Schiffer, Samml. aussereur. Schmett. f. 439; Boisduval, H.

n, Lép. Hét. p.531; Walker, List Lep. Het. B. M. p. 1581.

Hab. America meridionalis, insula Haiti. In Mus. Hopeiano Oxonize et Boisduval.

9]. Castnta srricata. C.alis fuscis, metallico nitentibus, albo variis, scil. strigis duabus

longitudinalibus basalibus, fascia submedia, valde angulata et in medio interrupta,

dimidio antico integro, postico e maculis 4 subovalibus formato; inter hanc fasciam

GENUS CASTNIA AND SOME ALLIED GROUPS. 175

et apicem striga obliquae maculis 3-4 formata serieque subapicali macularum, in

medio interrupta et versus apicem interdum fere obliterata; alis posticis nebula

media costali fulvescente, fascia maculari pone medium alteraque submarginali ¢

maculis minoribus formata albis. Long. corp. unc. 1. Expans. al. ant. unc. 23

(6-63 centim.).

Casinia strigata, Walker, List Lep. Het. B. M. i. p. 30; Boisduval, H. n. Lép. Hét. p. 519.

C. Godartui, Ménétriés, Enum, Corp. Anim. Mus. Petrop. pl. 11. fig. 4, p. 130.

Hab. Brasilia, Pernambuco. In Mus. Britann., Hopeiano Oxoni, et Boisduval.

22. CASTNIA DECUSSATA. C. alis fuscis viridi nitentibus, fascia angulata (literam magnam

V supinam simulante) alteraque valde sinuata e maculis 5 composita subapicali,

maculis alteris duabus versus angulum internum positis albis; alis posticis fascia

irregulari pone medium, e maculis 5-7 oblongis composita, strigisque nonnullis fere

obsoletis marginalibus prope angulum analem. Long. corp.unc. 1%. Expans. alar.

ant. unc. 2-3 (6-6 centim.).

Castnia decussata, Godart, Enc. Méth. ix. 799; Gray, Trans. Ent. Soc. Lond. ii. p. 146; Walker, List

Lep. Het. B. M.i.p. 29; Hiibner, Zutrage, iii. f. 689, 640; Boisduval, H. n. Lép. Hét. p. 519.

A figure of this species is contained in Jones’s ‘ Icones,’ pl. 89. f. 1, without any

reference or locality.

Var. a alis magis maculatis, = C. melaleuca, Boisduval, MS. in Mus. Parisiensi.

Hab. Brasilia. In Mus. Britann., Hopeiano Oxoniz, et Boisduval.

23. CASTNIA PAPILIONARIS. (Tab. XX XI. fig. 3.) Alis anticis obscure fuscis metallico

nitidis, puncto parvo ad apicem cellule discoidalis, fasciaque tenui recta inter

medium et apicem ale pallide flavis, fimbria alba; alis posticis nigris, fascia pone

medium, versus angulum ani attenuata, cerulescente. Long. corp. unc. 13. Exp.

alar. antic. unc. 4.

Castnia papilionaris, Walker, List Lep. Het. B. M. part xxxi. p. 42; Boisduval, H. n. Lép. Hét. p. 500.

Obs. The ochraceous colour of the body described by Mr. Walker is simply the result

of the rubbed condition of the specimen in the British Museum.

Hab. Bogota. In Mus. Britann., et Hopeiano Oxonie.

24. CASTNIA SATRAPES. (Tab. XXXI. fig. 4.) Alis anticis viridi-fuscis, macula magna

marginis interni (guttam fuscam arcuatam includente) fasciaque lata subapicali

albidis fusco irroratis, squamisque nonnullis subapicalibus albis; alis posticis

aurantiacis, venis radiantibus limboque nigris, hoc albido maculato. Long. corp.

une. 1 lin. 5. Exp. alar. antic. une. 4 lin. 2 (74 centim.).

Var. a maculis pallidis alarum anticarum intus connexis, ut in figura Kollari.

Var. 8 maculis pallidis alarum anticarum separatis, alis posticis aurantiacis nigro

limbatis.

Var.-y maculis pallidis alarum anticarum separatis, alis posticis flavidis nigro limbutis,

limbo serie macularum rubrarum intus notato. (Tab. XX XI. fig. 5.)

176 MR. J. 0. WESTWOOD ON THE LEPIDOPTEROUS

Casima satrapes, Kollar, Lep. Bras. Ann. Wien. Mus. ii. 216, pl.12. £.3; Walker, List Lep. Het. B.

M.1i. p. 33; Boisduval, H. n. Lép. Hét. p. 5388.

Hab. Brasilia. In Mus. Hopeiano Oxoniz et Boisduval.

25. CASTNIA PALATINA. C. alis anticis rufo-brunneis medio pallidioribus, macula magna

subovali e medio coste versus medium disci oblique extensa, gutta parva hyalina

subapicali alis posticis basi fuscis, medio flavidis versus costam aurantiacis, limbo

nigro striga submarginali flavo-maculari. Long. corp. unc. 13. Expans. alar. ant.

une. 33 (8 centim.).

Papilio palatinus, Cramer, ii. pl. 159. f. B,C ; Godart, Ene. Méth. ix. p. 799 (Castnia palatina) ; Dalman,

Monog. p. 18; Act. Holm. 1824, p. 403; Gray, Trans. Ent. Soc. Lond. ii. p. 147; Walker,

List. Lep. Het. B. M.i. p. 25; Boisduval, H. n. Lép. Hét. p. 513.

Hab. Cayenne, Surinam, Demerara, Para. In Mus. Britann., Hopeiano Oxoniz,

Boisduval.

26. CASTNIATHERAPON. Alis elongatis, anticis supra fulvis, striga tenui nigricante obli-

quae basi marginis postici fere ad apicem ducta, macula media disci fusca versus

basin vitta nigricante marginata, margine interno apiceque fuscescentibus, hoc

obscure biguttato ; alis posticis rufo-aurantiacis, limbo apicali nigro, hoc guttis septem

albidis notato. Long. corp. lin. 18. Expans. alar. ant. unc. 3 lin. 4 (74 centim.).

Castnia therapon, Kollar, Lep. Bras. Ann. Wien. Mus. ii. 218, pl. 13. f. 3; Walker, List Lep. Het. B. M.

i. 23; Boisduval, H. n. Lép. Hét. p. 540 (Orthia th.).

C. paradoxa (Boisduval, MS.), Herr.-Schaffer, Samml. aussereur. Schmett. f. 16, 17.

Hab. Brasilia. In Mus. Hopeiano Oxonie et Boisduval.

27. Casrnta cuirarcHa. (Tab. XX XI. fig.1, 2.) Foem. alis anticis magnis obovalibus

apice subacutis, stramineo-albis, brunneo-fusco variegatis, maculis in medio e

costa literam h magnam reversam valde obliquam rude simulantibus, striolaque

tenui denticulata fere ex apice versus basin marginis interni extensa, apiceque

ipso fuscis, hoc punctis tribus albis (intermedio multo majore) notato, limbo etiam

fusco; alis posticis leete fulvis basi magis rufis, fascia rufescente inter medium et

apicem alze seriem macularum nigrarum includente limboque denticulato nigro ;

corpore toto fulvo thorace saturatiore, subtus pallidiore.

Mas multo minor, alis anticis saturate fulvis notulis foeminze multo minus distinctis ; alis

posticis basi paullo obscurioribus, aliter ut in foemina notatis. Long. corp. maris

fere unc. 13, fem. unc. 13. Expans. alar. antic. maris unc. 33, foem. une. 5.

This species is closely allied to C. inca, the figures of which hitherto published appear

to represent female specimens. The female of the present species has the fore wings

much more strongly variegated ; and the broad black border of the hind wings in C-. inca,

with its row of pale straw-coloured spots, is replaced by a richly coloured and marked

margin in C. elitarcha. The cell and branches of the postcostal vein of the fore wing

are arranged as in V. Latreillii, the third and fourth branches separating at a consider-

able distance beyond the cell.

Hab. Panama (Salvin), Nicaragua (Belt). In Mus. Hopeiano Oxonie.

GENUS CASTNIA AND SOME ALLIED GROUPS. 177

28. CASTNIA PHALARIS (Fab. nec Godart). (Tab. XXX. fig. 3.) (C.alis anticis fuscis basi

et margine externo obscurioribus, disco nonnihil pallidiore, fascia lata obliqua nigri-

cante e medio costz ultra medium disci extensa, inde in plagam magnam rotundam

angulum posticum apicalem occupantem (medio pallidiorem) dilatata, maculaque

magna trigona subapicali (guttas tres subhyalinas includente) obscura; alis posticis

nigris limbo lato aurantiaco-fulvo, intus irregulariter undulato, striga submarginali

fusca margineque ipso interdum nigro dentato. Long. corp. lin. 10-15. Expans.

alar. antic. une. 24-84.

Papilio phalaris, Fabricius, Ent. Syst. ii. 1, p. 45; Jones, Icones, iti. pl. 75. f. 3, 4; Donovan,

Naturalist’s Repos. ii. pl. 47. f.1, 2 (nee Godart, nec Boisduval, =mygdon, Dalm.) ; Dalman, Monogr.

Castnia, p. 22; Act. Holm. 1824, p. 405; Gray, Trans. Ent. Soc. Lond. ii. 145; Walker, List Lep-

Het. 1. p. 19.

Castnia subvaria, Walker, op. cit. p. 25.

C. musarum, Boisduval, MS. in Mus. Parisiensi.

Var. a striga submarginali fusca alarum posticarum fere obliterata, margine ipso fulvo.

C. dionea, Hopfier, Neue Schm. pl. 5. f. 3; Boisduval, H. n. Lép. Hét. p. 523.

Hab. Minas Geraes Brasiliec. In Mus. Britann., Hopeiano Oxoniz, Berolini,

Boisduval.

C. phalaris was described by Fabricius from the cabinet and Icones of Mr. Jones,

whose figures were published by Donovan in his ‘ Naturalist’s Repository.’ The Fabri-

cian description is simply “alee supra fusce, anticee immaculate, postice fasciis duabus

fere marginalibus e maculis exalbidis.” Godart applied the description of Fabricius to

the insects subsequently described and figured by Dalman under the name of P. mygdon,

with which, indeed, the two rows of spots on the outer margin of the hind wings

described by Fabricius agree both in form and colour.

29. CASTNIA EUDESMIA. C. alis anticis glauco-fuscis, costa tenui margineque interno

late albo pulverosis, fasciisque duabus postice acuminatis e costa ultra medium

disci extensis; alis posticis nigris, fasciis duabus macularibus inter medium et mar-

ginem posticum sanguineis albo irroratis, margine ipso anguloque anali griseis, hoc

fulvo squamoso. Long. corp. une.13. Hxpans. alar. ant. unc. 34 (9-11 centim.).

Castnia eudesmia, Gray, Trans. Ent. Soc. L. ii. p. 145 ; Herrich-Schiffer, Samml. aussereur. Schm. f. 140,

141; Gay, Historia fisica de Chile, Lep. pl. 5. f. 8, p. 47; Walker, List Lep. Het. B. M.i. p. 19;

Boisduval, H. n. Lép. Hét. p. 517.

Hab. Chili; Brasilia? In Mus. Britann., Hopeiano Oxoniz, Boisduval.

30. CasTNra cocHRuUS. C.alis nigris; anticis in medio subpallidioribus, venis nisris,

striga obliqua tenui nigra pone medium disci versus apicem alee extensa, vittaque

subalbida postica adjecta; alis posticis nigris macula media alba transverse ovata

in sex partes divisa, venis guttisque nonnullis plus minusve obsoletis versus angulum

externum albis; abdominis segmentis 5 apicalibus rubris, nigro et albo marginatis.

Long. corp. unc. 13. Expans. alar. antic. unc. 4-42.

Papilio cochrus, Fabricius, Ent. Syst. 11. 1, p. 42; Donovan, Nat. Repos. pl. 156; Jones, Icones, iii.

pl. 77. f.1; Godart, Enc. Méth. ix. p. 798 (Castnia c.) ; Gray, Trans. Ent. Soc. Lond. ii. p. 147;

Walker, List Lep. Het. B. M.i. p. 20; Boisduval, H. n. Lép. Hét. p. 517.

178 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

Castma maris, Dalm. Monogr. p. 16; Act Holm. 1824, p. 401, pl. 1. f. 4.

Prometheus casmilus, Hiibner, Samm]. exot. Schmett. ii. f. 1, 2.

Hab. Brasilia. In Mus. Britann., Hopeiano Oxonie, Boisduval, &c.

31. CasTNIA RUTILA. C. alis anticis testaceo-fulvis, macula obscuriore in medio cost,

intus bifida, ramis curvatis, ad venam mediam extensis, serieque irregulari punc-

torum obscuriorum inter medium et marginem apicalem, scil. 2 versus apicem, supe-

riore majore nigricante guttam albam includente, 3 in medio majoribus, et. 2 versus

angulum internum multo minoribus, serieque marginali macularum cuneiformium

obscuriorum; alis posticis testaceo-rufis, fascia submarginali e maculis 8 nigris

formata, margineque ipso tenui nigro intus undulato. Exp. alar. antic. cire. une. 5.

Castnia rutila, Felder, Reise Novara, pl. 79. f.1; Boisduy. H. n. Lép. Hét. p. 496.

Hab. Amazonia. in Mus. Felder.

32. CasTnta Fasrici. C-.alis anticis luteo-fulvis (“ ferrugineis ”’), nebulis obscurioribus

fuscescentibus, scil. macula seu circulo juxta medium coste, fasciaque lata inter

medium et marginem apicalem, cum margine parallela, versus costam et marginem

internum dilatata, hujus fasciz marginibus undulatis et presertim postice obscu-

rioribus; alis posticis rufis (‘‘ pallide testaceis’’), fasciis tribus nigris, scil. prima ante

medium abbreviata, secunda mediana, tertia pone medium, his duabus crenatis,

serieque submarginali macularum sex nigrarum ovalium notatis. Expans. alar.

une. 4¢ (54 Gray, an recte P).

Castmia Fabricii, Swainson, Zool. Ill. iii. pl. 149; Thon, Ent. Archiv, 1829, f. 1236; Gray, Trans. Ent.

Soc. L. ui. p. 144; Walker, List Lep. Het. B. M.i. 19 (nec C. Fabricii, Godart, Ene. Méth. ix. 800,

=C. Marcel Serresi feem., nec C. Fabricii, Boisduval, H. n. Lép. Hét. p.539,=C. chremes.

Hab. Brasilia (“ diamond district ’’).

33. CASTNIA JaAPYX. C. alis rufo-brunneis, fascia obliqua pone medium obscuriore, in

mare macula parva pallidiore ad apicem cellule discoidalis punctoque hyalino inter

cellulam et marginem externum, in feemina maculis 5 rotundis inter se distantibus

discoidalibus flavescentibus ; alis posticis fulvis extus magis testaceis, basi fusces-

centibus, in foemina fasciis duabus submarginalibus e maculis parvis nigricantibus

formatis, his maculis in maribus fere obliteratis (nonnullis versus angulum ex-

ternum tantum apparentibus). Long. corp. unc. 14-1. Expans. alar. ¢ une. 33

(10 centim.), @ une. 44 (114).

Athis japyx, Hubner, Samml. exot. Schmetterl. ii. 362, f. 1, 2, ¢ ; Boisduval, H. n. Lép. Hét. p. 497.

Castnia Kirstenti, Thon, Archiv, ii. p. 7, f. 6-10, g¢ ; Walker, List Lep. Het. B. M. p. 33.

C. Fonscolombei, Godart, Enc. Méth. ix. p. 799; Gray, Synops. Trans. Ent. Soc. ii. p. 144; Walker, List

Lep. Het. B. M. i. p. 20.

Hab. Brasilia. In Mus. Britann., Hopeiano Oxonie, Boisduval, &c.

34, CASTNIA HEGEMON. C. alis anticis rufo-badiis, macula media costali fasciaque

obliqua pone medium ale (ad medium marginis interni et prope apicem dilatata

ef male determinata) magis brunneis; alis posticis basi nigricantibus, disco flaves-

GENUS CASTNIA AND SOME ALLIED GROUPS. 179

centi, margine anali albo, limbo lato rubro, strigis duabus undulatis nigris versus

angulum analem latioribus et ibi coalitis. Fcemina (sec. sententiam Boisduvalii)

differt fasciis alarum anticarum magis distinctis, postica seepe interrupta ; alis posticis

disco niveo, strigis nigris marginis postici magis determinatis. Long. corp. unc. 14.

Expans. alar. ant. unc. 33.

Castnia Hegemon, Kollar, Lep. Brasil. Ann. Wien, 1. 217, pl. 13. f. 2; Walker, List Lep. Het. B. M. i.

p- 26; Boisduval, H. n. Lép. Hét. p. 514.

C. Dalmannii, Boisduval, H. n. Lép. Hét. p. 512 (nec Gray =Grayi, Bdv. p. 498).

Dr. Boisduval has mistaken this very distinct species for C. Dalmannii of Gray, and

has applied the name of Grayi to the latter insect.

Hab. Brasilia, Rio Janeiro. In Mus. Britann., et Hopeiano Oxoniz.

35. CasTNIA creLA. “Ailes supérieures d’un gris-blanchatre chatoyant en vert ou en

violet, avec la base, le bord terminal, une grosse tache sous-costale et une bande trans-

versale de couleur brune; la bande transversale située sur le tiers postérieur est

sinuée; elle se dilate sur la céte et sur le bord interne, de sorte qu’elle est un peu

étranglée dans son milieu. Ailes inférieures d’une belle couleur orange, avec un arc

discoidal noir; leur tiers postérieur est marqué d’une rangée de taches un peu

oblongues, un peu plus pales que le fond, séparées par des nervures noires et ren-

fermées entre deux lignes ondulées d’un noir foncé.” Expans. alar. antic. 8 centim.

Casinia ciela (Boisduval, MS.), Herr.-Schaffer, Samml. aussereur. Schmett. figs. 486, 487; Boisduval,

H. un. Lép. Hét. p. 532; Walker, List Lep. Het. B. M. pt. xxxi. p. 43.

Hab. Bahia, Penambuco. In Mus. Boisduval.

36. CastNIA areus. “Ailes supérieures pointues au sommet, d’une teinte brune

obscure, qui oblitére en partie des bandes d’une couleur plus foncée, disposées comme

dans les espéces précédentes (Phalaris et Mygdon) mais beaucoup moins distinctes ;

cependant on voit toujours dans la fissure de la bande bifide trois petites taches

blanches plus ou moins marquées, un peu confluentes. Ailes inférieures noires, avec

trois rangées paralleles de taches mal alignées, dont celles des deux premiéres séries

sont blanches, plus ou moins cerclées de roux vif et celles du bord marginal, qui

sont presque pontiformes, d’un fauve orangé ainsi que la frange de langle anal:

chez la femelle, on voit sur les secondes ailes une tache fauve entre le dise et la

base.” Expans. alar. antic. 12 centim.

Castnia argus, Boisduval, H.n. Lép. Hét. p. 522.

_ This species, which I examined in Dr. Boisduval’s collection, appears to me to be

scarcely different from C. mygdon, being only of a larger size, and with a few more spots

towards the base of the hind wings.

Hab. Brasilia. In Mus. Boisduval.

37. CASTNIA MYGDON. C. alis anticis olivaceo-fuscis viridi nitentibus ; basi, faseia irre-

gulari obliqua e medio costz ad angulum internum extensa (ramum internum paullo

‘pone medium emittente ad medium marginis interni currentem), maculaque magna

SECOND SERIES.—ZOOLOGY, VOL. I. | 2B

180 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

costali subapicali (guttas tres subhyalinas includente) margineque apicali obscurio-

ribus ; alis posticis nigricantibus, inter medium et marginem posticum fasciis duabus

irregularibus e maculis subovalibus albidis aut flavidis formatis maculisque nonnullis

plus minusve obscurioribus versus medium disci positis. Long. corp. unc. 14. Exp.

alar. antic. unc. 25-3 (8-12 centim.).

Castnia mygdon, Dalm. Mon. p. 20, pl. 1. f. 2; Act. Holm. 1824. p. 403; Gray, Trans. Ent. Soc. ii.

p- 145; Walker, List Lep. Het. B. M. 1. p. 28.

C. phalaris, Godart, Enc. Méth. ix. 799 (nec Fabricius, &c.); Dalman, Monogr. Castn. p.22; Act, Holm.

1824, p.405; Gray, Trans. Ent. Soc. Lond. 11. p. 145; Boisduval, H. n. Lép, Hét. p. 521.

Sympalamides mimon, Hiibner, Samml. exot. Schmett. figs. 1, 2.

Hab. Brasilia, Parana, Papagaya. In Mus. Britann., Hopeiano Oxoniz, Boisduval.

38. CASTNIA CHELONE, “ C. alis anticis maris cinereis, foeminze fusco-cinereis, fasciis

duabus, maris murinis obsoletis, foeminze albis ; posticis aterrimis, cyaneo-micantibus,

fascia sinuato-angulata, fusca albido-marginata.”’ Exp. alar. antic. ¢ une. 2 lin. 9

(7 centim.), $ 3” 5” (9 centim.).

Castnia chelone, Hopffer, Neue Schmett. Heft ii. tab. iv. figs. 1, 2; Boisduval, H.n. Lép. Hét. p. 530.

Hab. Mexico. In Mus. Berol.

39. CastniA MENneErrigst. “Ailes supérieures pointues au sommet, d’un brun noir assez

obscur, avec deux bandes transverses d’un noir plus foncé, dont l’antérieure bifide

et la postérieure sinuée ; outre cela, ces deux bandes paraissent réunies par un rameau

latéral ; le long de la céte on voit aussi deux taches plus pales que la teinte générale.

Ailes inférieures noires, marquées vers leur milieu d'une grande tache jaune trans-

versale, sinuée en avant, suivie en arriére, du cété de ’angle externe, de quatre 4

cing points orangés, alignés et d’un autre point semblable rejeté en dehors. La

goutticre abdominale est blanche.”

“A peu pres de la taille de la femelle de C. Boisduvalii, dont elle a le port.”

Castnia Menetriesi, Boisduval, H. n. Lép. Heét. p. 511.

Hab. Brasilia. In Mus. Boisduval.

49, Castnia Humpoupti. “ Ailes supérieures d’un brun noirdtre, avec un reflet olivatre

tres-prononcé; elles sont traversées du milieu de la céte a langle anal, par une

bande blanche, atténuée inférieurement; elles ont, en outre, du coté du sommet,

une bande courte, flexueuse, maculaire, bien nette, composée de cing taches un peu

oblongues. Ailes inférieures d’un brun roux & leur extrémité, traversées par une

bande d’un blanc pur, sinué en dedans et s’élargissant insensiblement, pour arriver

4 Vangle anal; leur frange est entiérement blanche. Un peu plus grande que

V Atymnius, avec les ailes plus pointues au sommet.”

Castnia Humboldti, Boisduyal, H. n. Lép. Hét. p. 528.

Obs. Fascia 5-macularis subapicalis alarum anticarum, in individuo Boisduyalliano

guttas duas subcostales albas supra gerit; heec fascia in individuis Muszei Gallici e ma-

culis fere obsoletis constat.

Hab. Nova Granada. In Mus. Parisiensi et Boisduval.

GENUS CdASTNIA AND SOME ALLIED GROUPS. 181

41. Castnra sazasta. “Ailes supérieures d’un brun noirdtre, 4 reflet verdAtre: elles

sont marquées d’une bande linéaire trés-étroite, ou plutét d’une raie enfumée, allant

du milieu de la cdte a angle interne; ou voit, en outre, vers le sommet, mais trés-

faiblement, l’empreinte de la bande sinuée du dessous. Ailes inférieures noires vers

la base, avec un reflet vert, brunes 4 l’extrémité a reflet rougeatre, elles sont tra-

versées, comme dans les espéces du méme groupe, par une bande d’un blane pur,

mais & reflet violatre, légérement sinuée en dedans, et s’élargissant insensiblement

pour gagner l’angle anal. Frange des quatre ailes entiérement noire.”

De la taille de ? Atymnius avec les ailes plus pointues au sommet.

Castnia salasia, Boisduval, H.n. Lép. Hét. p. 529.

Hab. Mexico. In Mus. Boisduval.

42. CASTNIA EVALTHE. - C. alis latis, nigro-fuscis, anticis fascia media obliqua ad angu-

Tum analem extensa (ibique attenuata) alteraque subapicali tenuiore (interdum obso-

leta) albis aut flavis; posticis fascia media maculari vel macula magna discoidali

flava (interdum externe rufo late marginato) fasciaque maculari submarginali

anguloque anali late rufis. Expans. alar. ant. unc. 24-22 (7-10 centim.).

Var. a alis anticis bifasciatis, alis posticis fascia media submaculari flava, angulo anali

et maculis 5 vel 6 submarginalibus rufis. (Cramer, pl. 17. f. E, F.) Surinam;

Keuador. In Mus. Hopeiano.

Var. 8 maculis discoidalibus in alis posticis in maculam integram majorem coalitis.

(C. euphrosyne, Perty, Delect. pl. 31. f.1; Walker, List Lep. Het. B. M. i. 15.)

Brasilia et Demerara. In Mus. Hopeiano.

Var. y differt e var. 8 macula discoidali flava in alis posticis extus rufo late marginata,

colore rufo cum maculis submarginalibus plus minusve coalito. (Dalman, tab. i.

f. 3.)

Var. 6 similis var. a, at fascia subapicali in alis anticis obliterata maculisque fascize

medize posticarum angulo externo proximis fere obsoletis.. (C. Viryi, Boisduval,

p. 515, pl. 11. fig. 8.) Mexico et Nicaragua. In Mus. Hopeiano. Sore

Papilio evalthe, Fabricius, Syst. Ent. p. 480 ; Ent. Syst.iii.1, p.45; Donovan, N. Hist. Ins. India, Lep. pl.1;

Herbst, Icon. pl. 137. f. 1, 2; Boisduval, H. n. Lép. Hét. p. 514 (Castnia e.) ; Dalman, Monogr.

p- 14, tab. i. f. 3; Act. Holm. 1824, p. 401; Godart, Enc. Méth. ix. p. 798; Gray, Trans. Ent.

Soc. L. u. p. 143; Walker, List Lep. Het. B. M. i. p. 21.

P. dardanus, Cramer, i. pl. 17. f. E, F.

C. euphrosyne, Perty ut supra; Gray, op. cit. p. 144.

E. Viryi, Boisduval ut supra.

Hab. Brasilia, Para, Cayenne, Mexico, Ecuador. In Mus. Britann., Hopeiano Oxonize

(individua 8), Boisduval. .

43. CASTNIA AMycuS. C. alis anticis nigro-fuscis pallidius variegatis, squamis magnis

griseo-albis indutis, puncto seu fasciola subobsoleta ad apicem cellule discoidalis

fasciaque plus minusve obsoleta alba vel albida pone medium cost ad angulum

analem extensa margine interno dentato, fasciolaque subobsoleta abbreviata sub-

apicali, margine postico ante medium fasciculo elongato squamarum instructo ; alis

2B2

182 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

posticis nigris, medio disci ferrugineo fasciaque pone medium e maculis parvis plus

minusve obsoletis maculisque nonnullis submarginalibus rufis. Long. corp. lin.

91-14. Exp. alar. antic. unc. 13-3.

- Variat disco alarum posticarum omnino nigro, maculisque rufis multo minoribus fere

obsoletis.

Papilio amycus, Cram. iii. pl. 227, D, E.; Dalman, Monogr. p. 19 (Castnia a.); Act. Holm. 1824, p. 408 ;

G. R. Gray, Trans. Ent. Soc. L. ii. p. 145; Godart, Enc. Méth. ix. p. 798; Walker, List Lep.

Het. B. M.i. p. 27; Boisduval, H.n. Lép. Hét. p. 504.

Obs. As this species differs in the greater or less amount of red markings on the hind

wings, [am much inclined to consider the C. meditrina of Hopfier to be an extreme

variety in which the red has entirely disappeared. The less acute apex of the fore

wings relied upon by Boisduval appears to me to be the sexual (female) character.

Hab. Surinam, Brazil, Papagaya. In Mus. Britann., Hopeiano Oxonie (7 individua),

Boisduval.

44. CASTNIA MEDITRINA. “ C. alis anticis virescenti-nigris, fascia undulata obliqua gut-

tisque ad costam tribus albis ; posticis aterrimis cyaneo micantibus.” Expans. alar.

ant. une. 2 lin. 5.

The hind margin of the fore wings has the dilated patch of elongated scales noticed

in the description of the preceding, and to which neither the description of Hopffer

nor that of Boisduval allude.

Castnia meditrina, Hopfier, Neue Schmett. Heft i. tab. v. f. 4; Boisduval, H. n. Lép. Hét. p. 505.

An var. C. amycus ?

Heb. Brasilia. in Mus. Berolini.

45. Castnta HvBNERI. Calis anticis fulvo-fuscescentibus, fascia maculari plus minusve

continua e medio coste ad angulum internum extensa alteraque parallela e ma-

culis sex magis distinctis inter medium et apicem albis; alis posticis basi mar-

gineque lato abdominali fuscis, dimidio costali rufo-fulvo; dimidio externo nigro,

maculis 7 cuneiformibus pone medium irregulariter dispositis, serieque submarginali

macularum rufo-fulvarum; fimbria lata pallide fusca.

Casinia Hiibneri (Boisduval, MS.), Latreille in Cuvier, R. An. 2d ed. iii. pl. 20. f. 2, p. 489; Gray,

Trans. Ent. Soc. Lond. i. p. 146; Walker, List Lep. Het. B. M.i. p. 30; Boisduval, H. n. Lép.

Hét. p. 516.

Castnia Sternbergii, Kollar, Lep. Bras. Ann. Wien, 1. 216, pl. 12. f. 4.

Synemon Sternbergii, Mikan, Delect. Fl. et Fr. Brasil. ; Schaufuss, Nunquam Otiosus, p. 11.

Hab. Brasilia. In Mus. Britann., Hopeiano Oxonie, Boisduval.

46. CASTNIA JUTURNA. C. alis anticis fuscis, basi et margine interno griseo-squamosis,

fasciis 4 albidis, duabus anticis obliquis, e costa, marginem posticum et externum

haud attingentibus, duabus posticis brevioribus magis transversis, 3ta e maculis

obcuneatis composita, 4ta venis rufis in maculas divisa; alis posticis basi fuscis,

raacula magna nigra sanguineo cincta, medio fascia alba valde sinuata sanguinco

GENUS C4STNIA AND SOME ALLIED GROUPS. 183

extus marginata; dimidio externo alarum nigro, fascia maculari subapicali san-

guinea, maculis versus angulum externum albo irroratis. Long. corp. unc. 13. Ex-

pans. alar. antic. unc. 3/4.

Castnia juturna, Hopffer, Neue Schmett. pl. iv. f.3; Boisduval, H. n. Lép. ee. p. 508.

Castnia lorze, Boisduval, MS. olim.

Hab. Brasilia (Minas). In Mus. Hopeiano Oxoniz, Boisduval.

47, CASTNIA CRONIS. C. alis anticis nigris, costze basi maculisque thoracis rufis vel

albis, plaga magna discoidali postice dilatata, alteraque subcuneata in medio coste,

fascia abbreviata obliqua e maculis tribus composita pone medium, altera e maculis 9

ovalibus submarginalibus albis, imargine ipso albo guttato ; alis posticis abdomineque

albidis, margine apicali tenui nigro, intus crenulato. Expans. alar. antic. unc. 3-34.

Papilio cronis, Cramer, i. pl. 60. f. C. ??, pl. 178. f. A; Oliv. Enc. Méth. Ins. pl. 12. f.7; Dalman, Mo-

nogr. Castn. p. 23 (Castnia cr.) ; Act. Holm. 1824, p. 504; Godart, Enc. Méth. ix. 801; Gray,

Trans. Ent. Soe. L. ii. p. 146; Walker, List Lep. Het. B. M.i. p. 29. n. 34; Boisduval, H. n. Lép.

Hét. p. 534.

Obs. M. Boisduval, H. n. Lép. Hét. p. 534, considers that the figure given by Cramer,

i. pl. 60. f. C, as the male of C. cronis (Pieris cronissa, Godart, Enc. M. ix. Suppl. p. 814;

Pieris cronis, Boisduval H. n. Lép. i. p. 548) is manufactured out of the head, abdomen,

and hind wings of a Pieris attached to a mutilated thorax and fore wings of a Q. cronis.

Hab. Surinam. In Mus. Britann., Loddige et Herrich-Schiffer (valde mutilata).

48. CasTNIA cronipDA. “ Ailes supérieures noires, avec des taches blanches disposées

a peu prés comme dans la cronis, savoir, une trés-grande longitudinale sur le milieu,

une médiocrement grande sur la cdte et les autres plus petites 4 l’extrémité;

ces derniéres sont alignées au nombre de quatre ou cing seulement, et sont pré-

cédées vers le sommet de trois autres taches blanches allongées, il y a, en outre, sur

les nervures, une série de petits points blancs, trés-peu indiqués; Torigine de la

céte est rouge comme dans la cronis. Ailes inférieures d’un blane un peu jaunatre,

avec une large bordure noire, divisée par un cordon de gros points d’un blanc un

peu jaunatre. Le corselet est noir, marqué de deux points rouges, avec le collier

blane interrompu; l’abdomen est d’un blanc jaundtre avec l’extrémité d’un jaune

d’ocre.”’

“La femelle ne differe du male que par sa taille plus grande et par la bordure des ‘ailes

inférieures plus large avec des points plus gros.”

- Expans. alar. antic. g 8}, 9 103 centim.

Castnia cronida, Herr.-Schafler, Samml. aussereur. Schm. fig. 142; Boisduval, H. n. Lép. Hét. p- 535 2

Walker, List Lep. Het. p. 1581.

Except in the less complete development of the subapical row of spots on the fore

wings, and the wider black margin of the hind wings, this supposed species is evidently

identical with C. cronis, which appears, from Gray’s description of Mr. Loddige’s spe-

cimen, to be subject to a certain amount of variation.

Hab. Guyane francaise. In Mus. Stuttgard et Boisduval.

184 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

49. CASTNIA LicoipEs. “Elle a tout-a-fait le port et la taille de lespdce précédente

(Zicus) dont elle est trés-voisin, mais la massue de ses antennes est jaune. Ses ailes

supérieures sont de méme d’un brun-noir chatoyant en vert, avec une bande linéaire

blanche dentée en dehors, allant du milieu de la céte 4 l’angle anal. Il n’y a de bande

sinuée & l’extrémité; la frange est blanche. Les ailes inférieures sont d’un noir-brun,

traversées dans leur milieu par une bande blanche a reflet un peu violet, dentée en

dehors, s’élargissant insensiblement pour arriver 4 langle anal; le bord terminal est

divisé par un cordon de sept taches rouges, dont l’anale est double et plus petite.”

Castnia licoides, Boisduval, H. n. Lép. Hét. p. 527.

Urbanus licus, Hiibner, Samml. exot. Schmett. v. i. Celeb. a, fig. 1. 2.

Hab. Brasilia, Santa Catherina. In Mus. Boisduval.

50. CASTNIA INORNATA. “ Ferrugineo-fusca, subtus cervina, abdominis dorso nigricante

maculis luteis ornato, alis anticis fasciis duabus fuscis obliquis indeterminatis, ma-

cula reniformi obscure cinerea fusco marginata.”’

«Female ferruginous-brown, fawn-colour beneath. Frontal tuft porrect lanceolate,

extending rather beyond the head. Antenne subclavate; tips pale cinereous. Abdomen

blackish above, shorter than the interior border of the hind wings, each segment with a

transverse luteous spot. Fore wings with two irregular oblique brown bands, which are

diffuse on the outer side, and have between them a reniform dark cinereous mark, which

is bordered with brown; underside with two irregular dark brown bands beyond the

middle, the first composed of patches, the second much abbreviated hindwards, marginal

space fawn-colour. Hind wings with a fringe of very long hairs, which extends across the

disk from near the interior border. Hind wings beneath with an irregular abbreviated

ferruginous line across the disk, including at its fore end a whitish dot.” Length of the

body 10 lines, of the wings 31 lines.

Hab. ? In Coll. T. Chapman, Glasgow.

Castnia inornata, Walker, Cat. Lep. Heteroc. p. 106.

This species belongs to the group of which Chremes is the type.

51. CastNIA PENELOPE. ‘ Dem Icarus in Form und Zeichnung ganz ahnlich, die Ober-

seite aber in allem blasser, die Zcichnung auf den Hinterfliigeln braun statt schwarz,

die fussere Querbinde der Oberfliigel sowie weissen Tropfen der Unterfitigel fast:

verloschen : auf der Unterseite der Oberfliigel alle drei Binden bestimmt ausgepragt,

im Uebrigen opalisirend wie Zcarus und veilleicht nur Varietét.” Lat. 78 mm.

Hab. Brasilien.

Castnia penelope, Schaufuss, Nunq. Otios. p. 9.

52. CASTNIA ALBOFASCIATA. ‘“ Grosser als mimon, oben ebenso gezeichnet, die Oberfliigel

jedoch heiderseits der braunen Querbinden mit weisser, schmaler Kante eingefasst

und mit verwischtem Weiss ausgefillt. Unterfliigel mit drei weissen Flecklagen,

dieselben mit Schwarz und Zimmetbraun eingefasst, ahnlich wie bei Phalaris ; doch

GENUS CASTNIA AND SOME ALLIED GROUPS. 185

ist das Weiss uberwiegend ausgebreitet; unten dem letateren sehr ahnlich (vedlleicht

nur dessen 9 ?).” Lat. 88 mm.

Hab. Brasilia. In Mus. Kaden olim.

Castnia albofasciata, Schaufuss, Nunq. Otios. p. 10.

| Obs. The three following species form the genus CHRETES of Boisduval. See

supra, pp. 166, 167.]

53. CASTNIA CHREMES. C. alis anticis apice acutis, fuscis aut brunneo-fuscis, basi parum

pallidioribus, fascia obliqua pallidiore e medio costze ad medium disci extensa inde

dilatata, parte basali versus medium marginis postici extensa, parte externa ad

angulum posticum extensa, fascia altera abbreviata subrecta pallida inter priorem et

apicem, strigaque valde indistincta undulata prope marginem posticum: alis pos-

ticis maris nigris purpureo lete nitidis, punctis duobus (vel 1) costalibus, margine

ad angulum externum punctisque nonnullis submarginalibus late aurantiaco-rufis ;

foeminze rufo-aurantiacis, fasciis duabus maculisque marginalibus nigris: alis anticis

subtus pone medium fascia obliqua recta nigra in utroque sexu. Expans. alar.

antic. ¢ unc. 23-84, 2 unc. 3-34.

Papilio chremes, Fabricius, Ent. Syst. 111.1. 47 9 ; Jones, Icones, iii. pl. 72. f. 2; Walker, List Lep. Het.

B. M.1. p. 31 (Castnia chremes) ; Boisduval, H.n. Lép. Hét. p. 536 (Ceretes chremes) .

Corybantes nicon, Hiibner, Samml. exot. Schmett. ii. Versic. 1. f. 1, 4; Gray, Trans, Ent. Soc. L. ii,

p. 148.

Castnia thalaira, Godart, Enc. Méth. ix. 800 (2).

Castnia Fabricii, Swainson, teste Bdv. op. cit. p. 539.

Papilio thats, Drury, Il. ii. pl. 16. f. 4 (aot of Walker).

Foeem. var. major (exp. alar. antic. unc. 33). “‘ Ferrugineo-fusca, aurato viridique micans ;

pectus et abdomen aurato-ochracea; ale antic fasciis tribus abbreviatis fuscis ;

postice late ochraceze, subpurpurascentes, postice atre fasciis duabus arcuatis

ochraceis, 1ma submaculari, 2da maculari.”

Castnia morphoides, Wik. List Lep. Heter. B. M. pl. 31. f. 48, = C.nicon, Hib. Exot. Schmett.

Length of wings 44 lines. “ Nearly allied to C. icarus; but the markings of the fore

wings are different.”

Brazil. In Mus. Brit.

Foem. var. minor. (exp. alar. antic. unc. 2-24).

Castnia thats, Boisduval, op. cit. p. 537.

Hab. Brasilia, Minas Geraes, Rio Janeiro, between Sapezal and Conceicao (Burchell,

30 October, 1827). ‘‘ Umbroso sylvz dense, volatu et coloribus Phalene,” Burchell,

MS. In Mus. Britann., Hopeiano Oxonie, Boisduval.

From the great diversity of the sexes of this and the following species, and the variable-

ness in the intensity of their markings, there has been great confusion in their nomen-

clature.

The description of C. chremes (female) given by Fabricius is scarcely sufficient to

distinguish it from the smaller species, although his words “ fasciis duabus” would imply

simple and not irregular stripes, and his description of the fore wings, “ subtus fere con-

186 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

colores et dilutiores,” apply to it better than to the female of C. Marcelserresii. Mr.

Walker, from an inspection of the original drawings of Jones’s ‘ Icones.’ Danai, pl. 72,

referred the Fabrician insect to the C. nicon of Hubner. A copy of the figure of Jones

was made by Donovan, and is preserved in the Hopeian collection at Oxford. By the

kindness of Mr. Drewitt, the relative of Mr. Jones, and the present possessor of the

volumes of his ‘ Icones,’ I have been enabled to verify the figure and insect, the fascia

beyond the middle of the fore wings being quite straight, and extending to the anal

angle; the apical margin of the fore wings is but slightly rounded; and the black edge

of the hind wing does not extend to the outer angle. iS

The figure of Drury of his Pap. thais (iii. pl. 16. f. 4) exhibits several characters

hitherto unnoticed, which clearly identify it with Hitbner’s figure of the female of

C. nicon. These are the subtruncate (instead of strongly rounded) apical margin of the

fore wings, the curved (instead of quite straight and transverse) inner edge to the

central pale fascia of the fore wings, and the fulvous outer angle of the hind wings, towards

which the second black fascia emits two or three short strigze or small oval spots. In

the females of C. Marcelserresii these strigee or spots are united to the marginal fascia,

which is extended to the outer angle.

The males of the two species are at once distinguished by the undersides of their fore

wings being broadly black in the male of C. chremes and pale brownish grey in that of

C. Marcelserresi.

ITecannot but think the unique female described by Boisduval as agreeing with

“la figure donnée par Drury trés-exacte,” is a very small female of ¢hais, measuring

only 5 centimetres in the expanse of the fore wings, whereas the female which he

describes of thais (his C. chremes) measures 9 centimetres.

The Oastnia morphoides of Walker is certainly only a large female of C. thats, agreeing

closely with Hiibner’s figure (wicon 2).

The following is the description of an insect from Georgia, in North America, which

Herr Schaufuss considers may be a variety of this species.

“ (. thais, var.? Schaufuss {[Nunq. Otios. p.10]. Viel kleiner als Thais, Dr. Die Binden

oben fast verloschen oder schmaler, unten orangegelb: Oberfligel mit nur einer

aus dreieckigen Flecken zusammengesetzen Binde, die in die Randflecken, welche

oben nur angebraunt erscheinen, verlauft.” i

Hab. Georgia, America.

54. CASTNIA MARCELSERRESII. C. alis anticis ovatis (margine apicali versus apicem

rotundato), livido-fuscis aut fusco-fulvescentibus, fascia recta obscura e medio costz

ad medium marginis interni extensa, macula magna obliqua obscura intus subito

dilatata et interdum cum fascia irregulari e costa versus angulum analem extensa

connexa, apice obscuriore et parum angulato: alis posticis maris fuscis, serie

subapicali lunularum pallidiorum vix distinctarum (singulis punctum nigrum in-

cludentibus) : alis posticis foeminz rufo-fulvis, fasciis tribus undulatis nigris (ultima

marginali intus dentata): alis anticis subtus in utroque sexu pone medium macula

magna bipartita costali fasciaque subapicali plus minusve distincta nigris. Long.

corp. une. 1. Expans. alar. antic. unc. 23-3.

GENUS CASTNIA AND SOME ALLIED GROUPS. 187

Mas. Castnia Marel-serresii, Godart, Enc. M, ix. 800; Geyer, in Hiibner, Zutrige exot. Schm. iii. f. 711,

712; Gray, Trans. Ent. Soe. ii. p. 148 (excl. syn. P. thats, Drury).

Fem, C. Fabricii, Godart, Enc. M. ix. 800; Boisduval, H. n. Lép. Hét. p. 538 (nec Swainson).

C. thats, Walker, List Lep. Het. B. M. i. 31 (not of Drury).

Hab. Brasilia. In Mus. Britann., Hopeiano Oxoniz (5 mares, 6 foem.) &c.

The male of this species was described by Godart under the name adopted above, and

the female under that of Fabricii, with the observation that “ce Lépidoptére parait étre

Pun des sexes de celui que M. Swainson a figuré sous le nom de C. Fabricii.”” This sup-

position being erroneous, I have not thought proper to follow M. Boisduval in retaining

the name of C. Fabricvi for this species, but have adopted the usual plan of preferring

the name given to the male insect, C. Marcel-serresit. Gray, followed by Walker, erro-

neously considered the P. thats of Drury to be identical with the female of this species.

55, CasTNIA Diva. C. alis anticis apice acutis, supra nigro-fuscis, fascia latissima albida

ferrugineo-squamosa, e medio coste ad angulum analem extensa, pone medium sub-

interrupta, macula parva discoidali, alterisque parvis ovalibus pone medium albis

aut albidis brunneo circumdatis squamisque nonnullis subapicalibus ferrugineis :

alis posticis basi leete purpureis, fascia lata pone medium nigra, margineque postico

versus angulum externum aurantiaco intus undato. Expans. alar. antic. unc. 23-44.

Castnia diva, Butler, Lepid. Exot. pl. 17. f. 1, 2.

C. tricolor, Felder, Reise Novara, Zool. Bd. ii. Lep. lxxix. fig. 3 (sine descriptione) ; Boisduval, H. n. Lép.

Hét. p. 531.

Hab. Nicaragua, Columbia. In Mus. Britann., Hopeiano Oxonie, Felder, Boisduval.

Scetion Il, Species with the fore wings elongate-ovate. (See supra, p. 167, and

Plate XX XT.)

= Genus GAZERA, Boisduval, in H.-Schaffer, Samml. aussereur. Schm. ; Hist. nat. Lép. Hét. p. 546.

56. (1) Casrnia srmunans. “ Ailes entiéres, les supérieures d’une teinte roussdtre

enfumée depuis la base jusqu’au-dela du milieu, avec une tache oblique d’un brun

noiratre & lextrémité de la cellule discoidale: la partie postérieure de ces mémes

ailes est d’un brun noir, coupée par une bande oblique, droite, partant de la céte et

n’arrivant pas jusqu’au bord externe, outre celail y a, & partir de angle interne, une

rangée de trois ou quatre points jaunatres qui ne remonte pas jusqu’au milieu du

bord extérieur; le bord interne, la nervure médiane et la cdte sont plus ou moins

brunatres. Ailes inférieures d’un fauve roussitre, traversées au-delA du milieu par

une bande noire maculaire, elles ont en outre a l'extrémité une espéce de bordure

noiratre formée des taches cunéiformes ; cette méme bordure offre vers l’angle anal

deux points jaunatres et un point semblable vers l’angle externe. Taille de

Lycorea ceres.”

Gazera simulans, Boisduval, H. n. Lép. Hét, p. 547, pl. 11. fig. 4.

This species is the representative of Zycorea ceres, one of the Heliconian butterflies.

It is closely allied to, if not a variety of C. personata, Wk.

Hab. Columbia. In Mus. Boisduval.

SECOND SERIES.—ZOOLOGY, VOL. I, 2¢

185 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

57. (2) CAsTNIA PERSONATA. “ Fusca, aureo viridi micans, subtus flava; antennze apice

flavescentes ; abdomen subauratum vittis duabus ochraceis : ale cervine, sat angustz,

lituris nigro-fuscis, fascia marginali flavescente maculata; anticee vitta discali,

strigas quatuor emittente, postice vittis duabus parallelis extus abbreviatis. Nearly

allied to O. linus. Length of wings 36 lines.”— Walker. ~

The above description is unintelligible, as may be seen by M. Boisduval’s attempt to

translate it into French, and by his inability to perceive its apparent identity with his

Gazera simulans. The general colour of the four wings is obscure luteous, with black

markings; these consist of an oblique patch, arising on the costa before the middle,

dilated towards the middle of the wing into an elongated patch; beyond the middle of

the costa is a broad oblique band reaching to the middle, followed by four more distinct

luteous longitudinal spots preceding the broadly black apex ; the inner margin is dusky :

the hind wings are rather lighter luteous in the middle of the disk; the costa, a central

bar, and the apical margin black, the latter with four white dots on its inner edge near

the outer angle. Expans. alar. antic. lin. 36.

Castnia personata, Walker, List Lep. Het. B. M. xxxi. p. 48; Boisduval, H. n. Lép. Hét. p. 546 (Gazera

personata).

Hab. Guyaquil. In Mus. Britann.

58. (3) CasTNIA cononiA. (Plate XXXII. fig. 5.) Alis anticis elongato-ovalibus apice

rotundatis, dimidio basali obscure luteo-fulvo, striga postcostali plagaque magna

prope marginem internum nigris, hac striolam pallidiorem includente; macula

media (prope costam) postice angulata ejusdem coloris fasciaque obliqua (e maculis 5

ovalibus constante) inter medium et apicem alz albida; alis posticis saturate fulvis,

fascia postcostali maculis quinque ovatis pone medium, ad marginem analem majori-

bus et connexis, limboque apicali nigris ; capite albo punctato, collo postice fulvo ;

fascia transversa media thoracis flavida; abdominis fusci lateribus anguste fulvis,

segmentis ventralibus basalibus et apicalibus fulvis, intermediis nigris. Long. corp.

unc. 14. Exp. alar. antic. une. 33.

Hab. Ecuador. In Mus. Hopeiano Oxonie (olim Saunders).

This species is nearest allied to C. ecuadoria, with which the arrangement of the cell

and branches of the postcostal vein of the fore wings nearly agrees; but the transverse

vein closing this cell is longer, in consequence of which the branches 3 and 4 arise

wider apart.

59. (4) Castnza cratiInA. (Plate XXXII. fig. 4.) Alis anticis oblongo-ovatis apice

subrotundatis fulvis; striga costali prope basin, altera subcostali tertiaque multo

majore prope marginem internum, striga undulata obliqua discoidali, macula rotunda

media cum margine antico connexa, plaga magna obliqua inter medium et apicem,

limboque apicali dentato nigris ; alis posticis aurantiaco-rufis, striga subcostali, fascia

lata media e margine anali fere ad angulum externum extensa limboque dentato

nigris ; capite albo maculato, collo lato fulvo fascia transversa tenui flavida ; abdomine ©

GENUS CASTNIA AND SOME ALLIED GROUPS. 189

nigricante lateribus segmentisque 5 apicalibus subtus fulvis; thorace subtus albido

vario; antennis albidis. Long. corp. unc. 13. Expans. alar. antic. unc. 43.

Hab. Amazonia. In Mus. Hopeiano Oxoniz (olim Saunders).

This species is closely allied to C. cycna, but has the fore wings less rounded; the

arrangement of the cell and branches of the postcostal vein agrees with that of

C. cycna.

60. (5) Castnra EcuaDoRIA. (Plate XXXII. fig. 6.) Alis anticis oblongo-ovatis apice

- rotundatis; e basi ultra medium obscure lutescentibus, apice lato fusco; cellula

basali fusco marginata, macula media rotunda subcostali, striga obscura media,

alteraque majore marginis interni fuscis ; alis posticis obscure fulvis nebula magna

ex angulo anali versus medium ale extensa fimbriaque fuscis ; corpore fusco ; collari,

fascia media transversa thoracis, lateribusque abdominis lutescenti-albidis ; antennis

lutescentibus: alis subtus magis rufescenti-fulvis, apice fusco, posticis stria tenui

transversa subbasali fusca. Long. corp. unc. 14. Expauns. alar. antic. unc. 34.

Hab. Ecuador. In Mus. Hopeiano Oxonie (olim Sawnders).

This species so closely resembles C. ¢ruailla in general form and appearance that it

might easily be considered to be a variety of that species; but the arrangement of the

veins proves them to be totally different, there being a very distinct elongate-ovate cell

in C. ecuadoria, formed at the base of the branches of the postcostal vein, the 2nd, 3rd,

4th, 5th, and 5*th being emitted at nearly equal distances apart at the extremity of the

cell (Plate XXVIII. fig. 10).

61. (6) Casrnta zAGRmA. “ Ailes supérieures noires avec une bande longitudinale fauve,

sinuée, un peu irréguliére, coupée par les nervures partant du milieu de la base et

se prolongeant vers le bord postérieur ; outre cela, il y a sur le milieu de la céte un

petit trait, et au-dessous de celui-ci une tache un peu triangulaire de la méme

couleur, divisée par les nervures, l’extrémité postérieure offre deux rangées paralléles

de taches blanches lavées de jaune; mais la premiére rangée ne descend pas au-dela

du milieu de Vaile.” Expans. alar. antic. une. 44.

Gazera zagrea, Felder, Reise d. Novara, Zool. Bd. ii. Abth. 2, pl. 79. f. 2 (sine descr.) ; Boisduval,

H. n. Lép. Hét. p. 547.

Hab. Panama. In Mus. Felder.

This is the Heterocerous counterpart of Papilio zagreus, which isitself the counterpart.

of one of the Heliconian butterflies.

. Nearly allied to C. cratina, Westw., but with the fulvous markings of the fore wings

and the black markings of the hind wings much less developed in size. Tore wings

black, with a longitudinal fulvous vitta of irregular width, extending from the base of

the wing along its middle, rather deflexed and narrowed at its extremity; considerably

beyond the middle of the wing an oblique cuneated patch of the same colour arises at

the middle of the costa, followed by a row of six suboval spots, the lowest of which ranges

with the extremity of the central bar; beyond these is a submarginal row of nine white

2c2

190 | MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

spots, the middle ones being the smallest ; hind wings fulvous, costa narrowly black,

with a row of semiovate black spots beyond the middle, not extending to the outer

angle; apical margin black and strongly dentate within, the teeth near the anal angle

uniting with the spots of the inner row. Described from the figure of Dr. Felder.

62. (7) CASTNIA TRUXILLA. (Plate XXXII. fig. 3.) Alis fuscis, anticis oblongo-ovatis

apice rotundatis macula magna media lutescente e costa oblique versus angulum

internum extensa, guttam parvam obscuram fuscescentem versus costam ineludente ;

eutta parva lutea prope angulum internum nebulisque duabus oblongis pallidioribus

prope marginem internum; alis posticis ovalibus fuscis, basi ipsa ad marginem ana-

lem squamis fulyis notata; corpore fusco, capite albo-maculato, patagiis fulvo-

marginatis ; abdomine fusco, lateribus anguste fulvis; abdomine subtus basi et apice

fulvis, segmentis intermediis nigris, griseo marginatis ; antennis fuscis, apice fulvis ;

alis subtus pallidioribus, costa posticarum fulva. Long. corp. une. 14. Expans.

alar. antic. unc. 33.

Hab. Columbia. In Mus. Hopeiano Oxoniz (olim Saunders).

This obscurely coloured species is remarkable for its elongated fore wings (which are

rounded at the apex) and its oval hind wings. In the fore wings there is no cell formed

by the basal portion of the branches of the postcostal vem; nor is there any transverse

connexion between the 3rd and 4th branches of the same vein. (See Plate XXVIII.

fig. 12.)

68. (8) CAsTNIA coLUMBINA. “ Ailes supérieures d’un noir foncé; elles ont ala base, deux

bandes longitudinales, d’un roux un peu enfumé, dont V’inférieure se termine par deux

rayons blanchatres: outre cela, il y a sur le milieu de la céte un petit trait blane et

au-dessous de celui-ci, prés de la principale nervure, une tache blanche arrondie;

plus bas et plus en dehors deux taches blanches cunéiformes, ’extrémité postérieure

offre deux rangées paralléles de taches blanches inégales, les unes rondes, les autres

allongées: ailes inférieures d’un fauve vif, avee une bordure noire crénelée, et une

rangée de points blancs; outre cela elles offrent au-dela du milieu deux ou trois

points noiratres disposés transversalement.”’ Hxpans. alar. antic. 12 centim.

Gazera columbina, Boisduyal, H. n. Lép. Hét. p. 546.

Hab. Nova Granada. In Mus. Boisduval.

Obs. This species is nearly allied to C. salvina, but differs in the markings of the hind

wings.

64. (9) CasTNIA SALVINA. (Plate XXXII. fig. 1.) Alis anticis oblongo-ovatis nigrican-

tibus, basin versus vittis duabus longitudinalibus obscure rufescentibus, maculisque

novem, magnitudine variis, discoidalibus, serieque submarginali macularum novem

albidarum (maculis apicem appropinquantibus longioribus) ; alis posticis fulvis, costa

limboque lato integro nigris, hoc maculas sex submarginales parvas albas includente :

capite et thorace nigris, illo albo maculato, patagiis albo marginatis : antennis fulvis,

GENUS CASTZNIA AND SOME ALLIED GROUPS. 191

basi obscurioribus: alis subtus similiter ac supra coloratis, posticarum costa et

striga parallela subcostali nigris: abdomine fulvo, apice obscuriore; subtus medio

sterni abdomineque albidis. Long. corp. unc. 13. Exp. alar. antic. unc. 43.

Hab. Veragua (Salvin). In Mus. Hopeiano Oxoniz (olim Saunders).

The arrangement of the branches of the postcostal vein in this species is peculiar, there

being no basal cell formed by the transverse junction of the middle branches; the third

branch springs from the second at a considerable distance from the base of the latter;

and the 4th and 5th branches arise at a considerable distance from the transverse base of

the 6th branch (i. e. 65*), which forms the termination of the discoidal cell.

65. (10) Castnia cyona. (Plate XXXII. fig. 2.) Alis anticis subovatis fusco-nigris,

basi, costa, striga media (e basi ad medium alz extensa) margineque postico obscure

fulvis, striga media subcostali obliqua, macula ovali discoidali, fascia obliqua inter

medium et apicem maculisque 9 magnitudine variis albidis; alis posticis lete au-

rantiacis, extus in flavum pallescentibus, limbo intus acute dentato nigro: capite

albo-punctato; antennis albidis basi nigricantibus: thorace fascia antica transversa

fulva maculisque duabus posticis albidis: abdomine fusco, lateribus ventreque

fulvis. Long. corp. unc. 1}. Expans. alar. antic. une. 4.

Var. minor (exp. al. ant. unc. 34) fascia basali alee antics et striga marginis postici

obsoletis, coloribus etiam magis vividis. ~ ;

Hab. Columbia. In Mus. Hopeiano Oxoniz (olim Saunders).

The broadly ovate fore wings and the richly coloured hind wings, with their strongly

dentated black border, distinguish this species, which agrees with O. ecwadoria in the

arrangement of the cells and branches of the subcostal vein of the fore wings.

66. (11) Castyia mimica. (Plate XXXII. fig. 7.) Alis fusco-nigricantibus, certo situ

parum metallicis, anticis elongatis basi obscurioribus, squamis nonnullis rufis juxta _

tegulas; posticis macula magna transverso-oyali inter medium et angulum analem

flavescente: capite nigro, punctis duobus ante antennas, linea tenui postica pone

oculos, thoracis macula minuta antica albis; tegulis rufis; abdominis segmentis

intermediis lateribus rufo angulariter maculatis, squamisque albis strigas breves

laterales ad basin segmentorum formantibus: antennis nigris, clava subtus fulva:

coxis anticis et 2 posticis geniculisque pedum 4 anticorum rufis: abdomine subtus

utrinque albo maculato, segmentis intermediis macula parva rufa in medio notatis.

Long. corp. unc.1j. Expans. alar. antic. une. 3 lin.

Hab. Amazonia. In Mus. Hopeiano Oxonie (olim Saunders).

C. mimica, Felder, Reise d. Novara, Ins. Lep. pl. 79. f. 4: (sine descript.).

This remarkable species is, as to its colours, unique in the group; its wings are also

of peculiar form, giving it an aspect quite alien to its allies; it possesses the elon-

gated cell at the base of the branches of the subcostal vein; but the erd and 4th

branches arise at some distance beyond the cell. (Plate XXIX. figs. 11, 12.)

192 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

67. (12) Casrnra tinus. C. alis nigris hyalino-albido variis, venis nigris; anticis macula

magna subtrigona, venis divisa, fascia media e costa ad angulum internum api-

calem extensa in duas partes interrupta, macula magna ovali subapicali in sex

partes venis divisa, serieque submarginali punctorum alborum: alis posticis macula

maxima discoidali transverso-ovali, serieque externa macularum conoidearum magni-

tudine decrescentium versus angulum analem, ubi in puncta alba evadunt. Long.

corp. une. 14-14. Expans. alar. antic. unc. 83-4.

Variat venis in disco pallido alarum posticarum concoloribus nec nigris, maculis prope

angulum externum minoribus punctisque versus angulum analem fere obliteratis.

Papilio linus, Cramer, iii. pl. 257. fig. A; Herbst, Ins. pl. 78. f. 4.; Dalman, Monogr. p. 24; Act. Holm.

1824, p. 405 (Castnia linus) ; Gray, Trans. Ent. Soe. L. ii. p. 147; Walker, Cat. Lep. B. M. p. 32;

Boisduval, H.n. Lép. Hét. p. 544, f. 15.

Var. C. heliconioides, Herrich-Schaffer, Samml. exot. Schmett. pl. 4; Walker, op. cit. p. 1582; Bois-

duval, H. n. Lép. Hét. p. 545 (Gazera hel.).

Hab. Brasilia, Surinam, Cayenne, Guatemala. Mus. Britann., Hopeiano Oxoniz, ©

Boisduval.

Dr. Boisduval has separated two specimens in his collection from Cayenne from the

more ordinary Brazilian ones, considering the former the true representatives of

Cramer’s figure of C. linus from Surinam, and giving to the Brazilian insects a new

name, G. heliconides (recte Heliconioides, as previously published by Herrich-Schaffer).

I cannot, however, regard these insects otherwise than as belonging to one species, dif-

fering both individually and geographically, and even sexually: variations of the latter

character are exhibited in the form of the wings, which modifies to a certain extent

the form of the markings. Thus a female specimen in the Hopeian Collection from

Cayenne has the fore wings broad and almost ovate; and a male from the Amazons

(also with the veins of the disk of the hind wings concolorous) has the fore wings re-

markably long, as will be seen from the admeasurement of one of the fore wings of

these individuals as compared with those of two Brazilian specimens with black veins.

Veins of disk of hind Veins of disk of

wings concolorous. hind wings black.

Cayenne 9. Amazon. Brazilian 9. Brazilian 3.

Lengthofforewing .... .. . 48centim. 50 AT AS

\WVACRAN OVE Chg), Poe GBA Vat Ia AAD. bo 23 223 22

Length of outer margin from extreme

Apex tOnmMnerane |e. aael Ne eran siekime ome, 30 28 26

Length from base toinnerangle . . . 30 ,, dl 29 30

It will thus be seen that instead of the Brazilian individuals being “un peu plus grands

avec les ailes plus larges” than those of the Cayenne ones, as stated by Boisduval, the latter

are the largest in every respect, and more especially in the width of the wing.

The Cayenne female before me has the large spot of the hind wings more ovate,

the black anal margin extending further towards the base than in the Brazilian spe-

‘cimens; the veins in this large spot, in this female and in three other specimens in the

GENUS CASTNIA AND SOME ALLIED GROUPS. 193

Hopeian Collection, are concolorous, and not clothed with black scales as in the Brazilian

specimens; and the marginal spots near the outer angle of the hind wings are much

shorter than in the latter, whilst the small dots near the anal angle are reduced to a very

few scales. In the four Hopeian specimens, with pale veins in the disk of the hind wings,

the pale markings are less strongly yellow than in the Brazilian ones; and in the Ama-

zonian male above noticed the two apical spots of the terminal macular fascia are much

larger than the three following, whilst the three next are reduced to minute dots, the

one close to the inner angle being as large as the 8rd to 5th. Thus neither in size,

shape, nor markings can we specifically distinguish between the specimens of this

species.

68. (18) CasTNIA ACRMOIDES. C-alis anticis fuscis basi magis lutescentibus, medio fulvo

tinctis; venis omnibus dilute nigris, strigis longitudinalibus obscurioribus inter

venas fasciaque obliqua albido-ochracea e costa ultra medium, versus medium

marginis apicalis extensa; alis posticis aurantiaco-rufis, venis limboque apicali intus

dentato nigris, hoc serie macularum parvarum (subtus multo majorum) flavarum

notato. Long. corp. unc. 13. Expans. alar. ant. unc. 3 lin. 1.

Casinia acreoides, Boisduval, Sp. Gén. Lép.i. pl. 14. f.1; H. n. Lép. Hét. p. 540; Guérin, Icones R.

An. Ins. pl. 83. f. 4; Griffith, An. K. Ins. pl. 53. f.4; Gray, Trans. Ent. Soc, L. 1. p.148; Walker,

List Lep. Het. B. M. i. p. 32. ‘

Castnia actinophorus, Kollar, Lep. Brasil., Ann. Wien. i. p. 215, pl. 12. f. 2; Walker, List Lep. B. M.

i. p. 29; Boisduval, H. n. Lép. Hét. p. 520; “Gray (mn Ann. Kingd.) as C. acreoides,” Hrichs.

Berichte, 1839.

Hab. Brasilia. In Mus. Britann., Hopeiano Oxoniz, Boisduval.

This species bears so close a resemblance to Acrea thalia as often to be mistaken for

it. Boisduval states that its larva feeds on certain species of Bromeliacez.

It is surprising that Mr. Walker should not have perceived from Kollar’s excellent

figure that C. actinophorus (placed by Walker in his section 1 of subdiv. 2) is identical

with C. acreoides of Boisduval, which he places in his section 8. Deceived by this error,

Dr. Boisduval, who, like Walker, simply repeats or translates Kollar’s character of

actinophorus *, has also given the new supposed species as distinct, placing C. actino-

phorus in the genus Castnia, and C. acreoides in that of Orthia.

Genus IJ. Orruta, Boisduval (pars).

Ale magne, late ; antice subtrigone, cellula discoidali aperta, loco ven disco-cellularis tantum crassitie

minuta nodiformi ramorum 05* et c3* indicato, ramo supero discoidali (65*) e vena postcostali

pone cellulam emisso, cellula nulla accessoria postcostali: alee postice cellula angusta dimidiata vena

mediana connexa, cellula ordinaria aperta: abdomen alas imferiores haud postice superans.

(Plate XXIX. figs. 16, 17 +.)

* «¢ Alis anticis nigris radiis plurimis viridi-fuscis, basi fasciaque abbreviata dilute flavis: posticis luride rubris,

radiis limboque nigris, hoc flayo maculato. Long. corp. lin. 9-12. Exp. alar. ant. lin. 27-33.”

+ In figure 16 the small lettering of the two branches 6 5* and ¢3* is placed too high—the former branch, which

looks like-the terminal branch of the postcostal vein, and the latter branch, which looks like a fourth branch of the

194 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

Having restored C. therapon and acreoides (forming Boisduval’s first section of

Orthia) to Castnia, and O. augias and nexa belonging to the Agaristides, I have been

compelled to regard C. pelasgus, Fabr., and wnifasciata, Felder (which appears to me to

be identical with pelasgus), as a separate genus, for which I have, unwillingly, retained

the name of Orthia.

OrtHIA PELASGUS. Alis anticis nigris bronzeo-viridi tinctis, fascia lata lutescente aut

alba e medio costee ad angulum internum extensa, medio ejus paullo latiore apiceque

infero paullo incurvo ; alis posticis CL aoa, margine externo parum cerules-

cente. Expans. alar. ant. unc. 275-23.

Papilio pelasgus, Fabricius, Sp. Ins. ii. p. 54; Ent. Syst. i. pt. 1. p. 46 ; Herbst, Ins. pl. 136. f. 6; Oliv.

Ene. M. Ins. pl. 12. f. 5; Cramer, iii. pl. 202. fig. D; Dalman, Monogr. p. 21; Act. Holm, 1824,

p- 404 (Castnia p.) ; Godart, Enc. Méth. ix. p. 801; Gray, Trans. Ent. Soc. L. 1. p. 146; Walker,

List Lep. Het. B. M.i. p. 22; Boisduval, H. n. Lép. Hét. p. 54 (Orthia p.).

Castnia unifasciata, Felder, Reise d. Novara, Zool. Band ii. Abth. 2, pl. 79. f. 5 (sine deser.) ; Bois-

duval, H. n. Lép. Hét. p. 542 (Orthia unif.).

H.-Sch., Samml. exot. Sch. pp. 56 and 78, gives O. augias, H.-S., as = pelasgus, F.

Except in the white colour of the fascia of the fore wings, I can see nothing to separate

Dr. Felder’s figure from C. pelasgus of Cramer.

Hab. Brasilia, Guiana, Surinam, Ega, Bolivia. In Mus. Britann., Hopeiano Oxonize,

Boisduval.

Genus III. Synemon, E. Doubleday. (Plate XX XIII.)

In Lord’s Discov. in Australia; Klug, Abh. k. Akad. Berlin; Walker, List Lep. Heter. Brit. Mus. part i.;

Boisduval, H.n. Lép. Hét. 2

1. Synemon Sopuia. Nigro-fusca, alis anticis maris fuscis griseo-brunneoque nebulosis,

nebula obscura baseos alteraque submedia puncto albo adjecto; venis griseis, famine

nigricantibus squamis plumbeis irroratis, fasciam subapicalem maculis nigris divisam

formantibus, macula parva submedia fasciaque abbreviata costali maculari pone

medium : alis posticis maris basi nigris macula ovali ante medium, fascia maculari

pone medium maculisque submarginalibus flavis; famine nigris, macula magna

aurantiaca baseos, fasciaque lata abbreviata transversa marginis analis, macula

magna versus angulum externum, alterisque 3 vel 4 submarginalibus fulvis: alis

subtus luteis, fasciis ramosis nigris. Expans. alar. antic. lin. 18-22.

Hesperia? Sophia, White, in Grey’s Exped. Australia, App. p. 474, fig. 7; Doubleday, in Lord’s Discov.

Australia, App. i. p. 516, pl. 3. £. 5 (Synemon Sophia) ; Walker, List Lep. Het. B. M.1. p. 34, vii.

p. 1583; ma Abhandl. kon. Akad. Berlin, 1850, p. 248, pl. annex. f. 1,2; Boisduval, H. n. Lép.

Hét. p. 550.

Hab. Australia, King George’s Sound. In Mus. Britann., Hopeiano Oxonie,

Boisduval.

median vein, being separated from each other by the slenderly indicated fold running into the discoidal cell (the letters

c 3* in the plate being improperly placed opposite to its extremity), the closing of which by the transverse disco-

cellular yein is indicated by two small swellings, omitted by the engraver, opposite to each other in the middle of the

wing.

GENUS CASTNIA AND SOME ALLIED GROUPS. 195

2. SYNEMON THERESA. Griseo-fusca, disco subalbido, marginibus omnibus fuscis, venis

nigricanti-fuscis, in medio et ad marginem apicalem dilatatis, macula obliqua media

fusca albo extus marginata fasciaque subapicali e maculis ovalibus 4 et 3 fuscescenti-

bus inter venas obscuras formata; alis posticis fulvis basi fuscis, macula parva ante

medium fulva, margine externo tenui serieque macularum submarginalium fuscis.

Expans. alar. antic. lin. 18.

Synemon theresa, Doubleday, in Lord’s Discov. Australia, Append. i. p. 517, pl. 3. f.6; Walker, List Lep.

Het. B. M.i.p. 35, vi. p.84; Klug, Abhand. kon. Akad. Berlin, 1850, p. 249, pl. 3. £.5,6; Boisduval,

H.n. Lép. Hét. p. 552.

Hab. Nova Hollandia. In Mus. Britann., Hopeiano Oxoniz, Boisduval.

3. SyneMoN Lava. Nigro-fusca, subtus albida; antennis albo annulatis: alis anticis

fuscis, albido conspersis, maculis duabus irregularibus (una majore ante medium,

altera minore pone medium subcostali) albido separatis fasciaque apicali maculari

nigro-fuscis ; subtus aurantiacis, fusco maculatis : posticis aurantiacis, macula basali

fasciisque duabus nigris, prima arcuata, mediana, irregulari, secunda lata marginali,

serie guttarum fulvarum ornata; subtus albo guttatis. Expans. alar. antic. lin. 20.

crete leta, Walker, List Lep. Het. B.M.i. p. 36; Boisduval, H.n. Lép. Hét. p.553.

Hab. Nova Hollandia. In Mus. Britann.

4, SYNEMON SELENE. S. corpore fusco, subtus albido, abdomine supra griseo; alis

anticis fusco-griseis, medio late fuscis, ante apicem fusco maculatis, fascia maculari

transversa interrupta ante medium lunulaque media albis; posticis fuscis luteo tri-

fasciatis ; omnibus subtus albido-testaceis fusco maculatis: ¢ 2. Expans. alar. ant.

lin, 203-21.

Synemon selene, Klug, Abhandl. kén. Akad. Berlin, 1850, p. 249, pl. annex. f. 3,4; Boisduval, H.n. Lép.

Hét. p.553; Walker, List Lep. Het. B. M. vii. p. 1584.

Hab. Nova Hollandia. In Mus. Berlin.

5. Synemon ais. JS. corpore griseo, subtus pallido; alis anticis supra fusco-griseis,

fascia tranversa sinuata ante medium maculaque media griseis: posticis luteis fascia

transversa medio profunde sinuata baseos, maculis ad marginem margineque fuscis,

omnibus subtus luteis fusco-maculatis 9. Expans. alar. antic. 12+ lin.

Synemon nais, Klug, Abhandl. kén. Akad. Berlin, 1850, p. 250, pl. annex. f. 7; Walker, List Lep. Het.

B. M. vii. p. 1584; Boisduval, H.n. Lép. Hét. p. 553.

Hob. Nova Hollandia. In Mus. Berlin.

6. SYNEMON caTocaLoipEes. Nigyo-cinerea, subtus alba; antennis albo annulatis ; alis

anticis squamis plurimis pallide cinereis, basi maculis tribus alteraque versus

medium marginis interni nigris, macula magna pone medium nigro-ferruginea,

fascia arcuata guttarum albarum strigisque quatuor brevibus subapicalibus nigris,

margine apicali fusco intus crenulato; alis posticis lzete rufis, fascia parva media

SECOND SERIES.—ZOOLOGY, VOL. I. 2D

196 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

transversa versus marginem analem, costa Jate et irregulariter nigra, limboque

apicali nigro guttis nonnullis subapicalibus rufis. Expans. alar. antic. lin. 22.

The above description is made from the type specimen in the British Museum.

Synemon catocaloides, Walker, List Lep. Het. B. M. xxxi. p. 44; Boisduval, H. n. Lép. Hét. p. 551.

Hab. Swan River. In Mus. Britann.

7. SYNEMON PARTHENOIDES. Alis anticis brunneis obscurius nebulosis, fascia lata sub-

maculari e medio coste fere ad angulum internum extensa, venis divisa, altera

breviore bifida literam v simulante, inter precedentem et apicem, albis; alis posticis

nigris, macula rotunda aurantiaca prope basin, seriebus duabus macularum ejusdem

coloris inter medium et apicem, interiore in medio interrupta, altera versus angulum

externum abbreviata. Hxpans. alar. antic. fere une. 2.

Synemon parthenoides, Felder, Reise d. Novara, Zool. Bd. ii. Abth. 2, tab. 79. f. 7,8; Boisduval, H. n.

Lép. Hét. p. 550, pl. 11. fig. 5.

Hab. Noy. Hollandia. In Mus. Boisduyal.

8. SYNEMON mopsa. Nigro-fusca; alis subtus fulvescentibus, maculis testaceis, anticis

supra pallide fuscis vittis duabus albis basalibus, gutta alba ad apicem areole

discoidalis, pone medium fascia transversa maculari alba seriem macularum fus-

carum includente, margine apicali subgriseo; alis posticis pallide castaneo-fuscis,

nebulis nonnullis submarginalibus fuscescentibus. Hxpans. alar. antic. 15 lin.

Synemon mopsa, Doubleday, in Lord’s Discov. Australia, App. i. p.518, pl. 3. f. 7; Walker, List Lep. Het.

B. M.i. p. 35; Boisduval, H. n. Lép. Hét. p. 552.

S. theresa, var., Klug, Abh. kon. Akad. Berl. 1850, p. 249.

Hab. South Australia. In Mus. Britann., Boisduval.

9. SYNEMON ICARIA. Alis anticis griseo-albido nebulosis, basi obscuriore, fascia e medio

costee ad marginem internum (ibique dilatata) fusca, fasciis duabus inter medium

et apicem sinuatis obliquis; interna versus angulum internum seriem lunularum

formante, externa (subapicali) abbreviata, lunulis submarginalibus fuscis: alis pos-

ticis flavo-aurantiacis, macula magna basali marginis abdominalis cum striga un-

data submedia e margine costali fere connexa nigris, limbo apicali angusto nigro,

aurantiaco guttato. Expans. alar. antic. unc. 14.

Synemon icaria, Welder, Reise d. Novara, zool. Th. Bd. ii. Abth. 2, pl. 79. f. 6; Boisduval, H. n. Lép. Hét.

p. 554.

Hab. Nova Hollandia, Swan River. In Mus. Boisduval.

10. SyNEMON PLANA. (Plate XXXIII. fig. 8.) Fusca, abdomine pallido depresso: alis

anticis fusco-badiis, macula ovali inter basin et medium albido cincta, strigaque

tenui conica postice adjecta, macula indistincta pallida ad apicem cellule discoidalis

maculisque duabus majoribus inter medium et apicem nigricantibus, lineis albidis

marginatis; margine apicali tenui nigro, intus serie punctorum albidorum notato:

’

GENUS CASTNIA AND SOME ALLIED GROUPS. 197

alis posticis leete fulvis, guttis 3 et 1 pone medium margineque apicali tenui nigris :

alis subtus albis; margine tenui apicali omnium fusco, squamis luteis irrorato ;

anticis prope apicem punctis duobus, posticis cum 4 pone medium nigris. Expans.

alar. antic. unc. 13.

Synemon plana, Walker, List Lep. Het. B. M. i. 37, Boisduval, H.n. Lép. Hét. p. 554.

Hab. Nova Hollandia, Hunter River. In Mus. Britan., et Hopeiano Oxonie.

11. Synemon pirecta. (Plate XX XIII. fig.6.) Fusca, alis anticis concoloribus, fascia

recta alba, e medio costz fere ad angulum internum extensa, lunula abbreviata inter

medium et apicem minus distincta, spatio inter fascias plagam ovalem fusco-nigram

formante; alis posticis nigris, macula curvata inter basin et medium, fascia macu-

lari pone medium valde sinuata guttisque nonnullis ovalibus submarginalibus leete

aurantiacis ; alis subtus rufo-fulvis nigro maculatis. Hxpans. alar. une. 2.

Hab. Australasia. In Mus. Hopeiano Oxonie.

12. Synemon NupTA. (Plate XXXIII. fig.7.) Plumbeo-cinerea, macula oblique trans-

versa ad medium cellule discoidalis extus squamis albis marginata, fascia media

obliqua pone medium e costa ad medium disci extensa, altera abbreviata costali inter

hance fasciam et apicem strigaque subapicali e maculis conoideis nigris formata ; alis

posticis nigris extus cyaneo nitidis, plaga magna basali trigona, maculis duabus

pone medium prope marginem internum alteraque prope angulum externum san-

guineis: alis anticis subtus nigris, fascia abbreviata media costali alteraque sub-

apicali rufis ; posticis nigris guttis tribus albis discoidalibus in triangulum dispositis,

alteraque parva rufa inter duas posticas. Expans. alar. unc. 13.

Hab. Australasia occidentalis. In Mus. Hopeiano Oxonie.

13. SyneMon NorHA. (Plate XX XIII. fig.10.) Fusca, alis anticis cinereis, nigro variis,

macula magna indistincta inter medium disci et marginem posticum alteraque

subcostali, inter medium et apicem, in strigam punctorum nigrorum postice ad

marginem internum extensam terminante, strigaque maculari submarginali, margi-

neque ipso tenni nigricantibus ; alis posticis lete aurantiacis, basi fascia media (ad

costam paullo dilatata) limboque tenui marginali nigris, hoc serie guttarum auran-

tiacarum notato: alis subtus rufo-fulvis, anticis striga interna basali fasciaque brevi

(pone medium costze) extus guttis albidis notata; alis posticis macula ante medium,

fascia media limboque nigris, omnibus guttis albidis postice notatis. Expans. alar.

antic. une. 14.

Hab. Australasia, Albany (Brewer). In Mus. Hopeiano Oxonie.

14. SYNEMON OBSCURELLA. (Plate XXXIII. fig.9.) Fusco-nigricans, alis anticis striga

tenui abbreviata obliqua media alba, lunulisque duabus inter medium et apicem

pailide luteis (externa obscuriore); alis posticis nigricantibus, gutta parva lutea

inter basin et medium, dimidio apicali fulvo-luteo, striga undata subapicali, in medio.

interrupta, fuscescente: alis anticis subtus basi nigris, dimidio apicali fulvo, macula

2D 2

198 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

rotunda versus costam nigra; alis posticis nigris, ultra medium fulvis striga sub-

undata submarginali fusca. Hxpans. alar. antic. unc. 1}.

Hab. Australasia. In Mus. Hopeiano Oxonie.

15. SYNEMON HESPERIOIDES. (Plate XX XITI. fig. 11.) Fusca, alis anticis versus basin

squamis albidis variis, macula parva conoidea ad apicem cellule discoidalis, maculis

duabus (lineis tenuibus marginatis) strigaque apicali lunularum parvarum albidis;

alis posticis brunneis, pone medium striga fere indistincta macularum nigrarum,

margine apicali fusco: alis subtus brunneis, anticis in medio et ad apicem, posticis

in medio, serie irregulari circularum margineque apicali squamis albido-sericeis

irroratis. Expans. alar. antic. une. 15.

Synemon hesperioides, Felder, Reise d. Novara, zool. Th. Bd. uu. Abth. 2, pl. 82. fig. 12; Boisduval, H.

n. Lép. Hét. p. 555 (regarded as one of the Hesperiide).

Hab. Nova Hollandia. In Mus. Hopeiano Oxoniz.

16. SynemMon vacans. (Plate XXXIII. fig. 12.) Alis anticis fusco-cinereis, fascia

transversa nigra ante medium e costa, postice acuminata, marginem posticum vix

attingente, fascia altera abbreviata subcostali venis divisa nigra pone medium, extus

pallidius margivata, guttis tribus ovalibus nigris subapicalibus alterisque quinque

submarginalibus margineque ipso tenui nigris; alis posticis nigris, fasciis duabus

arcuatis aurantiacis: alis subtus fulvo-luteis, anticis macula irregulari media alte-

raque pone medium subcostalibus nigris, apice albido nigroque vario; alis posticis

magis fulvescentibus, medio circulo fere integro nigro notato, limbo tenui nigro,

fulvo guttato. Hxpans. alar. antic. lin. 19.

Hab. Nova Hollandia (Strange). In Mus. Hopeiano Oxonie.

Obs. Heec species cum sequente a reliquis differunt alis anticis angustioribus, margine

antico in medio parum concayo-sinuato nee convexo.

17. Synemon GRratiosa. (Plate XX XIII. fig.13.) Alis anticis fuscis, dimidio apicali

pallidiore, macula irregulari transverse obliqua submedia, intus squamis pallidi-

oribus irrorata, macula altera subcostali conoidea pone medium nigra, serieque

submarginali punctorum nigrorum (in medio interrupta), margine ipso tenui ob-

scuro; alis posticis aurantiacis, macula parva nigra versus costam et pone medium

posita limboque tenui nigro, in medio aurantiaco guttato: alis anticis subtus obscure

fulvis apice albidis, maculis duabus mediis duabusque inter medium et apicem fuscis,

margine fusco, albido guttato; alis posticis fulvis, maculis tribus mediis, externa

majore et ad costam extensa, limbo nigro, extus albido guttato. Hxpans. alar. antic.

mmbae, dees

Hab. Nova Hollandia. In Mus. Hopeiano Oxoniz.

Genus IV. Tascina. (Plate XX XIII. fig. 5.)

Insectum orientale Castnias Americe meridionalis mundo antiquo representans, cum quibus antennis

necnon numero venarum alarum congruit, his autem aliter dispositis ; palpis exiguis, lingua spirali

GENUS CASTNIA AND SOME ALLIED GROUPS. 199

obsoleta, alisque anticis apice acuminatis e Castniis facile distinguitur. Caput magnum ; oculi maximi,

laterales, antice approximati, spatio angusto inter se separati; palpi tenuissimi, breves, erecti, ad

medium frontis vix attingentes, articulo penultimo longo, ultimo minuto acuto. Vestigium nullum

lingue spiralis detexi. Antenne circiter 50-articulate, clavatz, articulis primis 32 filiformibus ; apice

acuto, parum incurvato, fasciculo parvo setarum terminato. Thorax robustus. Abdomen mediocre.

Tibie antic breves, in medio calcari armate. Tarsi antici elongati, pulvillo mediocri, unguibus

acutis, curvatis, simplicibus, setisque duabus elongatis apicalibus instructi. Ale sat magne, antice

trigonz, margine apicali fere recto, apice acuminato producto ; postice ovali-trigone, tenaculo (seta

crassa curvata prope basin costz) anticis ligatz: antic vena postcostali paullo ante medium ale

bifurcata, ramo antico ramulos tres emittente, quorum tertius (6 3) apicem ale attingit, ramo

postico etiam tres emittente ; vena mediana ramulos quatuor emittente, quorum superior (seu disco-

cellularis, c 3*) cum inferiore (65*) ven postcostalis venula nulla transversa conjungitur : ale

posticse vena postcostali bifida, ramulis hberis et cum venulis mediane haud conjunctis, vena mediana

e basi bipartita, singula parte etiam bifida, inde rami 4 mediani; vena anali et subanali simplicibus.

TASCINA ORIENTALIS. (Plate XX XIII. fig. 5.) Alis anticis saturate fuscis, fascia fere

recta obliqua pone medium costz fere ad angulum internum extensa albida, alis

posticis obscure aurantiaco-rufis, imbo lato fusco; corpore nigricante, abdominis

apice rufescente. Long. corp. unc. 14. Expans. alar. unc. 34.

Hab. Singapore (Wallace). In Mus. Hopeiano Oxonie.

Genus V. Hecatresta. (Plate XX XIII. figs. 1—4.)

Hecatesia, Boisduval, Monogr. Zygen. p.11; Walker, List Lep. Het. B. M. i. p. 54.

Corpus sat crassum, abdomine in maribus barbato. Caput parvum, ocellis duobus instructum. Antenne

elongatz, cylindricze, pone medium in maribus fusiformes vel subfusiformes, in foeminis minus

incrassate. Palpi ultra medium faciei assurgentes, dense hirsuti, articulis tribus, primo crassiore

curvato, tertio dimidium secundi paullo superante. - Lingua elongata spiralis. Thorax valde hirtus.

Ale, insecto quiescente, tectiformes, breves; anticee in maribus spatio magno reniformi vitreo

transverse striolato, prope costam, instructz, hac in medio in tuberculum corneum elongatum dila-

tata, ramos venz postcostalis dislocante, scil. ramo primo curvato marginem posticum arez vitre

formante, ramis 2, 3 et 4 subapicalibus ad basin inter se zque distantibus, vena quinta prope basin

arez vitreze emissa, vena cellulam discoidalem claudente fere indistincta at curvata; vena mediana

4-ramosa, ramis tribus ultimis ad basin valde approximatis: ale anticee in feemina area vitrea

earentes; cellula magna discoidali, apice fere recta transversa, cellulaque parva angusta subcostali

distincta, ad apicem ejus ramos tres emittente, medio bifido: ale posticz cellula discoidali ad

trientem alee extensa, ibique transverse et indistincte desinens, vena 3 ¢* e medio ejus emissa. Pedes

satis graciles, tibiis barbatis ; tibiee antice: calcari forti medio instructee.

This curious Australian genus was first described by Boisduval, and placed by him in

the family Zygzenidze, but removed to the Agaristides in his recent memoir on that tribe

published in the ‘ Revue de Zoologie,’ 1874, in which position, between Husemia and Aigo-

cera, it was placed by Walker. It appears to me, however, to be more nearly allied to

Castnia by means of the genus Synemon, agreeing with Castnia in the possession of the

small narrow subcostal cell formed by the branches of the postcostal vein, the fourth

of which arises from the third at some distance beyond the extremity of this small cell,

in the normally formed wing of the female. Until, however, we become acquainted with

200 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

the transformations of Hecatesia, and have learned whether the larva is xylophagous,

radicivorous, or phyllophagous, and the pupa armed with rows of recurved spines or

simple, we cannot decide on the true relations of the group.

1. HecatTesiaA FENESTRATA. (Plate XXXIIT. fig. 1.) Hem. thorace nigro-fusco, albo

trivittato; alis anticis nigro-fuscis, fasciis duabus transversis parum macularibus,

prima ante, secunda pone medium alarum; alis posticis luteo-fulyis, limbo lato

punctoque medio subcostali nigris; abdomine fulvo-luteo, maris fusco-barbato, serie

dorsali punctorum nigrorum. Expans. alar. antic. lin. 14-18.

Obs. The female, here figured for the first time, differs from the male only in wanting

the vitreous spot on the fore wings. |

Hecatesia fenestrata, Boisduval, Monogr. Zygen. p. 11, pl. 1. f. 2; Voy. de lAstrolabe, Ent. p. 171;

Spec. Gén. Lépid.i. pl. 10 B. fig. 7; Mon. Agaristides, Rev. Zool. 1874, and sep. issue, p. 23;

Chenu, Encycl. Hist. Nat. Pap. 236, f. 403 ; White, in Grey’s Journ. Exped. Austral. Append. p. 475,

pl. 8. f. 2; Walker, List Hep. Het. B. M. 1. 54.

Hab. Nov. Hollandia. In Mus. Britann., Hopeiano, &e.

2. HECATESIA THYRIDION. horace fusco-nigro, albo trivittato; alis anticis fusco-nigris,

fasciis tribus transversis macularibus guttisque nonnullis subapicalibus albis; area

vitrea in maribus angustiore quam in precedente specie; alis posticis aurantiacis,

limbo lato albo guttato punctoque subcostali mediano nigris ; abdomine supra auran-

tiaco, maculis nigris dorsalibus et lateralibus nigris ; subtus albo; palpis et pedibus

nigris, albo variis; antennis in utroque sexu pergracilibus. Expans. alar. antic.

bin, IP

Hecatesia thyridion, Feisthamel, in Voy. de la Favorite, v. Suppl. p. 19, pl. 5. fig. 1; White, Append. Capt.

Grey’s Journ. Exped. Austral. p. 475, pl. 8. f. 1 (m. andf.) ; Boisduval, Mon. Agaristides, p. 24;

Walker, List Lep. Het. B. M. i. p. 55.

Hab. Nova Hollandia, Albany (Brewer), Swan River (Curtis), West Australia

(Damell). In Mus. Britann., Hopeiano Oxonie.

3. HECATESIA EXULTANS. (Plate XXXIII. figs. 2m, 3/f.) Atra: capite et thorace flavo

suttatis; abdomine serie dorsali punctorum fulvorum, lateribus fulvo nigroque

variis : alis anticis nigris, maris margine antico in medio dilatato, pone medium pro-

funde emarginato, feeminze forma ordinaria, in utroque sexu guttis numerosis

flavis et griseis, fascias septem irregulares maculares alternatim flavas et griseas for-

mantibus ; fimbriis omnibus griseis nigro maculatis: alis posticis aurantiacis, limbo

lato maculaque trigona subcostali mediana nigris: capite subtus orbitu oculorum

et basi palporum flavo hirtis: pedibus nigris, tibiis fulvo hirtis, apicibus et tarsorum

artic. 1 et 2 apicibus albis; calcaribus longis nigris: abdomine pone medium ma-

culis sex transversis albis (per paria dispositis) : alis anticis subtus absque squamis

griseis; posticis guttis nonnullis albis in limbo nigro. Expans. alar. antic. cire.

une. 1,

Hecatesia exultans, Walker, List Lep. Het. B. M.1. p.58; Boisduval, Mon. Agarist. p. 24.

GENUS CASTNIA AND SOME ALLIED GROUPS. 201

Obs. The typical specimen of this species, described by Mr. Walker as a male, is a

female, wanting the vitreous spot of the fore wings, which exists in the true male, the

wings being of a remarkable form in the latter sex.

Hab. Swan River, Nov. Holl. (Diggles). In Mus. Britann., Hopeiano Oxonie.

The insects described in the remaining portion of this memoir have been regarded by

various authors as belonging to the group of which Castnia is the type—from which,

however, they are entirely distinct, belonging to other Heterocerous groups.

Genus OTHria, Westw.

Dr. Boisduval’s species of Orthia, O. augias and O. nexa, must be referred to the

Agaristides, the veining of their wings agreeing exactly with that of Agariste. I propose

to place them therefore as a separate genus of Agaristides, under the name of Othria.

From Zgocera they are distinguished by their geographical distribution being con-

fined to South America.

In O. augias (Plate X XIX. figs. 18, 19) the fore wings have a small postcostal cell,

the branches 62 and 43 arising far beyond the cell, as does also 05, whilst 05* springs

from a little branch from the middle of the hinder margin of the postcostal cell, and the

discoidal cell of the hind wings is imperfectly closed by a very slender arched disco-

cellular vein or, rather, fold.

1. OruRia avetas. “ Ailes noires sans aucun reflet; les ailes supérieures, dont les

nervures forment des stries grisatres longitudinales sont traversées, du milieu de

la céte a l’angle interne, par une bande d’un jaune @ocre, trés-droite. Ailes infé-

rieures sans taches, avec les nervures un peu plus claires.” Exp. alar. ant. une. 22.

Orthia augias (Boisduval, MS.), Herr.-Schaffer, Aussereur. Schmett. f. 18; Boisduval, H. n. Lép. Hét.

p- 542 (incorrectly referred in his text, but with doubt, to Orthia pelasgus).

Hab. Brasilia (Rio Janeiro), Minas Geraes (Rogers), Brasilia occidentalis (Rogers).

In Mus. Hopeiano Oxoniz, Boisduval.

2. OTHRIA NEXA. “ Hlle a le port et la taille de O. augias; ses quatre ailes sont d’un

noir terne, sans aucun reflet, avec l’extrémité des nervures grisatre comme chez

augias. Lesailes supérieures sont traversées, du milieu de la céte a l’angle interne,

par une bande d’un jaune d’ocre trés-pale un peu recourbée a son extrémité comme

dans pelasgus ; les ailes inférieures sont sans taches. Cette espéce pourrait bien étre

une variété femelle d’augias. Hlle ne différe que par la bande des ailes supérieures

notablement plus pale, formant un tant soit peu le crochet a son extrémité inférieure,

et par le dessous des ailes supérieures dont la céte est piquée de gris.”

Hab. ——? In Mus. Boisduval.

Orthia neva, Boisduval, H.n. Lép. Hét. p. 543.

202 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

3. OTHRIA AMAZONICA, Westw. Alis brevioribus, latioribus, anticis subtrigonis; atra,

alis fascia fulva e medio costee fere ad angulum analem extensa, postice nonnihil

incurva, alis subtus omnino concoloribus. Expans. alar. ant. une. 1, lin. 11.

Hab. Amazonia. In Mus. Hopeiano Oxonie.

Had not Dr. Boisduval described an insect as the probable female of O. augias, with

erey veins at the extremity of the fore wings as in the male, I should have been induced

to regard this Amazonian insect as the female of O. augias. The veins of its wings are

arranged exactly as in O. augias.

4. Orarta Linpier. Atra, collo, abdominis lateribus, alis anticis, macula magna basali

trigona alteraque transversa ovali pone medium alisque posticis (limbo lato nigro

excepto) fulvis, subtus magis stramineis. Expans. alar. 273 une.

Agarista Lindigii, Felder, Lep. Voy. Novara, pl. 107. f. 6.

Hab. Villa Nova Amazonize. In Mus. Hopeiano Oxonie.

Obs. Individua e Nova Granada et Ecuadoria allata paullo minora (exp. unc. 1g) macu-

lisque fulvis minoribus.

Obs. Individuum majus (exp. unc. 24%;) maculis albidis in alis anticis, alisque posticis

pallidioribus, limbo nigro angustiore.

In this species the second branch of the postcostal vein arises near the upper end of

the small accessory postcostal cell. In the hind wings the veins are arranged as

in O. augias.

5. OraRIA AMALTHEA. Atra, annuio colli, lateribus abdominis, puncto trigono in medio

fasciaque tenui abbreviata pone medium alarum anticarum, alis posticis (limbo

lato nigro excepto) fulvis. Expans. alar. antic. unc. 144.

gocera amalthea, Dalman, Analecta Entom. 49. 24; Walker, Cat. Lep. Het. B. M. i. p. 58.

Hab. Brasilia, Peru, Bogota? In Mus. Hopeiano Oxoniz.

“In this species the fore wings are more elongated, and the second branch of the post-

costal vein of the fore wings arises exactly at the tip of the small accessory post-

costal cell.

6. OTHRIA coLUMBINA. Tota atra, fascia obliqua parum flexuosa e medio costx fere

ad angulum analem extensa, fulva, postice parum latiore. Expans. alar. antic.

une. 2-(5-

Hab. Columbia (Chesterton), In Mus. Hopeiano Oxonie.

The wing-veins are arranged as in O. amalthea.

7. OTHRIA ECUADORINA. Corpore cum collo nigris, alis latioribus atris; anticis fascia

tenui recta obliqua pone medium nec marginem costalem nec angulum analem

attingente, maculaque magna ovali discoidali alarum posticarum aurantiacis, margine

anali nigro. Expans. alar. antic. unc. 2.

Hab. Eouador. In Mus. Hopeiano Oxoniz.

GENUS CdASTNIA AND SOME ALLIED GROUPS. 203

This species agrees. with the preceding in its black and orange colours; but it wants the

small accessory postcostal cell, and the branch 6 2 arises at half the distance between the

discoidal cell and the tip of the wing, followed at a short distance by 63, whilst 55 arises

from the hind edge of the main stem, slightly in advance of the origin of 62. (See Plate

XXIX. fig. 23.) In other respects the veins agree with O. augias.

Several other species, evidently congeneric with the above, are figured by Dr. Felder

in the 137th plate of his work on the Lepidoptera of the ‘Novara’ voyage, under the

names of Agarista Doleschallii, A. Semperi, A. Moorei, A. alethe, and A. Batesii.

The following insects must also be removed from their proximity to Castnia and

referred to the Agaristides.

Genus Damas, Boisduval, Voy. Astrol. pt. i. p. 259.

The insects of this genus are black and orange like the Othrie, but are natives of the

Kastern Archipelago.

1. Dantas ELEGANS, Boisduval, Voy. Astrol. pt. i. Lép. 259. 1.

Hab. New Guinea.

2. DAMIAS MELAXANTHA, Boisduval, Voy. Astrol. pt. i. Lép. 260. 2.

Cleis posticalis, Guérin, Voy. Duperrey, Atlas Ins. pl. 18. f. 5.

Hab. Port Praslin, New Ireland.

3. Damas DicHROA, Boisduval, Voy. Astrol. pt. i. Lép. 260. 3.

Hab. Offack and Bourou.

4, Damas varia, Walker, List Lep. Het. Brit. Mus. i. p. 15.

Hab. Richmond River, Australia.

I cannot distinguish this insect specifically from an insect collected in Batchian by

Mr. Wallace, now in the Hopeian Museum at Oxford, described by Mr. Walker under

the name of Burgena transducta.

Genus BureGEna, Walker.

BuURGENA TRANSDUCTA, Walker. (See Plate XXIX. fig. 21.)

The fore wings are narrow; but in the veining of its wings it agrees with Agaristu,

the small accessory postcostal cell emitting only one branch at its extremity, the branch

6 2 arising at a considerable distance beyond the cell, whilst 65 arises from the hinder

edge of the stem close beyond the accessory cell.

Genus Hrspagarista, Walker.

HESPAGARISTA INTERJECTA, Walker, List Lep. Het. B. M.i. p. 14.

Brought by Mr. Angus from Caffraria, and figured by him in his fine work on the

Kaffirs, pl. 30.f.10. Agrees with the Othrig in general form and antenne ; but the wings

are much more maculated, the extremity of the body is tufted, and the veining of the

SECOND SERIES.—ZOOLOGY, VOL. I. 25

204 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

fore wings differs with respect to the small postcostal cell, which emits one branch, 05, at

its extremity, whilst the main stem of the postcostal vein arises from its anterior edge

near the extremity, b 2 arising at a little distance beyond the cell, followed at a very

short distance by 03, 04 extending to the extremity of the wing. (See Plate XXIX.

fig. 22.)

Genus RorHta.

Another group of species, also closely allied to the preceding, are natives of Mada-

gascar, and have the small postcostal cell very narrow (thus differing from the typical

Eusemie, in which the small cell is wanting, Pl. XXIX. f. 20). They also have the

branch 6 2 arising from its upper extremity, and running free nearly to the tip of the

wing, 03 arising halfway between the cell and the tip of the wing, whilst 0 5 arises

close beyond the extremity of the postcostal cell; such at least is their arrangement

in C. pales and simyra. They may advantageously be formed into a separate genus,

to which the name of Rothia may be applied. Here belong

1. RovHiA PALES.

Agarista pales, Boisduval, in Guérin, Icon. R. An. pl. 83. f. 1; Faun. Ent. Madag. pl. 10. f. 1, 2.

Eusemia pales, Boisduval, Mon. Agarist. p. 75. no. 85; Walker, List Lep. Het. B. M. 1. p. 54.

Hab. Vicinity of Tananariva, Madagascar. In Mus. Hopeiano Oxoniz.

2. ROTHIA ERIOPIS.

Agarista eriopis, Boisduval, in Herrich-Schaffer, Auss. Schm. f. 31; Mon. Agaristides, p. 71 (Eusemia er.).

3. ROTHIA AGRIUS.

Agarista agrius, Boisduval, in Herrich-Schiffer, op. cit. f. 83; Mon. Agaristides, p. 72 (Husemia agr.).

4, ROTHIA PEDASUS.

Agarista pedasus, Boisduval, in Herrich-Schiffer, op. cit. f. 32 (nec 34 ut in Bdv.); Mon. Agaristides,

p- 73 (Eusemia ped.).

5. Rorura stmyra. Atra, capite et collo flavo-guttatis: alis anticis elongatis nigris,

cutta minima humerali fasciaque transversa parum sinuata (postice paullo latiore) e

medio costee fere ad angulum analem extensa flavescentibus; alis posticis atris,

fimbria ad angulum externum alba; corpore infra, pedibus, et abdominis lateribus

lutescentibus ; alis posticis subtus dimidio basali rufo. Expans. alar. une. 23.

Hab. Madagascar. In Mus. Hopeiano Oxonie.

The arrangement of the wing-veins of this species was noticed above.

Genus AIGIALE, Felder. (Wien. ent. Mon. iv. 110, 1860.)

Type Agiale Kollari (Castnia hesperiaris, Walker, List Lep. Het. B. M. pt. vii. p. 1588).

This Mexican insect is no other than the large Hesperian butterfly first mentioned by

me (in the ‘ Genera of Diurnal Lepidoptera,’ p. 506) as remarkable for wanting spurs to

GENUS CASTNIA AND SOME ALLIED GROUPS. 205

the hind tibie*, and which was subsequently formed into a separate genus by Dr.

Felder, under the name of dgiale Kollari. The arrangement of its wing-veins (Plate

XXIX. fig. 25), and, indeed, the whole structure of the insect, are entirely Hesperideous.

Specimens are in the Hopeian collection at Oxford, presented to me by Mr. Coffin.

Genus MEGATHYMUS.

Megathymus, Scudder, Syst. Revis. Amer. Butterflies, p.12 (Salem, 1872).

The type of this genus is the large butterfly figured by Boisduval and Leconte (Iconogr.

Lépid. Amér. Septentr. pl. 70), under the name of Hudamus? yucce, by whom also the

larva and pupa were represented, but in an unsatisfactory manner.

Walker, List Lep. Het. B. M. pt. 7, p. 1583, gavea short and very insufficient description

of this insect under the name of Oastnia yucce, which, he says, “ connects the Castnie

with the Hesperie. Had he been aware of the figures of the preparatory states given in

Leconte and Boisduval’s work, he would have seen that in all respects (and most especially

in the larva form) this is a true Hesperian. In its wing-veins it agrees with Agiale

Kollari, except that the angulated vein closing the discoidal cell is very slender.

Since the present memoir was presented to the Linnean Society, an admirable memoir

on this insect, with figures of its various characters in the preparatory and perfect states,

has been published by Dr. Charles V. Riley, in the ‘ Transactions of the Academy of

_ Sciences of St. Louis,’ vol. 11. January, 1876, and subsequently introduced into the

‘ Highth Annual Report on the Noxious, Beneficial, and other Insects of the State of

Missouri, by the same author, who has arrived at the opinion that the imsect is a

large-bodied Hesperian, and that it has no real relation with Castnia.

DESCRIPTION OF THE PLATES.

Puare XXVIII. |

Details of Castnia.

Fig. 1. Larva of C. eudesmia.

Fig. 2. Pupa of C. eudesmia, dorsal view.

Fig. 3. Pupa of C. eudesmia, with one of the pterothece raised up, showing the fore wing and its marks.

Fig. 4. Cocoon of C. eudesmia.

5. Venation of fore and hind wings of C. papilionaris.

6. Venation of fore wing of C. cacica.

Fig. 6 a. Venation of hind wing of .C. cacica.

Fig. 7. Characteristic portion of the veins of fore wing of C. mygdon.

Fig. 8. 7 i i of C. Boisduvallii.

* In ‘H. paniscus the two spines in the middle of the two hind tibie are wanting (H.-Schifter, Syst. Bearb.

pl. vi. f. 17).

2E2Z

206 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

Fig. 9. Fore wing of C. veraguana.

Fig. 10. Characteristic portion of the veins of fore wing of C. ecwadoria.

Fig. 11. +3 » of fore wing of C. cochrus

Fig. lla. e », of hind wing of C. cochrus,

Fig. 12. : » of fore wing of C. truvilla.

Fig. 13. 33 5, Of fore wing of C. tcarus.

Fig. 14. 5 »,. of fore wing of C. Hiibneri.

Fig. 14a. ss » of hind wing of C. Hiibneri.

Fig. 15. & » of fore wing of C. Latreillei.

Note.—The same numbers and small letters are used in the above figures of the venation of the wings, as

well as in the following Plates, to express the corresponding parts in the different species.

a, costal vein; 6, postcostal vein with its five branches, 61, 62,63,64, and 65, together with

6 5*, the upper disco-cellular vein, which is here regarded as portion of the postcostal system ;

c, median vein and its branches, c 1, ¢ 2, and ¢ 8, together with c3*, the lower disco-

cellular vem, which is here regarded as portion of the median system; y and z represent the

bases of 6 5* and c3* within the discoidal cell; d, the postmedian vein; pc.c, postcostal cell ;

de, discoidal cell ; de*, hinder division of discoidal cell ; pmc, premedian cell.

PLATE XXIX.

Fig. 1. Characteristic part of the hind wing of Castnia juturna.

Fig. 2 a » fore wing of C. Marcel-Serresii male.

Fig. 3 i » hind wing of ditto ¢, and annexed partial diagram of 2.

Fig. 4 3 » fore wing C. chremes, male.

Fig. 5. A » | hind wing of C. chremes, male and female.

Fig. 6 Ss » fore wing of C. diva.

Mig. 7 Bs », hind wing of ditto.

Fig. 8. Fore wing of C. cononia.

Fig. 9. Characteristic part of the fore wing of C. linus.

Fig. 10. » hind wing of ditto.

Fig. 11. a » fore wing of C. mimica.

Fig. 12. 4 » hind wing of ditto.

Fig. 13. Fore wing of C. acreoides.

Fig. 14. Characteristic part of hind wing of ditto.

Fig. 15. 55 » hind wing of C. therapon.

Fig. 16. Fore wing of Orthia pelasgus.

Fig. 17. Hind wing of ditto.

Fig. 18. Fore wing of Othria augias.

Fig. 19. Hind wing of ditto.

Fig. 20. Fore wing of Husemia.

Fig. 21. Characteristic part of fore wing of Burgena transducta.

Fig. 22. Fore wing of Hespagarista interjecta.

Fig. 23. Characteristic part of the fore wing of Othria ecuadorina.

Fig. 24. % eS fore wing of Agarista Lindigii.

Fig. 25. Fore wing of Agiale Kollart.

Fig. 26. Fore wing of Huschemon Rafflesii.

27. Fore wing of Hepialus humili, fem.

Fig. 28. Hind wing of ditto.

Fig.

GENUS CASTNIA AND SOME ALLIED GROUPS.

. Castnia veraguana.

. C. ardalus, var.

. C. phalaris.

Castnia clitarcha, ¢ .

. C. chtarcha, @.

. C. papihonaris.

cycna.

PLate XXX.

Fig. 4.

PuatTe XXXII.

Fig. 4.

Puate XXXII.

Fig

Puate XXXII.

207

Castnia amycus, var.

C. Dalmanni, 2 from Brit. Mus.

C. papagaya.

Castnia satrapes, var.

C. satrapes, var.

. 5. C. (G.) cononia.

6. C. (G.) ecuadoria.

7. C. (G.) mimica.

1. Hecatesia fenestrata, 9 ; 1a. Head and antenna of male; 1%. Palpus; 1c. Palpus denuded;

1d. Fore leg; 1e. Fore wing of male.

2. H. exultans 3.

3. H. exultans 2.

4. Wings of H. thyridion.

5. Tascina orientalis; 5a. Head, seen in front; 5 6. Ditto, sideways; 5 ¢. Hxtremity of antenna;

5 d. Fore legs; 5e. Extremity of fore tarsus; 5/f. Hind leg; 59. Venation of fore wing;

5h. Ditto hind wing.

Fig. 6. Synemon directa.

ihe

S. nupta.

8. S. plana.

9. S. obscurella.

Fig. 10. Synemon notha.

ll. S. hesperioides.

12. S. vagans; 12 a. Venation of wing of same.

13. S. gratiosa.

14a. Fore wing of Synemon Sophia; 146. Hind wing of ditto; 14c. Head and base of antenna;

14d. Palpus of S. Sophia; 14 e. Palpus, denuded, of ditto; 14,f. Fore leg of ditto; 14.9. Middle

leg of ditto; 14. Hind leg of ditto.

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a VENATION & DEVELOPEMENT OF CASTNIA.

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VENATION, SPECIES OF CASTNIA &c.

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NEW SPECIES OF CASTNIA.

JOWestwood del. TRANS. Linn. Soc.SER.2.Zoon Vou.1 Tas 31.

CBerjeau lith Hamhart imp.

NEW SPECIES OF CASTNIA

JOWestwood del. Trans Livw Soc Ser.2.Zoon Vou.1 Tas. 32

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NEW SPECIES OF (GAZERA.) CASTNIA.

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MS GLLELIDOD Sophia.

Hanhart mp.

[ 209 ]

IX. On a New Genus of Trematoda, and some new or little-known Parasitic Hirudinei.

By Joun Denis Macponatp, U.D., F.R.S., Deputy Inspector-General R.N.,

Professor of Naval Hygiene Army Medical School. (Communicated by G. E.

Dosson, M.A., M_B., F.LS.)

(Plate XXXIV.)

Read April 6, 1876.

MANY striking points of resemblance are traceable between the Trematoda and the

Hirudinei; but this is merely indicative of a representative relationship, or one of

analogy. The numerous errors into which our forefathers fell in grouping together

really incongruous things may be referred to the deceptiveness of primd facie resem-

blance, and the failing to recognize the intrinsic difference between the kind of relation-

ship here alluded to and that of a genuine affinity. This, I am quite sure, is one of the

greatest difficulties with which modern evolutional biology will have to contend. An

analogous property of gemmation in certain animals, in common with plants, being the

innocent cause, so to speak, of the dendritic or tree-like form, so deceived the earlier

observers that even their very word Zoophyte is accepted by us as a merely descriptive

term, in a sense which is precisely the converse of that which its etymological construc-

tion would indicate. The balance was so far in favour of the plants, that the doubt

gave rise to the additional complication of confounding this group with the Corallines.

Now, we know that there are also other representative relationships between the two

great organic kingdoms, such as conjugation, fission, contractility, and motion, in con-

nexion with which errors of judgment are continually creeping into the dicta of science.

I might further say that the light available to Linnzus and his immediate followers with

regard to these matters was scarcely less than that which we have at present to guide us

in the more difficult study of the primary modes of evolution observable in the ovum,

with the view of establishing the affinity or antipathy of the resultants, however different

or identical in type these latter may seem to be. Indeed, even up to the present, facts

are not wanting to show that very similar beings originate in a very dissimilar way, and

vice versd. Nevertheless we look forward with pleasure to the results that appear to be

promised to the labourers in this line of research.

- If we look upon the Trematoda as representing the Hirudinei, coupling also the

Oligocheta with the Polycheta, and the Rotifera with the Articulata, Prof. Huxley’s

interesting classification, embodying a large proportion of the results just alluded to,

would seem to sanction the idea. Thus the corresponding part of his table may be

arranged, without material change, in the following manner :—

1. ARCH ZOSTOMATA. 2. DEUTEROSTOMATA.

Trematoda. Annelida.

Oligocheta. Polycheta.

Rotifera. Arthropoda.

210 DR. J. W. MACDONALD ON A NEW GENUS OF TREMATODA,

There is surely something more than coincidence in this. There may be misgivings

as to the position of the Oligocheta; but perhaps something will soon turn up to recon-

cile it.

One or two additional points of analogy between the Trematoda and the Hirudiner

will be seen in the description of the forms which I have now to present to your notice.

The little Trematode represented in the first figure of the plate was found by me

creeping about in the respiratory siphon of a large Melo, or melon-shell, in Shark Bay,

Western Australia. It was between an eighth and a tenth of an inch in length, tallowy

white (to use an American expression), and so much resembled a leech as to deceive me

at first sight; for I had already seen white leeches, both parasitic and free, and I did not

expect to find a Trematode walking abroad. The fore part of the body was slender, taper-

ing, and depressed ; the oral sucker was subterminal, and of the usual cup-like form ; and

after a slight constriction, a bulbous pharynx was distinctly seen, from which the ceso-

phagus passed backwards for some length, to what would appear to be its bifurcation ;

and the appearance of the czeca could be traced to within a short distance of the posterior

extremity. But the ventral disk constituted the most striking feature in this little animal,

on account of its great size and complex structure. It occupied more than the two

posterior thirds of the inferior surface of the body, which was laterally compressed

instead of depressed in this locality. The disk, or virtually the foot, might therefore be

said to be narrow and elongated in form, and not unlike that of Aspidogaster, though

very different in structure. Thus, it presented four rows of alveoli, or deep sucking-pits

(fig. 3), the marginal series being the largest, with their long diameter running trans-

versely, while that of the intermediate ones was longitudinal. All the contiguous

borders of these little pits were so connected by straight bands of muscular fibres as to

divide the whole of the intervening area into triangular spaces, the centre of each of

which was ocupied by a curious tentaculiform cirrus, capable of complete retraction from

the apex, as in the common instance of the finger of a glove (fig. 4). Six rows of these

cirri, with thirty in each row, would make up 180, and four rows of alveoli, thirty in

each row, 120; so that the disk altogether presented a very remarkable appearance.

The water-vascular system, as in the Zrematoda generally, opened posteriorly. The

aperture, however, was subterminal and dorsal, or just above and within the hinder

border of the disk. The tortuous vessels of this system were seen running forwards on

either side of the body as far as the space between the pharyngeal bulb and the oral

disk, whence they turned backwards upon themselves, as shown in fig. 5. The genital

opening, as in most other cases, was situated in front of the ventral disk (fig. 1, 6).

About the same time I found a large black leech on a species of Myliobatis, presenting

at least two characters which were quite new to me—namely, a row of branchial leaflets

on each side of the body (figs. 8 & 10), and a caudal sucker alveolated (figs. 11 & 12

very much after the manner of the Tremotode already described.

The head of this creature was depressed, spatulate, and pointed in front (fig.8). The

anterior third of the body was without appendages, constituting a kind of neck, but the

rest was furnished on either side with a row of heart-shaped leaflets on short pedicles.

Over the base of these, on every fourth segment, was situate a pulsating projection,

AND SOME NEW OR LITTLE-KNOWN PARASITIC HIRUDINEI. 211

connected, no doubt, with the water-vascular system, as in Pontobdella. The rich black

body, moreover, was ornamented with six rows of white spots (fig. 10).

The alveoli of the caudal disk were disposed in rays rather than in circles, with

radiating muscular bundles between them ; they were circular and cuplike, with a central

retractile portion, reminding one of the mechanism of the suckers of the Cephalopoda.

Attached to the body in a very irregular manner, but chiefly at its fore part, were

several of the double tubular spermatophora shown in fig. 9. These curious bodies

I have also found on other marine Hirudinei, but always with some characteristic dif-

ference. Fig. 6, for example, represents a small black leech with white tubercles, refer-

able, apparently, to the genus Pontobdella, found on Rhinobatis in the same seas ;

and fig. 7 is its double-barrelled spermatophore, which is quite different from fig. 9,

though obviously of the same nature. Very little is positively known of the generative

processes of the marine leeches ; but the facts here mentioned may one day meet with a

satisfactory explanation. i

Fig. 13 is a small white leech, with two eyes, a retractile proboscis, and, in short, the

general anatomy of Glossiphonia, but characterized by possessing seven pairs of ramose

lateral branchize, taking up about the middle third of the body. It was found on the

conjunctiva of a green turtle at Huon’s Island, on the reef to the northward of New

Caledonia. Fig. 14 is another specimen of the same species, but much larger, taken in

Shark Bay; both figures are about double the natural size. Fig. 15 shows one of the

branchize as seen with a power of a quarter of an inch.

In systems of classification both the Oligocheta and the Hirudinei come under the

head of Abranchiata. But this is evidently a mistake; for, in the first place, it is difficult

to deny the branchial function of the ciliated finger-like organs surrounding the vent

in Oken’s genus Prato, of which I have found good examples in the Taro beds of the

Fiji Islands ; and, from what has been above advanced, there can be no doubt of the

existence of external branchize in the Hirudinei.

I have now only to say, by way of apology, that I have written this paper without

having the opportunity of consulting any recent researches on the subject; so that, if I

have mentioned things already known to science, they may at least be taken as corro-

borative matter.

DESCRIPTION OF PLATE XXXIV.

Fig. 1. A little Trematode, with large and complex ventral disk, found in the respiratory siphon of a

species of Melo, Shark Bay: a, oral sucker; 6, genital opening; c, ventral disk; d, external

opening of water-vascular system. x 20 diameters.

2. Posterior extremity of the disk, with cirriexserted. x 150.

3. Central portion of the disk, x 300: a, alveoli of the border; 6, intermediate alveoli; c, openings

of the cirri.

4, Small portion of the disk, x 350, to show the cirri in different conditions : a, completely retracted ;

b, partially exserted, with the apex still inverted ; c, completely protruded.

SECOND SERIES.—ZOOLOGY, VOL. I. 25

oOo CON DS Ot

ON SOME NEW OR LITTLE-KNOWN PARASITIC HIRUDINEI.

. Tortuous course, loop, and return of water-vascular canal.

. Small black Pontobdella, parasitic on Rhinobatis.

. Spermatophore of the same.

. Black leech, with branchial leaflets and alveolated caudal disk, found on Myliobatis.

. Spermatophora of the same.

10.

I).

12.

13.

14.

15.

Portion of the body of the same, slightly magnified.

Inferior surface of the caudal disk, magnified to the same extent.

Marginal portion of the disk, more highly magnified.

White leech, with lateral branchie, on conjunctiva of Green Turtle, New Caledonia.

Larger specimen of the same species, Shark Bay, W. Australia.

One of the branchiz. x 300 diameters.

Trans. Linn. Soc. Ser. Z.Zoon Vou I. Tas. 34

JD. Macdonald del.

g Ota petases beat

Hanhart imp

TREMATODA & HIRUDINEI

CBerjean lith

Bullen

X. On the Osteology and Pterylosis of the Spoon-billed Sandpiper (Burynorhynchus

pygmeeus, Linn.). By JOHN ANDERSON, I.D., .R.S.E., F.L.S., F.ZS., §¢.. Curator

of Indian Museum, and Prof. Comp. Anatomy at Medical College, Calcutia*.

(Plate XX XV.)

Read June 15, 1876.

AS Mr. Harting has already fully stated + all that has been ascertained regarding the

bibliography and life-history of this remarkable little bird, the following observations

will be confined to a description of the particulars indicated in the title of this paper.

These are founded on a specimen procured many years ago at Chittagong in Hastern

Bengal, on the mud flats at the mouth of the Karnafulie river }.

The Skull.—The distinguishing feature of the skull, as is well known, is the expansion

of the distal end of the beak into a quadrangular-shaped spoon, which nearly equais in

breadth the greatest transverse diameter of the cranium. In the specimen on which these

remarks are based all the sutures of the skull have disappeared, the palatines having also

united at their postero-internal angles. The orbit is very large, as in Tringa, its

longitudinal diameter equalling the interval between its posterior margin and the oc-

ciput. The skull has the giobular and smooth character of the pluvialine group of birds.

There is a feeble lambdoidal ridge, large occipital imperfections of ossification, and a

triangular foramen magnum, with the apex of the orifice directed backwards and slightly

upwards, as in Tringa minuta. The occipital and basicranial regions are somewhat

tumid, which is also a character of the Tringa skull; and the palatine and interorbital

regions also conform to it in their general characters. The latter region, however, is

nearly flat, and is more depressed than in any species of Tringa I have examined ; and

the groove of the frontal suture is not prolonged on to it, although the interlacrymal

depression is the same as in Tringa.

The occipital area of the skull is directed upwards and backwards, and, as defined by

the lambdoidal ridge, is of very limited extent compared with the parieto-frontal capacity

of the cranium. The two supraoccipital imperfections of ossification form its most

_ posteriorly prominent part, as is the case with the generality of skulls which have these

fontanelles—so much so, that they are continuous from side to side when the skull is

viewed in profile. They are symmetrical, and are narrower above than below; and

from their lower margins a groove is continued forwards along the sides of the foramen

magnum, defined externally by the tumidity of the exoccipitals. Inside the skull this

* T am indebted to the Trustees of the Indian Museum for their permission to illustrate this paper from the

materials in their possession.

+ ‘The Ibis,’ 1869, new ser. vol. v. p. 427.

+ This is one of the “three specimens in spirits” mentioned by Mr, Harting in the paper referred to (p. 433) as

haying been sent by Mr. J. E. Bruce from Chittagong in 1856.

2F2

214 DR. J. ANDERSON ON THE OSTEOLOGY

venous groove is seen to run along the upper surface of the inward projection of the

periotic; and at its highest part it is partially bridged by an outwardly and upwardly

projecting spicule from the epiotic. In Zringa minuta there are no occipital fontanelles ;

but the groove occurs along the lower half of the side of the foramen magnum, and opens

into the skull by a small foramen ; whilst in 7. salina the fontanelles and groove are as in

Eurynorhynchus. The squamous region is slightly concave, and the postorbital margin

is the same as in Tringa, there being a stylet-like postorbital process projecting down-

wards and forwards, separated by a very shallow notch from the descending process of the

squamosal, which is deeply bifurcated and more strongly developed than in Tringa—

although, as in that genus, the temporal fossa is very feebly marked. Immediately before

the foramen magnum there are from three to four minute foramina for the cranial

nerves ; and slightly anterior to these a larger one, separated from the paroccipital con-

tinuation of the previously mentioned groove. ‘The paroccipital process in this, as in the

Limicoline skull generally, is thrown much forwards onto the base of the skull, and is

directed forwards and downwards. Its inner side is perforated by a small foramen,

which, however, does not occur in Tringa minuta or T. salina; and the apex of the

process nearly touches the lateral expansion of the basisphenoid; the upper wall of the |

imperfect ring so formed is partially divided into two separate channels. Although the

orbit has the same general characters as in Tringa, the septum is more complete, having

a greater expansion superiorly. The ethmoidal ale are also more strongly developed

than in that genus, their superior and inferior external angles being more marked and more

outwardly projecting. The lacrymal is also more pronounced and formed somewhat

differently from that of Tringa, in which the downwardly prolonged portion that abuts

against the ethmoidal alee is a very fine osseous thread; while in Hurynorhynchus the

curve is less and the bone is stouter, and a sharp, but short, process is given off from the

outer margin, where it turns into the ethmoidal plates. The superior or orbital angle

of the laerymal is also stronger and more outwardly projecting than in Tvringa. The

nasals have a small anteriorly projecting lamina developed from their external margins

in a line with the posterior border of the nostril, and which does not occur in 7. salina

and 7. minuta. The quadrate bone, although proportionally larger, has the same form

asin Tringa; but it differs in having the articular surface for the mandible more deeply

concave, and, in the inner: side of the same surface having a much shallower notch.

The air-cells open behind, and between the proximal facets, by a round orifice, which

appears to be occasionally supplemented by a smaller one, and on the inside of the bone

by two or three such openings. The pterygoid has the general form and relations of the

Schizognathous. skull, but is much stronger than in Tringa. The basipterygoid process

is more produced than in that, genus; and there is a small process posterior and internal

to it, which, does. not occur, or is only feebly developed, in Tringa.

The palate is essentially Schizognathous; but, as already mentioned, the palatines have

ankylosed in the mesial line, and their vomerine processes have amalgamated with the

vomer, which is short and bifurcate at its tip, and does not extend forwards beyond the

origin of the nasal process of the maxilla. The palatines have much the same form as

in Charadrius pluvialis. The external lamina is at a slightly lower level than. the

AND PTERYLOSIS OF EURYNORHYNCHUS PYGMAUS. 215

internal, although the under surfaces of the palatines are but slightly concave. A notch

occurs behind the middle of their internal margins; and from it posteriorly a small de-

pendent lamina is prolonged backwards, gradually diminishing in size towards their

postero-internal angles. he concavity of the palatines is continuous with the notch.

In Tringa the notch is more strongly marked, and there is a contraction of the external

margin of the palatine in its neighbourhood, the dependent plates also in that genus

being more developed than in Hurynorhynchus; and the portion before the notch is like-

wise more concave. An anteriorly directed foramen occurs near the inner border a

short way before the notch. The maxillo-palatines in no way differ from those of Tringa.

The internasal septum has a large imperfection of ossification in its centre; and its

inferior border is jagged, terminating under the middle of the aforementioned lateral

lamina of the nasals. It has much the same capacity as in Tringa, in which, however,

its lower border slopes upwards in an unbroken line. The jugals and maxille call for

no remark.

As already stated, the distinctive feature of the skull is the great development of the

‘distal extremities of the preemaxille into a broad four-sided spoon—two anterior and

two lateral borders, the former meeting at an obtuse angle with

each other, and at an acute angle with the latter. It is

marked by a longitudinal mesial groove on its under surface,

and by a corresponding low ridge on its dorsal aspect. At

the point of divergence, where the nasal and palatine pro-

cesses are given off, there is an orifice, which, penetrating

between the plates of the premaxille, divides into a great

number of canals for the transmission of the nerves and blood-

vessels. These canals appear on the surface as so many fine

ridges, radiating forwards and outwards from the point of

entrance, and open along the anterior and lateral margins of fig. 1. Bill of Eurynorhynchus

the spoon, more especially the former. On the under surface ipo gpocers ta puoi,

there is one well-marked regular row, with a few somewhat Fig. 2. Upper view, head of

scattered foramina behind it; and on the same surface, on a ee sve een ePite

slight eminence, external and separated from the mesial .

furrow by a slight concavity, there is, on each side, a row of from 6 to 8. foramina.

A similar structure prevails in the mandible; but the positions of the ridges. and orifices

are reversed. A similar but less-intensified structure also prevails in Tringa..

Vertebral Column.—There are 15 cervical vertebree, as in Tringa. The atlas is a

narrow ring, with the odontoid ligament strongly ossified, and a pair of. obscure processes

on the under surface of the segment. In Tringa minuta the ligament is unossified, and

the two hypapophyses are much more developed. The axis has a thick nodose spinous

process, with a falcate backwardly projecting under process, and with two nodose processes,

Fig. 1.

* Through the kindness of Mr. Harting and the Editor of ‘ The Ibis’ the use of the above woodcuts has been

obtained. In the paper in ‘ The Ibis,’ 1869, Mr. Harting mentions them as being from the pencil of Mr. Swinhoe.

They are here introduced that the head with feathering of this singular bird may conveniently be compared with.

the bones of the same, given in Plate KXXXY.

216 DR. J. ANDERSON ON THE OSTEOLOGY

one on either side of its base. In the same species of Tringa the two spines have much

the same form as in Hurynorhynchus, but the nodose processes are absent in the inferior

one. All the remaining vertebrae of Hurynorhynchus, in their number, even to the

minute details of their structure, conform to the same parts in Zringa. The ribs also

call for no remark.

Shoulder- and Wing-bones.—The scapula is rather more dilated in its posterior half

than in Tringa minuta; and the suprascapular border is bevelled off to a point from its

lower margin, whereas in the latter species it is less pointed. The coracoid has much

the same proportion to the scapula and to the total length of the sternum as in Tringa ;

the head of the bone has the same form as in 7. minuta; similarity also prevails in the

shafts and distal ends; and both birds have hooked epicoracoid processes of one type.

The clavicles are the same as in the species of Sandpiper compared.

The sternum only differs from the sternum of Zringa in the anterior border of the

keel being less forwardly curved. The notches are the same in both forms; and a

similar imperfection of ossification exists at the anterior superior border of the keel.

The humerus is proportionally longer to the antibrachium than in 7. minuta. From

the humerus of the latter it is also distinguished by the feeble ridge occurring on the

preaxial side of the proximal extremity of the humerus of that bird being developed in

Eurynorhynchus into a strong triangular anteriorly curved process. The antibrachium

and manus have the same relative proportions as in Trimga.

Pelvis and Hind Limb.—The chief difference that exists between the sacrum of the

Spoon-billed Sandpiper and that of 7. minuta is the slightly greater dimensions of the

ilio-sciatic foramen. In Hurynorhynchus there is narrowing of area immediately below

the foramen, and the long free portion of the pubis is given off nearer to the acetabulum,

whilst it is more divergent from the ischium. The femur is very little more than

one half as long as the tibia, being proportionally shorter than in 7. minuta, the femur

of which it otherwise closely resembles, although somewhat stouter. The cnemial

process of the tibia is the same in both birds; and the perfect fibula of Hurynorhyn-

chus is more than one half as long as the tibio-tarsus. In all other respects the hind

limbs of the two genera are alike.

The Hyoid Arch.—The entoglossal cartilage is considerably longer and stouter than in

Tringa salina; and the entoglossal bone is shorter and broader, with a much larger

imperfection of ossification than in that species; but im other respects the remaining

elements are alike, only larger in Hurynorhynchus, owing to its greater size.

The Viscera.—in Tringa salina the intestine measures 7°75 inches, and in Huryno-

rhynchus pygmeus 8°75 inches. In the former the large intestine is 0°78 inch, and its

cecal appendages 0°62 inch long, while in the latter the great intestine is 0°88 and its

excum 0:70 long. The stomach and proventriculus are the same in these two forms.

The liver of Hurynorhynchus appears to be somewhat smaller, and differs from that

of Lringa salina in having the right lobe flattened externally, and much more pointed at

its lower end, which terminates in a sharp projection, this being somewhat the character

of T. minuta. Much importance, however, cannot be attached to such slight hepatic

differences, because these are liable to occur in individuals of the same species. The

AND PTERYLOSIS OF EURYNORHYNCHUS PYGMAUS. 217

tongue is longitudinally grooved, and fringed at its extremity in the same way as in

Tringa; but it is nearly thrice as broad as in that genus. It rapidly diminishes in

breadth towards its extremity, and contracts to a sharp point.

Pterylosis.—There are 12 rectrices and 24 remiges, of which the first two are the

longest and of equal length, the pinion adhering to the Tringa type in having ten

remiges. The wing, however, is more deeply bifid than in Tringa minuta or T. salina;

this is owing to the greater length of the posterior secondaries, and to the slightly shorter

character of the feathers intervening between them and the first primaries, the 23rd and

the 24th feathers of the wing being of nearly equal length.

The spinal tract follows the Limicoline type. The interhumeral clefts are much more

strongly marked than in 7. salina; and the posterior part is feeble at its commencement,

and its two halves meet at the tail. The spinal space is thus long and lanceolate. In

the anterior portions of the posterior clefts the feathers are arranged obliquely from

before backwards and outwards, in rows of from 3 to 4 feathers, which become more

and more longitudinal in their course, till at last, in the undivided extremity of the

spinal tract the feathers are in transverse rows of generally 5 feathers—the rows

being slightly backwardly convex, and the feathers diminishing in numbers as the tract

is traced backwards.

The oil-gland is pyriform and bare, with the exception of a circlet of feathers around

its orifice. The ventral tract is divided by the ventral space immediately before the

sternal keel; but in Tringa salina the division occurs slightly anterior to that. Its

branch, or pterylosis lateralis, is broad and strong, as in 7. salina, and is prolonged.

aleng nearly two thirds of the chest. The truncal branches are rather feeble at first,

but are stronger and broader on the abdomen, as in 7. salina. The humeral tract is.

strong, and at its origin unites with the last mentioned. The femoral or lumbar tract

originates from the posterior extremity of the spinal tract, as in Tringa, and has the

same general character. The lateral neck-space extends two thirds up the neck; and the

other spaces are the same as in Vringa and its allies.

Concluding Remark.—The foregoing examination of the osteological and other cha-

racters of Hurynorhynchus reveals only one important feature wherein this bird struc-

turally differs from the genus Tringa, namely the singular expansion of the bill, the

structural equivalent of a similar modification in Platalea leucorodia. Indeed this

species, as pointed out by Mr. Harting (/. c.), was originally placed in the genus Platalea

by Linnzeus *.

* Mus. Ad. Frid. i. Prodr. p. 26 (1764).

218

Fig. 1.

OSTEOLOGY AND PTERYLOSIS. OF EURYNORHYNCHUS.

DESCRIPTION OF PLATE XXXV.

Skeleton of the Spoon-billed Sandpiper (Eurynorhynchus pygmeus),in profile. Of natural dimen-

sions; and all the other illustrations are also drawn to the same scale.

. The skull, seen from above, and displaying the expansion of the premaxillaries and their

foramina.

. The palatal or lower base of the same cranium.

. An upper view, or oral surface of the (lower) mandible, its terminal width corresponding in

outline to the premaxillaries.

. A side view of the hyoid bone.

. The bones of the right wing, on their outer or dorsal surface.

. The right humerus, its imner aspect.

. Femur of right side, seen from in front.

. Tibia and fibula of same leg, also their anterior aspect.

. Tarso-metatarse of right limb, front surface.

. Pelvis and caudal vertebre, viewed from above. ‘The posterior rib of the right side remains.

. An inferior view of the sternum, furcula, coracoids, and costo-vertebral parts of the chest of

Burynorhynchus pygmeus.

S ANLETSO?, TRANS. LINN. Soc. SER. 2 Zoou. Vou I, Ph 35.

OSTHOLOGY OF THE

a, Lith THE SPOON-BILLED SANDPIPER, Banks & Co, B

HURYNORHYNCHUS PYGMABUS.

THE

TRANSACTIONS

THE LINNEAN SOCIETY

LONDON.

SECOND SERIES.—ZOOLOGY.

VOLUME I.

PART THE FIFTH.

LONDON:

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET:

SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON-HOUSE;

AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER-ROW.

M.DCCC.LXXVII.

TOUS 28 Lire noose

oP a a ae et

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ROU) CLG IU RRRRRI be te a em ase

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CONTENTS.

PART V.—Ocrosun, 1877.

/XI. On Steere’s Sponge, a new Genus of the Hexactinellid Group of the Spongida.

By James Movriz, WD., FLS., F.GS., Fe. (Plates XXXVI. & XXXVII.)

page 219

/XII. Notes upon the Oxystomatous Crustacea. By EpDwarp J. Mirrs, 7.L.S., F.Z.S.,

Assistant in the Zoological Department, British Museum. (Plates XX XVIII.

IN MY ee JRE Es OE TG cute CRO UM Ie baenng halt 1.935

vXIII. On the Domestic Pig of Prehistoric Times in Britain, and on the mutual Rela-

tions of this variety of Pig and Sus scrofa ferus, Sus cristatus, Sus andaman-

ensis, and Sus barbatus. By Grorce Rouixeston, UD. PRS, F.L.S.,

Linacre Professor of Anatomy and Physiology, Ozford. (Plates XLI.-

XLII.) MR iit Rg Nee id tee ag Sol me e OONL

vXIV. On two new Forms of Deep-sea Ascidians, obtained during the Voyage of H.MU.S.

‘ Challenger. By H. N. Mosstzy, IA., F.R.S., Fellow of Exeter College,

Oaford. (Communicated by Dr. J. Murtsz, F.LZ.8.) (Plate XLIV.) . . 287

/XV. On new Forms of Actiniaria dredged in the Deep Sea; with a Description of

certain Pelagic Surface-swimming species. By H.N. Mosetey, V.A., F.R.S.,

Fellow of Exeter College, Oxford, late Naturalist on board H.M.S. ‘ Chal-

lenger. (Communicated by Dr. J. Muniz, F.L.S.) (Plate XLV.) . . 295

teh) RE Ay bee

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| hy Sor ery IBS aa i vi ty urs pig eh if KR: Lay L hae 1M radish sat

; Dre lena Teer rhc oenteng s if. aon Dewey weed: % cotai hh tii Rag

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Al oo) Woes LA EATS LL DROME era M A ee? ail ie ih OH 43

Fe 8

XI. On Steere’s Sponge, a new Genus of the Hexactinellid Group of the Spongida.

By James Muniz, ICD., F.LS., F.GS., Se.

(Plates XXXVI. & XXXVI.)

Read January 20, 1876.

THE Order of Sponges now designated Hexactinellida has only of late years been

prominently brought forward to the notice of naturalists. As early as 1833 by Quoy and

Gaimard, and again more definitely in 1841 by Stutchbury, at least two or three forms

of the group had been registered in zoological literature ; and later on a few new members

were occasionally added. From about 1860, or even a few years later, up to the present

time a number of new and most curious genera and species have been minutely described.

'The issue of the monographs of Bowerbank, Oscar Schmidt, and Hackel, among others,

have greatly helped to advance and give prominence to the history of the Spongida; but

the study more especially of the Siliceous Sponges, however, received an unexpected

impetus when the first results of the deep-sea explorations became known. Besides

Dr. Gray, ever active in many fields, the labours of Carter, Wyville Thomson, Saville

Kent, and Perceval Wright, among our own countrymen, have borne good fruit so far as

the group in question is concerned. Not only has the deep-sea dredging yielded a partial

resolution of some of the difficulties of structural organization of certain of the more

singular kinds, but another item of considerable importance has become evident, as fresh

faunal ground has accrued: I allude to distribution. The geographical dispersion, then,

of the Siliceous or, as some prefer to call them, Vitreous Sponges, where unmistakable

closely allied kinds are widely severed, suggests many possibilities, geological and other-

wise, though no explanation is capable of easy proof from our as yet limited data. Every

new genus or species entered into the catalogue thus has a cumulative significance,

bearing ratio, moreover, to its peculiarities of build, where these either exhibit novelty,

intermediate characteristics, or set at rest moot points. On grounds such as these the

noyel specimen I herewith call attention to I believe possesses unusual interest.

History or Spectmen.—Professor Joseph B. Steere, M.A., LL.B., of the University of

Michigan, U.S., a few years ago resolved on enriching the Museum collections of his

alma mater. Young, enthusiastic, and fond of adventure, after an exploration and

somewhat lengthened stay in the regions of the Amazon, sending home valuable material,

he traversed the continent, crossing the Andes. Thence by the Pacific Ocean towards

China, he remained awhile in Formosa. Afterwards making his way to the Philippines,

he resided for longer or shorter periods in several localities of the group. In devious.

voyages, visiting the more notable islands of the Malay archipelago, he gathered a vast

array of objects of Natural History. Time speeding, he betook himself homewards, and

en route made a hasty stay in London, during the beginning of the month of October 1875.

Circumstances did not permit of his boxes and cases of material being unpacked here,

SECOND SERIES.—ZOOLOGY, VOL. I. 26

220 DR. J. MURIE ON STEERE’S SPONGE,

a few loose packages only being opened to gratify the curiosity of friends. Mere accident

in conversation led me to a sight of a sponge the subject of the present communication.

Its appearance and history in a trice convinced me of its rarity, and I advised the

owner to take advantage of the extensive collection at the British Museum, and after _

comparison &e. to describe the specimen. If ultimately deposited in the distant Michigan

Museum, the chances were its value and uniqueness might temporarily be lost sight of

among the plethora of his collections, and a long while elapse ere opportunity offered to

bring it before the scientific public. Neither time nor indeed inclination to a special

study of the group rendered Professor Steere desirous thereupon to enter on the task;

but still, knowing the interest attached to the object, he requested my drawing out a

notice of it for publication. This could have been little else than a mere superficial

survey had he not given me possession of the tempting specimen for a brief period, to

facilitate closer inspection and to procure a photograph of it. He still more generously

gave me a portion, quite enough for a thorough examination of its microscopic structure,

as the accompanying illustrations bear witness.

The following is Professor Steere’s verbal account to me of how and where the sponge

was obtained. When in the Philippines, 1875, his attention one day was called to what

at first sight he deemed to be a branching coral, hanging in the sun near one of the

native dwellings. On purchasing it, he learned from the fisherman, its first possessor,

that it had accidentally been dragged up by his hooks* from a depth of what he supposed

to be 400 to 600 feet, 7. e. roughly, then, between 70 and 100 fathoms. The locality where

originally obtained is the narrow deep channel between the strip-like and parallel islands

of Negros and Zebu. This channel is said to be some 100 miles long, and varies from

2 to 3 miles in breadth. The water is comparatively still, or with only a slightly percep-

tible current, never, generally speaking, being boisterous. This condition in the main

results from the construction of the channel, which to some extent is land-locked, as

reference to a chart of the district shows.

As regards the nature of the sea-bottom, Professor Steere believes it to be sandy. As

to the geological aspect of the surrounding country, according to him the island of Zebu

is chiefly characterized by a limestone formation ; but the larger island of Negros bears

apparently marks of volcanic origin.

He further informed me that the Huplectella and other peculiar sponges hitherto

* Dr. Meyer (Ann. & Mag. Nat. Hist. (4) 1874, vol. xiii. p. 67) has drawn attention to the mode of fishing for

sponges adopted by the natives of the Philippines. A number of small hooks are fixed to a light framework, and this,

sunk by a stone, is dragged on the bottom in depths quite 50 fathoms or more, and the hooks catch the objects as the

boat drives with the current. A confirmation of this has been published by one of the staff of the ‘ Challenger’

(the ‘ Times,’ 30th April, 1875, and quoted by Bowerbank, P. Z. 8. 1875, p. 506). But see also Mr. T. J. Moore’s

letter to the late Dr. J. EK. Gray on the mode of dragging for the “‘ Regadera ” (Huplectella) with a bamboo and nails

at Talisay, in the island of Cebu, illustrated (Ann. & Mag. Nat. Hist. 1869, vol. i. p. 196). Itagrees substantially with

what Professor Steere relates of how the fisherman obtained his specimen of sponge. In many of the Euplectellas sent to

this country, ragged holes or other slight injuries to the spicular framework indicate where jt has been caught by

the fisherman’s hooks. Since this paper was read, I have myself seen in Edinburgh one of the fishing implements in

question, brought home by the officers of the ‘Challenger’ Mxpedition; and I may state, moreover, that a capital

drawing representing a similar native instrument was exhibited at one of the meetings of the Linnean Society by

Capt. Chimmo, F.L.S.

A NEW GENUS OF THE HEXACTINELLID SPONGID. 22M

brought from the Philippine group are obtained by the natives in plenty on the opposite,

eastern, side of the island of Zebu, but that no sponges are said hitherto to have been

got from the sound or channel above mentioned.

One day, while in the forest, Professor Steere was suddenly apprised of the arrival of

the ‘Challenger’ Expedition, by a note from Professor Wyville Thomson, who had heard

of him and his whereabouts. He hastened on board, passing, as he assures me, a most

agreeable day in the company of our countrymen. He doubts if the English naturalists

obtained any other specimens of this sponge (Dendrospongia), though he told them of his

one, and where and how acquired*.

I may only further add to the history of the specimen that its transport has been an

adventurous one. Its great size and fragile nature prevented packing up in the boxes

with the other objects collected. Safe carriage then depended on its being carried about

by hand. To this end a native grass cord was fastened to the stronger portion of the

root, the more delicate branches were partly wrapped in cotton-wool, and the object

inserted in a paper bag, with a loop of the cord out. Thus slimly protected it afterwards

was borne for hundreds of miles on foot and on horseback; afterwards on shipboard it

was slung to the cabin roof, and there freely dangled during the subsequent long voyage

homewards. The wonder is such a delicate brittle organism escaped injury and smash, or

was preserved tolerably intact through such a journey.

EXTERIOR AND Sections.—This example of Steere’s sponge may well be compared in

outward figure to a shrub, or even with greater propriety, so far as superficial appearance

is concerned, to some of the branching corals. The above remark of course only applies

to the siliceous skeleton, the sarcode and dermal membrane being quite unknown to me.

The lower basal end is laterally expanded and to a certain degree flattish, the thickness

being less by one fourth than the breadth. The inferior left end (see fig. 1, Pl. XXXVI.)

has a depending, slightly curved, broken stump or kind of resemblance to a root, which,

however, may have been part of a horizontal branch; in this case the whole sponge might

have had a greater tilt or inclination to the right than the drawing exhibits. To the

inferior right corner a smaller broken branch projects upwards and slightly outwards.

Above this the basal portion mounts as a gently sinuous stem, giving off three irregular,

short, stoutish main branches about equidistant above each other. The middle one has

a sinistral obliquity, but only a short and considerably worn part of it remains. The

right main branch is given off lowest, and has received rather rough usage and consi-

derable abrasion ; about 2 inches from its commencement a knob only shows evidence

of a branch now absent. As much again higher up is a trifurcation, and each of these

secondary branches bifurcates, terminally producing the long upright but curvilinear

whorled surface branches, one, if not two, of these exhibiting short forked ends. The

left main branch also divides into three secondary branches expanding outwards; each

of these again bifurcating and trifurcating, and some terminally being forked like those

on the right. When in the living state doubtless this sponge has a somewhat more

* Since reading this communication, Sir Wyville Thomson’s return has enabled me to show him the accompanying

Plates ; the species he does not recognize as being among their collections, although it is possible a less perfect spe-

cimen or fragmentary portions may be found among the stores of material as yet imperfectly worked out.

262

222 DR. J. MURIE ON STEERE’S SPONGE,

arborescent outline than that now represented (its dendritic characters suggesting the

generic term), but I have preferred abiding by the photograph rather than adding branches

where fractured portions indicate their former presence.

Dimensions of sponges are so exceedingly variable that this character is only of very

subsidiary importance. Withal in the original description of a type full mastery of detail

is needful, for one never knows on what trivial peculiarities important issues may after-

wards be raised. Admeasurements in the present instance can be but rough approxi-

mations and not mathematical verities. On comparing the great Askonema of the

Portugal coast with its relatively diminutive representative Crateromorpha of the

Philippines, Carter*, in his excellent commentary on the group, says :—“ In short, like

most things in the west, if the Hexactinellidee do not surpass in beauty, they do in size,

for the most part, those of the east.” Steere’s sponge, however, has an area equalling

the large vase-like Askonema, its branched character, though, differing widely.

Its greatest vertical height is 3linches; from tip to tip of the most distant upper

branches is a trifle over 28 inches. Some 8 or 9 inches may be allowed for the length of

the broad lower, trifid stem portion; and the several longer branches vary, three measured

being respectively 16, 20, and 23 inches in their entire length. Taken by compasses, the

breadth quite at the flattish base, or owhat for convenience’ sake may be termed the root,

is fully 54 inches, but the same in the opposite diameter or thickness is barely more than

12 inch. Higher up and just below the broken middle main branch the stem lessens in

breadth to 3 inches across. The girth taken round each of the three so-called short main

branches, where about to divide into their numerous long upper branches, is from 44 to

5 inches. The long whorled branches have a circumference, at what further on will be

explained as the frills, of from 3 to 34 inches, that is, a diameter of about an inch; the

erooved portion intervening between the whorls, however, very little exceeds half that

diameter.

In allusion to the superficial aspect as it looks to the eye and under a hand-lens, the

way in which the specimen has been tossed and carried about should not be forgotten.

Whilst many of the top twigs and branches here and there have wonderfully escaped

injury, and the surface can be made out in detail with tolerable accuracy, other portions

are greatly abraded, and particularly so the stem part. he latter seems to have been

covered by a dermal membrane, now almost entirely rubbed off, at spots only a kind of

fluffy residue proving its former presence. But the abrasion notwithstanding eliminates

other features, especially the marked nature of the pores and oscula as connected with

the internal tubular and canal system. The inferior cracked and fractured pieces also

clearly point out that the said root- and stem-spicules are to a certain extent stronger and

more solid than those higher up.

The colour of the entire siliceous skeleton save lower parts is frosty white, not quite so

lustrous and pearly asis the “glass rope” of Hyalonema, nor so dull as Dactylocalys ; but

yet the marginal spicules are translucent, and in spots, or under certain lights, some parts

are more brilliant than others. The lower stem and so-called root is as a whole to the

eye of a browner tint, its spicular meshwork with a pale yellowish hue. But this coloration

* Ann. & Mag. Nat. Hist. (4) 1878, vol. xii. p. 369.

A NEW GENUS OF THE HEXACTINELLID SPONGID. 223

is evidently due to the presence of a greater amount of dried-up sarcode mingled with

interstitial foreign substances. This darker shade, moreover, brings out in relief specks

and lines of pure solid white, viz. sparsely scattered minute shells of Serpula. These

partially are superficial and partially are deeper seated in the sponge network. In one

hole I observed a small Crustacean, seemingly an Amphipod.

The exposed surface of the stem has been rubbed down moderately even, but never-

theless is far from smooth, being drilled by innumerable holes—in fact resembles shaven

cancellous texture, as is its real nature. The orifices are irregular in size and pattern,

though the most prominent are either circular or ovoid. The larger have a diameter of

nearly 3/5 inch, and are more widely scattered than the greatly more numerous and much

smaller intervening ones. Here and there also the former are so obliquely cut across or

worn as to form short shallow grooves or canaliculi; whereas the finer more angular holes

spring from a delicate lacework, of which more hereafter. 'The said perforations evidently

correspond to the oscula and pores of other sponges. The general surface, moreover,

exhibits several broad but very shallow impressions or concavities which traverse the stem

obliquely. These slightly scooped areas are seemingly traces of the deep hollows so

well marked between the whorls of the upper branches presently to be spoken of.

The distinguishing feature par excellence of the branches is a series of tufts or rosettes,

so continuous and interwoven in the main as to present a whorl running successively

round from base to apex. Hach branch in this particular may be likened to a thick

stranded coil of rope, or, still better, resembles the spiral twist of the horn of the Indian

Antelope (Antilope cervicapra). In some instances where two forks or branchlets arise

close together from a branch, before they tend to divaricate widely apart, their spiral fringes

in passing each other so interlock as to form bridges of union (see fig. 2, Pl. XXXVI.).

To the naked eye, but still more manifest with a pocket-lens, the spiral tufty elevation is

seen to be composed of a bunch of long parallelly placed spicules, which issue from the

axis of the branch at an oblique angle, and slightly spread out at their free ends. The

radiation, however, tends upwards or chiefly towards the direction of the spire; and thus

in each turn the upper border of the fringe or frill gently overarches the intervening

depression, its lower border proportionally less so. The apex of each branch is closed,

ending with a semitwist of the spicular frill. The said spicules are like so many glass

bristles; but their true character is considerably masked by their being bound together

throughout by short transverse and oblique secondary spicula, which produce a network

further to be described under “ Microscopic Structure.’’ What is more plainly visible in

this outward examination is a superficial frosty-like network of fine spicules, these, where

best preserved, causing a woolly appearance; their precise nature will again be discussed.

The spiral hollow between the frill differs from the latter rather in degree than in kind,

the relative absence of long protuberant spicula being its characteristic. The depth

varies, but averages a quarter of an inch. Its bareness brings out two features never-

theless, which are subdued in the frills. One is multitudinous oscula and pores, already.

sufficiently alluded to in the stem; the second, even more deeply interesting, is the presence

of a most lovely, delicate gossamer lacework, composed of minute spicules forming a

rectangular chequer. This is not everywhere intact, but sufficient traces of it exist as to.

224: DR. J. MURIE ON STEERE’S SPONGE,

render it likely that it extended over a great part of the branch, probably excepting the

summits of the whorled frills. Whether it extended over the stem it would be hazardous

affirmatively to assert, though the chances are it may have had a partial covering, basing

judgment upon what we know of other forms possessing a veil. Dr. Millar first drew my

attention to this structural point, and then we thought it only stretched over the large

orifices of the stem, a view we afterwards found erroneous. The significance of this

surface layer, both in a morphological and physiological point of view, I am inclined to

think is of considerable importance, but, as will be seen, it is not confined alone to this

form of hexactinellid sponge.

Having run over the most noteworthy points of the external features of our specimen,

I come to the interior. In studying this, pieces were cut across and in the long direction

of the axis. The transverse sections of both stem and branches disclose not a hollow

cylinder as in Aphrocallistes and Aulodictyon, a funnel-shaped or narrowed fusiform

cloacal cavity as in Rossella and Meyerina, nor yet quite such close areole as in Dactylo-

calyx, but rather one may say a kind of compromise between the last two. While tubular,

these openings are unequal, the larger corresponding to the exterior oscular apertures,

the lesser to the pores aforementioned. The latter, angular and all sorts of shapes, are

bounded by the small cross and tangental spicules; the former, rounder or ovoid, are the

open passages between the larger-sized bundles of vertical and obliquely radiating spicules.

In the centre of the body of the stem the open tubular character is best marked towards

the periphery; radii here are more visible, between which the short secondary spicula

intrude, while quite at the edge the frostwork of dermal sexradiate spicula encircles.

The longitudinal median sections still further explain the preceding, and morever, in |

some respects, agree with Meyerina, though in the general appearances and relative

thickness of the spicula an approach is made possibly nearer to Dactylocalyx. Longi-

tudinal fibrous-looking spicula appear to run up the entire length of the axis, but are in

fact a continuous repetition of stout sexradiate spicules, the cross bars being the’ short

arms. These median vertical pillars, moreover, most obviously branch off curvilinearly

towards the margins; and it is evidently the production of these which gives rise to the

exterior whorls. Thus there is a certain plumular arrangement, the obliquely directed

spicules corresponding to the barbs, the smaller annectent spicules to the barbules, and

the dermal appanage of terminal sexradiates to the so-called extradivisional fluff of owl-

feathering.

. Mrcroscoric Srructure.—Before adverting to this it behoves me to mention the in-

valuable assistance derived from my friends Dr. John Millar and Mr. H. J. Carter. To

the former I am indebted for making my best illustrative microscopical specimens, and

also with liberal hand placing others of different genera at my disposal for comparison,

besides discussing points connected with the interpretation of structure. Mr. Carter’s

labours and knowledge of Sponges, and especially of the group in question, are well

known and justly appreciated. When I add, then, that he has carefully examined portions

of the sponge, compared these with the figures in my Plates, and, moreover, measured the

spicula, drawing out a descriptive analysis of same for my use, it will be admitted I have

been amply helped and stimulated in investigating Professor Steere’s specimen. In my

A NEW GENUS OF THE HEXACTINELLID SPONGIDA. 225

own examinations of the structural peculiarities I have used object-lenses of both low

and high defining powers, according as delineations of the coarser parts or of the minute

spicula required.

The different constituents of the skeleton whereof Dendrospongia is built up I enume-

rate as under-mentioned; the measurements of spicules are after Mr. Carter.

1. The Vitreous or Glassy Fibre (figs. 4 to 18 inclusive)—This composes the basal

structure of the sponge, forming its tolerably compact root, stem, and cylindrical branches.

In the living state it supports the sareode, which is charged with other minute spicula,

to be described in the sequel. The glassy fibre is thickest in the central or oldest portions

of the root, stem, and branches, and thinnest towards the circumferential parts of same.

Admeasurements of that of the centre yield a thickness of +4, inch, whilst the smaller

fibre is about z 55 of an inch in diameter. -Hvery intermediate thickness between these

is met with, and, as will be understood, the earliest condition of the growing spicula may

even be less than the lowest dimension above mentioned. As has previously been adver-

ted to in remarks on the sectional naked-eye view, the fibre presents main longitudinal

and innumerable intercrossing branches—that is, extending upwards and outwards round

the axis, leaving room for the canals of the excretory system, which take the same

direction. These complicated anastomosing reticulations, however, are products of hex-

actinellid spicula, the various arms of which may either retain their straight character,

or, bending to a lesser or greater extent (figs. 12 and 22), give rise to sinuosities as the

case may be—their overlapping union and growth, with here and there ultimate coales-

cence of adjoining arms, producing all manner of configuration, and, indeed, the appear-

ance of fibre throughout. The whorls, to some extent already explained, are due to the

extension towards the circumference of the long, pointed, terminal filaments of, so to say,

certain strands of the spicular fibre. Thus the protruding pitchfork character of the

outermost hexactinellid fibres (over which the dermal network, fig. 9, is attached) results

from the central and horizontal arms of the spiculum being elongated and turned

outwards (fig. 12) towards the free surface of the sponge-branch, while the sixth or

inner arm is in continuation with the rest of the spicular glassy fibre internally. The

fibre itself is less or more roughened, that is, covered with short spines, very pronounced

and prominent in the younger spicular fibre.

With regard to these spiny productions on the fibre, Mr. Carter has called my attention

to a point concerning them, the importance of which I

was not then aware of. Under a high magnifyine-

power the said elevations are found not to be simple,

conical, and pointed, but, instead, “‘mucronate,”’ that is,

with the summit divided into two, four, or more expan-

ded points (see woodcut, fig. 1). He further observes

that the only other instance he knows of in which this

form of spine is present is in Parrea densa (‘ Annals,’

1873, vol. xii. p. 463, pl. xvii. figs.5 and 6). The accom- Diagram of spinous processes on the

panying woodcut is a reproduction of his sketch, kindly vitreous fibre of Dendrospongia, on the

forwarded in a note to me.

Fig. 1.

scale of 4, to =, inch. After Carter

226 DR. J. MURIE ON STEERE’S SPONGE,

2. Acerate spined Spicula (figs. 17, 18, and 19)—These are of two sizes, large and

small. Both sorts are long, acerate, fusiform, inequilateral, one half being thicker than

the other. The thicker moiety is rather more abruptly pointed than is the attenuated

one. The larger-sized spicula average yo inch long by gop inch at their thickest part.

The smaller acerate spicula in most respects agree with the preceding, but are not more

than one fourth or one fifth the size of the others. More frequently I have met with

them as represented (fig. 19), in bundles, whereas the larger spicula are usually single and

apart when obtained. The spines covering the large spicula (very manifest in fig. 18)

point away from the thick end, and they all slope in the same direction, closely applied

and almost parallel with the shaft. Hach spine is simple, smooth, and conical, supported

on a projecting basal portion of the shaft. These acerate spined spicula, and particularly

the big kind, may be regarded as subskeletal, being situate throughout the structure of

the sponge, though chiefly towards the surface. They appear to be a very common form

among the Hexactinellid sponges.

According to Mr. Carter, when elongated they apparently pass into the anchoring

spined spicules of the bearded species of the Hexactinellida. Moreover such spicula he

regards as “subskeletal,” because he avers they are often drawn into the vitreous fibre,

with which they become indistinguishably incorporated durimg life; he even holds that

the larger scopuline spicules, presently to be described, are occasionally similarly amal-

gamated.

3. Scopuline Spicula (figs. 13 to 16).—Two forms of these obtain, likewise differing

in dimensions. (a) The larger of the two spicula (fig. 13) consists of a straight shaft and

terminal arms from two to four in number. The shaft is microspined, inflated and pointed

at the free extremity, and the opposite end terminates in a palmate, bifid, or quadrangular

inflation, springing therefrom two to four and occasionally five arms. These latter are

opposite, extended fork-like almost in a line with the shaft itself, and are microspined

and indistinctly capitate, or with convex head. The shaft of these large scopuline spicules

is 2; inch long, and the arms yygq inch long. (4) The smaller scopuline spicula

(fig. 15) have also a straight shaft and from two to four terminal arms. They differ from

(a), besides size, in the arms being opposite and expanded laterally or flower-like. Both

shaft and arms are microspined, the arms very indistinctly capitate. In dimensions these

second kind of scopuline spicules have a shaft z3e inch long, the arms yy'95 inch long.

Besides their terminal arms, both kinds of the spicula under consideration are often more

or less provided with shorter armlets, which spring from the body of the shaft at right

angles, and vary in number as well as in length (figs. 14 & 16). While they are often

hardly traceable in the large scopuline form (a), they on the other hand are occasionally

profuse in the smaller kind (2), numbering in the latter frequently as many as twelve,

which usually are longest towards the middle of the shaft. These different figured sco-

puline spicula pass by almost imperceptible gradations into each other, though in most

slides under the microscope the two sorts are readily distinguished even when lying in

heaps, as is shown in fig. 11.

Mr. Carter remarks that the said scopuline spicula belong to the sarcode inside the

dermal meshwork, where their original arrangement has been destroyed by the contracting

A NEW GENUS OF THE HEXACTINELLID SPONGIDA. 227

of this sarcode in drying and subsequent partial destruction of it by mucedinous fungi.

They thus now appear in little detached masses without any definite arrangement,

although, here and there, in the dry sarcode which holds them together, a circular hole

may be seen indicative of their position and office in supporting the internal porous sar-

codic expansions during the lifetime of the sponge.

Indeed the sarcode generally has been destroyed

throughout this sponge in this way, and now only

hangs in small fragments here and there in the

otherwise naked fibre (consult portions figs. 11 &

12). Such, he observes, is the fate of most sponges

under these circumstances, especially if they are

exposed to dampness owing to the salt-water not

having been entirely soaked out of them at the

commencement.

A. Spicular Rosettes (figs. 24.&25).—Flesh-spicula

of the rosette type obtain, and, as in the preceding,

are likewise of two forms, a small and a large kind. Diagrammmatieal representation! Ofehe pros

(a) In this the small rosette or staple form (fig. 24) able mode of arrangement of the scopuline

the figure is relatively globular, and is g{¢ inch in spicula of Steere’s sponge, after a pen-and-ink

diameter. It consists of six straight, smooth arms ‘etch by Mr. Carter. Sarcode membrane,

parting at right angles from the centre, each arm Beret ihy, pore mea ye sede

terminating in a little discoid inflation from which

spring four to eight rays spreading in what Carter

terms a “ flewr-de-lis” (“* Hexactinellidee and Lithistide,” Ann. & Mag. Nat. Hist. 1873,

vol. xii. p. 355; the spinulo-multifurcate hexradiate stellate spiculum of Bowerbank,

Monog. Brit. Spong. vol. i. p. 259, pl. vill. fig. 192). Hach ray terminates in an inflation

which is expanded into a circular convex head bordered by four spines opposite and

recurved. These rosettes (a) are chiefly congregated about the angle of the dermal

meshwork and relatively numerous.

(6) The large rosettes (fig. 25), contrariwise, are very rare, and still more globular in

appearance, whilst they range from ;g to gz inch in diameter. Their centre or, so to

say, nucleus appears to consist of six short, stout arms, each arm terminating in a qua-

drangular inflation, from which radiate several long straight rays. The rays themselves

(very rarely met with bifurcate) end in a quadrangular or more often pentangular cap or

head (fig. 26), with free convex surface, but provided with recurved spines at the angles

of the opposite surface. This kind of rosette resembles the so called “ multiradiate

spicula of the sarcode” of Askonema setubalense, Kent (Monthly Microsc. Journ. 1870,

p- 246, pl. lxiv. fig 9).

5. Sexradiate or Dermal Hexactinellid Spicula (figs. 20 and 23, and in position figs. 3

and 9).—These forming by the coalescence of the adjoining spicula the delicate veil

latticework or rectangular dermal meshwork, have already had passing allusion called

to them when speaking of the exterior characteristics. The six arms part at right angles

from the centre, the horizontal ones being 735 inch long; the vertical are shorter.

. SECOND SERIES.—ZOOLOGY, VOL. I. 24H

tween the spicular bars and quadrangular

spaces, that is, when the sponge was alive.

228 DR. J. MURIE ON STEERE’S SPONGE,

Towards the fixed end (or centre) the arms are thickest, and throughout are minutely

tuberculate or subspined; the arm extremities are smoothish and pointed. It is by the

overlapping of the horizontal arms (fig. 23, a), and this to a considerable extent, that the

squares are produced, whilst the horizontal arms respectively project outwards and inwards.

The outermost tip frequently bears the small rosette (fig. 23, 6) already described.

Besides the foregoing spicula (save the large rosette), all present in abundance, I met

with, by chance, at least in one of the slides of objects examined, another form of minute

but not measured spiculum (fig. 27). Whether this had accidentally got amongst the

débris shaken out of the Dendrospongia, or otherwise forms a constant part of the latter’s

skeleton, I cannot say positively. Possibly this spiculum in question forms but a seg-

ment of a “rosette with short arms and long, straight, pointed rays,” similar to what Carter

figures from Huplectella aspergillum (Ann. & Mag. Nat. Hist. ser. 4, vol. xii, pl. xii. fig. 3);

only one or two examples were observed by me. I have figured it rather to arrest future

attention than to insist on its being an integral part of the sponge under consideration.

REMARKS ON THE FOREGOING.—From what precedes it will readily be admitted’ that

Steere’s sponge is one among the many already singular forms of the Vitreous Sponges

of considerable interest. In size, shape, and structural detail it differs sufficiently from

others of the known Hexactinellida, as I presume, to warrant generic and specific dis-

tinction, while it no less combines certain peculiarities assimilative and explanatory, to

some extent, of moot points in the construction and history of this altogether remarkable

group. Its dendroid or shrub-like branching aspect has suggested the generic appellation

I submit; and the collector’s name, to whom I am so much indebted, I convert into its

specific title. Its habitat but adds another to the many strange and delicate marine

organisms hitherto obtained in the Philippine seas; and doubtless when hereafter studied

in its live conditions some curious traits in its history may be unfolded.

DENDROSPONGIA STEERII, gen. et sp. nov.

Hexactinellid sponge characterized by its dendritic or shrubby contour ; occasionally

attaining a height of 3 feet or possibly more. Branches forking or dichotomous, with

continuous whorled series of spicular tufts from base to apices. Skeleton only known;

basework composed of relatively stoutish glassy fibres of coalesced, sexradiate and spino-

mucronate spicula, disposed in tolerably compact trabecule. Main direction of fibre

longitudinal to axis, in parallel straightish or slightly bent lines where continued into

exterior whorls; intercrossing fibres more irregular, as are the very numerous excretory

canals. Oscula and pores of moderate size distributed all over the free surfaces. Flesh-

spicula abundant, and of scopuline, acerate, and rosette shapes. A dermal veil of

‘slender, interwoven hexactinellid spicula probably clothes the major portion or possibly

the entire sponge.

Hab. Philippines, Channel between Negros and Zebu, from 70 to 100 fathoms ?

In discussing and assigning the affinities of this choice, indeed elegant, sponge, one

naturally first looks to the kindred stock from the Eastern Archipelago. To the Huplec-

tela group, including the genus Habrodictyon (Corbitella and Heterotella, Gray), their

A NEW GENUS OF THE HEXACTINELLID SPONGIDA. 229

open tubular unbranched and widely netted structure, besides basal cushion or root of

great long anchoring spicula, sufficiently distinguish and separate them from our specimen

in question, apart from the shape of the various spicula themselves. In Deyerina the

more approximated condition of the glassy fibres composing the stem, the somewhat

spiral and therefore to a certain extent whorled nature .of outer, pronounced elevations

of the same, and marked veil or latticework suggest closer ties with Dendrospongia; but

Meyerina is unbranched, and besides has a birotulate form of flesh-spicule entirely absent

in that compared. But, moreover, the Huplectellide, spite of their looser build of vitreous

fibre, have frills bearing some relation to the said whorls, and an exceedingly delicate

veil, though their true spicular skeleton, as well as that of Meyerina, has not such rough-

ened nor spino-mucronate anastomotic fibre. In Aphrocallistes its branching character

makes a closer approach, so far as mere outward configuration is concerned; but in the

Malaccan as well as Atlantic species the widish tubular branches, relatively thin walls,

absence of exterior whorled elevations, and other differences of framework stamp sepa-

ration from Dendrospongia. The former genus notwithstanding has scopuline and flesh-

‘spicula, and, at least in A. beatriz, such echinate closely reticulate main fibre as to show

relationship with the latter, though not of the closest tie. The massive subglobose

species of Dactylocalyx from Malacca, and even the branching forms of this genus from

other regions, though possessing an anastomotic network of glassy fibre bearing a certain

resemblance to some parts of Dendrospongia, nevertheless in other particulars strikingly

depart from the latter. Such genera as the goblet-shaped Crateromorpha and Labaria,

and the “sculptor’s mallet”’-like Hyalonema, in most aspects show recedence of type

from our Zebu representative.

Again, other genera of the Atlantic and American seas, the purse-figured Holtenia,

Rossella, and nut-like Sympagella, the bird’s-nest Pheronema,and the felt or blanket sponge

Askonema,'to wit, present relationships rather with one another than in any specially

marked degree to that under consideration. To Awlodictyon and Furrea, however, there

undoubtedly seems to be a nearer affinity, if much weight is alone to be placed on the

flesh-spicular identity. Examples of neither of those two genera, so far as I am aware,

have ever been obtained in a perfect and complete form, save the diminutive 7. infundi-

buliformis, Carter ; but still sufficient is known to yield a useful comparison. Both are

‘to some extent diminutive branching Hexactinellid sponges, but even in this respect widely

different from the great, handsome, dendritic Steere’s sponge. They are, besides, hollow

-‘and less or more tubular, with very thin walls and no trace of whorled tufted branches.

In one species of Farrea the closely reticulate vitreous fibre is thickly spined (and mucro-

nate according to Carter); there also obtains scopuline, acerate?, and rosette-shaped

spicules, bearing considerable likeness to those of Dendrospongia.

With all this similarity to species of Farrea in spicular form, considerably departed from

“in other features, it seems to me that Steere’s sponge in reality presents intermediate skeletal

‘characters between Meyerina, Aphrocallistes, and Furrea, while yet so characteristic

‘from all in several respects, that it is hard even to admit positive close affiliation to one or

“other.- I expect the future may reveal some form from the eastern seas that may modify

‘the difficulties of their relationships.

Dig

230 DR. J. MURIE ON STEERE’S SPONGE,

With regard to the homology of the so-called root, body, beard, and dermal spicules of

at least the major section of the sexradiate sponges, I venture to offer a few mere

passing remarks. I apprehend that the presence of a veil or thin dermal layer of lattice-

work is far more constant among the Hexactinellida than is generally suspected, its deli-

cacy in many instances, I fear, causing its destruction in whole or part ere examination

of specimens is made. In Meyerina it is a very beautiful structure, readily apparent to

the eye even at a distance. It exists like a spider’s web on the surface of the Huplectel-

lide. A fine sheet lies upon the Askonema’s felt. Labaria and Hyalonema both possess

such latticework. Holtenia and Rossella characteristically are veiled. Even Dendro-

spongia I am inclined to believe has such a covering spread all over its whorled branches

during life. In other genera the evidence borne by fragments leads to the conclusion of

its being a structural characteristic common to all. It seems to be for the support of the

dermal sarcode in which the pores are situated.

The beard and whiskers, so termed by some authors, of such forms as Pheronema,

Rossella, Holtenia, &c., evidently are the homologues of the spicula, shorter, it is true,

composing the elegant sinuous frills of the Huplectella series. These latter frills, however

much differing in aspect, moreover correspond essentially with the whorled tufts of Den-

drospongia, and with the elevated obliquely ranged spicular vent-ridges in Meyerma. In

these and doubtless other forms they appear to be more or less modified spicules in direct

relation with the true skeletal vitreous fibre, and by or to which the finer dermal lattice-

work is attached.

The long knitting-needle-like bundle of the glass rope in Hyalonema, the anchoring

spicules so well known in Holtenia &c., and the hairy-like bunch of lengthened spicula

at the root of Huplectelia, have all strictly a homological relationship. Indeed nearly all

the Hexactinellida that have been procured in any thing like a perfect condition offer

some remnants, representing spicula specially adapted to retaining a hold of foreign

bodies to which they are rooted, or the so-termed roots are sunk in the sand or mud.

Thus, again, however unlike may he the pattern of the vitreous fibre composing the

body or basis of such forms as Dyliusia and Iphiteon, as contradistinguished from that

of Orateromorpha, Habrodictyon, Euplectella, Dendrospongia, and Hyalonema, whether

the sponge be shallow, cup-shaped, tubular, branched, or more solid, this, so to speak,

basal or body-fibre of simple, fused, or compactly built sexradiate spicula has a common

homological significance.

However much modified, then, I am inclined to think the whole of the Hexactinellid

group, besides their bond of unity in six-armed spicula, are each provided with well-deve-

loped or traces of veil and root, superadded to such glassy fibre as more properly constitutes

their basal or body skeleton.

Postscript.—It was not until some time after I had laid my paper before the Society

that two contributions on the hexactinellid sponges by Dr. William Marshall, of Weimar,

in the Zeitsch. fiir wissensch. Zool.*, came under my notice. The first published of these

* “Untersuchungen iiber Hexactinelliden,” Z. f. w. Z. 11875, vol. xxv. (Supplementheft 2), pp. 142-243, pls.

xi-xvii.; and “Ideen iiber die Verwandtschaftsverhiiltnisse der Hexactinelliden,” Z. f. w. Z. 1876, vol. xxvii. pp.

113-136,

A NEW GENUS OF THE HEXACTINELLID SPONGIDA. 231

bore a special interest to me, as containing a photograph of a new genus and species of

sponge—Sclerothamnus Clausii, which I perceived had close affinity with Steere’s sponge.

Their identity at the first glance I certainly did not appreciate, possibly from his evi-

dently being a much smaller imperfect specimen, and also doubtless owing to the blurred

ill-defined photographic representation of it. A further careful study of his descriptions

and close comparison of figures has, however, convinced me that in the said Sclerotham-

nus and Dendrospongia, without a doubt, we have the same generic type, and as regards

specific distinction between them I am really at a loss to point out differences. 4 for-

tiori, I regard them as identical; and therefore Dr. Marshall’s name to this remarkable

form has the claim of priority, and my own will hereafter necessarily be sunk to that of a

synonym. Hence Sclerothamnus Clausii, Marshall,=Dendrospongia Steerii, Murie. The

examples on which Marshall founds his description and diagnosis are two odd pieces

got in an old cabinet of the Leiden Museum, locality and history unknown. My data,

therefore, clears habitat, and in many other respects supplies points wanting in his com-

munication. Dr. Marshal], notwithstanding, has observed and figured the ‘“‘ mucronate”

character of the glassy fibres, on which Mr. Carter has laid some stress, and he moreover

calls attention to certain phenomena connected with the growth and coalescence of the

_ glassy spicular fibre, besides what he regards as monstrous spicula.

In his ideas upon the relations of the Hexactinellida he constitutes a division SYNAv-

Loip#=— Das Lumen der Réhren der verschiedenen Nadeln hangt, wie diese selbst,

continuirlich miteinander zusammen, so dass das ganze Zittergewebe des Schwammes von

einen gleichfalls zusammenhingenden Rohrsystem durchzogen ist ”—in which the only

generic type is Sclerothamnus, whose characters are :—“ Zu den zusammenhaingenden

Zittern gesellen sich als frei bleibende Nadelformen Besengabeln.” ‘'The single species

as yet known, Sel. Clausii, according to him, has the following diagnosis :—“ Polyzoisch,

buschférmig, mit dichotomisch verzweigten, nicht in einer Ebene gelegenen Aesten. Zwei

Formen von Besengabeln. Zitterwork mit Hockerchen besetzt.”

Into his second division AsyNAULOID# and its subdivisions, and with his general con-

clusions and remarks on the relationships &c. of the Hexactinellida, neither space nor

other circumstances permit me herein to enter.

Apprnpix. By H. J. Carrer, F.RB.S.

With Dr. Murie’s description of Dendrospongia Steerii I should hesitate to interfere, even if I were not

well acquainted with the general and microscopic structure of the more durable parts of the sponge itself,

such is the descriptive power and truthfulness of the author. :

Professor Steere’s specimen is certainly the largest and noblest mstance of the Hexactinellida on record,

and from its branched form contributes to show that the Hexactinellida are subject to the same influences

in this respect as those which determine the variation in form and size of all other sponges, which appears,

to a certain extent, to be almost unlimited.

It is not a little remarkable, too, that a similar specimen, only not half so large (16 inches high), should

have been deseribed and figured by Dr. William Marshall, of Wiemar (Zeitschrift f. wiss. Zool. xxv.

Suppl., 20th November, 1875), just two months before Dr. Murie brought Prof. Steere’s specimen to the

notice of the Linnean Society. Still more remarkable is it that just nineteen years before this, and before

the same society (viz. 17th February, 1857), large fragments of this species should have been uncon-

232 DR. J. MURIE ON STEERE’S SPONGE,

sciously brought, and subsequently figured among the root-detritus of Prof. Owen’s Euplectella cucumer,

still where it was then, viz. in the possession of Dr. Arthur Farre (Trans. Linn. Soc. vol. xxu. pl. xxi.

fig. 1), with a microscopic portion, highly magnified im fig. 8, which had been “discovered by Dr. Farre”

himself, and thus had been thought worth figuring independently of the Huplectella—showing, as the

sequel will prove, that the peculiarities of the structure had not passed unnoticed, although the sponge

itself had not been discovered.

Lastly, it is worthy of remark that Dendrospongia appears to exist in the neighbourhood of the Euplec-

tellas both at the Seychelles and among the Philippine Islands.

Besides the fragments of Dendrospongia Dr. Farre also discovered another structure, which Prof, Owen

likened to a “harrow” (l.c. figs. 9,9 a). After this Dr. Bowerbank obtained permission of Dr. Farre to

examine the enéire specimen of Euplectella cucumer, and (among other fragments) brought away with him

specimens respectively of those discovered by Dr. Farre, to which I have above alluded. These Dr.

Bowerbank examined, described, and figured respectively, under the general name of Furrea occa (Proc.

Zool. Soc. 1869, pl. xxiv. figs. 1 and 7), conceiving the “ harrow-like tissue” to have belonged to “the

dermis,” and the other structure to the body of this sponge.

We know, however, now, from enéire specimens of the sponges represented by these two structures,

that they belong to two distinct species, and that the ‘“harrow-like tissue” is the dody-structure of

Farrea occa, and not identical with the dermal layer of the Hexactinellida, probably existing (according

to Dr. J. Millar, who first directed especial attention to it) throughout all the species, although often,

from its delicate structure, more or less rubbed off.

In 1873, just after having, by the kind assistauce of Dr. Farre, examined his justly celebrated specimen

of Huplectella cucumer myself, I also brought away specimens of the structures figured both by Professor

Owen and Dr. Bowerbank (J. c.), and, having examined them microscopically, came to the conclusion that

the “ harrow-like tissue” should remain as Farrea occa, and the other structure be designated Farrea

densa, as belonging undoubtedly to a different sponge (Ann. & Mag. Nat. Hist. 1873, vol. xii. p. 454,

pl. xv. fig. 5). Hence a fragment of the latter with its peculiar spines was figured (J. c. fig. 6), mention-

ing also (p. 463) that among the spicules boiled off from this fibre was a scopuline one like that .of

Aphrocallistes beatriz (which is nearly identical with that of Dendrospongia) “in great abundance.”

As soon, therefore, as Dr. Murie, through Dr. J. Millar, kindly put me in possession of an actualfragment

of Dendrospongia, I saw that the body-fibre was identical with that of Warrea densa, and that the scopulme

spicules were also identical, but I had observed no rosette. However, on turning to a mounted specimen

which had not been boiled in nitric acid, I found not only the rosette but the peculiar head of one of the

scopuline spicules also present with the body-fibre. It may be here remarked, respecting the rosette, that

Dr. Marshall states (Zeitschrift, /. c. p. 185) “ bei Sclerothamnus und Aphroeallistes findet sich keine

analoge Nadelform.” Thus we have traced Dendrospongia back to the Linnean Transactions of 1857.

Whether Farrea densa, Sclerothamnus Clausii, or Dendrospongia Steerii is to take precedence in nomen-

clature I care not. Dr. Murie’s description and illustrations take precedence with me as to accuracy and

fullness of detail. This lasts for ever ; but a name may be disputed to the end of time.

Dendrospongia Steerii evidently belongs to that group of sponges which possesses for one of its pecu-

‘liarities the scopuline spicule, which must not be confounded with the rosette that, so far as my obser-

vation extends, always accompanies it. The scopuline spicule is often taken into the skeleton-fibre, and

therefore may be regarded as a subskeléton spicule ; while the rosette, bemg essentially a flesh-spicule

(that is, confined to the sarcode) is seldom or never included in the vitreous fibre. Hence, if the skeleton-

fibre alone remains, the rosette (as was probably the casein Dr. Marshall’s specrmen) may have fallen

through it on the decomposition of the sareode, since, as I have before stated, when this takes place the

rosette passes out through the skeleton almost as easily as small pebbles through the meshes of a fishing-

net. In such instances, therefore, the absence of the rosette may not indicate that’ there never was any.

The rosette, again, often presents itself under a variety of forms in each species of the Hexactinellida,

as described and figured by Dr. Murie in Dendrospongia Steerit.

Fig. 9.

A NEW GENUS OF THE HEXACTINELLID SPONGIDA. 233

DESCRIPTION OF THE PLATES.

Prata XXXVI.

. Steere’s Sponge (Dendrospongia Steerii). Drawn on stone from a photograph, aided by a piece

of the specimen itself, and reduced ? nat. size. In accordance with the text the lettering

used signifies:—ro, root; s, stem; 7, right, m, middle, and /, left main branches; 0, secondary

branch; f, fork.

. Portion of a terminally forking branch, of the natural dimension. w, whorl or spiral tuft; or,

interlocking branch of same; g, intervening groove or spiral depression between the tufts.

. Asmall segment of a branch of Dendrospongia at one of the spiral grooves. Enlarged about

twice nat. size. This shows the appearance of the veil or delicate latticework of square

meshed spicules, spread over the surface and partially extending to the raised whorls; openings

of the oscula or vents are seen beneath the veil.

. Fragment of the interior of a branch, illustrating mode of skeletal arrangement of the large glassy

fibres. Enlarged about twice nat. size.

PuaTe XXXVII.

. Sketch of the foreshortened summit of a branch, exhibiting termination of spines &c. Of

nat. size.

. A transverse section (somewhat enlarged) of the stem at the base of one of the branches, and where

the ruff or whorled spicula have been considerably rubbed away. The central framework has

many irregularly contoured canals and of different sizes; towards and at the peripheral margin

the glassy fibres appear to radiate, the most marked of which constitute what remains of

the abraded whorls.

. Another cross-section made higher up, and where branching occurs; hence its more compressed

oval form, also in part due to the exterior spicula being worn off. Nat. size.

. A median longitudinal section of part of arubbed branch of D. Steeriz, and where only indications

of the spicular whorls are present. Enlarged about once and a half nat. size. This illustrates

the exact disposition of the internal main body of the glassy fibres or basal structure of the sponge..

The long, large, strong, and vertically directed bundles of spicula incline outwards or with

slightly sinuous curve follow a certain spiral arrangement, the marginal elevations cor-

responding to the whorls and the depressions to the grooves. A network of short cross or

oblique spicula bind together the axial and longitudinally directed stouter sorts. Canals and

oscula permeate freely.

A greatly magnified representation of a portion of the framework or glassy fibre as seen on its

superficial aspect. It exhibits in position the so-called veil-layer or lattice-lke squares of the

dermal hexactinellid spicula (see fig. 28, a, 6), also here and there deep-seated hazy patches of |

the flesh-spicula &c.

Fig. 10. Another portion of the glassy fibre, from the interior, still further enlarged, the dotting showing

its spmy roughening. At a one of the larger rectangular interspaces between the reticulated

vitreous spined fibre is partly shown; 4 is within a similar small circular area; ¢ indicates a

thin spined spicule, so bent and united at tips as to form a complete circle; springing from its

middle is a still more delicate needle-shaped spicule (d), which lies across ¢; below is a mass

“ of the common fibre variously agglutinated.

ON A NEW GENUS OF HEXACTINELLID SPONGIDA.

. Another piece of the vitreous fibre, wherein a, b, two main longitudinal branches, are seen, by

degrees, to approach, and further on, at c, to unite, and ultimately, by coalescence, to con-

stitute but a single thickened fibre. The areas between the glassy fibre are charged with loosely

arranged scopuline and other flesh-spicula. Specimen also very considerably enlarged.

. A couple of what may be compared to sprays of the terminal filaments of the stout glassy fibre,

greatly magnified. These, though compound branched, are characteristically pitchfork in

appearance, the free tips towards the right (corresponding in part to what is shown in fig. 9)

being those that, so to say, protrude through the veil latticework, and hold it in place even in

the dried condition of the skeleton. In some of the terminal single fibres the sexradiate con-

dition is marked, but the axial fibre, nevertheless, ends in a tapering point. Minute dermal

and flesh-spicula abound in this specimen.

. One of the longer scopuline flesh-spicules whose shaft is microspined.

Fig. 14.

Another spicule of the same kind, with secondary or spinal arms, set at right angles to the long

axis of the shaft.

. Scopuline spicule, with microspined shaft and head, bearing expanded arms.

. Variety of the last, with secondary or spinal arms.

. One of the large (subskelatal) acerate spimed spicula.

. Portion of same, still further enlarged, to demonstrate arrangement of the spines on its shaft.

. A group of the smaller-sized acerate flesh-spicula.

. A hexactinellid spiculum, with short axial arms, all the arms being slightly roughened or

microspined.

. Sketch of the pomt of junction of a sexradiate spicule, the four arms of which persist, but only

a rudiment of one of the axial arms 1s apparent in a central knob.

. A couple of the circumferential younger sexvadiate glassy fibre, the arms being unequal in

length, the longer ones tending to curvature.

. A few of the dermal hexactinellid spicula, 7. e. those composing the lattice-lke veil. At a the

mode of union of the squares is displayed, and at 6 a single sexradiate spicule is shown, with

a rosette in position tipping the free extremity of the axial arm.

4. A highly magnified representation in perspective of one of the small rosettes of the fleshy,

dermal, rectangular meshwork. (See fig. 236 for its position with respect to the free axial

arm of the sexradiate spicule.)

. The larger kind of multiradiate rosette, sparsely met with among the débris of the sponge.

. A foreshortened view, or from above, of the cap or head of one of the rays of the rosette, fig. 25.

. A spicule of a minute kind, of which only one or two were observed among the loose substance

shaken off the Dendrospongia.

All the figures of the spicula, from 18 to 27 inclusive, are enormously enlarged ; and the relative propor-

tions as to size of the different sorts have not been strictly adhered to, though in the main approximated.

For their accurate measurements &c. consult description in body of text.

Berjeau & Iterson lith,

ENDROSPONGIA STI

levine! lit

Trans Linn. Soc:Si

ER.Z.Z00L Vou.l Tas 36.

M& N.Hanhart imp.

Trans.Linn. Soc. Ser.2.Zoou Vou lTas 37

Berjeau lith. Hanhart imp

SKELETAL DETAILS OF DENDROSPONGIA.

[ 235 J

XII. Notes upon the Oyxstomatous Crustacea. By Epwarp J. Mizrs, 7.L.8., £.Z.8.,

Assistant in the Zoological Department, British Museum.

(Plates XX XVITI.—XL.)

Read June 15th, 1876.

I. Family LEUCOSIID A.

Remarks on the Literature of the Subject.

SINCE 1855, when Mr. Bell published, in the 21st volume of the Society’s ‘ Trans-

actions,’ his excellent monograph of the Leucosiide, several important memoirs have

appeared, adding considerably to our knowledge of the genera and species of this

family, which includes some of the most striking and beautiful forms in the whole range

of the Brachyura. In the ‘ Proceedings of the Academy of Natural Sciences of

Philadelphia’ for 1858 Mr. W. Stimpson published diagnoses of several new genera

and species of Zeucosiide, collected during the United States’ Expedition to the

N. Pacific, chiefly in the Chinese seas. In the ‘ Journal of the Boston Society of

Natural History,’ vi. 1857, the ‘ Annals of the Lyceum of Natural History of New York’

for 1860 and 1871, and in the ‘ Bulletin of the Museum of Comparative Zoology,’ ii.

1870, the above-mentioned author has described a considerable number of new genera

and species from different parts of the North-American coast. M. Alph. Milne-Edwards,

in the ‘ Annales de la Société Entomologique de France,’ v. 1865, the ‘ Nouvelles Archives

du Muséum d’Histoire Naturelle,’ x. 1874, and the ‘ Journal des Museum Godeffroy,’

part iv. 1873, has described and beautifully figured many new and interesting genera

and species from different parts of the world. Species of this family have also been

described or noticed by Gibbes, Herbst, Hess, Kinahan, and others.

In the present essay the species of this family in the collection of the British Museum

which do not appear to have been hitherto recorded, are described; and remarks are

added upon a few other species in the national collection.

Descriptions of New Species of the Family of Leucosiidee.

Leucosta, Fabricius.

The species of this, the typical genus of the family, are remarkable in many instances

for the beauty of the coloration and markings of the polished carapace. ‘They are found

throughout the tropical and warmer temperate zones of the oriental region, from the Red

Sea to the islands of the Pacific.

Twenty species are enumerated by Bell in his monograph of the family. Stimpson, in

1858, added three species from the Chinese seas to the list. A species from the

Australian coast was characterized in 1865 by Hess; M. Alph. Milne-Edwards in 1874

described and figured three species from New Caledonia; and in 1875 I described one

_ SECOND SERIES.—ZOOLOGY, VOL. I. 21

236 MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

from the Samoa Islands. There are in the national collection eight species of this

genus, apparently distinct from any that have been hitherto recorded.

1. LevcosIA FUSCOMACULATA, sp, n. (Pl. XX XVIII. figs. 1-3.) Allied to ZL. wrania,

Herbst, but less convex, with a more distinct, marginal beaded line; the carapace

distinctly punctated on the sides; anterior part of the thoracic sinus much

narrower, and its cavity, above the point of insertion of the thoracic anterior legs,

filled by a confluent series of but three or four closely placed, flattened tubercles,

whereas in L. urania the tubercles are at least twice as numerous, and remote from

one another. Abdomen of male smooth, 4-jointed. Colour light, yellowish brown,

with very indistinct spots and patches of dusky grey on the carapace. Length % in.;

breadth 2 in.

Hab. Wed Sea, Gulf of Suez.

2. LevucosIA PULCHERRIMA, sp. n. (Pl. XXXVIII. figs. 4-6.) Carapace smooth,

convex, sparsely punctate towards the front and anterolateral margins, which are

defined by a beaded line which terminates above the point of insertion of the first

pair of ambulatory legs. Hepatic regions concave in their front portion, and

slightly convex behind. Arm with a line of large tubercles on its anterior and

posterior margins, upper surface with a group of small tubercles at base, followed

by four larger; lower surface with many closely placed tubercles at base; wrist .

and hand smooth, fingers not quite meeting at base when closed. Thoracic sinus

deep, narrow, and well-defined in front, with a beaded margin; its cavity, above

the base of the anterior legs, filled with a series of about half-a-dozen small

tubercles. Colour stone-grey, inclining to brown on the back and white on the

ambulatory legs, a circle of rose-pink upon the back of each branchial region, a

trilobate pattern formed by three confluent semicircles of the same colour upon each

side of the anterior half of the carapace (these are confluent in the middle line, and

together somewhat resemble in outline a butterfly with the wings extended); there

are three white spots in the centre of each trilobate pattern; the basal half of the

fingers is orange; and there are rings of the same colour surrounding the joints of the

ambulatory legs and tubercles of the arms, also small spots of orange near the

lateral margins and upon the inferior surface of the carapace. Length 1), in.;

breadth +4 in.

Hab. Lizard Island. —

In all structural characteristics this species is nearly allied to ZL. neocaledonica,

A. M.-Edw.; but it is preeminently distinguished from this and all other species of the

genus by the strikingly beautiful coloration and markings of the carapace. The thoracic

sinus is also deeper, narrower, and better-defined in front, with a beaded margin. Only

a single specimen (2) in the collection.

3. LEUCOSIA ORNATA, sp. n. (Plate XXXVIII. figs. 7-9.) This species, in all struc-

tural characteristics, very closely resembles L. pulcherrima; but it is more strongly

MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA. 237

punctated, the concavity and convexity of the hepatic region is more decided, and

there are but four clearly distinct tubercles in the cavity of the thoracic sinus. The

colour is dark brown; there are three white spots upon each side of the gastric

region, and a small red circle on the back of each branchial region, The bases

of the fingers, tubercles of the arms, and joints of the ambulatory legs are banded

with orange, as in many species of the genus. Length 1 in.; breadth % in.

Hab. Ceylon,

This species was presented to the Trustees of the British Museum by H. W. H. Holds-

worth, Esq., with a valuable series of Crustacea from Ceylon. The specimen is a female.

4, LEUCOSIA BRUNNEA, sp.n. (Plate XX XVIII. figs.10-12.) Carapace convex, smooth

and polished, but very slightly punctated towards the anterolateral margins, and

with a finely beaded line continued to the middle of the posterolateral margins.

Front very slightly projecting beyond the eyes; behind it there is a short longi-

tudinal median ridge between the concave hepatic regions, Thoracic sinus wide, its

anterior margin straight, not beaded, and forming nearly a right angle with the

anterolateral margin of the carapace. Arm with numerous, small, crowded tubercles

upon its infero-anterior surface, upper surface with a group of very small tubercles at

base, beyond them four tubercles placed in a quadrangle; anterior and posterior

margins with a line of tubercles; wrist smooth; inner margin of hand with

a granulated line; fingers straight. Colour brown, inclining to red on the back of

the carapace. Length 13 in.; breadth 1 in.

Hab. Singapore.

This species is distinguished from its congeners by the form of the thoracic sinus &c.

5. LeUCOsIA RETICULATA, sp. n. (Pl. XX XVIII. figs. 18-15.) Carapace subglobose,

smooth ; sides with a beaded line extending from the front of the hepatic regions

along the anterolateral and about halfway along the posterolateral margins. Front

flat above, and not produced beyond the eyes; anterior margin straight. Thoracic

sinus short; anterior margin smooth, not beaded, and not defined at either ex-

tremity. The cavity of the thoracic sinus, above the insertion of the anterior legs,

is filled by a series of about half-a-dozen flattened tubercles. Anterior legs strong;

arm tubercular on the proximal half of its upper surface, the distal half smooth and

polished; on the anterior and posterior margins the tubercles extend nearly to the

extremity of the arm; on the anterior surface, and below at base, the tubercles are

numerous and crowded; wrist smooth, hand compressed, outer margin smooth,

inner with an obscurely beaded line. Abdomen of male smooth, without any

tubercle on the penultimate joint; terminal joint very small, the others of about

equal width throughout. Colour greyish green, yellowish on the sides and back of

the carapace, which is marked with reticulating lines ofa darker green. Ambulatory

legs yellowish brown. On each side of the carapace, above the point of insertion of

the anterior legs, there isa group of from six to a dozen small red granules. Length

11 in.; breadth 1 in. i

212

238 MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

Hab. Western Australia, Shark’s Bay.

This species is at once distinguished by its colour and the group of red granules on

each side of the carapace. Specimens of both sexes are in the collection.

6. Lrvcosta WHITMEE! (Plate XX XVIII. figs. 16-18.)

Leucosia whitmeei, Miers, Ann. & Mag. Nat. Hist. xvi. p. 342 (1875).

This species was described by me in 1875 from a specimen from Samoa, presented by

the Rey. 8. J. Whitmee to the Trustees of the British Museum. Specimens are in the

national collection, from Ovalau in the Fiji Islands, which probably belong to this

species, with which they agree in structural characteristics ; but all traces of the original

coloration and markings of the carapace have disappeared.

7. Leucosia PERRY, sp.n. (Plate XX XVIII. figs. 19-21.) Carapace rather depressed,

with a continuous beaded line defining the lateral and posterior margins, and with

a slight longitudinal median ridge between the hepatic regions. Front tridentate,

not concave above. Thoracic sinus entirely absent. Anterior legs rather slender ;

arm closely tuberculated above and below at and towards the base; wrist and hand

smooth, hand rather compressed, but not keeled or beaded; fingers, when closed,

meeting only at tips. Abdomen of male narrow, triangular, with the sides straight,

and witha slight tubercular prominence on the penultimate segment. Colour of

carapace deep brown, with two or three small, irregular, paler spots in front of the

branchial regions ; arms greyish; hands and ambulatory legs paler brown. Length

3 in.; breadth about 4 in.

Hab. Western Australia, Shark’s Bay.

This species is distinguished from all others of the genus having a tridentate front by

the entire absence of the thoracic sinus. In this respect it resembles L. orbicularis, Bell ;

but this species has the front much less produced forward, with the anterior margin

concave.. I have named it in honour of its discoverer, W. Wykeham Perry, Esq., by

whom it was presented to the Trustees of the British Museum, with three others of the

new species described in this paper *.

8. LEUCOSIA PUBESCENS, sp. n. (Plate XXXVIII. figs. 22-24.) Carapace convex,

smooth and polished; anterolateral margins with a finely beaded line, which does

not extend beyond the commencement of the posterolateral margin. Hepatic

regions concave. Front tridentate, flat above. Thoracic sinus with its anterior

* The following is a list of the species of Crustacea (besides several interesting specimens too young for deter-

mination) that have been presented by Mr. Perry to the collection of the British Museum :—

Lambrus nodosus. Geograpsus crunipes. Matuta obtusifrons, sp. 0.

Neptunus sanguinolentus. Sesarma obesum. Calappa hepatiea.

Caphyra rotundifrons (young). S. quadratum. Renuiipes pacificus,

Gelasimus annulrpes, var. Leucosia reticulata, sp. D. Squilla armata,

G, gaimardi. L, perryi, sp. .

Grapsus pictus. Matuta crebrepunctata, sp. D.

MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA. 239

margin smooth, not beaded, and not defined at either extremity, its cavity above

the base of the anterior legs filled with a short close pubescence. Upper surface of

arm with a group of small tubercles at base, followed by two larger, and with a

patch of dense short pubescence. Anterior and posterior margins with a line of

tubercles; lower surface densely tuberculated; wrist and hand smooth, fingers

straight. Colour greenish grey, with a series of eight marginal spots of darker

green, two spots of the same colour on the gastric region, and four on the cardiac

region. There are also two white spots on each side of the gastric region. Ab-

domen of male smooth, with all the joints except the last coalescent. Length nearly

1 in.; breadth 3 in.

Hab. Western Australia, Shark’s Bay.

This species is most nearly allied to Z. pallida, Bell, but differs from it in the form of

the thoracic sinus, which is wider in front; the arm of L. pallida has not the patch of

pubescence on its upper surface at base that always exists in specimens of L. pubescens ;

the hand is slenderer, with a granulated line on its inner margin.

Myra, Leach.

Myra mMAmMitiARts, Bell. (Plate XXXVIII. figs. 25-27.) Some specimens from

Australia, in the British-Museum collection, differ in several particulars from the

adult specimens of IZ. mammillaris, but are, I think, the young of that species. In

these examples the carapace is scarcely longer than broad, and there is a faintly

marked longitudinal median carina. The hepatic regions are more strongly convex,

the pterygostomian regions more decidedly angulated ; the tubercles on the posterior

margin of the carapace are more conical (though not acute), the anterior legs are

much shorter, and the tubercle on the penultimate segment of the abdomen of the

male much more prominent than in adult specimens of WM. mammillaris. Length -

inch; breadth 2 inch.

It is probable, if this determination be correct, that both If. carinata and W. elegans,

Bell, are founded on specimens that have not attained quite their full size, and that the

median carina, characteristic of these species, will be found to disappear entirely in fully

matured animals. Should, however, the examination of a larger series of specimens

prove the examples above described to be specifically distinct from IL mammillaris, they

may be designated by the name of WZ. angulata.

Noursta, Leach.

1. NuRsIA stNuATA, sp.n. Carapace moderately convex, broader than long, with three

tubercles placed on the cardiac region in the form of a triangle, a longitudinal

median ridge extending from the front to the posterior margin, and a transverse

curved ridge that extends from the cardiac region to the second tooth of the pos-

terolateral margin. There is also a slight elevation in front of each anterior tubercle

of the cardiac region. There are well-marked depressions between the ridges

towards the margins. Posterolateral and posterior margins deeply sinuated, with

240 MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

seven broad, triangular, nearly equal teeth. Anterior legs slender; arm granu-

lous and rounded on its anterior and inferior and the proximal half of its posterior

margins ; wrist and hand smooth; hand slightly compressed, but scarcely carinated.

Hab. Australia, Moreton Bay.

In both NV. plicata and N. abbreviata the anterior legs are smooth, the arm trigonous,

and the hand angulated and carinated. In N. plicata the teeth of the posterolateral

margins are unequal; in IV. abbreviata they are nearly ol)solete.

2. Nursta pLicaTa? (Plate XXXVIII. fig. 28.)

? Cancer plicatus, Herbst, Naturg. Krabben u. Krebse, iii. pl. lix. fig. 2 (1803).

Nursia hardwickii, Leach, Zool. Miscell. i. p. 20 (1817); M.-Edw. Hist. Nat. Crust. ii. p. 137 (1887).

Nursia plicata, Bell, Trans. Linn. Soc. xxi. p. 307 (1855), no figure,

Hab. Indian Ocean.

Mr. Bell observes of this species, that the only specimens with which he was acquainted

(those in the British-Museum collection) were brought from India by General Hard-

wicke. He evidently was unaware of the true habitat of the specimens of NV. plicata,

and confounded them with NV. hardwickii. These specimens were in the collection

in 1852, three years prior to the date of the publication of Mr. Bell’s essay; and one of

them is undoubtedly figured by him as W. plicatus (pl. xxxiv. fig. 4). In the British-

Museum copy of Herbst’s work the description and plate containing the figure of his

Cancer plicatus are unfortunately wanting; and therefore I cannot tell to which of the

undoubtedly distinct species, NV. sinuata and N. hardwickii, it is to be referred. JV.

hardwickii of all ages is distinguished from the WN. sinwata by the anterior legs, which

are flat, smooth on the upper surface, with the anterior and posterior margins of the

arms thin and sharp-edged, not rounded, as in NV. sinuata,

The specimen I have figured is the typical example of NV. hardwickii.

ARcANTA, Leach.

ARCANIA GRANULOSA, sp.n. (Plate XX XVIII. fig. 29.) Carapace shining, subglobose,

uniformly and distinctly granulated, with ten short, acute, equidistant, smooth

marginal spines—and one on the intestinal region, above the two posterior marginal

spines. The frontal spines short, projecting but slightly over the eyes. Anterior

legs slender, about twice as long as the carapace; arm distinctly, wrist and hand

very finely granulated. Length nearly 4 in.; breadth + inch.

Hab. Australia, Moreton Bay.

This species is distinguished by the granulated carapace (which is not at all tubercu-

late or spinulose on its upper surface) from the other species of the genus.

CRYPTOCNEMUS, Stimpson.

This genus was established by Stimpson in 1858 for a Crustacean discovered in Japan,

and remarkable for the extension of the carapace laterally over the ambulatory legs, to

which he applied the specific name of C. pentagonus. A second species, C. grandidierii,

was described by M. A. Milne-Edwards in 1865, from Zanzibar.

MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA. 241

CRYPTOCNEMUS HOLDSWORTHI, sp.n. (Plate XX XVIII. figs. 30-32.) Carapace nearly

transversely oblong, with the frontal region projecting; the lateral margins nearly

as long as the anterior margins, and forming nearly a right angle with the straight,

entire, posterior margin; smooth, without granules or tubercles, convex in the

centre, and slightly concave towards the margins. There is a longitudinal median

ridge between the front and cardiac region, and a less distinct, oblique ridge between

the cardiac region and posterolateral angles of the carapace. Anterior legs (in

female) smooth, with the arm trigonous, hand slightly compressed and cristate

above, the margins straight, not sinuated; fingers nearly as long as palm, meeting

throughout when closed. Ambulatory legs small, and entirely concealed beneath

the carapace. Abdomen of female broad, punctulated, 4-jomted; the terminal

joints coalescent. Length } in.; breadth 3 in.

Hab. Ceylon.

This species is named after its discoverer, H. W. H. Holdsworth, Esq. In C. gran-

didieri, A. M.-Edw., from Zanzibar, the front is broad and truncate, the elevated regions

of the carapace marked with granulated lines, and its posterior border emarginate. In

C. pentagonus, Stim., from Japan, the carapace is differently proportioned, the entire

lower surface of the body smooth, and the fingers only one third the length of the hand,

which has greatly expanded horizontal crests and deeply sinuated margins.

If. Family MATUTID A.

Historical Summary.—Although the remarkable crustaceans of the Oxystomatous

genus Matuta are among the commonest of the species of Brachyura inhabiting the

Oriental region, the differences in the armature and coloration of the carapace and anterior

legs are so slight and so numerous that their determination is a matter of great difficulty ;

and most of the later naturalists have been content to admit one, or at most but two

species, distinguished mainly by the coloration of the carapace. I have, after careful

examination of the large series of specimens in the British Museum, arrived at the con-

clusion that there are at least nine well-marked species in the national collection.

In the following condensed summary of the history of the genus, the principal authors

are mentioned who have contributed to our knowledge of the classification of the species.

Fabricius, in the Supplement to his ‘ Entomologia Systematica’ (1798), established the

genus Matuta for his previously described Cancer victor, and added another species,

M. planipes, which he characterizes simply as “thorace postice striato.’ It is impos-

sible, from this brief diagnosis, to identify the species with any of those described below

with any degree of certainty.

In 1817, Dr. Leach, in the third volume of his ‘ Zoological Miscellany,’ p. 12, pub-

lished an article on the ‘‘ Characters of the Genus Matuta, with Descriptions of the

Species.” In this essay four species are described, and two figured ; the Matuta vicior

of Fabricius is referred to, but not described; no attempt is made by Dr. Leach to

identify with his own the species of Herbst, Desmarest, Latreille, and earlier authors.

The characters employed in the definition of the species are unfortunately such as vary

249 MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

with the age and sex of the specimen ; and (in the case of I. peronii and WZ. lunaris)

examples of widely separated species are brought together, and exist in the nationa

collection under the same specific name. In the case of these species I have therefore

considered the specimen figured by Dr, Leach the type of the species. Of IZ. banksia

no labelled specimens are now in the collection; and I have been guided by the descrip-

tion alone.

In 1825, M. Latreille, in the tenth volume of the ‘Encyclopédie Méthodique d’ His-

toire Naturelle,’ published an excellent article upon the genus Matuta, in which he indi-

cates, for the first time, what I believe to be the natural subdivisions of the genus, based

upon the sculpture of the external surface of the hand. He describes a new species,

M. doryphora, and in a footnote points out distinctive sexual characters. His essay is

not noticed by M. Milne-Edwards in ‘ L’Histoire naturelle des Crustacés.’ This natu-

ralist, in his second volume of the work in question (1837), admits but two species of

this genus, WZ. victor and WV. lunaris, Herbst, characterized respectively by the carapace

being marked with numerous scattered dots or reticulating lines.

De Haan, in the fifth fasciculus of his elaborate work on the Crustacea of the ‘ Fauna

Japonica’ of Von Siebold (1841), reunited under one name, Matuta victor, all the species

of the genus, but distinguished six varieties, to which no original names are ‘applied —

although he quotes as synonyma the names of several of the species of the earlier

authors, as referring to them, whether rightly I am not in every case able to determine.

Dana, in 1852, in the first part of the ‘Crustacea of the United-States Exploring Expe-

dition,’ xiii. p. 395-6, notes the sexual differences in the striation of the ridge on the

mobile finger in the specimens collected by the Expedition, but adds nothing to our

knowledge of the species.

Hess (1865), in the ‘Archiv f. Naturgeschichte,’ xxxi. p. 158, pl. vi. f. 18, described

and figured a new species, MZ. picta, from Sydney. I. Hilgendorf, in the ‘ Reisen in

Ost-Afrika’ of Baron van der Decken (1869), figures details of a species he refers, I

believe, correctly to JZ. victor (op. cit. Crustaceen, p. 93, pl. iii. fig. 2), and adds remarks

upon the striated areas upon the inner surface of the hand (which, he thinks, serve the

purpose of producing sound), and upon the specific distinctness of I. victor and I.

lunaris.

Lastly, in 1874, M. Alphonse Milne-Edwards, in the ‘Nouvelles Archives du Mu-

séum,’ x. p. 54, reunites, under the name of I. victor, all the species of earlier authors,

remarking that it is impossible to admit the numerous species proposed by different

authors, every intermediate degree being found between forms which at first might

appear distinct.

I have endeavoured to retain the names of the early authors for the species hereafter

described, wherever possible; but their identification is often a matter of considerable

difficulty, owing to the insufficiency of the descriptions and the inaccuracy of the figures.

The species of the genus Watuta are distributed throughout the whole Oriental region ;

but I am not aware that any have been recorded from any other of the great geogra-

phical areas.

MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA. 243

A Revision of the Species of the Genus Matuta.

Marotta, Fabricius.

In the species of this genus the carapace is usually slightly broader than long, broadest

and with a strong, conical, acute, lateral spine at the junction of the anterolateral and

posterolateral margins. The front is about equal to the orbit in width, slightly con-

cave above, with a projecting median lobe, which in some species is notched, in others

entire. The anterolateral margins are rather convex, with about six small tubercles

behind the external orbital angle, and following them usually three granulated, larger

tubercles. The posterolateral margin is straight, with a tubercle, that is sometimes

obsolete, placed at some distance behind the lateral line. In the central region of the -

carapace are six tubercles, of which some, or all, may be obsolete; three of these are

placed in a median transverse series, one anterior on each side of the gastric, and one

posterior in the centre of the cardiac region. The anterior legs are robust, and may be

closely applied to the body; the arm is smooth, the wrist slightly tuberculated upon its

external surface; the upper margin of the hand has three prominent, acute lobes or

spines; on the upper part of the external surface are three transverse ridges, or two

parallel series of three tubercles; below these, on the outer surface, a ridge, or series of

tubercles and spines, the direction and form of which varies in the different species of

the genus; on the inner surface two small striated areas. The external surface of the

mobile finger in the male is usually transversely striated or beaded. The four following.

pairs of legs have the two terminal joints more or less expanded and compressed; the

expansion of the terminal joint is greatest in the first and fourth, and least in the third

pair of legs; and this joint is acute at the extremity in the first three, and rounded in

the last pair of legs.

(A) Hand of male with an interrupted ridge or series of spines and tubercles crossing its outer surface

obliquely, and continued along the outer surface of the lower (immobile) finger. Upper (mobile) finger

with a strongly beaded ridge extending throughout its whole length.

(a) Carapace covered with numerous dark-red spots.

1. Marura vicrrix. (Plate XX XIX. figs. 1-3.) site a Berk]

Cancer victor, Fabr. Spec. Ins. 1. Append. p. 502 (1781) ; Ent. Syst. ii. p. 449 (1798)?

Matuta victor, Faby. Ent. Syst. Suppl. p. 369 (1798) ; ? M.-Edw. Hist. Nat. Crust. ii. p. 115, pl. xx.

figs. 3-6 (1837) ; Régne Animal de Cuvier, Atlas, Crustacés, pl. vii. fig. 1.

Matuta peronii, Leach, Zool. Miscell. m1. p. 18, pl. 127. figs. 1, 2 (1817) @.

Matuta lesueurii, Leach, Zool. Miscell. mi. p. 14 (1817) 3. he

Carapace finely but distinctly granulated; two anterior tubercles of the carapace obso-

lete, the remainder, as well as that upon each posterolateral margin, faintly marked ;

lateral marginal spine long, and directed slightly forward. Front slightly emar-

ginate, with the lobes rounded. Hand not granulous upon its external surface, tie

‘tubercles on the upper half sharply defined; a small tubercle, followed by a very

strong, conical spine at the base of the oblique ridge, which is interrupted at base,

SECOND SERIES.—ZOOLOGY, VOL. I. 2K

24,4) MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

and obscurely defined. Ridge upon the external surface of the mobile finger strongly

beaded. Spots upon the carapace numerous, minute, simple, scattered, and without

any tendency to form reticulating lines. Length 1} in.; breadth nearly 2 in.

Hab. Red Sea; Zanzibar; Indian Ocean; Borneo; Australia.

This is the commonest and most widely distributed species of the genus. In the

females the ridge upon the external surface of the palm is replaced by a series of two

acute spines of unequal length, alternating with three small tubercles. The ridge upon

the external surface of the mobile finger is very obscure or obsolete.

I adopt the name of IZ victrix for this species, as it is certainly figured by M. Milne-

Edwards as Matuta victor in the Atlas of the ‘ Histoire Naturelle des Crustacés;’ and

it is also the commonest species of the genus. Perhaps the Matuta doryphora of La-

treille is also to be referred to this species. The description hardly suffices for its deter-

mination with certainty.

1a. MATUTA VICTRIX, var. CREBREPUNCTATA. (Plate XX XIX. fig. 4.) __

Cancer lunaris, Herbst, Naturg. der Krabben und Krebse, i. p. 140, pl. vi. fig. 44 (1790) ?

This differs from the preceding in having the carapace more coarsely granulated, the

spots very numerous, crowded, and forming small compound spots, each having the form

(under a lens) of a small curved line or circle with paler centre. In the females the

form of the hand is similar to that of IZ. victria.

Hab. Japan; Fiji Islands; New Hebrides.

This variety, on account of its crowded, dark-red spots, presents a very different ex-

ternal appearance from the preceding; and I am in some doubt whether it ought not to

be separated as a distinct species.

(b) Carapace with numerous looped reticulated or anastomosing lines.

2. MATUTA RUBRO-LINEATA, sp.n. (Plate XX XIX. figs.5 & 6.) Carapace finely but

eu distinctly granulated. Two anterior tubercles of the carapace obsolete, the others,

Ne as well as the tubercle upon each posterolateral margin, faintly marked. Lateral

| marginal spine long and acute. Front distinctly emarginate, the lobes more acute

than in MW. victriv. Hand of male nearly as in that species, but with the spine at

base of the oblique ridge shorter. The lines upon the carapace light red, often

interrupted and more crowded anteriorly, rarely forming complete loops and circles.

Length 14 in.; breadth 14%, in.

Hab. Indian and Pacific Oceans; Cheefoo.

A specimen presented by the Smithsonian Institution is named Jf. lunaris.

Tn the females the oblique ridge upon the palm is replaced by a series of five promi-

nences, the second of which is an acute spine, the others small and tuberculiform. The

ridge upon the mobile finger is obsolete. In coloration this species resembles Maiwia

lunaris, but differs in the oblique ridge upon the external surface of the hand, and the

beaded ridge upon the external surface of the mobile finger.

MR. E. J. MIERS ON THE OXYSTOMATQUS CRUSTACEA. 245

3. MATUTA LINEIFERA, sp.n. (Plate XX XIX. fig. 7.) Nearly allied to the preceding,

but with the carapace more finely granulated, the tubercles more strongly marked,

the two anterior distinct, as are also those of the posterolateral margins. Lines

upon the carapace dark red, very fine, thread-like, usually uninterrupted, and

forming complete but irregular loops and circles. Length 13 in.; breadth 14 in.

Hab. N.W. Australia, Nicol Bay.

Only a single male adult specimen is in the collection. A larger series of specimens

might show that it is but a variety of the foregoing. |

(B) Hand of male with an interrupted ridge, or series of tubercles and spines, crossing its external surface

parallel to the lower margin, and terminating at the base of the upper margin of the lower (immobile)

jinger. The ridge upon the outer surface of the mobile finger obsolete, or smooth, or very finely striated at

base, and beaded only towards the distal extremity.

(a2) Carapace marked with spots or blotches.

A, MATUTA GRANULOSA*, sp.n. (Plate XX XIX. figs. 8&9.) Carapace rather coarsely

granulated, especially towards the lateral marginal spines. ‘Two anterior dorsal

tubercles obsolete, the others and those on the posterolateral margins very indistinct.

Lateral marginal spines short, conical, and rather obtuse; front slightly but

distinctly emarginate, with the lobes rounded. Hand of male with the upper and

lower part granulous between the tubercles; the ridge upon the outer surface in

the male divided into five acute tubercles, of which the second is the largest. Ridge

upon the external surface of the mobile finger thin, smooth, not beaded. Spots

upon the carapace brownish red, of the same shape as in J. victrix, var. crebre-

punctata, but less numerous. A brownish-red patch upon the basal half of each

lateral spine. Length and breadth nearly 13 in.

Hab. Hastern seas.

In the female of this species the tubercles of the ridge upon the external surface of

the palm are disposed as in the males. The ridge upon the external surface of the

mobile finger is obsolete.

5. Matura BANKsII. (Plate XL. figs. 1 & 2.)

Matuta banks, Leach, Zool. Miscell. ii. p. 14 (1817) ?

Carapace very coarsely granulated, especially in the centre and towards the lateral spines ;

the tubercles, as well as those of the posterolateral margins, very strong and distinct.

Lateral marginal spines of moderate length, acute; front obtuse and rounded, or

* Since the description of this species was drawn up (from three specimens, of which the exact locality is unknown,

in the collection of the British Museum) I have had the opportunity of examining a series of seventeen specimens of

both sexes and different ages in a collection made by Mr. Mac Farlane on the islands of the Torres Straits. These

examples show that M. granulosa is never to be confounded with its nearest allies, M. banksi and M. maculata,

from which it is always distinguishable by its coloration, the absence of the anterior pair of dorsal tubercles, dc.

The characters generally are subject to little variation: the only points I have observed are, that in some specimens

the tubercles are rather more clearly defined, and the ridge on the outer surface of the mobile finger faintly striated

at its distal extremity, as in other species of this section of the genus.

2K 2

IAG MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

very obscurely emarginate. Hand slightly granulous in its lower half, the tubercles

_very strongly marked, ridge in the male divided into three acute tubercles alter-

nating with two spines. Ridge upon the outer surface of the mobile finger very

finely striated in the greater part of its length, and obscurely beaded towards

its distal extremity. Spots upon the carapace pink, more numerous and crowded

upon the anterior portion, but not disposed in reticulated lines. Length and

breadth about 1¢ in.

Hab. Indian Ocean; China Seas; Philippine Islands.

A specimen from the Philippines has the spots more scattered, and a large pink blotch

upon the penultimate and last joints of the second and fifth pair of legs.

In the female of this species there are two or three conical, acute tubercles and spines

upon the external surface of the hand. Ridge of mobile finger obsolete.

6. MATUTA MACULATA, sp.n. (Plate XL. figs. 3 & 4.) Carapace distinctly granulated,

the granules coarser upon the raised parts of the carapace, but the tubercles, both

of carapace and posterolateral margin, nearly obsolete. Lateral marginal spine

very long and acute; front obtuse, or but very faintly emarginate. Hand granulous

upon its inferior surface, the tubercles upon its upper part distinct, the ridge in the

male with five prominences, of which the first, third, and fifth are indistinct tuber-

cles, the second and fourth larger, acute. Ridge upon the mobile finger as in the

preceding species. Spots upon the carapace small and crowded anteriorly, pos-

teriorly larger, scattered, and each in the form of small curved lines and circles.

Length and breadth about 1,4); in.

Hab. Kastern and Chinese Seas; Panagatan Shoal.

In the female the tubercles of the ridge upon the hand are nearly asinthe male. The

ridge of the mobile finger is obsolete.

7. Marura prcta. (Plate XL. figs. 5—7.)

Cancer lunaris, Leach, Zool. Miscell. iii. p. 18, pl. 127. figs. 8-5 (1817), nec Herbst.

Matuta planipes, Desm. Consid. Crust. p. 102 (1825), nec Fabricius.

M. picta, Hess, Archiv f. Naturg. xxxi. p. 158, pl. vi. fig. 13 (1865).

Carapace distinctly granulated. Tubercles of the carapace and posterolateral margins

stronglymarked. Lateral marginal spine of moderate length, directed slightly forward..

Front obtuse and rounded or slightly emarginate. Hand not granulous upon its

external surface; tubercles upon the upper part rounded and obtuse, ridge on

external surface divided into four or five lobes, of which the second is an acute

spine, the remainder small, obtuse, or truncate. Ridge upon the mobile finger

distinctly striated in the greater part of its length, and beaded at its distal extremity.

Spots upon the carapace very numerous and minute, often disposed in reticulated

lines. Length 13 in.; breadth 144 in.

Hab. Ceylon; Indian Ocean; Zanzibar; Mauritius; Seychelles ; Australian Seas.

In the single adult female of this species in the collection of the British Museum, the

tubercles upon the lower part of the hand are stronger and interspersed with granules.

MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA. 24,7

The second and fourth of the series of prominences on the external surface of the hand

are acute, spiniform. The ridge upon the outer surface of the mobile finger is obsolete.

The example figured by Hess under the name of Jf. picta is almost certainly the young

of this species ; and I have therefore adopted his specific name. It is easily distinguished

by the tendency of the spots upon the carapace towards a disposition in reticulating

lines.

The Matuta distinguenda, Hoffmann, Recherches sur la Faune de Madagascar, &c.,

Crust. p. 27, pl. vi. figs. 49-52, vii. 538-57 (1874), may, I think, be this species, although

in the figure, which is evidently much exaggerated, the front is represented as entire,

the tubercles much more prominent than in any species of the genus known to me,

and the immobile finger of the hand of the female much abbreviated.

(6) Carapace marked with looped, reticulating, or anastomosing lines.

8. MATUTA OBTUSIFRONS, sp. n. (Plate XL. figs.8 & 9.) Carapace rather coarsely

granulated, especially in the centre and towards the lateral spines; tubercles all

strongly marked and distinct. Lateral marginal spines of moderate length, acute.

Front small, obtuse, rounded, and entire. Hand very faintly granulous upon its

external surface, tubercles upon the upper part not strongly marked; ridge in the

male nearly as in the preceding species. Ridge upon the mobile finger very finely

striated for the greater part of its length, and obscurely beaded towards the distal

extremity. Lines upon the carapace dark red, forming irregular loops and circles,

which are smaller and more crowded anteriorly, and interspersed with spots. Length

and breadth 1} in.

Hab. Fiji Islands; New Hebrides.

In the female the same differences in the sculpture of the hand are observed as in the

preceding species.

9. Matura LunaRis. (Plate XL. figs. 10 & 11.).

Cancer lunaris, Herbst, Krabben u. Krebse, iii. p. 43, pl. xlviii. fig. 6 (1799). Jvo!

Carapace very finely granulated; tubercles of the carapace and posterolateral margins

obsolete (except in quite young specimens). Lateral marginal spines acute, of

moderate length. Front divided by a triangular emargination into two sharply

defined acute lobes. Hand not granulous, or very faintly granulous below; tuber-

cles upon the upper part strongly marked; ridge upon the external surface in the

male divided into five lobes, of which the second is acute, spiniform, and the others

small, tuberculiform. Ridge upon the mobile finger obsolete. Lines upon the

' carapace pale red, more interrupted than in the preceding species. Length and

breadth about 13 in.

Hab. Yndian Ocean, Ceylon.

The form of the hand is almost exactly the same in the maie and female of this

species.

248

MR. BE. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

DESCRIPTION OF THE PLATES.

Puate XXXVITL.

Illustrations of Species of the Family Leucosiide.

Fig. 1. Leucosia fuscomaculata, sp. n., male: natural size.

Fig. 2. The same: lateral view of its thoracic smus, enlarged.

Fig. 8. The same: outline of the abdomen, natural size.

Fig. 4. L. pulcherrima, sp. u., female: natural size.

Fig. 5. The same: lateral view of its thoracic sinus, enlarged.

Fig. 6. The same: outline of the abdomen, natural size.

Fig. 7. L. ornata, sp. u., female: natural size.

Fig. 8. The same: a lateral view of its thoracic sinus, enlarged.

Fig. 9. The same: outline of the abdomen, natural size.

Fig. 10. L. brunnea, sp. n., female: natural size.

Fig. 11. The same: inferior view, showing angular sinus, enlarged.

Fig. 12. The same: outline of the abdomen, natural size.

Fig. 18. ZL. reticulata, sp. u., male: natural size. ;

Fig. 14. The same: lateral view of its thoracic sinus, enlarged.

Fig. 15. The same: outline of the abdomen, natural size.

ee ee

ge de oq oi

da da de

eo et aS

es ee ie eo

oe 0a og de

Fig

. 16. L. whitmeei, Miers, male: natural size.

g. 17. The same: lateral view of its thoracic sinus, enlarged.

g. 18. The same: outline of the abdomen, natural size.

.19. L. perryi, sp. u., male: natural size.

. 20. The same: lateral view, showing absence of thoracic sinus, enlarged.

21. The same: outline of the abdomen, natural size.

. 22. L. pubescens, male: natural size.

23. The same: lateral view of thoracic sinus, enlarged.

. 24. The same: outline of the abdomen, natural size.

. 25, Myra mammillaris, Bell, young male: natural size.

. 26. The same: lateral view, enlarged.

. 27. The same: outline of the abdomen, natural size.

. 28. Nursia plicata, Herbst, male: natural size.

. 29. Arcania granulosa, male: enlarged.

. 80. Cryptocnemus holdsworthi, female: enlarged.

Fig.

31. The same: outline of its abdomen, enlarged.

32. The same: arm of female specimen, greatly enlarged.

Puate XXXIX.

Illustrations of Species of the Genus Matuta.

. 1. Matuta victrix, Fabr., male: natural size.

. 2. The same, hand of maie: natural size.

. 3. The same, hand of female: natural size.

MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

. M. victrix, var. crebrepunctata, sp. n.: natural size.

. M. rubrolineata, sp. n.: natural size.

. The same: hand of male, natural size.

. M. lineifera, sp. u.: natural size.

M. granulosa, sp. n.: natural size.

. The same: hand of male, natural size.

Prats XL.

Illustrations of Species of the Genus Matuta.

. Matuta banksii, Leach: male, natural size.

. The same: hand of male, natural size.

. M. maculata, sp. n.: male, natural size.

, The same: hand of male, natural size.

. M, picta, Hess: male, natural size.

. The same: hand of male, natural size.

. The same: hand of female, natural size.

. M. obtusifrons, sp. n.: natural size.

. The same: hand of male.

. 10. M. lunaris, Herbst: female, natural size.

. 11. The same: hand of female, natural size.

249

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SPECIES OF LI

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SPECIES OF MATUTA

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[ 251 ]

XIII. On the Domestic Pig of Prehistoric Times in Britain, and on the mutual Relations

of this variety of Pig and Sus scrofa ferus, Sus cristatus, Sus andamanensis, and

Sus barbatus. By Grorce Roxiestoy, W.D., F.R.S., F.L.S., Linacre Professor

of Anatomy and Physiology, Oxford.

(Plates XLI.—XLIII.)

Read June 15th, 1876.

PORTIONS of two skeletons of domestic pigs having been put into my hands by the

Rev. William Greenwell, F.S.A., from an interment of the so-called late Celtic period, 2. e.

of the ultimate or penultimate century before the Roman conquest of this country,

I determined to compare them with such other specimens of Swide as might by any

possibility be genetically connected with them. Among these other specimens I may

mention, first, several specimens of the Wild Boar, Sus scrofa, var. ferus, from the alluvial

deposits of this neighbourhood, and now in the Geological Series of the Oxford Museum,

under the charge of Professor Prestwich, F.R.S.; secondly, five specimens of the Indian

Wild Hog, Sus cristatus, kindly lent me by Sir Walter Elliot, K.C.S.1., F.L.S.; thirdly,

two skulls of Sws andamanensis, presented to the Oxford University Museum by my friend

Prof. J. Wood-Mason, of the Indian Museum, Caleutta; and fourthly, four skulls

of Sus barbatus from Borneo. The extensive series of skulls of Suidze contained in the

British Museum, those in the Royal College of Surgeons of London, and the specimens

of wild- and domestic-swine skulls contained in our own collection were also used for

this comparison.

It may be well at the outset to specify the several points of wide and general interest

upon which such an inquiry as the ensuing may be brought to bear. First among these

I would mention its bearings upon the now so commonly discussed questions relating to

the early migrations of our own species. The pig was one of the earliest, possibly the

very earliest, of animals which man domesticated; and the question of the source or

sources whence it was derived has consequently an “ethnographisch-archzeologische Bedeu-

tung” (to use the words of Fischer *, in his analogous investigation as to the sources

whence the jade and nephrite of early European times were procured) of the first import-

ance. Gibbon + has remarked that “man is the only animal which can live and multiply

in every country from the equator to the poles ;” and he has proceeded to aver that “ the

hog seems to approach nearest to our species in that privilege.’ Asa matter of fact,

Gibbon here, as so often elsewhere, was very nearly though not quite exact: the north-

ward limit of the range of the Wild Boar may perhaps be taken as somewhere between

* H. Fischer, ‘ Nephrit und Jadeit nach ihren mineralogischen Eigenschaften.’? Stuttgart, 1875. For similar

investigations as to the sources of the cultivated plants and the weeds of prehistoric times, see Keller’s ‘ Lake-

Dwellings,’ translated by Lee, pp. 303 and 343.

+ ‘ Decline and Fall of the Roman Empire,’ chap. ix. note 9, p. 352. Smith’s edition.

SECOND SERIES.—7ZOOLOGY, VOL. I. 21

252 PROF. ROLLESTON ON THE DOMESTIC PIG

55° * and 60° N., and that of the tame Pig as 64° N., whilst that of the Common

Fowl + extends probably a little, and that of the Dog certainly much, further northward.

The mention of the Australian Dingo eee another amendment of the somewhat

cynical remark of our great historian.

Secondly, a very useful light is thrown, or may be thrown, upon the question of the

extent to which the influences of civilization act upon our own species, by the analogous

inquiry into the effects which domestication has been able to produce upon an animal

linked so closely with ourselves in a many-sided commensalism. The least pleasant

aspect of that commensalism, that, namely, which is presented to us by the facts of our

solidarity with swine in the maintenance of the alternation of the forms of life of Tenia

and Trichina, must, it may be remarked, upon any view of the origination of the four

species concerned, force upon our attention considerations of the very greatest gravity.

To take somewhat lower ground, from such a study as that of the variations of the Pig

under domestication, we may obtain safe criteria for estimating the relative effects of

food, whether scanty or abundant, of early or late exercise of the sexual functions, and

of intercrossing, upon the formation of facial characteristics. The importance of not

overlooking the influences of sex and of age is nowhere more forcibly pressed upon our

attention than in an examination of a series of skulls of Suide. It has often been over-

looked in disquisitions on the skulls of Hominide.

Thirdly, whether the question at issue as regards man between the Polygenists and the

Monogenists will, as has been predicted (Darwin, ‘ Descent of Man,’ ed. 2nd, p. 180,

1874), “die a silent and unobserved death” or not, there can be no doubt that illus-

trations of the argumentations whereby that question has been, or ought to have been,

dealt with, can be furnished nowhere more fitly and fully than in an inquiry into the

distinctness or non-distinctness of the various races of swine.

Three distinct views have been advocated as to the relationship of the domestic to the

wild swine, Sus scrofa, var. ferus. We may take as the first of these that advocated by

Professor Steenstrup t, and stated by him in the following plain words :—‘ Il n’y a pas

de transition 4 observer entre les sangliers et les plus anciens cochons domestiques.”

Tn this view Professor Steenstrup will find Mr. Samuel Sidney coincide in the opening

sentences of his work ‘On the Pig;’ and a view very closely similar to it was put

tentatively forward in the year 1821 by a savant who combined the functions of a Pro-

fessor of Materia Medica with those of Director of the Botanical Garden at Berlin,

Professor Link, in the following words, to be found in his work ‘ Die Urwelt und das

Alterthum,’ i. p. 192.

« Das zahme Schwein stammt nach allen Naturforschern von den wilden Schweinen

ab, und auch die Alten waren schon dieser Meinung §; doch scheint mir die Sache

* For the northward range of the Wild Boar and the tame Pig, see Brandt und Ratzeburg, Med. Zool. p. 89;

Fitzinger, Sitzungsberichte d. Acad. d. Wiss. Wien, 1864, p. 887; Radde, Reise im Siiden von Ostsibirien, i. 236;

Zimmermann, Geographische Geschichte, i. 189, 1778. Middendorff, in his ‘Sibirische Reise,’ p. 1062, 1867, gives

56° N. lat. as the extreme actual northward limit of the Wild Boar. (Since I wrote as above, Mr. H. N. Moseley has

procured for me a skull of a tame pig from Stene i B6, Lofoten Islands, 2° above the arctic circle.)

+ For the northward range of the domestic Fowl, Gallus domesticus, see Brandt and Ratzeburg, J. c. p. 150.

+ Bulletin du Congrés International d’Archéologie préhistorique 4 Copenhague en 1869, p. 163.

§ Varro, ‘ De Re rustica,’ 1. ii. c. 1.

OF PREHISTORIC TIMES IN BRITAIN. 253

keinesweges entschieden. Die Starke, Grosse und Farbe des wilden Schweines wiirden

keinen Unterschied machen, da die wilden Thiere starker, grésser, und dunkler gefarbt

sind, als die zahmen, aber die grossen Hauer des wilden Ebers scheinen doch nicht bloss

Vergrosserung zu sein. Die Feltdecke des zahmen Schweines findet sie niemals auch

bei dem fettesten wilden Schweine, die gestreifte Farbe des Frischlings ist sogar merk-

wurdig, die Stirn der wilden Schweine ist mehr gewélbt, die Ohren sind kiirzer, mehr

zugerundet, der Riissel langer, andere Verhaltnisse an den innern Theilen zu geschweigen.

Hs fehlt ganz an Beispielen, dass die Zihmung solche Veranderungen hervorgebracht.

Vielleicht stammt das zahme Schwein von einer orientalischen Art ab, welche gross aber

unschadlich sein soll, und hin und wieder in einigen Reisebeschreibungen erwahnt wird *.

Doch erfordert die Sache noch eine genauere Untersuchung. Das siamische Schwein aus

dem ostlichen Asien abstammend, ist ohne Zweifel eine besondere Art.”

Precisely the opposite view was held by Mr. Youatt, who at p. 35 of his work on “ The

Pig,” London, 1847, says, when speaking of the Wild Boar, “ no one can for a moment

doubt that itis the parent stock from which the domesticated breeds of swine originally

sprang.” Blasius (Saugethiere, 1857, p. 509), Fr. Cuvier (Hist. des Mamm., 1824), and

Giebel (Saugethiere, p. 225, 1859), would I apprehend, agree with the extreme view

enunciated by Mr. Youatt. An intermediate view is put forward by Riitimeyer in his

inestimable work, ‘Die Fauna der Pfahlbauten,’ pp. 186-190: according to him, what,

owing to the slow spread in Germany of improved breeds, may still be called das gemeine

Hausschwein, has originated from Sus scrofa, var. ferus ; whilst the Berkshire breeds, he

thinks, may owe their origin to Sus celebensis; and his Sus scrofa, var. palustris, ‘das

Torfschwein,” a domestic pig known as “ das Biindtner Schwein,” and Sus indicus may

represent a distinct stock, if not species. Nathusius, in his ‘Schweineschadel,’ p. 175,

agrees with Riitimeyer as to the origin of the large-eared race common in Central Europe,

but suggests Sus vittatus, of the islands of Java, Borneo, Amboyna, and Batchian, as the

parent stock of the widely spread domestic breed known as Sus indicus. In this latter point

he agrees with S. Miller. Fitzinger (/. c.) differs from Nathusius in supposing, without,

perhaps, adequate reasons, Sws leucomystax to be the parent stock of the Chinese, Cape,

Portuguese, and Cleveland breeds, whilst Sus cristatus, the Wild Pig of Hindostan, he

suggests as the parent stock of the Siamese and Sardinian races. Temminck (‘ Fauna

Japonica,’ p. 57, pl. xx., cit. Nathusius, /. ¢. p. 167) suggests that Sus leucomystax may

be the parent of the domestic Sws indicus; and though Nathusius, p- 167, demurs to

the enunciation of this statement as being definitely proved, his objection amounts to

little more than saying that Sus lewcomystax is probably not specifically distinct from

Sus vittatus, which Nathusius himself holds to be the parent stock of Sus indicus.

My own views, as based upon the data available to me, are to the following effect :—-

The prehistoric domestic swine which have come into my hands appear to me to be more

nearly affined to Sus scrofa than to any of the Asiatic wild swine with which I am

acquainted ; secondly, without wishing to affirm absolutely that too much weight has

been laid by Nathusius upon the shortness of the lacrymal for differentiating Sus indicus,

as we now see it, from Sws scrofa and its progeny, I am inclined to think that sub-

* Otter, Voyage en Persie, t. i. p. 207. D. Maillet, Description de l’Egypte, t. ii. p. 176.

254 PROF. ROLLESTON ON THE DOMESTIC PIG

equality, if not actual equality, between the malar and the orbital borders of the

lacrymal bone may be found in prehistoric skulls of the Sws scrofa, var. ferus, and

especially in the female skulls of that variety of Sus which, in other points, such as the

slenderness of the snout, differ from the “‘ Torfschwein,” the representative in those

times of Sus indicus, according to Riitimeyer ; and, thirdly, I think it is possible to show

that, whilst Sus cristatus, Sus leucomystax, Sus vittatus, and Sus timorensis form a closely

connected group of Swide non verrucosi, with which, again, Sus andamanensis and Sus

papuensis are to be allied, all these subspecies differ in points of considerable if not of

specific value from Sus verrucosus of Java, from Sus celebensis, and, finally, from Sus scrofa

of the Palearctic region as well as from the non-verrucose Sus barbatus of Borneo,

It may be well to begin with this latter point first, and to show that the group of

Eastern Pigs, of which the Wild Pig of India, the Malay peninsula, and the Lancay

Islands, Sus cristatus, may be taken as a type, is always distinguishable from Sus scrofa,

var. ferus, of Europe, and Asia north of the Himalayas. ‘This view is not equivalent to

one which should lay it down as certain that they are specifically distinct, a question

which it is not proposed to raise here. I shouldagree, however, with Mr. Jerdon, whose

book on the Mammals of India, 1874, came into my hands subsequently to the formation

of my opinion, in holding that the Indian Wild Hog was “ as worthy of specific distine-

tion as many other recognized species”’ (/.¢. p. 241), though this is not to say much.

I should not, however, entirely accept his statement to the effect that the head of the

Indian Wild Hog was longer and more pointed than that of the Kuropean Wild Boar,

though this has been laid down by Colonel Sykes (Proc. Zool. Soc. 1831, p. 30) as being the

state of the case; for in measuring the relative lengths of the nasal and fronto-parietal

regions of the Indian Wild Hog, I have come to think that precisely the reverse of this

statement is usually, though not invariably, the case, the Indian Hog having the nasal

bones shorter relatively to the rest of the roof-bones of the skull than Sus scrofa, var.

ferus. Neither do I agree with Colonel Sykes in holding that the straightness of the plane

of the forehead will differentiate the Indian at least from modern European Wild Boars:

some concavity is produced in the mesial contour-line of large Prehistoric Wild Boars,

not by any angulation at the junction of the facial with the cranial bones, as in modern

tame swine of highly cultivated breeds, but by the upgrowth of the back portion of the

skull roof-bones and the occipital transverse crest ; modern European Wild Boars, how-

ever, which are much inferior in size to their prehistoric and, indeed, to their medizeval

predecessors, have the fronto-parietal and nasal lines forming one unbroken straight line.

Colonel Sykes’s words, “ Tail never curled or spirally twisted,” appear to me (J. ¢. p. 11)

to be said of the tame variety of the Indian Hog; but though Fitzinger (Sitzungsbericht.

Akad. Wiss. Wien, 1864) specifies the form in which the caudal vertebree are carried as

one of the specific marks in each of his descriptions of the Swid@, and though Linnzeus

uses the words (Hd. xiii. Syst. Nat. p. 65) “Cauda sinistrorsum recurvata” for differen-

tiating Canis familiaris from Canis lupus, s. Canis cauda imcwrvata, | am inclined to

think too much weight may be laid upon this point.

Thoroughly trustworthy figures of the European Wild Boar, such as that given by

Schreber (Saugethiere, taf. cccxx.), or that in Buffon’s Hist. Nat. v. pl. xiv., and Fréd.

OF PREHISTORIC TIMES IN BRITAIN. 255

Cuvier, Hist. Mamm., represent it as having the root and tip of its tail lying evenly

between two points; though the Vienna zoologist just referred to says of this appendage

in this animal, “‘Schwanz geringelt, kurz, nicht ganz bis zum Fersengelenke reichend.”’

And as an indication of the trifling value of such a point as this, it may be remarked that

of two female specimens of the very well-marked species, Sus barbatus, one young, the

other old, figured by S. Miller, Verhand. i. taf. 30, one has the tail curled, and the other,

the elder one, has it straight.

Mr. Blyth (cit. Jerdon, 1. ¢.) holds 8. cristatus to be only a variety of the Wild Boar of

Europe, but still to be a well-marked race. De Blainville (Ostéog. Sus, p. 129) sees no

differences of morphological importance between any of the Asiatic Swine and the Euro-

pean Wild Boar, and says, “‘ La premiére espéce que le squelette nous permette de distin-

guer par des caractéres susceptibles d’étre lus et exposés est celle quise trouve dans toute

l Afrique au dela de |’ Atlas et jusqu’a son extrémité la plus méridionale et méme au dela

dans la grande ile de Madagascar, et qui est connue sous le nom de Sus larvatus”’ (Pota-

mocherus africanus, Porcus madagascariensis, nobis). Giebel follows De Blainville in

this. Dr. Gray, in Proc. Zool. Soc. 1852, p. 180, said that, after examining ten skulls of the

European Wild Boar and its offspring from this country, from the Gambia, and from the

Cape, as also twelve skulls of the Wild Hog from Continental India, he could not discover

any constant easily described character by which the European and the Indian kinds could

be distinguished. And, he adds, “ this is the case in the many other genera allied to

the Pigs.” It is true, no.doubt, that many animals, such as “ the lion and the tiger,

the fox and the jackal, the ass and the zebra, are far more strikingly differentiated by

their pelage than by their skulls,’ as Professor Huxley (‘ Prehistoric Remains of Caith-

ness, p. 132) has taught us; still it yet remains to be proved that differences which,

though only skin-deep, are constant and permanent, will not ultimately be found to be

correlated with more or fewer differences of the deeper-lying parts, either of a purely

qualitative or of a quantitative kind.

In a disquisition the ultimate object of which is the attainment of clearer views as to

the origin of our tame pigs, the question meets us at the outset whether there exists

any marked difference between the wild stocks under comparison as regards their suscep-

tibility of domestication. Upon this point I have to say that I find, in opposition to

what Mr. Samuel Sidney has laid down in the first chapter of his work on the Pig,

that Sus scrofa, var. ferus, is credited by most trustworthy authors with as great a

capacity for domestication as any wild animal, including its wild Asiatic congeners, upon

which observations are recorded as to this particular susceptibility. Pallas’s words in his

* Zoographia,’ i. p. 269, as to the Wild Boar of the Palearctic Region, called by him, not

inconveniently, Sus ewropeus, in contradistinction to the China or Siam Pig, called by

him Sus indicus, says plainly and emphatically “ Porcelli cicurari assuescunt facile et cum

domesticis generant.” Radde’s utterances (Reisen im Siiden von Ost-Sibirien, Bd. i.

p- 286) are even more to the point, as they affirm the like of wild swine of greater age.

They run thus “So muss ich gestehen dass sie sehr friedlicher Natur sind, und es mir

mehrmals passirte mittelalte Wildschweine sich mir bis auf vier Faden Weite nahen zu

sehen.”

256 PROF. ROLLESTON ON THE DOMESTIC PIG

The Asiatic pigs, secondly, of the group represented by Sws cristatus, though not, within

my knowledge, those known as Sus verrucosus, nor those known as Sus barbatus, have very

_similar and very numerous testimonies borne to their educability and capacity for attach-

ment to man. Fitzinger, indeed, says of the domesticated Chinese Pig, his “ Sus leuco-

mystax sinensis,” that it resembles the domesticated European Pig generally in its habits

and character, but that it shows much more attachment than the European farm-pig to

the persons who take care of it, and will even follow them about, although it is other-

wise troublesome and obstinate. (Sitzungsberichte d. Akad. Wiss. Wien, 1858, Bd. xxx.

p. 235). In Formosa, when the Dutch first became acquainted with it, in the beginning

of the 17th century, every native woman, we are informed, on the authority of Ogilby

(Atlas Chinensis, ii. p. 8, ci¢. by Swinhoe Proc. Zool. 1870, p. 643), had “a great pig

running after her, as we use to have a dog.” A closer intimacy than this has been

observed to exist between Homo sapiens asiaticus, Linn., and Sus sinensis, Linn., by Pro-

fessor Huxley (cit. Galton, Trans. Ethnol. Soc. vol. iti. 1865, p. 127), who has seen

sucking-pigs nursed at the breasts of women, apparently as pets, in islands of the New-

Guinea group. As regards the wild races, Sir Walter Eliiot tells me, in a letter of date

May 15, 1876,—“ I have seen the young of Sws cristatus, which had been captured by

some of the Indian nomad communities, and reared by them, running about among the

domestic stock ; so that it would be hard to say where the line should be drawn”’*. ‘The

wild Sws papuensis has been found by Europeans, as we are informed by M. R. P. Lesson

(Voyage de la Coquille, 1826, vol. i. p. 176), to be more thoroughly domesticable than the

half-wild state in which some of the natives are content to leave it would have led us to

expect, and to set up relations of mutual amity, not only with the human, but also

with canine companions.

_ As regards the Aithiopian region, the Wild Hog is reported to us by Dr. Barth + as

consorting on terms of perfect amity, and, indeed, intimacy, with other domesticated

animals than the dog, and also with the natives. These are his words relating to a

district in Central Africa :—‘*‘ Naked young lads were splashing and playing about in

the water together with wild hogs in the greatest harmony; never in any part of

Neegroland have I seen this animal in such numbers as here about the Shari. Calves

and goats were pasturing in the fields with wild hogs in the midst of them.”

It is impossible to be perfectly certain what wild hog this may have been; still it can

scarcely have been any other than Sus sennaariensis, which has been supposed (see Darwin,

‘Domestication,’ i. p. 71, 2nd ed.) by J. W. Schiitz to have been the parent stock of

Sus scrofa, var. palustris, of Rittimeyer, but underneath the entry of which, in the

British-Museum Catalogue of 1869, p. 338, I find the following note:—‘ Dr. Murie

says he has often seen and eaten the true Wild Boar of the genus Sus in Africa, as

well as the Potamocherus, on the west coast. I have never seen any, or the skull

of one.” Like Dr. Gray, I have had no quite satisfactory opportunities for forming an

* Tt would appear that this difficulty has been felt by others to be a very real one. Colonel Walter Campbell

tells us, at p. 325 of his ‘Indian Journal,’ 1864, that he fears “the young gentlemen of the present day have taken

to spearing village pigs instead of wild boars,” and that he has “seen the thing done before now.”

+ Referred to by Nathusius, p. 147, /.c.; or see his ‘ Travels in Central Africa’, vol. iii. p. 311, 1857.

OF PREHISTORIC TIMES IN BRITAIN. 257

opinion, such as inspection of authentic skulls gives, as to the relations of Sus sennaariensis

to Sus scrofa; but whilst going over the series of skulls of Suidz in the British

Museum, I came upon one which, though entered under the head of Potamocherus

africanus, ought, I make no doubt, to be entered under the head of Sus sennaa-

riensis, or, aS I should prefer to call it, Sus scrofa, var. africanus. This skull is

numbered 715 a; and of it we have the following history from Dr. Gray (Proc. Zool. Soc.

1868, p. 35; Brit. Mus. Catalogue, 1869, p. 342) :—* A skull without its lower jaw

(715 a) was brought home by Captain Alexander from his Expedition to Damara, and

presented to the British Museum. It is recorded in Mr. Gerrard’s ‘Catalogue of the

Bones in the British Museum’ as Sus capensis (p. 277). It is the skull of an adult

animal, with the crown of the grinders much worn. It is probably the skull of a

female, as it agrees with all the characters of Potamocherus; but it has only a well-

marked ridge across the upper part of the base of the sheath of the upper canine, and

the upper margin of the nose is not dilated nor swollen.”” When I took this skull into

my hands I was uncharitable enough to suggest to Mr. Gerrard that it was a skull of

Sus cristatus wrongfully assigned to Damara as its habitat. Leaving this low ground, I

came to think that it might have come from some descendant of pigs of the Sus indicus

breed which had run wild at the Cape and reverted to the Sus cristatus form. But I

have now no doubt that this isa skull of such a Wild Boar as those of which Dr. Murie

speaks; and if this be so, the Sus of Africa is not readily to be distinguished from Sus

eristatus, at least by cranial characters. Dr. Gray, under the heading Sus sennaariensis

(Proc. Zool. Soc. ¢. c. p. 32, and British Mus. Catalogue, 1869, p. 338), suggests that the

skull figured by De Blainville (Ostéographie, tab. v.) may have belonged to a Sus sen-

naariensis. It is described as “ Sus scrofa egypti” by De Blainville; but, as far as I

can judge from the drawing, it has all the characters, to be hereinafter detailed, which

distinguish Sus scrofa, var. ferus, from Sus cristatus, and consequently from such a

skull as that labelled 715 a, and brought home by Captain Alexander. Of course there

is no @ priori difficulty in the way of our supposing that the Wild Boar either of

Palestine or of Algiers, both well-known animals, may have extended into Hgypt, a

country which has so much both of the Palearctic and of the Aithiopian fauna in oceu-

pation of its territory. Anyhow Dr. Murie and Dr. Barth are sufficient witnesses to the

fact that a true Sus is found in Africa south of the Atlas and Sahara. I cannot, therefore,

accept Mr. Wallace’s statement (‘Geograph. Distrib. of Animals,’ vol. i. pp. 253, 286-322)

to the effect that a true Sus is not to be found in the Athiopian region. The mistake

made, and handsomely acknowledged, by Fitzinger (Sitz. Akad. Wiss. Wien, 1864, Bd. 49,

i. p. 389), in naming certain young specimens of Sus sennaariensis by the name which

Fr. Cuvier had given to the Masked Boar, supposing them to be the young of that species,

would not have occurred with adult specimens. What Fr. Cuvier called Sus larvatus, we

know as Potamocherus africanus ; to save further confusion of names and errors of fact,

it would be well to drop the name Sus larvatus altogether. Potamocherus, which I

observed to resemble Sus cristatus in having a large lacrymo-frontal ridge, is, of course,

specifically distinct from it. Sus sennaariensis, if,as I think is most likely, closely

allied to Sus cristatus, is another instance of the wide distribution of ‘“‘ Pachyderms,” a

258 PROF. ROLLESTON ON THE DOMESTIC PIG

point on which Riitimeyer insists, in a different tone (Herkunft unserer Thierwelt,

p. 34) from that adopted by Gibbon, but not less categorically, nor, indeed, less strikingly,

remarking, as he does, that at the present day the Hyrax and the Hippopotamus are the

only genera of ‘‘ Pachyderms” confined to one quarter of the globe.

The Athiopian region therefore must be held to possess a true Sus; and as to domestica-

bility, the Palearctic, the Oriental, and the Mthiopian Suidze have possibly equal claims*.

There would be little weight to be laid upon mere differences in size, even if much

ereater differences did exist between Sus scrofa, var. ferus, and Sus cristatus than a

comparison of the measurements given in the British-Museum ‘Catalogue of Carni-

vorous Pachydermatous and Edentate Mammalia,’ 1869, pp. 334, 337, 338, or a compari-

son of the skulls themselves shows to be the case; for Pallas’s+ words and Nathusius’s ¢

are both clear to the effect that differences of as much as 50 per cent. do exist between

individuals of the Sws scrofa, var. ferus; and a comparison of what I believe to be the

skull of an old Wild Sow, from the alluvium of the Thames, near Oxford (Pl. XLI. fig. 2),

with that of a modern Wild Boar (Pl. XLII. fig. 5), will put the matter beyond all

question.

According to Nathusius }, the differences in size between the largest and the smallest

wild swine are so considerable as to have caused them to be distinguished by such names

* Pigs fulfil excellently well the six conditions enumerated by Mr. Francis Galton, 1. c., as necessary for domesti-

cation, viz.:—1. That the animal should be hardy; 2. That it should have an inborn liking for man; 3. That it

should be comfort-loving; 4. That it should be found useful; 5. That it should breed freely; 6. That it should be

gregarious. Adsop, Ailian, and Lactantius (cit. by Bochart, Hierozoicon, i. 698) have, in various ways, remarked on

the peculiarity of the pig as contrasted with other domestic animals, in that it is usetul only when dead, giving

neither milk, as does the cow, nor wool as does the sheep. With this peculiarity is connected the fact, useful for the

often difficult task of deciding whether a particular skull came from a wild or a domestic breed, that domestic pigs

are usually made useful while young. Riitimeyer, indeed (J. c. p. 52), gives it as one of his reasons for supposing Sus

scrofa, var. palustris, to be represented by a wild as well as by a tame stock, that its remains are usually those either of

yery old or quite young individuals. A pig will father while quite young; and whilst gaining nothing in its capacity

of manufacturer of food in its own body, it loses in its capacity of a breeding animal with increase of age. This is

not the case with the cow; and the discovery, therefore, of remains of very old individuals of this species only jns-

tifies us in inferring that the cow was a scarce and valuable animal in the period and place to which it belonged.

See Riitimeyer, ‘ Fauna der Pfahlbauten, p. 10.’

The special value of the pig as a domesticated animal is commonly expressed in an estimate that “twice the weight

of tood may be obtained from hogs than can be obtained from the same cost of food by means of any other animals ”

(Richardson, 7. c. p.42). In a little more detail, it is to be remarked that the pig, as a meat-producer, stands at an

advantage (to the consumer), first, in the smaller relative weight of its ‘‘ offal” as compared with the entire weight

of its body, but secondly, and chiefly, in the large proportion of fat, the kind of food which is eminently the hardest

for a savage or for the poor to procure, which it will store up upon almost any dietary. For this, see Lawes and

Gilbert’s invaluable Paper in the Royal Society’s Transactions for 1859, ‘‘ On the Composition of some of the Animals

fed and slaughtered as Human Food,” page 565 for relative proportion of offal, pages 513 and 5438 for storing up of

fat. It may be here remarked that De Blainville (Ostéographie, G. Sus, Introduction, p. 107) may very likely be

right in suggesting that the pig may have furnished animal food to the earliest races of man before either cow or

sheep, but that he could not have had our knowledge of the very various kinds of animals which, even in these days,

furnish lower races of men with animal food, when he supposes that animal food must have been supplied by one or

other of the three animals named. And his reason for such truth as his conclusions contain is, it is to be feared, but

a poor one; it runs thus, “le Cochon étant Vanimal qui sympathise le moins par ses qualités affectives avec l’espéce

humaine adi étre celui qu’elle aura le moins répugné 4 tuer de sangfroid pour s’en nourrir ; ce qui aura eu lieu plus

tard et avec bien plus de répugnance pour le Mouton et pour le Beeuf.”

+ Zoographia, 1831, p. 267. + Schweineschidel, 1864, p. 65.

OF PREHISTORIC TIMES IN BRITAIN. 259

as “ Hauptschweine”’ and “ Kiimmerer”’ respectively ; the latter of which terms has an

equivalent in the word ‘“‘ Wreckling,” applied in some parts of England to the super-

numerary pig in a litter, 7. e. to the one which makes the litter exceed the number of ayail-

able teats, and fares accordingly. Two or three of the commonly reported facts * as to the

pairing and period of reproduction of the wild swine account very sufficiently for these

great differences in their size. Though the males are monogamous, severe battles never-

theless take place between them for the possession, it is said, of the largest females; the

smallest females consequently are left for the vanquished, which will usually be the

smallest males. Hence a great difference in the two sets of offspring would be reasonably

expected. But, further, it is known that the wild, like the tame + swine, will breed long

before the period of maturity ; and that the offspring of such unions, whether both of the

parents or only one be immature, are likely to be smaller in size as well as fewer in

number, needs no argument.

Whilst no & priori probability can be gathered from any greater domesticability in

favour of the claims of either Huropean, Asiatic, or African Sus to be the exclusive source

of our domestic pigs, and whilst mere size equally fails to differentiate these races,

the point of the relation between the length and the height respectively of the lacrymal

bone on which Nathusius has laid such weight (Schweineschidel, passim et pp. 9, 10, 838,

91, 92, 175), and to which Mr. Darwin has assigned so much importance (‘ Animals and

Plants under Domestication,’ vol. i. p. 70, ed. 1875), though in the immense majority of

cases enabling us at once to differentiate the skulls of Sus cristatus, as indeed of the other

Asiatic pigs without facial warts, from those of Sus scrofa, var. ferus, also sometimes fails

us. Having measured a very considerable number of skulls of Sus cristatus from very

various parts of India, and having invariably found them to have the orbital border of

the lacrymal shorter than, or at most only just equal in length tothe malar, and bearing

in mind the constant reference made by Nathusius to the morphological and classificatory

value of this proportional difference in the tame variety, the so-called Sws indicus, I was

entirely unable to understand how that author could say (/. ¢. p. 185) that two skulls of

Sus cristatus furnished to him by Mr. E. Gerrard differed from skulls of Sus scrofa, var.

ferus, only in being smaller altogether. But after measuring the skulls from Sir Walter

Elliot’s collection and those in the British Museum, with the result of feeling certain

that, from the contour, proportions, and, in adult males, the texture and sculpturing,

together with the lacrymal of Sus cristatus, it was always possible to distinguish such

skulls from those of our Wild Boar, I came, in the Royal College of Surgeons, upon a

skull which, whilst possessing certain other peculiarities (to be hereafter detailed) as

distinctive, more or less, of Sus cristatus, did combine with them the long lacrymal of

Sus scrofa, var. ferus. This skull is numbered 3251 a, and was pointed out to me by

Professor Flower, with his often experienced kindness, as being a skull of Sus crzstatus,

* See Richardson, ‘ Domestic Pigs,’ pp. 18, 19 (Warne, London); Samuel Sidney, ‘The Pig,’ p. 4 (Routledge,

London) ; Blasius, ‘ Siiugethiere,’ p. 509, 1857 ; Wagner, ‘Saugethiere, p. 426, 1835; Brandt & Ratzeburg, ‘ Medi-

cinische Zoologie,’ p. 88, 1829.

+ For period at which tame sows will breed, see Sidney U. c. p.61; Low, ‘On the Domesticated Animals of the

British Islands,’ p. 415.

SECOND SERIES.— ZOOLOGY, VOL. I. 2M

260 PROF. ROLLESTON ON THE DOMESTIC PIG

which, together with two others, with the same appearance as to textural condition,

had been brought out of the stores of the Museum into its series. Both these other

skulls possessed the short lacrymal so characteristic, as the subjoined tables of measure-

ments will show, of Sus cristatus and its allies, together with the other points usually

observed in that animal’s skull. The principal points, besides those of general facies

and proportions, which appeared to me to justify this assignment of skull 32514 (Royal

College of Surgeons) were, in the absence of the shortness of the lacrymal:—first, the

great prominence of what may be called the lacrymo-frontal ridge, that part of the

frontal bone, to wit, which les between the channel for the supraorbital nerve mesially

and the upper border of the lacrymal bone outwardly ; secondly, the great relative deve-

lopment of the part of the third molar which is posterior to the two anterior bicus-

pidate lobes of that tooth; thirdly, the absence of convexity backwards in the naso-

frontal suture. These three points are usually present in Sus cristatus, as seen in

figures 4 and 6, appended to this paper, and taken from two Indian specimens; and they

are usually found to be accompanied by the fourth peculiarity of a short lacrymal. One

or other of these characters may be absent; but in an undoubted specimen of an adult

male Sus cristatws | have never seen more than one of these missing; whilst it is rare

for the second, and very rare for either the first or the third, to be found in undoubted

specimens of the Palearctic Wild Boar. Ina skull figured by Mr. Richardson (/. ¢. p. 50),

from “an excavation in an island on Loch Gur, a lake in the neighbourhood of Limerick,”

and ‘“‘ found in company with skulls of oxen, goats, sheep, red deer, reindeer, and our

extinct gigantic deer, sometimes erroneously styled the Irish Elk,” but considered by

Nathusius (p. 150) to have belonged to a domestic animal, it is true that the lacrymo-

frontal ridge is represented as of great size ; but we must set against the assigning of much

importance to this fact the considerations that the drawing is taken from a reconstructed

skull, that it is obviously inaccurate in some points, as, for example, like a drawing in

S. Miller’s Verhandel. (Taf. 28 bis, fig. 3), im having an extra tooth posteriorly to its

canines, and that it may consequently be supposed to he likely to be inaccurate in other

particulars also. The fronto-lacrymal ridge is, of course, in the adult underlain by a pro-

longation of the frontal sinuses; it is, however, visible enough in very young specimens

of domestic pigs, which show other points of affinity to the Sws indicus long before the

frontal sinuses are fully developed; and I am ixclined to think it may sometimes,

though certainly not always, be detected in very young specimens of Sus scrofa, var.

ferus, such as the one figured by Nathusius, J. c. (Taf. 1. fig. 1, Taf. 1. fig. 13). Though

the fulness of this region is due in the adult partly to its beimg underlain by frontal

sinuses, which are relatively small in the early stages of the animal’s life, there

is still some justification for regarding this structure as an instance of the retention by

the adult of an early structural arrangement; for it is easy to understand that the

contour described by the external tables in early youth may be carried out conform-

ably by the bloodvessels of the scalp as the animal grows older. A parallel to

such a process is furnished to us very frequently, though by no means so nearly

universally, by the retention in the adult Sus eristatus of that fulness and convexity of

the vertical aspect of the fronto-parietal region which is characteristic of Sws scrofa

OF PREHISTORIC TIMES IN BRITAIN. , 261

both wild and domestic, as well as of Sus cristatus and Sus indicus, at birth*. Lieu-

tenants W. E. Baker and H. V. Durand, in their paper on “ Subhimalayan Fossil

Remains of the Daduptr Collection,” in the Journal of the Asiatic Society of Bengal,

vol. v. 1886, p. 664, observe of the two fossil skulls which they describe, that in both

“there is in the frontal plane a total absence of convexity. As this plane ascends

there is a tendency to concavity, in consequence of the parietal crests being more

strongly marked than in the existing species, and thus producing the appearance of

a gentle hollow where, in the common Wild Hog, there would be a gentle swell.”

Further on, in the same paper, the authors remark :—“ From the form of the cranium,

the shape of the canines and incisors, and the other points in which the fossil differs

from the existing species of the country, a specific difference may be inferred; for the

dissimilarity, although less than that which occurs between the Babyrussa, the 8. larvatus

and the Sus scrofa, or Common Hog, is too remarkable, particularly in the shape of the

canines of the lower jaw, to admit of the fossil bemg considered as a mere variety of

the Sus scrofa.” Sir Walter Elliot, however, to whom I owe this reference as well as

other things, writes me to the effect that the skulls sent by him to me “do not seem to

differ much from the Subhimalayan fossil specimens figured and described,’ as above

specified. And it is worthy of being put upon record that of five skulls of modern Wild

Indian Hogs thus sent by Sir Walter Elliot, three show the upgrowth of parietal crests,

which Lieutenants Baker and Durand had supposed to be characteristic of the fossil

animal, and to contribute towards justifying its claim to be considered specifically dis-

tinct. These three skulls have the following labels and histories appended to them by

Sir Walter Elliot, which, when coupled with the localities assigned below to the British-

Museum specimens (specimens not, so far as I can see, different in any essential point

from Sir W. Elliot's), bear importantly on the question of the unity of Sus cristatus :—

No. 71. Large Boar, killed near Rajkote, in Kattywar, June 4, 1832. He was with

a large sounder, and ripped two horses severely. Rajkote is in the extensive

open plains of the Kattywar peninsula.

No. 330. Niigherry Hills, 1840.

No. 428. Jaggiapettah, 1851. On the east side of the Madras Presidency, in the

Masulipatam district, on the high road from Masulipatam to Hyderabad.

These three skulls agree in having their third molars considerably worn and their

canines large, their muscular insertion-surfaces marked with polygonal reticulations + in

some places, and with arborescent markings in others, and, thirdly, in the spar-like hard-

ness and density of the bones generally ; and they must be supposed consequently to have

belonged to old and powerful male animals. In all of these points they differ more or

less from the other two skulls, also of male but of younger and less powerful animals.

But such differences as these are far from being of specific value, either in comparison

of modern races with fossil ones, or in comparison of modern races inter se. All the five

skulls, however, lent to me by Sir Walter Elliot possess the lacrymo-frontal ridge

developed into a very considerable prominence; and though every now and then I have

* See Nathusius, pl. i. figs. 1 & 3, pl. ii. fig. 13, and pp.8 & 18. Compare the mesial fulness in the frontals of

S. papuensis. T For similar reticulation in Bos primigenius see Rutimacyer, Fauna der Pfahlbauten, Taf. iii. fig. 3.

2m 2

962 PROF. ROLLESTON ON THE DOMESTIC PIG

had occasion, in going over the extensive series of skulls of Indian Wild Hogs which

have been available to me, to note an almost, I have never noted an entirely complete

disappearance of it. The fronto-lacrymal ridge is not, even in the aged hog, a mere

expansion or dilatation of the frontal bone: it is underlain, it is true, by an arm of the

frontal sinuses; but it has thick and independent walls of its own. Though not quite

exactly homologous with the supraciliary ridges of human anatomy, it is nevertheless

very closely comparable with them. In some varieties of the human species the supra-

ciliary ridges, whether underlain by sinuses or not, are very constantly developed into great

prominences. In a classification, indeed, of human crania proposed by Dr. Williamson *,

“skulls with the supraciliary ridge so prominent as to overshadow the face”’ formed one out

of four classes into which all crania were divided. It is not necessary to go so far as this

for acknowledging that this character may become, in the Suidee at least, a race charactery.

The second in order of the three points which, taken together, as they usually exist

together, with the fourth, that relating to the lacrymal, help us to identify Sus cristatus, is

complexity of its third molar, at least in male specimens, and specially the complexity of

that part of the tooth which lies posteriorly to the two primary bicuspidate lobes.

A reference to my description of fig. 8 (where the third lower-jaw molar of Sws

andamanensis represents this often complex tooth reduced to great simplicity) will enable

the reader to understand this second point of distinction between Sus cristatus and Sus

scrofa more readily than any verbal description, however long, without such reference.

It may be sufficient to say here that the division of the third molar which lies posteriorly

to the two anterior lobes, is, in the male Sus cristatus, equal sometimes in size to and

sometimes even greater than both those other lobes taken together. This is not often,

though it is occasionally, the case in Sus scrofa, var. ferus, of modern times. Nor is it

the case in the females of Sus cristatus; so that the greater development observed in the

males may perhaps be due to the working of the law of sexual battle. The large size of

the canines postulates a large determination of blood to the jaw; and the large size of the

third molar, a tooth evolved at the period of sexual maturity, when the animal “ venerem

et preelia tentat,” may be, to use a word suggested long ago by myself }, “ tautogeneous ”

with it. Still the fact that the third molars are small both in Sus barbatus and Sus bar-

birussa shows that this smallness cannot always be explained on physiological grounds.

The third point, that of the straightness of the naso-frontal suture, appears to have

some classificatory value, whether we look at it with the light furnished by its

importance and validity in the classification or identification of other animals, or in that

furnished by the facts of its own developmental history.

The nasal and the frontal bones together form a roof over the ethmoid and the turbinal

bones ; and as there is no apparent physiological reason why they should contribute in

* Observations on Human Crania in Museum, Fort Pitt, Chatham, 1857.

7 It is observable in the pigmy Nepal pig, Porcula salviania, as also in the African Potamocherus. It does not

appear to me that male Suidz possess it more markedly than females; and herein it differs from the supraciliary-

ridge development as seen in our own species, as also from the race-mark to be next mentioned. An analogous

eminence, rudimentary in S. papuensis, occupies the middle frontal line over an area homologous with the human

glabella in each of four pig’s skulls brought from the Admiralty Islands by H.M.S. ‘ Challenger.’

~ Nat. Hist. Rey. Oct. 1861, p. 486.

OF PREHISTORIC TIMES IN BRITAIN. 263

different proportions towards the securing of this end, the fact that they do so is of so

much the greater morphological value. In Sus cristatus the naso-frontal suture very

ordinarily ruus, as may be seen in fig. 4(and in Sus andamanensis, in fig. 3), straight across

the roof of the ethmoid, at right angles to the long axis of the skull; or the frontals may

intrude themselves mesially between the nasals, making thus the contour of the suture to

be convex forwards. Precisely the reverse is the case in the adult European Wild Boar.

Some weight has been laid * upon a similar conformation in the skull of the Tiger, as

being of service in differentiating it from the skull of the Lion; and though it is not

pretended that an equally great distinctness can be supposed to exist between the two

animals now under comparison, still the structural differences in the two sets of cases

are analogous.

But when we come to look at the skulls of developing Pigs, we see that real value

attaches, from this point of view also, to the relatively greater or less extension back-

wards of the nasal bones, and the contour described consequently by the naso-frontal

suture. A tape crossed at right angles to the long axis of the skull from one infra-

orbital foramen to the other, passes very closely in front of, and often parallel with the

naso-frontal suture in the very young pig ; the suture gets removed further and further

away from it as the pig increases in age. Nathusius (Taf. iii. figs. 11 and 13) has

figured the skulls of a young Wild Boar and of an adult Wild Boar upon the same plate ;

and the straight line of the suture of the former contrasts most instructively with the

backwardly arching contour of the latter. The straightness, therefore, of the naso-frontal

suture may be supposed to illustrate the principle that climatic or other conditions may

cause structural arrangements to be permanently retained in certain races whilst they

are obliterated in others. The retention of the prolongation of the sagittal suture over

the frontal region is believed, with much reason, to be hereditarily transmitted in our own

species; and I incline to think that the persistence of the frontal tubera with very

much of their infantile eminence, which we see not rarely in adult men, may be taken as

furnishing another parallel to the retention in some degree, by the adult Sus cristatus, of

the characters of the young animal’s naso-frontal relations.

In the fourth place, as regards the value of the relative shortness of the lacrymal

bone as a means for differentiating the skulls of Sws cristatus and its allies from

those of Sus scrofa, var. ferus, I was for a long time of opinion that the same might

be said of this all but invariably observed peculiarity of Sus cristatus, which I have

already said, however shortly, of the straightness of its naso-frontal suture; and

that all this may, mutatis mutandis, be repeated as to the lacrymal’s peculiarities

in the two subspecies there can be no doubt. Anybody who will examine the figures

given in Spix’s ‘ Cephalogenesis, or the reproduction of them in Erdl’s ‘Tafeln der

vergleichenden Anatomie des Schidels, 1841, can convince himself of the fact that

the malar border of the lacrymal is very short as compared with its orbital border in the

young Pig +; and if he extend his observation to the various adult Suide, he will find

* Ost. Catalog. Royal College of Surgeons of England, 4506. p. 706.

+ Though I do not suppose thatit would be possible to say in 1876 what Dr. Gray said (Proc. Zool. Soe. 1868, p. 19)

m 1868, to the effect that Nathusius’s works were not to be had either in the Library of the British Museum, or in

264) PROF. ROLLESTON ON THE DOMESTIC PIG

that this side of the bone goes on growing from a condition of permanent inferiority in

Sus andamanensis till it reaches subequality or entire equality with the orbital length in

some Suidee—such as Sus cristatus, Sus vittatus, Sus leucomystar, Sus taiwanus, Sus

timoriensis, Sus papuensis, aud Sus barbatus, all Suidee without facial warts—and finally

exceeds the orbital border in length considerably in Sus scrofa, var. ferus, and. ordinarily

in Sus verrucosus (and Sus celebensis ?), and disproportionately, it may be added, in the

African Wart-Hog, Phacocherus.

There is no doubt that the younger the pig the greater is the distance separating its

lacrymal’s length from equality with or superiority to its lacrymal’s height; but the

subjoined tables and the description of my figures show that there is no constant relation

to be observed between the growth of the entire facial skeleton out of the short proportion

of early days into the elongated muzzle of the adult, and the longitudinal evolution of the

lacrymal factor of that snout. Thelongest snout, such as that of the adult Sus barbatus,

may, as Nathusius has remarked with surprise (p. 167), show a lacrymal with the

proportions of the domestic Sus indicus, or the immature Sus scrofa, var. ferus; and

I venture to suggest that the elongation of the lacrymal, which Professor Owen taught

us to call a “ mucodermal bone,’ may be correlated with the evolution of the facial

warts which are found in all the Suidee with such lacrymals. If these two structures are

thus correlated, we come to be able to explain how it is that the length of the lacrymals

is, though not very variable, still as variable as we have found it to be; for the facial

warts themselves are a variable structure, as we should from several analogies expect them

to be. If they were not so, it would be difficult to explain how it is that in many

zoological descriptions * of our European Wild Boar, no mention is made of the presence

of warts (small ones, it is true) immediately below the eyes; and inasmuch as they are

so variable, and probably more liable + to disappear in the female sex, we can, on the

hypothesis of the evolution of the Jacrymal in length being correlated with their

the library of either the Zoological, Royal, or Linnean Society, it may nevertheless be convenient to give here the

results of his measurements of the borders of the lacrymal bones at different periods of the life of Sus serofa. They

stand thus at p. 10 of his work of 1864 :—

Newly born. 2 months old. 6 months old. Adult.

Lacrymal bone. millim. millim. millim. millim.

Ieleiohitiy ena eet ai 13 18 21

Frontal border ...... 11 20 42 60

Malar border........ 3 6 25 35

Ratio of height...... 1 Eg 2-6 3

Ratio of frontal border i 18 3:8 5:5

Ratio of malar border 1 2 8:3 11:6

* De Fatio, in his ‘ Faune des Veridbrés de la Suisse,’ 1869, p. 354, goes, if I understand him rightly, further

than this, by using the words “ Pas de saillies sur la face en dessous des yeux,” in his definition of “Le Sanglier ordi-

naire, das wilde Schwein.”

+ Dr. Gray, in his description of Potamocherus africanus (P.Z.8. 1868, p. 34), says of the male animal’s face,

that it is “ swollen and often warty on the sides in front,” and of the females, that “ the side of the nose is simple.”

Fitzinger, however, whose descriptions of the external characters appear to be carefully done, does not say that any

such sexual difference exists in this species or in any other of the Suide with warts. There are other reasons,

however, for the suggestions in the text, of which the sexual limitations of the facial callosities in the Orangs may be

taken as an example.

OF PREHISTORIC TIMES IN BRITAIN, 265

presence, understand how the lacrymal is sometimes found to be shorter than we should

have expected it to be in skulls such as those of the wild sows from prehistoric deposits

in this country, measurements of which are given below.

As the interpretation I suggest of the proportions observed to exist between the length

and height of the lacrymal bone depends upon the method of gradations, as applied not

only to several varieties, but indeed to several species of Suidee, as well as upon a history

of the development of the bone, it will be well now to give a short list of those species—

pointing out the particulars which justify their claim to that rank, and contrasting them

with the aggregate of characters which do enable us, as a matter of fact, to differentiate

such forms as Sus cristatus from Sus scrofa, possibly without justifying us in considering

them distinct species.

And first of Sws barbatus. I am in no way inclined to give too much weight to dif-

ferences in colouring or in character of hair or bristles; still a glance at the drawing of

this animal in 8. Miler’s Verhandl. Taf. 30, showing its half black, half tawny, wavily,

not crisply, curling beard, its ochraceous dorsal stripe, and its tail ending in a consider-

able brush, impresses one with the idea that it is impossible that the bony substructures

should not make some approach at least to a similar diversity from other forms of Suide.

This anticipation was fully borne out by an examination of four skulls, two of which are

in the British, and one in the Oxford Museum—and of Schlegel’s figure of the skull

described by him, and 8. Miiller, Verhandl. pp. 173, 179-181, and Taf. 31. figs. 4 &5.

The skull of Sus barbatus is absolutely longer than the skulls either of Sus scrofa, var.

Jerus, or Sus cristatus; and relatively to the body its length is considerably greater,

being, as given by Fitzinger, no less than # of the length of the trunk. The contour

described by its sagittal suture is, as my figures show, strikingly different from that

-deseribed by the corresponding suture in most other Suidze; its highest point is some

way in front of the plane of the occipital squama, and occupies a level far above the plane

occupied by the anterior half of the frontal or by the nasal bones; and these latter bones

make up more than one half of the entire length of the skull, resembling herein the

typical Suws scrofa, whilst the naso-frontal suture resembles that of Sus cristatus. Its

maximum interzygomatic width is in the middle of that arch, not at its posterior

border as in mest other Suide.

In addition to these larger points, the following may be mentioned as having a great

morphological importance, though relating to smaller structural peculiarities. The third

molar consists of three lobes; but, large as is the jaw and the canine armature of this Sus,

the most posteriorly placed of the three lobes is more simple than even the very simple

posterior lobe of the pigmy Sws andamanensis, having only one cusp prominently marked

in the upper, and four in the lower jaw, whilst the entire posterior lobe is little, if at all,

greater in antero-posterior extent, and much smaller in transverse, than either of the

anterior, differing thus altogether from Sus eristatus.

A second point, relating to a small structure as measured by the callipers, which is a

very large one, however, to the morphologist, is the permanent retention by Sus barbatus

of the mesopterygoid of Parker (Phil. Trans. 1874, p. 324, pl. xxxvi. fig. iv. ms.pg) as a

distinct bone. ‘This peculiarity was observed in all the four skulls of Sus barbatus

266 PROF. ROLLESTON ON THE DOMESTIC PIG

examined by me; and its obvious general significance is increased by the fact that it is

in the area of the mesopterygoid that the great basicranial cavities are excavated in the

Babirussa and Phacocherus, and are represented rudimentarily in Sus vitiatus.

Knowing what we do* of the affinity of the fauna of the subregion of Ceylon and

South India to that of Malaya, there is no @ priori improbability in a view which

should accept the Sus ceylonensis of Blyth + (Journal of the Asiatic Society of Bengal, xx.

p- 178) as identical with the Sus barbatus of Borneo. Mr. Blyth’s words, J. c., are as follows:

—“ Sus zeylanensis. Skull longer than that of the Indian Boar, nearly straight in profile,

very much contracted at the vertex. Palate contracting posteriorly to less than 1” from

the magnitude of the last molar, which is considerably larger in both jaws than in the

Wild Boar of India, the upper measuring 13” long by +8” broad anteriorly. Vertex nar-

rowing to 1” only in breadth. Total length of skull from vertex to tips of nasals 162”.

Altogether the skull approximates closely in contour to the figures of the skulls of Sus

barbatus by Dr. 8S. Miller and M. Temminck.’’ Dr. Gray appears to have had access,

which I have not had, to a photograph of this Sws zeylanensis, and says that, judging

from it, ‘the skull is much shorter and thicker than the skull of 8. barbatus. The pho-

tograph is much more like that of Sus verrucosus.” For my own part, I cannot think

Mr. Blyth would ever have spoken of a real male Sus barbatus as having a skull with a

straight profile ; Miller and Schlegel figured a female skull, whereas Mr. Blyth’s, I am

sure, was a male’s. As regards the contraction of the vertex, which shows Mr. Blyth’s

skull to have been a male’s, this is sometimes exceeded by the old and strongly muscular

Indian Wild Hog, such as Sir Walter Elliot’s No. 71 (for history of which see p. 261),

where it is only 8”, whilst in the skull No. 72 it is 1:8”, showing an oscillation, owing to

the varying action of the temporal muscles which entirely deprives it of any morphological

value; and the measurements of the molars, finally, assure me that there is no need to

add the words “ und wahrscheinlich auch Ceylon” to the word “ Borneo” as the “ Vater-

land” of Sus barbatus, as Fitzinger has done, J. c. p. 393. Miller and Schlegel, p. 179,

give Borneo, if I rightly read their words, as the “ habitat’ of this well-marked species.

Secondly, of Sus verrucosus. The soft and perishable parts of Sus verrucosus are even

more interesting and of greater importance than the bones; for they show us that wild pigs

do have appendages—warts, to wit, covered with long bristles, and attached to the corners

of the lower jaw, like those of the Irish Greyhound Pig, once so plentiful in Galway ; and

they thus do away with one of the objections to Mr. Darwin’s views, stated fairly (by him-

self) in the work on Animals and Plants under Domestication, i. p. 79, ed. 1875. If it

is a profitable thing to lay Midler and Schlegel’s figure of Sus verrucosus (tab. 28. J. c.)

alongside of Richardson’s figure of the “ Old Irish Greyhound Pig” (/. c. p. 49; Darwin,

l.c. p.79), it is profitable also to read the Dutch letterpress | of the two former authors,

* Tennant, Nat. Hist. of Ceylon, pp. 61-68, 186; Wallace, Geographical Distribution, i. p. 328.

7 See Gray, Proc. Zool. Soc. 1868, p. 24; Brit. Mus. Cat. 1869, p. 331.

+ The words of the two Dutch naturalists are, Verhandel. p. 177. ‘‘ De deer jonge voorwerpen dezer sort zijn niet

gestreept, en onderschieden zich daardoor van de jongen van Sus vitéatus en yan de meeste, ja, misschien van alle overige

soorten.” IJtis much to be regretted that Dr. Gray did not enter a note of this most important fact in his papers

above referred to, in which the ‘ Verhandelingen’ are often cited.

OF PREHISTORIC TIMES IN BRITAIN. 267

from which we learn, when helped by the translation given by Fitzinger, that the young of

Sus verrucosus are never striped, and hereby have it suggested to us how it may be that

so many of our domestic breeds never have their young striped—none, in fact, except the

Westphalian and the Turkish. Rittimeyer (J. c. p. 187) has suggested that the so-called

“« Berkshire breed” of domestic pigs may have its parent stock in Sus celebensis, as

figured by Schlegel and S. Miller, which resembles it in the great transverse develop-

ment of its skull, the verticality of its forehead, of its occiput, and its temporal fosse,

in the height of its zygomatic arch, and specially in the remarkable height and massive-

ness of its lower jaw. There is no doubt that Sus celebensis is very closely allied to

Sus verrucosus, having not only the same general facies but the same peculiarities,

though on a smaller scale, as those whence the Java pig has taken its name. The chief

point of difference which Riitimeyer points out between the figured Sus celebensis and the

Berkshire breed depends upon the artist, 1 apprehend—being, as it is, a greater length of

the molar series, which, however, consists of 8 teeth in the figure of Sus celebensis (Ver-

handelingen, pl. 28 dis, fig. 3); and it is possible enough that the young of Sus celebensis

(the contrary not having been definitely recorded) may be like the young of Sus verru-

cosus in not being striped, and that they may thus have resembled the young of the

Irish Greyhound Pig, as when old they actually do resemble it in the development

of the mandibular warts.

As regards the skulls of Sws-verrucosus, figured tab. 32 by Miller and Schlegel, the

lacrymal in fig. 4 appears to have the favourable relation of length to height which is

characteristic usually of the other Suide with facial warts; whilst in fig. 1, which was

taken from a very old individual, the height and length would appear to be subequal, as in

the skull (fig. 6) of Sus vittatus, on the same plate. I should have set less store by this

variation in the two drawings than I do, if Nathusius had not distinctly recorded (p. 179

l. c.) that this latter is the relation in the Sws-verrucosus skull in his possession. On the

other hand, all the undoubted Sws-verrucosus skulls which I have been able to measure

have presented the elongated form of this bone; and the closely allied species Sws cele-

bensis, as figured in Miiller and Schlegel’s fig. 8, tab. 28 b¢s, may or may not have the

long lacrymal usual in that pig. But, on the whole, I should certainly accept the position

that the elongation of the lacrymal was not constant in Sus verrucosus, though it has

been so in the skulls [ have seen; and with this position I should hesitate to assign to

this peculiarity all the value which Nathusius’s insistence might lead one to assign to it.

Taking, in the second place, this variability in the proportions of the lacrymal bone,

and coupling with it, first, the absence of striping in the young, and secondly the

presence of mandibular warts, such as one of our domestic breeds, viz. the Galway

Greyhound Pig, still retains, or did till quite lately retain, we may be tempted to think

that some form allied to Sus verrucosus or Sus celebensis may have been the single

parent stock of all our domestic breeds, except, of course, such as the Westphalian and

Turkish, which are striped in their youth, and would be referred to the indigenous Sus

scrofa, var. ferus, as their parent stock. In favour of sucha view, we have, of course, the

general principle “ Entia non sunt multiplicanda preeter necessitatem.”

But the wide range* of Sus vitiatus, over Java, side by side with Sus verrucosus,

* Tt is interesting to compare with these statements, as to the geographical distribution of the Asiatic Suide, the

SECOND SERIES.—ZOOLOGY, VOL. I. 2N

268 PROF. ROLLESTON ON THE DOMESTIC PIG

over Borneo with Sus barbatus, and, as stated, also over Amboyna, Macassar, Banka, and

Sumatra, gives it special claims to attention. This Pig appears to me to be very closely

allied indeed to Sus cristatus, and to be similarly and readily distinguishable from our

European Wild Boar. In this latter point I differ from Riitimeyer (J. c. p. 187), whilst I

should agree with him in considering it all but identical with Sus leucomystax. The

claims of Sus vittatus and Sus lewcomystax, and Sus taivanus of Formosa (occurring, as

they do, in an area comprising Japan as well as Java), to have given origin to Sus

indicus, the domestic Chinese pig, in days sufficiently far off to have allowed the tendency

to striping of the young to become eliminated, are very strong, and can scarcely be

considered antagonistic.

Closely allied as Sus cristatus is to these races, its severe struggle for existence entailed

by its habitat on a continental area tenanted by the tiger (Felis tigris), as well as other

Carnivora unfriendly to Suidee, appears to me to have specialized it, especially as regards

its dental armature and the bones which carry it, into divergence from the probable line

of parentage of the inconveniently so-called Sus indicus. Sus andamanensis, on the

other hand, and, I am inclined to think, one or two other of the Asiatic Swide, show, from

a precisely opposite cause, that of restriction to a very confined area, the same divergence

from what I have imagined the present stock of the Chinese breed may have been.

But against any such speculations about what we do not see in the darkness of past

ages we have to set what we can see by travel in the broad daylight of the present, viz.

that almost all Swid@ are readily domesticable by savages in almost every quarter of the

globe; and what savages do now they may very well have done formerly.

I have not been able to institute any satisfactory examination into the relations of the

Athiopic tothe Asiatic Suide; and I should welcome an opportunity of examining skulls

and skins of the true Sws seen by Dr. Murie and Dr. Barth in Central Africa. It would

be additionally satisfactory if investigations could be set on foot as to the existence or

non-existence in this Sws of the Cystic form of Tenia soliwm, which certainly exists in

Sus cristatus. Dr. Cobbold informs me that Tenia mediocannellata is the common tape-

worm of Indian as of other patients ; but I apprehend that, as it has been so very definitely

laid down by others + that the Pig is at least one source whence inhabitants and sojourners

in India become infested with tapeworm, it would be premature to conclude the reverse

even from his authoritative statements. It must be very difficult to prove a negative here.

The facts of most direct importance, however, in investigations as to the relationships

following words from M. Gabriel de Mortillet’s memoir in the ‘ Revue d’Anthropologie,’ iv. 4, 1875, p. 653, as to

the origin of bronze:—* Reste le groupe de V’extréme Orient Asiatique. C’est la évidemment ou il faut chercher

Yorigine du bronze. Les principaux gisements sont dans la presqu’ile de Malacca et surtout dans Vile de Banca,

mais ils s’étendent dans d’autres iles de la Sonde et remontent jusque dans l’empire Birman ou I’étain est encore

exploité actuellement dans le district de Merguy. Ce minéral, dans tous ces gisements se recueille de la maniére la

plus simple et la plus facile dans les alluvions. Ce sont bien certainement les alluyions les plus riches du monde en

étain et celles qui occupent la plus grande étendue. Il est donc tout naturel que ce soit celles qui les premiéres

aient attiré Vattention de Vhomme. Le cuivre se rencontre dans les mémes régions. Tout le monde connait les

gisements de cuivre des iles de la Sonde, Timor, Macassar, Borneo. Ja Birmanie anglaise présente des mines de

cuivre 4 cdté des ses exploitations d’étain. Le pays se trouve donc dans les meilleures conditions pour avoir yu

naitre Vindustrie dw bronze.” Gallus bankiva, the parent stock of our Common Fowl, is found over the same area.

tT Notably by Dr. Charles A. Gordon, Medical Times and Gazette, May 2, 1857, p.429; and by Dr. T. R. Lewis,

Appendix B. to Highth Annual Report of the Sanitary Commissioner with the Government of India, 1871.

OF PREHISTORIC TIMES IN BRITAIN. 269

of Prehistoric to modern races are those of the great variations observed—first, in the

entire size of the individual animals, and, secondly, in the proportions of particular bones,

and notably of the lacrymal bones, in specimens from the same species or subspecies of

Suide. In view on the one side of this twofold variability which is explicable upon

acknowledged physiological principles, and affects the wild races both of Europe and

Asia, and, on the other, of the aptness for domestication possessed by Sus scrofa, var.

ferus, it appears to me to be unsafe to postulate for Prehistoric British Swine any other

parent stock than the one just named. On the other hand, such is the diffusibility and

transportability of Saws that it is not impossible, nor inconceivable, that the domestic

Buropean Pig, even of the Stone Age, may have had an Asiatic or African origin. As

regards the Bronze Age, indeed, if its tin and copper did really come from the East, such

a view cannot be said to be even improbable. But the acceptance of it does not seem to

me to be necessitated by the facts.

Weasurements of the Orbital and Malar Borders of the Lacrymal Bone*.

I. Sus eristatus and its allies.

Skull of Indian Wild Boar, Sus cristatus, no. 72. Sir Walter Elliot’s collection. Killed about 1830,

at Haugul, S.W. corner of Dharwar province, in South Mahratta country.

Height of lacrymal 1in.; malar border 1 in.

Skull of Indian Wild Boar, Sus cristatus, no. 46. Sir Walter Elliot’s collection, shot at Dendelly

12th April, 1831, in the great forest tract between Dharwar and Goa.

Height of lacrymal 1in.; malar border 1 in.

Skull of large Indian Boar, Sus cristatus, killed near Rajkote, in Kattywar, June 4, 1832. He was

with a large “sounder,” and ripped two horses severely. No. 71 in Sir Walter Elhot’s collection.

Height of lacrymal 1-0 in.; malar border 0°8 in.

Skull of Indian Wild Boar, Sus cristatus, no. 428 in Sir Walter Elliot’s collection. From Jaggia-

pettah, on the east side of the Madras Presidency, in the Masulipatam district, on the high road from

Masulipatam to Hyderabad. An old male with some obliteration of sutures.

Height of lacrymal 0°95 in.; malar border 0°95 in.

* To this paper I append a number of measurements, taken from various Sude, of the length and height of the

lacrymal bone, this point haying had great weight laid upon it by Nathusius, and holding good within very con-

siderable limits. I have not given any measurements relating to the widening of the palate (a second point insisted

upon by Nathusius), because, as regards the wild races, I have found that this widening of the interpremolar, as

opposed to the intermolar transyerse diameter of the palate, is sometimes found in specimens which undoubtedly

belong to wild European boars, whilst, on the other hand, the two measurements are usually subequal in the other

wild races, such as Sus cristatus, which I have measured. In taking the length of the lacrymal along its malar

border, some little ambiguity is caused by the fact that in many specimens the lip of the orbit is a little everted, so

as to resemble to some extent the spout of a mortar rather than the rim of a cup at the line of j unction of the malar

with the orbital border of the lacrymal. Where the differences between the ‘heights and lengths are measured by

tenths or twentieths of an inch or by millimetres, this structural arrangement may make the measurements vary

importantly. There is, however, always a line separating the part of the lacrymal which is to be considered as

belonging to the inner aspect of the orbit from that which is strictly facial ; and from this line the measurements have

been taken. The frontal border of the lacrymal, again, often bends downwards just before reaching its orbital edge,

just posteriorly to the plane of the lacrymal canals, thereby curtailing the height of the bone for the distance corre-

sponding with this deflection. My measurements have been taken in the plane occupied by the lacrymal canals, so

as to ayoid this source of fallacy.

(>)

2N 2

270 PROF. ROLLESTON ON THE DOMESTIC PIG

Skull of young female, Sus cristatus. British Museum, 716 m. “Terai, Nepal; Dr. Oldham.”

Height of lacrymal 1-1 in. ; malar border 0°75 in.

Skull of adult female, Sus cristatus,716n. British Museum. “India; Professor Oldham.”

Height of lacrymal 0:1 in.; malar border 0:7 in.

Skull of young female, Sus cristatus, about 9 months old. British Museum, 716g. “ Nepal; B. H.

Hodgson.”

Height of lacrymal 0:9 in.; malar border 0°75 in.

Indian pig, Sus cristatus, British Museum, 716g, under 5 months of age, very convex frontals, slightly

tumid lacrymo-frontal ridges.

Height of lacrymals 0°5 in. ; malar border 0:25 in.

Indian pig, Sus cristatus, British Museum, 716 v. “Nearly adult female, Nilgherris.”

Height of lacrymal 0-95 in. ; malar border 0°6 in.

Young male of Sus indicus; im collection of British Museum, not catalogued; with 3rd molar just

coming into use—that is to say, about 18 months old.

Height of lacrymal 0:9 in.; malar border 0°9 in.

Skull of Indian Wild Hog, Sus cristatus, no. 3251 a, Royal College of Surgeons of England. ‘This is

the skull of an adult male probably, the complexity of the posterior lobe of the 3rd molars being very

great. The naso-frontal suture resembles that of Sus scrofa rather than that of Sus cristatus. ‘The

lacrymo-frontal ridge is very largely developed.

Height of lacrymal 0°] in. ; malar border 1:25 in.

Skull of Indian Wild Hog, Sus cristatus, no. 3251 6, R. C. 8S. England.

Skull of Indian Wild Hog, Sus cristatus, no. 3251 c. R, C. 8. England.

Height of lacrymal 0:1 in., malar border 0:95 in., in both.

These two skulls were probably from the same district as no. 3251 a; but they differ from it, and agree

with all the other specimens of Sus cristatus,in having the height and the malar length of the lacrymal

subequal.

Skull of Wild Sow from Andamaa Islands, Sus andamanensis, no. 1514.0, Oxford University Museum.

Height of lacrymal 17 mm.; malar border 8 mm.: adult.

Skull of Sus vittatus from Amboyna. Alfred Russel Wallace, Esq. British Museum, “1362 ec.”

An adult male with a nearly straight naso-frontal suture, and a convex fronto-parietal region.

Height of lacrymal 0°55 in.; malar border 0°8 in.

Sus vittatus. British Museum, 1362 f. “Skull of animal developing the hinder molar.” The

lacrymo-frontal ridges are convex, the frontal region is flat, the naso-frontal suture, however, is straight,

as in S. cristatus.

Height of lacrymal 1:05 in.; malar border 0°6 im.

II. Sus scrofa, var. ferus, and its allies.

“« Sus libycus.” British Museum, 713 a. ‘Hab. Asia Minor, Xanthus (Sir Charles Fellowes). ‘The

skull is very distinct from all the skulls of the Wild Boars from Germany in the British Museum.” The

animal was just about 18 months old: its interectorbital region is still convex; but it has no very marked

lacrymo-frontal ridge; its naso-frontal suture is convex backwards, as in Sus scrofa, var. ferus ; its nasals

are 8 in. long. as against 142 of total skull-length; and the height of the lacrymal is less than its

inferior length. Here it resembles Sus scrofa rather than Sus cristatus.

Height of lacrymal 1in.; malar border 0:15 in,

OF PREHISTORIC TIMES IN BRITAIN. 271

Wild Boar from Germany, Sus scrofa, var. ferus, 3253a: Royal Coll. Surg. England, adult.

Height of lacrymal 0:9 in.; malar border 1:5 in.

Wild Boar from Germany, Sus scrofa, var. ferus. Oxford University Museum, 1513 a. All teeth in

place.

Height of lacrymal 1 in.; malar border 1:5 in.

Wild Boar from Germany, Sus scrofa, var. ferus. Oxford University Museum, 1518. Presented by

Prof. Max Miiller ; with all teeth, except 3rd molar, in use; 16 months old.

Height of lacrymal 0°75 in.; malar border 1:05 in.

Young Wild Boar, Sus scrofa, var. ferus. Royal College of Surgeons of England. With first molar

only through, therefore between five and six months old, with frontals still very convex in vertical aspect,

but with flat lacrymo-frontal ridge.

Height of lacrymal 0°7 in. ; malar border 0-7 in.; frontal border 1-5 in.

Skull of old Wild Sow, Sus scrofa, var. ferus, from alluvium near Oxford. Figured Plate XLI. fig. 2,

from the Geological Collection, Oxford University Museum.

Height of lacrymal 17 millims.; malar border 18 millims.

Skull, from peat, of Wild Sow, Sus scrofa, var ferus. British Museum. With 3rd molar through.

Height of lacrymal 0°8 in.; malar border 1-05 in.

Skull from silt of old bed of Lea river, shown to me in British-Museum Geological Collection by

Mr. Davis. Under 18 months.

Height of lacrymal 0-6in.; malar border 0°5 in.

* Second skull from same locality, and in British Museum, with all the teeth in place, and the third

molar with considerable complexity in its posterior lobe.

Height of lacrymal 0:6 in.; malar border 0°8 in.

Skull of Sus scrofa, var. ferus? British Museum, 7132. Skull of a young animal from the Zoological

Society’s Museum. The animal was about 16 months old, all the teeth being in place except the

third molars above and below. ‘There is still some convexity across the frontals, though flattening is

beginning on either side towards the ectorbital processes. The lacrmyo-frontal ridge is not markedly

convex; the nasals are convex backwards in the middle line.

Height of lacrymal 0-9 in.; malar border 0°75 in.

Skull of Sus scrofa, var. ferus? British Museum, 713%. Skull of young animal about 6 months old,

from the Zoological Ssciety’s Museum. This skull was entered in the Catalogue of the Zoological Society’s

Museum under the label A. 763; but no locality was recorded under this heading. The frontals are

convex from the level of the supraorbital foramina backwards ; the fronto-nasal suture is straight, nearly

as it is represented in the young German Wild Boar (Nathusius, /. c. Taf. iti. 12). Lacrymo-frontal

ridge is slightly convex.

Height of lacrymal 0°75 in.; malar border 0°5 in.

IIL. Sus verrucosus, var. ceranica.

Brit. Mus. 712d. ‘Skull, adult. A Wild Boar from Ceram. Collected by Mr. Wallace.” Of this

skull [have noted that its third molar is comparatively simple, whereas the other specimens of Sus ver-

rucosus in the British Museum have it complex. All the other specimens of Sus verrucosus catalogued

p. 330, Catalogue 1869, as Sus verrucosus, with the exception of no. 1862 d, which was received from

Mr. Wallace as Sus vittatus, have been noted by me as having the lacrymal long in relation to its height,

and as having complex molars. As, however, Nathusins, /. c. p. 179, has noted the lacrymal to be short in

a specimen obtained by him from Java, and as, of two skulls figured by Solomon Miiller (Verhandel.

tab. 32. figs. 2 and 4), one has the lacrymal short and the other long, it is clear that this species varies

272 PROF. ROLLESTON ON THE DOMESTIC PIG

in this matter. The closely allied species Sus celebensis cannot be said quite certainly, from Miller’s

pl. 28 dis, fig. 3, to have, or not to have, the long lacrymal usually found in the pigs with facial warts.

Height of lacrymal 1-0 in.; malar border 0-75 in.

IV. Sus barbatus, BoRNEO.

Skull of Sus barbatus. 1519 d, Oxford University Museum. Figured Plate XLIII. fig, 7.

Height of lacrymal 1:2 in.; malar border 1-2 in.

Skull of Sus barbatus. Procured in Borneo by the Rey. C. Spencer Bubb.

Height of lacrymal 1:2 in.; malar border 1:2.

Skull of Sus barbatus. Brit. Mus. 712a; called Euhys barbatus, Hand-list, p.58, 1873.

Height of lacrymal 1:15 in. ; malar border 1 in.

In Sus barbatus the lacrymo-frontal ridge is not very prominent, though recognizable ; in the shortness

of the lacrymal, which is 4 constant character in this species, so far as recorded in 8. Miller’s figures

and elsewhere, and in the characters of the naso-frontal suture, it resembles Sus cristatus and its allies

rather than Sus scrofa.

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OF PREHISTORIC TIMES IN BRITAIN. 273

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Radde, ‘ Reisen im Sitiden yon Ost-Sibirien,’ i. p. 236.

Richardson, ‘ Domestic Pigs,’ London, Warne.

Riitimeyer, ‘ Fauna der Pfahlbauten,’ 1861.

Schreber, J. C. D. von, ‘ Die Saugethiere,’ 1835, Taf. 320 &e.

Sclater, Proc. Zool. Soc. 1861, p. 63; 1863, p. 122.

Sidney, S., ‘The Pig,’ London, Routledge.

Spix, ‘ Cephalogenesis.’

Swinhoe, Proe. Zool. Soc. 1862, p. 361; 1864, p. 382; 1870, p. 369.

Sykes, Proce. Zool. Soc. 1831, p. 104...

Temminck, ‘ Fauna Japonica,’ Taf. 20.

Wagener, A., Gétting. Gel. Anzeigen, Ix. p. 535, 1839.

Youatt, ‘The Pig, 1847.

Zimmermann, Geogr. ‘ Geschichte,’ i. p. 189, 1778.

Postscriet, March 22, 1877.

Srnce the publication of an abstract of the foregoing paper in ‘Nature’ for July 20.

1876, I have been enabled to compare the undermentioned skulls and figures of Swide,

and to consult the undermentioned volumes and memoirs which had not previously been

accessible to me. I have also been favoured with valuable information by the letters

of a number of scientific correspondents named below. The information thus gained, in

addition to that upon which my paper was based, has been in the main confirmatory of

the views I had come to independently of it; the very important fact, however, that the

young of Sus celebensis are striped, as communicated to me by Dr. A. B. Meyer, of the

Royal Zoological Museum at Dresden, is directly contradictory of the suggestion which

TI threw out at p. 267, as is also the fact published by him as to the striping of the young

of the nearly allied §. verrucosus directly contradictory of the statement of Miller and

Schlegel given above, p. 266.

List of Sources of Information on Suidee made available to me since July 1876.

I. A skull of S. celebensis °, kindly lent me by Dr. A. B. Meyer, Director of the

Royal Zoological Museum at Dresden.

II. A skull of a Sus, sp.?, 2, from Ternate ; also lent me by Dr. A. B. Meyer.

III. Two skulls of the Domestic Pig of Bengal (S.cristatus, var. domesticus), pre-

sented to the Oxford University Museum by E. Lockwood, Esq., Monghyr,

Bengal.

IV. Two skulls of 8. scrofa, var. ferus, presented by Dr. Fiedler, of Dresden.

V. ‘A skull of S. scrofa, var. ferus, presented by M. le Marquis de l’Aigle, Chateau

de Compiégne, Pise, France.

VI. Askull of §. scrofa, var. domesticus, from Stene i Bé, Lofoten Islands, 2°above Arctic

Circle; presented by H. N. Moseley, Esq., Fellow of Exeter College, Oxford.

VII. Two bronze statuettes of S. scrofa, var. ferus, from the Gallo-Roman period in

' France; presented by John Evans, Hsq., F.RB.S.

274. PROF. ROLLESTON ON THE DOMESTIC PIG

VIII. A Memoir by Prof. L. Riitimeyer, “Neue Beitrage zur Kenntniss des Torfschweins,”

in the ‘ Verhandlungen der naturforschenden Gesellschaft in Basel,’ iv. 1, 1864,

pp- 182-186.

IX. The Inaugural Dissertation “ Zur Kenntniss des Torfschweins,” by Johann Wil-

helm Schutz, Berlin, May 4, 1868, referred to by Mr. Darwin in ‘ Animals and

Plants under Domestication,’ vol. i. p. 71, ed. 2nd, 1875.

X. A memoir by Prof. R. Hartmann on the ‘“ Verbreitung der in nordéstlichen Afrika

wild lebenden Saugethiere,” in the ‘ Zeitschrift fir Hrdkunde zur Berlin,’ Bd. iii.

1868, in which (at pp. 349-352) there is an account of §. sennaariensis, which

the Professor and Dr. Schutz consider to be the parent stock of S. scrofa,

var. palustris.

XI. Letter from Dr. A. B. Meyer, of the Royal Zoological Museum, Dresden.

XII. Letter from Professor R. Hartmann, of Berlin.

XIII. Letter from Professor Fitzinger, of Vienna.

XIV. Letter and drawing from Professor Busk, of London.

XV. Letter from the Rey. C. Spencer Bubb, late Missionary in Borneo.

XVI. Letter from G. Dobson, Esq., B.A., M.B., F.LS.

A comparison of the skull of the old male of S. celebensis lent me by Dr. A. B. Meyer,

and the measurements of it given below, with the skulls of S. andamanensis and with the

descriptions of the skull of S. verrucosus given by Nathusius, Riitimeyer, and Miller and

Schlegel, will show that while agreeing with these skulls in general contour and propor-

tions it is intermediate between them in actual size. The highest point in the sagittal

contour is, asin the Swide above named, a little way anterior to the upper edge of the

occipital squama ; the transverse arc from one orbital process of the frontal to the other is

even more convex than in any other Asiatic Pig; the zygomatic arch is short, deep, and

roughly sculptured; the lower canine’s alveolus reaches back to a prominent external pro-

tuberance in the plane of the anterior lobe of the third molar, a point further backwards

than it usually attains to. The posterior lobes of the third molars, though consisting

of 4-5 cusps, have not the great antero-posterior length relatively to the rest of the tooth

which we find in specimens of the same sex of S. cristatus.

As in S. verrucosus, the bony palate is nearer to the basicranial bones than in most

other Swide.

_ Its measurements (being those adopted by Professor Riutimeyer, ‘ Verhandlungen

Gesellschaft Basel,’ 1864, p. 163, from Nathusius, ‘ Abbildungen von Schweineschadeln,’

1864, p. 6) are as follows :—

Measurement of Skull of Sus celebensis.

millim. inches.

Length from foramen magnum to tip of intermavxillaries ......... 281 = 103

Length of occipital crest to tip of intermaxillaries ............... 325 = 12.8

Length from occipital crest to tip of masals ....................6.-- 320 = 12°55

Meneth cof masal sMiiiancs. sues snaps seceiseonoy os se aRReRe eee elise is ih == Gl

Anterior part of frontal and sagittal sutures ..............000:000e i = Be

OF PREHISTORIC TIMES IN BRITAIN. 275

millim. inches.

Posterior part of frontal and sagittal suture .....................05 y= Bs

From occipital foramen to VOMET ..............-.ceseeeenec eee ee eee Ae — nel,

Brom occipital foramen to palate 272.20... -. 2... eee erence we 2 iO

Dentizerousypart Or maradllaeseseeseeeesese ce eseee sacar tee sea 20 — ee,

THRONES FOIE CHF JOBINS ons egonsecnsoopsossooousbecsacss0ass090R0EFogoRR: Cl ee:

Greatest interzygomatic width. ..................-ss20ceeeeerec gene eee 1440 = 55

Greatest trondalllavaditeeeaee peeves sata sses eee stsctls sates -ctace er 3 = Bh

Shanalligaty texerell WIGAN, 5. asosnosoesqcsboboosenansoeaneesonaussHeceopenseces ol = ie

Intermaxillary width ................... SNe SESERS OSA ODER EOROROEDCAATNG 37 = 145

Ibigast iaeisalll wally 645 45d nodesasdsesoosanaseeteseudes souesogecesenouse so8Ds Ales

Itatkemn@lere Tmalidiy 1. sous asongoHansdesbaeseoseoossuscusovasseucdscosbosens 43 = 17

InntieryomermnGllaie WrGHHd, 225000 secacogonosacosodacocsooncasnoapanesendn 800 an ss INS

Qreqijoriivall NEVE, ascoocroradepsonsesoosuedscosccksbnvasbosscergnnagoeRBODS 1 ee

millim 6 3

Lacrymal orbital border ............ 22 = -9 compare 1 in Sus verrucosus, Riitim.

Lacrymal malar border............... aa Pe rh ee lalicss

Lacrymal upper border ............ 31 = 14 » 184

millim te

Length of S. celebensis, sec. Dr. A: B. Meyer............... 1160 = 45°67

is. ee Miller & Schlegel ............ 950 = 37-40

Length of tail of S. celebensis, sec. Dr. A. B. Meyer...... 250 = 984

5 3 Mill. & Schleg........ 150 = 59

Height of S. celebensis, sec. Dr. A. B. Meyer ............ 550-570 = 21-22

33 53 Miller & Schlegel ......... 560 =

Dr. A. B. Meyer’s specimen was more brown than Miiller’s and Schlegel’s figure,

pl. 280is; the tuft at the angle of the mouth red-brown, and the hair on the nape

longer. As to his measurements of S. celebensis, Dr. A. B. Meyer observes that they are

taken from a full-grown animal, and, being larger than those of Miller and Schlegel,

bring S. celebensis more nearly on to a level with S. verrucosus. Its smaller size suggests

the dwarfing action of a confined insular habitat undergone in some of the upheavals or

subsidences of Celebes.

Dr. A. B. Meyer informs me that the young of S. celebensis are striped, in contradic-

tion to the suggestion which I (see p. 267, supra), being impressed with the closeness

of the affinity between this Sws and S. verrucosus (which Miiller and Schlegel had de-

elared not to be striped when young), had thrown out. He informs me also that he

has the authority of General von Schierbrant, who has lived thirty years in Java and is

a first-rate sportsman, for saying that Miiller and Schlegel (/.c. p. 177) are not correct in

what they say is the case with S. verrucosus. If this be so, the claim which S. verrucosus

would have had to be considered the parent stock of our improved breed of pigs falls to

the ground. On the other hand, I cannot, knowing the great modificatory power which

domesticating influences of one kind or other have been proved to possess over the highly

plastic porcine organism, and bearing in mind the similarity between the Irish Grey-

hound, Pig, and §. verrucosus, agree with Professor Riitimeyer (p. 184, 7. c. 1864) im ex-

cluding this Svs from consideration when we are speculating as to the parentage of

our domestic Pigs.

SECOND SERIES.—ZOOLOGY, VOL. I. 20

276 PROF. ROLLESTON ON THE DOMESTIC PIG

The skull of the Sws from Ternate lent me by Dr. A. B. Meyer belonged to a young

male, the third molars of which were just coming into place, and which may be supposed

therefore, according to the analogy of S. scrofa, var. domesticus, if that may be taken as

any guide, to have been about eighteen months old (see Nathusius,/.c. p. 21). It differs

from skulls of S. cristatus in the flatness of its lacrymo-frontal ridge, in the convexity

backwards of its naso-frontal suture, and in being somewhat smaller than male skulls of

that variety of Sus usually are. The posterior lobes of its third molars are not quite so

large as they usually are in S. eristatus ; but to this point I am not inclined to assign as-

much importance as some other writers do (see Riitimeyer, ‘Fauna der Pfahlbauten,’ pp. 33

and 188, and, per contra, Nathusius, J. c.pp.49 and 103). In the straightness of its vertical

contour it contrasts with WS. celebensis, S. andamanensis, S. vittatus, as figured by Muller

and Schlegel, /.c. plate 32. fig. 6, and with S. barbatus as figured here, Plate XLIII.

fig. 7; and resembles S. cristatws, as it does also in its general facies. Dr. A. B. Meyer

thinks that this Saws may possibly deserve to be considered a new species, S. ternatensis ;

but having had abundant experience of the facility with which the wild pigs of that

region cross considerable arms of the sea, he would, I gather, express himself with much

caution as to its relations to S. papwensis and the -Suidze of the neighbouring islands, vol-

canic and other, tenanted by swine. ‘The colouring,. however, of the head of this Sus

differs from that of any other Sws seen by Dr. A. B. Meyer, or figured by Schlegel—the

head being covered all over with long black hair, except in the region occupied by a broad

yellowish brown streak beginning between the eyes and descending to the snout, where

it broadens. .

Measurements of skull of Sus ternatensis.

millims.’ inches,

Length from foramen magnum to tip of intermaxillaries ......... 290 = 114

occipital squama 5 Sail Mihalis 355 = 141

a n THD OL MASS s5o00scesdoonsoa00 B15 = ss

Inen’e tivo feniasalls eee eee Vo ats SEES AR Ee pet ee cman eons

Anterior part of line of frontal and sagittal sutures ............... 8 = 32

TROANSTAONE JORNAG Oil SEVENG INE) .oooocona0eag0sedadandaaoanangacaocagnecs0aed 9 = 37

Mio lant Merial. ates ae tgeduoiaetaroeoslen ene see semer eee berien das bepasepretaiaseek 1443 = 5:65

Incisive sbomevlenetinii. veer yes teenie reli = saa ei Ae Le emanate i —

Greatest interzysomatic width ...............csenseeeeeee eee scese snes 3a osoo

Greatesttaciall awaditlssenomencesmretece: var aa or yseee oer ceee teen cas % = 88

east sacial-wid thes ss. de shen sels naecni se eeee ea pm eae an EE ET 86 = 1:45

Juaneracnarailehar (Gli dhl eas dedusnaanusasdudandecadsosaauavscudaedsdauaaseus 40 = 16

Weds tanasalawadthy ex gccee saree ee a Acerca sani ean erat Rae eanyaa Oa —

Ibayrsrsaavolere Aili eas asda osadebe Me naneetronee aon ordradé stiadadsersasdse¢ A) == hy

Tbniterprem olla swa dit nes main a seh eres eee eee aa eee rep eee aoe 44. = 1°75

(Ojoroyamhiel levered ohh aneamae ancoouatadainansocendaseddesicare asians bate coecooees Wl = 4r4

Ibeereyamen!l Guelomitaall WORCIE 5. sovo0vesconosenoescononbasoecopcdno or vscoce 2 = i

Pe HIT DIATMOOTOSL lent ayes: es oncck oa REET Oa 8

55 LUPPCEMDOLUER crap wnacastere anise aaiaccen cate a Cece MPR Eee 34 = 13

A comparison of the two skulls of the domestic Pig of Monghyr, Bengal, which I owe to

the kindness of H. Lockwood, Hsq., and which, he informs me, are kept in large numbers

OF PREHISTORIC TIMES IN BRITAIN. 277

by the pariah castes of his district, féed on human ordure, and are sold out of the district

to the Chinamen at Calcutta, with the skulls of the wild Pig of Hindostan (S. eristatus)

shows that they differ from them in little else than their smaller size. They may be

spoken of, therefore, as S. cristatus, var. domesticus; and they are as distinguishable

from any variety of S. scrofa, var. domesticus, or indeed of S. indicus, as is the wild S.

eristatus from other wild Suide.

It is noteworthy that one of the skulls belonged toa very old sow. In these, as in the

days when Juvenal spoke (Sat. vi. 160), of another eastern country as a place where

“ Vetus indulget senibus clementia porcis,”

it is a rare thing for a domestic Pig to be allowed among western nations to live long

enough to wear down its third molars. Mr. H. N. Moseley, however, informs me that

domestic Pigs are kept in the Chinese Buddhist monasteries till they die of old age and

infirmities ; and of India, Meiners, in his ‘Allgemeine kritische Geschichte der Re-

ligionen,’ 1806, Bd. i. p. 193, says, ‘‘In Asien war von jeher Hindostan, wie in Afrika,

Heypten, der Thron des Thierdienstes.”’

It is noteworthy, secondly, that with the worn-down molars of this aged domesticated

Pig were correlated abscesses on both sides of the lower jaw, much as might have been

the case (see J. R. Mummery, Esq., Trans. Odont. Soe. ii. 2, 1869, p. 72) in ill-fed human

beings with similarly worn-down teeth.

Mr. Lockwood informs me that the young of the domestic Pigs of his district are

striped like the young of wild Pigs.

The skull of a wild boar, from France, presented by the Marquis de l’Aigle, shows that

the posterior lobe of the third molar in the lower jaw may attain the same proportions

in S. scrofa, var. ferus, that it does in S. cristatus.

The skull of §. scrofa, var. domesticus, procured for me by H. N. Moseley, Esq., from

the Lofoten Islands, two degrees within the Arctic circle, has the long, low, lacrymal

characteristic of S. scrofa, var. ferus, a fact of particular interest when coupled with the

information, also procured for me by him, to the effect that the young domestic pigs of

that region are occasionally born with stripes, and with his observation that the old pigs

have a very wild-boar-like appearance. These facts should be borne in mind as telling

against the views propounded by M. André Sanson, in his memoir “Sur la prétendue

transformation du Sanglier en Cochon domestique,” in the ‘ Journal de l’Anatomie et de

la Physiologie,’ tom. iv. 1867, p. 38.

In Professor Hartmann’s memoir above referred to, it is stated (p. 850) that certain

Negro tribes, who disobey Mahommedan precepts by eating as well as domesticating S.

sennaariensis, excuse themselves by saying it was formerly the custom to do so, a fact which

goes some little way to disprove the view that this true Sws can be merely a feral variety

of S. scrofa, imported by Huropeans. Professor Hartmann, in a letter of date Sept. 28,

1876, says, “ Sus sennaariensis ist ein kleines dem europaischen Torfschwein (der Pfahl-

bauten) ahnliches Schwein, echtes Sus, welches wild durch einen grossen Theil von

Mittel-Afrika vorzukommen scheint . . . Sus scrofa ferus in der Sahara und in Algypten

nicht selten, findet sich angeblich ebenfalls in Sennaar, indess weiss ich hieritber nichts

vollig Sicheres.”

202

278: PROF. ROLLESTON ON THE DOMESTIC PIG

Professor Busk, in a letter to me of Dec. 17, 1876, informs me that in the Etruscan

Museum at Florence, amongst numerous little bronze articles extracted from the ancient

Etruscan tombs, there were many figures of animals, one in particular being a very well

made statuette of a pig, which to his eye very closely resembles the Berkshire breed, the

only point in which it differed being the comparatively large eye, whilst the rest of the

contour was quite what we might expect to see at an ordinary cattle-show. The animal

was represented apparently as having a close curled tail. With it were a good many

statuettes of stags, the horns of which were of the type of the phocene Cervus ctenoceros.

In the very perfectly restored Htruscan tomb erected in the Museum, with all its original

contents and frescoes, Professor Busk noticed, amongst other figures of animals, one very

well drawn of a monkey climbing up a tree or pole.

I should suggest that the monkey and the pig, both alike, are representations of

animals from the same quarter of the globe as that whence the kinsmen of the Htrus-

caus in the time of Solomon brought, every three years, into Mediterranean regions, the

ivory, apes, and peacocks, the Sanskrit names of which still remain to speak to their

habitat. See Max Miller, ‘ Lectures on the Science of Language,’ 1861, ser.i. p. 190, and

ser. li. p. 234, for the source whence these animals and copper came to Europe.

Professor Busk sent me by the same post odontograms of the teeth of S. eristatus,

S. scrofa, var. ferus, and S. scrofa, var. domesticus, which show that in S. crisiatus the

antero-posterior length of the third molars is much greater in relation to that of the rest

of the molar series than it is in either of the other Suide named. I have above, p. 262,

noted that the sexual differences in this matter are very considerable in S. cristatus ;

and those produced by domestication are also not insignificant—points which somewhat

impair the value of these statements of relative proportions, though no less an authority

than Riitimeyer avers, ‘ Fauna der Pfahlbauten,’ p. 188, that “mit viel grésserer Zihigkett

das Gebiss den Species-Typus ausseren Linflussen gegeniiber aufrecht halt als die Schddel-

bildung’’ (cf. however, Nathusius, /. c. pp. 49, 103, and Studer, Mittheil. Ant. Gesell.

Zurich, xix. 3, 1876, p. 67). Professor Busk also observes that in S. crista/us nearly all

the teeth, except molar 8, are wider in both jaws in proportion to their length than they _

are in the other two, and that from this we may suppose that the Indian pig is more

exclusively herbivorous than the tame or wild animals with which he has compared it.

Jerdon speaks of S. cristatus, 1. c. p. 248, as being in general almost entirely “ vegetable

feeders.” Captain Baldwin’s views (‘The Game of India,’ pp. 154-5, 1876) are to the

same effect.

Professor Busk’s drawing and description of this Etruscan pig, coupled with a similar

figure of a pig found in the ruins of Herculaneum (see Nathusius, /. c. p. 142, fig. 38,

and Darwin, /. c. p. 71), furnishes a good illustration of Riitimeyer’s saying (Fauna der

Pfahlbauten, p. 190), that though the modern breed of domestic pigs is recent enough

to have been introduced by steamboats, it nevertheless had been represented con-

tinuously from former ages by the Biindtner-Schwein in the valleys of the Grisons.

A figure of a sow suckling three young ones may be seen on an Umbrian medal of

probably the third century 8.c. at latest, figured by M. Sambon, in his ‘ Monnaies

Antiques de l’'Italie,’ 1870, pl. v..5, with the short snout bent on the chanfrin, the

OF PREHISTORIC TIMES IN BRITAIN. 279

pendent ears, and the mane limited to the nuchal region, which justify us in considering

it to have been intended to represent a tame variety. A pig, figured ibidem, pl. iv. 4,

from Etruria has a snout of such slenderness as to correspond very closely with the

description given of §. scrofa, var. palustris, whilst it contrasts in other points very

strikingly with the wild boars represented from Apuleia and Lucania, on the other side

of Italy (¢didem, plates xiv. 1, xv. 17 and 37, xix. 4), which have as close a zoological as

legendary connexion with the wild boar of Calydon, on the other side of the Adriatic

(see ‘ Thesaurus,’ Brandenburg, i. pp. 318 & 464, 1696, and ‘Thesaurus Numismaticus,’

1. p. 400, and tab. xl.).

Of the two bronze statueties given me by Mr. John Evans, one has the long slender

snout, and the mane reaching the whole length of the convex back, from the prominent

ears to the curled tail, which may justify us in considering it as intended for a wild boar;

the other combines the sturdy straddles and the long and large erect mane, beginning on

the forehead and in front of the erect ears, characteristic of a wild boar, with a snout as

disproportionately short, and tusks as reduced as we ever see them in the highest-bred

modern Chinese pig. The characters of the wild and tame varieties, however, being thus

inaccurately and inartistically combined in these statuettes, causes them to contrast dis-

advantageously with the Italian works of art just mentioned; but they furnish us with

a conclusive answer to the weak reasoning of De Blainville (‘ Ostéographie,’ Sus, p. 170),

expressed in the following words :— Du temps de César, il parait cependant qu’elle (la

culture du cochon) n’était pas encore parvenue dans les Gaules, car il n’est nullement

question de cet animal dans ses Commentaires ; elle s’y est donc propagée depuis la con-

quéte, d’ou elle a passé en Angleterre, qui ne possédait pas méme de sanglier dans ses

foréts!” If further answer were required to this astounding statement, a reference to

Mr. Evans’s work on British Coinage would furnish it—figures of the boar, some of

which are exceedingly characteristic, being given there on pls. vi., vill., xi., Xil., and XiiL.,

from those ancient coins. But all well-informed antiquaries are aware that the wild

boar is one of the earliest animals figured in Celtic works of art (see ‘Hore Ferales,’

p- 185, pl. xiv.; Montellier, ‘Mémoires sur les Bronzes Antiques,’ Paris, 1865; and

Stephens, ‘ Literature of Kymry,’ p. 250).

Professor Riitimeyer’s paper in the ‘ Verhandlungen der naturforschenden Gesellschaft

in Basel,’ iv. 1, 1864, is, I apprehend, referred to by Mr. Darwin when he says (‘ Animals

and Plants under Domestication,’ 1. p. 71, 2nd ed. 1875) that “‘ Riitimeyer himself seems

now to feel some doubt” as to whether the “ 'Torfsechwein”’ existed as a wild animal during

the first of the stone period. No reference is given /.c. to any memoir of Professor Riiti-

meyer’s; and I became, after writing the foregoing paper, acquainted with the one in

question from a mention of it made by Herr Edmund Naumann in ‘ Archiv fir Anthropo-

logie,’ Bd. viii. 1, 1875, p. 19, in a discussion on “‘ Die Fauna der Pfahlbauten im Starnber-

ger See.” From a perusal of this paper of Professor Ritimeyer, I am inclined to think that

he would regard the skull which I have figured (Plate XLI. fig. 2) asa skull of S. serofa,

var. palustris. I have spoken of it as the skull of a wild sow, considering, as said above,

p- 259, that early breeding may, in a species admitting of such a wide range of structural

oscillation, and notably in the matter of mere size, account for a very great distance be-

280 PROF. ROLLESTON ON THE DOMESTIC PIG

tween its male and female representatives (see for similar view as regards our own species

‘Journal of Anthropological Institute,’ vol. ii. 1875, p. 122). Iam not inclined to with-

draw from this view even after reading Prof. Riitimeyer’s and Herr J.W. Schutz’s memoirs.

For in the former of these I find (p. 151) that the jive skulls used for description are

acknowledged to be skulls of sows, and four of them to have been skulls of o/d sows, and

the measurements given at p. 163 have been taken exclusively from skulls confessedly

female (see p. 161). And I learn from Herr Schutz’s essay (p. 44) that Steenstrup (cit,

Wiegmann’s Archiv, xxvii. n. 112) had distinctly stated that the “Torfschwein ” was (as

I had hinted without any knowledge of his views, see p. 254, supra) simply the female repre-

sentative of 8. serofa, var. ferus. Professor Riitimeyer had not at the time of writing his

‘Fauna der Pfahlbauten,’ 1861 (see p. 33), a single perfect skull of the “ Torfschwein”’

available to his comparison ; the almost perfect skulls treated of in his memoir of 1864 (see

p- 150) have caused him to modify the view put forward at p. 190 of the earlier work,

and in the later one he allows (pp. 158-160) that, both in the matter of the length of the

lacrymal and in the absence of widening of the palate anteriorly, the 8. scrofa, var.

palustris, was more nearly affined to 8. scrofa, var. ferus, than to S. indicus. To show this

was the reason for giving a great number of my measurements, supra, pp. 269-272. I take

this opportunity of saying that the very small breed of the Scottish highlands and islands,

with suberect ears, usually of a dusky brown colour, with an arched back and coarse

bristles along the neck and spine, spoken of by Professor Low in his ‘ Domesticated

Animals of the British Islands,’ p. 429, may perhaps be such a breed as Professor Riiti- -

meyer suggests, in his later paper (p. 168, see also p. 148), should be looked for in

Eastern Europe or Western Asia or in fossil forms*.

Mr. G. EH. Dobson, F.L.S., ina letter to me of date Jan. 15,1877, informs me that the

young of the Pig of the Andaman and of the Nicobar Islands are striped.

The Rev. C. Spencer Bubb informs me that the young Borneo domestic Pig is some-

times striped and sometimes not, whilst the young Chinese Pig is never striped; and he

adds that there are certainly two domestic breeds in Borneo.

* T suspect that latitude has more to do with the production of such varieties than longitude, and still operating

causes more than geological. One of these causes is suggested by the words of Varro, ii. 4. 13, cited by Dureau de la

Malle, ‘ Economie politique des Romains,’ ii. p. 149: “Porcique nati hieme, fiunt exiles propter frigora.”

OF PREHISTORIC TIMES IN BRITAIN. 281

DESCRIPTION OF THE PLATES.

Puate XLI.

Fig. 1. Side view of partially reconstructed skull of Sus scrofa, var. domesticus, wanting nasals and inter-

maxillaries, from late Celtic interment at Arras, Hast Riding of Yorkshire. Oxford Museum.

It is, as Nathusius has well pointed out, /. c. p. 147, by no means always easy to be absolutely

certain as to the question whether a particular pig’s skull belonged to a wild or to a domesticated

individual. The difficulty is increased when the animal is young, as in this case, the last molar

having only just come into use, and the animal consequently being only about 18 months old,

and when as the small size generally, and especially thesmall size of the third molar and the

canine, may be taken, I believe, to indicate that it is of the female sex. It is true that the

paucity of cusps in the molars has been taken as indicating the wild state; a comparison,

however, of the male and female molar series in Sus cristatus has suggested to me that the

greater size of the molars really depends upon a greater supply of blood, such as the male

molars would get, by virtue of sharing the greater supply lavished on the canines, and such as

a well-fed domesticated animal’s molars would get im common with all its other structures and

organs. The comparatively vertical occipital squama is one main anatomical point in favour

of this skull having belonged to a domesticated specimen; the pterygoid, on the other hand,

has much of the obliquity characteristic of the wild Sus scrofa, var. ferus. When we consider,

however, that this skull was found in an interment containing a human body, together with

portions of another skull of a pig of the same age, the probability that it belonged to a tame

individual appears to be very great. The subjoining measurements show that the lacrymal bone,

though not so long relatively to its height as is often the case in the non-domesticated Sus

scrofa, is yet longer than it is im Sus indicus, or in any of the Eastern pigs from which Sus

indicus can with any probability be supposed to have descended.

inches.

Length from anterior external angle (apex) of frontal to middle of occipital ridge 5:9

eneth from apex of frontal toyapex ofmaxilla) 592 9 5 6 5 5 4 ns 4

Length from anterior border of orbit to temporal ridge. . . ..... . 38

henethifromyanterior border of orbit to‘posterior 9. . 5). ae «oy

Greavestpwidubgotocemput “esc jo es o, ie || | eptctud b tbs cen neug iy acdsee

hen et hyor molar Series) viene ypaceesen be hah Gace. eat ice Brekeiiiecteh ehidoie

ers HG OCCiPUti rae Aare tes ae Ren Cee Brit ce Coc tea de 378

Evers nigomlaenyanale ia ener oe eee ee a OR ed eens ise OTT.

Tengthvor lacrymo=malar sninre ey nue Welland yi tly yi OI

The difference here noted between the height of the lacrymal and the length of its lower

border amounts but to a tenth of an inch ; but in the female of Sus indicus, with the same stage

of dentition, Pl. XLII. fig. 4, the height of the lacrymal, instead of being one tenth of an inch

less, is 0°35 inch greater in length than its lacrymo-malar border.

Fig. 2. Skull of Sus scrofa, var. ferus, old 2, from alluvium of Thames valley, obtained for the Oxford

University Museum from the cutting for the drainage works near Iffley, 1876, by Professor

Prestwich, F.R.S.

This skull combines the general contour and the slender snout of Sus scrofa, var. ferus, with

a lacrymal bone differing little in its proportions from the lacrymals so characteristic of the

Asiatic pigs, less Sus verrucosus (and celebensis). ‘The fronto-parictal region does not form one

282

PROF. ROLLESTON ON THE DOMESTIC PIG

continuous slope in the same plane as is the case in the Wild Boar of Germany (Pl. XLII. fig. 5);

but this difference may be observed in skulls certainly of Wild Boars from the Thames-valley

deposits ; whilst the great wear of the teeth (see p. 258, note * above) and the slenderness and

length of the naso-facial region are much in favour of considering this specimen to have

belonged to the wild race. I have placed side by side with the measurements of this skull the

measurements given by Riitimeyer, /. c. pp. 45 and 183, of his “ Torfschwein,” Sus scrofa, var.

palustris. Unhappily, Rtitimeyer has never been able to procure (see pp. 43 and 45, note 1) a

skull of this variety of Sus with the facial bones in connexion with the braim-case, nor has

Nathusius (see p. 149, 7. c.) ever been able to see an uninjured lacrymal bone from the same

animal. These facts, whilst making the value of this skull (the opportunity of figuring which ~

I owe to the kindness of Professor Prestwich) greater, make the value of the comparison of its

measurements less.

The instructive observations of Nathusius (J. c. pp. 99-101), to the effect that ill-nourished

pigs have the entire length of their skulls greater as measured from the occipital crest to the

apex of the snout, whilst the portion of that length made up by the frontal and parietal is

somewhat shorter, and the nasal portion proportionally longer, when coupled with the fact of

the great wear of the teeth in this specimen, enable us to explain the one great point of

inferiority, that of the length of the fronto-parictal region, which this skull’s measurements

show us, compared with those of the “Torfschwein.” It may be added, that the true explana-

tion of Dr. Gray’s statement (Brit. Mus. Catal. 1869, p. 329), that the nasal bones of the skull

elongate as Suid increase in age, ‘and especially as they reach adult and old age,” is probably

that he had im his mind’s eye skulls of old and ill-fed wild pigs, such as this specimen., On

the other hand, when we are comparing such skulls as this with the “Torfschwem” of

Riitimeyer, we must recollect that he represents this latter variety of pig, and what he supposes

to be its still surviving representative, the “ Biindtner-Schwein,” as having a short snout (see

pp. 42, 45, 181-185). The classificatory value, however, of such a peculiarity is much reduced

by the results of such experiments as those of Nathusius just referred to.

Measurements of Iffley Skull and of Riitineyer’s “ Torfschwein,” pp. 45, 183, 1. c.

Iffley Skull. Torfschwein.

inches. inches.

Greatest frontal width between ectorbital processes . . . . . 32 37

Lease WHO OR TOMER 6 s 6 5 6 6 6 2 8 2 ob wo op) OS 0-9

Greatest interzygomatic width . . . . . ot eevee, TAG 4 7—4e9

Height of occipital from inferior border of fame magnum . . 35 38-45

Length of vertex from level of supraorbital foramen to ridge of

OUSIDE 6 56 6 : : ORS An

Horizontal distance com anterior Bonde of oie to oueeenee of

temporal fossa. . . : ‘ is lade Se Me aol le ae eo 31

Length of intermaxillary aoe Resin hordes pr acorns anh ao coed 1:9-2°4.

Maximum length of skull from apex of intermaxillaries . . . . 10°9 —

Length of nasal hones (approximately). . . ..°. . =. . . O55 —

Length from apex of intermaxillaries to inferior border of foramen

magnum . . 5 ; - 103 —

Length from anterior een Pls ened 0 of fr intel to eae of

occipital ridge. . . . 5 : 5:9 =:

Length from middle of ponte suture to male of sicelinttl

TCE Fat ata me eas cull ict ates No (MES aictg io ae, QoaMi RU uN Oa mae od iS _

OF PREHISTORIC TIMES IN BRITAIN. 283

Tffey Skull. Torfschwein.

inches. inches.

Length from anterior border of orbit to posterior. . . . . . 4I4 —

Greatest width of occiput . . . + hire Ppeie —

Breadth of nasal at commencement of eeeteontal Peck which

is point of maximum width ofnasals . . . . ..... 4d _

Maximum width of intermaxillaries . ......2.. =. «18 —

Height of lacrymal along rim oforbit . .... . . . . 17 millims.

Length of lacrymal along malar suture. . . ... =... =. IJ8wmillims.

Fig. 3. Sus andamanensis. 15146, Oxford University Museum, adult 9.

The skull from which this drawing was taken was that of a Wild Sow from the Andaman

Islands, procured for me by my friend J. Wood-Mason, F.G.S., of the Calcutta Museum. It is

about the same size as the two prehistoric British skulls, figured figs. 1 and 2, and as the

“ Torfschwein ” of Riitimeyer—resembling this latter im the one important particular, that of

its long fronto-parietal region, in which it differs from the British skull (fig. 2). It differs

from all the pigs’ skulls here figured in the exaggeratedly disproportionate shortness of the

malar border of its lacrymal bone, which is, as in the newly born European pig, little more

than half the length of the orbital border. In its convex frontal region we have, again, a

character retained in the adult Asiatic which is transitorily represented in the European pig. This

convexity of the fronto-lacrymal ridge is well marked and, like the straight naso-frontal suture

and the two other peculiarities specified, serves to differentiate Sus andamanensis from specimens

of similar size from the Palearctic region, and to show that it is a member of the group repre-

sented by Sus cristatus. Again, this skull resembles Sus cristatus and differs from Sus scrofa

in bemg somewhat shorter, and having the paroccipitals, temporal bulle, and pterygoid processes

more crowded together. The length of the nasals is a little less than that of the fronto-

parietal region in the vertical aspect, as is often, though not always, the case in Sus cristatus,

though very rarely, if ever, in Sus scrofa.

Neasurements of Skull of Sus andamanensis, female.

inches

Greatest frontal width between ectorbital processes. . . . .... =.=. 33

Meas tawidthnoumvervex ssa st ro mater eee to er ee a sy) ee OS

Greatest interzygomatic width. . . . rach TUT cee sa eee mercer Ae OD,

Height of occiput from inferior border of adteh magnum .. . . 37

Length of vertex from level of supraorbital foramina to middle of eee Heth 4:5

Horizontal distance from anterior border of orbit to temporal ridge . . . . 3:05

Length of intermaxillary along alveolar border . . ....2.2.2.~. 20

Maximum length of skull from apex of itermaxillaries. . . . .... 10: 8

Length ofnasals . . . . é db SME RS

Length from apex of stir ee erticid to ee ae of anaes magnum . . 91

Length from apex of frontal to middle of occipital ridge . . . 6 Jo ra BS

Length from middle of fronto-nasal suture to middle of occipital eo sees om ORO)

Length from anterior border of orbit to posterior . ...:. .. =.=... «CLS

Greatest width of occiput . . . : : 2°5

Breadth of nasals at commencement of naso- ee sate aie is me peadeet

partofnasals. . . . BEB dy! CST heaters ochre maces end eet cocaine 60)

Maximum width of as tate) LEMON Pee Ree 2 ey eee et soe eee ve

Heizhtiotlacryndl-alonetrim! of orbit’. 9. 2): UR OS SOB 2? 17 enim

SECOND SERIES.—ZOOLOGY, VOL. i. 2P

284

Fig.

PROF. ROLLESTON ON THE DOMESTIC PIG

Length of lacrymal along malar suture BR enn SNE gs Gling. Tease Neen peevadnli ns

[GRIN OE uOleGEE) 5 6 5 4 56 9 0 0 50 9 0 8 9 6 oe cs 4 6 ADCS,

Thensthvor premolar Series) ir itll ss i-inr a cE

Pratt XLII.

4, Skull of Wild Indian Sow, Sus cristatus, British Museum. See ‘Catalogue of Carnivorous,

Pachydermatous, and Edentate Mammalia,’ 1869, p. 334 (where this skull is catalogued as

“716 m. Skull of young female. Length 10}, height 74 inches. Terai, Nepal; Dr. Oldham”),

or Hand-List, published 1878, p. 64.

This skull has been figured to show the points in which it contrasts with the British pig’s

skull (figure 1) on the one hand, as to racial characters, and with the Indian boar’s skull (figure

6) as to sexual characters, agreeing, as it does, with the one as to sex, and with the other as

to age.

It is, as may be seen by constructing two triangles with the three first measurements given

of it and of that of Sus cristatus (fig. 6) respectively, a very much smaller skull than that ;

the principal points, however, enumerated in the description of that skull, as differentiating

it from Sus scrofa, can be recognized in its smaller contours. The third molar in the upper

jaw has the part posterior to its second principal lobe much smaller relatively and absolutely

than is the case in Boars of Sus cristatus; this posterior factor, however, effloresces into as

many as 7 cusps in the upper and 5 in the lower jaw of this specimen.

Measurements of Skull 716 m, British Museum Collection. Sus cristatus, from

Terai, Nepal.

inches. inches.

Vertes lemme 9 6 5 0 0 oo 6 Alle Length of lacrymal along malar suture 0°75

Rixtreme hercht.- ani EHiS Interpremolar transverse diameter of

Bee 5 6 oo eo 0 ep 6 oo NO (HAWS soe) oo Go 6 oo LG

Height of lacrymal along rim of orbit iil iumiemim@kie 6 6 56 5 6 6 5 « o lve

5. Skull of Wild Boar, Sus scrofa, var. ferus, from Germany. No. 1513 a, Oxford University Museum.

The entire dentition was in place; the portion of the third molar which is posterior to the four

primary cones is very small as compared either with the rest of the tooth or with its homologue

in the Indian Wild Boar (fig. 6). It is, however, considerably in front at a line drawn as a

perpendicular to the plane of the cutting-edges of the molars from the lacrymal canal. This

skull differs markedly from the skull just named in the greater length, relatively to the

height, of its lacrymal bone, in the greater length of its nasals relatively to the length of the

fronto-parictal region, in the absence of any great convexity in the part of the frontal abutting

upon. the lacrymal, and in the convexity backwards of its fronto-nasal suture.

Measurements of Skull of German Wild Boar, no. 1513 a, Oxford University Museum.

inches. inches.

Extreme length. . . ... . . 165 \Wichin GP meRAI), 5 6. 6 oo oo CS

Taine Inerlng 5 oc 0 6 6 9 9 a | Ol Maximum interzygomatic width . . 59

Base-line from anterior margin of fora- Maximum frontal (imterectorbital)

men magnum to apex of intermaxil- Wilda 6 6 2 o blo 6 0 o 5 ay

laricsmeyte Gee! ca ee LAO Height of lacrymal along rim of orbit 1:0

Length of nasals, approximatively . 80 Length of lacrymal along malar suture 1°2

Fronto-parietal length in vertical aspect 6°8

OF PREHISTORIC TIMES IN BRITAIN. 285

Prats XLITI.

Fig. 6. Skull of Indian Wild Boar, Sus cristatus, killed at Haugul, Dharwar Province, from the collection

of Sir Walter Elliot, K.C.S.I. No. 72.

In this animal the third molar is only just coming into place, and its very large multicuspi-

date posterior lobe is not so far forward relatively as in the European Wild Boar (fig. 5)

nor in other Indian hogs of greater age. It shows, however, very plainly the poiuts, several,

if not always all of which have, by their presence in every specimen of Sus cristatus which I

have examined, enabled me to distinguish it from Sus scrofa, var. ferus. It has the relatively

short lacrymal—a tape, stretched as an arc across the long axis of the skull, from the anterior

inferior angle of the bone on one side to the homologous point on the other, passing over the

frontals, and not over any part of the nasals; the naso-frontal suture, which lies entirely in

front of such an arc, is straight, and not convex backwards; the portion of the frontal which

is bounded internally by the supraorbital channel, and externally by the lacrymal bone, is

markedly convex. The nasals are broader, as is nearly always the case, and shorter also, which

is not by any means always the case, relatively to the fronto-parietal region of the vertex,

than in Sus serofa, var. ferus.

Measurements of Indian Wild Boar, no. 72, Sir Walter Elliot’s collection.

inches.

Extreme length , 16:9

Toa Inara 4G Gc ee ee el

Base-line from ant. margin of foramen

magnum to apex of intermaxillaries 13

Henethyofmosalsisne mylulal eur uae

Length of fronto-nasal suture to

middle of occipital ridge . 58

Width of nasals at apexof frontals . 2

Maximum interzygomatic width . 6

Maximum frontal (interectorbital)

Width: Uo \yaaiewecs ors His Ue et Al

Fig. 7. Skull of Sus barbatus, Borneo.

inches.

Minimum vertical width . . . . 41:8

Height of lacrymal along rim of orbit 1

Lengthof lacrymalalong malar suture 0:9

Interpremolar transverse diameter of

palate soe vunyn eat toon cee

Intermolar . tian yeti 2°05

Length of posterior upper molar . 1°35

IDFRENOMIN Seo a REOMPON Guu Beach Hub nor MES)

Length of third lower molar . . . 1:6

peccadt hy. eee, oe la eke §©6(OF8

1519 d, Oxford University Museum.

This skull, like the other three skulls of the same species examined by me (of which two are

in the British Museum), differs from those already described in large points as well as in small

ones; and there can be little reason for hesitating to accept it as specifically distinct from

them and, indeed, from all other Suidz.

The contour described by the middle line of its nasal and fronto-parietal regions superiorly,

the relations of the greatest width and greatest lengths both of the entire skull and of the nasal

bones, the position of the plane of its greatest interzygomatic width, not posteriorly, but in the

middle of the zygomatic arch, are points of large difference. The exceeding simplicity of its

third molars and the persistence of the mesopterygoid as a distinct bone, are points of small

difference, but yet of great morphological importance.

resembles the other Suid without facial warts.

Its naso-frontal suture and lacrymo-frontal ridge are more like those of these pigs than those

of Sus scrofa.

In the shortness of its lacrymal bone it

286 ON THE DOMESTIC PIG OF PREHISTORIC TIMES IN BRITAIN.

Measurement of Skull of Sus barbatus, 1519 d, Oxford University Museum.

inches. inches.

xtreme lensthy alc seni) ere LOW Maximum frontal (interectorbital)

DAIS eee a Bene circle. cua gh OF width . 45

Base-line utes Ala bark asic G37: Minimum vertical . 1°4

Length of nasals fio pan of postero- Lacrymo-malar line 1)

lateral tips . . ; AL, enrliOr Height of lacrymal 1:2

Fronto-parietal region in eel aspect Length of posterior upper molar 1:4

from same point as preceding mea- Breadth of anterior, which is much the

surement to ea ridge in straight widest lobe acy i 0:9

lime... Meld ice Menem nese ogi O30: Length of posterior lower molar 1:5

Width of batty Nias Meantitye oti (5 Breadth of anterior lobe . 0:8

Maximum (interzygomatic) widih 5 OS Intermolar space at narrowest point 11

Fig. 8. Left lower third molar of Sus andamanensis, female, 24 times the natural size.

This tooth shows the three divisions of the third molar of the true Suidz in great simplicity.

There are two bicuspidate lobes corresponding to the two principal lobes of molar 1 and molar 2,

and, like them, enclosing a single azygos lobe in the middle line between them. This azygos

lobe is developed from the second bicuspidate lobe. There is no ridge developed in Sus anda-

manensis on the anterior part of the tooth, é.e. on the face in contact with molar 2. Poste-

riorly to two bicuspidate lobes, which already show signs of wear, are seen five smaller cusps,

occupying in all a much smaller space than the rest of the tooth. In the males of Sus cristatus

these five smaller cusps would, as in well-fed domestic pigs, occupy a very much larger space

relatively and absolutely than they do here, or even in Sus scrofa, var. ferus. In Sus barbatus

the third lower molars are as simple as they are in Sus andamanensis, and, like this Sus, have the

third molars of the upper jaw simpler still than the lower. The five posterior cusps consist of

one placed mesially in the interspace between the primary cusps of the second pair, of three

placed like the dots in the sign -.-, and of a fifth, not constant, accessory cusp placed on the

inner side. The four latter are represented in the upper jaw of this pig by two, and in Sus

barbatus by one main cusp.

Trans. Linw. Soc. Ser.2.Zoon Vou.) Tas. 41

D.SCROFA wan FERUS. ?

Thames valley.

Caldo

Fig. 1

S.SCROFA va: DOMESTICUS. 2

Colite aierment, Yorkshire.

alba

Fig. 3.

S_ANDAMANENSIS. 2

C.Berjeau hth. sus Hanhart imp

dur yaretpap}y

Un] nealseg9

Aouisap

SOeddd WaVIOHOS Ss

9 bag

‘6 SNIVLSIHO'S

p bay

ch a¥] [TOK 1007 2 ass ‘00g NNIT SNVvH],

lrans. Linn Soc Ser 2 Zoo. Vor! Tan 43

Low. 3 molar

§.BARBATUS ”

D, ee

{SOT RCO ail

colt

C Berjeau itl Hanharl imp

‘ t fe 5 “y 5

7 a

E " & =~

. rt

sad

Z rf i

" 2

> iy

XIV. On two new Forms of Deep-sea <Ascidians, obtained during the Voyage of

HM.S. ‘ Challenger” By H.N. Mosstey, I.A., F.R.S., Fellow of Exeter College,

Oxford. (Communicated by Dr. J. Muniz, F.L.S.)

(Plate XLIV.)

Read December 21st, 1876.

THE two Ascidians herewith described and delineated are most unusual and aberrant

in several particulars and quite new to science.

The first form was obtained from the great depth of 2900 fathoms. I propose to

name it Hypobythius calycodes, in allusion to its occurrence at a vast depth and its cup-

like form. It was unfortunately imperfect when obtained. It is remarkable for

having a series of cartilaginous plates developed in its otherwise rather soft and

yielding test, which plates are arranged in symmetrical patterns, as will be seen by the

figures.

The other form, which I propose to name Octacnemus bythius, is most remarkable, on

account of its stellate eight-rayed form, and abnormal in the details of its structure. It

was obtained from 1070 fathoms. It appears to be entirely without near allies in the

group of Ascidians. The gill-sac is flattened out so as to be nearly horizontal, and

there is no gill-network. Peculiar developments of the muscular tunic are prolonged

into the eight curious conical protuberances of the test. The principal viscera are con-

tracted within a small rounded mass or nucleus, as in Salpa.

As only a single specimen of each of the forms was dredged by H.M.S. ‘Challenger,’

the specimens themselves are, I presume, unique; and since they are highly interesting

in many respects, are well worthy of the Linnean Society’s notice.

I. HyPoBYTHIUS CALYCODES, gen. et sp.nov. (Plate X LIV. figs. 1-6.)

On July 5th, 1875, the trawl was used in 2900 fathoms in the North Pacific Ocean,

lat. 37° 41’ N., long. 177° 04’ W. ‘The bottom consisted of a red clay with numerous

concretions of peroxide of manganese. A gigantic Hydroid, a Monocaulus* seven feet in

the length of its hydrocaulus, fragments of Hyalonema, a Holothurian, and a few other

animals were obtained, and with them the Ascidian now to be described in detail. The

specimen was considerably lacerated, but almost all its essential structures were pre-

served. The animal has the form of an inverted cone, compressed laterally so as to have

an oval transverse section. From the apex of the inverted cone is continued a cylindrical

stem, which is enlarged towards its inferior extremity (Pl. XLIV. fig. 1). That side

of the animal on which the ganglion lies will be denoted the dorsal. The animal is

viewed from the ventral surface in fig.1. The test is hyaline and extremely transparent.

* See a letter by G. J. Allman, F.R.S., in ‘ Nature,’ vol. xii. Oct. 28, 1875.

SECOND SERIES.—ZOOLOGY, VOL. I. 2 Q

288 MR. H. N. MOSELEY ON TWO NEW

Where it is simple it is thin and flexible, but in certain spots it is strengthened and

rendered stiff by the presence in it of rounded or plano-convex masses or plates of denser

tissue, which are tough and cartilaginous in consistence, and which are disposed over the

surface of the test in a nearly symmetrical manner. ‘These plates are extremely con-

spicuous when the test is held up to the light and viewed by transmitted light, because

they refract the light strongly; and the pattern formed by them on the test when

thus viewed has a very peculiar appearance. The disposition of the plates on the

ventral surface’ of the body will be seen from fig. 1, that on the dorsal surface from

fig. 2.

A series of globular lobes range on either lateral margin of the body, and give it here

considerable rigidity, and a ridge of slightly condensed tissue runs across the body at

the upper margin of its dorsal surface. A series of flattened plates is disposed over the

inferior region of the ventral aspect of the body, whilst the superior region of the same

aspect, covering the gill-sac and perforated by the inhalant aperture, is entirely devoid of

plates, very thin and flexible, and most perfectly transparent. On the dorsal aspect of

the body a very large plate occupies the middle line inferiorly, whilst immediately above

it two pairs of oblong plates (fig. 2,d) form a stiff shield for the principal viscera, which

lie upon them. ‘Two series of oval plates range on either side of the larger median ones,

and extend up as far as the exhalant aperture. The test-tunic is continued downwards

from the upper region of the body to form the outer wall of the cylindrical stem, thus

forming a tube. ‘The lower end of this tube is widened out into a funnel-shaped mouth,

and in the specimen had apparently been torn away from some object of attachment. -

The substance of the test is composed of transparent hyaline tissue, in which are

embedded the small bodies represented in fig. 4, the larger of which have a length of

from 007 to 014 millim. They are irregular in form, sometimes crystalline, or with

apparently crystalline contents. They are not sensibly altered in appearance by the

action of acetic acid, and no effervescence is produced in the test-tissue by that re-

agent. The bodies are present in the greatest abundance in the test-tissue at the base

of the stem. In the plates of denser tissue they are rather less abundant than else-

where. The test forms a simple sac continuous with the tubular cavity of the stem.

The exhalant orifice is an aperture in the test situate at the end of a short tube projecting

externally on the dorsal aspect just below the nerve-ganglion. Into it the ducts of the

generative glands and the rectum open. The inhalant aperture was entirely obliterated

in the only specimen obtained; it must have lain on the ventral aspect of the body,

since the dorsal wal! was intact. The arrangement of the muscular fibres and remnants

of attachment of the gill-sac seemed to indicate the position for it given in the figure,

where it is introduced conjecturally.

Closely attached to the inner surface of the test-wall is a delicate tunic (the mantle)

containing muscles. The muscles occur in the form of very fine bands, which have

a nearly parallel course. The series of bands springing from near the region occupied

by the heart follow the curved inner surface of the test-cavity towards its superior

margins. ‘The muscles are disposed most thickly in the lateral regions. The mesial

region of the dorsal surface is entirely devoid of them, but they extend over the whole

FORMS OF DEEP-SEA ASCIDIANS. 289

ventral wall. Only the lateral fibres are indicated in the figure; they are prolonged

superiorly in a horizontal direction along the upper margin of the dorsal wall of the

test-cavity. The tubular cavity of the stem is filled by a core composed of muscular

fibres embedded in gelatinous tissue, a prolongation of the mantle.

Of the gill-sac only a small portion remained intact in situ, but fragments here and

there attached indicated an arrangement as shown in the figure. The small portion

im situ lay over the nerve-ganglion. The fenestrations in the membrane are small,

simple, and irregular (fig. 3).

The mouth is situate nearly in the middle line (m, fig. 1). It leads by a short trans-

parent cesophagus to a stomach (s), which has opaque walls corrugated externally ; and

this viscus leads into a rectum which curves up to end at the exhalant aperture.

Beneath the stomach is a tubular heart with a wide vessel leading from it downwards

towards the stem.

In the loop formed by the cesophagus, stomach, and rectum is the large ovisac or

ovary, which is circular in outline. The elongate tubular testis running parallel to the

rectum in the middle line terminates posteriorly internally to the ovisac, its lower end

spreading out into a series of ramifications, which appear very like large nerve-fibres.

The testis-tube opens into the exhalant aperture close to the rectum, as does also the

oviduct, which lies to the dorsal side of the testis-tube and in the same line with it.

The ovisac was full of very large ova, measuring as much as 1°5 millim. in diameter

(fig. 5). These ova, on slight pressure being applied, were discharged from the oviduct.

They were found to consist of a very thick transparent test, with contents composed of

oily yelk-globules, without a germinal vesicle.

The testis was tumid, and full of an opaque white matter, which was discharged on

pressure in tenacious threads composed of spermatozoa.

The spermatozoa are very small. They were examined under a Hartnack No. 10

(immersion system). They consist of an elongate rod-like head, measuring ‘005 millim.

in length, and an excessively fine tail, the length of which could not be determined.

All the spermatozoa examined had a small transparent vesicle attached to one side of

their heads, as shown in the figures (fig. 6).

This Ascidian appears to be allied to Boltenia; but it is especially remarkable because

of the occurrence in its test of symmetrically arranged cartilaginous plates, which form

apattern. The name Hypobythius calycodes is proposed for it.

II. OcTACNEMUS BYTHIUS, gen. et spec. nov. ' (Plate XLIV. figs. 7-13.)

This stellate Ascidian was trawled March Ist, 1875, in 1070 fathoms, lat. 2° 33’S., long.

144° 04’ E., about 40 miles north of Rossy Island, Schouten Islands. From its peculiar

appearance, due to the presence of the eight long radiating conical processes of the test,

the animal was at first supposed to be a Medusa. The single specimen was considerably

injured, the muscular networks maintaining their attachments in only three of the conical

processes, but the test was entire.

The test of the animal is gelatinous and hyaline. On the under surface the body

presents a flat area of a nearly oval form (fig. 7). The border of this base is thickened

2Q2

290 MR. H. N. MOSELEY ON TWO NEW

into a slightly prominent, rounded ridge running round the periphery of the entire basal

area, and, further, is indented slightly opposite the interspaces between the long conical

processes, so as to have an undulating outline. Towards one end of the base (which

end of the animal will be termed anterior, since it is that in which the nerve-ganglion

lies), and in the middle line, is a prominence, also oval in outline (fig.7, pa). This pro-

minence is formed of a process of the basal part of the test. It terminates outwardly

in a tangled mass of rootlets, massed amongst which was found much sand and shell-

particles from the bottom. The Ascidian was evidently attached by this process or

pedicle.

Above the margin of the base the body of the animal is somewhat contracted, but its

walls then again spread outwards and extend into eight wide conical processes. The

processes terminate in abruptly narrowed tentacular-like tips (fig. 7, ¢, ¢), which are im-

perforate, and in which no sense-organ or any special structure could be discovered.

On the upper aspect of the body the eight conical processes are directly continuous

with the upper surface, which is somewhat hollowed or saucer-shaped.

Towards its anterior part in the middle line the upper surface is perforated by a trans-

verse slit, the inhalant aperture (fig. 8, 7a). The exhalant aperture is situated in front of

this, and at a lower level, proceeding from the wall of the body just above the base as

a short cylindrical tubular process (figs. 7 & 8, ea). The inhalant aperture is enclosed bya

pair of simple rounded lips, and is without tentacles.

The test forms a wide cavity, which extends freely into the capacious hollow conical

processes. The two apertures, inhalant and exhalant, form the only communication

between this cavity and the exterior.

A flat horizontal membrane is stretched across the test-cavity in such a manner as to

separate off an upper chamber communicating with the inhalant aperture from a lower

communicating with the exhalant. This membrane, in the central region of the body, is

thick and of an opaque white colour (fig. 9, 7). This thickened central portion runs out

peripherally into eight processes, directed to the intervals-between the long conical pro-

cesses of the test. Opposite these intervals the processes become attached, or give origin,

to bands of muscular fibres, which bands, after a short radial course, in which their

fibres remain parallel, split into two halves (fig. 7, 7m). The halves of the bands diverge

at an angle from one another, and proceed to the tips of the long conical processes, where

each is joined by the tip of a corresponding half-band from the next adjoining process.

The half-bands, as they run towards the tips of the conical processes, give off a series of

transverse muscular threads, which, passing from band to band, form a series of loops

one beyond the other, continued almost to the tip of the conical processes. These radial

muscles are apparently the homologues of the longitudinal muscles of ordinary Ascidians.

They would become longitudinal were the upper part of the discoid body of the animal

drawn upwards, so 4s to make the respiratory cavity tubular instead of saucer-shaped.

A second series of muscular threads lies beneath the radiating bands just described

about their points of bifurcation, and extending thence almost to their points of origin.

This second set of muscles takes a circular direction (fig. 7, em), and is continued round

the entire circuit of the animal, the several strands, of which there are about twelve,

FORMS OF DEEP-SEA ASCIDIANS. 291

appearing to be continuous throughout their length. These circular muscles lie beneath

the radial ones; and were the respiratory cavity elongated into a cylinder, the radial or

longitudinal muscles would thus be internal in position, the circular external. In order

to prevent confusion, and because of the difficulty of drawing them clearly, not nearly

the entire number of transverse and circular strands is inserted in the figures in the

accompanying Plate (figs. 7 & 8). Their arrangement is shown in detail in fig. 12.

Over the muscular meshwork thus formed, and extending from it to be continuous

in all directions with its thickened central portion, the horizontal membrane is continued

as a thin and transparent lamina. Opposite the indentations in the margin of the

thickened central portion of the membrane, 7. e. between the processes or thickened

folds attached to the muscular bands, this thin lamina is loose and hangs in bags or

depressions.

In the bottom of each of these depressions is a slight three-cornered elevation (fig. 8, a),

at the tip of which a perforation or aperture may possibly be present; but in the

specimen examined such could not be made out. At the tip of each small elevation were

seen only three minute closely apposed villous folds of the membrane, between which no

aperture could be detected. The central horizontal opaque white membrane is merely

a thickened part of the general membrane, which is spread over the muscular meshwork,

and reaches to the tips of the large conical processes. It is probably respiratory in

function, and represents a gill.

The membrane was observed to be attached to the inner surface of the test-wall at the

intervals between the conical processes ; but the specimen was too much injured to allow

of the investigation of the extent and manner of its attachment within the conical pro-

cesses. It appeared to be attached laterally on either side to the inner walls of these

processes, and is probably reflected so as to line their cavities. No normal perforated

gill could be discovered in any part of this membrane; but probably the central

thickened portion has a respiratory function, and possibly the eight small elevations may

prove to be openings. No reflection of the membrane over the inner surfaces of the upper

and lower walls of the test was observed.

The membrane was composed of an irregular mesh of fibrous tissue, with numerous

rounded gland-like cells, and numerous nerves proceeding to the radial muscular bands.

The mouth lies in the anterior part of the thickened portion of the membrane above

described, and at a short distance behind the inhalant aperture. Behind it is the endo-

style. The mouth isasimple oval aperture, encircled by a few sphincter muscular threads.

It leads directly into the digestive tract, which is embedded in a compact nucleiform mass,

which is seen conspicuously through the transparent test on viewing the animal from

beneath. The nucleus lies entirely beneath the horizontal membrane, which is attached

round the margin of themouth. ‘The exact arrangement of the viscera inside the nucleus

was not determined. A short tubular rectum projects from the anterior and inferior

extremity of the nucleus.

Posteriorly to this, on the inferior surface, is a well-developed ovary, and behind this,

again, the testis. The ovary is a racemose gland, whilst the testis is composed of short

cecal tubes. In minute structure these organs agree closely with the testes and ovaries

of other Ascidians.

292, MR. H. N. MOSELEY ON TWO NEW

Between the mouth and rectum, in the middle line, lies the heart-shaped nerve-

ganglion, immediately beneath which is a spherical body, which had the appearance of a

thin-walled sack full of a milky fluid. The wall of this sae is composed of a layer of

cells of rectangular outline, from the inner surface of which fine hair-like processes

depend (fig. 13). The hairs are not straight, but wavy, and appeared stiffer than

ordinary cilia. This sense-organ was not in sufliciently good preservation to allow of

more accurate histological investigation. A large nerve from the ganglion passed directly

over it in the middle line.

. A pair of strong muscular masses is developed at the posterior part of the nucleus,

one mass lying on either side. These muscles appear to take origin from the under-

side of the thickened horizontal membrane above, and to terminate on the sides of the

posterior part of the nucleus. The terminations of the muscular slips composing the

muscles are bifurcate (fig. 10, mm). Other narrow transverse muscular bands are present

between the mouth and rectum, embracing the fore part of the nucleus (fig. 10, m0).

The use or homologies of these muscles are not apparent.

The endostyle lies in the middle line, at the back of the nucleus, between the pair

of posterior muscles of the nucleus (fig. 10,2). It is very short. It showed the cha-

racteristic structure of the Ascidian endostyle—long, fine, granulated, spindle cells, packed

close side by side, with their long axes at right angles to the length of the organ.

The relation of the endostyle to the horizontal membrane was not observed, since the

organ was only discovered after the nucleus had become detached from the membrane.

The endostyle was then found tucked in between the paired posterior muscles of the

nucleus. It is placed in the position given to it in the figures, because this seems to be

the necessarily correct one. The organ, at all events, is on the ventral side of the

animal, or on the side of the mouth opposite to that on which the nerve-ganglion lies,

which is its normal seat.

The length of the base of the Ascidian was 5°5 centims., breadth 4°75 centims.; extreme

length between tips of the protuberances 7'5 centims.

On the whole this very perplexing animal appears to be an Ascidian in which the

respiratory sac is flattened out so as to become nearly horizontal, and in which no gill-

network is present. In Cystingia (Bronn, KI. und Ord. ii. p. 181) a gill-network cannot

be distinguished.

The radial muscles belong to the longitudinal set of other Ascidians and are internal ;

the circular are external in relative position. I can find no homologues of the muscles

of the nucleus. In having the viscera contracted into so small a nucleus, the animal

resembles Salpa. The nerve-ganglion is abnormal in position, in being situate on the

nucleus. It, nevertheless, is normal, in lying between mouth and anus, whilst the endo-

style is on the opposite side of the mouth as in other Ascidians.

The name Octacnemus bythius is proposed for this curious eight-rayed deep-sea form.

The above account is the result of an examination of the animal in the fresh condition.

A further investigation of the specimen as preserved in spirit is much to be desired.

FORMS OF DEEP-SEA ASCIDIANS. 293

DESCRIPTION OF PLATE XLIV.

Figs. 1-6 inclusive illustrate the structure of Hypobythius calycodes.

Fig. 1. The animal viewed from the ventral surface (i. e. the surface opposite to that on which the nerve-

ganglion lies). The viscera are seen through the transparent body-wall lying on the opposite

side of the cavity of the test. At the lateral margins the test swells into a series of lobes

composed of masses of compact gelatinous tissue, which lobes give a certain rigidity and

support to the test. Similar masses or plates of compact tissue are seen on the ventral

surface of the test in its lower region. In the single specimen obtained a longitudinal rent

divided the ventral wall of the test, and of the mantle and gill-saes, along the middle line down

to about the point indicated by the letter h.

This figure represents the animal of half the natural size, and the lettering applies as

follows :—

a, a. Plates of dense test-tisswe, similar to the other plates also seen, which are unlettered.

6. Muscular core of the stem of attachment of the animal. c. Prolongation of test forming the

outer tissue of the stem of attachment, here, at the end of the stem, widened out into a funnel-

shape. za. The inhalant aperture. No trace of this aperture remained in the specimen as

obtained ; its most probable position only is given. It may have lain somewhat more nearly

opposite the mouth on the ventral surface. ea. The exhalant aperture. ng. Nerve-ganglion

with sense-organ seen through the test and across its cavity. o. The ovary; od, the oviduct;

and ¢ the testis. gs. Margin of gill-sac attached by a series of fibres to the membrane lining

the interior of the test. Externally to the gill-sacs the directions of the lateral muscular bands

and their anterior prolongations along the dorsal margin are indicated. m. The mouth; s, the

stomach; and /, the heart. 6v. Large blood-vessel leading from the heart towards the stem.

Fig. 2. Sketch of the upper portion of the animal, much reduced in size, and as viewed from the dorsal

aspect, to show the symmetrical arrangement of the plates of thickened test-tissue.

ea. The exhalant aperture. d. Set of four plates covering the generative and digestive

organs. Beneath there is a large elongate oval plate.

Fig. 8. Portion of the gill-sac, to show its small and irregular fenestrations, which have a diameter of

about *10 millim.

. Bodies, partly crystalline or with crystallime contents, occurring in the tissue of the test.

. Ovum from the ovary; actual diameter 1°5 millim.

D> oO

. Spermatozoa, one of them viewed sideways, so as to show the attachment of the vesicle to the head.

Actual length of the heads of the spermatozoa 005 millim.

Figs. 7-18, inclusive, illustrate the structure of the Ascidian Octacnemus bythius,

trawled March 1, 1875, from 1070 fathoms.

Fig. 7. The animal viewed from below: natural size. The central part of the figure is occupied by the

flat irregularly oval base, with undulating outline and thickened margin (6). Beyond the

margin of the base the eight large conical processes are seen to radiate outwards, and terminate

in (¢) abruptly narrowed tentacle-like tips. The short pedicle of attachment (pa) projects

from the anterior part of the base ; the other parts (nucleus, respiratory membrane, and muscles)

are seen through the transparent wall of the test constituting the base. The nucleus is turned

over slightly to the left hand.

ea. Exhalant aperture. 6. Thickened border of the base. pa. Pedicle of attachment.

“7. Therectum. n. The nucleus. cm. Circular muscles. rm. Radiating muscles. g. Thickened

horizontal part of mantle-membrane.

294 MR. H. N. MOSELEY ON TWO NEW FORMS OF DEEP-SEA ASCIDIANS.

Fig. 8. The animal viewed from above ; also of the natural size. The long conical processes are seen as

in fig. 7, but their upper surfaces are flatter than their lower, and pass directly into the smooth,

somewhat hollowed, general upper surface of the animal.

This upper surface is perforated towards its anterior end by the transverse slit-like inhalant

aperture (¢a). Anteriorly part of the lateral wall and base are seen projecting beyond the out-

line formed by the union of the bases of the tentacles, and projecting from it the tubular exha-

lant aperture (ea). The mouth, endostyle, muscles, and membrane are seen through the

transparent upper part of the test. The rectum and nerve-ganglion are seen through the test

and horizontal membrane as well; they are somewhat directed to the left, being normally

in the median line. In both drawings the transverse muscular slips within the long. conical

processes are represented far too stout, and are not numerous enough. ‘These derivatives of

the radiating muscles are far less stout than the circular muscular slips.

za. Inhalant aperture. ea. The exhalant aperture. m. Themouth. e. Endostyle. r. Rec-

tum. ng. Nerve-ganglion. «a. Pyramidal eminences in the bottoms of the bag-like depressions

of the gill-membrane ; possibly apertures.

Fig. 9. Schematic, vertical, and longitudinal section, through the animal, along the middle line, z.e.

following the direction of the arrows in fig. 8.

ia. Inhalant aperture. ea. Exhalant aperture. m. Mouth. mn. Nucleus. mn. Muscles of

nucleus. 7. Rectum. g. Respiratory membrane. rm. Radiating muscles. ¢. Thickened

margin of the base. pd. Pedicle of attachment with its fibrous roots.

Fig. 10. The nucleus, enlarged. The horizontal membrane is attached round the mouth, which opens

above it, whilst the whole of the remainder of the nucleus lies below it. The cut edge of the

membrane (g*) is seen to pass between the nerve-ganglion and mouth, and (g) between the

posterior muscles of the nucleus and the endostyle.

m. Mouth with sphincter muscles. e. Endostyle, with membrane alongside. rm. Radiating

muscle (the median anterior band). g*. Edges of cut membrane. ng. Nerve-ganglion. so.

Sense-organ. m6. Transverse muscular bands on the anterior aspect of the nucleus. 7. Rec-

tum. mn. Posterior muscles of the nucleus. s. Stomach. ¢. Testis. 0. Ovary.

Fig. 11. The nerve-ganglion and sense-capsule, much enlarged.

ng. Nerve-ganglion. n,n. Nerves passing round the mouth, one on either side. m. Mouth.

so. Sense-capsule, across which runs a large nerve-stem, n*, proceeding from the inferior extre-

mity of the ganglion. The two filaments proceeding from the sense-capsule are of uncertain

nature.

Fig. 12. Muscular arrangement at one of the points of bifurcation of the radiating muscles, enlarged.

rm. Radiating muscles, appearing as a broad flat band towards their inner part, or part nearer

the centre of the discoid animal. The fibres composing the band commence at a, where their

pointed extremities are embedded in the thickened radial process of the thickened central part

of the horizontal membrane. Externally the band bifurcates, and the two halves of it, passing

into the long conical processes, give off thin transverse looping fibres on their outer side. cm.

The circular muscular fibres which lie behind the radiating ones. The inner ones curve inward,

the outer outward.

Fig. 13. Structure of the wall of the sense-capsule.

a. Part of the layer of cells composing the wall of the sense-capsule, seen in perspective.

b. Single cell, with its hairs or cilia.

TRANS. Lm. Soc Ser.Z. Zoot Vor.) Pi 44

HN Moseley. del?

OBerjean hth. Hanhart imp.

NEW DEEP-SEA ASCIDIANS

[ 295 ]

XV. On new Forms of Actiniaria dredged in the Deep Sea; with a Description of

certain Pelagic Surface-swimming Species. By H. N. Mosztny, WA., F.R.S.,

Fellow of Exeter College, Oxford, late Naturalist on board H.U.S.* Challenger,

(Communicated by Dr. J. Murine, £.L.8.)

(Plate XLV.)

Read February 15th, 1877.

THE deep-sea Actiniaria herewith to be described were obtained on various occa-

sions by the dredge or trawl during the voyage of H.M.S. ‘ Challenger.’ Actiniaria of a

variety of forms were obtained in deep water frequently during the Expedition and in

very various localities, and only a small part of these are accounted for in the present

paper. Several circumstances, such as rough weather, and the fact that, as frequently

was the case, the dredge or trawl was only got up to the surface at nightfall, combined

to prevent all the Actiniaria obtained being worked at in the fresh condition, and very

often, when other circumstances were favourable, some more important subject called

for immediate attention. A few of what appeared to be the most interesting forms were,

however, sketched and partly worked out whilst the animals were fresh, and the

results are here given. The pelagic Actinias ((/imyadine) were obtained either with

the surface-net or in the trawl.

Family ACTINTIAD &.

Subfamily Minyapinaz.

NAUTACTIS PURPUREUS, n. sp. (Plate XLV. figs. 1 & 2.)

A small floating Actinia was found amongst the contents of the towing-net on

August 26th, 1874, lat. 13° 39’ S., long. 150° 53’ H., between the New Hebrides and the

north-east coast of Australia. The animal was very small, not more then 7 millims. in

height, when measured with the parts about the mouth protruded. The body is cylin-

drical, with the aboral surface much flattened. The costz are slightly elevated rounded

ridges, 24.in number, nearly equal in size, and without tubercles. The tentacles are 12 in

number, disposed in two series, 6 primary larger ones lying nearer the mouth than the

6 somewhat smaller secondary ones; they are simple, very short indeed, reduced

in fact to rounded conical tubercles. Between the tentacles at the margin of the disk

are a series of small tubercles of triangular outline. The mouth is linear. The cosial

and aboral surfaces of the animal are of a chocolate colour with a tinge of lilac. Near

the margin of the disk is an irregular line of bright orange-yellow. The margin of the

mouth is of a dark burnt-sienna colour. From it radiate out streaks of chocolate

colouring, which fork to embrace the bases of the transparent white tentacles and the

intertentacular tubercles. These tubercles are of a glistening but opaque white, with

bright orange colouring on their lateral borders, and a dark spot in their centre.

SECOND SERIES.—ZOOLOGY, VOL. I. 2k

296 MR. H. N. MOSELEY ON DEEP-SEA ACTINIARIA

The animal on retracting the protruded parts of the mouth became of a simple cylin-

drical form with the disk concave.

The absence of tubercles on the costa, and their presence on the calicinal margin

only, and the extreme shortness of the tentacles, places this Minyad near the genus

Nautactis. It differs, however, from the Nawtactis olivacea*, as described by MM.

Milne-Edwards and Haime, in having 24 costs instead of 22, and in the nature of the

tentacles, which in J. olivacea are tuberculated, simple and very small towards the

centre, and multilobate towards the circumference of the disk.

Larva Minyap. (Plate XLV. fig. 3.)

On Nov. 2, 1874, off the Philippines, in lat. 12° 48’ N., long. 122° 10' B., a larval

Minyad was obtained with the towing-net from the surface.

This larva was about 1°5 millim. in height. The body was ciliated over its entire outer

surface. No tentacles were as yet formed in the larva, but the mouth was maintained

everted, and showed six distinctly marked lobes. Six primary mesenteries were well

developed, and six intermediate secondary just indicated. Upon the larva being

compressed, a single pair of mesenterial filaments was emitted from its mouth. The

filaments and the ectoderm were crowded with simple, elongate, ovoid nematocysts.

OcEANACTIS, gen. nov.

Characters.—Body transparent in the contracted condition, smooth and spherical in

shape; when expanded hemispherical, provided with rounded costal ridges and a single

row of costal tubercles. Tentacles simple, elongate, conical, and in two rows. Base

very small, entirely invisible in the contracted condition, with an aperture in the centre

communicating with the body-cavity.

- OCEANACTIS RHODACTYLUS, n. sp. (Plate XLV. fig. 4.)

On July 10th, 1874, the trawl used in 700 fathoms in lat. 37° 36’ S., long. 179° 24' E.,

about forty miles off New Zealand, came up full of a tenacious blue clay. In the clay

were found two Actinias in a contracted condition. When thus contracted they were

nearly spherical, with a tense and perfectly smooth transparent external tunic, through

which the mesenterial septa were plainly visible, as also the ovaries and the stomach-

wall. No trace of any base or basal aperture could be made out in this condition of the

animals. The oral orifice appeared as a small, slightly protuberant reddish cone. Hach

animal on the whole looked not unlike a small onion. Placed in fresh water for the

night the animals expanded to the form shown in the accompanying figure.

The body of the animal in the expanded condition is nearly hemispherical. The upper

part of the stomach is protruded as a semitransparent cone beyond the circle of ten-

tacles. There are twenty mesenterial septa, the spaces between which rise up as rounded

ridges or costee on the wall of the expanded body. ‘Ten of these bear, just above the

origin of the tentacles, rounded prominent tubercles.

* ¢ Histoire Naturelle des Coralliaires,’ Milne-Edwards et J. Haime, tome i. p. 230.

AND PELAGIC SURFACE-SWIMMING SPECIES. ; 297

The tentacles, 20 in number, are simple, elongate, conical in shape, and disposed in

two series. The tentacles of the outer series are the larger, and correspond to those

costze which bear tubercles. The inner are in connexion with the intermediate coste,

and arise from the edge of the disk just internally to the larger tentacles. At the aboral

pole of the animal is a small aperture, surrounded by a small circular area with tumid

folds representing the base. This aperture is in direct communication with the body-

cavity, for one of the ovaries became protruded through it without any violence having

been used. There is hence no separate chamber for the air-vesicle as in Minyas, but the

condition is like that in Peachia. The general transparent appearance of the animal,

however, and absence of base sufficient for attachment in an animal of this short hemi-

spherical shape, point to its being a free-floating and not a deep-sea form. Perhaps the

condition is here somewhat as in Nawtactis, where the base may be reversed, the air

extruded, and the body attached from time to time to floating bodies (M.-Hd. & H. /.c.

p- 230). The body of the animal is transparent and bluish coloured, with a slight

dusting of light red on the costee. The base is red. The costal tubercles are transparent

and marked with red, especially at their lines of origin from the costz. The tentacles

are of the same light red colour and somewhat transparent. The margin of the mouth,

which presents ten slight indentations, is also tinged with red. In the contracted con-

dition of the animal the ovaries show through the body-wall as yellow bodies near the

aboral pole. The stomach also is seen in the same manner as a transparent whitish

cylinder. The tentacles and tubercles are full of nematocysts of the common elongate

form, and about ‘04 to ‘05 millim. in length.

The animal apparently belongs to the group of the Minyadine (M.-Edwards & Hatate,

1. c. tome i. p. 227), but does not seem to come exactly within either of the three genera of

that subfamily, viz. Winyas, Plotactis, or Nautactis. Erom Nautactis it is distinguished

by the simplicity of its tentacles, from Plotactis by its single row of costal tubercles, and

from Minyas by not possessing a series of tubercles on each costa. It is further

distinguished from all three genera by the perforation of its base.

Subfamily ActTininaz.

ACTINIA ABYSSICOLA, sp.n. (Plate XLV. fie. 5.)

This animal, found adherent to the stem of a species of Mopsea, as depicted, is drawn

out and elongated in a direction transverse to the central axis of the body in relation

with the slender cylindrical form of its support.

The adherent surface of the base is extremely elongated, and of the same width

throughout, and is closely applied to the Mopsea-stem lapping round it, the region of

junction of the opposite edges being marked by a perfectly straight line. Along this

line of union the two edges are somewhat coalescent, but separable without the appli-

cation of much force. At either end of the elongated base the body swells out into

rounded masses, narrowed at the extremities, and evidently representing the fleshy

lobes present around the bases of normal Actinias, which are thrown into this form in

this instance by the extreme contraction and elongation of the attaching surface. Within

2R2

298 MR. H. N. MOSELEY ON DEEP-SEA ACTINIARIA

the rounded masses is a very stout and prominent muscular ring, pulled out into an

oval form, and somewhat contracted laterally; and within this is a smoother space

rounded from side to side, and with its inner margin bounding an oval opening,

immediately within which le the tentacles. This region of the body is coloured bright

pink, with a few radial streaks of slightly darker colour, and contrasts with the muscular

ridge and lobes external to it, which are of a pale pinkish yellow. The muscular ridge

is especially pale in colour. This ridge and the coloured space being immediately

internal to it, represent the vertical wall of the body of normal Actinias. The tentacles

are small, about 60 in number, disposed in two rows, and of a very pale pink colour.

The disk is dark pink in colour and of an oval form. The mouth very irregularly oval.

The greatest length of the largest specimen obtained was 34 millims. Another

specimen measured 23 millims.

Actinia abyssicola was obtained on two occasions from deep water, viz.:—On May 6th,

1873, in lat. 40° 17’ N., long. 66° 48’ W, from 1350 fathoms; and also on May 29th,

1878, to the south-west of Bermuda, from 1075 fathoms.

The two specimens procured on May 6th came up alive, but never expanded them-

selves more than as seen in the figure.

Very similar Actinias were dredged frequently from deep water attached to rod-like

bodies, such as Aleyonarian stems. I find in my notes, e.g., that one came up from

1400 fathoms, attached to a Hyalonema-spicule (lat. 12° 8’ S., long. 145° 10° H.), near

Raine Island, N.E. coast of Australia, August 29, 1874. I identified the species, how-

ever, only in the two above-cited instances.

ACTINIA GELATINOSA, n. sp. (Plate XLV. fig. 6.)

The body of the single specimen of the animal obtained was gelatinous and transparent.

The base was closed tightly around the stem of a dead Gorgonoid. The opposite

margins of the basal surface were approximated beneath this stem, and closely united

and coalesced along a linear suture. The basal surface thus formed a tightly fitting

sheath, in which the supporting Gorgonoid stem was held. The sheath and stem reached

only to a point opposite the mouth of the animal, the base having been drawn towards

the stem in forming its attachment, so that part of its margin here, instead of its central

point, lay beneath the mouth. Nearly one halfof the mass of the body thus lay in front

of the end of the stem of attachment. This free half was movable on the remainder of

the body, as if on a hinge (fig. 6,@); and as the specimen was turned about, it fell over

so as to shut over or enclasp the other firmly attached portion of the body, and thus

close the cavity of the animal as with a lid. The tentacles appeared to be numerous,

and disposed around the margin of the disk; but owing to their extreme softness were

too much injured to allow of their being counted. The body of the living animal was of

a transparent bluish colour. The tentacles were of a light brick-red colour, the mesen-

teries of a dark madder tint, being coloured with what I have elsewhere termed

“ polyperythrin’”’*.

* Consult H. N. Moseley, ‘‘ On the Colouring-matters of various Animals,” &c., Quart. Journ. of Microsc. Science

(new ser., January 1877), vol. xvii. p. 2. '

AND PELAGIC SURFACE-SWIMMING SPECIES. 299

It is possible that I may have been deceived as to the folding over of the body, for it

may be noted that the animal was injured by the dredge; but it appeared to me that the

hinge-like folding place was natural, and not artificially produced.

This animal was dredged between the Banda group and Amboyna, in lat. 4° 21’ S.,

long. 129° 07’ E., on Oct. 3, 1874.

EDWARDSIA CORIACEA, sp. n. (Plate XLV. figs. 11-14.)

The body is cylindrical, and when fully extended of nearly equal: diameter throughout,

and worm-like in appearance and movement. When the animal is at rest with the

tentacles fully expanded, as shown in fig. 11, the posterior portion of the body is dis-

tended and somewhat ovoidal. The entire median region of the body is covered with a

dense coriaceous test, thickly beset with grains of sand, shells of Foraminifera, &c., the

whole forming a tough brown tube, into which the anterior region of the body is

completely retractile. In the contracted condition the animal is pointed at the anterior

extremity, whilst the posterior thin-skinned region is distended and pushed out into

the form of a rounded disk (fig. 12). The animal thus in the act of retraction of its

anterior extremity must obtain firmer hold in the sand by distension of the posterior

part. The base is imperforate and of a pinkish vioiet colour. The general colouring

of the anterior retractile region of the body and disk is a pale pinkish white. The ten-

tacles are 16 in number, disposed in two series, of which the external are most brightly

coloured. They are marked by equidistant transverse bands of reddish brown, the outer-

most placed on the tips. On the disk are yellow streaks between the bases of the ten-

tacles, and on the light external tentacles reddish-brown streaks placed internally to

these on either side. The oral protuberance is uncoloured. The remainder of the

anterior retractile region of the body, for a short interval beneath the tentacles, is un-

coloured. Lower down it has short longitudinal streaks of reddish brown, corresponding

to the tentacles in position, the streaks corresponding to the external row of tentacles

being broader than those opposite the intermediate ones. The streaks are absent

opposite four of the tentacles of the external row. Thus, passing round the body, are

seen a colourless space, then three coloured streaks, then another colourless space, and

so on (fig. 14). The length of the specimen as obtained, when at rest and contracted

(fig. 12), was about 275 centims.; when fully extended, with the tentacles expanded,

about 4 centims. JZ. coriacea is allied to H. beautempsis (M.-Hd. & Haime, Hist. Nat.

des Coralliaires, tom. i. p. 285) in having only 16 tentacles, but differs from it in the

fact that the tentacles are disposed in two series, in the details of colouring, and in the

structure of the test.

This Actiniad was procured with the dredge from 600 fathoms off Cape St. Vincent, on

January 16th, 1873. It was closely contracted when found in the dredge, but expanded

on being placed in fresh sea-water, and lived several days.

CoRALLIMORPHUS, gen. nov.

Characters—Body rigid, smooth, gelatinous, not contractile, without pores, but with

an adherent base; disk circular and large; tentacles non-retractile, elongate, conical,

300 MR. H. N. MOSELEY ON DEEP-SEA ACTINIARIA

with a rounded terminal knob, of several sizes, disposed in regular series at the margin of

the disk and in two circlets on its surface.

CoRALLIMORPHUS PROFUNDUS, n. sp. (Plate XLV. figs. 7, 8, & 8a, 80.)

The animal is in form a short cylinder, with flat upper surface or disk, and is rounded

below, where the base takes the form of an excavation, its margins being turned inwards.

Attached to the centre of the disk, but occupying not nearly its whole area, was, in the

single specimen obtained, a small nodule of peroxide of manganese, of which many were

brought up by the trawl. The wall of the body of the Actinia is firm and rigid, and the

animal is unable to contract so as to shut in the disk and tentacles, 60 tentacles are

present, of which 48 are attached to the margin of the disk, and 12 spring from its

surface. The marginal tentacles are of three sizes—24 being very small, 12 much larger,

and 12 intermediate in dimensions. The disk, which has a diameter of about 4 centims.,

shows a series of radially disposed rounded ridges, corresponding to the summits of the

intermesenterial chambers. Of those corresponding to the 12 larger tentacles, 6 are

broader than the others, and terminate towards the mouth in broader ends; these may

be considered as primary. A pair of them coincide in direction with the long axis of the

elongate mouth. These primary rays of the disk bear each a tentacle at a distance from |

the margin of the disk of about one third their length. The 6 secondary rays bear

smalier tentacles, which are placed much nearer to the margin of the disk. The tentacles

are elongate conical, with a small knob at their extremities. The mouth is of an elongate

oval form, and is sunk in a slight eminence in the centre of the disk. The animal is

covered with a hyaline transparent integument, through which the ochre-yellow coloured

inner tunics and viscera show themselves. The ovaries are especially conspicuous,

being of a dark ochre-yellow, of which colour also is the mouth. ‘The base is of a dark

brown.

The animal has three kinds of nematocysts or thread-cells. One of these is exactly of

the same structure as figured by Mébius as occurring in Caryophyllia Smithii*, being

extremely large and of an elongate cylindrical form, with narrowed ends. These cells in

the present species are remarkable for their great size, being apparently the largest

nematocysts known. They measure about ‘18 millim. in length, whereas the cells of

Caryophyllia Smithit measure only -102 to ‘071 millim. They are thus specially favour-

able objects for examination, and I clearly saw in them the peculiar triple invagination of

the wide first part of the thread figured by Mobius; and I also saw the various stages of

development of the cells exactly as figured by him.

The second kind of nematocysts is of nearly the same form as that just described, but

a little shorter. It contains a thread which is of uniform dimension, and beset through-

out its length with short spines: the thread is simply coiled in a somewhat irregular spiral

within the cell. These two kinds of thread-cells occur together in the tentacles of the

animal. The third form of nematocysts is shown (figs. 8 a, 8 0) as seen in the expanded and

* Mobius, “ Ueber den Bau, den Mechanismus und die Entwicklung der Nesselkapseln,” Abh. nat. Ver. Hamburg,

1866. (Cited by Allman, figures copied by Dana.)

AND PELAGIC SURFACE-SWIMMING SPECIES. 301

unexpanded conditions. These cells are smaller than the others, being of about one fourth

the length of the larger ones. They are of an ovoid form, but slightly bent. They have

a thread which, when protruded, has a very wide, fusiform commencement, round which

is wound a spiral line, bearing spines, the succeeding fine portion of the thread-being

unarmed. In the contracted condition a triflmg invagination takes place in the first

broad part of the thread, as in the largest cells; and the spines are of course reversed in

inclination, being turned backwards towards the base of the cell. These cells seem to

form an interesting transition towards the peculiar three-spined thread-cells of Hydroids ;

they occur on the base of the animal.

A single specimen of this interesting form was dredged in the South Pacific Ocean in

lat. 39° 4 S., long. 105° 5’ W., on Noy. Ist, 1875, from 2025 fathoms. A closely similar

specimen was likewise obtained which differed only in having darker, madder-coloured,

radial streaks on the disk, and in being much larger, measuring 10°5 centim. in diameter

of the disk. It was abnormally irregular in having 52 instead of 48 marginal tentacles,

caused apparently by the intercalation of an additional half-system of 4 tentacles; but it

may evidently be referred to the same species. It was dredged, together with a large

number of other Actininze, in lat. 33° 42’ S., long. 78° 18’ W., on Dee. 17th, 1875, from

1375 fathoms.

It seems necessary to refer the above-described form to a new genus. It resembles

the genus Discosoma* in being rigid and unable to contract the disk and cover the

tentacles, but differs from it in that the tentacles are here not very numerous, and that

they are provided with a knob at the tip, as in Corynactis. Similar or kindred forms

were often dredged in deep water by the ‘Challenger.’ They seem closely allied to the

simple disk-shaped corals, such as Stephanophyllia, in which the tentacles are also

knobbed and disposed at the margin, and on the surface of the disk, in a closely similar

manner. They lave further similar thread-cells to these solitary corals, and, indeed, ap-

pear to differ from them only in having no calcareous corallum developed. Indeed, in one

species of Stephanophyllia, obtained in deep water, the calcareous skeleton was in a most

rudimentary condition, being a mere delicate network of calcareous trabecule. So like

in appearance were these forms of Actininze to the living corals, that it was only by

feeling them as they were found in the dredge that I became convinced that they had no

calcareous skeleton, and I macerated several in caustic potash in the hopes of finding

traces of a corallum present in them. They seem to approach these disk-like corals very

closely, and possibly to represent forms from which the corals were developed.

CoRALLIMORPHUS RIGIDUS, n. sp. (Plate XLV. figs. 9 & 10.)

A form allied to that just described, and evidently to be referred to the same genus,

was obtained amongst the Moluccas in 1425 fathoms. The body of the animal is some-

what cartilaginous, tough and firm, maintaining its shape, and not contractile. Its form

is that of a truncate cone, the base being of less than half the diameter of the oral disk.

The lateral walls of the body present smooth, slightly projecting, rounded ridges or costee,

* Vide MM. Edwards & Haime, J. c. tome i. p. 255.

302 MR. H. N. MOSELEY ON DEEP-SEA ACTINIARIA

corresponding in position to the intervals between the attachments of the mesenteries.

The base is hollowed out into a cavity (fig. 10), which cavity had probably been occupied,

as in C. profundus, by some object to which the animal had been attached. The cavity

is surrounded laterally by a thin but firm wall, formed of the obliquely projecting wall

of the body, the margins of the cavity being thus somewhat undercut, so as to afford

increased holding power. The bottom of the cavity, or actual base, is smooth and

imperforate. The oral disk is elevated and rounded towards the centre. The tentacles

are 72 in number: of these, 12 larger or primary are disposed at equal intervals along

the margin of the disk, with 12 secondary and 24 still smaller tertiary ones placed

symmetrically between them. On the surface of the disk itself, at about half the distance

between the margin of the mouth and that of the disk, is a circle of 12 tentacles, placed

opposite the primaries, and intermediate in size between these latter and the secondaries.

Halfway, again, between the line of origin of this inner circle of tentacles and the disk

margin is a further circle of 12 smaller tentacles, placed opposite the secondaries. The

tentacles are elongate-conical in form, tipped by very distinct knobs. The bases of the

tentacles are very firm and cartilaginous. The general colour of the animal is a very

pale pinkish yellow. The disk has a dark madder colouring * in parts, and rings of the

same colouring girdled the tentacles just beneath their knobs. The single specimen

obtained was much rubbed, and hence the exact distribution of the pigment was

uncertain.

The specimen was dredged between the Banda group and Amboyna, in lat. 6 21’ S.,

long. 129° 7’ H., from 1425 fathoms, on Oct. 3rd, 1874.

Family CERIANTHID&.

CERIANTHUS BATHYMETRICUS, sp. nov. (Plate XLV. figs. 15-20.)

This species is contained within a tube formed of the felted threads of its large

nematocysts, as is the case with the other species of the genus. The tube is 11 centims.

long, and about 14 centim. broad in the broadest part. It is represented of the natural

size in the figure (fig. 15). ‘The opening at the upper extremity is in the form of a longi-

tudinal slit. The tube is beset with foraminiferous shells. Embedded in its substance

were found spicules of a Hexactinellid sponge, and also amongst its fibres were nume-

rous small nematoid worms (fig. 20). The animal contained within the tube is about

91 centims. long. There is an opening at the apex of the conical posterior extremity of

the body as in other species of the genus. The tentacles are disposed in two series. ‘Those

of the outer series are 5 millims. in length, and are, in the one perfect specimen available

for examination, 28 in number. Those of the inner series are about 1 millim. in length

and 14 in number. The upper part of the digestive cavity corresponds exactly to that

figured in Bronn’s ‘ Klassen und Ordnungen des Theirreichs,’ 2te Bd. Taf. vii. fig. 4,

after Haime, from Cerianthus membranaceus, and termed ‘“‘ Magensack;’’ but the digestive

* The madder colouring on both species of Corallimorphus here described was due to the substance 1 have termed

polyperythrin (Quart. Journ. Micros. Science, J. ¢.).

AND PELAGIC SURFACE-SWIMMING SPECIES. 303

cavity in C. bathymetricus does not terminate, as in C. membranaceus, above the line of

commencement of the mesenteries, but is continued downwards for some distance, its

walls consisting of a thin and delicate membrane, supported by the mesenteries and

having an irregular lower border. This lower sac is evidently in this case the true

stomach, since it was filled in the specimen examined with food, consisting of Globigerina-

shells together with small Crustacea. The main part of the mesenteries are not con-

tinued to the apex of the body, but only extend down about 3 millims., and from these

mesenteries are prolonged the ovaries, beyond the terminations of which is a clear space

of about 7 millims. in length. There is no canal formed of the opposed mesenteries and

leading from the mouth to the posterior part of the body-cavity ; but this is represented by

three or four stout muscular threads derived from the mesenteries, which are attached

to the inner surface of the body-wall near the posterior opening. These muscular

threads are branched superiorly, and thus each is derived from two or more mesenteries.

The animal was not examined until it had been put with its tube in spirit, under the

belief that it was an annelid, and its colouring, if it had any, was therefore destroyed.

Only two kinds of nematocysts were found in it, EAE portions of the epidermis,

the tentacles, and the test were examined.

The large, characteristic Cerianthus thread-cells, which form the main bulk of the

animal’s tube, resemble those of C. membranaceus in all respects except that the thread

is wound in an even spiral around the periphery of the internal cavity of the cell, the

axis of the spiral being at right angles to the longer diameter of the cell. In C. mem-

branaceus the thread is, as figured in Bronn (J. c. Taf. 1. fig. 7), irregularly doubled up

inside the cell. These cells are in C. bathymetricus ‘06 millim. long by ‘03 millim. broad.

The other kind of thread-cell observed was of the common elongated ovoidal form,

with the thread wound in a spiral about the longer axis of the cell (fig. 5). The

largest of these cells are of the same length as those forming the bulk of the test, but

they are very much more slender. They are not very abundant in the test, but occur

in large numbers on the tentacles. Thread-cells of all sizes and stages of development

are to be found in various regions of the body, but all of them are referable to these two

forms.

C. bathymetricus differs from the hitherto described species in the number of tentacles

arrangement of the digestive cavity, absence of a canal formed by the mesenteries,

the form of the thread-cells, and in its very small size. Two specimens were obtained

on June 19th 1873, in 2750 fathoms, in lat. 35° 29' N., long. 50° 53’ W., between the

Bermudas and the Azores.

GENERAL REMARKS.

The occurrence in the deep sea of representatives of well-known shallow-water forms

of Actiniaria is of considerable interest. The Hdwardsia here described seems to have

undergone no modification of any moment in a descent to 800 fathoms. The Ceri-

anthus, which is from the great depth of 2750 fathoms, is much dwarfed, and differs

in some details of its anatomy from the shallow-water forms; but it is astonishing

that the same genus should be able to adapt itself to shallow tropical seas and the ice-

SECOND SERIES.—ZOOLOGY, VOL. I. 28

304 MR. H. N. MOSELEY ON DEEP-SEA ACTINIARIA

cold water of the deep sea. A Cerianthus occurs at the Island of Mactan, opposite Zebu,

in the Philippines, which has a tube 1 foot 4 inches long, the animal itself being

6 inches in length. I found this Cerianthus abundant in water only 1 foot in depth at low

tide about the reefs, with its tentacles expanded, in the full glare of the tropical sun.

Yet a species of the same genus can exist at a depth of three miles, where no solar

light exists, and in water at a temperature close to freezing-point. .

The occurrence of a Nematoid worm in abundance in the substance of the tube of the

Cerianthus at this great depth is a very striking fact. The fact that some of the deep-

sea Actinias still retain a vivid colouring in the dark depths is also of great interest and

importance. I have dwelt on such facts and on the colouring-matters of deep-sea

animals generally elsewhere *. The usual absence of rocks or other objects offering a

wide surface for attachment seems to have led to the development of the peculiar elon-

gated base in Actinia abyssicola and A. gelatinosa, adapted to adhere to dead Alcyo-

narian stems, and to the hollowing out of the base of Corallimorphus for attachment to

small bodies, such as manganese nodules.

The genus Corallimorphus is of especial value, as appearing to be the nearest ally

amongst Actiniaria of many simple discoid corals, and for the excessively large size of

one form of its thread-cells or nematocysts ; these nematocysts seeming to be the largest

hitherto observed in any animal. There can be little doubt that Oceanactis is a pelagic,

and not a deep-sea form. Pelagic animals, such as Pyrosoma, Meduse, &c., were con-

stantly found in the trawl, caught by it on its way up or down, and mixed up with the

deep-sea animals from the botom. Pelagonemertes was obtained in this way.

DESCRIPTION OF PLATE XLV.

Fig. 1. Nautactis purpureus: side view, magnified. Actual height of the animal 7 millims.

Fig. 2. Enlarged view of portion of the disk of the same, to show arrangement of the colouring.

Fig. 3. Larval Minyad from the surface-net at the Philippines. Enlarged 15 diameters. Actual height

1:5 millim.

Fig. 4. Oceanactis rhododactylus. Twice natural size.

Fig. 5. Actinia abyssicola, attached to a Mopsia stem. Twice natural size.

Fig. 6. Actinia gelatinosa. The animal is shown of half the natural size. The line of continuation of

the rod to which the Actinia was attached within its sheath is shown by dotted lines: a, spot at

which the body doubled upon itself.

Fig. 7. Corallimorphus profundus. Side view, showing the attachment of the base to a small manganese

nodule.

Fig. 8. The same. Direct view from above of the disk. Figs. 7 and 8 both of natural size.

Fig. 8a. Nematocyst of same in quiescent, and 84 in active condition. x 510. Actual length of

cell 044 millim.

Fig. 9. Corallimorphus rigidus. Diagram of the disk, to show the arrangement of the tentacles and

their proportionate sizes.

* «On the Colouring-matters of various Animals, and especially of Deep-sea forms dredged by H.M.S. ‘Chal-

lenger.’” By H. N. Moseley. (Quart. Journ. of Microsc. Science, vol. xvii. p. 1, 1877.)

Fig.

AND PELAGIC SURFACE-SWIMMING SPECIES. 305

. 10. The same, seen in vertical section, to show the form of the oral disk, the mode of origin of the

tentacles, and peculiar cavity in the base. Figs. 9 and 10 both half natural size.

g. 11. Edwardsia coriacea, at rest, with the tentacles expanded. Twice the natural size.

. 12. The same, when completely contracted.

. 13. Portion of the disk, with three tentacles enlarged, to show the colouring.

. 14, Arrangement of the coloured streaks on the body-wall of the same, immediately beneath the

edge of the disk.

. 15. Cerianthus bathymetricus. Tube of the animal viewed intact, of natural size.

. 16. The same, tube laid open, to show the animal én situ.

. 17. The same, animal laid open and dissected. T, Outer tentacles; ¢, ner tentacles; or, mouth

and plaited upper region of the digestive tube; s, stomach, or lower portion of the digestive

tube, in which were found the remains of a crustacean ; m, mesenteries, from which proceed in

one region three muscular threads, which are prolonged to the apex of the body near the

posterior opening, where they are attached; g, generative glands.

18. Large nematocyst of same, and of the kind of which the tube or test is mainly composed.

19. Second form of nematocyst of same.

20. Small Nematoid parasite occurring in the wall of the tube.

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TRANSACTIONS

THE LINNEAN SOCIETY

LONDON.

SECOND SERIES.—ZOOLOGY.

VOLUME I.

PART THE SIXTH.

LONDON:

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET:

SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON-HOUSE;

AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER-ROW.

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CONTENTS.

PART VI.—NovemBeER, 1877.

J v1. On the Birds collected by Professor J. B. Steere in the Philippine Archipelago.

By R. Bowpter Swarrt, F.L.S., 7.Z.8., &c., Senior Assistant, Zoological Depart-

ment, British Museum. (Plates XLVI-LIV.).... .. . . page 307

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[ 307 ]

XVI.—On the Birds collected by Professor J. B. Steere in the Philippine Archipelago.

By R. Bowpier Sarre, £.L.8., F.Z.S., §c., Senior Assistant, Zoological Depart-

ment, British Musewn.

(Plates XLVI.-LIV.)

Read November 16th, 1876.

ON his rettirn from the Philippines, Prof. Steere passed through London in the autumn

of 1875, and delighted me with an account of his travels in the Hast. At that time his

collections were packed up, and it was not possible to examine them ; but this year (1876)

he returned to England, and was so good as to place his birds in my hands for description.

This would have been more difficult to accomplish had it not been for the very elaborate

memoir published in the spring of last year by the Marquis of Tweeddale (then Viscount

Walden). Previously to this the ornithology of the Philippines was not concisely

described; and I must record my great obligations to his memoir on the birds of the

Philippine Archipelago. I must further acknowledge Lord Tweeddale’s kindness

in allowing me free access to his fine collection of Philippine birds, thus enabling me

to make comparisons of many rare and difficult species. He has seen all the specimens

obtained by Dr. Steere, and agrees with me as to the correctness of my determinations.

Knowing that Dr. Steere had visited several islands hitherto untrodden by the

naturalist, I was prepared to see several new species; but I was indeed surprised at the

large number of novelties which he has brought home with him. Forty new species

were discovered by him; and there is no doubt that even this number would have been

inereased but for the severe attacks of a fever which he contracted in Balabaec. As it is,

however, the results are most considerable, and Dr. Steere will receive the hearty

applause of all naturalists for the vigorous way in which he combated the difficulties of

climate and personal danger in pursuit of science, and reaped such an abundant harvest

in face of these trials. The danger from pirates alone may be estimated from the fact

that he collected many of his specimens in the company of native hunters and their

dogs—the latter being indispensable companions on an expedition, to give warnings

of the approach of piratical Malays. Again, in Basilan it was impossible even to

bathe under the shelter of the fort without keeping firearms at hand for personal

defence.

The following sketch of his expedition is extracted from an article communi-

cated by me to ‘Nature’ of the 3rd of August, 1876 :—‘ Leaving Hongkong for

Manila, in May 1874, Dr. Steere crossed the island of Luzon by way of Mauban and

Lucban to the Pacific, passmg some time on the mountain of Ma-hay-hay, near the

Laguna de Bay. In July he went by steamer to the colony of Puerto Princesa, on the

east side of the island of Palawan, where he stayed a month. Thence he crossed to

the island of Balabac and remained a month, afterwards visiting the south-east corner

of the island of Mindanao and resting for a month and a half at Zamboanga and the

SECOND SERIES, ZOOLOGY, VOL. I. 27

308 MR. R. B. SHARPE ON BIRDS COLLECTED BY

Indian village of Dumalon in the same province. The island of Basilan, lying between

Mindanao and the Sooloo group, was next visited; and here he stayed two weeks, after

which he returned to Zamboanga, and thence to Manila. In the month of December he

again went south, stopping at Tlo Ilo, on the island of Panay, and visiting the mountains

in the interior. After a short stay at the neighbouring island of Guimaras he crossed

over to Negros, journeying on horseback round the north end of the island; thence in a

native boat he traversed the sea to Zebu, which he crossed, till he arrived at the town of

the same name, where he took horse again and rode southward, crossing the island once

more and passing over the strait to the town of Dumaguete, on the island of Negros.

Dr. Steere then went back to Zebu and crossed to the island of Bohol ; after passing

round part of this island he returned to Zebu and afterwards to Manila, where he

visited the Negritos on the north side of the Bay of Manila, leaving finally in April for

Singapore.”

In the preparation of this paper I have followed, as far as possible, Lord Tweeddale’s

memoir, in order to make it supplementary to the latter; and, before commencing the

detailed account of the species, it may be as well to compare the results obtained by

Dr. Steere with those recorded by Lord Tweeddale in 1875.

The number of birds included by the last-named author was 219. From this total

I am only aware of one species that should be deducted, viz. Circus wruginosus. For

this error I myself am responsible, as I recorded in the ‘Catalogue of Birds’ a young

Marsh-Harrier, which I have since come to the conclusion is only an immature Circus

spilonotus (cf. Sharpe, ‘ Ibis,’ 1876, p. 30). Dr. Briiggemann has added a Pitta from

Luzon, in the Darmstadt Museum. :

The result of Dr. Steere’s expedition has been to add more than sixty species to the

avifauna of the Philippines; and by restoring to its position as a species Dicewm papuense

from Mindanao, and by adding the few birds known from the Sulu Islands, we have now

atotal of 286 Philippine species. The following remarks give some idea of the actual

alterations made in our knowledge of the different islands.

PHILIPPINE ISLANDS.

In the above-mentioned paper of Lord Tweeddale’s no less than 57 species are cata-

logued as being of Philippine origin, without any exact locality being known for any of

them beyond the above vague habitat. I am able to reduce this number by eight, as

follows :—

. Circus eruginosus is C. spilonotus, juv.

. Brachyurus erythrogaster is from Luzon (Mus. J. Gould).

. Zeocephus rufus is from Luzon (Mus. Brit.).

. Padda oryzivora is from Mindanao (Steere).

. Sturnia violacea is from Negros (Séeere).

. Calenas nicobarica is from Mindanao (Steere).

. Ortygometra cinerea is from Zebu (Steere).

. Gorsachius melanolophus is from Mindanao (Steere).

OTe oP CO DS

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 309

Many of the remaining 49 species are doubtfully Philippine, and I believe that a con-

siderable reduction will be ultimately made.

Inzon.—Dr. Steere remained so short a time in this island, and then only collected in

the neighbourhood of Manila, a locality already well worked, that he made no additions

to our knowledge of the ornithology of Luzon, the few birds obtained by him being

all well-known residents. Lord Tweeddale notices 133 species from Luzon. From this

total Totanus calidris must be deducted as being inserted in error, as it already appears

in the “ Philippine” column ; but, one species being omitted in counting up the total,

there remain still 133 species known from Luzon. Cataguan is treated by Lord Tweed-

dale as a separate locality; but it should really be considered as part of Luzon, as it is

situated on one of the irregular peninsulas of the south-eastern corner of the island.

This will enable us to add 8 species collected by Cuming at Cataguan, and now in the

British Museum, viz. Penelopides panini, Zeocephus rufus, and Puffinus leucomelas.

In addition to this, Mr. Gould’s collection contains the Red-breasted Pitta (Brachyurus

erythrogaster), from Manila, which can therefore be removed from the “ Philippine”

column to the Luzon list. The Pitta recently described by Dr. Briggemann as P. kochi

(Abhandl. Bremen, v. p. 65, pl. ii. fig. 6) makes the present number of species known

from Luzon 138.

Panay.—The number of species included by Lord Tweeddale was 9; and to these

Dr. Steere has added 22, making a present total of 31 species known from this island.

Of the acquisitions fowr are new, viz. Craniorrhinus Waldeni, Humyias panayensis,

Xantholestes panayensis, and Dicewm dorsale. The others are chiefly well-known

Philippine forms occurring to the notice of naturalists for the first time in Panay.

Guimaras.—To the 27 species collected by Mr. L. C. Layard and Dr. Meyer in this

island Dr. Steere has only added 4, viz. Philentoma albiventris, Dendrophila enochlamys,

Diceum hematostictum, and Cinnyris speratus. Of these, all but the last are new to

science. The total number of species now recorded from Guimaras is therefore 31.

Negros.—To the 38 species of birds recorded from this island by Lord Tweeddale,

Dr. Steere adds 21, the following being new species :—Oriolus Steerit, dithopyga mag-

nifica, Anthreptes chlorigaster, Phabotreron nigrorwm, the latter being the P. leucotis

of Lord Tweeddale’s list. The total number of species noticed from Negros is now 59.

Zebu.—Highteen species are enumerated by Lord Tweeddale; and: Dr. Steere only adds

one, Ortygometra cinerea, a well-known bird from the Malayan islands. It was stated

to occur in the Philippine archipelago; but hitherto no particular island had been

assigned as its exact habitat.

Camiguin.—This island was visited by Mr. Cuming; but I cannot say whether his

collections of birds made here were large, as, unfortunately, the bulk of his ornithological

consignments were dispersed without a complete record of their contents having been

made. I have discovered, however, two species from the present island in the Museum,

viz. Craniorrhinus leucocephalus and Brachyurus sordidus.

Mindanao.—Dr. Steere obtained 28 species in this island, in addition to the 19

included by Lord Tweeddale; to the latter should also be added Philentoma cyaniceps,

which is in the British Museum from Mindanao. The new species obtained by Dr. Steere

27 2

310 . MR. R. B. SHARPE ON BIRDS COLLECTED BY

were Brachywrus propinguus and B. Steer. This large Pitta, discovered near Zamboanga,

is one of the most interesting of Dr. Steere’s discoveries.

Basilan.—This island lies to the south of Zamboanga, and has apparently never before

been visited by a naturalist. 23 species were obtained by Dr. Steere; and of these

the following were new:—Chrysocolaptes maculiceps, Halcyon Winchell, Hypothymis

superciliaris, Setaria ruficauda, Zeocephus cinnamomeus, Macronus striaticeps, Dendro-

biastes basilanica, Irena melanochlamys, Hypsipetes rufigularis, Orthotomus cinereiceps,

O.frontalis, Diceum hypoleucum, Kudrepanis pulcherrima, Sarcophanops Steer. Con-

sidering that all the collecting done on this island was accomplished within a short

distance of the Spanish fort and settlement, it may be confidently hoped that a good deal

still remains to be discovered in Basilan.

Malamauwi.—Dr. Steere informs me that this little island is really nothing but a con-

tinuation of Basilan, from which it is separated only by a narrow channel about 100

yards across. This strait, however, is very deep, allowing a large ship to pass in; and it

is a favourite halting-place for the Spanish gun-boats. The distance between these two

islands is so small that doubtless the same species of birds are common to the two.

Malamaui is a low flat island, but very heavily wooded. Of the 8 species obtained by

Dr. Steere, only one, Swrniculus velutinus, did not occur in any other of the Philippine

Islands ; but three of the new species discovered on Basilan were also found on Malamaui.

Cujo.—Dr. Steere did not collect any birds here; so that at present we know only of

the 5 species recorded by Lord Tweeddale.

Palawan.—On this island Dr. Steere broke new ground; and of the 31 species obtained

by him the following were new :—Chrysocolaptes erythrocephalus, Zeocephus cyanescens,

Nixornis Woodi, Ptilocichla falcata, Phyllornis palawanensis, Irena Tweeddalii, Criniger

frater, Cittocincla nigra, Arachnothera dilutior, Aithopyga Shelleyi. Of the remaining

21 species recorded, 13 at least are well-known Bornean and Malayan forms, showing

the strong Bornean element in the ornithology of the island, as might almost have

been expected from its proximity to Borneo. At the same time the presence of

such birds as Tanygnathus luzoniensis, Leucocerca nigritorquis, and Brachywrus sordidus

(not B. Muelleri), shows great affinity to the true Philippine avifauna. We must wait

therefore for thorough exploration of the island before we can determine for certain its

zoological affinities ; but there can be little doubt that it holds an intermediate position

between Borneo and the other islands of the Philippine archipelago. !

- Balabac.—Dr. Steere was so prostrated with fever during his residence here that he

was unable to do as much work as he would have done had his health been good. He

met with only 9 species, two of which, Dryococcyx Harringtoni and Parus amabilis, were

new, and not found elsewhere. Most of the remaining ones are known from Borneo or

from the other islands of the Philippine group.

Sulu Islands—Myr. Hugh Low has been so good as to give me some birds which he

had obtained from Si Butu ; and although they consisted of only two species, they clearly

show the nature of the Sulu avifauna, and its close relations to that of the Philippines.

One of them is an apparently new species of Sarcops; and the other is a new Oriole

(Oriolus suluensis, Sharpe). As far as I am aware, only two other species, Pelargopsis

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 311

gigantea and Carpophaga Pickeringi, have been recorded from the Sulus; and the blockade

maintained by the Spaniards renders exploration in this archipelago difficult.

At p. 128 Lord Tweeddale alludes to a number of Indo-Malayan genera which are

wanting in the Philippines. The researches of Dr. Steere enable us to supply the

following—Anthreptes, Aithopyga, Arachnothera, Prionochilus, Mixornis, Macronus,

Phyllornis, Tora, Criniger, Buchanga, Hulabes, Tiga, Dendrophila. Besides these the

Eurylemidz are represented by a new genus, the Timeliidze by two, and the Cuculide

by one. All of these new genera are very closely related to well-known Indo-Malayan

forms. It should be noted, moreover, that Palawan, the Bornean character of whose

avifauna has already been alluded to, supplies us with most of the missing genera.

In the present paper 45 species are added, which are presumably peculiar to the

Philippines ; so that out of 285 species, 151 are peculiar (¢f Walden, J. ¢. p. 128).

Following Lord Tweeddale’s example, I give a table “‘ showing by families the proportion

of species peculiar to the Philippine Islands.”

sélses sélsek 38328

Families. S 2 3 By g Families. ee o8e Families. BE\ 8 3 "3

A | 4 oto AA oe ABA oS

Psittacides .......... 9 9 Mundi dsoenneteere 2 Gourideaeneeeeeeene 4

Falconide :......... 14 4 Rittidss) Sa ssnee tse 5 5 Phasianide ........ 1

Sirieidecsts Meee tte cao. 4 3 || Timeliide .......... 4 4 || Tetraonide.......... 2 1

Bicidee cities cy tas's0 9 7 || Crateropide ........ 2 INTIRACHD. co 50005000 1 1

Trogonide .......... 1 il Trenides oss ake 3 3 || Rollulide .......... 1 1

Meropide .......... 2 1 Brachypodide ...... 8 6 || Megapodide ........| 2 1

Coraciide .......... 1 Saxicolidease eee 6 3 || Charadride ........| 8

Alcedinide.......... | 15 G) ||| Siyllvattebsis Gon6occc5n 10 6 || Glareolide.......... 1

Capitonide.......... 2 Motacillide ........ 5 1 Gallinulide ........ 4 1

Cypselide .......... 2 er RaridasnesraSerecy ta ae 4 3 || Rallide ............ 8 2

Caprimulgide........ 3 3 || Meliphagide ........ 1 Dens || eParridse espe a seu 1

Cuculide .......... 12) 11 Nectariniide ........ 16/ 13 Scolopacide ........ 9

Bucerotide.......... 5 5 Certhiide .......... 1 1 Ciconidee .......... 1

amit dea ys 22 heer er cncrste 3 1 Conyideeanrreeie see il 1 ISRUISGED 5 5 50 Ganac0 oc 11 1

Artamide .......... il SUIICED oooce cose 7 3 PSTERONGEDS 3.5 oe Cac oee 4 2

Campephagide ...... 8 4 || Kurylemide ........ 1 1 Procellariide ........ 1

Dicruride .......... 3 Py }| VsaiarelGE}e5 Sooecaes 1 IGAGES Gade apoolee 5

Muscicapide ........| 14 | 11 IRMETCED.6 oadcooccas 6 | odicipidear ye ene 1 il

Hirundinide ........ 3 Treronidz mere 13 9 Pelecanide.......... 6

OME sosccoscadut 4 4 || Columbide.......... 4 3) | THERES. 5 cong edso ox 1

I now proceed to a detailed catalogue of Dr. Steere’s collection; and in this I follow

almost entirely the arrangement adopted by Lord Tweeddale, though in some few

instances not agreeing with the order of the different families: this, however, is a small

matter, my object being to render the presen paper as far as possible a supplement to

the memoir above mentioned.

312 MR. R. B. SHARPE ON BIRDS COLLECTED BY

PSITTACT.

PLYCTOLOPHID.

Cacatua, Vieill.

1. CaAcATUA HH{MATUROPYGIA (Mill.): Walden, Tr. Z. S. ix. p. 132.

a. 2. San Bernardino, Negros, Jan. 1875. Eyes pink.

[I saw a small species of Cockatoo in Balabac apparently of the same kind as the _

Negros bird; but, owing to a severe attack of fever, the three I shot were not skinned.

In Palawan these birds are also plentiful.—J. B. 8. |

PSITTACID A.

Prioniturus, Wagler.

2. PRIONITURUS DISCURUS (Vieill.): Walden, ¢. ¢. p. 132.

a. & ad. Isabella de Basilan. Beak nearly white; iris grey.

b. 2 juv. San Bernardino, Negros, Jan. 1875. Eyes dark.

c. juv. Balabac, Sept. 1874.

[In little flocks near San Bernardino, among the wild plantains. Owing to their

similarity of colouring, they were not easy to see among the foliage.—J. B. 8S. ]

T have carefully examined the above specimens, and have compared them with Lord

Tweeddale’s series. I can recognize but one species, although great difference exists in

the form of the middle tail-feathers. The one from Basilan is an adult, with a long

shaft and broad racket, whereas the other two have these plumes only slightly elongated,

and the spatulee a mere widening of the attenuated apex of the feather. The species was

not before known from any of the above-mentioned islands.

TANYGNATHUS, Wagler.

3. TANYGNATHUS LUZONIENSIS (L.): Walden, ¢. ¢. p. 133.

a. Puerto Princesa, Palawan.

6. 2. Dumalon, Mindanao, Oct. 1874. Eyes nearly white.

c. 9. San Bernardino, Negros, Jan. 1875. yes light yellow.

d. Negros.

[In flocks near Dumalon, coming into the fruit-trees in the town. The natives eat

them.—J. B. 8.]

Negros was known as a habitat for the species, it having been obtained by Mr. L. C.

Layard; but Palawan and Mindanao are both new localities for it.

Cyctorsitra, Hombr. & Jacquinot.

4, CYCLOPSITTA LUNULATA (Scop.): Walden, é. c. p. 133.

a. Manila.

Only one specimen collected, and that without the scale-like collar.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 313

Loricuxts, Blyth.

5. LoricuLus curysonotvs, Scl.: Walden, ¢.¢. p. 137.

a. Zebu.

Agrees with Zebu (typical) examples in Lord Tweeddale’s collection.

6. LoRIcULUS REGULUS, Souancé: Walden, z. ¢. p. 135.

a. Panay.

[The native names for all these little Parrakeets are Cust and Culacissi, both being

derived from the curious whistle of the birds. They seem to reside amongst the cocoa-

nut palms, and are frequently captured and kept by the natives.—J. B. 8.].

Lord Tweeddale and I carefully compared Dr. Steere’s specimen, and came to the

conclusion that it was an individual of L. regulus not adult. Panay is therefore a new

habitat for the species.

ACCIPITRES.

FALCONIDA.

ACCIPITRIN &.

ACCIPITER, Briss.

7. ACCIPITER ViRGATUS (Temm.): Sharpe, Cat. B. i. p.151. Teraspiza virgata, Walden,

t.c. p. 141.

a. ad. Negros.

Tn rich adult plumage with red breast. It is in all likelihood a male, as it has the fol-

lowing measurements :—Total length 11:8 inches, culmen 0°85, wings 6:9, tail 5-5, tarsus

21. The above dimensions, however, do not tally with those of either sex given in my

Catalogue; and it is probable that a further revision may prove that more than one race

is included under the title of A. virgatus. Negros is a new habitat for this Sparrow-

Hawk.

AQUILIN A.

SPILORNIS, G. R. Gray.

8. SPILORNIS HOLOSPILUS (Vigors): Walden, 7. c. p. 142.

a. ad. Mount Ma-hay-hay, near Manila.

Burastur, Hodgs.

9. BurastuR iInpicus (Gm.): Walden, ¢. ¢. p. 143.

a. 3 juv. Zamboanga, Mindanao. LHyes greenish yellow.

6. ad. Panay.

This species, a migrant from the China coast, visiting the Moluccas in winter, is now

first recorded from the above two islands.

314 MR. R. B. SHARPE ON BIRDS COLLECTED BY

PICARLA.

PICIDA.

THRIPONAX, Cabanis.

10. THRIPONAX JAVENSIS (Horsf.): Walden, 7. c¢. p. 146.

a. g. Zamboanga. LHyes yellow.

6. 9. San Bernardino, Negros, Jan. 1875. Eyes yellow.

c. 2. Negros.

d. 9. Puerto Princesa, Palawan.

[Climbing about the dead trees, generally in pairs.—J. B.8.]

I agree with Lord Tweeddale that we cannot separate the Philippine Black Wood-

pecker from the Javan bird. It was not known before from Palawan and Negros, but

had been recorded from Mindanao by Wagler.

CHRYSOCOLAPTES, Blyth.

11. CHRYSOCOLAPTES XANTHOCEPHALUS, Walden & Layard: Walden, ¢.c. p. 147.

a. 6. San Bernardino, Negros, Jan. 1875. yes carmine.

[I saw more of this Woodpecker, which was not rare; but I had the greatest difficulty

in getting about the forests in Negros, owing to the obstruction caused by the trees

blown down in a recent typhoon.— J. B.S. ]

This species was originally described from the island of Negros; but only the female

was known. Dr. Steere has now procured the male, which shows that the species is an

excellent one. The description of the male is as follows :—

Adult male-—General colour above ruddy crimson, all the feathers olive-brown at the

base, the upper tail-coverts entirely of the latter colour, with a crimson mark on their

margins; scapulars and wing-coverts exactly resembling the back; the bastard wing,

primary-coverts, and quills brown, externally yellowish olive, the secondaries brown,

inclining to olive on the outer webs, which are externally crimson like the back, all the

quills spotted with white on the inner web, much more largely on the secondaries; tail-

feathers dark brown, with a slight wash of olive on the outer webs; crown of head bright

crimson, with yellow bases to the feathers ; lores, eyebrow, and entire sides of face bright

yellow, as also the throat, which is bordered on each side with a narrow black mousta-

chial line, and has also another down the centre of the throat; sides of neck yellow, with

three lines of black running down from behind the ear-coverts; neck all round, and

fore neck, scaly in appearance, the feathers being yellowish buff fringed with black ;

remainder of under surface yellowish buff, the sides of the body somewhat streaked with

olive-brown; under wing-coverts dull fulvous, with olive-brown margins to the feathers,

which are also slightly washed with red ; bill stone-brown (in skin) ; legs yellowish, claws

black; “iris carmine’’ (S¢eere). Total length 10°5 inches, culmen 1°6, wing 5:6, tail

3°8, tarsus 1:05,

12. CHRYSOCOLAPTES MACULICEPS, sp. n. (Plate XLVI. fig. 2.)

Ad. supra coccineus, plumis ad basin olivascentibus et celatim albo maculatis; tectri-

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 315

cibus alarum dorso concoloribus nec maculatis; tectricibus primariorum nigrican-

tibus extus olivascenti-aureis; remigibus nigricantibus interne albo ovatim macu-

latis, extus olivascenti-aureis gradatim coccineis; supracaudalibus nigris, versus

apicem olivaceis et fulvo maculatis; cauda nigricanti-brunnea olivaceo lavata, rec-

tricibus externis basaliter fulvo maculatis ; pileo summo olivascenti-brunneo, fulvido

transversim maculato, coccineo adumbrato, maculis fulvis hoc colore nitentibus;

loris et facie laterali fulvescenti-flavis, regione parotica postica genisque anguste

nigro striatis; gula etiam fulva, vitta malari distincta nigra; collo undique marmo-

rato, plumis lete fulvis nigro marginatis, torquem collarem squamosam forman-

tibus; corpore reliquo subtus ochrascenti-fulvo, plumis pectoralibus nigro margi-

natis, reliquis nigro irregulariter transfasciatis; subalaribus pectori concoloribus ;

rostro corneo, mandibula ad basin flavicante; iride coccinea. Long. tot. 10, culm.

1:55, alee 5°55, caudee 3°55, tarsi 1:05.

a. Isabella de Basilan, Oct. 1874. Eyes carmine.

The nearest ally to this species is C. hematribon; but the Basilan bird differs in its

paler red and more spotted head, buff-coloured ear-coverts, spotted mantle and upper

tail-coverts. Dr. Steere marked on his label that one bird was “full-grown,” not being

young, as it might have been supposed to be.

13. CHRYSOCOLAPTES ERYTHROCEPHALUS, sp. n. (Plate XLVI. fig. 1.)

Ad. supra auratus vix coccineo lavatus; remigibus brunneis extus auratis, secundariis et

tectricibus alarum interscapulio concoloribus; pileo cristato undique coccineo;

gutture paullo pallidiore, vitta mediana angusta nigra, altera utrinque malari an-

gusta nigra; preepectore et torque collari ochrascenti-fulvo, plumis nigro anguste

circumdatis ; corpore reliquo subtus fulvescenti-albo, nigro variegato, plumis an-

guste nigro marginatis, hypochondriis et subcaudalibus irregulariter nigro transfasci-

atis; axillaribus albidis nigro maculatis; subalaribus nigris albo ovatim maculatis,

margine alari aureo lavato; supracaudalibus nigricantibus, olivaceo terminaliter

lavatis; cauda nigra; rostro virescenti-corneo; pedibus saturate plumbeis; iride

coccinea. Long. tot. 10-0, culm. 1°7, alee 5°65, caudee 3°4, tarsi 1:15.

a. 6 ad. Puerto Princesa, Palawan.

This species appears to be distinguished from all the other members of the genus by its

red throat and head.

Trea, Kaup.

14. Ties JAVANENSIS (Ljungh): Salvad. Ucc. Born. p. 55.

a. §- Puerto Princesa, Palawan. Lyes black.

fam unwilling on the strength of one specimen only to institute a new species; but

certainly this three-toed Woodpecker differs from all the others with which I have com-

pared it in the uniform sides of the fore neck, which are brown, like the rest of the

under surface, the latter being barred across. It appears to agree best with a Bali bird

obtained by Mr. Wallace, and is probably the same in worn plumage.

SECOND SERTES.—ZOOOLGY, VOL. T. 2uU

316 MR. R. B. SHARPE ON BIRDS COLLECTED BY

TROGONID A.

HARPACTES, Swains.

15. HARPACTES ARDENS (Temm.): Walden, ¢.c. p. 149.

a. 2. Isabella de Basilan.

[I found plenty of Trogons in the forests some little distance inland, but I only suc-

ceeded in preserving one female bird.—J. B. 8.]

The single bird obtained being a female renders it difficult to identify the species ; but

as 7. ardens is known from Mindanao, it is possible that it extends also into Basilan.

MEROPIDA.

Meroes, Linnzeus.

16. Mrrors pHinippinus, L.: Walden, ¢. ¢. p. 149.

a,b. Negros.

c. Panay.

d. Dumalon, Mindanao, Oct. 1874. yes red.

e. Zamboanga, Mindanao.

[At Zamboanga I found them flying over a wet rice-field taking dragon-flies on the

wing, and returning to stakes to- devour them. In the other islands I saw them in the

open.—J. B. 8.]

Panay and Mindanao are new habitats for this species.

17. Mzrors Bicotor, Bodd.: Walden, ¢. c. p. 150, pl. xxvi. fig. 1.

a, b. Panay.

c. 2. Dumalon, Mindanao, Oct. 1874. Eyes carmine.

[Found in swamps, perching in the low trees at Dumalon. In Panay it was shot

more in the open, settling on weeds and sticks a little way from the ground.—J. B. S.]

Not known before from either of the above-mentioned islands.

CORACIIDA.

Eurystomvs, Vieillot.

18. EURYSTOMUS ORTENTALIS (L.): Walden, @. c. p. 152.

| a, b, c. Dumalon, Mindanao.

d, e. . San Bernardino, Negros, Jan 1875. Eyes hazel.

{Found in abundance in the Indian village of Dumalon, in the island of Mindanao, as

well as in Negros. In the former place they were perched on limbs of dead trees in the

edgings of clearings; and in Negros they were seen along the sides of the streams on

limbs overhanging the water.—J. B. 8.]

Although recorded from Negros, Mindanao is a new locality for the species.

PROF, STEERE IN THE PHILIPPINE ARCHIPELAGO. 317

ALCEDINID A.

ALCEDININE.

Atcrpo, Linneeus.

19. ALCEDO BENGALENSIS, Gm.: Walden, ¢. c. p. 152.

a. juv. San Bernardino, Negros, Jan. 1875. Eyes dark.

6. $. Dumalon, Mindanao. Eyes hazel.

[Very generally distributed throughout the Philippines, and found on nearly every

stream.—J. B.S. ]

_ Previously to Dr. Steere’s visit, we only knew of this species from Luzon.

PELARGOPSIS, Gloger.

20. PELARGOPSIS LEUCOCEPHALA (Gm.): Sharpe, Monogr. Alced. pl. 31.

a. 6. Puerto Princesa, Palawan, July 1874.

_ [J found this species in mangrove swamps in the island of Palawan.—J. B. 8.]

The above specimen belongs to P. lewcocephala of Borneo, not P. Gouldi, which is

probably confined to Luzon.

21. PELARGOPSIS GIGANTEA, Walden, Ann. N. H. (4) xiv. p. 123 (1874): Meyer, ¢. ec.

pp. 401, 475.

a. Dumalon, Mindanao.

[Frequently seen flying along the sea-beach, fishing over the open sea, and perching

upon the low trees on the sandbars.—J. B. S.]

Lord Tweeddale described this distinct species from Sulu in the Sulu archipelago, where

Dr. A. B. Meyer obtained it; and now Dr. Steere has procured it in Mindanao. Itisa

representative of P. melanorhyncha of Celebes, but has a reddish bill, like P. leucoce-

phala &c. I compared the Mindanao bird with Lord Tweeddale’s type specimen.

Cryx, Lacépéde.

22. CEYX MELANURA, Kaup: Walden, ¢. ¢. p. 153.

a. 6. Dumalon, Mindanao. LHyes hazel. |

[I only saw the one specimen of this pretty Kingfisher, in deep thick jungle, perching

low. Its food consisted of small crabs.—J. B. 8. |

The single bird collected by Dr. Steere differs from every other Philippine specimen

which I have examined in having the wing-coverts black with rufous edgings, whereas

all the others which have fallen under my notice have had them black spotted with

blue; and Lord Tweeddale has a nestling resembling the adult in this respect. At the

same time I do not like to found a new species on a single example.

HALcyon, Swainson.

23. Hancyon @uLARIS (Kuhl): Sharpe, Monogr. Alced, pl. 70. Entomobia gularis

(Kuhl), Walden, ¢. ¢. p. 154.

2U 2

3818 MR. R. B. SHARPE ON BIRDS COLLECTED BY

a. 6. Maasin, Panay, Jan. 1875. yes dark.

b,c. 3; d. 2. San Bernardino, Negros, 1875. Eyes dark.

[There is scarcely any difference in the sexes; but the female seems a little lighter

below, and to have rather a more greenish tinge on the scapulars, back, and tail.

I noticed this species along the streams, sometimes fishing, but as often feeding on

the ground, where its diet seemed to consist of earth-worms. I saw this, or a closely

allied species, near Zamboanga, in Mindanac, but failed to secure it.—J. B. S. |

Mr. Layard obtained this Kingfisher in Negros; but Panay is a new locality for it.

24. Hatcyon HomBRONI, Bp.: Sharpe, Monog. pl. 115. Actenoides hombroni, Walden,

t.c. p. 155.

a. 3. Dumalon, Mindanao. Eyes hazel.

[Only met with on one occasion, in low thick jungle which was covered every day by

the sea.—J. B. 8.]

This Kingfisher is only known from the original type specimen in the Paris Museum,

which was obtained in Mindanao by Messrs. Hombron and Jacquinot. I have never

seen a skin before; and the plate in my ‘Monograph’ was drawn from a coloured

sketch sent over to me from Paris. I therefore give a full description of the adult

male in Dr. Steere’s collection :—

Adult male.—Head and nape bright blue, more brilliant on the sides of the head,

over the eye, and on the nape; round the latter a narrow line of deep black; ear-

coverts tawny chestnut; along the lower line of the lores a streak of black feathers,

reaching below the eye, and widening behind the latter, being here shaded with blue;

cheeks bright blue, forming a broad band; sides of neck and hinder part of the latter

deep tawny, varied with narrow black edgings to the feathers; mantle blackish, mottled

with tawny spots, these being subterminal with a narrow black frmge; middle of back

and scapulars and wing-coverts green with a slight shade of verditer, each feather

having a distinct subterminal spot of ochraceous buff; quills blackish, externally washed

with greenish, the primaries edged with ochraceous, the secondaries with the same sub-

terminal spot of ochre as on the wing-coverts; lower back, rump, and upper tail-coverts

bright silvery cobalt, the sides of the back and the lateral coverts blackish washed with

blue; tail-feathers deep blue, with black shafts; throat white, slightly washed with

tawny; rest of the under surface deep tawny, whiter on the centre of the abdomen; the

breast-feathers with narrow nearly obsolete blackish margins; thighs externally black-

ish, internally deep tawny; feathers at side of vent, adjoining sides of lower back, deep

blue, the outer web more or less ochraceous; under wing-coverts and axillaries deep

tawny ; the quills blackish below, edged with pale tawny-buff along the inner web; bill

coral-red, the culmen black (in skin); iris hazel. Total length 11°83 inches, culmen 2-0,

wing 4°95, tail 4°15, tarsus 0°75.

25. Haucyon WINCHELLI, sp.n. (Plate XLVII.)

9 ad. supra nigra, purpureo lavata; dorso postico, uropygio et supracaudalibus argenteo-

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 319

cyaneis, his purpurascentibus;. loris et torque collari postico castaneo-fulvis ;

pileo nigro, lateraliter cyaneo, supercilium latum ab oculo summo usque ad

nucham productum exhibente; facie laterali cum regione parotica et colli lateribus

nigris purpureo vix lavatis; alis nigris, vix extus purpurascenti lavatis; cauda

purpurascenti-nigra ; genis et corpore subtus leete cervino-fulvis, abdomine albicante ;

subalaribus et remigibus intus cervinis; rostro nigro, ad basin mandibulze carnoso ;

iride brunnea. Long. tot. 10-0, culm. 2°05, alee 4:1, caudee 3:2, tarsi 0°55.

a. ©. Basilan (Isabella). Eyes hazel.

[This Kingfisher inhabits the forest; and its food consists of beetles, dragon-flies, Ke.

I met with it at a long distance from water.—J. B. S.]

This species is named, by Dr. Steere’s request, after his friend and old tutor, Mr.

Winchell.

26, Hatcyon cHoris (Bodd.): Sharpe, Monogr. pl. 87. Sauropatis chloris, Walden,

t. c. p. 155.

a.ad. Negros.

6. 2 juv. Dumalon, Mindanao, Oct. 1874. Eyes dark.

ec. g ad. Isabella de Basilan.

{General in the islands of Mindanao, Panay, Negros, and Zebu, perching on low

trees, and searching for worms on the ground; they also fish occasionally. They

usually frequent cane-fields and open plains, perching on low timber.—J. B. S.]

Not before recorded from Mindanao, nor from Basilan.

CYPSELID.

Macropreryx, Swainson.

27. MAcropTERYx comATus (‘Temm.): Walden, ¢. c. p. 158.

a, 6. Negros.

c,d,e. 6. Dumalon. Eyes dark.

[In great abundance along the sea-beach at Dumalon, sitting towards evening in

clusters on the limb of a tree. In Negros I shot the pair right in the woods.—

eal bears

The birds from Mindanao belong to the small race, or true WZ. comatus of Tem-

minck, and agree in their measurements with Bornean examples. The specimens from

Negros, on the contrary, are much larger, as Lord Walden found to be the case with

the Luzon birds. The following measurements exhibit these differences in size :—

Long. tot. ale. caude. rect. ext. tarsi.

a. 6 ad. Negros (Steere) . . . . 6:6 5:45 2:0 315 0.3

boRads 5 x Speer en 6:4 5°35 2:0 315 0:3

ce. g ad. Dumalon ,, <i 5:8 5-1 2-0 2°95 0°25

dG ads)" ,, 35 = TORS 5:8 4:8 1:8 3°0 0°25

e. ¢ ad. 5 5 ffs) cee 6:0 4°95 18 29 0°25

ip & ath Ibelonern (hoy) 6 Give ss 6:0 4-9 16 30 0°25

qed re a 61 4:95 1:85 31 0°25

320 MR. R. B. SHARPE ON BIRDS COLLECTED BY

Contocata, G. R. Gray.

28. CoLLOCALIA TROGLODYTES, Gray: Walden, ¢. c. p. 158.

a. Panay.

[Shot at some distance from the sea, in the hills.—J. B. 8.

Compared with the typical specimens, the Panay bird is more blue above, with a

purplish gloss on the wings. Underneath the throat and breast are washed with silvery

white. The types are much browner, with very little purple gloss above, and scarcely

any of the silvery white below. They are also a little larger (wing 3°65-3:75 inches).

These differences, however, are only due to age, Dr. Steere’s Specimens being older;

the measurements are as follows—total length 3:7 inches, culmen 0:15, wing 3°65,

tail 1-65, tarsus 0°35. Y

The two specimens in the Museum have no exact locality to them; for they are

registered as from “ Malacca,” at the end of a Philippine collection, both received from

the late Mr. Cuming. It is probable that they are really from the Philippines; but

there is no proof of this.

CUCULID®.

CUCULIN &.

CAcoMANTIS, S. Miller.

29. CACOMANTIS MERULINUS (Scop.): Walden, ¢.c. p. 160.

a. 3. Malamaui, Oct. 1874. Eyes and inside of mouth cherry-red, legs yellow.

b. 2. Maasin, Panay, Jan. 1875. Eyes grey.

Originally described by Sonnerat from Panay, and recorded by Prof. Schlegel from

Luzon (Govers). It also occurs in the southern Philippines, as Dr. Steere shot it in

Malamaui; and it is probably found throughout the archipelago.

Curysococcyx, Boie.

30. CHRysococcyx BASALIS (Horsf.): Salvad. Ucc. Born. p. 62.

a, Negros.

Agrees with a Flores specimen collected by Mr. Wallace. Total length 5-7 inches,

culmen 0°55, wing 3°7, tail 2°6, tarsus 0°6.

Eupynamis, Vigors and Horsfield.

31. HUDYNAMIS MINDANENSIS (L.): Walden, J. ¢. p. 162.

a. 2. San Bernardino, Negros, Jan. 1875. Eyes carmine.

[I found this in thickets of low trees about the sugar-cane fields and along a little

river near San Bernardino.—J. B. 8. ]

The specimen collected is in the rufous and black plumage.

SuRNIcULUS, Lesson.

32. SURNICULUS VELUTINUS, Sp. 0.

3 ad. supra velutino-nigra vix purpureo lavata; scapularibus alisque metallice viri-

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 321

descenti-nigris vix chalybeo nitentibus ; uropygio, supracaudalibus caudaque metal-

lice virescenti-nigris, illis chalybeo nitentibus; facie laterali et corpore subtus

velutino-nigris haud metallicis; tibiis niveis, plumis longis obtectentibus externe

albis, interne nigris; remigibus subtus nigris, albo transfasciatis, pogonio interno

albo ovatim maculato fasciam unicam formante; rostro et pedibus nigris, his subtus

flavis; iride saturate brunnea. Long. tot. 8°3, culm. 0°8, ale 4°55, caude 4:1,

tarsi 0°65.

a. 2. Malamaui, October 1874. Eyes dark.

This pretty little Cuckoo is closely allied to S. dugubris (Horsf.), but is at once to be

distinguished by the velvety black colour of the upper and under surface, and by the

absence of metallic gloss on these parts, what little colour there is being of a purplish

shade. The size is also somewhat smaller.

PHENICOPH HINA.

DASYLOPHUS, Swainson.

33. DASYLOPHUS SUPERCILIOSUS (Drap.): Walden, ¢. c. p. 162.

a. 3g. Mount Ma-hay-hay, near Manila.

[Only found in Luzon, and never seen by me in any of the other islands. It was

plentiful in the neighbourhood at the foot of Ma-hay-hay in the thickets about the

villages—J. B. 8. ]

Dryococcyx, gen. n.

Closely resembling Rhinococcyx, Sharpe (P. Z. 8. 1873, p. 604), but distinguished at

a glance by the shape of the nostril, which is a small circular hole situated in a deep

perpendicular groove which separates the lores from the upper mandible.

Fig. 1.

Profile of Dryococcya Harringtoni.

34. Dryococcyx HARRINGTONI, sp. n.

¢ ad. supra metallice viridis vix olivascens; alis dorso concoloribus, primariis extus

chalybeis ; cauda viridi, terminaliter 4 castanea, rectricibus externis 3 castaneis;

capite summo olivascenti-brunneo; collo postico castaneo lavato, torquem collarem

vix formante; fronte et genis anticis mentoque cinereo lavatis; facie laterali et

corpore subtus toto lete rufis, abdomine et subcaudalibus castaneis; regione oculari

nuda coccinea; rostro pallide viridi; iride coccinea. Long. tot. 18, culm. 1°65, ale

6°35, caudee 9°3, tarsi 1°55.

a. 3. Balabac, September 1874. Eyes currant-red, as well as the excrescences on the face.

322 MR. R. B. SHARPE ON BIRDS COLLECTED BY

[I saw others of this species when I reached the jungle on the hill-sides during an

excursion I made into the mountains.—J. B. 8. |

This new form of Coucal is most nearly allied to Rhinococcyx curvirostris, from which

it differs not only in the structural peculiarities of the nostrils but also in colouring.

It has an olive-brown head and partial rufous collar round the neck, and has none of the

greenish black colour on the abdomen and thighs, these being rufous like the breast.

By Prof. Steere’s desire I have named this species after Mr. Harrington, Professor of

Botany in the University of Michigan, and an old schoolfellow and college friend of the

discoverer’s.

CENTROPODINS.

CEnTROCOCCYX, Cabanis and Heine.

35. CENTROCOCCYX VIRIDIS (Scop.): Walden, ¢. c. p. 168.

a. Zamboanga, Mindanao.

PyrRHocentorR, Cabanis and Heine.

36. PyYRRHOCENTOR MELANOPS (Less.): Walden, ¢.c. p. 164.

a. 3. Malamawi, October 1874. yes cherry-red.

BUCEROTID.

Buceros, Linnzeus.

37. BucrRos HyDROcORAX, L.: Walden, t. c. p. 164.

a. juv. Head only. Mount Ma-hay-hay, Manila.

(This Hornbill was plentiful on the above-named mountains.—J. B. S.]

CRANIORRHINUS, Cabanis and Heine.

38. CRANIORRHINUS LEUCOCEPHALUS (V.): Walden, ¢. ¢. p. 165, pl. xxvii. figs. 1, 2.

a. Zamboanga, Mindanao.

39. CRANIORRHINUS WALDENI, Sharpe, Journ. Linn. Soe. Zool. vol. xiii. p. 155; Elliot,

Monogr. Bucerot. part 1.

Ad. supra niger, ubique metallice olivascenti-viridi nitens; alis dorso concoloribus ;

cauda fulva, rectricibus quatuor intermediis pallide castaneis, omnibus late nigro

terminatis et dimidio basali nigro; pileo et collo undique saturate castaneis, plumis

supraocularibus et regione parotica antica fulvescentibus ; regione periophthalmica,

genis et gutture toto nudis; corpore reliquo subtus nigro viridi nitente; rostro

rubro, mandibula et crista rostri maxime sulcatis. Long. tot. 28, alee 14°6, caudze

10°8, tarsi 2:1.

a. 3. Panay.

[I shot this Hornbill on the highest ridge of the mountains west of Ilo-Ilo. This is

the only place where any of the virgin forest is left; and here alone I saw these birds.

They were not very rare; but I could only get a single specimen, as they flew so high in

the trees that ry gun could scarcely reach them.—J. B. 8.]

PROF, STEERE IN THE PHILIPPINE ARCHIPELAGO, 328

PENELOPIDES, Reichenbach.

40, PENELOPIDES PANINI (Bodd.): Walden, ¢.c¢. p. 166, pl. xxviii. figs. 1, 2.

a. Negros.

PASSERES.

LANIIDA.

Lantus, Linneeus.

41. Lantus tuctonensis, L.: Wald. ¢. ¢. p. 171, pl. xxix. fig. 1,

a. Adult. Panay.

6. ¢ imm. Basilan (Isabella). Eyes hazel.

[In the cultivated country of Panay, in the lower country. In Basilan they were

found about the clearing in the vicinity of the fort at Isabella.—J. B. S.]

Dr. Steere adds two more islands to the known Philippine habitats of this species ;

but, as this Shrike is a migrant from China, it probably visits every one of the group.

Mr. Low and Mr. Everett have lately discovered it in North-western Borneo.

ARTAMIDA.

ARTAMUS, Vieillot.

42. ARTAMUS LEUCORHYNUSs (L.) : Walden, ¢.c. p. 174,

a. Dumalon, Mindanao, October 1874.

[Very plentiful at Dumalon and also in Negros, flying much like Swallows over open

ground, and lighting upon stumps.—J. B. 8.]

This Swallow-shrike was not before known from Mindanao.

CAMPEPHAGIDA.

GRAUCALUS, Cuvier,

43, GRAUCALUS SUMATRENSIS, Mill. ; Salvad. Uce. Born. p, 150.

a, 6. Puerto Princesa, Palawan.

[Found in similar situations to G. striatus.—J. B. 8.]

This is a well-known Bornean and Sumatran bird; and its occurrence in Palawan is

one of the evidences of a Bornean element in that island.

44, GRAUCALUS STRIATUS (Bodd.): Walden, ¢. ¢. p. 175, pl. xxx. fig. 1.

a. 3. San Bernardino, Negros, Jan. 1875. yes reddish.

6. 3. Malamaui. Eyes light yellow, nearly white.

c. Zamboanga, Mindanao.

[The difference in the colour of the eyes in specimens a, 0 is curious, as they are nearly

identical in plumage. ‘They were found in low trees in oper country.—J. B. 8.]

A very full account of this species is given by Lord Tweeddale (/. ¢.), describing all

the changes of plumage. Malamaui is a new locality.

SECOND SERIES.—ZOOLOGY, VOL. I. 2X

o24 MR. R. B. SHARPE ON BIRDS COLLECTED BY

LALAG#, Boie.

A5, LALAGE poMINIcA (Miuill.): Walden, 7. c. p. 178.

a. Panay.

b. ¢. Dumalon, Mindanao. Hyes dark.

c. 9. Malamaui, Oct. 1874. Eyes dark.

This species was added to the Philippine avifauna by Dr. Meyer, who met with it in

Zebu and Guimaras. All the islands mentioned above are new localities for it.

Prricrocotes, Boie.

46. PpricrocoTus 1GNEUS, Blyth: Sharpe, Str. F. 1876, p. 209.

a,b,c. ¢ ad. Puerto Princesa, Palawan. Eyes black.

d,e. 2 juv. Puerto Princesa.

- [The first specimen I shot in a mangrove tree; and the other four I killed from the door

of my house as they were eating fruit on a small clearing. The iris in all was black.—

J. B.S.)

This species of Pericrocotus is known from Malacca and Borneo, being very common

in the north-western part of the latter island. The island of Palawan is therefore an

entirely new locality for it. Prof..Steere’s series comprise a pair of fully adult males,

a third in full plumage, excepting a few yellow feathers on the throat, and two young

males. One of the latter is in the dress of adult P. flagrans, showing that at first the

young male resembles the old females; for there can be no doubt that P. flagrans is only

the hen of P. zgneus.

DICRURIDA,

Dicrurvs, Vieillot.

47. DickURUS MIRABILIS, Walden: id. ¢. ¢. p. 181.

a. 6. San Bernardino, Negros, Jan. 1875.

6, Negros.

[Always in low thickets, and feeding on fruit. Rather hard to shoot, owing to their shy

habits.—J. B.8.] geen: .

BucHanea, Hodgson.

48, BUCHANGA CINERACEA (Horsf.): Sharpe, Cat. of Birds, iil. p. 250.

a. 2. Puerto Princesa, Palawan. yes hazel.

[Procured in the deep woods a good way from the sea.—J. B. 8. ]

Compared with (typical) Javan examples, the present specimen shows complete iden-

tity. It isnot quite in full plumage, but is assuming the grey coloration of the adult, the

young feathers being downy in texture and not of such a blue-grey colour as in the old

bird, the shade of the feathers being of a dull ashy. The measurements are as follows

—total length 9 inches, culmen 0:9, wing 4°7, tail 3°75, to tip of outer feather 4°85,

tarsus 0°7,

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. By)

MUSCICAPIDA.

PuitEntomA, Hyton.

49, PHILENTOMA ALBIVENTRIS, sp. n.

Ad. schistaceo-cyanescens, plumis frontalibus medialiter argentescenti-albido striolatis ;

dorso scapularibusque leete castaneis, his summis vix schistaceo lavatis; tectricibus

alarum schistaceo-cyanescentibus, majoribus intimis extus castaneis; remigibus

nigris, primariis vix schistaceo limbatis, secundariis extus castaneis; rectricibus

duabus centralibus nigris, duabus proximis extus castaneis, intus nigris, reliquis

omnino castaneis; facie laterali sordide cyanescente ; gutture et pectore toto schis-

taceo-cyanescentibus, plumis anguste argentescenti-albo medialiter striolatis, abdo-

mine toto albo; crisso et subcaudalibus pallide castaneis, hypochondriis imis fulvo

lavatis; subalaribus cinereis, majoribus albicantibus. Long. tot. 6:4, culm. 0°65,

alze 3°05, caude 3°45. tarsi 0-7. |

a. ad. Guimaras.

[Found in the thick woods on low trees.—J. B. S.]

This species closely resembles P. cyaniceps, which is in the Museum, from Mindanao,

but is distinguished at once by its white belly. It is not improbable that it may be a

sexual plumage of P. cyaniceps.

Ruipipura, Vigors & Horsfield.

50. RHIPIDURA NIGRITORQUIS, Vigors. Leucocerca nigritorquis, Walden, t. c. p. 182.

a. &. Puerto Princesa, Palawan. yes black.

b. 9. Zamboanga, Mindanao. : Eyes dark.

[Found by me springing about over the knees of thé mangrove-trees, feeding appa-

rently on insects.—J. B. 8.]

I cannot see the slightest difference in the sexes of this bird, which seems also to be

the same specifically from the different islands of the Philippines.

Cyornis, Blyth.

51. CYORNIS PHILIPPINENSIS, sp. n. Cyornis banyumas, Walden, ¢. c. p. 152.

C. similis C. banyumati, sed abdomine et subcaudalibus albis pspaepead Long. tot.

5°8, culm. 0°45, alse 2:95, caude 2°6, tarsi 0-7.

a-c. Panay.

This is the C. banyumas of Lord Tweeddale’s list ; but he does not omit to notice certain

differences. On comparing Dr. Steere’s specimens with a large series from Labuan and

other localities, I notice that the belly and under tail-coverts are white in the Philippine

birds, while they are always rufous in those from Borneo. I think, therefore, that the

former must be kept distinct.

Hypotuymis, Boie.

52. HypoTHyMIs AZUREA (Bodd.): Walden, 7. c. p.182.

“a, 6. 2. Basilan (Isabella). Eyes dark brown,

[Pretty abundant in the island of Basilan.—J. B. S. |

326 ; MR. R. B. SHARPE ON BIRDS COLLECTED BY

This wide-spread species appears to be found in most of the islands of the archipelago,

having been noted from Guimaras, Negros, and Luzon.

53. HypoTHYMIS SUPERCILIARIS, sp. n.

¢ ad. cyanescenti-cinereus, pileo saturate ceruleo; fronte et linea angusta super-

ciliari argentescenti-cyaneis; loris et plumis periophthalmicis cinereo-nigris ; facie

laterali, gutture et pectore summo cyanescentibus ; corpore reliquo subtus albicante,

cyanescenti lavato ; tibiis cyanescenti-cinereis ; tectricibus alarum dorso concoloribus,

plumis intus nigricantibus; remigibus nigricantibus, extus leete cyaneis, secundartis

intimis omnino cyaneis; rectricibus intermediis cyaneis, reliquis nigricantibus extus

cyaneis; subalaribus pallide cyanescenti-cinereis ; rostro nigro; pedibus brunneis;

iride saturate brunnea. Long. tot. 6:2, culm. 0-6, ale 3:1, caudie 3:0, tarsi 0°65.

a, 6. Isabella de Basilan, October 1874. Eyes dark brown.

[A true Flycatcher in habits, flying from limb to limb, and catching flies on the wing.

—J. B.S8.] .

The species nearest to this new one seems to be ZZ. indigo; but the latter is a much

smaller bird. The general tint of the upper surface is of a much greyer blue in

H. superciliaris, while the bright cobalt wings and tail also distinguish it in addition to

the silvery blue frontal line and eyebrow. The under surface is greyish blue, not greenish,

the colour extending over the throat and chest, leaving the rest of the under surface

bluish white. In H. indigo the breast is also greenish, leaving only the abdomen

white. .

Eumytias, Cabanis.

54. EUMYIAS PANAYENSIS, sp. n.

Ad. sordide cyanescens, pileo vix leetiore; tectricibus alarum minimis dorso concolori-

bus, majoribus cum remigibus rectricibusque nigricantibus dorsi colore marginatis ;

facie laterali et corpore subtus vix leetius cyanescentibus, abdomine medio albicante

cyaneo lavato; crisso et subcaudalibus subalaribusque albis. Long. tot. 6-0, culm.

0°55, alee 2°95, caudee 2°45, tarsi 0°7.

a. ad. Panay. .

Closely allied to #. sordida, Walden (£. ceylonensis, Gray, Handl. B.i. p. 326. no. 489,

deser. nulla), but distinguishable by its broader bill and by its brighter cyaneous under

surface. In #. sordida the breast is grey, with a brighter blue shade on the chin, and

the lores and feathers in front of the eye and at the base of the lower mandible are

black. In #. panayensis the breast is rather bright greyish cyaneous, a little more

silvery on the throat, lores, and base of mandible, the latter parts not showing any

black.

Burauis, Boie.

55, BUTALIS MANILLENSIS, Bp.: Walden, ¢. c. p. 183.

a. Panay.

The single specimen collected by Dr. Steere proves that 6. manillensis of Bonaparte

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. B27

is the species more familiarly known as B. griseosticta of Swinhoe; and of course

Bonaparte’s name will take precedence over the latter title. This identification was to a

great extent predicted by Lord Tweeddale (J. ¢.).

XANTHOPYGIA, Blyth.

56. XANTHOPYGIA NARCISSINA (Temm.): Swinh. P. Z.8. 1871, p. 380.

a. &. Dumalon, Mindanao. Eyes hazel.

A lovely specimen in full plumage. This is the first time that the species has been

known from the Philippines, where it is doubtless only a winter visitant.

XANTHOLESTES, gen. n.*

57. XANTHOLESTES PANAYENSIS, sp. 0.

é. supra olivascenti-flavus, pileo saturatiore; uropygio leete flavo; tectricibus alarum,

remigibus et rectricibus brunneis flavido marginatis; loris flavis; subtus leetissime

flavus, gutture vix obscuriore. Long. tot. 4°7, culm. 0°45, alee 2°2, caude 2:2,

tarsi 0:5.

a. ad. Panay.

Closely allied to XY. helianthea, Wall. (Walden, Tr. Z. 8. viil. p. 66, pl. vii. fig. 1), of

which it is a representative species, differing in its smaller size and olive-greenish head

and face, the latter being yellow in X. helianthea.

Srrarra, Blyth.

58. SETARIA RUFICAUDA, Sp. D.

$ ad. supra fulvescenti-brunneus, pileo vix obscuriore; supracaudalibus caudaque

castaneis, rectricibus vix apicaliter brunnescentibus; tectricibus alarum dorso

concoloribus, majoribus saturate brunneis extus dorsi colore lavatis; tectricibus

primariorum nigricantibus extus angustissime olivaceo lavatis; remigibus saturate

brunneis extus fulvescenti-brunneo lavatis, secundariis intimis dorso concoloribus ;

loris cum genis et facie laterali cinereis, regione parotica fulvescenti lavata ; cula et

pectore abdomineque toto albis, pectore summo olivaceo lavato; hypochondriis

olivaceis; tibiis cinerascenti-brunneis; subalaribus subcaudalibusque albis, illis

extus olivaceis; rostro nigro; pedibus pallide brunneis; iride saturate brunnea.

Long. tot. 5°4, culm. 0°6, alee 2°85, caudz 2°35, tarsi 0°7.

a, b. 8. Isabella de Basilan, October 1874. Eyes hazel.

| Has the habits of a Flycatcher; found in rather low deep woods, and only met with

in Basilan.—J. B. 8. |

This new species is similar to S. pectoralis, Salvad., which it strongly resembles below ;

but it differs in its grey cheeks and bright-chestnut upper tail-coverts and tail. In the

latter character it comes very close to S. affinis, which also has the upper tail-coverts and

tail red; but then the latter bird has the lores and sides of face white, not grey.

* Mzyiolestes, in which genus these birds have been placed, is a synonym of Hemipus; cf. Sharpe, Cat. B. iii. p. 305.

328 MR. R. B. SHARPE ON BIRDS COLLECTED BY

ZEOCEPHUS, Bonaparte.

59. ZEOCEPHUS CINNAMOMEUS, sp. n. (Plate XLVIIT. fig. 1.)

3 ad. omnino cinnamomeus, subtus paullo letior, abdomine albicante; remigibus nigri-

canti-brunneis, extus et intus cinnamomeis; cauda saturate castanea, rectricibus

quatuor medianis versus apicem brunnescentibus. Long. tot. 7°5, culm. 0°75, ale 3°4,

caudee 3°35, tarsi 0°7.

a. &. Isabella de Basilan. Eyes dark.

Although not possessing such a strongly graduated tail as Z. rufus*, this bird is

apparently congeneric, and may be distinguished by its brighter and clearer cinnamon-

red colour, and especially by its white belly. Like Z. rufus it has a wattled ring round

the eye.

[Of this new bird I only saw one specimen, which I distinguished by its bright red

colour among the green leaves. It was shot in a thick forest in the interior of the island

of Basilan, at some distance from the sea.—J. B.8.]

60. ZEOCEPHUS CYANESCENS, sp.n. (Plate XLVIII. fig. 2.)

9 ad. schistaceo-cyaneus; loris et margine angusto frontali, mento summo et macula

ad basin mandibule posita, nigris; remigibus nigricantibus dorsi colore extus

lavatis; cauda dorso concolori, sed saturatiore; rostro ¢yaneo; palpebra carun-

culata cyanea; iride nigra. Long. tot. 85, culm. 0°85, ale 3°85, caudze 4:25,

tarsi 0°75.

a. &. Puerta Princesa, Palawan, July 24. Eyes black ; inside of mouth and tongue leaf-green ;

eyelid and bill light blue.

[I saw a few more of this species in the island; but it did not appear to be very

abundant.—J. B. 8.]

This species is totally distinct from the other two rufous-coloured ones.

HIRUNDINID A.

Hirunpo, Linneeus.

61. Hirunpo rustica, L.: Dresser, B. Hur. part XXX.

a, 6. & juv. Dumalon, Mindanao. LHyes dark.

[Found in some numbers; not seen far from the beach.—J. B. S.]

These birds seem to be our Common Swallow and not H. javanica. The specimens

procured were both young birds; but I do not doubt the identification.

ORIOLIDA,

G2. OnroLus CHINENSIS, L.: Sharpe, Cat. B. iii. p. 208. Broderipus acrorhynchus

(Vig.), Walden, ¢. ¢. p. 185.

a. 2 ad. San Bernardino, Negros, Jan. 1875. Baill flesh-colour.

b. g juv. Dumalon, Mmdanao. Hyes dark.

c, ¢ ad. Balabac, Sept. 1874. Eyes “ currant-”red.

* Tord Tweeddale states (¢. c. p. 183) that the precise locality of Z. rufusis not known. We have two specimens in

the Museum collected by Cuming at Cataguan, in Luzon.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 329

[Pretty generally distributed through all the islands of the Philippine group, but

rather shy. At Bernardino it seemed to be common, feeding on the fruits of the

wild bananas.—J. B. 8. ]

Between the adult male and female there is scarcely any perceptible difference beyond

a slightly more greenish tinge on the back and scapulars of the latter.

The young bird hasa blackish bill; and the yellow of the back has a very strong wash of

greenish; the wings and tail are browner, the centre tail-feathers being olive; the horse-shoe

mark is blackish washed with yellow; and on the breast are several fine streaks of black,

63. ORIOLUS SULUENSIS, Sharpe, Cat. B. ili. p. 205.

a, 6, c, d. Si Butu, Sulu archipelago.

Mr. Low very kindly presented to the Museum four specimens of an Oriole which

at first appeared to me to resemble Dr. Cabanis’s description of his Oriolus formosus ;

but, thanks to the kindness of Dr. Briggemann, who procured me the loan of some

specimens of the latter bird, I find that the species is really very distinct, as may be

seen by consulting Mr. Dawson Rowley’s plate (Orn. Misc. pl. lvi.).

64. OrtoLus SteERt, Sharpe, Cat. B. ii. p. 218, pl. x.

g ad. supra sordide aureo-flavus, regione parotica et colli lateribus dorso concolori-

bus; loris, genis, gutture toto et preepectore cinereis; corpore reliquo subtus albo,

plumis medialiter nigro late striolatis; subcaudalibus lete aureis; tectricibus

alarum minimis et medianis dorso concoloribus, majoribus remigibusque nigricanti-

bus extus flavis, secundariis intimis dorso concoloribus; tectricibus primariorum

et primariis nigris extus cinereo limbatis; subalaribus cinereis, externis flavo extus

limbatis; rectricibus nigris aureo apicatis versus basin pogonii interni flavicantibus,

pennis duabus centralibus flavicantibus, apicaliter nigricantibus; rostro rubro ;

iride coccinea. Long. tot. 7-5, culm. 0°9, alee 4°45, caude 3°1, tarsi 0°9.

a. &. Basilan (Isabella). Eyes cherry-red.

6. Negros.

[In Negros I found these Orioles feeding on the wild bananas, everywhere rather shy,

—J. B.8.] ;

The Negros example is rather larger and darker in colour, with broader black ovate

streaks on the belly; the yellow spot on the outer tail-feathers too is rather larger.

Total length 8°3 inches, culmen 1:05, wing 4°7, tail 3:45, tarsus 0°9. Although this

indicates a larger race, I do not propose to give it a specific name. .

This new Oriole resembles at first sight O. wanthonotus, which it equals in size and in

the broad black streaks on the belly; but it is quite distinct from this and from every

other Oriole by reason of its grey throat and fore neck and yellow upper surface. In form

it is a true Oriolus, approaching in colour some of the species of Mimeta.

PITTIDA.

Bracuyvrts, Thunberg,

65. BRACHYURUS STEERII, sp.n. (Plate XLIX.) Sharpe, ‘Nature,’ August 1876.

2 ad. supra olivascenti-viridis; uropygio leetissime cyaneo; supracaudalibus et cauda

330, MR. R. B. SHARPE ON BIRDS COLLECTED BY

nigris; pileo summo et capitis lateribus nigerrimis; tectricibus alarum letissime

cyaneis, majoribus nigris cyaneo terminatis; remigibus nigris, secundariis intimis

dorso concoloribus, externis extus viridibus, primariis 4°, 5°, 6° extus prope basin

macula alba ornatis; gula alba; corpore reliquo subtus lete cyaneo; pectore et

abdomine medio nigerrimis; crisso et subcaudalibus scarlatinis; rostro nigro;

iride saturate brunnea. Long. tot. 7°5, culm. 1:1, alee 4°75, caude 1°7, tarsi 1°7.

a. 8 ad. Dumalon, Mindanao.

b. 8 juv. Dumalon, Mindanao.

[This bird was probably not very rare in the neighbourhood of the village of Dumalon,

as I had one or two more specimens of it, which unfortunately were destroyed.—

dig 18s tS

There is no species with which this beautiful Pitta can be directly compared, as it

differs so thoroughly from all the described forms. Its bright blue breast and white

throat distinguish it from all the black-headed members of the genus.

A second specimen is apparently a younger male from the same locality. It differs

only in the colouring of the belly, which is pale scarlet all the way up, becoming blacker

in certain parts, and showing that the whole is becoming black as in the full-plumaged

bird described.

66. BRACHYURUS PROPINQUUS, Sp. 0.

B. affinis B. erythrogastro, sed tergo cyanescente, plaga transversali dorsali tantum

viridi; pileo castaneo-rufo; collo postico lete cyaneo; gutture toto nigricante,

mento tantum rufo maculato; preepectore nigerrimo ; plaga pectorali lete cyanea,

pectoris lateribus viridescentibus ; corpore reliquo subtus latissime scarlatino ; sub-

alaribus brunneis sordide cyanescenti lavatis, alis ut in B. erythrogastro coloratis.

Long. tot. 6°5, culm. 0°75, ale 3°6, caude 1°5, tarsi 1°35.

a. & ad. Balabac, Sept. 1874. Iris grey.

6. g juv. Dumalon, Mindanao. Iris hazel.

As might be expected, this bird is very closely alliefl to B. erythrogaster, which is

apparently a Manila species. We have compared the Balabac skin with four examples

of the true B. erythrogaster, and find that it cannot be united to that species. The

differences may be summarized as follows :—

B. erythrogaster. B. propinquus.

Back entirely green, including the scapulars, the Back entirely cobalt, meluding the scapulars ;

lower ones only of the latter washed with blue at only the middle of the back’ green washed with

the tip; rump and upper tail-coverts and tail blue, forming a band across the back.

blue.

Throat brown, gradually shading into black on Throat entirely brownish black, becoming jet-

the fore neck. black on the fore neck. ;

Chest green, with a slight wash of blue on the Chest bright blue, the sides only greenish.

central portion.

Under wing-coverts greyish brown. Under wing-coverts dull blue.

This species is intermediate between B. erythrogaster and B. cyanonotus of Ternate,

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 6881

differing from both in its blackish throat and blue under wing-coverts. The other

characters are duly enumerated above. The young bird from Mindanao seems to belong

to this species; but being immature, it is rather difficult to say.

67. BracHyurus sorpipvs (Miull.). Melanopitta sordida, Walden, J. c. p. 187.

a,b. 3 Q. Puerto Princesa, Palawan.

c. g. Basilan.

[Common in both the above-mentioned islands. They alight on the low stumps and

logs in the woods, and emit a curious shrill note at intervals of about a minute. On

being disturbed they fly off rapidly and disappear in the jungle, but very soon return

to the same spot, and commence whistling again directly the danger appears to have

passed. They are said by the natives to raise a large heap of leaves and nest on the top

of the mound.—J. B. S.]

Considering the number of Bornean forms shown by Dr. Steere to inhabit Palawan, it

is interesting to note that the Black-headed Pitta of that island is the true Philippine

species, and not the Bornean one, B. Muelleri. These two birds are closely related ;

but the Bornean species almost entirely wants the black patch on the belly, and has the

thighs blackish instead of ochraceous brown.

TIMELIIDA.

Macronvts, Jardine and Selby.

68. MacRONUS STRIATICEPS, sp. n.

¢ ad. supra fulvescenti-brunneus, scapis pallide fulvescentibus ; uropygii plumis longis,

laxis, scapis basaliter albis ; tectricibus alarum dorso concoloribus et eodem modo

striatis, majoribus autem fulvescenti-brunneis concoloribus; remigibus brunneis

extus castaneis ; cauda brunnea, rectricibus extus castaneo lavatis; pileo nuchaque

nigris, clare medialiter albo striatis; facie laterali alba, plumis utrinque nigro

marginatis, quasi striolatis; gula pure alba; corpore reliquo subtus albido, plumis

pectoralibus olivaceo striatim marginatis, corporis lateribus, tibiis subcaudalibusque

olivascenti-brunneis albido striolatis; subalaribus olivascenti-brunneis ; remigibus

subtus cineraceis, intus rufescentibus; rostro nigricante; pedibus brunneis; iride

brunnea. Long. tot. 5:2, culm. 0°6, ale 2°15, caudze 2°15, tarsi 1°8.

a. 2. Isabella de Basilan. Eyes hazel.

6b. g. Malamaui, October 1874. Eyes hazel.

[The sexes are alike. Shot in exactly the same localities as Mixornis Woodi.—

J. B.S.]

5 Mrxornis, Hodgson.

69. Mrxornis Woopl, sp. n.

Ad. similis WY. pileate, sed facie laterali cinerea, mento albo et dorso olivascenti-

cineraceo distinguendus. Long. tot. 5:2, culm. 0°6, ale 2°35, caude 2°25, tarsi 0°8.

a, 6. S. Puerto Princesa, Palawan.

c. 6. Puerto Princesa. Eyes flesh-colour.

‘d. &. Puerto Princesa. Eyes hazel.

[Only found in the deep brush.—J. B. 8. ]

SECOND SERIES.—ZOOLOGY, VOL. I. Pas

332 _ MR. R. B. SHARPE ON BIRDS COLLECTED BY

PTILOCICHLA, gen. n.

Allied to Turdinus, but distinguished at a glance by the long drooping sickle-shaped

plumes on the sides ofthe lower back. The type is :—

70. PYILOCICHLA FALCATA, sp.n. (Plate L. fig. 3.) .

g ad. pileo summo nuchaque rufescenti-brunneis; fronte laterali et striga postoculari

cervino-rufis ; loris anguste albis; macula anteoculari et regione parotica nigris;

genis gulaque albis, vitta malari angusta nigricante; dorso toto nigricanti-brunneo,

plamis late medialiter fulvis, dorsi postici plumis longis, laxis, falcatis, dependenti-

bus, brunneis vel nigricantibus late albo striatis; alis rufescenti-brunneis, remigibus

intus nigricantibus ; cauda omnino rufescenti-brunnea; corpore subtus nigricante,

late albo medialiter striato; tibiis, subalaribus et subcaudalibus brunneis, fulvo

striolatis ; rostro flavicanti-corneo, mandibula flavicante; pedibus saturate brunneis.

Long. tot. 6°8, culm. 0:95, alee 3:2, caudee 2°4, tarsi 1:3.

a. &. Puerto Princesa, Palawan.

[When first I saw this curious bird he reminded me at once of the Ant-birds

(Formicariide) which I had seen in South America. The little fellow was in the

thickest wood, hopping about on the ground, and puffing himself out like a ball of

feathers, presenting a most remarkable appearance.-—J. B. 8.]

DENDROBIASTES, gen. 0.

Closely allied to Alcippe, but distinguished by its long wings, which reach to the end of

the tail. The type is :—

71. DENDROBIASTES BASILANICA, sp. n. (Plate LITT. fig. 1.)

2 ad. supra rufescenti-brunnea, uropygio paullo fulvescenti lavato; supracaudalibus

castaneis; pileo magis fuscescenti-brunneo; loris, supercilio indistincto et facie

laterali fulvescentibus ; annulo periophthalmico letius fulvo; regione parotica ful-

vescenti-brunnea, auguste albido striata; corpore subtus albicante, gutture vix

fulvido lavato; pectoris lateribus, torque pectorali, hypochondriisque fulvescenti-

brunneis ; tibiis et subalaribus rufescentibus, margine alari rufescenti-brunneo; alis

caudaque supra saturate brunneis, tectricibus dorsi colore extus lavatis, remigibus

et rectricibus extus castaneis; rostro nigricanti-brunneo; pedibus flavicantibus ;

iride brunnea. Long. tot. 4°2, culm. 0°55, ale 2°4, caudz 1°6, tarsi 0°8.

a. 9. Isabella de Basilan. yes hazel.

[Shot in open ground near the fort at Isabella, inhabiting the low bushes in this

neighbourhood.—J. B. 8.]

This seems to be an entirely new form of Timeliine bird, remarkable for its long wings.

Wing of Dendrobiastes basilanica: nat. size.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 300

PHYLLORNITHID”.

PuyLiornis, Boie.

72. PHYLLORNIS PALAWANENSIS, sp.u. (Plate L. figs. 1, 2.)

s ad. supra lete viridis, pileo antico magis flavicante; tectricibus alarum dorso con-

coloribus basaliter cyaneis, minimis late purpureis plagam distinctam exhibentibus,

extimis et tectricibus primariorum purpurascenti-cyaneis; remigibus nigris, extus

cyaneis, secundariis viridibus versus basin cyaneis; cauda sordide cyanea, viridi plus

minusve lavata; loris et plumis ad basin mandibule positis viridi-cyaneis; facie

laterali aureo-flava, regione parotica postica virescente; oculo plumis purpureis

circumdato ; genis albis, fasciam mystacalem formantibus; gutture toto late aureo-

flavo ; corpore reliquo subtus viridi; subalaribus cineraceis, viridi lavatis, margine

alari cyanea; rectricibus intus cinerascentibus, pogonio interno versus basin albi-

cantibus; rostro nigro ; iride saturate brunnea. Long. tot. 7:0, culm. 0°85, ale 3°5,

caudee 2°8, tarsi 0:8.

a. 6; 6,¢. 2. Puerto Princesa, Palawan. July 1874. Eyes dark.

[Shot in low bushes and trees in some numbers, eating fruit about the borders of the

clearings. They were generally in small flocks.—J. B. S.]

The sexes seem to be alike in every respect excepting as regards the moustache, which

is blue instead of white in the male.

The present species bears some resemblance to the female of P. Sonnerati, but is

' distinguished by the blue eye-ring and the blue on the wings and tail. It seems to be

quite a distinct species from any yet known.

Tora, Horsfield.

73. [oRA scAPULARIS, Horsf.: Salvad. Ucc. Born. p. 190.

a. g. Puerto Princesa, Palawan. Eyes black.

b,c. g;d. 2. Puerto Princesa. Eyes grey.

[Shot in the same locality as the Jrena, and found always in pairs.—J. B. 8.]

This is an interesting addition to the birds of the Philippines; but, like so many of

the Palawan birds, it is a well-known Bornean species.

IRENID.

Trena, Horsfield.

74. IRENA TWEEDDALII, Sharpe, Cat. B. iii. p. 268. (Plate LI. fig. 1.)

I. similis F. puelle, sed colore cyaneo vivide argentescenti-turcoso minime purpurascente

distinguenda. Long. tot. 8-0, culm. 1:05, alee 4°85, caudze 3°85, tarsi 0°7.

a. ad. Puerto Princesa, Palawan.

[This bird was eating fruit in low bushes near Puerto Princesa. A number of convicts

were hard at work making a road; and two or three of these birds were feeding in

close proximity, quite undisturbed by the noise of the men at work.—J. B. 8.]

This undoubtedly new Irena has the short tail-coverts of J. pwella, and agrees exactly

2x2

Bes MR. R. B. SHARPE ON BIRDS COLLECTED BY

with that species in the distribution of the black and blue colours ; the latter, however,

instead of being purplish, is of the most brilliant silvery turquoise-blue. I have named

this species, which bears the same relation to J. puella that J. turcosa does to I. cyanea,

after Lord Tweeddale, who has so carefully worked out the species of this genus, and

to whom we are indebted for his most excellent paper on the birds of the Philippines.

75. IRENA MELANOCHLAMYS, sp. n. (Plate LI. fig. 2.)

3. ad. similis I. cyanogastri, sed interscapulio scapularibusque nigris haud purpureo

lavatis distinguenda. Long. tot. 10-0, culm. 1:05, alee 4°8, caudze 41, tarsi 0°7.

a. &. Isabella de Basilan. Eyes carmine.

[Only seen on one or two occasions. It is apparently rather rare in the part of the

island which I visited.—J. B. 8.]

BRACHYPODIDA.

POLIOLOPHUS, gen. n.

76. PoLroLoruus vrosricrus.—Ivus ? wrostictus, Walden, ¢. c. p. 191, pl. xxxii. fig. 2.

a. 3. Negros. yes hazel; a yellow ring round the eye.

The combination of an eye-wattle and a crest seem to me to indicate that this bird

should be placed in a distinct genus from Pycnonotus or Otocompsa; and I therefore

propose the name of Poliolophus. The colouring of the bird is very badly rendered

in the plate accompanying Lord Walden’s paper—the greyish crest not being defined,

and the eye-watile entirely omitted.

CriINIGER, Temin.

77. CRINIGER FRATER, Sp. 0.

g ad. affinis C. gutturali, sed pectore ochrascenti-flavo et subcaudalibus ochrascentibus

distinguendus. Long. tot. 8°8, culm. 0-95, ale 4:0, caudee 3-7, tarsi 0°85.

a. 3. Puerto Princesa, Palawan, July 1874.

This new species is very closely allied to two others, viz. OC. gutturalis and C. gularis,

all of them having brown heads and olive-green backs. It differs from C. gularis in

being larger, with a longer beak, and in having the under surface from the throat

downwards ochraceous yellow,, varying towards pale fawn. Other characters are the

brown tail with olive-green edges, and the greyish white cheeks with silvery white

shaft-streaks. In (. gularis the ear-coverts are uniform ashy brown, the tail is dull

chestnut-red with pale ochraceous brown edgings; the under surface, with the excep-

tion of the throat and fore neck, is bright yellow, including the under tail-coverts.

To C. gutturalis the Palawan bird is very similar, but it differs in the colouring of the

under surface. C. guttwralis has the chest ashy brown washed with olive, the under

surface pale yellowish with tawny under tail-coverts; C. frater has the throat white,

and the entire rest of the under surface ochraceous yellow, including the under tail-

coverts. The flanks in both birds are olivaceous, and they are both alike in the grey

facial markings and the white-streaked ear-coverts.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 335

Hypsiprtes, Vigors.

78. HYPSIPETES PHILIPPENSIS (Gm.): Walden, #. c. p. 191.

a. 3. San Bernardino, Negros, Jan. 1875. Hyes reddish hazel.

[Feeding on the wild bananas at the foot of Malaspina volcano.—J. B. S.]

79. HYPsIPETES RUFIGULARIS, sp. 0.

3 ad. H. philippensi persimilis, sed genis brunneis haud striatis et gutture toto rufo con-

colori distinguendus. Long. tot. 9°8, culm. 1:1, alee 4°45, caudze 4°25, tarsi 0°85.

a. 3. Malamaui, Oct. 1874. yes hazel.

b. Sg. Isabella de Basilan. Eyes cherry-red.

[Feeding on fruit, and resembling Thrushes in their actions. Very abundant both in

Basilan and Malamaui, which is a flat island, only separated from Basilan by a narrow

deep channel about 100 yards broad.—J. B. S.]

This species differs from H. philippensis in its uniform rufous throat and white breast

without the conspicuous white shaft-streaks. It is also a decidedly larger bird.

TURDIDA.

Monricona, Boie.

80. Monricona sonirartia (Miill.): Walden, ¢. c. p. 192.

a. Panay.

[I got this Rock-Thrush on the top of the ranges of mountains in Panay. They were

scattered about on the mountain-sides, but were extremely shy, hopping about the fallen

timber, and disappearing on the first alarm of danger—J. B. 8. ]

Pratincona, Koch.

81. Pratincona cApRrata (L.): Walden, ¢. c. p. 198.

a. &. Panay.

[Shot in a field about halfway up the mountains in Panay, as it was flying about from

one bush to another.—J. B. 8.|

The discovery of this species in the Philippines is very interesting, as since Brisson’s

time it had not been met with there, and Lord Tweeddale appeared almost inclined to

doubt its real occurrence there.

Crrrocincia, Gould.

82. CITTOCINCLA NIGRA, sp.n. (Plate LII. figs. 1 & 2.)

3 ad. nigra, indigotico nitens; alis nigris, tectricibus alarum omnino dorso concolori-

bus; cauda nigra, quadrata, rectricibus quatuor intermediis nigris, reliquis albis,

basaliter nigris; facie laterali et corpore subtus omnino indigotico-nigris; corporis

lateribus et hypochondriis albo maculatis; tibiis nigris; subalaribus nigris; sub-

caudalibus albis; rostro nigro; pedibus nigricanti-brunneis; iride nigra. Long.

tot. 9°0, culm. 0°7, ale 3°4, caudee 4°6, tarsi 1:05.

336 MR. R. B. SHARPE ON BIRDS COLLECTED BY

g. mari similis, sed minor et abdomine albo, hypochondriis rufescentibus distinguenda.

Long. tot. 7-2, culm. 0°6, alee 0-15. caudze 3°6, tarsi 1:0.

a, 6. 3. Puerto Princesa, Palawan, July 1874. Eyes black.

c. 2. Puerto Princesa, July 1874.

The chief difference between the genera Cittocincla and Copsychus seems to lie in the

long tail of the former, this being much longer than the wing in Citfocinela but about

equal to it in Copsychus. 'Therefore the present bird is a new species of the greatest

interest, as being a true Cit/ocincla in structure with the plumage of a Copsychus; and of

course its black breast distinguishes it from C. swavis, C. macrura, &c. The difference

in the sexes is very extraordinary, and quite unlike any thing else known to occur in

this genus.

SYLVIID A.

Puyiioscorvs, Boie.

83. PHYLLOSCOPUS LUGUBRIS, Blyth: Seebohm, Ibis, 1877, p. 78.

> a. 8. Dumalon, Mindanao, Oct. 1874. yes hazel.

IT am able to state, with pleasure, that my friend Mr. Henry Seebohm, who has so

thoroughly worked out the Phylloscopi, has identified the above specimen for me as

P. lugubris. He considers that it is a young male, a bird of the year, not haying

quite completed its moult.

CIsTICOLA, Kaup.

84. CISTICOLA, sp. ?

a. San Bernardino, Negros, Jan. 1875. Eyes light hazel.

[Shot in grass in fields of sugar-cane near 8. Bernardino.—J. B.8.]

OrtHoromus, Horsfield.

85. ORTHOTOMUS FRONTALIS, Sharpe, Ibis, 1877, p. 112, pl. u. fig. 1.

g ad. supra olivascenti-viridis, uropygio paullo flavicante; tectricibus alarum dorso

concoloribus; remigibus saturate brunneis, extus dorsi colore layatis; rectricibus

olivascenti-brunneis, pallidius terminatis et macula saturatiore brunnea subter-

minali notatis et anguste flavido limbatis; fronte et supercilio angusto castaneis ;

pileo summo cum nucha et colli lateribus cinereis; facie laterali albida, regione

parotica cinereo lavata; genis et gutture toto sericeo-albis; corpore reliquo subtus

fulvescenti-albo, hypochondriis et subcaudalibus flavicanti-viridibus; tibiis pallide

castaneis; subalaribus albis flavicanti lavatis, margine alari lete flavo; rostro

saturate brunneo, mandibula flavicante; pedibus flavicanti-brunneis; iride pallide

brunnea. Long. tot. 3°7, culm. 0°55, ale 1°6, caudee 1:5, tarsi 0°75.

a. 3. ad. Zamboanga, Mindanao. yes light hazel.

b,c. 2 juw. Basilan (Isabella). Eyes grey. |

[This small Tailorbird was very abundant in the grass near Isabella de Basilan.—

J.B.8.]

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 307

The two young specimens seem to me to belong to the same species as the adult

male: but this is by no means certain; for in one specimen I can detect a slight rufous

shade on the sides of the crown, the adult O. frontalis of course not having this

colouring.

86. ORTHOTOMUS CINEREICEPS, Sharpe, Ibis, 1877, p. 113, pl. ii. fig. 2.

é ad. pileo toto cum nucha et colli lateribus saturate cinereis; loris fulvescentibus ;

facie laterali et vitta angusta malari schistaceo-nigris; regione parotica sericea

alba; dorso toto et tectricibus alarum leete olivaceo-viridibus, his majoribus letius

flavo marginatis; remigibus saturate brunneis, extus late dorsi colore lavatis;

rectricibus saturate brunneis, anguste olivascenti-flavo limbatis et terminatis, ma-

cula subterminali nulla; mento et gule lateribus albis; gula media et jugulo

nigris, plumis paucis vix albido marginatis; pectoris lateribus et hypochondriis

subcaudalibusque olivascenti-flavis; corpore reliquo subtus medialiter sericeo albo;

tibiis olivascentibus; subalaribus flavidis, margine alari lete flavo; rostro corneo-

nigricante, mandibula flavicante; pedibus pallide brunneis ; iride pallide brunnea.

Long. tot. 5, culm. 0°65, alee 1°85, caude 1:9, tarsus 0°9.

a. 6 ad. Basilan (Isabella). Eyes light hazel.

[Found in the grass near the fort at Isabella de Basilan.—J. B. 8.]

87. ORTHOTOMUS CASTANEICEPS, Wald. Zz. c. p. 195.

a. ad. Panay.

[Shot in the long grass on the mountains at a height of 2000 to 3000 feet.—J. B. 8.]

This is a perfectly good species. I have compared Dr. Steere’s skin with the type in

Lord Tweeddale’s collection from Guimaras, and find them identical.

88. ORTHOTOMUS RUFICEPS (Less.): Salvad. Ucc. Born. p. 248.

a. ad, Puerto Princesa, Palawan.

I have compared this specimen with Bornean and Sumatran examples, and can find no

difference in plumage. Total length 4°7 inches, culmen 0°65, wing 1:95, tail 1:85, tarsus

1:85. It is the first occurrence of the species in the Philippines.

MOTACILLIDA.

CorYDALLA, Vigors.

89. CoRYDALLA LUGUBRIS, Walden, ¢. c. p. 198.

a. Negros.

[Found in the open grassy plains.—J. B.S. |

Lord Tweeddale having kindly lent me the typical specimen from Guimaras, I have

been able to compare Dr. Steere’s Negros skin ; and I find them quite the same.

300 MR. R. B. SHARPE ON BIRDS COLLECTED BY ©

Antuus, Bechstein.

90. Anruus eusTavi; Corydalla gustavi, Swinhoe, P. Z. 8. 1871, p. 366.

a. Isabella de Basilan.

Having carefully compared the Basilan skin, I came to the conclusion that it was

the same as the Chinese species, which is, therefore, recorded from the Philippines for

the first time.

PARID A.

Parus, Linn.

91. PARUS AMABILIS, sp. n. (Plate LIII fig. 2.)

? ad. pileo undique cum regione jugulari nitide indigotico-nigris; collo postico et inter-

seapulio leete sulfureis; dorso reliquo pulchre cinereo; supracaudalibus indigotico-

nigris; cauda nigra, rectricibus omnibus albo terminatis, reliquis etiam extus

omnino albis; alis indigotico-nigris, tectricibus alarum late albo terminatis, fascias

alares formantibus ; remigibus albo apicatis, primariis extus cinereo lavatis, secun-

dariis intimis etiam versus apicem pogonii interni albis; corpore reliquo subtus

leete flavo; subalaribus albis, macula nigra prope marginem tectricum inferiorem

posita; rostro nigro. Long. tot. 4°5, culm. 0°45, ale 2°85, caudee 1°75, tarsi 0°75.

a. ad.; b. juv. Balabae.

The young bird is very different from the adult. It is brown above, with an olive-

greenish tinge, the head and face browner. Below it is yellow, duller than in the

adult, the throat olive; wings and tail browner than in the adult, with the same white

spotting, but not so pronounced.

92. PaRuUS ELEGANS; Machlolophus elegans (Less.), Walden, ¢. c. p. 199.

a,b. Palawan; ¢. Guimaras.

SITTID A.

93. DENDROPHILA FRONTALIS, Horsf.: Salvad. ¢. ec. p. 161.

a,b. 6. Puerto Princesa, Palawan. Eyes light chrome-yellow.

[A genuine Nuthach in its habits. I shot my two specimens during a little sojourn

that I made among the herdsmen on the opposite sides of the bay from Puerto Princesa.

They were running about on the timber logs exactly as our own Nuthach would do at

home.—J. B. 8.]

Not to be distinguished from Bornean examples.

94. DENDROPHILA @NOCHLAMYS, sp. nu. (Plate LIII. fig. 3).

2 ad. similis D. frontali, sed ubique letius colorata; collo laterali et postico cum inter-

scapulio colore pectoris vinaceo adumbratis; sed preecipue vitta angusta lorali alba

et macula parva alba ad basin mandibule posita distinguenda. Long. tot. 5-0,

culm. 0°65, alee 3:0, caudee 1°7, tarsi 0°7.

a. ad. Guimaras.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 339

{Found running up and down the trees in the manner of Nuthatches. I saw a pair,

but only managed to secure one.—J. B. 8. |

This beautiful new species resembles D. frontalis, but is larger and altogether more

richly coloured; the vinaceous colour of the underparts overspreads the sides of the

neck, and the hinder neck and mantle, the rest of the back being bluish. The chin

alone is white on the under surface; and the species is further distinguished by a narrow

white line running along the base of the lores to the eye from the beak, and by a second

small white spot at the base of the lower mandible.

NECTARINIIDA.

DIC HINA.

Diczum, Cuvier.

95. DicmuM HaMAtTOosTIoTUM, Sharpe, ‘ Nature,’ Aug. 1876, p. 297.

3 ad. supra indigotico-niger, dorso postico magis schistaceo; supracaudalibus nigris ;

tectricibus alarum dorso concoloribus, majoribus cum remigibus et rectricibus

nigricanti-brunneis, extus virescenti limbatis; facie laterali nigra, genis et corpore

subtus albis, hypochondriis cinereo lavatis; fascia preepectorali transversali schis-

taceo-nigra ; pectore et abdomine medialiter lete coccineis, illo utrinque schistaceo-

nigro maculato; subcaudalibus et subalaribus albis, remigibus intus albo margi-

natis; rostro et pedibus nigris. Long. tot. 3°7, culm. 0°45, ale 11, caude 1°2,

tarsi 0°55.

a, b. ad. Guimaras.

c, d, e. ad. Negros.

fg. &- Negros. Hyes hazel.

[Found in all cases in the cocoa-nut trees, flying about and perching on the bamboo

receptacles which are hung under the blossom-stems to catch the juice. The natives

said they were feeding on the latter; but whether this was the case, or whether they

were eating the insects which were attracted by the juice, I was unable to determine.—

J. B.S. |

Two young birds differ from the adult in being slaty grey above, the wings and tail

blacker, with greyish margins to the feathers; under surface of body ashy whitish,

washed with yellowish on the abdomen, and with greyish on the sides of the body. In

one specimen the red colour on the under surface is appearing, and the blackish feathers

on the upper surface: in both the lower mandible is yellowish at the base.

Guimaras and Negros specimens are identical; but the slaty black on the fore neck

and breast varies in extent. It is, however, well developed in every specimen, and

shows clearly that the species is not Diceum papuense of Mindanao, which is the bird

Mr. Gould has figured as the female of his D. retrocinetwm.

96. Dicatm HYPoLEUCUM, Sharpe, ‘ Nature,’ Aug. 1876, p. 298.

3 ad, supra niger, pileo vix viridi nitente; facie laterali collique lateribus nigris; genis,

et corpore subtus toto cum subalaribus subcaudalibusque albis; als caudaque

SECOND SERIES.—ZOOLOGY, VOL. I. - 2 2

340 MR. R. B. SHARPE ON BIRDS COLLECTED BY

nigris ; rostro et pedibus nigris; iride saturate brunnea. Long. tot. 8-6, culm. 0:5,

alee 2°1, caudee 1:05, tarsi 0°55.

is Be MValanaau October 1874. Eyes hazel.

b. $. Isabella de Basilan. Eyes hazel.

[Found in the mangroves in the same localities as the Nectariniide.—J. B. 8.]

This species is distinguished at a glance by its uniform white under surface.

97. DiczuM DORSALE, Sharpe, ‘ Nature,’ Aug. 1876, p. 298.

3 ad. supra lete schistaceo-cinereus, pileo clariore; plaga interscapulari aurantiaco-

miniata; tectricibus alarum dorso concoloribus, extimis anguste olivaceo lavatis;

remigibus nigricantibus, extus dorsi colore marginatis, secundariis paucis olivaceo

limbatis, intimis dorso concoloribus; supracaudalibus schistaceo-cinereis; cauda

nigra, rectricibus schistaceo limbatis; loris nigris; facie laterali saturate schistacea :

subtus leetissime aurantiacus, gula et abdomine imo flavis; subalaribus albis, axilla-

ribus pallide flavis; tibiis intus cinereis, extus flavis; rostro nigricante, mandibula

versus basin pallida; pedibus saturate brunneis. Long. tot. 3°5, culm. 0°5, ale 1:9,

caudee 1:05, tarsi 55.

? a mari diversa: supra olivascenti-viridis, uropygio magis flavicante, facie laterali

pileo concolori; subtus leete flavicans, pectore et subalaribus leetioribus. Long. tot.

3°7, culm. 0°5, alee 1:9, caudee 0°9, tarsi 0°55.

a.g. Panay.

[Shot in the highest part of the island, in the remains of the virgin forest on the highest

range of the mountains west of Ilo Tlo.—J. B. 8.]

The bird which I consider to be the female of this species has unfortunately lost its

label. The only Dicewm with which this new species can be confounded is D. trigono-

stigma; but it is at once distinguished by its yellow throat and blue lower back and

rump.

98. DicmuM TRIGONOsTIGMA (Scop.): Salvad. ¢. c. p. 166.

a. 6 ad. Negros.

The appearance of this species in Negros is somewhat remarkable. As the specimen

entirely agrees with others from Borneo, it might have been expected to occur in

Palawan.

PRIONOCHILUS, Strickl.

99. PRIONOCHILUS, sp.

a. 2 juv. Puerto Princesa, Palawan. Hyes dark.

The only specimen obtained by Dr. Steere is a young bird which I'am unable to refer

with certainty to any species. It is evidently the young of a bird belonging to the same

eroup of Prionochilus as P. percussus and P. santhopygius; and, from its yellow spot

on the rump, it must be closely allied to, if not identical with, the latter. The occurrence

of the genus is interesting; but the exact species can only be determined on the exami-

‘nation of the adult.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 3841

ARACHNOTHERA, Temm.

100. ARACHNOTHERA DiLuTIOR, Sharpe, ‘ Nature,’ August 1876, p. 298; Shelley,

Monogr. Cinnyr. part 3.

ad. supra sordide olivascenti-flavus, uropygio paullo cinerascente,

brunneo, collo postico vix olivascenti lavatus; alis cinerascenti-brunneis, tec-

tricibus vix olivaceo limbatis, remigibus extus olivaceo lavatis, secundartis latius ;

cauda saturate brunnea, rectricibus olivascenti-flavo marginatis, extimis versus

apicem cinerascentibus, tribus externis pogonio interno albo apicatis; facie laterali

saturate brunnea, loris nigricantibus, plumis ad basin mandibule positis et mento

albicantibus; gutture albido, cinerascenti lavato; corpore reliquo subtus flavi-

canti-albido, subcaudalibus albis; fasciis pectoralibus leete aurantiacis ; subalaribus

albis vix flavido lavatis; remigibus subtus cinerascentibus, intus albo marginatis ;

rostro nigricante, mandibula basaliter albida; pedibus saturate brunneis; iride

saturate brunnea. Long. tot. 5-7, culm. 1:5, alee 2°7, caude 1:9, tarsi 0°75.

Eyelids yellow.

pileo saturate

a. 3. Puerto Princesa, Palawan, J uly 1874. Eyes dark. Breeding.

[Shot in the mangroves, but not very abundant.—J. B. 8. ]

Cinnyris, Cuvier.

101. Cryyyris sppratus: Nectarophila sperata (L.), Walden, ¢. c. p. 200.

a,b. § 2. Negros.

c-t. g ad.; k. § juv. Guimaras.

[The pair obtained in Negros were shot together flitting about a dead tree about

20 feet in height, and were apparently nesting. In Guimaras it was very common, hover-

ing about the bamboo cups which are hung up to catch the sap in the cocoa-nut

blossoms.—J. B. 8.]

102. CrynyRis sUGULARIS (L.): Arachnechthra jugularis, Walden, ¢. c. p. 200.

a,b,c. g. Negros.

d,e,f.3 2. Panay.

g. 2. Zamboanga, Mindanao. Eyes hazel.

EUDREPANIS, Sharpe.

Allied to Hthopyga,- but at once distinguished by its rounded tail, which has not the

elongated central feathers of the above-named genus. The type is

103. EUDREPANIS PULCHERRIMA, Sharpe, in Shelley’s Monogr. Cinnyr. part 3.—ditho-

pyga pulcherrima, Sharpe, ‘ Nature,’ 1876, p. 297.

g ad. supra saturate olivascenti-viridescens, plumis basaliter griseis quasi marmoratis ;

plaga frontali letissime metallice violacea; capite laterali nigricaate, linea supra-

parotica metallice violacea; dorso postico et uropygio lete sulfureis ; supracauda-

libus metallice chalybeo-viridibus, rectricibus nigris hoc colore layatis: subtus

lete sulfureo-flayus, macula magna preepectorali miniata; subalaribus albis ; sca-

222

BA2 MR. R. B. SHARPE ON BIRDS COLLECTED BY

pularibus dorso concoloribus; tectricibus alarum metallice viridibus; remigibus

nigricanti-brunneis, extus olivaceo lavatis: rostro nigro; iride coccinea. Long.

tot. 3°5, culm. 0°75, alee 1:8, caudze 1-05, tarsi 0°55.

a. $. Basilan, Isabella. Eyes carmine.

Shot in the mangroves near the town of Isabella.

104. dAitHopyeA MAGNIFICA, Sharpe, ‘ Nature,’ August 1876, p. 297; Shelley, Monogr.

Cinnyr. part 3.

44, similis 4. flavostriate, sed major, rostro fortiore, abdomine et subalaribus nigris

distinguenda. Long. tot. 5:0, culm. 0°95, alee 2°3, caude 2:0, tarsi 0°6.

a-e. Negros.

This is a large representative of Athopyga flavostriata from Celebes, but is distin-

guished at once by its large size and blackish under wing-coverts and belly.

105. AirHopyeA SHELLEYI, Sharpe, ‘Nature,’ August 1876, p. 297; Shelley, Monogr.

Cinnyr. part 3.

g ad. pileo circumscripte chalybeo; loris nigris; facie laterali tota cum colli laterali

et postico et interscapulio saturate scarlatinis; dorso olivascenti-viridi, tectricibus

alarum minimis dorso concoloribus; tectricibus majoribus remigibusque brunneis

extus olivaceo marginatis; plaga uropygiali lete sulfurea; supracaudalibus et

rectricibus duabus centralibus lete chalybeis, reliquis nigris extus chalybeo mar-

ginatis: subtus lete sulfureus, vitta malari chalybea; gula quoque flava, utrinque

searlatino marginata; pectore scarlatino striolata; subalaribus albis. Long. tot.

4-1, culm. 0°65, ale 1°85, caudz 1°85, tarsi 0°55.

a, b, c, d. Puerto Princesa, Palawan, July 1874. yes black.

e. Balabac.

[I met with this species on both the above-mentioned islands. The first one I shot

was in the town of Puerto Princesa, in the mangroves, fluttering about among the

blossoms. I afterwards fell in with it a few miles inland in the forest. In .Balabac I

shot it also in the mangrove.—J. B. S.]

106. ANTHREPTES MALACCENSIS (Scop.): Salvad. ¢.c. p. 178.

a,b,c. 8; d,e. 9. Puerto Princesa, Palawan. Eyes carmine or cherry-red.

~ The specimens collected agree with Bornean examples.

107. ANTHREPTES CHLORIGASTER, Sp. n.

A. similis A. malaccensi, sed major, capite colloque lete metallice viridibus nee purpu-

rascentibus, dorso reliquo et plaga magna humerali leetissime violaceis, tectricibus

alarum majoribus clare castaneo marginatis; remigibus et gutture ut in 4. malac-

censi picturatis; corpore reliquo subtus viridiscenti-flavo ; cauda nigricante, metallice

violaceo late lavata. Long. tot. 5-2, culm. 0°75, alee 2°75, caudee 1:95, tarsi 0°7.

a,b. 3. Negros.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 343

[Not found in mangroves, like 4. malaccenis, but feeding on the blossoms of the

cocoa-nut trees.—J. B. 8.]

These two specimens, collected by Dr. Steere, are precisely similar the one to the

other; and as they differ from a series of at least twenty specimens of 4. malaccensis,

I have no doubt that the green breast is a constant character, and that the species is

therefore distinct.

CORVIDA.

Corone, Kaup.

108. CoRONE PHILIPPINA (Bp.): Sharpe, Cat. B. iii. p. 42.—Corvus philippina, Bp.:

Walden, ¢. ¢. p. 201.

a. Negros.

[Crows were found in most of the islands; but I only preserved one specimen. The

native name is ‘ Quawk,’ from the note. I saw birds apparently of this species in Negros,

Zebu, Panay, Mindanao. In Negros they were very plentiful about the house where I

was staying, and very impudent, coming down into the yard and stealing the jerked

beef which was being cured at the time.—J. B. 8.]

Dr. Steere’s specimen measures—total length 18°5 inches, culmen 2°45, wing 11:9,

tail 8:0, tarsus 2°05.

STURNIDA.

STURNIN A.

StTuRNIA, Lesson.

109. STURNIA VIoLACEA (Bodd.): Walden, ¢. c. p. 203.

a. Negros.

6. 6. Dumalon, Mindanao.

[In large numbers in flocks in both the above localities.—J. B. S.]

This Starling was not before known from either of the above islands.

Catornis, G. R. Gray.

110. CALORNIS PANAYENSIS (Scop.); Walden, ¢. c. p. 205.

a. Panay.

b,c. 6 9. Maasin, Panay, Jan. 1875. yes light carmine.

[In large flocks in the open fields, feeding in the cultivated grounds. They were tearing

to pieces the flowers of a leguminous tree.—J. B.S.]

111. Catornis cHALYBEUS (Horsf.): Salvad. Uce. Born. p. 271.

a. °.Puerto Princesa, Palawan. Hyes carmine.

b. $ juv. Puerto Princesa. LHyes grey.

The only adult specimen procured by Dr. Steere is a female in rather dull plumage.

It seems to me, however, to agree sufficiently well with a hen bird sent by Mr. Low

from Labuan; but it appears to be a little larger in its measurements than the Bornean

birds, the dimensions of Dr. Steere’s specimen being as follows—total length 7°6 inches,

culmen 0°85, wing 3°95, tail 2°65, tarsus 0°9.

B44 .. MR. R. B. SHARPE. ON BIRDS COLLECTED BY: :

EULABETINA.

Sarcops, Walden.

112, SARCoPS CALVUS (L.): Walden, ¢.c. p. 205.

a, b, c. Negros.

d. 8. San Bernardino, Negros, Jan. 1875. Eyes reddish hazel ; bare parts flesh-colour.

[ I met with this bird in great abundance in the upper part of the island of Negros,

frequenting the groves of wild bananas, and feeding on the fruit of these trees.—J. B. 8.]

118. Sarcops Lowit, sp. n.

S. similis S. calvo, sed interscapulio argentescenti-griseo nec nigricante, et corpore

subtus argentescenti-griseo distinguendus. Long. tot. 12°5, culm: 1:15, ale 5:1,

caude 5:0, tarsi 1:3. |

a. Si Butu, Sulu archipelago.

This species appears to be distinguished from the ordinary Sarcops calvus by its grey

mantle, which resembles the rest of the back, and by the colour of the under surface,

which. is silvery grey with a narrow blackish line down the centre of the body,

whereas in S. calvus the mantle is black, and the under surface also, ouly the flanks

being grey.

Gracuna, Linneus.

114, GRACULA JAVANENSIS (Osbeck): Salvad. ¢. c. p. 274.

a, Puerto Princesa. ;

6, c. 8 ¢. Puerto Princesa, Palawan. yes black, bill red, wattles bright orange-yellow.,

[These Minas were not common in Palawan, but were found in the high trees in the

jungle, making discordant noises. The natives catch them and cut their tongues, in

order to make them 'talk.—-J. B. 8.]

Identical with Bornean specimens.

EURYLAMIDA.

‘SARCOPHANOPS, gen. 0.

Exactly resembling Serilophus in general character, but differing at once in the large

wattle round the eye. The type is

115. SARCOPHANOPS STEERI, sp.n. (Plate LIV.)—Hurylemus Steerii, Sharpe, ‘ Nature,’

August 1876, p. 297. |

é ad. supra saturate cinereus ; uropygio et supracaudalibus nitide purpurascenti-vinaceis ;

cauda leete castanea ; capite summo nitide et saturate purpurascenti-brunneo ; torque

collari et colli lateribus albis; loris albis capitis colore apicatis; facie laterali et

gula tota nigris; corpore reliquo subtus lilacino, hypochondriis vix vinaceis, ab-

domine imo albido; subcaudalibus fulvescentibus; tibiis cinereis; axillaribus albis;

subalaribus nigricantibus; alis nigris, secundariis intimis basaliter albis, extimis

extus aurantiaco-flavis, fasciam alarem transversalem exhibentibus; remige primo

PROF. STEERE IN THE. PHILIPPINE ARCHIPELAGO. o45-

extus versus basin auguste albo; caruncula opthalmica pallide indigotica; rostro

pedibusque pallide indigoticis. Long. tot. 6:8, culm. 0:9, ale 3°45, caudse 2°75,

tarsi 0°85.

? ad. mari similis, sed pectore albo nec lilacino distinguenda.

a,b. 3; c. 9. Isabella de Basilan. Bill, wattle about eyes, and feet light indigo-blue. yes

colour of gold-stone.

- [I only fell in.with this species on one single occasion in the island of Basilan, when I

found a little flock of about seven individuals in the thick deep jungle towards the

centre of the island. They were feeding on fruit, and were very tame, having probably

never been disturbed before by a gun, and did not take flight until I had secured three

of theirnumber. Of all the birds I ever saw, this one has the most beautiful eye. I can

only describe it as being of the colour of “ gold-stone;” that is to say, it was like a piece

of clear crystal crowded with specks of gold. All three specimens had the iris exactly

the same.—J. B. 8. ]

This is the gem of Dr. Steere’s discoveries. The eye-wattle, the peculiar coloration,

the difference in the sexes, render the genus entirely distinct from all the other members

of the Hurylemide.

PLOCEIDA.

Pappa, Reichenbach.

116. Pappa oryztvora, L.: Walden, ¢. ec. p. 207.

a. Zamboanga, Mindanao.

[I saw plenty of these in little flocks of from five to seven, perching on the roofs and

nesting in the latter.—J. B. 8.]

Identical with Javan examples.

Munta, Hodgson.

117. Munta arricaPrLua (V.): Salvad. Ucc. Born. p. 265.

a, 6. Zamboanga, Mindanao. LHyes reddish hazel.

Dr. Steere’s example is not referable to If, Jagori, Cab., according to the diagnoses

given by Lord Tweeddale (¢. ¢. p. 207), as it has a black head. The black on the belly

extends right up to the throat in a continuous patch, this perhaps being a character of

the male only. After examining a large series of these chestnut-and-black Finches in

the British Museum, I have come to the conclusion that Lord Tweeddale’s identifications

are correct; but I believe that with increased material the number of species will have to

bereduced ; at present, however, as Dr. Steere’s specimen is clearly not referable to

true W. Jagori, it must be kept as M. atricapilla, with Labuan examples of which it

agrees best, having the same black head.

118. MuniA LevcocAsTRA (Blyth) : Salvad. Uce. Born. p. 267.

a. %. Puerto Princesa, Palawan. Hyes dark red.

[In large flocks in the rice-fields, where they made great havoc.—J. B. 8.]

I have compared the above specimen with Malaccan individuals, and have found them

identical.

346 MR. R. B. SHARPE ON BIRDS COLLECTED BY

COLUMBA.

TRERONIDAN.

TRERON, Vieill.

119. TrERON Nastca, Schl.; Salvad. Ucc. Born. p. 288.

a. 9. Puerto Princesa, Palawan. Eyes orange.

I have compared this specimen with a Sumatran pair collected by Mr. Wallace, and

find them identical.

OsmoTRERON, Bonaparte.

120. OSMOTRERON VERNANS (L.): Walden, ¢. c. p. 210.

a, b. Maasin, Panay.

[Shot in cane-fields and open thickets.—J. B. 8. ]

Panay is a new locality for this species.

121. OsMOTRERON AXILLARIS (Gray): Walden, é.c¢. p. 211.

a, Zamboanga, Mindanao.

b. Negros.

[Very abundant in both the above places, and called by the Spaniards “ Llorosa,” or

the “ weeper,” from their mournful note. ‘They were flying about in flocks and feeding

on fruit.—J. B. 8. |

This Fruit-Pigeon was not before known from Mindanao.

PHABOTRERON, Bonaparte.

122. PHABOTRERON NIGRORUM, sp. 0.

P. similis P. leucoti, sed gula et plaga faciali pallide cervina distinguenda. Long.

tot. 9:2, culm. 0°7, alee 4°95, caude 49, tarsi 08°5.

a. &. San Bernardino, Negros, Jan. 1875. yes with a narrow rig of carmine;, crop full of

Chil peppers.

[A Ground-Pigeon, by no means uncommon, but difficult to preserve on account of

the looseness of the plumage. It was found in the corners of the cane-fields, where the

red peppers had grown up wild.—J. B. 8.]

On comparing Dr. Steere’s adult male bird with those from Luzon in Lord Tweeddale’s

collection, it appears to differ in having the throat pale fawn-colour instead of rufous,

and the cheek-stripe is fawn-colour instead of white. As other Negros birds in Lord

Tweeddale’s cabinet exhibit the same differences, we may fairly suppose them to be con-

stant; and in that case the species requires a name.

CarpPoPHaGa, Selby.

123. CARPOPHAGA NEA (L.): Walden, ¢. c. p. 215.

a. Negros.

b. 9. Sau Bernardino, Negros, Jan. 1875. Eyes carmine.

c. Puerto Princesa, Palawan.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 347

[Pretty generally distributed, as I saw it in most of the islands, Mindanao, Palawan

and Balabac, as well as in Negros. It was feeding in most instances on the fruit of the

wild fig. —J. B. S.]

This wide-spread species probably occurs on all the islands.

Myristictvora, Reichenbach.

124. MyRIsTiIcIvora BICOLOR (Scop.): Walden, ¢. ¢. p. 217.

a. 6. Balabac. yes bright red.

[This species was plentiful in Palawan and Balabac. In both islands they were found

at certain times of the day coming down to the mangroves along the sea-shore in large

flocks. What they were doing there I could not find out, unless they found something

to feed upon along the beach. Off Balabac and Palawan were several sandy islands

where the larger Pigeons nested in large numbers; and during my stay at Balabac I saw

the natives returning from an expedition to one of these with a number of young birds

and eggs, probably of this and the preceding species.—J. B. 8.

Hemrenaca, Bonaparte.

125. HEMIPHAGA POLIOCEPHALA (Gray): Walden, ¢.¢. p. 217.

a. San Bernardino, Negros.

[Found in high woodlands in pairs or two or three together and not in flocks.—

J. B.S.]

COLUMBIDA.

Macropyeia, Swainson.

126. MAcROPYGIA TENUIROSYRIS, Gray, Walden, ¢.¢. p. 218.

a. &. Isabella de Basilan. Eyes hazel.

6. Negros.

[Shot near streams. They seem to frequent rocky ground in the neighbourhood

of water.—J. B.8.]

I have compared the above two specimens with others in Lord Tweeddale’s collection

from Luzon and Negros, and could not find any difference in the Basilan example.

TurtuR, Selby.

127. TurTUR DUSsUMIERI (Temm.): Walden, ¢. ¢. p. 218.

a. 2. Maasin, Panay, Jan. 1875. Eyes brick-red.

[This species is found in the open fields and roads in pairs upon the ground.—J. B. 8.]

Panay is a new locality for the species.

128. Turtur HUMILIs (Temm.): Walden, ¢. c. p. 219.

a. Mount May-hay-hay, Luzon.

SECOND SERIES.—ZOOLOGY, VOL. I. 3A

348 ' MR. R. B. SHARPE ON BIRDS COLLECTED BY

GOURIDA.

PHLoceNas, Reichenbach.

129. PHLOGaNAS LUZONICA (Scop.): Walden, 7. c. p. 221.

a, b, c. Manila.

[The natives snare these birds in some numbers; and all my specimens were procured

from a friendly priest, who gave them to me out of his aviary. I saw the same or an

allied species in numbers at the north end of the island of Negros.—J. B. 8.]

CHALCOPHAPS, Gould.

130. CHatcopHars tnpIca (L.): Walden, ¢. c. p. 221.

a. 9. Maasin, Panay, Jan.1875. yes dark.

b. g. Puerto Princesa, Palawan. THyes dark.

[This Dove seemed to me to be a common Pigeon in most of the islands.—J. B. 8.]

GALLIN &.

PHASIANIDA.

Gaus, Linnzeus.

131. GaLLus BANKIvA, Temm.: Walden, 7. c. p. 223.

a § ad. Panay.

[I also saw Jungle-fowl in Mindanao, where they were plentiful, and also in Palawan

and Balabac. They are difficult to shoot, as they are very wary; and I have hunted for

hours with birds crowing all about me without getting a shot. The Indians of the

forest-villages snare them, and often keep them alive, crossing them with the domestic

fowls.—J. B. 8.]

MEGAPODIIDA.

MeEcApopius, Quoy et Gaimard.

132. Mrcapopius Ltowtt, Sharpe, P. Z.8. 1875, p. 111.

a. @. Dumalon, Mindanao, Oct. 1874. Eyes hazel.

b,c. pull. Balabac.

[In Balabac I found large numbers of these birds, and procured both eggs and young.

They were nesting on the sea-beach on the edges of the brush, the nests being

about 12 feet across and about 8 or 4 feet in height, and made of sand and bits of

shells and sticks. The natives. say that sometimes 50 or 60 eggs are found in one

of them. In Mindanao, in the mountainous country, some way inland, I found

heaps of earth piled up against fallen logs, which the natives said were Megapodes’

nests; I dug into them myself, but did not find eggs. On one occasion the nest had

been burrowed by a huge land-lizard.—J. B. S.]

The Megapode collected by Dr. Steere is not the true MZ. Cumingi of Manila, but

appears to be identical with I. Lowit of Labuan. It is a very dark-coloured bird,

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 349

leaden grey underneath, and measures as follows—total length 12 inches, culmen 1°15,

wing 9°38, tail 3°6, tarsus 2°55. The dimensions do not agree with those given by me

(P. Z. 8. 1875, p. 111); but the dark leaden-grey under surface agrees exactly with

Labuan birds.

GRALLA.

CHARADRIIDA.

CHARADRIUS, Linneeus.

133. CHARADRIUS FULVUS, Gm.: Walden, ¢. c. p. 226.

a. Balabac, Sept. 1874.

RALLIDA.

OrTYGOMETRA, Linnzeus.

134. ORTYGOMETRA CINEREA (V.): Walden, ¢. ¢. p. 230.

a,b. g 9%. Naga, Zebu Island. Eyes reddish hazel; feet olive-yellow.

[These birds were found in a grassy wet marsh; they were’ actively running about

through the grass and water, flying short distances, and alighting on tussocks. They

were in great abundance.—J. B. 8. |

This is the first exact occurrence of the species in any of the Philippines; though

specimens were obtained by Mr. Cuming in the archipelago, no island was recorded as

the habitat.

CICONIID.

MELANOPELARGUS, Reichenbach.

135. MELANOPELARGUS EPIscoPUS (Bodd.) : Salvad. Uce. Born. p. 356.

a. Negros.

[This Stork was killed out of a flock of fifteen or twenty individuals as they flew low

over the mangroves towards the high lands in the interior, and apparently on their way

to roost.—J. B. 8.]

This is the first time that the species has been noticed in the Philippines.

ARDEIDA.

Busuucus, Pucheran.

136. BuUBULCUS CoROMANDUS (Bodd.): Walden, ¢.c¢. p. 237.

a. Puerto Princesa, Palawan.

[White Egrets were noticed in the rice-fields in nearly every island.—J. B. S.]

Buroriwes, Blyth.

137. Buroripes savanica, Blyth: Walden, ¢. c. p. 287.

a. Negros.

Gorsacuivs, Bonaparte.

188. GoRSACHIUS MELANOLOPHUS (Rafil.): Walden, ¢. ec. p. 288.

a. Dumalon.

oAQ

390 MR. R. B. SHARPE ON BIRDS COLLECTED BY

[Found at some distance from the water, running along a road in the forest, where

there were some shallow puddles of water.—J. B. S.]

LARIDA.

STERNINZ.

STerNA, Linn.

139. STERNA BERGII, Licht.: Saunders, P. Z.S. 1876, p. 657.

a, b. Balabac, Sept. 1874.

[In abundance, flying round the coast of Balabac in flocks.—J. B. 8.]

The specimens were seen by Mr. Howard Saunders; and he pronounced them to be

of this species.

The following Table is drawn up on the model of Lord Walden’s, and embraces all the

species known to inhabit the Philippines. The new species discovered by Dr. Steere are

here printed in small capitals; and those which he has added for the first time to the

Philippine avifauna are printed in italics.

Philippines.

| Polynesian Region.

| Malay peninsula,

| Australian Region.

|| Papuan Region.

| Panay.

| Zebu.

| Camiguin.

| Mindanao.

| Basilan.

| Malamaui.

| Cujo.

| Palawan.

| Balabae.

| Sulu archip.

| Celebes.

| Borneo.

| Jaya,

| Sumatra.

| Timor.

| Tndia.

1. Cacatua hematuropygia...............- a

2. Prioniturus discurus.................-. 5

3. Tanygnathus luzoniensis ..............-+ no

Cyclopsitta lunulata..................6 30

Loriculus philippensis.................- ae

9 TEAWUS)s os oo cGo0dD000000000G0 Pe Bloc

3 ORO, oo00000900000000000 oA[o allo olfoo

99 WATT 6 cong o0g00n0n0000008 sallsailorolloallae

i chrysonotus).))..)ece le - ee o.dlloa|lolle alloo

Wal cousevertisn was sececnaisctoetee tater: aol een lteol cles

. Microhierax erythrogenys is

> ZNSE THMIVAUSFNTS) 5 6505G0000000000000bnN0 #

pm. SOWIE. cosccccansca0go00005000 %

5 ANCE WARM 3 2g00550000000000000 .

. Spizaetus philippensis.................-; As

. Spilornis holospilus

p IBMT THON 55 4a56509000Gn00b00900

. Haliaétus leucogaster

. Haliastur intermedius

. Klanus hypoleucus

21. Baza magnirostris

22, Circus melanoleucus :

23. ,, spilonotus

24, Bubo philippensis...................00-

25. Scops megalotis

26. Ninox philippensis

27. Scelostrix candida

28. Thriponax javensis

29. Mulleripicus funebris

30. Chrysocolaptes heematribon o

31. ” xanthocephalus.......... 30

32. oy lucidus

x x

5 ee

Eom

Lok * | Guimaras.

: & *% * | Negros.

> & & & | Luzon.

SSRN STR ONE SOON OU

LR Re eT

Ho RO

34, ERYTHROCEPHALUS ......|.. dolla alle allo.alla allo olla allo ollo offo- off 3

BD. Tig@ Javanensts ..... 0... cece cee neeenac cles

36. Jungipicus maculatus ..

Oipmulannactesiard cuore rence niet rerit te

38. Merops philippinus

32. 4, _ bicolor

ae =

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. abl

Philippines. a & é| <

A A 2 a a %

a EWEN lel VAI SIEW ei ay Wel 8.2/5

25/2) 2) s\2)5| 2) 2) .|2lS| 2/218) [Eel slel ele):

5/5 /2|2/3/8/2| Zl eislalelals leis lais|alzlelsie

ARS) aisl/Olalalalolalalalolals jaja lala |4| ala

. Eurystomus orientalis ..........4..--. wel xe x |e le ele | oe * wlelaelalelelele

. Alcedo bengalensis........ siotehaied Bs sper: walle n]e [ae ]e feo] x as weleelelalalelel se |

. Aleyone cyanipectus ........... dsio.o ce %

3. Pelargopsis gouldi ............+....... ~-|%

> gigantea ......... Uboonouda cafeefe fe c|e ete te [ae fe cle ele ole ole ol xe

on leucocephala......seveeeeees calls alla c[oalle-ollo allsolfo clle alla allo alles |

pCeyxc rE, 58o55cqub0duub 00000000 calks ile olocleolloalloalies |

oy TEORCGUD, cogannosdon Usa oonagee Halen cts cect alle ate os cle lente ele fool «lle | ae |x |. «tae

» philippinensis ...... vooosodsgbee - «| x

. Halcyon gularis .......... bon oodocd ot ela fale

ULE coanacooUudo Arevads cored’ *

» coromanda ...... Eiainrlahe renee *

59 chloris ....... Boisetitancte ge tieh ony eeloe leet lala}. ele |e

. SWANCHELIL | cial sveyerctals stele a siete wefecfe ete efecto ete ol= «| se

Ps ombroninemm mer iecireyastr colle aiiagiio dliaciloollac| |e

: is JOHAGRENT, oon eugcdncncaogeccus Bae |

. Santholema hemacephala ...........- sly vale] x

: a TOSCAM Eira rtene heleysloraloetobst ate cclleulercllo oles

. Macropteryx comatus.................. wef fee} ole * |e]. .|*] %

. Collocalia troglodytes ................ ole |e

. Lyncornis macrotis......... oobuooooeaD ole 1

. Caprimulgus manillensis .............. Beles “ #

$n ise Gegoqooou50s00d5 *

. Cacomantis merulinus ........ RO ores Oalltd Pa\les|lo allo alloallcallaallo alle»

. Chalcococcyx amethystinus ............|-- *

. Chrysococcyx basalts .......0e eevee eens |e n allo olloall es

. SURNICULUS VELUTINUS .............-|++ sallecliocle llealle alla-alioolles

. Hierococcyx strenuus .......-.......- aoe

b op [eee] Fasccoocogonge0o *%

. Eudynamis mindanensis .............. wale efaefae fae fe |e eda

. Desylophus superciliosus .............. os | xe

Lepidogrammus Cumingi..............- «le

. Centrococeyx viridis .................. welacle el} | elle ele

. Dryococcyx HARRINGTONI .......... oro slate oallo elloal lal ection lana cea eretle3

Pyrrhocentor melanops.............+.. Sallartic olla clic olla olla oll eslts oll es

. Buceros hydrocorax ..........00++005 calles j

. Craniorrhinus leucocephalus............ 6 lle allo ollo cllo ollo ol [Eales i

Ry INV/ATED ENT ae ater lal-eietelel oes colon |e

. Penelopides panini.................-5- ele lela x

} op MEMES 6 GooeGanddcocec wa feede

} ILENE REBT! Soocboo0a0g6000000C0 Gallo oljoallfalle olle alles |

9 schach . tie tle alten eee ee eeee nee #

», lucionensis ..... DO AOE TOO RCS tio Se ae ee ae ea (ae

. Artamus leucorhinus.................- colacde-lalade ete -lale-le> wxlelaelale-le dels %

. Graucalus striatus ......)..02.-ce eee es edlesloaic slesleeioelesio alle

i ue SUMAETENSIS occ e cece scenes Jalloollodieclloale alloca atloolle clio alles

. Wolvocivora cerulescens .............. alse jlo

. Lalage dominica eee wee ew tere ewe eee eof] HR] sede Le fe ef fee] ele ele fe ele ele ele Paes Kyle sfe ole Ve

. Pseudolalage melanoleuca.............. ole

. Hyloterpe philippinensis .............. --|%

. Pericrocotus cimereus.............20005 Alls alloallactio alle ole clla allo elle allaolle allo clo aloo eatlaniea les

é + UGNEUS: tet AA areh ah sPayecchaveeecn seat: delle aloollecleollocleclscloelaciocl fala cloniocllestaicila cles

. Dicrurus balicassius ..........0.0+000- Boies

Bs mirabilis ...... araedarshisrats aeelocatacs dollo olloclfe allie

« Buchanga cineraced 1... eevee veeneeeee salle cl[ealla clio clloollectlaa|is alle clo alles

. Philentoma cyaniceps ..............-- sallestoollocie ooo cles

b PP ABINAORABS coacoasconodoo Ballo alls oles

- Leucocerca nigritorquis..............-.. sllfsllodleale olleslleel|eello ello allo oll es

. Cyornis philippinensis ................ wef alate e]e | ae

. Hypothymis azurea .................. Salts |loolles lee lle clloclle alles|locilacilaclioollo alloallealtes||ealESilo all £2

% SUPERCILIARIS............ pollocicollo cic clac|laelloclies

. EUMYIAS PANAYENSIS .......0.000.002 solle sllee

. Butalis manillensig..................6% Bela

. AanthopyGid NArctssind . 1... .veeesaeees Jellocle ile clocliaallocl|e3

. XANTHOLESTES PANAYENSIS .......... Be) loo

A SE PARTAWR UTC AUDA) se) </dsteleyo1 4 siete crosses BallNollbollecoiio allaallen| tac lee

b ADENINE HUNKS 5 GogopeoguuHueoudouoc le

MR. R. B. SHARPE ON BIRDS COLLECTED BY

Philippines.

z a/8|.|3 Ae

Jel 3/3) 18/8] 8l 2 slslzleis| |é

aE: SBA HEBEAEOE

AJAIS(A\8|S5/4/S ls l6lalalalslals|a

107. ZEoCEPHUS CINNAMOMEUS ...........-- calleelico|lp ilo dlloallerg|loolle

108. op CYANESCENS .........-.--- selloHoolloclle sto Wo elloolle olla allo alles

WO9. Berundo rustica . 00.5. c en cence sec csee dol latallaiollacollpalloval enlle™ ;

110. ” BMV), 6 cog ood oocnd05n0e. Solloa| alo oliodipclnelaoloclieciioaissllsallaallesi|=s|lasiies

111. Cecropis daurica...........-. soo0eccoe «lig Seale

112. Oriolus chinensis........ goaderegabende Solles|le allsl celica hall ealle allo allo ollo cll =

1B 5) SMIIMEDTs cong congeaagdoxcce00c Melleclloaleellerlecleellectiealloellecte cls nes

114, oj PUMAIDIGRL oS Hacamgodooecoadeee oellb olla oleic lo alle pllo oles

1B, og, olay Sep oo obconax0¢00K0 %

116! Durdus obsemrus). shes... eden eee snl Re

117. > chrysolaus).-. 2... Cee Gece *

118. Brachyurus erythrogaster ..,........... soll es

119. 9) PROPINQUUS ...+.ssseeeee- pollee|io clic clo clic cla cll Sailo olla allo cic oltex

120. n SOWGWGWS ooaoaondooggenoace salaeiia als alae twstae te cl als ale ll ae

121. 5 STEER ...... Aeharic Chunagsts arllaRerateraltteltexa em itae

122. MACRONUS STRIATICEPS ..........+... 00) o6|o alo alio alle ole el[o of ell ee

TOR WooxorENAs| NWWOOWE ooocosgapac eo cascade 2 ele le PP ka

TOM, [Parra Oyonerenc/A TVUE(OYNIWN 5 oc coon oD OCaeOO aaliaaliociiaceeliccleelcallo alls allaciles

125. DENDROBIASTES BASILANICA ......-... Pelle ollacheciecllec|iacloolles

126. Megalurus palustris ...........-..5-.. > all

127. Crateropus caudatus ....... ocononroseS %

198 ImEwAUDWEEDDAGIE . $0... cece h ee rle celle gical celica colle elloalloollas|lo oll =

IPE, 5 OVRIMOERISIMES goo accuocens0no0KG ~ ole |e

130, ,, MBLANOCHLAMYS ...........- Salamon) ele dio dices oloolle3

131. PHYLLORNIS PALAWANENSIS .....0----|--\..}e-|..fe-Jeelecfee[e ele ede |

132. Tora scapularis ...... He OOITOE Ge Btothe oe calla alioctla cle alle lle elle alle ollo olfo ols

133. Ixus golavier ........ non0D0CCUoONGDCS sll

B45 0) SWeRSIS Beale gnoonace o900000n)oa||

135. Poliolophus urostictus ......0..0- odo o}]-a\lo ollo all =

136. CRINIGER FRATER........ aocacooce Daelloollo alice allo allo cle cia lf olla ollo olla oll ex

137. Hypsipetes philippinensis.............. sof ae le el ae fae | ae

138. RUFIGULARIS ..... Re cpt ne laa a neallee | xe

139. Monticola solitaria SA ede CaREE OG oe oc wl el oe Sflecllec|[antto olfeclloalfe oll alle of elle

140. Pratincola caprata aes aren ates cra adeeotel stars bis Ps ele. Se PS gel aal aera a ae iy males

141. Cittocincla luzoniensis ................ ciel

142. 9 NL GURVAG ole fey artustey ets aane tenc eecuetele oollacliogl ouallbelleeleoalGrallor alle ells alte

143. Copsychus mindanensis.............. oalecle elle *

144, Calliope camtschatkensis .............. *

145. Gerygone simplex .............. on0000 > oll es

146. Phylloscopus magnirostris.............. allo lol's alle clic lic cloalic elle olla cle cilaalle ole nlic ol ca lloolle

147. 9) UMHS osoonacoovevedos Prllnelleetie alinelie cler| (2

148. Acrocephalus orientalis................ 3.0) e olleallo oll 22

149. Cisticola semirufa ........ gaootorceace ues %

150. Orthotomus derbianus ..............-. *

151. 9) CHUANG IS Gooocenoceeoce wf ed ae | ae

152. 9 TUMGTES. Geos ouedecoessane echo clecte cle ele cleelioolie clic allooll es

153. s CINEREICEPS.............. B.a[s clo o}\s allo oleeli= ale ollcs

154. a INZONAY NTI Alc bGanosconcoat a ollc clio olla allele clic ell ail ee

155. Budytes viridis .......... La nee Heya ce Neve RecA TTR Te 1 Ra get ag ad

156. Calobates melanops .................. ool ss * eles

157. Motacilla luzonensis ...........0++.0. *

158. Corydalla lugubris...............00005 celecllachealles

WSO Anthus Gustav. v0. secs cue neces ve ches Soon Sole le aleeheioelles

AGO;sBarnselepamsy s 2) e)siaevatec seman delehels %

IGN, gy ARUBA co san one ELS net LE A Bs OR Te Ve eat ll aca etal

162. Dendrophila frontalis................0% pee ltimetion o

163. H GNOCHLAMYS ............ palle alles i

164, Zosterops meyeni ....... Sere ece ee tera welled slates

165. Myzanthe pygmea.................... wl gelln of ge

166. Diceeum retrocinctum ........ dgrooece Selle eels alles

167. D9 jAYMUEWES ooocoupanneonocaauge a ea ee a eg *

168. y H@MATOSTICIUM ............ ello ole alleles

169. A | UrarOysmOCWENe Cagodor padoueouD aA tea bee %|

170. yy WON 5 soneo0 ag 0c aga0accas oolle olla ;

171. UGCROLUTIO oaocaccaccovccoo PASS ME

172. Prionochilus, Ro soaodvegaadatbodsHeces|(on Pee Maa eel el Salle cllsctlecalles

173. ARACHNOTHERA DILUTION ............|.. ARR be Pasa lec i s|loulls

| Malay peninsula.

| Australian Region.

| Polynesian Region.

| Papuan Region.

| Timor.

| India.

PROF. STEERE IN THE

PHILIPPINE ARCHIPELAGO.

Philippines. a

: | 3S 3 & A

| lelel (eidlel@l lelellalel (2/2).

3/812] £)2| 2\4|4|21s/8|2|2|2| 2 <#lZ/414

4/8 /s [A/S 6/5 |4/4]6/8)4/2/6]/8|3|a) 4/6) 4

174. ANTHOPYGA SHELLEYI ................).0).5 calla * |

175. = MAGNIFICA. 22.0. .ce eee wee lee lee ole

176. EUDREPANIS PULCHERRIMA............).-J.-].. bleed *

7c. Cinnyris speratus) (os. «Jacl > | - x|--| ale

178. 5 FURIES cose oosocogussasonalles pale sleallealleslloniles

179. Anthreptes malaccensts .........00.+000)--|e-].- pallesilocticailon ls

180. es CHLORIGASTER) 2... 500..5.]s-|-s]. 0]. - re

181. Rhabdornis mystacalis ................].. %

182. Corone philippima ...............-0..0/.- Pa ee ee | x

183. Acridotheres cristatellus ..............).. *

14> Sturmia: violaceay A nlss 5 cider. pss aicin’ cin oelaelioe Slo siies|lodllooilesiie |x

185. Calornis panayensis ..................|-- xl e|--|ele

186. s GiHh|ETS 55-s580aceresoses cog: allo {baile oltosal le Aes

TES /, SPSS GENIE co Suton nesocococude se. sla] ele]

188 3 OweHtide sesotoouuaocen seu O65 Iealealle 35 atoll a ode |g

189. Eulabes javanensis 2.0... ce eee veces eee eee salle mal oO ioral 2

190. SARCOPHANOPS STEERT ..............].. alle Bille x

HOls Passer*montanus.%..% 6.0. cde we eee oe z|--|. BeGolleallad thine lagi ore #

NODS Pade ay OryZiVOEM) wc las aleiclalsicieiem <loereselae fie -|. salle oll silo lelelalels

GB, Wikies Feo Soo soe goonsbonUsoNee soso x]. a

TDs by elmer, coo cose uo ooooet aH ae aesllas calle ol|ee

195. ,, TMNT) 5 Sar od boeine OUT SeNS SO so *

GS ee LCUCOR AS LIeum ielteteyovereitereleleleleled-lo!| ee jae -|x

WOVE OxyCerca Jago. . hase. oes sows ce eeen| a> *

IGS, IraWe TEOSIe a Gen decocsooouDosacdao ae slodlo ale 5 eee se *

199. Osmotreron vermaDS ..........0.0+-00- elelad--|.. as 50 zlelele|..|=

200. ” cpa Peat heels Seases EAS ace acto ais Ea od ie oe “|x

OOIS Weucotreron pironiert. <.).. sje ~ 2 ee oe = xl..lx

202. Rhamphiculus occipitalis..............).. %

203. Phabotreron amethystina ..............|.- %

204. 0 IGIGONG Co ocanondoseceDcusos &

205. or NEG HORMONE | Pyeleseyorcies ster ekeeee| Seni Jee |e

206. Carpophaga cenea .........0..0 eee eeeles PA slo allele ol ger lets es « glelxl.-|x

pA | Bike ee SIAL Slecbgde ciel eqectanie ai

208. Ptilocolpa griseipectus ................|.. *

209. Myristicivora bicolor..................|-- Ne slersl esl clxlael. ele] a] e|--lel-

210. Hemiphaga poliocephala ..............].. Pao eee

211. Jantheenas eriseigularis................|.. #{--|e-]

212. Macropygia tenuirostris................/.. x|--|--[e|- we

AIG, Aime GiechiniOul oe po stub bouoc ue bbl lan xlael--l

D4.) 5, InpiemiiGy SoGoodeencaopouerougs os }e ede

215. Foleo raezonr cays 154.2 pce stancetots wet eile x

216. Chaleophaps indica ..............2..5[-: xlel--|xl- % a ae

Bis Calisstdseotiarieal ots ects sle 1c RL PMI HN Lk aa S.

218. Geopelia striata .... 0... .ceeese sees ees #|--|..]..1. =|

POG allusybanlkdivene sey etcrcvcnetajcce lop: sivebax siete ever ee xlels|- --|*

DI0s Arharophita tsps pee tala: let-teiclafercieiata see % 2

221. Excalfactoria chinensis................ *% wlale|aele-|s

OVP eurnixocell atari tose teleice cee % -

22E. IOI, A, oo SereooadocHodaduccdood %

294, Mecapodius Cumingi...............00. x

225. 5 It dork oS a=ppo bose sede coldalla 4 as sles

2G, Chimchisuwlias) joouaddocusaaeacdc0sllse Salsllesy (6 Aa fee fae]. fae] ae | se |e

327, Sam EnOMINSH OAD bop. caneusgeavn lodliecltclk let & al sae

298, Aigialitis Geoffroyi............2.ee00-s = salle Re dleelles|les

OMIM dubia; 0050 as theca atom ome: y alld

230 or WOME, coooocoapoocecaccc Palas ne wefaelafee|- |. fae

231. Esacus magnirostris ..............000- False : Rolls ¥ #| 3 Dalle

232. Himentopus autumnalis ..............]. aly ce sale Ba lelleal incl eslealles

233. Himantopus leucocephalus ............|..]..].. Pas «le. x lel «|e % Q

OpuL, Cibmenik, Quemalis! saogoouooconeaeueuKc F %* all salleatleallecllocleslle

235. Porphyrio pulverulentus .............. x RNG

236. Gallinula chloropus .............+....|.. * Me salle aio alles

Bevin (Colliers GME, “GogcooncnuegonooendG x Ae sollodljo alte:

238. Hrythreea phoenicura .............+...)-- te aal{sol| es x |e]

939. Ortygometra cinerea ...........2-++---|--]--|-- eNealloc salle:

240; -Porzana pygMea .....-.-.cceeeeesaee x. 2c :

353

:|¢

5.2

81) 3

21 2|8

alle

2).2\8

s| 2

= a

E|als

Ellie:

4) |

a |

= * |

vole el ae}

MR. R. B. SHARPE ON BIRDS COLLECTED BY

* * *

Philippines. g

: A

|| |ele]./8 Sl el alc alle

és) alE/s/Slel2(zlelEla eiclelelzia

oa Q Ala] slolsa io ie | s

E lelsl2is|é|alalSe Ale G lelaelz|alé

PML, Vetoes WEEE 5g oooueobODeoHNDOODDDO # | oes eae

242. Rallina fasciata ........... baddoncce | ox || | ae

243, Amaurornis olivacea ............+005 +] x

244, Typotenidia torquata .............. *

245 os philippensis............ Foor xlecfeferfe-

246, a HAPEND soooncacgo00da0e. <- || x «|e | |e] Rae |e

247, Hydrophasianus chirurgus........ pooa)ooll . sefeetae fects

248, Numenius pheeopus ................ wef ee +| % % | ele]

249. Rhyacophilus glaveola ........++..--/++| % a Paella

250. Tringoides hypoleucus .............. Boller x lee | ele].

251. Totanus calidris ................ pocolt3l] oc x fae laelec|ael-

252. n CamescensS .........06- Rikieie Tats Boll 6% lal a&lelecde-

28. Dring rutCOllisysys ese) iclo+ slelelels ss e\e\e)° | x sfoe | ae | ae |e

254. Gallinago scolopacina............+ on ol[ool| & safeafe de

259. megala ...... SodOdOOHUGG 4 ++] & *

256. Rhynchiea capensis ...........- oagalesliiios

201. Ardea PULPULED . oe sence ocicie ewan ot |

258. Ardetta flavicollis .................. #I oo.

259, 4, += clmmamomea..............-- +]

260. on... HIGHS Sha bono Rd 0b OaCK0 000 ey 36 s+. 0 in

261. Bubuleus coromandus ..........+.+- -o| x +l ae]. * |: #

262. Herodias garzetta ....... poodooeca0s | x Sol aig .

263. 3 Intermedia .......... i nreiene Paleo . Pal -

264. Butorides javanica ................ ef] # etfs | ae

265. Nycticorax manillensis ...... seeeeee eles] oe

266. 8 BRMGINS oconos000000000000 o| sla le] e |

267. Gorsachius melanolophus ............ ..[ e- sof ete] a [el alls

268. Melanopelargus episcopus ......+. Gooooo! & +|

269. Querquedula multicolor.............. ++] x

270. Anas luzonica........ badooodccodedg)ool 6

271. Dendrocygna vagans ............+... +] % “hee

| 272. Nettapus coromandelianus ..........|-+| ae ap

273. Puffinus leucomelas ................ | x

| 274, Larus, sp..............-% sonoo0d00GS | #

275. Hydrochelidon leucopareia .......... | x H/e |e

276. 90 Ss o0a00000 seseeceesailnsl|

277. Sterna Bergit ............ SHaoKodey S safe fret ete ete eta ate cle e[e ale cleo] a

| 278. Onychoprion anesthetus ............ wef we fate cpa ete ef cde fees ele ete ele o]e olae Lae |e oe] ae].

| 279, Anous stolidus...... apooacumoAGBACOD %

280. Podiceps philippensis....... pae000000 #

281. Pelecanus roseus........ g0000000000 sfeo]

282. = philippinensis ............ ef ae Jee fe efecto e |e ete ete cds cfeefe efecto cle ate el ae dee] ae |e «

283. Dysporus piscator .................. * safe ete cfe efecto ete cle ete fae ]e ete -]e

284. | BWBs.ccoopssocac0s0sdesqan (O° + el ele |e | *

| 285. Phalacrocorax, sp........+--s+..0es+ *

286. 9 oes OSU eo eE | #

287. Plotus melanogaster ................]-- ge fe cfe [ae fe cde ete cfe cde cle ele efe alee] ae | ae | ae [ae fe ete

Motallyrrrteeie 49) 187 |31)31)59|19) 2 |48,23) 8) 5 |31) 9 | 4

* To the above add Brachyurus Kochi from Luzon, making therefore 138 from that island.

| Australian Region.

| Polynesian Region.

| Papuan Region.

| India.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO.

DESCRIPTION OF THE PLATES.

Puate XLVI.

Fig. 1. Chrysocolaptes erythrocephalus, n. sp.

Fig. 2. C. maculiceps, n. sp.

PiatE XLVII.

Halcyon Winchellii, n. sp.

Pirate XLVI.

Fig. 1. Zeocephus cinnamomeus, n. sp.

Fig. 2. Z. cyanescens, nu. sp.

PLATE X LIX.

Brachyurus Steerii, adult and young, n. sp.

Puate L.

1. Phyllornis palawanensis, male, n. sp.

Fig. 2. The same, female.

3. Pitilocichla falcata, u. sp.

Puarte LI.

Fig. 1. Irena Tweeddalii, n. sp.

cw)

. I. melanochlamys, u..sp.

Puate LIT.

Fig. 1. Cittocinela nigra, male, un. sp.

Fig. 2. The same, female.

Puatre LILI.

Fig. 1. Dendrobiastes basilanica, n. sp.

Fig. 2. Parus amabilis, n. sp.

Fig. 3. Dendrophila enochlamys, u. sp.

Prats LIV.

Sarcophanops Steerii, male and female, n. sp.

SECOND SERIES.—ZOOLOGY, VOL. I.

ARTI. ant ,

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é i ae iran Seen a Aue risen car

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RB Sharpe. Trans.Linn.Soc.s

1. CHRYSOCOLAPTES ERYTHROCEPHALUS.

GF calemans hth Z.C MACULICEPS.

TRANS Linn S¢

fs ad

IER Z

ZOOL VOL

HALCYON WINCHI

M&NHanhart imp

RB.Sharpe.

JG Keulemans hth.

EPHUS CINNAMOM!

2.4.CYANESCENS.

Trans.Linn.Soc.SeR.Z.Zoou Vou. |. Pu 48.

Use

M&N Hanhari up.

walpy TIAA LS SQOUNAHOVE

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MELANOCHLAMYS.

M&N Hanhart imp

.B.Sharpe. Trans. Linn.Soc. 5eR.2.Zoon Vou lPu 52

J.GKeuylernans lith. CITTOCINCLA NIGRA, é&? Mé&NHanhart imp

RBSharpe.

JG@Keulemans hth.

Trans. Linn.Soc.SER 2. Zoo. |

Z.PARUS AMAI

DENDROBIASTI

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HS BASILANICA ; M&N Hanhart amp

DENDROPHILA NOCHLAMYS

RB Sharpe. Trans Linn.Soc.Ser.2.Zoon Vou 1.Pu.54

J.&Keulemans hth SARCOPHANOPS STEERII. ¢ &2. M&N Hanhart imp.

THE

TRANSACTIONS

THE LINNEAN SOCLETY

LONDON.

SECOND SERIES.—ZOOLOGY.

VOLUME I.

PART THE SEVENTH.

LONDON:

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET:

SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON-HOUSE;

AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER-ROW.

“ M.DCCC.LX XVIII.

Pet APT 3B)

Pa AS ie TS Ste TS

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Oe, CANE ASE ETE: Oe PRY eae ly

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CONTENTS.

PART VII.—Aveust, 1878.

XVII. On the Male Genital Armature in the European Rhopalocera. By F. Bu-

CHANAN WuitE, U.D., F.L.S., ¥c. (Plates LV.-LVII.) . . . page 357

XVIII. Morphology of the Mammalian Ossicula auditis. By AtBan H. G. Doran,

F.R.C.S. (Communicated by Professor Fiower, F.RB.S., F.L.S., Fe. (Plates

TEAVALLI IN 3 DD. St NY) |e eek SS haere Cree uae aS ora Rl) Sr /

XIX. On the Annelida obtained during the Cruise of H.U.S. ‘ Valorous’ to Davis

Strait in 1875. By W.C.M‘Intosn, WD., LL.D., PRSS. L. & £., FL.S.,

Cj; (elate IXeVve and. 2 pWOodcuts)iu ys en a) 2 ge 4) ees 2 499

XX. On the Sacral Plexus and Sacral Vertebre of Lizards and other Vertebrata.

By St. Grorce Mivarr, Sec. L.S., and the Rev. Roperr CiarKke, F.L.S.

(lanes XeVily Cee Vvalieeand OkWoodeuts\Ma seas 42 2 =. = ols

| vs ol OS ee A Oe

‘ k Jae h Jen 17 5 ii ; f j ELEN

x A F u A

«th Wo MT ane ell piety Uh Sah, et REE hae Dai lta aay

\ bi tps ; ' ‘ , ro x

i AES ONE.” PET eatethe cr SAA i eee OANA :

‘ : Me

: Qh 8 Pe ree ar SOARES a. Sovak ea Barth) ,

HaAy MLCRO UE Re OS eR ORR Via ies AS ‘a iNhaae a

eas hy AEA AAS CO choke Oe ae

ae : Cah aeimanty % bine 4 ae ee el og

r eS t ay NE ANCE ’ Y Sea Ny 4 3 tes

TO Ue RR MTT ACR CCS A ME eT 6

ne ty svohiao te. Hh Mine 14 aS V2) ae ;

[ 357 ]

XVII. On the Male Genital Armature in the European Rhopalocera. By

F. Bucuanan Waite, I.D., F.LS., Sc.

(Plates LV.—LVII.)

Read December 21st, 1876.

IN several orders of the Insecta the structure of the organs upon which the perpetua-

tion of the species depends has afforded in very many cases the best and surest cha-

racters for the discrimination of species. But in the Lepidoptera, one of the largest

of the orders, and that on which perhaps the greatest amount of attention (scientific or

otherwise) has been bestowed, the structure of these parts has not only not received

the consideration it deserves, but has been almost entirely ignored *. Moreover those

lepidopterologists who have not altogether neglected to examine this structure have been

of opinion that it affords generic rather than specific distinctions.

- With this opinion, however, my observations prevent me agreeing; for I think that

not only generic, but, in many (if not in every) species, good specific characters are to be

found; and to show that this is the case is my object in laying before the Society the

following notes.

In them I have confined myself to the European Rhopalocera. Whether every family

of the Lepidoptera will show equally good characters my researches have not yet been

extensive enough to enable me to decide; but from what I have seen I am inclined to

think they will. The distinctive characters do not always lie in the same parts, nor are

they equally perspicuous in every group; but in one part or other, if not in all, some

distinctive character is to be found, and, allowing so much for individual variation, the

difference of structure is constant and appreciable.

An examination of the hind body of a male Trichopteron will show that it is furnished

with certain corneous appendages, protruding from the interior of the apical segment,

which appendages have been termed the ‘ appendices anales.” Of these there are

normally three pairs—the appendices superiores, inferiores, and intermedie.

If the hind body of a male Lepidopteron is denuded of the scales and hairs with which

it is clothed, somewhat similar appendages (more or less covered externally with scales

and hairs) will be seen protruding, to a greater or less extent, from the orifice of the

seventh or apical segment (PI. LV. figs. 2 & 3). In many cases the appendages protrude

far enough to permit of a good view of their form; but frequently it is necessary to

remove part of the apparent apical (7th), and occasionally of the penultimate (6th) or

even of the antepenultimate (5th) segment, before they can be properly seen.

On making the necessary dissection the appendages will be found to be three in

* A notable exception is afforded by Mr. S. H. Scudder’s various Memoirs on the Hesperide of North America

(* Proceedings’ and ‘ Memoirs’ of the Boston Soc. of Nat. Hist. &c.).

SECOND SERIES.—ZOOLOGY, VOL. I. 30

358 DR. F. B. WHITE ON THE MALE GENITAL

number—one superior, and two lateral. The lateral are the equivalents of the appendices

inferiores in the Trichoptera, and in this paper will be termed the “' harpagones”

(harpdgo -onis, a grappling-hook). The upper appendage may be the equivalent of the

pair known as the appendices superiores in Trichoptera; but though sometimes deeply

bifurcate, it yet consists of only one piece, which I will call the “ tegumen”’ (¢égimen,

a cover).

On dissecting the parts more deeply, it will be seen that the tegumen, though most

largely developed in the dorsal are (or tergite) of the segment, is continued as a chitinous

ring more or less round the ventral arc (or sternite) (Pl. LV. figs. 4a, 10 ¢, 11 ¢, 12¢, and

18c). The harpagones arise either from within the ventral portion of the tegumen, or

are superimposed on that part (PI. LV. figs. 4d,10d, 11d, 12d, 13d).

I will not at present say any thing further about the deeper anatomy, or concerning

the development of these structures, which probably represent the eighth abdominal

segment.

More or less immediately below the tegumen is situated the anal opening of the

alimentary canal (Pl. LV. figs. 10 a, 11 a, and 12a); and below that, again, is the intro-

mittent organ with its sheaths (Pl. LV. figs. 4c, 106, 116, and 126). In the latter good

characters will probably be found to exist; but as very frequently a good view thereof

cannot be obtained in a dry specimen, and as the soft parts are more or less altered by

desiccation, further allusion thereto will not be made in this paper. Part of the structure

is incidentally figured in Pl. LVI. fig. 26a and Pl. LVI. fig. 18 p.

In the female Lepidopteron the armature is much less developed, and the parts suffer

so much in desiccation that an examination of fresh specimens is absolutely necessary.

In some cases, however, the structure of the apical and penultimate abdominal segments

present interesting structural characters.

Before proceeding to give some details of each part of the armature, I may state that

I have examined many species in addition to those mentioned and figured.

The drawings have been made by the camera lucida, except the diagrams Pl. LY.

figs. 10, 11, 12, & 18,

They have been made from dry specimens, which are better suited for the purpose than

fresh ones. In the latter the necessary operation of removing the hairs and scales is

apt to result in causing the parts to assume unnatural positions, while the parts are

sufficiently hard not to be altered in form by desiccation.

_In giving side views I have only figured the parts next the eye, though frequently the

corresponding parts on the opposite side are more or less visible; nor have I figured

the sheaths or other parts of the intromittent organ that were visible in some cases.

The drawings (and descriptions) only attempt to give an idea of what is visible without

an alteration of the position of the object under the microscope. The same part viewed

in a very slightly different position will often present quite a different form; so that to

arrive at a correct idea of the structure the part itself must be examined. As much care

as possible, however, has been taken to place in identical position the parts figured in the

different species, in order that a just comparison may be made.

ARMATURE IN THE EUROPEAN RHOPALOCERA. 359

1. Structure of the Apical Segment.

In discussing this and other points of the genital armature, it will be convenient to

take one species as the type or standard with which to compare all others; and in

selecting Hpinephele Hyperantus, L., as the type or standard species, I wish to remark

that I do not claim for it any special position amongst the butterflies.

The typical structure, then, is that in which (looking at it from the side) the hind

margin of the segment is perpendicular, or nearly so, for about half the distance (less or

more) from the upper margin to the lower margin, and then slopes obliquely forward

(te. towards the head of the insect)-till, at the lower margin, it nearly meets the hind

margin of the penultimate segment (Pl. LY. figs. 1 & 2). This form I shall ferm the

typical or angular. The oblique portion may be either straight, or slightly concave, or

convex. The upper margin, or dorsal surface, is slightly sloped backwards, so that the

height of the orifice is rather less than that of the penultimate segment. The lateral

surfaces are gently rounded towards the longitudinal furrow (pleurite) which marks on

each segment the division between the dorsal and ventral ares. or this reason the orifice

viewed from behind presents a long oval outline, slightly constricted at about one third

the distance from the lower margin. Viewed from above, the hind margin of the segment

presents a flatly rounded outline (PI. LV. fig. 3).

2. Modifications of the Apical Segment in the various Families.

I must first premise that for this and other parts of the structure the appearance is

described as seen in profile from the side, unless when stated otherwise.

Paprtionip#. Structure various. In Papilio Machaon (P\. LVI. fig. 40 4) only the

upper half is visible, and is triangular in form, assuming the character of the tegumen,

for which I mistook it. In P. Alexanor there is a similar modification. In P. Poda-

lirius (Pl. LVI. fig. 394) the hind margin is perpendicular. In Thais Polyxena

(Pl. LVII. figs. 1,4, B) and 7. Medisicaste (Pl. LVII. figs. 2, a, B) the hind margin is

concave, and the segment assumes to a certain degree the character of the tegumen. In

Parnassius Apollo (Pl. LVI. figs. 3, A, B, c) and P. Delius there is a considerable departure

from the type, the segment being long oval in outline, excavated for half the distance above,

and nearly closed below. In P. Mnemosyne the departure from the type is not so great.

Prerip#. Hind margin more frequently oblique throughout than angular (Pl. LVI.

figs. 27 a, 28 A, 29, 30.4, 31 A,B, 33.4, 37 A,B, and 38,8). In Colias there is an oblong

prolongation in the direction of the dorsal mesial line (Pl. LVI. fig. 36 a from side, 8B

from above). In Leucophasia the structure is very aberrant. From the side the

outline is oblong, rounded at the apex; from above oblong, rounded, and deeply emar-

ginate at apex; from below it appears widely split to the base, with a triangular piece

partly filling the aperture (Pl. LVI. figs. 34.4 side, B above, c below).

Lyca/nip#. Angular (Pl. LVII. figs. 4 a, 12 a, 13 a), or perpendicular (Pl. LVILI.

figs. 17 4, 18 a, 194), more rarely concave (Pl. LVII. fig. 144).

Eryctntip#. Angular, approaching oblique (Pl. LVI. fig. 26 a).

LisyTtHEID#. In the single European species, Libythea celtis, the structure is aberrant.

3c 2

360 DR. F. B. WHITE ON THE MALE GENITAL

The dorsal surface of the segment is prolonged into two stout terete horns (Pl. LVI.

figs. 25 a side, B above), which, seen from the side, are slightly curved downwards ;

there is a tooth on the margin of the segment below the horn; and the remainder of the

hind margin is concave.

ApatuRiIp#. Angular, approaching concave (Pl. LVI. figs. 24, 34, 4 a).

NymMpHaLip#. Frequently concave and the orifice contracted (Pl. LV. figs. 264,

29 4), or concave and the orifice not markedly contracted (Pl. LV. figs. 14 a, 17 a), or

angula (Pl. LV. figs. 22a, 24a, 38). In some species of Argynnis the structure is

peculiar. In 4. Miobe (Pl. LV. figs. 184 side, Babove) the apex is deeply emarginate in

the dorsal mesial line. In A. Adippe (Pl. LV. figs. 19 a side, B above) the same part is

toothed and is not emarginate. Some doubts have been expressed as to the distinctness

of these two species. Thus, in the ‘Entomologist’s Monthly Magazine,’ vol. viii. p. 166,

Mr. A. G. Butler gives it as his opinion that they are only forms of one species; and at

p- 206 he reiterates this opinion, and cites Freyer in support of it. I think, however,

that the structural difference now pointed out is sufficient to show that the views of

those who consider both to be good species are correct.

DanatpmH. I have not sufficient material to enable me to give the structure in this

family.

Satyrip#. Frequently angular or oblique. In several species aberrant structure

occurs in the form of stout spines, arising from the hind margin. In their simplest

form these are hairs, not much stouter or persistent than the other hairs which clothe

the segment, and may be seen in Satyrus Fidia (Pl. LVI. fig.11). In Satyrus Tolaus

(Pl. LVI. fig. 10) they are more spiny, and each is emarginate at the tip. In Satyrus

Hermione (P1. LVI. figs. 12 « side, B above) they are considerably stouter, are reduced

to two or three in number on each side, and arise from a special process of the segment.

In Epinephele Janira (Pl. LVI. fig. 14.4) there is on each side a single spine, which

appears to consist of a mass of smaller ones, slightly unequal in length, agglutinated

together. In several other species there is a trace of these appendages.

HesPerip#. Angular (Pl, LVII. fig. 28 a, &c.), or nearly perpendicular (Pl. LVII.

figs. 25 a, 314, &c.).

3. The Tegumen.

‘Viewed from above, the tegumen is oval-acuminate in outline, but truncate at the

base ; the basal half is ovately spherical; and the acuminate apex is somewhat terete,

and curved slightly downwards. Nearly halfway between the base and apex a slender,

curved, spine-like lobe (which will hereafter be termed the “‘side lobe’’) is given off on

each side, and curves downwards, inwards, and backwards, so that it is not very well

seen from above (Pl. LV. fig. 3, undissected, and fig. 8, dissected out).

Viewed from the side, the outline is somewhat triangular, with the hind margin concave,

and the upper margin convex ; from about the middle of the hind margin the side lobe

arises and curves backwards alongside and below the apical half of the tegumen (Pl. LV.

fig. 4b, dissected).

Viewed from below, the tegumen is seen to be concave, with the edges slightly

ARMATURE IN THE EUROPEAN RHOPALOCERA. 361

incurved, and gradually approaching backwards till they meet and join in the apical half

(Pl. LV. fig. 9, dissected).

The standard form of the tegumen is that in which, looking from above, the organ is

much longer than broad, and tapers from a moderately broad base to a moderately sharp

apex, and, looking from the side, when the apical half gently curves downwards and

the side lobes are well but not inordinately developed. The standard length is that in

which the tegumen slightly exceeds the harpagones, when viewed from the side and the

parts are in their normal position. As will be seen, there is often a very considerable

departure from this typical form. ,

4, Modifications of the Tegumen im the various Families.

PAPILIONID#. Structure very abnormal. In Papilio it is not visible without dissec-

tion, and is shortly triangular and deeply emarginate—in fact, almost bilobed (P1. LVI.

fig. 39 B). As pointed out above, the upper margin of the last segment assumes the

character of the tegumen, and the tegumen proper appears in the character of the side

lobes (Pl. LVI. fig. 404 & B). In Thais the construction is somewhat on the same plan as

in Papilio, but the last segment is less tegumen-like (LVII. figs. 1&2). In both these

genera I at first was deceived by the form assumed by the apical segment. In Parnassius

the tegumen is larger, but various in form. In P. Apollo (Pl. LVII. figs. 3 B & c) and

P. Delius the apex is short and slightly developed, and the side lobes (or what appears

to be them) are exceedingly developed, and on the same plane as the apical lobe. In

P. Mnemosyne the structure is rather different: the apex is bilobed, resembling the

side lobes in the other species; and the side lobes proper are smaller and situated

below.

Prerip#. Structure various. Side lobes obsolete or very slightly developed. Tegumen

rather short, acuminate-triangular, with entire apex (Pl. LVI. figs. 273, 308, & 373),

or somewhat bifurcate (Pl. LVI. figs. 31c, 345, & 38 B).

Lycznipm. Structure still further modified, the development of the part being rather

in a lateral than a longitudinal direction. In Thecla (Pl. LVII. figs. 4-8), Awrotis (Pl.

LVII. fig. 9), and Thestor (Pl. LVII. fig. 10), the tegzumen may be regarded as decidedly

transverse and broadly developed laterally; and in these genera it is relatively much

larger than the other parts. In Zycena (Pl. LVII. figs. 11, 12, & 13) the development

is still decidedly but more narrowly lateral. In Polyommatus (Pl. LVII. figs. 14-24),

while in some species the development is transverse, in the majority it is more longi-

tudinal, the sides, however, and not the centre, being produced. P. Alsus (Pl. LVII.

fig. 24) is an exception, as in this species the greatest development is in the direction of

the mesial dorsal line, as we find in the type.

In this family the side lobes are frequently present, but, instead of arising from each

side of a central prolongation, spring from the under and inner side of the lateral prolon-

gation (Pl. LVII. figs. 7B, 104,128 & c, 19 B, &.). In P. Alsus (Pl. LVII. fig. 24 4)

their situation is more normal.

Erycinip#. Short and broadly triangular. Side lobes developed, but nearly hidden

(Pl. LVI. figs. 26 B & c).

362 DR. F. B. WHITE ON THE MALE GENITAL

Lisytnerpa2. Somewhat typical, but short, and side lobes either not developed, or

hidden (Pl. LVI. fig. 258).

Apaturip®. In Apatura (Pl. LVI. figs. 2 & 3) somewhat typical, but shorter, and

no side lobes. In Charaxes (Pl. LVI. fig. 4B), structure very abnormal. Viewed from

above and dissected, the tegumen is transversely oblong, four times as broad as long,

and the outer angles developed into short acute tubercles. No side lobes visible.

NympHatips#. In the majority approaching the standard form, but rather short, and

base rather broad. Side lobes usually obsolete or not apparent, but in some cases well

developed. Apex of tegumen usually entire, but occasionally bifurcate (Pl. LV. figs. 16 c,

21 8B, 23.8), or abnormally formed (PI. LV. figs. 18 c, 19 c, 26c, 318).

Satyrip#. Structure typical, and side lobes well developed (Pl. LVI. figs. 5-24).

In Canonympha (Pl. LVI. figs. 16 & 17) there is a tendency in the apical prolongation

to become a little wider before the tip, and again narrowed. In Melanargia (Pl. LVI.

fig. 5 a) the apex of the side lobe is abruptly bent downwards.

Hesperip#. Structure more or less aberrant. Viewed from the side, the tegumen

is often more or less abruptly bent downwards, frequently very abruptly (Pl. LVII. figs.

28 B, 30, 338A).

Viewed from above, it is often longly triangular, the apex slender and produced

(Pl. LVIL. figs. 25 B, 268, 278), or more or less longly bifurcate (Pl. LVII. figs. 28 c,

31 B, 320,338). In Hesperia linea (Pl. LVII. figs. 268 & c), and H. lineola (Pl. LVII.

figs. 27 4 & B) there is a process underneath the apical half resembling the side lobes

coalesced. In dineola this process forms a groove in which the apical half lies, the process

being prolonged beyond the apex. ‘The side lobes are not otherwise apparent.

5. Structure of the Harpagones.

Viewed from the side, the harpago appears as a somewhat triangular convex plate

(more convex on the lower than on the upper margin), extending beyond the hind

margin of the last segment. The apex is prolonged, rather incrassated, and toothed and

curved upwards and inwards towards the tegumen (Pl. LV. fig. 2). On dissection, the

shape of the harpago is seen to be a long oval, with the free end prolonged and curved

(Pl. LV. fig. 4). Viewed from below, the shape is long oval, with a convex surface,

and the apex prolonged and curved inwards towards the other harpago (PI. LV. fic. 5).

Viewed from the inside, the harpago is seen to be deeply concave, with the lower margin

~ much incurved, and joined to the upper margin near the apex, to make the long tubular

tip (Pl. LV. fig. 6), which is toothed on the upper margin (Pl. LV. fig. 7, more highly

magnified).

This is perhaps one of the simplest forms that the harpago assumes; but, as we shall

see, it is often, in other species, provided with lobes or processes, generally arising from

the upper margin, which is also, especially towards the apex, frequently more or less

toothed. The interior surface is also sometimes furnished with spines or processes ;

but, except where these are visible when the harpago is viewed from the outer side,

frequent reference will not be made to them on the present occasion.

ARMATURE IN THE EUROPEAN RHOPALOCERA. 363

6. Modifications of the Harpagones in the various Families.

Papiiionip#. Usually well extruded, large and broad, without many or large pro-

cesses (Pl. LVI. figs. 39a, 404; Pl. LVII. figs. las, 2438). Parnassius forms an

exception, the harpagones not being visible without dissection (Pl. LVIL. figs. 3 4,D) in

Apollo and Delius, and only partly visible in Mnemosyne.

Prerm#. Usually well extruded, large and broad, and without conspicuous processes

or spines (Pl. LVI. figs. 27-33, and 36-38). Lewcophasia is (as in other respects) very

exceptional. The harpagones require dissection to render them visible to any extent,

are very small, and situated rather inferiorly than laterally. In Z. sinapis (Pl. LVI. figs.

34D, F) they seem to be coalesced ; in Z. lathyri (Pl. LVI. fig. 35) they appear to be less

closely united.

Lycamyip#. Usually requiring more or less dissection before a good view can be

obtained; often not larger than the tegumen, and frequently much smaller; shape

various ; with or without processes or lobes. In Thecla (Pl. LVILI. figs. 4-8), Aurotis

(Pl. LVII. fig. 9), and Thestor (Pl. LVITI. fig. 10) usually small, not very conspicuous,

and without processes. In Zycena (Pl. LVII. figs. 11-13) rather larger, and sometimes

furnished with a spine. In Polyommatus (Pl. LVI. figs. 14-24) as large or larger than

the tegumen, and frequently with the apical half bilobed or subbilobed. P. Alsus (Pl.

LVII. fig. 24.4) has a long narrow curved process arising from near the base of the

upper margin.

Erycinip#. Requiring dissection; short and broad, and without processes (Pl. LVI.

fig. 268).

Lisyrunip#. Well extruded; large and somewhat typical in form; without pro-

cesses (Pl. LVI. fig. 25).

Apvaturip&. Well extruded; large and broad, without processes (Pl. LVI. figs. 2—4).

Nympnatip#. Variously extruded; large, and often furnished with large processes.

In some genera there is an approach to the typical form, and no processes (PI. LV. figs.

36-38); in others the harpago is broader, and also without processes (Pl. LV. figs. 32

& 33); in others there is a large process arising from the upper margin at the apex,

near the middle, or near or at the base (PI. LV. figs. 14-31 & 35, 36). In many dissec-

tion is necessary for a proper view of the parts.

Satyrip#. Usually well extruded; large but not broad, and often with prolonged

apex; without- processes like those so frequent in the preceding family, but often with

a large tooth or tooth-like lobe about the middle of the upper margin (Pl. LVI. figs. 6,

8, 9, 11, 12). The apex and upper margin (especially towards the apex) is often toothed

(Pl. LVI. figs. 5, 20, 22-24).

Hespermp#. Moderately well extruded; rather large, without large processes. Apex

often somewhat bilobed (Pl. LVII. figs. 25-29), upper margin with a large tooth

(Pl. LVII. fig. 30).

The following Table shows the characteristic variations of the structure of the different

parts in the various families :—

DR. F. B. WHITE ON THE MALE GENITAL

364

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ARMATURE IN THE EUROPEAN RHOPALOCERA. 365

7. Value of all the Armature in showing Affinities and Differences between Families,

Genera, and Species.

The arrangement of the families given in this paper is that adopted by Dr. Staudinger

in his ‘Catalog der Lepidopteren des Europzischen Faunengebiets,’ not as necessarily the

best classification, but as that of the most recent list of the European Lepidoptera. My

observations have been confined to the species inhabiting Europe proper.

PaPinionipz. Affinity with Lyczenidz and Hesperide, and with Pieride. Generic

differences great ; specific differences sufficiently well marked *.

Prerip#. Affinity with Papilionide and Erycinide, and with Nymphalide (Nym-

phalides). The genera rather closely allied, especially Aporia and Pieris, and Antho-

charis and Zegris, which latter two have scarcely generic differences. Colias is well

separated by structure of apical segment. Lewcophasia is very aberrant in all respects.

The specific differences are strongest in the structure of the tegumen in Pieris, Antho-

charis, and Gonepteryx, and of the harpagones in Leucophasia and Colias.

LycznibD#. Affinities with Papilionide and Hesperide, and perhaps Charawes in

Apaturide.

Thecla, Aurotis, and Thestor form one group, and have scarcely generic differences.

Lycena is well differentiated from them, and approaches Polyommatus, which connects

the family with Hesperide. In Polyommatus there are at least four groups of species :

—Ist, Beticus and its allies; 2nd, Arion and Argiolus (perhaps forming each the type

of a group); 8rd, Corydon, Adonis, Icarus, Astrarche, &c., which are rather closely

allied; and, 4th, minimus (Alsus), which diverges considerably from the type of the

family. (I have not yet succeeded in detecting any difference between Astrarche and

Artaxerxes, which, indeed, are usually united.)

Erycinipa. Affinity with Pieridee and Nymphalide (Nymphalides). ,

Lisytaeip#. Affinity with Nymphalide (Nymphalides), Apaturide, and Satyride.

* In his “Systematic Revision of some of the American Butterflies” (Rep. Peabody Ac. Sc. for 1871, pp. 24-82),

Mr. 8. H. Scudder has revived many old (Hiibnerian, &c.), and founded several new genera. Without expressing

any opinion as to the validity of these genera (which seem to have been somewhat unnecessarily multiplied), or of

Mr. Scudder’s changes in nomenclature, it may not be amiss to point out how the structure of the genital armature

(the characters derived from which are only once or twice used in the work in question) supports, in some instances,

Mr. Scudder’s views.

Papilio Podalirius, L., and P. Machaon, L., are given as the types of the genera Iphiclides, Hb., and Amaryssus,

Dalm., respectively—a generic separation which the structure, especially of the apical segment, rather confirms.

Of the genus Vanessa, polychloros, Antiopa, and urtice are referred to three different genera. The structure of

the armature supports the separation of wrticew (genus Aglais, Dalm.), but not of the other two.

The genus Argynnis is split up into several, two of which include the species we have had océasion to treat of.

Aglaia and other of the large species belong to the restricted genus Argynnis; the smaller species are contained in

the genus Brenthis, Hb. The character on which those and some other genera are founded by Mr. Scudder (it need

hardly be remarked that the genera, as Hiibnerian, are of no yalue) haye been shown by Mr. T. L. Meade (Canadian

Entomologist, viii. 232, Dec. 1876) to be for the most part so inconsistent that their value is very questionable.

Attention may therefore be called to the difference of structure of the tegumen in at least some of the European

species of Argynms and Brenthis, as giving, at any rate, subgeneric characters. In the Hesperide the generic separa-

tion of Thaumas, lineola, &c., from sylvanus &c., seems to be warranted.

SECOND SERIES.—ZOOLOGY, VOL. I. 3D

366 DR. F. B. WHITE ON, THE MALE GENITAL

Probably an examination of more than the single European species belonging to this

and the preceding family would reveal greater affinities.

Apaturip&. Affinity with Nymphalide and Libytheide. The two genera are very

different in the structure of the tegumen; and the species of Apatura are sufficiently

well separated by the form of the harpagones.

NyMPHALIDm. Related to the Apaturide and Satyridz by the Nymphalides, which,

again, are connected with the Vanessides by Araschnia and Pyrameis.

LTimenitis, Neptis, and Nymphalis have scarcely generic differences.

Araschnia and Pyrameis are generally different from Vanessa; but Grapta is scarcely

so. Vanessa is nearer to Argynnis than to Melitea.

The specific differences are often great. In the larger genera the species arrange

themselves in groups. Thus in Vanessa, wrtice and Jo form one group, and polychloros,

Aanthomelas, L-album, and Antiopa another. In Argynnis the large and small species:

form each a group, containing several subgroups.

Satyrip#. Affinities with Nymphalide (Nymphalides) and Libytheide. Generic

differences not strong, except in Melanargia and perhaps Gineis. Pararge, Epimephele,

and Satyrus scarcely separated. Species sufficiently well differentiated.

Hzsperrp#. Close affinity to Lyceenide, less close to Papilionid, and more remote

to Melitea, in Nymphalide. Genera and species fairly well separated *.

I regret very much that, from want of material, I have been unable to examine,

amongst the Rhopalocera, any but European species. It is much to be desired that

some one who has at his command a large collection of the butterflies of all regions

should investigate more extensively than I have been able to do, the structure of the

genital armature.

DESCRIPTION OF THE PLATES.

Note.—Where the outline of the hind margin of the last segment is ragged, part has been cut away

(e.g. Pl. LV. f. 168).

Puate LY.

Fig. 1. Apex of hind body of Epinephele Hyperantus, before denudation.

. The same denuded : side view.

. The same: viewed from above.

. A side view of the same, with the integuments of the apical segment removed. a, the tegumen ;

b, side lobes; c, intromittent organ; d, harpago.

5. The same: viewed from below ; the harpagones only.

6. View of the inside of the harpago.

7. Apex of harpago, more highly magnified.

8. Tegumen from above, with the side lobes.

9. Tegumen from below, with the side lobes.

EB Oo 0

* Cf. Scudder and Burgess, ‘On Asymmetry in the Appendages of hexapod Insects, especially as illustrated by

the Lepidopterous genus Wisoniades’ (Proc. Boston Soc. Nat. Hist. xii. pp. 282-306) and other works of Mr. Scudder

on the Hesperide.

Fig. 10.

Fig.

ARMATURE IN THE EUROPEAN RHOPALOCERA: . 367

Diagrammatic plan of parts, from behind, in Epinephele Hyperantus. a, anal opening; 6, intro-

mittent organ; c, tegumen; d, harpagones. The same lettering applies to figs. 11, 12.

. The same in Argynnis Aglaia.

. The same in Pieris brassice.

. Side view (the apical half of harpago is cut away).

NyYMPHALIDz.

. Argynnis Lathonia, L. a, from the side; 3, from above.

. Argynnis Paphia, li. a, from the side; 3, from above.

. Argynnis Euphrosyne, L. a, from the side; 3B, the same dissected ; c, tegumen, from above,

dissected.

. Argynnis Aglaia, Lu. 4, from the side; 8, from above.

. Argynnis Niobe, L. a, from the side ; B, from above ; c, from the side; D, from above, dissected.

. Argynnis Adippe, Lu. a, from the side ; B, from above; c, from the side, dissected.

. Argynnis Laodice, Pall. a, from the side; 8B, tegumen, from above: both dissected.

. Argynnis Amathusia, Hsp. a, from the side; B, tegumen, from above.

. Melitea Phebe, Knoch. a, from the side; 8, from above.

. Mehitea Athalia, Rott. .«, from the side; s, from above: both dissected.

. Melitea Britomartis, Assm. A, from the side; 8, the same, dissected ; c, from above, dissected.

. Melitea Didyma, O. 4, from the side; B, from above.

. Vanessa urtice, L. a, from the side; 8, the same, dissected ; c, from above, dissected.

. Vanessa polychloros, u. From the side, dissected.

. Vanessa L-album. From the the side, dissected.

. Vanessa xanthomelas, Esp. A, from the side; 8, the same, dissected.

. Vanessa Antiopa. a, from the side; 8, from above: both dissected.

. Vanessa Jo. a, from the side; 8, from above: both dissected.

. Pyrameis Atalanta, L. a, from the side, dissected; 8, appendage on inner side of harpago;

Cc, apex of tegumen.

. Pyrameis card, Lu. a, from the side, dissected ; 8, appendage on inner side of harpago;

Cc, apex of tegumen.

. Grapta Egea, Cr. A, from the side; 8, from above.

. Grapta C-album, L. a, from the side; 3, from above.

. Araschnia prorsa, L. (=Levana, L., var. gen. 1.). a, from the side; 3, from above.

. Neptis Lucilla, ¥. From the side.

. Limenitis Sibylla, L. From the side.

Prats LVI.

NympPxHaip# (continued).

. Nymphalis populi, L. From the side.

APATURIDE.

. Apatura Iris, l. a, from the side; 3s, from above.

. Apatura Ilia, Schiff. a, from the side ; 3, from above.

. Charaxes Jasius, L. a, from the side; 8, tegumen, from above, dissected.

SATYRIDZ.

. Melanargia Larissa, H., var. Herta. a, from the side; 8, tegumen, from above.

. Pararge 4igeria, L. 4, from the side; 8, tegumen, from above.

3D 2

Fig.

DR. F. B. WHITE ON THE MALE GENITAL

7. Pararge Rowellana. a, from the side; 8, tegumen, from above.

8. Satyrus Arethusa, Esp. 4, from the side; B, tegumen, from above.

9. Satyrus Statihnus, Hufn. From the side.

10. Satyrus Iolaus, Bon. From the side.

11. Satyrus Fidia, L. From the side.

12. Satyrus Hermione, L. a, from the side; 8. apex of last abdominal segment, from above.

13. Epinephele Lycaon, Rott. From the side.

. Epinephele Janira, . 4, from the side; 8, tegumen, from above.

15. Canonympha Iphis, Schiff. From the side.

16. Canonympha Typhon, Rott. Tegumen, from above.

17. Cenonympha Pamphilus, L. Tegumen, from above.

18. Cenonympha Hero, L. Tegumen, from above.

19. Gineis Aello, Hb. From the side.

20. Erebia Stygne, O. (= Pirene, Hb.). a, from the side; 8, tegumen, from above.

21. Hrebia Goante, Esp. From the side:

22. Erebia Tyndarus, Esp. From the side.

23. Erebia Melampus, Fuessl. 4, from the side; B, inner side of harpago.

24. Erebia Epiphron, Ku. 4, from the side, parts protruded more than naturally ; 8, mner side of

harpago.

LisyTHEID2.

25. Libythea Celtis, Esp. a, from the side; 8B, from above.

ERYCINID A.

26. Nemeobius Lucina, L. a, from the side; s, the same, dissected; c, tegumen, from above,

dissected.

Pirrip#.

27. Pieris napi, L. A, from the side; 8, tegumen, from above.

28. Pieris Daplidice, L. a, from the side; 8, from above.

29. Pieris brassice, LL. A, from the side; 8, tegumen, from above.

30. Aporia crategi, L. a, from the side; B, tegumen, from above.

31. Anthocharis cardamines, lu. a, from the side; B, from above; c, tegumen, from above.

32. Anthocharis Damone, Feisth. Tegumen, from above.

33. Zegris Eupheme, Esp. a, from the side; 8, tegumen, from ahove.

34. Leucophasia sinapis, . A, from the side; 8, from above; c, from below; pv, from the side;

dissected ; n, tegumen, from above, dissected; F, harpagones, from below, dissected.

35. Leucophasia Duponcheli, Stgr. (= Lathyri, Dup.). Harpagones, from below.

36. Colias Edusa, F. a, from the side; 8, from above; c, tegumen, from above, dissected.

37. Gonepteryx rhamni, L. a, from the side; B, from above.

38. Gonepteryx Cleopatra, L. a, from the side; B, from above.

PApiLiIonIpz.

39. Papilio Podalirius, L. a, from the side; 8, tegumen, from above, dissected.

40. Papilio Machaon, L. a, from the side; 8, from above, dissected.

Pruate LVI.

1. Thais Polyxena, Schiff. a, from the side; 8, from above; c, tegumen.

2. Thais Rumina, L., var. Medisicaste, I. a, from the side; 8B, from above.

“EO o1

ARMATURE IN THE EUROPEAN RHOPALOCERA. 369

. Parnassius Apollo, L. a, from the side; s, from above; c, from below; p, from the side,

dissected.

Lycanip2.

. Thecla ilicis, Esp. a, from the side; 8, the same, dissected; c, tegumen, from above, dissected ;

D, harpagones, from below, dissected.

. Thecla quercus, Li. a, from the side; 8, tegumen, from above: both dissected.

. Thecla pruni, Lu. a, from the side; 8, tegumen, from above : both dissected.

. Thecia betule, L. a, from the side; c, tegumen from above; B, harpagones, from below: all

dissected.

. Thecla rubi, i. a, from the side; B. tegumen, from above; c, harpagones, from below: all

dissected.

. Aurotis roboris, Esp. 4, from the side; B, tegumen, from above: both dissected.

. Thestor Ballus, F. a, from the side; 8, tegumen, from above: both dissected.

. Lycena Alciphron, Rott., var. Gordius, Sulz. a, from the side; c, tegumen, from above; ., har-

pagones, from below : all dissected.

. Lycena Eurydice, Rott. a, from the side; B, from the side, dissected; c, tegumen, from above,

dissected.

. Lycena virgauree, LL. A, from the side; B, from the side, dissected; c, tegumen, from above,

dissected ; p, sheath of intromittent organ, from above.

. Polyommatus Beticus, L. 4, from the side; 8, from above, dissected.

. Polyommatus Telicanus, Lang. a, from the side; B, the same, dissected.

. Polyommatus Arion, L. a, from the side; 8, from above : both dissected.

. Polyommatus Corydon, Poda. 4, from the side; 8, tegumen, from above.

. Polyommatus Adonis,Hb. 4, from the side; 8, tegumen, from above.

. Polyommatus Icarus, Rott. a, from the side; 8, tegumen, from above.

. Polyommatus Optilete, Knoch. a, from the side; 8, tegumen, from above: both dissected.

. Polyommatus Astrarche, Bgstr., var. Artaxerxes, L. a, from the side; 8, tegumen, from above:

both dissected.

. Polyommatus Afgon, Schn. a, from the side; B, tegumen, from above: both dissected.

. Polommatus Argiolus, L. a, from the side ; 8, tegumen: from above, both dissected.

. Polyommatus minimus, Fuessl. (= Alsus, F.). a, from the side; B, tegumen, from above: both

dissected.

HEsPERIpz.

. Nisoniades Tages, Lu. a, from the side; 8, tegumen, from above, dissected.

. Hesperia Thaumas, Hufn. (=Linea, F.). a, from the side; c, the same, cissected; s, tegumen,

from above, dissected.

. Hesperia lineola, O. A, from the side ; 8, tegumen, from above, dissected.

. Hesperia Sylvanus, Esp. a, from the side; 8, the same, dissected; c, tezumen, from above,

dissected.

. Syrichtus serratule, Rbr. 4, from the side; 8, tegumen, from above, both dissected.

. Spilothyrus alcee, Esp. (= Malvarum, Hffg.). a, from the side, dissected.

. Syrichtus malve, L. (= Alveolus, Hb.). a, from the side; B, tegumen, from above.

. Carterocephalus Palemon. Pall. (=Paniscus, ¥.). , from the side ; B, the same, dissected ; c, tegu-

men, from above, dissected.

. Cyclopides Morpheus, Pall. (=Steropes, Schiff.). a, from the side; B, tegumen, from above : both

dissected.

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GENITAL ARMATURE EUROPEAN RHOPALOCERA

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GENITAL ARMATURE EUROPEAN RHOPALOCERA

(ff 74

XVIII. Morphology of the Mammalian Ossicula audits. By AuBan H. G. Doran,

F.R.C.S. (Communicated by Professor Fiowsr, F.B.S., F.L.S., §c.)

(Plates LVITI.—LXIV.)

Read December 21st, 1876.

AMONG John Hunter’s original collection are some preparations in spirit, repre-

senting the tympanic region of mammals, with the small ear-bones i situ, but only a

very few imperfect dry specimens of the internal ear-bones themselves, though those of

man are better represented. Little was added to this department at the Royal College

of Surgeons’ Museum until 1866, when Mr. Toynbee’s valuable pathological auditory col-

lection showed the want of a corresponding series in the physiological department. For-

tunately, about the above date, the nucleus of such a collection offered in that of Dr. Max

Hibrich, of Munich, which the College authorities purchased. The Conservator, Prof.

Flower, wisely availed himself of opportunities to increase this; and at the present date

a very complete series of most of the genera of Mammalia is the result.

In drawing out a catalogue of these, I was led to study minutely and compare the

various groups one with the other. I soon found that, excepting the labours of Prof.

Hyrtl, of Vienna, on the osteology of the entire auditory apparatus in the Mammalia

(‘ Vergleich. Anat., Untersuch. iiber das innere Gehororgan des Menschen und der

Saugethiere,’ Prague, 1845), little or nothing in a connected form had been done to

elucidate the characters of the small ear-bones of Mammalia. Occasional desultory

remarks are to be found scattered in papers on human and comparative anatomy; but

these in most cases are entirely lost sight of. This suggested to me, in preparing and

arranging the Museum series, to take notes thereon and the critical comparison of

the objects themselves and collation of the literature on the subject, have resulted in the

following descriptive and comparative account of the ossicula auditis in the class

Mammalia.

In such small objects as the internal ear-bones it is at first difficult to apprehend the

value that may be attached to them as regards physiological influence on audition, and

still less as to their morphological importance; but if, for instance, those of man be

compared with those of marsupials, the wide difference is at once apparent. Again, in

such groups as the Carnivora or Cetacea, the family characters derived from these

diminutive ossicula are in the main very distinctive.

As the small ear-bones of the human being, for various apparent reasons, have neces-

sarily had most attention called to them, and moreover as man, in other respects, is con-

sidered by some a standard of comparison, I have consequently taken the human ossicula

- first into consideration. Moreover I have described their peculiarities in such a way

that the homologies of the parts of each ossicle in the lower mammals may be more

clearly understood.

372 MR. A. H. G. DORAN ON THE MORPHOLOGY

The drawings forming the accompanying Plates are from nature, and in all cases

enlargements. By their representation side by side in series a much better idea of the

family and generic distinctions are obtained than by mere verbal descriptions. Indeed

the latter have grown inordinately in my hands; and but for the circumstance that the

full data whereon my conclusions are founded are absolutely necessary, pending the fact

of this series of ossicula being unique, or certainly the most extensive now extant,

I should gladly have dispensed with this portion of the labour, and curtailed my text

accordingly.

In the Journal of this Society (Zool. No. 68, vol. xiii. p. 185) and in the Proc. Roy.

Soc. (No. 172, vol. xxv. p. 101) I have already indicated briefly a few of my results, and

in the latter referred to those gentlemen to whom I have been indebted for objects,

advice, and assistance.

Tue AUDITORY OSSICLES IN MAN.

The Malleus.—In the human malleus (Pl. LVIII. figs. 1 & 2, taken from a fully

developed foetus) the portion of the head above the articular surface is prominent,

smooth, and convex, projects more on the outer than the inner side, and is broadest

in its lateral measurement. Posteriorly it bears the articular surface for the incus,

which has very elevated borders and is about three times as broad as it is deep vertically.

This surface lies very obliquely compared with the mallei of most mammals, so that

its external extremity is much higher than its internal. It is generally described

as “ saddle-shaped,” though, from a functional point of view, Helmholtz + has objected

to this expression. In most works on human anatomy it is regarded as if it were one

single facet; but on comparing it with the same surface on the malleus of a Cat

(Pl. LVIII. figs. 30 & 31), it will be seen that it should be considered as made up of two

facets, much less distinct, it is true, than in Felis and most other mammals, but far

better-marked than in certain monkeys. A very faint groove, more marked in some

human mallei than in others, and not reaching either extremity, divides them, running

in the very oblique long axis of the whole surface. The more internal and upper facet

(Pl. LVIII. fig. 30, a@ * a), or portion of the articular surface above the groove, repre-

sents that facet which is almost completely superior in position in most lower mammals

(fig. 31, a* a), the more external (d* 0) below this groove corresponding to the lower

facet in many other mammals. Both rise into a high vertical convexity about the

middle of the whole surface, which is there much contracted ; their planes slope down-

wards towards the groove, so that a concavity is formed, plainly visible when the arti-

cular surface is viewed sideways. Now it may be remarked that of all the portions of

the three ossicula there is none that requires so minute and careful an investigation as

the articular surface of the malleus; and although the results of such labour are often

puzzling, and not invariably satisfactory, still, as a whole, many distinctive points of

value will be found in this region among the different orders of mammals. It will be

shown, when speaking of the ruminants, that the characters of this surface undergo

+ See his remarks on the malleo-incudal articulation in his work, ‘The Mechanism of the Ossicles and the

Membrana Tympani,’ translated by J. Hinton, New Sydenham Society, 1874.

OF THE MAMMALIAN OSSICULA AUDITUS. 373

conspicuous changes during growth in some genera, whilst in others no such alterations

take place. Hence the advisability of procuring young as well as adult ossicula, before

coming to conclusions with regard to peculiarities in any group,.is manifest.

Between the head and the manubrium is the short, constricted “ neck.” This is

rather flattened laterally, so as to be narrower in its vertical than in its horizontal measure-

ment. On its external aspect is a sharp sigmoid ridge *, convex forwards at first, where

it is near the anterior border of the articular surface, then concave forwards, where its

anterior end loses itself on the root of the manubrium (PI. LVIII. fig. 30). I call

particular attention to this ridge, as it is almost constant in those animals where the

malleus has a distinct neck, and is very plain in the mallei of the fissiped Carnivora and

most Ungulates, appearing as the sharply curved compact neck itself, the remainder

being a thin lamella of bone joining the processus gracilis. In the Common Badger

(Meles taxus) it will be seen to curve so sharply as to form a distinct projection.

Observe that this sigmoid portion—evidently the essential part of the neck, if we give

that name to the bony isthmus which seems made to unite the head to the manubrium—

runs away from the processus gracilis, but is lost on the base of the manubrium. This is

easily accounted for when we remember that the neck is originally developed from the

ineurved dorsal end of the mandibular arch, the apex of which forms the manubrium itself,

Professor Parker has recently shown 7.

On the anterior aspect of the neck, close to the root of the manubrium, is the root or

place of origin of the processus gracilis. Now, after examining numerous mallei from

human foetal and adult skulls, I have hardly ever found a trace of the wide lamina of

thin bone filling up the angle that process forms with the neck as in the fissiped

Carnivora. But as I believe the sigmoid posterior part of the neck in Homo to

represent the similar curved isthmus which constitutes the whole neck (independently

of the lamina) in these animals, I consider it correct to say that the stout compact

portion of the ‘neck’ of human anatomists in front of that sigmoid ridge is really

the same ossification as the upper part of the lamina, when such exists—the lower

being absent in any form, whether compact or papery. In good specimens from a

fetus at full term, it will be seen that a little fringe of friable bone extends as far

upwards as the front of the head, which latter, in these cases, sends forward a slight

projection, identical with a far more developed process seen in some lower animals,

especially in many of the Rodentia with laminated mallei, and notably m the Musquash

(Fiber zibethicus); in these extreme cases that projection may readily be taken for

the processus gracilis, if the anatomist overlook the fact that its root is not in the right

position for the origin of the latter.

The form of the human processus gracilis is well known. It cannot be seen entire

unless taken from a feetal or very young human skull, not only on account of its fra-

gility, but also because it actually atrophies to a mere stump before adult life ¢; so that

after removing a malleus from the temporal bone of a full-grown subject, the anatomist,

* Henle describes this as “a sharp border” (“ Kante’’), but he does not mention its sigmoid form.

+ “ On the Structure and Development of the Skull in the Pig,” Phil. Trans. 1874.

+ See Helmholtz, op. cit.

SECOND SERIES.—ZOOLOGY, VOL. I. 3E

374 MR. A. H. G. DORAN ON THE MORPHOLOGY

unaware of this fact, has frequently searched in vain along the Glaserian fissure for this

process supposed to have been broken off. It forms with the neck an angle of 125°, and

runs almost horizontally forwards to the fissure. It is generally bowed regularly, and

is concave forwards; sometimes it is somewhat wavy; but in all cases the degree of cur-

vature is slight. When well preserved, the processus gracilis at birth is longer than the

manubrium.

Below and internal to the processus gracilis the insertion of the tensor tympani muscle

is sometimes marked by a faint elevation, close to or upon the manubrium representing

the “ processus muscularis” of Hyrtl, which attains a great development in some lower

animals. In the human malleus, however, an elevation is occasionally seen as marked

as its homologue in the Bears, presently to be described.

That important process the manubrium, or handle, now claims our attention. In

Homo it is below the average length, far shorter than in ruminants or terrestrial Car-

nivora, though longer than in some Seals or Whales. It forms with the neck an angle of

about 140°; Hyrtl gives 150°; but he must have taken the axis, not of the upper and greater

part of the manubrium, but of the recurved portion at the tip, from strongly curved

examples. It is of a stout make, and broader at the base than in the Simiide; it is

much flattened laterally, the sides looking respectively antero-superiorly or postero-

inferiorly ; they are slightly convex, notwithstanding their compression. ‘The extremity

is always slightly recurved, though more in some specimens than in others, and per-

ceptibly spatulate and smooth towards the tympanic membrane. The edges of the

manubrium are both thick and blunt; and the outer edge, though of a certain breadth,

and giving attachment to fibres along its entire length, is rarely separated from each side

by sharply defined borders, so as to constitute a true and distinct outer surface, as seen

in many lower animals, and particularly in the Canide *. This edge ends below in the

spatulate dilatation, above in a very distinct sharp projection, well curved outwards.

This is the “ processus brevis:” its good development is a prominent feature in the

human malleus; and it will be seen to be better marked in the higher than in the lower

Quadrumana, though it occasionally reappears in the latter. In most mammals it is

merely an angle at the outer aspect of the base of the manubrium ; in many it does not

exist even in that form, especially where the neck of the malleus is very short.

Such is the human malleus. It may be seen as a good central type of the bone in

mammals, a type which very frequently reappears as we go downwards in the scale—a

malleus characterized by a well-developed,head, a moderately long, constricted neck, and

a broad-based, stout manubrium. It holds an intermediate position between the almost

or quite neckless type seen in the Cebide, in Tupaia, and in the true Squirrels, and the

broadly laminated form so well seen in the terrestrial Carnivora and the artiodactylate

Ungulata. It will be seen to be repeated, with modifications, in the higher monkeys, re-

appearing among Carnivora in the Ichneumons, and certainly approached (with some

important distinctions) in the Seals. It is also imitated in some of the Insectivora,

* Tn a recent contribution to the ‘ Archiy fiir Ohrenheilkunde,’ Trautmann describes and figures this condition as

if constant in Homo. It is only in a very few specimens of the human malleus, however, that I haye distinguished

two distinct borders running completely from the processus brevis to the tip.

OF THE MAMMALIAN OSSICULA AUDITUS. 375

Rodentia, &c. As for its comparison with the malleus of the Anthropomorpha, such a

consideration may be more conveniently deferred until the latter have been described

individually. The same remark applies to the other ossicula.

The Incus.—In the embryo this little bone holds a very prominent position, as the

proximal end of the hyoidean arch. In comparative anatomy it has hardly as yet

received the attention it deserves, on account of its having been supposed to vary less

in different mammals than do the other ossicles. My own observations lead me to con-

sider that in reality the chief objection to the incus as a distinctive feature in the skeleton

is the frequency with which it varies in form in different individuals of the same species.

Still it possesses important distinctive features in many orders of mammals. The incus

of Homo is of even more central type than the malleus.

The “‘ body” is well developed both in length and in height, and is considerably com-

pressed laterally. Its posterior part, or that which lies over the processus brevis, is more

developed than the anterior. The articular surface looks upwards and forwards, and is

shaped to fit that of the malleus; its more external limit is on the posterior segment of

the body, which is higher and more external than the anterior, in accordance with the

high position of the external extremity of the opposing surface on the malleus, already

described. On the other hand, that boundary of the surface on the incus opposing the

inner, which is the much lower extremity of the corresponding area on the malleus, lies

upon the lower and more internally placed anterior segment of the body. Into the

deep groove between the two extremities fit the convexities of the two facets on the

malleus. The very oblique position of the incus renders any purely verbal explanation

of the relations of this surface somewhat confusing; and the difficulty is increased in

this case chiefly because, in describing the malleus, I have acted in conformity with its

condition in the lower animals, where, as I have already remarked, that which is almost

the upper end of the articular surface in Man is in them rather the outer extremity, the

boundary which lies almost quite superiorly in most mammals being almost internal in

our species ; hence the homologue of their upper facet in Homo has an interno-superior,

and that of their lower facet an externo-inferior position.

Around the inner margin of this articular surface of the incus is a distinct groove, like

that which is often so strongly developed in the Seals (especially in Cystophora and

Phoca); and there is a distinct concavity on the inner surface of the body, at the bottom

of which is a minute nutrient foramen, often, however, filled up, even in the incus of a

new-born child.

The “processus brevis”’ of a human incus is, as a rule, not much shorter than the so-

called “processus longus.” It may, with more reason, be called the “ posterior crus.”

In Man, however, it lies as much superior to the rest of the ossicle as posterior; and in

some animals (the Dugong, &c.) it is almost absolutely superior; but the term given

above is less clumsy than “ postero-superior crus.” This posterior crus is broad at the

base and blunt at the point, where, on the inner aspect, is a minute pit or concavity,

generally admitted to be a true articular surface. It is moderately divergent from the

body. The processus longus, or stapedial crus, is slender and slightly convex inwards at

first, then forms a gentle curve, suddenly turning inwards at the extremity, which bears

3E 2

376 MR. A. H. G. DORAN ON THE MORPHOLOGY

a flat elliptical disk on a narrow pedicle. This is the “ os orbiculare,” better termed the

“Sylvian apophysis’ *. Among anatomists, Sappey gives reasons for considering it a

separate bone; but as it is now recognized as being formed on the proximal end of the

hyoidean arch, and, on the other hand, is separate from the long crus in an early stage of

development, it would appear correct, on such premises, to consider it an epiphysis of

the incus.

The Stapes—Whilst the ossicles already described take, in Homo, a rather central

type, this little bone assumes in the Bimana an extreme and highly developed form.

The beautiful bicrurate stapes of Man differs strikingly from the simple columelliform

representative of that ossicle in the Ornithorhynchus; and the study of the types inter-

mediate between those two extreme forms is highly instructive.

The “head” is broad and shailow; its free aspect is elliptical and a little concave, to

articulate with the os orbiculare. The crura are long and widely divergent, so as to

leave a large aperture between them and the base. The anterior crus is the more

slender and the straighter of the two; it is widely grooved towards the aperture in its

upper half; but the channel generally narrows close to the base. The posterior crus has

a curve or shoulder, often very marked, not far from the head; it is deeply channelled

towards the intercrural aperture by a groove almost as deep near the base as near the

head. ‘This sulcus joins, under the head, that which exists on the opposite crus.

Sometimes the anterior crus is more curved than usual, but seldom so sharply as the

opposite limb of the stapes. In such cases it is still the more slender, and the groove is

less marked near the base than it is on the posterior crus. I have examined several

specimens of short, stumpy stapedes, from both infants and adults, where it was found

very hard to distinguish the anterior from the posterior crus by their own characters

alone +.

The base, or foot-plate, is reniform, its upper border being a wide arch, its lower

slightly concave in the middle. Its posterior extremity is always well rounded off; and

its anterior is usually much sharper; but this distinction is often the least marked where

the crura are most alike.

The base is slightly convex towards the vestibule. I have never seen a true umbo

like that which is found on the stapes of many monkeys. On the tympanic aspect it is

formed into a shallow tray by the prolongation along its margin of the high edges of the

channel within the crura. This condition is very frequent in animals where that ossicle

is well developed and lightly made. A bony ridge}, traversing the long diameter of

the tympanic aspect of the foot-plate, is sometimes seen, but is not constant. The aper-

ture between the crura of the human stapes is larger, when measured vertically as well

* A beautiful description of this structure and the incudo-stapedial joint is to be found in Kysell’s “Beitrige zur

Anatomie des Steigbiigels,” Arch. f. Ohrenheilkunde, vol. v.

+ But the straightness of the anterior crus is generally much more apparent in the stapes of an infant than in that

of an adult. A specimen from an adult Bushwoman in the College collection is of a very marked infantile type

in the form of its crura. ‘This agrees with the characters of the rest of her skeleton, described in Flower and Murie’s

“ Account of the Dissection of a Bushwoman,” Journal of Anatomy and Physiology, vol. 1.

+ The “ crista stapedis.” See Hysell, loc. cit.

OF THE MAMMALIAN OSSICULA AUDITUS. 377

as horizontally, than in the lower animals, even including the Elephant, as Hyrtl justly

observes; but in the Golden Mole (Chrysochloris) and a few other fossorial animals

the width of the aperture at the base is proportionally greater, though the head is much

nearer the foot-plate.

The presence of a minute tubercle on the inner side of the head of the stapes does not

seem constant in our species. The occasional ossification of a part of the stapedius tendon

in Homo is well known; it is a constant feature in many animals.

For the most recent minute observations on the anatomy of the human ossicula, the

works of Magnus *, Helmholtz +, Eysell }, Rudinger §, Trautmann ||, Urbantschitsch {,

and others should be consulted.

THE OSSICULA OF THE SIMIIDA.

In a set of ossicles from a large mature fetus of a Gorilla (Pl. LVIITI. fig. 3) the incus

much resembles that of Homo, the stapes is not so large as in the human fcetus at birth,

and the crura are a little more slender and straighter. The tympanic ring from the

same foetal Gorilla has a considerable part of the floor of the external meatus already

developed, which is not the case in Homo at birth. Both rings are preserved in the

College of Surgeons’ collection.

I have examined a single malleus of the adult Troglodytes gorilla. It is above

the average size of the same bone in Man. The free upper portion of its head is

more developed than in Homo or Simia, and is somewhat flattened vertically, at the

same time protruding considerably forwards, as in the Chimpanzee. The articular sur-

face remains oblique, so that, as in our species, its outer extremity, or the homologue of

that extremity in other animals, lies very much higher than does the inner. The

outer and upper segment of the articular surface is far larger than the inner **—another

point in which this genus differs from Man and the Orang. The upper and lower facets

are saddle-shaped; the vertical convex ridge is very near the inner margin of the whole

surface; and hence the external, concave, sloping portions of both facets are very wide.

A very faint groove divides them, indeed, as in Man; it is only relatively that they can

be spoken of as two separate facets. This is the case with the remaining Quadrumana.

The neck is rather short and very narrow; the size of the head and the length of the

handle make it look less developed than it really is. The sigmoid ridge described in

Man is well marked. In the mallei from a fetal Gorilla the processus gracilis is very

slender, especially at the root.

The manubrium is very long, resembling in that respect the Orang’s, and differing

from the rather shorter manubrium of the Chimpanzee and Man. It is somewhat narrow

at the root, and well recurved at the tip, which is more thoroughly spatulate in Homo and

* « Beitrage zur Anatomie des mittleren Ohres,” Virchow’s Archiv, vol. xx. + Op. cit. = Op. eit.

§ ‘Atlas des menschlichen Gehérorgans,’ Munich, 1875.

|| “Der gelbe Fleck am Ende des Hammergriffs,” Archiv f. Ohrenheilk. vol. xi.

@ “Zur Anatomie der Gehérknéchelchen des Menschen,” bzd. vol. xi.

** This condition is not nearly so marked in specimens from new-born Gorillas.

378 MR. A. H. G. DORAN ON THE MORPHOLOGY

the other anthropoid apes; but the processus brevis is quite as well developed as in our

species.

In the Chimpanzee (Pl. LVIII. fig. 4) the malleus is much more human than in the

Gorilla or Orang-utan, and is as big as most of the numerous mallei removed by me

from adult male skulls in the dissecting-room of St. Bartholomew’s Hospital. In the

shape of its head, which projects markedly forwards, and in the nature of its articular

surface, of which the outer segment is much the widest, it approaches 7. gorilla more

than Homo or Simia; but in neck and manubrium it is very human, the only differ-

ence being that the latter, in this ape, is rather narrower at the base, and more curved

than in Man, and its well-developed processus brevis is directed upwards, and hardly

outwards. In length the handle does not exceed that of our species—another prominent

distinction from the other two apes.

The body of the incus resembles that of Homo; the processus brevis is more slender,

and ends in a sharp point, with no trace of any depression on it. The processus longus

is rather stouter and shorter than in Man; it forms, with the posterior crus, a right

angle.

The stapes of an adult Chimpanzee is smaller than the smallest specimen of that

bone from Homo in the College collection. The crura are almost equally curved; they

are shorter and more slender than in Man, but wider apart at their insertion. They

are well grooved towards the aperture, which is very wide. The base resembles that

of Homo, though less distinctly reniform, and equally rounded off at both extremities.

In the Orang-utan (Pl. LVIII. fig. 5) the head of the malleus is rather less developed,

and less regularly globular than in Man, and does not project conspicuously forwards

as in Lroglodytes. The whole bone, however, is of about the average size of its human

representative. The articular surface is shaped as in Man, but is less extensive; it is

hardly wider at its external or externo-superior extremity than internally. The neck is

as narrow as in the last genus, but rather longer; and it seems as if the sigmoid ridge

had become almost effaced in the process of elongation, being hardly perceptible, though

so marked in Troglodytes.

The manubrium is as long and slender as in the Gorilla; but the processus brevis is

very ill developed, though distinctly bent outwards. The extremity of the manubrium

is less dilated than even in Homo. .

In the incudes of Simia, in the College collection, removed from a young male Orang,

the body is much more developed *, particularly behind, than in Man, though not as

wide horizontally. Hyrtl asserts that the body is markedly smaller. The posterior

crus is as small and pointed as in Troglodytes; but the processus longus is short and

slender, and more incurved than in Homo; the crura too are more divergent.

The stapes has a very slightly developed head, and much more slender crura than in

Troglodytes. ‘The anterior is quite straight, the posterior slightly curved. The base

is distinctly reniform, and equally rounded at each extremity.

* Less of the body is taken up by the articular area than in the incus of Homo, as that surface is smaller abso-

lutely and proportionally in this ape.

OF THE MAMMALIAN OSSICULA AUDITUS. 379

In Aylobates (Pl. LVIII. fig. 6) the malleus (which is rather small even in a large

Gibbon’s skull) resembles in many respects that of the Chimpanzee. The upper part

of the head projects forwards; its articular surface is very wide externally, after the

Troglodytes type. From the fore part of the head projects a thin fringe of bone, which

does not join the root of the processus gracilis as in the Cynomorpha, and is repre-

sented, as in this genus, in some mallei from new-born children. The neck is short

and constricted ; the manubrium is as short as in Man, and much curved; it has a very

prominent, outwardly directed processus brevis, and a distinctly spatulate dilatation at

the tips; the whole angle forms, with the neck, an angle about as wide as in Homo, not

being so obtuse as in the Cercopithecide. Moreover the neck, though short and con-

stricted, is longer than in Semnopithecus, which has the most distinctly necked malleus

of all the next group. Altogether the affinities of the malleus of a Gibbon tend more

towards Homo than to the tailed monkeys.

In H. syndactyla and H. leuciscus (and probably in other Gibbons, only not con-

stantly) a distinct “processus muscularis” for the attachment of the tensor tympani

exists. In the latter it is seen as a distinct projection from the inner edge of the manu-

brium, at some distance from its root. In H. syndactyla it is reduced to a very small

tubercle on the root, almost on the site of the attachment of the tendon in Homo.

The incus in the Siamang (H. syndactyla) has a body of the same form as in Homo ;

in a specimen of that ossicle from H. leuciscus the body is higher and narrower, whilst

in an incus from another Hylobates the body is as narrow as in some of the Cebide.

The stapes of Hylobates has always curved, divergent crura, and a wide aperture *.

This is particularly marked in H. syndactyla. In this ossicle the Gibbons are allied to

Homo, Simia, and Troglodytes, in a manner not nearly so apparent in the malleus, and

still less in the incus. The very straight and not very divergent crura of the stapes in

all the tailed monkeys, even Semnopithecus, distinguish that group of the Quadrumana

not less from Hylobates than from Homo+. With regard to the ossicula of the Simiide,

in general, it may be briefly stated, in conclusion :—

I. That the ossicula of Troglodytes niger are, taken as a whole, most like those of

Homo. T. gorilla closely resembles Homo in its incus and stapes, but less in its malleus ;

Simia more resembles Homo in the head and articular surface of the malleus.

IJ. Though the incus of some species of Hylobates exhibits a tendency to low type, in

the malleus, and particularly in the stapes, this genus is quite anthropoid.

III. In their ossicula, but most markedly in the stapes, these apes are much more allies

to Homo than to the lower monkeys.

* In the Hylobates of uncertain species just referred to, the stapes bore on its foot-plate a distinct umbo, or central

‘convexity on the vestibular aspect.

7 From want of a sufficient number of good specimens when I contributed the communication already quoted to

the Royal Society, I was under the impression that the Gibbon’s stapes was of a low type; but I am now, after

more fayourable opportunities, compelled to modify that assertion.

380 ' MR. A. H. G. DORAN ON THE MORPHOLOGY

Tur AUDITORY OSSICLES OF THE CYNOMORPHA, OR TAILED OLD-woRLD MONKEYS.

The study of the ear-bones of the Old-world monkeys yields interesting results; for

they all possess certain types in common; but in comparing them with the Anthropo-

morpha we find definite grades of distinction, particularly between Semnopithecus and

Macacus.

In examining the malleus of an HEntellus (Semnopithecus entellus, Pl. LVIII. fig. 7),

the free upper portion of its head will be seen to be well developed and globular, dis-

tinctly projecting forwards. In the majority of the mallei of this species and genus a

distinct fringe of bone, of papery consistence, is found on the front border of the head,

joining the root of the processus gracilis (which is stout, stumpy, and very brittle in

the Cynomorpha). This (which is the exception in Man, and when existent the frmge

never joins the root of our processus gracilis, and is as little indicated in the anthro-

poid apes, excepting, perhaps, Hylobates) is a clear approach to the laminated type of

malleus seen in the Carnivora, &c. The articular surface is shaped much as in Man

and Simia, and is as little divided into the two facets of the typical mammalian malleus

as is the case in Homo. The neck is very short, but, being much constricted, is quite

distinct. The manubrium forms with the axis of the neck and head an angle of at

least 155°, distinctly wider than in any of the last family. This process is of moderate

length, though the shortness of the neck makes it look rather longer than it actually

is. It is almost of equal breadth laterally, till near the tip, where it turns up rather

_ sharply, and is there discoidal rather than spatulate.

Immediately above this dilatation is a distinct convex projection, giving a wavy look

to the manubrium seen in many lower monkeys. There is always a well-developed

and everted processus brevis in this genus; and on the inner edge of the handle, above

the projection just described, on the opposite or outer margin, is a distinct ‘ processus _

muscularis ” (as Hyrtl styles it, in describing the carnivorous malleus, and as it will be

henceforward designated in this paper), to which, in the recent skull, the tensor tympani

muscle will be found attached, as I have verified by dissection. Its position in this

and in neighbouring genera, well down the manubrium, reminds the observer of the

same process in Herpestes and in many Rodents.

In its very distinct though very short neck, and well-developed processus brevis, the

malleus in the genus Semnopithecns resembles the Simiidze more than any of the fol-

lowing genera except Colobus, where we find similar characters; but there the form

of the head is different. But the wide angle between the body and manubrium, the

peculiar form of that process, with a tendency to a discoidal termination, and the

presence of a distinct processus muscularis, as well as the trace of a lamina between

the head and processus gracilis, are points that ally this genus far more closely to other

Old-world monkeys.

The incus, on the other hand, in all the skulls of this genus which I have examined,

more resembles that of the anthropoid apes than in the succeeding genera, being broad

in the body, measured horizontally, and not very deep, with a considerable distance

between the origin of each crus, which diverges as much as in Man.

OF THE MAMMALIAN OSSICULA AUDITUS. 381

But the stapes of Semnopithecus assumes to perfection the type almost constant in the

lower monkeys. The head is deep rather than wide, the crura practically quite straight,

the base rather broad and almost plane, and the aperture much smaller than in the

Simiide. Thus, when we take into consideration all three ossicula in this genus, we

* see at once that the majority of their salient features ally them, as a whole, to lower

genera. The nature of the incus, and the few points given above regarding the malleus,

show a decidedly high,tendency in Semnopithecus, compared at least with Macacus.

In the malleus from a large skull of Colobus satanas (Pl. LVIII. fig. 8), propor-

tionally smaller than the same ossicle in a Semnopithecus of the same size, the head is

rather ill-developed and flattened laterally—a condition seen, but never to so marked a

degree, in Macacus and Cynocephalus. But there is a distinct, short, constricted neck,

very like that of the last genus, with a well-developed processus brevis ; only the processus

muscularis is abortive, and the tip only slightly spatulate. The incus is distinctly higher

and narrower than in the genus Semnopithecus ; and its crura arise nearer to each other.

Hence, as to these bones, Colobus must take a lower place than Semnopithecus, the ill-

developed head of the malleus being apparently a generic peculiarity, and the incus

allying it decidedly to succeeding genera. But the well-developed processus brevis, and

the still distinct neck, are marked affinities to the genera already considered.

Cercopithecus (P|. LVIII. fig. 9) is central among these Old-world monkeys in its

ossicula. The head of the malleus is fairly developed; the articular surface, though

shaped as in Homo, assumes the position usually observed in lower animals, the outer

extremity lying but little higher than the inner. A trace of lamination, as described in

Semnopithecus, exists; the root of the processus gracilis is very thin and fragile. The

neck is always extremely short (in C. cynosurus, however, I have found it nearly as long

as in S. entellus); and the manubrium runs almost in its axis. This is similar in

form to that of the last two genera, and bears a distinct processus brevis, never so well

- developed as in any higher monkeys, excepting Simia. In C. diana it is more prominent

than in C. cynosurus or in C. lalandit. The extremity is always spatulate, often dis-

coidal (as in a Diana-monkey, of which the ossicula are in the College collection). But

there does not seem to be any convexity above the tip, as described with Semnopithecus.

A more or less distinct trace of a processus muscularis is always seen upon the inner

border of the manubrium*. The incus is very high and narrow in C. cynosurus, but in

the other species resembles that of Colobus; the processus longus is very straight and

slender. The stapes is as described with Semnopithecus; in one from a fine C.diana the

base is so broad that the whole bone forms almost an equilateral triangle.

In Wacacus (Pl. LVITI. fig. 10) the malleus assumes an extreme condition in this

group. The head presents a surface flattened laterally, and projecting considerably for-

wards. This is best seen in WU. nemestrinus and WZ. erythreus; in I. cynomolgus it is

nearly as small as in Colobus. There is a trace of lamination and a stout processus

* This is represented in a sketch of the malleus of C. sabeus in Owen's ‘Anatomy of Vertebrates, vol. iii.

fig. 177 c.

SECOND SERIES.—ZOOLOGY, VOL. I. 3F

382 MR. A. H. G. DORAN ON THE MORPHOLOGY

gracilis, of which a great part is persistent in adult Macaques. Although the neck does

exist, it is extremely short, especially its anterior wall, the manubrium running almost

in its axis. The latter seems to spring from immediately below its articular surface ; its

extremity is often quite discoidal; the processus brevis may be completely obsolete, but

it is fairly marked in some species; the processus muscularis is in the same position as in ~

Semnopithecus, and is generally well developed.

The body of the incus is always either square or higher than broad; the anterior

segment is nearly as high as the posterior. Hence it approaches (in the first pecu-

liarity) the form seen in some specimens from Hylobates, and (in both the above points)

many of the Cebide. Both processes are rather long and slender; the long crus is

very straight till close to the tip. The stapes has straight crura and a broad base, as.

in all the past genera.

In Cynopithecus (Pl. LVIII. fig. 11) the head of the malleus is as ill-developed as in

Colobus, the neck and manubrium closely resemble the same in the Macaques. The

incus has an extremely stout square body; the anterior segment is very well developed.

The crura are very much stouter and shorter than in any Macacus of the same size,

and shorter, though of the same thickness as in Cynocephalus. 'This incus, with its

square body and short stout crura, is as different from the anthropoid type, and as

strikingly reminds the observer of the shape of the bone in Cebus and Ateles, as the high

narrow incus of Hylobates and Macacus diverges from the type of Homo in a direction

towards the form prevailing in Mycetes and in other less typical Cebide.

The stapes, too, though very similar to the prevailing form in this group see Semno-

pithecus), has a narrower base; nor are the crura so far apart as usual.

In Cynocephalus (Pl. LVIILI. fig. 12) the characters of the malleus are like those of

Macacus, but rather more marked. The head is not so developed in proportion to the

size of the bone, is extremely flattened laterally, and projects very sharply forwards. The

neck is also extremely shortened, especially in its anterior wall; the manubrium runs —

almost in its axis. This process has a very well developed processus brevis, and a sharp,

extremely prominent muscular process almost halfway down the inner margin of the

handle.

The incus has a rather shallow body, and very stout crura; the anterior is rather long.

It is clearly of a higher type than in Macacus.

The stapes of the Baboons does not differ from the common type in this group, and is

in outline almost an equilateral triangle.

In conclusion, it may be said of the Old-world tailed-monkeys :— —

I. That they all depart from the apes and Man, and resemble lower monkeys and

most other mammals in the straight and little-divergent crura of the stapes. A pro-

cessus muscularis on the malleus is almost constant.

II. In the distinctly-necked malleus, with a well-formed head, and in the incus,

broad between the crura, Semnopithecus approaches the Simiide, especially Hylobates ;

but the stapes is not at all anthropoid.

III. In Cercopithecus the malleus is nearly as high in type, but the incus is either

square-bodied or high and narrow, as in lower monkeys.

OF THE MAMMALIAN OSSICULA AUDITUS. 383

TV. In Colobus (and Cercocebus) the head of the malleus is almost as ill-developed

and sometimes more ftattened than in Macacus, and the incus is of the same form.

V. In Macacus, Cynopithecus, and Oynocephalus the malleus has a very short and

constricted neck; the manubrium, which bears a processus brevis and muscularis, forms

a very wide angle with the rest of the bone; the incus and stapes exhibit constantly

the tendency to low types, already seen occasionally in higher Quadrumana.

THE OssICULA OF THE CEBIDA.

The ossicula of the Spider-monkeys aman present some singular peculiarities not

observed in neighbouring genera.

The head of the malleus (Pl. LVIII. fig. 18) is extremely flattened vertically as well as

laterally ; seen from above it appears as a smooth, convex, almost quadrilateral surface.

The articular area is very wide, far more extensive than in any higher monkey; the

upper segment or facet is not so wide as the lower, and they meet each other at a very

open angle without a trace of agroove. The upper lies almost vertically above the lower

segment, so that the whole surface resembles the condition seen in the lower Mammalia,

and is unlike the type of Homo. Anteriorly a little friable layer of bone fills up a

part of the angle between the head and the processus gracilis, the stump of which is

stout. This homologue of the lamina seen in some mammals is often broken off and

absorbed in the mallei of old Spider-monkeys. The head never bulges prominently

forwards.

So far the malleus bears characters common to the other Cebide; but it differs from

them all in possessing a singular neck, quite unlike that of the Anthropomorpha, though

almost as long. Through the kindness of Prof. Garrod I have been enabled to examine

numerous skulls of Ateles melanochir, A. hybridus, and A. ater, from specimens once

living in the Zoological Gardens; and I find these characters best marked in A. mela-

nochir. Immediately below the articular surface the neck is very stout, and it becomes

narrower close to the handle, though even there it is much less constricted than in the

apes and Old-world monkeys. The very short and curious manubrium forms with the

neck an angle often not more than 120°, though occasionally much wider. Constricted .

near the base and tip, it bulges in a conspicuous manner in the middle, and terminates

in a wide discoidal enlargement. The processus brevis is represented by a prominent

rounded knob. There is no processus muscularis; and the tendon of the tensor tympani

muscle is inserted into the root of the manubrium close to the neck. This malleus is

certainly the most specialized among all the Quadrumana.

The incus has avery well-developed and very square body; in a specimen from 4. hybridus

I find it slightly longer horizontally than vertically. The anterior segment is almost as

high as the posterior; and the articular surface, looking rather outwards, appears unusally

deep from the outer surface of the body, and unusually shallow from the inner aspect.

The incus, on this account, will be seen to be most like that of Cebus, except in the last

feature, not showing that tendency to great height and narrowness so frequent amongst

other American monkeys.

The stapes has a well-developed head, and Ise slender crura, not very divergent, and

3F 2

384 MR. A. H. G: DORAN ON THE MORPHOLOGY

both inserted some distance from the extremities of the base, which is slightly convex

towards the vestibule.

The cruraare longer than in most Cebide, where the stapes closely resembles that of the

Old-world monkeys, only differing from them, as a rule, in the greater length of their

limbs; hence this distinctive feature of the entire family is best marked in Afeles.

In the Howlers (Mycetes) the head of the malleus is of the same form as in A/eles,

but still more flattened vertically and still less developed, the height of its convexity

Yhardly rising above the level of the upper border of the articular surface (Pl. LVIII.

fig. 14). This latter is extremely wide in all directions and very shallow; both facets

are very saddle-shaped. There is absolutely no neck; but immediately from below the

articular aspect springs the manubrium, which is tolerably long, forms an even, rather

wide curve, and becomes gradually narrower towards the extremity, which ends in a

perfect disk. There is not a trace of a processus brevis; and the processus muscularis is

almost obsolete. In IZ. seniculus the manubrium runs almost in the axis of the body;

in 1. laniger and WH. ursinus it forms a distinct angle.

The malleus of Wycetes, so different from that of Ateles, where there is a thick neck

and a manubrium bulbous in the middle and possessing a tubercular processus brevis,

may be considered a good type of this bone among the Cebide, and to a certain extent

among the less-typical Lemurs. It differs strikingly from the same bone in Man, Apes,

Old-world monkeys, and the genus Lemur.

The incus is very deep vertically, and narrow horizontally, with thick and short crura

arising close to one another. The articular surface has the peculiarity noticed in Afeles.

This. bone, too, is very typical among the Cebide.

The stapes is of the ordinary monkey type, the base concave laterally, with a trace of

an umbo ina specimen of I. laniger in the College collection, but not in one of JZ. wrsinus,

where the crura are almost fused; a minute aperture which exists between them is, I have

reason to believe, accidental *.

In Cebus (Pl. LVITI. fig. 15) the malleus differs little from that of Mycetes; the head

is almost identical in form. The manubrium runs still more closely to the axis of the

_ body; and no rudiments of a processus brevis or muscularis can be traced. The handle

does not form the free and regular curve seen in most Howling-monkeys (Mycetes), but

is slightly bent near its root, and straighter towards its extremity, which is a little turned

up and almost a perfect disk. é

. The incus departs from the type of Mycetes, Pithecia, Nyctipithecus and the Marmosets,

the body being generally longest horizontally, or quite square. The posterior crus is

rather thick at the base; it diverges, as in most Cebide, very little from the body. The

processus longus is often long and thin, and always divergent.

The stapes is of the usual monkey form; but in most specimens the posterior crus is

always slightly bowed and thicker than the anterior.

In the Douroucoulis (Wyctipithecus, Pl. LVIII. fig. 16) I do not find any distinction

* « Bei den Briillaffen ist die Basis konkayv, aber in der Mitte als Umbo gegen das Vestibulum vorgetrieben. Bei

einem Sapajou dessen Species unbestimmt ist, finde ich in beiden Ohren die Crura des Stapes verwachsen.”—Hrrrt,

op.cit. But I have noticed an Umbo in an ape (see Hylobates).

OF THE MAMMALIAN OSSICULA AUDITUS. 385

of importance in the malleus from the same bone in Cebus, except that a rudimentary

processus muscularis is to be found halfway down the inner border of the manubrium in

Nyctipithecus. The incus, on the other hand, is always high and narrow; the posterior

erus hardly diverges from the body, the slender processus longus diverging considerably.

The stapes has long slender crura and a comparatively narrow base.

In the Squirrel Monkey (Saimaris sciurea, Pl. LVIII. fig. 17) the malleus has the

general characters described with Cebus; but the articular surface is still shallower, and

the manubrium runs much less nearly in the axis of the body, often forming an angle

of not more than 150°. The incus is higher than broad, resembling that of Myctipi-

thecus in its crura, as does the stapes.

In a single specimen of the malleus and incus of the Black Saki (Pithecia satanas,

Pl. LVIII. fig. 18) the former ossicle has a manubrium rather thicker at the base, and

more distinctly flattened laterally (as in most mammals) than in the above-described

genera. ‘This process, too, forms the same wide curve seen in Mycetes; and the incus

has the same extremely high narrow body as in the Howlers; the vertical measurement

of the posterior segment is even greater than in those monkeys.

After careful observation, I consider that the malleus in the Hapanip# (Marmosets)

is intermediate between the same in the Cebide and in the Nycticeboid lemurs. The

description given of the malleus in the genus Cebus will apply to Hapale with two important

distinctions—namely, that the articular surface is always narrower and deeper in the

Marmoset, and in that animal the whole body is bent more inwards on the manubrium,

a character much more prominent in the Nycticebide, but in Hapale the head of the

malleus is longer in its vertical measurement than in Nycticebus. The body of the

incus is quite square in H. jacchus; but in a specimen of H. melanura in the College

collection it is as high and narrow as in the Douroucoulis (NMyctipithecus felinus and

N. trivirgatus). The stapes is of a different form from the type prevalent in the last

family. The head is very long; or, more correctly speaking, the crura are fused half-

way to their insertion, where they suddenly diverge towards the base. This is very

- like the condition seen in many of those marsupials where the crura are not fused

throughout. Hence in the genus Hapale may be found the lowest type of ossicle seen

in all the Primates (Pl. LVIII. fig. 19).

In Midas (Pl. LVIII. fig. 20) the malleus is of much the same form as in Hapale, the

articular surface also is rather deeper and narrower than in the Cebide; but the ten-

dency of the body to be bent inwards on the manubrium is not so plainly seen, at least

in YN. edipus, rufimanus, and geoffroii, as in Hapale. In all these species the incus is

high and narrow. The stapes in WZ. edipus has a small head with free crura and a wide

aperture; but one from UW. geoffroyii in the College collection has the peculiarities of

that ossicle in Hapale.

The absence of a neck to the malleus of most of the New-world monkeys seems not to

be due to an absolute suppression of that part, but rather to its being bent down below

the articular surface, and lying actually in the axis of the manubrium. In many mallei

from the genus Cebus a slight rounded projection may be detected high up on the outer

edge of the handle. In such cases, if the small segment of the manubrium above that

386 MR. A. H. G. DORAN ON THE MORPHOLOGY

prominence were raised to the exact axial line of the head (for even in these cases the

manubrium is never literally in that axis, but always slightly bent on the head), the rest

of the handle retaining its angle, we should have an ossicle almost exactly like the

malleus of Ateles. The processus muscularis, placed so low down as the inner edge of

the manubrium, would then be brought close to the neck, its normal or, at least, most

usual position.

THE OSSICULA OF THE LEMURID.

In the genus Lemur (Pl. LVIII. fig. 21) the malleus has a well-developed head, rising

considerably above the level of the articular surface, which is tolerably deep, and much

like the same in the Old-world monkeys. A faint trace of a neck exists ; the manubrium

forms an angle of 150° with the rest of the ossicle, and ends in a broadly spatulate or

discoidal termination. In the specimens of ZL. varius and L. albifrons in the College

collection, the manubrium is much flattened near the base, as in Homo; and in ZL, albi-

frons a distinct processus brevis exists; but in L. catta, L. nigrifrons, and L. xanthomystax

the base of the handle is narrow, and hardly perceptibly flattened; but probably in any

species of this genus a processus brevis may be occasionally developed. A small area of

lamellar bone is generally to be seen between the head and the stump of the processus

gracilis. The malleus of Lemur bears more resemblance to the same in the Old-world

monkeys than to the Ceboid type—not so, probably, from any nearer relation of the Lemurs

to the Macaques &c., which is most unlikely, but rather from the fact that the malleus

of the American monkeys is highly specialized, while it happens that the same in Lemur

and the tailed Old-world monkeys is of a far more generalized type in all cases.

The incus decidedly varies in form in different species of Lemur. In the College

specimens of that bone in Z. varius the body is wide horizontally and shallow vertically,

with stout widely divergent crura. In ZL. nigrifrons and L. xanthomystax the body is

both shallow and narrow, the crura stout and but little divergent, so that the ossicle has

avery tooth-like appearance. In Z. albifrons and L. catta the body is square, and the

processus longus is slender and very divergent. cH

The stapes has always long and extremely slender crura, very slightly bowed outwards,

with a wide aperture. The base is invariably convex towards the vestibule, and rather

narrow vertically. The space between the crura is in this genus always occupied by a

bony canal, which runs between two projections of the petrous bone above and below

the fenestra ovalis, and transmits a small blood-vessel—a frequent condition among some

Insectivora and Rodents. The same condition is seen in a skull of Hapalemur in the

Museum of the Royal College of Surgeons.

Among the Indrisinee, Propithecus (Pl. LVIIL. fig. 24)* has a malleus more like

that of Cebus than Lemur. The head is of much the same shape, and the articular

surface of the same depth, as in the latter; but there is the same absence of all trace of

neck, so frequent in the American monkeys. The manubrium is sharply curved in the

middle, and terminates in a very large disk. The incus is wide horizontally; the pos-

* Sketched from a young specimen. The presence of the processus gracilis, with some friable tissue between it and

the neck, alters its aspect. :

OF THE MAMMALIAN OSSICULA AUDITUS. 387

terior segment is much deeper than the anterior; and the crura are thick, and both

divergent. The stapes resembles that of Lemur; but the crura are stouter. In both

the skulls of Propithecus in the College Museum a bony canal passes between the

crura of the stapes, as is the case in a skeleton of Indris (Lichanotis) brevicaudatus. In

a skull of the Avahi (Jndris laniger) I have found the trace of a bare vessel, not supported

by bone, instead of an ossified intercrural canal.

The malleus of Galago (Pl. LVIII. fig. 22) strikingly resembles that of the Hapalide ;

but the upper part of the head is better developed, the articular surface is still narrower

and deeper, and the head is much shorter vertically than in Hapale or Midas. It is

quite neckless, and bent very much inwards on the manubrium—an exaggeration of a

condition frequent in the mallei of Primates where the neck is suppressed. The manu-

brium is slender throughout, even at the base, and ends in a very distinct disk; there

is a processus muscularis halfway down the inner margin. The incus is very high and

narrow, as in Wyctipithecus, Pithecia, &c.; and the anterior segment is actually as high

as the posterior, and projects forwards conspicuously. The processus brevis runs in the

long axis of the posterior part of the body, with hardly a trace of divergence, the pro-

cessus longus diverges extremely. This arrangement of the crura recalls Nyctipithecus,

and is usual in the Lorises. The crura of the stapes are slender, and perfectly straight ;

the intercrural space is moderate; and no bony canal appears to occupy it in the

recent skull.

In a specimen of the mallei (Pl. LVIIT. fig. 23) and incudes of-Cheirogaleus cookii,

the malleus, although resembling Galago in its head and articular surface, has a

manubrium which springs from immediately below the articular area of the head, yet

bears a distinct projection corresponding to the processus brevis of Man. This pecu-

liarity is seen in some squirrels, a family where gradations between the necked and

the neckless malleus are as numerous as in the Primates. The handle, too, in Chei-

rogaleus is rather flattened naturally. This ossicle altogether is related not distantly

to the same in the genus Lemur; but the incus assumes those conspicuous features seen

in Galago; the divergence of the anterior crus is extreme. There appears to be no

bony intercrural canal supporting the stapes in this genus.

In studying the Nycticebide or Lorisine, I find in the malleus of Wycticebus

(Pl. LVIII. fig. 26) that the characters of the neckless type of that ossicle are all

prominent. The head is very short in its vertical measurement, very much bent

inwards on the manubrium, and bears a deeply cut but not very wide articular surface.

All these features are seen in the Galagos, although the last two are approached in the

Marmosets. The singular manubrium is long and very straight; its outer border is

somewhat wavy; and there is a distinct projection and processus muscularis for the

tendon of the tensor tympani halfway down the inner margin. The handle terminates

in a large disk. The strong resemblance of the malleus of Nycticebus and of some other

Primates to the same ossicle in the Tupaiidee and Squirrels is interesting, especially as

they are all small thickly furred arboreal animals.

The incus has a shallow body, quite unlike that of Galago; but the anterior segment is

nearly as high as the posterior. Its articular surface is completely on the top of the

388 MR. A. H. G. DORAN ON THE MORPHOLOGY

body (strictly expressed, on the antero-superior aspect), and does not lean outwards, as

described in Ateles, and as found more or less in all the Cebidze and Hapalide. More-

over this surface is in Nycticebus very saddle-shaped, indeed almost convex. The pos-

terior crus is stout, short, and slightly divergent; the stapedial crus is stout, straight,

long, and very divergent, supporting an “os orbiculare”’ or Sylvian apophysis, which is

small as in all Lemurs.

The stapes has slender crura, not very divergent ; the base is rather thick and almost

plane. I have never found any intercrural bony canal in any skull of Nycticebus, Arcto-

cebus, Loris, and Perodicticus.

In a malleus of V. javanicus the articular surface is deeper cut than in WV. tardigradus.

The characters of the ossicula of Loris gracilis are almost identical with those of

Nycticebus ; only the head of the malleus is even still shorter vertically, and the anterior

part of the body of the incus projects forwards somewhat as in Galago. In Arctocebus

calabarensis (Pl. LVIII. fig. 27) the extreme characters of the neckless malleus are even

better-marked than in Zoris; the head of the malleus is shorter than in any of

the Platyrrhine monkeys, and the articular surface is far deeper cut than even in

Hapale. :

Perodicticus potto (Pl. LVIII. fig. 25) resembles Nycticebus in its malleus and

stapes; but the incus is higher than broad, its articular surface being, however, similar

in position to the same in the tardigrade Loris.

In Tarsius spectrum (Pl. LVIII. fig. 28 *) the malleus, which is quite neckless, has a

head of square outline, not nearly so compressed vertically as in the last family, and

projecting considerably forwards. The articular surface is deep, and rather wide; the

manubrium is slender, ending in a spatulate extremity; there is a trace of a processus

brevis ; and the tensor tympani is inserted into the inner border of the manubrium midway

between the tip and the root. The incus has a square body; both crura diverge con-

siderably. The stapes is small, even in proportion to the size of the animal; its erura

are very thick, and its base almost plane.

- Whilst the characters of the ossicula of the aberrant Tarsiws are somewhat negative,

the sole representative of the Chiromyide presents very salient features in its ear-

bones. :

All the ossicula of the Aye-aye (Chiromys madagascariensis, Pl. LVIII. fig. 29) are

much larger in proportion to the size of the animal than inany tailed monkey. The mallei

in the College collection are larger than any from A‘eles, and as large as that ossicle in

Cynocephalus. The head is much flattened vertically ; its upper free surface is broad

and concave, far more developed than in the other Lemures. The articular surface is

deeply cut and wide vertically; the two facets are well-marked: the upper is very wide,

and the lower is very small. There is a distinct short neck, twisted, together with the

head, somewhat inwards; the manubrium is long, rather narrow laterally at the base,

and forms a similar curve and angle to what is seen in the anthropoid apes. There is a

small angle corresponding to the processus brevis, and a distinct processus muscularis

* Taken from a new-born Tarsius. The remark made in the foot-note to the description of Propitheeus applies

to this as well.

OF THE MAMMALIAN OSSICULA AUDITUS. 389

on the inner edge of the manubrium nearer the root than the tip. The outer aspect

of the manubrium facing the tympanic membrane, though very narrow, is distinctly

bordered off from both sides by sharp edges, till it dilates very gradually into the narrow

spatulate, recurved extremity. This distinct outer surface to the manubrium is seen

in the Carnivora, Ungulata, and many Rodents. As it is in the Rodentia that the

processus muscularis is so frequently placed completely on the manubrium, the extensive

lamina being generally absent in that order, it follows that the malleus of Chiromys is

more allied to the Glires than to the flesh-eating and ungulate Mammalia.

The incus is remarkable for the sharp projection backwards of its posterior segments ;

the erura are stout, short, but neither are strongly divergent; the Sylvian apophysis

is extremely elliptical, or rather key-hole-shaped in outline, being broad horizontally

and very thin vertically. The crura of the stapes are stout, the posterior much bent,

as in Homo; the intercrural space is wide, and not occupied by a bony canal in the

recent skull.

The above researches into the ossicula of the Platyrrhini and the Lemuride will now

be briefly summarized.

I. In Afeles the malleus is of peculiar shape. It has a short neck and a rounded

prominence corresponding to the processus brevis. The articular surface is shallow, as

in all the Cebide. The incus has not a very high body; the crura of the stapes are

longer than in other monkeys.

IT. In Mycetes and Pithecia the neck of the malleus is quite suppressed (together

with the processus brevis), and the manubrium at its root runs in the long axis of the

head. The incus has a very high narrow body. The stapes, in these and the genera of

the Cebidze yet to be mentioned, ‘has not so long crura as in Afeles, but is of the

same form as in the Old-world monkeys.

III. In Cebus, Nyciipithecus, and Saimaris the malleus much resembles that of

Mycetes; but the head is bent on the root of the manubrium. In Cebus the body of

the incus is not higher than in Afeles; but in the other genera it is almost as high as in

Mycetes or Pithecia.

IV. In the Hapalidz the malleus has a rather deeper articular surface than in the

Cebidze; it is quite neckless; and the head is bent on the manubrium. ‘The incus is

generally high in the body. In the Marmoset the crura of the stapes are often fused for

some distance below the head, as in mammals of a very low grade.

V. Among the Lemuridx, the Nycticebide, Galaginide, and Propithecus (at least in

young specimens of P. diadema) much resemble the Hapalidze and Cebus in the malleus ;

but the articular surface is deep, deeper than even in Hapale; the head is very

short, and strongly bent on the manubrium. The incus has a very high body in the

Galagos, but not in the Nycticebide. Tarsius and Perodicticus resemble Nycticebus.

VI. But in Lemur the malleus loses the specialized form common in the other

Lemuride and the New-world monkeys. There isa constricted neck, and a more or less

developed processus brevis; the manubrium forms a distinct angle with the neck,

instead of the head being bent on it as in the Galagos &c. The incus is not high and

SECOND SERIES.—ZOQOLOGY, VOL. T. 36

390 MR. A. H.G. DORAN ON THE MORPHOLOGY ~

narrow. ‘There is always a bony canal between the crura of the stapes. This is seen in

the Indrisinz as well, but does not appear to exist in Galago.

VIL. In Chiromys the ossicula are large, and their affinities indistinct. The malleus

hasa much flattened head and a trace of processus brevis; in most respects it is of the

rodent type. The incus is of a somewhat peculiar shape; the stapes has curved crura,

a character seen in the higher monkeys, but also in many of the Sciuromorpha.

THE OSSICULA OF THE CARNIVORA.

The Fissipedia are almost as remarkable for the very trifling variations in the ear-

bones of the different families, genera, and species as the Pinnipedia are for the boldly

defined individual characters of all three ossicula in closely allied genera. The study of

these bones in the latter suborder is rendered all the lighter on account of their great size.

Yet while the Pinnipedia maintain, in the malleus at least, a certain adherence to the

common and most typical form of ossicle, the Fissipedia possess a type of malleus

unusual except among certain Ungulates and Rodents.

The Fissipedia must first be considered; and the study of their ossicula shows that

distinctive features of any clear value in classification are chiefly confined to the malleus,

the variations in the form of the ineus being more limited, although of interest in some

families.

The divisions of the suborder used here are those given by Flower* (P. Z. 8.

1869, p. 4).

The typical malleus of a fissiped carnivorous mammal may thus. be described :—It has

a small head, with a deeply cut articular surface, of which the facets are divided by a deep

well-marked groove; the neck is thin, long, cylindrical, and more or less sharply curved,

bearing on its inner side a muscular process for the tendon of the tensor tympani. The

manubrium forms with the neck an angle seldom over 100°; it has an angular pro-

jection at the outer side of its base, which may be dignified by the name of processus

brevis, although never developed to the extent seen in Man. It is narrow laterally;

but the side towards the membrana tympani is constantly separated from the two other

sides by sharp borders, and is often very broad,

But the great and conspicuous feature is the broad lamellar expansion between the

head and neck, on the one hand, and the very slender processus gracilis, which often

appears as merely a thickened lower border of that bony lamina, undoubtedly a minor

modification of the condition existing to an extreme degree in the Hedgehogs, Shrews

and Bats. This is observed to a corresponding extent in the Ruminants and larger

Muride.

The incus in the Fissipedia is always small in proportion to the malleus, directly con-

trary to what obtains with the Seals. Excepting extreme shortness of the processus

brevis in some families, there are few distinctions of importance between the incudes of

the various members of this group. The same applies to the stapes; and such slight

differences as exist will be noted in the description of individual genera or species.

* Tn his paper “ On the Value of the Characters of the Base of the Cranium in the Classification of the Order

Carnivora, and on the Systematic Position of Bassaris and other disputed Forms,”

OF THE MAMMALIAN OSSICULA AUDITUS. 391

In the Frnip#& the malleus (Pl. LVIII. fig. 32), a bone of light construction, exhibits

the general characters already described. The head is moderately developed; the arti-

cular surface is narrow vertically, but very deeply cut; the upper facet is nearly flat,

the lower quite convex; they are separated by a groove, which is very sharply marked

towards the inner limit of the articulation, but fades completely before reaching the

outer border. In the characters of this surface the smaller Cats may be known from the

Civets. The manubrium is not only narrow at the base, but its outer aspect, though

sharply bordered anteriorly and posteriorly, is extremely narrow throughout; and the

whole process, though well recurved at the tip, is not strongly bowed, with its concavity

forwards. The last two points distinguish the feline malleus from that of the Dogs,

the Hyzna, and Proteles. The extremity is only slightly spatulate, as in most Carnivora.

From the inner side of the thin, curved neck proceeds a long sickle-shaped processus

muscularis, as in the Canidze and the Weasels, much more slender than in Profeles and

Hyena, and better developed than in Cryptoprocta, the Viverride, Procyonide, and

Ailuride ; in the Bears it is absent. The lamina is well developed, though never as

wide as in the smaller families; even in the largest Lions and Tigers it remains papery

and semitransparent, not so compact as in Hyena.

The body of the incus (Pl. LVIII. fig. 42) is deep; its posterior crus is always well-

developed; the Sylvian apophysis is small.

The stapes (Pl. LVIII. fig. 56) has straight crura, with a wide aperture between them ;

and the thin but broad base is more or less convex towards the vestibule.

All the above characters are best seen in the Lion, where the lower facet of the arti-

cular surface of the malleus is unusually concave. In all the animals this facet lies

rather external to the upper, the outer extremity of the articular surface is higher

than the inner, but never to the extent seen in Man and in many monkeys. In the

Tiger, the manubrium is rather more slender than in the Lion. In the Cat (Pl. LIX.

fig. 1) the angle at the outer aspect of the base of the manubrium projects so much as

to form a true “processus brevis.” In the Leopard, the groove between the facets

of the malleus almost reaches the outer border of the articular surface, and there is a

distinct concavity of both facets external to the point where the groove ceases, an

approach to the Viverrine type. In the Cheetah (felis jubata) the manubrium is some-

what shorter than in the above cats, and more distinctly spatulate at its termination.

The Serval has a stapes with long slender crura, much less divergent than in most

Carnivora. In F. lynx and F. caracal the malleus has a stouter processus gracilis, and

is altogether more solid than in other cats of the same size. This is not the case with

F. caligata, which has a very broad lamina to the malleus, as in the European Lynx.

These smaller Felidz retain the characters of the articular surface of that bone as seen

in the Lion.

In the Hymyip# (Pl. LVIII. fig. 34) the malleus is of stouter build than in any

other family of the Fissipedia, excepting, perhaps, certain large Canidee; the head is

greater than in the Cats, yet not large in proportion to the development of the rest

of the bone. The articular surface is wider than in Felis; and the groove, well marked

internally, fades in the middle of the articulation; the facets it divides are flatter.

3G 2

392 MR. A. H. G. DORAN ON THE MORPHOLOGY

The neck is of the character seen in most Carnivora, but proportionally rather’ thicker.

The manubrium is much thicker than in Felis, though not broad at the base laterally.

Tts outer surface is moderately broad above, but very broad towards the point; thus,

lying exactly intermediate between Fels and Proteles, the whole process is rather’

more curved, with the concavity forwards, and less recurved at the tip than in the

Cats. The great feature in this and the next family is the processus muscularis, which

is very stout at the base, blunt at the tip, and almost straight. The lamina is denser

than in the Cats and Dogs. The incus (PI. LVIII. fig. 48) resembles that of a large Felis.

The stapes (Pl. LVIIT. fig. 58) is rather larger than in the other Carnivora; its crura

are slender, as in the greater Felide, not thick, as in Wolves.

In the Aard-Wolf (Proteles, Pl. LVIII. fig. 35) the head of the malleus is smaller and

less convex than in Felis or Hyena; its articular surface approximates to the Viverrine

type: both the facets are convex internally; but they run into one another, forming a marked

concavity external to the point where the groove ceases, the latter only running half across

the surface, as in Hyena. The manubrium has its outer surface broad throughout, as

in the Dogs; this character distinguishes it from Hyena, which it resembles in having

an almost equally straight and solid processus muscularis. The body of the incus is

shallow, as in Viverra and Canis; both crura are very stout and well developed.

Among the Viverride two distinct varieties of malleus are to be found. One, unlike

any other among the Fissipedia, is seen in Herpestes and Suricata, and in all probability

in allied genera; the other, which will be described first, is found in the genera Viverra,

Paradoxurus, Nandinia, Arctictis, Cynogale, and Hupleres.

The malleus of the Civet and its allies possesses very negative characters, or may be con-

sidered to present, together with those of the Cryptoproctidz, Procyonidz, and Ailuride,

a central type, from which those of the Cats, Dogs, and Bears are much specialized, the

above-named families exhibiting in this bone very slight tendencies towards the modi-

fications observed in those more important groups, which they respectively approach in

their general anatomy. Taking the most prominent point of distinction between the

mallei of terrestrial Carnivora, namely the form and development of the processus

muscularis, we find that process long in the Cats, and absent in the Bears. Now,

as it is of moderate length in the Civets and Racoons, from a quite natural moditication

of the characters of the bone in the two former families, it comes to pass that the

latter animals, very different from each other in the whole skeleton, come close to each

other as far as the malleus is concerned. F

The head of this bone in Viverra (Pl. LVILI. fig. 37) is small, its free surface pro-

jecting but little above the upper margin of the articular area. The articular surface

is narrow vertically and deeply cut; both facets are convex; the groove between them

is very deep, and, as in the Cats, it runs from the inner border to the middle of the

surface, where, however, it does not fade, as in Felis, but bifurcates; the two branches

run to the outer surface, and enclose a triangular, somewhat concave space, which may

be considered a third facet, and is clearly a further modification of the condition de-

scribed in Proteles, where the two branch grooves do not exist, but the facets run,

externally, into a concavity. In many faintly marked Viverrine mallei the bifurcation is

OF THE MAMMALIAN OSSICULA AUDITUS. 393

effaced; then the articular surface appears precisely as in Proteles. The neck is always

thin, and much curved towards the manubrium, which latter process is very long and

slender in all directions, even near the root; the angle corresponding to the processus

brevis is blunt; the outer surface, though well bordered, is extremely narrow throughout,

the terminal dilatation being almost imperceptible; and the point is little recurved.

The whole manubrium is very straight, as in the Cats, never forming the curve seen in

the Dogs and, to a less degree, in Hyena and Proteles. The processus muscularis is a

small mammillary projection, never quite absent. The processus gracilis forms a wide

angle with the neck, so that the lamina is extensive.

The incus (Pl. LVIII. fig. 45) is a small bone with a shallow body, bearing a pro-

portionally deep articular surface. Both crura are well developed; the posterior diverges

little from the body.

The stapes (Pl. LVIII. fig. 60) has always very slender crura, with a large space between

them, and a thin, broad base ; the former are inserted close to each extremity of the latter.

The Viverrine malleus strikingly resembles that of certain ruminants ; but the articular

region in those ungulates is of a different character, and the other two ossicles are

unlike those of the Carnivora.

Viverra cwetta exhibits all these characters. The processus muscularis of the malleus

is well developed, and, in a small way, approaches the straight conical type seen in Hyena

and in Proteles. In V. malaccensis that process is very short, and the manubrium is

very slender and almost styliform at the tip. In Genetta vulgaris the processus muscu-

laris is short, but still prominent; otherwise the malleus is as in Viverra; but the crura

of the stapes, though very thin, are not so extremely slender in this animal and in the

remaining Viverride as in the true Civet Cat. In Nandinia binotata and Paguma

grayit, as well as in Cynogale bennettii (Pl. LIX. fig. 13 a), the processus muscularis is

still short; but in Paradoxurus bondar and musanga it is reduced to a conical tubercle.

In Hupleres goudotiu (Pl. LIX. fig. 10) the head of the malleus is proportionally larger

than in the Civet; but the processus muscularis is as long, and the neck as well deve-

loped, as in that animal. The incus has a slender and well-developed processus brevis,

as in most of the Cats, Dogs, and Viverridze. The stapes is larger in proportion to the

incus than even in Herpestes. In Arctictis binturong (Pl. LIX. fig. 2) the malleus is

much more distinct. The neck makes a very short curve before joining the root of the

manubrium, so that the whole bone appears “hog-backed.” The manubrium is much

broader at the base and more recurved at the tip than in any of the preceding genera

—a distinct approach to Herpestes; but the processus muscularis, a mere tubercle, as in

Paradoxurus, still arises from the neck, and the lamina is well developed*.

One cannot fail to be struck by the great difference between the malleus of Herpestes

(Pl. LVITII. fig. 88) and that of the preceding Viverridee. The Ichneumons are the only

Carnivora where that auditory ossicle approaches the somewhat central type seen in

Man. The head is large, with a free orbicular surface; it bears an articular surface

sinilar to that of the Civets. The neck is short and stout; the manubrium is much

broader at the base, and more recurved at the extremity, than in any of the past genera

* The figure is from a specimen where the lamina was mutilated.

394 MR. A. H. G. DORAN ON THE MORPHOLOGY

except Arctictis, and forms a very obtuse angle with the neck; the processus brevis is

generally well marked. The short, stout, conical processus muscularis does not spring

from the neck, but from the inner edge of the manubrium, a perceptible distance beyond

its root, approaching a characteristic of some rodents. ‘The lamina is very little deve-

loped, indeed is not perceptible until close examination.

The incus (Pl. LVIII. fig. 47) is rather larger than in the Civets; the stapes is large

(Pl. LVITII. fig. 61), and resembles that of Genetta and Arctictis.

The species do not differ much in their ossicula. The head of the malleus in Herpestes

ichneumon and in H. paludosus is particularly large. In the manubrium of the latter, Prof.

Flower has detected a minute foramen which perforates it close to the root of the pro-

cessus muscularis. In the above species and in H. nepalensis, where the head is smaller,

the processus brevis is well marked; but it is much blunter in H. griseus and H. pulve-

rulentus. In the latter (Pl. LVIII. fig. 38) the processus muscularis is a more con-

siderable distance down the inner edge of the manubrium than in the malleus of any

other ichneumon ; its incus is proportionally larger.

In Suricata zenik (Pl. LIX. fig. 3) the ichneumon type of malleus is maintained. The

head is large, the neck extremely short. ‘The manubrium leaves the neck at a wider

angle than in Herpestes, and is distinctly shorter, broader at the base, and more spatulate

at the tip. The processus muscularis is situated on the inner side of the manubrium, a

perceptible distance from the neck; and the lamina is very little developed.

In the Cryptoprocta the malleus closely resembles that of Viverra; the head is a little

larger, the articular surface similar; the manubrium is even more slender; the processus

muscularis is almost as short as in Paradoxurus. The incus (Pl. LVIII. fig. 44) is a

larger bone than in the Civets; and the stapes (Pl. LVIII. fig. 59) has thicker crura than

in Viverra.

Thus Cryptoprocta ferox is incontestably more allied, as far as its ossicula are con-

cerned, to the Viverride than to any other family; and the latter are clearly more

related to the Cats by the type of their ear-bones, as Hyena and Proteles are related

to the Dogs.

In the Canidz the ossicula have far more marked distinctive features than they bear

in the Viverridee.

The malleus (Pl. LVIII. fig. 33, and Pl. LIX. fig. 5) often rivals that of the Hyzna

in stoutness of build. The head cannot be said to be remarkably developed ; the articular

surface is narrow, and deeply cut; both facets are concave; the groove dividing them

never bifurcates, as in well-marked viverrine mallei; but the facets join in a concave area

external to the point where the groove fades, as in Proteles. The neck is only mode-

rately curved; the manubrium, the most distinctive feature in the canine malleus, is

long, rather thin at the root, and very strongly curved, with the concavity looking

forwards. The outer aspect is a distinct surface, not only sharply bordered from the

sides, as in all Carnivora, but likewise very broad throughout, though rather broader

towards the extremity. Thus the Hyznide, as has been already remarked, resemble the

Dogs in their manubrium ; but Proteles is still more canine. On the other hand, the

Canidee have a long sickle-shaped processus muscularis projecting from the inner side of

OF THE MAMMALIAN OSSICULA AUDITUS. 395

the neck, as in the Felide; it is never straight and conical asin Hyena. The lamina

is broad, and bounded below by a well-marked splinter-like processus gracilis. The body

of the incus (Pl. LVIILI. fig. 43) is shallow; and the crura, which are both of nearly equal

length, are conspicuously far apart, even at their origin. The stapes resembles that of

the Cats. Hyrtl considers that the aperture between the crura is much smaller in the

Dogs; but it is more correct to say that in the large Canide the crura are more massive

than in the Lion or Leopard, and encroach more on that space, which, in small dogs, is

often as wide as in the domestic cat. The anterior crus is often curved.

In large specimens of the Dog the outer surface of the manubrium is so expanded at

its borders that the rest of that process appears as a sharp ridge springing from the inner

side of that external aspect. In the Wolf the same surface of the manubrium is even

broader than in the malleus of a Newfoundland or a Mastiff (although I have prepared

a malleus from a small dog where it is nearly as broad); and the whole process is

extremely curved; the extremity, however, is not much turned up, as in the straighter

manubria of many other mammals. The crura of the Wolf’s incus are extremely far

apart; and the anterior crus of the stapes is always well curved, and thicker than the

posterior. In the Dog the crura are far more similar in straightness and bulk. (See

Pl. LVIII. figs. 33, 48, 57, & Pl. LIX. fig. 5.)

The other species in this genus have ill-marked specific characters. In the European

Fox the neck of the malleus is shorter than in the Dog or Wolf; the outer aspect of

the manubrium is nearly as broad as in Canis lupus; but in C. vulpes (Pl. LIX. fig. 6) it

is nearly of equal breadth throughout. The crura of the incus diverge as much as in

the Dog, and hence less than in the Wolf; the anterior crus of the stapes is slightly

curved, but not much thicker than the posterior. In a specimen of C. fulvus I found

the manubrium rather longer than in the Fox. The crura of the stapes were both

straight, and the opening between them very small. In the Arctic Fox (C. lagopus) the

malleus has a much longer neck than in the two foxes above named, and a longer pro-

cessus muscularis than in any of the preceding Canide ; the incus is wolf-like. In the

Cape Hunting Dog (Lycaon pictus, Pl. LVIII. fig. 41) the processus muscularis is stouter

and straighter, and proportionally larger than in any other of the Canide; it closely

resembles that of Proteles; but the outer surface of the manubrium is quite canine, being

very wide, and almost of equal breadth throughout ; the incus is most like a dog’s in aspect.

In the Long-eared Fox ( Ofocyon) the neck and manubrium mallei are both rather short ;

and the processus muscularis is as long as in Canis (Pl. LIX. fig. 4).

In Canis lupus and familiaris the lamina of the malleus is wider than in the Foxes,

Iycaon or Otocyon.

The MustELIp# possess auditory ossicles which are not so remarkable for any single

distinctive feature or prominently developed process as for their very characteristic generai

appearance. ,

The malleus of the Glutton (Gwlo, Pl. LVIII. fig. 39) has a rather elongated head; the

free globular portion, fairly developed, projects considerably upwards. The articular sur-

face is always wide, contrary to other land-Carnivora ; but the facets are of the same pattern

asin Viverra. The neck is long, and only slightly curved, except in the Badger ; the manu-

396 MR. A. H. G. DORAN ON THE MORPHOLOGY

brium is always short, broad at the base, much recurved at the tip, though not curved

as a whole, with the concavity forwards, as in Dogs; and it forms an extremely obtuse

angle with the neck. There is always a rudimentary processus brevis. The muscular

process is always stout at the base, and tolerably long. The lamina. is very long and

narrow, owing to the very acute angle formed between the neck and the somewhat stout

processus gracilis—a condition diametrically opposite to that seen in the Procyonide,

and a great modification from most other Carnivora.

Careful comparisons seem to lead to the conclusion that the musteline malleus is

intermediate between the typical carnivorous form and that seen in Herpestes and

Suricata. By slightly enlarging the head in this family, and fancying the lamina ab-

sorbed, and shifting the processus muscularis to the root of the manubrium, we have

an ichneumon’s malleus without altering the manubrium or the angle it forms with

the neck. But by widening the lamina, lengthening and thinning the handle, and

placing it at nearly a right angle to the neck, the weasel’s manubrium may be brought

to the type prevailing in the other Carnivora already described. The thick curved

processus muscularis reminds the observer of Canis, and completely separates it from

the Procyonide and Urside. In some foxes there is a marked tendency to shortening of

the manubrium, and to a wide angle between that process and the neck; but it curves

more, as a whole, with the convexity backwards, and less at the tip; then, on the other

hand, the weasel’s manubrium is never so broad at its outer surface as in these vulpine

Canidee.

It may seem at first remarkable that the malleus in the Mustelide is unlike that of

the Bears, except perhaps in the head and articular surface; but after all, there are

considerable differences in the tympanic bullae beneath them. (See Flower, P. Z. S.

1869, p. 11, figs. 2 & 4.)

The incus (Pl. LVIII. fig. 55) is unusually distinctive in the weasels, having a very

short, slightly curved processus brevis, as pointed out by Hyrtl, at least in Mustela and

Lutra. The tendency to shortness of this process is as marked in most bears as in many

weasels, and nearly as great in the Mluride and Procyonide, except Nasua. The pro-

cessus longus stretches well forwards, and bears a rather large pedunculated Sylvian

apophysis.

The stapes (Pl. LVIII. fig. 62) has straight crura and a stout base; the former are

inserted each close to the extremity of the latter. The base is very bullate towards the

vestibule in small weasels, as it is in Hyrax, Geomys, and Phalangista cookie.

In the Pine Marten (Wartes abietum, Pl. LIX. fig. 7) all the above characters are well

marked in all the ossicles. The outer surface of the manubrium is broad at the base and

tip, which is well recurved and spatulate. The processus longus of the incus is shorter than

in other weasels; the stapes is larger, with a well-formed bulla. In Mustela vulgaris,

M. putorius, and WM. erminea the neck of the malleus is generally rather shorter than in

Martes and Gulo. The base of the stapes of the Mink (Mustela vison) is very bullate.

In Jetonyx zorilla the malleus has a very stout deflected muscular process; the base of

the stapes is slightly inflated (Pl. LIX. fig. 8). In the Glutton (Guwlo luscus) the large

malleus has a very elongated head and neck, a wide articular surface, and a short diver-

OF THE MAMMALIAN OSSICULA AUDITUS. 397

gent manubrium. The incus has a very short processus brevis, but a long, far-stretching

processus longus ; the whole bone is very like a Bear’s. The stapes has thick crura; and

the base is absolutely concave in the horizontal direction.

In the Badger (Meles tazus) (Pl. LVIII. fig. 40) the malleus is remarkable. The

head is as long as in Gwlo; the articular surface is even wider; and the facets are almost

flat, whilst in the above-described genera of weasels they are both quite convex*. The

most distinctive feature is a conical projection, formed by the neck making a sharp

angular bend, instead of a curve, close above the manubrium. It appears at first sight

to be the processus brevis, and thus makes the handle look longer than it really is. But

on close examination, the true representative of the processus brevis may be seen, pro-

jecting outwards, a perceptible distance below the conical projection. The incus resem-

bles that of Gulo. A very slight representative of the above-described projection exists

on the neck of the malleus of the American Badger (Taxidea americana). As regards MW.

taxus, this process (Pl. LVIIT. fig. 40) is equally prominent and well developed in five pairs

of mallei from skulls in the College collection. Its evident relation to the orbicular promi-

nence on the malleus of the Shrew (Sorex) and other mammals will be discussed later on.

The malleus of Helictis nipalensis resembles that of Meles in general aspect; but there

’ is no projection from the neck; the muscular process is not as long as in Mustela.

In the Otter (Zutra) (Pl. LIX. fig. 9) the malleus resembles the Weasel’s; the neck

is much constricted close to the manubrium. The incus has an extremely small pro-

cessus brevis ; the processus longus is stout and divergent. This character is better seen

in the Sea-Otter (Hnhydra), where that crus supports a large flask-shaped Sylvian

apophysis. -Aonyx (Pl. LIX. fig. 18) closely resembles Lutra in its ossicles.

In the Procyonip# the malleus is devoid of good distinctive features, and is of

slender make, with a broad lamina, as in the Civets. The head is not much developed ;

the articular surface is always narrow, and shaped very much like the same in the

Viverride ; the manubrium is generally long and slender, but the outer aspect, though

narrow, is broader than in that family. The muscular process is always present and

always short. The processus gracilis leaves the neck at a very wide angle, so that the

lamina suddenly becomes broad to an extent seldom imitated in the Civets. In a well-

preserved malleus of a member of this family this feature is very prominent; but the

lamina is often mutilated.

The incus has generally an ill-developed processus brevis, never so small as in

Mustela; the crura of the stapes are straight, and, together with the base, are thicker —

than in Cats, Dogs, and Civets. :

In the Raccoon (Procyon lotor, Pl. LIX. fig. 11) these characters are well marked : the

processus muscularis of the malleus is as large as in Viverra ; the end of the manubrium is

distinctly spatulate ; and the whole process is long and slender. The lamina, too, widens

suddenly from the great divergence of the processus gracilis. The processus brevis of the

incus is very much more slender than the stapedial crus, which is very broad, and turns

up sharply at the tip. The base of the stapes is very convex. In the Coati (Naswa nasica,

* Tn this and many other Mustelide the lamina is sometimes perforated, probably from atrophy or ill develop-

ment. Owen describes such a condition in Gulo and Lutra (Anatomy of Vertebrates, vol. iii.).

SECOND SERIES.—ZOOLOGY, VOL. I. 3H

398 MR. A. H. G. DORAN ON THE MORPHOLOGY

Pl. LIX. fig. 12) all these features in the malleus and incus are very feebly marked: the

neck of the malleus is rather short; the processus muscularis is a mere tubercle; the ma-

nubrium is very slender, and the lamina wide but not wider than in a typical Civet’s

ossicula. The processus brevis of the incus is more developed than inthe Raccoon. The

stapes is of stouter make than in Procyon, or in any Felis or Viverra; and its thick base

is very convex. In the Kinkajou(Cercoleptes caudivoloulus, Pl. LIX. fig. 14) the malleus

is much more distinct in type: the head is more developed upwards than in the other Pro-

cyonidee, and resembles that in many bears and weasels; the articular surface is deeply cut

and very narrow ; the neck is long, the manubrium is rather short, broad at the base, and

well recurved at the tips, as in the Mustelide ; but it does not form nearly so wide an angle |

with the neck, and the lamina has the characters of the Raccoon’s. The incus (Pl. LVIII.

fig. 53) has avery small sharp-pointed processus brevis, and a stout processus longus, as in

Procyon. It may be here noticed that, while the tendency to great shortness of the former

process exists in the Mustelidee, Procyonide, Aluridee, and Ursidé alike, in the first two

families the processus longus is stout and short; in _dlurus and the Bears it stretches out

far,asin Man. The base of the stapes in Cercoleptes (Pl. LVIII. fig. 63) is almost flat.

In the Cacomix! (Bassaris astuta) the malleus (Pl. LIX. fig. 15) more resembles that of

Viwerra and Procyon; but its head projects more than in either of those animals, and

the processus gracilis is extremely divergent, so that the procyonine type of a wide

lamina is best marked in Bassaris. This rare animal has a peculiarity already described

and better marked in Meles. The neck of the malleus forms a sharp conical projection

just above the manubrium (Pl. LIX. fig. 15, a, »), which being readily mistaken for the

angular upper extremity of the base of the latter, representing the processus brevis,

makes the whole manubrium look unusually long. In reality the handle is as short as

in Cercoleptes, and nearly as broad at the base; the extremity is distinctly recurved and

only slightly spatulate. The incus (Pl. LVIII. fig. 52) has a very small processus

brevis ; the stapedial crus is stout, and rather longer than in Procyon ; it supports, as in

the rest of the family, a sessile elliptical Sylvian apophysis.

In the Panda (#lurus fulgens) the malleus (Pl. LIX. fig. 16), asa whole, somewhat re-

sembles that of Procyon. 'The head forms a narrow projecting knob above its articular

surface, which is broader than in the last family. The neck is rather short, and almost as

sharply curved as in Viverra. The lamina has the same depth as in the Procyonide,

owing to the divergence of the processus gracilis. The manubrium is rather short, not

very broad at the base, and but little recurved at the tip; it does not form a very broad

angle with the neck : thus it is intermediate in characters between those of the Weasels

and the typical Procyonide, and resembles its representatives in most Bears. The

processus brevis of the incus (Pl. LVIII. fig. 51) is very short and sharp-pointed; the

processus longus is rather thin, and stretches forwards considerably, as in the Ursidze ;

it bears at its extremity a distinctly pedunculated and discoidal Sylvian apophysis.

‘The Ursip# are the only terrestrial Carnivora where a large malleus is to be seen with

an obsolete processus muscularis (Pl. LVIII. fig. 36). The head of that ossicle is well

developed, often projecting considerably upwards; the articular surface is rather wide,

as in the Mustelide; but the groove between the facets is generally faintly marked.

OF THE MAMMALIAN OSSICULA AUDITUS. 399

The neck is somewhat stout and boldly curved. The manubrium is not unlike the

same in Felis, but generally shorter and broader near the base; it turns up at the tip,

which is slightly spatulate; but the outer aspect is rather narrow. It never forms the

bold curve seen in the Canide, Hyzenide, and Protelide. The processus muscularis is

a mere tubercle or almost imperceptible elevation close to the root of the manubrium.

The processus gracilis always diverges considerably from the neck, so that the lamina

is very broad, as in Procyon, whereas the same is narrow in Distela.

The incus (Pl. LVIII. fig. 49) has a very small processus brevis, whilst the stapedial

crus stretches far forwards, and supports a Sylvian apophysis in the form of a disk on a

very narrow peduncle. The stapes (Pl. LVIII. fig. 64) has a well-formed head and

rather slender crura.

In the Polar Bear ( Ursus maritimus, Pl. LVIII. figs. 36, 49, & 64) all these characters

are well marked. The neck forms a sharp curve close to the manubrium, rather ap-

proaching the condition described in Cercoleptes; but it is placed lower, and does not give

an appearance of false length to the manubrium. This process is stout, sharply angled

at the base, and well recurved at the tip. The processus muscularis is represented by

a tabercle nearly as large as in Paradoxurus. The shortness of the processus brevis

of the incus, and the length and divergence of the other crus, are very marked; the

latter bends, forming a prominent knuckle, before contracting to give attachment to the

peduncle of the Sylvian apophysis. There isa rather wide aperture between the crura of

the stapes; the base is broad and plane.

In the Spectacled Bear ( Ursus ornatus, P]. LIX. fig. 17) hardly a trace of the processus

muscularis can be found on the malleus ; the manubrium is rather long and slender, little

recurved at the tip, and not sharply angled at the base. In an incus in the College col-

lection there is a slight tubercular projection close to the extremity of the processus longus,

just above the Sylvian apophysis. The crura of the stapes are rather thick, with a

small aperture between them.

The malleus of the Himalayan Bear (U. tibetanus) is more like that of U. maritimus,

especially in the manubrium, which is not very long, and is broad at the root, sharply

angular at the base and well recurved at the tip; but there is hardly a trace of processus

muscularis. The anterior crus of the stapes is distinctly curved.

In the Sloth Bear (Welursus labiatus) the head of the malleus is less developed than in

an Ursus of the same size. The manubrium is less stout and recurved than in the large

species of the typical genus, but is as angular at the base. There is no trace of any

processus muscularis; the lamina is very wide, and bounded in front by a prolongation

ofthe head, which joins the processus gracilis, a condition seen less plainly in other Car-

nivora, but constant in some other mammals.

THE OssICULA OF THE PINNIPEDIA.

The ossicula of the majority of the Seals and their allies are not only very large but

also-well-marked and readily distinguishable in the different genera or even species.

The ossicles of the Otariidz differ from those of the other families in their absolutely

3H 2

4.00 MR. A. H. G. DORAN ON THE MORPHOLOGY

and proportionally small dimensions. The ear-bones of an adult Otaria jubata are very

much smaller than those of any small Seal, such as Phoca hispida.

The malleus from the skull of a Sea-lion (Otaria stellert) (Pl. LIX. fig. 20) that

measured 0°305 metre from the premaxilla to the front of the foramen magnum, is itself

only 0-008 metre in length from the summit of the head to the tip of the manubrium ;

but this and the other ossicles, though small in these eared-seals, are always of stout

make. ‘The head of the malleus, instead of projecting well above the articular surface,

as in Man, or forming a smooth convexity in front of it, as in most of the Fissipedia, is

hollowed out anterior to that surface, and forms a shallow pit as in all other Pinnipedia.

The articular surface is very wide laterally and rather narrower vertically ; each facet has

so high a vertical convex ridge meeting its fellow that the whole area is quite convex,

forming a prominent angle in a horizontal view, though, viewed perpendicularly, the

planes of the facets do slope towards each other, and form a distinct concavity. The

groove dividing these prominent facets is but faintly marked. The neck is thin,

much constricted towards the manubrium, and never long. The manubrium is longer

than in the true Seals, and shorter than in the Fissipedia; it is broad, and flattened

laterally at the base, narrowing laterally, and turning well up at the point, which is only

slightly spatulate; it does not curve as in the Dog and Hyzna. There is a sharp angle

at its base; and its outer aspect forms a true well-bordered surface, very narrow, except

at the base and at the slight dilatation near the extremity There is no trace of a pro-

cessus muscularis, nor of any lamina, the processus gracilis running close under the

head and neck, to which it is fused by a medium of very compact, not lamellar, bone.

All the characters of the Otarian malleus are thoroughly phocine, putting aside its

small dimensions. Yet Hyrtl says,—“ Bei Otaria ist er (der Hammer) nicht grosser als

bei der Fischotter, und hat den gemeinsamen Typus aller Fleischfresser.’ Had that

distinguished anatomist said ‘“ common type of all aquatic Carnivora,” he would have

been correct; but he means the reverse, as the context shows.

But the incus does not approach any of the singular forms observed in the other

Fissipedia. It assumes very much the form seen in Man, though smaller in proportion

to the malleus, having high compressed sides, a rather short, but stout, blunt-pointed

and only slightly divergent processus brevis, and a long, slender, far-stretching stapedial

erus. ‘This is a modification of the forms seen in the arctoid Carnivora (compare figs. 49

& 50, Pl. LVIII.), where the posterior crus is short and not divergent. The extended or

widely divergent character of the processus longus is, it wili be remembered, very

marked in Ursus maritimus, in Alurus, and to a less extent in Bassaris. The little

stapes has stout crura, often, but not invariably, fused; they are not inserted precisely

into the extremities of the base, which is very thick, and quite concave horizontally.

In Otaria jubata the ossicula are largest : the incus is as large as in Homo, and similar

in form; the posterior part is much higher than the front of the body; the sides are

almost flat. The whole bone is wonderfully light in make compared with the incus in some

true Seals. ;

In Otaria stelleri (Pl. LIX. fig. 20) the concavity on the front of the head of the

malleus is very shallow, the neck and manubrium are moderately long ; the posterior part

OF THE MAMMALIAN OSSICULA AUDITUS. 401

of the body of the incus is not much higher than the anterior, the latter being well’

developed ; the anterior or long crus stretches well forwards, and supports, as in the

other Eared Seals, a Sylvian apophysis in the form of a perfectly circular sessile disk.

The crura of the stapes are quite fused, and the base is very thick; the bone strongly

resembles the same in the Dolphins. In Otaria falklandica (Pl. LIX. figs. 18, 19)

the depression in front of the head of the malleus is deep and well defined, the neck is

very short and constricted, the manubrium shorter than in Steller’s Sea-lion, though,

from the ill-development of the neck, it looks as long. The body of the incus is not

quite so broad as in the last two species, the posterior part is much higher than the

front, the processus longus does not reach so far forwards as in O. stelleri and jubata.

The two crura of the stapes are very thick and straight, with a distinct space between

them, and a base like that in O. stelleri. It is more regular in form than in any true

Seal; but a stapes of Phoca grenlandica in the College collection makes a near approach

to it, excepting that the ossicle in the Greenland Seal is much larger.

In the Walrus (7richechus) (Pl. LIX. figs. 21, 22) the malleus has a large broad head,

of a rhomboidal form, its anterior and posterior aspects being very much flattened, and

very wide in the middle, tapering at their upper and lower extremities. The former

surface bears a deep concavity, as in Otaria and the Phocide; but, unlike either, there is

asmooth rounded surface, of small extent, above the level of the articulation, repre-

senting the globular part of the head seen in other mammals. Posteriorly lies the

articular surface, as broad horizontally and as narrow vertically as in the Seal, but

concave in both directions; the facets are only slightly saddle-shaped, and divided by a

very distinct but shallow groove. The neck is short, not broad near the head, and

extremely constricted close to the manubrium; it is also flattened antero-posteriorly.

It bears no vestige of a muscular process; and the root of the processus gracilis is very

blunt. The manubrium, which forms little more than a right angle with the neck,

is extremely short; it is proportionally broad at the base and at its outer aspect; there

is a distinct everted processus brevis at its upper extremity. The whole bone is not so

large in proportion to the whole skeleton as in the true Seals, but is larger than in any

Otaria. It is plainly a mere modification of the Seal-type of malleus, the articular

surface being the most distinct feature ; the shortness of the malleus is only a slight

exaggeration of the condition seen in Stenorhynchus, where, on the other hand, the neck

is shorter than in Trichechus. It far less resembles the Otarian malleus.

The incus is a large bone, with the posterior part of the body well developed and

bulging externally ; the processus brevis is long, thin, and rather divergent. The pro-

cessus longus is stout, stretches far forwards, and is broadest in the middle, without a

trace of a groove on its broad inner aspect. It forms a gentle sigmoid curve, and bears

on its extremity a thick, sessile, discoidal Sylvian apophysis. This bone, too, is very

seal-like in the thinness of the processus brevis. The long, far-reaching processus

longus allies it to O¢aria; but I have examined one incus of Trichechus where that crus

was rather short; and moreover divergence and length is seen in the same process in

the Cystophorine Seals, but masked by the huge over-developed body of the incus.

The stapes is a high narrow bone, with a small head, thick crura, completely fused,

402 MR. A. H. G. DORAN ON THE MORPHOLOGY

without a trace of an aperture between them in any specimen I have seen, and a narrow,

rather thin hase, concave horizontally. The whole is shaped like a bottle with a short

neck, and very unsymmetrical on account of the outline of the anterior crus being by

far the most convex. This stapes strongly resembles that of MJacrorhinus, while Otaria

closely assumes in its stapes the type of the Dolphin. These two animals approach,

though less nearly, the Sirenian form of that bone.

The Cystophorina, the first subfamily of the Paocipm, are remarkable for the great

difference between the auditory ossicles of two well-known species otherwise closely

allied, and often included in the same genus, viz. the Hooded Seal ( Cystophora cristata)

and the Elephant Seal (Macrorhinus proboscideus). The latter most resembling Tri-

chechus in its ear-bones, it will be described first.

The head of the malleus of Macrorhinus proboscideus (Pl. LIX. figs. 23, 24) is flat-

tened laterally, as in the Walrus; it is still larger in proportion to the rest of the hone,

but is rather triangular than lozenge-shaped, the upper aspect being rounded off and

very broad, not produced to a blunt point. Anteriorly the head is marked by a deeply

concave pit, quite triangular, its basal boundary being uppermost—that is, corresponding

to the broadest or upper part of the whole head. The articular surface is very small,

not extending to the periphery or upper part of the broad posterior aspect of the malleus.

It exhibits the extreme of vertical narrowness and horizontal breadth so general among

the Phocidz. Both facets have a very prominent vertical convex ridge; so that the

whole surface is horizontally quite convex, though laterally slightly but distinctly con-

cave; the groove dividing them is extremely faint. The neck is very short; and the

processus gracilis springs from it as a prominent splinter of bone, connected with the

head by a thin and very brittle layer of compact, not lamellar bone. There is no trace

of a muscular process. The manubrium is very short, though not quite as short as in

Trichechus and Stenorhynchus ; it forms almost a perfect right angle with the neck; the

base is very broad; it is much recurved at the tip, which is spatulate; the outer aspect

is very wide at the base, corresponding to the processus brevis; it widens again near the

termination.

In general configuration the malleus of IZ. proboscideus closely resembles that of

Stenorhynchus ; indeed the only differences I can find are in the wider articular surface

and the still shorter manubrium, forming a wide obtuse angle with the neck in the latter

Seal. In Zrichechus the articular surface is different from that of Macrorhinus, and the

manubrium is still shorter. In Cystophora and Phoca the neck is long, and the manu-

brium forms a very wide angle with the rest of the bone.

The incus is of unique form. It is more than double the bulk of its own malleus, and

exceeds in size its representative in Balena and in most specimens of Manatus. The

oreat feature is the immense development of the posterior part of the body, which leans

over to the outer side of the crura to an extraordinary degree. The anterior part

projects forwards like a lip, and is also inclined somewhat outwards at its uppermost

part. The articular aspect is very small, occupying, to an equal extent, both the pro-

jecting posterior and the more level anterior part of the body; internal to it is a deep

pit, divided by a groove, which runs in a line with the ridge dividing the facets. This

OF THE MAMMALIAN OSSICULA AUDITUS. 403

latter peculiarity, also seen in Stenorhynchus, distinguishes the incus of Macrorhinus

from that of Phoca and Cystophora. The processus brevis is a short process, very stout

at the base and blunt at the tip; it does not diverge from the body; the long crus is

short compared with the dimensions of the whole incus, rather thin, quite cylindrical, and

stretches well forwards from the body, with a wide curve, as in the Bears, the Sea-lions,

and the Walrus. This bone is only a caricature, so to speak, of the same condition seen

in Phoca (compare fig. 23, with fig. 27, Pl. LIX.); but whilst the posterior part of

the incus of Macrorhinus assumes and exaggerates the Phocine type, the long crus is,

as just noticed, rather like the same in the last families, Otariide and Trichechide ;

so that the incus of Macrorhinus is a truly central form.

The stapes strongly resembles that of Trichechus, only itis of much stouter build; the

head is even smaller; the crura widen close below it, and then slowly approximate to the

base; they are entirely fused ; but on the front aspect of the bone is a deep concavity,

representing the normal aperture. The base is a thick bordered facet, and is quite saddle-

shaped, being very convex vertically, though nearly plane horizontally.

In Cystophora cristata (Pl. LIX. fig. 25) the head of the malleus is flattened antero-

posteriorly, and somewhat oval, being broadest horizontally; there is a concavity in

front and a facet behind, convex laterally and concave vertically, as in Phoca and

Wacrorhinus, &c.; but it occupies nearly the whole of the posterior aspect, being far

wider than in the latter, while the whole head is smaller. The neck is extremely long,

more so than in any other Seal, and almost perfectly cylindrical, being but little con-

stricted near the handle; it is, moreover, very straight. The manubrium is short,

rather narrow at the base, well recurved and dilated at the extremity; it forms a very

obtuse angle with the neck, and bears a prominent blunt processus brevis. Altogether

this bone resembles the malleus of Phoca, from which it may at once be known by the

length of its neck.

The posterior part of the body of the incus is well developed, and bulges internally,

but not nearly to the extent seen either in Stenorhynchus or Macrorhinus. There isa

shallow groove round the inner side of its facets, as seen ina more developed form in

Phoca. The processus brevis is short and stumpy; the longer crus is shorter and

stouter than in Macrorhinus; still it stretches well forward, and in that process

alone the incus of this animal resembles that of the last-described species rather than

Phoca.

_ The stapes has an ill-developed head and very thick crnra, both croofed. but the

anterior more so than the posterior. There is a considerable round aperture between

them; the base is broader than in the last family, and slightly concave.

This stapes is truly intermediate between that of the Pinnipedia already noticed and

those to follow. It differs from the former in having an intercrural space and a wider

base, and from the latter in that the space between the crura is very much more limited,

and, the crura being almost parallel, the base is therefore narrower.

Tn the SrenorHyNCHINA (see Pl. LIX. fig. 26, Lobodon carcinophaga*), the head of the

* Syn. Stenorhynchus serridens, Owen, P.Z.8. 1843, p. 131, & Cat. Ost. Mus. Coll. Surg u. p. 641.

404 MR. A. H. G. DORAN ON THE MORPHOLOGY

malleus closely resembles that of Macrorhinus. The excavation in front of its large, flat,

triangular head is deeper, the articular surface is similar but wider, the neck and the

manubrium are even shorter; the latter differs from that of the Elephant-seal (Macro-

rhinus) in being more recurved and spatulate at the tips; its outer aspect is very

broad.

There is uo trace of a processus muscularis, or of the special articular process seen

in Phoca vitulina and hispida; but I find the tendon of the tensor tympani is inserted

in a depression on the exact site of the latter process in a malleus of L. carcinophaga.

This would seem to indicate that the extra articulating process of some Seals is a

modified processus muscularis, though nearer to the head than that process is usually

placed; but, on the other hand, it will be seen that in Phoca barbata, where no trace

of the second articulating process can be seen, there exists a sharp, tubercular processus

muscularis in the exact position of the same in the Felide and Canide.

In the incus the posterior part of the body is more developed than in any Seal ex-

cepting Macrorhinus. Internal to the facets is a deep pit, with a deep groove, which

runs in a line with the groove on the articular surface ; this peculiarity is better marked

in Lobodon and Stenorhynchus* than in the Elephant-seal. The posterior crus is very

short and stout; the processus longus is short, thick, and sharply curved; it is almost

plane, not channelled, on its inner aspect, and the Sylvian apophysis is placed a little

to the posterior side of its extremity.

Hence this bone resembles that of Macrorhinus in its peculiar depression internal to

the articular surface, but Phoca in its processus longus, which, however, is not grooved

as in that animal.

The stapes is rather aberrant. In the Crab-eating Seal (L. carcinophaga) it has long,

very divergent, and much flattened crura, with or without an aperture. 'The base is wide,

thick, and concave horizontally. Inthe Sea Leopard (8. leptonyx) the flattening of the

crura is very marked ; both limbs have rather crooked outlines.

Among the true PHoctna, the ossicula are remarkable for marked differences in closely

allied species. These will be enumerated after describing the ear-bones of the common

Seal (Phoca vitulina) (Pl. LIX. figs. 27, 28, & Pl. LX. fig. 1).

The head of the malleus is flattened, as in the Seals already described; it is rather

small, and nearly circular in outline. Its upper border, the site of the globular part

of the head in most mammals, is quite thin and sharp. The anterior aspect is com-

pletely occupied by a deep concavity (Pl. LIX. fig. 27.¢). The articular surface occupies

more than half the posterior aspect; so that it is more extensive than in Macrorhinus,

but not quite so large as in the other Seals already described. As in most of them, it

is narrow vertically, and very broad horizontally; and from the sharp, perpendicular

ridge on the facets, it is convex in the latter sense, though vertically it is distinctly con-

cave; even the ridges slope downwards to meet each other, and are divided (or, rather,

* Prof. Flower informs me that Prof. Peters (Monatsb. Akad. Berlin, 1875, p. 393), in a footnote, has notified that

the term Stenorhynchus applied by F. Cuvier (1824) to the Small-nailed Seal had been preoccupied by Lamarck (1819)

for a genus of Crabs; hence Prof. Peters proposes to substitute the term Ogmorhinus for the Seal-genus in question.

T haye preferred, however, to use Stenorhynchus as that at present best known to naturalists.

OF THE MAMMALIAN OSSICULA AUDITUS. 405

merely indented) by a faint groove. The neck is thin and subcylindrical, rather long,

but never so regularly cylindrical or so prolonged in this species and genus as in Cysto-

phora cristata. Close to and a little below the articular surface there springs from the

neck a little projection in the form of a much-truncated cone, directed inwards and

slightly downwards and forwards; its extremity is quite plane and round, and articulates

with a similar projection from the incus (Pl. LIX. fig. 28, & LX. fig. 1, eas). This

peculiarity, as well as the concavity in the head, is already well known to anatomists.

The processus muscularis is absent. The manubrium closely resembles that of Cysto-

phora, but is slightly longer ; besides, it has a distinct processus brevis, which is curved

upwards towards the neck in well-preserved specimens.

The incus closely resembles that of Cystophora; the articular surface is surrounded

posteriorly and internally by a deep ear-shaped depression; the posterior crus is very

short, and generally thick; the stapedial crus is short, rather thin, boldly curved

inwards, and deeply grooved on its internal aspect, bearing on the outer and interior

aspect of its base a projection as well developed and similar in form to that on the back

of the neck of the malleus, with which it articulates (Pl. LIX. fig. 28). When these

ossicles are articulated, a small foramen exists between the ordinary and the peculiar

phocine articulations (Pl. LX. fig. 1). The extremity of the processus longus, which is

rather thin, bears a Sylvian apophysis, happily compared in its form by Hyrtl to a house-

fly’s proboscis.

The stapes has very stout, rather short and crooked crura; the anterior is most bent;

they diverge considerably towards the base, and have an almost circular aperture between

them ; the base is also thick, and quite concave in its long diameter.

Such are the ear-bones of Phoca vitulina; and no essential variations have been found

in numerous specimens examined by me at the Royal College of Surgeons. In a

specimen of the left ossicula of the Harp-seal (Phoca grenlandica) (Pl. LIX. fig. 30)

the neck of the malleus does not bear a trace of the singular extra articular process ;

but Hyrtl has observed such a projection in this species. There is a faint protuberance

at the back of the neck, close to the manubrium, which may resemble the processus

muscularis seen in P. barbata. The neck is nearly as long (though not nearly so regu-

larly cylindrical), and the manubrium as short, as in Cystophora. The incus (Pl. LX.

fig. 3) wants the extra articular projection, as might be expected ; its crura are otherwise

exactly similar to P. vitulina; the limbs of the stapes, though rather straighter, are quite

as divergent. .

In the Bearded Seal (P. barbata, Pl. LIX. fig. 29) the extra articular process of the

malleus is quite absent; but on the back of the neck, near the root of the manubrium, Is

a distinct processus muscularis (absent in the def¢ malleus of one specimen in the College

collection, though present on the right side). In describing the Stenorhynchina, I dis-

cussed the relationship of the true processus muscularis to the extra articulating process

and the deep depression found on its site in that Seal. The fact that in P. barbata there

is a true processus muscularis much lower down the neck than the articulating process

is placed in P. vitulina, is against the probability that the latter is a modification of the

former; but the insertion of the tendon of the tensor tympani into a deep pit situated in

SECCND SERIES.—ZOOLOGY, VOL. I. 31

406 MR. A. H. G. DORAN ON THE MORPHOLOGY

the malleus of Stenorhynchus and Lobodon, on the exact site of that articulating process,

is in favour of that theory, which cannot be disproved till a malleus is found with both

the processes coexisting. |

The incus of P. barbata (Pl. LX. fig. 2) most resembles that of P. grenlandica, as

both necessarily want the extra articulation; but the groove around the inner side of the

facets is faint. The anterior crus of the stapes is very crooked.

In a complete set of the ear-bones of the Ringed Seal (P. hispida, Pl. LIX. fig. 31)

the malleus and incus closely resemble those of P. vitwlina, having the extra articular

process on both bones; there is no trace of a processus muscularis. The stapes has a

very large space between the crura, which are very divergent; and the anterior is much

bent; the base is very broad; and, in fact, this bone in this little Seal has all the

distinctive features of this subfamily well pronounced.

I have only procured a single incus of the Grey Seal (Halicherus grypus, Pl. LX.

fig. 5); it resembles that of P. vitulina, except that the extra articulating process is

much less developed. .

The foregoing observations on the auditory ossicles of the Carnivora may be con-

veniently abridged in the following manner :—

I. The ossicula of the Fissipedia differ from those of the Pinnipedia very strongly, in

type as well as in consistence.

Il. The laminated form of the malleus prevails throughout the Fissipedia, excepting

in Herpestes and allied genera, where the lamina is nearly obsolete.

III. The Felidz possess a malleus which bears a long, curved, and slender processus

muscularis, and a long manubrium, with a narrow, but distinct and sharply bordered

outer aspect. The processus brevis of the incus is slender and rather long. The crura

of the stapes, as in all these Carnivora, are straight and moderately divergent.

IV. In the Viverride the malleus (except in Herpestes and Swricata, whereof the

peculiarity is mentioned above; moreover the processus muscularis is ox the manu-

brium, away from the root, in these genera) has a very ill-developed processus muscularis,

and the outer surface of the manubrium is even narrower than in the Cats, yet still

distinctly bordered. The incus and stapes are very similar to those of Felis.

V. The extreme narrowness of the outer aspect of the manubrium and the very

small processus muscularis of the tolerably large malleus, of Cryptoprocta ally it more

closely to Viverra than to Felis.

VI. In Hyena the processus muscularis of the malleus is very stout and straight.

The outer surface of the manubr’ wm is very broad near the tip.

VII. In Proteles the processus muscularis is as stout and as straight as in Hyena;

but the outer surface of the manubrium is very broad throughout, as in Canis.

VIII. In the Canidz the processus muscularis of the malleus is as long, curved, and

slender as in the Cats, but the outer surface of the manubrium is very broad throughout.

IX. In the Mustelide the handle of the malleus is much shorter, and forms a wider

angle with the neck than in any of the above Fissipedia. The processus brevis of the

incus is very short, as in the Bears.

OF THE MAMMALIAN OSSICULA AUDITUS. 4.07

X. In the Procyonide the manubrium of the malleus is long, its outer aspect as

narrow as in the Viverride, and the processus muscularis as ill developed as in that

family. The processus brevis of the incus is shorter (except in Masa) than in the already

named groups, excepting the Weasels.

XI. In Bassaris and Aluwrus the malleus has the same characters asin Procyon. The

lamina is very wide, as in most Bears; but the processus muscularis appears never to

be absent. The processus brevis of the incus is very short.

XII. In the Urside the manubrium is long, with a very narrow outer surface; the

processus muscularis is very short, or quite obsolete, the lamina wider than in the Cats

and Dogs. The processus brevis of the incus is very short, and the stapedial process

long and divergent from the body. In the latter ossicle it is allied to the Weasels, Pro-

cyonide, Bassaris, Hlurus, and to the Kared Seals.

XIII. In all the Seals the ossicles are of very dense consistence; and in all but the

Otariide they are very large, absolutely and proportionally.

XIV. In the Otariidz the ossicles are small; the incus is of the ursine type, but the

malleus and stapes decidedly phocine.

XV. In TPrichechus the malieus resembles that of Phoca, excepting that the manu-

brium is extremely short. The incus is very like that of the Phocina, but the stapes

like that of Macrorhinus.

XVI. In the Cystophorina, the ossicula of Cystophora are very similar to those of

Phoca; but in Macrorhinus the malleus and ineus more resemble those of the Steno-

rhynchina, whilst the stapes is of the Walrus type.

XVII. In the Stenorhynchina the malleus has a very large head, and a manubrium

almost as short as in Trichechus; the body of the incus is very bulky. The stapes is of

slender make compared with the same in the Walrus and Macrorhinus. Lobodon and

Stenorhynchus are very similar in their ear-bones.

XVIII. In the Phocina the head of the malleus is excavated, and the articular sur-

face of a distinctive form, as in all the Fissipedia. The neck is not as long as in Cysto-

phora cristata, but longer than in the other Seals; the manubrium is of moderate

length. The incus has often a special process, articulating with a corresponding deve-

lopment from the malleus. The stapes has rather stout crura; an aperture is constant,

being never effaced as in Zrichechus and Macrorhinus. Neither is that ossicle nearly

so bulky as in those genera; but it is a little stouter than in the Stenorhynchina.

Tue OssICULA OF THE RODENTIA.

This order is remarkable for extreme variety in the form of the auditory ossicles in

different families: nearly every type of malleus may be observed among Rodents; a

peculiar kind of ankylosis between that ossicle and the incus is exclusively found in

certain groups of this order; and there is likewise great variety in the form of the

stapes.

To study the peculiarities of the ossicula of the ScruRIDA, a set should be selected from

a skull of one of the fine large foreign. species, such as the Malabar Squirrel (Sciwrus

312

4.08 MR. A. H. G. DORAN ON THE MORPHOLOGY

maximus, Pl. LX. fig. 6). The head of the malleus of that animal is large, rising well

above the articular surface, but much flattened laterally. The articular area is deeply

cut; each facet has a vertical ridge running down the middle, meeting its fellow at the

groove of separation. Of the two segments thus formed in each facet the outer are most

concave, the inner almost flat. There is absolutely no neck; and the manubrium runs

almost in the axis of the body, and springs from close under the lower border of the arti-

cular surface. This process in S. maximus is narrow throughout, but flattened laterally,

and terminates in a well-developed disk. Halfway down its inner border is a slight

projection ; this is the processus muscularis (Pl. LX. fig. 6, pm),—very frequently in that

position among Rodents. The processus gracilis, which I have found entire ina large

adult skull, is very long, thin, and straight; and a small area of friable opaque bone

extends from the head to its root. The incus has a moderately developed body, a short,

pointed, and non-divergent posterior crus; and the processus longus is broad at the root

vertically, long, very straight, and widely divergent; it turns up very sharply at the

extremity, which is tipped by a minute elliptical Sylvian apophysis. The stapes is

large, with extremely slender crura; the anterior is the thinnest ; both are slightly curved

and very divergent. There was no bony canal between the crura in the skull whence

the stapes was extracted for the College collection ; but an intercrural canal was found in

S. vulgaris, S. bicolor, and S. carolinensis. The base is thin and slightly bullate towards

the vestibule; its extremities, especially the anterior, extend considerably beyond the

attachment of the crura.

In the Jelerang Squirrel (S. bicolor) the ossicula are very similar, but the termination

of the manubrium is spatulate rather than discoidal. This process, moreover, is broader

at the root; and the processus muscularis, on the inner border, is very prominent. In

the Common Squirrel (S. cwlgaris) the manubrium mallei is even broader, but the

termination is an almost perfect disk. The processus muscularis is in the same position

as in S. mazimus. In a young British Squirrel’s skull I found the processus gracilis

of the malleus nearly five millimetres in length, running free for two fifths of its length

from the ossicle, beyond which it becomes a little dilated, and is fused by its inner border

to the neighbouring bone; its extremity is a little rounded.

In Tamias striatus (Pl. LX. fig. 9) the malleus resembles closely that of S. vulgaris in

every respect, especially in the manubrium; but at the point where that process leaves

the body of the ossicle there is a blunt angular projection, a rudimentary processus brevis

The body of the incus is very shallow. The stapes has a bony canal between its

crura *.

In the European Souslik (Spermophilus citillus, Pl. LX. fig. 10) the malleus, in general

respects like that ossicle in Sciwrus vulgaris, has a still more distinct trace of a processus

brevis (pb) than has Tamias. The incus is of the Zamias form; and there is a bony

canal between the crura of the stapes.

The malleus of Anomalurus fraseri (Pl. LX. fig. 8) is of about the same size as that of

Sciurus bicolpr; the manubrium is broader and more flattened laterally than in the true

* In the Bristly Ground Squirrel (Xerus setosus) the ossicula are very similar.

OF THE MAMMALIAN OSSICULA AUDITUS. 409

Squirrels, and ends in a very narrow, spatulate, and not discoidal termination. There is

no trace of a processus brevis. The body of the incus is well developed, the anterior

part as much as the posterior. The stapes differs from that of the true Squirrels ; its

crura are short, stout, and almost straight, not very divergent. There is no intercrural

canal in the recent skull. The base is rather thick, quite flat towards the vestibule, and

projects more than a millimetre beyond the insertion of both crura. In the general

characters of its ossicula, Anomalurus is as nearly a true Squirrel as it has recently been

shown to be, in its general anatomy, by Mr. H. R. Alston *.

In the Flying Squirrels (Pteromys, Pl. LX. fig. 7) the malleus, in general resembling

the British Squirrel’s, has a more distinct trace of the processus brevis on the manubrium

even than Spermophilus. The termination of the handle is spatulate. The body of the

incus is as shallow as in Tamias, Spermophilus, and many of the smaller species of Sciwrus.

The stapes is of the same form as in the above-named genera, and unlike that of Anoma-

lurus, having slender, widely divergent crura, occupied in lifetime by a bony canal; the

anterior crus is inserted into the base some distance from its extremity, the posterior

joining the base almost precisely at the opposite end.

The above descriptions show that Sciwrus is the most specialized of the genera already

examined, particularly in the absence of any processus brevis to the malleus, which is

not as broad as in most Rodents. The distinctions from Sciwrus in the other genera

already discussed will be seen to be truly transitional when allied families are

considered.

In the Marmots (Arctomys, Pl. LX. fig. 11) the sides of the malleus are flatter than in

Sciwrus, and the head rises into a conspicuous blunt point high above the articular surface,

which is extremely deep and narrow. All this gives a peculiar aspect to the ossicle.

The manubrium is much broader at the base than in any true Squirrel, though other-

wise of the same type; the extremity is very faintly spatulate, the inner border very

angular. The neck is quite wanting; and there is hardly any trace of a processus

brevis. ‘The incus has a very wide shallow body; the crura are very similar to those

in Sciwrus. The stapes is a proportionally very large bone, with slender, bowed, and

widely divergent crura. There is a dense bony canal between them during lifetime; and

they are inserted almost exactly into the corresponding extremities of the base, which is

very narrow vertically, and slightly convex towards the vestibule. The characters of the

malleus and stapes separate Arctomys from the Squirrels: in the former bone this genus

approaches the Beaver, where, however, there is a neck and a very prominent processus

brevis. The Marmot’s malleus, with its broad, flattened manubrium, is of the type most

prevalent in this order.

In the Dormice, Myoxip# (Pl. LX. fig. 18, Myoaus glis), the head of the malleus is as

compressed laterally and as flattened at the top, above the articular surface, as in Sciurus,

and the articular surface is as deep and narrow vertically. So far it differs from

Arctomys, where the head rises high and the articular surface is wide and shallow. How-

ever, the malleus of Myoxus resembles families yet to be described in having a faint

“On Anomalurus, its Structure and Position,” Proc. Zool. Soc. 1875.

410 MR. A. H. G. DORAN ON THE MORPHOLOGY

trace of a neck and a characteristic manubrium, very thin, very broad at the base, and

still broader at the middle, where an angular projection from the inner border constitutes

the processus muscularis. There is a distinct angle corresponding to the processus brevis ;

and the outer aspect of the manubrium, though very narrow, is distinctly bordered off

from the sides, as in the Carnivora; the extremity is distinctly spatulate. The malleus of

both Myoxus glis and M. muscardinus much resembles the same among certain Muridee,

where the lamina, conspicuous in Jus, is nearly suppressed.

The incus in the Dormice has a longer and more slender processus longus than in

other families of Rodents; the posterior crus is short and very thick.

The stapes has very slender and straight crura, neither of which is inserted into the

extremity of the base, which is quite plane. No bony canal exists between the crura in

the two species above mentioned.

In the Beavers, Castorip# (Pl. LX. fig. 12), the head of the malleus is still large in

proportion to the size of the whole ossicle, as in all the above families; it is also rather

flattened laterally: the articular surface is very wide vertically, as in the Marmots; but

the head does not rise high above it. There is a narrow lamina between the head and

the processus gracilis, and a distinct, much constricted neck. The manubrium is very

broad laterally ; at the outer aspect of the base is a very prominent angle (pd); it is

recurved and spatulate at the tip; but the processus muscularis (pm) is placed close

to its root, so that there is no angular projection on the inner border. Such is the case

in many Rats and Hares; but in the form of the head of the malleus the Beaver more

resembles the Squirrels.

The processus brevis of the incus is very short and pointed, it diverges distinctly, as in

most of the Muride. The body is square and well developed; the processus longus is

rather long, and supports a well-developed Sylvian apophysis. The stapes of the Beavers

is peculiar, it is large, with very slender, widely divergent, and rather short crura, both

inserted some distance from the corresponding extremity of the base, which is very large,

being long horizontally and wide vertically. There is no bony intercrural canal.

In the Murnipa, Rats and Mice, the malleus retains the singular form seen, with modi-

fications, in Arctomys, Castor, &c., in its less typical genera; but in the genus Mus and

allied genera the broad lamina seen in Sorex, the Carnivora, and the Ungulata reappears.

Even in more aberrant genera the lamina is always developed, though to a less degree.

In Mus decumanus (Pl. LX. fig. 14) the head is little developed and deeply excavated

by the articular surface; the neck is long, thin, and much curved, and, together with

the head, is united to the processus gracilis by a broad area of lamellar bone (/); this

lamina is bordered above by a linear prolongation of compact bone from the front

of the head (phm), the processus gracilis bordering it anteriorly and below. From

the outer aspect of the neck projects an orbicular apophysis (oa), a prominent

spherical knob of bone, as seen in the Shrews. It displaces the manubrium considerably

outwards; and, as in Sorea, it is probably an extreme development of the boss sent

outwards from the shoulder of the incurved capitular portion of the first postoral

bar *. The tensor tympani is not inserted into it, but into a minute tubercle close to

* «On the Structure and Development of the Skull of the Pig,” by Prof. W. K, Parker, F.R.8., Phil. Trans.

1874, p. 301,

OF THE MAMMALIAN OSSICULA AUDITUS. ALL

the root of the neck, and just below it. The relations of the processus muscularis to

the orbicular apophysis will be considered when the ossicula of the Shrews are described,

as certain interesting homologues are seen among the Insectivora, throwing much light

upon the subject.

The manubrium is much flattened laterally, very broad at the base; and, as in Castor,

the processus muscularis is at its root, not halfway down its inner margin. The pro-

cessus brevis is an angular projection concealed by the orbicular apophysis, which lies

above and internal to it.

The incus has an ill-developed, slightly divergent, short crus. The processus longus is

stout, and projects far forwards.

The stapes, as in other Muride, has slender, long crura, not very divergent towards

the base, which is thin and narrow; there is no intercrural canal.

us ratius closely resembles the above in its ossicles; in IZ. musculus the orbi-

cular apophysis is very well developed, and was first described in that animal by

Hagenbach.

In the Golden-bellied Beaver-Rat (Hydromys chrysogaster) the malleus, being larger,

shows the murine characters very well (Pl. LX. fig. 16); the spherical knob of bone is

proportionally less developed than in the smaller mice. Below this orbicular apophysis

a tubercular projection can be detected; this is the real processus muscularis; I have

seen the tendon of the tensor tympani still attached to it. The processus brevis of

the incus is very short.

In Hapalotis mitchelii (Pl. LX. fig. 15) the orbicular apophysis is ill iewclepen and

the tubercular processus muscularis, close to the root of the manubrium, is plainly

visible with the aid of a pocket-lens. The manubrium itself is rather shorter than in

Mus. The anterior crus of the incus is far stouter than the posterior. The stapes is of

the same form as in the past genera of the Muride; and, as in them, there is no bony

canal between the crura.

In the Water Voles (Arvicola, Pl. LX. fig. 17) the lamina of the malleus is less deve-

loped, the orbicular apophysis is quite absent, and the manubrium is rather short and

very broad at the base. The processus muscularis is tubercular, and is placed on the

inner side of the neck, close to the manubrium, as in Mus. In A. cmphibius I have

found a bony canal between the crura of the stapes, but not in 4. arvalis.

In the Musquash (Fiber zibethicus, Pl. LX. fig. 18) the head and lamina of the malleus

resemble those of Arvicola; but from the front of the head a very stout process of bone

(ib. p hm) runs to join the upper border of the lamina: as the true processus gracilis is

very slender, the former might, on hurried observation, be mistaken for it. This process

from the head exists in the mallei of most mammals, but rarely to so highly developed

an extent as in Fiber. Its homologue will be seen to form a conspicuous feature in the

Cetacean malleus. The manubrium is extremely broad and flattened, even for a Rodent ;

the processus muscularis is at: the root of the manubrium, asin Mus. The incus has an

ill-developed processus brevis, and a long, divergent processus longus.

The stapes has the same form as in ws and the other Muride, the crura being long,

412 MR. A. H. G. DORAN ON THE MORPHOLOGY

slender, and not very divergent near the base. There is a bony canal between them in

the recent skull. — .

In the Hamster (Cricetus frumentarius) the malleus has nearly as well-developed a

lamina as in Jus; there is no orbicular apophysis; and the manubrium is not so broad

at the base as in the other Muride. The processus brevis of the incus is very ill-

developed, a character more pronounced in Castor and Fiber than in Mus. There isa

bony canal between the crura of the stapes.

Asan example of the Siphneinze, the malleus of Hilobius talpinus is somewhat rat-like

in form, having a small head and a distinct lamina; the manubrium bears a processus

brevis close to its root. It is sometimes partially ankylosed to the incus. The incus

itself has a very ill-developed body with well-developed crura, a feature seen in the

fossorial rodent families distinct from the Muride, and about to be considered. ‘The pro-

cessus longus is much longer than the posterior crus. The stapes has very slender crura,

between which runs a very dense bony canal in the recent skeleton.

Coming to the SpaLaciD#, considerable variety will be found in the ossicula of pro-

minent genera. In Spalax typhius (Pl. LX. fig. 19) the head of the malleus is ill-

developed and laterally flattened, with a very shallow articular surface, hardly any trace

of alamina, and a very constricted neck. The manubrium runs almost in the axis of the

neck; it is short, and is very broad at the base, much laterally compressed; there is a

prominent angle representing the processus brevis. In its malleus Spalax bears the

same relation to Geomys that Castor does to Arctomys, the mallei in Geomys and

Arctomys being quite neckless ; otherwise, in the flattened heads, with wide and shallow

articular surfaces, they all bear a certain relation to one another; and all seem inter-

mediate between Sciurus and Mus.

The incus of Spalax much resembles that of Hllobiws; but the body and articular

surface are still shallower *; the crura are very similar in form and proportion. The

stapes is of an extraordinary and quite unique type. It is very large, measuring from

head to base two millimetres, the base of the skull measuring from the preemaxilla to the

occipital foramen not quite five centimetres, whilst in a skull of Capromys, measuring

over eight centimetres, the stapes is only a millimetre and a quarter in height. The

head is a small narrow rod; the anterior crus is very straight, and runs forward at a great

angle to the posterior, which is much bowed, and inserted into the corresponding

extremity of the base almost vertically under its origin. The base is well developed,

convex, both vertically and horizontally, and very reniform, its lower border being con-

cave. There is no bony intercrural canal in the recent skull.

In the Bay Bamboo Rat (Rhizomys badius, Pl. LX. fig. 20) the head of the malleus is

well developed and not much compressed laterally : the articular surface is much deeper

than in Spalax or Hllobius; and the neck is long and constricted. There is a trace of

a lamina; and the head is prolonged into a point forwards as in Fiber; near the root

of the manubrium is a small tubercular processus muscularis. The manubrium is longer

than in Spalax, and forms a sharper angle with the neck; it is of similar shape, and

very broad laterally, The malleus of Rhizomys bears a strong resemblance to that of

* The posterior part of the body is much deeper than the anterior.

‘OF THE MAMMALIAN OSSICULA AUDITUS. 413

some of the less-typical Muride, especially Fiber. The ossicle is decidedly not so

specialized as that of Spalax. The incus has a very shallow body; the posterior crus is

less developed than in the other genera, though the processus longus is stout and long

—a murine character. The stapes has straight crura, more divergent than in the Muride,

and a very broad base; there is no canal between the crura in lifetime.

In the subfamily Bathyergine the genus Bathyergus exhibits in its ossicles a feature as

distinctly allying it to the Cavies and Porcupines as the past fossorial rodents are allied

by other peculiarities to the Muride, and more distantly to the Sciuride.

The head of the malleus (Pl. LX. fig. 22) is much elongated anteriorly, but not so

much as in the Hystricide. The neck is thin and constricted; and there is a distinct

laminar expansion beneath it. The manubrium is rather short ; it is tolerably wide near

the base, but gets very narrow towards the tip, and has a distinct processus brevis, and a

small stout tubercle on its inner border, representing the processus muscularis, as in many

Squirrels. The head of the malleus is completely fused to the body of the incus by their

articular surfaces, the contour of which is deeply marked all round the seat of ankylosis ;

the bony union involves the neck and a part of the inner border of the manubrium,

from the root to close above the tubercle for the tendon of the tensor tympani, the manu-

brium being fused to the processus longus of the incus nearly down to the point where

it turns inwards towards the head of the stapes. This ankylosis was observed by Hyrtl.

Whilst the malleus differs from the hystricine malleus in the head being less produced

forwards, the incus is distinguished from that of the Porcupines in having the very

shallow body and proportionally large crura seen in the rest of these fossorial animals.

The posterior crus is particularly long. The processus brevis is as well developed as in

Spalax or Ellobius, and hence different from the same in Khizomys.

The stapes (Pl. LX. fig. 22) is a large bone, though not proportionally as great as in

Spalax. Both crura are straight, not very long, and widely divergent. The base is very

broad, and extremely convex towards the vestibule ; the crura are inserted some distance

from both extremities. There does not appear to be any bony canal between the crura ;

at least such does not exist in the skulls of Bathyergus in the College collection.

Among the Saccomy1p®, in the Pouched Gopher (Geomys bursarius, Pl. LX. fig. 21)

the malleus is much flattened laterally, and the articular surface is extremely shallow,

long yertically and narrow in horizontal measurement ; there isno lamina, no trace of a

neck ; and the manubrium, which runs almost in the long axis of the head, is little com-

pressed laterally, and has no processus brevis. It more resembles the malleus of a

Beaver or Marmot than the same in Sciwrus or Mus.

The incus is of singular form; its body is extremely shallow; so is the articular

surface; the processus brevis is very stout and blunt, and disproportionately large,

whilst the anterior crus is very slender throughout, bearing a well-developed pedun-

culated Sylvian apophysis at its extremity. The stapes has rather stout well-diverging

crura; the aperture is occupied, in the recent skull, by a very delicate ossified canal for an

artery. On its base there projects towards the vestibule a very large bulla, better

developed than in Mustela or Hyraz, and rivalling the same condition already described

by Hyrtl as existing in Phalangista cookii.

SECOND SERIES.—ZOOLOGY, VOL, I. BE

414 MR. A. H. GZ. DORAN ON THE MORPHOLOGY

It is evident that Geomys is more nearly allied to the families already described than

to the Cavies, and is not in any way so related to the latter, in the characters of its

otosteals, as is Bathyergus.

In the Jerboas, Drpoprp#, very interesting transitional characters may be observed.

In one adult skull of Dipus egyptius I have found the malleus and incus unanky-

losed (see Pl. LX. fig. 23) on both sides; in other crania of this Jerboa these ossicles were

firmly united. The head of the malleus is extremely small, and deeply excavated by its

articular surface. The neck is slender and constricted; there is a broad and distinet

lamina, along the front of which a portion of the head is prolonged as in Fiber (phm).

The manubrium is very thin and fragile; itis compressed laterally and broad at the base,

which has a curved upper border, ending externally in a true processus brevis (p 6).

The outer aspect of the handle dilates into a very broad, spatulate, slightly recurved

extremity ; its inner border is stout, and has a sharp angle not far from the tip; but the

tensor tympani muscle is not inserted into that projection, being attached to a little

tubercular processus muscularis where that border joins the neck (pm). The incus

of Dipus has a moderately developed body, a stout posterior crus, and a slender very

divergent processus longus. The stapes has long, exceedingly slender crura, not widely

divergent ; the base is narrow horizontally. I have found a canal between the crura in

some skulls of Dipus, but not constantly.

Altogether the malleus of Dipws approaches to that of the Muride, having a small

head and wide lamina, as in ws, and avery broad manubrium, as in Fiber. In its

frequent fusion with the incus it allies itself to the Porcupines and Cavies, where, how-

ever, the head is always well-developed at the expense of the lamina.

The malleus of Pedetes caffer (syn. Helamys capensis) is more Hystricine, since the

head is much developed at the expense of the lamina, and the bone is firmly fused to the

incus. The stapes, too, has well-divergent crura.

Among the Porcupines, Hystricip#, the fusion of the two outer ossicles is extreme,

and the line of separation is barely perceptible externally, in Hystrix; but in none of

the rodents to be described does this bony union extend beyond the normal articular

area as in Bathyergus.

The malleus of Hystrix cristata (Pl. LX. fig. 24) has a very well-developed head,

which projects very prominently forwards, taking the shape of a Phrygian cap, the

convex border being above, the concave below; the blunt apex points forwards and

downwards. In a perfect specimen the processus gracilis (pg) may be seen running close

to the lower border of the head, a very limited area of lamellar bone (/) connecting

them. ‘The manubrium, much flattened laterally and very fragile, is trapezoidal rather

than triangular, on account of the sharp angle on its inner border, which is here a true

processus muscularis, the tendon of the tensor tympani being actually inserted into it.

The broad upper border of the base, well-developed here, as in Fiber or Dipus, forms

a very obtuse angle with the outer border, so that the site of the processus brevis

(pb) is not so prominent as the angular processus muscularis which is readily

mistaken for it in a detached bone. The outer border dilates very gradually into a

narrow, spatulate, slightly recurved extremity. The incus, which is firmly ankylosed

_OF THE MAMMALIAN OSSICULA AUDITUS. : A415

to the malleus in quite young skulls, has a well-developed body and stout crura, the

posterior diverging as much as the processus longus. The crura of the stapes are very

straight, rather short, and very divergent; the posterior is inserted into the base at a

marked distance from its extremity. There does not ever appear to be any bony canal

between the crura, either in Hystrix or Cercolabes. The base is very broad vertically as

well as horizontally, and rather convex towards the vestibule.

In Hystriz javanica the angular projection on the inner border of the manubrium

mallei is even better-marked than in H. cristata. In the Guiana Tree-Porcupine (Cerco-

labes insidiosus, Pl. LX. fig. 25) the manubrium is like that of Hystriz; but the head is

more massive than in the common Porcupine’s, and still more than in the elongated

mallei of Cavia, Chinchilla, &c. The stapes has a rather narrow but very stout base,

convex towards the vestibule; and the crura, as in M/us and Hydrocherus, are not widely

divergent. There is no bony intercrural canal.

Among those Rodents arranged by Mr. Alston (/. c.) under the family Ocropontipa,

in the Ground-Rat (Awlacodus swindernianus, Pl. LX. fig. 27) the head of the malleus is

still more produced forwards than in Hystriz, and there is as little trace of a lamina; but

in the manubrium it differs from the Porcupine’s in having no projection on the inner

border, the processus muscularis being a prominent tubercle close to the root of that border.

The outer aspect ends in a very broad spatulate dilatation. The fusion of the malleus and

incus is complete; the line of junction is very faint. The processus brevis of the incus

does not diverge so much as in Hystrix. The crura of the stapes are slender and very

divergent ; both are inserted into the base a distinct distance from each extremity; and

there is no bony intercrural canal in the recent skeleton. The base is very broad

vertically and horizontally.

In Fournier’s Capromys (Capromys pilorides, P|. LX. fig. 26) the head of the malleus

is as much produced forwards as in Aulacodus ; the manubrium, however, resembles that

of Hystriz, as the processus muscularis (pm) is placed on the middle of the inner border*.

There is the same obtuseness of the angle corresponding to the processus brevis (p 6) as in

all these Hystricomorpha. The fusion of the malleus and incus is so complete that the line

of demarcation is often quite effaced, especially posteriorly. The stapes is very small in

proportion to the other ossicles, as is seen in Dasyprocta; the crura are thick, short, and

not widely divergent ; the base is very broad ; and an intercrural bony canal exists in some,

but not all, of the skeletons of this rodent in the Museum of the Royal College of Surgeons.

Hyrtl states that in the Coypu (Myopotamus) the malleus and incus are not anky-

losed ; no ossicula of this animal have yet been prepared for the College collection.

In a skull of an Octodon cumingii, six weeks old, the author found the malleus and

ineus quite unankylosed (Pl. LX. fig. 28). The former ossicle is rather different from

the same in the above animals. The upper part of its head is ill-developed, and hardly

projects above the articular surface; the latter is narrow and deeply cut; the neck is

very short; the manubrium is long, and not so broad at the base as in the Porcupines

and Rats; it bears a trace of a processus brevis (pd); and the processus muscularis

(pm) is halfway down the inner border of the handle. There is, as usual in this

* The letters pm in the lithograph are placed too near the processus longus of the incus.

3K 2

416 - MR. A. H: G. DORAN, ON THE MORPHOLOGY

section of the Rodentia, a narrow rudimentary lamina (/). The ineus of Octodon has

‘a well-formed body and slender crura. The crura of the stapes are very thin; its base is

wide; there is a dense bony intercrural canal in the recent skull*.

In the CHINcHILLIDA, the malleus of Chinchilla lanigera (Pl. LX. fig. 29) has a head

extremely produced, so as to look like a slightly flexed index finger, a very charac-

teristic feature. The manubrium is very broad, resembling that of Aulacodus rather

than that of Hystrizx, the processus muscularis being on the inner border close to its

root and not further down. ‘This ossicle isvery completely fused by its articular surface

to the ineus, of which the body is well developed; and both crura are stout and short.

The crura of the stapes are very slender; and there is a distinct bony canal between them

in the recent skeleton.

Among the Dasyproctip#, the Spotted Cavy (Celogenys paca) has the malleus and

incus completely fused. I have found, in a mutilated specimen prepared for the

College collection, that the line of demarcation between the ossicula was best-marked

above and internally; Hyrtl describes it as best-marked below and only partially indi-

cated above. The head of the malleus is not so prominent forwards as in Hystria.

There is no canal between the crura of the stapes in the recent skull.

In the typical genus Dasyprocta (Pl. LX. fig. 30) the fused malleus and incus (anky-

losed even in very young skulls) closely resemble the same in Hystrix. 'The head of the

malleus, however, is less produced, and the two otosteals are more flattened than in the

Porcupine. The stapes is small in proportion to the other ossicles ; the crura are short,

straight, and not widely divergent, so that the aperture is small. The base is very broad,

projecting far beyond the insertions of each crus. In one out of three skulls of D. aguti

I have found a bony intercrural canal; none existed in a cranium of D. antillensis.

In the Patagonian Cavy (Dolichotis patachonica, Pl. LX. fig. 31) the malleus and

incus are completely ankylosed. The head of the malleus is stouter and straighter, and

at the same time more produced than in Dasyprocta. The boundaries of the articular

surface are clearly marked on the outer, but very faint on the inner aspect of the fused

bones. The manubrium of this representative of the Caviip# is trapezoidal as in Hystria.

The angle at the outer aspect of the base is very obtuse, but still it represents the

processus brevis (p06); the well-developed and tubercular muscular process (pm) is

seated on a sharp angle on the inner border, which is very thick above and quite sharp

below that process. The outer border broadens regularly from the processus brevis to

the tip, which is slightly everted and spatulate, its edges spreading out widely from the

thin compressed sides of the manubrium. The body of the incus is both broad and deep ;

and its crura are of almost equal thickness and length. The stapes is proportionally

large, the crura very straight, rather long, and divergent; they are deeply grooved

within; and in the recent skull a bony canal runs between them. The base is broad,

extending considerably beyond the insertions of the crura; it is convex towards the

vestibule ; the convexity is separated from the periphery by a deep channel, so as to

form a large umbo. |

* Tn the figure of the malleus of Octodon the letters pm haye been placed where pg should have been written ;

and J is marked too close to the head of the malleus.

OF THE MAMMALIAN OSSICULA AUDITUS. 417

In the Guinea-pig (Cavia caprera, Pl. LX. fig. 33) the malleus and incus, which are

ankylosed in new-born specimens, resemble strongly those of Dolichotis; but the head of

the malleus is more produced and more slender. The manubrium has the same general

characters; only its thin flattened sides and spatulate extremity are much broader than

in the Patagonian Cavy; there is a distinct trace of a processus brevis. The stapes has

all the characters seen in Dolichotis; there is sometimes a bony canal between its

rura.

In Hydrocherus capybara (Pl. LX. fig. 32) the malleus and incus are ankylosed in the

adult, but often free in young skulls. The head of the malleus is stout and less produced

than in Cavia and Dolichotis; and the manubrium mostly resembles the same in the

latter. The crura of the stapes are long, rather crooked, and slightly divergent; the

base is somewhat narrow, stout, and extends beyond the insertions of the crura. No

bony intercrural canal exists, or at least it is not constant.

Among the Glires Duplicidentati, the LEporiD& possess a malleus which does not

assume the extreme types seen in different Rodents, such as the Sciuride and the

Hystricomorpha. It assumes the less-specialized characteristics of the Marmots, Bears,

and some of the Muride.

In the English Hare (Lepus ewropeus, Pallas) the head of the malleus is ill developed,

rising hardly at all above the articular surface; however, it sends a narrow projection

towards the trace of the lamina. The articular surface is rather deeply cut, and rather

narrow laterally; the facets have a vertical convex ridge across their middle as in many

animals in other orders. Between the body and the processus gracilis is a very fragile

and rather narrow bony lamina, generally damaged in the removal of the bone, but plainly

visible if the malleus of a new-born hare or rabbit be prepared ém st¢w with the tympanic

bone. I have examined it in its natural position in the adult hare and rabbit,

and certainly have found the lamina partially absent. That it should atrophy during

- adolescence is highly probable, seeing that the processus gracilis itself is partially absorbed

in Man and in many lower animals. —

The neck of the Hare’s malleus is very distinct, beimg much constricted and. rather

short. The manubrium is of the sword-shaped form seen in the Beaver, being very thin

and flattened laterally, broad at the base, and well recurved. ‘There is a trace of pro-

cessus brevis (Pl. LX. fig. 34, Z. cuniculus, pb); the muscular process (pm) is situated

on the inner border very close to the neck. The outer edge is very narrow, though

distinctly bordered off from the sides and hardly dilated at the tip. This broad manu-

brium, without an angle on its inner margin, may be considered a central type among

Rodents, differing, on the one hand, from the narrower manubrium of the typical

Squirrels, and, on the other, from the trapezoidal handle of the malleus in the Hystri-

comorpha.

The incus has an ill-developed posterior crus and a very divergent stout processus

longus, bearing a well-developed elliptical and pedunculated Sylvian apophysis. Of

course the articular area is deeply cut; otherwise, particularly in the crura, this ossicle

resembles the same in most Rats and Squirrels.

A418 MR. A. H. G. DORAN ON THE MORPHOLOGY

The stapes has perfectly straight and well-diverging crura; the base is broad, and

projects beyond their insertions. There is no bony intercrural canal in the recent skull.

In the Rabbit (Lepus cuniculus, Pl. LX. fig. 34) the ossicles are identical in form with

those of the Hare. In both these animals the processus muscularis may be situated

absolutely on the neck of the malleus, and not at all on the inner edge of the manu-

brium as is usually the case.

The conclusions arrived at, after the study in detail of the ossicula of the Rodentia or

Glires, is as follows :—

I. In the Sciuromorpha there is a tendency in the malleus to suppression of the neck

and processus brevis. This is complete in Sciurus and Anomalurus; but the neck and

the process are gradually developed in Xerus, Tamias, Spermophilus, Pteromys, Myoxus*,

and Castor, the process being perceptible as a slight projection in the first four genera,

whilst it is well developed and there is a distinct neck in the Dormouse and Beaver.

In Arctomys there is no neck, and the processus brevis is as ill developed as in the

Ground-squirrels and Pteromys; the malleus is extremely flattened, and projects above

the articular surface. As in all other rodents where the incus is free f, it has a rather

square solid body, with the posterior crus but little divergent. The stapes is large, with

wide, slender, divergent crura (least so in Anomalurus) ; a bony canal runs between the

crura in lifetime.

The great breadth of the manubrium, so distinctive of Rodents, is well marked in

Castor and Arctomys, least in Sciwrus. The processus muscularis is situated on its

inner border.

II. In the Myomorpha the laminated malleus reappears in its best-developed form in

Mus, Hydromys, and allied species; in some of these Muride there is an orbicular

apophysis, so frequent in the Chiroptera and the Shrews. In the less-typical genera the

lamina is narrow, the manubrium extremely broad, with a prominent processus mus-

cularis on its inner border. The stapes has long slender crura, little curved or diver-

gent. A bony intercrural canal is rare. ,

III. In the fossorial Rodents remarkable varieties are seen in all the ossicula, especially

in the stapes, which is bullate in Geomys and of unique type in Spalax. Hillobius, Spalaz,

Rhizomys, and Geomys} show distinct affinities to the Muride in the form of the

malleus, whilst in Bathyergus we find that the malleus and incus are fused even to a

srveater extent than in the Hystricidee.

TV. In the Hystricomorpha the malleus is almost invariably fused to the incus in the

adults, but often freein young subjects. In Dipus § the malleus is more like those of

some of the Muride than is that ossicle in the rest of the group. The head of the

malleus is produced forwards moderately in Hystria and its allies, also in Cavia,

* Myowus is rather Sciuromorphic than Myomorphic in its ossicula.

+ Excepting Spalax and its allies.

+ The absence of a neck to the malleus in Geomys allies it to the Squirrels.

§ As the malleo-incudal ankylosis is frequent in Dipus, and probably constant in Pedetes, these genera are not

placed here among the Myomorpha.

OF THE MAMMALIAN OSSICULA AUDITUS. 419

Dasyprocta, and Hydrocherus, but to an extreme degree in Awulacodus, Capromys, and

especially in Chinchilla. The stapes is small, with well-developed crura, not very diver-

gent. A bony canal runs between the crura in some specimens from several species, but

is not constant. t

V. In the Duplicidentati the characters of the ossicles are rather central. The malleus

is of much the same form as in the less-typical Muride; but the head is less developed.

There is never any fusion of the malleus and incus in Lepus.

THE OssICULA OF THE UNGULATA.

The different families of Perissodactyla offer more sharply defined characters in their

auditory ossicles than do the even-toed Ungulates.

Suborder PERISSODACTYLA.

Among the RurNocerorips, the ossicula are all very small in proportion to the

size of the animal. In Rhinoceros bicornis (Pl. LXI. fig. 1) the malleus, from the top

of the head to the top of its long manubrium, measures one centimetre, the incus is

two millimetres high between the crura, and its body is barely four millimetres broad,

all much inferior measurements to the same in a Horse, and hardly above the pro-

portions in a Tapir.

The malleus has an extremely deep and wide articular surface, reducing the head to a

mere cup-shaped process, with none of the free globular portion seen more or less in

most mammals. The upper facet (very prominent from the ill-development of the head)

is not as large as the lower; both are slightly saddle-shaped, and run into each other at

the bottom of the deep excavation without any sharp dividing ridge. The neck is cylin-

drical, curved, and rather narrow, bordered anteriorly by the lamina. The manubrium

is very long and slender: it is bent somewhat inwards as it leaves the neck, curving

boldly outwards near the extremity, which is distinctly spatulate. It forms nearly a

right angle near the neck; a well-rounded tuberosity occupies the site of the processus

brevis; there is a small tubercular processus muscularis on the inner aspect of its root.

The sides of the manubrium are flattened and narrow throughout, the inner border is

rounded, not sharp; the outer angle is still bordered, though extremely narrow. In

front of the neck is a laminar expansion (Pl. LXTI. fig. 1, 2), which is continued (rather

than merely blended below, as in Carnivora and Ruminants) into a very large lamelliform

processus gracilis (pg), shaped like a streamer or pennant, and measuring, in a speci-

men in the College, twelve millimetres, the manubrium being only nine millimetres long.

A process from the head (pm) bounds the anterior and upper border of the

lamina.

Comparing this bone with the corresponding ossicle in other Perissodactyla and

Artiodactyla, the presence of a lamina might lead the observer to the assumption that

the malleus of Rhinoceros is more like that of the Pigs and Ruminants than that of the

Equide; the same remark applies to the very similar malleus of Tapirus. Buta very

little consideration must convince him that the lamina in the Rhinoceros is of a different

type from that of the even-toed Ungulates. It is not, as in them, a broad lamella of bone

420 MR. A. H. G. DORAN ON THE MORPHOLOGY

stretched from the root and adjacent part of the processus gracilis to the neck and head,

but a narrow strip of lamellar bone running with that process far in front of the head

into the Glaserian fissure. Indeed the thin sickle-shaped processus gracilis of Equus

is probably a modification of it. The Rhinoceros’s malleus approaches a type very

frequent in the Insectivora, the Bats, the Marsupials, and some Edentata, as Priodon,

where the processus gracilis is elongated in a very similar manner. But more will be

said on the subject when the ossicles of the latter groups are discussed.

The incus of Rhinoceros is of a very peculiar form, the body being very broad and

shallow, and the crura both widely divergent ; the posterior is long, stout, and blunt-

pointed; the processus longus is rather short, boldly curved, deeply grooved on its inner

aspect, and bears on its extremity a greatly developed Sylvian apophysis, pedunculated

with a slightly concave oval surface towards the stapes. It is absolutely and, as I

believe, proportionally the largest Sylvian apophysis existing, greater than the same in

Galeopithecus, to which Hyrtl gives the palm; but he does not describe the ossicula of

the Rhinoceros.

The stapes has a very large head, slightly convex, to fit the apophysis; the posterior

crus is quite straight, the anterior curved forward just before its insertion. In the spe-

cimen in the College collection there is a thin lamina of bone between the upper halves

of the crura. The base is broad and convex.

The ossicula of R. sondaicus and R. sumatrensis are almost identical in character.

Among the Tarrrips, the malleus of Tapirus bairdi (Pl. LXI. fig. 2) resembles con-

siderably that of the Rhinoceros, particularly in the large excavation for the incus, and

the very slight development of the upper part of the head; but the neck is shorter, more

curved, and not so closely defined in front from the lamina; the manubrium is not so

much bent inwards, and is very little curved outwards at the tip, which is less broadly

spatulate. Besides the processus muscularis there is a small tubercle on the inner side of.

the neck, not on the root of the manubrium as in Rhinoceros. The lamina is broad, and

runs into a similar processus gracilis, which is slender and fragile.

The incus has a very broad body; its interior segment is much less developed than the

posterior, by which it differs from Rhinoceros and approaches Hquus.

In the Equip, the typical Equus caballus (Pl. UXT. fig. 3) has a malleus with a well

developed globular head, exceeding, in that respect, other Ungulates, and a wide, deeply

cut articular surface, whose edges are raised above the level of the rest of the head, upon

which it encroaches very little. The facets are almost plane, but slightly saddle-shaped ;

the groove between them is not well marked. The neck is rather long, somewhat

flattened, and twisted a little outwards near the root of the manubrium, which leaves

it at a right angle, is long, but broader at the base than in the Tapir and Rhinoceros,

The manubrium is only slightly spatulate and recurved at the tip, and has a narrow but

well-bordered outer surface. ‘The processus brevis is represented by a blunt tubercle,

much bent forwards; the muscular process is a small tubercle on the inner side of the

neck, close to the root of the manubrium. The processus gracilis is long and sickle-

shaped, not nearly so broad as in Rhinoceros; and there is little trace of a lamina:

hence the equine malleus is a less specialized bone than that of the other odd-toed

OF THE MAMMALIAN OSSICULA AUDITUS. 421

Ungulates, and far more removed from the Artiodactyle form. The incus is very much

larger than that of the Rhinoceros, both absolutely and proportionally. Its body is rather

broad, and cannot be distinguished posteriorly from the stout, short, and little-divergent

processus brevis; the anterior part is very little developed ; the processus longus is more

slender than the processus brevis, though still rather broad at the base; unlike the pos-

terior crus, it diverges considerably from the body. The stapes is quite triangular, has

a well-developed head, with extremely straight crura, not very divergent towards the

base, and inserted a perceptible distance from each extremity. The base itself is broad

and almost plane.

In Hquus asinus, and still more in #. zebra, the processus muscularis of the malleus

is more developed than in #. caballus, appearing as a slightly curved blunt-pointed pro-

cess. The incus in both the former has a slender and more divergent processus brevis ;

in the Zebra the whole body is more developed. The Sylvian apophysis is a flat sessile

disk; in adults it fuses more or less completely with the processus longus, as in Man.

The stapes of #. zebra is very horse-like, the head being very broad and stout, the crura

straight, and the aperture triangular, with sharp angles; the base is thick, with an

umbo towards the vestibule. The stapes in £. asinus has a smaller head; the anterior

crus is distinctly curved ; and the intercrural aperture is round rather than triangular.

A tubercle, with an ossified piece of the stapedius tendon attached, is often seen in

the stapedes of the Equide and also in many Artiodactyles.

SUBFAMILY ARTIODACTYLA.

This group is interesting for the frequent recurrence of the broadly laminate form of

malleus which prevails among the fissipede Carnivora. ‘The incus, as will be shown, is a

bone of great interest and distinctive value in these animals.

In the Hippopotamus (Pl. LXI. figs. 4 & 5) the malleus has a large head, with a well-

developed spherical upper aspect, and a very broad articular surface, of which the upper

facet is of great breadth, almost plane, but has a vertical convex ridge near the outer

border ; while the lower facet is very narrow, and vertically much elongated, so as to be

tongue-shaped and quite convex in both directions. ‘They blend together undivided by

_ any sharp groove; and the entire surface is somewhat shallow. The neck is short, very

broad towards the neck, and constricted close to the manubrium; it is considerably

flattened laterally; and from its inner aspect, a considerable distance from the root of

the handle, springs a long, straight processus muscularis, of equal thickness throughout,

and hence blunt-pointed. The manubrium is long, it leaves the neck at an obtuse angle,

and is flattened laterally, but not very broad at the base. The processus brevis is a

sharp angular projection bent rather backwards ; the outer aspect of the handle is sharply

bordered from the sides, and spatulate at its extremity, which is but a little recurved ;

the whole process is almost straight. The processus gracilis is very slender and united

for some distance to the neck and head by spongy, not lamellar bone.

This bone is very like the same in the pig, the chief distinction being in the cha-

racter of the articular surface; the spongy bone between the neck and processus

SECOND SERIES.—ZOOLOGY, VOL. I. 3L

422 MR. A. H. G. DORAN ON THE MORPHOLOGY

eracilis represents the lamina so marked in the other Artiodactyla; it exists in the

same modified form in the still larger malleus of the Elephant.

The incus has a very high and rather narrow body; the upper part of the articular

surface is very broad, the lower part forms a deep, narrow groove, to fit the lower facet

of the malleus. Both crura are very short and divergent; there is a small tubercle

over the outer aspect of the processus longus, which crus is concave on its inner side.

This high-bodied incus is thoroughly porcine. The stapes has a comparatively small

head, with a flat tubercle on its posterior side for the stapedius tendon. The crura

are both straight, the posterior is much thicker than the anterior, and they diverge very

considerably, leaving a large space between them. The base is thin, broad, and rather

convex; the whole bone forms an equilateral triangle. Hyrtl considers the aperture

between the crura to be small; but it is very wide in three stapedes of Hippopotamus

in the College collection.

In the Wart-Hog (Phacocherus ethiopicus) the malleus resembles that of Sus, about to

bedescribed ; but the following distinctions exist : 1, the upper part of the head is narrower,

and projects more prominently forwards; 2, the articular surface is wider and shallower,

the lower facet is nearly as large as the upper, both are only slightly and evenly

saddle-shaped, and the groove between them is very faint instead of being sharply cut.

This is a distinct approach to the Hippopotamus. The manubrium is almost straight

towards the extremity. The incus is like that of Sus, except that the processus ‘ brevis”

is absolutely longer than the stapedial crus, and is slender and blunt-pointed. ‘The

stapes is pig-like; its crura are extremely slender.

In the Surpm (Pl. LXI. fig. 6) the malleus is of the broadly laminate type. The head

is well developed, though not so large proportionally as in the Hippopotamus, and

projects forwards rather than upwards; the articular surface is narrow and deep. The

groove dividing the facets is very sharply defined ; the upper is larger than the lower; and

both are saddle-shaped, a vertical convex ridge running near their outer borders, leaving

to each a very narrow outer segment,—a very different condition from that which exists

the Carnivora, where the malleus is of a very similar form. The neck is very narrow,

and boldly curved; a stout lamina connects it with the thin processus gracilis. From

the inner aspect of the neck, a slight distance from the root of the handle, springs a long,

stout, and straight processus muscularis, similar to that of the Hippopotamus. The

manubrium leaves the neck almost at a right angle; it is not very broad laterally at the

base, and has a sharp processus brevis pointing vertically upwards. Its outer aspect

has well-defined borders, but is very narrow; the extremity turns up slightly, but dis-

tinctly, and is but little dilated.

The incus has a well-developed exceedingly square body, with short processes, as in

Phachocherus and Hippopotamus, and contrasts strongly with the incus of the Ox and

Deer, and still more with that of the Perissodactyla. Its articular surface is deeply cut,

to fit that of the malleus; the lower facet is narrow and shallow, a natural and necessary

variation from the same in the Hippopotamus, corresponding to a similar difference in

the malleus. The stapedial crus is scarcely longer than the processus brevis, and sup-

ports a small, discoid, pedunculated Sylvian apophysis. ‘The stapes tends to the quadri-

OF THE MAMMALIAN OSSICULA AUDITUS. 4.23

lateral type frequent in the Ruminants, the head being very broad ; the crura, wide apart

at their origin, are slender, and diverge little towards the base; the aperture is large,

and almost circular. The base is stout, with a very convex umbo towards the vestibule,

as Hyrtl has previously observed.

In the genus Sus all the above characters are strongly marked; the lamina, however,

is much encroached upon even ina feetal pig. In Babirussa alfurus the articular surface

of the malleus has but a faint line between the facets, the lower of which is considerably

narrower than the upper. This approaches the type of Phacocherus. The processus

muscularis is very long and stout; the end of the manubrium is slightly but distinctly

recurved.

In the Peccary (Dicotyles tajacu, Pl. LXI. fig. 7) the malleus has a longer and a

thinner neck than in Sws, much less abruptly curved, and a wide, perfectly papery lamina

between it and the processus longus; the manubrium, which forms a more obtuse angle

with the neck than in the other pigs, has a rather wider outer surface, and a distinctly

broader and more spatulate extremity.

The more compact and less laminar expanse of bone between the neck and the pro-

cessus gracilis in the Pig, as compared with the Peccary, might be explained by its

greater size; and, indeed, in the larger animal in any Order there is often a tendency

to such thickening of that area; still, in the large malleus of the Lion this lamina is

as thin, in comparison with the rest of the bone, as the corresponding structure in the

Cat and Genet; on the other hand, it is denser in the Hyzena, which is intermediate

in size between them. Hence the degrees of density of the lamina constitute fair

generic distinctions.

The incus of Dicotyles is even higher and narrower than in Sus; the processus longus

is rather longer. The stapes has a broad head, very straight crura, and a very thick

base, with an umbo convex towards the vestibule, as in Sus.

In the Chevrotains, Tracunip# (Pl. LXI. fig. 8), the head of the malleus is small;

and the articular surface is deeply cut, with a sharp groove between the facets, which are

both saddle-shaped, a sharp convex ridge running vertically down their surfaces, near the

outer margin, as in the Pigs, whilst in most of the Bovidee the facets are nearly plane.

The lower facet is almost as wide asthe upper. The neck is shorter than in Sws, and con-

sequently much shorter than in most Ruminants; but the malleus resembles that of the

latter group of Artiodactyla in the bluntness of the angle at the outer side of the base of

the manubrium, which, moreover, forms a very obtuse angle with the neck; the processus

muscularis is short, but stout. The incus invariably retains the solid square form seen

in the Pigs; and the crura are as short. The stapes has a small head and more divergent

crura than in Sus; the base is very thick and quite plane; the crura are both inserted a

perceptible distance from its extremities. Hence this bone, with its divergent limbs,

has the form of an almost equilateral triangle, as seen in some Antelopes, particularly in

Gazella, differmg from the almost quadrilateral stapedes of both Bos and Sus. The

corresponding ossicle in the Cervide, having a narrow base, with straight, equal crura,

is intermediate in form.

In Tragulus javanicus the manubrium does not form quite so obtuse an angle with

oL 2

ADA MR. A. H. G. DORAN ON THE MORPHOLOGY

the neck as in Tragulus napw*. On the inner side of the neck of the malleus of 7. hanchil

is a small tubercle-like processus muscularis. The Sylvian apophysis on the end of the

long process of the incus is a thick disk, wider in diameter than the end of the processus

longus, and pedunculated.

In the CamreLipm or Tyxopopa (Pl. LXI. figs. 10 & 11, Camelus dromedarius) the

upper part of the head of the malleus above the articular surface is not much developed ;

and, as in the Pigs, it projects forwards, but does not rise above the level of the articular

area in an upward direction. This surface is very wide vertically ; its edges are much

raised above the level of the rest of the head. The facets are almost plane; the lower is

as long as, but narrower than, the upper; they run into each other without any trace of a

groove. The neck is very short, rather broad towards the head, and very constricted

near the manubrium ; there is only a very small fragile lamina stretched between it and

the processus gracilis. The manubrium is extremely long, rather slender even at the

base, and does not form a very obtuse angle with the body; a bold rectangular projection

corresponds to the processus brevis; the outer aspect is narrow, but sharply bordered,

the extremity distinctly spatulate and only a little recurved.

The incus is very singular and distinctive in form (Pl. LXI. fig. 9, Lama peruana).

It is extremely high and narrow; and the anterior part of the body is nearly as deep as

the posterior—a rare condition in any Order}. The articular surface is equally distinctive ;

it is very wide, corresponding to that on the malleus, and absolutely convex from above

downwards; its margins are very prominently raised above the level of the body; its

lower and anterior extremity stops very far short of the root of the processus longus.

Hence the incus in the Camelide is quite distinct from that of other even-toed Ungu-

lates, even from those where it is high and narrow. The stapes has a small narrow

head, long and rather slender crura, not very divergent; and both are inserted some

distance from the extremities of the base, which is very thin, particularly in proportion

to the same in other Ungulates; it bears a base of an umbo.

The malleus of the Camel and Llama are very similar, in the former the head projects

more forwards than in its American representative. In Camelus and Lama pacos there

is a small, tubercular processus muscularis on the inner side of the neck of the malleus,

hardly perceptible in LZ. peruana.

In the Prong-buck, or American Antelope (Antilocapra americana, Pl. LX1I. fig. 12),

the head of the malleus is very little developed, the upper portion projecting neither

upwards nor forwards; the articular surface is very wide and rather deep, its borders

are much raised; both facets are almost plane; the upper is very broad; the lower is

narrowed, and is sometimes prolonged into a short tail-like extremity, running onto

the neck; but this feature is not constant. I have noticed the same peculiarity

in individual mallei of Goats and Antelopes. The groove between the facets is

distinct. The neck is very long, cylindrical, and only gently curved, it is joined to the

slender processus gracilis by a well-marked lamina. From the inner aspect of the neck

projects a very stout, straight, blunt-pointed processus muscularis. The slender manu-

* The processus muscularis is sometimes obsolete in these species.

+ In Camelus the body of the incus is not so high and narrow as in Lama.

OF THE MAMMALIAN OSSICULA AUDITUS. 425

brium has a well-bordered outer aspect of almost equal breadth throughout; there is a

sharp angular projection at its base; and it is but slightly recurved and very little

dilated at the tip.

The incus is high and narrow; its articular surface is deeply cut ; and when there is a

tailed prolongation of the lower facet on the malleus, a corresponding peculiarity is seen

on that facet on this bone. Both crura are very short; the lower is very divergent and

boldly curved. The stapes is almost quadrilateral; the head is very broad, with a pro-

minent process for the stapedius tendon; the crura are stout and almost parallel; the

anterior, which is the thinnest, is inserted into the extremity of the base, which is rather

thin and almost plane.

In general characters the ossicula of Antilocapra closely resemble those of the Bovide.

The body of the incus is almost as stout on square as in the Pigs and Tragulus; but

this feature is seen in Gazella.

THE Bovyip2.

In this family it is hard to find distinctive features in the auditory ossicles of different

species, while, to add to the difficulty, in many instances these bones undergo considerable

_ changes during the growth of the animal; so that little reliance can be placed on any

conclusions based on the observation of one specimen alone from any species. It is

necessary to examine duplicates as well as examples from different species of the

same genus. Through the kindness of Sir Victor Brooke, Bart., the author has been

enabled to collect a large series of the ear-bones of this order from the skulls in that

gentleman’s fine collection of skeletons of Ruminants.

Beginning with the Common Sheep (Ovis aries), it is highly important to select

complete series of the ossicula from individuals of all ages, for reasons immediately to

be explained.

Taking a set from the skull of an adult sheep (PL LXI. fig. 16), the malleus will

be seen to have the upper and anterior part of its head ill developed, as in Antilocapra,

and unlike Sws and its allies. The articular surface (a most important region to observe

closely) is wide vertically as well as laterally, and also deeply cut; each facet is dis-

tinetly saddle-shaped, with a sharp vertical ridge down the middle. The neck is cylin-

drical and sharply curved, as in most mallei of this type; the lamina is of moderate

breadth. The processus muscularis is curved and prominent, but not very long. The

manubrium is long and slender, compressed laterally, but not broad even at the base,

on the outer extremity of which is a rectangular projection, hardly to be called a pro-

eessus brevis. The outer aspect is sharply bordered and extremely narrow, hardly dilating

at the extremity, which is but little recurved.

From repeated observations, I find that the malleus of an adult domestic Ovis closely

resembles that of a foetal Bos tawrus of about the sixth week (Pl. LXI. fig. 15); but at

birth the characteristic, extremely shallow articular surface, and strongly curved manu-

brium, are always found in the ox; the outer aspect of that process, too, is very broad

in foetal calves, though never in adult sheep. ‘The same articular surface is very deeply

cut in the mallei of lambs.

4.26 MR. A. H. G. DORAN ON THE MORPHOLOGY

The incus in an adult sheep has a well-developed body, which is by no means shallow,

but never as square and solid as in the Pigs and their allies. ‘The great development of

the crura in Ovis makes the body look smaller than it really is. The processus “ brevis ”

is very long and straight, and almost cylindrical in form, though a little compressed

vertically. The processus “longus,”’ well divergent from the other crus, is absolutely

shorter; it bends very abruptly above its termination, which bears a very well-formed

pedunculated and perfectly elliptical Sylvian apophysis. In the incus of a new-born lamb

(Pl. LXI. fig. 17) the body is much shallower, as in many less typical Bovide; that part

of the same ossicle in the adult sheep, on the other hand, clearly approaches the high

well-developed type seen in adult oxen.

The stapes in a large adult skeleton is quite trapezoidal; the head is large and square,

bearing a slight trace of a process for the insertion of the stapedius muscle on its inner

aspect. The crura are thick and deeply channelled towards the aperture ; they are wide

apart at their origin and almost parallel; the anterior is the thinner and longer; so that

base is further from the head in front than behind. The base is thick, its upper border

is strongly arched, and both extremities project very perceptibly beyond the insertion of

the crura.

But in the new-born lamb the stapes is generally quite triangular, the head being

much less developed, and the crura close together at their insertion. It will be seen to

retain features observable among many adult Antelopes—that ossicle in the Sheep, when

full-grown, approaching, as does the incus, the type seen in Bos.

The Ox will now be considered ; and it will be demonstrated afterwards how every grade

of transitional features in all the ossicula, between Ovis and Bos compared, are seen in

every kind of combination in Goats and Antelopes.

In a malleus taken from a large Ox’s skull (Pl. LXI. fig. 14) the head appears no more

developed above the facets than in Ovis; but the articular surface, wide horizontally, is

very wide vertically, the facets gaping far apart. The upper is almost plane, though

still slightly convex; the lower is more perceptibly elevated, and is divided from the

upper by a very faint groove. The neck is short and very strongly curved; the lamina

is moderately wide; the processus muscularis projects from the inner side of the neck

some distance from the manubrium, and is rather short, almost straight, and blunt-

pointed. The manubrium is boldly curved, with the concavity forwards; its outer

aspect is rather broad and sharply bordered, dilating but little and very gradually

towards the tip. The base projects upwards, outwards, and a little backwards, and may

be said to bear a true processus brevis. .

The malleus of a foetal calf at the sixth week (Pl. LXI. fig. 15), on the other hand,

is much more like that of an adult Sheep, and is almost as large. The articular surface

is wide and deep vertically, as in Ovis, but not so deep as in the adult Bos; and the

facets are more perceptibly convéx. The manubrium too is almost straight, and slightly

recurved at the tip, instead of being well bowed forward. In the mature foetus the

malleus has assumed the adult Ox’s character.

The incus in the full-grown Bos has a very well-developed body, its outer and inner

aspects both showing a wide, convex, bony surface; the postero-superior border over

OF THE MAMMALIAN OSSICULA AUDITUS. 427

the processus brevis is twice as long as the antero-internal; and the articular surface,

well to the front, is of necessity wide. The processus brevis is as long and cylindrical

as in Ovis, and much curved outwards; the processus “longus” is not so long, but it

stretches out further than in the Sheep, and does not turn up so sharply near the Sylvian

apophysis, which is similar in form. In the fcetus of a calf at the sixth week the body

is about as well developed, proportionally, as in an adult domesticated Sheep.

Taken as a whole, the Ox has the best-developed incus of all the Artiodactyla, inas-

much as the body is in all directions larger than even in Sws, where the crura are charac-

teristically short, whilst those processes in Bos are even longer than in Sheep and

Antelopes (where the body is shallower), and as long as in the Deer (where it is generally

very shallow and broad).

The stapes of the adult Ox is absolutely oblong. The head is very broad, with a pro-

minent tubercle for the stapedius tendon, which latter is partly ossified. The crura are

quite parallel, or even nearer together at the base than the head in very large spe-

cimens; they are well channelled towards the aperture; the anterior is very thin and

straight, and a little longer than the posterior, which is thicker; the base is very stout,

and projects well beyond the crura. The whole bone is only an extreme modification of

the form seen in Ovis; indeed in the feetal calf that bone is still trapezoidal, or almost

triangular, the head being less broad, and hence the crura nearer together at their

origin.

In the adult Zebu (Bos indicus) the ossicula are more slender than in the Ox; the

manubrium of the malleus is not so boldly curved; the stapes is even more oblong than

in B. taurus; its crura are very long and close together.

The ossicula of the Burrhel (Ovis nahoor) are more like those of a new-born lamb than

those of a full-grown domestic sheep. The body of the incus is shallower, though the

crura are as long; nor is the stapes so quadrilateral as in O. aries.

In the adult Goat (Capra) the malleus has an articular surface as wide as in Ovis, and

hence not so wide or so plane as in Bos. The manubrium is distinctly curved, almost as

much as in the Ox in C. bez, less so in C. hircus, C. megaceros, and C. egagrus, judging

from specimens in the College collection. The body of the incus is sometimes shallow in

this genus; but in the skulls of large goats it becomes as high as in the adult Sheep.

The posterior crus is not very long; the anterior is sharply bent, as in Ovis. But even in

large adults the stapes remains triangular, though the head is always thick, and the crura

a distinct distance apart at their origin; they diverge considerably towards the base,

which is very wide and thick, projecting beyond their insertions, and is quite plane

towards the vestibule. The crura are almost of equal thickness; but the anterior is

rather the longer.

In the Chamois (Rupicapra tragus, Pl. LX1. figs. 22-34) the articular surface of the

malleus is wider than in Ovis or Capra, and almost as wide and as level as in Bos; but

the manubrium is slender and almost straight, as in the Antelopes. The incus resembles

the full-grown Sheep’s in the development both of its body and crura; and the stapes is

of the same trapezoidal form.

428 MR. A. H. G. DORAN ON THE MORPHOLOGY

In the malleus of Nemorhedus (Capricornis) swmatrensis (Pl. LX1. fig. 29) the articular

surface is as in Ovis; the manubrium is slightly curved; and the lamina is extremely

wide. The stapes is, on the other hand, positively triangular, as in most Antelopes, the

head being rather small, the crura close together at their origin, and very divergent ; they

are rather thick; the base projects but little beyond them, and is not very wide. Ina

single malleus of Oreotragus saltatrix, from an adult skull in Sir V. Brooke’s collection,

the articular surface is narrower vertically than in Ovis; the manubrium is long, straight,

and very slender. The malleus in Saiga tartarica (Pl. LXI. fig. 23) is very sheep-like

in the nature of its articular area and in the form of its manubrium; the incus (PI.

LXI. fig. 35) has a very shallow body, with widely separating and diverging crura; the

stapes is quite triangular. .

In Gazella (Pl. LXI. fig. 18) the malleus retains most of the characters of Ovis; but

the articular surface is often wider, the processus muscularis is almost always shorter

and smaller, the angle at the base of the handle is so prominent as to form a true pro-

cessus brevis, and the extremity of the manubrium is well recurved and spatulate, the

whole outer aspect of that process being rather wide. The body of the incus is always

well developed in both directions, so as to take from the length of the crura (which, on

actual measurement, are fairly long) and to make the whole bone resemble the same in

the Pigs and Tragulinee, where the processes are very short, especially the processus

brevis. ‘The stapes is a large bone, of very delicate make, and almost an equilateral

triangle in form; the head is small, the crura very slender and divergent, the base very

wide and plane, extending well beyond the insertion of the crura. The ossicula of

G. dorcas, G. levipes, and G. bennettii are very similar.

The malleus of Antilope cervicapra is of the Gazelle type; but the stapes has stouter

and less-divergent crura. The malleus of Procapra picticaudata (Pl. LXI. fig. 27) is

similar; the incus (fig. 38) has as large a body as in any Gazella. The same remarks

apply to Nanotragus pygmaeus and N. montanus (fig. 13), though even in adults of the

latter species the body of the incus remains shallow; at least such is the case in the

College series. The stapes approaches the quadrilateral form of that of Ovis, the head

being larger and the crura more nearly parallel than in Gazella.

In Cephalophus (fig. 28) the articular surface is almost as wide on the malleus as on

that ossicle in Bos, the processus muscularis is very short, and the manubrium is of the

long, straight, and slender form frequent in Antelopes yet to be noticed. The incus

(fig. 39) in C. maawellii is very like a Sheep’s; the stapes is triangular, though the crura

are very divergent. The malleus of Nesotragus moschatus is of the Cephalophus type ;

in the incus from an adult Tetraceros quadricornis (fig. 37) the body is as shallow as ina

lamb; the malleus (fig. 25) and stapes resemble those of Cephalophus ; the same applies

to the malleus of Nemorhedus goral (fig. 30), where, however, the processus muscularis is

longer than in C. mergens or C. maxwellii; the incus has a shallow body and very long

divergent crura. In the adult Kobus ellipsiprymnus (fig. 24) the articular surface of the

malleus is quite as wide as in Bos, but the manubrium is well recurved and not very

slender, altogether more like that of Gazelia than of the other Antelopes. The incus

has a very high body, quite as high as in the Ox; thecrura are rather short. This ossicle

ON THE MAMMALIAN OSSICULA AUDITUS. 429

exceeds the incus of the Pigs in height as much as the incus of some other Antelopes

exceeds that of Sws in breadth.

In the Harnessed Antelope (Tragelaphus scriptus) the malleus has a wide articular

surface and a very long, slender manubrium, which is almost straight. In Damalis luna-

tus, D. pygarga, and Alcelaphus bubalis (Pl. LX1. fig. 19), the articular surface is always

wide, asin Bos, but the manubrium is rather broad at the base and more curved than in

Tragelaphus. In all these genera the incus has a very shallow body, with very large,

thick, and long crura, somewhat as in the Cervide. The only stapes in the College col-

lection (from .4. bubalis) is triangular.

Catoblepas gnw and OC. gorgon are remarkable for the great length and slenderness of the

manubrium mallei, which is slightly turned up at the tip (fig. 26). The head is very

small, the articular surface is wide and smooth, and the processus muscularis is mode-

rately developed. The stapes of C. gorgon is triangular, of C. gnu more oblong from the

crura being more parallel. In both the anterior crus is the thinner.

In the Musk Ox (Ovibos moschatus) the malleus (Pl. LXI. fig. 31) resembles that of

Bos, but the manubrium is straighter, the body of the incus (fig. 40) is not so high, and

the processus longus is as sharply curved as in the Sheep. As the crura of the stapes

diverge a little, that ossicle takes the form of an isosceles triangle with a narrow base.

The left ossicula from an adult Anoa (figs. 32, 41) all resemble those of Bos in the

broad articular surface and well-curved manubrium of the malleus, the high body and

long erura of the incus, and the slender anterior crus of the stapes, which bone, how-

ever, is triangular from the divergence of the crura. The head bears a large tubercle for

the stapedius.

In Camelopardalis giraffa the auditory ossicles (fig. 21) are remarkably like those of

the Ox, and undergo similar changes in the growth of the young animal.

The articular surface of the malleus in an adult and a young male is still wider and

more plane than the Ox; the manubrium is strongly curved. The incus is all but iden-

tical in form with the Ox’s; the body is very well-developed; the stapes is almost rhom-

boidal; the anterior crus is much longer than the posterior, and more slender (fig. 21).

In the ossicula of a new-born or very young Giraffe, the articular surface of the

malleus is deeper, and not so wide vertically nor so plane as in the adult, the manu-

brium is much straighter, the incus has a far shallower body, and the stapes is almost

triangular. All these characters remind the observer of the changes which occur in the

development of Bos.

The ossicula of the Deer are very small and light, a set from an adult Cervus (Pl. LXI.

fig. 20) of large size being all smaller than a series from Gazella dorcas. The foetal

type of Bos prevails more strongly and persistently among adult Deer than even in

Ovis. The head of the malleus in the Red-deer is very little developed : and the articular

surface is deep, very saddle-shaped, and narrow vertically; the manubrium is long and

slender, only slightly bowed and recurved at the tip; the processus muscularis is a short

stout tubercle; the lamina is very broad. The incus has a very shallow, ill-developed

body, with a long, stout, diverging posterior crus, and a short processus “ longus,” sharply

SECOND SERIES.—ZOOLOGY, VOL. I. 3M

430 MR. A. H. G. DORAN ON THE MORPHOLOGY

curved, and bearing a distinct, pedunculated, elliptical Sylvian apophysis. The stapes

is quite triangular, with a small head and slender crura, which are almost equal and not

very divergent. There is generally a trace of a tubercle on the head of the stapes for

- the stapedius tendon.

There is no distinctive feature between any of the ossicula in Cervus elaphus, C. aais,

C. mexicanus, O. pudu, Dama vulgaris or Capreolus caprea. In Cervulus muntjac the

lamina of the malleus is very wide, and, contrary to Hyrtl’s assertion, the stapes (in the

College collection) is by no means thoroughly triangular ; nor can any trace of the pear-

shaped tuberosity described by that author be seen in the specimens examined by me.

In Hydropotes inermis the ossicula are identical in form with those of Cervus; the

same applies to two sets of the ear-bones of Moschus moschiferus (figs. 33, 42) from two

different skulls in the College collection. The shallow-bodied incus, with its long, stout,

divergent processus brevis, is strikingly distinct from the square-bodied, short-limbed

incus of Tragulus (Pl. LXI. fig. 8).

THE OssICULA OF THE PROBOSCIDEA.

The malleus of Elephas (Pl. LX. figs. 38, 39) is the largest seen in any terrestrial

animal, measuring nearly an inch in length from the head to the tip of the manubrium.

The portion of the head above the articular surface is not much developed, either

superiorly or anteriorly ; it is smooth and globular, and widest from behind forwards.

The articular area is shallow and wide vertically; its facets, which are only slightly

convex, run into each other even more insensibly than in Man, not a trace of a groove

dividing them. The neck is of a very peculiar form. It is very broad and flattened

laterally, and bent strongly outwards on itself. The segment above the bend is formed

of rather spongy bone, with a very irregular surface, generally marked with more than

- one nutrient foramen; posteriorly is a sharp ridge of the same sigmoid form as that seen

in Homo; and anteriorly a thin splinter-like processus gracilis is given off without a

lamina. The lower half of the neck is of very compact bone, smooth on the surface;

it gives off the manubrium at a little more than a right angle. The handle is of the

same stoutness as the lower portion of the neck; it is rather short, broad at the base

which bears a prominent everted processus brevis, flattened laterally, and moderately

recurved at the tip, There is a distinct sharply bordered outer surface to the manu-

brium ; but it is narrow, and very little dilated towards the extremity.

This bone is the most distinctive by far of the three ossicula in the Elephant. The wide,

shallow articular surface is seen among the larger Ruminants; but the breadth and short-

ness of the manubrium distinguish it from that of any Ungulate. There is no lamina;

but the anterior part of the less dense upper segment of the neck probably represents it *,

especially as the sigmoid ridge behind that part represents the entire neck of a laminated

malleus. The manubrium recalls a form not rare in the Rodents; but in this great

animal there is no trace of any processus muscularis, the tensor tympani being

attached to a tubercular elevation on the angle formed by the bend of the neck. TI had

the advantage of dissecting the internal ear of the old female Indian Elephant that died

* See remarks on the malleus of Sus, Phacocherus, and Hippopotamus.

OF THE MAMMALIAN OSSICULA AUDITUS. 431

in the Zoological Gardens in the summer of 1875. The tendon of the tensor tympani

was on both sides attached to the elevation referred to above; yet asimilar tubercle may

be seen in other mammals where a processus muscularis also exists. The great size and

peculiar development of this ossicle seems to mask all features which might relate it to

any other Order in particular.

The malleus is as large in Z. africanus as in H. indicus; but in the former it is of

somewhat lighter make, and the neck is not so broad nor so abruptly bent, while the

manubrium is longer.

The incus of the Elephant has a well-developed body, much deeper posteriorly than in

front, where it is excavated for the articular surface. The crura are both short; the

processus brevis diverges but little from the body, and is rather slender for the bulk of

the whole bone; the processus longus is more divergent, and turns up very slightly at

the tip, which bears a small, round, flat and sessile Sylvian apophysis. This bone,

taking even its large development into consideration, cannot be referred to any type

prevailing among the Ungulates. An approach to it is seen in many Rodents, espe-

cially as regards the much greater divergence of the processus longus compared with

the posterior crus.

The stapes in Hlephas indicus (Pl. LX. fig. 36) is quite triangular; the head is

well developed, and bears a tubercle on its posterior aspect for the stapedius tendon; the

posterior crus is quite straight the anterior is slightly curved; the base is broad, and

convex towards the vestibule; it projects considerably in most specimens beyond the

insertion of the posterior crus.

Tn a single specimen of the stapes of E. africanus, in the College collection, the head is

broad, but shallow, and both crura are quite straight; the bone is triangular, but trun-

cated at the apex (fig. 37).

The peculiarities of the Ungulate ossicula may thus be summarized :—

I. In this Order the central* laminated type of malleus is the rule; the processus

muscularis is generally present, but is not often of any length; the incus is rather large,

and variable in form; the stapes has a broad head, and is sometimes quadrilateral.

II. In the Equide the malleus is most generalized; the head is well developed, and

the lamina almost obsolete. Inthe Rhinoceros and Tapir the malleus is of a very low

type, almost recalling that of the higher Marsupials. The stapes in all Perissodactyla is

triangular, with a wide head.

III. In the Tylopoda the malleus possesses characters seen both in the Artiodactyla

and Perissodactyla, the incus is of peculiar form, the stapes is triangular.

IV. Among the Artiodactyla the Suide are distinguished by the production forwards

of the head of the malleus and its deep articular surface, even in the larger species. The

incus has a very square body, with short crura; the stapes is quadrilateral. Hippo-

potamus resembles the Pigs, excepting that the stapes is perhaps more triangular.

V. In Tragulus the incus precisely resembles that of Sus, but the malleus and stapes

are like the same in the smaller Antelopes.

* My reasons for using this term when speaking of the laminated malleus will be found in the description of the

mallei of the Marsupials.

3M 2

432 MR. A. H. G. DORAN ON THE MORPHOLOGY

VI. In the Bovidee the head of the malleus is not produced forwards, as in the Pigs ;

the articular surface is wide and shallow in the adults of the large species. The crura

of the incus are always long; the body is solid, and very high in the Ox, shallower in the

Sheep: the stapes is quadrilateral in the larger members of this family. The feetal

ossicula of the great Bovidze much resemble the ear-bones of the full-grown smaller kinds.

The gradations of type between Ovis and Bos, and the intricate distinctions between

the ossicula of different genera of Antelopes, have already been discussed.

VII. The ossicula of the Giraffe are very similar to the same in the adult Ox.

VIII. In the Cervide the malleus is of the type seen in the Calf, or in the adults of

smaller Bovidee. The processus muscularis is stout, but short in the genus Cervus; the

incus has a very shallow body, with very large crura, even in Moschus, which may thus

' at once be distinguished from Zragulus. The stapes is always triangular.

Tur OssicULA OF THE HYRACES.

In Ayrax (Pl. LX. fig. 35) the upper part of the head of the malleus is a wide,

smooth, spheroidal surface; the articular area is extremely shallow, wide in both direc-

tions; and there is not a trace of any groove dividing the facets. The neck is narrow and

rather long; there is nolamina. The manubrium is well recurved at the tip, which is

slightly spatulate; close to its root a small tubercular processus muscularis is generally

perceptible. The outer aspect of the whole manubrium is sharply bordered and narrow ;

there is a rudimentary processus brevis.

The incus is remarkable for a peculiarity already noted by Hyrtl: the body is ex-

tremely small, whilst the crura are both long and divergent. That anatomist describes

those processes as thin; but in both Hyraa dorsalis and H. capensis they are stout

and well-developed.

The stapes is perfectly triangular, with a small head and very straight crura. The base

is thick and bullate towards the vestibule *.

The ossicula of the Hyraces recalls rather those of the Equide than their repre-

sentatives in any Artiodactyle or Rodent. The malleus is not unlike that of Hgwus,

where also the processes take considerably from the size of the body of the incus; the

triangular form of stapes prevails, too, among the Perissodactyla.

THE OSsiIcULA OF THE INSECTIVORA.

In the different families that compose this Order we meet with great variations in the

form of the ossicula, which are sometimes of almost as low a type as their representatives

in the Marsupials; in other instances they assume characters seen even in the higher

Primates. Tt is in the Insectivora that we so often find a bony canal between the crura

of the stapes, giving support to a small vessel, the “ Steigbiigelarterie” of Hyrtl. This

canal has already been described as existing in the genus Lemur and in some Rodents.

It reaches, however, the highest degree of development in the Order now under con-

* According to Prof. Owen, “ the base of the stapes is rarely ossified beyond the circumference.” In the speci-

mens in the College collection a bulla exists.

OF THE MAMMALIAN OSSICULA AUDITUS. 433

sideration ; Hyrtl devotes a long and interesting chapter (J.c.) to the vessels of the

mammalian tympanum, chiefly dwelling on the stapedial artery and its. branches.

According to that anatomist, this vessel seems, in Hrinaceus and other Insectivora, to

end ultimately in branches distributed to the orbit, whilst the homologue of the internal

carotid is given off before the stapedial artery enters the canal between the crura of the

stapes; but the distribution of these vessels, and the extent to which they are sur-

rounded by bony canals, differ in certain families. Still that is a subject. which does

not pertain to the distinctions of the auditory ossicles in the same animals.

The ossicula of the Shrews (Sorex) are best studied in the crania of some of the large

American species (Pl. LXITI. figs. 1, 2).

The malleus is readily detached from the skull by maceration, together with the

tympanic bone, which is almost identical in its ring-like undeveloped character to its

homologue in the Marsupials. By the aid of a fine needle the ossicle may be separated

from the tympanic without fracturing the processus gracilis.

The head is small, as in most laminated mallei. It bears a rather deeply cut articular

surface, with two facets, of which the uppermost is the larger, and the lower the more

convex. The neck is long and thin, and midway between the head and manubrium it is

bent on itself, forming a sharp angular projection. At the point where it joins the

manubrium is attached a prominent globular process of bone (Pl. LXII. figs. 1, 2, oa),

which projects downwards and backwards in the natural position, and seems, as it were,

to push aside the handle outwards and forwards. The manubrium itself is as thin and

slender as in most Marsupials. A distinct angle, corresponding to the processus brevis,

exists; it may be seen at the base of the manubrium, projecting anteriorly to the

globular process. The extremity of the handle is almost styliform, although, with the

aid of a strong lens, I have detected not only a slight spatulate dilatation, but also an

outer surface to the manubrium, eeincateliae narrow, yet sharply bordered from the very

thin and compressed sides *.

The lamina is as wide as in a typical terrestrial Carnivore, and runs forward with the

processus gracilis without receiving a thin process from the front of the head, such as is

seen to join the processus gracilis in most laminated mallei. That process is flattened,

narrow near the root, but wide in the middle, tapering to a point at the Seer: ; it is

closely applied to the tympanic bone.

A very small tubercular elevation on the inner side 6f the neck, close to where it joins

the manubrium, represents the processus muscularis. To study the relation between

this process and the orbicular projection from the neck, a malleus of a Sorex should be

placed side by side with one from a Gymmnura, where both exist well developed + (PI.

LXII. figs. 2 oa & 14).

The exact morphological signification of the orbicular process is a difficult point to

decide. It exists in many Insectivora andin many of the Muride. It clearly cannot be

* For a fuller explanation of this expression, “ sharply bordered from the sides,” see descriptions of the malleus in

the terrestrial Carnivora.

+ This elevation can only properly be seen by placing a very clean malleus of Sorex on a piece of black paper, and

viewing it under the microscope with a 1-inch object-glass,

434 MR. A. H. G. DORAN ON THE MORPHOLOGY

the processus brevis of Man, as the sharp angle at the base of the manubrium, which

represents it in the Shrew, may be plainly distinguished (with the aid of a lens), and

seen to be pushed outwards by this very orbicular apophysis under discussion. Nor

is it the processus muscularis; for such a process exists in Sorex. In the Bulau

(Gymnura rafflesii), as in the Shrew, a large, curved, and conspicuous processus

muscularis exists, as well as a large, well-formed orbicular apophysis. The tolerably

large malleus of this animal (fig. 14) is very favourable for examination to eluci-

date the question. I consider it highly probable that this orbicular process is the

homologue of the sharp projection seen in the neck of the malleus of the common

Badger, and already described in this paper. It is true that a sharp projection is

also to be found on the neck of the malleus of Sorex; but in that little animal the

projection lies midway between the head and the handle, whereas in Meles the sharp

projection is placed close to the manubrium, in the exact position where the orbicular

process is to be found both in Sores and in all other animals where it exists.

Hence it is not the homologue of any named portion of the malleus in any of the

Mammalia; nor is it represented, except as an orbicular process, in any member of that

class, putting aside the angular projection on the Badger’s malleus*. In all probability

embryology will be able to show that it represents some part better developed in the

homologue of the mammalian malleus among the Sauropsida and Ichthyopsida.

The incus of Sorex has a fairly developed body ; the processus brevis is very short and

non-divergent ; the stapedial crus is rather long, and diverges considerably from the

body; it supports a Sylvian apophysis, very minute (even proportionally), and mounted

ona slender pedicle. Altogether this ossicle is of a type very frequent among the Mar-

supials; yet it will be remembered that extreme shortness of the processus brevis

and marked divergence of the processus longus are seen in Mammals as high as Ursus

and Mustela.

The stapes of Sorex has a minute vessel running between its crura in the recent skull,

but unsupported by any bony canal; so that it drops out and is readily lost in maceration.

The head is ill-developed, broad, and shallow ; the crura are much curved, especially the

posterior ; and the aperture is very wide, as in Homo. The hase is narrow horizontally,

and very narrow vertically. The stapes of Sorex is evidently of a much higher type

than are the two outer ossicles.

The malleus of the Daesman (I/yogale moschata) must hold an important share of the

-anatomist’s attention when he undertakes the study of the orbicular apophysis, so

frequent in this Order (see Pl. LXII. figs. 3, 4).

The upper portion of the head is even less developed than in Sorex, projecting but

little above the upper edge of its articular surface, which is narrow in its horizontal, but

extremely wide in its vertical measurement. The incus in the recent skeleton lies, I

have observed, as much posteriorly as in the higher Primates. Both facets are of about

the same size; hence the lower is proportionally larger than in most animals; they are

nearly plane, and not divided by a sharp groove. The neck is short and curved. Ata

first glance the outline of the lower facet of the articular surface looks like a portion of

* And in the malleus of Bassaris and a few other animals.

OF THE MAMMALIAN OSSICULA AUDITUS. 435

the neck; and a sharp angle is seen where the latter commences, very like the projection

which actually exists in the middle of the neck of the malleus of Sorer. But, by

examining these ossicles in the Shrew and in Wyogale by means of a lens, it will

be seen that in the former the articular surface is not deep vertically, but the neck is

long and sharply angled, whilst in the Daesman the articular surface is long, and the

neck short. The lamina is wider than in Sorex; the processus gracilis is slender and

partially ankylosed to the tympanic bone in the adult. Close to the root of the manu-

brium there springs from the inner side of the neck a round-headed process (Pl. LXII.

fig. 3, ow) mounted on a long pedicel, to which I find that the tendon of the tensor

tympani is attached in the recent skull.

The manubrium is long, and forms a very obtuse angle with the neck; it is much

flattened laterally, and rather broad at the base, so as to be of the sabre-like outline

frequent among Rodents. The outer angle of the base is blunt and slightly bent towards

the neck ; the same aspect of the rest of the manubrium is sharply bordered from the

sides, and distinctly dilated at the extremity.

It is instructive to compare the malleus of Myogale with the corresponding ossicle in

Gymnura, which possesses an undoubted orbicular apophysis homologous to that of

Sorex, and an equally distinct processus muscularis. The latter process, moreover,

actually coexists with the former in many Shrews, or at least is represented by a

tubercle.

The attachment of the tensor tympani tendon to the single process in Myogale makes

that projection a functional analogue of the processus muscularis; but since it projects

backwards, as the orbicular apophysis does in Gymnura, where the muscular process

points inwards and forwards, it seems to me that this solitary process of the Daesman

must be considered as a homologue of the orbicular apophysis of the Shrew and Gymunura,

serving functionally as a muscular process, the true representative of the latter being

suppressed in this animal.

The incus much resembles that of Sorex; but the body is better developed, especially

posteriorly. The processus brevis is very short and non-divergent; the stapedial crus is

long, straight, and ends in a rather large discoidal Sylvian apophysis.

The stapes has a very wide aperture, occupied in the recent skeleton by a large bony

canal. The head is broad and shallow; there is a small tubercle for the insertion of the

stapedius tendon. Both crura are very slender and divergent ; the anterior is consider-

ably the more curved. The base is of necessity wide horizontally ; it is distinctly concave

towards the vestibule, without any bulla or umbo.

The ossicula of If. pyrenaica are very similar to those of the Russian Myogale. In

this animal and in the skulls of the latter and larger species I have always found a very

stout bony canal between the crura of the stapes. Hyrtl (op. cit.) observes :—

“In Sorex and Myogale the vessels run exactly as in Tulpa, without, however, being

enclosed within the tympanum in bony canals.” In describing the vessels of the tym-

panum in the Mole, he had previously remarked that “the bony canal does not fill the

three-cornered aperture of the stapes; on that account the stapes can be moved up and

down on the canal like a ring on the finger.” If this be “exactly” the case in spe-

436 _ MR. A. H. G. DORAN ON THE MORPHOLOGY

cimens of Myogale examined by Hyrtl, or if the stapedial intercrural artery was not, in

his examples, enclosed within the tympanum in a bony canal (whereas in the College

specimens that osseous tube nearly blocks up the whole aperture), it follows that there is

great variety among individual specimens in this respect, as I have actually BOSE eL in

the crania of Squirrels and Cavies.

In the Mole (Zalpa europea, Pl. LXII. fig. 5) the head of the malleus is rather large

for an ossicle of the laminated type; it forms a distinct globular prominence above the

level of the articular surface, also projecting forwards. The area for articulation with

the incus is wide laterally on account of the head being hardly flattened at all in that

direction ; but it is not long vertically; the facets form a sharp angle with one another,

and the uppermost is the larger. The neck is short and curved, the lamina wide

and rather opaque for so small a malleus; the processus gracilis is thin and very fragile.

here is no orbicular process; and the processus muscularis is reduced to a scarcely

perceptible tubercle.

The manubrium is short, and forms an angle of about 120° with the neck. The outer

angle of the base, corresponding to the processus brevis, is considerably bent forwards ;

the whole manubrium is rather broad, laterally, at the base, but narrows rapidly, so as

to be almost styliform at the extremity.

The incus of the Mole has an ill-developed body ; and the processus brevis is almost

obsolete. The processus longus is most characteristic, and has been long familiar to

comparative anatomists. It is deeply channelled out on its front and inner aspect, and

very long. It terminates in a proportionally small, pedunculated Sylvian apophysis.

The stapes is similarin form to that of Myogale; the crura are very slender and well

arched. ‘The intercrural canal is very small, and justifies Hyrtl’s remark quoted in the

above description of the Daesman’s stapes.

The extreme difference between all the ossicles of this animal and those of the Golden

Mole (Chrysochloris) is most instructive and significant; the great distinctions between

the malleus of Zalpa and that of Condylura,; where the incus is very similar, are also

interesting, but less to be expected.

The malleus of Scalops has a large head, as in the last genus; its lamina too is of the

same form. ‘There is a small tubercular processus muscularis close to the root of the

manubrium, which latter fragile process is very readily broken in examining the attach-

ment of the tensor muscle of the tympanum in the recent skull.

‘The incus (Pl. LXII. fig. 6) is more generalized than in either Talpa or Condylura, as

the processus brevis is not so rudimentary ; and though the stapedial crus is long, it is

not so remarkably broad as in those animals. As in them, however, it is deeply

channelled. P

The malleus of Condylura (Pl. LXII. figs. 7, 8) is doubly interesting :—first, because

it is so different from that of Zalpa; secondly, because it possesses, like Myogale, a pro-

cessus muscularis remarkably like the orbicalar apophysis seen in the Shrew.

The head of the malleus in the Star-nosed Mole is very small; nor is the articular

surface deep vertically ; its upper facet is wider than the lower. The neck is very long

and narrow, and is bent into a very sharply projecting angle midway between the head

OF THE MAMMALIAN OSSICULA . AUDITUS. 437

and manubrium ; the lamina is very wide and thin. Here it may be observed that in all

the above characters this ossicle resembles the same in Sorex. It has not either the

well-formed head of alpa or the vertically wide articular surface and short neck of

Myogale. The processus gracilis is broader at the root than in the Shrew; and in form,

being of equal breadth for the greater part of its length, it resembles the same in the

Phalangers and other Marsupials *.

The manubrium is rather short, it is broad at the base, with a very prominent ex-

ternal angle, not produced into a processus brevis. Its outer aspect, though extremely

narrow, is nevertheless separated from the much-flattened sides by distinct borders; the

extremity is very faintly dilated, so as to appear styliform+. The whole manubrium is

separated from the neck by a very deep constriction, so as to be readily broken off.

From this constriction, on the inner aspect, arises a very prominent globular process,

attached by a very narrow peduncle.

It is as difficult to decide whether in this animal the above is the analogue of the true

globular apophysis of the Shrew and Gymnura, or whether it is the processus muscu-

laris, as is the case in the malleus of Myogale. In Condylura the process certainly

points backwards, as in Myogale, like the globular apophysis of Gymnura; but (likewise

as in Myogale) the tendon of the tensor tympani is attached to the root of the process

in the Star-nosed Mole, which may hence be either considered the homologue of the

processus muscularis, or else as the homologue of the globular apophysis made use of, so

to speak, by the tensor-tympani tendon as a point of attachment.

The incus in Condylura is almost identical in character with that of Talpa in the

smallness of its body, the extremely slight development of its processus brevis, and the

large deeply channelled processus longus.

Chrysochloris possesses a malleus of a perfectly unique pattern (Pl. LXIT. figs. 9, 10).

Otto and Rudolphi first appear to have observed the extraordinary development of the

head of the malleus; but the former repeated the error of the latter, who took the

enormously enlarged head for a fourth ossicle. Hyrtl discovered their mistake, and

settled the true nature of the singular pear-shaped prominence from the malleus of the

Cape Golden Mole. The error of the first two anatomists above named must have arisen

from their having attempted to remove the ossicle by pulling at its manubrium with

forceps without enlarging the aperture of the meatus; in doing so the bone is readily

broken. If the malleus be extracted by chipping away some of the bone forming the

prominent convexity in the temporal fossa behind which the enlarged head is lodged,

then there is no fear of its breaking at the neck ; but it cannot be drawn safely through

the meatus. When removed entire, it may be examined separately with little fear of

damage, as it is by no means fragile.

The head of the malleus of Chrysochloris is extremely elaine so as to be over a

quarter of an inch in length. MHyrtl describes it as pyriform, although, being rather

thick and cylindrical at its root, convex behind, and slightly concave in front, it

* This process is united, partly by bone, to the tympanic ring in the adult Condylura.

+ [have never found the extremity of the manubrium to be absolutely styliform in any mammal except in its

modified homologue in the Dolphins.

SECOND SERIES.—ZOOLOGY, VOL. I. 3N

438 MR. A. H. G. DORAN ON THE MORPHOLOGY

may be better compared in shape to a grenadier’s bearskin. It is formed of dense and

brittle bone, and lies in a concavity which bulges into the temporal fossa. There can

be no doubt of its nature as a simple though extreme elongation of the ordinary

caput mallei. On its root posteriorly is seated, very obliquely, the equally modified

articular surface, which is long and narrow, slightly concave, without any indication

of division into facets, and prominently raised above the level of the contiguous parts

of the ossicle.

The neck is long, stout, and cylindrical; and a small depression on it extends towards

the root of the head, and probably represents the site of the lamina, the ridge of bone

below it being the root of the processus gracilis. I have found the ossicle slightly

adherent to the tympanic bone at that ridge, and have reason to believe that part of the

processus gracilis is modified and enlarged, and assists in forming a portion of the pro-

cess chiefly developed from the head. The manubrium (mm) is very short, being hardly

the length of the neck; it bears a very prominent processus brevis, and is scarcely

dilated at the extremity. Hyrtl does not appear to recognize either neck or processus

brevis in the Golden Mole.

T have detected a small tubercular processus muscularis in the neck of the manubrium

of C. trevelyani, where the head does not form so great a prominence in the temporal

fossa.

The incus of Chrysochloris is as modified in form as is its malleus. The body is of the

long and narrow shape of the articular surface of the outer ossicle, and is much con-

stricted before giving off its crura; its long axis intersects a line drawn from the tip of

the long to that of the short process, instead of being parallel to the latter as in other

mammals. Both crura are long, and stout, and rounded; the stapedial crus ends in a

very minute Sylvian apophysis.

The stapes has a small head, and short, very divergent crura; a stout bony canal runs

between them in the recent skeleton. The base is long horizontally, and very broad

vertically in the middle; it is convex towards the vestibule without being bullate. In

many respects this stapes resembles that of Spalaz.

In the Hedgehog (Hrinaceus europeus, Pl. LXII. fig. 13) and its allies the malleus

assumes a very characteristic form. The head of that ossicle is very small, and bears

a perfectly saddle-shaped articular surface, the two convex facets insensibly merging

into each other, without any groove to separate them. The neck is extremely long, and

forms a wide arch without any angular prominence as in Sorex. From its inner side, a

perceptible distance from the root of the manubrium, springs a straight, narrow, and

prominent processus muscularis; and nearer to the handle there projects from the neck a

considerable prominence, not so large as in the Shrew, or even as in Gymnuwra, but

clearly the homologue of the orbicular apophysis in those and in other Insectivora.

The manubrium is slender, and but slightly dilated, and recurved at the tip; the angle

corresponding to the processus brevis is very blunt, and pushed outwards and forwards

by the rudimentary orbicular apophysis.

The lamina is wider than in any other animal; the processus gracilis is reinforced

by a distinct process from the front of the head; and they together form an extremely

OF THE MAMMALIAN OSSICULA AUDITUS. 439

broad process, ankylosed to the tympanic ring in the adult. This process is broadest

towards its extremity, which is well rounded; it is much flattened laterally; and as

it stands at right angles to the plane of the lamina, it presents a very characteristic

appearance. Close to its root, at about the point of junction of the processus gracilis

with the process from the head, it is perforated for the passage of the chorda tympani

nerve.

The malleus of #. dealbatus is very similar to that of the common British species.

The prolongation of its processus gracilis is even wider and longer; but the lamina is not

quite as broad.

_ That the rod-like projection from the inner side of the neck of the Hedgehog’s mal-

leus is the true processus muscularis there can be no doubt, as the tendon of the

tensor tympani muscle will be found attached to it m the recent skull; nor, from the

nature and position of the prominence nearer to the malleus, can it very well be other-

wise than the homologue of the orbicular apophysis of Sorex, which is not utilized in

this animal for the attachment of the muscle, a distinct processus muscularis existing

as well. It seems extraordinary that the malleus of the common Hedgehog, broken in

half, should ever have been taken for two ossicles; yet Hyrtl has corrected that erro-

neous opinion enunciated by a previous observer. Such a mistake could scarcely have

been made if the homologies of the fragments of that or any other ossicle had been care-

fully made out.

The incus of the Hedgehog lies almost vertically above the malleus in the recent

skeleton. Its body is wide and shallow; the processus brevis is very short, but stout,

and pointed. The stapedial crus is long, stout, rounded, and very divergent from the

body; near its root, to the inner side, it bears a small tubercle, and at its extremity a

well-developed Sylvian apophysis on a narrow pedicle. In their general characters the

ineudes of the Hedgehog and of most of the musteline and arctoid Carnivora agree; but —

the remaining ossicles are totally dissimilar.

The stapes of the Hedgehog has a very: small head, with a small tubercle for the

stapedius-tendon on its posterior aspect. Both crura are very slender, grooved towards

the aperture, and much curved, so as to be furthest apart midway between the head and

the base. The posterior is often the straightest. A large vessel passes through the wide

aperture in the recent skull, not enclosed in a bony canai. The base is narrow and

almost plane towards the vestibule.

In the Bulau (Gymnura Rafflesii, Pl. UXII. fig. 14) the head of the malleus is better

developed than in Hrinaceus, but still small; the articular surface is of similar character .

The neck is much shorter and more sharply curved. From its inner aspect, near the

handle, there projects a very conspicuous slightly curved processus muscularis ; and

closer behind the manubrium is a prominent, but sessile, convex process, eviden tly the

homologue of the orbicular process in the Shrew. The presence of both these projections,

equally well developed in this animal, is of great value in considering, by comparison,

the homologues of the single process in Myogale and in other Insectivora. The manu-

brium is much laterally compressed, but not very broad at the root; the site of the pro-

cessus brevis is rounded off, and not prominent. The lamina is much narrower than in

3N 2

440 MR. A. H. G. DORAN ON THE MORPHOLOGY

the Hedgehog ; and the processus gracilis is produced into a much more slender appen-

dage, which bears, however, a foramen for the passage of the chorda tympani.

In the specimen in the College series of ossicula, the incus of Gymnura is extremely

like that of Hrimaceus, but the processus brevis is longer.

The stapes in Gymnura has rather straighter crura and a broader base than in the

Hedgehog. I have not found any bony canal between the crura in the skulls examined

by me.

In the Tenrec (Centetes ecaudatus) the head of the malleus (Pl. LXII. fig. 11) re-

sembles that of the Hedgehog; but the neck is shorter even than in Gymnura, and is

bent on itself into a sharp angle, as in Sorex. It bears a short processus muscularis,

and a very large, convex, but sessile, orbicular apophysis. The manubrium is quite

hedgehog-like, the lamina still narrower than in Raffles’s Gymnura; the processus

gracilis is perforated near the root of its free part beyond the lamina by a foramen

for the chorda tympani nerve. This ossicle in the Tenrec decidedly shows affinities to

Sorex.

But the incus, otherwise like that of Hrinaceus, has a well-developed and divergent

processus brevis. This is not the case in the Shrews and Moles, though it is observed in

Ericulus and, to a certain extent, in Gymnura. In the Macroscelide and Tupaiidee this

process is more developed than in the Hedgehog; but the body is higher than in that

animal and in Centetes.

The stapes is of the Hrinaceus type; the crura are slender and bowed, and not sepa-

rated by any bony canal in the recent skeleton; the base is moderately broad.

In Hriculus nigrescens (Pl. LXII. fig. 12) the malleus is more like that of Centetes

than that of Hrinaceus, the neck being short and sharply curved, and the lamina narrow.

The orbicular apophysis is not well developed; and the processus muscularis is reduced

' to a tubercle. The processus gracilis is perforated by a very distinct foramen for the

chorda tympani.

The incus, as in Centetes, has a body and processus longus similar to that of the

common Hedgehog; but, as in the former animal, the processus brevis is rather long,

thin, and divergent.

The stapes has a very broad base compared with that of Hrinaceus ; the head and crura

are very Similar in character.

The malleus of Solenodon cubanum is much like that of Centetes; it has a prominent

orbicular apophysis and a much more developed processus muscularis than has Hriculus.

The free part of the processus gracilis is unfortunately broken off in the College

specimen.

In the Macroscelide the malleus assumes a characteristic form widely differing from

the types of Hrinaceus and Sores.

In the Elephant Shrew (Wacroscelides alexandri, Pl. UXII. fig. 15) the caput mallei

is ili-developed and much taken up by the articular surface, which is composed of two

very distinct facets separated by a sharp groove. The lower is nearly the size of the

upper and more anterior; both have a high vertical convexity running across them.

The neck is rather short and curved; it is extremely constricted near the manubrium ;

OF THE MAMMALIAN OSSICULA AUDITUS. 441

the constriction appears all the greater on account of the narrowness of the lamina.

Close to the narrowest part, on its inner aspect, the neck bears a distinct, curved pro-

cessus muscularis. I can detect no trace of any orbicular apophysis.

The manubrium is long and rather narrow at the base, which bears externally a trace

of a processus brevis. Its outer aspect is not only sharply bordered from the sides, but

remarkably and almost uniformly broad; the extremity is but slightly dilated and

recurved. ;

The lamina is extremely narrow, owing to the processus gracilis joining rapidly a

process from the head, and thus leaving little room for any area of lamellar bone. The

processus gracilis thus reinforced forms a thin straight bar of bone, which runs half

across the front of the tympanic cavity, to fit into the Glaserian fissure, quite different

from what prevails in the preceding families of the Insectivora, where that process runs

downwards at once, and is curved so as to fit accurately to the tympanic ring.

The incus lies almost vertically above the malleus in the recent skull; it has a

well-developed square body; the part above the processus longus overhangs that pro-

cess. The processus brevis is stout, blunt-pointed, and does not diverge from the body.

The stapedial crus is straight and freely divergent; it supports a very small Sylvian

apophysis.

The stapes has an ill-developed body, and long, slender, extremely straight crura. The

aperture is wide and triangular, though not as wide as in the Shrews and Hedgehogs.

A very stout bony canal fills it during lifetime. The base is rather broad, and convex

towards the vestibule.

The malleus of Rhynchocyon cirnet (Pl. LXII. fig. 16) resembles that of Macroscelides

in general characters, particularly in the narrow neck and singular processus gracilis ;

but the head is better-developed, and the processus muscularis is still stouter and more

curved. The incus is very like that of the Elephant-shrew ; its stapedial crus bears a

fairly developed Sylvian apophysis. The crura of the stapes are very slender.

Itis remarkable that in Petrodromus (fig. 19) (the ossicula of which I have been able

to examine through the kindness of Dr. Giinther) the malleus is much more Shrew-like

than in the Hlephant-shrew. The lamina is wider; and there is a trace of an’ orbicular

apophysis. The processus muscularis and the manubrium are of the same form as in

Macroscelides. As for the incus of Petrodromus, it is not only quite unlike that of Sorea

or Hrinaceus, but in its high and narrow body it diverges also from Muacroscelides and

Ehynchocyon, and tends towards Tupaia. The stapes has straight crura, with a bony

canal between them in the recent skeleton.

In Tupaia the malleus differs from that of any other insectivorous mammal (Pl. LXII.

fig.17). The head of the malleus is broad antero-posteriorly, but flattened laterally. It

supports a very shallow articular surface, saddle-shaped, and only indistinctly divided

into two facets. There is not a trace of any neck or lamina; the tensor-tympani tendon

is inserted into the root of the manubrium. ‘This latter process arises from the head

directly, and has no angle at the outer side of its base corresponding to the processus

brevis. It is not very broad at the base, and is very little dilated at the tip; its outer

aspect is distinctly bordered from the sides.

442 MR. A. H. G. DORAN ON THE MORPHOLOGY

In being neckless and devoid of lamina, the malleus of Twpaia much resembles that

of some of the lower Primates, especially IJidas or Hapale, or certain Lemurs, the

shallow articular facets being a Ceboid feature. It cannot be said to resemble the mallei

of the Sciuridze so closely; for though the latter have a similar head, and are neckless, the

manubrium is of a different type.

The incus has a high and narrow body; the processus brevis is rather long, thin, and

pointed, but hardly diverges from the body; the stapedial crus is scarcely longer than

the short process; it is curved and divergent, and bears a very small Sylvian apophysis.

In general characters this bone is very like the same in many Monkeys and Lemurs. The

author has never seen the long process so divergent as to form (with the body and

short process as the top-piece) a letter T out of the incus, as Hyrtl states of Tupaia

javanica.

The stapes has a very small head and extremely slender, straight crura; the base is

rathernarrow. ‘The aperture is filled by athick bony canal in the recent skeleton, whence

this frail ossicle can with difficulty be removed.

In the Colugo (Galeopithecus volans, P|. LXII. fig. 18) the head of the malleus is much

flattened antero-posteriorly, and is produced a little forwards and upwards above the

articular area. The surface for articulation with the incus is of the type of Homo and

the higher Primates, being narrow vertically and wide laterally ; the outer extremity is

placed much higher than the inner ; and the two facets are faintly indicated, the upper

lying rather internal to the lower; both have a vertical convex ridge internal to the

middle line of the whole surface. The neck is very short and-constricted; and a very

narrow lamina (which seems to me to be absorbed in the adult skull, as the processus

gracilis is in Man) connects it to the slender processus gracilis. It bears a small tuber-

cular processus muscularis on its inner aspect.

The manubrium forms little more than aright angle with the neck, and is long, broad,

flattened laterally, and recurved ; it possesses a distinct trace of a processus brevis. Its

outer aspect is distinctly bordered from the sides, but very narrow, and prominently

spatulate at the recurved extremity.

The affinities of the malleus of Galeopithecus are puzzling. At first, judging from its

articular surface, narrow neck, and large flattened manubrium, with a trace of a pro-

cessus brevis, it seems allied to the highest Primates. But, on the other hand, in the

Manis the articular surface is shaped much as in Homo; and in Galeopithecus the head is

flattened, and the narrow lamina and slender processus gracilis connected with it also

resemble the corresponding parts in the scaly Edentata; and the manubrium still more

nearly resembles that of the Manis than that of Man, having an outer aspect separated

by broad borders from the sides. It approaches the form of the malleus in Lemur more

than that of any other of the Lemuridz ; but its head is not so well developed, nor of the

form seen in any Lemur or Monkey, and its neck is more constricted and distinct than in

any of these animals except the Apes and highest Cercopithecide. It differs widely from

the malleus of the Zarsius, the Nycticebide, and Galagos ; but it is more like that of

Chiromys, where the head is better-developed, though the articular surface, neck, and

manubrium are very similar.

OF THE MAMMALIAN OSSICULA AUDITUS. 443

Tt differs, comparing it with the other Insectivora, very completely from the Shrews’

and Hedgehogs’ in the smallness of the lamina and general form, and as widely from

that of the Tupaiide in possessing a constricted neck and in other respects; but it cer-

tainly has points of resemblance to the malleus of Rhynchocyon and Macroscelides, the

main distinctions being the form of the articular surface and the character of the lamina

and processus gracilis, though the latter feature is not very different in this animal from

that of the Elephant-shrew, except that the processus gracilis is more slender.

The incus is very high and narrow, especially over the short crus; in this respect it

exceeds the Zupaia incus and approaches that of Mycetes and Galago, though it does

not project over the processus longus as in the latter. The processus brevis is very short,

and curved backwards. The stapedial crus is stout, not very long, and bears a well-

developed very elliptical Sylvian apophysis on a narrow pedicle ; it does not diverge from

the body so much as in Tupaia; but the incus altogether resembles that of the latter

animal more than that of any other insectivorous mammal.

The stapes of a young Galeopithecus has an ill-developed head, with slender bowed

erura, and a large aperture not occupied by a bony canal in the recent cranium. The

base is broad and very convex; it projects far beyond both crura. In a pair from an

adult skull in the collection the aperture is filled by a thin lamina.

In most respects both incus and stapes of this animal more resemble those of the

Macroscelidze and Tupaias than of the Shrews, Moles, and Hedgehogs. They are not

strongly allied to the same ossicles in any definite group of the Primates, though strik-

ingly like them in one or two characters above given. Hyrtl considers that the incus of

Galeopithecus bears a larger Sylvian apophysis than in any other animal; but I find it

larger in Myogale, Rhinoceros, Ovibos, and several other mammals.

The above researches on the ossicula of the Insectivora lead to the following con-

clusions :—

I. These bones all offer great variations in the different families of this Order ; and

there is no constant positive character to be foundin any of the three ossicula. The

stapes is never Sauropsidan or columelliform, as in Manis and the Marsupials; but the

very frequent wideness of its intercrural aperture is perhaps not so much a sign of high

type, as an accidental feature in relation to a bony canal passing through it, or, at

least, on account of a large vessel, unsupported by such a canal, running between

the crura *.

Il. Chrysochloris has the most specialized ossicula in this Order. The extraordinary

malleus appears as if rather modified from the highest or anthropoid type than from the

more central laminated form of the ossicle. The almost equally peculiar incus, in the

development of its body and short crus, must be considered as of rather high type than

otherwise ; and the stapes is quite the reverse of Sauropsidan in all its characters.

IIT. The ossicula of Galeopithecus are very generalized in character. The malleus

has several characters seen in the highest Quadrumana and in some Lemurs, most of

which, however, are reproduced in certain mammals as low as the Hdentata. The incus

* More will be said, however, on this topic in the description of the stapes in the insectivorous Bats.

AAA MR. A. H. G. DORAN ON THE MORPHOLOGY

is of a type hardly seen except in certain Cebide and Lemuride; the stapes has no par-

ticularly high affinities. In the malleus Galeopithecus is decidedly allied to the Macro-

scelidee among Insectivora, and to no other family in this Order; but in the incus it more

approaches Tupaia.

IV. In the Tupaiidee the malleus assumes to an extreme degree the neckless and non-

laminated type common in most Cebide and some Lemurs, as well as in Sciwrus; but the

manubrium is rather of the form prevailing in the above-named Primates than of that

seen in the Squirrels; and so in every respect is the incus. All the ossicula differ from

the varied forms in other families of the Insectivora, except that the incus somewhat

approaches in type that of Galeopithecus. Taken as a whole, the ear-bones of Tupaia are

higher in type than in any other family belonging to this Order.

V. The Macroscelide have a more specialized malleus than can be seen in any other

insectivore except Chrysochloris; the extremely constricted neck and the narrow lamina

and processus gracilis running straight forwards to the Glaserian fissure are highly

characteristic. The processus muscularis is well-developed. The head is better-deve-

loped in Rhynchocyon than in Macroscelides. In Petrodromus the malleus is far more

like that of Sorex than in the two former genera; but the incus is not in any sense

Shrew-like.

VI. In the Shrews, the Myogalidze, Talpidee, Centetidee, and Hedgehogs proper one

common feature exists—a malleus with a wide lamina and a processus gracilis united to

the tympanic ring after the fashion of the Marsupials—in short, a low type of malleus.

The processus brevis of the incus is always ill-developed except in the Centetidze and

Gymnura. The intercrural aperture of the stapes is always wide.

VII. In the Soricide the malleus has a moderately wide lamina, an orbicular apo-

physis, and a more or less distinct processus muscularis, to which, and not to the

spherical protuberance, the tendon of the tensor tympani is inserted. Asa whole the

malleus is very much like that of a Marsupial. The stapedial crus of the incus is not

very broad.

VIII. In the Myogalide the malleus differs in several respects from that of the Shrews

proper: the manubrium is of stouter make ; and an orbicular process exists, which pro-

bably is the homologue of that in the malleus of Sorez, but. certainly functionally

replaces the processus muscularis, absent in Myogale. The incus resembles that

of Sorex.

IX. In the Talpide the malleus of Condylura most resembles that of Sorex, but has an

orbicular process serving as a processus muscularis, as in Myogale; its incus resembles

that of Talpa. In Scalops the incus has not the chief distinguishing feature seen in

Talpa and Condylura; but its malleus is of the form seen in the common Mole, where

the head is better-developed than in the Shrews and Hedgehogs, and the lamina not so

broad. The incus of Talpa has a very broad and channelled processus longus.

X. The Erinaceide and Centetidee have a more specialized malleus than the Shrews,

Daesmans, and Moles. The great feature is the very broad processus gracilis perforated

to transmit the chorda tympani nerve; the processus muscularis and an ill-developed

orbicular process coexist ; the lamina is very broad. In Centetes and Hriculus the pro-

OF THE MAMMALIAN OSSICULA AUDITUS. 445

cessus brevis of the incus is well developed; that crus is smalier in Gymnura and

abortive in Hrinaceus.

XI. The orbicular apophysis of the Shrews, Daesmans, Myogale, and Hedgehogs allies

them to the Muridz, where also the malleus is laminated.

THE OssICULA OF THE CHIROPTERA.

The malleus of Péeropus (Pl. LXII. figs. 26, 27) is attached to the ring-like tympanic

bone much in the same manner as among the Insectivora and Marsupials. The anterior

or upper extremity of that bone is fissured, and the processus gracilis fits into the cleft,

beyond which is a flat surface, to which the extremity of the process is ankylosed. The

portion of the tympanic external to the cleft is very thick, and it runs over the outer

surface of the lamina and neck of the malleus, a distinctive feature not observed in the

non-placental mammals or in the Insectivora, the tympanic ring reaching, in those

animals, not further backwards than to the root of the processus gracilis.

The head of the malleus is small, and projects very little beyond the articular surface,

which is placed as much posteriorly as superiorly, and is shallow, but deep vertically. It

is distinctly divided into two facets, although they are not separated by a sharp hori-

zontal groove; the upper is larger, and lies a little more internally than the lower, and

both are faintly convex. The neck is narrow and well curved, as in most laminated

mallei; the lamina is rather narrow, and formed of opaque and brittle, not semitrans-

parent, papery bone: the processus gracilis is broad and thin at its root, the lower

border being, however, well defined, and the lamina is prolonged upon it; this process

ends in a point as in most Insectivora, not in a foliaceous dilatation as in many

Marsupials.

T have found, in dissecting the ear of a fresh Bat (Pteropus), that the tendon of the

tensor tympani is inserted into the point of junction of the neck and manubrium, at the

most usual site, in fact, of the processus muscularis, which does not exist; but there is

a distinct prominence close to the manubrium at the place where a large orbicular

apophysis is seen in the Shrew (Sorex), Mouse (Mus), &c. The relations of that process

to the processus muscularis have been already fully discussed in describing the ossicula

of the Shrew and other Insectivora. Suffice it to say here, that in Péeropus the rudiment

of the orbicular apophysis does not replace functionally the processus muscularis as in

Myogale and Condylura. ‘This rudiment is better marked in the College specimen from

P. hypomelanus than in that from P. edulis in the same collection.

The manubrium is slender and rather long, broad at the base and flattened laterally ;

in fact, sword-shaped. Its outer and upper angle (not to be confounded with the rudi-

mentary orbicular apophysis which pushes it aside) is somewhat blunt, so that no pro-

cessus brevis exists; the outer aspect is very narrow, but bordered sharply from the

sides, and ends in a distinct spatulate dilatation.

It is evident that a verdict of “low type” must be passed on the malleus of

Pteropus. It has no anthropoid, ceboid, or lemuroid characters. It is not the typical

laminated malleus of the Carnivora and the Ruminants. It is decidedly lower than the

SECOND SERIES.—ZOOLOGY, VOL. I. 30 .

446 MR. A. H. G. DORAN ON THE MORPHOLOGY

malleus of the Muride, where the lamina is broad and semitransparent as in the

Carnivora. Its general resemblance to the malleus of the Marsupialia is very evident.

The incus of Pteropus has a well-formed body, not deeply excavated. The processus

brevis is distinct, but very short; the stapedial crus is rather long, and ends in a pedun-

culated almost discoid Sylvian apophysis. In relation to other animals its characters

are negative.

The stapes has a small head with slender crura; the posterior quite straight, the

anterior generally a little curved *. They do not diverge so much as in Man; but there

isa free aperture not occupied in the recent condition by any intercrural canal. The

base is rather narrow horizontally, and almost plane towards the vestibule; it projects

but little beyond the insertion of each crus.

The stapes of Pteropus must be considered of rather central type, for it shows no sign

of any tendency towards the columelliform stapes of most Marsupials; but its crura do

not diverge as in the anthropoid Apes and Man, nor even to the extent seen in Ruminants

and Carnivora. It most resembles the stapes of those Insectivora where the crura are

not widely bowed to admit the passage of a bony canal, or the same ossicle in the Mice.

This variety of stapes, with its straight crura, diverging but little, and its consequently

narrow base and narrow but distinct aperture, lies between the carnivorous type, where

the aperture is larger, but the crura as straight, and the form seen in Didelphys, where

the crura are free throughout, yet leave a very narrow aperture between them.

I have experienced great obstacles in the study of the ossicula of the insectivorous

Bats, as in museum specimens the bones of the auditory region are frequently lost in

macerating the skull; still more generally are the incudes and stapedes wanting, and

the delicate malleus is frequently mutilated. Nevertheless I have succeeded in procuring

specimens of at least the latter ossicle from Bats belonging to all the chief divisions of the

group, nor, judging from duplicates, does there seem to be great variety among them.

In studying the malleus of Asellia tridens, as an example of the RHINOLOPHIDA, a

very strong general resemblance to the same bone in the Shrew (Sorex) may be observed

ataglance. It is absolutely necessary, in studying the ossicula of the smaller Chiroptera,

to place thoroughly clean white specimens upon a piece of black paper; they may then

be viewed by direct illumination under the microscope. A 1-inch object-glass is suffi-

ciently strong for this purpose.

The head of the malleus in Asellia tridens + (Pl. LXII. fig. 20), though small com-

pared with the area of the entire ossicle, as in all laminated mallei, is well formed

and convex. ‘The articular surface is rather deeper cut than in Pferopus; but the facets

are of similar character, the upper and anterior being the larger. The neck is narrow,

longer even than in Sorex, and resembles its homologue in the Shrew in being sharply

bent midway between the head and the manubrium. At the apex of this angle I

find, projecting internally, a curved process (pm), remarkably like a second process

(acpm).

* Hyrtl (op. cit.) states that the crura “in Pieropus edulis and P. vulgaris [sic] are convergent before their insertion

into the base.” This is not the case in the College specimens.

¢ In Mr. Dobson’s ‘ Monograph of the Asiatic Chiroptera,’ Asellia is reduced to a subgenus of Phyllorhina.

OF THE MAMMALIAN OSSICULA AUDITUS. 447

At first sight the anatomist may be inclined to consider that pm (fig. 20) is accessory,

and acpm the homologue of the processus muscularis; for acpm lies in the usual

position occupied by that process in laminated mallei; and though the muscular process

is often found lower down, as in the Cavies, where it is seated on the manubrium, it is,

on the other hand, rarely seen higher up than acpm. But whilst I have never

found both processes coexisting in the malleus of any other Bat*, the upper of the two

observed in Asellia (pm, fig. 20) is constantly observed in the remaining insectivorous

Chiroptera (Pl. LXIJI. figs. 22, 24, 25, pm). As, then, the lower process, acpm, in

Asellia, is the peculiarity of this (and probably of a few allied Bats), and the upper is the

most frequent in others, it is reasonable to consider the latter, pm, as the homologue of

the processus muscularis, so common in other orders, the lower acpm being an acces-

sory process. From the base of acpm a sort of splint or buttress runs for a short

distance across the lamina; it must not be mistaken for the thin processus gracilis below

i, though it is most probably ossified as a part of that element.

Besides the structures just described, a large orbicular apophysis exists, identical in

position to the same in Sorex and Mus, and as large in proportion to the entire bone as

in those animals, but less conspicuous, because it is sessile, not pedunculated. It is far

better developed than in Pteropus.

The processus gracilis is slender, and appears as the thickened free border of the

lamina; it is joined by a process from the head, the homologue of what has been

already very frequently described in this memoir; the two form a rather broad curved

process, which fits into a cleft in the ring-like tympanic bone. The lamina is very

broad, and formed entirely of semitransparent bone; a buttress from the lower processus

muscularis runs across it, but is lost before the processus gracilis joins the process from

the head.

The manubrium is slender and long, well compressed laterally, and narrow, in that

sense, even at the root. It is not triangular, as is the case in most laminated mallei, but

truly quadrilateral ; for the inner aspect as well as the outer forms a narrow surface

sharply bordered from the sides almost to the point. A tendency to this condition may

be observed in the malleus of the Shrew and in some Carnivora, but never to so complete

an extent ; for in Asellia this inner surface is as broad as the outer. In the great majority

of animals the inner aspect of the manubrium is a simple border, sharp or blunt as the

case may be. The outer border or, rather, surface is very narrow, but ends below ina

distinct spatulate dilatationt. Above it forms a rather sharp angle, slightly incurved

towards the neck, in fact a rudimentary processus brevis.

In the well-formed lamina and orbicular and muscular process the malleus of Asellia

tridens resembles that of certain Insectivora and Rodents, and differs from the less

specialized type observed in Péeropus. It must be borne in mind that the characters of

the malleus in Asellia and other insectivorous Bats almost coincide with the distinguish-

ing features of the same ossicle in two tribes of mammals belonging to different Orders, but

* Excepting Phyllorhina fulva and P. speoris, where the malleus in all other points precisely resembles that of

Asellia tridens. + This dilatation is shown in fig. 21, the malleus of Plecotus.

30 2

448 MR. A. H. G. DORAN ON THE MORPHOLOGY

alike remarkable for the small size of the species, namely the Muridz and Soricide.

There may be some functional explanation of this similitude.

Among the VESPERTILIONIDA, the malleus of Plecotus awritus (Pl. LXITI. fig. 21) closely

resembles that of Asellia ; but the lower process (fig. 20, ac pm) is absent. The extremity

of the manubrium is rather widely spatulate.

The stapes has long crura, both almost straight; unlike most Bats, they diverge but

little; the aperture is on that account long and narrow; itis not occupied, in the recent

skull, by any bony canal.

In Vesperugo serotinus (Pl. LXII. fig. 25) the malleus also much resembles that of

Asellia; the upper or true processus muscularis alone exists, so that we miss the lower,

which in Asellia sends a conspicuous buttress from its base across the lamina. The

orbicular apophysis is smaller and more pointed in Vesperugo.

The incus of Vesperugo has a well-formed body, deeply excavated by the articular

surface; the processus brevis is short, but stout; the processus longus is very long and

divergent; it is slightly grooved on its upper and anterior aspect, and ends in an almost

discoid Sylvian apophysis, mounted on a very slender peduncle. In general characters

this incus is of a low type; the same form is frequent among Marsupials. In all the

insectivorous Bats the body of the incus is deeper and better developed over the pro-

cessus longus than it is over the shorter crus. The minute stapes of Vesperugo has a

small broad head, with a tubercle on its posterior aspect. The crura are slender and

very divergent; the posterior is quite straight, the anterior a littlecurved. The aperture

is very wide; the base has a rather thick border, so as to be slightly concave towards the

vestibule. It is much narrower anteriorly than behind—a well-marked feature here,

seen to a less extent in the stapes of Homo, and still less distinctly in most mammals.

The completely bicrurate stapes of these small Chiroptera, with a moderate or wide

aperture, is interesting when we consider the low type of the other two ossicula; but the

wideness of the aperture may not so much indicate high type, but may be considered as

merely due to the passage of an artery * between the crura; the same remark, as I have

already observed, applies to many Insectivora where the stapes has a wide aperture.

Whether a vessel passes through the opening free or unsupported by a bony canal is of

small moment ; still some may consider the wideness of the aperture as a sign of high

type, since in Lemur, where a bony canal exists, the crura do not diverge widely ; so that

in the Bats and Shrews the wideness may not be entirely for the benefit of the canal or

artery, especially as these structures do not always completely block up the aperture.

In many Rodents I have found the blocking complete without the crura being much

bowed or very divergent, as in some skulls of Oapromys.

The malleus of Vespertilio nattereri (Pl. LXII. fig. 22) is very similar to that of Ves-

perugo. The processus muscularis, too, in this Bat, seems in position to be the homo-

logue of the upper process in Asellia, especially as, in the place of the lower distinct

process in the latter, I observe in Vespertilio a faint eminence which sends a pro-

* Seo Hyrtl, op. cit. pl. ii. fig. 13. This artery is given off from the carotid together with the occipital.

OF THE MAMMALIAN OSSICULA AUDITUS. 449

longation across the lamina. The processus longus of the incus is very long. The stapes

is identical in form with that of Vesperugo*.

Among the EMBALLonuripz WMolossus obscurus shows in its malleus (Pl. LXII.

fig. 24) all the typical appearances above described in Asellia; but the processus mus-

cularis in Yolossus, as in most Bats, represents the upper process described as existing

in Asellia. The orbicular apophysis is not so large as in the last-mentioned genus. The

quadrilateral condition of the manubrium is very wel! marked.

Among the Puyiuostomip# I fail to find any distinctive characters. In the malleus

of Chilonycteris osberni the processus muscularis is very stout and long, and I have

preserved the tensor tympani muscle actually inserted into it in a specimen in the

College collection; but the process is the same in position as that in Molossus, hence

corresponding to the upper process in Asellia. The reader is referred to the description

of the malleus in the latter genus for arguments on the homologies of the two processes.

The mere fact of the upper process being found the most constantly throughout the

Chiroptera is, as already remarked, in favour of its being the true processus muscularis:

the lower being so very rarely found.

The incus of Chilonycteris has an extremely long processus longus, but the processus

brevis is much less developed than in the Bats already described; this in itself is

decidedly a sign of low type.

In Mormops blainvillii the malleus has a similar character to that of Chilonycteris; the

processus muscularis is in the same position (Pl. LXII. fig. 28).

I regret I have been unable to procure ossicula from the crania of Bats belonging to

other genera. Sufficient material, however, has been examined to warrant certain definite

conclusions on the characters of the ear-bones of the Chiroptera.

I. In all the Chiroptera the ossicula much resemble those of the Soricide and allied

Insectivora, particularly in the malleus.

Il. In the frugivorous Bats the malleus does not resemble that of Sorex so strongly

as does the same ossicle in the remaining section of this Order; indeed it approaches the

marsupial type im general characters, but bears a strongly marked rudiment of the

orbicular apophysis seen in the Shrew.

III. In all the insectivorous Bats the malleus bears a long, curved processus mus-

cularis, and in some genera (at least in Asellia) a process apparently accessory to it.

A well-formed orbicular apophysis is constant, the lamina is always wide, and the

manubrium is quadrilateral, the outer and inner aspects being sharply bordered off from

both the sides.

IV. The incus in all the Chiroptera has a very short processus brevis and a very long

divergent processus longus, as in: Sorex and the Marsupials. In this point the Shrews

and Bats differ from the Muridze, where the malleus is similar in character to the same

ossicle in the former animals.

V. The stapes is never columelliform, or even approaching that low type, but, on the

contrary, the aperture is generally wide; this is partly necessary, at least, for the

* Prof. Owen describes in Vespertilio noctula ‘‘a retention of the columelliform confluence of stapes and incus *

(Anat. of Verteb. vol. iti.).

450 MR. A. H. G. DORAN. ON. THE MORPHOLOGY

transmission of an artery unsupported by any bony canal, but may be considered as

well (for reasons given above at length) to indicate an approach to the Primates.

VI. Judging from their ossicles it may be said that the Chiroptera are very closely

allied to the Insectivora far more than to any other order of mammals.

THE OSSICULA OF THE CETACEA.

The highly specialized tympanic and petrous bones of the Whales and Dolphins are

now familiar, not only to the anatomist, who confines his studies to recent animals, but

also to the student of paleontology, on account of their great density and consequent

durability. The already known peculiarities of the soft parts in the auditory apparatus,

as they exist in this Order, may be conveniently studied in the fine preparations made by

Hunter, and now preserved in the Museum of the Royal College of Surgeons*. His

observations, quoted in the Catalogue +, are full and satisfactory as far as concerns the

common species described therein, but preparations of the tympanum and labyrinth of

rarer types remain unrepresented in that series. Further researches in the anatomy of

the cetacean tympanum and membranous labyrinth are much to be desired.

But, in accordance with the scope of this paper, the morphological characters of the

ossicula must alone occupy our attention. I will describe them relatively, as touching

the distinction of each ossicle in the different genera, and comparatively as regards the

homologies of the several parts of the highly modified cetacean ear-bones to correspond-

ing portions in the more familiar types of terrestrial or partially terrestrial mammalia.

In the College collection there is one fine specimen of the malleus of Balena mys-

ticetus (Pl. LXII. fig. 28), and another, precisely similar in character, attached to the

tympanic bone (No. 2488, Osteol. Ser., Pl. LXIII. fig. 13), probably from B. australis. At

the first glance, no part of this bone, as it is seen in other animals, can be distinguished

in these specimens, excepting the articular surface; after a little observation, the pro-

cessus gracilis will be recognized; and by further careful study of the specimen, the

observer will find that the modifications are not quite so great as they at first appear.

The peculiarities of this ossicle may be rendered more intelligible by first examining

it i situ, as in Pl. LXIII. fig. 18, which includes part of the (left) tympanic and petrous

bones in B. australis and is viewed from in front. The malleus is seen to be firmly

ankylosed to the tympanic (tb) by its processus gracilis (pg) and by certain appendages

(7., phm), presently to be explained. That ossicle is, in Balena, anterior to the membrana

tympani; and the processus gracilis runs, not forwards, but outwards, to a special pro-

jection of the margin of the tympanic bone to which it is attached. This projection

probahly represents the anterior extremity of the annulus tympanicus of other mammals,

the part in front being an extension of that ossification to surround the beginning of the

‘Eustachian canal. The petrous bone (pet) lies much above the tympanic; hence the

unusual position of the articular surface of the malleus, as Cuvier has observed. It

follows that the homologous parts of the malleus of Balena and the same bone in an

ordinary mammal will not be found in the same respective aspects of each bone.

® Nos. 1582-1598 8, Physiological Series. + Physiology, vol. iii. p. 115.

OF THE MAMMALIAN OSSICULA AUDITUS. 451

The malleus of B. mysticetus (Pl. LXII. fig. 28) measures about half an inch or a little

more in length. The head is very large; its intero-inferior aspect is smooth and quite

convex; externally and a little anteriorly it is very deeply excavated, the upper border

of the cavity extensively overhanging it, especially towards the front, whilst behind there

is a deep notch in that border. Hyrtl justly compares the excavation to that seen in the

anterior aspect of the malleus of the Seal (Phoca); and from the extremely overhanging

border, the bone assumes a form seen in certain shells, as Van Beneden has observed ;

indeed, putting aside the articular region and manubrium, the head much resembles the

tympanic bone to which it is attached. On its upper aspect is situated the articular

surface, which looks a little inwards and is deeply concave. It is made up of two

facets, both of a semicircular form; the upper and external is almost plane, and by far

the most extensive; it is separated from the lower, which is convex, by a sharp groove.

In front of the head, on the outer aspect, is a deep channel, distinct superiorly, but

absent below. This cuts off a portion of the malleus, which, close to the channel, is

broad and flattened, bearing a distinct tubercle (Pl. LXII. fig. 28, & Pl. LXIIL. fig. 18,

pm), which projects upwards and inwards; but beyond the tubercle it gradually nar-

rows as it runs forwards, ending in a hook-like point (mz), which curves downwards and .

outwards, and is longer and narrower in B. australis than in B. mysticetus. The

tubercle (pm) is the processus muscularis, as may be seen in preparation No. 1598 B in

the Physiol. Ser. Coll. Surg., where the tensor tympani muscle remains still attached to

it. The hook-like process (mz) is without doubt the true manubrium ; into it is inserted

the apex of the curious prolongation of the membrana tympani characteristic in this

Order. :

The homologous parts of the malleus in Balena (Pl. LXIII. fig. 13) may be com-

pared with the same in the more familiar malleus of Felés (Pl. LXIII. fig. 14).

Cuvier states, in his ‘ Lecons d’Anatomie Comparée,’ that in the Dolphin there is no

manubrium, and that the malleus of the Whale “resembles it in every respect, but is

double its size.’ Hyrtl (J. c.) correctly denies this assertion, and remarks of the Ceta-

cean manubrium :—“ It is only very short, and reduced to a rounded bony tubercle,

separated by a deep incision from the head. . . . The processus minor [sic] is represented

by two small pointed appendages, to which a triangular process of the membrana tym-

pani attaches itself.” But Van Beneden remarks* of Balena mysticetus :—“< The

malleus possesses neither any distinct apophyses nora manubrium.” Lastly, Huxley +

asserts, in describing the anatomy of the Porpoise:—‘“‘ The tympanic membrane is

concave externally; and, as is usual in the CuTacra, is connected by a ligament

with the handle of the malleus.” I cannot see how the curved process of the malleus

of Balena can represent any thing otherwise than the inflected splinter of bone on the

blunt extremity of the malleus of the Dolphin {, which is the true manubrium ofall other

mammals; and, judging both by analogy and by the preparation above referred to, the

* ¢ Ostéographie des Cétacés,’ Yan Beneden and Gervais.

+ ‘ Manual of the Anatomy of Vertebrated Animals,’ 1871.

+ Pl. LXIII. fig. 15, mn, from a Globiocephalus.

452 MR. A. H. G. DORAN ON THE MORPHOLOGY

tubercle can hardly be fairly denied to be the homologue of the processus muscularis.

Taking these assertions for granted, and recognizing henceforth in this treatise the

curved process as the handle of the malleus, it will be shown that the manubrium is far

better developed in Balena than in the Dolphins or Sperm-whales, and still more than

in the Ziphioid Physeteride.

From the malleus to the tympanic bone runs, almost directly outwards, a process over

half an inch in length. It is made up of two stout splints or columns, one forming its

posterior, the other its anterior border, and a broad flooring of thin bone connects

them by their inferior borders ; this latter is continuous with the floor of the excavation

in the head of the malleus, the posterior raised and everted margin of that cavity

running to join the posterior splint, the anterior coming distinctly from the corresponding

margin of the depression in the malleus of B. australis; in that of B. mysticetus the

anterior splint is given off in front of the groove, at least so I find it in the spe-

cimens I have examined; but this appears to be a trifling point, probably subject to

variety.

The whole of this process, highly characteristic of the Cetacea, has for years been

‘recognized by most authorities as the processus gracilis. One anatomist, Buchanan, who

has carefully studied the anatomy of the ear in this Order, and written a most lucid

description of the Whale’s membrana tympani, represents this bony connecting medium

in a Plate *, and there calls it “the long handle of the malleus ;”’ whilst the process mn

in Pl. LXIT. fig. 28, & Pl. LXIIT. fig. 18, at the end of this paper,’is represented in

Buchanan’s illustration as very much longer than I can find it in any specimen in the

College Museum, and styled “the short [sic], slender, or gracilis processus of the mal-

leus.” Here is a great confusion of terms, as other anatomists make the “long” and

“‘eracilis’? process synonymous; then, too, the expression “long handle” is very vague.

Hunter (Phil. Trans. 1787) merely says that the process from the membrana tympani is

inserted “into that bone” (@. e. the malleus), and speaks of the other process, that is the

bony attachment of the malleus, without naming the homologies of those parts of the

ossicle in question. Home (Phil. Trans. 1812) figures the ossicles very correctly, but calls

the manubrium the “ short handle,” and the processus gracilis the “long handle.”

But when we come to compare once more the process connecting the malleus to the

tympanic bone in Balena with the more typical malleus of Helis (Pl. LXIII. fig. 14), we

- shall find that, more strictly speaking, the anterior splint (fig. 13, pg) is the homologue

of the processus gracilis, whilst the posterior (phim) is the prolongation towards it from

the head of the malleus seen so frequently in terrestrial mammals; then the floor of

thin bone between them (2) represents the lamina (fig. 14, 7) of the Carnivora, Ruminants,

&c. The whole of the above connecting process between the malleus and tympanic bone

is rather longer in Balena, and perhaps in Balenoptera, than in the Dolphins and

Ziphioid Whales.

Tn its large excavated head and short curved manubrium, the malleus of Balena bears

a distant resemblance to the same in Macrorhinus and Stenorhynchus among the Seals.

* «Physiological Tlustrations of the Organ of Hearing.’ London, 1828.

OF THE MAMMALIAN OSSICULA AUDITUS. 453

The incus of B. mysticetus (Pl. LXII. fig. 29) and of B. australis, which exactly

resembles it, is more generalized than in any other cetacean (excepting, perhaps, Wega-

ptera and Neobalena). From the position of the articular surface of the malleus, the

incus lies above and internal to it, with the corresponding surface downwards. The body

is very solid and square. The posterior crus is short; stout, and recurved at its extremity.

The processus longus, not so oyer-developed as in other Cetacea, is still very stout and

broad to its extremity; it forms a bold sigmoid projection. A faint groove cuts off the

Sylvian apophysis from the rest of the processus longus; it does not expand or project in

any way from the crus itself. The articular surface on the body is very prominent; its

upper facet is almost plane and larger than the lower, as in the malleus; a sharp ste:

which separates them, fits into the groove between the facets of the malleus.

The stapes (Pl. LXII. fig. 29) is nearly half an inch long and very narrow. The head

is well developed, and bears a rough surface posteriorly for the insertion of the stapedius

tendon. The crura are long, straight, and slender; they diverge but little, and are con-

nected by a lamella, which has only a very small aperture in it; this lamella is evidently

an ossification of the thin membrane thrown across the crura in the recent stapes of a

typical mammal. ‘The base is generally immovably soldered to the fenestra ovalis *.

The stapes of Balena bears a certain resemblance to that of Stenorhynchus, and differs

from all other Cetacea in the length and slenderness of its crura. )

Tn describing the process which unites the malleus to the tympanic bone, the anterior

column or border of that process (Pl. LXIT. fig. 28, Pl. LXIII. fig. 18) was assumed to

represent the true processus gracilis, and the posterior (phm) as homologous to the

prolongation from the head of the malleus, which in other mammals generally joins it.

This posterior column runs as an elevated ridge two or more inches over the outer

surface of the scroll-like tympanic bone, ending in a broad lamina, which is closely

applied to the upper part of the deep oblique groove which separates the anterior from

the posterior portion of that bone, which would seem to indicate that the posterior

column was the processus gracilis. But the anterior, which seems, from its position

between the head and manubrium, to be the true processus gracilis, is soon lost on the

immer side of the tympanic bone, where it makes a sharp ridge in some whales (Phy-

seter &c.), as if the root of Meckel’s cartilage had become embedded in that bone during

ossification. Altogether, then, arguments are in favour of the anterior column being the

homologue of the processus gracilis. This can alone be verified by the examination of

very small foetal Cetacea.

In the Pike Whale (Balenoptera rostrata) the auditory ossicles assume much more

thoroughly the type most frequent in the Cetacea, excepting Balena.

Indeed the malleus (Pl. LXIT. fig. 30) is as extreme in its modifications as that of

Globiocephalus, Phocena, and Delphinus, being only exceeded in specialization by the

Ziphioids. It is less like the malleus of Balena than is the same ossicle in Physeter or

even Orca.

* In Balena mysticetus; but I have since found this not to be invariable, and in an adult petrous bone of

B. australis in the College Museum the stapes has fallen out.

SECOND SERIES.—ZOOLOGY, VOL. I. 3P

454 MR. A. H. G. DORAN ON THE MORPHOLOGY

The head is less developed than in Balena, especially below and internally *; it has a

deep depression externally, of which the borders are elevated, but not nearly so over-

hanging as in the Greenland Whale. The articular surface is very similar, and the

upper facet is the larger. In front of the head is a very prominent well-rounded tubercle,

on the under and inner surface of which is a sharp splinter of bone (mm), with its point

directed backwards; it lies flat against the malleus.

Hyrtl, in denying Cuvier’s assertion that the manubrium is absent in Dolphins and

Whales, continues :—‘“ It is only very short, and reduced to a rounded bony tubercle,

separated by a deep incision from the head.” But in describing the tympanum of Mo-

nodon he remarks :—‘‘ The fleshy process of the membrana tympani is half an inch long,

and attached to a ledge [ Leiste| on the outer border of the malleus.” Now the malleus

of Balenoptera is very similar to that of Monodon, and still more to that of Delphinus

and Phocena. By carefully dissecting the tympanum of a Monodon, and examining the

preparations of that cavity from all the remaining above-mentioned genera as seen in

the College Museum (Nos. 1590-3 and 1596), I find that the tensor tympani tendon is

almost invariably inserted into the most prominent and anterior part of the rounded

tubercle of the malleus, whilst the process from the membrana tympani + is inserted, as

Hyrtl says, into the splinter of bone described above; it is, however, on the under, not

the outer surface of the malleus. Hence it must be correct to consider the bony splinter,

Hyrtl’s “ Zeiste,” as the manubrium, whilst the bony tubercle } should be considered

the homologue of the processus muscularis of other mammals. The short curved process

in the Greenland Whale, well divergent from the body, is evidently an intermediate

condition between the long manubrium of terrestrial quadrupeds and the flattened

splinter of Balenoptera and the Dolphins.

The process joining the malleus to the tympanic bone is very similar to that in

Balena, but shorter. The anterior pillar, or true processus gracilis, is very stout, and

the lamina is rather denser than in Balena. i

The incus (Pl. LXII. fig. 31) assumes, like the malleus, the form prevailing in genera

presently to be referred to. The body (which bears an articular surface similar to that

of Balena) is very ill developed, and the posterior crus is almost atrophied ; it measures

about a line in length, though tolerably stout. The processus longus is enormously

developed, and incurved so abruptly as to present anteriorly a knee-like projection. On

its broad extremity it bears a smaller and narrower, thick and elliptical Sylvian apo-

physis, of about the size and shape of the base of a Squirrel’s stapes. Hence the incus

of Balenoptera differs from that of Balena more than the latter is different from that

of most terrestrial Mammalia.

The stapes (Pl. LXII. fig. 31) is not so long as in the Greenland Whale, though its

crura are longer than in the Dolphins. The head is well developed, and bears a rough

* Tt is much flattened on its outer and inner aspects.

+ See Pl. LXIII. fig. 16, where the tendon of the tensor tympani and the process from the membrana tympani

(in Monodon) are both indicated.

+ The exact site of the insertion of this tendon is marked by a rough surface in Balenoptera, Orca, and most

other Cetaceans.

OF THE MAMMALIAN OSSICULA AUDITUS. 455

tubercle for the insertion of the stapedius tendon. The crura are very stout, straight,

and more divergent than in Balena. The base was not soldered to the fenestra ovalis

in the stapes removed from a skull of an adult male in the College collection, but could

be displaced uninjured without any force being required to detach it; the same was the

ease with the stapes of an adult Balenoptera in the British Museum. ‘The base is

quite concave towards the vestibule. The aperture between the crura is small and

circular. :

The malleus of the small New-Zealand Fin-Whale (Balenoptera huttonii, Gray) in the

British Museum almost precisely resembles that of B. rostrata; the tubercle is propor-

tionately less developed, and the splint-like manubrium of the northern species is, in

B. huttonii, reduced to a mere rough surface, as in many Dolphins.

A stapes from the gigantic Steypireythr (Balenoptera sibbaldii), kindly presented to the

College by Professor Turner, has longer crura than that of B. rostrata, but is only half

as large again *.

The malleus of Megaptera nove-zelandi@ (Pl. LXII. fig. 32) has its head as flattened

as in the last described genus; but in front, instead of a short round tubercle, is a stout,

long, blunt-pointed process curved downwards. On the inner aspect of the latter is a

sharp spine (mn), directed backwards, and representing, as in Balenoptera, the manu-

brium. The blunt tip of the process (pm) has a rough surface, evidently for the insertion

of the tensor tympani; and posteriorly, close to the head, is a prominent tubercle, only

faintly indicated in Balena and Balenoptera, the analogies of which are not evident,

though it decidedly is not a muscular process in the last-named genera, as preparations

in spirit, already quoted, clearly show.

I have not had the fortune of examining the incus of a Megaptera, and cannot

find a detailed description of it in any work on the Ear or on Cetacea. Van Beneden

states that it is small, and that one of its articular surfaces is ill developed (in I. boops) ;

in the illustrations to his work, that ossicle is represented with a body almost as well

developed as in Balena, and the “short crus” is depicted rather stout, long, and pointed,

the “long” process not very thick.

The stapes of Megaptera nove-zelandia, and also that of MW. longimana, Gray, has

as long crura and as wide an aperture as in Balena; in a specimen of the former

it was found ankylosed to the petrous bone; in the latter species (judging from an adult

specimen in the British Museum) such ankylosis is at least not invariable.

In the greater length of the process bearing the homologue of the manubrium and the

insertion of the tensor tympani, in the form of the stapes and probably of the incus,

Megaptera tends more towards Balena than to Balenoptera; but, on the other hand,

the flatness of the caput mallei relates it very nearly to the latter; and the whole process

from the head is produced and shaped not so much in-the manner seen in Balena, but

rather as in Platanista.

The malleus of Neobalena marginata + (Pl. LXII. fig. 33) is highly interesting, since,

* See Appendix to this section.

+ Dr. Hector (Trans. N. Z. Inst. 1872) considers that this species, known to Dr. Gray under that name through .

oP 2

456 MR. A. H. G. DORAN ON THE MORPHOLOGY

like that of Megaptera, it exhibits intermediate characters between Balena and Baleno-

ptera. It is larger than the malleus of an adult Balena; its head is less flattened than

in Balenoptera. The upper facet of the articular surface is extremely wide and almost

plane; the lower is small, and as convex as in most Cetaceans; it must be remembered

that in the Greenland Whale and Southern Whale the former facet is rather larger than

in Balenoptera. From the front of the head a process projects, which bears on its inner

side a sharp spine (mn), directed more downwards and less compressed against the body

of the malleus than in the last-mentioned whale, but neither so long nor directed so much

downwards as the stout curved manubrium of Balena, which it represents (Pl. LXII.

fig. 28, Pl. LXIII. fig. 18, mn).

In the plates illustrating Camper’s ‘Observations Anatomiques sur la Structure et le

Squelette des Cétacés’ the ossicula of Physeter are very correctly figured. He speaks .

of “la manche, ou lapophyse de Rau”’ as if those terms were synonymous ; the site of

the insertion of the tensor tympani is not mentioned or indicated.

The malleus in the Cachalot and in the Ziphioid Whales possesses highly distinctive

characters.

This ossicle in Physeter macrocephalus (Pl. LXII. fig. 34) is very small in proportion

to the bulk of the animal. In general form it may be compared to a half-split pea,

which it hardly exceeds in size. The head is small and convex; it does not bulge or rise

high over the upper facet. The depression on its outer aspect exists, but is not so

extensive as in any of the Baleenoidea. The articular surface is very wide, considering

the ill-development of most of the elements of the bone. Its upper facet is very broad

and almost plane, the lower is much smaller, convex, and everted, like the mouth of a

pitcher.

The process connecting the malleus to the tympanic bone is extremely short, but the

same elements can be distinguished in it as in Balena.

But the most distinctive feature in the malleus of Physeter is the suppression of the

projecting tubercle which in other Cetacea bears the homologue of the manubrium and

the insertion of the tensor tympani. On the anterior aspect of the head is a sharp, pointed

tubercle (pm) looking directly upwards; directly below it is a very small blunt projec-

tion, pointing, in some specimens, a little forwards; in most examples it is hardly per-

ceptible. This (mz) is the representative of the manubrium, and is much better deve-

loped in some of the Ziphioid Whales shortly to be described. In specimen No. 1584,

Physiological Series, in the Museum of the Royal College of Surgeons, showing the

auditory apparatus of Hyperoodon, where the malleus is very similar in form to the same

ossicle in Physeter, the insertion of the membrana tympani to the tubercle (mn,

Pl. LXI1. fig. 35), and of the tensor tympani muscle into the upper process (pm), is well

displayed. Hence mn in figs. 34 & 35 is the representative of the manubrium in Phy-

seter and Hyperoodon, whilst pm is the homologue of the processus muscularis.

piece of whalebone only, is identical with the Caperea antipodurum, described in the B. M. catalogue from a solitary

tympanic bone. Gervais and Van Beneden consider Caperea antipodarum to be the same as their Balena antipo-

darum, which they distinguish from B. australis, and rightly, so far as the mallei are concerned.

OF THE MAMMALIAN OSSICULA AUDITUS. 457

The incus of Physeter has a much smaller body than that of Balena; but this

portion of the ossicle is, on the other hand, not so reduced in size as in Balenoptera and

the Dolphins. The articular surface is very wide; the upper facet is the larger and

planer, the lower is very concave, to fit the corresponding parts on the malleus. The

ridge dividing them is most distinct anteriorly. The processus brevis is slender, but

rather long; the processus longus is very stout, but not so big as in the Delphinoidea

and in Balenoptera; it bears at its extremity a small, sessile, elliptical Sylvian

apophysis.

The stapes has extremely stout crura, not so divergent as in the Dolphins ; the aperture

is sometimes obliterated. The footplate has a very thick border, and is a little con-

cave towards the vestibule. I have never found this ossicle ankylosed to the petrous

bone.

In the Ziphioid Whales the ossicula much resemble those of Physeter.

In Hyperoodon rostratus (Pl. LXII. fig. 35) the malleus has a larger head than in the

Cachalot ; that portion of the ossicle is also more convex, and rises a little above the

upper border of the articular surface.

There is hardly any trace of a depression on the outer aspect of the head; it seems

rather as if the bone above had grown into it, and reduced it to a horizontal slit. The

process holding the malleus to the petrous bone is extremely short, but its homologies

can be easily made out. The articular surface is, proportionately to the rest of the

ossicle, considerably smaller than in Physeter, nor is the upper so much larger than the

lower facet.

The processus muscularis (Pl. LXII. fig. 35, pm) is even less developed than in

Physeter; the manubrium (mz) is a still more insignificant tubercle.

The incus also in general characters resembles the same in the Cachalot, but possesses

a character which it shares with the two other Ziphioid Whales whose ossicula the author

has examined, and in which it differs from Physeter. This feature is the long, flattened,

and very divergent processus brevis, which is narrowest in the middle. The articular

surface has its facets divided by a very sharp ridge, equally defined in its whole course ;

the upper facet is plane, the lower concave, as in Physeter, but consistently with the

characters of the articular area in the malleus of Hyperoodon the upper facet is, in its

incus, not so much larger than the lower, as is the case in the Cachalot. The body is

developed to the same extent as in Physeter; the processus longus is as stout but

shorter than its representative in that Whale.

The stapes in Hyperoodon (Pl. LXII. fig. 35) has very stout crura, with hardly a

vestige of an aperture. The base is narrower than in Physeter, and not nearly so thick at

its margin ; towards the vestibule it is quite concave.

The stapes, but not the other ossicula, of Ziphius cavirostris is figured in a plate

illustrating Prof. Turner’s paper on that Cetacean*. It resembles, apparently, that of

Hyperoodon.

The malleus from the fine adult specimen of Berardius arnuaii in the College Museum

(Pl. LXIII. figs. 1 & 18) bears, to a well-marked degree, all the chief peculiarities seen

* Trans, Royal Soc. Edinb. 1872.

458 MR. A. H. G. DORAN ON THE MORPHOLOGY

in Physeter and Hyperoodon, The head is better developed than in either of those

Whales. It is half as long again horizontally as vertically, whilst in Hyperoodon

this portion of the ossicle, though not more prominently convex than in Berardius, is

almost a hemisphere. It rises very prominently above the facets. The depression on its

outer aspect is small and slit-like; the process holding the ossicle to the tympanic bone

is as short as in the Physeteride already described, and made up of the same elements

(compare Pl. LXIII. fig. 1 with figs. 13 & 14). The specialized modifications in the

processus muscularis and manubrium are very marked in the malleus of Berardius. The

processus muscularis (fig. 1, pm) is nearly as long as in Physeter, so that it is better

developed than in Hyperoodon. Here it may be observed that it is only in the Phy-

seteridee that the processus muscularis can be found as an actual process among the

Cetacea, sincein the Whalebone Whales and Dolphins its homologue is either a rough sur-

face or a depression on the tubercle, from which also springs the manubrium; but itis this

very tubercle which is wanting in the Cachalots and Ziphioid Whales, where the processus

muscularis is distinct.

In Berardius the manubrium is much better developed than in Hyperoodon or

Physeter. It is seen as a small, sharp spine *, arising close to the root of the processus

muscularis, and running directly downwards, whilst the other process runs straight

upwards. The two processes together lying close against the head, resemble the con-

trivance seen on board ship for belaying cable. The articular surface is extensive even

in proportion to the rest of the malleus; its upper facet is of the same form as in

Hyperoodon, plane, and very elevated at its margin; the lower is smaller and convex,

everted as in the Ziphioid Whale just mentioned.

Henee, in its malleus, Berardius is more perfectly specialized than either Pleat

or Hyperoodon. This ossicule is certainly, among these three Whales, largest and

bulkiest in Berardius, smallest, proportionately, in Physeter, owing to the ill-develop-

ment of the portion of the head external to the facets in that animal.

The incus (Pl. LXIIT. fig. 1) resembles that of Hyperoodon rather than Physeter; the

processus “‘ brevis” is still longer, more divergent, and flattened and expanded at its

extremity. Close to its root the body (as is the case in Hyperoodon) is not so reduced

and shallow as in Physeler; the groove dividing the facets is not very sharp in

Berardius. The processus longus is quite similar in character to that of Hyperoodon.

- The stapes is longer than in the other Physeteridee here described. In the College

specimens there is a prominent tubercle on the inner aspect of the head for the tendon

of the stapedius. The cruraare long, stout, and straight, a slit-like but pervious aperture

divides them. The base is narrow, its margin is rather thick, and it is quite concave

towards the vestibule.

I have examined the malleus and incus from a skull of Mesoplodon hectori in the

British Museum. These bones are much smaller than their representatives in the above-

described Physeteride. The malleus has a head as small as in Physeter, moderately

convex, and produced backwards to a prominent blunt point. The head projects slightly

above the facets, as in Hyperoodon; the articular facets are proportionately still smaller

* Vl. LXIII, fig. 1, where this spine is lettered im in place of mn, its true homologue.

OF THE MAMMALIAN OSSICULA AUDITUS. 459

than in that Whale, nor is the upper much larger than the lower. The processus mus-

cularis is a short blunt point. I found a few shreds of the tendon of the tensor tympani

adherent toit. The manubrium is represented by the faintest trace of a tubercle; there

is a ridge above it, into which a part of the process from the membrana tympani is pro-

bably inserted as in Platanista.

The processus gracilis and its accessories eonnectiae the malleus to the tympanic bone

is extremely short, and the depression in the head above it almost effaced. See Pl. LXIII.

figs. 19 & 19 a, where this bone in the closely allied Wesoplodon grayii is shown.

The incus is of the same character as in Hyneroodon and Berardius; it is rather

smaller in proportion to the malleus. The ridge separating its facets is almost as sharp

as in Hyperoodon.

Taken as a whole, and judging from the incus as well as the malleus, Mesoplodon

resembles Hyperoodon rather than Berardius. In the ill-development of the head of the

malleus we have a character which should place this Whale between Physeter and

Hyperoodon. In the smallness of the articular surface of the malleus it differs alike from

the Cachalot and Berardius, and approaches Hyperoodon.

The stapes has no aperture, and in form coincides with the same DSelele | in Hyperoodon,

confirming the remarks in the last paragraph.

In the Delphinidee the ossicles may be said to be central in character among the

Cetacea, the Baleenoidea and Platanistidee departing from them towards other mammals,

the Physeteride diverging, on the other hand, from the Dolphins in the highly specialized

characters of the malleus and incus. It is remarkable that among the Whales proper

the genus Balenoptera should so closely resemble the Dolphins in its ossicula.

In Delphinus the malleus (Pl. LXIII. figs. 2 & 3) in general form reminds the observer

strongly of the same in Balenoptera*. The outer aspect of the head is less flattened,

the articular facets are similar, the upper being the larger and planer, the lower the most

convex; but instead of the large rounded tubercle seen in the Whale just mentioned, a

small, blunt, slightly excurved process projects from the head in Delphinus. On its

inner surface may be observed a small spine (mm), directed backwards, hardly perceptible

to the eye, yet readily detected by rubbing the finger forward over the internal aspect of

the malleus. This represents the very perceptible homologue of the manubrium

described as existing in the Pike-Whale, and equally or more developed in other genera

of Dolphins rt.

The body of the incus is very feebly developed, being, as it were, absorbed by the

stapedial crus. Its articular surface has facets divided by a sharp ridge, as in the

Baleenoidea, and unlike Physeter and Berardius. The processus brevis is very slender

and much shorter than in the Ziphioids; the processus longus is very stout, and shaped

like its representative in Balenoptera; the Sylvian apophysis is of the same nature.

The stapes is about a line in length, with fairly divergent crura and a very thick base,

quite concave towards the vestibule. ‘The aperture is closed, but always represented in

* The ossicula of a fetal Dolphin, four inches in length, are all of precisely the same form as in the adult.

+ The homologous parts of the malleus in the Dolphins are indicated in Pl. LXIII. fig. 15, representing that

ossicle in Globiocephalus (see also figs. 3 to 12).

460 MR. A. H. G. DORAN ON THE MORPHOLOGY

this genus by a minute pit; and in no specimen of any Dolphin of any age in the Col-

lege Museum was it found ankylosed to the petrous bone, nor in any other Cetacean except

Balena mysticetus, and then not invariably. Hyrtl, who has described the ossicula of

the Cetacea (op. cit.) far more extensively than any anatomist before him, states that all

trace of the aperture is effaced “in Delphinus longirostris, braziliensis, tursio and

rissow ;” but he asserts that it is also absent in Monodon, though a distinct trace is seen

in the two College specimens from different skulls; and although he finds that the

aperture “ reappears in the Porpoise,” it does not seem more distinctly indicated in the

examples in the College collection than in Delphinus, and is decidedly not pervious.

The distinguished anatomist continues :—“ Possibly this depends on the age of the

individual specimen ; a conjecture, however, not justified by the persistence in all other

animals of the form of the stapes in different degrees of age.” I have not found the

aperture more blocked up or less clearly indicated by a depression in adult than in

young Delphinoid crania; and, on the other hand, a specimen of the stapes of Bradypus

tridactylus in the College collection, from an adult Sloth, has a much smaller aperture

than another stapes from a cranium of a younger animal of the same species.

In Globiocephalus (Pl. LXIITI. figs. 4 & 15) the head of the malleus is rather larger

than in Delphinus, and the projecting tubercle rather smaller; but it is marked at its

apex by a distinct rough surface for the insertion of the tensor tympani tendon (pm),

and the manubrium (mm) is represented by a very sharp spine pointing backwards and

flattened against the inner side of the ossicle. The bony process connecting the malleus

to the tympanic bone is rather long, and the same elements as described in Balena

may be observed in it ; the processus gracilis (pg) or the anterior column is thick. The

incus has the same characters as in Delphinus ; the stapes, too, is very similar, but some

specimens have a pervious aperture.

In a Lagenorhynchus in the British Museum the ossicula are of the same form as in

the last genus, and the manubrium is as well developed (fig. 11). In Phocena the

malleus (fig. 5) most resembles that of Globiocephalus; the incus is very similar, and

the stapes has generally a very minute but pervious intercrural aperture. The malleus

is connected by bone to the tympanic, though by a somewhat thin processus gracilis,

contrary ts Hunter’s experience. .

In Delphinapterus albicans (Pl. LXIII. fig. 8), the Beluga or White Whale, the

malleus much more resembles that of Delphinus than the same ossicle in G'lobiocephalus,

the process in front of the head being rather long and curved outwards. The recurved

splinter representing the manubrium in other Dolphins is quite absent, and represented

by a rough ridge (mn), hardly visible except by aid of alens*. ‘There is no excavation

on the outer aspect of the head of the Dolphins as in Balena; it seems as if the edges

so prominent in the Greenland Whale had rolled in and filled up a cavity, and to a less

extent in Physeter.

The incus of Delphinapterus is of the form prevailing among the Delphinide. The

stapedes in the College collection have neither so much as a trace of an aperture.

* As the malleus of this Dolphin is larger than that of most Globiocephali, the absence of the spine-like manu-

brium (as in the smallest Delphint) is not due to diminished proportions or development of the entire ossicle.

OF THE MAMMALIAN OSSICULA AUDITUS. 461

The malleus of a large Orca from Tasmania (Pl. LXIII. fig. 7) in general aspect

appears like a magnified malleus of Delphinus; but some of its elements are propor-

tionally more highly developed than in the typical genus. The head is large and convex ;

the depression on its outer aspect is very deep, and almost recalls Balena. The size of

the processus gracilis and the elements associated with it, connecting the ossicle to the

tympanic bone, contrasts strikingly with the reduced proportions of the homologue of that

process in the malleus of the Physeteride. The articular surface is as small as in Delphi-

nus, and the facets are of similar type; but the tubercle from the head of the malleus is

far larger than in any of the Dolphins of which the ossicula have been already described.

it is much more prominent than in Balenoptera. At its upper extremity, looking

forwards, is a conspicuous rough depressed surface for the insertion of the tensor-

tympani tendon, Below is a sharp splinter-like process representing the manubrium.

This is much more developed than in Delphinus, or even Globiocephalus; and instead of

being directed backwards, and pressed, as it were, against the head, it is directed con-

siderably in a downward direction. ‘The whole tubercle assimilates to its homologue in

Balena, as the same in Megaptera recalls Platanista in general features.

The incus of Orca is quite Delphinoid; its short crus is thin and rather long, the

stapedial crus very stout. The stapes is similar to that of Delphinus; it has a minute

aperture.

In a smaller Grampus (Pseudorca meridionalis, Flower) all the ossicula much resemble

those of Orca. The malleus has the tubercle bearing the manubrium a little less

developed (fig. 6), so as to approach Globiocephalus.

The homologies of the splinter of bone inflected from the process projecting from the

bead of the malleus in most Cetacea, and of the rough surface or actual tubercle seen on

the point of that process, have been discussed in the description of the ossicula of

Balenoptera; and specimens in the College Museum illustrate the relations of the soft

parts in Dolphins, Whales, and Hyperoodon. I have dissected several ears of Dolphins,

Porpoises, and of a Narwhal, and the malleus from the latter dissection (Pl. LXIII.

fig. 16) is mounted in the special collection of ear-bones on which this treatise is

founded. The tendon of the tensor tympani is left attached to the apex of the process

from the head; and the whole membrana tympani is also retained, dried and varnished ;

its broad, flat, and pointed process may be seen attached by the point to the splinter-like

manubrium, Hyrtl’s “Leiste.” All the above preparations conform, on the whole, to

the descriptions given by that distinguished anatomist, and by Buchanan, concerning

the anatomy of the Cetacean tympanum. ‘The dissection above described was suggested.

by a figure of the malleus of Monodon in Hyrtl’s work.

The malleus of the Narwhal (Monodon, Pl. LXIII. figs. 9 & 16), though essentially

the same in type as that of Globsocephalus, is much elongated antero-posteriorly, and the

head merges into the broad process in front without any constriction or trace of sepa-

ration. The articular surfaces are of equal size(as Hyrtl observes), the upper being not

larger than the lower, as in all other Cetacea. The processus gracilis is of the same

type as in Balena and the rest of the Order; it is rather long. The incus has an even

SECOND SERIES.—ZOOLOGY, VOL. I. 3Q

462 MR. A. H. G. DORAN ON THE MORPHOLOGY

thicker stapedial crus than in other Delphinoidea*; the posterior is short and small; and

the facets are, of necessity, of equal size; they are divided by a sharp and prominent ridge.

The stapes is of the ordinary Dolphin type, with a very thick base, generally (but not

always) very concave towards the vestibule. It was found unankylosed to the fenestra

ovalis in a large adult skull in the College Museum. ‘The ossicle in this case had a

minute almost pervious depression between the crura.

In the ossicula of the PLATANISTID# we find interesting modifications. In the Gangetic

Dolphin (Platanista gangetica) the head of the malleus (Pl. LXIII. figs. 12 & 17) is

of the same form as in Delphinus, and the upper facet is larger than the lower; but the

process in front is extremely elongated, so as to be longer than the head itself, from

which it is hardly more distinct than in Monodon, except that a faint groove, generally

observed in the true Dolphins, divides them above and internally. Instead of a rough

and only slightly concave surface at the very extremity of the process, as is found in

most Cetacea to mark the insertion of the tensor tympani, there is a very deep pit on

the external aspect of the projection, actually nearer its root than its pot; this is

highly characteristic (fig. 17). The point itself is sometimes slightly hooked; and

from it to close under the articular surface, all along the inner aspect of the process,

runs a very distinct groove, ending posteriorly in a faintly marked tubercle representing

the manubrium (fig. 12, mn) It seems as if the groove were homologous with the outer

surface of that process in other Orders of the Mammalia, the hooked or blunt point

corresponding to the processus brevis. Many of the fibres of the fleshy prolongation of

the membrana tympani to the malleus are attached to the bottom of that groove, and

are often seen still adherent in imperfectly macerated specimens. In Monodon and the

Dolphins proper some of the fibres are in the same manner inserted along the back of the —

anterior projection from the head of the malleus, as well as to the splinter-like more

definite homologue of the manubrium +.

The processus longus of the incus (fig. 20) isas thick as in Monodon, and the articular

surface for the head of the stapes isas small; the posterior crus is even less developed

than in Delphinus. The stapes is of the same form as in the smaller Dolphins, without a

trace of any intercrural aperture; there is sometimes a large tubercle for the stapedius-

tendon on the inner aspect of the head.

By the kind permission of Dr. Giinther I have been enabled to examine the ossicula

from the skull of Inia geoffroyit belonging to the British Museum. ‘The general shape

ofthe malleus (Pl. LXIII. fig. 10) reminds the observer more of Delphinus than Pla-

tanista. The head and facets resemble the same in the Gangetic Dolphin ; but the process

in front is not nearly so produced as in Platanista. The tensor tympani is inserted into

-a pit (at pm) as in the Gangetic genus, which, however, is seated in Inia quite at the

tip of the process from the malleus, on the site of the same insertion in Delphinus. 'The

manubrium is a very distinct reflected spine (mm); and there is a groove running

* Hyrtl (op. cit.) directs attention to the small size of the articular surface for the head of the stapes on the end of

this process. :

+ In making a fresh dissection of the tympanum of Plataiista, I found a strong band of fibres inserted into the

groove, and the tensor tympani was inserted into the depression above described (Pl. LXIII. fig. 17).

~OF THE MAMMALIAN OSSICULA AUDITUS. 463

forward from it. Hence the malleus of Jnia allies it, on the whole, more to Delphinus

than Platanista. The articular surface for the stapes on the stout processus longus

is as wide as in Delphinus; the processus brevis is rather long, but slender. The

stapes is of the type prevailing among all the Dolphins ; it has an intercrural aperture.

Tn a full-sized adult Inia these ossicula are less than half as large as their representatives

in a Platanista of the same age and proportions.

In taking a general concluding view of the auditory ossicles of the Cetacea, it may be

remarked :—

I. The ossicula of these animals differ from those of other Mammalia rather in their

solidity than in any great size in proportion to the whole skeleton, and more in extreme

modifications of detail than in solidity of composition; for in size they are exceeded by

the Seals and Sirenia, in density by the latter, but in specialization of form by no other

mammals.

II. The general diagnostic features are :—in the malleus, constant, firm bony ankylosis

to the tympanic bone through the medium of the processus gracilis, with ill-development

or complete suppression of the manubrium; in the incus, great development of the

stapedial crus ; in the stapes, thickness of the crura contracting or obliterating the aperture.

This latter bone always fits firmly into the fenestra ovalis, to which, however, it is in

no species constantly ankylosed.

III. The genus Balena possesses undoubtedly the most generalized ossicula among

all the Cetacea, particularly in the case of the incus. The malleus has the least-modified

form of manubrium, the whole bone distantly resembling that of certain large Seals

(Macrorhinus, Stenorhynchus), as does also the stapes. |

IV. The malleus of Neobalena is the next least specialized. That of Megaptera is

much more modified in regard to the aborted condition of the manubrium.

V. Both in the malleus and incus Balenoptera is far more modified than any of the

above genera*. ‘The incus is quite Delphinoid, unlike either the Balena or Physeter

type. The malleus is even more modified than in the Dolphins ; its manubrium is of more

tubercular form, and the whole bone is more flattened.

VI. In the Physeteride the malleus is extremely modified. The tubercle projecting

from the head in other Cetacea is here obsolete; the manubrium and processus muscu-

laris are reduced to two small spines. That representing the manubrium is almost sup-

pressed in Physeter, Hyperoodon, and Mesoplodon, but is well developed in Berardius.

The head is largest in Berardius; in Physeter the articular surface is very extensive.

The incus is intermediate in type between the same in Balena and Delphinus. In the

the Ziphioid Whales the short crus is long and of peculiar shape. The stapes differs a

little in character from that of the true Dolphins. ,

VII. In the Dolphins the malleus is never so modified as in the Ziphioid Whales: the

tubercle projecting from its head is pointed, not rounded as in Balenoptera; and the

seat of insertion of the tensor tympani muscle is indicated by a roughened, depressed

surface. The manubrium may be quite absent (Delphinapterus) or reduced to a small

* See Appendix to this scction.

3Q 2

464 MR. A. H. G. DORAN ON THE MORPHOLOGY

spine, depressed against the side of the ossicle (Globiocephalus, Lagenorhynchus, Pho-

cena); or the spine may be well developed and directed downwards, away from the rest

of the malleus (Orca, Pseudorca). Part of the process from the membrana tympani is

inserted along a narrow groove on the tubercle. Monodon differs from other Dolphins

in the characters of its articular surface, and the length of the head and tubercle of the

malleus.

VIII. All the Dolphins agree in the characters of the incus, which has an ill-developed

body and processus brevis, but a very large stapedial crus. The stapes, too, is very

similar in character in all these Cetacea.

IX. Platanista differs remarkably from the true Dolphins in the great length of the

process from the head of the malleus, as well as in minor particulars. Such distinctions

are hardly perceptible in Inia.

Appendix to Remarks on Cetacean Ossicula—The malleus of a foetal Balenoptera

sibbaldii in the Museum of the University of Edinburgh bears a long process in front of

the head, with the same homologues, apparently, as are seen in Megaptera. Whether

or not this process is arrested, whilst the head increases during the growth of B. sib-

baldii is a question only to be solved by the examination of the malleus in an adult of

this species. If the process remains long, then it is truly remarkable, since it allies this

Balenoptera to the genus Megaptera, and separates it from B. rostrata and B. huttoni.

The incus from the same footus is of the Balena type, as the body is very well deve-

loped, the long crus by no means so large as in Balenoptera rostrata. For this observa-

tion I am indebted to the kindness of Prof. Turner. I may further add that the malleus, in

the skeleton of Macleayius (Hubalena) australis at the British Museum, is of the Balena

type.

THE OssICULA OF THE SIRENIA.

On account of their large size and conspicuous peculiarities, the ossicula of the Dugong

and Manatee have already become familiar to most comparative anatomists; and,

through their great durability, even the same bones in Rhytina and Halitherium have

been described respectively by Claudius * and Krauss +; but the most prominent cha-

racters which have rendered them interesting to zoologists are due to their structure and

consistence, the homologies of their various segments and processes being, on the whole,

much easier to distinguish than is the case with the ear-bones of the Cetacea.

he malleus of the Dugong (Halicore australis, Pl. LXIII. figs. 26, 27) is a heavy

pone, weighing on an average 40 grains, and measuring about an inch from the root of

the processus gracilis to the tip of the manubrium; the largest specimen never attains

the size and weight of the malleus of species of either Manatus or Rhytina, though, from

Krauss’s account, it appears to be longer than in Halitherium schinzi.

It is important to bear constantly in mind the position of this bone in the Sirenian tympa-

num. In ordinary mammals the head of the malleus lies far back in the tympanic cavity,

* «Tas Gehérorgan von Rhytina stelleri,’ Mém. Acad. Imp. des Sciences de St. Pétersbourg, 1867.

+ Leonhard u. Bronn’s N. Jahrb. 1862, p. 385.

OF THE MAMMALIAN OSSICULA AUDITUS. 465

so as to be often invisible from the external meatus; the processus gracilis stretches well

forwards from the rest of the malleus ; and the articular surface is placed almost wholly pos-

terior. In the Cetacea, on the other hand, the head of the malleus is much less posterior in

position, the processus gracilis runs from it directly outwards to join the tympanic bone,

and the facets look inwards. In the Dugong and Manatee the malleus holds an inter-

mediate position, as may be seen by examining a skull of Halicore in the College

Museum (No. 2633), where all the left ossicula are in situ. The head lies quite anterior

to the posterior horn of the dense annulus tympanicus, its manubrium projecting directly

outwards from it; the processus gracilis runs as completely forwards as in terrestrial

mammalia; but, again, the articular surface is as internal as in the Whales and

Dolphins.

The massive head of the malleus of Halicore australis is of irregular shape, about as

long antero-posteriorly as vertically, and slightly flattened from within outwards. Its

uppermost portion is wide, slightly convex, and does not project above the upper facet

as in most Mammals. The articular region lies entirely on the very broad inner aspect

of the ossicle. It is made up of an upper facet and two lower, which latter represent

the single inferior facet in other mammals, the extero-inferior segment of the articular

surface of the human malleus; they form almost a right angle with each other, and an

obtuse angle with the upper facet, a very distinct groove dividing them. The superior

facet is reniform, with its long axis vertical, and its concavity forward; its surface is

distinctly convex from above downwards. The smaller inferior facets* are oval; and

their edges generally coalesce along the high ridge which divides them, and whence

they slope. On the outer aspect of the malleus is a very deep groove running forwards

and slightly downwards; its upper border is thick and prominent; its lower is formed by

the root of the manubrium. This projecting upper border is homologous with the sigmoid

ridge on the neck of the human malleus, as may be seen by laying an ossicle of Homo

and of Halicore side by side, with the manubrium forwards. Hence it must represent

the solid curved neck of the laminar type of malleus in the Carnivora, Ruminants, and

other Mammals+. Completely in front of the malleus of the Dugong is the processus

gracilis (Pl. LXIIT. figs. 26, 27), which is broad at its root, but narrows rapidly.

Owing to the whole ossicle being placed unusually forwards this process is far shorter

than in terrestrial Mammals. Its upper surface is smooth and convex, being con-

tinued from the upper surface of the head; its lower is flat and very short, separated

by a deep horizontal groove from a prominent tubercle on the front of the head below

it. The processus muscularis is a blunt tubercle, looking forwards from the inner

aspect of the malleus, and is continued backwards and upwards as a very promi-

nent convex ridge, which is lost on the head of the malleus behind the inferior facets,

near the ridge on the anterior aspect already described. Strictly speaking, the whole

must be considered as the processus muscularis, having its root on the true neck (the

* These evidently represent the inferior facet of other animals, the convex ridge being here much exaggerated,

even to the point of dividing that facet into two.

tT See the account of the human malleus, antea.

466 MR. A. H. G. DORAN ON THE MORPHOLOGY

outer and anterior projection), as in many other animals. The manubrium stands out

from the head completely external; and its point is directed rather backwards: it is

separated from the rest of the malleus above by the groove on the outer aspect of the

ossicle; below it is blended with the base of the ridge continued from the processus

muscularis. It is wide at the root, and triangular in form; its upper border is very

broad, particularly towards the groove, and sharply bordered from the sides ; externally

it runs into the outer aspect by forming a prominent bold convexity, instead of a pro-

cessus brevis; the outer aspect is also sharply defined, its borders being continued from

above, and towards the extremity it is spatulate: in some specimens this dilatation is

very marked, in others hardly perceptible. The sides of the manubrium are flattened,

and somewhat concave near the root; the inner border is rather sharp, and quite concave ;

it blends with the head as aforesaid below the processus muscularis. It is wide at the

root, and triangular in form: its upper border is very broad, particularly towards the

groove, and sharply bordered from the sides; externally it runs into the outer aspect by

forming a prominent bold convexity instead of a processus brevis; the outer aspect is

also sharply defined, its borders being continued from above, and towards the extremity

it is spatulate; in some specimens this dilatation is very marked, in others hardly per-

ceptible. The sides of the manubrium are flattened and somewhat concave near the —

root; the inner border is rather sharp, and quite concave, it bends with the head, as

aforesaid, below the processus muscularis.

The incus of Halicore (Pl. LXIII. figs. 28, 29) is singularly placed, its “ processus

brevis” being absolutely superior in position to the rest of the bone, instead of being

postero-superior, whilst the stapedial crus is equally inferior; the facets look directly

outwards on account of the internal position of those on the malleus. The body is twice

as wide as it is deep; its inner aspect, which is wide, smooth, and convex, fits into a

depression in the petrous bone; its articular surface consists of a concave reniform facet,

corresponding to the upper one on the malleus, separated by a sharp projecting angle

from two oval lower facets, which look towards each other, and are divided by a deep

eroove, into which the projecting angle between the lower facets on the malleus is locked

in the recent skeleton. The posterior of the two lower incudal facets is borne on a very

prominent tubercle projecting from the outer and more posterior surface of the body of

the incus—a most characteristic feature, by which this ossicle in Halicore may be at once

distinguished from the same in Manatus, and apparently in Rhytina and Halicore.

The processus brevis is very short, and lies above and somewhat anterior to the rest of

the ossicle; it is firmly ankylosed to the petrous bone; but being thin and brittle, the

whole incus is generally detached if the skull is dropped on the ground, or struck

violently after maceration. The processus longus arises far from the outer crus on

account of the great width of the body, to which it lies inferiorly and slightly posteriorly.

It is short, slightly curved, flattened extero-internally, and of almost equal thickness

throughout ; the oval disk at the extremity is generally separated from the rest of the

crus by a faint groove, so as really to represent the os orbiculare, the existence of which

in Halicore seems to be denied; as a rule, the larger the mammal the less developed is

its Sylvian apophysis,

OF THE MAMMALIAN OSSICULA AUDITUS. 467

The stapes of the Dugong (fig. 30) measures about half an inch in length, and is of

very solid consistence; as the crura diverge but little, the base is narrow; the anterior

extremity of the latter and of the fenestra ovalis is placed much lower than the posterior,

although, after examining five skulls of Halicore in the College Museum, I do not find

the long axis of the base of the stapes or of the fenestra vertical, as has been asserted ; the

same remark applies to Manatus. 'The head is well developed, and the articular surface

is concave; the site of insertion of the stapedius is generally indicated by a slight depres-

sion. The anterior crus is half the thickness of the posterior, and both are rather

crooked; the aperture is circular, generally just large enough to admit a small sewing-

needle, but in some specimens it is of the calibre of a No. 1 English catheter. The base

is convex, most so posteriorly, and is narrowest towards the front ; it never appears to be

ankylosed to the fenestra ovalis, or even tightly fitted imto it, as in the Cetacea. The

broad well-formed head and comparatively thin anterior crus somewhat remind the

observer of the stapes of Bos; but in the other ossicula of Halicore and the other Sirenia

few, if any, distinct analogies to the same bones in any particular order of Mammalia can

be made out.

Since the extinction of the Rhytina the American Manatee (W/. americanus) has borne

the largest and heaviest malleus known in any animal, as Hyrtl has already stated.

Indeed, to judge from Claudius’s account of that ossicle in Rhytina, of which the head,

he remarks, is of about the size of an ordinary hazel-nut, it hardly can have exceeded,

as a rule, the malleus of a large Manatee, though I have never seen the processus

gracilis of the latter “at least 13!" in diameter,” which is the case in the specimen from

the extinct Sirenian he describes. The specimen in the collection of ossicula in the

College Museum (Pl. LXIII. figs. 21, 22), from a skull of WZ. americanus, in the Com-

parative-Anatomy series, measures over an inch in length, and weighs over sixty-one

erains; the head is quite as large as a middle-sized hazel-nut. This bone, as well as the

other ossicula, lies, in the recent skull, in the same position as has already been described

in Halicore; but it offers many points of difference in its structure from the malleus of

the latter animal.

The head is much longer antero-posteriorly than vertically. All the facets are larger ;

and the upper, though reniform, has its concavity backwards, and its surface is distinctly

concave above. In the College specimen there is a prominent peg-like process (Pl. LXIII.

fig. 21) projecting immediately below the inferior border of this facet, bearing supe-.,

riorly another small facet; not a trace of this can be seen in any specimen from -

M. senegalensis in the same series. The lower facets in Manatus americanus are of

much the same form as in the Dugong; but at the top of the prominent angle their

borders do not blend, but are nearly a line apart. The outer aspect of the malleus is

very wide and quite convex; the groove is very shallow anteriorly, and quite effaced

behind, in which direction, however, is a very sharp and prominent ridge analogous to a

similar projection in Halicore, the homologies of which have already been discussed.

The processus muscularis is a stout incurved process, continued backwards as a con-

spicuous elevation towards the former ridge, precisely as in the Dugong.

468 MR. A. H. G. DORAN ON THE MORPHOLOGY

The manubrium differs in form from that of Halicore. The upper border is very sharp

and narrow; it forms a semicircle, and projects against the membrana tympani in the

live subject; the outer border is short, and its margins, united above, diverge but little

towards the extremity ; its surface is concave vertically, and slightly convex horizontally.

The inner border is much blunter than the outer in Manatus.

The processus gracilis is not relatively so well developed as in Halicore, and projects,

as a fragile splinter of bone, from the massive head; it is firmly ankylosed to the

tympanic ring; and, from its very nature, the whole malleus is easily broken off from

the ring.

The incus (figs. 23, 24) is placed in the tympanic cavity precisely as in the Dugong,

the processus longus lying above the body, and the other crus running downwards from

it. The body is even shallower and wider between the crura than in Halicore ; its inner

surface is less extensive and less convex, and much overhung by the upper facet of the

articular area, which facet is very wide, reniform, and faintly concave; the two lower

facets look towards each other, and are separated by a deep groove, into which the cor-

responding prominence between the opposing surfaces on the malleus are fitted during

lifetime. There is not nearly so marked a prominence behind the upper and more

posterior of these facets in the Manatee, like that which projects so conspicuously on the

posterior side of the Dugong’s incus. The processus longus is rather long, slightly

curved, and flattened extero-internally; it widens out towards the point, where it is

ankylosed to the petrous bone. The processus brevis is very short, stout, and curved;

the articular surface for the stapes is very wide, almost circular, and nearly plane; a

faint trace of a rim can be seen above it, separating it from the crus, as the homologue

of the os orbiculare.

No stapes of this Manatee exists in the College collection; but that of I, senegalensis,

of which there are several examples, will be shortly described. The malleus from an

adult of the latter species is rather smaller than in the American kind ; the upper facet

is not so wide, and there is no peg-like projection internal to it. The processus muscu-

laris is extremely stout and prominent; but the manubrium is not so broad as in

M. americanus.

The incus of Manatus senegalensis very closely resembles that ossicle in the American

Manatee; the processus longus is of similar form and length; the articular surface is

slightly different, in accordance with the characters of that surface on the malleus of the

African species; hence the upper facet is smaller, and does not overhang the inner side of

the body as much, and the two lower facets gape wider than in the American species.

In none of the existing Sirenia does the incus reach the size of that of the Seal

(Macrorhinus proboscideus), where it is larger than in any other animal.

But as is the case with the malleus, so is the stapes of Manatus (fig. 25) largest among

mammalian stapedes. It is over an inch in length, and weighs about one drachm; it is

of the dense consistence of the other ossicula in this Order. In shape it differs from the

stapes of Halicore, being over twice as long as it is wide. The head is well developed,

and bears an almost cireular articular surface. Both crura are long and hardly

ON THE MAMMALIAN OSSICULA AUDITUS. 469

divergent; both are crooked; but the anterior is the thinner and straighter; the

posterior is very broad inferiorly, often bulging prominently just above the base. The

aperture appears as an oblique slit, a quarter of an inch long from above; but on the

lower aspect it is more circular: in a large specimen it is of the calibre of a darning-

needle. The sides of the crura bounding it are slightly concave, but not absolutely

grooved. The base is rather narrower than are the crura a little above it, but it is

wider than the head; it is convex towards the vestibule; and its margin is generally

rather concave above, in accordance with the form of the fenestra ovalis in this animal.

According to Claudius *, the malleus and incus of Rhytina closely resemble those of

Manatus. In the work already referred to, these ossicles are figured and described ; and

that anatomist does not name any point in which the extinct differs from the above-

named existing Sirenian as far as these bonesare concerned. ‘The inner border of the

manubrium, he says, is shorter, narrower, and sharper than the outer (that is, the upper

and outer surface of our description); but he does not state whether the latter is broad,

with sharply defined margins, as in the Dugong, or sharp and narrow, as in the Manatee.

The incus is placed, according to his account, precisely as in the existing species; and

the short crus is ankylosed to the petrous bone; the processus longus is described as

long and strong, though in the plate it does not appear to be longer than in Manatus.

Dr. Krauss + has had the good fortune of procuring and examining all three ossicula

of Halitherium schinzi, which he figures both separate and articulated. From the

letterpress and the plates it appears that the malleus is intermediate in form between

the Manatee’s and Dugong’s. The body is figured as short antero-posteriorly, asin Hali

core ; but the manubrium is represented more of the Manatws-type; the articular surface_

though not distinctly described or figured, must, judging from the incus, be as in the

latter animal; the processus gracilis is rather long, thin, and brittle. As for the incus,

it is both described and figured as much more like that of Manatus than Halicore; the

articular surface is represented as large as in the Manatee, and similar in its facets; more-

over there is no prominent tubercle on the outer aspect of the body, backing up one of

the facets, as I have observed and described in Halicore. 'The stapes is described as

very similar to that of Halcore, “‘ whilst that of the Manatus is much larger, thicker,

- and rounder.” The illustration confirms this statement, but represents the crura as

rather longer than in the Dugong; the anterior crus is much the thinner ; the head is

broad; and the whole bone decidedly, as far as one can judge from a plate, approaches

the quadrilateral type of the larger Ruminants.

In conclusion, it may be said of the ossicula of the Sirenia :—

I. That their dense structure and clumsy form, consistent with the characters of the

whole skeleton, distinguish them at a glance from their homologues in all other Orders.

IT. That their modifications lie much more in their peculiar general conformation than

* Mém. Acad. Imp. des Sciences de St. Pétersbourg, 1867.

t As already quoted in the Jahrb. 1862.

470 MR. A. H. G. DORAN ON THE MORPHOLOGY

in the suppression or peculiar development of any of their processes, which are so remark-

able in the Cetacea.

III. Not only do they differ clearly from the Cetacean ossicula, but their modified

general structure may be said to mask or obliterate any homologies to the ossicula in other

mammals, which otherwise might be observable—the only exception being the approach

to a quadrilateral type of stapes in Halicore, apparently more marked in Halitheriwn,

which, whatever be its value, cannot fail to remind the anatomist of the bovine form of

that ossicle.

IV. Halicore differs distinctly from Manatus in the antero-posterior shortness of the

body of its malleus, in the smallness of its articular facets, in the form of its manubrium,

in the tubercle on the outer and posterior aspect of the body of the incus supporting the

articular surface, and in the shorter incus with more divergent crura. —

V. In Rhytina the malleus and incus appear very like those of Manatus. But in

Halitheriwm the characters of the ossicula are extremely intermediate—the incus being

most like that of Manatus, the stapes like the Dugong’s, and the malleus possessing

features seen in both.

VI. The distinguishing feature by which WM. americanus may be known, by its ossi-

cula, from IM. senegalensis, is a slight difference in the characters of the malleo-incudal

articulation.

Tab OSSICULA OF THE EDENTATA.

As is the case with the rest of the skeleton, the ear-bones in this Order exhibit con-

siderable variety of type; the characters of the malleus and incus are very general;

whilst the stapes assumes the Sauropsidan form in one group, and approaches it in

several others.

In the two recent genera of the PayropHaGa the malleus offers points of difference too

marked to be overlooked. ‘This ossicle in the Ai, or Three-toed Sloth (Bradypus tridac-

tylus, Pl. LXTV. fig. 1) has a rounded, well-developed head, projecting a little upwards, but

more forwards, and rather flattened laterally. The articular surface is deeply cut, the facets

lying almost at right angles to one another ; they are narrow horizontally; the upper is

almost plane, the lower perceptibly convex ; the neck is long, slender, and almost straight.

By laying the malleus of Homo by the side of one from this Hdentate, the identity of the

sigmoid ridge of the former with the much less curved neck, just described, in the latter

will at once become very evident. ‘The processus gracilis forms a very acute angle with

the neck, leaving a narrow space between them, homologous with the lamina, but filled

up with thick and friable rather than lamellar bone. There is no trace of any processus

muscularis, either on the neck or the handle. The manubrium forms an angle of over

150° with the rest of the malleus; it is of medium length, very thin, and compressed

laterally, and broad at the base, where, externally, is seen a sharp angle, representing the

processus brevis. The external aspect is bordered from the sides by sharp edges, as in

the Carnivora, &c., but very narrow; the extremity is slightly spatulate, and perceptibly

recurved.

OF THE MAMMALIAN OSSICULA AUDITUS. ATL

Thus the malleus of Bradypus has very general characters. It cannot be considered

a low type, like that of any marsupial; but its affinities are rather in the direction of

Mammals undeniably higher than the Edentata, such as the higher Primates, the

Ichneumons, and the Horses, not from any positive characters, but rather from the

absence of salient modifications so prominent in animals much nearer to them in other

respects.

The incus (Pl. LXIV. fig. 1) has a rather shallow body, with proportionally large

crura. The processus brevis is very stout and rather long, sharply pointed and very

divergent from the body; the processus longus is well developed and broad. I have

not observed any bifurcation on its inner surface in three incudes from different

individuals in the College Museum, although Hyrtl names that peculiarity in describing

the incus of Bradypus. The Sylvian apophysis is small, but very distinct, being attached

to the crus by an extremely narrow pedicle. The affinities of this ossicle are very

uncertain ; the stout divergent processus brevis recalls the type of medium-sized Bovide ;

but in the Sloth the incus is smaller in proportion to the malleus than is the case with

the Ungulata.

The stapes in Bradypus tridactylus (fig. 1) is a very interesting bone, remarkable for

having occasional (not constant) features which relate it very decidedly to the Didelphia

and Sauropsida. In one pair belonging to the College series the head is moderately

developed, the crura are stout, rather long, and but little divergent, leaving between

them a narrow but very distinct aperture. In another pair in the same collection

the right stapes has no aperture, whilst the left has a minute foramen between its

erura. In all, the base is very broad, recalling its form and proportions in the Marsupial

stapes.

Yet Hyrt! remarks (op. cit.), “It is most remarkable that Bradypus tridactylus has

only one crus, which is broad; Bradypus didactylus (sic), on the other hand, has two

very thick crura, which, on that account are separated by a space hardly the width of a

hair. The simple crus of the stapes of B. tridactylus is very thin and transparent, whilst

the stapes in B. didactylus is massive and conical.” He then remarks that he has only

examined the stapes cf B. tridactylus in an embryo.

From Hyrtl’s observation, and from the distinct and complete fusion of the crura in

the stapes on one side only in the specimen already referred to, we see in the Ai a

singular wavering between the higher and lower type of this ossicle in the Vertebrata

that possess it. The intercrural aperture in none of the College specimens is as large as

it is in the columella of the Golden Hagle (Aquila chrysaetos), so that the presence of that

space must not be considered unreservedly as a tendency to higher type.

The ossicula of the Unau (Cholepus didactylus) differ from tlie above in some respects

(see Pl. LXIV. fig. 2). The head of the malleus is even more developed, and is some-

what produced forwards, as in the Hystricide. The articular surface is similar; the neck

is rather longer, but more curved, assuming the sigmoid form of the ridge on the human

malleus. The lamina is similarly represented by a narrow tract of rather thick bone.

Indeed the malleus of both the Sloths is distinctly intermediate between the non-lami-

3R 2

472, MR. A. H. G. DORAN ON THE MORPHOLOGY

nated form of Homo and the broadly laminated type of the Carnivora and Artiodactyla*.

The manubrium of Cholepus is longer than in Bradypus, and forms little more than a

right angle with the neck; this alone gives the whole malleus a different aspect from

that of the Ai. The processus muscularis is represented by a distinct tubercle on the

inner side of the root of the manubrium; the latter is even broader than in Bradypus.

The incus of Cholepus does not differ from that of the Ai; the Sylvian apophysis is

of the same character.

The stapes of the Unau has even less divergent crura than that of Bradypus; ‘the

aperture is either very small or quite obliterated: these two conditions may be seen in

different examples in the College collection.

Turning to the ENromopHaGa, the ossicula in the genus Dasypus are larger than in

the Sloths. In the College collection all three in Dasypus villosus are absolutely

larger than in Bradypus.

In the fine malleus of the 6-banded Armadillo (Dasypus sexcinetus, P|. LXIV. fig. 4)

the head is well formed and globular, without being produced forward or in any par-

ticular direction. The articular surface is deeply cut, the facets lying at right angles to

each other; the upper is very wide and nearly plane, the lower is barely half its area

and more convex. The neck is short and bent; the processus gracilis is arched towards

the body of the ossicle, and then curved downwards and forwards; the space thus formed

is narrow, and occupied by a true lamina of really lamellar bone. The processus mus-

cularis is a bony tubercle on the inner side of the neck, close to the root of the handle.

The manubrium is long and straight; it forms an angle of about 140° with the neck ;

it is compressed laterally, but not very broad at the base. A true processus brevis

exists—the angular projection at the external aspect of the base being considerably

everted, and hence is not merely the simple apex of an angle formed by the upper edge

of the base and the outer side of the manubrium. This outer aspect, as in Bradypus, is

very narrow, yet distinctly bordered from the sides; the extremity is very slightly

spatulate and but little recurved. In the well-formed head, decurved processus gracilis,

narrow lamina, and long manubrium this malleus recalls that of the Horse (Hquwus).

The body of the incus (Pl. LXIV. fig. 4) is shallow and broad; the processus brevis

(which lies much more superiorly than posteriorly) is short and sharp-pointed, it

diverges very little. ‘The processus longus is thin, long, and very divergent; the pro-

cessus Sylvii is absolutely sessile, and hardly separated in the adult by any trace of a

groove; but a distinct channel is to be seen in incudes from new-born Armadillos. The

characters of the crura very plainly and completely divide Dasypus from the Sloths.

The stapes is a large bone of very light and elegant form (fig. 4). The head is long

and flattened, with a small tubercle for the insertion of the stapedius. The crura are

long, straight, and conspicuously slender ; they are well grooved towards the aperture,

which is very wide. The base is broad horizontally, but not very deep vertically ; it is

distinctly convex towards the vestibule.

* Refer to the remarks on the human malleus (wnted), particularly the description of the sigmoid ridge and its

homologies.

OF THE MAMMALIAN OSSICULA AUDITUS. 473

In every respect this stapes approaches the type of the higher Mammalia, as does that

ossicle in the Ant-eaters and Orycteropus, contrasting strongly with the Sauropsidan

stapes of Manis and the tendencies in that direction already described in the Sloths.

In Dasypus villosus and D. minutus the ossicula are almost identical with the same in

D. sexcinctus. The stapes in the Hairy Armadillo is even more slender than in the six-

banded species ; the processus muscularis is more prominent in the malleus of D. mimutus

than in the larger species.

In the Peba Armadillo (Zatusia peda, Pl. LXIV. fig. 3), the ossicula resemble greatly

those of Dasypus. The processus muscularis mallei is obsolete in the College specimens ;

the crura of the incus are of similar character ; the stapes is of slender form with a wide

aperture. I have not found the head of this ossicle to be unossified in any young or

adult Zatusia, as Hyrtl asserts of T. hybrida; but that portion is readily broken

off from the fragile crura. The malleus in the new-born Zatusia (fig. 7) has an ill-

developed head and a similar manubrium to that of Priodon (fig. 5). It is remarkable

that whllst the malleus of the adult Zatusia or Dasypus resembles that of Hquwus, the

same ossicle in the young of those genera or in the adult Priodon reminds the observer of

Rhinoceros. ‘This is at least remarkable when we remember the relationship of the

ossicula of the adults in some of the Ungulata to those of the young in others.

In Priodon gigas (Pl. LXIV. fig. 5) the malleus has a much less developed and less

globular head than in the above-described Armadillos: the articular surface is of similar

form; the neck is thinner, longer, and straighter. The lamina is rather wide, and pro-

longed a little on the strongly decurved processus gracilis, which is joined by a styliform

process from the front of the head, which has its homologue in the Carnivora, Rodentia*,

Ungulata, Cetacea t+, and most other Orders. The processus muscularis is wanting.

The manubrium forms little more than a right angle with the neck; it is much more

slender than in Dasypus, and bent outwards on the rest of the malleus; there is no pro-

cessus brevis, but, on the contrary, a very blunt projection at its site. The ill-developed

head, peculiar lamina, and the form of the manubrium give it a general resemblance to

the malleus of Rhinoceros.

The incus of Priodou (fig. 5) has a shorter processus brevis than in Dasypus; the long

crus is much stouter, and has a notch or fork on its upper margin. Hyrtl has observed

a similar appearance in the Sloths. The processus longus bears a distinct, sessile, and

very elliptical Sylvian apophysis.

The stapes (fig. 5) has much stouter crura than in Dasypus or Tatusia, with a small

intercrural aperture. The base is rather narrow; and in no wise does this ossicle bear

any resemblance to that of Bradypus or Manis.

In the Three-banded Armadillo (Volypeutes tricinctus, fig. 6), the malleus of the

adult much resembles that of Priodon; but the lamina is very wide, so that the ossicle

is modified from a low type in the same direction as in the Carnivora, Artiodactyla, and

Muride. The neck remains straight, and the outer angle of the base of the manubrium

* See remarks on the malleus of the Musquash (Fiber).

+ Description of the malleus of Balena, anted.

AGA, MR. A. H. G. DORAN ON THE MORPHOLOGY

blunt, as in the Giant Armadillo; but there is a distinct tubercular processus mus-

cularis in the same position as in Dasypus.

The incus of Tolypeutes resembles that of Dasypus and Tatusia much more than

Priodon, the processus longus being long, slender, straight, and very divergent.

But the stapes from the adult specimen in the College Museum is of more Sauropsidan

type than in any other Edentate except Manis. ‘The head is long and narrow; and the

slender crura keep together till close to the foot-plate, where they diverge, leaving a small

aperture betweenthem. The base is broad vertically as well as horizontally. It is the

manner of divergence just described, and not merely the smallness of the aperture, which

relates this stapes of Tolypeutes to obviously lower forms. As Manis in its stapes with

absolutely fused crura allies itself to Dasypus, Phalangista, Perameles, the Ornitho-

delphia, and most of the Sauropsida, so does Tolypeutes tend, in the character of the

aperture in that ossicle, to Macrapus, Didelphys, and some Birds.

In the squamate Edentata all three ossicles present positive points of peculiarity.

The malleus of Manis dalmanii (Pl. LXIV. fig. 8) has a large head, much flattened

antero-posteriorly ; the front is but slightly convex, and it does not project beyond the

upper margin of the articular surface. This latter has a very prominent border, and in

form is not unlike the same articular surface in Man and the Anthropoid Apes. It is

placed very obliquely, so that its outer extremity * lies much higher than the inner

(fig. 10); it is also divided into two facets, one superior and internal, the lower

being the more external. Hach presents a vertical convexity; and they slope towards

each other, so that the whole articular surface appears concave when viewed late-

rally. But this surface is far wider in proportion to the entire head than it is in

Homo, nor does the head in Manis project prominently above it as in Man; moreover

the division into facets is even less distinct +. The neck is very short and curved; the

~ lamina is of medium width, and formed of not very thin or transparent bone; there is no

processus muscularis. The manubrium forms an angle of about 130° with the neck, and

resembles that of Dasypus; a distinct processus brevis exists; the manubrium is rather

broad at the base, and much flattened laterally ; the extremity is distinctly recurved.

The character of the articular surface and the form of the head distinguish the malleus

of Manis alike from the same in the Sloths and the Dasypodide.

The incus (fig. 8) lies in the skeleton with its articular surface looking directly

forwards; it is rather broader than deep in the adult Manis daimanii. The postero-

superior part of its body (which is really absolutely superior in Janis) is well developed

and bulging, but not so markedly deeper than the other part of the body over the pro-

cessus longus as is the case in some species of Manis. The crura are very short, and

diverge but little ; the stapedial crus bears an ill-developed elliptical Sylvian apophysis,

which is pedunculated.

The stapes of the Pangolin (Manis) is most remarkable, being absolutely Sauropsidan ;

% This is actually the wpper extremity in Manis; but being the homologue of the outer in most mammals, that

term is retained, as in describing a similar condition in the human malleus.

+ On the subject of the facets, I must again refer to my observations on the homologies of the articular surface in

the human malleus.

OF THE MAMMALIAN OSSICULA AUDITUS. 475

the head is very ill developed; the column does not exhibit the least trace of any division

into crura ; the foot-plate is almost circular, and in diameter almost equals the length of

the column. <A tendency to this form of the base has already been described in Bradypus ;

and the gradations from the completely bicrurate stapes of Dasypus, through the form

seen in Tolypeutes, to that in Manis are most instructive to observe.

In Wanis tricuspis, M. longicauda, and WZ. pentadactyla, the malleus is identical in form

with that of I. dalmanii; but in the incus (Pl. LXIV. fig. 9) the portion of the body over

the processus brevis is extremely high, so that the whole ossicle assumes a form seen in

Mycetes and Pithecia; the processus brevis is thinner and slightly more divergent than

in YW. dalmanii, though both processes are as short. The stapes has a shorter column,

without any trace of division into crura, and it is absolutely shorter than the diameter

of the foot-plate, which is not so circular as in the species first described.

In the Great Ant-Eater (Myrmecophaga, P|. LXTV. fig. 11) the malleus has a large head,

very much flattened laterally and produced forwards; its upper portion, very wide antero-

posteriorly on account of the lateral flattening, does not project at all above the level of

the articular surface, which is rather deeply cut, and much wider vertically than from

without inwards. The lower facet is the most concave, and is divided from the upper

by a very faint groove. The neck is long and almost straight; there is a very narrow

lamina; the manubrium forms an angle of about 150° with the neck. The handle is

long and slender; there is no processus brevis, but a very blunt angle in its. place; the

extremity is very slightly spatulate and scarcely recurved. The sides are flattened, and

not broad near the base ; there is no trace of any processus muscularis. This malleus of

the Great Ant-eater bears a certain resemblance to that of Priodon, from which it may

at once be distinguished by the form of the head.

The body of the ineus of Myrmecophaga (fig. 11) is almost square, although rather

broader than deep. The processus brevis is moderately long, divergent, and blunt-

pointed; the processus longus is well developed without being very stout, and bears a

distinct elliptical Sylvian apophysis on a broad pedicle. The whole bone is large in

proportion to the malleus.

The stapes (fig. 11) has an ill-developed head and long very slender crura, with a

wide aperture between them. The base is thin, but very broad vertically as well as

horizontally. Thus this bone in the Great Ant-eater approaches the type of Dasypus

rather than Bradypus, Tolypeutes, or Manis. The vertical breadth of the foot-plate

(that is to say, the tendency to roundness in that portion of the ossicle) is the oniy

approach towards the Sauropsidan columella, the crura tending towards much higher

types.

In Tamandua tetradactyla (Pl. LXIV. fig. 12) the head of the malleus, though flat-

tened laterally, projects much more above the upper margin of the articular surface

than in Myrmecophaga, so as to be very convex above. The neck is a little more

curved, and the lamina wider; the manubrium is very similar in form. The incus has

an even squarer body ; its processus longus bears a similar Sylvian apophysis.

In Cyclothurus didactylus (P\. LXIV. fig. 18) the head of the malleus is as prominent

_ above asin Tamandua; the neck is shorter, and the manubrium is much broader at the

A76 MR. A. H. G. DORAN ON THE MORPHOLOGY

base, than in the two larger Ant-eaters. The whole bone somewhat resembles the same:

ossicle in Bradypus. The incus has a shallow body ; and the end of its stapedial crus bears

a very elliptical and sessile Sylvian apophysis, so narrow vertically as to be almost

rod-like. The stapes in a young specimen in the Collection has no trace of any division

between the crura, and in general resembles that ossicle in the Sloths.

In the Ant Bear (Orycteropus ethiopicus, Pl. LXIV. fig. 14) the head of the malleus

is ill-developed, the articular surface is shallow and wide vertically, both facets are but

faintly convex. The neck is long and curved ; it bears internally and nearly one line

distance from the root of the handle a prominent tubercular processus muscularis. There

is a wide and transparent lamina. The manubrium is very long, and forms a right angle

with the body; it is rather broad at the base, and presents an angle corresponding to the

processus brevis ; the extremity is very little dilated or recurved. In the characters of the

head the malleus of the Aard-vaark (O. capensis) resembles Priodon ; but the greater

length of the manubrium, the broad lamina, and the very distinct processus muscularis

distinguish it readily from the latter.

The body of the incus is shallow ; both crura are long and divergent; the Sylvian apo-

physis is almost obsolete, being only separated from the long crus by an indistinet

groove.

The stapes has an ill-developed head, and two long slender crura bowed outwards; a

thin bony lamina is spread between them below, closing the lower part of the aperture,

which otherwise would be wide. The foot-plate is not very wide horizontally, and is narrow

vertically. Hence the stapes of Orycteropus, on account of the latter feature, is of a

still higher type than that of Myrmecophaga.

Reviewing, then, the ossicula of the Edentata as a whole, it may be asserted :—

I. That in the Bradypodide the characters of the malleus and incus are fairly gene-

ralized, whilst the stapes assumes to a certain extent, though never completely, Sau-

ropsidan characters. Cholepus differs from Bradypus in the form of the malleus.

II. That in the Loricata the genera Dasypus and Tatusia present much higher cha-

racters in their ossicula than are found in Priodon and Tolypeutes. In the first two

genera the stapes is of a high type, both as to its crura and its base; in Priodon that

ossicle is more Sloth-like; in Yolypeutes it resembles that of the Kangaroos and other

Marsupials, or even some Birds, but not the complete columelliform type of Manis and

most Aves.

III. Manis has the most positive characters of interest to be found among all Hdentata :

the malleus is more specialized than in the other groups; the incus has peculiar though

less unusual characters; and the stapes is more absolutely Sauropsidan in every respect

than in any other placental mammal.

IV. Inthe Ant-eaters the malleus may be known from that of other Edentata by the

form of its head. The stapes is of as high type, as to the free separation of its crura, as

that of Dasypus; but the base has a tendency to assume the circular rather than oval

form seen in the columella of Birds. Cyclothurus, in its malleus, tends more towards

OF THE MAMMALIAN OSSICULA AUDITUS. ATT

Bradypus and Dasypus than do Myrmecophaga and Tamandua; and its stapes, In one

young specimen at least, approaches much more to the Sloth type.

V. In Orycteropus the malleus, quite unlike that of any Ant-eater, Sloth, or Manis,

approaches to a certain extent that of Priodon, from which it has, on the other hand,

very distinctive features. The stapes is of as high a type as in Dasypus.

VI. That many instances are observed in this Order of the ossicula of the adults of

some genera resembling those of the young of others; especially does the malleus of

Priodon or Tolypeutes, however old and well developed the specimen it be removed

from, remind the observer of the malleus of a new-born or fcetal Dasypus or Tatusia.

The highest degree of development is not necessarily seen in the largest species, as is

at once evident on comparing the malleus of Priodon gigas with that of Dasypus

mmutus.

THE OSSICULA OF THE DIDELPHIA.

The study of the ear-bones of the Marsupials is of necessity peculiarly interesting, as

it is on them that we must depend to a great extent for elucidation concerning the

obscure homologies of the proximal elements of the mandibular and hyoidean arches in

the Mammalia and Sauropsida. Considering how undoubtedly inferior these animals

are to placental mammals, it must be clearly understood that their auditory ossicula

are, on the whole, of proportionally high type; for though these little bones in adults of

this subclass resemble, as a rule, their representatives in the foetus of many higher

animals, still hardly one persistent embryonic feature observable in any Order of the

Didelphia is not also to be found in many highly organized Monodelphia. Indeed, in

large Orders, as the Ruminants, or smaller ones with families very specialized and

different in important features from each other, as the Hdentata, certain genera, as

already shown, retain in a most remarkable and instructive manner characters which

disappear during advance to maturity in genera closely allied to them, where the species

may be either larger or of distinctly higher conformation.

Those families will first be described where the stapes has divergent crura, as this

character is manifestly a high one. But it must here be remarked that the divergence

is slight, and only commences a little above the base-plate; never has any Marsupial

been found to possess a stapes with a wide aperture extending freely in the direction of

the head as in Man, most Insectivora, and many Rodents; nor can examples be found

where, as in the Cetacea, the smallness of the aperture is due rather to thickness and

over-development than to non-divergence of the crura. On the other hand, the inter-

mediate condition seen in this section of the Marsupials exists already in many mono-

delphous mammals, as in Priodon, Tolypeutes, and to a certain extent in Rhinoceros and

the Marmosets, as the author has already indicated. The families where this type of

stapes is found are the Macropodide and the Didelphyide.

Secondly the ossicula of those families will be described where the stapes is absolutely

columelliform, without any aperture or any indication of divergence of crura. Even this

still lower type is represented in higher mammals, namely in Manis. The stapedes of

SECOND SERIES.—ZOOLOGY, VOL, I. 38

A478 MR. A. H. G. DORAN ON THE MORPHOLOGY

many Cetaceans and Seals are in no way homologous with that of this section, for a reason

siven in the last paragraph. This group will include the remaining families of the

Didelphia—to wit, the Peramelide, Phalangistide, Dasyuride, and Phascolomyide.

Among the larger species in these groups a small aperture occasionally exists.

The malleus of an adult Kangaroo (Macropus major, Pl. LXIV. fig. 31) is of light con-

struction, though in no wise so thin and papery as in the Bandicoots. The head is small,

and only moderately developed, as in the majority of laminated mallei in the Monodelphia ;

though much flattened laterally, it is not broad in that sense, nor does it project in any

direction beyond the articular surface. This latter is placed more superiorly than behind,

as in most Hdentates, owing to the position of the incus. It has a raised and prominent

border; owing to the lateral flattening of the head, it is rather narrow, but deeply cut.

Its facets are of the character seen in most of the higher Mammalia, and hence differ

from the type of Man, Monkeys, and Manis. The upper or, rather, the anterior is wide

and almost flat; it is twisted a little more inwards than the lower, which is narrow and

convex; the groove between them is well marked. The neck is rather short and slightly

curved; the lamina is very narrow. The long and slender manubrium forms a right

angle (or even a few degrees less than 90°) with the neck ; the site of the processus brevis

is very obtuse ; and the sides are compressed, though not broad even near the root; there

is a distinctly bordered outer surface, as in Carnivora, Ruminants, and others, very

narrow, and but slightly spatulate near the tip, which is hardly perceptibly recurved. A

very slight trace of a tubercle at the junction of the neck with the root of the manubrium

represents the processus muscularis.

From the front projects the remarkable processus gracilis, already well known to ana-

tomists*. It is much longer than the manubrium, and curved downwards and back-

wards, and also somewhat outwards. It is attached by membrane (or, in some individuals,

actually ankylosed) to the tympanic ring, and is flattened and papery in character, rather

broader below than at the root, but ending in a point; near the body of the malleus it is

separated from the lamina by a thin ridge; and the narrow lamina itself is prolonged on

the anterior edge of the processus gracilis. The anterior part of the head of the bone is

not prolonged on to the front of the lamina, as in most laminated mallei; on the con-

trary, it is separated from the lamina by its very high border 7.

Such is the malleus of Macropus. Let us review its peculiarities, and compare them

with those of the same ossicle in higher mammals; for the more the latter possess cha-

racters in which the Macropodian malleus differs from that of the Primates, the lower,

in that respect, must such Monodelphia be considered.

In the lateral flattening and general form of the head the mallei of Carnivora and

Ruminants, and other Monodelphia where the lamina is well developed, all resemble the

malleus of Macropus; so does that of Myrmecophaga and Tamandua; only in the Ant-

* See Hyrtl, op. cit.; Peters, ‘Berichte der Berliner Academie,’ 1867; his conclusions in his paper for that year on

this subject were admitted to be incorrect by himself in an article in the same publication, 1868. See also, on the

latter, Huxley, Proc. Zool. Soc. 1869.

+ This high border is most marked on the external aspect of the malleus. It is very distinct in all the Kangaroos

and Wombats. (See Pl. LXIV. fig. 31.)

OF THE MAMMALIAN OSSICULA AUDITUS. 479

eaters just named the flattening is greater and the head itself larger. In the narrow

curved neck the same animals resemble the Kangaroos rather than Man and the higher

Monkeys, and differ in having a distinct neck from the Cebide. But as to the lamina,

which is absent in Homo and the Quadrumana and Lemures, it is well developed in the

small-headed malleus of a Carnivore or Ruminant, but ill developed, though existent, in

Macropus. A similar degree of partial development has already been noted in some

Edentata, as in Cholepus or Myrmecophaga; and it is to be remembered that the same

condition may be said to exist in Priodon, whilst Tolypeutes possesses a lamina as wide

as in many Carnivora; so that the latter genus differs from Priodon in one way, as

Dasypus does in a much higher direction, the head of the malleus being almost

anthropoid in development. On the other hand the lamina is again suppressed in Wanis,

and the articular surface is singularly like that of Homo, though the head is ill deve-

loped, and the stapes of a lower type than in Macropus, much lower than in Dasypus.

From the above facts, it appears to me to be justifiable to consider that the narrow-

necked and narrow-laminated type of malleus, as seen in Macropus, is the lowest form,

which has become developed, in higher mammals, into two higher forms :—one where

the lamina is more developed, as in the Carnivora; the other where the neck is stouter

or even merged into the head, as in Homo and the Quadrumana. The latter form

must be considered the highest, not only because it exists in Man, but also because

the former, or laminated type, most resembles the Kangaroo’s malleus in the form

of the head, and in the neck being equally long, narrow, and slightly curved; indeed,

in the very Orders where the laminated type prevails, the occasional partial or complete

suppression of the characteristic lamina is often associated with an increased develop-

ment of the head, and a greater stoutness or shortness of the neck, as I have already

observed in Herpestes and Suricata.

On the other hand, a form of malleus exists in other Monodelphia where not only are

the head, neck, and even the manubrium strikingly like the same in Macropus, and the

lamina of like small development, but the latter is prolonged upon a well-developed pro-

cessus gracilis in a similar manner. This is the case in Rhinoceros and Priodon.

Hence uot only is the WMacropus type of malleus the lowest of the three, but the lami-

nated type is clearly the second or the intermediate form, the large-headed and thick-

necked form being, therefore, unquestionably the highest; indeed it is almost as much

above the laminated as it is above the Macropoid variety. These observations suggest

an easy test of the grade of any mammal when judged by its malleus; and, as a rule,

the other ossicula agree with it.

In a young Halmaturus, measuring two inches without the tail, I have found the

malleus of similar form; but the head is much larger in proportion to the body than in

Macropus*; thé lamina is the same in character and similarly prolonged on the pro-

cessus gracilis.

Supposing, then, that both head and lamina, as existent in the young Halmaturus,

remain in a comparatively arrested state of development, the adult must necessarily

* The same, it is worth recording, is the case in the malleus of the adult Phalanger (Phalangista).

38 2

480 MR. A. H. G. DORAN ON THE MORPHOLOGY

have a malleus of the type of Macropus, Priodon, or Rhinoceros; let the head remain

undeveloped, and the lamina take an extra growth, then the laminated type will be seen

in the full-grown animal; but if the head and neck grow, they must necessarily encroach

on the lamina, and the adult will have a malleus like that of Man.

The foetal malleus of Dasypus or Tatusia, very Macropoid in character, undergoes the

latter change, whilst in Tolypeutes it grows into a laminated hammer, and in Priodon

remains unchanged in the adult. Almost similarly the mallei of the adult Rhinoceros

and Hquus differ, whilst that of the foetal Horse is remarkably like the adult Rhinoceros’s.

Again, the mallei of foetal Ruminants ultimately overshoot the latter animal in type,

but do not ultimately reach the high form of Hquus.

The incus of Macropus (Pl. LXIV. fig. 27) lies in a very similar position to that of

many Hdentates, the processus brevis being almost superior; and the stapedial crus is

directed rather backwards as well as inwards. The body is very small in proportion to

the crura, and deeply excavated by the articular surface. The processus brevis is stout

and blunt-pointed, rather long, and little divergent from the body. The processus longus

is long, and rather slender, it is not much curved, and supports a very distinct almost

circular Sylvian apophysis on a narrow pedicle ; altogether it more resembles the singular

ineus of Perameles than that of any Monodelphian mammal.

The stapes (figs. 27 & 42) has a small head and two slender crura united for halfway

towards the base, where they suddenly diverge, leaving a distinct triangular aperture ;

the outline of the crura is slightly concave. There is frequently a spur-like process for

the insertion of the stapedius-tendon, rather on the inner crus than on the head. The

base, which projects considerably beyond the insertion of the crura, is much more nearly

circular than in most placental mammals; the upper border particularly is very. much

curved.

All these characters are seen almost literally imitated in Tolypeuies. I deem it hardly

necessary to waste time in showing at length how this type is intermediate between the

bicrurate stapedes, with large apertures, as in Homo, and the columelliform type of the

Ornithodelphia; still less is there any occasion to demonstrate which is the lowest, and

which the intermediate form, that being seen at a glance if a Man’s, a Kangaroo’s, and

an Echidna’s stapedes be put in a row. The Macropoid type of stapes is approached, it

must be remembered, in animals as high as the Marmosets.

All the ossicula of other species of Macropus, as well as those of Hypsiprymnus and

Lagorchestes, resemble the same in M. major. The crura of the stapes divide in a

similar manner, and the base is broad and flat; nor has it the least tendency to become

bullate in the smaller Kangaroos, such as Hypsiprymnus murinus (Pl. LXIV. figs. 27,

28, 29, & 30).

The DipELPHYID# will now be described, as they resemble the Kangaroos in that the

stapes is not completely columelliform.

The malleus of the Opossum of the United States (Didelphys virginiana, P|. UX1V.fig.15)

is a little smaller than the same ossicle ina small Hypsiprymnus ; in its characters it closely

resembles that of Wacropus. The head is ill developed, and the articular surface deeply

cut, with facets of the same character as in the family just described. The neck is of the

OF THE MAMMALIAN OSSICULA AUDITUS. 481

same slender build; it is very straight until close to the manubrium, where it curves

suddenly. The lamina is decidedly wider than in any Kangaroo, and prolonged upon

the processus gracilis, forming a curved process applied, as in other marsupials, to the

tympanic ring very closely, though readily detached from the latter even in the oldest

male specimens, without the aid either of a lens or of any great skill in dissection. This

process is of more regular form in Didelphys than in Macropus, being of nearly equal

breadth throughout except at the extremity, which is slightly dilated. No foramen can

be detected in it, as in Hrinaceus, which possesses a somewhat similar but still more

uniformly broad processus gracilis.

The manubrium is even more slender and straight than in Macropus; but at the outer

angle of its base, the site of the processus brevis in Homo, it is peculiarly broad and flat-

tened, projecting towards the neck on the outer aspect. ‘The extremity is almost styli-

form; still there is a distinctly bordered though extremely narrow outer surface*, which

may be seen by a magnifying glass to be slightly dilated towards the tip. This malleus

may be said to be a trifle higher in type than that of the Kangaroo, exhibiting certain

affinities to some Insectivora.

The incus of the Virginian Opossum has a deeper body than in the Kangaroo, with the

articular surface as deeply cut ; the processus brevis is thin, long, and rather divergent

from the body, lying almost vertically above the latter in the recent skeleton. The pro-

cessus longus is very long, and bears at its extremity a circular Sylvian apophysis on a

very long pedunele, which is broad, and fiattened where it joins the extremity of the

apophysis +. A high development of this process is frequent among the Insectivora; but

it is also seen in other families of the Marsupialia, where the peduncle is of similar form.

The stapes has a small head, sometimes bearing a tubercle for the stapedius-tendon.

The slender crura diverge rather higher than in Macropus, so that the aperture is more

distinct and more triangular. The base, which is thin and flat, projects far beyond the

erura, is more oval, and hence not so deep vertically as in the Kangaroo (Pl. LXIV.

figs. 15 & 41).

The stapes of Didelphys is certainly of a higher type than that of Macropus or any

other marsupial, both in the greater divergence of the crura and in the characters of the

base. It much resembles that of some small American Monkeys and of certain Rodents

and Insectivora. Classifying the different types of stapes on the same principle upon

which the mammalian malleus was classified after describing that ossicle in Macropus,

it may be asserted that we find two extreme types:—on the one hand the stapes of

Homo, the Simiidee, and some Rodents (as Arctomys) and Insectivora {, where the crura

are freely divergent and curved with a large aperture; and on the other hand the

completely columelliform stapes of the Ornithodelphia, Phalangistide, and Manidee.

* See the description of the ossicula of the Carnivora, anted.

+ This apophysis is, of necessity, very easily broken off in preparing the ossicula for anatomical observation.

+ The stapes in these latter and clearly inferior animals has a much less-developed head than in Homo; and the

wideness of the aperture is necessary for the transmission of a bony canal. It may be said to constitute another

and a rather high type.

482 MR. A. H. G. DORAN ON THE MORPHOLOGY

Between these extremes will be found :—first, the stapes with straight crura and a

moderate aperture, as in the Carnivora and Muride, including the Cetacean form,

where the opening is filled more or less by the thickness of the crura; then the stapes,

with thin straight crura and a very small aperture, as in Bradypus; and lastly the

variety seen in Midas, Didelphys, Tolypeutes, and Macropus, where the crura are fused

for a considerable way from the head, diverging near the base, the separation beginning

much earlier in the first two than in the last two genera just named, so that the former

possess a subvariety of higher type; from this last we find examples, in several Orders,

of insensible gradations towards the still higher straight-limbed type, with a free aper-

ture, as in the Carnivora.

In Didelphys opossum all the ossicula are of very similar character; the lamina of the

malleus is even wider; and there is a little tubercle on the inner side of the neck,

representing the processus muscularis, very feebly indicated in D. virginiana. In both

species the processus gracilis is held in a chink between two forks of the tympanic ring

during lifetime, as in most of the Marsupialia. |

In the remaining families of marsupials the divergence of the crurais either completely

absent in the stapes or exceptional and inconstant.

In the ossicula of the Tasmanian Wolf (Thylacinus cynocephalus) (P1. LXIV. fig. 17),

so like a Canis in its external appearance, there is hardly one point to be found in which

they resemble the ear-bones of a dog.

The head is very ill-developed, though not much flattened laterally, and distinctly

convex upwards. ‘The articular surface is deeply cut, with prominent margins; the lower

facet is very narrow. The neck is very thin, long, and narrow, forming a very prominent

humplike bend midway between the head and the handle. The lamina is very broad and

prolonged upon the processus gracilis, to form with it an extremely long process, which

is very narrow, except near the club-shaped, curved, and flattened extremity. The pro-

cessus muscularis is indicated by a very faint elevation on the inner side of the neck.

The manubrium is slender, and very similar in character to that of Macropus*.

Altogether, even in the character of the lamina, the malleus of the Tasmanian Wolf

more resembles that of certain Insectivora than that of any Carnivorous mammal.

The incus has an ill-developed body with a very short processus brevis ; the stapedial

crus is long and very divergent; it bears a small elliptical Sylvian apophysis on a rather

long pedicle.

-The stapes (figs. 17 & 43) is absolutely columelliform. The head is very small, and

the column long, and rather broader near the base; there is not a vestige of an aperture.

The base is wide horizontally, but shallow vertically, with the upper border well curved,

contrasting with the almost circular base of the Ornithodelphia and Sauropsida, and

constituting a higher type of the columelliform stapes, in that the footplate assumes the

same form as is observed in the higher mammals, where that ossicle is bicrurate.

* The front and lower part of the head ends in a point, ‘and is not prolonged on to the lamina joining the pro-

cessus gracilis in the Carnivora &c, With regard to the manubrium in Dasywrus, consult my observations on Myr-

mecobius further on.

OF THE MAMMALIAN OSSICULA AUDITUS. 483

In the smaller species of Dasyures, as, for instance, Dasyurus maugei (Pl. LXIV.

fig. 16), the malleus is of very similar character to that of Thylacinus, though the neck

is not so boldly curved, and the lamina is narrower. The ineus and stapes are identical

with the corresponding ossicles in the Tasmanian Wolf. Not a trace of any aperture

exists in the stapes.

In Dasyurus ursimus the malleus has a larger and rounder head, a much narrower

lamina, and, in particular, a much stouter and stronger manubrium, than in the above-

~ described species; its sides are of considerable breadth near the root. The processus

longus of the incus bears a small Sylvian apophysis on an extremely thin and fragile

pedicle. The stapes is absolutely columelliform. ‘The two inner ossicles are smaller in

an adult D. ursinus than in a full-grown Thylacinus; but the malleus of the former is of

nearly the same size and of more solid make than in the Tasmanian Wolf.

The malleus from a small Phascogale (Pl. LXIV. fig. 18) in the collection resembles

that of Dasywrus in every respect; the lamina is wide. The processus brevis of the

ineus is very short; the stapedial crus bears a similar but proportionally larger Sylvian

apophysis compared with the same in Thylacinus. The little stapes is columelliform

without an aperture; the stapedius-muscle attached to it is very large, as in most of these

animals.

In Uyrmecobius the malleus is of the same light build as in the Thylacine. The head

projects considerably above and in front of the lamina. The neck is very long, but

not niore curved than in Dasywrus. The lamina is as wide and smooth as. in the Tas-

manian Wolf, and hence more extensive than in the Dasyure and Phascogale ; it joms the

processus gracilis to form the usual long curved process applied to the tympanic ring.

The manubrium is of the same form as in the other Dasyures ; but its outer angle (which

in this family is generally sharper than in the Kangaroos) is conspicuously bent out-

wards so as to form a genuine processus brevis. The incus has an extremely short pro-

cessus brevis, a deeply cut articular surface as in other Dasyures, and a distinct Sylvian

apophysis on a long pedicle. The stapes is perfectly columelliform.

Among the PHaLANGIsTIDz the malleus of Phalangista vulpina (Pl. LXIV. fig. 20)

may be selected as a typical example. The head, though small, is better developed than

in the Dasyures; and though it does not rise high above and in front of the facets, its

upper surface is wide antero-posteriorly, projecting backwards over the articular surface.

This latter is rather deeper cut than in the Dasyures; and the upper facet is very wide,

especially vertically. The neck is rather short, but curved, the lamina rather narrow ;

and the processus gracilis is of equal or almost equal breadth throughout, so as to look

like a curved piece of steel ribbon. This is much more like the form of that process

in the Didelphyidz than in the Dasyures. The manubrium is almost straight, and is

slender and little recurved at the tip, as in the Kangaroos; the angle corresponding to

the processus brevis is rather obtuse.

A notable feature in this family is the large size of the incus. In Phalangista vulpina

it is decidedly slightly larger than in Macropus major, and larger than in any Dasyure,

nearly twice the size, in fact, of the same ossicle in Thylacinus; yet the malleus of au

adult Vulpine Phalanger is hardly larger than that of Dasyurus maug@i, where the incus

ASA MR. A. H. G. DORAN ON THE MORPHOLOGY

is hardly ever one third the size. The body of the incus of P. vulpina is well developed

and not much encroached upon by the articular surface, which has well-defined edges.

The processus brevis is stout and blunt-pointed ; it diverges little from the body. The

processus longus is long, rather straight and divergent; it bears a small pedunculated

Sylvian apophysis. |

The stapes in large well-formed skulls of this species has a small aperture as in

Macropus; its base is much longer horizontally than vertically ; and there is a small umbo

on the centre of the base.

In Phalangista cookii the distinctive characters above given are still better marked.

The upper part of the head is well developed in the malleus, projecting well backwards

over the articular facets. The lamina is very narrow, but produced on to the processus

gracilis; they together form a curved ribbon-like process of very even breadth through-

out. Theincus is as large and well-developed a bone as in P. vulpina. The stapes

(Pl. LXIV. fig. 21) is most remarkable for a character already noted by Hyrtl(/.c.): the

crura are united throughout without a trace of any aperture, forming a stout pillar, much

shorter than in any other Marsupial ; and the base is inflated into a large bulla projecting

prominently into the vestibule. I have not found this peculiar feature in any other

Phalanger; and since Hyrtl represents it precisely as it appears in a specimen recently

procured from a Cook’s Phalanger preserved in spirit in the store-room of the College

of Surgeons, it may be concluded that the bulla is a constant feature in this species.

Its morphological value does not appear to be great, as a bullate stapes occurs spo-

radically, so to speak, among higher mammals—for instance, in animals not closely allied

to each other, such as Mustela, Geomys, and Hyraz.

In Petawrus (Pl. LXIV. fig. 22) the malleus has even still more pronounced features

than in Phalangista cookii. The head is markedly of the form already described in

Phalangista vulpina; the neck is shorter, and the lamina rather wider; but in the

Flying Phalanger the manubrium is of stouter make in the above-described species, and

in the larger species (as in the fine and typical malleus of the Taguan, Petawrus

taguanoides, Pl. LXIV. fig. 23) it ends in a very marked spatulate dilatation *.

The incus of Petaurus is a large bone with similar characters to that of Phalangista.

The stapes often has a distinct perforation, though otherwise it is columelliform; the

base is much wider horizontally than vertically.

In the characters of the malleus and of the large incus, the Flying Phalangers show

certain affinities to the Sciuromorphic Rodents.

In the Koala (Phascolarctos fuscus, Pl. LXIV. figs. 24, 34) the malleus does not

possess the distinctive features so well marked in other Phalangers.

The head is very small and ill-developed, projecting little either upwards or backwards ;

it is not much flattened laterally ; the articular-surface is rather wide; and the upper

facet is planer but much larger than the lower. The neck is long, thin, and curved, the

lamina rather narrow ; and the processus gracilis forms with it a similar process to that

in Phalangista, but more prolonged. The manubrium presents a great peculiarity

* This dilatation is marked in Phalangista cookii, though the end of the manubrium is almost styliform in

P. vulpina.

OF THE MAMMALIAN OSSICULA AUDITUS. 485

(fig. 34): it is rather broad at the base and flattened laterally, very slightly curved,

and dilated at the tip; but above, at the site of the processus brevis, it is not only

broad, but presents a distinct concavity; the raised margins (of which the outer appears

continued from the neck), very wide apart above, almost coalesce below, but keep slightly

apart towards the extremity, forming a very narrow outer surface to the manubrium *.

The incus of the Koala is smaller in proportion to the malleus than in the Phalangista;

and the processus brevis is very short and ill-developed ; the stapedial crus bears a pedun-

culated Sylvian apophysis.

The stapes is absolutely columelliform, without any trace of an aperture; it bears a

small bony process for the attachment of the stapedial tendon.

Hence in all three ossicula Phascolarctos exhibits characters in which it differs from

Phalangista, and resembles rather more the majority of Marsupials.

In the Wombat (Phascolomys vombatus, Pl. LXIV. tigs. 32, 33) the most interesting

feature is the enormous development of the processus gracilis of the malleus.

The head of the malleus is even less developed and more compressed laterally than in

the Dasyuridz ; it is much encroached upon by the articular surface, which is both

deeply cut and wide; the lower facet, though more convex, is not much narrower than

the upper, from which it is divided by a well-marked groove. The neck is thin and

curved, as in the Kangaroos and Dasyures; and the lamina is narrow. The manubrium,

as in most Marsupials, is slender, and forms little more than a right angle with the

neck. The outer aspect of the base is very broad, though not so modified as in the

Koala. The extremity of the handle is narrow but distinctly spatulate. The processus

gracilis, joined by the narrow lamina, forms a curious process three times as long as

the manubrium, which is closely applied to the tympanic bone, and can readily be

detached from it even in the adult. At first it is narrow and almost of as even breadth -

as in the Phalangers; but its distal two thirds form a broad sabre-like or leaf-like

process, sharp-edged, much flattened, and ending in a tapering point. The whole pro-

cessus gracilis is only an extremely developed form of the type seen in the Rhinoceros

and in many of the Insectivora and Marsupials. The foliaceous extremity is readily

broken off if the specimen be not carefully prepared.

In the incus the body is fairly developed, especially above the processus brevis, though

the whole bone is not so large in proportion to the malleus as in Phalangista. The pro-

cessus brevis is rather short, but stout at the base; it diverges but little from the body.

The stapedial crus is long and thin, and supports a large elliptical Sylvian apophysis on

a slender pedicle. The stapes is quite columelliform ; the head forms a narrow elliptical

capital to the fused crura; just beneath it is a spur-like process, to which the tendon of

‘the stapedius may be found attached in the recent or even in the imperfectly macerated

‘skull. The base is not of the Sauropsidan character, being as narrow vertically as

in Homo.

The ossicula of P. latifrons are very similar in character. They are all rather larger in

the Wombats than in the largest Dasyures (such as D. wrsimus) or Thylacinus.

* The blunt outer aspect of the base of the manubrium in the mallei of large. Dasyures and other Marsupials

exhibits an approach to the above-described peculiarity in the Koala. ~

SECOND SERIES.—ZOOLOGY, VOL. I. 3T

486 MR. A. H. G. DORAN ON THE MORPHOLOGY

In the PrramMeLip# the malleus is of light and fragile construction, although wholly

osseous and not very different in character from the same ossicle in many older Mar-

supials (Pl. LXIV. fig. 25).

The head is very small, with a deeply cut articular surface, which has two facets, the

upper and anterior being the widest and plainest of the two, The neck is long and

thin; it forms, by suddenly bending on itself midway between the head and manubrium,

asharp projecting angle, as in the equally fragile malleus of the Soricide. The slender

manubrium is rather broad and flattened at the base, and is blunt externally at the point

corresponding to the processus brevis. It ends in an almost styliform extremity. The

lamina is narrow, and joins the processus gracilis to form a curved process slightly broad

near the tip, as in the Dasyuride ; this is fitted into the tympanic bone as in other

Marsupials.

The incus of Perameles has a rather high and narrow body. ‘The processus brevis is

rather long and thin, and not divergent from the body. ‘The processus longus is thin,

long, and very divergent ; it ends in a very conspicuous and perfectly elliptical Sylvian

apophysis, supported on a pedicle which is extremely thin at its attachment to the

stapedia] crus, but becomes broad and flattened before it ends in the extremity of the

apophysis. This is but an extreme development of the condition so frequent in the

Didelphia.

The stapes is completely columelliform ; and the column isas short as in the Passerine

birds; the base, too, is rather wide vertically. This ossicle is generally imperfectly

ossified even in the adult Bandicoot.

In Cheropus all the ossicula are of the type seen in Perameles (Pl. LXIV. fig. 26).

All the ear-bones of this family must be considered as of a very low type, the fragile

. malleus of the adult recalling that ossicle in the foetus of higher mammals. The ill-

development of the processus brevis incudis and the persistent distinct character of the

Sylvian apophysis, as well as the perfectly columelliform stapes, combine to force upon

the observer the conclusion that, in respect of the ossicula, the Peramelide are the

lowest of the Didelphia; for in no other family are the little bones to be found all

three so thoroughly of low development in every detail.

In concluding this review of the Ossicula of the Marsupialia, it may be said, by way of

a summary :—

I. That the ear-bones in this subclass are always of a low type, yet not as a rule so

inferior to those of the Monodelphia as might be inferred, judging from the rest of

their skeletons.

II. That no point showing a low grade of development in any ossicle in any Mar-

supial is not occasionally met with in higher mammals, excepting that in the latter the

ear-bones are never seen to retain to so high a degree the foetal consistence as may be

observed in the adult Perameles.

III. That the Bandicoots or Peramelidz are, in respect of their ossicula, the lowest

family in the whole subclass, both in the type of all three bones, and in their frail

papery consistence compared with the rest of the cranium.

OF THE MAMMALIAN OSSICULA AUDI'TUS. 487

IV. That in the same respect the American Opossums or Didelphyide must be con-

sidered the highest of the Marsupials. The malleus is, in type and consistence, similar

to that of many Insectivora; the incus has the body and short crus as well as the pro-

cessus longus well developed; and the stapes is more perfectly bicrurate than in any other

allied family.

V. The Macropodidee are very central among Marsupials in their ossicula. The

malleus is of low type, but tolerably firm osseous consistence; it bears the peculiar

highly-developed foliaceous processus gracilis aiso seen in the Wombats; the incus is

fairly developed; and the stapes is always partially bicrurate.

VI. The Phalangistidee come next in order. They are decidedly inferior to the

Kangaroos in that the stapes is generally (though not invariably) columelliform; more-

over the malleus is of the form seen in a new-born Macropus; but in the size and

good development of all parts of the incus they are as high or even higher than the

American Opossums. Phascolarctos is lower in all respects than Phalangista; its

malleus is of a very distinctive character.

VII. The Wombats are slightly lower than the last family, the stapes being always

columelliform, and the ineus having a rather short processus brevis, but a stapedial

crus rather of the Perameles type. The malleus is not so much of high type as an

extreme form of the specialized corresponding ossicle in Macropus, the processus gracilis

being very large, curved, and flattened.

VIII. The Dasyures have the stapes invariably columelliform, and the incus of low

type; but all their ossicles are of more solid consistence than in the Bandicoots

(Peramelidee).

THe OssIcuLA OF THE MoNOTREMATA.

Next to the ear-bone of Man, those of Hchidna and Ornithorhynchus are of all other

mammalian ossicula the best known to the anatomist. From the conformation of the

base of the skull in the Monotremata, the malleus cannot fail to obtrude itself on the

glance of the student cursorily examining the cranium. ‘Then these little bones are of

profound interest to the morphologist when studying this subclass, which is so un-

-doubtedly allied in many respects to the Sauropsida, where two out of three ossicula

are suppressed. Good general descriptions of the ear-bones of the Monotremata, with

clear figures, are to be found in the works of Huxley *, Hyrtl +, Peters ¢, and others.

I leave such important questions as What bones are the homologues of the mam-

malian malleus and incus in the Sauropsida? to distinguished authorities in the united

branches of morphology and embryology. Next, I need hardly remind the anatomist

that, as far as the ossicles are concerned, the Monotremata wear a perfectly mammalian

uniform, having malleus, incus, and stapes. If any theorist consider this Order more

Sauropsidan than mammalian, on account of the most marked feature in the first two

* Proc. Zool. Soc. 1869; see woodcut, p. 404. t+ Op. cit.

+ “ Ueber das os tympanicum und die Gehérknochelchen der Schnabelthiere &c.” Monatsberichte der Akad. der

Wissensch. Berlin, 1867.

3T 2

488 MR. A. H. G. DORAN ON THE MORPHOLOGY

bones, he must, for the same reason, place the Porcupines and Cavies nearly as low in

the scale, and decidedly in an inferior position to the Marsupials; and as for the stapes, it

is of a type precisely imitated in the placental Manis.

It remains for me to describe the ear-bones of this Order as I have detailed the same

in higher milk-secreting vertebrata, investigating carefully their exact homologues in

the former.

In Echidna hystrix the two outer ossicula are proportionally much larger, better

developed, and more firmly osseous than in Ornithorhynchus. It is on that account

advisable to commence with the description of the ear-bones of the so-called Porcupine

Ant-eater.

Let the united malleus and incus of an Hchidna be removed from the skull. In so

doing observe that the processus brevis of the incus lies anterior as well as superior to

the stapedial crus in the natural condition, an exaggeration of what is seen in the

Sirenia, Marsupials, and other mammals. Even in Homo the short crus is almost as

superior as it is posterior to the rest of the incus. Observe also, if the skuil be im-

perfectly macerated, how large the tensor tympani muscle is, as Professor Huxley *

has already remarked. Were it attached to the inner edge of the manubrium, as in

the Rodents, it would require still greater care and preparation, to remove the malleus

entire, than even is necessary in those animals. Now let the united malleus and incus

be examined from the inner aspect (Pl. LXIV. fig. 36), and the complete outline of the

incus may plainly be seen. The inner aspect of its body is free, square, and faintly.

concave, whilst antero-inferiorly it is limited by a convex border fitting into a concavity

in the malleus. The whole, at this region, has precisely the same appearance as may be

seen in articulating the incus and maileus in a Marsupial, or in any mammal where the

latter ossicle has a small ill-developed head.

But view the bones from without (fig. 35), and it will then be observed that the

malleus encroaches on the whole of the outer surface of the body of the incus, so that

only the processus brevis (pd) and the tip of the processus longus (pl) can be seen. As

it covers the greater part of the long process, on turning to the inner aspect of the

ossicles the singular appearance of a lamina of bone, reflected from the outer border of

the stapedial crus towards the manubrium, is at once accounted for, that lamina being

simply an integral part of the malleus appearing from behind at the point where, from

this aspect, the incus ceases to cover it.

‘Hence the incus is articulated to the malleus by its whole inner surface, including the

greater part of its long crus, and also by its convex inferior border. It is firmly anky-

losed to the malleus; and even in a young skull the ossicles cannot be separated unin-

jured without difficulty.

Bearing in mind the above remarks, the description of the malleus by itself will now

be more comprehensible. The head is narrow and concave; externally it is prolonged

upwards into a thin scale (Pl. LXIV. fig. 35, shm), of which the wide convex free

border reaches to the base of the short crus, and conceals still more of the long process

* Proc. Zool. Soc. 1869, “« On the Representative of the Malleus and Incus of the Mammalia in the Sawropsida.”

OF THE MAMMALIAN OSSICULA AUDITUS. 489

of the ineus. This scale hides the concave excavation of the head, which, on the other

hand, is plainly visible from the inner aspect of the united bones (fig. 36, hm).

From the inner aspect, too, it may be seen that the head sends forward a long, pro-

minent, and very straight process to join the processus gracilis *. The analogue of this is

already familiar to us in most laminated and in other mallei. The neck is rather long,

and not strongly curved; its free border appears from in front of the incus when the

' bones are viewed from within, as just described; in the opposite direction it blends

insensibly with that part of the bone representing the lamina in higher mammals.

- From the outer aspect the neck is more clearly defined. Close to the root of the

manubrium a small tubercle rises from the inner side of the neck, to which the tensor

tympani muscle will be found attached in imperfectly macerated specimens. Hence

this tubercle is the processus muscularis.

The manubrium of Lchidna is of a type very frequent among the lower mammalia.

It is of moderate length, compressed laterally, but not so thin and broad at the root as

in some Rodents. The outer angle of its base (the site of the processus brevis, which is

absent) is very blunt, broad, and well rounded; the outer aspect is very narrow, though

sharply bordered from the sides, and ending in a narrow but distinct spatulate dilata-

tion. The entire process, in slenderness, bluntness of its outer angle, &c., recalls the

same in Rhinoceros, Priodon, and the Marsupials. It cannot be said to bear any clese

resemblance to any process of the os quadratum or any neighbouring bone in the

Sauropsida.

The processus gracilis is readily distinguished, as it starts off from the malleus, forms

a deep curve, and joins the process from the head, to form the long straight splinter of

bone which articulates with the tympanic. This condition is well known; but anatomists

hitherto have described the united part as if it were solely and entirely the processus

gracilis, ignoring the process from the head, which I have frequently described in

animals where it is well marked, notably in Fiber. Whilst this latter process, as above _

stated, is traced along the united portion from the inner aspect, the continuation of the

processus folianus proper must be looked for in viewing the malleus from without; then

it can be seen to terminate in a broad lamellar expansion, on which the process from the

head loses itself posteriorly.

Between the neck and the free curved portion of the processus gracilis (and hence best

seen from the outer aspect of the ossicle) the analogue of the lamina so frequent in the

higher mammals may be distinguished : it is neitWer broad nor thin, and runs on towards

the united processes, with which it blends insensibly. This, it may be remembered, is the

same as in the Marsupials.

The body of the incus has already been referred to, and the free inner aspect described,

as well as the manner in which it is wholly covered externally by the malleus+. The

processus brevis is short and flattened extero-internally; it projects from the outer

* See Pl. LXIV. fig. 36. The proccss from the head (phm) is here distinguished from the processus

gracilis (pq).

+ Excepting its anterior edge, which is just left uncovered.

490 MR. A. H. G. DORAN ON THE MORPHOLOGY

aspect above the scale-like process from the head of the malleus, which covers the

body. The processus longus is not much longer than the short crus; it is straight and

flattened, and wholly concealed from without by the malleus, except its extremity, which

is bent inwards, and bears a very small but distinct Sylvian apophysis.

It will be seen at a glance that the kind of ankylosis between the two outer ossi-

cula in Echidna is totally different from their mode of union in Porcupines and Cavies,

where the malleus and incus are joined on the same level externally as internally ;

and, besides, there is always in the latter animals a peculiar development of the head

of the malleus forwards, away from the incus, instead of the scale-like prolongation of

the head on to the outer side of the latter ossicle. .

The incus of Echidna is, after all, not so very much smaller in proportion to the

malleus than in many Marsupials; only it is more flattened; flatness is also seen in the

same ossicle in those Rodents where it is fused with the malleus. I have never found

it to be so loosely attached to the malleus as in young skulls of the Duck-bill.

The stapes is absolutely columelliform. Its head is very small, with no trace of any

tubercle for the stapedius-tendon, which does not appear to exist in this animal*. The

column bears not a trace of any subdivision into crura; it is rather long, and the foot-

plate is almost perfectly circular.

What relation does the stapes of this avowedly low mammal bear to the columelliform

representative of the same in higher animals on one hand and in the Sauropsida on the

other? Is the column of the stapes of Hehidna formed of the same element or elements

as in Aves ?

By referring to the description of the stapes in the Marsupialia, it will be seen that

this bone is of the same perfectly columelliform type in the Dasyures, Wombats, Bandi-

coots, and most Phalangers, as in the Echidna; and among the Edentata, Manis has a

similar type of stapes. In some of the above animals the base of the ossicle in question

- is less perfectly circular than in Echidna, and hence more like the foot-plate of the

bicrurate stapes of Homo. Then the column is shorter in some of these animals than in

the Porcupine Ant-eater.

But it happens that all the above characters rather strengthen the alliance between

Marsupials, the Manide, and the Monotremata, than show that those animals are parti-

eularly ornithic or reptilian.

In some birds where the column is almost as short as in these mammals, it divides

distinctly into two crura, as distinctly as in Macropus and Tolypeutes. This is the

case with the Nightingale (Pl. LXIV. fig. 45) and Redstart; or at least it is so in the

specimens in the College collection. In others, where the column is very long, it may

also divide into two crura near the foot-plate, as is very clear in the Hagle (Aquila,

Pl. LXIV. fig. 46). Besides, the foot-plate in many Finches and other birds, and in

Crocodiles and other reptiles, is not of the circular outline seen in the Echidna, but as

long horizontally, and narrow vertically, as in the stapes of the highest mammals.

Then, without plunging into unsettled questions of development, does the column of

* Huxley. I have never seen a vestige of any representative of that muscle.

OF THE MAMMALIAN OSSICULA AUDITUS. 491

the stapes of these inferior mammals represent the same or only a part of the elements

making up the column of a bird’s columella ?

In all these mammals we have an incus with a processus longus and Sylvian apo-

physis articulating with the stapes, which is columelliform; but the column is clearly

identical with the two crura of the higher Mammalia, as the transitional condition

seen in Macropus &c. clearly proves.

In birds the column of their stapes or “columella” is certainly the same, whether

short (as in the Finches, Pl. LXIV. fig. 44) or very long (as in the Hagle, Albatross, &c.,

figs. 46, 47); for the same “ extrastapedial” and “suprastapedial” elements are found

attached to its outer extremity in the same manner in both varieties. But at least part

of the column is recognized by the latest investigators into embryology * as the “ medio-

stapedial’’ element which represents the long crus of the incus in mammals, and

begins as the proximal incurved extremity of the hyoid arch in both the Mammalia

and the Sauropsida.

Now, this long crus of the incus always existing in mammals, it follows, if the most

advanced embryologists have not erred, that the column of the Marsupial and Mono-

trematan stapes (as well as that of Manis) represents only a part of the columella in

Aves. It is evidently formed entirely from the mass of cells which grow from the

periotic capsule to meet the proximal end of the hyoidean arch (Parker, J. c.) ; for since

that end becomes the processus longus of the incus and its Sylvian apophysis in these

mammals (identical with the mediostapedial element in birds), no segment of that

arch can take any share in the formation of their stapes, as if is seen to do in the

Sauropsida.

In examining the ear-bones of Ornithorhynchus paradoxus, the first thing that strikes

the observer is the relatively small size of the malleus and incus compared with those bones

in Hehidna, and.their imperfectly ossified semimembranous texture, in which they exceed

those of any other mammal. The stapes in the Duck-bill is nearly as large as in the

Porcupine Ant-eater (Pl. LXIV. fig. 37).

If the united malleus and incus of Ornithorhynchus be more closely investigated, it

will be found that in ossicles from young skulls they may be readily disarticulated ;

but in specimens from the adult this is not invariably the case. Professor Peters +

asserts that these bones remain free throughout life in this animal; but in a set which

I prepared for the College collection, from a full-grown skull, the malleus and incus

remained ankylosed on the right side. ee

The manner of union between these two ossicula is precisely the same as in Hehidna.

The incus is seen from the inner aspect (Pl. LXIV. fig. 38), but almost completely

concealed in viewing the malleus from without (fig. 37) {.

The portion of the malleus representing the head in higher mammals (fm) is ex-

* Consult summary at the end of Professor Parker’s memoir “ On the Structure and Development of the Skull of

the Pig,” Phil. Trans. 1873.

+ Loe. cit.

+ In Pl. LXIV. figs. 37 & 38, the same abbreviations are used as in the figures of the corresponding ossicles

in Echidna.

492, MR. A. H. G. DORAN ON THE MORPHOLOGY

tremely small; and hence the greater narrowness of the malleo-incudal articulation, as

seen from within, compared with Echidna. But it sends up from without a scale-like

process (sim), as in that animal, which conceals the body of the incus and the root of

its long process; so that on viewing the bones from within, the same appearance of a

lamina of bone, reflected from that process to the malleus, may be noted. When the

incus can be detached, a smooth slightly concave space on the process from the head

comes into view. This has been already figured in Peters’s memoir already quoted.

From the head a very stout and straight process is sent forwards, blending with the

very thin papery processus gracilis. The neck is longer, and much better defined than

in Lchidna, and forms a very wide curve. The true processus folianus is merely the

slightly thickened free border of the narrow but distinct lamina; it joins the well-deve-

loped process from the head. It is only by a careful comparison of these elements in

Ornithorhynchus with the same in laminated mallei of placental mammals that the error

can be avoided of taking the process from the head for the processus gracilis *.

The processus muscularis is of a unique form. It is a wide angular projection from

the inner side of the neck above the manubrium. The processus gracilis (and its adjunct

just referred to) is united with the tympanic bone in the same manner as in Hehidna.

This bone is here less developed and more sharply curved than in that animal. Hence,

united with the malleus, it bears a very characteristic distinctive appearance.

The manubrium is short, ill developed, and so imperfectly ossified, as to shrink when the

malleus is dried. Its outer angle is blunt, very broad, and slightly concave, somewhat

as in Phascolarctos. Its shank is very slender, and slightly dilated at the extremity ;

the outer aspect is excessively narrow, but distinctly bordered from the sides.

The incus is small and flattened, as in Hehidna. Its inner surface is more concave, and

narrower where it joins the malleus. The crura are short, stout, and straight, and almost

of equal length; the stapedial crus bears a very faint indication of a Sylvian apophysis.

The stapes is absolutely columelliform, and not so firmly osseous as in Hehidna ; the

column is rather longer, the head broader, and the almost circular footplate is not

quite as large. As a processus longus of the incus exists, the column must be the

exact homologue of the same in Echidna, and of the two crura of the stapes of the

higher mammals, and, for the same reason, represents only part of the column of the

Sauropsidan columella. In fact the presence of the processus longus of the incus throws

the stapes of the Monotremata both towards other mammals, and away from the birds.

The result of the above considerations shows :—

1. That, in their ossicula, the Monotremata are as low as in the rest of their skeleton,

yet, as might be expected, thoroughly mammalian.

2. That their distinguishing features are a peculiar form of articulation between the

malleus and incus by means of a scale-like development from the head of the former,

and the presence of an absolutely columelliform or unicrurate stapes. Otherwise the

three ossicula are not much modified from their representatives in the lower Marsupials.

* Compare phm and pg in Pl. LXIV. figs. 36 & 38, with the same in Pl. LXIII. fig. 14.

OF THE MAMMALIAN OSSICULA AUDITUS. 493

3. That a definite incus has long been admited to exist, articulated, as above noticed,

with the malleus, and ankylosed in the Hehidna, and generally in the adult Ornitho-

rhynchus. The method of ankylosis is quite different from that seen in certain Rodents.

4. That as a processus longus of the incus exists, and hence in all probability is the

proximal end of the second postoral arch, it follows that the column of the stapes of

these animals (and of all other mammals where it is of similar shape) represents the two

crura of the stapes of higher mammalia, but only a part of the column of the Sauropsidan

columella, which is generally admitted to be formed partly from the ‘‘ mediostapedial ”’

element of the hyoid arch. But a further careful study of the embryo of these animals

is necessary to settle this point otherwise than by analogy.

5. That the malleus and ineus of Ornithorhynchus differ from those of Echidna in

matters of detail, but in no essential features which might ally either animal more or

less than the other to higher mammals or to the Sauropsida, excepting, some might say,

the semimembranous consistence of the malleus of the Duck-bill.

DESCRIPTION OF THE PLATES.

Pirate LVIII.

Throughout the Plates the deft malleus and incus are represented as viewed from the inner side, that

position being the most convenient for displaying the homologous elements. The “ natural position,”

z.e. that in which the manubrium of the malleus is turned towards the observer, is objectionable, because

foreshortening of the ossicles interferes with a ready comparison of their various processes and relative

dimensions as seen in different animals. The ossicula are enlarged 8 times natural size, unless otherwise

specified. The letters used im the various figures bear the following signification :—as, articular surface

of malleus; , neck ; pb, processus brevis ; pg, processus gracilis; a, a*, upper facet, and 6, b*, lower

facet of malleus; sr, sigmoid ridge of malleus ; emn, external surface of manubrium; pl, processus

longus ; ¢, concavity head of malleus ; eas, extra articulating surface on malleus and incus ; gr, groove

on incus; an, angular projection on neck of malleus; oa, orbicular apophysis ; mall, malleus; Im, fig. 22,

Pl. LX., in error for pm; pm, processus muscularis; hm, head of malleus (fig. 1, Pl. LXIII., lettered

wrongly for mn, see footnote p. 458) ; pg, processus gracilis; J, lamina; phm, process head of malleus ;

acpm, accessory processus muscularis ; mn, manubrium; pet, petrous bone; 7b, tympanic bone; shim,

scale head of malleus ; ms, medio-stapedial element.

Figs. 1-17. Left ossicula: 1,2, Homo*; 8, Troglodytes gorilla; 4, Troglodytes niger ; 5, Simia satyrus ;

6, Hylobates syndactylus ; 7, Semnopithecus entellus ; 8, Colobus satanas; 9, Cercopithecus cephus ;

10, Macacus erythreus ; 11, Cynopithecus niger ; 12, Cynocephalus anubis ; 18, Ateles melanochir ;

14, Mycetes ursinus ; 15, Cebus; 16, Nyctipithecus trivirgatus; 17, Saimaris sciurea.

Fig. 18. Left malleus and incus: Pithecia satanas.

Figs. 19-22. Left ossicula: 19, Hapale jacchus; 20, Midas edipus ; 21, Lemur albifrons; 22, Galago

crassicaudata.

* The remaining figures of the left ossicula show the malleus placed as in fig. 2, and the incus as in fig. 1. I had

intended that the human malleus and incus should be placed together in the desired position in the same group—

namely, fig. 2; but by an oversight the object was not reversed on being drawn on stone.

SECOND SERIES.—ZOOLOGY, VOL. T. 3U

494 MR. A. H.G. DORAN ON THE MORPHOLOGY

Fig. 23. Left malleus and incus: Cheirogaleus cookii.

Figs. 24-26. Left ossicula: 24, Propithecus diadema (young) ; 25, Perodicticus potto; 26, Nycticebus

tardigradus.

Fig. 27. Left malleus: Arctocebus calabarensis.

Figs. 28, 29. Left ossicula: 28, Tarsius spectrum (young); 29, Chiromys madagascariensis.

Figs. 80, 31. Articular surface of right malleus: 30, Homo; 31, Felis, much enlarged ; a, a*, upper facet;

6b, b*, lower facet.

Figs. 32-41. Right mallei of various Carnivora, arranged so as to show the principal distinguishing

features seen in the different families, more especially the processus muscularis and the relative

breadth of the outer aspect of the manubrium: as, articular surface of malleus; n, neck ;

1, lamina; pg, processus gracilis; pm, processus muscularis; pd, homologue of processus

brevis; emn, external surface of manubrium:—82, Felis leo; 33, Canis lupus; 34, Hyena

maculata; 35, Proteles cristatus; 36, Ursus maritimus; 37. Viverra civetta; 38, Herpestes

ichneumon ; 39, Gulo luscus; 40, Meles taxus, showing angular projection on neck; 41. Lycaon

pictus.

Figs. 42-55. Left imcus: 42, Felis domestica; 43, Canis familiaris; 44, Cryptoprocta ferox ; 45, Viverra

cwetta; 46, Paradoxurus musanga; 47, Herpestes pulverulentus ; 48, Hyena; 49, Ursus mari-

timus ; 50, Otaria falklandica; 51, Alurus fulgens; 52, Bassaris astuta; 53, Cercoleptes caudi-

volvulus ; 54, Lutra vulgaris ; 55, Mustela erminea.

Figs. 56-64. Left stapes: 56, Kelis domestica; 57, Canis lupus: 58, Hyena; 59, Cryptoprocta feroz ;

60, Viverra civetta; 61, Herpestes pulverulentus; 62, Mustela erminea ; 63, Cercoleptes caudi-

volvulus; 64, Ursus maritimus.

Puate LIX.

Figs. 1-3. Left ossicula: 1, Felis domestica; 2, Arctictis binturong ; 3, Suricata zenik.

Fig. 4. Left malleus and incus: Octocyon lalandii.

Figs. 5-12. Left ossicula: 5, Canis familiaris; 6, Canis vulpes; 7, Martes abietum; 8, Ictonyx zorilla ;

9, Lutra vulgaris **; 10, Hupleres goudotit ; 11, Procyon lotor ; 12, Nasua nasica.

Fig. 13. Left malleus and incus: Aonya indigitata.

Figs. 13a-16. Left malleus : 13a, Cynogale bennettii ; 14, Cercoleptes caudivolyulus ; 15, Bassaris astuta ;

an, angular projection on neck; 16, Hlurus fulgens.

Figs. 17, 18. Left ossicula: 17, Ursus ornatus; 18, Otaria falklandica (c in this figure and in the mallei

of others of the Pinnipedia, represents the concavity on the anterior surface of the head of the

malleus). j

Fig. 19. Right ossicula: Otaria falklandica.

Fig. 20. Left ossicula: Otaria stelleri.

Vig. 21. Right ossicula: Trichechus rosmarus.

Fig. 22. Left ossicula: Trichechus rosmarus.

Fig. 23. Right ossicula: Macrorhinus proboscideus; c, concavity of malleus.

Fig. 24. Left ossicula: Macrorhinus proboscideus.

Fig. 25. Right ossicula: Cystophora cristata; c, concavity of malleus.

Fig. 26. Left ossicula: Lobodon carcinophaga.

Fig. 27. Right ossicula: Phoca vitulina; c, concavity of malleus.

Fig. 28. Left ossicula: Phoca vitulina; eas, extra articulating surface on the malletis and incus;

gr, groove on the incus.

Figs. 29-31. Left malleus: 29, Phoca barbata; 30, Phoca greniandica; 31, Phoca hispida.

* The processus muscularis was broken off in the specimen from which this figure was taken.

OF THE MAMMALIAN OSSICULA AUDITUS. 495

Puate LX.

Fig. 1. Left malleus and incus: Phoca vitulina (articulated).

Figs. 2-5. Left incus: 2, Phoca barbata; 3, Phoca grenlandica; 4, Phoca hispida; 5, Halicherus grypus.

Figs. 6-35. Left ossicula: 6, Sciwrus maximus; 7, Pteromys nitidus; 8, Anomalurus fraseri; 9, Tamias

striatus ; 10, Spermophilus citullus; 11, Arctomys marmotta; 12, Castor fiber; 13, Myoxus

glis; 14, Mus decumanus (oa, orbicular apophysis) ; 15, Hapalotis mitchellii; 16, Hydromys

chrysogaster ; 17, Arvicola amphibius ; 18, Fiber zibethicus (p.h.m, process from head of malleus) ;

19, Spalax typhlus ; 20, Rhizomys badius ; 21, Geomys bursarius ; 22, Bathyergus maritimus ; .

23, Dipus egyptius ; 24, Hystrix cristata; 25, Cercolabes insidiosus ; 26, Capromys pilorides ;

27, Aulacodus swindernianus ; 28, Octodon cumingii; 29, Chinchilla lanigera ; 30, Dasyprocta

agutt; 31, Dolichotis patachonica; 32, Hydrochwrus capybara; 33, Cavia caprera; 34, Lepus

cuniculus ; 35, Hyrax capensis.

Figs. 36, 37. Stapes: 36, Elephas indicus (adult) ; 37, Elephas africanus.

Fig. 38. Right ossicula: Hlephas indicus.

Fig. 39. Left ossicula: Hlephas indicus.

Puate LXI.

Fig. 1. Left ossicula: Rhinoceros bicornis.

ig. 2. Left malleus and incus: Tapirus bairdi.

Figs. 3, 4. Left ossicula: 3, Equus caballus ; 4, Hippopotamus amphibius.

Fig. 5. Right ossicula: Hippopotamus.

Figs. 6-9. Left ossicula: 6, Sus scrofa: 7, Dicotyles tajacu ; 8, Tragulus javanicus ; 9, Lama peruana.

Fig. 10. Right malleus: Camelus dromedarius.

Fig. 11. Left malleus : Camelus dromedarius.

Figs. 12-21. Left ossicula: 12, Antilocapra americana ; 13, Nanotragus montanus ; 14, Bos taurus (adult) ;

15, Bos taurus (foetus 6th week) ; 16, Ovis aries (adult); 17, Ovis aries (foetus, at birth) .

18, Gazella dorcas (adult) ; 19, Alcelaphus bubalus ; 20, Cervus elaphus (adult) ; 21, Camelo-

pardalis giraffa (adult) .

Figs. 22-33. Left mallei of various Ruminants* (young) : 22, Rupicapra tragus; 23, Saiga tartarica ;

24, Kobus ellipsiprymnus ; 25, Tetraceros quadricornis; 26, Catoblepas gnu; 27, Procapra

picticaudata ; 28, Cephalophus maxwellit ; 29, Nemorhedus sumatrensis ; 30, Nemorhedus goral ;

31, Ovibos moschatus ; 32, Anoa depressicornis ; 33, Moschus moschiferus.

Figs. 34-42. Incudes of various Ruminants (left): 34, Rupicapra tragus; 35, Saiga tartarica; 36;

Kobus ellipsiprymnus ; 37, Tetraceros quadricornis; 38, Procapra picticaudata; 39, Cephalo-

phus maxwellii ; 40, Ovibos moschatus; 41, Anoa depressicornis ; 42, Moschus moschiferus.

Puate LXII.

As many of the ossicles are here represented much magnified, a smaller figure is, m all such cases,

placed to the left of the greater, the smaller being drawn on the same scale as in all the other plates, 2. e.

three times the natural size.

Fig. 1. Right ossicula of a large American Sorex: oa, orbicular apophysis; pb, processus brevis.

Fig. 2. Left malleus from the same Sorer. Fig. 6. Left meus: Scalops aquaticus.

dig. 3. Left malleus: Myogale moschata, Fig. 7. Right malleus; Condylura cristata.

Fig. 4. Right ossicula: Myogale moschata. Fig. 8. Left malleus and incus: Condylura.

Fig. 5. Left ossicula: Talpa europea. Fig. 9. Right malleus: Chrysochloris capensis,

* The laminz were more or less mutilated in the originals from which several of the figures in this group were

taken.

3U 2

496

Figs

Fig.

Tig.

Hig.

Fig.

Wig.

Fig.

MR. A. H. G. DORAN ON THE MORPHOLOGY

. 10-18. Left ossicula: 10, Chrysochloris ; 11, Centetes ecaudatus ; 12, Ericulus nigrescens ; 13, Eri-

naceus europeus; 14, Gymnura rafflesit ; 15, Macroscelides alexandri; 16, Rhynchocyon cernet ;

17, Tupaia tana; 18, Galeopithecus.

19. Left malleus: Petrodromus.

20. Right malleus: Asellia (Phyllorhina) tridens ; acpm, accessory processus muscularis.

21. Right malleus: Plecotus auritus (outer aspect).

22. Left ossicula: Vespertilio nattereri.

23. Right malleus: Mormops blainvillic.

24. Left malleus: Molossus obscurus.

25. Left ossicula: Vesperugo serotinus.

26. Right ossicula: Pteropus edulis.

27. Left ossicula: Pteropus edulis.

28. Left malleus: Balena mysticetus.

29. Right incus and stapes: Balena mysticetus.

30. Right malleus: Balenoptera rostrata.

31. Right incus and stapes: Balenoptera rostrata,

. 82. Left malleus: Megaptera nove-zelaendiz.

. 33. Right malleus: Neobalena marginata.

. 84. Left malleus, right incus and stapes: Physeter macrocephalus.

35. Left ossicula: Hyperoodon rostratus.

Pratre LXIL.

.1. Left malleus and incus: Berardius arnuxit.

. 2. Left ossicula : Delphinus.

gs. 8-12. Right malleus: 3, Delphinus; 4, Globiocephalus ; 5, Phocena; 6, Pseudorca meridionalis ; 7,

Orca; 8, Delphinapiterus albicans ; 9, Monodon; 10, Inia geoffroyit; 11, Lagenorhynchus ; 12,

Platanista.

, 13. Left malleus: Balena australis, in situ (pet, petrous bone ; tb, tympanic bone).

. 14. Left malleus: Felis, for comparison of the homologous parts in figs. 13, 15, 16,17, 18, &e.

Figs. 15-17. Left malleus: 15, Globiocephalus ; 16, Monodon (with tendon of tensor tympani and process

Spe ste ete see eS

Set Ge Se Se (So Se Sa So So

3 oe of Ue Fe Ge Je de de ihc}

pas

from membrana tympani attached) ; 17, Platanista.

. 18, 19. Left malleus: 18, Berardius arnuxii; 19, Mesoplodon (Oulodon) grayii.

. 19a. Right malleus: Mesoplodon grayii (inner aspect, showing the facets, the groove for the process

from the tensor tympani, and the pointed posterior extremity of the head).

. 20. Right incus: Platanista,

91. Left malleus: Manatus americanus.

.22. Right malleus: ditto.

23. Right incus: ditto.

24. Left incus. ditto.

. 25. Stapes: Manatus senegalensis.

26. Left malleus: Halicore australia.

27. Right malleus: ditto.

28. Right incus: ditto.

¢. 29. Left incus: ditto.

g. 30. Stapes ditto.

OF THE MAMMALIAN OSSICULA AUDITUS. 497

Prats LXIV.

Figs. 1-6. Left ossicula: 1, Bradypus tridactylus ; 2, Cholepus didactylus ; 3, Tatusia peba ; 4, Dasypus

sexcinctus ; 5, Priodon gigas; 6, Tolypeutes tricinctus.

Fig. 7. Left malleus: Tatusia hybrida (new-born).

Fig. 8. Left ossicula: Manis dalmanit.

Fig. 9. Left meus: Manis tricuspis.

Fig. 10. Left malleus: Manis dalmanii. Head, much enlarged, to show articular surface.

Fig. 11. Left ossicula, Myrmecophaga jubata.

Fig. 12. Left malleus and incus: Tamandua tetradactyla.

Figs. 13-22. Left ossicula: 13, Cyclothurus didactylus ; 14, Orycteropus capensis; 15, Didelphys virgi-

nana ; 16, Dasyurus maugei; 17, Thylacinus cynocephalus; 18, Phascogale ; 19, Myrmecobius

fasciatus ; 20, Phalangista vulpina; 21, Phalangista cookii; 22, Petaurus flaviventer.

Fig. 23. Left malleus: Petaurus taguanoides.

Figs. 24-27. Left ossicula: 24, Phascolarctos fuscus; 25, Perameles; 26, Cheropus castanotis ; 27,

Macropus laniger.

Fig. 28. Left malleus: Halmaturus Bennettii (in situ).

Fig. 29. Left malleus and incus: Lagorchestes.

Fig. 30. Left ossicula: AHypsiprymnus.

Fig. 31. Right malleus: Macropus major.

Fig. 32. Left ossicula: Phascolomys vombatus.

Hig. 33. Right malleus: Phascolemys vombatus.

Hig. 34. Right malleus: Phascolarctos, showing great width; upper portion of the manubrium.

Fig. 35. Right malleus and incus: Echidna (Tachyglossus) hystrix, from without; shm, scale-like

process from head of malleus, concealing body of incus; pd, pl, processus brevis and longus

of incus.

Hig. 36. Left malleus and incus: Echidna, from within ; hm, head of malleus; pg, processus gracilis ;

phim, process from head of malleus, blending with processus gracilis.

Fig. 37. Right malleus and stapes: Ornithorhynchus paradozus.

Fig. 38. Left malleus: Ornithorhynchus.

Figs. 39-48. A series of stapedes of mammals, “ columellz ” of Sauropsida, displaying the gradations of

type in the Mammalia of the occasional reappearance of the bicrurate stapes in Aves: ms,

medio-stapedial element in columella of Sauropsidan, equivalent, according to Professor Parker,

to the processus longus of the mammalian incus.

Figs. 39-43. Srapes: 89, of Man; 40, White Bear; 41, Virginian Opossum ; 42, Great Kangaroo ; 43,

Tasmanian “ Wolf” (Thylacinus).

Figs. 44-48. Columella: 44, of House-Sparrow; 45, Nightingale; 46, Golden Eagle; 47, Wandering

Albatross (Diomedea exulans) ; 48, Nilotic Crocodile (adult).

N.B. The stapes of Homo, 39, is drawn on the scale maintained as the rule throughout these plates ;

so likewise are 46,47, and 48. The remaining stapedes are represented of the same size as fig. 39, for a

better comparison of their general forms.

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r 499 |

XIX.—On the Annelida obtained during the Cruise of H.M.S. ‘ Valorous’ to Davis

Strait in 1875. By W. C. M°Invoss, ICD., LL.D., FRSSL.S #., PLS, §e.

(Plate LXV.)

Read June 21st, 1877.

FOR the opportunity of examining the collection made during the voyage of the

‘ Valorous,’ I am indebted, as on many similar occasions, to Dr. Gwyn Jeffreys, the

scientific chief of the expedition. The specimens were not quite in such good pre-

servation as those collected, for instance, in the dredging expeditions of H.M.S. ‘ Por-

cupine, and Dr. Jeffreys’s yacht ‘Osprey,’ since specially prepared bottles and spirit

were sent on these occasions. No group of marine Invertebrates is more difficult to

preserve in a satisfactory manner than the Annelida; and it may not be inopportune

again to draw attention to the necessity of pursuing, as far as possible, the method

indicated below *.

From the days of Otho Fabricius onward the Annelids of the Greenlandic seas have,

in comparison with those of other parts, received unusual attention. Thus, besides the

mention of between 40 and 50 marine Polycheeta by the foregoing author in 1780,

A.S. Cirsted published, in 1843, his ‘ Gronlands Annulata Dorsibranchiata,’ a work

illustrated by eight good lithographic plates. He described about three dozen species,

and calculated that at least fifty Greenlandic dorsibranchs were known. Moreover, in the

year 1818, it was in the same region (Baflin’s Bay) that Sir John Ross discovered the great

depth at which marine animals lived, by dragging up from 1000 fathoms a fine Astro-

phyton and mud containing “* Lumbricus tubicola” (? Hyalinecia tubicola, O. F. Miller).

The laborious papers of Dr. Malmgren of Helsingfors, which include the Greenlandic

collections made by O. Torell and Amondsen, give us, with the exception of the remarks

of Mobius (which I have not seen) the most recent enumeration of the Annelidan

* On removing the Annelids from the dredge or collecting-jar they should be immersed for some hours in large

flat vessels containing the strongest methylated spirit, or, if this is inconvenient, placed sparingly in capacious bottles

-of the same fluid. After the lapse of two or three hours they should be taken out of the large flat vessels and put

in separate bottles (few in each) of the same spirit. In the case of those immersed in the latter at first, the liquid

should be poured off and replaced by fresh spirit. It is during the first twenty-four hours that the greatest care is

necessary ; for if by inattention the specimens are allowed to soften, their subsequent preservation is always

unsatisfactory. The bottles should be systematically surveyed again within a day or two; and, as a rule, in any

case in which the colour of the spirit is deep or its transparency clouded, the fluid should be removed and fresh spirit

or ahsolute alcohol again added. In some collections the annelids are only represented by their cuticular coats

enclosing, as in a bag, a mass of pulp: much time is lost in specific determination, while for thorough investigation

they are worthless. Further, there is generally no difficulty in preserving the majority of the hardy forms alive for

many months in deep bottles half filled with sea-water, and, in the case of those which bore in sand, a layer of the

latter on the bottom. One or two specimens only should be placed in each jar; and all should be supervised daily

in hot weather, for the removal of the injured and dead and the renewal of the seawater.

500 DR. W. C. MSINTOSH ON THE ANNELIDA

fauna of the region. From the works of Dr. Malmgren the list published in the ‘ Arctic

Manual’ for 1875 was drawn up by Dr. Liitken, of the University Museum, Copenhagen.

The list comprises 101 species of marine Polychzta. The remarks on the Annelids of

Novaia Zemlia by Dr. Ehlers *, and those of the Siberian Seas by Prof. Grube +, show

that not a few forms have a wide range in the arctic waters {.

Though an accident which happened to the ‘Valorous’§ prevented the extensive

dredging operations contemplated by Dr. Jeffreys, and limited the species of marine

Polycheta to about 68, yet of this comparatively small number 13 are new to the

Greenlandic area, and, in addition, about 9 are new to science. Finally, so far as our

present knowledge of the Annelids goes, there is no doubt that Dr. Jeffreys’s opinion

that the Greenlandic Invertebrates are more European than American, is substantiated.

EUPHROSYNE BOREALIS, Cirst. Station 1, 70° 30’ N., 54° 41’ W., off Hare Island,

Disco, 175 fathoms, on a bottom of sandy mud. The specimens measured about an

inch in spirit, with 25 body-segments, besides the first and last. The branchize in the

rows agreed with the description of Prof. M. Sars (viz. 7 or 8 in each row), though

sometimes one or two were deficient. Some are simple, the majority bifid, and that

nearest the median line generally trifid. The bristles are much more prominent organs

than in #7. foliosa, Aud. & Kd.; and the spur on the smooth kind is much shorter. The

bifid bristles, again, have bolder transverse serrations. TZ. foliosa occurs abundantly

between tide-marks in Herm, whereas the arctic species frequents deep water, and

ranges to Spitzbergen and Finmark. No example has yet been found in British waters.

Amongst the bristles are many specimens of a small red mite.

The form of the branchize seems to be very characteristic in the British forms, viz.

EL. foliosa, Aud. & Kd., and HL. armadillo, Sars.

Nycuia crrrosa, Pallas. Fine specimens from Holsteinborg Harbour, from 7 to 35

fathoms. This form has a wide range, from British waters to Scandinavia, Iceland,

Spitzbergen, and the Gulf of St. Lawrence.

Nycu1a AmonpsEeNI, Malmgren. Station 3, 69° 31’ N., 56° 1’ W., sandy mud, 100

fathoms, and Godhavn Harbour, Disco, in 5-20 fathoms. A common species in the

Gulf of St. Lawrence, Canada.

Eunoa Girstepi1, Malmgren. Holsteinborg Harbour, in 35 fathoms. The specimens

measure an inch and three quarters long, and are larger than those from Canada. It

ranges to Iceland, Spitzbergen, and Finmark.

HarmMornok impricata, L. Station 4, 67° 50’ N., 55° 27’ W., 20 fathoms on shell-

sand; station 5, 66° 59’ N., 54° 27' W., 60 fathoms on sand and shells; Holsteinborg

Harbour, 7-35 fathoms; outside Godhayn Harbour, in 80 fathoms; station No. I.

H.MLS. ‘ Alert,’ 65° 00’ N., 53° 00’ W., 30 fathoms, on a rocky and gravelly bottom. This

ubiquitous form is abundant; and there is nothing peculiar in size or coloration; indeed

* Ann. Nat. Hist. 4th ser. vol. xi. p. 464.

+ Mém. de Savants étrangers (St. Petersburg), t. viii. 1858.

+ The forthcoming ‘ Report on the Annelida of the Austrian North-Polar Expedition, by Dr. Emil von Maren

Faller of Vienna, will afford further valuable information on the subject.

§ See “ Reports on the ‘ Valorous’ Expedition,” Proceed. R. 8. vol. xxv. No. 1738, p. 182. <A preliminary

notice of the Annelids also appears in the latter.

OBTAINED DURING THE CRUISE OF H.M.S. ‘ VALOROUS.’ 501

much larger examples occur in my collections from Exeter Bay, Greenland. Some

varieties approach Hvarne, but there is no difficulty in discriminating them.

EVARNE IMPAR, Johnston. Station 4, 20 fathoms; Station 6, 64° 5’ N., 56° 47' W.,

410 fathoms, in sandy mud; Holsteinborg Harbour, in 10 fathoms; Godhavn Harbour,

5-20 fathoms.

EUPoLyNoé occipENTALIS, M°I. Station 1, 175 fathoms, in sandy mud. The frag-

mentary specimen has scales, which were absent in the Canadian example*. In spirit

these have a ferruginous-brown hue, and the surface is almost smooth, the minute

spines being insignificant. On the outer border they have rather numerous, long, slender

cilia (PJ. LXV. fig. 1), the tips of which are very slightly dilated. Parasitic Pedicelline

abound on the bristles. The Lepidonotus sublevis of Prof. Verrill + may be this or an

allied form, but his description is too lax for determination.

55° 10’ N., 25° 58’ W., in 1785 fathoms, on a bottom of Globigerina-ooze. This is repre-

sented by a minute and fragmentary example, which at first sight was considered to be

a Leanira. In Sthenelais and Stgalion the ventral bristles are more or less boldly

bifid ; in Husthenelais, while the bristles lean to the former types in regard to terminal

segmentation and outline, yet many have perfectly tapered tips without a trace of the

secondary process. In the present form every one of the inferior bristles (Pl. LXV.

figs. 2 & 2a) has a filiform tip, but the upper group (fig. 2) more nearly approach the

characteristic terminal portions in Leanira; it needs but the shortening of the tips of

the processes and the better definition of their transverse markings to convert them into

those of an ordinary Leanira, for the same gradation between the superior and inferior

series of the ventral bristles is maintained in that group. Moreover, the steps

between Sigalion and Sthenelais on the one hand, and Leanira on the other, are

further indicated by the diminution of the serrations on the distal end of the shaft of the

bristle, such being distinct in Husthenelais hibernica}, very rudimentary in the present

form, and altogether absent in Leanira.

The injured fragment is devoid of scales and is eyeless. The palpi resemble those of

Eusthenelais hibernica; the tentacular cirri (and probably the tentacle also) are short.

_A few lobate papillee occur on the feet, as in the last-mentioned species, and the ventral

cirrus does not seem to reach the tip of the setigerous division.

PxHoto# minutTA, Fabr. 7-10 fathoms in Holsteinborg Harbour; 5-20 fathoms in

Godhavn Harbour. The specimens are large.

NEPHTHYS cHcA, Fabr. Station 4, 20 fathoms, on shell-sand; 10 fathoms in Hol-

steinborg Harbour; 5-20 fathoms in Godhavn Harbour. Many Infusoria are attached

to the bristles.

NEPHTHYS CILIATA, O. F. Miller. Station 1, 175 fathoms, off Hare Island, Disco, in sand

and mud; Station 3, 100 fathoms, in sandy mud; 10 fathoms in Holsteinborg Harbour,

5 to 20 fathoms in Godhavn Harbour, Disco. Swarms of a commensalistic form, re-

sembling at first sight a little Leech, occur between the feet, and quite differ from similar

-* Ann. & Mag. Nat. Hist. 1874, vol. xiii. p. 264. + Invert. Anim. of Vineyard Sound, p. 581.

+ Trans. Zool. Soc. vol. ix. pt. 7, p. 407, pl. 73.

SECOND SERIES.—ZOOLOGY, VOL. I. 3X

502 DR. W. C. MSINTOSH ON THE ANNELIDA

forms (Zoxosome) occurring in the British species in having a shorter pedicle, with a

larger basal portion for attachment, while the tip of the body is bluntly rounded and

presents only an aperture on one side.

PHYLLODOCE GR@NLANDICA, Cirst. Station 4, in 20 fathoms, on shell-sand; Godhavn

Harbour, 5-20 fathoms. The large specimens exceed in size those from British waters.

Eurawia viripis, O. F. Miller. 7-12 fathoms, Holsteinborg Harbour.

EULALIA PROBLEMA, Malmgren. Station 5, 66° 59’ N., 55° 27’ W., 60 fathoms, on

sand and shells. In spirit the specimen is dusky brown dorsally, and the lamelle are of

_the same hue; ventrally it is paler. The tips of the bristles are comparatively short,

and the enlarged distal end of the shaft finely spinulose. Dr. Malmgren received only

a single injured example from the same (western) shores of Greenland.

Hreone arctica, Malmgren. Station 5, 60 fathoms, on sand and shells. A minute

species, in which the proboscis (in the present examples) forms a clavate process, smooth

throughout the greater part under a lens, but toward the terminal dorsal region of the

partly extruded organ a distinctly papillose area occurs. In some there is a division of

the proboscis into a firm basal region and a more membranous distal portion. It seems

to come close to #. pusilla, the chief differences between the species, according to Dr.

Malmgren, being that in the latter the body tapers equally anteriorly and posteriorly,

and the head is longer than broad; whereas in E. arctica the body tapers less, and the

antero-posterior and transverse diameters of the head nearly correspond. The superior

lamella of the foot touches the next division in the anterior and central regions of the

body in #. pusilla, but is separate in #. arctica. ‘The anal cirri are elongate (linear-

fusiform) in the former, broadly ovate in the latter. The condition of the proboscis in

FE. pusilla is not noticed. In the present form the inferior lamella is not longer than

the bristle-papilla. It was formerly procured in Spitzbergen and Finmark.

ETEoNE FLAVA, Fabr. 5-20 fathoms, Godhavn Harbour, Disco.

ANCISTROSYLLIS GR@NLANDICA*, n. sp. A remarkable form, apparently allied to

the Syllide, was dredged at Station 6, 64° 5’ N., 56° 47’ W., in 410 fathoms, on

a bottom of sandy mud. ‘The specimen is about 3 of an inch long. The head

forms a short blunt cone, divided by a somewhat shallow fissure into two frontal

lobes (Pl. LXV. fig. 20), each of which has a short tentacle at the base, while

toward the posterior border another short conical tentacle occurs in the middle line.

Immediately below the great lateral eye on each side are two somewhat larger

tentacular processes, which resemble those on the adjoining segments of the body.

The latter is slightly tapered anteriorly, and more distinctly so posteriorly. The

segments are deeply cut, especially behind the anterior third and for some distance

along the wide part of the body. They are 57in number. Each foot, in the anterior

third, has a short dorsal and a smaller ventral cirrus (Pl. LXV. fig. 3), a single large

and characteristic hook, which somewhat resembles those on the first body-segment

of Paramphimome pulchella, Sars, placed above the former, and a slender longitudinally

striated bristle (generally enclosed in the tissues). The inferior division bears, in a promi-

nent lobe, a spine and a series of simple bristles, with a slight wing on one side—best

* dyxiorpov, hamus.

OBTAINED DURING THE CRUISE OF H.M.S. ‘ VALOROUS.’ 503

developed in the lower group, which also have a claw at the tip. The latter process is

indistinct in the delicately tapered superior series. The cuticle of the foot has many

cylindrical papillee; and similar organs occur on the dorsal and ventral cirri, those

on the latter, however, being less marked. The relationships of this form are not easily

made out; for while agreeing in some respects with the form of the head in such as

Microsyllis, it wholly differs in other points, so that it is only provisionally ranged

amongst the Syllide.

NEREIS ZONATA, Malmgren. Station 1, 175 fathoms, in sandy mud, off Hare Island,

Disco ; Station 8, 100 fathoms, on a similar bottom. In a large male example a peculiar

change occurs in the bristles; for besides the ordinary kind, toward the anterior third

of the body, a dense tuft of translucent bristles exists in each division of the foot. In

shape these resemble the tips of the Heteronereis forms, though they are only partially

developed, the tips alone projecting through the skin, while the shafts form a dense

mass in the neighbourhood of the spine. The distal division of these bristles (fig. 4)

resembles the blade of a sword, with a finely serrate edge, and tapering to an acute point,

which is slightly prolonged in a filiform manner. Moreover, a tendency to the foliaceous

condition of the feet accompanies the appearance of the Heteronereis-bristles, processes

appearing at the base of the dorsal cirri, a foliaceous growth at the inferior bristle-

papilla, and a process on each side of the ventral cirrus. Such, as usual, are best

marked posteriorly. The specimens are tinted with fine reddish-brown bars.

NEREIS PELAGICA, L. Station 4, 20 fathoms, shell-sand; Station 5, 60 fathoms, on sand

and shells; Station 6, 410 fathoms, in sandy mud; Godhavn Harbour, 5-20 fathoms;

H.M.S. ‘ Alert,’ Station 1, 65° 00’ N., 58° 00’ W., 30 fathoms, on a rocky and gravelly

bottom.

LUMBRICONEREIS FRAGILIS, O. F. Miller. Station 1, 175 fathoms, off Hare Island,

Disco; Stations 3 and 6, wt anted; Station 7, 63° 9 N., 56° 43’ W., 1100 fathoms, on a

bottom of mud; Station 16, in 1785 fathoms, in Globigerina-ooze; outside Godhavn

Harbour, 5-20 fathoms. This species, common in Britain, has hardly been differentiated

in the Catalogue of the. British Museum from the abundant southern type Notocirrus

tricolor, Mont.

NoTHRIA CONCHYLEGA, Sars. Stations 1, 3, 5, wt anted. The specimens are large and

in very firm tubes of small pebbles, very similar to those from the Gulf of St. Law-

rence. The British examples have tubes composed of flat bivalves, fragments of Echino-

derm-tests, &c.

GLYCERA CAPITATA, Cirst. Stations 4 and 7; Station 9, 59° 10'N., 50° 25’ W., in 1750

fathoms, and on a muddy bottom.

GLYCERA SETOSA, (rst. On the surface of the sea at Ritenbenk Kulbrud, Waigat

Strait; a fragment from Station 4. This form is readily distinguished by the great

length of the feet and bristles and by the structure of both. The nerve-trunks are

rounded and more distinctly separated than usual in the group. The longitudinal

ventral muscles do not seem quite to cover the trunks in the soft preparation, though

a thin layer of tissue does.

3x 2

504 DR. W. C. MSINTOSH ON THE ANNELIDA

ARICIA GRENLANDICA, n. sp. (Pl. LXV. figs. 5-9). A fragmentary specimen (anterior

region) was dredged at Station 4, in 20 fathoms. Many of the examples of the genus

procured elsewhere by the dredge are in the same condition (7. e. represented by the

anterior region), while those procured by digging are not uncommonly devoid of the

same part. The general form of the head and body corresponds to that of its allies,

and especially to Aricia Kupfferi, Ehlers *, a form described from the collection of the

‘ Porcupine.’ It is at once distinguished, however, from the latter, 4. Cuvieri, and

the abundant Scotch 4. Latreillic by the absence of the pectinate rows of papillee on the

ventral surface. The snout is elongated and pointed, and a little pigment exists at the

base of the region dorsally, but there is no evidence of distinct eyes in the preparation.

The lateral folds of the mouth turn smoothly round to form the posterior border of

the first bristle-bearing segment. The succeeding sixteen bristle-bearing feet (2nd to

17th) constitute the anterior region, characterized by the pectinate rows of papille

along the lateral border of the foot and the numerous strong bristles. The papille are

few in number, generally about six, and the upper is considerably longer than the others :

ail are simple, conical, hypodermic papille. The stout bristles of the inferior division of

the foot are somewhat like those of M. Claparéde’s 4. feetida, consisting of thick shafts

with peculiarly curved tips (Pl. LXV. fig. 5a), the striated brownish region ending in a

fine point, beyond which the translucent terminal wings proceed to the tip. The edges of

the latter (fig. 5, seen from behind) are finely serrated. Besides the foregoing, there are

numerous short tapering bristles of the same type as the dorsal fascicle, and a few large

hastate brown spines (fig. 6), which posteriorly sometimes project in a conspicuous

manner. The dorsal fascicle consists of a series of stout tapered bristles, having the

characteristic transverse markings so common in the group, and apparently due to

minute rows of spines transversely arranged. The tips of the bristles are much attenu-

ated, and the distance between the transverse rows (which become mere lateral serrations)

increases with the diminution of the bristle.

After the form of the foot alters, the bristles of the dorsal fascicle are much more

slender and the distal spinous rows are further separated. Amongst these are some shorter

forms (also with spinous shafts) which terminate in a fork, one of the prongs being

longer than the other and both slightly thickened at the tip (fig. 7). The web of the

fork has fine chitinous hair-like processes. A distinct curve of the shaft occurs below

the fork. The ventral bristles are much larger, with evident serrations and finely

tapered tips, the peculiar narrowing which takes place being less easily noticed than n

A. Latreillvi and others.

The branchize commence on the 5th bristled segment (fig. 9).

SCOLOPLOS ARMIGER, O. F. Miller. Stations 4 & 7, ut antea; Holsteinborg Harbour,

30 fathoms.

NAIDONEREIS QUADRICUSPIDA, (rst. A fragment from Station 7, 1100 fathoms, on a

muddy bottom.

AMMOTRYPANE AULOGASTER, H. Rathke. 5-20 fathoms, in Godhayn Harbour.

* Zeitsch. f. w. Zool. Bd. xxv. 1, p. 57.

OBTAINED DURING THE CRUISE OF H.M.S. ‘ VALOROUS.’ 505

OPHELIA Limactna, H. R. Station 4, 20 fathoms, in shell-sand. Of good size, but

not larger than at St. Andrews.

TACHYTRYPANE JEFFREYSII, n. sp. (Pl. LXV. fig. 10). Station 9, 59° 10'N., 50° 25’ W.,

1750 fathoms, on a bottom of mud. The body of this species is elongate, being upwards

of 2 inches long in spirit and about +5 of an inch in breadth, rounded in front, but marked

by the usual ventral ridges throughout the rest of its extent. There are about thirty

segments, besides the first and last, the anterior being short, those in the middle very

long. The head forms a short cone, with a minute filiform process at the tip. No

bristles are visible under a lens, the body being smooth and iridescent, like that of Lino-

trypane apogon, for the fine transverse lines are not conspicuous; very minute simple

bristles are, however, present. There are no cirri. The last caudal segment is separated

by a well-marked furrow from the rest of the body, and terminates in a slender recurved

process bent downward and forward. In the structure of the body-wall Tauchytrypane

Jeffreysti forms an intermediate link between Linotrypane and the other forms, con-

necting both with Ammotrypane and Ophelia. The cuticle (fig.11) is greatly thickened

(as in Limotrypane), and a special process passes inward from the median line on the

ventral surface toward the nerve-cords; it is further marked by fine cross strie. From

each side of the nerve-trunks the oblique muscle proceeds to the body-wall, and cuts off a

segment of the longitudinal muscular layer. The intestinal canal is filled with GJodi-

gerina-mud. .

AMMOTRYPANELLA ARCTICA, n. sp. Station 16, 1785 fathoms, on Globigerina-ooze.

A somewhat small form, measuring about ? of an inch long, and more slender than a

young Ammotrypane aulogaster of the same length, while the contour is different. The

cephalic lobe (Pl. LXV. fig. 12) especially diverges, being bluntly rounded, and furnished,

on each side of the base dorsally, with a considerable pigment-speck. The succeeding

segments are somewhat narrow, as in allied forms (e. g. Ammotrypane), and each has a

tuft of simple silky bristles, which, in a dorsal view (as in fig. 12), project prominently

outward and backward in the first seven or eight segments. The following twelve or

thirteen segments are larger, but their bristle-tufts are minute, and being sunk in the

groove opposite the attachment of the oblique muscle, do not show laterally in ordinary

views. ‘Toward the posterior end the bristles again become somewhat more distinct, and

slender cirri still remain on several feet. The caudal segments are short, and the last is

funnel-shaped and terminates ina smooth rim, The ventral sulcus commenees anteriorly

at the buccal orifice, and is well marked in front of the caudal process. The cuticle is

dense. The condition of the specimen is unfavourable for minute work. The total

number of segments seems to be about thirty.

A still more minute example comes from the same station. It presents no cirri;

but the state of the specimen (which appears to have been dried) is unfavourable for

the preservation of such organs. A papilla, considerably larger than in 4. aulogaster,

exists at the tip of the snout. The bristles are long in front of the caudal process,

which is cylindrical and appears to have an even rim. This may be an injured young

example of 4. aulogaster, though the snout and contour differ.

506 DR. W. C. M‘INTOSH ON THE ANNELIDA

Like Tachytrypane, the two foregoing had the intestinal canal filled with fine Globi-

gerina-rud.

TRAVISIA GLANDULOSA, n. sp. (Plate LXV. figs. 15, 16).

From the same depth (1785 fathoms, Station 16) is a Travisia, which, while agreeing

in form, presents certain differences from the ordinary 7. Forbesit, Johnst. The example

has been partially deprived of cuticle. The greater part of the basement tissue of the

body-wall is marked by prominent rows of punctures or dots, resembling those in the

Sipunculide. In the central region a larger row runs transversely across each seg-

ment, towards the posterior border. In comparing these with similar specimens of

T. Forbesii, the markings are much more uniform in the latter; and though here and

there a large papilla occurs, there is none of the uniformity characteristic of the new

form.. The dotted appearance is caused by the projection of the papille inward, as

observed in transverse section (Pl. LXV. fig. 16); and each has a tubular connexion

with the outer surface, like that noticed in the ventral papille of Aphrodita and Her-

mione. The cirri near the tail are much shorter and less distinct than in 7. Forbesi.

The specimen is so imperfect that some hesitation is felt in naming it; but the structure

of its body-wall is very different when contrasted with 7. Forbesii.

ScALIBREGMA INFLATUM, H. R. A fragmentary (small) example in 80 fathoms, out-

side Godhavn Harbour.

TropHonta PLUMOSA, O. F. Miller. A fragmentary specimen, in 175 fathoms, off

Hare Island, Disco (Station 1). The inferior hooks have their points somewhat less

curved than in the ordinary British forms, and the mud-and pebble-coated warts are a

little coarser.

FLABELLIGERA AFFINIS, Sars. Holsteinborg Harbour, 30 fathoms ; Godhavn Harbour,

5-20 fathoms. The specimens are considerably larger than those usually met with in the

British seas.

ScOLECOLEPIS CIRRATA, Sars. Station 1, 175 fathoms; Station 6, 410 fathoms. The

fragment is large.

From the first-named station is also the anterior region of a form which resembles

the foregoing in general outline. The head, however, instead of presenting the pair of

large, brownish, and somewhat faleate eyes in front of the median (occipital) tentacle,

exhibits only two small brown specks, placed close together. The foliaceous dorsal

pinne are also larger; and no trace of branchie occurs on the fragment, which con-

sists of about twelve segments. The bristles present no feature of interest.

ScoLECOLEPIS (?) JEFFREYSII, n.sp. Station 4, 20 fathoms, on shell-sand. This is repre-

sented by a fragment (about } of an inch in length) of the anterior end, and is closely

allied to Scolecolepis, but distinguished by the form of the snout, the arrangement of

the branchize, and other particulars. The head (Pl. LXV. fig. 13) is bluntly pointed

and divided into three regions by two antero-posterior curved lines, which are marked

with dark pigment posteriorly, and therefore very distinct. he central region is also

tinted with dark pigment posteriorly ; and in front of this is the short and somewhat

clavate tentacle, which has a constricted base, the whole rising from an elevated portion

of the snout. The body is flattened from above downward, and narrowed posteriorly

OBTAINED DURING THE CRUISE OF H.M.S. ‘ VALOROUS.’ 507

in the fragment. The first segment bears only the dorsal and ventral bristle-tufts ;

the second has, in addition, a dorsal cirrus; while the fourth possesses a well-marked

branchial process. The branchize remain of considerable size (¢. e. nearly meet in the

middle of the dorsum) for about eight segments, and then diminish to the fifteenth, in which

they are very minute. The bristles are simple striated structures, with a slight wing, as in

S. cirrata; and they increase in length so as to project outward and backward at the pos-

terior part of the fragment. In transverse section (Pl. LXV. fig. 14) the body-wall presents

(at the end of the fragment) an ovoid outline. The cuticle is well marked, and the hypo-

derm thick, especially along the entire ventral surface. The dorsal longitudinal muscles

are considerably less than the ventral, and a large gap occurs in the median raphe.

The ventral longitudinal muscles are separated by a central interval, at the bottom of

which the nerves lie. The oblique muscles pass over to the raphe at the summit of the

latter. The circular muscular coat is not much developed, but the lateral vertical bands

supplement it. The alimentary canal is rounded, has a thick circular muscular coat,

and the glandular inner surface is richly ciliated.

PrionosPio STEENSTRUPI, Malmgren. Fragmentary examples from Stations 3 & 16.

The stout anterior bristles show peculiar dotted markings, which are also present in the

short and curved inferior bristles of the median segments. The hooks have more than

one tooth above the great process.

CIRRATULUS crRRaATUS, O. F. Miller. Holsteinborg Harbour, 30 fathoms.

CAPITELLA CAPITATA, Fabr. Station 3, 20 fathoms. An injured fragment of a small

example.

MaLDANE Biceps, Sars. A single injured fragment in 175 fathoms, off Hare

Island, Disco (Station 1).

Ma.pane Sarst, Malmgren. Holsteinborg Harbour, 30 fathoms. Fragment.

Roprneé Lovent, Malmgren. Holsteinborg Harbour, 30 fathoms. A fragment in a

tube sparsely coated with clear quartzose sand-particles and rather brittle.

NIcOMACHE LUMBRICALIS, Fabr. Station 1, 175 fathoms, off Hare Island, Disco. The

size, like that of the Canadian examples, is large. The tubes are composed of sand or

sandy mud. on

AXIOTHEA CATENATA, Malmgren. Godhavn Harbour, 5-20 fathoms; and a fragment

from Station 1. Tubes composed of sandy mud. .

PRAXILLA ARcTICA, Malmgren. Station 7, 1100 fathoms, in mud. The species is

smaller than P. pretermissa, and the hooks agree with those figured from a Zetlandic

example in Trans. Roy. Soc. Edinb. xxv. pl. xvi. fig. 18. The tubes are composed of

very fine sand, and have a hyaline inner coat.

OwENIA FILIFoRMIS, D. Ch. Station 5, 60 fathoms; Station 16, 1785 fathoms; out-

side Godhavn Harbour, 80 fathoms. The tubes at the latter station closely resemble

the stalks of Alcyonidium, growing in proximity, while a fragment from 1785 fathoms

has a tube composed of Globigerine united by the tough secretion.

CISTENIDES GRANULATA, L. Holsteinborg Harbour, 10 fathoms; Godhavn Harbour,

5-20 fathoms. The tubes offer a noteworthy character, since they are in most cases con-

508 DR. W. C. MSINTOSH ON THE ANNELIDA

structed of transparent quartzose sand-particles set in the opaque secretion, so that, at

first sight, they seem pitted all over.

CISTENIDES HYPERBOREA, Malmgren. Station 1, 175 fathoms, off Hare Island, Disco.

The tubes of this species generally present a darker hue and less regular masonry. The

cement in both is more abundant than in Pectinaria belgica or Amphictene auricoma, so

that the architecture in the latter is more finished.

AMAGE AURIcULA, Malmgren. Station 4. A small fragment seemed to approach

Samytha sexcirrata, Sars, but its condition negatived satisfactory results.

AMPHITRITE CIRRATA, O. F. Miiller. 5-20 fathoms, Godhavn Harbour, Disco.

NicoLEA ARcTICA, Malmgren. 5-20 fathoms, Godhavn Harbour. The specimens are

larger than the following, and the pedicles of their branchiz are longer.

NICOLEA ZOSTERICOLA, Cirst. & Gr. Holsteinborg Harbour, 30 fathoms.

LEZNA ABRANCHIATA, Malmgren. Stations 5 & 6. A fragment also from the latter

presents certain affinities with Lanassa Nordenskidldi, Malmgren; but its condition is

such that a degree of doubt remains. The body is grooved along the ventral aspect

behind the ventral scutes. The bristles resemble those shown by Malmgren in Laphania

Boecki, that is, they have wings at the tip, and the latter in some is short and broad.

The hooks, again, have elevated crowns, with about 5 teeth above the large process, as

in Lanassa.

THELEPUS CIRCINNATUS, Fabr. A small example from Station 5; fragment from Hol-

steinborg Harbour, 10 fathoms; large specimens from Station 1 (30 fathoms), 65° 00' N.,

53° 00' W., H.M.S. ‘ Alert.’ It is well to remember that the posterior hooks show two

teeth above the large fang.

TRICHOBRANCHUS GLACIALIS, Mgrn. Station 12, 1450 fathoms, on a bottom of Glo-

bigerina-ooze and stones. ‘The tubes are chiefly composed of Globigerine.

TEREBELLIDES StrRemiI, Sars. Station 6, 410 fathoms; Station 7, 1100 fathoms;

Godhavn Harbour, 5—20 fathoms; and outside the latter in 80 fathoms.

SABELLA CRASSICORNIS, Sars. Fragments from Godhavn Harbour appear to belong

to this species. The occurrence of the branchiz in the tube showed that they did not

pertain to S. pavonia, as first supposed. The present species and S. spetsbergensis, Mgrn.,

seem to be closely allied, but the condition of the specimen was such that absolute accu-

racy was not attainable. The pigment on the branchie resembles that found in the

latter form. The tube is thin and tough.

EUCHONE ANALIS, Kroyer. Station 4, 20 fathoms on shell-sand ; Holsteinborg Harbour,

10 fathoms. One was in a fine hyaline tube coated with coarse quartzose sand and

minute fragments of shells.

CHONE INFUNDIBULIFORMIS, Kroyer. Station 4; 5-20 fathoms in Godhavn Harbour.

The hooks (Pl. LXV. fig. 17) differ from Dr. Malmgren’s figure. One was in a tough

brownish tube. .

Another Chone, evidently belonging to a different species, was dredged in the latter

locality, but it had apparently been dried before being placed in spirit. The collar is

large and of a similar form to that in C. infundibuliformis. The branchie (which are

OBTAINED DURING THE CRUISE OF H.M.S. ‘ VALOROUS.’ 509

about twenty in number on each side) are fragmentary, and appear in process of develop-

ment, as they scarcely exceed in length the diameter of the body. The long anterior

crotchets (Pl. LXV. fig. 19) slightly differ (in curves) from those of C. infundibuliformis,

while the posterior hooks (fig. 18) still more diverge, a comparison with one from the

same region in the former being made by referring to fig. 17. Whether this is the

Chone Duneri, Mgrn., or another species, the brief remarks and somewhat unsatisfactory

figures given by the author prevent determination. The posterior hooks especially

disagree.

The remarkable differences in the structure of the muscular system which distinguish

Euchone and Chone, on the one hand, from Sabella and Dasychone on the other, are like-

wise associated with corresponding differences in regard to the arrangement of the nerve-

cords. The fasciculi of each dorsal and ventral longitudinal muscle in Chone infundi-

buliformis form a complex double spiral in transverse section; while in Huchone the

dorsal also form a double spiral, less elaborate than in the foregoing, and the ventrals

each form asimple spiral. The nerve-trunks in both are much more closely approximated

than in Sabella and Dasychone.

SrrpuLta? A fragmentary tube of a minute, shining, quinquecostate species, with four

transverse ridges anteriorly, and irregularly twisted; from Station 11, 1860 fathoms.

DirRYPA GRENLANDICA, n.sp. (Proc. Roy. Soc. vol. xxv. no. 178, p. 219, woodcuts 1 & 2.)

Station 12, 1450 fathoms, on G‘lobigerina-ooze and stones.

A. Tube of Ditrypa grenlandica, enlarged. 3. The aperture, still more highly magnified.

The tube is about half an inch in length, not much thicker than a thread, and curved

from end to end like a bow (see woodcut, fig. 1,4). It tapers very gradually from the

anterior to the posterior end, contrasting in this respect with the more decided diminu-

tion in D.arietina. The oral aperture (fig. 1, B) forms a smooth and slightly constricted

rim, which is narrower than the tube almost by the depth of the ridges. The latter are

eight in number, and run from the anterior to the posterior end of the tube, though, it °

must be added, none of the specimens are quite perfect. It differs from any other

Ditrypa known to me in its slender form and the well-marked longitudinal ridges. In

the most perfect specimen the latter coalesce to some extent posteriorly, so that the tip

of the tube is somewhat quadrangular.

SPIRORBIS BOREALIS, Daud. Godhavn Harbour, 5-20 fathoms. Tube only.

SPIRORBIS VERRUCA, Fabr. Station 5, 60 fathoms. Tube only.

SPIRORBIS SPIRILLUM, L. Station 5.

SECOND SERIES.—ZOOLOGY, VOL. I. 3Y

510 DR. W. C. MSIINTOSH ON THE ANNELIDA

A remarkable tube (woodcut, fig. 2) occurred at Station 16, 1785 fathoms, amongst

Globigerina-ooze. It consists of a minute

tube (about the thickness of a slender silk

thread) of fine greyish mud, and having an

enlargement at one end, the blunt tip being

partly formed by a Globigerima-test. The

enlarged portion is tufted with what at first

sight (under a lens) appear to be minute

and finely ramose alge; but the micro-

scope shows that these delicately branched

processes are composed of the same elements This (ann Sinton 106), bey ee ahs

as the tube.

The Nemerteans are represented by a species from Stations 4 & 5, Holsteinborg Harbour

and Godhavn Harbour. The colour in spirit is brownish purple on the dorsum, whitish

beneath. The short body and large proboscis distinguish it from Nemertes Neesii; but it

may be related to the Amphiporus grenlandicus, Girst.—the more so, as the pigmentary

portion of the skin is apt to fall off in spirit-specimens. The longitudinal muscular coat

is cut into numerous and somewhat regular fasciculi by the radiating fibres. The base-

ment-tissue outside the circular coat is largely developed. The proboscis conforms to

the typical structure. If Mirsted’s NV. Neest was like this form, the British species widely

diverges. Another species, probably a Tetrastemma, comes from Station 5, but its con-

dition prevents accurate determination.

DESCRIPTION OF PLATE LXV.

Fig. 1. Portion of the outer edge of an anterior scale of Eupolynoé occidentalis. x 210 diam.

Fig. 2. One of the upper series of the ventral bristles of Husthenelais abyssicola. 2a, another of the

same series more closely resembling those in Sthenelais. x 350 diam.

Fig. 8. Foot from the anterior third of Ancistrosyllis grenlandica. x 210 diam.

Fig. 4. Heteronereis-bristle from a posterior foot of Nereis zonata, Mern., g. It is not completely

developed, and only the tips project beyond the fleshy part of the foot. x 210 diam.

Fig. 5. Anterior strong bristles (from the lateral pectinate row) of Aricia grenlandica, seen from behind.

5 a, another from the same part, viewed laterally. x 350 diam.

' Fig. 6. Large bristle from the anterior papillose feet (inferior division) of the same form. x 210 diam.

Fic. 7. Two of the bifid bristles from the region behind the pectinate feet (in the same animal). The

curve of the shaft is indicated in that on the right. x 350 diam.

Fig. 8. Tip of the inferior group of bristles after the change of type. The peculiar narrowing of the

organ towards the tip is apt to escape notice. x 350 diam.

Fig. 9. The 20th foot of the same species (not well preserved). @, branchial process. x about 20 diam.

Fig. 10. Tachytrypane Jeffreysti, nu. sp. Enlarged under a lens.

Fig. 11. Transverse (vertical) section of the body-wall of the same, about the anterior third. The speci-

men was not well preserved. x 20 diam.

OBTAINED DURING THE CRUISE OF H.M.S. VALOROUS. 511

. Ammotrypanella arctica, nu. sp. Enlarged under a lens.

. Scolecolepis Jeffreysii, n.sp. Enlarged under a lens.

. Transverse section of the anterior region (shortly behind the branchiz) of the same. The right

side is somewhat distorted from imperfect preservation: em, vertical muscle; om, oblique

muscle; dm, dorsal longitudinal muscle; ac, alimentary canal ; v, blood-vessel ; n, nerve-cords.

x 90 diam.

. Papille on the inner surface of the cuticular tissues of Travisia glandulosa. x 90 diam.

. Large papille of one of the transverse rows, as seen in transverse section of the body-wall of the

same. x 350 diam.

. Hook from posterior third of Chone infundibuliformis, Kroyer, from Godhayn. x 350 diam.

. Hook from the posterior third of the other Chone. x 350 diam.

. Crotchet from the anterior region of the latter species. x 350 diam.

. Anterior end of Ancistrosyllis grenlandica. Enlarged under a lens.

aie

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ANNELIDS FROM WN. ATLANTIC & DAVIS STRAIT.

finden

XX. On the Sacral Plexus and Sacral Vertebre of Lizards and other Vertebrata.

By St. Gtorce Mivarrt, Sec. L.S8., and the Rev. RoBert CuaRKE, /.L.S.

(Plates LX VI., LX VII.)

Read May 3rd, 1877.

INTRODUCTORY REMARKS.

It has of late been recognized that in any attempt to answer the question, which

vertebra of any lower animal answers to the first sacral vertebra of Man, the nervous no

less than the osteological relations of the parts should be carefully investigated; and it

has been considered that the nervous rather than the osteological relations should be

deemed the more important—in fact it has been sometimes assumed that the nerves

must be taken as the fixed points, and that the bones must rather have their homology

decided by the nerves than vice versd. Should it be possible to show that in any group

of reptiles both the nervous and the osteological relations of any vertebra constantly

agree with the nervous and osteological relation of Man’s. first sacral vertebra, the

homology between two such parts may well be taken as thereby established; but if

either of these sets of relations exhibit discrepancy, then of course such homology

cannot be considered satisfactorily determined. Nor can we justly set aside osteo-

logical in favour of nervous resemblances if it should turn out that the nerves

themselves exhibit notable variations of condition as we pass from one allied form to

another—a@ fortiori if there should be variations in this respect even within the limits

of a species. It might surely be anticipated that more or less variation would be found

to exist in nervous as well as in skeletal structures; and in the event of such

anticipations being justified, the determination of sacral homology must depend upon a

comparison of the values of the conflicting claims of differing degrees of resemblance in

both the osseous and nervous systems—unless we prefer to consider the osteological

sacrum and the nervous sacrum as distinct structures, which may or may not com-

pletely coincide, and may or may not widely diverge. .

Professor Gegenbaur, in his “ Beitrage zur Kenntniss des Beckens der Vogel”, has

come to the conclusion? that there is such a complete agreement between the anterior

of those two Saurian vertebree the expanded and elongated transverse processes of which

abut against the ilium and the first sacral vertebra of Man. In both such transverse

process abuts against the ilium, and in both (as he represents it) such vertebra is that

one which is directly postaxial to the issuing forth of the most preaxial of those nerves

the fibres of which go to the sciatic plexus and not at all to the lumbar plexus.

With respect to the second vertebra abutting against the ilium, however, he represents

1 See the ‘ Jenaische Zeitschrift, Band vi. Heft 2.

* As shown by his diagram fig. 2, at page 201.

514 MESSRS. MIVART AND CLARKE ON THE SACRAL

an osteological but not a nervous agreement with Man’s second sacral vertebra. The

divergence consists in this, that whereas in Man the nerve issuing preaxially to every

sacral vertebra, except the last, goes entirely to the sacral plexus, Professor Gegenbaur

represents the nerve in Saurians which issues preaxiad of the second ilium-joining

vertebra as only contributing but a very small part of its fibres to that plexus. The

arrangement thus generally indicated by him is supported by descrip-

tions of the nervous conditions which he found to exist in a certain Fig. 1.

number ef different species.

Professor Hoffman! has also given brief notices and rough diagrams

of the arrangement of the lumbar and sacral plexuses in a variety of

Batrachians and Reptiles.

These two authors, however, do not agree in their account, nor

have we been fortunate enough to entirely agree with either of them ;

and, indeed, the discrepancies manifested by the species we have

examined have seemed to us somewhat noteworthy, and such as to

render it desirable that we should go somewhat further into detail as

to nerve-distribution than has been done in the memoirs referred to.

Amongst the Lacertilia, Professor Gegenbaur has investigated the Veyves of Lizard (from

composition of the lumbar and sacral plexuses in Chameleo vulgaris, Gegenbaur).

Lacerta viridis, Lacerta agilis, Uromastix spinipes, and Grammato- er, crural; 1s, ischi-

phora barbata. Professor Hoffman has examined it in Chameleo, atic nerves; “other

Monitor, Gecko, Lacerta viridis, Iguana tuberculata, Urothropus, and i.

Polychrus marmoratus. We have made use of, for examination, Cha-

meleo vulgaris, Monitor arenarius (two specimens), Lacerta viridis, Teius tequexin, Iguana

tuberculata (two specimens), Agama colonorum, and Grammatophora barbata.

In our descriptions we shall make use of the following nomenclature :—The nerve

coming out in Lizards between the elongated transverse processes which abut against

the ilium we shall speak of as the intersacral nerve ; the nerve issuing out in front of

the more preaxial of these elongated transverse processes we shall call the first presacral

nerve; the nerve still more preaxial we shall name the second presacral, and so on.

The nerve issuing out postaxial to the more postsacral of the two elongated transverse

processes we shall call the jirst postsacral nerve; the nerve still more postaxial we shall

call the second postsacral, and so on.

NERVES OF THE LUMBAR, SACRAL, AND PRECAUDAL REGION OF LACERTILIA.

Chameleo vulgaris. (Plate LXVL. fig. 1.)

According to Professor Gegenbaur?, the sciatic nerve in this animal appears to be

formed by three roots, one intersacral and two presacral, the first presacral being the

largest. The intersacral nerve gives off branches to the postsciatic nerves, while the second

presacral nerve bifurcates, giving one branch to join the third presacral, so forming the

crural nerve, which thus has but two roots.

' See ‘ Niederlindisches Archiy fiir Zoologie,’ Band iii. Heft 2, p. 143 (1876). * L.c. pp. 199, 200.

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 515

Professor Hoffman! finds the sciatic nerve of the Chameleon to be formed by four roots,

having, in addition to those given by Gegenbaur, a postsacral root also, formed by a

branch from the first postsacral nerve. He also represents the nerve formed by branches

from the second and third presacral nerves (and thus having but two roots) as dividing

into two important branches, one the obturator nerve, the other the crural.

In our specimen of Chameleo vulgaris we found that the sciatic nerve might be said

to have either three or four roots, according as the branch (41a) connecting together

the first and second postsacral nerves should be considered a branch going from the

former to the latter, or from the second postsacral nerve to the first one. But our

specimen differed importantly both from the descriptions of Gegenbaur and Hoffman in

that, in the first place, the crural (15) and obturator (5) nerves were formed by three

roots instead of two, and, in the second place, that the position of all the roots of the

sciatic and crural nerves was different.

According to both Gegenbaur and Hoffman the sciatic nerve in this animal is formed

by the intersacral and the two presacral nerves, with or without the addition of fibres

from the first postsacral nerve, while of these roots the first presacral is the largest.

We, however, found the sciatic nerve (33) to be formed by the intersacral (29), first

presacral (12), and first postsacral (80) nerves; while of these roots the intersacral was

the largest. Again, both Gegenbaur and Hoffman found the first presacral nerve to be

undivided and to go wholly to the sciatic plexus, while we found the first presacral nerve

to bifurcate, one branch (12) descending to the sciatic, the other (11) ascending to the

lumbar plexus.

According to both Gegenbaur and Hoffman this lumbar plexus is formed from the

second and third presacral nerves, while we found it to be made up from the firsts

second, and third of these nerves (2, 9, and 11). Having united into a single trunk, we

found that this trunk almost immediately divided (as represented by Hoffman) into the

crural (15) and (5) obturator nerves, whereof the latter traversed the small pubic

foramen, called obturator foramen by Professor Hoffman?.

Lacerta viridis. (Plate LXVI. fig. 2.)

Professor Gegenbaur represents the sacral plexus of Lacerta as formed of three roots,

the middle one of which (that formed by the first presacral nerve) is the largest, and

sends no branch to the lumbar plexus; while the second presacral nerve bifurcates,

sending one branch to the last-mentioned plexus and one to the sacral plexus. The

lumbar plexus is formed by but two roots, one being part of the second presacral nerve,

and the other part of the third presacral nerve.

Professor Hoffman® quite agrees as to Lacerta viridis with Professor Gegenbaur, save

that he represents.no nervous connexion whatever between the intersacral nerve and the

first postsacral.

In one specimen of this species we found four roots to the sciatic nerve, there being a

distinct contribution to it from the first postsacral. Moreover we found as many as

three roots to the lumbar plexus, the fourth presacral nerve (2) concurring in its

1 Z.¢. pp. 174, 175, fig. 6. * L.c. tab. xi. figs. 9 and 10, fo. $ L.c. pp. 175, 176, fig. 9.

516 MESSRS. MIVART AND CLARKE ON THE SACRAL

formation with the whole of the third (9) and the preaxial bifurcation (11) of the

second presacral nerve. ‘The crural (15) nerve (that passing over the brim of the

pelvis) we found to be entirely formed by the preaxial part of the bifurcation of the

second. presacral, giving off, moreover, a filament to join the obturator nerve (5), which

has thus three roots :—1, the filament just mentioned; 2, the third presacral nerve; and

3, the fourth presacral nerve. The disagreement between the conditions found by us

and those described by the authors referred to was thus less striking in Lacerta than in

Chameleo. The sacral plexus was formed from the lumbo-sacral nerve (12), the first

presacral (21), the intersacral (29), and the first postsacral (30).

Teius tequexin. (Plate LXVI. fig. 3.)

A specimen of Teius teguexin was (with other specimens, as will be hereinafter

mentioned) most liberally placed at our disposal by Professor Flower, the Curator of the

Museum of the Royal College of Surgeons, from the stores of that Institution.

Unfortunately, in spite of its magnificent external appearance, it proved to be, internally,

in a state very unfavourable for dissection, while it was the first of all save one

examined by us. We cannot therefore speak with confidence as to the number of nerves

which contributed to form the lumbar plexus; but we did not observe that more than

two did so, the third (9) and part of the second (11) presacral; but we are certain that

three nerves contributed to the sacral plexus in addition to the lumbo-sacral nerve (12)

or postaxial bifurcation of the second presacral. The first postsacral (80) certainly

contributed to it, as well as the first presacral (21), thus differing from the conditions

described and figured by Professor Gegenbaur; while of the sacral roots the ¢ntersacral

(29), and not a presacral root, was the largest.

Grammatophora barbata. (Plate LXVI. fig. 4.)

This species is described by Gegenbaur! as only differing from his other Lizards in the

equality of the two branches of the bifurcating second presacral nerve. Its sacral nerve

therefore, according to him, is not reinforced by the first postsacral nerve.

In our specimens we did find it so reinforced, the sciatic nerve having thus four roots,

of which the first presacral (21) is (as Gegenbaur says) the largest. The lumbar plexus

we found to be formed, not by two but by three nerves—the fourth (2), third (9), and

second (11) presacral, the last bifurcating, and so giving off (12) the lumbo-sacral nerve

(or most preaxial root of the sciatic) as well as (11) the most postaxial root of the crural ;

but the two last-mentioned nerves we did not find of equal size, and the lumbo-sacral was

the more slender. On the right side we found the three roots of the lumbar plexus all

unite together, and then give off two branches, viz. the obturator (5) and crural (15)

nerves. On the left side, however, the fourth and third presacral nerves united and then:

gave off two branches; while the preaxial bifurcation of the second presacral also

bifureated, its bifurcations joining the two branches just mentioned, and then running

on as the obturator (5) and crural (15) nerves.

1 L.c. p. 200.

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 517

Agama colonorum. (Plate LXVI. fig. 5.)

This animal was the only one found by us to present that character figured’ by

Professor Gegenbaur, and described by him as existing in all Lizards”, namely, the non-

junction with the sacral plexus of the first postsacral nerve (80). The most postaxial of

the sciatic roots was the intersacral nerve (29), which gave off one or two branches (411),

going to the cloaca and tail respectively. The first presacral nerve (21) formed the

stoutest root; but besides a rather stout lumbo-sacral nerve (12) (forming the postaxial

bifurcation of the second presacral) the sacral plexus also received a slender branch (46)

from the third presacral, which bifurcates at its origin like the second presacral. In

no other Lizard whatever have we found the sacral plexus receive a branch from the

third presacral. Professor Gegenbaur mentions no such condition, while Professor

Hoffman has only found it? in one specimen of Polychrus marmoratus. This was not

the case in another specimen‘ of his, thus showing (what our own dissections have

convinced us of) that two individuals of the same species may present considerable

nerve-differences. The sciatic nerve (38) gave off, as usual, a branch high up to the

obturator internus muscle (28).

In Agama the lumbar plexus was formed by three roots. The most preaxial was the

fourth presacral nerve (2); the second root consisted: of the greater part of the third

presacral nerve (9), the rest of that nerve (46) going to the sacral plexus. The most

postaxial root consisted of a slender branch of the second presacral nerve (11). This

slender branch passes over the before-mentioned outer slender branch (46) so as to forma

marked cross on each side of the body. The obturator nerve (5) was principally formed

by the fourth presacral (2), but received an accession from the third presacral; it gave off

a branch (7) to the iliacus. The crural nerve was formed mainly from the third presacral

nerve (9), but received a branch from the fourth presacral and also the slender branch

before mentioned (11) from the second presacral.

Iguana tuberculata. (Plate LXVII. fig. 1.)

Professor Gegenbaur does not appear to have examined the sacral nerves of this

species; but Professor Hoffman? describes the sciatic as arising from three roots, namely,

1, a bifurcation of the second presacral nerve (i. e. the ilio-lumbar nerve); 2, the whole

of the first presacral nerve; and 3, a branch from the intersacral nerve. No branch is

described or figured as going to it from the first postsacral nerve. The lumbar plexus

is described as formed by four roots:—1, the fourth presacral nerve; 2, the preaxial

bifurcation of the third presacral; 3, the postaxial bifurcation of the same third pre-

sacral; and 4, by the preaxial bifurcation of the second presacral. The first and second

roots unite together to form the obturator nerve, and the third and fourth roots unite

together to form the crural nerve.

We have examined two individuals of Iguana tuberculata, for one specimen of

1 L.¢. p. 201.

2 « Vor allem ist die Betheiligung postsacraler Nerven am Plexus ischiadicus ginzlich aufegehohen.”.

3 L.c. p. 178, fig. 13. 4 Luc. fig. 12.

“§ L.c. pp. 176, 177, fig. 11. His figure does not agree with his description as regards the obturator nerve.

SECOND SERIES.—ZOOLOGY, VOL. I. 3Z

518 MESSRS. MIVART AND CLARKE ON THE SACRAL

which we were indebted to the great kindness of Professor Flower, and to Professor

Parker for the other. We found both sides to agree as to the arrangement of the roots

of the lumbar and sacral plexus, and at the same time to differ from the description

above taken from Hoffman, as also from that given by Gegenbaur as existing in Lizards

generally. We found the sacral plexus and sciatic nerves to be formed by four roots,

the most postaxial of which comes from the first postsacral nerve. We found the lumbar

plexus to be formed by three roots, the most preaxial of which is the fourth presacral

nerve. Fibres from all the roots of the lumbar plexus appear to go to form the obtu-

rator nerve; but the crural nerve seems to be formed from the second and third presacral

nerves only.

The main difference between our specimens and the figure and descriptions of the

authors referred to, however, consists in the concurrence of the first postsacral nerve in

the formation of the sacral plexus.

In order the better to determine the essential nature of the nerve-roots, we thought it

desirable to follow out their branches to the muscles which are supplied by them; being

favoured by the large size and fine condition of the specimens examined.

The fifth presacral nerve (1) we found to have no connexion with the lumbar plexus,

but to send its branches to the pyramidalis! and other abdominal muscles.

The fourth presacral nerve (2) bifurcated, the more preaxial branch (3) going to the

internal oblique? and perhaps other abdominal muscles. The postaxial bifurcation (4)

went to form the obturator nerve, but sometimes, before doing so, gave off a branch (7) to

the iliacus?. The muscle is, however, also supplied by branches given off (7) from the

obturator nerve itself, as on one side of the specimen figured.

The third presacral nerve (9) joins a branch (11) from the second presacral nerve, and

it may communicate with the last-mentioned nerve by an additional short neryous

connexion (14).

The second presacral nerve (11+12) bifurcates, the preaxial bifurcation (11) joining

the third presacral nerve, with or without the aid of the short connecting branch (14).

Hither at this junction (13), or from the second presacral nerve before the junction, the

crural nerve (15) is given off. The postaxial bifurcation joins the sacral plexus. The

second and third presacral nerves having thus jgined together, they are continued on

by a single branch (10) which joins the postaxial bifurcation (4) of the fourth sacral

nerve, before described, to form the obturator nerve (5), which, after giving off a branch

(6) to the iliacus, perforates the pubic foramen and. then divides into branches, which

respectively go to the pectineus, obturator externus, tibial adductor, adductor magnus, and

gracilis muscles. or

The crural nerve (starting, as has been said, either directly from the preaxial bifur-

cation of the second presacral nerve, or else from the junction of this bifurcation with the

third presacral) passes over the brim of the pelvis, and, giving off two or three branches

‘Tn naming the muscles, we have followed the nomenclature adopted in the paper “ On the Myology of Iguana

tuberculata,” published in the ‘ Proceedings of the Zoological Society’ for 1867, p. 766. The pyramidalis is there

described at p. 772.

? This muscle is described at p. 771 of the paper referred to. * For zliacus see U. ¢. p. 787.

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 519

(16) to the iliacus, then continues on to be distributed to the crwreus (20) and rectus

femoris (19), giving off in its course branches to the tsascalted) gluteus medius (17) and

maximus: (18).

The first presacral nerve (21), which is the largest of all, goes entirely to join the

postaxially extending bifurcation (12) of the second presacral nerve already mentioned ;

and these then, having effected a junction, are continued on by a single trunk (22) to

join the intersacral nerve (29).

The intersacral nerve (29) gives off, in the male, very close to its root, a very small

branch (81) to the penis, and then receives accessions, in the shape of two slender

branches (32 and 32), from the first postsacral nerve (30). The intersacral nerve, before

joining the conjoined trunk formed by the two more preaxial nerves, may give off a

branch (86 a) to the femoro-caudal muscle’, or a branch to this muscle may be given

off (86) from the great sciatic nerve itself.

The three roots of the sacral plexus (the intersacral and two presacrals) having thus

joined into one whole mass, the said mass divides into three unequal branches. The

smallest of these branches bifurcates and goes to the obtwrator internus®? muscle (23).

A second branch, which is rather slender, curves round the tendon of the femoro-caudal,

and gives off branches to the gracilis (24), the semimembranosus (25), the semitendinosus

(27), and the pyriformis (28) and biceps femoris muscles.

The main branch from the sacral plexus, the great sciatic nerve (33), gives off three

small branches to the gluteus medius (34) and maximus, to the ilio-peroneal (35), and to

the femoro-caudal (36) muscles. It then continues on down the thigh, just where the

deep edges of the gluteus maximus and ilio-peroneal are adjacent to each other, and,

distally, close to the delicate tendon of the femoro-caudal muscle*. When about two

thirds down the thigh, the sciatic nerve divides into the internal (37) and external (38)

popliteal; and later the latter again divides into two branches (89 and 40).

The first postsacral nerve (30), much smaller than the three preceding nerves, very

soon gives off a very long branch (44), which descends into the femoro-caudal muscle

and passes down the tail. It then bifurcates, but before or after bifurcating supplies

the slender branches (32 and 32) before mentioned which go to the intersacral nerve.

Its preaxial bifurcation ends by small branches (45), which supply the skin, transversus

perinei, and external, lateral, and caudal muscles. Its postaxial bifurcation (41) passes

under the tendon® from the caudal muscles which is attached to the tuberosity of the

ischium. It becomes connected by a very short branch (41 a) with the second postsacral

nerve, and then ends in delicate branches, which go to the external skin, transversus

perinet’ and sphincter cloace muscles.

The second postsacral nerve (42), more delicate than the first, gives off a branch (43)

1 This muscle is described at p. 791 of the paper referred to. Therein doubt was expressed as to correct naming

of the muscle, the name of which was chosen from the resemblance of the muscle in question to the gluteus maximus

of the Echidna. Its neryous supply coming from the crural is a strong argument against its being really the gluteus

maximus. 2 For this muscle see p- 794 of the paper before referred to.

* See Lc. p. 793. * For this tendon see J. c. p. 794, figs. 17 & 18 y.

° See lc. p. 774. ® See 7. ¢. p. 773.

382Z 2

520 MESSRS. MIVART AND CLARKE ON THE SACRAL

postaxial to the cloaca, and, after becoming connected, by the short branch (41 a) already

mentioned, with the first postsacral, it ends in delicate fibres whieh supply the sphincter

muscle and mucous membrane of the cloaca.

Monitor arenarius. (Plate LXVILI. fig. 2.)

The genus Monitor does not appear to have been examined by Professor Gegenhaur ;

but Professor Hoffman has described and figured! the arrangement of its lumbar and

sacral nerves. He represents its sacral plexus as formed from four roots, the most

postaxial of which is the first postsacral nerve, while the most preaxial is the lumbo-

sacral nerve coming off postaxiad from the second presacral nerve. The root consisting

of the first presacral nerve is the thickest, and does not bifureate, the whole of it going

to the sacral plexus. The second presacral nerve is shown dividing at once into three

branches, one of which is the lumbo-sacral nerve, another joins a branch of the third

presacral to form the crural nerve, while the remaining branch continues on as the

obturator nerve.

We have been fortunate enough to obtain two fine and large specimens of Monitor

arenarius, for one of which we are again indebted to the great kindness of Professor

Flower. In both these specimens we found on both sides of the body the same arrange-

ment of the nerve-trunks, which differed considerably from the conditions described by

Professor Hoffman. We found the sacral plexus to be formed of but three roots, the most

preaxial root consisting of the postaxial bifurcation of the first presacral nerve, while the

intersacral nerve was slightly the largest, and the first postsacral much the slenderest.

The first presacral nerve bifurcated, one branch going to the sacral plexus as already

mentioned, the other continuing on, as the crurai nerve, over the brim of the pelvis, but

being connected on each side with the second presacral nerve by a pair of delicate

nervous cords. The second presacral nerve having received (or given off) these nervous

cords, received an accession from the postaxial branch of the third presacral nerve, and

is then continued on as the obturator nerve.

We have thought it well to investigate the distribution of the nerves in this species,

being a species sufficiently remote from Jguana. We have, however, found a very close

similarity in this respect between these two animals.

The fourth presacral nerve (1) is like the fifth of Zguwana; it goes to the pyramidalis

and other abdominal muscles, and takes no part in the formation of the lumbar plexus.

The third presacral nerve (2) is like the similarly numbered fourth presacral of Jgwana,

except that we did not note that its postaxial branch (4) gave off any nerve to the

iliacus, although on one side of our specimen it gave off a branch which joined another

(10 a) from the second presacral, and thus continued on to the iliacus. The second

presacral nerve (9) is like the third of Jguana, except that it is less intimately fused

with the nerve postaxiad to it, with which it is only connected by commissural fibres

(14, 14, and 14.0). Having joined the postaxial branch (4) of the nerve before it, it is

continued on as the obturator nerve (5), giving off branches to the iliacus (6) before it

1 ZL. c. p. 175, fig. 7.

PLEXUS AND SACRAL VERTEBRZ OF LIZARDS. 521

passes through the pubic foramen. Having passed through the foramen, it divides (8),

and is distributed to the same muscles as in Zywana. On one side we found that minute

branches (16) went to the last-named muscle from the outermost of the commissural

fibres above referred to (14a). The first presacral nerve is like the second presacral of

Iguana, bifureating similarly, and sending one branch to the lumbar and one to the

sacral plexus. The former branch (11) is continued on (18) to become the crural nerve

(15), receiving on its way the commissural fibres before mentioned (14, 14 a, and 14d)

from the second presacral nerve. Having passed over the brim of the pelvis it divides

and supplies the very same muscles as in Jywana. The postaxial bifurcation (12) joins

the sacral plexus; but before doing so gives off a branch (23), going to the obturator

internus, the gracilis (24), and the semimembranosus (25) muscles. As in Zguana, the

branch to the obturator imternus (23) is isolated, while the fibres going to the gracilis

and semimembranosus (24 and 25) come off from the branch which also supplies the

semitendinosus (27) and the pyriformis (28). This difference is explicable by the fact

that the connexion between the sciatic and first postsacral nerves is less in Wonitor than

in Iguana. Therefore if the obturator internus, part of the semimembranosus, and the

gracilis are supplied by fibres coming from the first presacral, while the semitendinosus

and pyriformis ave supplied with fibres from the first postsacral, the whole set might be

supplied by the same main branch in J/guana (on account of the greater fusion above

mentioned), while they could not be so in Monitor. The intersacral nerve (29) is like

the intersacral nerve of Jgwana, save that it is connected with the first postsacral (30) by

only one connecting branch (82) instead of two. It thus appears that in Monitor there

is ne nerve answering to the first postsacral of /gwana, which seems entirely suppressed.

The sciatic nerve (33) was as in Jgwana, save that it gave no special branch to the femoro-

caudal muscle and no branch to the gracilis, semimembranosus, semitendinosus, and

pyriformis muscles. The first postsacral nerve (80) is like that of Jgwana, save that it is

(as has been said) less united with the intersacral nerve and the sciatic plexus, and

that it gives off the branch going to the semitendinosus (27) and pyriformis (28) muscles

as well as to part of the semimembranosus (26), which latter comes off from the sciatic

nerve in Jguana. This branch, like the branch supplying most of the same muscle in

Iguana, carves round the tendon of the femoro-caudal muscle. The same branch which

supplies the last-mentioned muscles also sends nerves (36) to the femoro-caudal itself.

As in Iguana, it gives off a branch (41) which passes under the tendon going from the

caudal muscles to the tuberosity of the ischium, and supplies the transversus perinei and

sphincter cloace muscle. It is also connected by a commissural branch (41 a) with the

second postsacral nerve; but in Monitor it is the commissural branch which is continued

(44) into the tail. The second postsacral nerve answers to the second postsaeral of Jgwana,

and gives off branches (42) to the mucous membrane of the cloaca and skin; but the

second branch to the cloaca which exists (43) in Jguana was not found in Monitor.

Thus the differences between these two very divergent Lacertilian forms may be thus

stated :—In Jguana four nerves contribute to form the sacral plexus; in Monitor only

three. In Jgwana the second presacral nerve bifurcates and goes to the two plexuses:

in Wonitor it is the first presacral that does so. In Zyuana the second, third, and fourth

522 MESSRS. MIVART AND CLARKE ON THE SACRAL

presacral nerves unite to form the lumbar plexus; in Monitor it is the first, second, and

third which doso. In Jguwana there is a branch to the iliacus (7) which was not detected in

Monitor. In Iguana the semimembranosus is not supplied from the first postsacral nerve ;

in Monitor it is so supplied. In Jguana the intersacral and first postsacral nerves unite

by two commissures; in Monitor by only one. In Jguana the femoro-caudal is supplied

by the sciatic nerve; in Monitor by the first postsacral. In Jguana more of the first

postsacral nerve joins the sciatic plexus than in Monitor, and the semitendinosus and

pyriformis are, in Monitor, supplied by the first postsacral instead of, as in Jgwana, from

the sciatic. In Iguana the obturator internus has a branch to itself, which comes from

the root of the sciatic. In Monitor it is supplied, in common with the gracilis and

semimembranosus, by a branch which is given off by the most preaxial root (12) of the

sacral plexus.

GENERAL CONDITION OF THE NERVOUS AND OssEOUS STRUCTURES OF THE SACRAL

REGION IN LIZARDS.

It appears, then, that in Lizards generally (judging from the specimens we have

examined and from the observations of Professors Gegenbaur and Hoffman), the

lumbar plexus may be formed by from two to three roots, and that the most preaxial of

them is never in advance of the fourth presacral nerve, while the most

postaxial root is never more postaxial than the first presacral nerve. Fig. 2.

Only in Monitor and Chameleo have we found this plexus to have

a root as postaxiad as the first presacral nerve, and neither of the jam

authors referred to represents it with so postaxiad a root.

The sacral plexus may have from three to four roots, but we have

generally found four. The most preaxiad of these is the lumbo-sacral

nerve (12), formed from the third presacral nerve (as sometimes in

Polychrus), or from the second presacral (as in Grammatophora, Lacerta,

Teius, Agama, and Iguana), or from the first presacral (as in Monitor).

The most postaxiad root is from the second postsacral, as in our spe-

cimens of Chameleo.

marmoratus 1m two

Generally, according to our examinations, the roots are the first jnaiviguals (from

(or first and second) presacral, the intersacral, and the first postsacral. Hoffman).

According to the authors referred to, it is generally the first and second

presacral and the intersacral, and never any postsacral. It cannot be — ‘schiatic nerve; o,

doubted, however, but that the roots preaxiad to the first vertebra aves f ie

with enlarged transverse processes may vary in number from one to Bie Be hy,

three, and that there may be two, one, or none behind the posterior postsacral vertebra;

vertebra with enlarged transverse processes. In all the Lizards re- s,s, vertebre with

ferred to there are two such vertebree with enlarged transverse pro- _—“"/arged transverse

cesses abutting against the ilimm. Are both these vertebrae sacral or 9 OS™

neither ; or if only one, which ?

The relation of the nervous roots to these transverse processes varies from species to

Nerves of Polychrus

crural nerve; 2,

is)

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 523

species, and even in different individuals (apparently, from Gegenbaur’s and Hoffman’s

works) of the same species. Shall we, then, consider that the nerves are the fixed points ?

If so, it is evident that we must extend our survey, and consider the nervous conditions

of the sacral region in other orders and classes of animals.

Crocodiles present no marked differences from Lizards, according to Gegenbaur,

who describes the sciatic nerve as formed from three roots—one intersacral, one pre-

sacral, and one postsacral, the intersacral one being much the largest. Professor Hoff-

man + agrees in this.

In Tortoises, according to Gegenbaur?, there are four roots to the sacral plexus—1,

one postsacral, as large as the next preaxiad, ¢.e. 2, the intersacral, and 3 and 4, two

presacrals, one of which is the largest of all. According to Hoffman? also there are four

roots; but he represents the postsacral root as by far the most slender, and the inter-

sacral root as larger than either of the two presacral nerves.

Fig. 4.

Nerves of Alligator (from Hoffman). Nerves of Tortoise (from Hoffman).

¢, crural nerve ; 2, ischiatic; 0, obturator; s, s, €, crural nerve; 2, ischiatic nerve; 0, obturator

vertebrae with expanded transverse pro- nerve; s,s, vertebrae with expanded trans-

cesses; 1 pr—3 pr, presacral vertebra ; verse processes; 1, 2, 3 and 4, presacral

1 po, postsacral vertebra. vertebra.

In all the Reptilia enumerated, the transverse processes which abut against the ilium

are wholly or in part parapophysial, and are in serial relation to (serial homologues of) the

capitular processes (or the capitular parts of the whole transverse processes) of the more

preaxial vertebra. The junction of the sacral vertebra with the ilium is much post-

acetabular in Saurians, but in Crocodiles and Tortoises (some at the least) it is about

opposite the acetabula.

THe Sacral REGION oF BATRACHIANS.

Here we have almost always but a single vertebra with enlarged transverse pro-

cesses abutting against the ilium. .

In the tailed Amphibia we have in Cryptobranchus japonicus, according to Professor

Humphry 4, a sciatic nerve formed from three roots, two of which are preaxial of the

_ | L.c. p. 187, fig. 14, * L.c. p. 200. ° L.c. p. 167, fig. 5.

* Journal of Anat. and Physiol. 1872. The muscles and nerves of C. japonicus, p. 51, woodcut.

524 MESSRS. MIVART AND CLARKE ON THE SACRAL

enlarged transverse process, while the third is postaxial to it, the obturator and crural

nerves being formed by two roots, one preaxiad to, the other

coincident with, the most preaxial root of the sacral plexus. Fig. 5.

The third spinal nerve, preaxial to the ilium (fig. 5, a), bifur- ary

cates, one branch forming the obturator nerve (fig. 5, 00), the

other contributing to form the crural (a.cr). The second spinal ,..,°

nerve, preaxial to the ilium (6), also bifurcates, one branch

joining the postaxial bifurcation of the preceding nerve to form Neves of Cryptobranchus (from

the crural, the other joining the spinal nerves, pre- and post- ae

axial to the ilium (¢ & d), to form the sciatic (sc). Hoffman merely cites! Professor

Humphry, and he gives a wrong reference.

In Salamandra maculosa and Triton cristatus, according to Hoffman?, the sacral nerve

is formed by one root preaxiad and two postaxiad of the transverse processes joining the

ilium. Dr. J. G. de Man? agrees with this representation, save that he places all the

roots one vertebra more preaxiad. We found, in Zriton cristatus (Pl. LXVI. fig. 6),

the same conditions as those described by Dr. de Man; but there was an interesting

difference between the two sides of the individual examined, inasmuch as there was a

lumbo-sacral nerve (12) only on one side, so that the sciatic had two roots (21 and 30)

on one side and three upon the other.

According to Professor Gegenbaur*, Salamandra maculosa

has three roots to its sacral plexus—(1) a delicate branch from

the nerve coming out preaxially to the large transverse pro-

cess; (2) one large root formed by the whole of the nerve,

i.e. coming out postaxially to the large transverse process ;

and (3), a delicate branch from the nerve next postaxiad.

Thus Gegenbaur agrees with Hoffman and differs from

Dr. de Man.

We found that a specimen (woodcut, fig. 6) of this species,

dissected by us, agreed as to its sacral roots with the obser-

vations recorded by Dr. de Man and figured by him in his nerco pvith tose of eaeceie

Table vii. fig. 4; but the branch connecting the second post- _ the Plates and as described in

sacral (9) nerve with the next postaxial nerve (12) appeared _ the text.

to us to be rather formed of fibres going preaxiad to the

crural nerve (15) than of fibres going thence postaxiad to the sacral (33). If we were

right in this view, the sacral nerve had but two roots, one issuing preaxiad (12) and the

other postaxiad (30) to the ilium-joining vertebra. Moreover, in our specimen, a branch

from the third preaxial nerve (2) joined, but very slightly so, the lumbar plexus on one

side, while on the other side it approximated to, but did not join, that plexus, and then

went on its way.

In the tailless Batrachia, the sciatic nerve appears, according to Hoffman, to be

a.cr

Fig. 6.

Lumbo-sacro-caudal vertebree and

nerves of Salamandra maculosa.

The lettering and numbers

1 ZL. ¢. p. 149, fig. 1. ? L.¢. p. 149, fig. 2.

* « Myologie comparative de extrémité postérieure chez les Amphibies,” Niederlindisches Archiv, ii. p. 58.

‘ L.c. p. 200. 5 L. ¢. pp. 156, 157, figs. 3, 4.

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 525

formed of three roots, one preaxiad and another postaxiad to the ilium-supporting ver-

tebra, while the third root is still more postaxiad. In Batrachians the transverse pro-

Fig. 7. Fig. 8.

Nerves of Dactylethra, fig. 7, and of Rana and Hyla, fig. 8 (from Hoffman).

¢, crural nerve; 2, ischiatic nerve; 0, obturator nerve; 1 pr—3 pr, presacral vertebrae; s. vertebra with enlarged

transverse processes.

cesses abutting against the ilium are parapophysial, not diapophysial, in nature, thus

agreeing with those of Reptiles.

THE SACRAL REGION OF MAMMALS.

In Man the great sciatic nerve has six roots, the most preaxial of which (the fourth

lumbar nerve of anthropotomy) may be called the second presacral of the nomenclature

here adopted, since the next postaxiad (the fifth lumbar nerve) comes out preaxial

to the first transverse process abutting against the ilium, and would thus be our first

presacral. The next postaxiad (the first sacral nerve of anthropotomy) may be reckoned

as equivalent to what we have termed intersacral in Lizards, while the remaining three

may be reckoned as equivalent to our first, second, and third postsacral nerves.

Now if we take the nervous relations as our exclusive test, then the vertebra postaxial

_ to the first sacral root of Lizards must be the homologue of the first sacral vertebra of

Man; and in that case this vertebra would, in Monitor and Chameleo, be the first with

expanded processes abutting against the lium. In Iguana, Teius, Lacerta, and Gramma-

tophora, however, it would be the vertebra preaxiad of such first ilium-joining vertebra,

while in Agama and, at least sometimes, in Polychrus', it would be the second vertebra

in front of such ilium-joining vertebra.

In an inferior mammal, such as the Cat (Pl. LXVI. fig. 7), we found the sacral plexus

to be formed by four roots, two very much larger than the others. The most preaxial (12)

root (the largest but one) is formed by part of the nerve coming out between the pen-

ultimate and ultimate lumbar vertebre; the next (the largest) comes out (21) in front

' See ante, woodcut, fig. 2, p. 522.

SECOND SERIES.—ZOOLOGY, VOL. I. AA

526 MESSRS. MIVART AND CLARKE ON THE SACRAL

of the first sacral vertebra; the third (the smallest but one) comes out (29) from the

most preaxiad sacral foramen; the last (the smallest) comes forth from (29 a) the second

sacral foramen. Thus here the sciatic nerve is formed from nerves issuing out of the

sacrum in a less proportion than in Man, and more by nerves issuing preaxiad to the

sacrum—not that the nwmber of the latter is greater than in Man, but that their relative ~

size, compared with those from out the sacrum, is greater. Compared with Lizards, it would

seem, with respect to nerves, that the first and second sacral vertebree of the Cat answer

very well to the two vertebree with enlarged transverse processes of Lizards, while osteo-

logically they, of course, also answer very well to them. There can be little doubt, how-

ever, but that the two first sacral vertebree of the Cat are to be considered homologous

with the anterior human sacral vertebrae; and therefore it would seem that the two

iliam-joining vertebree of Lizards should be considered homologous with the anterior

human sacral vertebree.

In Man, the Cat, and also in other mammals, down to the Hchidna, the transverse

processes abutting against the ilium are parapophysial, like those of the other classes

before described—viz. Reptiles and Batrachians. In all mammals which we have ex-

amined, however, the junction of the sacral transverse processes with the ilia is pre-

acetabular, although that junction is much less preacetabular in position in Man than it

is in most mammals.

Altogether, from the osseous and nervous conditions considered jointly in the groups

hitherto referred to, we would venture to propose the following definition of a “sacral

vertebra” in Mammals, Reptiles, and Batrachians:—“A vertebra with parapophysial

transverse processes which abut against the ilium preaxiad, or postaxiad, or opposite to the

acetabulum, and having a root of the sciatic plexus coming forth either immediately pre-

axiad or postaxiad of it.” This definition will exclude from the sacrum of Man, as not

abutting against the ilium, the more posterior vertebree called “sacral” in anthropotomy.

But in the lower mammals (even already in Apes) the number of so-called “sacral”

vertebree augments more or less with age by the ankylosis of the sacral vertebrz, so as

to render the extent of the ‘‘sacrum” very variable. It would surely be well, then, to

distinguish the human sacral vertebre (like the ribs) into true and false, those being the

true sacral vertebree which abut against the ilium.

THE SACRAL REGION IN BrrRDs.

The determination of the homological relations of the different parts of the postdorsal

part of the spinal column of Birds is a matter of much difficulty. It might be antici-

pated, however, from the close relations and probable genetic affinity between Birds and

Reptiles, that the true sacral vertebra of the former class might be readily determined

by the aid of reptilian characters. But however close may be the morphological afti-

nities, they are so strangely disguised by great physiological differences, that the results

of comparison are much too unsatisfactory to justify the anticipation above referred to.

The enormous forward prolongation of the iliac bones and the great power and activity

of the erect and vigorous legs of birds, compared with the backwardly extending ilia and

the sprawling and feebly pushing legs of Saurians, combine to produce in Birds a redun-

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 527

dancy of both osseous and nervous structures compared with those of the existing Rep-

tilia. Could we dissect the different forms of Pterosawria and Dinosauria, these diffi-

culties and obscurities would no doubt disappear; but the hiatus is too great between

the most Sauroid of existing Birds and any form of Lizard for the homological rela-

tions, in this respect, to be readily and easily solved.

In Birds generally, we find that a greater or less number of the vertebre which are

postaxial to those bearing elongate ribs, develop short parapophyses which abut against,

and ankylose with, the preaxiaily extending, preacetabular part of the ilium. Postaxial

to these come vertebree (which have been called1+ lumbo-sacral) which develop no para-

pophyses at all, or only minute rudiments of such, a fossa (the acetabular or renal

fossa) resulting from their absence.

Behind these, again, there are generally vertebree with elongated parapophyses abutting

against the ilium; and the first one, two, or three of these have often a more or less

decidedly different aspect from those which postaxially succeed. These vertebrae, which

we will provisionally call, for distinction, “posterior parapoyhysial,”’ abut against the

postacetabular part of the ilium in the Struthionide and in most other birds. In the

Merganser they are placed very much postaxiad to the acetabulum. In Otis they

are opposite the middle of the acetabulum, and they are even more preaxiad in Tetrao

tetric and Gallus, while in the Peacock (Pavo) they abut against the preaxial part of

the acetabulum. Thus in birds these posterior parapophysial vertebre (whatever may be

their true nature) sometimes have a position relative to the acetabulum nearly approach-

ing that which is found in the sacral vertebre of some mammals, though generally their

relation to it is that of the sacral vertebree of Saurians.

Ina few birds none of the vertebre postaxiad to the acetabulum develop parapophyses.

This is the case, é. g., in the Hornbill (Buceros), in which the most postaxiad vertebra, with

parapophyses abutting against the ilium, abuts against the front margin of the aceta-

bulum. In the Woodpecker (Pica) the position of the most postaxiad of these parapo-

physes is still more removed preaxially from the acetabulum.

In some Parrots the most postaxial of these vertebree with parapophyses is even more

preaxiad to the acetabulum than in Pica; but in other Parrot-forms there are also one or

two vertebree with parapophyses opposite the middle of the acetabulum, and separated

by an interval from the preacetabular vertebrae with parapophyses.

Judging, then, from the skeleton alone, we should be inclined to consider these posterior

parapophysial vertebre abutting, in most Birds, against the postacetabular part of the

ilium as the homologues of the lacertian sacral vertebre which also abut against the

postacetabular part of the ilium. And just also as we should consider the sacral vertebree

of Alligators and Tortoises, in spite of their more preaxial position with relation to the

acetabulum, as really homologous with the sacral vertebre of Lizards, so we should con-

sider the more preaxially situated posterior parapophysial vertebre of Otis, Tetrao,

Gallus, and Pavo as homologous with the postacetabular posterior parapophysial vertebrze

of the Struthionide and of most birds. What, then, is to be said as to the vertebrz of

1 Trans. Zool. Soc. vol. x. p. 9.

4a 2

528 MESSRS. MIVART AND CLARKE ON THE SACRAL

this region in such forms as Buceros and Pica and the Parrots? Osteologically they

are indeterminable, but they do not stand alone; the similar vertebrae of the Diver

(Colymbus) are also osteologically indeterminable from their ankylosis and lateral com-

pression, and the corresponding vertebrae of the Penguins can hardly be discriminated.

Quite osteologically indeterminable, again, is such a sacrum as that of the Frigate Bird

(Fregata), in which none of the vertebree of this region are devoid of parapophyses, and.

in which there is therefore no differentiation into anterior and posterior parapophysial

vertebrae within the pelvic part of the vertebral column.

We must turn, then, for aid to the nervous conditions of the parts.

Now in birds (fig. 9) the sciatic plexus and nerve has (as repre-

sented by Professor Gegenbaur) as many as six roots!; and if

we are to take nervous conditions as our test, and if we are to

consider the most preaxial parts of the sciatic nervous mass

in Birds as corresponding with the most preaxial parts of the

sciatic mass in Lizards, then we must consider that some or ae,

other of the avian vertebree without transverse processes (an-

terior to Gegenbaur’s vertebrae a and 0) must answer to those

vertebrae of Lizards which have enlarged transverse processes

to abut against the ilium.

But we have seen that the amount of variation in Lizards

is such that, if nerves be taken as our test, we must deny that

the ilium-joining vertebree of one species answers to the ilium-

joining vertebrae of other species—nay, even that, in indi-

viduals of the same species, we ought (according to this view)

to call the ilium-joining vertebree “sacral” in one and

“lumbar” in another. So to do, however, would surely be Nerves of Birds (from

ings alse b i Ho Be longated

If Professor Gegenbaur is, as he doubtless is, right as to Datimece ot aa

the sciatic nerve-roots in Birds, then, as there are six such against the ilium, opposite

roots, and the most preaxial of them comes forth in front of the acetabulum; er, crural

the most postaxial of those vertebrae with parapophyses which — °tve 00¢, obturator nerve;

are preaxiad to the vertebrae without parapophyses, it follows |” wo

that such vertebra would be included within that definition peeve

: 1'-5', vertebree postaxiad to

of “sacral”? which we have above proposed as applicable to a5.

Mammals, Reptiles, and Batrachians. But, at the same time, :

at least the first of the posterior parapophysial vertebre (7. e. the first vertebra postaxiad

to those without parapophyses) would also enter into that definition; and thus in most

birds we should have sacral vertebre separated from each other by interposed vertebrze

not sacral, which would be a very forced and unnatural conception.

But it might, perhaps, be suggested that as the ilia are so antero-posteriorly extensive

in Birds, the preacetabular and most preaxial of the vertebree giving out sacral roots in

transverse processes abutting

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 529

Birds answer to the sacral vertebree of mammals, which are preacetabular ; while the post-

acetabular and most postaxial of the vertebree giving out sacral roots in Birds answer to

the sacral vertebra of Reptiles, which are postacetabular in position. . But, as has been

said, in some Birds the posterior parapophysial vertebre abut beside the acetabulum ;

and there is so little difference in this region between Man and the Peacock that it

would seem to be very unreasonable not to consider the more preaxial of the posterior

parapophysial vertebree of that bird as being the homologues of the anterior sacral ver-

tebre of Man—and this the more so, seeing that the difference is bridged over by the

Tortoises and Crocodiles, which have their sacral vertebree beside the acetabulum. May

we not, then, take the nervous conditions as our guide, and say that every vertebra which

has a sacral root coming forth, either pre- or postaxiad of it, is “sacral’’? But then the

last two lumbar vertebree of at least many mammals would be sacral; while if we deter-

mine that only those vertebree shall be sacral which give forth from their postaxial end

a root going entirely to the sacral plexus, then the last lumbar of Man and the Cat would

be “sacral,” as also the first presacral vertebra of Teiws and Agama, while the second

sacral vertebra of no Lizard we have examined would be “ sacral’”’ at all !

If we were to add to our definition of a sacral vertebra, as proposed for Mammals, Rep-

tiles, and Batrachians, the words “ together with any or every vertebra without parapo-

physes and interposed between vertebree abutting against the ilium, and having a root

of the sciatic plexus coming forth either immediately preaxiad or postaxiad of it,’’ we

should have a definition which would include all the vertebree which might be considered

“sacral” in most Birds; but it would leave out in an arbitrary and unnatural manner the

vertebree in such forms as Buceros and Pica, which vertebree are plainly homologous with

those vertebree which in most Birds are without parapophyses, but are interposed between

vertebrae which have parapophyses. Neither, moreover, would it apply to Fregata.

According to Professor Gegenbaur, the most preaxiad root of the sacral plexus is, in

some species at least, the fifth spinal nerve in front of the posterior parapophysial ver-

tebree; and if he has correctly determined his vertebree! a and J in Pica and Buceros,

then the vertebra which he considers the fourth presacral will alone enter into even our

enlarged definition of a sacral vertebra as above given, and which will therefore be a

manifestly incomplete one for birds. The same would be the case with those Parrots

which have no parapophysial vertebree in this region, save greatly preaxial to the aceta-

bulum ; while in those which have such vertebre, five or six vertebree would be included

under it.

On the whole, then, seeing, on the one hand, the manifest homology between the

sacral vertebre of Man and Lizards (by the help of those of Crocodiles and Tortoises), and,

on the other hand, the manifest homology between the sacral vertebra of Lizards and

the posterior parapophysial vertebrae of most Birds, we think it better to consider the

latter vertebree in Birds as alone truly sacral, and to regard such forms as Buceros, Pica,

and certain Parrots as differing from the rule of the class by the suppression of their

parapophysial processes, and Fregata as differing from the same rule by the development

of parapophyses in all the vertebre of this region.

> L. c. plate vii. figs, xxix. and xxx.

530 MESSRS. MIVART AND CLARKE ON THE SACRAL

The sacral vertebree in Birds may be defined, then, as “vertebre having one of the

more postaxial roots of the sciatic plexus coming forth either immediately preaxiad or

postaxiad, having parapophysial transverse processes abutting against the tlium, and placed

mmediately postaxiad to vertebre which are devoid of such parapophyses, or else being

the homologues of vertebre so conditioned in most other Birds.” By the combination of

these two definitions, the former one for Mammals, Reptiles, and Batrachians, and this

latter one for Birds, it seems that the sacral vertebre may be defined in all Vertebrata,

above Fishes, which have pelvic limbs.

What, then, are we to consider those vertebree of Birds to be which are preaxial to the

sacrum—those, namely, which are generally more or less entirely devoid of transverse

processes? It appears to us that in Birds this region of the spine is really augmented in

extent and in the number of its vertebree in unison with the augmentation in the number

of the sciatic nervous roots. Such vertebre may, of course, be considered as corre-

sponding with the human lumbar vertebre; but inasmuch as they are so constantly

developed in Birds, and are so peculiar in that class, it may be as well to distinguish

them by a distinct appellation, and to call them, as one of us has elsewhere* before sug-

gested, ‘“lumbo-sacral vertebree.”

At first sight it might be well supposed that these vertebra answer to the sacral ver-

tebree of Man; but it is evident which of the vertebrze of Reptiles answer to the human

sacral vertebree ; and we agree with Professor Gegenbaur in thinking that the sacral ver-

tebree of Birds may be determined by the sacral vertebree of Reptiles. Upon this view the

normal type must be taken as existing in Reptiles and Batrachians, compared with which

we have in Man a multiplication of vertebre, postaxial to the true sacral vertebra, giving

their nerves to the sacral plexus, while in Birds we have a similar multiplication of the

vertebree which are immediately preaxial to the sacrum. The sciatic plexus in Man is

mainly furnished from the true sacral vertebree and the posterior vertebree which have

coalesced with them: it is furnished mainly from the presacral (lumbo-sacral) vertebree

in the Bird. That there should be this difference is, however, very little surprising when

we consider that even in the Cat it is the presacral and not the sacral vertebrae which give

forth the greatest part of the fibres of the sciatic plexus. Yet no one would dream of

calling the last lumbar vertebra of the Cat “sacral” on account of such increase in

volume of the presacral part of its sacral plexus. As to augmentation in the number

of roots, the variations we have found to exist in Lizards afford abundant ground for

declining to change the designation of a vertebra from “lumbar” to “sacral,” or vice

versd, on this account alone.

1 Trans. Zool. Soc. vol. x. p. 9.

Gt o

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 5381

DESCRIPTION OF THE PLATES.

Prats LXVI. fig. 1. Lumbo-sacral plexus of Chameleo vulgaris.

2. Ditto, Lacerta viridis.

3. Ditto, Teius teguexin.

4. Ditto, Grammatophora barbata.

5. Ditto, Agama colonorum.

6. Ditto, Triton cristatus.

7. Ditto, Cat (Felis catus).

1. Lumbo-sacral plexus and nerves of Iguana tuberculata.

2. Ditto of Monitor arenarius.

Priare LXVII. fig.

The following lettering applies consecutively to the different figures.

NERVES.

. Spinal nerve, third preaxial to the one giving off the lumbo-sacral nerve (. e. fifth presacral nerve

in Lguana, fourth in Monitor).

. Spinal nerve, second preaxial to the one giving off the lumbo-sacral nerve (fourth or third pre-

sacral).

. Branch to abdominal muscles.

. Branch continued on into the obturator nerve.

. Obturator nerve.

. Branch or branches to éhacus.

. Other branch or branches to ilacus.

. Obturator nerve dividing to supply the pectineus, obturator externus, tibial adductor, adductor

magnus, and gracilis muscles.

. Spinal nerve, next preaxial to that which gives off the lumbo-sacral nerve (third or second pre-

sacral nerve).

. [ts connexion with the obturator nerve.

. Preaxial bifurcation of the nerve, the postaxial bifurcation of which is the lumbo-sacral nerve

(second or first presacral nerve).

. Lumbo-sacral nerve (generally the postaxial bifurcation of either the second or the first presacral

nerve, but in Salamandra the entire first presacral) .

. Continuation of nerve 11 towards the obturator nerve.

. Connexion between nerves 11 and 9.

. Crural nerve.

. Branches to iiacus.

. Branch to gluteus medius.

. Branch to gluteus maximus.

. Branch to rectus femoris.

. Branch to crureus.

. Spinal nerve intermediate between the spinal nerve giving off the lumbo-sacral nerve and

22.

the more preaxial of the vertebree with enlarged transverse processes (either Ist presacral

nerve or absent).

Trunk formed by 12 and 21, and going to sciatic nerve.

Branch to obturator internus.

532

24.

25.

26.

27.

28.

29.

ON THE SACRAL PLEXUS AND SACRAL VERTEBR# OF LIZARDS.

Branch to gracilis.

Branch to semimembranosus.

Another branch to semimembranosus.

Branch to semitendinosus.

Branch to pyriformis.

Intersacral nerve. _

29 a. Second intersacral (in Cat).

30.

3l.

32.

33.

34.

35.

36.

First postsacral nerve.

Branch to penis.

Commissure or commissures between 29 and 30.

Sciatic nerve.

Branches to gluteus maximus and medius.

Branch to tlio-peroneal.

Branches to femoro-caudal.

36 a. Extra branch to femoro-caudal.

37.

38.

Internal popliteal nerve.

External popliteal nerve.

39 and 40. Divisions of external popliteal nerve.

Al.

Branches to ¢ransversus perinei and sphincter cloace.

41 a. Branch connecting first and second postsacral nerves.

42.

43.

44,

45.

46.

Second postsacral nerve, ending in branches to mucous membrane of cloaca and to skin.

Branch to cloaca.

Branch to femoro-caudal, and extending down to the tail.

Branches to skin and lateral caudal muscles.

Branch in Agama (i.e. in fig. 5, Plate LXVI.), extending postaxiad from the third presacral

nerve to the lumbo-sacral nerve.

VERTEBRA.

A. Most preaxial vertebra articulating with ilium.

B. Second (or more postaxial) vertebra articulating with ilum.

I. Vertebra preaxiad to A.

II, III, IV, and V. Vertebrz successively more and more preaxial.

Ik,

Vertebra postaxial to the more postaxial vertebra articulating with ilium, or to the only vertebra

which so articulates.

TI*, ILI*, and IV*. Vertebrz successively more and more postaxial.

Trans Linn.Soc.5zr.2.Zoon Vou. Pu 66

Agama colonerunv Triton cristatus.

Chameleo vulgaris

Granmatophora barhata

TZ. re Pe = ae

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Lacerta viridis. 2

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THE

TRANSACTIONS

THE LINNEAN SOCIETY

LONDON.

SECOND SERIES.—ZOOLOGY.

VOLUME I.

PART THE EIGHTH.

EON DON:

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET:

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XXI.

XXII.

XXIII.

XXIV.

CONTENTS.

PART VIII.—June 1879.

The Butterflies of Malacca. By ARTHUR GARDINER Borurr, F.L.S., F.Z.S.,

Senior Assistant in the Zoological Department, British Museum. (Plates

MODAN EINES. TXT re PE RET CRESS ar CUB SRS OS ORNS spageld533

On certain Organs of the Cidaride. By Cuaries Stewart, Esq., F.L.S.,

M.R.C.S., Lecturer on Comp. Anat. St. Thomas’s Hospital. (Plate LXX.) 569

On Hypsiprymnodon, Ramsay, a Genus indicative of a distinct Family (Pleopo-

dide) in the Diprotodont Section of the Marsupialia. By Prof. OwEn, C.B.,

F.RS., L.L.S., §c.. Superintendent of the Natural-History Departments in the

British Monscums (2 lates) IAXeXele SIXOXCLL, Pa 8.) ER oS

Descriptions of some minute Hymenopterous Insects. By J.O. Wxstwoop, W/.4.,

ee Sirece: (Plater XOMUUP er ee i on Sy ee ee eur oeS

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[ 583 ]

XXI. The Butierflies of Malacca*. By Anvuur Garpiver Burimr, F.L.S., ZS,

Senior Assistant in the Zoological Department, British Museum.

(Plates LXVIII., LXIX.)

Read December 21st, 1876.

THE generosity of Captain Stackhouse Pinwill, who has presented to the British

Museum the whole of the Lepidoptera collected by himself during some years’ residence

in Malacca and Penang, has enabled me to study (with, I trust, some degree of profit to

others as well as to myself) the Lepidopterous fauna of the Malayan peninsula. As I

was obliged to go over the whole of the species hitherto recorded from Malaysia, and as

Captain Pinwill’s collection is the most complete one hitherto received from the penin-

sula, I determined not to lose the present opportunity of making a complete list of the

Butterflies as yet known to occur at Malacca, comparing them at the same time, by means

of a table, with the species hitherto received from India, China, Siam, and the islands

to the south of the peninsula.

Of the 258 species now registered from Malacca, 36 appear to be endemic; of the

remainder rather more than a fourth occur either at Assam or Nepal, more than a

seventh at Moulmein, less than a seventh at Ceylon, nearly two fifths (apparently) in_

the island of Penang, about two elevenths at Singapore, about three sevenths in Borneo,

about three sixteenths in Sumatra, more than a third in Java, about two thirteenths in

Siam, rather more than a tenth in China, two species in the New Hebrides, and six in

Australia. We see therefore that, with the exception of the last-mentioned eight

species, the Butterflies of Malacca are limited to the Indian Region ; there are, however,

several forms occurring in the Australian Region which some Lepidopterists would not

regard as specifically distinct, such as the various allied forms of Danuais, Melanitis,

ycalesis, Doleschallia, Neptis, Diadema, Cynthia, Junonia, Lampides, Amblypodia,

Catopsilia, and Papilio. All of these are, nevertheless, characterized by slight but con-

stant differences, and consequently have a right to be regarded as distinct species.

It appears to me to be important to those who desire conscientiously to study the

geographical distribution of animals, to discriminate between even the most nearly allied

species. I believe that few things have more retarded zoological geography than the

reckless association together of so-called “local varieties” under the same specific name.

I will cite an instance in illustration of what I mean. Some Lepidopterists still assert

that Diadema bolina ranges from Northern India to New Zealand, whereas that species

does not occur outside the Indian Region. Allied forms, indeed, are common throughout

the Moluccas, Australia, and the South Pacific; and one of them (D. nerina), strange

* Two of the new species mentioned in the present paper, and an abstract of the whole, have already appeared in

the Society’s Journal (Zool. vol. xiii. p. 115 and p. 196 respectively).

SECOND SERIES.—ZOOLOGY, VOL. I. AB

534 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

enough, is found both in Java and the Australian Region, whilst a dwarfed form of it is

taken in the Philippines; yet D. nerina, although more widely distributed than D. bolina,

exhibits even less individual variation.

The fact of two species of Terias, discovered in the New Hebrides, turning up in

Malacca, is somewhat startling; but it is quite possible, in a genus where only a prac-

tised eye can at once detect the specific differences, that the same form of imago may

have been simultaneously developed in both localities ; moreover, breeding might (as it

often does) reveal constant and well-marked distinctive characters whereby the earlier

stages could at once be separated.

The following Note, which appears to me to be worthy of the consideration of natu-

ralists, has been communicated by my friend Mr. W. L. Distant, who was for some years

resident at Province Wellesley :—

** All catalogues of the Penang fauna must be accepted with some qualifications. The

dependencies of Penang and Province Wellesley are usually made more or less into one

zoological province, especially as regards the insects. This I found to be the case when

collecting there a few years ago, all collections being then made indiscriminately from

both localities.

*“The island of Penang is about 20 miles long by 9 miles broad, comprising an area

of about 107 square miles. <A group of granite hills occupies about two thirds of its

extent, running through its centre from north to south, bounded by a plain on their

eastern and western sides. These hills are densely covered with a beautiful forest on all

sides ; and their highest point, West Hill, is about 2600 feet above the sea.

“Province Wellesley is situated on the Malayan peninsula, and is separated from

Penang by a strait from 2 to 10 miles broad; it is about 35 miles in length, averaging

8 miles in width. Although it possesses many hills of a considerable elevation (yet not

equalling those of the island of Penang), its general character is more of a plain, in which

jungle tracts and cultivated lands of rice and sugar-cane are its characteristics. Man-

grove swamps, often of great extent, exist in the neighbourhoods of its many creeks and

rivers.

“There is little doubt that, when the floras and faunas of these two districts are

worked out separately and distinctly, a common character will be found to pervade them

both, but that many species will be found locally distinct and constant. Species may be

expected to exist in the lofty wooded regions of Penang that are absent in the plains and

less elevated hills of Province Wellesley. At present, however, when mixed collections

are made from both regions, the habitat ‘ Penang’ must be accepted as not representing

the insular fauna alone, but comprising also that of a portion of the Malayan

peninsula.”

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 539d

List of Species collected by Captain Pimwill.

RHOPALOCERA.

Family NYMPHALIDA.

Subfamily Danan, Bates.

Genus Evpia@a, Fabricius.

1. Evpr@a PHa@sus, Butler, Proc. Zool. Soc. p. 270. n. 3 (1866). od ?, Malacca.

This is clearly the species described by Felder as HZ. Castelnaut.

2. EupL@A OcHSENHEIMERI, Lucas, Rev. et Mag. Zool. p. 315 (1853). ¢ 2, Malacca.

3. Eupt@a Groret, Felder, Reise der Nov., Lep. i. p. 339. n. 470, pl. 41. fig. 7

(1865). & 2, Malacca.

4, EuPL@A MARGARITA, Butler, Proc. Zool. Soc. p. 279. n. 84 (1866). ¢ ?, Malacca;

2, Penang.

5. EUPL@A CHLO#, Guérin, in Deless. Souv- Voy. d. l’Inde, ii. p. 71 (1843). ¢ ?, Malacca.

6. HurpL@a Bremert, Felder, Wien. ent. Monatschr. iv. p. 398. n. 16 (1860). 3, Ma-

lacea. Mr. W. L. Distant took this species at Province Wellesley; he says that all

the species of Huplea which he captured were flying in the months of August and

September.

7. Eupt@a Ménérriést, Felder, Wien. ent. Monatschr. iv. p. 398. n. 15 (1860).

Malacca.

8. Eupt@a Prnwitti, n. sp. (PI. LXIX. fig. 9.) Primaries brownish piceous,

purple-shot, slightly paler along the external border: a long sericeous interno-median

streak : secondaries paler brown, deepest at base; two marginal series of whitish spots,

clear at anal angle, obsolescent and decreasing towards apex ; costal area greyish. Pri-

maries below paler than above, with a subcostal spot and an elliptical interno-median

spot pinky white; a spot in the cell and two beyond it bluish white, two or three white

dots at external angle; secondaries with the submarginal spots clear whitish; a spot in

the cell and seven in an arched series beyond it lilacine whitish; base and pectus

white-spotted. Expanse of wings 4 inches. ¢ 2, Malacca. ZH. Pinwilli is allied to

LE. Meénétriési, but larger, darker, and shot with purple.

9. Evpn@a MipAMus, (Papilio midamus) Linneeus, Mus. Lud. Ulr. p. 251 (1764).

$ 2, Malacca; ?, Penang. ‘ This species is very common in gardens ” (WV. L. Distant).

Genus CALLipL@a, Butler.

1. CALLIPL@A DIOCLETIANUS, (Papilio diocletianus) Fabricius, Ent. Syst. iii. 1, p. 40.

n. 118 (1793). ¢ ¢%, Malacca. ‘Common in gardens” (W. L. Distant).

2. CALLIPL@A VESTIGIATA, (Luplea vestigiata) Butler, Proc. Zool. Soc. p. 288. n. 58,

fig. 1 (1866). 3 ¢, Malacca.

4B 2

536 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

3. CALLIPL@A LEDERERI, (Huplea Ledereri) Felder, Wien. ent. Monatschr. iv. p. 397.

n. 14 (1860); Reise der Nov., Lep. ii. p. 317. n. 481, pl. 40. figs. 5, 6 (1865).

3 ?, Malacca.

Genus SALPINX.

1. SALPINX LEUCOGONIS, n. sp. Nearly allied to E. vestigiata, but smaller, the costa

of primaries not so strongly arched, the outer margin slightly inarched in the centre,

the submarginal spot on second median spot wanting, the subcostal spot above the

end of the cell much smaller, the spot on interno-median area widened into a notched

blotch, all the spots lilac; secondaries paler, only three lilacine whitish spots placed

obliquely near apex, no submarginal spots, anal angle white. Primaries below with

only two or three submarginal .and three or four marginal white dots; secondaries with

no white blotches beyond the cell. Expanse of wings 3 inches 2 lines, ¢, Malacca.

Genus Danats, Latreille.

1. Danats PLEXIPPUS, (Papilio plexippus) Linneeus, Mus. Lud. Ulr. p. 262 (1764).

3 ¢, Malacca; 3, Penang. “Common in August and September” (W. L. Distant).

2. DANAIS MELANIPPUS, (Papilio melanippus) Cramer, Pap. Exot. ii. pl. 127. figs. a, B

(1779). 3 2, Malacca; 3, Penang.

8. DANAIS VULGARIS, Butler, Ent. Mo. Mag. p. 164 (1874). 3 %, Malacca.

4. DANAIS SEPTENTRIONIS, Butler, Ent. Mo. Mag. p. 163 (1874). 3, Malacca ;

2, Penang.

5. DANAIS GRAMMICA, Boisduval, Sp. Gén. Lép. i. pl. 11. fig. 10 (1886). 3 ?, Malacca.

6. DANAIS MELANEUS, (Papilio melaneus) Cramer, Pap. Exot. i. pl. 30. fig. p (1775).

?, Malacca.

7. Danats crocHa, Butler, Proc. Zool. Soc. p. 57. n. 53, pl. 4. figs. 5, 6 (1866).

3 %, Malacca. “ Occurs at Penang in August and September” (W. L. Distant).

Genus IpzEopsis, Horsfield.

1. Ipropsts Daos, (Idea daos) Boisduval, Sp. Gén. Lép. i. pl. 24. fig. 3 (1836).

3 2, Malacca; ?, Penang.

Genus Hest1a, Hibner.

1. Hestta LyNncEvs, (Papilio lynceus) Drury, Ill. Ex. Ent. ii. pl. 7. fig. 1 (1778).

3, Malacca. ‘ The species of Hestia go by the local name of ‘the widow’ in Province

Wellesley; the genus was very abundant near the top of Penang Hill” (W. L. Distant).

2. HESTIA LINTEATA, n. sp. (Pl. LXIX. fig. 6.) Nearly allied to H. belia, but much

larger, the veins less broadly black-bordered; spots larger, excepting towards the costa

of primaries at apex; discoidal spot of secondaries very large; clavate markings ter-

minating the veins much longer, more slender in the middle. Hxpanse of wings 5 inches

6 lines to 6 inches 8 lines. o ?, Malacca (six examples).

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 537

Subfamily Satyrina#, Bates.

Genus Mexaniris, Fabricius.

1. MELANITIS LEDA, (Papilio leda) Linneeus, Syst. Nat. i. 2, p. 773. n. 150 (1766).

$ ?, Malacca, Penang. “Common in jungle-grass” (W. L. Distant).

Genus Letae, Hiibner. .

1. Levis evRopA, (Papilio europa) Fabricius, Syst. Ent. p. 500 (1775). 3,2, Malacca;

6, Penang. “In grassy places” (W. L. Distant).

Genus Mycaxzsts, Hiibner.

1. MycanEsis MINEUS, (Papilio mineus) Linneeus, Syst. Nat. i. 2, p. 768. n. 126 (1766).

d %, Malacea; 2, Penang.

2. MYCALESIS POLYDECTA, (Papilio polydecta) Cramer, Pap. Exot. ii. pl. 144. figs. H, F

(1779). 3, Malacca, Penang.

3. Mycatusis JANARDANA, F. Moore, Cat. Lep. E.I. C. i. p. 284 (1857). 3, Malacca.

4. MycaLEsis HESIONE, (Papilio hesione) Cramer, Pap. Exot. i. pl. 11. figs. C, D (1779).

3d ¢, Malacca, Penang.

5. Mycazsis Fusca, (Dasyomma fuscum) Felder, Wien. ent. Monatschr. iv. p. 401.

n. 27 (1860). Malacca.

YrrHima, Westwood.

1. Ypruima PHILOMELA, Hiibner, Zutr. ex. Schmett. figs. 83, 84 (1806). 3, 2, Malacca;

?, Penang.

2. YPHTHIMA (sic) METHORA, Hewitson, Trans. Ent. Soc. 3rd ser. ii. p. 291, ( Ypthima)

pl. 18. figs, 20, 21 (1865). ¢ %, Malacca; 2, Penang.

3. YPTHIMA CORTICARIA, n. sp. Wings above greyish brown, an inner submarginal

streak and an outer submarginal line blackish diffused; primaries with a large subapical

oval black ocellus, with two plumbaginous pupils and a testaceous iris surrounded by a

dusky line; a similar smaller unipupillated rounded ocellus near anal angle of second-

aries. Wings below white, densely reticulated with brown, two ill-defined, interrupted,

central, subparallel lmes; a submarginal streak and line as above: primaries with the

ocellus brighter and better defined than above; outer border pale brown, bounded by

the inner submarginal streak: secondaries with three ocelli, one at apex (below which

is sometimes another extremely small ocellus) and two placed obliquely at anal angle,

the lower one small and irregular. Expanse of wings Linch 8 lines. Allied to Y. nareda.

3 ¢, Malacca.

Subfamily ExyMnriin#, Kirby.

Genus Enymnias, Hiibner.

1. HiyMnras NIGRESCENS, Butler, P. Z. S. p. 520, pl. 42. fig. 1 (1871). 3 2, Malacca,

Penang. A long series of this species, exhibiting no variation.

538 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

2. ELYMNIAS MEHIDA, (Melanitis mehida) Hewitson, Exot. Butt. iii. Wel. pl. 1. figs. 2,3

(1863). Malacca.

Subfamily Morruin#, Butler.

Genus AMATHUSIA, Fabricius.

1. AMATHUSIA PHIDIPPUS, (Papilio phidippus) Linneeus, Syst. Nat. i. 2, p. 752 (1767).

3 ?, Malacca.

Two forms (probably seasonal varieties) occur in Malacca, the one with dark bands

below, the other with pale bands; in the female of the dark form the subapical yellowish

patch on the upper surface of primaries is more distinct. I have seen three examples in

Mr. Distant’s collection from Penang. It occurs in August and September.

Genus ZEvuxrip1A, Hibner.

1. ZEUXIDIA AMETHYSTUS, Butler, P.Z.S. p. 485. n. 5 (1865). 3 2, Malacca. The

female differs from the male much in the same way as in the sexes of Z. doubledayi.

Genus DiscopHora, Boisduval.

1. DiscopHorA MENETHO, (Papilio menetho) Fabricius, Ent. Syst. iii. 1, p. 83. n. 260

(1793). &, Malacca.

2. DIscOPHORA TULLIA, (Papilio tullia) Cramer, Pap. Exot. i. pl. 81. figs, A, B (1779).

& ?, Malacca. The typical male differs from that of the Indian form in the unspotted

character of the upper surface.

Genus XANTHOTZNIA, Westwood.

1. XANTHOTANIA BUSIRIS, (Clerome busiris) Westwood, Trans. Ent. Soe. ser. ii. vol. iv.

p- 187 (1858). 2, Malacca.

Genus THAUMANTIS, Htibner.

1. THAUMANTIS NOUREDDIN, Westwood, Gen. Diurn. Lep. p. 337. n. 6, note (1851) ;

Trans. Ent. Soe. ser. i. vol. iv. p. 175. pl. 20 (1858). ¢, Malacca. ‘‘ Occurs at Penang

in August and September” (WV. L. Distant).

2. THAUMANTIS PSEUDALIRIS. (Pl. LXVIII. fig. 1.) Zhauwmantis aliris 8, Westwood,

Trans. Ent. Soc. n.s. vol. iii. p. 176 (1856-58); Butler, Journ. Linn. Soc. xiii. p. 115.

This species differs from the male of Z. aliris from Borneo in having the band of

primaries above half the width, not notched, and yellower in tint; the basal area of all

the wings ferruginous. Below, the area beyond the band is not striated, and is of the

same rusty red colour as the broad outer border, the latter is also paler at the edge, and

is cut much shorter by the obliquity of the transverse band ; the basal spots are rusty

red instead of red-brown and black; the irregular band of secondaries is paler and much

more constricted in the centre; the external and discal areas are altogether paler and

more uniform in tint; the ocelli are considerably smaller and paler; the spots between

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 539

the ocelli and the yellow submarginal spots are obsolete. Expanse of wings 4 inches

7 lines. &, Malacca.

As Professor Westwood figures the Bornean female, I am obliged to rename the

Malacca male.

Subfamily NympHatin», Bates.

Genus DoLESCHALLIA, Felder.

1. DoLESCHALLIA PRATIPA, Felder, Wien. ent. Monatschr. iv. p. 399 (1860). ¢ 2,

Malacca. This species has hitherto been represented in the Museum collection by only

one female; it differs considerably from D. disaltide in both sexes.

Genus CHARAXES, Ochsenheimer.

1. CHARAXES JaLysus, Felder, Reise der Nov., Lep. iii. p. 438, pl. 59. fig. 5 (1867).

$, Malacca. Mr. Distant gave me an example of this species taken by: him at Province

Wellesley.

2. CHARAXES HEBE, Butler, P. Z. 8. p. 634, pl. 37. fig. 3 (1865). ¢ %, Malacca. The

male does not differ at all in pattern from the female.

3. CHARAXES SCHREIBERI, (Mymphalis Schreiber) Godart, Enc. Méth. ix. Suppl. p. 825

(1823). ¢, Malacca.

Genus Apottas, Boisduval.

1. ApoutAs aponta, (Papilio adonia) Cramer, Pap. Exot. iii. pl. 255, figs. C, D (1782).

3 3, Malacca.

2. ADOLIAS GARUDA, Moore, Cat. Lep. E.I. C. i. p. 186. n. 374(1857). &, Malacca.

3. ADOLIAS JAMA, Felder, Reise der Nov., Lep. ili. p. 431 (1867). ¢ 2, Malacca. This

species appears to be common.

4. 2 ADOLIAS LAVERNA, Butler, Cist. Hnt. i. p. 29 (1870). ¢ 2, Malacca. Now

that both sexes of this species have come from the same locality, I find that the male

from Borneo figured in my ‘ Lepidoptera exotica’ is a distinct species, the secondaries

of typical A. laverna being totally unrelieved by bright colouring, and much resembling

those of A. garuda.

5. ADOLTAS ASOKA, Felder, Reise der Nov., Lep. iil. p. 433, pl. 58. fig. 1 (1867). 3 ¢,

Malacca. ‘This species was not previously in the collection of the British Museum.

6. Apottas MoNINA, (¢ Papilio monina) Fabricius, Mant. Ins. ii. p. 51 (1787). 3 2,

Malacca. The female, although much like 4. puseda, is rather different. The following

is probably only a variation of 4. monina.

7. ADOLIAS LUDEKINGII, Vollenhoven, Tijd. Ent. v. p. 189, pl. 10. fig. 3 (1860).

3 2, Malacca.

540 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Genus SyMPHzADRA, Hiibner.

1. SYMPHZDRA DIRTEA , (Papilio dirtea) Fabricius, Ent. Syst. iii. 1, p. 59. n. 184

(1793). 32%, Malacca, 3, Penang.

“ Slices of cut pineapple placed along a road that ran by this jungle, were generally

sure in a short time, at the proper season, to be visited by a good supply of both males

and females.” “Old and fallen fruits of most kinds were attractive; but sliced pine-

apple was mostly used as bait” (W. LZ. Distant Hunt. Mo. Mag. vol. xii. p. 207).

Genus Tanatcra, Butler.

1. Tawaiticta arnuNA, (Adolias aruna) Felder, Wien. ent. Monatschr. iv. p. 400. n. 24

(1860). ¢, Malacca. Not previously in the Museum collection.

2. TANACIA PULASARA, (Adolias pulasara) Moore, Cat. Lep. E.I. C. i. p. 190 (1857).

3 %, Malacca. The male is very closely allied to 7. varwna of Vollenhoyen, but has

the apical area of secondaries white, and the band between the zigzag lines of primaries

broader.

Genus AtHyMA, Westwood.

1. ATHYMA LARYMNA, Westwood & Hewitson, Gen. Diurn. Lep. pl. 35. fig. 1 (1850).

3, Malacca.

2. ATHYMA LEUCOTHOH, (Papilio leucothoé) Linnzeus, Mus. Lud. Ulr. p. 292 (1764).

36, Penang.

3. ATHYMA KRESNA, Moore, P. Z.S. p. 12. n. 6, pl. 50. fig. 4 (1858). 3 2, Malacca.

4, ATHYMA SABRATA, Moore, P. Z.S. p. 18. n. 10, pl. 51. fig. 1 (1858). ¢ 2, Malacca.

5. ATHYMA NIVIFERA, n. sp. (Pl. LXIX. fig. 4.) Athyma nefte, Moore (ex parte),

P. Z. S.1858, p. 18. This species may be at once distinguished from A. nefte of Cramer

by the much narrower central band, which in the male is much more distinctly blue at

the edges; by the longer and narrower trifid subapical band on the male, and the deeper

colour of the underside. Expanse of wings 2 inches 5-9 lines. ¢ ?, Malacca. It is

also common in Borneo, and occurs at Assam.

6. Arayma amHarA, Druce, P. Z.8. p. 344, pl. xxii. fig. 2 (1873). ¢, Malacca. In

the examples from Malacca the discoidal white streak of primaries is reduced to one or

two white dots, connected by a pale brown streak.

7, ATHYMA CLERICA, n. sp. (Pl. LXIX. fig. 5.) Nearly allied to A. abiasa, but larger,

the spots of primaries more oblique and larger, the subbasal transverse white band of

secondaries narrower; the discal series of spots much larger, more inarched above anal

angle, forming a waved band, divided by the nervures; a well-marked greenish-grey

submarginal streak. Below much paler, with the differences of the upper surface; sub-

marginal series of lituree replaced by the submarginal streak, which is rosy greyish (not

eveenish, as above). Expanse of wings 2 inches 7 lines. 6, Malacca.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 541

8. ArHyma rp1Ta, Moore, P. Z.8. p. 16. n. 16, pl. 51. fig. 8 (1858). ¢ 2, Malacca.

9. ATHYMA PRAVARA, Moore, Cat. Lep. EI. C. i. p. 173, pl. 5a. fig. 4 (1857).

3, Malacca.

Genus Eurrpeus, Westwood.

1. Euripus EUpL@omES, Felder, Reise der Nov. Lep. iii. p. 415. n. 688 (1867).

3 %, Malacca. This species is a mimic of Calliplea diocletianus.

Genus EvLacura, Butler.

1. EULACURA OSTERIA, (Apatura osteria) Westwood, Gen. Diurn. Lep. p. 305, note

(1850). 3 ?, Malacca.

Genus LEBADEA, Felder.

1. LEBADEA ALANKARA, (Aconthea alankara) Horsfield, Cat. Lep. E.I. C. pl. 5. fig. 6

(1829). 3, Malacca. This appears to be the male of the succeeding species.

2. LEBADEA MARTHA, (Papilio martha) Fabricius, Mant. Ins. ii. p. 56. n. 555 (1787).

?, Malacca.

Genus Limenitts, Fabricius.

1. Limeniris PRocris, (Papilio procris) Cramer, ii. pl. 106. figs. E, F (1779).

$2, Malacca; ¢, Penang. ‘‘ Very common at Province Wellesley” (W. L. Distant).

Genus Neptis, Fabricius.

1. NEPTIS MAMAJA, n. sp. (Pl. LXIX. fig. 3.) Nearly allied to WV. ewrynome, but

always to be distinguished by the narrower externo-discal band of white spots in secon-

daries, colouring below rather brighter than in NV. ewrynome. Expanse of wings 1 inch

11 lines to 2 inches 4 lines. ¢ 2, Malacca and Penang. This appears to be the com-

monest Neptis in Malacca.

2. NEPTIS LEUCONATA, n. sp. (Pl. LXIX. fig. 1.) Wings above olive-brown, banded

and spotted with cream-colour; arrangements of markings as in WV. nata, but the bands

of secondaries wider and nearer together; pale brown intermediate streaks better defined.

Wings below slightly redder; bands and spots as above; intermediate streaks white.

Expanse of wings 2 inches. 6, Malacca.

3. NEPTIS GONONATA, n. sp. (PI. UXIX. fig. 2.) Wings of the same shape and with

the same spots and bands as NV. soma; but all these markings pure white instead of

sordid yellowish white. Wings below chocolate-brown; bands below straighter; sub-

marginal line of secondaries below white. Expanse of wings 2 inches 1 line. 3, Ma-

lacca. Intermediate in character between NV. nata and N. soma.

4. Neptis Nata, Moore, Cat. Lep. E.I. C. i. p. 168, pl. 4a. fig. 6 (1857).. 32,

Malacca.

Vikast Group.

5. Nepris viKasi, Horsfield, Cat. Lep. E.I. C. p. 168, pl. 5. figs. 2, 2a (1829).

3, Malacca.

SECOND SERIES.—ZOOLOGY, VOL. I. 4¢

5A2 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Columella Group.

6. Nupris CoLUMELLA, (Papilio colwmelia) Cramer, Pap, Hxot. iv. pl. 296. figs. A, B

(1782). & 2%, Malacca.

Tiga Group.

7. NEPTIS DORELIA, n. sp. (Pl. LXVIII. fig. 3.) Papilio hetiodore, Fabricius (nec

Cramer nee Moore), Mant. Ins. ii. p.52. n. 516 (1787). 5 %, Malacca.

The type of the Fabrician species differs from the pretargles of WV. tiga in the Horsfield

Cabinet from Java in having a single instead of a double central arched line across the

under surface of the secondaries; this line is, in some examples, slightly thickened.

8. Nepris T1r¢A, Moore, Proc. Zool. Soc. p. 4. n. 3 (1858). 3%, Malacca. This

species agrees with an example in the Banksian cabinet, but not with the Fabrician

type; there are two central brown curved lines across the underside of the secondaries,

partially or wholly filled in with paler brown.

9. Nupris PARAKA, n. sp. (Pl. LXVIII. fig. 2.) Almost exactly like W. hordonia

above, but brighter in colour, and with the tawny bands rather broader ; the submarginal

tawny streak of primaries deeply bisinuate so as to form two wide arches, and intersected

by a black line. ‘Wings below ochre-yellow, the black portions of the upper surface .

represented by brownish stains, and partially blotched with grey: the basal area of

secondaries crossed by two black oblique liturze; the centre band bordered by two inter-

rupted black lines; the outer border intersected (above the indication of the submarginal

tawny line of the upper surface) by a slightly waved black line. Expanse of wings 1 inch

7 to10 lines. & 2, Malacca. The typical N. heliodora very nearly resembles this

species on the upper surface ; but below it is very different.

10. NEPTIS DINDINGA, n. sp. (Pl. LXVIIL. fig. 6.) Larger than W. heliodora, the

bands broader, particularly the central band of secondaries, which is twice as broad and

extends nearly to the base; subapical patch of primaries twice as wide, and deeply

indented in the centre of its inner border; submarginal tawny streak replaced by pale

brown (but intersected by the black line) in the primaries, tawny but extremely slender

in the secondaries. Differences below much as. above; discal band of secondaries broad

and black, and intersected by a whity-brown streak. Expanse of wings 2 inches,

?, Malacca.

We have a specimen in the British Museum from Moulmein, which measures 2 inches

2 lines in expanse. It is a well-marked species,

11. Nepris norDoNIA, (3, deck hordonia) Stoll, Suppl. Cram. pL 33. figs. 4, 4 D

(1790). ¢, Malacca,

Genus PANnpITA, Moore.

1. Panprra stnore, Moore, Cat. Lep. E.I. C. i. p. 182, pl. 6a. fig. 3 (1857). ¢, Ma-

lacca; Penang.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 543.

Genus HypoLimnas, Hiibner.

1. HYPOLIMNAS BOLINA, (Papilio bolina) Linneeus, Mus. Lud. Ulr. p. 295 (1764).

3 ?, Penang.

2, HyPoLIMNAs INCOMMODA, n. sp. ¢& very similar to the male of ZH. bolina, but with

the subapical band of primaries straighter on the underside, and the pale brown sub-

marginal spots narrower and darker: expanse of wings 3 inches 1 line. ? differs from

the female of H. bolina in having a broad oblique subapical white band on the upperside

of primaries, a large diffused sordid white patch just beyond the cell of secondaries, and

the submarginal spots all separated, small, and pale brown: expanse of wings 3 inches

ll lines. 3& 2, Malacca.

This form seems to replace H. bolina in Malacca; in some respects it approaches the

Javan species H. nerina.

Genus CetTHostA, Fabricius.

1. CETHOSIA METHYPSEA, n. sp. Allied to OC. hypsina and C. penthesilea: from the

former it differs in its smaller expanse, the regularity of the subapical oblique whitish

band of primaries, in the presence of an apical submarginal series of four whitish spots

connecting the lower extremity of this band with the costa, and in the much less pro-

minent angulation of the outer edge of the red band on the under surface of secondaries ;

from the latter it differs in its smaller expanse, much brighter coloration, the yellower

tint of the subapical band of primaries, the more irregular and much more broadly

white-bordered red band below; the primaries are alse more produced. Expanse of wings

2 inches 9 lines to 3 inches 3 lines. ¢, Penang. The larger measurement is taken

from examples previously in the collection. We have a female from Assam.

2. CETHOSIA HYPSINA, Felder, Reise der Nov., Lep. iii. p. 885 (1867). 3 2, Malacca.

Also in the Museum collection from Nepal and Assam.

Genus CrrrocHroa, Doubleday.

1. CrRRocHROA oRISSA, Felder, Wien. ent. Monatschr. iv. p. 899 (1863); Reise der

Nov., Lep. iii. pl. 49. figs. 7. 8 (1867). 3°, Malacca; Penang.

2. CIRROCHROA JOHANNES, Butler, P. Z. 8. p. 221, pl. 17. fig. 10 (1868). 3 2, Ma-

lacca.

3. CIRROCHROA BAJADETA, Moore, Cat. Lep. E.I. C. i. p. 150, pl. 3a. fig. 3 (1857).

?, Malacca. This seems to be Felder’s C. malaya.

4, CIRROCHROA ROTUNDATA, n. sp. ¢. Nearly allied to C. mithila of Moore, the

wings more rounded, primaries less produced; markings of primaries above almost

obsolete; outer undulated line of secondaries much more distinct; below all the bands

and spots tawny instead of ochre yellow: expanse of wings 2 inches 11 lines. Malacca.

I have no doubt that this species is distinct from C. mithila; the coloration of the bands

below is strikingly different.

Ac 2

544 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Genus TrERINOoS, Boisduyal.

1. TERINOS TEUTHRAS, Hewitson, P. Z. 8. p. 89 (1862). 2, Malacca.

2. TERINOS ROBERTSIA, Butler, Ann. & Mag. Nat. Hist. ser. iii. vol. xx. p. 400, pl. 8.

figs. 2-4 (1867). 3%, Malacca. This species appears to be tolerably common. Mr.

Distant took both the above butterflies at Province Wellesley.

Genus Messaras, Doubleday.

1. MESSARAS ERYMANTHIS, (Papilio erymanthis) Drury, Il. Ex. Ent. i. pl. 15. figs. 3, 4

(1773). 3%, Malacca; Penang.

Genus ATELLA, Doubleday.

1. ATELLA PHALANTA, (Papilio phalanta) Drury, Ill. Ex. Ent. i. pl. 21. figs. 1, 2 (1778).

3 ?, Malacca; Penang.

2. ATELLA sInHA, (Terinos sinha) Kollar, Hiigel’s Kaschm. iv. 2, p. 438 (1848).

3, Malacca.

Genus Cyntuta, Fabricius.

1. CYNTHIA EROTELLA, n. sp. Allied to C. erota, but much smaller, brighter in colour;

the primaries generally comparatively shorter, the costal margin more arched, the ocelli

smaller in the male, and the discal lunulated line much less strongly defined; secon-

daries of the female with four ocelli, with broad bright orange irides. Wings below

slightly more reddish in colouring, with scarcely a trace of pearly grey clouding on the

external area. Expanse of wings, ¢ 3 inches 2-8 lines, ? 3 inches 7 lines. 3 ?, Ma-

lacca.

I have examined a good series of examples, and am therefore confident of the dis-

tinctness of this species. In general size and the indistinctness of the lunulated discal

line it agrees with C. deione; but it is altogether brighter in colour, with longer tails

to secondaries, and a different arrangement of the lines on the under surface of the

wings. Mr. Distant took it at Province Wellesley.

Genus PARTHENOS, Hiibner.

PARTHENOS LILACINUS, n. sp. (Minetra gambrisius, Hewitson, Gen. Diurn. Lep. pl. li.

fig. 2). Differs from P. gambrisius in the lilac internal area of primaries and basal area

of secondaries above; in the blue-green basal areas of secondaries below, and the much

better defined series of black liturze beyond the hyaline band of primaries. Expanse of

wings 4 inches 2 lines. 3o ?, Malacea ; Penang.

I have seen long series of both the Indian and Malayan forms, and am therefore

satisfied that the differences are constant. The Fabrician type agrees with our examples

from Silhet.

Genus Crrust1s, Boisduval.

1. CyrestTIs RAHRIA, Moore, Cat. Lep. H.I. C.i. pl. 8a. fig. 2 (1857). 3 2, Malacca;

?, Penang.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 545

2. CYRESTIS NIVEA, (Amathusia nivea) Zinken-Sommer, Nova Acta. Acad. Cur. xvi.

p. 138, pl. 14. fig. 1 (1831). 3, Malacca. ‘“ Not scarce at Province Wellesley ” (W. L.

Distant).

Genus Ereoris, Boisduval.

1. ERGoLIs ARTADNE, (Papilio ariadne) Linnzeus, Syst. Nat. i. 2, p. 778. n. 170 (1767).

3 ?, Malacca; ¢, Penang.

2. ERGOLIS MERIONE, (Papilio merione) Cramer, Pap. Exot. ii. pl. 144. figs. G, H (1779).

$, Malacca; Penang.

Genus JunoniA, Hiibner.

1. Junonta rpHita, (Papilio iphita) Cramer, Pap. Exot. iii. pl. 209. figs. C, D (1782).

$ ?, Malacca; ¢, Penang.

2. JUNONIA LAOMEDIA, (Papilio laomedia) Linneus, Syst. Nat. i. 2, p. 772. n. 145

(1767). 3%, Malacca. ‘Common at Province Wellesley ” (W. L. Distant).

3. JUNONIA LEMONIAS, (Papilio lemonias) Linnzeus, Mus. Lud. Ulr. p. 277 (1764).

36, Malacca; ¢? 3, Penang.

4, JUNONIA oORITHYA, (Papilio orithya) Linneeus, Mus. Lud. Ulr. p. 278 (1764).

3 2, Malacca. ‘‘Common at Province Wellesley” (W.L. Distant).

5. JUNONIA ASTERIE, (Papilio asterie) Linnzeus, Syst. Nat. i. 2, p. 769. n. 183 (1767).

3 2, Malacca; Penang.

Genus RHINOPALPA, Felder.

1. RurNopatpa FULYA, Felder, Wien. ent. Monatschr. iv. p. 399. n. 21 (1860).

3, Malacca.

Family ERYCINID A.

Subfamily NemeEosi1inz, Bates.

Genus ZmmeERos, Boisduval.

1. ZemEros EMESOIDES, Felder, Wien. ent. Monatschr. iv. p. 396. n. 10 (1860); Reise

der Nov., Lep. ii. p. 289, pl. 36. figs. 9-11 (“1865”). 32, Malacca. This species is

evidently very common.

2. ZeMEROS ALBIPUNCTATA (PI. LXIX. fig. 10), Butler, Cist. Ent. i. p. 236 (1874).

3 2, Malacca. We have the female of this species from Borneo. It seems to be a

common species.

Genus ABIsARA, Felder.

1. ABISARA sAvirRI, Felder, Wien. ent. Mon. iv. p. 897. n.12 (1860). 39, Ma-

lacea. This is quite distinct from the Indian species A. susa of Hewitson.

2. ABISARA KAUSAMBI, Felder, Wien. ent. Mon. iv. p. 397, n. 11 (1860). 392, Ma-

lacca. A common species, the male of which is very different from 4. echerius.

546 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

3. ABISARA HAQUINUS (Papilio haquinus), Fabricius, Ent. Syst. iii. 1, p. 55. n. 169

ie @, Malacca.

Subgenus Laxrra.

4. ABISARA TANITA, (Taxila tanita) Hewitson, Ex. Butt. ii. Tax. pl. i.(1861). ¢, Ma-

lacea. The rounded wings of this and the allied species distinguish them from typical

Abiscra; but Iam doubtful of its generic distinction. TZ. orphna, placed by Kirby as

type of Taxila, must, as I have shown elsewhere, give place to 7. egeon or T. fylla,

T. orphna having been indicated in the ‘ Genera of Diurnal Lepidoptera’ as not the type

of the genus.

Family LYCAINID A.

Subfamily Lycanina, Butler.

Genus Porrrra, Moore.

1. PortTIA SUMATRA, (Pseudodipsas sumatre) Welder, Reise der Nov., Lep. ii. p. 259,

pl. 86. figs. 24-26 (1865). ¢, Malacca; ¢, Penang.

Genus Grrypus, Boisduval.

1. Gerypus Horsrrerpi, (Miletus Horsfieldi) Moore, Cat. Lep. E.I. C. i. p. 19, pl. 1a.

fig. 21857). ¢ 2, Malacca; ¢, Penang.

2. GERYDUS NIVALIS ?, (Miletus nivalis) Druce, P. Z. 8. p. 348. n. 4 (1873). Malacca.

I cannot identify this species with certainty; our example measures only 10 lines in

expanse of wing.

3. GERYDUS SYMETHUS, (Papilio symethus) Cramer, Pap. Hxot. i. pl. 149. figs. B, C

(1779). 3, %, Malacca; ?, Penang.

Genus Anops, Boisduval.

1. Anops maLayica, Felder, Reise der Nov., Lep. ii. p. 221, pl. 28. fig. 18 (1865).

36 ¢, Malacca.

2. ANOPS SPERTHIS, (Anops sperthis) Felder, Reise der Nov., Lep. ii. p. 222 (1865).

3, Malacca; ?, Penang.

Genus Lipuyra, Westwood.

1. LipHYRA BRASSOLIS, Westwood, Proc. Ent. Soc. p. 31 (1864). ¢, Malacca.

Genus Lycmnopsts, Felder.

1. LycaNopsis HARALDUS, (Papilio haraldus) Fabricius, Mant. Ins. i. p. 82. n. 744

(1787). , Malacca. This is L. ananga of Felder.

Genus CasraLius, Hubner.

1. Casratius Rostmon, (Papilio rosimon) Fabricius, Syst. Ent. p. 523. n. 341 (1775).

3 2, Malacca; Penang.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 547

2. CASTALIUS ETHION, (d Lycena ethion) Westwood & Hewitson, Gen. Diurn. Lep.

pl. 76. fig. 8 (1852). 2, Above very like C. roxus, but with the white band extending

nearly to the costa of primaries, internally excavated and externally broadly truncate-

clayate at its upper extremity; below white, spotted with dark brown, as in the male.

Expanse of wings 11 lines. ¢?, Malacca.

3. CASTALIUS ROXUS, (Polyommatus roxus) Godart, Enc, Méth. ix, p. 659. n. 14.2 (1829),

Horsfield, Cat. pl. 2. figs. 4, 4.a (1828). & ?, Malacca,

Genus Lamprpes, Hiibner,

1. Lampipes Patan, (Lycena patala) Kollar, Hiigel’s Kaschmir, iv. 2, p. 419 (1848).

3, Malacca.

2. LAMPIDES MACROPHTHALMA, (Lycena macrophthalma) Felder, Verh. zool.-bot. Ges.

xii. p. 483. n. 115 (1862). 3, Malacca,

3. LAMPIDES BEROH, (Lycena beroé) Felder, Reise der Nov., Lep. ii. p. 275, pl. 34,

fig. 36 (1865). 3 2, Malacca.

4. Lamprpes atura, (Cupido aluta) Druce, P. Z. 8. p. 349. n. 16, pl. xxxii. fig. 8 (1878).

32, Malacca. Nearly allied to Z. nora of Felder, from Amboina, but smaller, with

less acuminate primaries.

5. LAMPIDES ALMOoRA?, (Cupido almora) Druce, P. Z.8. p, 349. n. 14, pl. xxxii. fig. 7

(1873). 3, Malacca.

6. Lamprpes BzxTIcUS, (Papilio beticus) Linneeus, Syst. Nat. i. 2, p. 789 (1767).

3 ?, Penang.

7. LAMPIDES CHRULEA, (Cupido cerulea) Druce, P. Z.8, p. 349. n. 18, pl. xxxii. fig. 6

(1873). 3, Malacca.

8. Lampipes AGNATA, (Cupido agnata) Druce, P. Z.8. p. 106, pl. xvi. figs. 2-4 (1874).

3 2, Malacca.

9. LAMPIDES PSEUDELPIs, n. sp. (Pl. LX VIII, figs. 7,8), Nearly allied to Z. elpis, which

it resembles above; below, however, with the white transverse bands narrower, irre-

cular, and broken up, the ground-colour more distinctly bluish opaline, and the large

submarginal black spot of secondaries less broadly encircled by ochreous. Expanse of

wings 1 inch 6 lines. 32, Malacca,

10. LAMPIDES KANKENA ?, (Lycena kankena) Felder, Verh. zool.-bot. Ges. xii. p. 481

(1862). 2, Malacca,

Genus CATAPACILMA, n, gen,

Allied to Lampides and Miletus, but differs from both in having three tails to the

secondaries; the antenne are long, slender, and acuminate; the wing-cells and neura-

tion generally are very like that of Lampides; the wings below are spangled with

silver, much as in Wiletus (Hypochrysops, part, Felder).

548 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

1. CATAPHCILMA ELEGANS, (Hypochrysops elegans) Druce, P. Z. 8S. p. 350, pl. xxxii.

fig. 12 (1873). 3, Malacca; Penang. The type was described from Borneo. The

figure, which is extremely rough, represents the species with three to four tails to

secondaries; but either the fourth has been broken off in all the examples which I

have examined, or does not really exist.

Genus Lycawna, Fabricius.

1. Lycmya cacaya, Felder, Reise der Nov., Lep. ii. p. 278, pl. 34. figs. 11-18 (1865).

2, Malacca.

2. LyCmNA LyYSIZONE, Snellen, Tijd. voor Ent. pl. 7. fig. 2 (1876). 3, Malacca;

Penang. Rather larger than L. karsandra; paler below.

3. LycmNA KARSANDRA, Moore, P. Z. 8. p. 505, pl. 31. fig.7 (1865). ?, Penang.

4. Lycm@NA SANGRA, (Polyommatus sangra) Moore, P. Z.8. p. 772, pl. xli. fig. 8 (1865).

3, Malacca; ?, Penang.

Subfamily Tueciinz, Butler.

Genus AmpBiyropiaA, Horsfield.

1. AMBLYPODIA CENTAURUS, (Papilio centaurus) Fabricius, Syst. Ent. p. 520. n. 329

(1775). 3, Malacca; ?, Penang.

2. AMBLYPODIA EUMOLPHUS, (Papilio ewmolphus) Cramer, Pap. Exot. iv. pl. 299. figs.

G, H (1872). 32, Malacca.

3. AMBLYPODIA ANTHELUS, Westwood, Gen. Diurn. Lep. pl. 74. fig. 6 (1852). 3, Ma-

lacca. |

4, AMBLYPODIA ADATHA, Hewitson, Cat. Lyc. B. M. pl. 4. figs. 29-31 (1862). ¢ 2, Ma-

lacca. ‘This is quite distinct from A. micale.

5. AMBLYPODIA ATosIA, Hewitson, Ill. Diurn. Lep. p. 9, pl. 2. figs. 8, 9 (1868).

36 2, Malacca.

6. AMBLYPODIA AMPHIMUTA, Felder, Wien. ent. Mon. iv. p. 396. n. 6 (1860).

3 ¢, Malacca.

7. AMBLYPODIA EPrmuTA, Moore, Cat. Lep. Mus. E.I. C. i. p. 42. n. 62 (1857).

3 2, Malacca.

8. AMBLYPODIA ANTIMUTA, (Arhopala antimuta) Felder, Reise der Nov., Lep. ii. p. 233

(1867). ¢, Malacca.

9, AMBLYPoDIA AROA, Hewitson, Il]. Diurn. Lep. p. 18, pl. 2. fig. 12 (1863). 3, Ma-

lacea. A species from India, labelled by Mr. Hewitson as 4. aroa, is more nearly allied

to A. epimuta.

10. AMBLYPODIA METAMUTA, $, Hewitson, Il]. Diurn. Lep. p. 18, pl. 2. figs. 154, 155

(1863). 3%, Malacca. The whole upper surface of the male is coloured deep purple

as in the primaries of the female, but without the black border.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 549

11. AmpLypop1a HYPoMUTA, Hewitson, Cat. Lyc. Brit. Mus. i. p. 11, pl. 6. figs. 63, 64

(1862). 32, Malacca; ¢, Penang, Quite distinct from 4. amphimuta.

12. AMBLYPODIA APIDANUS, (Papilio apidanus) Cramer, Pap. Exot. ii. pl. 187. figs. F, G

(1779). 3, Malacca.

13. Amspiypop1 ? D1arpi, Hewitson, Cat. Lyc. B. M. p. 9, pl. 5. figs. 41, 42 (1862).

$, Malacca. This species would, I think, be better placed in Uéica.

Genus Hypotycana, Felder.

1. Hypotyca#Nna ERYLUS, (Polyommatus erylus) Godart, Enc. Méth. ix. p. 638. n. 60

(1823). 32, Malacca; 2, Penang.

2. HypotycmNa? erotus, (Papilio etolus) Fabricius, Mant. Ins. ii. p. 66 (1787).

$, Malacca. This species appears to me to agree better with Wyrina.

Genus ApHnzvs, Hiibner.

1. APHN@US LOHITA, (Amblypodia lohita) Horsfield, Cat. Lep. Mus. E.I. C. p. 106.

n. 88 (1829). 3&2, Malacca; Penang.

Genus Drvporrx, Hewitson.

1. DEvDORIx PETOSIRIS, Hewitson, Ill. Diurn. Lep. p. 22, pl. 9. figs. 30, 31 (1863).

2, Malacca.

2. DEUDORIX PHERETIMA, Hewitson, Ill. Diurn. Lep. p. 21, pl. 9. figs. 27-29 (1863).

3, Malacca.

3. DEUDORIX JARBAS, (Papilio jarbas) Fabricius, Mant. Ins. ii. p. 68. n. 648 (1787).

3, Malacca. In D. jarbas the veins of the primaries are blackened, which readily distin-

guishes it from D. melampus.

4. DEuDoRIx pomitia, Hewitson, Ill. Diurn. Lep. p. 19, pl. 6. figs. 6, 7 (1863),

3, Malacca.

Genus Ionaus, Htibner.

1. Ionaus Lonernus, (Hesperia longinus) Fabricius, Ent. Syst. Suppl. p. 4380 (1798).

?, Penang.

Genus Myrina, Fabricius.

1. Myrina mecistrA (?), Hewitson, Ill. Diurn. Lep. Suppl. p. 5, pl. 3. figs. 77, 78 (1869).

Malacca.

2. Myrina TRAVANA, Hewitson, Ill. Diurn. Lep. p. 38, pl. 17. figs. 59, 60 (1865).

3 2, Malacca. The male of this is like a Dewdorix on the upper surface, the female

like an Amblypodia.

3. Myrina cuirra, (Zhecla chitra) Horsfield, Cat. Lep. E.I. C. p. 97, pl. 1. fig. 5

(1829). ?, Malacca.

4, MyrInA MARCIANA, Hewitson, Ill. Diurn. Lep. p. 34; pl. 12. figs. 12, 13 (1868).

3, Malacca.

5. Myriva Lapiruis, Moore, Cat. Lep. Mus. E.I. C. i. p. 48. n. 79 (1857). 3, Malacca.

SECOND SERIES.—ZOOLOGY, VOL. I. 4D

550 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

6. Myrina THARIS, (Oxylides tharis) Hubner, Zutr. ex. Schmett. figs. 883, 884 (1837).

3 2, Malacca.

7. Myrina amrita, Felder, Wien. ent. Mon. iv. p. 395 (1860). ¢ ?, Malacca.

8. Myrina FREJA, (Hesperia freja) Fabricius, Ent. Syst. iii. 1, p. 263 (1798). 2, Penang.

Genus Loxura, Horsfield.

1. Loxura atymnus, (Papilio atymnus) Cramer, Pap. Exot. iv. pl. 331. figs. D, EH

(1784). 32, Malacca; 3, Penang.

Family PAPILIONIDA.

Subfamily Prerina, Bates.

Genus DrxtiAs, Hubner.

1. DELIAS METARETE, n. sp. Nearly allied to D. hyparete, but longer in the wing, the

greyish apical area of primaries more uniform, in the female with a series of five or six

whitish (not yellowish) decreasing streaks between the veins; external border of second-

aries pale grey in the male, broadly blackish in the female, the red markings distinctly

visible. Apical spots of primaries below bounded within by a slender angulated greyish

line: internal area of secondaries of a much brighter yellow colour; scarlet submarginal

spots about twice as large at anal angle, not extending above the second subcostal

branch, bounded inwardly by a slender interrupted black streak. Expanse of wings

3 inches 2 lines. 3 ?, Malacca; Penang. ‘This species occurs also in Borneo.

2. Dewias pionE, (3, Papilio dione) Drury, Ill. Ex. Ent. ii. pl. 8. figs. 3, 4 (1778).

?, Malacca. The female resembles that of D. egialea, excepting that the bands are white

instead of orange.

Genus Trertias, Swainson.

1. TeRtas Formosa, (Hurema formosa) Hubner, Zutr. ex. Schmett. figs. 979, 980

(1837). 3 ?, Malacca.

2. TmrtAs sari, Horsfield, Cat. Lep. Mus. E.I. C. p. 186 (1829). 3%, Malacca.

“Occurs at Province Wellesley ” (W. L. Distant).

8. TeRIAs HECABEOIDES, Ménétriés, Cat. Mus. Petr., Lep. i. p. 85, pl. 2. fig. 2 (1855).

3%, Malacca; 3, Penang.

4, TERIASINANATA, Butler, P. Z. S. p. 617. n. 35 (1875). %, Malacca (3 examples).

5. TeRtAs pumIvaRts, Butler, P. Z. 8. p. 617. n. 36, pl. lxvii. fig. 7 (1875). 3, Malacca.

I cannot find any distinguishing characters to separate the two preceding Malayan forms

from their representatives in the New Hebrides.

Genus CatopsritA, Hiibner.

1. Caropsri1a crocate, (Papilio crocale) Cramer, Pap. Exot. i. pl. 55. figs. C, D (1779).

3%, Malacca; ¢, Penang.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 5d51

2. CATOPSILIA CATILLA, (Papilio catilla) Cramer, Pap. Exot. iii. pl. 229. figs, D, E

(1782). 3%, Malacca; 2, Penang.

3. CATOPSILIA CHRYSEIS, (Papilio chryseis) Drury, Ill. Ex. Ent. i. pl. 12. figs. 3, 4

(1773). 3&2, Malacca; Penang.

4, CATOPSILIA SCYLA, (Papilio scylla) Linneeus, Mus. Lud. Ulr. p. 242 (1764). 2, Ma-

lacca; 3, Penang. The specimens are all of the C. etesia type, having the secondaries

bright golden orange; one example nearly approaches the brighter specimens of typical

C. scylla.

Genus Hrpomora, Hiibner.

1. Hezomora GLAUCTIPPE, (Papilio glaucippe) Linneeus, Mus. Lud. Ulr. p. 240 (1764).

3, Matacca.

Genus Appras, Hiibner.

1. APPIAS PLANA, n. sp. 6d. Constantly differing from the Javan A. leptis of

Felder in the absence of the black border of secondaries, a trace only of which exists at

the apex of these wings; also larger, the primaries more produced; the apex of primaries

below and the secondaries of a clearer cream-colour: expanse of wings 2 inches 8 lines.

6, Malacca. This species is also common in Borneo.

2. APPIAS CARDENA, (Pieris cardena) Hewitson, Ex. Butt. ii. Pier. pl. 3. figs. 17, 18

(1861). ¢, Malacca. Also common in Borneo.

3. APPIAS ELEONORA, (Pieris eleonora) Boisduval, Sp. Gén. Lép. i. p. 481. n. 64 (1836),

3%, Malacca; 2, Penang. The female nearly resembles that sex of A. enarete.

4, APPIAS NATHALIA, (Pieris nathalia) Felder, Wien. ent. Monatschr. vi. p. 285 (1862).

$ %, Malacca. Occurs also at Singapore. The type described from the Philippines.

5. Appras PANDA, (Pieris panda) Godart, Enc. Méth. ix. p. 147, n. 102 (1819).

$, Malacca. This appears to be only the worn male of 4. nathalia; but the black border

seems narrower.

6. APPIAS NERO, (Papilio nero) Fabricius, Ent. Syst. rig) Ls 708 sayy ag Zl (Gl),

8 %,Malacca; ¢, Penang. Occurs also in Java and Siam.

7. ApprAas FIGULINA, (2 Pieris figulina) Butler, Ann. & Mag. Nat. Hist. ser. 3, vol. xx.

p- 399, pl. 8. fig. 1 (1867). 3%, Malacca. The male is brighter in colour than

A. nero &, and golden yellow, with a greyish streak connecting the first and second

branches of primaries, and usually a greyish indistinct nebula across the disk of second-

aries. The'type of A. figulina was from Singapore; the species occurs also in Borneo.

Genus BELENOIS, Hithbner.

1. Brnenors oynts, (Pieris cynis) Hewitson, Ex. Butt. iii. Pier. pl. 8. fig. 54 (1866).

3%, Malacca. The form figured by myself in the ‘Transactions of the Entomological

Society’ may, I think, turn out to be distinct.

4D 2

552 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Subfamily Papitioninz, Bates,

Genus ORNITHOPTERA, Boisduyal.

1. ORNITHOPTERA RUFICOLLIS, nu. sp. 6. Allied to O. flavicollis: wings smaller, com-

paratively narrower; outer margin of primaries more distinctly inarched; collar carmine.

Expanse of wings 5 inches 4 lines to 6 inches 1line. ¢ Malacca.

Unfortunately no female examples have come; the male seems to be not uncommon.

Mr. Distant has an example of the female from Province Wellesley. The primaries are

of a rather more rounded form and duller black colour than those of the male; the streaks

are more grey in colouring; the secondaries have a broad deeply indented marginal

border, six large discal spots, the veins and the base black. It was taken on the 21st of

February 1869, on the lower part of “ Batu Kawan.”

Genus PAPILIO, Linneus.

1. PAPILIo ANTIPHATES, Cramer, Pap. Exot. i. pl. 72. figs. A, B(1779). ¢ 2, Malacca.

2. PAPILIO SARPEDON, Linnzus, Mus. Lud. Ulr. p. 196 (1764). ¢ ¢, Malacca; Penang.

3. PaprLio EvEMON, Boisduval, Sp. Gén. Lép. i. p. 234 (1836). 3 2, Malacca.

4, PAPILIO AXION, Felder, Verh. zool.-bot. Ges. xiv. p. 305. n. 224, p. 350. n. 128

(1864). 3¢, Malacca. A vwariety of this species occurs in which the bands and spots

are cream-coloured.

5. Paprito BATHYCLES, Zinken-Sommer, Nova Acta Acad. Nat.-Cur. xv. p. 157, pl. 14.

figs. 6, 7 (1881). ¢, Malacca. ‘‘ P. bathycles is common round puddles in wet

weather ” (W. L. Distant).

6. Paprnio ARYCLES, Boisduval, Sp. Gén. Lép. i. p. 231. n. 51 (1836). o, Malacca.

One example was taken at Province Wellesley by Mr. Distant.

7. PapinIo AGAMEMNON, Linneus, Mus. Lud. Ulr. p. 202 (1764). 39, Malacca;

?, Penang.

8. PAPILIO MALAYANUS, Wallace, Trans. Linn. Soe. xxv. p. 59 (1865). od ?, Malacca;

$, Penang. A distinct species, like P. sthenetus above, but more like P. erithonius

below. “‘ Excessively common in gardens, August and September” (W. L. Distant).

9. PAPILIO DEMOLION, Cramer, |Pap. Exot. i. pl. 89. figs. A, B (1779). 3 2, Malacca.

10. Paritio DreLEsseRrtTI, Guérin, Deless. Souv. Voy. d. lInde, ii. p. 68, pl. 17

(1843). 3, Malacca.

11. Papriio ctytta, Linnzeus, Mus. Lud. Uly. p. 296 (1764). 3, Penang.

12. Papriio potytes, Linnzeus, Mus. Lud. Ulr. p. 186 (1764). 3 ?, Malacca; ¢, Pe-

nang. Var. P. stichiws, Hibner, Samml. exot. Schmett. (1806-16). 3%, Malacca.

The male of the latter form has the yellow spots well defined on the underside of

secondaries, just as the female has a greater amount of red colouring on the under-

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 553

side than than typical P. polytes. I suspect this variation to be seasonal, and

not polymorphic.

18. PApiILio HELENUS, Linnzeus, Mus. Lud. Ulr. p. 185 (1764). ¢ 2, Malacca.

14, Papriio PREXASPES, Felder, Reise der Noy., Lep. i. p. 107, pl. 15. fig. d (1865).

$¢ alacca. Not previously in the Museum collection.

15. Paprnio iswaRa, White, Entom. i. p. 280 (1842). o, Malacca; ¢ ?, Penang.

16. Paprnio sATURNUS, Guérin, Deless. Souv. Voy. d. l’Inde, ii. p. 70, pl. 18 (1848).

6, Malacca.

17. Pariuio EsPert, n. sp. (Pl. LXVIILI. fig. 7.) 3, Papilio protenor (part), Esper

[nec Cramer], Ausl. Schmett. pl. 29. fig. 2 (1785-98). ¢. Primaries above grey, with a

diffused broad subapical white band, the base, veins, and broad streaks between them

black, a red patch at base of discoidal cell; secondaries precisely as in the male, but with

a red spot on anal angle; differences below as above: expanse of wings 5 inches 10 lines.

$ 2, Malacca; ?, Penang.

18. Paprito mEstor, (3, Papilio androgeos), Cramer, Pap. Exot. i. pl. 91. figs. A, B

(1779). Iliades mestor, Hiibuer, Verz. bek. Schmett. p. 89. n. 931 (1816). ¢. Differs

from the female of P. Esper in having the white patch of primaries transferred to the

external angle, and the wings longer; below, the differences are the same as in the

male. Expanse of wings § inches 5 lines. ¢ ?, Malacca; 3, Penang.

19. Paprnio acHaTES, (? Papilio achates) Cramer, Pap. Exot. ii. pl. 182. figs. A, B

(1779). 3%. Wings much more uniform in colour than in the two preceding species, a red

basal streak as in P. androgeos; the red submarginal spots below much more restricted,

as in the female: expanse of wings 5 inches 6 lines. ¢ ?, Malacca.

Most Lepidopterists have united the three preceding forms in their collections as

polymorphic varieties of P. agenor, Linnzeus, noticing only the differences of the

females. I am satisfied that all three (with other forms referred to P. memnon, agenor,

and protenor) are distinct species.

20. PaPInio VARUNA, White, Entom. i. p. 280 (1842). ¢2 Malacca; Penang. The

type was described from Penang, where it is evidently common.

21. PAPILIO DIPHILUS, Esper, Ausl. Schmett. pl. 40 B. fig. 1 (1785-98). ¢ ?, Malacca;

Penang.

Family HESPERITD A.

Genus Casyapa, Kirby.

J. Casyapa THRAX, (Papilio thraz) Linneeus, Syst. Nat. i. 2, p. 794 (1767). 3, Malacca.

2. CASYAPA IRAVA (Hesperia irava), Moore, Cat. Lep. E.I. C. i. p. 254 (1857).

3 ?,,Malacca.

554 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Genus Hesperia, Fabricius.

1. HESPERIA HARISA, (Jsmene harisa) Moore, Proc. Zool. Soc. p. 782 (1865). 2, Malacca.

2. Husreria virra, Butler, Trans. Ent. Soc. p. 498 (1870); Lep. Exot. pl. lix. fig. 9

(1874). o&, Malacca.

The examples from Malacca have two minute transparent dots placed obliquely beyond

the cell of primaries. I have no opportunity of comparing the type at present, and

therefore do not know if there is any trace of these dots in the Bornean form.

3. HESPERIA BADRA, (Goniloba badra) Moore, P. Z.8. p. 778 (1865). 3%, Malacca.

Genus Copanus, Hiibner.

1. Copatus ELIA, (Hesperia elia) Hewitson, Trans. Ent. Soc. ser. 3, vol. ii. p. 489. n.9

(1866). o%, Malacca. The type was described from Sumatra.

2. COBALUS CILIATUS, n. sp. Dark chocolate-brown; primaries with two small spots

placed obliquely upon the median interspaces, and three dots inan angular line beyond the

cell hyaline white; secondaries with a broad abbreviated transverse band from the ab-

dominal margin, and the fringe (excepting at apex) white; body above greyish; head and

prothorax shot with golden green; abdomen irregularly white at the sides. Primaries

below as above; secondaries crossed by a broad angular white band, its lower half twice

as broad as its upper, connected with the white fringe by a white spur upon the second

median interspace; body below white, abdomen laterally banded with black lituree, anus

brown. Expanse of wings 1 inch 7 lines. ¢, Malacca.

Genus PampuiLa, Fabricius.

Section Gegenes, Hibner.

1. Pampnia arta, (Lsmene aria) Moore, P. Z. 8. p. 784 (1865). 32, Malacca. The

type was from Bengal.

2. PAMPHILA JULIANUS, (Hesperia julianus) Latreille, Ene. Méth. ix. p. 763. n. 99

(1823). <3. Malacca. Originally discovered in Java.

3. PAMPHILA MATTHIAS, (Hesperia matthias) Fabricius, Hut. Syst. Suppl. p. 483 (1798).

3%, Malacca; 3, Penang.

Section Pamphila (typical).

4, PAMPHILA AUGIAS, Linnzeus, Syst. Nat. i. 2, p. 794. n. 257 (1767). &, Malacca;

Penang.

5. PAMPHILA MASOIDES, n. sp. Above very like P. mesa of Moore, deep chocolate-

brown, banded and spotted with tawny; below similarly marked to P. mesa, but suffused

with deep tawny throughout. Expanse of wings 1 inch 1 line. 3 ?, Malacca. This

species is at once distinguished from P. mesa by the tawny colour of its bands (in

P. mesa they are ochreous); the outer band of primaries is also somewhat less irre-

gular; and the ground-colour of the underside is tawny instead of yellow.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 55d

6. PaMPHILA MARO, (Hesperia maro) Fabricius, Ent. Syst. Suppl. p. 482 (1798).

3, Malacca; 5 ?, Penang.

7, PAMPHILA NIGROLIMBATA, (Thymelicus nigrolimbatus) Snellen, Tijd. voor Ent.

p- 165, pl. 7. fig. 5 (1876). ¢ ‘Malacca.

Genus Piastinera, Butler.

1. PLASTINGIA CALLINEURA, (Hesperia callineura) Felder, Reise der Nov., Lep. iii.

p. 578, pl. 71. figs. 9, 10 (1867). 3, Malacca.

Genus CycnLoprpsEs, Hiibner.

1. CycLoPrpEs SALSALA, (Msoniades salsala) Moore, P. Z. 8.786 (1865). ¢ 2, Malacca.

Previously known from Bengal.

Genus Astictorrervs, Felder.

1. AstIcTOoPTERUS JAMA, Felder, Wien. ent. Mon. iv. p. 401. n. 29 (1860). 3, Malacca.

2. ASTICTOPTERUS XANITES. (Pl. LXIX. fig. 7.) %. Astictopterus xanites, Butler,

Trans. Ent. Soc. p. 510 (1870). 32, Malacca. The type was from Sarawak; the male

is rather smaller, and has the orange band of primaries abbreviated and of a deeper

colour.

3. ASTICTOPTERUS GEMMIFER, n. sp. Chocolate-brown; primaries with a broad post-

median bright orange band; end of cell and apical area of primaries and disk of

secondaries spotted, in certain lights, with shining amethyst-coloured spots. Hxpanse of

wings 1 inch 6 lines. ¢ ?, Malacca (5 examples).

4, ASTICTOPTERUS ARMATUS, Druce, P. Z.S. p. 359. n. 3, pl. xxxiii. fig. 7 (1878).

?, Malacca. The type was described from Borneo.

5. ASTICTOPTERUS DiocLES, (Nisoniades diocles) Moore, P.Z.S. p. 787 (1865).

g-%, Malacca. The type was-from Bengal.

. 6. ASTICTOPTERUS sinDU, Felder, Wien. ent. Mon. iv. p. 401. n. 30(1860). ¢, Malacca.

In this species the orange band is transverse, which at once distinguishes it from the

male of A. wanites. It is also smaller.

7. ASTICTOPTERUS STELLIFER, n. sp. Above much like a small 4. jama, deep choco-

late-brown. Below irrorated, particularly at apex of primaries, with bronzy golden

scales; internal area of primaries whity-brown; a small white spot at the end of the

cell in all the wings, and a small dull white spot close to the middle of the submedian

vein of secondaries; body below pale brown; palpi clothed with golden hair-scales ;

antennze black above, yellow below. Expanse of wingslinch. 6, Malacca (2 examples).

The under surface of this species reminds one of Cyclopides salsala of Moore.

556 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Genus PLESIONEURA, Felder.

1. PLESIONEURA FOLUS, (Papilio folus) Cramer, Pap. Exot. i. pl. 74. fig. F (1779).

3, Malacca; Penang.

2, PLESIONEURA ALYSOs, Moore, P. Z. 8. p. 789 (1865). 3 2, Malacca.

3. PLESIONEURA ASMARA (Moore in litt.). Similar to P. dan, but not tawny-tinted,

the three spots in the centre of primaries united, and hyaline white: expanse of wings

linch 8lines. ¢ 2%, Malacca (5 examples).

4, PLESIONEURA DAN, (Papilio dan) Fabricius, Mant. Ins. ii. p. 88. n. 798 (1787).

3, Malacca.

5. PLESIONEURA PINWILLI, n. sp. (Pl. LXVIIL. fig. 4.) Primaries black with a

bluish shot; a broad oblique shining pale-yellow subhyaline patch, separated by the median

nervure and its second and third branches into three spots; two small spots of the same

colour, placed obliquely below it, on the interno-median interspace : secondaries bright

orange, the base and the apical portion of external border chocolate-brown ; remainder

of outer border, a rounded spot at end of cell, a second near anal angle, and five, sub-

marginal, touching the outer border, black: head and thorax greenish grey, vertex of

head edged with sordid white ; abdomen orange banded with black. Primaries below as

above, excepting that there is a bifid whitish spot above the end of the cell, a whitish

spot at base of interno-median area, and that the inner margin is brown: secondaries

bright orange; the costal and outer borders irregularly purplish black ; fringe brown-

ish; a subcostal dash, a rounded spot at the end of the cell, and a reniform spot near

the anal angle black : body below and legs bright ochreous, palpi pale ochreous; neck

below white : antennz black above, testaceous below. Expanse of wings 2 inches 2 lines.

3, Malacca. Most nearly allied to P. tabrica of Hewitson from Darjeeling.

Genus TacrapEs, Hiibner.

1. TAGIADES RAVI, (Pterygospidea ravi) Moore, Proc. Zool. Soc. p. 779 (1865).

$2, Malacca. I have compared this with the type, with which it entirely agrees.

I have seen specimens of what is evidently a nearly allied species labelled as T. ravi; they

differ in having the hind borders of the secondaries above, and the abdominal half below,

white.

2. TAGIADES GANA, (Pterygospidea gana) Moore, P.Z.S8. p. 780 (1865). ¢ ?, Malacca.

Originally discovered at Bengal.

3. TAGIADES CALLIGANA, n.sp. (Pl. LXIX. fig. 11.) Closely allied to 7. atticus, but

the secondaries much narrower, and the two black spots towards anal angle wanting,

leaving only two submarginal black spots; submarginal spots below contiguous,

almost forming a border. Hxpanse of wings 1 inch 8 lines. ¢ ¢, Malacca

(2 examples).

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 557

A, TAGIADES LAVATA, n. sp. (Pl. LXIX. fig. 8.) Wings above dark smoky brown;

primaries with a sinuous series of five subapical hyaline white dots; secondaries with anal

half of external border snow-white, its inner edge sharply defined; body brown. Pri-

maries below paler; secondaries snow-white, with the costal area brown, its inner edge

diffused ; a small brown litura across the first median branch; body below whitish.

Hxpanse of wings 1 inch 6 lines. 6, Malacca. Allied to 7. pralaya.

HETEROCERA.

Family SPHINGIDA.

Subfamily Macroeiossin# (Grote), Butler.

Genus Macroeiossa, Ochsenheimer.

1. Macroeiossa PRoxIMA, Butler, P.Z.S.p. 4, pl. 1. fig. 1 (1875). ¢, Malacca. The

figure of this species in the ‘ Proceedings’ is not satisfactory, the yellow band being

left too broad, the diverging basal black streaks being also omitted.

Family ZYGAINID Ai.

Subfamily Zveaninz, Butler.

Genus Syntomis, Ochsenheimer.

1. SYNTOMIS BASIFERA, Walker, Journ. Linn. Soc. vi. p. 92. n. 37 (1862). %, Penang.

This species is intermediate in character between §. apicalis and S. detracta.

Family ARCTIID A.

Genus Spitosoma, Stephens.

1. Spriosoma,n. sp. A wholly white species. It is in very bad condition ; and there-

fore I prefer not to name it. ¢, Penang.

Family LITHOSIIDA.

Subfamily LirHostin a.

Genus ARrGiInA, Hubner.

1. ARGINA CRIBRARIA, (Phalena cribraria) Clerck, Icones, pl. 54. fig. 4. %, Penang.

Subfamily Hypsin.z, Butler.

~Genus Hypsa, Hubner.

1. Hypsa HELIconrIA, (Phalena-Noctua helicouia) Linneeus, Syst. Nat. i. 2, p. 839

767). 2, Malacca.

2. Hypsa susstminis, Walker, Lep. Het. Suppl. i. p. 212 (1864). 2%, Malacca;

3g, Penang.

SECOND SERIES.—ZOOLOGY, VOL. I. 4 is

558 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Subgenus Damatts, Hiibner.

3. Hypsa (DAMALIS) EGENS, Walker, Lep. Het. ii. p. 453. n. 12 (1854). ¢, Malacca.

Genus NrocuERA, Hiibner.

1. NEOCHERA MARMOREA, (Hypsa marmorea) Walker, Lep. Het. vii. p. 1674 (1856).

2, Penang.

Genus CaLLipuLA, Hiibner.

1. CALLIDULA ABISARA, n. sp. Nearly allied to C. sakwni from Java, but smaller,

duller in colour; the band of primaries narrower, more regular, and rounded at its

lower extremity ; wings below more densely reticulated with brown, and duller. Expanse

of wings 1 inch 2 lines. ¢, Malacca (1 example).

Apparently the sameas the Bornean form; but the type is notin good condition. Our

Bornean examples differ from C. sakuni in the colour of the band of primaries, which

is yellow with only the inner edge orange. This is probably the case with fresh

examples from Malacca.

Genus CiEostris, Horsfield.

1. CLEostRis caTamitA, (Tetragonus catamitus) Hitbner, Zutrige, figs. 653, 654 (1832).

3, Penang.

Family NYCTEMERIDA.

Genus Nyctemera, Hiibner *.

1. NYCTEMERA TRIPUNCTARIA, (Phalena tripunctaria) Linneus, Syst. Nat. i. 2, p. 864

n. 226 (1767). 3, Malacca.

2. NYCTEMERA CoLETA (Phalena coleta), Cramer, Pap. Exot. iv. p. 153, pl. 368. fig. H

(1782). o&, Malacca.

Genus Sucusio, Walker.

1. Secusio MuNDIPrIoTA, (Nyctemera mundipicta) Walker, Journ. Linn. Soc. iv. p. 184,

n. 7 (1860). ¢, %, Malacca; 2, Penang. The type was from Singapore. I have re-

ferred the above species, with NV. ¢rita (its nearest ally), WV. plagiata, N. annulata, and

NV. distincta, to the genus Secusio; they differ from the species of Nyctemera in the

Museum.

Family CHALCOSIID A.

Genus Mintonra, Walker.

1. Mrttonta Basatis, Walker, Lep. Het. Brit. Mus. ii. p. 365 (1854). ¢, Malacca

The bands, in the single example sent, are rather wider than in females from Java.

* Type NV. coleta, the first three species belong to Otroeda, the fifth to Amnemopsyche, the seventh to Secusio ;

the two remaining forms are congeneric..

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 559

Genus EvuscHEeMA, Hiibner.

1. EUSCHEMA SUBREPLETA, Walker, Lep. Het. ii. p. 406. n. 3 (1854). Hazis bello-

naria, Guénée, Uran. et Phal. ii. pl. 18. fig. 1 (1857). ¢ %, Malacca.

Genus AmegstA, Westwood.

1. AMESIA JUVENIS, n. sp. Most nearly allied to A. venusta: primaries rich purplish

chocolate; apical area, and a series of five marginal spots ultramarine blue; a series of

eight submarginal white dots; a sickle-shaped series of five discal white dots: secon-

daries red-brown, with two submarginal series of white spots, the inner or discal series

ill defined : body blue-black ; head and tegule spotted with blue. Wings below red-brown,

costal half of primaries purplish; base of costa metallic green ; two submarginal series of

white spots uniting into streaks at anal angle of secondaries, the inner series of prima-

ries deeply incurved, the three uppermost spots of the outer series edged internally

with blue; body below metallic green spotted with white. Expanse of wings 2 inches

9 lines. ¢&, Malacca. Hasily distinguished from its allies by its pale red-brown

secondaries.

2. AMESIA PEXIFASCIA, Butler, Journ. Linn. Soe. xiii. p.115. Primaries rich purplish

chocolate, external two fifths covered by a broad externally deeply digitate dentated

snow-white band, interrupted by the black nervures, the portion crossing the end of the

cell divided through the centre into two large spots; secondaries almost exactly as in

A. eupleoides of Herrich-Schaffer, but the anal angle dark greenish-grey; body deep

purplish chocolate, the three last abdominal segments blue-green ;, palpi, tegulee, and

thorax spotted with lilacine dots. Primaries below as above, but with a basal white

spot, two costal, two subcostal, two discoidal, and two interno-median blue-edged white

spots; secondaries below nearly as in A. ewpleoides, but the anal angle grey; body

below brown, with a lateral series of blue-edged white spots. Expanse of wings 3 inches

11 lines. ¢, Malacca. The most striking species in the genus.

Genus Latrion, Walker.

1. LAURION CORCULUM, n. sp. Sepia-brown; primaries crossed by an oblique narrow

sulphur-yellow band, beyond which the apical area is almost black. Wings below paler,

greyish towards the base; primaries with the costa as far as the band blue; secon-

daries with four apical marginal steel-blue spots; sides of pectus and anterior coxz

blue; venter banded with white. Expanse of wings 1inch1line. 4d, Malacca.

Genus CHaucosta, Hiibner.

1. CHALCOSIA COLIADOIDES (var. LATIFASCIATA). Chalcosia eoliadoides, Walker, Journ.

Linn. Soc. vi. p. 97 (1862). 2, Malacca. Differs from the Bornean form in the more

regular, non-lunated, broader, subapical, curved, cream-coloured band of primaries.

Genus Cyctosta, Hiibner.

1. CycLosIaA PANTHONA, (Phalena | Geometra| panthona) Cramer, Pap. Exot. iv. p. 68,

pl. 322. fig. C (1782). %, Malacca.

AE 2

560 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Family LIPARID A.

BIRNARA, 0. gen.

Allied to Pantana: wings rounded, semitransparent; costal nervure of primaries

terminating above the end of the cell; subcostal with four branches, the third and fourth

forming a fork to apex; radials emitted near together from the upper part of the cell;

discocellulars zigzag ; median nervure with four branches, the third and fourth emitted

near together : secondaries pyriform, costal nervure extending to apex ; subcostal emitting

two branches from a short footstalk; radial forming a fourth branch of the median;

discocellulars in an oblique slightly angulated line. Type B. nubdila, n. sp.

1. BrrnaRa NUBILA, n. sp. Primaries smoky grey, the veins white, base ochreous ;

secondaries greyish or sordid white; body ochreous, antenne brown. Wings below pale

grey, becoming smoky grey towards the outer margin; body ochreous. Expanse of wings

2 inches 1 line. 2, Malacca. Allied to Pantana bicolor from China, which, with Hloria

margimalis and Genusa discifera, may be referred to this genus.

Genus KETrELiA, n. gen.

Wings semihyaline, broad: primaries subtriangular; costal nervure extending to

beyond the end of the cell; subcostal emitting five branches, the first branch emitted

before the end of the cell, the second some distance beyond the end, immediately beyond

which the vein forks, the upper ray proceeding to near apex, and then forking again,

the lower ray simple; upper radial emitted from the upper part of the discocellular

line (which is zigzag); lower radial forming a fourth median branch: secondaries pyri-

form; costal nervure proceeding to apex; subcostal emitting two branches from the

superior extremity of the cell; radial emitted from the inferior extremity of the cell on

a level with the third median; upper discocellular acutely angular, its lower portion

twice as long as its upper, the lower extremity of the cell is consequently greatly in

advance of the upper: body slender; antenne rather short, broadly pectinated. Type

K. Lowi.

1. Kerrer1a Lowit, n. sp. Semitransparent white, the borders and veins opaque; a

pale brown straight line from the costa to the external angle of primaries, and from

the apex to the anal angle of secondaries; body white; palpi head, back of tegule,

base of primaries, sides of abdomen (more or less), and the greater portion of the legs

ochreous yellow; antennze pale brown. Expanse of wings2inches5 lines. 3, Malacca;

Sarawak (Low).

Our Malacca example being rubbed, I have taken as my type one of the three spe-

cimens kindly presented to the Collection by Mr. Low. The genus is allied to Penora.

Genus Oreyi1A, Ochsenheimer.

1. ORGYIA TURBATA, n.sp. Primaries ochreous, densely irrorated with ferruginous ;

the apex, a spot near external angle, and an ill-defined subbasal spot bright golden

yellow; subapical area red-brown; two transverse sinuated blackish lines, widely

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 561

diverging to costa; a black-edged reniform ferruginous spot at end of cell; a subbasal

transverse dusky line: secondaries reddish brown. Wings below sandy brownish;

primaries with a large dusky patch over the end of thecell. Expanse of wings 113 lines.

$, Malacca. I have seen a second example from Moulmein in Mr. Atkinson’s collection.

Genus Parasa, Moore.

1. ParasA BANDURA, Moore, Cat. Lep. E.I. C. ii. p. 417. pl. xia fig. 9 (1858-9).

3, Malacca.

Family SATURNIID A.

Genus Arracus, Hitbner.

1. Arracts atias, (Phalena-Bombyx Atiacus atlas) Linneeus, Syst. Nat. p. 808. n. 1.

3 ¢, Malacca.

Family HYBLAID A.

Genus Hyspuza, Fabricius.

1. HyBL#A CONSTELLATA, Guénée, Noct. ii. p. 391. n. 1261. ¢ ?, Malacca.

Family OPHIDERID A.

Genus OPHIDERES, Boisduval.

1. OPHIDERES FULLONIA,(<, fullonia) Clerck, Icones, tab. 42. figs. 3, 4 (1759-64) ;

Ophideres cajeta, Walker (nec Cramer), Lep. Het. xiii. p. 1223 (1857); ¢, Phalena-

Noctua pomona, Cramer, Pap. Exot. ii. p.122, pl. 77. fig. C(1779). 3 2, Malacca.

Clerck gives three figures on pl. 42, and names the central one O. fullonia; it may or

may not be the O. fullonia of Linneus. I have little doubt that the true O. cajeta

is identical with Walker’s O. bilineosa. According to the example in the Banksian

cabinet (which, however, is not the type), the O. dioscoree of Fabricius is a variety of

this species in which the silvery spot of primaries is developed into a streak be-

tween the median branches.

Family OMMATOPHORIDA.

Genus Nycrreao, Hiibner.

1. NycrIpao CREPUSCULARIS, (Phalena-Attacus crepuscularis) Linneeus, Mus. Lud.

Ul. p. 878; Clerck, Icones, pl. 53. figs. 3, 4 (mec 1, 2). Malacca.

Figures 1 and 2 of Clerck’s plate represent Guénée’s W. leucotenia. ‘* Comes into the

house at night; I have never known it to be taken in any other way” (W. L. Distant).

Family BENDID A.

Genus HuLopeEs, Guénée.

1. Hunopes caRANEA, (Phalena-Noctua caranea) Cramer, Pap. Exot. ii. p. 140,

pl. 269. figs. B, F (1782). Malacca.

562 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Family THERMESIID A.

Genus Coruza, Walker.

1. CoTUZA DREPANOIDES, Walker, Lep. Het. xv. p. 1552 (1858). o, Penang.

. Family URANIID A.

Genus NycTaALEMoN, Dalman.

1. NYcCTALEMON DOCILE, n.sp. Allied to WV. hector, from which it differs in its superior

size, deeper and redder coloration, narrower central white band, which is bordered

internally by a broad brown band, and externally by a narrow one; the costal border

not spotted with white; a broad oblique diffused discal brown band: secondaries with

the apical area much less streaked with black, the anal area much more streaked ; the

marginal interrupted black border narrower; the tails considerably longer and narrower

towards their extremities. Hxpanse of wings ¢ 5 inches 6 lines, ? 6 inches 4 lines.

3 ¢, Malacca (6 examples). This species is a well-marked local representative of

N. hector.

Family URAPTERIDA.

Genus Uraptpryx, Leach.

1. URAPTERYX MARGINIPENNIS, n. sp. Wings greyish brown, transversely streaked

with clay-colour, and reticulated throughout with transverse dark-brown hatchings:

primaries with the apex broadly, and the outer margin rather narrowly, nearly sulphur-

yellow; costa whitish; a yellow spot on the second median interspace: secondaries with

the outer margin rather narrowly sulphur-yellow; two decreasing submarginal black

dots above the angle of the wing: body greyish brown. Wings below grey, spot and

borders paler than above, reticulations obsolete; body below whitish. Expanse of wings

2 inches. 6, Malacca.

This curious species is not nearly allied to any species with which I am acquainted ;

it approaches most nearly to U. crocopterata; but the wings are very slightly angulated.

Family GHOMETRIDA, Guénée.

Genus AGATHIA, Guénée.

‘1. AGATHIA DISCRIMINATA, Walker, Lep. Het. xxii. p.591 (1861). 6, Penang.

Family ACIDALIID A.

Genus ACIDALIA.

1. ACIDALIA EXxcLUSA, ( Hemerophila? exclusa) Walker, Cat. Lep. Het. xxi. p. 320

(1860) ; Macaria obstataria, Walker, l.c. xxiii. p.928 (1861). do, Malacca. Walker

notes this species as Bremer’s Philobia cinerearia; but, as I have no Chinese examples

before me, I cannot certify the correctness of this determination. |

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 563

Genus ZANCLOPTERYX, Herrich-Schaffer.

1. ZANCLOPTERYX SAPONARIA, Guénée, Phal. ii. p. 16. n. 914 (1857). Malacca.

Family MICRONIID A.

Genus MicrontA, Guénée.

1. MicrontTA ASTHENTATA, Guénée, Phal. ii. p. 24. n. 925 (1857). Malacca.

Family ZERENIID A.

Genus Genusa, Walker.

1. GenusA BicuTTA, Walker, Cat. Lep. Het. iv. p. 818. n. 1(1855). 4, Malacca.

G. delineata and circumdata must be retained among the Liparide; they are not in any

way related to G. bigutta (which I consider the type of the genus).

Family MARGARODIDA.

Genus GLYPHODES, Guénée.

1. GLYPHODES AMETHYSTA, n. sp. Basal two thirds of wings hyaline white; external

third purplish pearly, bounded internally by a sinuated black line; primaries with a

tapering bronze band from costa to inner margin at basal third, a subquadrate spot of

the same colour at the end of the cell, costa brown-bordered, a bronze brown discal

diffused tapering streak; body probably brown, but too much rubbed for description.

Hxpanse of wings 1 inch 2 lines. 6, Malacca.

Family BOTIDIDA.

Genus Borys, Latreille.

1. Borys PrEROPHORALIS, Walker, Cat. Lep. Het. Suppl. iv. p. 1418 (1865). ¢, Ma-

lacca.

2. Botys, sp. —? Closely allied to B. contigualis. In bad condition. Malacca.

Therefore it appears that of the 280 species of determinable Lepidoptera collected by

Captain Pinwill, 41 are new or hitherto unnamed forms*.

* The following Table includes the whole of the species hitherto recorded from Malacca ; the species noted in this

paper from Penang only are necessarily omitted. The names of some of the genera formerly included under Huploa

haye been altered subsequent to the reading of this paper. Consult Journ. Linn. Soc. Zool. vol. xiv. p. 290 (1878),

“On the Butterflies in the Collection of the British Museum hitherto referred to the Genus Huplea of Fabricius.”

564 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Tabular View of the Butterflies of Malacca.

EF 3

wi | 3 3 3

Species from Malacca. 3 x ; S| a

# 5 E FI hb a S$ ie UO is ‘3 Remarks.

al2lElélelalalelalelalé

Euploea phoebus, Butler....... eee * * * | *

>, Ochsenheimeri, Lucas .. | ele bok | | ae. ee

50 Grotei, Felder .......... 26/50 los l\oel}o0 loo ]] sof} co lpoo |} ao ff ob |] on | Wpao taro Cocarin.

>» Margarita, Butler........ callooloa|l 3 oolloo | oollool| Ss

a «GUE, CGF caccnc000d Fee lPestcan Weesacia | seal h cases brewed og

» Meénétriési, Felder ...... eee Was, leer tod. Wee loo lee. [es

» Pinwilli, Butler.

mridernts! LANNE Pee aee Se IW Ges eel|s eee Sea lector repeal oe 9 enemy | ok

Calliploea diocletianus, Fabrictus ..| * |..|..|*% | * | %*|..| *]*

33 vestigiata, Butler ...... Reed lzesenl uate | tater incuced| aaaee|| Seen Se.

3 Ledereri, Felder ........

Salpinx leucogonis, Butler. *

Danais plexippus, Linn. ........ Elfen lfc [Rion litetee|ccte[tereg fo ( ea [ese

5, melanippus, Cramer ...... @ loo | ca |} &3 Joo lloo | ma ye I Ss

oy wallets, JW 5 20900000 * | *e].. fe. | & | ® *

», septentrionis, Butler ...... * |...) #1 *

» grammica, Boisduval...... oa | 43 |] ool} oo lool oo] oa] &

>, melaneus, Cramer........ reLibacdanel cs

crocea, Butler .......... Se cece ieezes Il Bet Ie *

Tdeopsis daos, Boisduval ........ * | * | x hs *

Hestia lynceus, Drury .......... * * | * | *

» Jlinteata, Butler.

Melanitis leda, Linn............- * | * | * | * * * | * | x

Lethe europa, Fabricius.......... * * * * *

Ceelites humilis, Butler.

Mycalesis mineus, Hewitson ...... ocd \Weserdl Reged lle eal neuen HOS

5 polydecta, Cramer...... eel iste: ere. seo arcade |

a janardana, Moore ...... Bred pecan liccea Ibevead ivcictel este (eevos | sss

on nautilus, Butler........ *

3 hesione, Cramer ......|..|+.|.-|*%]..|..]--|%

ie fusca, Felder .......... ited bvattecll vere. Meet eS aoa

Ypthima philomela, Mibner ...... Ae se oo | £3

of methora, Hewitson...... odhee lia ||. 3

corticaria, Butler.

Elymnias nigrescens, Butler ...... lela lliga las hea Iles

as Tutescens, Butler 000000 so ||oclloo| <3 <3 |] & |] &

mehida, Hewitson ...... weal gees lige aeoh| isaead a

Amathusia phidippus, Linn....... colfcallcol & loo | & loo | & | 3

Zeuxidia amethystus, Butler...... sca lhases) Ursseeelll essed hteceall BORE [RR

Discophora menetho, Hubricius.. ..| *

6 tullia, Cramer........ SE |e Aeon eestelll eae al ioe el | pesepll eek toda asks

Xanthotenia busiris, Westwood ..|..|..|..|..|*|*|*

Clerome gracilis, Butler.......... aie. IN ceo lpee aoa gterd |S)

» faunula, Westwood ...... syeclllageedllnes al Sa ao

Thaumantis noureddin, Westwood..|..|..|..|*|..|%*

sh pseudaliris, Buéler.

Doleschallia pratipa, Helder ...... BO ora |ieeo Il 3

Charaxes jalysus, Felder ........ Bee [deel Ree pon it. 3 *

55 hebe, Butler .......... a * | *

or Schreiberi, Godart...... oe * *

Prothoe caledonia, Hewitson...... Be kscot leceen || aren | eto lfc

Adolias adonia, Cramer.......... Br NeaSel hers |fearcah Incortad | beress cease fie

= gamle, WORe 2600500000 Ae (ea oS ee Tl ead eal aoe eee 3

», Jama, Melder............ ../| *

ee ee eee ee ee SS

MR. A. G. BUTLER ON THE

Species from Malacca.

Adolias layerna, Butler

ramada, Moore

asoka, Felder.

monina, Fabricius

ludekingii, Vollenhoven....

Sympheedra dirtea, Fabricius

i, ? emalea, Guérin.

Tanaécia aruna, Felder

a pulasara, Moore

Athyma larymna, Westwood

leucothoé, Linneus

kresna, Moore

> sabrata, Moore ....... 5

> wivifera, Butler ........

so eaaalngin?, 2 an oc cooc on

>» ¢lerica, Butler.

cy Mabie, WUE eeeene oadood

Peplavata,, Moore... 4. + ch.)

> urvasi, Felder.

Euripus eupleeoides, Felder.

EHulacura osteria, Westwood ......

Lebadea alankara, Horsfield ......

> Martha, Pabricwus........

Limenitis procris, Cramer........

Neptis mamaja, Butler ...... :

leuconata, Butler.

gononata, Butler.

nata, Moore .....

vikasi, Horsjield

columella, Cramer

dorelia, Butler

tiga, Moore ....

paraka, Butler.

dindinga, Butler

» hordonia, Stoll

Pandita sinope, Moore

Hypolimnas incommoda, Butler.

Cethosia hypsina, Felder

Cirrochroa orissa, Felder

johannes, Butler.

bajadeta, Moore

a rotundata, Butler.

Terinos teuthras, Hewitson

> robertsia, Butler

Messaras erymanthis, Drury

Atella phalanta, Drury

» sinha, Kollar

Cynthia erotella, Butler

Parthenos lilacinus, Butler ......

Cyrestis rahria, Moore

>» nivea, Zinken-Sommer

Ergolis ariadne, Linneus

>, merione, Cramer

Junonia iphita, Cramer

laomedia, Linneeus

lemonias, Linneus

orithya, Linneus

5, asterie, Jinneus

eudoxia, Guérin.

5S)

55 +tAOMCUlA, LANNEUS ......

| Assam & Nepal.

| Moulmein.

* * KF

* KK:

| Ceylon.

* KK:

SECOND SERIES.—ZOOLOGY, VOL. I.

* | Penang.

* KK KK KK KK KK K

* eK!

BUTTERFLIES OF MALACCA.

565

| Singapore.

| ‘Borneo.

KKK,

| Sumatra.

* *

| Jaya.

* * *K OK

* KK KK KOK KK

| iene

| Guna

* * *K *

| South Pacific.

| Australia.

Remarks.

Probably more than one spe-

cies.

Probably Felder’s C. nivalis.

566 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

F :

e g

i 1 ele g ales

Species from Malacca. 3B S)a| o/s} ¢ g a | moniaks:

elg/elel®/elalelalelelé

@D a) () =| ie) =| 3 |S | 9 5) 3

4/Sl/Ol/a la /MAlals/al/olal|<

Rhinopalpa fulva, Felder.

Zemeros emesoides, Felder ...... go Wiod-|| oo |I,oo Ilo) Es

», albipunctata, Butler...... .. | *¥

Albisara sayitri, Felder .......... *

Se kausambi, Felder........ a *|* | * | *

5 haquinus, Mabricius ....| * 23 || oo *

3) tamita, Hewson 3... . - * | *

» dumayjanti, Helder.

Poritia sumatre, Felder.......... * am || %

Gerydus Horsfieldi, Moore........ x |*|*|..|/*]..]..]..].. | Possibly more than one spe-

s nivalis, Druce .........- * * cies here.

3 symethus, Cramer ...... * *

Anops malayica, Felder.

» | Sperbthis, Welder .........- ao} oa|| ool]

Liphyra brassolis, Westwood.

Lycenopsis haraldus, Fabricius....)--|--]-+-]--|--|+-|*

Castalius rosimon, /% “Brien Soa eae x |x) *¥]xe]..] ee]. | Ee] &

45 ethion, Westwood ...... oo || £3] Selool £2 ico] cal] oo |] 3

sf elna, Hewitson ........ se ll art [ence] ten facie | tees) | rout a

3 roxus, Godart ........ sonllicn oo Woodlacd| call ool ts

Lampides patala, Kollar ........ jello TES foollosiias Wicolleeloolaolloo lao) thse strom IN, Ibagtisy,

p macrophthalma, Felder ..|..| * | * |..] *|*#]--]--|..]--]--]*

Bs peroé, elder .........- Soles lactic ll eolloolloc|laall gallon || oo fae || deyeeyiexman ina arith pyerines.

3 aluta, Druce ...,.....- eer) egoIL E.

565 elum@a, JOA sn 200006 da |loa|} oo *

Cranlen, IDFA sacoo50a\|c0 ||o0|| = | co]| 20 || &

oh agnata, Druce ........ oo lloolloollool| £3 | coil co} oa] 3

8 pseudelpis, Butler.

kankena, Felder........ solles eslicstico ll oolleci oot calico ao lloo || Jats chrom ng INigolnems.

Catapceci!ma elegans, Druce...... allan 23) e3lloo | <2 lee loo | sali oalloo || ao || AWE@ tite Somtinern Jimtie,

Lyceena cagaya, Felder .......... Balloo llco|loolloe|| <2 eo llesllae |colloo lea || damae wan tne Iai pmes.

», lysizone, Snellen ........ yo || oll eo || sa) oc ger Ge

GEMS, WOOO socodoosans se lloa|lood| £3 lloall oo) colds || delice oo |foo || Ayo tarorm Itemeill,

Amblypodia centaurus, Fubricius..|..| * | * | *|--|--| *|--| x

ae eumolphus, Cramer ..|..| *|--|..| ¥|--]--] *

3 anthelus, Westwood ..| ..| *

A adatha, Hewitson....|..|..|--|--| * | ¥

5 OSE, JADUSID d6o50\]50 lla | 00 |lc0 || oo || £3 |) 53

* amphimuta, Felder <.\\.. =. == |= || a || % | =

55 epimuta, Moore...... P \loollos|lroll eoilocllee|/oell as loo) co lloo | Atyqae som Ibnetin,

3 antimuta, Felder.

mj aroa, Hewitson ...... Pree (vee |ReOn ececaa eacallicoced leg

_ metamuta, Hewitson..|)..|..]--|-.-|--|--| *

an hypomuta, Hewitson ..|..|..|--|.-|.-] *

a3 apidanus, Cramer ....|..| ¥|--|--|.-| ¥]..] *

a IDreGhi, JEWS scoollcallosal|oo|)o0llaa]}o0 las |jo0 || &

a vihara, Felder.

inornata, Felder.

Hy polycena erylus, Godart ...... x) x] .. |e]... | &

etoluss LHAVRBUS 5550) |) £2 oo lao] do || $3 os] 8

Amphneus lohita, Horsf. ....:..... Caroll to Sale elles on letoneceliice [ae eulibe:

Deudorix petosiris, Hewson <2. .\\4.\| sa fe- |ae [es | aes | ina) oe te Also from Silhet.

a pheretima, Hewitson....)..|..|--]..| * | *

if jarbas, Mubricius ...... Pra | ere Wess orl cecal cts ec (stelle

aa doinitia, Hewitson...... ae MH) ape. W! Bee even RS Oe

Myrina megistia, Hewitson.

>, travana, Hewttson........ Oya Waren len ot] ecseenleo a) as, SmI

op Ga, ZOE on coanaco re | Pet call leo call eceead IS al owed ern ee

>» Marciana, Hewitson ..... 6 lNiceems| Need | ence nace cen ee genes

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 567

2 &

| é it)

. a |S aa Utyi Ieee || 1S, Sales

Species from Malacca. Bae) a ||P Es g/8 < FH "a Remarks.

slelele/P@lEleleisl/2/3ia

a Slol/Aalscoilsi/3]s8 lis |2)58

qi/al/ola/e2lalals |aijol3ass

Myrina lapithis, Moore .......... .. | ®]e- * | * *

ap Alitierch We Mdinae Benes 66 oc * . ..|% *

= enue, JUG ne canbuD O° *

>» anasuja, Felder.

Loxura atymnus, Cramer........ ..| |] *] * # * | * | *|..|--| Type from Coromandel.

Delias metarete, Butler.......... x Leelee) #1 &] &® *

PCO, DrUrg) ocr. «oto = acs | Sia] ec er Poy sea esr avert Pe

; ninus, Wallace.

Terias formosa, Hiuibner ........ 7 aes Hears | ese *

Pe SAITSPFLONS/(ELO, | ayes have =e 0 pip ficic || oe |) <3 * *

» hecabeoides, Ménétriés ....| * |--| * | *

PPINAnabaEw Eee... 4. =o Ee fea co Ihc ce, | ectese eto) ced] owe ol rel oal 3

Ba Puulaniswebutler 2242.2. Bon | Re | aia freer fet cM onl @ ll dew, fhdeee ear Hecke

senna, Felder.

Catopsilia erocale, Cramer ......| *| * | * | * *)*}*e |e] * *

= eatilla, Cramer........ x*|*|*/|* *|..| * * *

5 chryseis, Drury ...... *| * * 25 *

i scylla, Linneus ...... 3 * * *

Hebomoia glaucippe, Linneus ....| ¥ | * | *}-- é * * Possibly more than one

Appias plana, Builer............ oe [ee}eot--|--] ® species.

» cardena, Hewitson ....... J || Saalherel ly eve lretenl) 3

» leonora, Bowsd...........: ott Sem eer hess

» nathalia, Felder.......... .-]--|--]--| ®] ¥].-]..]-.]..]..]-- | Type from the Philippines.

~ jmmube Colin coccooue se Pea|| cio ool fee il clooeell bie Ihecs

Be ETICIOS -HULUIEGIUS) a1) < -tetterel™ PY bes teal lis ete I Sal fae PE 2d ed ae an Pe

figulina, UBULHery Siete totarar es fests eel ie onl es We Sele

Be lonce cynis, Hewitson ........ Pe ieescah feel (oe acetal aetna 3

Ornithoptera ruficollis, Butler ....|..|--|--| *

Papilio antiphates, Cramer ...... ce fee} ee] ee] ee | ® *

» Ssarpedon, Linn........... e | #]..) #]-. | *

» evemon, Boisd.....: ..../--]--]++| *]--| *

a) Gen, MWS. csoceoooees * | * || 3 *

» bathyeles, Zinken-Sommer .|..|--|--| * | *| * *

5, abycles, Bowsd. .......... * --| *¥| *| "=| *] *

» rama, Felder.

» agamemnon, Jinn. ...... *|*| *|* * *x | x

» malayanus, Wallace...... Seared ete || eatkcculine cates leer thse | ee \

» demolion, Cramer........ ..| ¥].-] *]-..] e]..] *| *{..]..]--| Am allied form came from

», Delessertii, Guérin ...... 5 lec | een Reveal fe S. India. }

3 polytes, Zinn. .........- x |.-| #1] *|.-| *]--| *| *]--|--|--]| & theseus type.

Pe belemusse Mem iaerte steers #].- |. * | * | ¥ *

3 prexaspes, Felder.

eis wara Voce meprte ere Peso | heb Me Sat epee 3

» saturnus, Guérin ........ ES | ere | ihe Ail echo cpodl ee 3 ie

A - Wispert Butler Vs.1).. oar we x

» mestor, Hiibner.......... SE ont! on] 3

pp 2eHabess Oi CmMe a 1-v-- lie aid (234 Weta ese Hols 4 Penal ted ll se etl 263

mm NERO Ae oo enone. eu ibare | calla

» diphilus, Hsper.....-.... *|--| *| * fa ||oo |! 2 || es|| £2

oO, CHIEU Goncacec 6 Peilleaoulhon *|*

erebus, Wallace ........ *

3 TDANR MOONE | ect cen 3 od ..|-.| ¥] *#] *

Hesperia harisa, Moore.......... Fae cers [lecae: || eon” || Carentan |, petal neces | ee Type from Sikkim.

x malayana, Felder ...... Sol (earn fee An allied species comes

35 yan: /35/td 17s eee eEBrS lole Beet lead Mee (ey asa | from Jaya.

Bare Mare enero |e lee en ices ee eee |e. eal ee seal ecsypertrom. Bengal:

es ciliatus, Butler.

4AEZ

568

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

: | :

a |g g a | 3

Species from Malacca. 2 =e so | $ £ | Remarks.

FlelSil al Slslsslalalele

#/S\e|s/e\ S12 | e\ 21/8) 2] 2

4/Sl/OlsA/e2\/Alalael/azl/o\;al<

Pamphila aria, Moore .......... 20 * ne Ne Type from Bengal.

“3 julianus, Latrelle...... 9 * |. * *

+ matthias, Fubricius ....| * * | * be * | & | x?

3 augias, Linneus ...... * * | * : Be ace al [ena *

r mesoides, Butler.

ws maro, Hubricius........ a * | * * |..|..|..]| Also from 8. India.

= nigrolimbata, Snellen ..|..|..|..|.-|--|--}-.| *

Plastingia callineura, Felder...... Soilleacoall odeetll | Reel Sea aa avog | ake

Cyclopides salsala, Moore ........ diced orca SeaHl saiecdl etedMlieeond leer: Type from Bengal.

Astictopterus jama, Helder ...... SSCA al as AO] coset ech all

= xanibes, Butler ....|..|:.)--|..|-- || *

3 gemmifer, Butler....)..|..|.-]..]--|--| *

3 armutus, Druce a All) ee *

5 diocles, Moore...... 4a * * Type from Bengal.

a sindu, Felder ...... ye *

ey stellifer, Butler.

Plesioneura folus, Cramer........ uae AM cry Ileana oie ltors * | x

a alysos, Moore........ -. | *¥ | #] |. | ® *

i asmara, Moore ...... yea ee * Also from the Khasia hills.

‘3 dan, Fubricius ....-. ue * Also from India.

ss Pinwilli, Butler.

Tagisdes ravi, Moore............ boll oe ll oa Nee ea lltoa || 3 * |..|..|.-. | Type from Bengal.

Heme, AWNOGRA ~sadsn00c00 BR eee meat Wee lleos |e a ils or |eael Male Type from Bengal.

a calligana, Butler.

3 lavata, Butler.

$5 trichoneura, Melder...... Doel cull aekee (ete chet all ieehod |e oe Also from Bengal & Silhet.

DHS HORNS oouaacaccc0g00 65 | 38 | 83 | 94 | 46 112) 41 | 87 | 39) 26) 2 | 6

. 1. Thaumantis pseudaliris (upper surface) 3. Fig. 1

2. Neptis peraka (both surfaces). 2

3. Neptis dorelia (both surfaces) . 3

4. Plesioneura Pinwilli (upper surface). 4,

5. Salpinw (Huplea) leucogoms (upper sur- 5

face) 9. 6

6. Neptis dindinga (both surfaces). 7

7. Papilio Esperi (upper surface) ? . 8

8. Lampides pseudelpis (upper surface) ¢. 9

2), v1 5 (both surfaces) 3. 10.

EXPLANATION OF PLATES.

Puate LXVIII.

11.

Puate LXIX.

. Neptis gononata (both surfaces).

. Neptis leuconata (both surfaces).

. Neptis mamaja (both surfaces).

. Athyma nivifera (both surfaces) g.

. Athyma clerica (both surfaces) ¢.

. Hestia linteata (upper surface).

. Astictopterus xanites (upper surface).

. Tagiades lavata (upper surface).

. Euplea Pinwillt (upper surface) 3.

Zemeros albipunctata (upper surface) 3.

Tagiades calligana (upper surface).

A.S Butler Trans. Linn. Soc. SER 2. Zoo. Vor 1.Tas.68.

Neptis peraka. Thaumantis pseudaliris.

Lampides pseudelpis . ?. Lampides pseudelpis. 3.

Papilio esperi.

ik Ba

eres owe iene 7 ae Meets j a

4 . yeh iy read NF,

nm b ‘ pe ; ; a em “

et a alate eine RM ou oh

‘i : Vs

aa ; ‘ Sibtict :

- 5

yan

nS) :

. ¥;

' é .

eee; A

‘ .

e Sa

ont hy

o) * -

* $

. ‘

‘

7 <

. Ley

¢ .

Z :

A.G. Butler

Neptis leuconata.

Euplea poowilli.d.

BUTTERFLIES OF MALACCA.

Trans. Linn. Soc.SER.2.Zoo1 Vou 1.TAB-69.

[ 569 ]

XXII. On certain Organs of the Cidaride. By Cuarius Stewart, Esq., F.L.S.,

M.R.C.S., Lecturer on Comp. Anat. St. Thomas’s Hospital.

(Plate LXX.)

Read December 6th, 1877.

SOME years since, whilst examining a spirit-preserved specimen of Dorocidaris papil-

lata, I first noticed the organs which it is the object of this paper to describe. An

undetected leak in the cell in which they were preserved unfortunately allowed them to

dry before a drawing had been made. ‘To the kindness of the Rev. A. M. Norman I am

indebted for two specimens, which have enabied me to confirm my observation, and

make the necessary drawings.

Of the regular Echinoidea the families Diadematide, Echinometrade, and Hchinide

have long been known to possess external branchie in the form of five pairs of conical

hollow processes of the outer margin of the buccal membrane. The branchie project

outwards into the surrounding water, their interior being bathed by the fluid of the body-

cavity, of which chamber they are diverticula. Their under surface is amplified by

secondary diverticula, the whole organ having its walls strengthened by small calcareous

plates, and sometimes spicules. Large cilia clothing its exterior, keep a constant current

of water over it. The bases of the branchiz are attached to the edges of the ten notches

which are usually present at the buccal margin of the corona. As might be expected,

these notches are most marked where the branchize are largest, e. g. Toxopneustes,

Spherechinus, &c.

The existence of such branchize in the Cidaride was denied by Miller; but in the

magnificent ‘ Revision of the Hchini’ of Prof. A. Agassiz, p. 694, he says:—‘‘In the

Cidaridz these gills are found in cuts of the buccal membrane itself, close to the acti-

nostome (Miller denies their existence ; yet he has himself figured the cuts through

which the gills pass in a species of Cidaris).”

I have carefully removed the spines one by one from the buccal membrane of Doroci-

daris papillata preserved in spirit, and then examined it under the microscope without

finding a trace of cuts or branchize (Pl. LXX. fig. 1); but in dry specimens the actinostome

appears notched from the shrinking of the membrane, which here occupies a gap left

between the interambulacral plates; this membrane shows a few creases, but nothing of

a branchial character.

If the abactinal or dorsal half of the corona be now removed, five remarkable organs

will be seen (fig. 2), which I believe must functionally take the place of the absent

external branchiz. They are situated immediately beneath the outer forked extremities

of the five compasses, and are diverticula of the chamber that occupies the intervals

between the jaws and their muscles, and which is bounded by the reflection of the inner

lining (peritoneum) of the test over them.

570 MR. C. STEWART ON CERTAIN ORGANS

In a full-grown specimen of Dorocidaris these branchiz are about half an inch in

length, are gracefully curled upwards, and have on their under surface a double row of

compound secondary diverticula, which, largest near the base of the branchiz, are

gradually reduced in size towards their tips. The walls of these organs are exceedingly

delicate, and contain similar spicular plates to those found in the neighbouring peri-

toneum. On either side of the base of the branchize external to the oblique ligaments of

the compasses there is a small group of bulla-like elevations of the peritoneum similar to

those found on the sides of the gills.

The size of the jaw-chamber can clearly be increased by the raising of the compasses

when the transverse muscles contract, especially if the jaws be protruded. Such move-

ments of the parts are frequently going on; so that if the chamber communicate with the

surrounding water, as it probably does near the tips of the teeth, water would then pass

in and out of the space, and bathe the interior of the gills. (I have not yet had an

opportunity of examining Asthenosoma, but should think it most probable that similar

gills are present there to those found in the Cidaride.)

If this hypothesis of one of the functions of the compasses, their muscles, and ligaments

be correct, some light is thrown on what has long been a puzzle to me, viz. of what use

they are in the economy of the regular Echini. The compasses have been said to be

used as levers by which the transverse muscles separate the points of the teeth ; and the

oblique ligaments have been thought by some to be muscles approximating the points of

the teeth. The ligamentous nature of the latter, however, is quite evident both under

the microscope and to the naked eye; and as such they are described by Valentin and

Prof. Agassiz.

There are two reasons why I consider that the compasses have nothing to do with

movements of the teeth. First, there are, exclusive of them, powerful muscles by which

all necessary movements of the jaws can be accomplished, viz. five pairs of retractors

passing obliquely downwards and inwards from the auricles to the tips of the jaws (pyra-

mids), which can also separate the teeth, five pairs of protrusors arising between the

auricles and inserted at the bases of the jaws, the whole of the contiguous faces of the

jaws giving attachment to a muscle which, passing from one to the other, would approx-

imate the teeth. Secondly, the mechanical arrangement of the compasses is unfavour-

able to their acting as teeth-movers. It seems, then, possible that the function of the

compasses is to raise the peritoneum from the base of the jaws; at the same time this

would stretch the oblique ligaments, and raise this membrane from their sides. Another

function might also be to remove injurious strain from the madreporic canal when the

jaws are protruded.

Although the pedicellarize of the Cidaridze have been described by many observers,

I have not seen the following peculiarity of the armed or chambered pedicellarize men-

tioned. In this form, when the jaw ends in a terminal fang, the chamber has a second

opening in addition to the large triangular tooth-armed orifice near the tip, the fang

itself being hollow, and perforated external to and in front of the tip, a place which

corresponds exactly with the opening of the groove in the terminal fang of the gemmi-

OF THE CIDARIDA. 571

form pedicellarize of the Echinidz, and which in them probably serves as a channel for

the injection of a poison secreted by two sac-like glands appended to each jaw-like piece

of the pedicellarian head. It would appear from this that the armed pedicellarize of the

Cidaridee are the analogues of the gemmiform ones of the Echinide. The position of the

opening in the fang is admirably adapted for the injecting of poison into a wound;

the flesh, which is compressed on its inner or convex side, has a cavity torn in it in

front, into which the poison could readily pass, from the opening of the fang being

situated in its anterior border and not at its point. A similar contrivance for a like

purpose is found in the falces of spiders, poison-fangs of most venomous snakes, &c.

Mixed with the ordinary forms of pedicellariz on or near the apical system of

D. papillata, 1 have often found the large armed variety possessing four jaws; this is,

I believe, the second instance of such an exception to the rule of there being three only,

avery remarkable form having been previously described by Sir Wyville Thomson as

existing in Asthenosoma. It differs, however, in all other respects from those of the

latter genus, only varying in the number of its jaws from the ordinary form found in

Doricidaris.

The next point to which I wish to call attention is that in Cidaris tribuloides and

Phyllacanthus baculosa, in addition to the minute calcareous arches which bridge over

the first three or four of the inner ambulacral pores, counting from the inferior opening

of the corona, and which have already been described by J. Miiller under the name

of vertebral processes, there are between all the inner pores delicate solid spines, some-

times two in number, between each pair of pores; they project downwards, and tend to

arch over the ambulacral vessel. The vertebral processes, of which these spines are

but a modification, have already been shown, in the paper by J. Miller before

referred to, to be the homologues of the auricule of the Diadematide, Hchino-

metradz, and Echinide, and of the vertebral ossicles of the Asteroidea, which

in like manner arch over the ambulacral vessel and nerve. The continuation of the

representatives of these arches throughout the whole length of the ambulacra in some

of the Cidaridze becomes therefore interesting. These spines remind one of similar pro-

cesses scattered over the inner surface of the corona in the Clypeastride.

The genital gland of D. papillata shows a spicular framework consisting of calcareous

triradiate spicula, some of which are of large size, and have their angles more or less

filled by an irregular network ; occasionally also one of the radii is prolonged, and projects

as a spine from the surface of the gland. Sir Wyville Thomson, in his ‘ Voyage of the

Challenger,’ vol. ii. pp. 227, 228, describes an interesting new species of Cidaris under

the name of C. nutriz. He points out many features of resemblance presented by it

to D. papillata, and others in which it differs from that species, amongst which is men-

tioned the paucity of spicula in the walls of its genital glands, whereas D. papillata is

said to have them in great abundance. Having examined a considerable number of

the latter species, I find great variation in the number of the spicula, although their

general form is constant in similar specimens, those having comparatively small genital

tubules having the spicula most abundant, whereas where the tubules are of large size

(females ?) the spicula are often very scanty and small.

572 MR. C. STEWART ON THE CIDARIDZ.

DESCRIPTION OF PLATE LXX.

Fig. 1. Portion of buccal membrane of Dorocidaris papillata. The tips of the five teeth are seen in the

centre. The spines have been removed from one side to show the ambulacra and the absence of

actinal cuts and external branchie. x 14 diameters.

Fig. 2. Internal branchia of D. papillata. x 8 diam.

Fig. 8. D. papillata, having upper half removed to show jaws and internal branchiz : slightly enlarged.

Figs. 4 & 5. Internal and side views of two jaws of an armed pedicellaria, showing perforation of terminal

fang. x 80 diam.

Fig. 6. Portion of ambulacrum of Cidaris tribuloides from near buccal membrane, showing vertebral

processes. x 12 diam.

Fig. 7. Side view of portion of an ambulacrum from near buccal membrane. Phyllacanthus baculosa.

x 12 diam.

Fig. 8. Portion of genital gland of D. papillata. x 60 diam.

Soc. See. 2. Zoor Vou], Pr

WN. DO

Trans. L

CStewart del.

ALATA.

LLACANTHUS BACULORA.

PAPIL

MBE

ARTIS

wv Lt)

ORO

6.CIDARIS TRIBULOIDES

Hanhart oop,

TPN

CBerjeau th

[ 573 J

XXIII. On Hypsiprymnodon, Ramsay, a Genus indicative of a distinct Family (Pleopo-

didee) in the Diprotodont Section of the Marsupialia. By Prof. Ownn, C.B.,

FRS., FLS., &c., Superintendent of the Natural-History Departments in the

British Museum.

(Plates LXXI. & LXXII.)

Read January 17th, 1878.

‘THE modification of the hind foot in the Marsupial series is peculiar in the way in:

which the digits depart from the typical mammalian number, 5. The peculiarity is.

chiefly shown by an atrophy of the second and third digits of a kind like that. which

affects the third digit of the fore foot in the Aye-aye—namely, attenuation without loss

of length, and this in relation to a special use distinct and apart from. the foot as a loco-.

motive or prehensile organ.

Not only are the slender metapodials of the digits 11 and 111 (Pl. LX XII. fig. 10) wrapped.

up with the others in a common sheath of skin, but also the proximal and middle

phalanges, leaving only the distal ones with their claws free, but in close contact and of

equal length; and these claws have been uniformly observed in the Marsupials with

the so-modified hind foot to be used as scratchers and cleansers of the fur.

This condition of hind foot is found in carnivorous, frugivorous, and herbivorous Mar-

supials, and is associated with as many distinct formule of the dental system.

The modification of digits 11 and 111, moreover, may coexist with the normal develop-

ment of the digits Iv and vy, and with an abnormal one of digit 1, in the form of a

powerful opposing hallux, as exemplified in the grasping hind foot of the scansorial

phalangers (woodcut, fig. 1); or it may be associated with an abnormal development of

the digits Iv and v in both breadth and length, especially of Iv, with a reduction of

digit 1 to rudimental insignificance, as in the gradatorial Bandicoots (Perameles); or it

may be associated with a still greater development of digits rv and y, and especially of

Ty, with total loss of digit 1, as in the hind foot of the saltatorial Potoroos and Kan-

garoos (woodcut, fig. 2).

But in the subject of the present paper a completely developed hallux (woodcut, fig. 3;

Pl. LXXII. fig. 7,1) is combined with the fur-cleansing and locomotive toes of the

Poéphaga* ; and this singular osculant condition of foot is associated with the dentition of

the Hypsiprymnide. Thus the transitional character of the mammal under consideration

would seem to place the genus between the scansorial and saltatorial groups of the great

diprotodont division + of the Marsupial order. The analogous approach in digits mand

tt to the Kangaroo type of foot made by the Bandicoots (Peramelide) may be a mere

adaptive one in relation to their fur, and at any rate is an affair of one of the Polypro-

* «Outlines of a Classification of the Marsupialia,’ Trans. Zool. Soc. vol. ii. 1839, p. 332.

+ ‘Researches on the Fossil Remains of the Extinct Mammals of Australia,’ 4to, 1876, vol. 1. pp. xii, 107.

SECOND SERIES.—ZOOLOGY, VOL. I. 4G

574 FROF. OWEN ON HYPSIPRYMNODON.

todont suborder *, and must therefore be set aside in the consideration of the affinities

of Hypsiprymnodon, a genus which I regard as the type of a distinct group of the

Diprotodontia, for which group, from the typical completeness numerically of the toes,

the name Pleopodidee + is proposed.

Fig. 1. Fig. 3. Fig. 2.

Fig. 1. Sole, hind foot, of Phalangista.

Fig. 2. Sole, hind foot, of Kangaroo.

Fig. 3. Sole, hind foot, of Hypsiprymnodon.

This group is at present represented by a single species, Hypsiprymnodon moschatus,

Ramsay, the description of which, unaccompanied by illustration, excited much interest,

and a strong desire with the students of the Marsupial order to make a further acquaint-

ance with the new form {.

I have recently been gratified by the reception of skins of mature male and female

specimens kindly transmitted to me by the describer of the species, the present accom-

plished Curator of the Australian Museum, Sydney. The skull and bones of both fore

and hind feet were included in the dried skins; and these, being extracted, have afforded

the materials for the following notes, which, with the accompanying illustrations (Plates

LXXI. & LXXIT.), may be acceptable to the Society, although they add little to the

accurate description in the ‘ Proceedings of the Linnean Society’ of the capital of one of

our most remote and prosperous colonies.

* ¢ Researches on the Fossil Remains of the Extinct Mammals of Australia, 4to, 1876, vol. i. pp. xii, 105.

+ mdéos, full; cows, foot.

+ “ Description of a new Genus and Species of Rat-Kangaroo, allied to the Genus Hypsiprymnus, now proposed to

be called Hypsiprymnodon moschatus. By E. Pierson Ramsay, F.L.S., C.M.Z.S.,” Proceedings of the Linnean Society

of New South Wales, vol. i. pp. 33-35, 1876.

PROF. OWEN ON HYPSIPRYMNODON. 575

From the presence, direction, and relative size of the hallux, the Hypsiprymnodon

might be inferred to be a grasper, perhaps an occasional climber, though not so strictly

arboreal as the Dendrolagues and Phalangers. Its nearest affinities, as shown by the

skull and dentition, are to the Hypsiprymnide.

Both male and female of Hypsiprymnodon, as exemplified by the transmitted skins, are

of equal size; each measures 1 foot 833 inches from snout to end of tail, this being 5

inches 9 lines in the female, 5 inches 4 lines in the male, of which the tip of the tail

appears to be entire. The skins indicate a rather long and slender-bodied quadruped,

with a circumference at the haunches of about 8 inches, the trunk not tapering forward,

as in the Potoroos, in which family also the smaller size of the female is notable, though

the sexual difference of size is less in some of the Potoroos than in the Kangaroos.

If to a skull of 2 inches 7 lines (65 millims.) in length 10 millims. be added for a

snout shrunk in the dried skin, and 2 millims. for occipital integument, we get a length

of head of 8 inches. The limbs are less unequal in length than in the Potoroos placed

by Mr. Waterhouse in his subgenus Potorous, the species of which have “the hinder

legs shorter and the head more slender and pointed”’ than in the Hypsiprymni proper.

In the relative length of the head to the body, and especially to that of the hind limb,

Hypsiprymnodon recedes from the Poephagous group, especially as exemplified in the

Kangaroos and in such Potoroos as Hypsiprymnus campestris, Gd., and Hypsiprymnus

platyops, Gd. In this proportion Hypsiprymnus apicalis, Gd., comes nearest to Hypsi-

prymnodon.

The pelage of Hypsiprymnodon moschatus resembles that of the Potoroos, and differs

from that of Bandicoots, in which the hair is of two kinds, the outwardly visible

and longest being coarse and harsh to the touch, that beneath forming a soft, somewhat

scanty fur. In Hypsiprymnodon the fur is of moderate length, pretty closely applied,

having numerous rather longer interspersed fine hairs, the visible portion of which is

black or blackish, and pointed; these are relieved by the dark- and light-barred colour

of the visible part of the shorter hairs, all the hairs at their basal portions showing a

leaden greyish tint. The tips of many of the shorter hairs are of a brighter brown than

the rest, approaching to yellow. Mr. Ramsey describes the upper surface of the body of

the living or recent animal as “clothed with a close and rather stiff fur of rich golden

colour mixed with black; the head, face, and lower part of the legs dark-brownish grey ;

along the centre of the throat and chest a few patches of white” *. These, in the skin

of the male transmitted, are of a light yellowish tint; and the abdomen is a light brown.

“The young are of a more golden hue, and less white on the under parts” +. The

pelage is continued by shorter hairs from the trunk over the base of the tail for half

an inch or more, where it abruptly ceases. The rest of the tail is nude and covered

by reticularly disposed scales about 14 or 1 millim. in size to the gradually attenuated

extremity. These scales are almost black on the transversely convex upperside, and of

lighter leaden colour on the less convex underside. The diameter of the tail at the

beginning of the scaly part is 8 millims. A few minute very short hairs project from

* Tom. cit. p. 34. . t Ibid.

462

576 PROF. OWEN ON HYPSIPRYMNODON.

the interstices of the scales, almost requiring a magnifier for detection. The caudal cuticle

is in the form of lacertian scales in many Potoroos; but in these, as in Lagorchestes, the

scales are partially hidden by short decumbent hairs, varying in position in different

species ; the scaly tail is most sparingly clothed, and with short stiff hairs, in the sub-

genus Potorous, Wth. On the head there are a few scattered supralabial, malar, and

superorbital vibrisse, black, fine in texture, about an inch in length.

The muffle is nude, as in Bettongia and Potorous, the black naked skin being con-

tinued a short way back from the upper part of the nose. The nostrils are lateral,

opening close to the terminal nasal disk; they are curved, with the convexity forward ;

the disk is vertically grooved, and projects beyond the mouth. The lateral extent of the

oral cleft is 1 inch (26 millims.). From the nose-tip to the eye is 1 inch 5 lines (86

millims.). The palpebral cleft is 11 millims. From this to the meatus auditorius is

22 millims. ‘The irides are dark hazel”*. The ears are rounded; 20 millims. long by

22 millims. broad; the upper margin is slightly deflected. The basal half of the outer

surface is clothed with short fine yellowish hairs; the rest of the auricle is naked.

The limbs, fore and hind, are less unequal in length than in the Hypsiprymnide; the

forearm and paw measure 3 inches-2 lines (80 millims.), the leg and hind foot 4 inches

9 lines (122 millims.). The trunk does not taper forward, as in the Kangaroos, but

maintains a more equable thickness in agreement with the more equal length and strength

of the fore and hind limbs.

The first and fifth digits of the fore feet, (Pl. lxxii. fig. 5) though shorter.and smaller than

the rest, are by no means so reduced as in Perameles; they retain the general type of those

in Hypsiprymnus. The pollex, 1, is the shortest; its claw does not extend beyond the base

of the fore finger 11. The little finger, v, attains the distal end of the proximal phalanx

of the finger, 1v. The mid digit, 111, exceeds the adjoining ones by nearly the length of

its claw. The claws are small, subcompressed, subdecurved, not broadened above, as in

Macropus. The short hairs of the hand terminate at the base of the digits, the dorsal

tegument of which is smooth and scaly. On the palmar side of the hand the hairs cease

at the proximal border of the palm. This presents a pair of unequal longitudinally

oblong callosities, and, in advance of these, three smaller ones near the base of the digits

II, Iv, and v respectively. The longer hairs of the fore limb shorten, but less abruptly

than in the hind limb, at about a third of an inch from the wrist. The callous pad

beneath the hind foot extends from the base of the toes backward to the end of the

heel, reducing the coating of short and adpressed dark brown hairs to a part of the

upper surface.

The chief gradatorial toe, woodcut, fig. 3, Tv, is not so disproportionately long and strong,

nor is the fifth toe so weak, as in the Kangaroos and the more Kangaroo-like Potoroos.

In the Hypsiprymnide the proportions are most nearly approached by the species Potorous

murinus. The claw of digit tv has not the three-sided hoof-like character of that of the

Kangaroo; it is subcompressed, shorter and deeper than the cleansing-claws of 11 and 111.

The longer hairs of the leg terminate abruptly half an inch above the tarsus, like

* Tom. cit. p. 34.

PROF. OWEN ON HYPSIPRYMNODON. 577

the legs of a pair of trousers, beyond which the hairs become abruptly very short. They

cease behind, leaving the heel bare, but are continued over the front and sides of the

tarsus, along the upper surface of the metatarsus, and along that of the phalanges of

toe Iv as far as the claw. The dorsum of the second phalanx of the hallux and that

of the same part of toe v, are covered by scaly integument, like birds’ toes. The

same integument covers the dorsum of the common sheath of the cleansing-toes

11 and 116.

The skull of Hypsiprymnodon (Plate LX XII. figs. 1, 2, 3) is smaller, narrower, more

elongated and attenuated anteriorly than in any known Potoroo—consequently departs

further in general form from that of the Kangaroos. The alisphenoidal bulla is rela-

tively smaller than in the Potoroos. The frontals are relatively narrower in proportion

to their length than in most Potoroos, showing rather the form of those bones in some

of the smaller Kangaroos and in the Dendrolagues. But the ridges continued back-

ward from the postorbital processes are very faintly indicated in both sexes of Hypsi-

prymnodon; in Dendrolagus they bound the temporal fossee above by a well-marked

elevation: Hypsiprymnodon resembles Hypsiprymnus in this difference. The nasal

bones are larger and narrower than in any Potoroo. Their pointed ends project 4 millims.

in front of the naso-premaxillary suture. A line dropped from their tip would clear the

front border of the foremost incisor. The length of the nasals is 1 inch 3 lines (31

millims.); their basal breadth 4 lines (9 millims.). The fronto-nasal suture is on a

transverse parallel with the fore part of the orbits, not in advance thereof as in Potoroos;

its position is that which obtains in Dendrolagus; but it is transverse, not angular. The

lacrymal advances a little way upon the face; it is grooved externally, and conveys the

duct by a canal commencing in advance of the orbit, as in other Marsupials and in Birds.

The maxillary is pierced by two antorbital foramina, one small, 6 millims. in advance

of the orbit; the other, which is larger, and answers to the normal foramen in Kan-

garoos and Potoroos, is 8 millims. in advance. In Potoroos* and Kangaroos + the upper

premolar descends more or less in advance of the antorbital foramen ; in Hypsiprymnodon

the foramen is in advance of the premolar. The Potoroos differ from most Kangaroos in

the longer and narrower facial part of the skull anterior to the zygomata, which part

also tapers more anteriorly. Hypsiprymnodon carries further this difference, the cor-

responding part of the skull being relatively longer and more pointed than in Hypsi-

prymnus cuniculus e.g. The extent of the maxillary in advance of the premolar

exceeds that of the premaxillary; in the Potoroos cited below reverse proportions

prevail. The course of the maxillo-premaxillary suture resembles that in the Potoroos:

the crown of the small canine projects from the lower end of the facial part of that

suture, as in Potoroos. The palatal plate of the maxillary has a pair of minute fora-

mina on the transverse line a little in advance of the premolar. The chief vacuities of

the bony palate are the two hinder ones, each half an inch in length, together 4 lines

(13 millims.) in breadth; the pair of premaxillary fissures are relatively longer and

* Waterhouse, ‘Mammalia ;’ Hypsiprymnus penicillata, plate vi. fig. 3a; Hypsiprymnus minor, plate viii. fig. 3.

+ Owen, ‘Foss. Mamm. of Australia,’ plate Ixvi. fig. 1 (Osphranter rufus) ; plate Ixxx. fig. 1 (Phascolagus erubes-

eens); plate Ixxxiy. fig. 1 (Halmaturus ualabatus).

578 PROF. OWEN ON HYPSIPRYMNODON.

narrower than in Potoroos. The palatal vacuities are bound behind by slender trans-

verse bars of the palatines, and extend forward to the interspace betweeen the premolar

and first molar teeth. Mr. Ramsay rightly notes of the posterior palatal openings

the character of the anterior margins meeting the exterior lateral at an acute angle*.

The breadth of the skull outside the zygomata is 1 inch 3 lines (82 millims.); the

vertical diameter of the zygoma at the back part of the orbit is 5 millims. Jt does not

increase posterior to this part, as in Kangaroos, but rather loses in that dimension as it

extends to the tympanic bone.

3—3 1—1

The dental formula of Hypsiprymnodon is that of the Potoroos, viz. 7. = ne

Dp. a m. i = 30. The molar series in each jaw, according to characters of develop-

ment, include the teeth symbolized as p. 4, d.3, m.1, m.2, m.3+. In some Potoroos

(Hypsiprymnus Gilberti e. g.) the crown of the premolar deviates from the longitudinal line

of direction of the four following teeth by a slight turn outwards, and rather more so

in the mandible t. A similar outward bend of the premolar is more marked in Hypsi-

prymnodon §, in which the crown of the tooth, p. 3, presents the proportions shown in

H. Gilberti. In most Potoroos the fore-and-aft extent of the crown of the grooved and

trenchant premolar is relatively greater. The molars in Hypsiprymnodon repeat the

proportions most common in the Potoroos, the last (m. 3) being the smallest, whilst in

H. Gilberti the first (d. 4) is the smallest.

The length ofthe forearm and hand from tip of elbow to the nail, inclusive, is 3 inches

2 lines (80 millims.), of leg and foot, 4 inches 9 lines (122 millims.), of fore foot, in-

cluding carpus, 1 inch 8 lines (80 millims.), of hind foot, including tarsus, 2 inches 5 lines

(61 millims.), of face from tip of nose to base of ear 2 inches 5 lines (61 millims.).

The ulna and radius are in contact along their distal halves (Pl. LX XII. fig. 5). The

radius is more slender and of more uniform breadth than in Hypsiprymnus Gilberti or

Bettongia penicillata (Pl. LX XII. fig. 9). In these and other Potoroos the bone is pro-

duced ulnad at its distal lower two thirds asa sharp plate. The length of the radius is

1inch 9 lines (45 millims.) in Hypsiprymnodon moschatus, but is only 1 inch 6 lines

(40 millims.) in Bettongia penicillata, with twice the breadth, viz. 5 millims. ‘These

differences are notable for the exigencies of paleeontology in reference to detached bones

in caves, breccias, and drifts. The ulna of Hypsiprymnodon is 2 inches 2 lines

(55 millims.). The metacarpal and phalanges are longer and more slender in Hypsi-

prymnodon than in Hypsiprymnus, save the ungual phalanges of 11, 111, and iv, which are

both longer and stronger than in the Potoroos.

The tibia and fibula, after their proximal coarticulation, keep -apart for more than

half their length distad; in the rest of their course they are in close contact but not

ankylosed. In Bettongia penicillata the interosseous space occupies but one third of the

proximal portions of the shafts of the two leg-bones; in the rest of their extent the tibia

and fibula are ankylosed, with a slight linear indication of their primitive distinctness.

* Tom. cit. p. d4. t ‘ Odontography,’ 1844; Proceedings of the Zoological Society, 1877, p. 358. .

t Plate LXXII. fig. 8. § Ih. figs. 3 & 4.

PROF. OWEN ON HYPSIPRYMNODON. 579

The length of the tibia in Bettongia penicillata is 34 inches; in Hypsiprymnodon it

is 2 inches 8 lines.

The length of the foot-bones in Hypsiprymnus Gilberti (Plate LX XII. fig. 10) is

4inches 6 lines; in Bettongia penicillata it is 3 inches 10 lines; in Hypsiprymnodon

(ib. figs. 6 & 7) it is 2 inches 5 lines.

The entocuneiform bone (ib. figs. 6 & 7, e) is proportionally larger, especially broader,

than in the Potoroos in relation to the articulation of the metatarsal of the hallux, 1.

The rest of the structure of the tarsus adheres, save in the shorter proportion of the bones,

to the Poephagous type, exemplified in plate Ixxii. of my ‘Fossil Mammals of

Australia.’

Mr. Ramsay states that the habitat of Hypsiprymuodon moschatus is the Rockingham-

Bay district, and that it frequents the dense and damp positions of the scrubs which

fringe the rivers and clothe the sides of the coast-range. Here it is by no means rare ;

but, from its retiring habits and the density of the forest, is difficult to obtain. The

habits are chiefly diurnal, and its actions, when not disturbed, by no means ungraceful.

It procures its food by turning over the débris in the scrubs in search of insects, worms,

and tuberous roots, frequently eating the palm-berries (Piychosperma Alexandre), which

it holds in its fore paws, after the manner of the Phalangers, sitting up on its haunches,

or sometimes digging. Seldom more than one or two are found together, unless accom-

panied by the young. “In March 1874 I obtained from Mr. Broadbent a female with

two young in the pouch, very small. During the same month a half-grown young one

was shot in company with the adult male and female. They evidently breed during

the rainy season, which lasts from February to May. Both sexes havea strong, although

not disagreeable, odour of musk, which appears to be stronger in the female’’*.

In the skins transmitted to me the scrotum of the male was large, and in the usual

marsupial prepudendal position t. In the female the orifice of the pouch was in the

corresponding part of the abdomen ¢. Both the size and position of the hind thumb

(woodeut, fig. 3) indicate the foot to be a grasper ina much lower degree than that of

the Phalangers and Opossums. Mr. Ramsay does not, indeed, adduce any instance

from his own or his correspondent’s observations of the opposition, or the mode of oppo-

sition, of the hallux to the other digits in the act of seizing or of climbing. Although it —

might be inferred, if the fruit of the Ptychosperma Alexandre was plucked from the

branch and not seized after falling to the ground, that Hypsiprymnodon attained it

by climbing.

Even in that case the act would seem to be occasional or exceptional; and the sum of

the evidence tells for the ground-dwelling habits of this small and interesting Marsupial.

The hallux here is, indeed, of more moment to the biologist in its homological than in

its teleological relations. It points, as in the case of more rudimental and useless parts

in other species, to the ancestral type.

Hypothetically, the earliest Marsupials are assumed to be pentadactyle; and this

assumption is supported by the following facts. The majority of the extinct Marsupials

* Ramsay, tom. cit. p. 34. + Plate LXXI. fig. 1. + Ib. fig. 2.

580 . PROF. OWEN ON HYPSIPRYMNODON.

from British mesozoic formations are polyprotodont *; and most of the existing species

of that dental division of the Order are pentadactyle f.

Among the Diprotedonts { the scansorial Marsupials retain the pentadactyle foot

with a maximized hallux §. In the gradatorial Diprotodonts, development has wrought

on the two outer toes (Iv and v), and especially on the fourth, with ultimate loss of the

innermost or first toe. This modification, characteristic of the Potoroos, reaches its

extreme in Macropus and Cheropus. The instructiveness of the foot of Hypsiprymnodon

lies in its indications of an early stage of modification of the pentadactyle type in the

direction reached by the great Kangaroos. The fourth toe is, indeed, the largest

(woodcut, fig. 3, Pl. LX XIT. figs. 6 & 7, 1v), but not so much larger than the fifth (ib. vy).

The size of both, being less exaggerated, shows the cleansing-digits (ib. 11 & 111) on a less

dwarfed scale. Finally, the hallux (ib. 1) is not yet lost: it is small, indeed, and so

placed as hardly to be opposable to any other digit, unless to the metatarsal part of m1.

As the habit of progression on hard ground increased, the share taken by the fourth

toe, as being most on a line with or most directly continuing groundward the tibia

or main pillar of the leg, would determine a greater flow of blood to the moving

powers of that toe; and as the exercise of these powers would become more strenuous

with the increase of weight to be so moved, especially when the motion was accelerated

by leaps, so the fourth toe has come, at last, asin the great Boomer Kangaroo, to be almost

exclusively the instrument applied to the resisting earth in the saltatory mode of

locomotion.

The reduction of the toes in terrestrial birds follows an analogous course. I need

hardly premise that the fifth toe in the foot of the Lizard is not developed in any bird.

The first toe, or hallux, shows its opposable character and grasping function in the

Perchers. When retained in a ground-bird, Apéferyz e. g. (Pl. LX XII. fig. 11,1), it is, as

in Hypsiprymnodon, minute, and placed above the level of the other toes. In the Cassowary

(PL LXXIT. fig. 12) it is lost; the second toe (1) is reduced in length and breadth, but

is provided with an unusually long and slender claw, which the bird may be seen occa-

sionally to apply in preening the plumage; the third toe (11) is that which has acquired

the largest size and takes the greatest share in the act of support and progression ; the

outermost toe (Iv) is smaller, but not so reduced as the innermost (1, Pl. LX XII. fig. 12).

Here we have an analogous reduction to that shown in the Potoroo.

Finally, in the Ostrich, besides 1, the next toe (11) is lost; and the swift and powerful

course of the bird is allotted, as in the Kangaroo, to the two outer toes, and mainly to

one of these (111), the outermost toe (Iv) being much reduced, but still characterized by

its five phalanges (Pl. LX XII. fig. 18).

A thaumatogenist || may, indeed, contend that the species Struthio camelus was

created, at the beginning, by primary power, with but two toes to each foot, in size,

supials of England,’ 4to, vol. i. pp. 12-75, plates i.-tii. (1877).

t Lom. cit. p. 105. + Ibid. p. 107.

§ Art. Marsupratra, ‘ Cyclopsedia of Anatomy,’ 8vo, 1841, p. 285, fig. 111.

|| ‘ Anatomy of Vertebrates,’ vol. ili, 1868, p. 814.

PROF. OWEN ON HYPSIPRYMNODON. 581

shape, and proportion best adapted to the need of this feathered denizen of the desert.

But the nomogenist *, in reply, asks for the significance, or an explanation, of the

retention of the metatarsal of the last toe (fig. 13, 11), the jointless pointed extremity of

which projects, like the splint-bone of the same.lost toe in the Horse, on the inner side

of the articular end of the huge metatarsal of the adjoining functional toe (111).

In speculating on the way of operation of the secondary law in the coming to be of

species and genera distinguished, among other characters, by the number and _pro-

portions of the toes, a nomogenist has hazarded the following supposition :—“ as the

surface of the earth consolidated, the larger and more produced mid hoof of the old three-

toed Pachyderms [ Paleotheriwm] took a greater share in sustaining the animal’s weight ;

and more blood being required to meet the greater demand of the more active middle toe,

it grew; whilst the side toes, losing their share of nourishment, and becoming more and

more withdrawn from use, shrank ; and so on, according to the hardening of the ground,

until only the hidden rudiments of metapodials [11 and Iv] remained, and one hoof became

maximized for all the work” +. The writer illustrates this notion by a subjoined

diagram of the structure of the foot in the ungulates of the Middle Hocene, the Miocene,

and the Pliocene periods {.

But since the publication of the volume above cited, the structure of the foot of an

older ungulate mammal, the Coryphodon of our plastic clay §, has been demonstrated by

the fossils of that genus discovered by Professor Marsh in the corresponding Tertiaries of

North America ||.

And here, as in the oldest known form of Marsupials, the primitive pentadactyle type

of foot is revealed, from which the successive perissodactyle hoofed forms have deviated

by concentration of size and force through strenuous use upon digit m1, and by

degeneracy through opposite physiological conditions affecting size and power, first of

digits 1 and v, which are lost in Palgotherium, next of digits m1 and tv, dwarfed in

Anchitherium and Hipparion, until finally these also disappear in Hquus.

An analogous modification under other functions of the hind foot is traceable in

certain extinct species of Bruta (Edentata, Cuwv.).

In Myloden digit 1 is obsolete; 1 is small, but retains the ungual phalanx;

development is concentrated on 111; the ungual phalanx is wanting and the others

stunted, in Iv and v J.

In Megatherium both digits 1 and 11 are gone; Iv and v are clawless, stunted, and

seem to have been developed in a common hoof-like callosity. The whole power of the.

foot as an uprooter is concentrated in digit 111**.

* ¢ Anatomy of Vertebrates,’ vol. iii. 1868, p. 814. t+ Tom. cit. p. 793. + Ibid. woodcut 614.

§ Owen, ‘ British Fossil Mammals,’ 8yo, 1846, p. 299, cuts 103, 10.

|| Marsh, “ Principal Characters of the Coryphodontide,” American Journal of Science and Arts, vol. xiv. July 1877,

p. 81, pl. iv. fig. 2. A more direct and continuous ancestry of existing Equines will shortly be described and

illustrated, in American Fossils, by the same author.

@ Owen, ‘ Description of Mylodon,’ 4to, 1842, p. 117, pl. xxii.

** Memoir on the Megatherium,’ 4to, 1860, p. 72, pls. xxv. andxxvi. The degree and course of mutilation and

of partial and exaggerated development of certain toes in the hind foot of other Bruta are detailed in pp. 74-77.

SECOND SERIES.—ZOOLOGY, VOL. I. 4H

582 PROF. OWEN ON HYPSIPRYMNODON.

In these ponderous slow-moving quadrupeds the exercise of the muscles and con-

comitant vigour of circulation and innervation in locomotion was inferior to those

affecting the main instrument in the swift course of the Horse and Kangaroo; the

claw-bearing toe mainly (if not exclusively) exercised in exposing the ramifications of the

root of the tree to be felled, was the subject of those influences which govern excessive

or exclusive development.

In conclusion, I may remark that the longest and strongest toe in the pentadactyle

hind foot of the Lizards is the fourth; and I deem it indicative of the nearness to

Saurians of the Implacentalia among mammals—an affinity so remarkably shown by

monotremes in other parts of the skeleton—that in the progressive reduction of the foot

in Marsupials the fourth toe, Iv, holds the preeminence.

In the march of Man the chief motive power is concentrated on the innermost toe

(1) as the main instrument, thence called the “ hallux,’”’ or “ great toe.” In this modi-

fication of the foot for bipedal station and locomotion the human kind stands alone in

the animal kingdom.

DESCRIPTION OF THE PLATES.

Puate LXXI.

Figs. 1 & 2. Hypsiprymnodon moschatus, Ramsay: nat. size. 1, male; 2, female.

Puate LXXTII.

Fig. 1. Side view of the skull of Hypsiprymnodon moschatus, Rms.

2. Upper view of the same.

3. Under view of the same. 1 1, 2, 3, incisors; c, canine; Pp 3, premolar; d4, milk-

molar; m 1, 2, 3, molars.

4.. Upper view of the mandible of the same. Letters as in fig. 8.

5. Bones of the forearm and paw of the same. 1, II, 111, Iv, V, digits.

6. Bones of the hind foot: £, entocuneiform ; 1 to v, digits.

7. The same, from the opposite (plantar) side.

8. Mandible of Hypsiprymnus Gilberti: c, canine ; p 8, premolar ; d 4, milk-molar; ™ 1, 2,3,

molars.

9. Bones of forearm and paw of the same. 1 to v, digits.

10. Bones of the hind foot, plantar aspect, of the same: 1 to v, digits. All the above

figures are of the natural size.

11. Bones of the foot of Apteryz.

12. The same bones of the Cassowary (Casuarius).

13. The same of the Ostrich (Struthio). The same letters are applicable to these three

figures, viz. e, epiphysis; 2, 3, 4, metatarsals; 1, 11, 111, 1v, digits.

ROwen. _ Trans. Linn. Soc. 5EeR.2.Zo00u-Vou.1.Pu72

; 17 AYP Ss TERY MN OD) On SRIOMENG Rio Even iaNiUiS7

CBerjean lith, TLAPTHRYX 1Z.CASUARIUS. 13.STRUTHIO Hanhart imp

[ 583 ]

XXIV. Descriptions of some minute Hymenopterous Insects.

By J. O. Wzstwoop, W/.A., F.L.S., Se.

(Plate LX XIII.)

Read April 4th, 1878.

THE fourth volume of the ‘Transactions’ of the Linnean Society (1798) contains the

figure and description of an almost invisible Hymenopterous insect, referred by the

author, Mr. Shaw, to the Linnean genus Jchnewmon, and described under the name of

Ichn. punctum, with a doubtful reference to the Ichn. atomos of Linnzeus. The specimen

is stated to have been taken on the surface of a window; and on various occasions I

have taken the same or a closely allied species creeping on windows, and of so minute

a size as to be only visible when seen against the light and in motion. The insect is

remarkable for its antennze terminating in an oval club (being a female), and the long

fringe of hairs with which the edges of the wings are furnished. It is at the present

day referred to the subfamily Mymarides among the Proctotrupide, and to the genus

Anaphes of Haliday (Entomol. Mag. i. 269).

The twenty-fourth volume of the ‘Transactions’ of the Linnean Society (1863) adds

materially to our knowledge of these minute insects by the publication of a memoir, by

Sir John Lubbock, ‘‘ On two Aquatic Hymenoptera, one of which uses its wings in swim-

ming.” In this remarkable memoir the author describes two different species—one

belonging to the Mymarides, and the other, as it seems to me, to an aberrant group of

the Eulophideous Chalcidide. Of the first of these two species as many as twenty-one

individuals were observed by Sir John Lubbock, each swimming in a basin of pond-water

with the assistance of its wings, and using its legs apparently only for walking, so that

its motion under water might almost be called a flight. This insect was at first regarded

as identical with Polynema fuscipes of Haliday, but was subsequently considered to be a

distinct species, and described under the name of Polynema natans, although that genus

possesses 13-jointed antennz in the males, whereas P. natans has only twelve joints in

its male antenne. As Sir J. Lubbock did not give any representation of the male

antenna, I have represented it in Plate LX XIII. fig. 1, from a specimen of the insect

communicated to me by Sir J. Lubbock. With regard to the genus to which this insect

belongs, it is to be observed that Dr. Arn. Foerster (Hymenopt. Studien, ii. Heft, pp. 118,

121) objects to the name Polynema, because there was a fish genus so named previously,

and to the synonymous name Hutriche of Nees von Esenbeck (Monogr. Ichn. Min.),

because Stephens had given the name of Hutricha to a Lepidopterous genus. He con-

sequently proposed for it the name of Cosmocoma. Considering that it is unnecessary

and inexpedient to alter such generic names because they happen to have been previously

employed in different classes of animals, I should have retained the generic name given

Au 2

584 PROF. WESTWOOD ON SOME

to the insect in Sir John Lubbock’s memoir, had it not been that the difference in the

number of joints in its antennz above noticed would not allow it to be arranged with

the other Polyneme, the number of joints of those organs in the Proctotrupide affording

the principal characters for generic classification. Hence I should prefer to arrange

Sir J. Lubbock’s insect in Haliday’s genus Anaphes, since it has 12-jointed male and

9-jointed female antennee, as in that genus, notwithstanding that Anaphes is placed in a

section having the abdomen sessile or subsessile, whilst it is pedunculated in Polynema;

but the figure of the female given by Sir J. Lubbock (pl. xxiii. fig. 2) shows so small

an amount of pedunculation as to throw doubt on this point; and it will be seen that

the wings of P. natans (figs. 4 & 5) agree exactly with those of Anaphes (figs. 8 & 9)*.

With regard to the natatorial habits of this curious insect, I may observe that the long

fringe of fine hairs along the margins of the wings is admirably adapted to retain a

bubble of air, needed for the respiration of the insect under water, which, as Sir John

Lubbock observes, takes place through spiracles in the usual manner. M. Victor

Audouin showed how this was effected in an insect belonging to the family Carabidee

and genus A/pus, which passed a considerable portion of its existence in salt water

beneath high-water mark, and which was clothed with long setee; and it is in like manner

owing to the sericeous covering of the Argyroneta that the diving-bell spider is able to

subsist under water.

It is a curious analogy that whilst these insects, the most minute of the Hymenoptera,

should be furnished with these beautifully fringed wings, the most minute species of the

Coleopterous order (viz. the Trichopterygia) should also have their wings similarly

fringed; no one, however, as far as I am aware, has ever observed a Trichopteryx under

water; and it may be suggested that the fringe of hairs in that genus is used as a supple-

ment to the narrow elongated wings in effecting the ordinary flight of the Beetle.

The generic name J/ymar has been retained for the singular little insect figured by

Curtis under the name of

Mymar PULCHELLUS (Plate LX XIII. fig. 3). Brit. Ent. pl. 411.

This insect has 18-jointed antennee in the male and 9-jointed in the female, with the fourt

joint remarkably elongated and slender ; the fore wings forming a long and filiform thread,

terminated by a dilated oval membrane (like a boatman’s paddle), which is deeply fringed

with hairs, whilst the hind wings are reduced to a short rigid bristle. The tarsi are

4-jointed; and the abdomen is fixed to the thorax by a long slender peduncle. As Mr..

Curtis figured the female of this insect, I have thought it advisable to give a figure of

the opposite sex (having taken specimens of both sexes by sweeping grass and low plants

on a hot bank near Richmond Park), as well as to show its difference from another

species of the genus, discovered in Ceylon by Mr. Staniforth Green, as well as from the

still more remarkable species collected in St. Helena by the late T. V. Wollaston.

* Tf this proposal as to P. natans being placed in the genus Anaphes be rejected, I beg to suggest that a new

generic name be given to it, and would carry out Sir J. Lubbock’s original intention, by giving to it the generic name

of Valkerella.

MINUTE HYMENOPTEROUS INSECTS. 585

MyMAR TAPROBANICUS (Plate LX XIII. fig. 4, ¢, and fig. 6, ?), Ward, in Ent. Mo. Mag.

Feb. 1875, p. 197.

In the work above referred to, the female only of this remarkable species is described.

I have now, however, the pleasure of describing both sexes, collected in Ceylon by

sweeping low herbage, by Mr. Staniforth Green, which are preserved as microscopical

objects in Canada-balsam slides; and the male is fortunately arranged so that its

structure can well be observed from above; but the female is fixed laterally. The head

is large, transverse, with a slight ridge in the male between the anterior part of the

eyes, the antennz being affixed on the fore edge of this ridge, considerably above the

mouth ; so that the geniculation of the antennz is quite of a different nature from that

of the Chalcididz. The antenne of the male are very long, 13-jointed, and quite filiform ;

the basal joint slightly curved, and the second joint short, the remaining eleven joints

nearly equal in length and slenderness; in the female the antenne are 9-jointed, the

first joint longer than in the male, the second joint rather short and thicker than the

apex of the preceding joint, the third joint slender and slightly longer than the second,

the fourth very long, slender, the fifth to the eighth small, and the ninth forming a large

oval mass. The thorax is elongate-ovate, the anterior part conical, and the hind part

truncated ; the abdomen is attached to the thorax in both sexes by a thickish peduncle,

slightly longer than the posterior coxe; the abdomen is very small in the male,

larger and pointed at the tip in the female, which is furnished with a short, sharply

pointed ovipositor, with two long lateral valves. The fore wings resemble those of IZ. pul-

chellus, except that they are not so strongly battle-door shaped at the tips; the disk of this

terminal part is furnished with two rows of short fine sete, andthe margins of this part

are furnished with strong bristles. The hind wings are about two thirds of the length of

the fore wings (thus differing from those of IZ. pulchellus), and are quite bristle-shaped ;

they have a few very short fine hairs on their margins and near the base; on the fore

margin they have three small sharp hooklets (fig. 7) for fastening them to the hind margin

of the fore wings. The legs are very long and slender, with 4-jointed tarsi and large

pulvilli. .

The whole insect is of an ochraceous colour, paler beneath, with the antennze and legs

concolorous; the apical half of the dilated portion of the fore wings is dusky, and the

tips of the antennze and tarsi are darker.

I also refer to the genus Mymar* avery singular species collected by the late lamented

T. Vernon Wollaston by sweeping low herbage in the Island of St. Helena, of which

fortunately, both sexes were procured, and which agree in having the fore wings of large

size, coriaceous, setose, convex, and, when shut, forming a semiglobular dome over the

abdomen, quite unlike any previously known Hymenopterous insect.

Mymar Wottastonit. (Plate LXXIII. fig. 8, d; fig. 9, 2.)

Niger, nitidissimus, glaberrimus, capite thorace paulo latiore, subquadrato-rotundato, convexo, postice

valde emarginato, thorace compresso, prothoracis lateribus subtus paullo dilatatis, pedunculo abdo-

* Tf it should be deemed necessary to separate this species from the genus Mymar, I beg leave to suggest that its

affinity therewith may be indicated by the employment of the generic or subgeneric name of Mymarilla.

586 PROF. WESTWOOD ON SOME

minis fere thoracis longitudine fulvo-testaceo ; abdomine oyali, nitido; antennis maris corpore toto

dimidio longioribus, nigris, articalis duobus basalibus fulvis ; pedibus elongatis, gracilibus, fulvis, apice

tarsorum obscuro ; tibiis posticis in utroque sexu setosis ; alis anticis ovalibus, convexis, semiglobosis,

coriaceis, nigris, valde nigro-setosis ; alis posticis fere linearibus, setosis. Long. corp. ¢ circ. ‘2 mill.,

9 -274 mill.; long. anten. maris °3 mill.

Habitat in gramimosis in Insula Sanctze Helene.

ALAPTUS Excisus. (Plate LX XIII. fig. 10, 3, and fig. 11, ?.)

I am indebted to Mr. Whitmarsh, of Wilton, near Salisbury, for an opportunity of

examining a very large number of glass slides, prepared for the microscope, containing

minute insects mounted in Canada balsam—an excellent plan for the examination of

such objects, so far as the observation of the general outline and of detached parts are

concerned, the gummy solution rendering the parts more or less transparent. Amongst

these specimens I found two insects belonging to the Mymarides, which I have no

hesitation in regarding as the male and female of the same species. Both specimens

had been reared from white blotches on oak leaves, evidently caused by the action of

the minute larva of one of the leaf-mining Vinee (Lithocolletes?). The blotches were

about 1} in. in diameter. The leaves were gathered on the 9th September, 1871; and

the little Mymars appeared on the 6th October; one of the moths appeared on the 16th

September, 1871, and two other kinds of parasitic flies on the 4th October following.

The action of the Canada balsam has destroyed the colours of the insects; so that the

following description is confined to structural characters; moreover the male insect

has unfortunately been fixed by the Canada balsam on its side, and the exceedingly

minute size of the creatures rendered any attempt at displaying them, by mee

the limbs in the usual manner, ineffectual.

The head in the male is of large size and of an oval form (seen laterally), transverse

in the female and widest behind; in this sex it appears to be furnished with two

large appendages, truncate at the tips, which may possibly be dilated palpi. The

antennee of the male are long and filiform, 10-jointed, the basal joint being the largest,

the remaining nine being nearly equal in size. The antenne of the female are 8-jointed,

the first joint large, the second smaller, the third considerably shorter and thinner than

the preceding, the fourth to the seventh gradually but slightly thickened, and the eighth

forming an elongated oval mass. ‘The details of the thoracic segments are not easily

determined, owing to the mode of preservation of the specimens; but the scutellum

seems to be of large size and semicircular. The abdomen is sessile, depressed, and

gradually pointed to the tip in the female, whilst it is more ovate in the other sex, with

the male ‘organ protruded. The wings are of equal size and shape in both sexes, the

posterior ones being as large as the anterior, which latter have a remarkable dilatation

near the base of the posterior margin, terminating in an acute notch; the remainder of

the margins of all the wings is fringed with long hairs; the legs are long, slender, and

terminated by 5-jointed tarsi with large pulvilli.

The 5-jointed tarsi, the number of joints in the antenne of the two sexes, the sessile

abdomen, and the very long narrow wings, agree with the characters of Haliday’s genus

Alaptus given by Walker in the‘ Annals of Natural History,’ vol. xviii. (1846) p. 50. Of

MINUTE HYMENOPTEROUS INSECTS. 587

this genus two or, more probably, only one species is recorded in this country, namely

A. minimus, “‘ferruginosus, antennis et pedibus pallidis ;” the supposed second species,

A. fusculus, “ Preecedente major colore obscurior antennis longioribus, vix revera species

distincta”’ (op. cit. p. 51). Another species, 4. pallidornis [? pallidicornis], is slightly de-

scribed by Foerster, found near Aix la Chapelle. It is scarcely half as long as 4. mini-

mus, with yellowish-white antenne (Hym. Stud. ii. p. 120). As these authors do not

mention the singular dilated and excised base of the fore wings, I consider the one before

us distinct, to which may be applied the name of ALAPTUS EXCISUS.

A, antennis maris corpore paullo longioribus, feminz corpori «qualibus ; alis anticis basi postice dilatatis

et subito excisis. Insecta minutissima. Long. cire. } mill.

The second insect described by Sir J. Lubbock, under the name of Prestwichia aquatica,

offers considerable difficulty as to its true position, in consequence of the loss of the

typical specimens (all six of which are described as having been females) and the different

descriptions given of the structure of the antenne. Thus, in the Latin description

(p. 140) supplied by Mr. F. Walker, we read “‘antennz graciles, subclavate, inarticulate ;

clava longi-fusiformis ;”’ and in his English description they are described as ‘‘10-jointed,

the first joint nearly as long as the flagellum; the second, third, fourth, fifth, and sixth

joints equal in thickness, successively decreasing in length; seventh fusiform; eighth,

ninth, and tenth forming an elongate-fusiform club.” In the specific description, by Sir

J. Lubbock, they are said to consist of six or seven segments, a conjectural second seg-

ment (shown by a trace of a division near the end of the basal joint) being short and

round, the third joint longer and increasing somewhat in size towards the apex, the

fourth similar in shape but smaller, the terminal part of the organ forming a club-like

mass, which appears to be composed of three segments, the last joint, however, being

very indistinctly marked. In his fig. 10, and in the upper antenna in fig. 11, the antenne

are represented as only 4-jointed, whilst in fig. 12 and in the lower antenna of fig. 11

they are figured as 5-jointed. From this diversity of description, however, we may conclude

that the first joint is considerably elongated, the antenna being geniculated at its extremity.

The real second joint (the supposed preceding joint being, I believe, an optical delu-

sion) is shown in fig. 12, followed by a shorter third joint, the remainder of the an-

tenna consisting of the clava, of which the first division alone is indicated in fig. 12,

whilst the second division, as described in the text, is more or less obsolete. As to

the intermediate third, fourth, fifth, and sixth joints mentioned in Mr. Walker’s English

description, they are as illusive as the “ antennz inarticulate” of his Latin description.

We are warranted, then, in concluding that the antennze were geniculate* at the end

of the long first joint, and that the terminal portion formed a more or less distinctly

8-articulated clava. Such characters, however, remove Prestwichia from the My-

marides, and associate it with smaller aberrant Hulophides. The structure of the

tarsi of Prestwichia also, in my opinion, removes them from the Mymarides, and

brings them, in like manner, to the Eulophides. It is true both Mr. Walker and Sir

* Tt is proper to observe that in most of the Chalcidide the antenne are inserted low down in the face, close to

the mouth, the basal joint extending upwards to the crown of the head, and the remainder of the antenna being bent

downwards at a sharp angle, and only reaching to the mouth when at rest.

588 PROF. WESTWOOD ON SOME

J. Lubbock distinctly describe the tarsi as 4-jointed, the latter adding that the fourth

segment (pl. xxii. fig. 15) is in all the legs very small. Referring to my genus Tricho-

gramma (as described in Mr. Taylor’s ‘ London and Edinb. Phil. Mag.’ 1838, p. 444), in

which the tarsi are described as 8-jointed,—a description which was adopted and

confirmed by Mr. Haliday, one of the clearest-sighted of entomologists,—Sir J. Lubbock

is “disposed to consider that the tarsus in Trichogramma is in reality 4-jointed, and

that Westwood and Haliday’s descriptions must be amended”’ (p. 141). Now, whoever

has studied these minute Hymenoptera is well aware that in many of them the

pulvillus of the tarsi is greatly enlarged, having the ungues affixed at the sides towards

the base, the tarsus itself being only 8-jointed (see, for example, the figures of the

tarsi of the Chacidideous genera published by Mr. Curtis in his ‘ British Entomology,’

especially those of Hulophus, pl. 183; Encyrtus, pl. 395; and Phagonia, pl. 427; and

T have not the least hesitation in affirming, from the inspection of Sir J. Lubbock’s

figures 10, 11, 14 & 15, that the tarsi of Prestwichia are only 3-jointed, with an enlarged

pulvillus, as more distinctly shown in fig.15. This character, therefore, also removes the

genus from the Mymarides—amongst which it is arranged by the Rev. 'T. A. Marshall in

his ‘Catalogue of British Hymenoptera Oxyura,’ between Cosmocoma (Polynema) and

Caraphractus.

The structure of the fore wing of Prestwichia is also indicative of a similar want of

relation with the Mymarides. As will be noticed by comparison of the figure of its wing

(fig. 18) with those of Polynema (fig. 4) and Anaphes (fig. 8), without dwelling on the shape

of the wing and its rounded extremity, it will be seen that the thick subcostal vein

extends half the length of the anterior margin of the wing, where, at its apical extremity,

“it turns inwards and ends abruptly ’’—a character never found in the Mymarides.

Sir John Lubbock, as above stated, has referred to my genus Trichogramma; and I

cannot but think that Prestwichia belongs to the little group of which the former genus

is the type, and which Foerster has formed into his family Trichogrammatoide (Hym.

Studien, Heft 2, p.87). In my original description of Trichogramma I described the

antenne thus :—“ breves, 6-articulate, articulo 1mo longo, 2do brevi, gracili, 3tio quam

2do majori, crassiori, 4to et 5to brevibus, 6toque maximo oblongo-ovato apice acuminato,”

accompanied by a figure both of the antenna and the 3-jointed tarsus. More recent

examination of the antenne leads me to think that the supposed second joint is only

the narrowed extremity of the long basal joint (as represented in the accompanying

Plate LX XIII. fig.12); nor am I clear that the large oval apical joint is not in reality

composed of two or three joints closely soldered together. The most remarkable cha-

racter of Lrichogramma consists of the curious mode in which the fine hairs of the

fore wings are arranged in lines, of which I am not aware of any similar instance

throughout the order Hymenoptera. Plate LX XIII. fig. 13, represents the fore wing of

Trichogramma evanescens, of which no figure has hitherto been published; and it will

be seen that Prestwichia differs from it in having the fine hairs scattered over the

wing irregularly.

In the Entomological Magazine (vol. i. p. 340, July 1833) Mr. Haliday published the

description of a new genus with trimerous tarsi, under the name of Calleptiles (giving

MINUTE HYMENOPTEROUS INSECTS. 589

as a synonym the name of Microma, printed without description in Curtis’s ‘ Guide’).

The type of this genus, C. latipennis, had been reared from subcutaneous larve in the

leaves of Aquilegia by Curtis, and is described as having “ alee hyaline radice obscuriores.”

To the description is appended an editorial note that the insect is probably identical with

my Trichogramma evanescens ; and in the ‘ Monographia Chalciditum,’ 1839, p. 18, the

last-named species reappears as belonging to my genus Péeroptrix, placed in a distinct

section (1), erroneously described as having 4-jointed tarsi (which is the distinguishing

character of Pteroptrix), with Calleptiles latipennis, Haliday, given as a synonym. In

the ‘Entomologist,’ 1840-1842, the insect again reappears, in the descriptions of Mr.

Haliday’s beautiful series of plates of Chalcididee, under the name of Trichogramma

evanescens mas ; but the figure of the insect (Entomol. Mag. pl. K. figs. 4, 4a—4d) differs

in several important respects from my insect and its details, and evidently represents

not only another species with a dark band across the fore wings, but probably a male

insect, whilst mine was most probably a female. In Mr. Haliday’s figures the number

of lines in which the hairs of the fore wings are arranged is considerably greater than in

my insect; and the antenne are evidently 7-jointed, and strongly setose, the first joint

of moderate size, the second about the length of the former and much thicker; the

third resembles the second, as does also the fourth, which, however, is larger than the

third, whilst the fifth, sixth, and seventh are evidently coalesced, together forming a

3-jointed oval mass.

A species of this group is described by Foerster in the ‘Verhandlungen’ of the

Rhineland Natural-Hist. Soc. vol. viii. p. 26, pl. 1. fig. 9, a, b,c, under the name of

Trichogramma Walkeri, which has the antennee 4-jointed or (if the last joint consists, as

the author supposes possible, of three joints soldered together) 6-jointed—the Ist of

moderate size, the 2nd and 8rd equal, subovate, and nearly as large as the basal joint,

and the terminal part of the antenneze forming an elongated oval mass, the whole

antenne being very slightly setose. In his ‘ Hymenopt. Studien,’ Heft 2, p. 89, Foerster

incorrectly alludes to this insect as having been described by him (loc. cit. supra) under

the name of Calleptiles Walkeri, and proposes, on account of the great length of the

ovipositor of the female, to form it into a separate genus Centrobia.

In Foerster’s figures of the antennz of Poropoea, those of the male are represented as

7-jointed, the first of moderate size, the 2nd small ovate, and the five following equal-

sized, twice as large as the 2nd, and shortly ovate. The female antenne are also 7-jointed,

the 3rd and 4th joints thicker than in the male, and the last three joints clearly uniting

to form a terminal oval mass. In Ratzeburg’s second species, Ophioneurus signatus, the

basal joint of the antennz is of moderate size, the 2nd as thick and nearly as large as

the first, the remainder (7 joints as figured) forming a large oval mass.

In the ‘Annals of Natural History,’ 2nd ser. vol. vii. pp. 211-213, Mr. Walker pub-

lished, from Mr. Haliday’s MSS., descriptions of several of these very minute insects

found in Ireland—namely, Myina annulipes and livens, Chetostricha (un. g.) dimidiata,

Trichogramma vitripennis, and Oligosita (n. g.) collina—accompanied by the following

note, also by Haliday.

SECOND SERIES.—ZOOLOGY, VOL. I. AT

590. PROF. WESTWOOD ON SOME

** Synopsis of the Trichogrammini.

«Tarsi trimeri: tibiz antice calcari apice inciso: antenne articulis 6, 3 extremis in clavam

coarctatis (an semper?) : abdomen subsessile: statura Apheline (Myine) fere et huic magis affines

videntur quam Hulophinis: Oligosita vero pedibus gracilibus, tarsis 2di paris elongatis, alis longe fim-

briatis Thysani speciem mentitur.

GENERUM CONSPECTUS.

* Ale antice seriatim pubescentes.

+ Vena costam sinu tantum attingens ad ortum radii. 1. Trichogramma, Westw.

tt Vena costam longius decurrens ante ortum radii. 2. Chetostricha, n. g.

** Ale vage pubescentes.

+ Ale: antice late, margine subtiliter ciliatee. 3. Brachista, nu. g.

tt Alse antice anguste longe fimbriate (plumatze). 4. Oligosita, n. g.”

Of the species of these minute insects having the hairs of the wings disposed in linear

series and 3-jointed tarsi, Foerster describes five genera, Viz. :—

1. Poropoea, Foerst. in Verhandl. nat. Vereins Rheinlande, 8th year, p. 28, pl. 1. fig. 10, de (1851)

= Ophioneurus, Ratzeburg, Ichn. der Forstinsecten, iii, p. 197 (1852), consisting of two species, which

are parasitic in the eges of Apoderus curculionoides and Rhynchites betule, which reside, in the larva state,

in rolled-up oak-leaves.

2. Trichogramma,Westw.

3. “ Chetosticha, Walker” [rectius “ Chetostricha, Haliday,” pace Foersteri].

4. Lathromeris, Foerster, and

5. Centrobia, Foerster.

But he adds to the family a section consisting of three other genera, in which the hairs

of the wings are not arranged in lines, viz. :—

6. Asynacta, Foerster ; 7. Brachista, Haliday ; and 8. Oligosita, Haliday.

And it is, I do not doubt, in this latter section that Prestwichia must be placed. ‘These

three genera are thus tabulated by Foerster :—

aa. Fligel nicht reihenweise behaart.

fe Filer siebengliederig. Asynacta.

jf. Fiibler sechsgliederig.

g- Vorderfliigel breit, am Rande mit zartem Haarsaum. Brachystica [rectius Brachista*, Hal.1.

gg- Vorderfliigel schmal, lang gewimpert. Oligosita, Hal.

The difficulty pointed out in a preceding part of this paper as to the precise character

of the antenne of Prestwichia prevents me from determining whether it does not in

reality enter into the genus Oligosita, Hal., of which the antennz are thus described :—

“ Subclavate, brown towards the tips, much more than half the length of the body, first

joint very long, second cup-shaped, third and following forming a spindle-shaped club.”

This description seems to indicate that the specimen described by Haliday was a male,

as these organs in the opposite sex would probably be differently formed; at all events,

in this uncertainty, I venture to describe and figure in this place a minute species, pro-

bably congeneric with Prestwichia (and of which I took several specimens on the 18th

of August, 1887, by sweeping grass in Richmond Park), under the name of

* Changed by Foerster as being too near to Brachistes, Wesmael.

MINUTE HYMENOPTEROUS INSECTS. 591

OLIGOSITA SUBFASCIATA. (Pl. LX XIII. figs. 14-19.)

Picea, capite et dorso thoracis fulvis, hujus parte postica picea, linea media longitudinali fulva, oculis

magnis, nigris ; antennis pedibusque fulvescentibus ; alis anticis nubila media pallide fusca notatis.

Long. corp. ? millim. LExpans. alar. 1} millim.

Habitat Richmond Park, comit. Surrey, mense Augusto capta.

The antennee (fig. 15) are about twice the length of the head, and consist of a long

basal joint, more slender in the middle, a second joint, not quite half the length of the

first, followed by a minute annular joint, the fourth joint being rather smaller than the

second, and three terminal joints forming an elongate oval setose mass.

The mandibles (fig. 16) are strong, horny, and terminated by four small obtuse teeth ;

the maxille (fig. 17) are large, and terminated by a large fleshy setose lobe, having a

very short cylindrical and apparently exarticulate palpus, with two terminal short sete ;

the lower lip (fig. 18) is nearly of equal size with the maxille, elongate-conic, rounded

at the extremity; the labial palpi very small, slender, cylindrical, and terminated by a

small seta. The tarsi (fig. 19) are 3-jointed, terminated by a large pulvillus, with two

small lateral ungues. The fore wings are dilated at the hinder extremity; they are well

clothed with fine setee, and their apical margin is fringed with very long hairs; the sub-

costal vein is free for a short distance from the base of the wing, where it unites with

the margin, and extends to the middle of the wing, terminating in a very short and

nearly triangular stigmatic branch extending into the disk. The hind wings are very

narrow, with a prominent hook before the middle of the costal edge, the margins with

_a fringe of long hairs, and a row of fine hairs on the disk.

Amongst the numerous minute Hymenoptera collected at or near Oolombo, in

Ceylon, by Mr. Staniforth Green, which he has mounted in Canada balsam and kindly

presented to me, are several Trichogramme and ailied species, of which I add descrip-

tions and figures as further illustrating the relations of Prestwichia,

Oxigosita StanrForTHi. (Pl. LX XIII. figs. 20, 21.)

Tota luteo-fulva, oculis fuscis; antennis perbrevibus, geniculatis, articulo primo magno, secundo semiovali,

tertio annuliformi, quarto parvo subrotundato, reliquis tribus clavam longe ovalem formantibus ; alis

anticis subangustis, angulo antico apicali rotundato-truncato, longe ciliatis, disco parum setigero,

ramo stigmatico distincto, apice angulato-truncato ; alis posticis angustis, utrinque longe ciliatis ;

abdomine elongato-conico ; oviductu longitudine abdominis, e basi hujus exeunte, apice serrato,

Long. corp. 0-017 une.

Habitat in graminosis in insula Taprobana (D. Staniforth Green).

The insect here described is a female, as shown by the exserted ovipositor (fig. 21),

which is finely serrated at the tip. The structure of its antennz and wings are very

clearly shown in the specimen, and lead me to infer that it cannot be generically separated

from Ol. subfasciata. The peculiar form of the fore wings, which have the hind

margin straight and produced, whilst the anterior apical angle is rounded off, gives the

wings a form quite unlike that of any other Hymenopterous insect.

At 2

592 PROF. WESTWOOD ON SOME

Oxicostta ? Nopicornis. (Pl. LXXIII. fig. 22.)

Tota fulvo-lutea, oculis obscuris, venis alarum pedibusque pallide luteo-albis ; capite lato; antennis

brevibus, geniculatis, articulo primo elongato, secundo brevi obconico, 4: proximis semiovalibus,

fere eequalibus, singulo secundo parum majore, setis longis antice subverticillatis, articulo apicali

(septimo) ovali, quam praecedens paulo minore; alis anticis magnitudine ordinaria, angulo antico apicali

rotundato-truncato, longe ciliatis, disco parum setoso, setis in lineas 5 irregulariter dispositis ; alis

posticis angustis utrinque longe setosis, angulo hamifero costali conico; genitalibus masculinis

porrectis quasi biarticulatis, parte basali subtus spma armata. An mas Ol. Staniforthi? long.

corp. 0014 une.

Habitat cum precedente in Taprobana (D. Staniforth Green).

Iam obliged to add a mark of interrogation to the generic name applied above to this

insect, from ignorance of the true structure of the male antenne of the type of Oligosita.

The specimen before me is a male, as shown both by the exserted genitalia and the

structure of the subverticillated antennee (fig. 23), with the terminal joints not forming a

triarticulated mass.

TRICHOGRAMMA (APROBOSCA) EROSICORNIS. (Pl. LXXIII. fig. 24.)

Corpus breve, contractum; abdomine sessili; capite magno ; antennis brevibus, geniculatis, articulo primo

longo, secundo paryo semiovali, tertio minimo annuliformi, ultimo (quarto) elongato, valde irre-

gulari, quasi eroso, subuodoso, nodulis verticillatis; mandibulis angustis, curvatis, apice bidentatis ;

maxillis cum labio massam ovalem apice breviter setosam formantibus palpisque brevibus exarticulatis

instructam; pedibus sat longis, tarsis 3-articulatis, pulvillo magno ; alis anticis latis, ciliis medioeriter

elongatis marginatis, ramo stigmatico elongato, apice crassso inflexo, setis disci in lineas circiter 10

dispositis ; alis posticis-angustis, fere linearibus, longius ciliatis, angulo hamifero costali prominente. |

Tota fulva, oculis obscuris, pedibus antennisque pallidioribus. Long. corp. 0:018 unc.

Habitat in insula Taprobana (D. Staniforth Green).

The form of the fore wings, which are relatively broad, and the linear arrangement of

the fine hairs on their disk, prove that this curious insect must be closely allied to our

English Trichogramma ; but the singular antennee (fig. 25), of which the terminal nodose

portion under my strongest power shows no trace of transverse articulation (although

the groups of verticillate hairs might be supposed to indicate so many joints which

have become ankylosed), render it necessary to form at least a separate subgenus for

its reception. In favour of the exarticulation of this terminal part, the genus Thysanus,

described by Haliday *, may be quoted, in which the terminal joint is exarticulated,

elongated, and cylindrical; it has been placed in the section Tetrastichoidee by Foerster.

The antennze appear, at first sight, to consist of only three joints, of which the last is

very long and linear, especially in the males; but between the second and third joints

there appear to be three very minute annuli (more distinctly shown in the figure of the

female than in the male antenne f). The tarsi are not described, except that the inter-

mediate pair are said to be “longissimi” (very long); the detached figure of one of the

tarsi represents it as five-jointed, with a scarcely visible pulvillus; but Foerster states

* Annals Nat. Hist. iy. 234, December 1839. This name has been unnecessarily (as it seems to me) altered by

Foerster to Triphasius (Hymen. Studien, p. 84).

+ ‘ Entomologist, No. 26, December 1842, plate K. fig. 3a male, fig. 36 female antenna; fig. 3c tarsus.

MINUTE HYMENOPTEROUS INSECTS. 593

that they consist only of four joints (J. c.), and hence the genus is placed by that author in

the section Tetrastichoide. The wings, however, are very strongly ciliated ; and notwith-

standing the difference in the number of joints of the tarsi, I should be inclined to place

the genus near Pteroptriz, Westw. (unnecessarily altered to Pterothrix by Foerster), and

not far removed from the Trichogrammatini.

EXPLANATION OF PLATE LXXIII.

The whole of these figures are highly magnified,

Fig. 1. Antenna of the male of Valkerella (Polynema) natans.

. Tarsus of the fore leg of the same species.

. Mymar pulchellus, male.

. Mymar taprobanicus, male.

5. Tibia and tarsus of the same insect.

6. Mymar taprobanicus, female.

7. Hooklets of wings of this species, necessarily represented much too coarse.

8. Mymar Wollastonii, male.

9. Mymar Wollastonii, female.

10. Alaptus excisus, male.

11. Alaptus excisus, female.

12. Antenna of Trichogramma evanescens.

13. Fore wing of the same species.

14. Oligosita subfasciata.

15 to 19. Parts of the same insect : fig. 15, antenna ; fig. 16, the mandible; fig. 17, maxilla;

fig. 18, labium ; and fig. 19, the tarsus.

20. Oligosita Staniforthii, female.

21. Extremity of ovipositor of the same insect.

22. Oligosita? nodicornis, male.

23. Antenna of O. nodicornis 3.

24. Trichogramma (Aprobosca) erosicornis.

25. Antenna of the same species.

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JOWestwood del. Trans. Linn Soc, See. 2 Vou.1Pu.73.

SPECIES OF MYMAR ALAPTUS, OLIGOSITA, & TRICHOGRAMMA.

Berjeau bth. With detacts all highly magnified. Hanhart imp.

—"

‘

hee

Abisara dumajanti, 566; haquinus,

546, 566; kausambi, 545, 566;

savitri, 545, 566; tanita, Hewit-

son, 546, 566.

Abranchiata, remarks on, 211.

Acanthozoon, n. g., Coll., 86.

armatum, el. (Pl. XVIIL.),

87, 95; papilio, 87, 95.

Accipiter nisus, skull of, 135.

virgatus, 313, 350.

Accipitres of Philippines, 313.

Accipitrine, 313.

Acidalia exclusa, 562.

Acidaliidee, 562.

Acrea thalia, 193.

Acridotheres cristatellus, 353.

Acrocephalus orientalis, 352.

Actinia abyssicola, sp. n. (Pl. XLY.),

297; gelatinosa, sp. n., 298.

Actiniade, 295.

Actiniaria, general remarks on, 303.

, on new forms of, dredged in

the deep sea, 295.

, Pelagic, surface-swimming

species of, 295.

Actinine, 297.

Adolias adonia, 539, 564; asoka,

539, 565; garuda, 539, 564;

jama, 539, 564; laverna, 539,

565; Iudekingii, 539, 565; mo-

nina, 539, 565; ramada, 565.

Agiale, 204.

Kollari (Pl. XXIX.), 205.

ABgialitis dubia, 353; Geoffroyi,

353 ; mongolica, 353.

Aigithognathe, facial morphology of,

107.

, palate, varieties of, 111.

A®gocera amalthea, 202.

INDEX.

Athopyga magnifica, Sharpe, 342,

353; Shelleyi, Sharpe, 342, 353.

Aetomorphee, structure skull of, 128.

Agama colonorum, sacral plexus of,

(Pl. LXVI.), 517.

Agarista, position of, 157.

agrius, 204; alethe, 203;

Batesii, 203; Doleschallii, 203 ;

eriopis, 204; Lindigii, 202; Moorei,

203; pales, 204; pedasus, 204;

Semperi, 203; simyra, 204.

Agathia discriminata, 562.

Aglaope infausta, venation wings of,

160.

Alaptus excisus, Westw., n. sp. (Pl.

LXXIII.), 586, 587; fusculus, 587;

minimus, 587; pallidornis, 587.

Alcedininze wanting vomer, 127.

Alcedinide, 317.

Alcedo bengalensis, 317, 351.

Alcyone cyanipectus, 351.

Alligator, sacral nerves of, 523.

Allman, Prof., on the structure and

systematic position of Stephanoscy-

phus mirabilis, the type of a new

order of Hydrozoa (Pl. XIV.), 61.

Amage auricula, 508.

Amathusia phidippus, 538, 564.

Amaurornis olivacea, 354.

Amblypodia adatha, 548, 566; am-

phimuta, 548, 566; anthilus, 548,

566; antimuta, 548, 566; apida-

nus, 549, 566; aroa, 548, 566;

atosia, 548, 566; centaurus, 548,

566; Diardi, 549, 566; epimuta,

548, 566; eumolphias, 548, 566;

hypomuta, 549, 566; inornata,

566 ; metamuta, 548, 566 ; vibara,

566.

Amesia juvenis, n. sp., Butler, 559 ;

pexifascia, 559.

Ammotrypane arctica, M‘Intosh, n.

sp. (Pl. LXV.), 505; aulogaster,

Rathke, 504.

Amphiporus lactifloreus, 73 ; hastatus,

73; spectabilis, 77.

Amphitrite cirrata, 508.

Anzeretes parulus, yomerine cartilages

of, 110.

Anas luzonica, 354. :

Ancistrosyllis greenlandica, M‘Intosh,

n. sp. (Pl. LXY.), 502.

Anderson, Prof. J., on the osteology

and pterylosis of the Spoon-billed

Sandpiper (Eurynorhynchus pyg-

meus, Linn.), (Pl. XXXY.), 213.

Anisopoda, members of, from Atlan-

tic, 23.

Annelida obtained during cruise of

H.MS. ‘ Valorous’ to Davis Strait

(1875), 499.

Annelids, on mode of preservation

(footnote), 499.

of Greenlandic seas, Dr. M<‘In-

tosh on, 499.

Anops malayica, 546, 566; sperthis,

546, 566.

Anous stolidus, 354.

Anthreptes chlorigaster, sp. n., 342,

353; malaccensis, 342, 353; ma-

laccensis, os uncinatum of, 111.

Anthus gustavi, 338, 352.

Apaturide, modifications apical seg-

ment of (Pl. LVI.), 360.

harpagones of, 363.

tegumen of, 362.

, values genital armature of, in

family, genera, and species, 364,366.

596

Aphneeus lohita, 549, 566.

Apical segment in European Rhopa-

locera, modifications in various fa-

mnilies of, 359, 364.

Apoderus curculionoides, 590.

Appendices anales of Lepidopteron,

357; of Trichopteron, 357.

Appias cardena, 551, 567 ; eleonora,

551, 567; figulina, 551, 567; na-

thalia, 551, 567; nero, 551, 567 ;

panda, 551, 567; plana, n. sp.,

Butler, 551, 567.

Apseudes cxca, Willemoes-Suhm, 24.

Aquiline, 313.

Arachnothera dilutior, 341, 352.

Arborophila, sp. ? 353.

Arcania granulosa, sp. n. (Pl.

XXXVIIL), 240.

Archzostomata compared with Deu-

terostomata, 209.

Archaster of W. Indies, 50.

Arctiidee, 557.

Arcturus of W. Indies, 50.

Ardea purpurea, 354.

Ardeide, 349.

Ardetta cinnamomea, 354 ; flavicollis,

354; sinensis, 354.

Argina cribraria, 557.

Argynnis, 159.

Aricia Cuvieri, 504; foetida, 504;

ercenlandica, M‘Intosh, n. sp. (Pl.

LXV.), 504; Kupfferi, 504; La-

treillii, 504. ;

Armature, male genital, of the Euro-

pean Rhopalocera, 357.

Artamide, 323.

Artamus leucorhynus, 323, 351.

Ascidians, on new forms of deep-sea,

287.

Askonema setubalense, spicules of,227.

Aspidogaster, comparison of 'Trema-

tode with, 210:

Astacus, deep-sea blind species, 48.

fluviatilis, 46 ; pellucidus, 49 ;

zaleucus, n. sp. (PI. X.), 49.

, anatomy of, 48.

, characters of, 49.

, colour and mode of life,

50.

, compared with the mam-

moth cave species, 49.

Astictopterus armatus, 555, 568 ;

diocles, 555, 565; gemmifer, n. sp.,

Butler, 555, 668; jama, 555, 568;

INDEX.

sindu, 555, 568; stellifer, n. sp.,

555, 568; zanites (Pl. LXIX.),

55 5,568.

Astrogonium of W. Indies, 50.

Astur soloénsis, 350; trivirgatus,

350.

Asynactia, 590.

Atella phalanta, 544, 565; sinha,

544, 565.

Athis japyx, 178.

Athyma amhaia, 540, 565; clerica,

n. sp., Butler (Pl. LXIX.), 540,

565; idita, 541, 565; kresna,

540, 565; laryma, 540, 565; leu-

cothoé, 540, 565; nivifera, n. sp.,

Butler (Pl. LXIX.), 540, 565;

pravara, 541, 565; sabrata, 540,

565; urvasi, 560.

Atlantic Crustacea from ‘ Challenger ’

Expedition, 23.

Attacus atlas, 561.

Atya, 27.

Auditory ossicula(Pls.LVIII.-LXIY.).

—— of Mluride, 392, 398;

of Anthropoidea as compared with

man, 379; of Antilocapride, 424;

of Antilopidea, 427; of Artiodac-

tyla, 421; of Baleenoidea, 456,464 ;

of birds, 490; of Bovide, 425,

432; of Camelide, 424, 431; of

Canide, 394, 395; of Carni-

vora, 390; of Castoride, 410; of

Cebide, 383; of Centetidee, 440,

444; of Cervide, 429, 482; of

Cetacea, 450-464; of Chevrotains,

423; of Chimpanzee, 378; of

Chinchillide, 416 ; of Chiromyide,

388, 390; of Chiroptera, 445—

449; comparison of monkeys and

lemurs, 389; of Cryptoproctide,

392, 394 ; of Cynomorpha or tailed _

Old-World monkeys, 380-383; of

Dasyproctide, 416; of Dasyu-

ride, 483, 487; of Delphinide,

459, 464; of Didelphia, 477-487 ;

of Didelphyide, 480, 487; of Dipo-

didwe, 414; of Duplicidentati, 419;

of Echidna, 488, 490 ; of Kdentata,

470-477; of Emballonuride, 449 ;

of Entomophaga, 472; of Equide,

420,431; of Erinaceide, 438, 444 ;

of Felide, 391 ; of Fissipedia, 390 ;

of Fossorial Rodents, 418 ; of Galeo-

pithecus, 442, 443 ; of Galaginide,

387, 389; of Giraffe, 429, 432;

of Gorilla, 377; of Hapalide or

Marmosets, 385; of Hyznide,

391, 392; of Hylobates, 379; of

Hyraces, 432; of Hystricide, 414;

of Hystricomorpha, 418; of Indri-

sine, 386; of Insectivora, 432—

444; of Lemuride, 386; of Lepo- .

ride, 417; of Loricata, 472, 476 ;

of Macropodide, 478, 487 ; of Ma-

eroscelids, 440,444; of man, 372;

of Manide, 474; of Marsupialia,

477-487 ; of Monotremata, 487—

492; of Muride, 410; of Muste-

lide, 395, 396 ; of Myogalide, 434,

444 ; of Myomorpha, 418; of My-

oxide, 409; of Nycticebide, 387,

389; of Octodontide, 415; of

Orang-utan, 378 ; of Ornithorhyn-

chus, 491, 493; of Otariide, 400,

407 ; of Peramelide, 486 ; of Peris-

sodactyla, 419 ; of Phalangistidiv,

483, 487 ; of Phascolomyidee, 485,

487; of Phocide, 404, 407; of Phyl-

lostomide, 449; of Physeteride,

456,463; of Phytophaga, 470; of

Pinnipedia, 399-407 ; of Platanis-

tide, 462, 464; of Proboscidea, 430;

of Procyonide, 392, 397; of Rhi-

nocerotidie, 419 ; of Rhinolophide,

446; of Rodentia, 407-419; of

of Saccomyide, 413; of Sciuride,

407, 408 ; of Sciuromorpha, 418 ;

of Simiide, 377; of Sirenia, 464—

470; of Soricide, 433, 444; of

Spalacide, 412; of Suide, 422,

431; of Talpide, 436, 444; of

Tapiride, 420; of Tragulide, 423,

431; of Trichechide, 401, 407;

of Tupaiide, 441, 444; of Ungu-

lata, 419-431; of Urside, 398,

399; of Vespertilionide, 448; of

Viverride, 392; of Ziphiide, 457,

464.

Axiothea catenata, 507.

Balabae, birds of, 310.

Basilan, birds of, 310.

Baza magnirostris, 350.

Belenois cynis, 551, 567.

Bendidex, 561.

Bermudas, on a Nebalia from, 26.

Birds’ palate, Aigithognathous varie-

ties of, 111.

Birds, sacral nerves of, 528; skull,

structure and development of, 99.

Birnara, n. g., nubila, Butler, 560.

Blind deep-sea Astacus, 48; Deca-

pods, 50; Tanaid, 23.

Blyth, Mr., on Sus cristatus, 255.

Botidide, 563.

Botys pterophoralis, 563; sp. ?, 563.

Brachista, 590.

Brachypodide, 334.

Brachysticta (syn.), 590.

Brachyurus erythrogaster, 308, 352 ;

propinquus, sp. n., 330, 352 ; sor-

didus, 309, 331, 352; Steerii, sp. n.

(Pl. XLIX.), 329.

Branchiz in Cidaride, 569; Echi-

noidea, 569.

Brown Linnet, facial arches of, 108.

Bryozoa of W. Indies, 50.

Bubo philippensis, 350.

Bubulcus coromandus, 349, 354.

Buceros hydrocorax, 322, 351.

Bucerotide of Philippines, 322;

structure face of, 126.

Buchanga cineracea, 324, 351.

Buntings, pterygoids of, 110.

Burgena transducta, 203.

Butalis manillensis, 326, 351.

Butaster indicus, 313, 350.

Butler, A. G., on the butterflies of

Malacca, 533.

Butorides javanica, 349, 354.

Butterflies of Malacca, 533.

Buzzards, structure face of, 136.

Cacatua hematuropygia, 312, 350.

Cacomantis merulinus, 320, 351.

Calappa hepatica (ftnote), 238.

Calleptiles latipennis, 559 ; Walkeri,

589.

Callidula abisara, n. sp., Butler, 558.

Calliope camschatkensis, 352.

Callipleea diocletianus, 535, 564; Le-

dereri, 536, 564; vestigiata, 535,

564.

Calobates melanops, 352.

Calcenas nicobarica, 308, 353.

Calornis chalybeus, 343, 353; pana-

yensis, 343, 353.

Camiguin, birds of, 309.

Campephagide, 323.

Cancer lunaris, 244; plicatus, 240;

victor, 243.

INDEX.

Canis familiaris, 254; lupus, 254.

Caphyra rotundifrons (ftnote), 238.

Capitella capitata, 507.

Caprimulgus europzeus, structure face

of, 113; double vomer of, 125.

griseatus, 351; manillensis, 351.

Caraphractus, 588.

Carcinas meenas, zoéa of, 47.

Cardinalide, pterygoids of, 110.

Cardisoma guanhumi, 46; sp. (San

Jago), develop. of, 46.

Cariama, structure face of, 129.

Carinella annulata, 78, linearis, 73.

Carpophaga senea, 346, 353 ; Picker-

ingii, 311, 353.

Carter, H. J., remarks on Dendro-

spongia Steeru, 231.

Castalius elna, 566; ethion, 547,

566; rosimon, 546, 566; roxus,

547, 566.

Castnia, charac. by Boisduval, 166 ; of

Westwood, 167.

, Monograph of, 155; acreoi-

des (Pl. XXIX.), 193; actino-

phorus, 193; actor, 169; albofas-

ciata, 184, 185; amycus?, 181,

182; ardalus, 171; argus, 179;

atymnius, 172; Besckei, 174; Bois-

duvalii, Becker, MS., 173; Bre-

courti, 171; cacica (Pl. XX VIII.),

169; chelone, 180; chremes (Pl.

XXIX.), 185; ciela, Botsduval,

MS.,179 ; clitarcha (Pl. XXXT.),

176; cochrus (Pl. XXYVIII.),

177; columbina, 190; cononia

(PI. XXXIT.), 188; cratina, 188;

cronida, 183 ; cronis, 183; cycna,

189, 191; Dalmannu (Pl. XXX.),

171, 179 ; daedalus, 167; decus-

sata, 175; dionea, 177; diva

(Pl. XXIX.), 187; ecuadoria (PI.

XXXII.), 189; eudesmia (Pl.

XXVIII.), 177; euphrosyne, 181;

evalthe, 181; Fabricii, 178, 185;

Fonscolombei, 178; futilis, 173 ;

galinthias, 172; geron, 168; Go-

dartii, 169, 175; Grayii, 171;

harmodius, 172; hecate, Botsduval,

MS., 174; hegemon, 171, 178,

179; Herrickii, Boisduval, MS.,

174; hesperiaris, 204; Hubner

(Pl. XXVITI.), 182; Humboldti,

180; icarus, 170; var. invaria,

170; inca, Boisduval, MS., 174;

SECOND SERIES.—ZOOLOGY, VOL. I.

597

inornata, 184; imvaria, 170;

japyx, 178 ; juturna (Pl. XXIX.),

182, 183; Kirstenii, 178; Langs-

dorfi, 170; Latreillii (PI.

XXVIII.), 161, 168; licas, 172,

173; licoides, 184; linus (Pl.

XXIX.), 192; lorze, Boisdu-

val, MS., 183; Marcelserresii

(Pl. XXIX.), 186; maris, 178;

meditrina, 182; Menetriesi, 180;

mimica (Pl. XXXIT.), 191; mor-

phoides, 185, 186 ; musarum, Bois-

duval, MS., 177; mygdon (Pl.

XXVIII.), 179, 180 ; orestes, 174 ;

palatina, 176; Pallasia, 171; pa-

pagaya (Pl, XXX.), 170; papili-

onaris (Pl. XXXI.), 175; para-

doxa, 163; paradoxa, Bozscduval,

MS., 176; penelope, 184; per-

sonata, 183; phalaris (Pl. XXX.),

177, 180; procra, 169; pylades,

169; rutila, 178; salasia, 181;

salvina (Pl. XXXIT.), 190; sa-

trapes (Pl. XXXI.), 175, 176;

Schreibersii, 168; simulans, 187 ;

Spixu, 172 ; Sternbergii, 182 ; stri-

gata, 174, 175; subvaria, 177;

thais, 185; thalaira, 185; the-

rapon (Pl. XXTX.), 176; tricolor,

187; truxilla, 162, 190 ; umbrata,

174; unifasciata, 194; veraguana

(Pls. XXVIII. & XXX.), 168;

Viryi, 181; zagrea, 189; zeryn-

thia, 170.

Castniz and allied genera, 164; ar-

rangement veins of hind wings,

162, 163.

as related to the Heliconiide,

158.

, Classifications of, 165, 166 ;

geographical range of, 164; Ma-

dame Merians and others, obser-

vations on transformations of, 164;

palpi of, 164; remarks on venation

wings of, 158, 161, 162, 163, 164;

transformations of, 164.

Casyapa irava, 553, 567 ; thrax, 553,

567.

Catapecilma, n. gen., 547; elegans,

548, 566.

Catopsilia catilla, 551, 567; chryseis,

561, 567; crocale, 550, 567;

scylla, 551, 567.

Cecropis daurica, 351.

598

Celeomorphe (Woodpeckers), rank as

a zoological group, 20; structure,

face of, 2.

Centrobia, 590.

Centrococeyx viridis, 322, 351.

Centropine, 322.

Centrostomum ocellatum (Pl. XIX.),

88, 97; punctatum, 88, 97.

Ceratiocaris, 28.

Ceretes chremes, 185.

charac. by Boisduval, 166.

Cerianthide, 302.

Cerianthus bathymetricus, sp. n. (Pl.

XLY.), 302; membranaceus, 303.

Cethosia, 157; hypsina, 543, 565;

methypsea, n. sp., Butler, 543.

Ceyx melanura, 317, 351 ; philippen-

sis, 851; tridactyla, 351.

Cheetostricha, 590; dimidiata, 589.

Chalaraspide, 45.

Chalaraspis, n. gen., type of new

schizopod family (Pl. VIIL.), 37;

anatomy of, 38; characters, 39;

range of, 37; systematic position

of, 39.

unguiculata, n. sp., 40; mode

of life and colour, 40.

Chalcophaps indica, 348, 353.

Chalcosia coliadoides, var. latifasciata,

559.

Chalcosiide, 558.

Challenger Exped. Actinaria, 295 ;

Atlantic Crustacea of, 23; Deep-sea

Ascidians, 287.

Chameleo vulgaris, sacral nerves of

(Pl. LXVI.), 514.

Charadriide, 349.

Charadrius fulyus, 349, 353.

pluvialis, skull compared with

Eurynorhynchus, 214.

Charaxes hebe, 539, 564; jalysus,

539, 564; Schreiberi, 539, 564.

Chauliodus, W. Indian fish, 50.

Chelidon urbica, stylo-hyal arches of,

106; postoral arches of, 105.

Chelonians, comparisons palate of,

with Birds, 130.

Cheluria terebrans, 67.

Chick’s Skull, researches on, 101.

Chone Duneri, 509; infunduliformis

(Pl. LXY.), 508.

Chrysococeyx basalis, 320, 351.

Chrysocolaptes erythrocephalus, sp. n.

(Pl. XLVI.), 315, 350; maculi-

INDEX.

ceps, sp. n., 314, 350; xanthoce-

phalus, 314, 350.

Chrysocolaptes hematribon, 350; lu-

cidus, 350.

Cicéntide, 349.

Cidaride, on certain organs of, 569.

Cidaris nutrix, 571; tribuloides (Pl.

LXX.), 571; of W. Indies, 50.

Cinnyris speratus, 341, 353; jugu-

laris, 341, 353.

Circus melanoleucus, 350; erugi-

nosus, 308; spilonotus, 308, 350.

Cirratulus cirratus, 507. ;

Cirrochroa bajadeta, 543, 565; jo-

hannes, 543, 565; orissa, 543,

565; rotundata, n. sp., Butler,

543, 565.

Cistenides granulata, 507; hyper-

borea, 507.

Cisticola, sp.?, 336, 352; semirufa,

352,

Cittocinela Iuzoniensis, 352; nigra,

sp. n. (Pl. LIL.), 335, 352.

Clarke, R., and Mivart, St. G., on

sacral plexus and sacral vertebra,

Lizards and Vertebrata (Pl. LXYVI.

& LYII.), 513.

Cleosiris catamita, 558.

Clerome faunula, 564; gracilis, 564.

Cobalus elia, 554, 567; ciliatus, n.sp.,

Butler, 554, 567.

Coccothraustes, pterygoids of, 110.

Coceygomorphe, face of, 126.

Coelites humilis, Butler, 564.

Collingwood, Dr., on marine Plana-

rians, collected partly by Dr.

Kelaart at Trincomalee, and partly

by Dr. Collingwood in Eastern

Seas (Pls. XVII. to XTX.), 83.

Collocalia troglodytes, 320, 351.

Columbee of Philippines, 346.

Columbide, 347.

Comparison of Trematoda and Hiru-

dinei, 209.

Copsychus mindanensis, 352.

Coraciide, 316.

Corallimorphus, gen. noy., 299 ; pro-

fundus, sp. n. (Pl. XLY.), 300;

rigidus, sp. n., 301.

Corone philippina, 343, 353.

Corvide, 343.

Corvus monedula, stapedials of, 107.

Corybantes nicon, 185.

Corydalla lugubris, 337, 352.

Cosmocoma, 588; =Polynema, 583.

Cossus, 158, 164.

Cotuza drepanoides, 562.

Craniorrhinus leucocephalus, 309,

322, 351; Waldeni, 322, 351.

Crateropus caudatus, 352.

Criniger frater, sp. n., 834, 352.

Crossurus vittatus, 67.

Crustacea, notes on Oxystomatous,

235; on some Atlantic, 23; tabu-

lar view, how often got in ‘ Chal-

lenger’ Exped., 56.

Cryptobranchus, sacral nerves of,

524,

Cryptocnemus Grandidieri, 241;

Holdsworthi, sp.n.(Pl. XX XYIIL.),

241; pentagonus, 241.

Cuculide, 320; on structure and de-

velop. of face in, 112.

Cujo, birds of, 310.

Cyclopides salsala, 555, 568.

Cyclopspitta lunulata, 312, 350.

Cyclosia panthona, 559.

Cynthia erotella, n. sp., Butler, 544,

565.

Cyornis philippensis, sp. n., 325, 351.

Cypselidee, 319.

Cypselus apus, interorbital fenestrze

of, 104.

Cyrestis rahria, 544, 565; nivea,

545, 565.

Cystisoma Neptunus, 24.

Cystisomidz, a new family of Crus-

tacea, 25.

Dactylethra, sacral nerves of, 525.

Damalis egens, 558.

Damias dichroa, 203; elegans, 203 ;

melaxantha, 203 ; varia, 203.

Danaidx, modifications apical seg-

ment of, 360.

Danaine, 534.

Danais crocea, 536, 564; grammica,

536, 564; melaneus, 536, 564;

melanippus, 536, 564; plexippus,

586, 564; septentrionis, 536, 564;

vulgaris, 536, 564.

Dasylophus superciliosus, 321, 351.

Deep-sea Ascidians, 287.

Deidamia crucifera= Willemoesia cru-

cifera, 50; leptodactyla, 49, 50.

Deidamia, n. gen. of Crabs, but a

synonym, 50.

Delias dione, 550, 567; metarete,

n. sp. Butler, 550, 567; ninus,

567.

Dendrobiastes basilanica, sp. n. (PI.

LITI.), 332, 352.

Dendrocygna vagans, 454.

Dendrophila frontalis, 338, 352 ;

cenochlamys, sp. n. (Pl. LIII.),

338, 352.

Dendrospongia Steerii, Muriec, gen. &

sp.n.(Pls. XXXVI. & XXXYVILI.),

228, acerate spined spicula of, 226 ;

affinities of, 229; sections of, 221 ;

history of, 219; microscopic struc-

ture of, 224; Mr. Carter’s remarks

on, 232; scopuline spicula of, 226 ;

sexradiate dermal spicula of, 227 ;

spicular rosettes of, 227; vitreous

fibre of, 225.

Dentalium of W. Indies, 50.

Descriptions of some minute Hyme-

nopterous insects, by Prof. J. O.

Westwood (Pl. LX XIII), 583.

Desmognathous palate, varieties of,

112.

Deudorix domitia, 549, 566; jarbas,

549, 566; petosiris, 549, 566;

pheretima, 549, 566.

Deuterostomata compared with Ar-

cheostomata, 209.

Development face of Yungidx, 17.

—— of Land Crabs (PI. XI.), 46.

Diadematidx, 569.

Diceum dorsale, 340, 352; hema-

tostictum, 339, 352; hypoleucum,

339, 352; papuense, 352; retro-

cinctum, 352; trigonostigma, 340,

352.

Dicholophus cristatus, structure skull

of, 128.

Dicruride, 324.

Dicrurus balicassius, 351; mirabilis,

324, 351.

Digonopora (or moncecious Turbel-

laria) subtribe, 86, 87.

Diprotodont Marsupial Hypsiprym-

nodon, Prof. Owen on, 573.

Discophora menetho, 538, 564 ; tullia,

538, 564.

Dithyrocaris, 28.

Ditrypa arietina, 509; greenlandica,

M‘Intosh, n. sp., 509.

Doleschallia pratipa, 539, 564.

Domestic Swine, Prof. Rolleston’s

views on, 253.

INDEX.

Doran, Alban H. G., on morphology

of mammalian ossicula auditus

(Pls. LVIII.-LXIY.), 371.

Dorocidaris papillata, branchie of,

569; compasses, muscles, and liga-

ments of, 570; genital gland of,

571.

Drepanophorus, 77, 79.

Dryococeyx, Sharpe, n. g., 321.

Harringtoni, sp. n., 321, 351.

Dysporus piscator, 354; sula, 354.

Eagles, on palatine bones of, 137.

Kar-bones, small, of Mammals, A. H.

G. Doran on, 371.

Echinide, 569.

Echinoidea, branchiz of, 569.

Kchinometridze, 569.

Kechinus of W. Indies, 50.

Edwardsia coriacea, sp. n. (Pl. XLY.)

299.

Elanus hypoleucus, 350.

Elasmodes obtusus, n. sp., 88, 93.

Elymnias lutescens, 564; mehida,

538, 564; nigrescens, 537, 564.

Elymniine, 537.

Encyrtus, 588.

Epinephele Hyperantus, as type geni-

tal armature (Pl. LY.), 359.

Ergolis ariadne, 545, 565; merione,

545, 565.

Erycinide, 545,

—— modifications apical segment of

(Pl. LYI.), 359.

, harpagones of, 363.

, tegumen of, 361.

, values genital armature of, in

fam., gen., and species, 364, 365.

Eryon arctiformis, 55.

Eryontide, gen. of fossil crustaceans,

to which Willemeesia, n. gen., deep-

sea, is probably allied, 50.

Erythrea pheenicura, 353.

Hsacus magnirostris, 353.

Estheria, 27.

Kteone arctica, 502;

pusilla, 502.

Euchone analis, 508.

Kudamus? yucce not a Castnia, but

a Hesperian, 205.

Kudamus (Megathymus) yucce, vena-

tion wings of, 160.

Eudrepanis pulcherrima, 341, 353.

Eudynamis mindanensis, 320, 351.

flava, 502;

599

Eulabes javanensis, 353.

Eulabetine, 344.

Kulacura osteria, 541, 565.

Eulalia problema, 502; viridis, 502.

Eulophus, 588.

Eumyias panayensis, sp. n., 326,

351.

Eunoa Crstedi, 500.

Kupalamides ctesiphon, 169. |

Euphausia simplex, 45,

Kuphausiide, 45.

Euphrosyne armadillo, Sars, 500;

borealis, 500 ; foliosa, 500.

Eupolynoé occidentalis, M‘Zntosh,

n. sp. (PI. LXY.), 501.

Kuplectella cucumer, 232.

Kuplea, 157; Bremeri, 535, 564;

chloé, 335, 564 ; Grotei, 535, 564 ;

margarita, 585, 564; Ménétriési,

535, 564; midamus, 535, 564;

Ochsenheimeri, 535, 564; phos-

bus, 535, 564 ; Pinwilli, n. sp. (Pl.

LXIX.), 535, 564.

Kuripus euploeoides, 565.

European Rhopalocera, on male geni-

tal armature of, 357.

Wild Boar, 254.

Eurylemide, 344.

Eurylepta (Pls. XVII.-XIX.) affinis,

87,96; atraviridis, 87, 95; cere-

bralis, 87, 96; dulcis, 87, 96;

fusca, 87, 95; Kelaartii, 87, 92;

purpurea, 87, 96; striata, 87, 97;

undulata, 87,95; violacea, 87, 96;

viridis, 87, 96; zeylanica, 87, 97.

Euryleptide, 87.

Kurynorhynchus pygmeus, 213; com-

parison skeleton of, with Tringa,

217; hyoid arch of, 216; pelvis

and limb of, 216; shoulder-girdle

of, 216; skull of, 213; vertebral

column of, 215 ; viscera of, 216.

Kurystomus orientalis, 316, 351.

Kuschema subrepleta, 559.

Kusemia pales, 204.

Eusthenelais abyssicola, M‘Intosh, n.

sp., 501; hibernica, 501.

Eyarne impar, 501.

Excalfactoria chinensis, 353.

Face, structure of, in Buzzards, 136;

in Caprimulgus europeus, 113; in

Coccygomorphe, 126 ; in Cuculine

Birds, 112; in Dicholophus cris-

4K 2

600

tatus, 128; in Falco tinnunculus,

132; in Giant Goatsucker, 124;

in Harriers, 136; in Kites, 136; in

Podargus, 124; in Scythrops novee-

hollandiz, 120; in Trochilide, 116.

Facial Arches in Passerine types, on

structure and development of, 104.

Falconidx, 313.

Falco severus, 350.

Fern-Owl, structure face of, 113.

Flabelligera affinis, 506.

Fringillide, preoral arches of, 108.

Galathea of W. Indies, 50.

Gallicrex cinerea, 353.

Gallinee of Philippines, 348.

Gallinago megala, 354; scolopacina,

B54.

Gallinula chloropus, 353.

Gallus bankiva, 348, 353.

Gastropacha repanda, 160.

Gavia ridibunda, structure skull of,

138; 1st stage, 142; 2nd stage,

145; 3rd stage 146; 4th stage,

148; 5th stage, 148; 6th stage,

150.

Gazera as charac. by Boisduval, 166.

columbina, 190; personata,

188; simulans, 187; zagrosa, 189.

Gecarcinus lateralis, 46.

Gelasimus annulipes, var. (footnote),

238 ; Gaimardi (footnote), 238.

Genital armature, male, European

Rhopalocera, 357.

Genusa bigutta, 563.

Geograpsus crinipes (footnote), 238.

Geometride, 562; wings of, 160.

Geopelia striata, 353.

Gerydus Horsfieldi, 546, 566; ni-

yalis?, 546, 566; symethus, 546,

566.

Gerygone simplex, 352.

Gigantic Hydroid, 287.

Glareola orientalis, 353.

Glycera capitata, 503; setosa, 503.

Glyphodes amethysta, n. sp., 563.

Gnathophausia, n. gen., 36; gigas,

N. sp., carapace of, 28; characters

of, 37; gnathopods, periopods, and

branchie of, 30; oral apparatus

of, 29; pleon and appendages of, 31.

gracilis, colour and mode of life,

34; systematic position of, 35;

zoéa, 0. sp., structure of, 32, 37.

INDEX.

Goatsucker, structure face of, 124.

Gorsachius melanolophus, 308, 349,

354.

Gouride, 348.

Gracula javanensis, 344.

Gralle of Philippines, 349.

Grammatophora barbata, sacral plexus

of (Pl. LXVI.), 516.

Grapsus cruentatus, 46; terrestrial

habits of, 46; pictus(footnote), 238.

Graucalus striatus, 323, 351; suma-

trenesis, 323, 351.

Green Woodpecker, skull of, 4.

Guimaras, Birds of, 309.

Gull, on structure skull of, 138.

Gymnorhina, palate of, 111 ; tibicen,

stylohyal of, 107.

Halcyon chloris, 319, 351; coro-

mandi, 351; gularis, 317, 351;

hombroni, 318, 351; Lindsayi,

351; pileata, 351; Winchelli, sp. n.

(Pl. XLVIL.), 318, 351.

Haliaetus leucogaster, 350.

Harmothoé imbricata, 500.

Harpactes ardens, 316, 350.

Harpagones, European Rhopalocera,

structure of, 362.

Harriers, structure face of, 136.

Hebomoia glaucippe, 551, 567.

Hecatesia, gen. as characterized and

defined by Westwood, 199; exultans

. (Pl. XXXIIT.), 200; fenestrata,

200; thyridion, 200.

Heliconiide as related to the Cast-

nie, 158.

Helotarsus ecaudatus, mesopterygoid

of (Pl. XXY.). 130.

Hemiphaga poliocephala, 347, 353.

Hepiali, 158.

Hepialus humuli (Pl. XXIX.), 163.

Herodias garzetta, 354; intermedia,

354.

Hespagarista, 203; interjecta, 203.

Hesperia badra, 554, 567; harisa,

554, 567; malayana, 567 ; sophia,

White, 194; vitta, 554, 567.

Hesperide, 553.

» modifications apical seements

of (Pl. LVII.), 360.

’ harpageones of, 363.

—, tegumen of, 362.

, value genital armature of, in

fam., gen., and species, 364, 366.

Hesperide, venation wings of, 160.

Hestia linteata, n. sp. (Pl. LXIX.),

531, 564; lynceus, 536, 564.

Heterocera, 557.

Heterocerous Lepidoptera, venation

wings of, 160.

Hexactinellid group of Spongida,

new genus of, 219.

Hierococcyx pectoralis, 351; stre-

nuus, 351.

Himantopus autumnalis, 353; leu-

cocephalus, 353.

Hind foot, Prof. Owen on its deyelop-

ment in yarious groups, 581.

Hirudinei, on some new or little-

known parasitic, 209.

Hirundinide of Philippines, 328.

Hirundo gutturalis, 352; rustica, 328,

352.

Hornbills, structure face of, 126.

House-Martin, postoral arches of, 105.

Hulodes caranea, 561.

Humming Birds, face of, 116.

Hyalincecia tubicola, 499.

Hybleea constellata, 561.

Hybleeidee, 561.

Hydrochelidon leucopareia, 354.

Hydrophasianus chirurgus, 354.

Hydrozoa, type new order of, 61.

Hyla, sacral nerves of, 525.

Hyloterpe philippensis, 351.

Hymenopterous insects, descriptions

of some minute (Pl. LX XIII.), 583.

Hypobythius, nov. gen., 287; caly-

codes, sp. n. (Pl. XLIV.), 287.

Hypochrysops elegans, synonym, 548.

Hypolimnas bolina, 543; incom-

moda, n. sp., Butler, 543, 565.

Hypolyzna erylus, 549, 566 ; etolus,

549, 566.

Hypotenidia philippensis,

striata, 353 ; torquata, 354.

Hypothymis azurea, 325, 351 ; super-

ciliaris, sp. n., 326, 351.

Hypsa egens, 558; heliconia, 557 ;

subsimilis, 557.

Hypsine, 557.

Hypsipetes philippensis, 335, 352;

rufigularis, sp. n., 335, 352.

Hypsiprymnide, 573.

Hypsiprymnodon moschatus, bones

&e., limbs, 578,579; dentition of,

580; descrip. exterior of, 575; feet

of, generally, 576; habits of, 579;

354 ;

male scrotum and female pouch,

579 ; skull of, described and com-

pared with other marsupials, 577 ;

sole of hind foot of, 574. _

Tanthenas griseigularis, 353.

Ichneumon atomos, 583; punctum,

--583.

Icteridee, pterygoids of, 110.

Ideopsis daos, 536, 564.

Iguana tuberculata, sacral nerves of

(Pl. LXVII.), 517.

Incus of Mammalia (see under Au-

ditory ossicula), 371-497.

Indian Wild Hog, 254.

Internal ear-bones of Mammalia,

’ Mr. Doran on, 371.

Intersacral Nerve, definition of, 514.

Tolaus longinus, 549.

Tora scapularis, 333, 352.

Trena cyanogastra, 352; melanochla-

mys, sp. n. (Pl. LI.), 334, 352;

Tweeddalii, Sharpe, 333, 352.

Trenidz, 333.

Isis of W. Indies, 50.

Ixus goiavier, 352; sinensis, 352.

Jungipicus maculatus, 350.

Junonia asterie, 545, 565; eudoxia,

565; iphita, 545, 565; laomedia,

545, 565; lemonias, 545, 565;

orithya, 545, 565.

Kangaroo, sole of hind foot of, 574.

Kelaart, Dr., Marine Planarians col-

lected by, 83.

Kettelia Lowii, n. gen. & sp., 560.

Ketupa ceylonensis, mesopterygoid of

(Pl. XXY.), 131.

Kites, on structure face of, 136.

Lacerta viridis, sacral nerves of, 515.

Lacertilia, nerves of lumbar, sacral,

and precaudal region of, 514.

Lalage dominica, 324, 351.

Lambrus nodosus (ftnote), 238.

Lampides agnata, 547, 566; almora,

547,566; aluta, 547, 566; beeticus,

547 ; beroé, 547, 566; cerula, 547,

566; kankena, 547, 566; macroph-

thalma, 547, 566; patala, 547,

566; pseudelpis, n. sp., Butler (Pl.

LXVIII.), 547, 566.

TLanassa Nordenskidldi, 508.

Land-Crabs, on development of, 46.

INDEX.

Lanice conchilega, 73.

Lanius lucionensis, 323, 351; nasu-

tus, 351; schach, 351.

Laomedea flexuosa, 65.

Laphania Boecki, 508.

Laridx, 350.

Larus, sp., 354.

Lathromeris, 590.

Laughing Gull, skull of, 140.

Laurion corculum, n. sp., Butler, 559.

Lezena abranchiata, 508.

Lebadea alankara, 541, 565; martha,

541, 565.

Lepidogrammus Cumingi, 351.

Lepidonotus subleyis, 501.

Lepidopterous genus Castnia, J. O.

Westwood on, 155.

Leptochelia, 68.

Leptoplana aurantiaca, Coll., n. sp.

(Pl. XVIII.), 88, 94; patellensis,

n. sp., 88, 93.

Leptoplanide, 88.

Lethe europa, 537, 564.

Leucocerca nigritorquis, 351.

Leucophasia sinapis, venation wings

of, 160.

Leucosia (Pl. XXVIII.) brunnea,

Miers, sp. n., 237; fuscomaculata,

sp. n., 236; neocaledonica, 236;

ornata, sp. n., 236; pallida, 239;

Perryi, sp. n., 238; pubescens, sp.

n., 238; pulcherrima, sp. n., 236;

reticulata, sp. n., 237; urania, 236;

Whitmeei, Miers, 238.

Leucosiide, descriptions new species

of, 235.

Leucotreron gironieri, 353.

Libytheidee, modifications apical seg-

ment of (Pl LVi.), 359.

; harpagones of, 363.

—, tegumen of, 362.

, values genital armature of, in

family, genera, and species, 364,365.

Limenitis procris, 541, 565.

Jimnoria terebrans, 67; xylophaga,70.

Linota cannabina, facial arches of, 108.

Linotrypane apogon, 505.

Liparidee, 560.

Liphyra brassolis, 546, 566.

ithosiide, 557.

Lizards, sacral plexus and sacral ver-

tebre of, 513; nervous and osseous

structures of sacral region in, 522.

Lophogaster typicus, 27.

601

Lophogastride, characters of, 36, 45.

Loriculus chrysonotus, 313, 350;

Hartlaubi, 350; occipitalis, 350 ;

philippensis,350; regulus, 313, 350.

Loxura atymnus, 550, 567.

Lubbock, Sir J., on aquatic Hymeno-

ptera, 583.

Luidia of W. Indies, 50.

Lumbriconereis fragilis, 503.

Lumbricus tubicola, obtained by Sir

John Ross, 499.

Luzon, Birds of, 309.

Lycena cagaya, 548, 566; karsandra,

548; lysizone, 548, 566; sangra

548, 566.

Lycenide, 546.

, modifications apical segment of

(Pl. LVII.), 359.

—, harpagones of, 363.

==F, tegumen of, 361.

, values genital armature of, in

family, genera, and species, 364,365.

Lyceenine, 547.

Lycenopsis haraldus, 546, 566.

Lycorea ceres, 187.

Lyncornis macrotis, 351.

Macdonald, Dr. J. D., anatomy of

Tanais vittatus, occurring with

Limnoria and Chelura terebrans in

excavated pier-wood (PI. XV.), 67.

, on new genus of Trematoda and

some new or little-known parasitic

Hirudinei (Pl. XXXIY.), 209.

M‘Intosh, Dr. W. C., on Annelida

obtained during cruise of H.M.S.

‘Valorous’ to Davis Strait (Pl.

LXY.), 499.

, on Valencinia armandi, a

new Nemertean (Pl. XVI.), 73.

Macroglossa proxima, 557.

Macroglossinz (of Malacca), 557.

Macronus straiticeps, sp.n., 331, 352.

Macropteryx comatus, 319, 351.

Macropygia tenuirostris, 347, 353.

Malacca, Butterflies of, 533.

Malamaui, Birds of, 310.

Maldane Sarsi, 507; triceps, 507.

Male genital armature in European

Rhopalocera, 357.

Malleus of Mammals (see under Au-

ditory ossicula), 371-497.

Mammalian internal ear-bones, A. H.

Doran on, 371.

602

Margarodide, 563.

Marine Leeches, branchie of, 210;

generative processes of, 211.

Planarians of eastern seas, 83.

Marsupialia, earliest forms pentadac-

tyle, 580; Hypsiprymnodon mos-

chatus, Prof. Owen on, 573; modi-

fication hind foot of, 573.

Matuta (Pls. XX X1X.—XL.) Banksii,

245; distinguenda, 247; doryphora,

244; eranulosa,sp.n., 245; Leseurii,

248 ; lineifera, sp. n., 245; lunaris,

247; maculata, sp. n., 246; ob-

tusifrons, sp. n., 247; Peronii, 243 ;

picta, 246; planipes, 246 ; revision

species of, 243; rubro-lineata, sp.

n., 244; victor, Mabr., 243; victrix,

243, victrix, var. crebrepunctata,

244,

Matutidee, historical summary of, 241.

Megalema asiatica, skull of, 122.

Megalurus palustris, 352.

Megapodiide, 348.

Megapodius Cumingi, 353; Lowi,

348, 353.

Megathymus, 205.

Melanitis leda, 537, 564.

Melanopelargus episcopus, 349, 354.

Meropide, 316.

Merops bicolor, 316, 350; philip-

pinus, 316, 350.

Messaras erymanthis, 544, 565.

Microhierax erythrogenys, 350.

Microma, 589.

Micronia astheniata, 563.

Microniide, 563.

Micrura fusca, 73.

Miers, HE. J., notes on Oyxstomatous

Crustacea (Pl. XXXVIII.), 235.

Milionia basalis, 558.

Mindanao, Birds of, 309.

Minyad, a larval (Pl. XLY.), 296.

Minyadineze, 295.

Mivart, St. G., and Clarke, R., on

‘sacral plexus and sacral vertebra

of Lizards and other Vertebrata

(Plates LXVI., LXVII.), 513.

Mixornis Woodi, sp. n., 331, 352.

Mca mirabilis, 73.

Monitor arenarius, sacral neryes of,

520.

Monocaulus, sp. ?, 287.

Monticola solitaria, 335, 352.

Mopsea of W. Indies, 50.

INDEX.

Morphine, 538.

Morphology of mammalian Ossicula

auditus, by A. H. G. Doran, 371.

Moseley, H. N.,on new forms of deep-

sea Ascidians obtained during

voyage H.M.S. ‘Challenger’ (Pl.

LXIYV.), 287.

, on new forms of Actiniaria

dredged in deep-sea; with descrip-

tion of certain pelagic surface-

swimming species (Pl. XLY.), 295.

Motacilla luzonensis, 352.

Motacillide, 337.

Mulleripicus funebris, 350.

Munia atricapilla, 345, 353; leuco-

gastra, 345, 853; jagori, 353;

minuta, 353.

Murie, Dr. J., on Steere’s Sponge, a

new genus of the Hexactinellid

group of the Spongida, 219.

Muscicapide of Philippines, 325.

Musophagide, os uncinatum of, 127.

Mycalesis fusca, 537, 564; hesione,

587, 564; janardana, 537, 564;

mineus, 537, 564; nautilus, 564;

polydecta, 537, 564.

Myina annulipes, 589; livens, 589.

Mymar pulchellus (Pl. LXXIIT.),

584; taprobanicus, Ward, 6 & 2

585; Wollastoni, Westw., n. sp.

(Pl. LXXIT.), 585.

Myra angulata, 239; carinata, 239 ;

elegans, 239; mammillaris (Fl.

XXXVIIL.), 239.

Myrina amrita, 550, 567; anasuja,

567; chitra, 549, 566; freja, 550 ;

lapithis, 549, 567 ; marciana, 549,

566; megistia, 549, 566; tharis,

550, 567; travana, 549, 566.

Myristicivora bicolor, 347, 353.

Myside, 45.

Myzanthe pygmea, 352.

Naidonereis quadricuspida, 504.

Nasal labyrinth of Woodpecker, 5, 9.

Nautactis purpureus, n.sp. (P1.XLY.),

295.

Nebalia, sp.?, from Bermudas, 26;

longipes, n. sp. (Pl. VI.), anat. of,

26; characters of, 27.

Nebaliide, 45.

Nectariniide, 339.

Negros, Birds of, 309.

Nemeobiine, 545.

Nemertean, on a new, 73.

Nemertes Neesii, 510.

Nemotois scabriosella, venation wings

of, 160.

Nephropsis Stewarti, 50.

Nephthys cxea, 501; ciliata, 501.

Neptis (Pls. LXVIII.—LX1X.) colu-

mella, 542, 565; dindinga, n. sp.,

542, 565; dorelia, n. sp., 542, 565;

gononata, n. sp., 541, 5€5; hor-

donia, 542, 565; leuconata, n. sp.,

541, 565; mamaja, n. sp., d41,

565; nata, 541, 565; paraka, n.

sp., 542, 565; tiga, 542, 565;

vikasi, 541, 565.

Neptunus sanguinolentus(ftnote), 238.

Neochera marmorea, 558.

Nereis pelagica, 503; zonata, 503.

Nettapus coromandelianus, 354.

Nicolea arctica, 508 ; zostericola, 508.

Nicomache lumbricalis, 507.

Ninox philippensis, 350.

Nothria conchylega, 503.

Notocirrus tricolor, 503.

Numenius pheopus, 304.

Nursia abbreviata, 240; Hardwick,

240 ; plicata? (Pl. XXVIIT.), 240;

sinuata, sp. n., 239.

Nychia Amondseni, 500; cirrosa, 500.

Nyctalemon docile, n. sp., Butler, 562.

Nyctemera coleta, 558; tripunctaria,

558.

Nyctemeride, 558.

Nycticorax griseus, 354; manillensis,

354.

Nyetipao crepuscularis, 561.

Nymphalide, 534.

——, modifications apical segment of

(Pl. LY.), 360.

: harpagones of, 363.

—-, tegumen of, 362.

, values genital armature of, in

family, genera, and species, 364, 366.

Nymphalinz, 539.

Oceanactis, gen. nov. (Pl. XLY.), 296;

rhodactylus, sp. n., 296.

Octacnemus, gen. noy. (Pl. XLIY.),

287 ; bythius, sp. n., 289.

Ocypoda hippeus, 46; rhombea, 46.

Oligosita, 590; collina, 589; nodi-

cornis, Westw., n.sp. (Pl. LX XIIZ.),

592; Staniforthii, n. sp.,591; sub-

fasciata, n. sp., 091.

Ommatophoride, 561.

On Atlantic Crustacea from ‘ Chal-

lenger’ Expedition, by Dr. R. v.

Willemoes-Suhm, 23.

On certain organs of the Cidaride,

by Charles Stewart, 569.

On Hypsiprymnodon, Ramsay, a new

genus indicative of a distinct family

(Pleopodidee) in the Diprotodont

section of the Marsupialia, by

Prof. Owen (Pls. LXXJ-LXXII.),

573.

On morphology of skull in Wood-

peckers (Picide) and Wrynecks

(Yungidee), by W. K. Parker, 1.

Onychoprion anesthetus, 354.

Ophelia limacina, 505.

Ophideres fullonia, 561.

Ophideridee, 561.

Ophiomusium of W. Indies, 50.

Ophioneurus signatus, 589, 590.

Orgyia turbata, n. sp., Butler, 560.

Oriolide, 328.

Oriolus chinensis, 328, 352; philip-

pensis, 352; Steerii, Sharpe, 329,

352; suluensis, Sharpe, 329, 352.

Ornithoptera ruficollis, n. sp., Butler,

552, 567.

Orthia, gen. as characterized by Bois-

duyal, 166, and Boisdv. pars, 193.

augias, 167, 201; nexa, 201;

pelargus, 194.

, 167, untenable as a genus, see

Othria, Westw., 201.

Orthotomus castaneiceps, 337, 352 ;

cinereiceps, 337, 352; derbianus,

352; frontalis, 336,352; ruficeps,

337, 352.

Ortygometra cinerea, 308, 349, 353.

Os uncinatum in Passeres, 110; in

Sun-birds, 111.

Osmotreron axillaris, 346, 353 ; ver-

nans, 346, 353.

Ossicula auditus, on the morphology

of the mammalian, 371.

Osteology of the Spoon-billed Sand-

piper, on the, 213.

Othria, gen. as characterized by

Westwood, 201.

amalthea, 202; amazonica, 202;

augias (P]. XXIX.), 201; colum-

bina, 202; eucadorina, 202; Lin-

digii, 202; nexa, 201.

Owen, Prof., on Hypsiprymnodon mos-

INDEX.

chatus, 573; development of hind

foot in vertebrates, 581, 582.

Owenia filiformis, 507.

Owls, structure skull of, 138.

Oxycephalus piscator, 25.

Oxycerca jagori, 353.

Oxystomatous Crustacea, EH. J. Miers

on (Pls. XXXVITI-XL.), 235.

, literature of subject, 235.

Pachyrhamphus, vomerine cartilages

of, 110.

Padda oryzivora, 308, 345.

Palate of Hemilophus fulvus, 13 ;

of Eagles, 137; Picus analis, 13; P.

major, 14; P. minor, 15; P. viridis,

7; Picumnus minutus, 16; of Vul-

tures, 137.

Palate, summary of characters of the

Saurognathous type of, 19; varie-

ties of Desmognathous, 112.

Palawan, Birds of, 310.

Pamphila aria, 554, 568; augias,

554, 568 ; meesoides, nu. sp., Butler,

554, 568; maro, 555, 568; mat-

thias, 554, 568; nigrolimbata,

555, 568; pilanus, 554, 568.

Panay, Birds of, 309.

Pandita sinope, 542, 565.

Papilio achates, 553, 567 ; agamem-

non, 552, 567; amycus, 182 ;

antiphates, 552, 567; archeus,

156; arruana, 156; arycles, 552,

567; axion, 552, 567; bathycles,

552,567 ; Boisduvalii, 156; brama,

567; chremes, 185; clytia, 552;

cochrus, 177; croesus, 156; cronis,

183; cronius, 156; cyparissias,

168; dardanus, 181; Delessertii,

552, 567; demolion, 552, 567;

diphilus, 553, 567; dedalus, 168 ;

erebus, 567; Esperi, n. sp., Butler

(Pl. LXVIII.), 553, 567; evalthe,

181; evemon, 552, 567; harmo-

dius, 172; helenus, 553, 567;

icarus, 170; iswara, 553, 567 ;

licus, 173; linus, 172; Lydias,

156; malayanus, 552, 567; mestor,

553, 567; oceanus, 156; palati-

nus, 176; panthous, 156; paris

and local varieties of, 156; pega-

sus, 156; pelasgus, 194; phalaris,

177; pirrha, 172; polytes, 552,

567; poseidon, 156; prexaspes,

603

558, 567; priamus, 156, and local

var. of, 156; pronomus, 156 ;

pylades, 170; rama, 567; rich-

mondia, 156; sarpedon, 552, 567 ;

saturnus, 553, 567; syphax, 172;

thais, 185; triton, 156; Urvil-

liana, 156; yvaruna, 553, 567 ;

zagreeus, 189.

Papilionide, 550.

, modifications apical segment of

(Pl. LVI.), 359.

harpagones of, 363.

tegumen of, 361.

——, value genital armature of, in

fam., gen., and species, 364, 365.

Papilionine (subfam.), 552.

Paradisea papuana, palate of, 111.

Paramphinome pulchella, 502.

Parasa bandura, 561.

Parasitic Hirudinei, 209.

Paride, 338.

Parker, W. K., on Morphology of Skull

in Woodpeckers and Wrynecks, 1.

, on the Structure and De-

velopment of the Bird’s skull

(Plates XX.—XXVIT.), 99.

Parthenos lilacinus, n. sp., Butler,

544, 565.

Parus amabilis, sp. n. (Pl. LIJIT),

338, 352; elegans, 338, 352.

Passer montanus, 353.

Passeres of Philippines, 323.

Passerine types, on structure and

development of facial arches in, 104.

Patagonia gigas, face of, 116.

Pedicellariz of Cidaride, 571.

Pelargopsis gigantea, 311, 317, 351;

Gouldi, 351; leucocephala,317,351.

Pelecanoides urinatrix, interorb. fe-

nestre of, 104.

Pelecanus philippensis, 354; roseus,

354.

Penang fauna, to be accepted with

modifications, 534.

Penelopides manille, 351; panini,

309, 323, 351.

Pericrocotus cinereus, 351; igneus,

324, 351.

Petalophthalmus, n. gen., a Mysidi-

form Schizopod, 40; characters of

the genus, 43 ; systematic position

of, 43, 44; armiger, n.sp. (Pl. VII.),

40, 41, 44; anatomy of, 40; habitat

of, 40 ; habits and colour of, 44.

604

Phabotreron amethystina, 353; leu-

cotis, 309, 353; nigrorum, sp. 2.,

346.

Phacocheerus, 264.

Phagonia, 588.

Phalacrocorax, sp. ?, 354.

Phalangista, sole hind foot of, 574.

Phasianide, 348.

Philentoma albiventris, sp. n., 325,

351; cyaniceps, 309, 351.

Philippine Archipelago, on birds col-

lected by Prof. J. B. Steere in, 307.

, bird families of, 311.

— , distribution birds of, 308.

, table genera and spec. birds

of, 350.

Phlogcenas luzonica, 348, 353.

Pholoé minuta, 501.

Phyllacanthus baculosa, 571.

Phyllodoce greenlandica, 502.

Phyllornis palawanensis, sp. n.

(Plate L.), 333, 352.

Phyllornithide, 333.

Phylloscopus lugubris, 336, 352;

magnirostris, 352.

Picarie of Philippines, 314.

Picide, 314.

Picus analis, palatal structure of, 13 ;

major, 14; minor, 15; viridis,

development of skull, 4.

Pieride, modifications apical segments

of (Pl. LVI.), 359.

——, harpagones of, 363.

—— ——.,, tegumen of, 361.

——,, values genital armature of, in

fam., gen., and species, 364, 365.

Pierinee (subfam.), 550.

Pieris cronis, 183; cronissa, 183.

Pig, on the domestic, of prehistoric

times in Britain, 251.

, lachrymal bone, distinctively

characteristic, 259, 263.

, nasals characteristic of, 263.

——,, naso-frontal suture of, 262.

, Skull, measurements of lachry-

mals in different species, 269.

, third lower molar distinctive

between Sus scrofa and S. cristatus,

262, 278, 284.

, value as a domestic animal, 258.

Pigs, Aithiopian species of, 257 ;

African do., 256; Asiatic varieties

of, 256, 261; Bornean young, oc-

casionally striped, 280; Chinese

INDEX.

young, never striped, 280; Irish,

260; of Asia, 254.

Pig’s skull, difficulty of distinguishing

European and Indian kinds, Dr.

Gray on, 255.

Pipra auricapilla, vomerine cartilage

of, 110.

Pitta Rochi, Bruggemann, 309.

Pittide, 329.

Planaria, application of term, 85.

Planarians, Mr. Collmgwood on ma-

rine species of, 83.

Planocera thesia (Pl. XIX.), 88, 98.

Planoceride, 88.

Planularia of W. Indies, 50.

Plastingia callineura, 555, 568.

Platalea leucorodia, 217.

Pleopodide, n. fam., Owen, 573.

Plesioneura alysos, 556, 568; as-

mara, 556, 568; dan, 556, 568;

folus, 556, 568; Pinwilli, n. sp.,

Butler (Pl. LXVIIL.), 556, 568.

Plexus and sacral vertebree of Lizards

and Vertebrates, 513.

Ploceidx, 345.

Plotus melanogaster, 354.

Plyctolophide, 312.

Podargus, on structure of face in, 124,

Podiceps philippensis, 354.

Poliolophus urostictus, 334, 352.

Polychrus marmoratus, sacral nerves

of, 522.

Polynema fuscipes, 583 ; natans, 583 ;

natatorial habits of, 584.

Pontobdella, black species of (Pl.

XXXIYV.), 34.

Poreus madagascariensis, 255.

Poritia sumatree, 546, 566.

Poropoea, 589, 590.

Porphyrio pulverulentus, 353.

Porzana fusca, 353 ; pygmea, 353.

Potamocheerus africanus, 255.

Pourretia coarctata, 164.

Pratincola caprata, 335, 352.

Prato, branchie of, 211.

Praxilla arctica,507; praetermissa,507.

Prehistoric times, on the domestic pig

of, 251.

Presacral nerves defined, 514.

Prestwichia aquatica, Lubbock, 587.

Prioniturus discurus, 312, 350.

Prionochilus, sp. ?, 340, 352.

Prionospio Steenstrupi, 507.

Procerus Busku, Coll., n. sp. (Pl.

XVII.), 87, 91; concinnus, Coll.,

n. sp. (Pl. XVII.), 87, 90; Han-

cockanus, Coll., n. sp., 87, 91.

Prometheus casmilus, 178.

Prothoe caledonia, 564,

Pseudolalage melanoleuca, 351.

Psittaci of Philippines, 312.

Psittacide, 312.

Pteroptrix, 589.

Pterylosis of Spoon-billed Sandpiper,

217, 218.

Ptilocichla, n. gen., 332.

falcata, n. sp. (P1.I1.), 332, 353.

Ptilocarpa griseopectus, 353.

Puffinus leucomelas, 309, 354.

Pyrrhocentor melanops, 322, 351.

Querquedula multicolor, 354.

Rallide, 349.

Rallina fasciata, 354.

Ramphastos toco, skull of, 127.

Rana pipiens, stapedials skull of, 106.

, sacral nerves of, 525,

Remipes pacificus ({tnote), 238.

Rhabdornis mystacalis, 353.

Rhamphiculus occipitalis, 353.

Rhinopalpa fulva, 545, 566.

Rhipidura nigritorquis, 325, 351.

Rhopalocera, 535 ; on the male geni-

tal armature in European, 357.

Rhyacophilus glareola, 354.

Rhyncheea capensis, 304.

Rhynchites betule, 590.

Rodine Loveni, 507.

Rolleston, G., on domestic Pig of pre-

historic times in Britai, and on

the mutual relations of this variety

of Pig and Sus scrofa, var. ferus,

S. cristatus, S. andamensis, and S.

barbatus (Pls. XLI.—XLIII.), 251.

Rollulus, sp. ?, 353.

Rothia, n. gen., Westw., charac. of,

204; agrius, 204; eriopis, 204;

pales, 204; pedasus, 204; simyra,

204.

Sabella crassicornis, 508; pavonia, .

508; spetbergensis, 508.

Sacral plexus in Lizards and Verte-

brates, 513.

region, nervous and osseous

structures of, in Batrachians, 523 ;

in Birds, 526; in Mammals, 525;

in Lizards, 522.

Sacral vertebre in birds defined, 530 ;

in Lizards and Vertebrates, 513;

in Mammals, Reptiles, and Batra-

chians, 526.

Salamandrina maculosa, sacral nerves

of, 524.

Salpinx leucogonis, n. sp., Butler,

536, 564.

Samytha sexcirrata, 508.

Sandpiper, Spoon-billed, skeleton of,

213; pterylosis of, 213.

Sarcophanops Steerii, sp.n. (Pl. LIV.),

344, 353.

Sareops calvus, 344, 353; Lowii,

sp. n., 344, 353.

Saturniide:, 561.

Satyride, modifications apical seg-

ment of (Pl. LVI.), 360.

harpagones of, 363.

tegumen of, 362.

, value of genital armature of, in

fam., gen., and species, 364, 366.

Satyrinz, 537.

Saurognathse (Woodpeckers), deve-

lopment face of, 2.

Saurognathous type of palate, sum-

mary of characters of, 19.

Sealibregma inflatum, 506.

Scardia mediella, venation wings of,

160.

Scelostrix candida, 350.

Schizopods, comparison deep-sea and

shallow-water forms, 45; on some

genera of, with a free dorsal shield,

28; synopsis characters of, 44.

Sclerothamnus Clausii, Marshall, =

Dendrospongia Steerii, Wurie, 231.

Scolecolepis cirrata, 506 ; ? Jeffreysii,

MIntsh., n. sp. (Pl. LXY.), 506.

Seoloplos armiger, 504.

Scops megalotis, 350.

Scudder, Mr. §., Syst. Revis. Amer.

Butterflies, remarks on, by Dr. F.

B. White (ftnote), 365.

Seythrops novee-hollandiz, on struc-

ture of face in, 120.

Secusio mundipicta, 558.

Serpula ?. sp. of, 509.

Sesarma obesum (ftnote), 238 ; quad-

ratum (ftnote), 238,

Sesia, 164.

Setaria ruficauda, sp. n., 327, 351.

Sharpe, R. Bowdler, on the birds

eollected by Professor J. B. Steere

INDEX.

in the Philippine Archipelago (Pls.

XLVI.-LIYV.), 307.

Siphonium of Crow’s and Crocodile’s

skull, 106.

Sipunculus of W. Indies, 50.

Sittide, 338.

Skull of Birds, on the structure of, 99.

Accipiter nisus, 35; in Aetomor-

phe, 128; Dicholophus cristatus,

128; Falco tinnunculus, 132.

Gavia ridibunda, Ist stage, 142 ;

2nd stage, 145; 3rd stage, 146 ;

4th stage, 148; 5th stage, 148 ;

6th stage, 150.

Megalema asiatica, 122 ; Spar-

row-Hawk, 135; Strigide, 138;

Woodpeckers and Wrynecks, 1.

Sphenodon, stapedial processes of, 106.

Sphingide of Malacca, 557.

Sphyngiceps, n. g., Coll., 86; lacteus,

Coll. (Pl. XVII.), 87, 90.

Spilornis holospilus, 313, 350.

Spilosoma, n. sp., 557.

Spirorbis borealis, 509 ;

509 ; verruca, 509.

Spizaétus philippensis, 350.

Sponges, mode of fishing for, in Phi-

lippines (ftnote), 220.

Spongide, on a new geuns of Hexac-

tinellid group of, 219.

Squatarola helvetica, 353.

Syuilla armata (ftnote), 238.

Stapes of Mammalia and Birds (see

under Auditory ossicula), 371-497.

Steatornis caripensis, structure face

ef, 123, 125.

Steere, Prof. J. B., on birds collected

by, in the Philippines, 307.

Steere’s Sponge, on, 219.

Sterna Bergii, 350.

Sternine, 350.

Stephanoseyphus, general remarks on,

64, 65; homology, 64; mirabilis,

n. sp. (Pl. XIY.), 61, 65.

Stewart, Charles, on certain organs of

the Cidaridse, 569.

Strigidee, on skull of, 138.

Strix stridula, mesopterygoid of,

130.

Struthionide, feet of, as compared

with some Marsupials, 580.

Sturnia violacea, 308, 343, 353.

Sturnids:, 343.

Stylochidee, 88.

spirillum,

SECOND SERIES.—ZOOLOGY, VOL. I.

605

Stylochoplana elegans (Pl. XIX.),

88, 98; meleagrina, 88, 98.

Stylochopsis malayensis, Coll., n. sp.

(Pl. XVIIT.), 88, 94.

Suidz, bronze statues of, 278, 279;

Etruscan, 278 ; odontograms teeth

of, 278 ; Prof. Riitimeyer on, 279;

Schutz, Herr J. W., on, 280; skull

measurements ‘of, 275, 276, 281,

282, 283, 284, 285, 286 ; twist in

tail of, 254.

Sulu Islands, birds of, 310.

Surniculus velutinus, sp. n., 320,351.

Sus andamanensis, 254, 262, 280,

283; barbatus, 254, 265, 272, 285,

286; barbatus, peculiarities skull

of, 265; barbirussa, 262; capensis,

257; celebensis, 253, 267, 274;

ceylonensis, 266; cristatus, 253,

254, 255, 256, 269, 284, 285;

europeus, 255; indicus, 253; lar-

vatus, 255; leucomystax, 253 ;

papuensis, 254; scrofa, var.

wgypti, 257; scrofa, var. ceramica,

271; scrofa, var. domesticus, 281;

scrofa, var. ferus, 252, 270, 281 ;

scrofa, var. palustris, 253; senna-

ariensis, 256, 277; sinensis, 256;

tawanus, 264; ternatensis, 276;

timorensis, 254; verrucosus, 254,

266, 367; vittatus, 253, 267:

zeylonensis, 266.

Swine, bibliography on, 272, 273;

parasites of, as elucidating spe-

cific distinctions, 268; relation-

ship of domestic to wild, views on,

252 species from Lofoten Islands,

Sylvicolidee, pterygoids of, 110.

Sylviide, 336.

Sympalamides mimon, 180.

Sympheedra dirtea, 540, 565; emalea,

565.

Synemon, gen. as used by Westwood,

194; catocaloides (Pl. XXXIIT.),

195, 196 ; directa, 197; gratiosa,

198 ; hesperioides, 198 ; icaria,

196; leta, 195; mopsa, 196;

nais, 195; notha, 197; nupta,

197; obscurella, 197; parthenoides,

196; plana, 196,197; selene, 195,

Sophia, 194; Sternbergii, 182;

theresa, 195; vagans, 198.

Syntomis basifera, 557.

606

Tachytrypane Jefireysi, M‘Intosh, n.

sp. (Pl. LXV.), 505.

Tenia in Swine, 268; solium, 268 ;

mediocannellata, 268.

Tagiades calligana, n. sp., Butler (Pl.

LXIX.), 556, 568; gana, 556, 568;

layata, n. sp., Butler, 557, 568;

rayi, 556, 568; trichoneura, 568.

Tanaécia aruna, 540, 565; pulasara,

540, 565.

Tanagride, pterygoids of, 110.

Tanaid, on a blind deep-sea, 23.

Tanais vittatus, anat. of (Pl. XV.),67;

cavolini, 67; dubius?, 68; habits

of, 70.

Tanygnathus luzoniensis, 312, 350.

Tascina, gen. as defined by West-

wood, 199; orientalis, 199.

Tegumen of European Rhopalocera,

structure of, 360.

Teius teguexin, sacral plexus of, 516.

Telphusa fluviatilis, carries young,

and development with metamor-

phosis, 48.

Terebellides Stroemi, 508.

Terias formosa, 550, 567; hecabe-

cides, 550; imanata, 550, 567 ;

pumilaris, 550, 567; sari, 550,

567; senna, 567.

Terinos robertsia, 544, 565; teu-

thras, 544, 565.

Thaumantis noureddin, 538, 564;

pseudaliris, 538, 564.

Thaumops pellucida, a synonym, 25.

Theclinee (subfam.), 548.

Thecomedusx, a new order of Hydro-

z0a, 65.

Thelephus circinnatus, 508.

Thermesiidx, 562.

Thriponax javensis, 314, 350.

Thysanozoon Alderi, Coll., n. sp., 87,

88; Allmanni, Coll.,n.sp., 87, 89 ;

auropunctatum, 87, 94.

Tiga javanensis, 315, 350.

Timeliide, 331.

Tinea (footnote), 157.

Torfschwein, 254, 279, 282.

Tortoise, sacral nerves of, 523.

Totanus calidris, 309, 354;

cens, 304.

Trayisia Forbesii, 506; glandulosa,

M‘Intosh, n. sp. (P1. LXY.), 506.

canes-

INDEX.

Trematoda, new genus of, 209; n. sp.

of, described, 210.

Treron nasica, 396, 343.

Treronide, 346.

Trichobranchus glacialis, 508.

Trichogramma, 588, 590; erosicornis

(Aprobosca), n. sp. (Pl. LXXIIT.),

592; evanescens, 588 ; vitripennis,

589; Walkeri, 589.

Trichogramini, genera of, 590.

Tringa minuta compared with Ku-

rynorhynchus, 213, 217 ; ruficollis,

354; salina compared with Hury-

norhynchus, 214, 217.

Tringoides hypoleucus, 354.

Trochilide, structure face of, 116.

Troglodytes vulgaris, vomer of, 109.

Trogonide, 316.

Trophonia plumosa, 506.

Tube from Globigerina-ooze, Green- |

land seas, 510.

Turbellaria dendroccela, 85, 86, 87.

Turdide, 335.

Turdus chrysolaus,351; obscurus, 351.

Turnix ocellata, 353.

Turtur Dussumieri, 347, 353; humi-

lis, 347, 353.

Tympanic ring of Gorilla, 377.

Typhlolepta Byerleyana, Coll., n. sp.

(Pl. XVIL.), 88, 92.

Typhloleptide, 88.

Urania, 157.

Uraniide, 562.

Urapteride, 562.

Urapteryx marginipennis, n.

Butler, 562.

Urbanus licus, 184.

Sp.,

Valencinia Armandi, M‘Intosh, n. sp.

(Pl. XVI.), 73; cutaneous system

of, 73; digestive system of, 78 ;

muscles of, 76; nervous system of,

78; lineiformis, 73.

¢Valorous,’ Annelida

eruise of (1875), 499.

Vertebrata, sacral plexus of, 513.

Volvocivora czrulescens, 351.

Vomers of Prcus viridis, 4.

Vultures, the palatine bones of, 137.

obtained in

Westwood, Prof. J. O., descriptions

of some minute Hymenopterous in-

sects, 583.

Westwood, Prof. J. O., monograph of

the lepidopterous genus Castnia

and some allied groups (Pls.

XXVIII-XXXIIT.), 155.

White, Dr. F. Buchanan, on the

male genital armature in the Eu-

ropean Rhopalocera (Pls. LY.—

LYII.), 357.

Wild Boar, from what stock derived,

253.

Willemoesia, noy. gen.(Pls. X.—XII1.),

50; crucifera, n. sp., anatomy of,

52, characters of, 54, colour and

habits of life, 53 ; leptodactyla,n. sp.

(Pl. XIII.), 50, characters of, 54,

Willemoes-Suhm, Dr. R., on some

Atlantic Crustacea from the ‘ Chal-

lenger’ Expedition, 23.

Woodpeckers, on skull of, 1.

Wren, facial structure of, 109.

Wrynecks, morphology skull of, 1.

Xantholema hemacephala, 351;

rosea, dol,

Xantholestes, gen. nov., Sharpe,

327; panayensis, sp. u., 327,

351.

Xanthopygia narcissina, 327, 351.

Xanthotenia busiris, 538, 564.

Ypthima corticaria, nu. sp., Buéler,

537, 564; methora, 537, 564;

philomela, 537, 564.

Yunx torquilla, face of, 17; palate

of, 17.

Zanclopteryx saponaria, 563.

Zebu, Birds of, 309.

Zemeros albipunctata, 545, 566;

emesoides, 545, 566.

Zeocephus cinnamomeus, sp. n. (Pl.

XLVIII.), 352, 328; cyanescens,

sp. n., 328; rufus, 308, 352.

Zereniide, 563.

Zeuzera, 164.

Zeuxidia amethystus, 538, 564.

Zoéa-stages of Crabs, observations on,

46, 47.

Zosterops Meyeni, 352.

Zygenide, 557.

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