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HARVARD UNIVERSITY.
LIBRARY
OF THK
MUSEUM OF COMPARATIVE ZOOLOGY.
\.l\t>Niij-rvCj<Lj
M.^J^A'iA^CiX —
\^ci\— VSjoioaXA^-^ ■
""■ ~ 100 4
THE
TRANSACTIONS
OP
THE LINNEAN SOCIETY
or
LONDON.
SECOND SERIES— VOLUME A^III.
ZOOLOGY.
Vx\\\^M*///
LONDON:
PlilNTED BY TAYLOR A>-D FEAMCIS. E£D LION COURT, FLEET STIiEET.
SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON HOUSE ;
AND BY LONGMANS, GKEEN, AND CO., PATERNOSTER-HOW.
1900-190:;
CONTENTS.
PAUT I.— September, 1900.
I. On a Collection of Braclvyura from Torres Straits. By W. T. Calman, B.Sc,
TJniversity College, Dundee. {Communicated by Prof. D'Arcv W. Thompsox,
C.B., F.L.S.) (Plates 1-3.) pages 1-50
PART II.— September, 1900.
II. Report 0)1 a Collection made hy Messrs. F. V. McCoxnell aiul J. J. Quelch at
Mount Boraima in British Guiana. [Communicated hy Prof E. Ray Laxkester,
I>.C.L.,F.R.S., Director of the Natural Sistory Museum.) (Plates 4-6.) . 51-76
PART III.— December, 1900.
III. On the Structure and Affinities of Echiurus unicinctus. By Alice L. Embleton,
B.Sc. [Communicated hy Prof. G. B. Howes, Sec.L.S.) (Plates 7-10.) . 77-97
PART IV.— May, 1901.
IV. The Terrestrial Isopoda of Neio Zealand. By Charles Chilton, M.A., D.Sc.
(N.Z.) ; F.L.S. London; Pesearch Felloio, Zliiversity of Fdintmryh. (Plates
11-16.) 99-152
PART v.— August, 1901.
V. Etude d^une Espece nouvelle de Lepadides (Scalpellum giganteum, n. sp.) et de
Poecilasma carinatum, Hoek. Par Monsieur A. Gruvel, Charge d'un Cours de
Zoologie a la Faculte des Sciences de Bordeau.r. (Communique par M. le
Professeur Howes, Sec. Linn. Soc.) (Plate 17.) 153-161
PART VI.— September, 1901.
VI. On the Affinities of zEluropus melauoleiicus, xl, Milne-Edwards. By E. Ray
Lankester, 3L.A., LL.D., F.B.S., F.L.S., Director of the Natural History
Depaj-tments of the British Museum. (Plates 18-20.) 163-172
[ iv ]
PART VII.— OcTOBEK, 1901.
A^'II. On the Intestinal Tract of Birds ; with Remarks on the Valuation and Nomen-
clature of Zoological Characters. Brj P. Chalmers Mitchell, M.A., D.Sc.
Oxon., F.L.S., F.Z.S., Lecturer on Biology at the London Hospital Medical
College. (Plates 21-23.) pages 173-275
PART VIII.— June, 1902.
VIII. Sur qii-elques Lepadides nouveaux de la Collection du British Iluseum. Par
A. Gruvel, Maitre de Conferences a la Faculte des Sciences, TJniversite de
Bordeaux. {Communique par 31. le Prof. G. B. Howes, Sec. Linn. Soc.)
(Plate 24 aud 5 Illustrations in the Text ) 277-295
PART IX.— October, 1902.
IX. On the Actinian Bunodeopsis globulifera, Vei^rill. By J. E. Duerden, Ph.I>.>
A.E.C.Sc. (Bond.), Bruce Fellow, Johns Hopkins University. {Communicated by
Prof. G. B. Howes, F.B.S., Sec. Linn. Soc.) (Plates 25 & 26.) . . . 297-317
PART X.— February, 1903.
X. On the Mophology of the Brain in the Mammalia, with Special Beference to that
of the Lemurs, Recent and Bxtinct. By G. Elliot Smith, M.D. {Sydney),
Fellow of St. John's College, Cambridge, and Professor of Anatomy, Egyptian
Government School of Medicine, Cairo. {Communicated by Prof. G. B. Howes,
D.Sc, LL.I)., F.B.S., Sec. Linn. Soc.) (With 66 Illustrations in the Text.)
319-132
PART XL— February, 1903.
XL On the New Zealand Pity II abranchiate Crustacea-Macrnra. By George M.
Thomson, F.L.S. (Plates 27 29.) 133-453
PART XII.— July, 1903.
XII. On the Morphology of the Cerebral Commissures in the Vertebrata, with Special
Beference to an Aberrant Commissure found in the Forebrain of certain Beptiles.
By G. Elliot Smith, M.A., M.D., Fellow of St. John's College, Cambridge ;
Professo J- of Anatomy, Egyptian Government School of Medicine, Cairo. {Com-
municated by Prof G. B. Howes, D.Sc, LL.D., F.B.S., Sec. Linn. Soc)
(With 36 Illustrations in the Text.) 455-500
PART X III.— December, 1903.
Titlepage, Contents, and Index 501-517
UWv'^ \ V \1 U I
VV1
2nd Ser. ZOOLOGY.
[VOL. VIII. PART 1.
THE
TRANSACTIONS
OP
THE LINNEAN SOCIETY OF LONDON.
ON A COLLECTION OF BRACHYUBA FfiOM TORRES STRAITS.
BY
W. T. CALM AN, D.Sc,
CNIVEESITY COLLEGE, DUNDEE.
(Communicated by Prof. D'Arcy W. Thompso.v, C.B., F.L.S.)
^
LONDON:
PRINTED FOR THE LINNEAN SOCIETY
BY TAYI.OK ANU FRANCIS, RKI> I.ION COURT. FLEET STREET.
SOLD A'V THE SOCIETY'S APARTMENTS, BURLINGTON-HOtlSE, PICCADILLY, W.,
AND BY LONGMANS, GREEN, AND CO., PATERNOSTER-ROW.
September 1900
TRANSACTIONS
OF
THE L I N N E A N SOCIETY.
I. On a Collection of Brachytira from Torres Straits. By W. T. Calman, D.Sc,
University College, Dundee. {Communicated by Prof. D'Akcy W. Thompson, C.B.,
F.L.S.)
(Plates 1-3.)
Eead IGth November, 1899.
Introductory.
I HIS paper deals with the Brachyurous Crustacea collected hy Prof. A. C. Haddon
during his first expedition to Torres Straits in 1888. The collection comprises about
87 species, three of which are described as new. Owing probal:)ly to the fact that
attention was given to collecting the smaller and less conspicuous forms, a number of
interesting and little-known species were obtained, some of which I have redescribed and
figured. From the same cause, however, the determination of many of the specimens
proved to be a matter of no little difficulty, and the identification of one or two of the
more obscure species is at best pro^dsional.
Of the three species here described as new, Cryptociiemus Iladdonl belongs to a genus
hitherto comprising only four species, all of which are known only from single
specimens. Pilummis cristipes is apparently very distinct from any known species, and
its title to inclusion in the extensive genus Pilumiius may perhaps be disputed.
Lamltrus confragosus belongs to a genus already overburdened with species, but, so far
as the means at my disposal enable mu to judge, it seems to deserve tlie distinction of a
specific name.
SECOND SERIES. — ZOOLOGY, VOL. VIII. 1
DR. W. T. CALMAN ON A COLLECTION OF
Among the species already known, concerning which I am able to furnish fresh details,
the most interesting is the parasitic Hapalocarcinus marsupialis. Briefly described forty
years ago by Stimpson, it appears to have escaped re-examination by carcinologists,
though the curious gall-like growths to which it gives rise on corals are well known.
The occiu'rence of the three known Inuo-Pacific species of Palmis [Cymopolia), two
of which have been recorded hitherto only from widely distant localities, has afforded an
opportunity for a detailed examination of their distinctive characters.
In the case of certain species described by Adams and White and by Miers, I have
been able, from an examination of the type specimens in the British Museum (Natural
History), to sui^plement the original accounts and to make one or two alterations in the
matter of nomenclature.
1 am much indebted to Prof. P. Jeffrey Bell and to Mr. R. I. Pocock for their
courtesy and kindness in facilitating my work among the collections under their charge.
My thanks are also due to Prof. E. L. Bouvier and Dr. J. G. de Man for giving me
information and advice on various points, and to Dr. Chas. Chilton for the loan of
literature relating to Australasian Crustacea*.
As regards nomenclature and classification, I have followed, where jjossible, the
important work of Dr. A. Alcock, ' Materials for a Carcinological Pauna of India,' now
in course of publication. The extensive collections upon which his revision of the
Indian species is based, and the exhaustive way in which the bibliography of the subject
is treated, render this work indispensable for the student of the Indo-Paciiic Crustacea.
List of the Species.
Tribe CYCLOMETOPA.
Family Xanthid^.
*CarpiHus conve.rus (Forsk.).
Carpilodes sp.
Liomera cinctimana (White).
*Atercjatis floridus (L.).
*Lophact(Ba granulosa (Riipp.).
* Lophozozymus octodentatus (M.-E.).
dodone (Herbst).
*Xantho [Leptodius) exaratus (M.-E.).
*Etisus Icevimanus, Rand.
*Etisodes frontalis, Daua.
* electra (Herbst).
*Actcea Riippellii, Krauss.
* calculosa (M.-E.).
granulata, var. carcharias, White.
Peronii, var. squamosa, Ilend. (?).
* hystrix, Miers.
Xanthodes Lanwrckn (M.-E.).
*Chlorodius niger (Forsk.).
*Phymodius ungulatus (M.-E.).
sculptus (A. M.-E.).
Chhrodopsis melanodactylus, A. M.-E.
spinipes, A. M.-E.
Cymo Andreossyi (Aud.).
melanodactylus, De Haan.
*Ozius guttatus (M.-E.).
Pseudozius dispar, Dana.
* Pilumnus cursor, A. M.-E.
* pulcher, Miers.
* seminudus, Miers.
* lanatus (Latr. ?), Miers.
longicornis, Hilg., var.
• cristipes, n. sp.
* Actnmnus sctifer (De Haan).
Trapezia f err uginea, var. areolata, Dana.
* cyinodoco (Herbst).
Tetralia glaherrima (Herbst).
* Since this paper was read I have been enabled, by the kindness of Mr. T. R. R. 8tebbing, to consult Paulson's
little-known work on the Crustacea of the Red Sea. Prof. Walker, of this College, has been good enough to translate
for me several passages to which reference will be made.
BEAOHIUEA FROM TOKRES STKAITS.
Family Portunid^.
Lissocarcinus orbicularis, Dana.
Caphyra rotundifrons, A. ]\I.-E.
*Nepttmus sanguinolentus (Herbst).
* jjelagicus (L.).
* {Acheluus) granidatiis (M.-E.).
* ( ) , var. unispinosus, Miers.
*Thalamita prymna (Herbst).
* adinete (Herbst).
* sima, M.-E.
Family CANCRiDiE.
*Kraussia nitida, Stimpson.
Tribe CATOMETOPA.
Family Grapsid^.
*Met.opograpsus tnessor (Forsk.).
*Varuna litter atu (Fabr.).
Family Ocypodid*.
*Ocypoda ceratophthalma (Pallas).
Uca ( = Gelasimvs) tetragonon (Herbst) .
Ceratoplax sji.
Tribe OXYSTOMA.
Family Calappid^e.
*Calappa hepatica (L.).
Family Leucosiid^e.
Cryptocnemus Haddoni, n. sp.
* Oreophorus frontalis, Miers.
*Myra fugax ( Fabr. ) .
* australis, llasw. (?).
*Leuvosia longifruns, var. pulcherrima, Miers.
* Haswe/Ii, Miers.
I'seudophilyra trideiitata, Miers.
Arcania gracVipes, Bell (?).
Family Dorippid^ (?).
*Falicus Jiikmii (White).
IVkitei (Miers).
serripes ( Aleock & Anderson) .
Tribe OXYRHYNCHA.
Family Maiid^.
*Ach(Bus affinis, Miers.
*Paratymolus sexspinosus, Miers.
*Oncinopus aranea, De Haan.
*Xenocarcvmis tuber ml alvs, White.
*Huenia proteus, De Haan.
*MencBt/m(s it/oiioceros, Latr.
Hyastenvs spinosus, A. M.-E.
* oryx, A. M.-E.
* coHvexus, Miers.
verrucosipes (Adams & White) .
Broc/di, De Man.
*Naxia serpulifera (Guer.).
* taunts, Pocock.
Tylocarcinus sty.r (Herbst).
*Paramit/>rax [Chlorinoides) Coppingeri, Hasw.
* ( ) aculeatus (M.-E.).
*Schizophrys aspera (M.-E.).
Cyclax suborbicularis (Stimpson) .
* Pseudomicippa vai'ians, Miers.
*Micippa philyra (Herbst).
'^Tiarinia ungusia, Dana.
Family Parthenopid.e. r„„j, 9
*Lambrus [Aulacolambrus) koplonotus, Ad. &Wh..,
* [Parthenolambrus) calappoides, Ad. & Wh.
( ) confragosus, n. sp.
Incertae sedis.
Family Hapalocakcinid^ (nom. nov.).
Hapalocarcimis nuirsupialis, Stimpson.
In the above list I have marked with an asterisk those species which have been
ah-cady recorded from the North or North-east of Australia. Of the species not so
recorded the majority are known to have a wide distribution within the Indo-Pacific
region, and the few^ cases where the range of a species is considerably extended
(e. g., Palio/is Whitci) merely help to emphasize the uniformity of the crustacean fauna,
whose range is coterminovxs with that of the coral-reefs over this extensive area.
4 BR. W. T. CALMAN ON A COLLECTION OF
Carpilitjs convkxus (Forsk.).
Carpilius convexus, H. Milne-Edwards, Hist. Nat. Crust, i. p. 382, pi. xvi. figs. 9-10; Alcock,
Jouru. Asiatic Soc. Bengal, Ixvii. (2) p. 80 (1898).
Three females and two males. One labelled " Found within hollow in Tubipora."
.Locality. " Mer."
Carpilodes sp.
A number of very small specimens belonging to this genus do not agree satisfactorily
with any of the described species, but in the absence of a larger series for comparison I
do not venture to describe them as new. They are identical with certain small specimens
in the British Museum determined as C. rugatus, Latr., but differ from larger specimens
of that species and from the figure given by A. Milne-Edwards (N. Arch. Mus. Paris, i.
pi. xii. figs. 3, 3 a-h) in the fact that the lobulations on the surface of the carapace,
and especially on the branchial regions, are more numerous and do not have the
regularly transverse direction so marked in C. rugatus.
The C. cariosm of Alcock (Journ. Asiatic Soc. Bengal, Ixvii. (2) 1898, p. 86, and
Illustr. Zool. ' Investigator,' Crust, pl. xxxvi. fig. 7, 1899) * resembles our specimens
rather closely ; but in that species the carapace is not quite so broad, the antero-lateral
borders less convex, and the lobules on the lateral regions of the carapace are arranged
in a slightly different manner.
Locality. "Murray Island, 15-30 fms."
LlOMEEA CINCTIMAKA (White).
Carpilius cinctimanus , White, in Jukes's Voy. 'Fly,' ii. Appendix, p. 336, pl. ii. fig. 3 (1847) ; Adams
& White, Voy. ' Samarang,' Crust, p. 37, pl. vii. fig. 4.
Liomera lata, Dana, U.S. Expl. Exp., Crust, i. p. 161, pl. vii. figs. Qa-d.
L. cindimana, Uana, op. cit. p. 161.
L. lata and L. clnctimana, A. Milne-Edwards, N. Arch. Mus. Paris, i. pp. 219-220 (1865).
L. cinctimana, A. Milne-Edwards, N. Arch. Mus. Paris, ix. p. 176, pl. v. fig. 4 (1873).
Carpilodes cinctimanus, Miers, Ann. & Mag, Nat. Hist. (5) v. p. 234 (1880) ; Henderson, Tr. Linn.
Soc, (2) Zool. V. p. 354 (1893).
L. cinctimana, Ortmann, Zool. Jahrb. Syst. vii, p. 450 (1893) ; Alcock, Journ. Asiatic Soc.
Bengal, Ixvii. (2) p. 88 (1898).
Two male and two female specimens.
Locality. " Murray Island."
Distr'ihutioii. Zanzibar to Tahiti.
The synonymy of this species and of the genus of which it is the type give striking
evidence of the intangible nature of the characters on which we are forced to rely in the
classification of the Xanthoid crabs. As defined by Dana, Liomera is stated to resemble
Carjiilius in the disposition of the antennae (that is to say, the basal anteunal joint enters
into the inner orbital hiatus), and the genus is separated from Carinlodes only by the
fact that the fingers are said to be sharp-pointed instead of spoon-shaped at the tip.
* Cy. also Nobili, Aun. Mus. Gcuova, (2) xx. p. I'oG (1S99X
BEACHTUEA TEOJI TOREES STEAITS. 5
A. Milne-Edwards in his monograph of the Cancridae {I. c. 1865) abandoned the use of
the character drawn from the shape of the finger-tips as a generic distinction, but he
retained the genus Liomera, placing it in the group of genera having the l)asal antennal
joint " s'unissant seulement au front par son angle interne," and separating it thus from
Carpilodes by a character in direct contradiction to the origmal definition of the genus.
Miers in 1880 {I. c.) referred our species to Carpilodes, accepting that genus in Milne-
Edwards's sense as having the basal joint of the antenna " produced along the exterior
margin of the infero-lateral frontal process so as to enter partly witliin the interior
orbital hiatus" (Chall. Rep., Brachyura, p. 133, 1886). Altliough thus deprived of its
type species, the genus Lioniem was retained by Miers, who defines it (Rep. Voy.
' Alert,' Crust, p. 528) very much as Milne-Edwards liad done, and refers to it certain
species wliich he later included (Chall. Hep. p. 125) in the genvis Xantho. Ortmann
and Alcock retu.rn to Milne-Edwards's position, including this species in Liomem and
defining it as having the basal antennal joint not entering the orbital hiatus, and the
first-named author gives a figure of L. ctiictinuina to illustrate this very point. It will
thus be seen that of the authors who have examined this species Dana and Miers regard
the basal antennal joint as entering the inner orbital hiatus, while Milne-Edwards,
Ortmann, and Alcock state explicitly that it does not. As a matter of fact, Liomera
occupies in this respect an intermediate position between two extremes, which are
connected by a continuous series of gradations. On the one hand, we have forms where
the basal joint lies nearly longitudinally and meets the posterior process of the front at
its tip, so that the short line of junction between the two is transverse to the axis of the
joint ; on the other liand, we may have the basal joint lying very obliquely to the axis of
the body, meeting the frontal 2>i'ocess with its inner edge, so that the prolonged line of
junction is approximately parallel to the axis of the joint, which thus lies more or less
comjiletely in the hiatus between the frontal process and the low^er wall of the orbit. In
the present species, however, the short trapezoidal basal joint meets the frontal process
at its tip, but the short line of junction between the tw'o is obliquely placed w'itli
reference to the long axis of the joint, so that a small portion of the latter may be
regarded as lying in the orbital hiatus between the frontal process and the suborbital
■wall. Moreover, some individual variation in this resjject is observed when a series of
specimens is examined, and, if we may judge by the analogous case of Actcea calciilosa
referred to below, it would seem that this character is liable to change with the growth
of the individual.
Atergatis flouidus (L.).
Atergatis foridus, Alcock, Jouru. Asiatic Soc. Beuji-al, Ixvii. (2) p. 98 (1898).
Two small specimens of this widely distributed and common Indo-Pacific species
The carapace of the larger is 10 mm. long and IG mm. broad, the relative breadth being
somewhat greater than in larger specimens.
Loc((Uty. " Murray Island, reef."
6 DE. W. T. CALMAN ON A COLLECTION OF
LOPHACT^A GRANULOSA (Riipp.).
Cancer limbatus, H. Milne-Edwards, Hist. Nat. Crust, i. p. 377, pL xvi. figs. 1-3.
LofhacUm gramilosa, A. Milne-Edwards, N. Arch. Mus. Paris, i. p. 247; Alcock, Journ. Asiatic Soc.
Bengal, kvii. (2) p. 101 (1898).
A female specimen.
Locality. " Torres Straits."
LOPHOZOZTMTJS OCTODENTATUS (Milne-Edwards).
Xantho octodentatus, H. Milne-Edwards, Hist. Nat. Crust, i. p. 398.
Lo})hozo:ymus octodentatus, Haswell, Cat. Austr. Crust, p. 58.
L. epheliticus (L.), Miers, Crust. Voy. ' Alert/ p. 207 ; De Man, Zool. Jalirb. Syst. viii. p. 518.
L. octodentatus, Alcock, Journ. Asiatic Soc. Bengal, Ixvii. (2) p. 106 (1898).
Three male and three female specimens. The largest, a male, has the carapace
75 mm. hroad, and in it, as in the other smaller specimens of both sexes, the chelse are
about equal in size. The difference between the sexes in the prominence of the last two
pairs of antero-lateral teeth noted by De Man (/. c. p. 519) does not hold good for our
specimens.
Locality. " Tringing reef, Mabuiag."
LoPHOzozYMus DODONB (Hcrbst).
Lophozozymus dodone (Hbst.), De Man, Arch. Naturg. liii. (1) p. 270, pi. x. figs. 2, 2 « (1887) ;
Alcock, Journ. Asiatic Soc. Bengal, Ixvii. (2) p. 108 (1898).
Two female specimens, about 13 mm. broad, agreeing perfectly with the figures and
short description given by De Man. The grooves on the outer and inner surfaces of the
dactylus of the chelipeds are very broad, and the upper margin between them is reduced
to a thin sharp crest. De Man notes that these grooves are deeper in young individuals.
Locality. " Channels between reefs, Mer."
Distribution. Mozambique to Tahiti.
Xantho (Leptodius) exaratus (Milne-Edwards).
Chtorodius exaratus, H. Milue-Edwards, Hist. Nat. Crust, i. p. 402.
Lejjtodius exaratus, A. Milue-Edwards, N. Arch. Mus. Paris, ix. p.222 (1873) ; De Man, Arch. Naturg.
lii. (1) p. 285 (1887).
Xantho (Leptodius) exaratus, Alcock, Journ. Asiatic Soc. Bengal, Ixvii. (2) p. 118 (1898).
A single small female specimen of this common and variable species. It appears to
differ from the typical form, as described by the authors quoted above, only in the fact
that the wrist and hand of the chelipeds are rugose above and the outer face of the hand
is granulated.
Locality. " Cockburn group (N. Queensland), shore."
BRACHYUEA FROM TORRES STRAITS. 7
Etistjs LJ3VIMANUS, Rand.
Etisus lavimanus, Rand., Dana, U.S. Expl. Exp., Crust, i. p. 185^ pi. x. fig. 1 ; Alcock, Journ. Asiatic
Soc. Bengal, Ixvii. (2) p. 131 (1898).
A single small specimen (17'5 mm. long) agreeing in general shape and proportions
of carapace with the still smaller specimens figured by Dana (/. e. fig. 1, /), the breadth-
ratio of the carapace being the same (1-46) as that given by him. In full-o-rown
specimens the carapace is much more transverse, the breadth-ratio being about 1-6.
Locality. " Fringing reef, Mabuiag."
Etisodes frontalis, Dana.
Etisodes frontalis, Dana, U.S. Expl. Exp., Crust, i. p. 187, pi. ix. fig. 3; Dc Man, Notes Leyden Mus.
xii. p. 8, pi. 1. fig. 2.
Our two specimens (about 11-5 mm. long by 17-25 mm. broad) agree more closely with
De Man's figure than with Dana's, which is taken froma-very small specimen only 8 mm.
broad. The slight emargination of the frontal lobes is even less marked than in De Man's
figure, these lobes being nearly transversely truncate. The posterior teeth of the antero-
lateral margin are rather less spiniform, and some other slight differences are to be
gathered from De Man's detailed description.
Locality. " Fringing reef, Mabuiag."
Etisodes electra (Herbst).
Etisodes sculptilis, Heller, SB. Ak. Wien, xliii. p. 333 (18G1) ; A. iMilne-Edwards, N. Arch. Mus.
Paris, ix. p. 236, pi. ix. fig. 2 (1873).
E. electra (Ilbst.), De Man, Arch. Naturg. hii. (1) p. 290 (1887).
Two specimens, agreeing well with Heller's description and Milne-Edwards's figure.
The larger, a female, measiu'cs 7 mm. in length by 10 mm. in breadth (breadth- ratio l'-J<2),
and is thus somewhat broader than is indicated by either of the authors named, whose
measurements give the breadth-ratio as 1'36 and 1'38 respectively. The smaller specimen
is a male 4 mm. long by 5' 25 mm. broad, giving a ratio of about ISl.
Localitj/. " Murray Island, reef."
AcT^A RiJPPELLii, Krauss.
Actcea Ruppellii (Kr.J, De Man, Zool. Jahrb. Syst. viii. p. 499 (1895) ; Alcock, Journ. Asiatic Soc.
Bengal, Ixvii. (2) p. 144 (1898).
Our specimens agree in most points with the description of this species as given by
De Man {I. c. and earlier papers there referred to). They show some variation in the
hairiness of the body and in the prominence of the regional divisions of the carapace.
One specimen, considerably larger than the others, differs from them in the greater
length of the hairs on the carapace and in the scantiness of the short dow^n which in the
other specimens covers the carapace below the long hairs. In this specimen tlie finger-
tips are slightly excavate. All our specimens can be matched from the series referred
8 DK. W. T. CALMAN ON A COLLECTION OF
to A. Buppellu in the British Museum collections, but iu none of the latter are the hairs
quite so long as in our largest individual.
Localities. '• Murray Island " ; " Thursday Island, fringing reef and shore " ; "Albany
Passage, 10 fath."
ACT^A CALCITLOSA (Milne-Edwards).
Cancer calculosvs, H. Milne-Edwards, Hist. Nat. Crust, i. p. 378 (1834).
Actma calculosa, A. Milne-Edwards, N. Arcli. Mus. Paris, i. p. 276, pi. xviii. fig. 3 (1865) ; Haswell,
Cat. Aiistr. Crust, p. 45 (1882).
Euxanthus tuberculosus, Miers, Crust. ' Alert,' p. 205, p. xix. fig. A (1884).
AcUea calculosa, Henderson, Tr. Linn. Soc. London, (2) Zool. v. p. 356 (1893) ; Alcock, Journ. Asiatic
Soc. Bengal, Ixvii. (2) p. 152 (1898).
Pive specimens of this species are in the collection. Prof. E. L. Bouvier has very
kindly taken the trouble to compare one of our specimens, sent to him for the purpose,
with the original type specimen of H. Milne-Edwards's Cancer calculosus in the Paris
Museum. He writes as follows : —
" J'ai compare minutieusement votre Crabe avec le type 6.'Actcea calculosa, Edw. II
appartient evidcmment a la meme espece. Les tubercules du test y sont beaucoup plus
saillants, mais ils sont partout disposes dans le meme ordre et sont entoures aussi
d'une aureole peripherique de ponctuations je vous le repete, on ne saurait douter
de I'identification."
The identification of our specimens being vouched for on such excellent authority, it
may be usefiil to give in some detail their characters as compared with Aetata granulata
(And.), since it appears that recent writers have not always successfully distinguished
the two species.
The breadth-ratio of the carapace varies from 1-33 to 1-4 without apparent relation to
the actual size. The three posterior lobes of the antero-lateral margin are prominent
and well-defined, and in front of these the first lobe is represented by a single tubercle.
In Actma ffranulata * the lobes are low% rounded, and indistinctly separated. The regions
of the carapace are much more distinct in A. calculosa, being separated by rather deep
grooves, which are in part smooth and free from granules. There is a marked and
generally smooth groove parallel to the hinder margin of the carajoace and separated
from it by two or, in one case, three rows of granules. The posterior margin is defined
at each end by a small spiniform tubercle which is not distinct in A. (jranulata. The
cardiac area is of a rather different shape from that shown in the figure given by
A. Milne-Edwards, being produced and much narrowed anteriorly. The tubercles on the
surface of the carapace are rounded, smooth, and surrounded each by radiating puucta-
tions. On the posterior part of the carapace in some specimens they become depressed
and confluent. In A. granulata the tubercles are more pointed and the radiating
* I have used lor comparison two specimens of A. <jranulata from Japan in the Museum of University CoUege.
These diifer slightly from Savigny's figure (l)escr. de I'Egypte, Crust, pi. vi. fig. 2) in the rougher appearance of the
carapace, but they iseem to agree in all essential points with this as well as with the descriptions of later writers.
BEACHTUEA EEOM TORRES STRAITS. 9
punctations mark off more or less prominent accessory tubercles which surround the base
of each large tubercle. The frontal lobes in A. ealculosa are smooth or indistinctly
granulated on the edge, while in A.yranulata they are edged with pointed granules. In
A. ealculosa the outer surface of the hands bears smooth, bluntly conical tubercles
arranged in longitudinal rows ; in ^. cjranulata the tubercles are lower, surrounded bv
prominent and pointed accessory tubercles, and the arrangement in roAAs is less reo-ular.
The walking-legs are covered with tubercles rounded at the tip, smooth, and larger in
size than in A.grannlata, where they are low, pointed, and more numerous. The merus
joint of the last pair of legs is strongly serrate above, the serrations increasing in height
towards the distal end, and its posterior face is nearly smooth. In A. granulata the
seiTations of the upper edge are small and irregular, and the posterior surface is covered
with granulations. The sternum and abdomen, especially in the male, are smooth, with
scattered pimctatious, while in ^. (jraunlata the same regions are usually much granulated,
at least posteriorly.
In his report on the Crustacea collected by H.M.S. ' Alert,' Mr. Miers has described
and hgured, imder the name of Enxanthns tuherculosm, a species which I believe, after
examination of his type specimens, to be identical with the present. Mr. Miers writes,
" As the basal antennal joint enters well within the inner orbital hiatus, this species must,
I think, be referred to the genus Enxanthns^ In his specific description the account of
the basal antennal joint is qiialified by the words " in the adult,'" and certainly the
difference in this respect between the largest and the smallest of the specimens he was
describing is conspicuous enough to have suggested a doubt as to the validity of a generic
distinction resting on this point alone. As a matter of fact it is easy to find specimens
both of the present species and of A. granulata in which the l)asal antennal joint enters
quite as far into the orbital hiatus as in any but the largest of Miers's specimens *. The
individual which he figures, and from which his description is mainly drawn, is a large
male, 23 mm. in length. The carapace is rather wider than in smaller specimens, the
breadth-ratio being about l"i3, and the tuberculation of the carapace is very strongly
developed. The smaller specimens associated with this by Mr. Miers, and obtained bv
the ' Alert ' in the vicinity of Torres Straits, are all but identical with the specimens in
the present collection from the same locality. Mr. Miers further states that the smaller
specimens "have much the aspect of certain Actceo!, e. g., A. f/raitnlata (Aud.) and
A. carcharhts, White ; from both of which species they may be distinguished upon the
most superficial examination by the smoothness of the sternum and post-abdomen." In
spite of the emphasis of the last sentence, I find in the British Museum collections
specimens determined by Mr. Miers as^. granulata (among others the one referred to in
his ' Challenger ' Report, p. 120) which resemble in every respect the smaller types of his
" Euxanthus tuberculosm." The few specimens referred to A. ealculosa in the British
Museum collection are rather different in appearance from our Torres Straits specimens,
* Paulson (whose work ilr. Miers had not .seen) had already established a new genus Eu.vanth,»/cs for the
reception of Actn'Ci granulata ou account of the structure of its antennal region, which he figures (■ (!ruritacea of the
Red Sea " (Russian), Kiev, 187;', p. 33, pi. vi. figs. 3 & 3 a).
SECOND SERIES. — ZOOLOGY, VOL. VIII. 2
10 DR. W. T. CALMAN ON A COLLECTION OF
the tubercles on the carapace being very niucb dejiressed, confluent, and smooth. I
believe, however, that they must be referred to the same species.
Localities. " Thursday Island " ; " S. of Orman's reef, 5-7 fath." ; " Channels between
reefs, Mabuiag " ; " Channels between reefs, Mer and Dara " (5 specimens).
ACT.a;A GRANULATA, var. CARCHARIAS (White).
Actaa carcharias, White, Proc. Zool. Soc. London, 1847, p. 224 ; A. Milne-Edwards, N. Arcli. Mus.
Paris, i. p. 276 (1865).
A single female specimen, perhaps sterile, the abdomen appearing unusually narrow.
It agrees perfectly with White's type specimen in the British Museum, save that the
under surface is rather smoother. I have no doubt that Miers is right in regarding this
as merely a variety of A. gramdata (Chall. Eep., Brachyura, p. 122). The Japanese
specimens of A. gramdata referred to above show a tendency towards this variety in the
rougher aspect of the carapace as compared with Savigny's figure.
Locality. " S. of Orman's reef."
Distribution. Swan Eiver, W. Australia ( White).
AcT^A Peronii, var. squamosa, Henderson (?).
Act<sa Peronii, var. squamosa, Henderson, Tr. Linn. Soc. London, (2) Zool. v. p. 357.
A male specimen, 10 mm. in length and 14 mm. in breadth. It differs much in
appearance from the few specimens of A. Peronii (all of smaller size) available for
comparison, but it agrees closely with the short description given by Prof. Henderson
of his var. squamosa. The tubercles of the antero-lateral margin are replaced by
sharj^ spines, of which there are five on each side behind the external orbital tooth.
In the typical A. Peronii there are only four tubercles on the antero-lateral margin.
LocalUy. " Torres Straits."
Distribution. India {Henderson).
AcTiEA HTSTRix, Micrs.
Acteea hystrix, Miers, Kep. ' Challenger ' Brachyura, p. 121, pi. xi. fig. 3.
Pour female specimens having the following dimensions : —
Breadth-ratio.
1*22
1-3
1-36
1-43
The measurements given by Miers are : —
6-0 mm. 8-0 mm. 1-33
Our smaller specimens agree very closely with the single type specimen. The
measurements given above show a regular increase in the relative breadth of the cara-
pace with increasing age. In the largest specimen the antero-lateral borders are more
curved, the front is narrowei-, and the carapace departs from the hexagonal outline which
it presents in the smaller specimens as in Miers's figure. The surface of the carapace is
BRACHTUEA FEOM TORRES STRAITS. 11-
raore closely granulated tlian is shown in tlie figure, the granules heing of different sizes,,
smooth and rounded on the posterior part of the carapace, and becoming spiniform in
front and at the sides. Tlie spinules on the haud are more thickly set and those on the
legs are longer than in Miers's figure.
Localitij. " Torres Straits."
With the above I may associate provisionally two specimens which I cannot identify
with certainty. The smaller (length 5 mm., breadth 7 mm.) resembles somewhat closely
the specimens of A. Injstrix, difTeriug chiefly in the Ijlunter armature of the I)ody and
legs. The granules of the carapace are less closely packed and are rounded, not spiniform
in any part ; on the chelipeds they are conical, and on the legs they have the form of
bluntly truncated spines. In the larger specimen (length 9 mm., breadth 12-5 mm.) the
granules of the carajmce are still more depressed and smoother, and the tubercles on the
limbs are less prominent. These specimens differ from Actica iiodnlosa, Ad. & White,
in the much narrower carapace, the breadth-ratio of which is about 1-4 as compared with
1-56 in the last-named species. The Chlorodius fracj'ifer of Adams and Wiiite, referred
to Actceodes by Miers, is apparently a somewhat similar species, but the carapace is still
narrower (breadth-ratio 1'19) than in our specimens and the lobes of the antero-lateral
margins are indistinct and spined. I think it not improbable that a larger series would
connect these specimens with A. hystrlx and possibly with some of the older species.
Xanthodes Lamarckii (Milne-Edwards).
Xantho Lamarckii, H. Mihie-Edwards, Hist. Nat. Crust, i. p. 391.
Xanthodcs granosomanm , Dana, U.S. Expl. Exp., Crust, i. p. 175, pi. viii. figs 10 a-c.
Xanthodes Lamarckii, A. Milne-Edwards, N. Arch. Mus. Paris, ix. p. 200, pi. vii. fig. 3 ; De Man, Arch.
Naturg. liii, (1) p. 2G3 ; Alcock, Journ. Asiatic Soc. Bengal, Ixvii. (2) p. 157.
A male specimen, 10-5 mm. in length by 15-5 mm. in breadth. The carapace is
relatively narrower than in other specimens of this species in the Museum of University
College.
Locality. " Torres Straits."
Chlorodius niger (Forsk.).
Chlorodius niger, Alcock, Journ. Asiatic Soc. Bengal, Ixvii. (2) p. 1(50 (1898).
Eleven specimens, four males and seven females, showing some variation in thei-elative
prominence of the lobules on the carapace and in the acuteness of the antero-lateral
teeth. Some of the specimens show very distinctly the spinulation of the upper edge of
the merusof the ambulatory legs which De Man finds in the type specimens (Zool. Jahrb.
Syst. viii. p. 520).
Localities. " Fringing reef and shore, Thursday Island " ; " Reef, Wyer."
Phymobius ungulatus (Milne-Edwards).
Chlorodius ungulatus, H. Milne-Edwards, Hist. Nat. Crust, i. p. 400, pi. xvi. figs. 5-8 ; Dana, U.S.
Expl. Exp., Crust, i. p. 205, pi. xi. figs. Ha-b.
Phymodius ungulatus, A. Milne-Edwards, N. Arch. Mus. Paris, ix. p. 218; Micrs, Kcp. ' Chall.'
12 DR. W. T. CALMAN ON A COLLECTION OF
Brachyura, p. 1-'5U ; OitiiKum, Zool. Jalirb. Syst. vii. p. 4(i4 ; De Man, Zool. Jabrb. Syst. vii.
p. 524; Alcock, Jouru. Asiatic Soc. Bengal, Ixvii. (2) j). 163 (1898).
Three female specimens.
Oi'tmann (I.e.) unites Daiui's F. iiionticulosus with this species, hut Alcock (7. c.)
retains it as distinct. The characters chosen liy Alcock as diagnostic do not seem qu^ite
consistent with the descriptions of former writers and do not enahle me to discriminate
hetwo(in the species in a series of eleven specimens from Samoa and Japan in our
Museum. The three specimens in Prof. Haddon's collection agree with the original
description of P. monticulosus in liaving the chelipeds " armed with very small pointed
tuhercles "; but as these tubercles beset the whole outer surface of the liand, the specimens
would by Alcock's definition be referred to P. ungulatiis. As regards the sculpture of
the carapace, I can observe no constant difference between these and specimens from
Samoa, in which the chelipeds are distinctly of Alcock's mouticnlosus-ij])e.
Locality. " Torres Straits."
Phymodius sculptits (A. Milne-Edwards).
Chloi-odius sculptus, A. Milne-Edwards, N. Arch. Mus. Paris, ix. 1873, p. 217, pi. viii. fig. 4.
Phijinodius sculptus, Alcock, Jonrn. Asiatic Soc. Bengal, ixvii. (2) p. 164 (1898).
Two male specimens, the larger measuring lO'ii mm. in length by 16-5 mm. in breadth.
They agree very closely with the figures and descriptions quoted above.
Locality. " Torres Straits."
Distribution. Hed Sea to Samoa.
Chlouodopsis melanodactylus, a. Milne-Edwards.
Chlorodopsis mdanodactylus, A. Milue-Edwards, N. Arch. Mus. Paris, ix. p. 229, pi. viii. fig. 7 (1873).
Two male specimens, agreeing well with Milne-Edwards's description and figures, save
that the hands, and especially the fingers, are rather longer and the tubercles on the
fingers are more numerous. The series of specimens in the British Museum shows some
variation in these respects, but in none are the hands so elongated as in our specimens.
Localiti/. " Murray Island, reef."
Distribution. New Caledonia.
Chlorodopsis spinipes (Heller).
FUodins .spinipes, Heller, SB. Ak. Wien, xliii. p. 340, pi. ii. fig. 22 (1861).
Chlorodopsis spinipes, A. Milne-Edwards, N. Arch. Mus. Paris, ix. p. 230, pi. viii. fig. 6 (1873);
De Mau, Arch. Naturg. liii. (1) p. 282 (1887) ; Alcock, Journ. As. Soc. Bengal, Ixvii. (2) p. 169
(1898).
A male and a female specimen are in the collection. De Man has given a redescrip-
tiou of the antero-lateral teeth, finding the descriptions of Heller and Milne-Edwards
obscure. Heller's account, however, is quite api^licable at least to the larger of the
two specimens before me. He mentions (1) two teeth at the outer end of the upper
and lower margins of the orbit respectively, enclosing between them a deep notch, the
external orlntal hiatus ; (2) the three spiniform teeth oP the antero-latsral margin ;
BEACH YURA FKOM TORKES STRAITS. 13
(3) in front of the first of these latter, " zwei ahnlicli gestaltete Ziihnchen ul:)er unci unter
dem "Ranclc unmittelbar hinter der Auii:enhohle." The fig'iire which he gives shows
clearly that the upper tooth of the last-meutioned pair (3) is simply the most anterior of a
row of spiniform tubercles running parallel to the antero-lateral margin on the upper
surface of the carapace. In the figure this tooth is acute and much larger than the
others ; in our sj^ecimens it is much less prominent and less acute, hut still eonsifler-
ahly larger than the succeeding tul^ercles. It is directly over the " suh-hepatic "
tubercle or tooth, which is ev-idently the second tooth of (3), and, looking at the
carapace from the front or the side, the two are naturally associated as in Heller's
description. Milne-Edwards's account is briefer, and does not mention the double extra-
orbital spine. The row of tubercles parallel to the antero-lateral margin is described, and
four antero-lateral teeth are counted, tliat wliich we have called " sub-hepatic" being
reckoned as the first. The figure given by Milne-Edwards resembles very closely
the larger of our two specimens. The smaller specimen diflFers in the fact that the
sub-hepatic tooth (the first antero-lateral of Milne-Edwards) is nearly obsolete, being
represented only by a minute granule. This seems to have been the case with the single
specimen examined l)y De Man, who was thus unal)le to reconcile the apparently
conflicting statements of Heller and Milne-Edwards.
Locality. " ^Eurray Island, reef."
Disfrihutiou. Red Sea to New Caledonia.
Cymo Andreossyi (And.).
Cyrno Andreossyi, Dana, U.S.Expl. Exp., Crust, i. p. 225, pi. xiii. figs. 2u-b; Alcock, Jouru. Asiatic
Soc. Bengal. Ixvii. (2) p. 173 (1898).
One male specimen.
Locality. " Murray Island, reef."
Distribution. Eed Sea to Tahiti.
Cymo melanodactylus, De Haan.
Cijmo melanodactylus, Dana, U.S. Expl. Exp., Crust, i. p. 225, pi. xii. lig. 1 ; Aloock, Jouru. Asiatic
Soc. Bengal, ixvii. (2) p. 174 (1898).
A male specimen.
Locality. " Torres Straits."
Distribution. Ceylon to Eiji and Japan.
Ozius 6UTTATUS ( Milne-Ed wards).
Ozius guttatus, H. Milne-Edwards, Hist. Nat. Crust, i. p. 4.06; A. Milne-Edwards, N. Arch. Mas.
Paris, ix. 1873, p. 239, pi. xi. fig. 1 ; Miers, Rep. Voy. ' Alert,' Crust, p. 228 ; De Mau, Arch.
Naturg. liii. (1 ) 1887, p. 291 ; Ortmann, Zool. Jahrb. Syst. vii. 1893, p. 476.
A female specimen, measuring 32 mm. in length by -iS'S mm. in breadth. The
frontal teeth are considerably more prominent than in Milne-Edwards's figure.
Locality. " Torres Straits.''
14 DE. W. T. CALMAN ON A COLLECTION" OF
PsETJDOZius DISPAE, Dana.
Pseudozius dispar, Dana, U.S. Expl. f]xp., Crust, i. p. 235, pi. xiii. fig. 9.
Sphmrosius dispar, Stimpson, Proc. Acad. Philad. 1858 (1859), p. 35.
Pilumnns nitidus, A. Milne-Edwards, N. Arch. Mus. Paris, ix. p. 249, pi. x. fig. 2 (1873) ; De Man,.
Arch. Naturg. liii. (1) p. 305 (1887).
Pseudozius dispnr, Ortmann, Zool. Jahrb. Sj'st. vii. ]). 433 (1893).
Two sj)ecimens, the larger of whicb, an ovigerons female, differs somewhat from
Milne-Edwards's figure as regards the outline of the carapace. The antero-lateral margin
is considerably shorter than the postero-lateral. The greatest width is in the line
of the penultimate antero-lateral teeth and well in front of the middle of the length.
In the figure the greatest width is at about the middle at the level of the last pair of
antero-lateral teeth. The description given by De Man applies accurately, in most points,
to our specimen, except as regards the supposed sexual difl"eronces. He found the
carapace narrower in a male specimen, the breadth-ratio being only 1"25 as against 1-37
in the female; and in the male the outer surface of the large hand was smooth with only
a few granules near the proximal end, while in the female the whole surface was covered
with granules as in the smaller chela. The latter difference between the sexes was also'
found by Ortmann. As regards tlie breadth of the carapace, our sjiecimen is intermediate
between the two examined by De Man, the ratio being I'Sl. The measurements given
by Milne-Edwards have evidently suffered from some misprint, but measurements takea
from his figure of an adult male give a breadth-ratio of 1"41, considerably greater than
that given by De Man for the female. As regards the granulation of the large cheliped,
our specimen presents exactly the condition figured by Milne-Edwards and described by
De Man as characterizing the male, the outer surface being smooth with a few granules
grouped near the proximal end. Our second specimen, a minute and immatm-e female.
Las more numerous granules on the large chela, which, however, is still much smoother
than the small chela. In both cases the right cheliped is the larger.
De Man suggested the possible identity of Milne-Edwards's species wiihihe Fseudozius
dispar of Dana, and this suggestion has been accepted by Ortmann. Dana's figure is
very similar to our specimens, and his description, though lacking in detail, applies
perfectly, save in the one point that the finger of the large hand is said to be " smooth
and round, and not channeled." In our specimens, as in De Man's account, the fingers
of both hands are grooved. Dana gives the breadth-ratio of the carapace as 1-21, rather
less than the narrowest individual examined by De Man. The table of generic characters
given by Dana [1. c. p. 229) states that the carapace in this genus is " fere planus " ; but
that this does not apply to the species in question may be gathered from the fact that
Stimpson refers it to his genus SphcBrozius, which is defined as having the body sub-
globose. Stimpson, however, gives as a generic character " Margo frontalis et supra-
orljitalis continuae nee sinu nee incisura separataD." In our specimens, as in De Man's
description, the frontal lobes are separated fi'om the supraorbital margin by a slio-ht but
BKACHYUEA FROM TOKKES STRAITS. 15
distinct notch. Ortmann retains the species in the original genus '• weil die Antero-
lateralziihne iindeutiich sind."
Locality. " Murray Island, reef."
Distribution. Sulu Sea to New Caledonia.
Plltjmnus cursor, a. Milne-Edwards.
Pilumnus cursor, A. Milne-Edwards, N. Arch. Mus. ix. p. 244, pi. ix. fig. 4 (187.3) ; Miers, Hep. Voy.
' Alert,' Crust, p. 223 ; De Man, Arch. Naturg. liii. (1) p. 29!) (1887) ; Alcock, Journ. Asiatic Soc.
Bengal, Ixvii. (2) p. 195 (1898).
Three of our four specimens agree pretty closely with the descriptions of the aiitliors
cited above. They show some variation in the hairiness of the carapace : in two male
specimens the body is covered with a scanty short pubescence, u ith wjiich, near the front
margin, a few longer hairs are intermixed ; in a female individual, however, the lono
hairs are much more numerous imd extend further hack on the carapace. The hands
are ornamented with rows of tubercles, not qviite so thickly set as in Milne-Edwards's
figure, interspersed with short hairs. The legs are a little longer than in the figure, and
the merus of the first three pairs has a few spinules on its upper margin.
One specimen, a male infested by a Pi,hizocephalan parasite, differs in certain points
from the others, but cannot, I think, be specifically separated from them. The outer
surface of the larger (right) chela is here for tlie most part quite smooth, with only a
few granules and hairs near the proximal end and along the upper margin, the groove
on the dactylus is represented by a line of punctations, the sub-hepatic tubercle is
wanting, and the ambulatory legs are even longer than in the more typical sj)ecimens.
In the armature of the cheliped this specimen resembles P. Audet^soni, De Man (Journ,
Linn. Soc, Zool. xvii. p. 59, pi. iii. figs. 5, 6), from which, however, it is distinguished
by the shape of the carapace, the antero-lateral margins being much shorter and the
postero-lateral more nearly parallel than in that species.
Localities. " Murray Island, reef " ; " Channel between reefs, Mer."
PiLTJMNUS PULCHER, Miers.
Pilumnus pule her, Miers, Rep. Voy. 'Alert,' Crust, p. 219, pl. xxii. fig. A.
Actumnus pulcher , Ortmann, Semon's Forsch. Reise Austr., Crust, p. ^'Z (1894).
The larger of our two specimens is only G mm., long, but it agrees pretty closely with
Miers's description and figiu-e and with the much larger type sjiecimens with which I
have compared it. The chief difference is the somewhat greater length of the legs ; in
Miers's specimens the penultimate leg is about as long as the greatest breadth of the
carapace, while in our larger specimen the carapace measures 6'5 mm. in breadth, and
the corresponding leg nearly 8 mm. in length. Ortmann has referred this species to
Actumnus, but the characters which he adduces hardly seem to justify its removal to that
genus.
Locality. " ^lurray Island."
16 DR. W. T. CALMAN ON A COLLECTION OF
PiLTJMNUS SEMINUDTJS, MierS.
Pilumnus seminudus, Miers, Rep. Voy. ' Alert/ Crust, p. 222, pi. xxi. fig. C.
Two female specimens, measuring about 6-25 mm. in length. Though little more than
half the size of Miers's type specimen they agree very closely with it. In one individual
the antero-lateral teeth are much less prominent than in the type, and the middle tooth
of the right side is wanting. In hoth specimens the pubescence on the front part of the
carapace is less developed, and there are a few longer hairs in a transverse row in front
of the protogastric region. The species bears a considerable resemblance to the
Psmdozius di>ipar referred to above, and perhaps the two should not be geneiically
separated.
LocaUtij. " Mabuiag."
PiiiTJMNTJS LANATUS (Latr. ?), Miers.
Pilumnus lanatus (Latr.), Miers, Rep. Voy. ' Alert,' Crust, p. 220, pi. xxi. fig. A.
A number of small and probably immature specimens are all but identical with tlie
' Alert ' specimens described under this name by Mr. Miers. The close pubescence
covering the body and limbs ends abruptly on the outer surface of the larger chela along
a diagonal line drawn from the base of the dactylus above to the proximal angle below,
the rest of the surface being smooth and polished. The antero-lateral teeth are smaller
than in Miers's figure, the last tooth in particular being much reduced. A small, but
distinct, sub-hepatic tubercle is present, the upper surface of the carapace is shghtly
uneven in front, and there is a well-marked tubercle on the hepatic region above. The
chelge are rather shorter, the lower finger is not hooked at the tip as in Miers's figure,
and I cannot detect the spinules which he describes on the carpus of the ambulatory
legs.
Locality. " Torres Straits."
Pilumnus longicoknis, Hilgd.
Pilumnus longicornis, Hilgendorf, Monatsb. Akad. Berlin, 1878, p. 794, pl.i. tigs. 8, 9 ; var., Miers, Rep.
'Challenger' Brachyiua, p. 157; Alcock, Journ. Asiatic Soc. Bengal, Ixvii. (2) p. 193 (1898).
A male specimen is referred with some doubt to this species. The carapace measui-es
17 mm. in length and 23 mm. in breadth (exactly the dimensions of Ililgendorf's
specimen), is strongly arched in the front part in an antero-posterior direction and
slishtly so from side to side. The three antero-lateral teeth are very prominent and
conical, the second and third with spiniform points curved forwards. The acute sub-
hepatic tuhercle, though small, is visible from above, causing the antero-lateral margin
to appear four-toothed. The external orbital angle is produced into a small triangular
tooth. The postero-lateral margin is longer than the antero-lateral, and is slightly
concave owing to the prominence of the last pair of antero-lateral teeth. The ^hole
surface of the carapace bears scattered and rather coarse granules, only some of the
broad, shallow, inter-regional grooves being smooth. The setae covering the carapace are
rather thick-set and moderately long, springing in pencils of four ur five, of which one
BBACHTURA FROM TOERES STRAITS. 17
is genei'ally much longer than the rest. The front is much tleflexed, the inner lohcs
broadly rounded, the outer small and spiniform. The upper orbital margin has two
hardly visible fissures, and is, like the lower margin, only faintly granulated. The
internal angle of the lower margin is acutely rounded. The sub-hepatic region bears a
few" granules near the outer margin besides the sub -hepatic tooth. The merus of the
chelipeds has a stout blunt tooth near the distal end of its upper margin. The carpus
has a number of granules on its outer surface, which is clothed with long setae, and there
is a sharp tooth at its inner angle. In the hand of the larger cheliped the greatest
breadth is about equal to the length of the palm measured in the middle line, and the
dactylus is about three-fourths of this length. The whole outer surface of the palm
bears longitudinal rows of acute granules, with one or tw^o smaller granules scattered in
the rather wide interspaces between the rows. On the upper margin some of the
granules become spiniform, and the whole surface bears numerous rather long setfe,
which extend with the granulation on to the bases of both fingers. The fingers are slightly
furroAved. The merus of the ambulatory legs has the upper edge terminating distally in
a spine, behiiid which there is a notch running down on both faces of the joint as a
short groove.
From Hilgendoi'fs accou.nt our specimen differs in the rather more concave postero-
lateral margins; in the more prominent sub-hepatic tooth (" nur angedeutet ") ; in the
much less distinct granulation of the orbital margins ; and in having the whole outer
surface of the hand covered with granules and setfe. The granules on the surface of
the carapace also aj)pear to be more numerous. Unfortunately both antennal flagella
are wanting in our specimen. The regions of the carapace are not so distinctly marked
as in Hilgendorf's figure, but they appear to be similarly disposed. The tooth on the
distal end of the merus of the ambulatory legs is not indicated by Hilgendorf.
The ' Challenger ' specimen described by Miers, and regarded by him as a variety of
this species, differs from the type and agrees with our specimen in the characters of the
orbital margin, of the larger cheliped, and of the ambulatory legs. The sub-hepatic
tooth, however, is said to be deficient and the chelipeds are nearly equal in size.
P. Sluiteri of De Man (Weber, Reise Niederl. O.-Ind. ii. p. 283, pi. i. fig. 2, and (as
P. ForskaUi, M.-E.), Arch. Naturg. liii. (1) p. 295, pi. xii. fig. 1) is a closely allied species,
but differs in the less prominent antero-lateral teeth and more granulated caraj>ace, and
in the absence of the notch and tooth on the merus of the ambulatory legs. P. scabri-
iisciiliis, Ad. & Wh. (Zool. Voy. ' Samarang,' Crust, p. 44, pi. ix. fig. 5) has the antero-
lateral teeth less prominent, wdde, and denticulated.
Locality. " Fringing reef, Mer, Murray Island."
Distribution. E. Africa (Inhambane, Hilgendorf), to Tongatabu (Jflifrs).
PiLTJMNUS CKiSTiPES, n. sp. (Plate 1. figs. 1-3.)
Carapace closely covered with a short fur, which does not conceal the ratlier prominent
regional areolte. On removing the fur, a few scattered granules are seen, each bearing
a tuft of short hairs. The front part of the carapace is strongly convex in an antero-
SECOND SERIES. — ZOOLOGY, VOL. YIII. 3
IS I»K. W. T. CALMAN ON A COLLECTION OF
posterior direction, while the posterior part is flat, rrom side to side, in the line of the
lateral teeth, the surface is only slightly convex. The strongly deflexed front is divided
by a rather deep incision into two rounded lobes, and the outer angles form sharp
downwardly directed teeth, not visible from above, separated by a groove from the
supra-orbital angle. The upper margin of the orbit is interrupted by two short open
fissures, and a third is present on the lower margin just below the external angle. The
external angle of the orbit is not very prominent, rounded, and produced backwards for
a short distance as a slight ridge above the level of the first antero-lateral tooth. The
antero-lateral margin is equal in length to the postero-lateral, and is cut into four thick,
bluntly rounded teeth, increasing in length from before backwards, covered with fur
intersjjersed with granules.
The flagellum of the antenna is nearly one fourth the length of the carapace.
The ridges of the palate are distinct but not very prominent, and become obsolete before
reaching the front margin of the buccal frame.
The chelipeds are very unequal ; the merus is short, trigonous, the upper margin
carrying a large rounded tooth separated by a narrow incision from the projecting
rounded distal angle ; the carpus has on its convex outer face scattered granules,
l^artly arranged in oblique lines and interspersed with fur, and a transverse groove
runs parcillel to the distal margin ; the hand of the larger cheliped (the right) has the
palm but little longer than broad, with longitudinal rows of granules nearly hidden
by the fur on its outer face ; the fingers are stout, aliout one-half the length of the
palm, with blunt rounded teeth on the inner edges ; the dactylus has a patch of
granules and hairs at its base, and both fingers are slightly grooved. In the smaller
hand the granules on the outer face are less regularly arranged in rows.
The ambulatory legs have the merus wdth a sharp crest on its upper edge, rising
gradually with a straight edge towards the distal end, where a narrow notch separates
it from the prominent rounded distal tooth. In the first three pairs the anterior and
posterior faces of the merus are smooth and nearly free from hair. In the last pair of
legs the jjosterior face of the merus is closely furred. The carpus and propodus of all
the legs are considerably expanded and flattened, covered with fur rather longer than
that on the carapace, and the carj)us has a deep longitudinal groove on the anterior and
posterior faces. The dactylus of all the legs is stout and nearly cylindrical. The
abdomen (female) has all seven joints free.
The form described above differs from all the species of Pilumnus known to me in the
crested merus of the ambulatory legs, the feebly developed endostomial ridges, and the
blunt teeth of the antero-lateral margin. In the first two of these characters it aj)proaches
the P. dilai/pes of Adams & White (Zool. Voy. ' Samarang,' Crust, p. 4i, pi. ix. fig. 4),
for Avhich Miers has proposed to constitute a separate genus Lophopilmniius (Rep. Chall.
Brachyura, p. 148) ; but in that species the antero-lateral teeth are broad and denticulated
and the meral crests are of very dilferent shape and are not divided by a notch near the
distal end. In the great convexity of the anterior portion of the carapace the species
has some resemblance to an Actumnus, in which genus, however, the carapace is not
BRACHTUEA FEOM TORRES STRAITS. 19
flattened posteriorly and is usually strongly arched from side to side. Dr. De Man has
suggested to me a possil)le relationship with the genus Lophoxmithns. In that "-enus,
however, the carapace is usually depressed and glabrous, and though Miss Eathlnin has
recently described a species L. frontalis (Proc. U.S. Nat. Mus. xvi. 1893, p. 23G) formino-
an exception in both these respects, it does not appear to draw appreciably nearer to
the present form. The whole aspect of our species, the shape of the front, and many
other small points are so suggestive of Filumnus that it seems best, for the present at
any rate, to include it in that genus, tliougli it certainly diverges considerably from the
more typical species.
Length of carapace 20 ram. Length of chela 18 mm.
Breadth „ 28 „ „ dactylus 8 „
Length of larger cheliped 31 ,, „ last leg 30 „
Locality. " Fringing reef, Mabuiag."
AcTUMjrus SETiFER (De Haan).
Pilumnus setifcr, De Haan, Faun. Japon., Crust, p. 50, pi. iii. fig. .3.
Actttmnus setifer, A. Milne-Edwards, N. Arch. Mus. P.iris, i. p. 287, pi. xv. figs. 5-.") 6 (18G5) ;
Alcock, Journ. Asiatic See. Bengal, Ixvii. (2) p. 202 (1898).
Our four specimens show considei-ablc variation in several points, l)ut must all, I think,
])e referred to this common and widely distributed species. The smallest specimen is a
female, and, although only 6'25 mm. in length, carries eggs. In this individual the
lobulation of the carapace is not very pronounced, and the antero-lateral teeth are
prominent and tipped with spiniform tubercles. In some of the larger specimens the
lobules of the carapace are more prominent, the carapace is more convex, and the antero-
lateral teeth are reduced to low, rounded lobes, on wliich the minute spiniform points
are completely hidden by the dense pubescence covering the whole carapace. These
differences, as well as slight variations in the relative breadtli of the carapace, appear to
be independent of age or sex. In a very large male, 17'5 mm. in length, from Sagami
Bay, Japan, in the Museum of University College, the antero-lateral spines are very
distinct, and the lobulation of the carapace is less pronounced than in a specimen only
7 mm. long in Prof. Haddon's collection.
Localities. " Flinders Entrance, near Mer, 20-30 fatli." ; " S. of Orman's reef,
5-7 fath."
Trapezia feretjginea, var. areolata (Dana).
Trapezia ferrvginea areolata, Ortmann, Zool. Jalirl). Syst. x. p. 206 (with synonymy) ; Alcock, Jour-;.
Asiatic Soc. Bengal, Ixvii. (2) p. 221 (1898).
One specimen, an ovigerous female about 10 mm. in length, having the lateral teeth
of the carapace acute. De Man has pointed out (Arch. Naturg. liii. (1) p. 317) that this
juvenile character is occasionally retained in adult individuals, though as a rule these
teeth become obtuse (var. iy/^;'>M«s, A. M.-E., N. Arch. Mus. Paris, ix. p. 259, pi. x. fig. (5).
8*
20 DR. W. T. CALMAN ON A COLLECTION OF
The reticulations on the carapace of our specimen are rather larger and more symmetri-
cally disposed than in Milne-Edwards's figure.
Locality. " Murray Island."
Distribution. Ceylon to Tahiti.
Trapezia crMODOCE (Herbst).
Trapezia cymuduce, Ortmaim, Zool. Jalirb. Syst. x. p. 203 (with synonymy) ; Alcock, Journ. Asiatic
Soc. Bengal, Ixvii. (2) p. 219 (1898).
Three specimens are referable to this species as defined by Ortmann. The pubescence
on the outer surface of tlie chela is rather scanty and not conspicuous in dried specimens,
and the lower margin of the chela is faintly granular. The carpus, of the chelipeds bears
internally an acute spiniform tooth ; only in one detached cheliped is this tooth blunt
{of. De Man, Arch. Naturg. liii. (1) p. 310).
Locality. " Murray Island, channels between reefs, 15-20 fath."
Tetralia GiiABERUiMA (Herbst).
Tetralia glaberrima. Ortmann, Zool. Jahrb. Syst. x. p. 209 (with synonymy); Alcock, Jouru. Asiatic
Soc. Bengal, Ixvii. (2) p. 223 (1898).
Two specimens, one of which resembles tlie form named T. nigrifrons by Dana (U.S.
Expl. Exp., Crust, i. p. 262, pi. xvi. fig. 2). The dark marginal band extends across the
wdiole front edge of the carapace and halfway down the sides.
Locality. " Reef, Wyer."
Distribution. Red Sea to Marquesas.
LissocARCiNrs ORBICULARIS, Dana.
Lissocarcinus orbiadaris, Dana, U.S. Expl. Exp., Crust, i. p. 288, pi. xviii. figs. 1 a-e; A. Milne-
Edwards, Arch. Mus. Paris, x. p. 418 (1861) ; Alcock, Journ. Asiatic Soc. Bengal, Ixviii. (2)
p. 20 (1899).
Six specimens presenting all the characters of Dana's species, but showing some slight
variation in the distinctness of the antero-lateral teeth and in the concavity of the
postero-lateral borders and consequent prominence of the lateral angles.
Locality. " Murray Island, reef."
Distribution. Mauritius to Fiji.
Caphyra rotundiprons, a. Milne-Edwards.
Camptonyx rotundifrom, A. Milne-Edwards, N. Arch. Mus. Paris, v. 1869, p. 156, pi. vii. figs. 11-12.
Cajihyra rotundifrons, A. Milne-Edwards, N. Arch. Mus. Paris, ix. 1873, p. 174.
Two female specimens of this very rare species are in the collection. They agree very
exactly with Milne-Edwards's description and figure, the only observable differences
being that the frontal lobes are slightly more prominent in the middle and are separated
from the supra-orbital margin on either side by a shallow notch ; the anterior margin of
the merus of the chelipeds bears three teeth, and the merus of the second legs has no
BEACHYUEA FEOM TOEEES STEAITS. 21
spine on its upper border. The carapace of our larger specimen measures 12'25 mm. in
length by 15"5 mm. in breadth.
Locality. " Torres Straits."
Distribution. New Caledonia and Samoa [Ililne-Edwards).
Xeptuntjs sangtjinolenttjs (Herl)st).
i?//ja «a«^«iwo/e«/o, Milne-Edwards, Hist. Nat. Crust, i. p. 451, &in Rl'gne Anim., Crust, pi. x. fig. 1.
Neptunus sanyuinolentus, A. Milue-Ed wards, Arch. Mus. Paris, x. p. 319 (18G1) ; Alcock, Jouru.
Asiatic Soc. Bengal, Ixviii. (2) p. 32 (1899).
Two small and. imperfect specimens, the larger only 15 mm. in length, belong
apparently to this common species, though the characteristic " ocelli " are very faintly
indicated on the carapace.
Locality. " Torres Straits."
Neptunus pelagicus (L.).
Lupa pelagica, Milne-Edwards, Hist. Nat. Crust, i. p. 450.
Neptunus pelayicus, De Haan, Faun. Jap., Crust, p. 37, pis. ix., x. ; A. Milne-Edwards, Arch. Mus.
Paris, X. p. 320 (1861) ; Alcock, Jouru. Asiatic Soc. Bengal, Ixviii. (2) p. 34 (1899).
Two specimens, about 22 mm. in length , are no doubt referable to this common species,
though they differ in some slight details from large specimens. In general shape and
in the character of the antero-lateral teeth they approach the N. armatus of A. Milne-
Edwards [1. c. p. 322, pi. xxxiii. fig. 2), but the external frontal teeth are not in our
specimens obtvise, and the spine on the median supra-orbital lobe is indicated, though not
so large as in full-grown specimens of N. pelagicus. As Ortmann has pointed out (Zool.
Jahrb. Syst. viii. p. 75), it is very doubtful whether N. armatus is a distinct sj)ecies.
Milne-Edwards says of it : " Cette espece est de tons les Neptunus connus la plus
elargie " ; while Miex"s, referi'ing to the vei*y specimen described by Milne-Edwards, states
that " the carapace is relatively somewhat narrower .... than in N. pelagicus of about
the same size" (Rep. Voy. 'Alert,' Crust, p. 229),
Locality. " Fringing reef, Mabuiag."
Neptunus (Achelous) granulatus (Milne-Edwards).
Lupa granulata, Milne-Edwards, Hist. Nat. Crust, i. p. 454.
Amphitrite gladiator, De Haan, Faun. Jap., Crust, p. 65, pi. xviii. fig. 1 [not pi. i. fig. 5).
Achelous granulatus, A. Milne-Edwards, Arch. Mus. Paris, x. p. 344 (1861).
Neptunus {Achelous) granulatus, Alcock, Journ. Asiatic Soc. Bengal, ixviii. (2) p. 45 (1899).
A female specimen, 12 mm. long, agrees well with De Haan's figure, except that, as in
all young specimens, the lateral spines are more elongated.
Locality. " Murray Island."
Neptunus (Achelous) granulatus, var. unispinosus, Miers.
Achelous granulatus, var. unispinosus, Miers, Rep. Voy. ' Alert,' Crust, p. 230, pi. xxiii. fig. B.
Neptunus [Achelous] unispinosus, Miers, Rep. Voy. 'Challenger/ Brachyura, p. 180; De Man, Zool.
Jahrb. Syst. viii. p. 558.
•22 DR. W. T. CALMAN ON A COLLECTION OF
Two males, 7'5 and 11 mm. long respectively, agi*ee witli Miers's type specimen in the
shape of the frontal lobes and in having only one spine on the posterior edge of the arm.
The antcro-lateral teeth, however, are not distinctly more spiniform, nor is the last tooth
longer than in specimens of A. grmiulatus of similar size. The second spine of the
posterior edge of the arm is represented by a slight rudiment, as it is, indeed, in the type
specimen. I do not think that the form can be ranked as more than a variety of
A. granulafus, as it was originally regarded by Miers.
Localities. " Sabai Channel " ; " Murray Island.''
Thalamita prtmna (Herbst).
The forms of Thalamita in which the front is divided into eight lobes were distributed
by A. Milne-Edwards among seven species and reunited by Kossmann into one, while
more recent writers haA^e expressed various views intermediate between these two
extremes. Alcock has recently affirmed his belief in the correctness of Kossmann's view,
while retaining, for the sake of convenience, sei^arate specific names for some of the
forms. Ten specimens collected by Prof. Haddon belong to this section of the genus and
fall into three groups, not one of w^hich agrees in all points with any of the described
species : —
{a) A large male, the carapace of which measures 37 mm. in length by 59 mm. in
breadth, agrees best on the whole wdth the descriptions of the typical Th. prymna, but
presents certain points of diff'erence. Comparing the frontal lobes wdth the figure given
by DeMan (Journ. Linn. Soc, Zool. xxii. pi. iv. fig. 5), the outer or fourth pair are much
more strongly arcuate and resemble the figure of Th. spinimana (I. c. fig. 7) ; the third
pair of frontal lobes are separated by an open fissure from the second or submedian, as
in Dana's figure of Th. crassimana (U.S. Expl. Exp., Crust, pi. xvii. fig. 9«) ; the sub-
median are slightly less prominent than the median lobes, which they distinctly ovei'lap
above, an arrangement which, according to A. Milne-Edwards (Arch. Mus. Paris, x.
p. 362), "ne se voit jamais chez le Th. prymna." De Man's description and figure of
Th. Cfpruleipes (Zool. Jahrb. Syst. viii. p. 568, pi. xiv. fig. 12 a) fits this specimen well as
regards the third and fourth frontal lobes, but the median pair are stated to be wider
than the submedian, while in the present instance the reverse is the case. The basal
antennal joint carries a row of about four sharp spines besides some smaller granules.
Milne-Edwards assigns to it only two or three spines, while Dana figures an irregularly
toothed crest. The fourth antero-lateral tooth is very small, and the greatest breadth of
the carapace is measured between the third pair of teeth. The cheliped differs from all
descriptions of Th. prymna in having three spines instead of two on the upper margin of
the hand, the additional spine being smaller than the other two and close to the
proximal end.
{h) A male specimen, 21 mm. long by 33 mm. broad, has a row of granules on the
basal antennal joint and a minute fourth antero-lateral tooth, and would therefore be
referred by Milne-Edwards's table (/. c. p. 367) to Th. Stimpsoni, which De Man regards
as a variety of Th. Bancv (Journ. Linn. Soc, Zool. xxii. p. 78). With De Man's figure
of Th. Dante {I. c. pi. iv. fig. 8) our specimen agrees in the nearly strtiight anterior edge
BKACHYURA FEOM TORRES STRAITS. 23
of the outer frontal lobes, but it differs in having the other three pairs of lobes separated
only by slight notches. The abdomen does not present the peculiar outline figured by
De Man (7. c. fig. 9), but he has since stated that this character is not constant (Notes
Leyden Mus. xv. j). 285). In most other points this individual agrees closely with the
above described specimen a, lacking, however, the third spine on the ujoper edge of the
hand. A larger female specimen (31 mm. long) agrees with this, except that the fourth
autero-lateral tooth is a little larger.
(c) Eight specimens, all of small size, including two ovigerous females of 8 mm. and
7 mm. in length respectively. The outer frontal lobes have a well-curved margin ; the
third lobes are narrow, rounded, and sej)arated from the second by an open notch; the
second or submedian lobes are very broad, about half as broad again as the median pair,
which they distinctly overlap ; the three inner j^airs of lobes reach to about the same level.
The fourth antero-lateral tooth is very minute or absent. The ridges of the hand have
the same arrangement as in Th. pri/mua, but on the lower half of the outer face the
ridges and the intervening spaces are quite smooth. The basal joint of the antenna is
markedly shorter than in the other specimens described above.
These small specimens depart more widely from the typical 27i. piyimia than do the
other specimens, but I have failed to identify them with any of the described species.
Localities. " Flinders Entrance, near Mer, 20-30 fath." ; " Channels between reefs,
Mui-ray Island, 15-20 fath."
Thalamita admete (Herbst).
Thalamita admete, Milne-Edwards, Hist. Nat. Crust, i. p. 459 ; Daua, U.S. Expl. Exp., Crust, i. p. 281,
pi. xvii. figs. 5«-c; A. Milne-Edwards, Arch. Mus. Paris, x. 1861, p. 350.
T. Saviffinji, A. Milue-Edwards, 1. c. p. 357.
T. admete, Alcock, Journ. Asiatic Soe. Bengal, Ixviii. (2) p. 82 (1899).
Three female specimens, the largest 10'5 mm. long by 11'5 mm. broad, carrying ova.
They appear to agree with the descriptions of Tk. Savignyi, and are very similar to
specimens so labelled in the British Museum. There appears, however, to be little doubt
that this form is only a variety of Th. admete.
Locality. " Channel between reefs, Mer."
Thalamita sijia, Milne-Edwards.
Thalamita sima, Milue-Edwards, Hist. Nat. Crust, i. p. 460.
Portunus {Thalamita) arcuatus, De Haan, Faun, Jap., Crust, p. 43, pi. ii. fig. 2.
Thalamita sima, A. Milne-Edwards, Arch. Mus. Paris, x. p. 359 (1861) ; De Man, Zool. Jahrb. Syst.
viii. p. 564 (1896) ; Alcock, Jouru. Asiatic Soc. Beugal, Ixviii. (2) p. 81 (1899).
Six specimens, including an ovigerous female only 8'5 mm. in length, are referred to
this species. In the larger specimens (20 mm. long) the antennal crest is smooth or
nearly so, but in the smaller it is minutely granulated. In none are the margins of the
median frontal lobes concave as described by De Man.
Localities. "Fringing reef and shore, Thursday Island"; " Channel between reefs,
Murray Island."
24 DE. W. T. CALMAN ON A COLLECTION OF
Kraussia nitida, Stimpson.
Kranssia nitida, Stimpson, Pr. Acad. Pbilad. 1858, p. 40 ; Miers, Rep. Voy. ' Alert,' Crust, p. 235 ;
Henderson, Tr. Linn, Soe., (2) Zool. v. p. 379, pi. xxxvii. fig. 9 (1893) ; Alcock, Journ. Asiatic
See. Bengal, Ixviii. (2) p. 98 (1899).
A single male specimen appears to agree well with this species as briefly characterized
by Stimpson and Miers and more fully by Henderson and Alcock. The frontal lobes are
rather less deeply subdivided than in the figure given by Henderson. The whole surface
of the carapace is covered with very minute granulations in short transverse rows. The
outer surface of the hand is faintly granidated distally and near the upper edge.
Locality. " Channel between reefs, Murray Island."
Metopogeapsus messor (Forsk.).
Metopoyrapsus messor, Kingsley, Proc. Acad. Nat. Sci. Philad. 1880, p. 190; Ortmann, Zool. Jahrb.
Syst. vii. 1894, p. 701.
A small specimen (8-6 mm. long) appears to belong to this widely distributed species.
In the relative length of the propodus of the ambulatory legs it approaches the variety
gracilipes of De Man (Notes Leyden Mus. xiii. p. i9), but the specimen is too immature
for precise determination.
Locality. " Cockburn group (N. Queensland), shore."
Varuna litterata (Fabr.).
Varuna litterata, Kingsley, Proc. Acad. Nat. Sci. Philad. 1880, p. 205; Ortmann, Zool. Jahrb. Syst.
vii. 1894, p. 713; De Man, Zool. Jahrb. Syst. ix. 1897, p. 112.
One female specimen.
Locality. " Torres Straits."
OCYPODA CERATOPHTHALMA (Pallas).
Ocypoda ceratophthalma, Ortmann, Zool. Jahrb. Syst. x. 1897, p. 364 (with synonymy).
Five full-grown males and one female and a number of immature individuals are in
the collection.
Localities. " Eeef, Murray Island " ; " Mer " ; " Mabuiag."
UCA TETRAGONON (Hcrbst).
GeJasimus tetragonon, Kingsley, Proc. Acad. Nat. !Sci. Philad. 1880, p. 143, pi. ix. fig. 11 (with
synonymy) ; De Man, Notes Leyden Mus. xiii. 1891, p. 24, pi. ii. fig. 6 ; Ortmann, Zool. Jahrb.
Syst. vii. 1894, p. 754.
Ilea tetrayona, Ortmann, Zool. Jahrb. Syst. x. 1897, p. 348.
A male specimen, in which the carapace measures 13 mm. in length, is referred to this
species. In the armature of the fingers the large chela agrees precisely with Kingsley's
figure [I. c), but the fingers themselves are much shorter, not equalling the palm in
length, and the outline of the chela therefore resembles the figure of G. variatus which
Kingsley (/. c. pi. x. fig. 32) copies from Hess. The last-named species is regarded by
BRACHTUEA FEOM TORRES STEAITS. 25
De Man and Ortmann as synonymous with the present. De Man's description and figure
(I. c.) agree closely witli our specimen, save that the orbits are more oblique and the
fingers of the chela much longer in the figure.
Locality. "Torres Straits."
Distribution. Eed Sea to Sandwich Islands.
Ceratoplax (?) sp.
An imperfect dried specimen resembles rather closely in general shape the Ceratopln.r
ciliata of Stimpson, as figured by Miers (Chall. Rep., Brachyura, p. 234, pi. xix. fig. 3).
It differs, however, in the broader meral and carpal joints of the walking-legs (Miers
states that in his specimen the legs " are i-ather more slender than in the description of
Dr. Stimpson ") and in the rather stouter fingers of the chelipeds. It differs, moreover,
not only from Miers's figure, but also from the generic diagnosis in the fact that the
antero-external angle of the merus of the third maxillipeds is roimded oil' and not
distinctly produced.
Locality. " Torres Straits."
Calappa hepatic a (L.).
Calappa hepatica (L.), Alcock, .Tonrii. Asiatic Soc. Bengal, Ixv. (2) p. 142 (1896).
Three female specimens, the carapace of the largest measuring 40 mm. in length by
60 mm. in breadth.
Locality. " Miu-ray Island."
CRTPTOCNEMrs Haddoni, n. sp. (Plate 1. figs. 4-8.)
Description. The margins of the lateral wings of the carapace are convex, presenting no
salient lateral angle such as is present in C. pentagonus, Stps., and passing, with scarcely an
indication of a poster o -lateral angle, into the posterior margin, where the curve meets its
fellow in the middle line in a slight re-entrant angle. The lateral margin is continued on
to the dorsal surface of the carapace beliind the hepatic region on either side as a faintly
marked ridge which soon dies out. The front is obtusely triangular and slightly reflexed,
the straight line of each side being continued beyond the orbit to the prominent hepatic
angle. On the flat dorsal surface a low longitudinal keel runs from the tip of the
rostrum to near the posterior edge of the carapace, being most prominent on the cardiac
region, and the branchial regions are very slightly inflated. The surface of the carapace
is perfectly smooth ; the posterior and lateral margins, as well as the faint ridges on the
hepatic regions, are microscopically beaded. The antennular fossse are transverse. The
third maxillipeds have the merus equal in length to the ischium, acutely triangular and
projecting well beyond the margin of the buccal area, though not so far as to be visible
from above. The exopod is equal in breadth to the ischium ; its outer edge is convex,
the tip truncate and very slightly concave. The chelipeds have the merus trigonous,
with two minutely granular lines on its lower margin ; the carpus has a slight keel
exteriorly ; the hand is compressed, the edges acute, the palm being one and a half times
SECOND SERIES. — ZOOLOGY, VOL. VIII. 4
26 T)R. W. T. CALMAN ON A COLLECTION OF
as long as broad ; the tingers are grooved and oue-tbird tlie length of the palm. The
walking-legs have the merus compressed and crested above and below, the carpus and
propodus with a double keel on the upper edge and the dactylus styliform. The abdomen
of the female is very nearly circular in outline, and the first, second, and last somites are
free. The whole of the under surface is quite smooth.
Length 5-5 mm. ; breadth 7 '25 mm.
Of the four described species of Cryptocnmius our new form approaches most closely
to the type, C. pentagonus of Stimpson (Pr. Acad. Philad. 1858, p. 161), figured by
Miers (Proc. Zool. Soc. 1871), p. 13, pi. ii. tig. 5), whose imperfect specimen I have
examined. In that species, however, the wings of the carapace are produced into acute
lateral angles, and the straight postero-lateral and posterior margins meet at an obtuse
angle ; the front is more sti'ongly reflexed and more acute, the branchial regions are more
convex, the lateral margins are not continued on to the dorsal surface in front, and the
granulation of the posterior and lateral margins is more pronounced than in the present
species ; the antennular fossae ai'e oblique and the palp of the external maxillipeds is
rounded, not distinctly truncate, at the tip. In C. Hokhicorthi, Miers (Tr. Lion. Soc,
(2) Zool. i. p. 241, pi. xxxviii. figs. 30-32, 1878), the lateral margins are at right angles
to the posterior margin, and there are two olilique carinse running from the cardiac
region to the posterior corners on the dorsal surface of the carapace. C. Grand tdicrl,
A. Milne-Edwards (Ann. Soc. Ent. France, (4) v. p. 155, pi. vi. fig. 4, 1865), resembles
Q. HoldswortJii in general shape, but has a broadly truncate front, the posterior border
is notched in the middle, and the surface of the carapace has three granulated carina?
diverging from the centre to the rostrum and the two posterior corners respectively.
Finally, C. oboliis, Ortmann (Zool. Jahrb. Syst. vi. p. 570, pi. xxvi. fig. 12, 1892), has a
nearly circular outline, the rostrum is bifid, and the hepatic prominences are acute,
almost spiniform, teeth.
Locality. " Channel between reefs, Mer."
Oreophoeus frontalis, Miers.
Oreuphorus frontalis, Miers, Rep. Voy. ' Alert,' p. 2o4-, pi. xxvi. fij^. B.
A comparison of our eight individuals with the unique type specimen leaves no doubt
as to their identity. Miers's figure does not represent quite accurately the proportions
of the carapace, the relative length, and especially the prominence of the whole frontal
region, being considerably exaggerated.
Localities. "Flinders Entrance, near ^ler " ; "Channels between reefs, Murray
Island."
Myra tug ax (Fabr.).
Myra fur/fix (Fa])r.), Alcock, .lourii. Asiatic Soc. Bengal, Ixv. [2) p. 202 (1890) (and syuouyms).
M. pentacantlia ?, Alcock, 1. c. p. 204.
Two male sp(>ciuiens, of small size and therefore difiieult to determine with certainty,
ar(^ probably young forms of this common and variable species. The smaller of the two
possesses five spines on the posterior margin of the carapace and a well-marked tubercle
BHACHYURA FROM TORRES STRAITS. 27
on the intestinal region. It agrees closely with the type specimens of M. dahia, Miers
(Proc. Zool. Soc. 1S79, p. 12), from Japan, whicli Miers suhsecpiently (Rep. ' Challenger '
Erachyura, p. 314) identitied with the A[. coaUta of Ililgeudort' (Monatsljer. Akad. Berlin,
1S78, p. 812, pi. i. figs. 6, 7), ranking it as a variety of 31. fugax. The 31. pentacautha
ol' Alcock, of which I have examined specimens, differs chiefly in the puhescence of the
frontal region. Tliis provisional species is regarded hy its anthoi- as heing prohablv
the young form of M. fugax.
Locality. " Channels between reefs, ^Murray Island."
Myra -\ustkalis, lias well (?).
Myra australis, Haswell, Proc. Limi. Soc. N. S. Wales, iv. p. 50, pi. v. tig-. ;i (1879) ; Haswell, Cat.
.^ustr. Crust, j). 122 ; Miers, Rep. ' Challeuger ' Bracliyura, p. 315.
A female specimen, 20 mm. in length, is provisionally referred to this species on
account of its general resemhlance to specimens so named in the collections of the
British IMuseum. Prom these and from Haswell's account, however, it differs in the
shorter neck-region, in the finer and closer granulation of the surface, and in the absence
of any distinct group of granu.les on the intestinal region.
Loralihi. " Channel between reefs, Mer."
Leucosia longifrons, var. pulchekrima, Miers.
Lenco.y'ni jnilclierrima, Miers, Tr. Linn. Soc. (2) Zool. i. p. 236, pi. xx.wiii. figs. 4-6 (1877); Haswell,
Proc. Linn. Soc. N. S. Wales, iv. p. 46 (1879).
L. spkndida, Haswell, 1. c. p. 47, pi. v. fig. 1.
L. loiiyifroHs, \ay. piikherrima, Alcock, Journ. Asiatic Soc. Bengal, hv. (2) p. 219 (1896).
A male specimen, agreeing minutely with Miers's type specimen, save that the
" tlioracic sinus " is rather more contracted. \\ydxi from the colour-markings, Alcock
states that this variety is distinguished from the typical L. longifrons by having the
surface of the cai apace slightly punctate instead of perfectly smooth, and by the stronger
dorsal and ventral keels on the jjropodites of the ambulatory legs. Neither of these
characters is so Avell marked in the Torres Straits specimen as in some specimens from
Yokoliama which I refer to L. longifroiis. The carapace of the present specimen is mucli
distorted, being swollen on one side, probably by the presence of an epicarid parasite in
the branchial chamber.
Locali///. " Channel between reefs, Murray Island, 15-20 fath."
Leucosia Haswelli, Miers.
Leucosia HasireUi, ]\Hers, Rep. 'Challenger' Bracliyura, p. 821, pi. .\xvii. tig. 2 ; Alcock, Journ.
Asiatic Soc. Bengal, Ixv. (2) p. 222 (1896).
Our specimen shows an almost precise agreement with the descriptions of Miers and
Alcock and with the type s))ecimens in the British Museum. The inner (or lower)
margin of the hand is defined hy two distinct rows of fine granules, the lower replacing
the row of punctations of Alcock's description.
Locality. "Pringing reef and shore, Thursday Island."
4*
28 DH. W. T. CALMAN ON A COLLECTION OF
PSEUDOPHILYRA TRIDBNTATA, Miers.
Psmdo/ihihjra tridentata, Miers, Proc. Zool. Soc. 1879, p. 11, pi. ii. fig. 4; Alcock, Journ. Asiatic Soc,
Bengal, Ixv. (2) p. 250 (1890).
Our specimen, a male, agrees closely with Miers's type specimen, which, though
described as a male, is apparently a sterile female. The inferior hepatic prominences
are in our specimen placed a little further forward, so that they are visible from above
in front of the superior prominences. The figure which accompanies Miers's description
is a very indifferent representation of the type specimen, the whole frontal region, for
instance, appearing relatively much too broad. The following are the approximate
dimensions of the type and of our specimeu : —
Type specimen. Torres Straits specimeu.
Length of carapace 10 mm. 13-5 mm.
Breadth „ 8-5 „ 11-5 „
Height „ 5-5 „ 7-0 „
Width of front between external orbital teeth ... 2'35 „ 2'75 „
Length of cheliped 210 „
„ palm 7-0 „
Width of palm 3-5 „
Length of fingers 30 „
Alcock's description applies very well to the specimen before me, except that he states
the hand to be about " half as long again as broad." As will be seen from the figures
given above, the palm alone, exclusive of the fingers, is twice as long as broad. Each
finger carries a low obtuse tooth on the inner edge about the middle of its length.
Locality. "Torres Straits."
Distribution. Persian Gulf {Alcock) ; S. Japan {Miers).
Akcania gkacilipes. Bell (?).
Arcania gracilipes, Bell, Trans. Linn. Soc. xxi. p. 310, pi. xxxiv. fig. 9 (1855) ; Alcock, Journ.
Asiatic Soc. Bengal, Ixv. (2) p. 270 (1896).
A male specimen, the carapace of which measures about 7 mm. in length and in
breadth, with chelipeds about 15 mm. long, is referred with some doubt to this species.
Compared with Bell's type specimen, it diff'ers in the greater excavation of the hepatic
regions above and the consequent greater prominence of the neck-region. The front
(between the orbits) is also more prominent at the outer corners, so that the orbits have
a more lateral position. The carapace, as a whole, appears much less inflated, and the
inter-regional grooves, especially the branchio-cardiac grooves, are deeper. The marginal
tvibercles are less prominent, and the whole surface is covered with closely-set depressed
and smooth granules. In the type the granules are more widely spaced and more or less
distinctly capitate or fungiform. Some phrases of Alcock's description, the " sunken "
hepatic region and the cju*apace " closely covered with flat discoidal granules," are more
suggestive of our specimen than of the type.
Locality. " Minders Entrance, Mer, 20-30 fath."
BRACK rUEA FROM TORRES STRAITS. 29
Genus Palicus, Philippi.
( = Cymopolia, Roux.)
The most diverse opinions have been expressed as to the systematic position of this
genus, and although it retains its place among the Dorippidoe in Bouvier's recent revision
of that family (Bull. Soc. Philomath. Paris, (8) ix. 1898), there appears to be considerable
reason to doubt the correctness of this view. Without attempting to enter on a
discussion of the question, I may note that the penial appendages of the male are (in the
single specimen 1 have examined as to tliis point) distinctly exserted from the sternum
at some distance from the bases of the last pair of legs. "With regard to the disposition
of these parts in the Dorippidse, the statements of authors are conflicting. Miers, for
instance, writes : " The sexual appendages in tlie male are exserted from the sternum "
(Rep. 'Challenger' Brachyura, p. 32G), while Ortmann has " miinnlicho (Tenitaloffmmg
stets coxal gelegen " (Bronu, Thier-Reich, Crust, ii. p. 1157). I find that in Borlppe
the latter statement is the more correct, although the penial tube lies, at its base, between
two processes of the sternum, which may in {D. sima) meet above and form a complete
ring. The greater separation of the place of emergence of the penes from the bases of
the legs in Palicus tends to support the view of those authors who would ally this genus
with the Catometopa.
The description and figures of Pleurophncus spinipes given by De Man (Arch. Xaturg.
liii. (1) p. 311, pi. XV. fig. 1, 1887) are strongly suggestive of close affinity between that
genus and the present. The general outline of the carapace, the relative length of the
four pairs of ambulatory legs, the shape of the third maxillipeds, and the very broad
sternum are among the points of i-esemblance between the two. Pleurophricus crisiatipes,
A. M.-E., the type and only other species of the genus, appears, from Milne-Edwards's
figure ( Journ. Mus. Godeffroy, Heft iv. pi. xii. fig. 6), to have less resemblance to Palicus,
the ambulatory legs being all of about the same length. De Man considers Pleui-o-
phricus to be more nearly allied to Corystoidea than to any other group of Brachyura.
Milne-Edwards had placed it among the Oxystomata, while Miers suggests that its place
is with tbe Schizophi'ysinse among the Oxyrhyncha (Journ. Linn. Soc, Zool. xiv. p. 660).
Palicus Jukesii (White). (Plate 1. figs. 9-13.)
Cymopolia Jukesii, White, Jukes's Voy. ' Fly,' ii. App. p. 338, pi. ii. tig. 1 (lSi7) ; .Miers, Zool. Voy.
'Erebus' and 'Terror,' vol. ii. no. xx. Crust, p. 3, pi. iii. figs. -1 a-c (1875) (1874?) ; Miers, Rep.
' Challenger ' Brachyura, p. 335 (1886).
C. carinipes, Paulson, Crustacea of the Red Sea (Russian), Kiev, 1875, p. 73, pi. ix. figs. 4-4 «.
Palicus Jukesii, Bouvier, Bull. Soc. Philomath. Paris, (8) ix. p. 12 (sep. copy) (1898).
Description of male. The carapace is subquadrate in outline, with the lateral margins
slightly convergent anteriorly. The surface is very uneven, being thrown into rounded
transverse ridges, of which two, crossing the cara2)ace at the level of the gastric and
cardiac regions respectively, are the most conspicuous. The prominent regions are
coarsely granulated and the whole surface is nearly free from hairs. The front is
divided into two rounded lobes defined from the orbital margin on either side by a
30 ]>K. W, T. CALMAN ON A COLLECTION OF
distinct notch. The upper margin of the orbit presents two fissures, of which the inner
is a A^-shaped notcli, while the outer is closed and inconspicuous. Tlie external orbital
tooth is blunt, and behind it on the lateral margin are two w^ell-marked teeth, with a
slight indication of a third. The two lobes of the lower orbital margin are sharply
triangular, the inner being the more prominent, and have the edges hardly or not at all
granulated. On the sub-hepatic region just behind the lower orbital margin there is a
blunt transversely elongated tubercle or short ridge.
The eye-stalk carries about three tubercles, the largest of which, close to the corneal
region on the anterior edge, is in the form of a flattened lobe with a rounded distal edge.
The basal joint of the antenna has a very prominent longitudinal keel on its ventral
surface, and externally a blunt laterally compressed lobe springs from near the base of
the joint and is directed forwards and outwards. The two succeeding joints of the
peduncle are narrow and cylindrical. The ischium of the third maxillipeds has two
marked diagonal ridges on its ventral surface. The merus is produced distally external
to the insertion of the carpus as a conspicuous rounded lobe extending to more than half
the length of the carpus.
The chelipeds in the single male specimen examined are rather feeble and are perhaps
not fully developed. The palm is subcylindrical and has faint longitudinal ridges on its
outer surface.
The ambulatory legs of the second and third pairs have the upper (or anterior)
edge of the merus cut into four teeth. The crest on the anterior margin of the carpus
has no distinct proximal lobe, but the distal lobe is a sharp tooth set a little way back
from the end of the joint. The propodus is much expanded, being three and a half times
as lono- as broad, and the anterior edge is strongly convex. In the second pair of anibu-
latorv legs (but in none of the others) there is, on the ventral surface of the merus at its
proximal end, a short longitudinal ridge, which is minutely and regularly granulated. The
abdomen of the male has all the somites free and each is crossed about the middle of its
length by a transverse ridge. The lateral margins are slightly concave and form a
distinct angle at the sixth somite. The sternum and abdomen are finely granulated.
The first abdominal appendages of the male are stout and the two lobes of the tip are
closelv approximated, the outer extending a little beyond the inner.
The specimen from which my description and figures are taken agrees minutely with
the specimens in the British Museum with which I have compared it. According to
Miers's figure, the abdomen of the female is subcircular in outline, with all the somites
distinct and transversely ridged as in the male.
CuviopoJia varinipes of Paulson is very likely identical with the present species. His
fio-iu-es show the general shape of the carapace to be very similar, though the lateral
margins arc more nearly parallel. The transverse grooves and ridges of the surface and
the granulation of the more prominent parts correspond Avith the sj^ecimen here
described and figured. Paulson's figure of the entire animal (/. c. fig. 4) is, ap^mreutly,
inaccurate as regards the shape of the frontal lobes, which his enlarged figure (fig. 4 a)
shows to differ but sliglitly from the present form. The outer of the two fissures in the
BRACllYU'KA FROM TORRKS STRAITS. 31
supra-orbital margin is represented as open and V-shaped. The lower orl)ital margin, the
tubei'cles on the eye-stalk, the basal joint of the antenna, and the external niaxillipeds
are all figured almost exactly as in our specimen. The tirst pair of walking-h^gs are
rather stouter. The most marked ditference, however, is that the tubercle on the under-
side of the hepatic region is more strongly developed, being represented by a curved
transverse ridge, fmni the outer end of wliich a row of granules (not found in our
specimen) runs backwards for a short distance pai-aliel to the lateral margin of the
carapace.
Locality. "Torres Straits."
DistrihHtioH. Sir C. Hardy Island (Torres Straits) [Jlli'de); Port Denison {I laswell) \
Celebes Sea (Jliers) ; Red Sea {Panlmii).
Palicus Wiiitei (Miers). (Plate 2. tigs, lt-19.)
Cymupolia IVhitei, Miers, Rep. Voy. 'Alert,' Crust, p. 551, pi. xlix. fig. C (1881).
Palicus IVhitei, Bouvicr, Bull. Soc. Philomath. Paris, (8) ix. p. 12 (sep. copy) (Ih'JS).
The lateral margins of the cai-apace are more nearly parallel than in /'. Jukesii.
The surface is less uneven, the transverse ridges being less marked, but the regions are
fairly prominent and well-defined, though there is some variation in these respects
among our siiecimens. The granulation of the surface is much finer and there is a
scanty short pubescence interspersed among the granules. The frontal lobes are
separated from the upper orbital margin on each side by a shallow concavity. Both the
fissures of the upper orbital margin are open and V-shaped, whih; a third, present in
some members of the genus, is slightly indicated by a notch at the base of the outer
orbital tooth. This tooth is generally more acute than in P. Jnkesii, but the two
succeeding teeth on the latex'al margin are less prominent than in that species. The
lobes of the lower margin of the orbit are low and rounded, and the edge is finely
granulated. There is no tubercle on the sulvhepatic region behind the orbital margin.
The basal joint of the antenna bears a slight longitudinal ridge on its ventral face and is
produced externally into a broad rounded lobe, flattened dorso-ventrally and projecting
into the orbit. The two succeeding joints are robust, the third joint especially being-
expanded and compressed.
The large prominence on the eye-stalk has a peculiar and characteristic form. It is a
crescentic or sickle-shaped blade, springing from a narrow base near the distal end of
the anterior edge of the eye-stalk, and curving over, close to but free from the corneal
surface, terminating externally in an acute point.
The ischium of the third maxilUpeds is only faintly ridged on its ventral face, and the
antero-external process of the merus is very small, not reaching to one-half the length
of the carpus. The chelipeds are feeble in both sexes, the palm cylindrical, without
ridges, but faintly granular and pubescent. The second and third pairs of walking-legs
have the merus pubescent and faintly granulated, but without teeth on the margins.
The anterior crest of the carpus has rounded proximal and distal elevatioirs, the latter
close to the end of the joint. The propodus is considerably narrower than in L\ Jnkesu,
32 DR. W. T. CALMAN ON A COLLECTION OF
being nearly four times as long as broad, and tbe anterior edge is only slightly convex.
The granulated ridge on the underside of the merus of the second pair is present as in
P. JuJcesii.
The abdomen of both sexes is smooth, beset with small scattered setae. The fourth,
fifth, and sixth somites are fused together, but the sutures are faintly visible. In the
male the sides of the abdomen are straight, convergent, and curve gently inwards from
the base of the last somite to the tip without any distinct angle.
The first abdominal appendages of the male are slender, and the terminal lobes are
divergent, the outer lobe being twice as long as the inner.
Our specimens differ from Miers's type specimens in the greater robustness of tlic
second and third pairs of ambulatory legs, the merus and, to a less extent, the propodus
being distinctly broader. In all other characters, however, the agreement is
complete.
Locality. " Torres Straits.''
Dislrihtitioii. Seychelles, 4-12 fath. {Miers).
Palicus serripes (Alcock & Anderson). (Plate 2. figs. 20-22.)
Cymopolia serripes, Alcock & Anderson, Journ. Asiatic Soc. Bengal, Ixiii. pt. 2 (1895) (? 1894), p. 208 ;
Illustr. Zool. ' Investigator,' Crust, pi. xxiv. fig. 7 (1896).
Palicus serripes, Bouvier, Bull. Soc. Philomath. Paris, (8) ix. p. 12 (sep. copy) (1898).
Carapace with lateral margins convergent anteriorly, the surface leaving the I'egional
areas well-defined Init not very prominent, beset with rather coarse granulations on the
more prominent parts. Posteriorly the granulations tend to l)ecome squamiform, and
there is a line of conspicuous scale-like elevations just within the posterior and postero-
lateral margins. Scattered hairs occur among the granules, more numerous on the
depressed portions of the surface. The front is four-lobed ; the inner lobes are acutely
rounded and depressed, and extend beyond the outer lobes, which are low, rounded,
slightly recurved, and hardly defined from the orbital margin. The upper ruargin of the
orbit has two deep V-shaped fissures separated by an acute tootli, and a shallower notch
at the base of the outer orbital tooth. The latter is acute, and is separated by a short
interval from the first of the four subequal antero-latei'al teeth. The lower margin of
the orbit is convex in its outer part, concave internally, and terminates in a shai-p
internal orbital tooth. Just behind and parallel to the infra-orbital margin is a curved
granulated ridge terminating internally in a small tubercle close to the tubercle which
forms the anterior corner of the buccal frame. The basal joint of the antenna is
pi'oduced extei-nally into a flattened lobe, similar to, but smaller than, that found in
P. TVhitei, while the ventral face of the joint bears a row of three or four small tubercles
in place of the longitudinal ridge found in that species. The two succeeding joints are
rather elongated and cylindrical. The flagellum extends well beyond the outer angle of
the orbit. The eye-stalk bears numerous, low, rounded tubercles, two of these on the
anterior margin being larger than the others. The ischium of the third maxillipeds is
nearly smooth on its ventral surface, and the merus has a well-developed antero-external
BEACHYURA FEOM TORRES STRAITS. 33
process with a small acute tooth at its tip. The exopod is broader than in the two
preceding sjiecies. The cheliped of our single imperfect specimen is rather short and
stout, the ])alm comjiressed, with rows of granules on its upper edge and outer face.
The first pair of legs liave the merus granulated and with a stout tooth at the distal
extremity of its u^^per (or anterior) eJgc. The propodus and dactylus have each a few
serrations on the lower edge. The merus of the second and tliird pairs is granulated,
the granules becoming stout teeth on the upper and lower margins. The anterior crest
of the carpus has two 2)rominent teeth, with smaller serrations between. The propodus
is about two and a half times as long as broad, the upjjcr edge nearly straight, the lower
edge serrate. The dactylus is broad and has three or four coarse and somewhat irregular
teeth on its lower edge. There is no granulated ridge on the vmderside of the merus
of the second pair.
The single, much injured, female specimen from which the above description is taken
agrees well with the description of Alcock and Anderson. The figure whicli they give,
however (taken from a larger specimen), diflFers in some details. The outer lobes of the
front are much less prominent, so that the front appears two-lobed ; the first lateral
tooth follows close upon the extra-orbital and is separated by a slight interval from the
second ; the granulation of the carapace appears to be less coarse, and mor(> restricted
to the prominent lobules.
Localilij. " Torres Straits."
Distribution. " Madras coast, sliallow " [Alcock ^' Aiulevson).
ACH.33US AFPiNis, Miers.
Acheeus affinis,W\ers, Kep. Voy. 'Alert,' Crust, p. 188; De Mau, Arch. Naturg. liii. (1) p. 218
(1887) ; Alcock, .Tourn. Asiatic Soc. Bengal, Ixiv. (2) p. 172 (1895).
A male specimen 55 mm. long is referred to this species. It is much smaller than
any of the specimens of A. affinis with which I have compared it, i)ut it agrees with
them in the characteristic points of the tuberculated eye-stalks and tlie bilobed cardiac
tubercle. As De Man points out, the eye-stalk carries a small tul)ercle near the tip in
addition to the large one at the middle of its length, and a small tubercle also lies behind
the bilobed eminence on the cardiac region. Our specimen has four granules on the
gastric region, two median and two lateral. The free part of the antenna is only a little
shorter than the carapace. The neck is rather longer than in tlie type specimen, and the
carapace as a w^hole is a little narrower. As in Miers's account, the merus of the cheUpeds
is " somewliat trigonous," rather than " fast cylindrisch " as De Man describes it.
Locality. " Channel between reefs, Murray Island, 15-20 fath."
Paratymolus sexspinosus, Miers.
Paratymolus sexspinoms, Miei-s, Rep. Voy. ' Alert,' Crast. p. 2G1, pi. sxvii. tig. B ; Heudersoa, Traus.
Linn. Soc., (2) Zool. v. p. 2,7)2 (1893).
In our single perfect specimen, which I have compared with Miers's type, the rostrum
SECOMD SERIES. — ZOOLOGY, VOL. VIII. 5
34. DE. W. T. CALM AN ON A COLLECTION OV
is hardly cmarginate and the second antero-lateral tooth is less prominent than in Micrs's
flgui'e.
This sjiecies is recorded from India (Tuticoriu) by Prof. Henderson, though it is
omitted (probably through inadvertence) from Alcock's revision of the Indian
Oxyrhyncha.
Localities. " Channel between reefs, Mer " ; " Mabuiag " (fragmentary specimen).
Oncinopus aranea, De Haan.
Inachus [Oncinopus) aranea, De Haau, Faun. Japon., Crust, p. 100, pi. xxxix. fig 2.
Oncinopus aranea, Adams & White, Zool. Voy. ' Samarang,' Crust, p. 3.
0. neptunus, Adams & Wliite, /. c. p. 1, pi. ii. fig. 1.
0. aranea, Alcock, Journ. Asiatic Soc. Bengal, Ixiv. (2) p. 183 (1895).
Five female specimens, showing considerable variation in the relative prominence of
the frontal lobes and in some other details. Alcock describes the antennte as "extremely
short, reaching only just beyond the tip of the rostrum " ; in our specimens, however,
as in the figure of Adams & White, the flagella of the antennae reach far beyond the tip
of the rostrum.
Locality. " Channel between reefs, Mer."
Xenocarcintjs tuberctjlatus, White.
Xenocarcinus tuber culatus.i White, Proc. Zool. Soc. 1847, p. 119; A. Milne-Edwards, N. Arch. Mus.
Paris, viii. p. 253, pi. xii. figs. 1 a-g (1872) ; Alcock, Journ. Asiatic Soc. Bengal^ Ixiv. (2) p. 192
(1895) ; Alcock & Anderson, Illust. Zool. ' Investigator,' Crust, pi. xxxiii. figs. 3, 3 a (1898).
A single, much damaged, male specimen, agreeing fairly well with Milne-Edwards's
fig. 1, save that the carapace is much narrower.
Locality. " Murray Island."
HuENiA PROTEUS, De Haan.
Maia [Huenia) proteus, De Haan, Fauna Japon., Crust, p. 95, pi. sxiii. figs. 4-6.
Hueniaproteus, Alcock, Journ. Asiatic Soc. Bengal, Ixiv. (2) p. 195 (1895).
The series of this well-knowji species comprises two young males, an adult and a
young female, and a " sterile " female in which only the 5th and 6th somites of the
abdomen are fused.
Localities. " Channel between reefs, Mer"; " South of Orman's reef, 5-7 fath."
Menaethitjs monoceros, Latr.
MenaeUiius monoceros, Latr., Milue-Edwards, Hist. Nat. Crust, i. p. 339.
M. anyustus, Dana, U.S. Expl. Exp., Crust, i. p. 120, pi. vf. figs. 5 a-b.
M. monoceros, Alcock, Journ. Asiatic Soc. Bengal, Ixiv. (2) p. 197 (1895).
Two specimens of this very variable species approach most closely to the form named
M. cmgustus by Dana, but the rostrum is rather shorter.
Localities. " Murray Island " ; " Sabai Channel."
BRACHYUliA FKOM TOEEES STEAITS. 35
Hyastenus spinosus, a. Milue-Edwards.
Hyastenus spinosus, A. Milne-Edwards, N. Arch. Mus. Paris, viii. p. 250 (1872) ; Miers, Kep.
' Challenger' Brachyura, p. 5G ; Alcock, Journ. Asiatic Soc. Bengal, Ixiv. (2) ]). 211 (1895).
A female specimen, about 25 mm. in total length, agrees witli the characters of this
species as briefly indicated by Milne-Edwards and by Alcock, except that the lateral
epibranchial spines apjiear rather small. It is identical with specimens in the British
Museimi referred to this species by Mr. Miers. Au ovigerous female 20 mm. long
diflfers in possessing a small acute granule on the cardiac region and three small tubercles
(besides the epibranchial sj)ine) on each branchial region. Tlie two gastric spines are
very small, but in other respects the specimen appears to agree with this species.
Localities. "Murray Island"; " South of Orman's reef, 5-7 fatb."
Hyastenus oryx, A. Milne-Edwards.
Hyastenus oryx, A. Milne-Edwards, N. Arch. Mus. Paris, viii. p. 250, pi. xiv. tig. l (1872) ; De Man,
Arch. Naturg. liii. p. 224, pi. vii. fig. 2 (1887) ; Alcock, Journ. Asiatic Soc. Bengal, Ixiv. (2)
p. 214 (1895).
Of the two specimens which I refer to this species, one, a female about 16 mm. in
total length, agrees closely with the example figured by De Man, the rostral spines
being rather less than one-fourth of the total length and strongly divergent. In a male
specimen about 18 mm. long the rostral spines are more than one-third of the total length
and quite parallel, and the whole carapace is rather narrower and not quite so much
contracted in front, though still much more so than in Milne-Edwards's figure. The
parallel rostral spines of the latter specimen give it a certain resemblance to H. Sehcc,
White (Seba, Thesaurus, iii. pi. xviii. tig. 12), in which, however, the distal parts of the
much longer spines are divergent. In the specimens of M. Sebcc which I have examined
the carapace is much less tuberculated than in the present specimens. The chelipeds of
our male specimen resemble Milne-Edwards's figure, but the hands are rather less
expanded distally, and the serrated edges of the fingers meet for about half their length
instead of only at the tip as Alcock states.
Locality. " South of Orman's Reef, 5-7 fath."
Hyastenus convexus, Miers.
Hyastenus convexus, Miers, Eep. Voy. 'Alert,' Crust, p. 19fi, pi. xviii. fig. B; Henderson, Trans.
Linn. Soc, (2) Zool. v. p. 344 (1893).
A male specimen of 10 mm. and a female of 13 mm. total length are referred with
some douljt to Miers's species, with which they agree in the very convex gastric region and
in the absence of spines from the carapace. They differ chiefly in the shorter rostral
spines, which in the female are less than one-fourth of the total length. In the male
there are minute acute tubercles representing the epibranchial spines as iu the specimen
described by Henderson.
LocaUtii. " Mer."
36 DE. AV. T. OILMAN ON A COLLECTION OF
Hyastenus Brockii, Dc Man.
Hyastenus Brockii, De Man, Arch. Naturg. liii. (1) 1887, p. 221, pi. vii. figs. 1 a-6; Henderson,
Trans. Liun. Soc, (2) Zool. v. p. 344 (1893).
A single very imperfect dried specimen ( <s ) appears to agree with De Man's description
and figures of this species. The total length is about 15 mm., half of which is taken up
by the slender rostral spines. The carapace is narrower and the rostral spines less
widely divergent than in the figure. The tubercles on the carapace are only very slightly
marked, but the arrangement, so far as can be seen, corresponds with De Man's account.
The structure of the orbital region agrees closely with the figure.
Locality. " Toi-res Straits."
Distribution. Amboina {De 3Ian) ; Gulf of Martaban {Henderson).
Hyastenus vekrucosipes (Adams & White). (Plate 2. figs. 23 & 24.)
Chorinus verrucosipes, Adams & White, Zool. Voy. ' Samarang,' Crust, p. 13, pi. ii. fig. 3 (1848).
Hyastenus verrucosipes, A. Milne-Edwards, N. Arch. Mus. Paris, viii. p. 250 (1852) (name only).
Paramithrax verrucosipes, Miers, Ann. Mag. Nat. Hist. (5) iv. 1879, p. 10 (name only).
This species, which does not ajipear to have been reobserved since its discovery, is
represented in the collection by two female specimens agreeing so closely with the
original type specimen of Adams and White as to leave no doubt of their identity. The
surface of the carapace is rather more uneven than is represented in the figure given by
these authors, and it is slightly more contracted behind the orbital region, so that the
postorbital processes appear more prominent. The ujoper hiatus of the orbit is rather
more open in our specimens than in the type, and the rostral spines are distinctly
knobbed at the tip. The first j)air of Avalking-legs are relatively longer than in the
figure, and the dactyls, especially of the last pair, are longer and more slender. The legs
in our specimens do not present the "wart-like tubercles" described by Adams and
White ; but I am inclined to think, after examining the now somewhat imperfect type
specimen, that these tubercles are simply tufts of hair agglutinated together by drying.
As regards the systematic jwsition of this species, Prof. A. Milne-Edwards doubtfully
includes it in his list of the species of Hijastenus, while Miers incidentally refers to it as
a Parcmiithrax. The type specimen in the British Museum is now labelled Acanthophrys
verrticosij)es, and I am informed that this name was given to it by Mr. Miers.
It seems plain, however, that this species cannot be referred to Farctmitlirax, since
there is only one hiatus instead of two in the upper margin of the orbit, or, in other
words, the spine which, in that genus, intervenes between the supra-ocular hood and the
postorbital process is here wanting. The condition of the orbital region is most closely
paralleled by Dana's figures of his Lahaina ovata (U.S. Expl. Exp., Crust, i. p. 93, pi. ii.
figs. 1 a-c). The svipra-ocular hood is very prominent, having the corners produced and
the anterior one acute and curved forwards ; the long postocular process is cupped, or
rather grooved, along its anterior face for the reception of the eye ; the basal antennal
joint carries a small spine distally on the ventral surface, and two smaller tubercles in a
BRACHTUEA FEOM TORRES STRAITS. 37
line immediately Leliind it, while a rectangular plate projectiag from the outer edge of
the joint forms the floor of the orbit. Faii-ly wide gaps are left between the postocular
process and the supra-ocular hood and basal antennal plate in the upper and lower walls
of the orbit respectively. Dana's genus is generally regarded as a synoaym of Hj/astcuus,
and I have accordingly followed Milue-Edwards in adopting that name for the present
form, though it differs considerably in the structure of the orbit from those species of
Hifcistemis which I have examined. Prom Acanthuplirijs it appears to differ in the fact
that the inner distal corner of the merus of the third maxillipeds is notched, and the
structure of the orbital region is very different from that shown in Milne-Edwards's
figure of A. cristimanus (Ann. Soc. Ent. France, (Jj) iv. 18(55, \)\. v. fig. 3 a).
Localiti/. " Murray Island, channel l)etween reefs, 15-20 fath."
Distribution. " Eastern seas " {Adams 8f JF/iite).
Naxia SERPiTLiFEKA (Guerin).
Nuxia serpulifera, Milne-Edwards, Hist. Nat. Crust, i. p. 313; Haswell, Cat. Austr. Crust, p. 21
(1882) ; Miers, Hep. Voy. 'Alert/ Crust, p. 19G (1881) ; Pocuck, Aun. Mag. Nat. Hist. (G) v.
p. 79 (1890) ; Ortmann, in Semon's Zcol. Forsch. Austral, v., Crust, p. 43 (189-1).
A large female specimen (83 mm. in length) of this characteristic Australian species
is in the collection.
Localiti/. " Torres Straits."
Naxia taurtjs, Pocock.
Naxia tavrus, Pocock, Ann. Mag. Nat. Hist. (G) v. p. 70 (1890); Alcock, Journ. Asiatic Soc. Bengal,
Ixiv. (2) p. 219 (1895) ; Alcock & Audersou, Illustr. Zool. ' Investigator/ Crust, pi. xxxiii.
figs. 5-5 a (1898).
N. cerastes, Ortmann, Semou's Zool. Forsch. Austral., v. Crust, p. 43, pi. iii. fig. 4 (1894) ; Alcock,
Journ. Asiatic Soc. Bengal, Ixiv. (2) p. 220 (1895) ; Alcock & Auderson, Illustr. Zool.
'Investigator,' Crust, pi. xxxiii. figs. 2-2 « (1898).
Two male specimens, differing from Pocock's type in the much shorter rostral spines
—about G-5 mm. in a total length of 17 mm., — with the accessory spinules well in front
of the middle of their length. The meral spines, large and conspicuous on the first pair
of walking-legs, are all but obsolete on the succeeding pairs. In other respects these
individuals do not differ materially from the type specimen. Of the two figures given
by Alcock and Anderson our specimens resemble most that named N. cerastes, Avithout
agreeing precisely with either. It can hardly be doubted, however, that, as Alcock lias
suggested, N'. cerastes is merely a variety of N. tauriis.
Locality. "Channels between reefs, Mer, 15-20 fath."
Tylocaecinus STYX (Herbst).
Pha siyx, ]\Iilne-Ed\vards, Hist. Nat. Crust, i. p. 308.
Microphrys stijx, A. Milue-Edwards, N. Arch. Mus. Paris, viii. p. 247, pi. xi. fig. 4 (1872).
Tylocarcirms styx, Miers, Ann. Mag. Nat. Hist. (5) iv. p. 14(1879); Alcock, Journ. Asiatic Soc.
Bengal, Ixiv. (2) p. 235 (1895).
38 DB. W. T CALMAN ON A COLLECTION OF
A single female specimen. The carapace is a little narrower than in Milne-Edwards's
figure, but in other respects the specimen agrees perfectly with this and with Alcock's
description.
Locality. " Murray Island, reef."
Distribution. Eed Sea to Fiji.
Paramithrax (Chlorinoides) Coppingeri, Haswell.
AcantJiopfirys acnJeatus, A. Milne-Edwards, Ann. Soc. Ent. France, (4) v. p. 140, pi. iv. fig. 4 (1865)
(not Cliorimis aculeatua, M.-E. Hist. Nat. Crnst. i. p. 316).
Paramitlirax Coppingeri, Haswell. Proc. Linn. Soc. N. S. Wales, vi. p. 750 (1881) *; Haswell, Cat.
Austr. Crust, p. 15.
Paramithrax (C/iloriiioides) Coppingeri, Miers, Rep. Voy. ' Alert/ p. 192.
Ctilorinoides Coppingeri, Miers, Rep. ' Challenger ' Brachyura, p. 53, pi. vii. fig. 3.
A female specimen, about 6"5 mm. in total length, differs from Miers's figure and from
specimens in the British Museum in having the rostral spines short (about one-fourth
the length of carapace) and deflexed, and in having the supra-ocular hood deeply cut into
three spiniform teeth, the middle one being reflexed at the tip. Miers's figure shows it
as obscurely divided into two lobes, but Haswell's original description reads : " upper
orbital border with three straight, acute, spinous teeth." Only one spine is present
behind the double spines on the cardiac region, as in Miers's description and figure, but
a small tiibei'cle represents the second spine which Haswell describes.
The shape of the supra-orbital border in our specimen is very like that shown in
Milne-Edwards's figvire of Acanthophrys aculeatus, which in other respects resembles
so closely Miers's figure of P. Coppingeri as to leave little room for the doubt which
Miers appears to have had as to the identity of the two species. Haswell's name for
the sj)ecies, however, still holds good, since that employed by Milne-Edwards is pre-
occujned by the next-mentioned species.
Locality. " Torres Straits."
Paramithrax (Chlorinoibes) acui,eatus (Milne-Edwards).
Cliorinus aculeatus, Milne-Edwards, Hist. Nat. Crust, i. p. 316.
Paramithrax {CIdorinoides) aculeatus, var. arinatus, Miers, Rep. Voy. 'Alert,' p. 193, ]}\. xviii. fig. A.
Clilorinoidts aculeatus, Miers, Rep. ' Challenger ' Briichyura, p. 53 ; Henderson f, Tr. Linn. Soc.,
(2) Zool. V. p. 345 (1893).
Paramithrax {Ctilorinoides) aculeatus, Alcock, Jouru. Asiatic Soc. Bengal, l.xiv. (2) p. 241 (1895).
A small female specimen appears to find its place among the variations of this species.
There are, as usual, five spines on the mid-dorsal line of the carapace, but the spine which
in the normal type occu2)ies the middle of the posterior margin appears to be wanting,
* This reference is given wrongly by Miers in the ' Challenger ' Report.
t Henderson's remarks on Miers's variety armatns are based on an oversight of the fact that the figure in
De Haan's great work, to which he refers, does not represent P. aculeatus, though so named on the plate, but
De Haan's species P. loiu/injiinus.
BEACHTUEA FEOlVr TOREES STRAITS. 39
tlie number being made up by the intercalation of a small spine just in front of the lar'>-e
one on the cardiac region. All the spines of the carapace are distinctly knobl)ed at the
tip. The supra-ocular hood is deeply divided into two teeth, of which the anterior is acute
and turned upwards and forwards while the posterior is truncate. The nieriis of the
ambulatory legs bears two spiniform tubercles distally. The rostral spines are strongly
divergent and hardly more than one-third of the length of the carapace.
Locality. " Torres Straits."
ScHizoPHRYS ASPERA (Milnc-Edwards).
Mithrax asper, Mihie-Edwards, Hist. Nat. Crust, i. p. .320.
Schizophrys uspera, A. Milne-Ed wards, N. Arch. Mus. Paris, viii. p. 231, pi. x. fig. 1 (1872) ; Aleock
Jourii. Asiatic Soe. Bengal, Ixiv. (2) p. 243 (18<J5) ; Aleock & Anderson, Illustr. Zool. ' Investi-
gator,' Crust, pi. XXXV. figs. 1, 1 a (1898).
One female and three male specimens, belonging to the typical form of this species as
described by A. ;Milne-Edwards, having only one accessory spiuule on each of the rostral
horns. The largest male, 2G mm. in length, is still immature, the chelipeds beiu"- no
longer than the succeeding legs. In the other two males, 17 and 12 mm. lon*^^ respectively
the carapace is relatively narrower, and in the still smaller female the distance between
the extra-orbital spines is hardly less than the greatest width of the carapace. In the
small specimens the surface of the carapace between the large tubercles is quite smooth.
Locality. "Murray Island."
Cyclax suborbicularis (Stimpson).
Cyclomaia maryaritatu, A. Milne-Edwards, N. Arch. Mus. Paris, viii. p. 23G, pi. x. figs. 2-3 (1872).
Cyclax suborbicularis, Aleock, Journ. Asiatic Soc. Bengal, Ixiv. (2) p. 215 (1895).
An ovigerous female, 24 mm. in length.
Locality. " Mvu'ray Island."
Distribution,. Red Sea to New Caledonia.
PsETJDOMiciPPA VARIANS, Micrs. (Plate 2. figs. 25 & 26.)
Pseudomicippe? varians, Miers, Ann. Mag. Nat. Hist. (5) iv. p. 12, pi. iv. fig. 8 (1879).
Pseudomicipjja ? varians, Miers, Rep. Voy. •' Alert,' Crust, p. 197 ; Miers, Rep. ' Challenger '
Braehyura, p. 68 ; Ortmanu, in Seinon's Forseh. Austr., Crust, p. 40.
Three male specimens, agreeing with Miers's types of this briefly described species, and
showing the characters assigned by him to the male sex, the gastric region being but
little elevated and the rostral spines not perceptibly deflexed.
The only point indicated by INir. Miers as distinguishing this species from the P. tenuipes
of A. Milne-Edwards (Ann. Soc. Ent. France, (4) v. p. 139, pi. v. figs. 2, 2 a, 1865) is the
character of the sternal surface, which in the last-named sj)ecies is stated to be
" remarquable par I'existence a la ligne de jonction de chaque anneau de cretes saillantes
et legerement granuleuses." In addition to this, how^ever, certain small differences in
40 DE. W. T. CALMAN 0^' A COLLECTION OF
the arrangement of the tuhercles on the carapace are apparent on comparing our
specimens with Milne-Edwards's account. He states that the gastric region carries a
median row of five tuhercles flanked hy two lateral pairs, but in the figure only four
median tubercles are distinctly seen, and this is the case also in our specimens. Milne-
Edwards further describes the cardiac region as " marquee de quatre petits tuberculcs
places de cliaque cote de la ligne mediane,'' but his figure shows two median and two
lateral tubercles. In our specimens there is only a bilobed median tubercle in the centre
of the cardiac area. Between the main tubercles, and more especially on the branchial
regions, the surface of the carapace is in our specimens rather uneven. The rostral spines
are stated by Milne-Edwards to be cylindrical : the figure shows them as rather broad
and apparently somewhat flattened and having the inner edge obtusely angled about the
middle of its length. In the Torres Straits specimens these spines are more slender and
cylindrical, tapering only very slightly to the bluntly rouaded tip. In the structure of
the orbital region, and in the presence of a large tubercle al)ove the orbit external to the
base of each rostral spine, our specimens agree closely with Milne-Edwards's species. In
spite of the differences above enumerated, the general resemblance between the two
species is so consideraljle that some doubt must still remain as to their distinctness.
Mr. Miers has indicated a doubt as to whether the present species is correctly referred
to the genus Fseudomicippa. In the type species, P. nodosa, Heller (SB. Akad. Wien,
xliii. (1) p. 301, pi. i. fig. 3), the rostrum is very strongly defiexed and the anterior angle
of the orbit is produced into a long spine, while the basal antennal joint is shaped rather
diflferently, its distal tooth being directed obliquely forwards instead of outwards as in the
present form.
As regards the systematic position of the genus, Ortmaun points out that it has been
wrongly placed among the Maiidje and has no affinity with Micippa. He would place it
among the Inachidce, either in the subfamily Iiiachincc or the Stenocinopina;. Miers,
followed by Ortmann, had suggested that the form briefly described by Haswell as the
tyj)e of a new genus under the name of Microhalimiis dejlexlfrous (Proc. Linn. Soc.
N. S. Wales, iv. p. 435, pi. xxv. fig. 2, 1879) might be identical with the present species.
Alcock, in his classification of the Oxyrhyncha (Journ. Asiatic Soc. Bengal, ixiv. (2)
pp. 164 &, 166), retains Ilicrohalimns and Pseudomicippa as distmct genera, j)lacmg the
former in the "Alliance Inachoida" of his sid)family Inachincc, while classing the latter
in the " Alliance Stenocionopoida " of the subfamily Mailnce. Alcock does not discuss
these genera further, nor does he indicate to which of them the present species should be
referred. It seems likely that P. varians should be generically separated from P. nodosa,
and that its most natural position is that which Alcock assigns to 3IicrohaUmits.
Haswell's figure shows, however, that 31. dcfiexifrons is at least specifically distinct.
Micippa philyea (Herbst),
MicipiJa masrari'iiica (Koi-sm.), Mieis, Aim. Mag. Nat. Hist. xv. p. 7 (1885).
M.phihjra (Hbst.), Alcock, Joiini. Asiatic Soc. Bengal, Ixiv. (2) p. 249 (1895).
Three specimens agree very closely with Haswell's description and figures of his
BRACHYURA FROM TORRES STRAITS. 41
M. superciliosa from Torres States (Pr. Linn. Soc. N. S. Wales, iv. p. 446, pi. xxvi. fig. 2,
1879), which is ranked by Alcock as a synonym of this variable and widely distributed
species.
Localities. " Channel between reefs, Mer " ; "Reef, Wyer."
TiARiNiA ANGUSTA, Dana.
Tiarinia angusta, Dana^ U.S. Expl. Exp., Crust, i. p. 11.3, pi. iii. figs. 7 a-b.
T. spinirostris, Hasncll, Cat. Austr. Crust, p. 28 ; Ortmann, Zool. .Talirl). Syst. vii. p. 02 (189.3).
T. angusta, De Man, Zool. Jabrb. Syst. viii. p. 492, pi. xii. fig. 2 (1895).
Our three specimens (2 s , 1 2 ) agree closely with Haswell's description of his
T. spinirostris (which De Man regards as synonymous witli Dana's species), save that in
the female specimen there are four spines instead of three on the outer margin of each of
the rostral horns. De Man figures a specimen with four spines on one side and three on
the other, and he describes also a female specimen witli only two spines on each side.
The carapace of our largest (femalej specimen measures 45 mm. in length by 27 mm. in
breadtli. De Man's largest specimen was 27 mm. long and 14-5 mm. wide. Dana's
specimen was only 6 lines long and 2\ lines wide. Haswell gives no dimensions.
Locality. " Murray Island."
Lambrus (Aulacolambrtjs) hoplonotus, Adams & White, var. ?
Lambnis hojAonutus, Adams & Wbite, Voy. ' Samaraug,' Crust, p. 35, pi. vii. fig. 3; Miers, Aun. Mag.
Nat. Hist. (5) iv. p. 22 (1879) ; Miers, Rep. Voy. ' Challeuger,' Bracbyura, p. 98 ; Alcock,
Journ. Asiatic Soc. Bengal, Ixiv. (2) p. 273 (1895).
A male specimen, 9 mm. in length, is probably to be referred to " this protean species,"
as Miers calls it. It differs considerably from the figm*e of Adams and White, the whole
anterior part of the carapace being much narrower and more produced, and the rostrum
projecting well in front of the orbits. There is a slight jiostorbital constriction, followed
by a sharp hepatic tooth, which is longer than the succeeding spiniform teeth of the
lateral margins. The granules on the surface of the carajjace are acute and spiniform.
Our specimen agrees pretty closely with some referred by Mr. Miers to his variety
longioculis, which " is best distinguishable by the subspiniform tubercles of the branchial
regions" ('Challenger' Bracbyura, p. 99), the greater jjrotrusion of the eyes being, I
believe, quite an accidental circumstance.
Locality. " Torres Straits."
Lambrus (Parthenolambrus) calappoides, Adams & White.
Parthenope calappoides, Adams & White, Re]). Voy. ' Samaraug,' Crust, p. 3-1, pi. v. fig. 5.
Lambrus [Parthenolambrus) calappoides, Alcock, Journ. Asiatic Soc. Bengal, Ixiv. (2) p. 275 (1895).
Our single specimen, a male, is only about 7"5 mm. in length, and the jjoiuts of
difference from the figure of Adams and AVhite are probably due to its small size. The
surface is rather smoother, the incision marking off the hepatic prominence posteriorly is
hardly perceptible, and the posterior edge of the carapace is more regularly crenated.
Locality. " Channel between reefs, Mer."
SECOND SERIES. — ZOOLOGY, VOL. VIII. 6
42 l>li. W. T. CALMAN ON A COLLECTION OF
Lambhus (Pakthenolambkus) confragosus, n. sp. (Plate 3. figs. 27 & 28.)
Bescriptioii of female. Carapace roughly triangular in outline, about as long as broad.
Rostrum truncated and considerably deflexed. Hepatic regions projecting in tbe form
of a vertically compressed, narrow, bluntly pointed tooth. In front this tooth is defined
by a deep notch separating it from the postorbital angle, and behind a shallower excava-
tion marks it off from the convex, crenated, lateral margin. Tlie postero-lateral corners
are produced into short, blunt spines. The posterior margin is irregularly toothed, with
a marked excavation on either side a little way from the outer end and an obtuse median
angle, which projects well beyond the line joining the postero-lateral corners. The
surface of the carapace is very uneven and irregularly granulated and pitted. The
gastric and cardiac regions rise each into a short, blunt spine, and a smaller spine or
large tubercle lies between the cardiac spine and the posterior margin. The branchial
regions are inflated, topped with oblique crests of granules, and separated from the
median regions of the carapace by well-marked grooves, which sink into deep fossae on
either side of the gastric and cardiac regions.
The chelipeds are very heavy, slightly unequal, and aliout one and a half times the
length of the carapace. The merus is irregularly toothed along its anterior and posterior
borders. The hand bears on its outer (or upper) margin two thin, rounded, cristiform
lobes, of which the larger is close to the proximal end. The upper surface carries an
oblique and irregularly toothed crest, and is elsewhere unevenly granulated. The fingers
are rather less than half the length of the palm, and the dactylus is toothed and granulated
on its outer (or upper) margin near the base. The ambulatory legs are much compressed,
with the upper and lower margins of the joints serrate.
Length of carapace 15 mm.
A second specimen, only 8 mm. long, probably belongs to the same species. The
carapace is a little longer than broad, the rostrum is tridentate and even more strongly
deflexed than in the specimen above described. The hepatic prominences, though very
well-marked, are not so much compressed and dentiform. The surface of the carapace
is smoother, and the crenatious of the lateral margins less numerous. The chelipeds
are rather more slender, and there is only one cristiform lobe on the outer edge near
the base.
The species described above, Avhich I suppose to be ncAV, resembles in general form the
L. tarpeius of Adams and White, but differs from it in the much more rugged surface of
the carapace, in the compressed dentiform shape of the hepatic prominences, and in the
armature of the chelipeds, which in the last-named species lack the flattened cristiform
lobes on the outer margin of the hand. Many species, however, of this extensive genus
are known to vary within wide limits, and it is possible that a larger series of specimens
than has been accessible to me would unite the present form with one or other of the
species already described.
LocuUUj. " Chanuel between reefs, Mer."
BEACHTUEA FEO:\I TOEEES STEAITS. 43
Hapalocakcintjs marstjpialis, Stimpson. (Plate 3. figs. 29-40.)
Hapalocarcinus marsupiafis, Stimpson, Proc. Boston Soc. Nat. Hist. vi. 185G-59, p. 412 ; Semper,
'The Natural Conditions of Existence as they affect Animal Life' (London, 1881: Internat.
Sci. Series), pp. 216 >^t seq., fig. 64 c.
Description of female. The carapace is soft and membranous, depressctl, broadly oval
in outline and truncated in front and behind. The breadth is equal to or a little less
than the length. The front is slightly dcflexed, obscurely tridentate, the median tooth
being sharp and separated by a shallow concavity on either side from the rounded lateral
corners. There are no true orbits, the space occupied by the eye being quite undefined
above and externally, while below the eye rests directly on the everted anterior margin
of the buccal frame. Tlie lateral margins of the carapace are rounded off dorso-ventrally
and evenly arcuate from before backwards. The posterior margin is concave and is about
three-fifths as long as the anterior margin. The surface of the carapace is perfectly even,
without any perceptible furrows, and is smooth except for a few minute and widely-
scattered setse.
The abdomen consists of seven distinct segments, of which the first is partly
concealed under the posterior margin of the carapace. The first three segments
are visible from above, and are hardly more than half the width of the carapace. The
remaining four segments are bent under the body, and form a, broad oval plate about
equal in size to the carapace. The middle part of this plate, formed by the terga of the
four segments, is of somewhat firm consistency and is surrounded by a Avide membranous
border, which at the sides is folded inwards to form the lateral walls of the capacious
egg-pouch. The surface of the abdomen, like that of the carapace, is beset with minute
scattered sette.
The ocular peduncles are relatively large, subconical, not lying in distinct orbits. The
corneal surface is facetted, but, in our specimens, devoid of pigment. The antennules ai-e
large and exserted, there being no fossettes for their reception. The basal joint is pro-
duced externally into a large conical process directed obliquely forwards and terminating
in a stout spine. The two succeeding joints are stout, cylindrical, and subequal in length,
tlie distal one carrying the two rudimentary flagella, each consisting of a few joints and
clothed with numerous long setae. The antennae consist of five joints, of which the first
is broad and triangular and the succeeding joints narrow, cylindrical, and successively
diminishing in diameter, the last being hardly thicker than the long seta which springs
from its tip. The buccal aiea is very large, extending across the whole width of the
carapace in front. Its anterior margm is sinuous, curving forwards on eithei- side below
the eye, and taking the place usually occupied by the inferior margin of the orbit. The
median part of the buccal margin approaches so closely to t\w base of the antennules
that an epistome can hardly be said to exist. The third maxillipeds do not nearly cover
the buccal cavity, and are widely separated from each other at the base by a semicircular
area of the sternum. The ischium is flattened, subtriangular in shape, widening
gradually from a narrow base, and having its antero-internal angle produced forwards,
rounded, and fringed with setse. The merus is articulated at the outer end of the distal
G*
44 DE. W. T. CALMAN ON A COLLECTION OF
margin of the ischium, and is less than half the width of the latter, hardly wider than the
succeeding joints, and hut little flattened. The exoj)od is rudimentary, heing a simple
lobe about half the length of the ischium. The epipod is well-developed. The second
maxillipeds have the basal part of the exopod much expanded. In the first maxillipeds,
also, the same part is very stout and much stronger than is usual in this appendage,
while the inner lobe or endopod is small and subtriangular. The chclipeds are rather
stout, about twice the diameter of the succeeding legs, and smooth save for scattered
setae similar to those on the carapace. The merus has a small spine near the distal end
of the inner margin. The hand is not much thicker than the preceding joints. Tiie
palm is less than twice as long as broad, nearly twice as long as the fingers. Thedactylus
has a single tubercle on its inner edge. Tiie walking-legs are not at all concealed by
the carapace. The first three pairs are about equal in length to the chelipeds and the
last pair is a little shorter. The dactyli are strong and curved, and bear a low tooth on
the inner edge near the tip. Tlie legs bear scattered set^e, which are larger and more
numerous than are those on the body. The abdominal appendages are reduced to three
pairs, corresponding to the second, third, and fourth abdominal somites. They are
uniramous with the exception of the first, which carries on the outer side near the base
a small unjointed aj)peudage which appears to represent the exopod.
The female generative aj)erturesare ci-escentic in form, and are situated on the sternum,
far apart from each other, close to the bases of the third legs.
The dimensions of our two sjjecimens are as follows : —
Stimpson's brief description applies very well, on the whole, to the specimens examined
by me. The most serious discrepancy is that the exopod of the third maxillipeds is
described as '' slender and palpigerous." It seems quite possible, however, that in
examining the entire animal the rudimentary exopod of this appendage may have been
overlooked, and the more conspicuous exopod of the second maxilliped may have
appeared to belong to the third. The antennules are stated to be " very short and
minute," but as they are said to be " jjlaced at the inner angle of the orbit," it seems not
unlikely that these epithets should be transferred to the antennae, to which they are more
applicable. Some other less important differences, such as the description of the front
as " straight," may reasonably be attributed to imperfect observation. On the other
hand, tlie description of the general shape, the large abdomen, the large buccal area
occupying nearly the whole breadth of the carapace, the third maxillipeds with " the
ischium large and dilated within, while the merus is very small and slender like the last
three joints," and several other details can apply to no other crustacean, and leave no
doubt that we are dealing with Stimpson's species.
BEACHYUKA FROM TOREES STEAITS. 45
So far as I can discover, no furtlier description of this remarkable form has been
published. Semper gives a figure of it, but it is on too small a scale to be of much use.
It represents the carapace as proportionately broader than in our specimens. All the
recorded specimens have been females.
The peculiar habitat of this Crab was unknown to Dr. Stimpson, who states that his
specimens were found " clinging to the branches of living madrepores at the depth of
one fathom in the harbour of Hilo, Hawaii."
Ehrenberg had long before noticed certain deformities on corals caused by the presence
of Crustacea, and had compared them to the galls formed by plants. In his work on the
Corals of the Red Sea he writes of the species Serlalopora subidatn, Lamk. : —
" Paguri parvi (P. comlUophilos) domicilia in ramis efiiorescentibus sibi pcirant, et veras
gallas fere eliciunt, quales plantse gerunt" (Beitr. z. Kennt. Korall. roth. Meeres, p. 123).
I am not aware that Ehrenberg's " Payurus " has since been recognized, or that any other
Pagurid has been found inhabiting coral-galls, so that it is possible that Ehrenberg's
remarks may really refer to the present species *. Be this as it may, Verrili was the first
to definitely associate these coral-" galls " with Stimpson's Rapalocannnus. In a paper
on "llemarkable Instances of Crustacean Parasitism " (Amer. Journ. Sci. (2) xliv. 1867,
p. 120) he writes : — " Another peculiar mode of parasitism I have observed in a singular
ernstacean {Kapalocarcinus marsupialls, Stimpsou) from the Sandwicli Islands. This
creature lodges itself among the slender branches (jf a coral [PociUopora cccspltosa,
Dana), and causes, probably by its incessant motions, the branches to grow up and
snrround it on both sides by fiat expansions of coral terminating in digitations which
often interlock above, leaving ojienings between them suitable for the uses of tlie parasite
but usually too small to allow of egress. ]Most specimens of the corals of this species
sustain one or more and often numerous examples of these curious enlarged bulbs among
the branches." In a subsequent paper " On the Parasitic Habits of Crustacea " (Amer.
Nat. iii. 1869, p. 239), Prof. Verrili adds that he had " observed similar cavities on
Pocillopjora elonguta from Ceylon, which are probably made by another species of the
same genus." In his " Synopsis of the Polyps and Corals of the North Pacific Exploring
Exi^edition " (Proc. Essex Inst. vi. (1S68) 1870, p. 91), the same writer records the
occurrence of galls on most of the sjoecimens of Pocillopora ccespitosa, Dana, from the
Hawaiian Islands, and he also writes of P. brecicoruis, Lamk. : — " One specimen has a
bulb similar to those made by Hapalocarcinus marsiiphdis, l)ut belonging prol)ai)ly to
another species of the same genus, since it differs con.siderably in form. The aperture is
closed except a few small openings above."
In the general account of his researches in the Philippine Islands (Zeit. wiss. Zool. xiii.
1863, p. 560), Semper has a note on a small " Porcellana " which he found living iu
cavities in a " millepore." In his work on ' The Natural Conditions of Existence as
* I leani from Mr. R. Kiikpatrick, who has been kind enough to find the above quotation forme, that Kluuzinger
refers to Ehrenberg's observations, and gives a figure of the galls on the same sjjecies of coral (which he refers
to -S'. siiiiiosa, M.-E. &. H.) without, however, giving any details as to the parasite (Korall. rjlh. iloer. ii. p. 72,
pi. vii. fig. 15).
46 DE. W. T. CALMAN ON A COLLECTION OF
they affect Animal Life ' (1881) this Crah is identified with the present species, and a
fuller account is given of the " galls " formed by it. These were found on the corals
Siderojiora digitata and S. palmata and on species of Seriatopora. He describes the
formation of the gall by the growth of two broad flattened branches, and notes that the
shape differs according to the species of the coral. " In the Seriatopora both the twigs
are leaf-shaped and beset with more or less numerous offshoots terminating in sharp
spines. In the more solid Focillopora the twigs also have spines, but they are more
massive. Finally, in Sideropjora spines are wholly absent, and the two twigs between
which the crab lives are altogether more massive." He describes the gradual closure of
the gall by concrescence of the edges, " till at length only two fissures, more or less wide,
are left, which plainly show, by their position opposite to each other, that it is through
them that the current for respiration passes : one fissure serves for the influx, the other
for the exit, of the water ; " and reasons are given for believing that these fissures are kept
open owing to the current of water checking the growth of the coral so long as the crab
remains alive.
Semper states that a Hapalocarciniis, " it woiild seem identical in species," occurs at
Reunion (p. 281), but I can find no other record of its occurrence there. He also
mentions (pp. 217 & 453) certain observations by Graeffe, but no reference is given,
and I have failed to trace the source from which he quotes *.
Bassett-Smith f has described galls formed by a crab on Serlatoporu iinhricata, B.-S.,
fx'om the Tizard Banlc in the China Sea. The crab is not described, but it probably
belonged to the present species.
Hickson $ has described and figured galls on a Jlillepora containing a crab which he
assumes to be Sapalocarclnns. The galls are unlike those described by the authors
quoted above, being inflated bulbs with a single, wide, terminal aperture.
The series of galls examined by me are formed on a sjiecies of Scriatopora §, and the
mode of growth agrees well with the descrij)tions of Verrill and Semper. The earliest
stage is represented by a specimen (fig. 38) in which the gall is beginning to be formed at the
point of bifurcation of a branch. A broad palmate process, slightly concave internally,
* As certain passages from Samper's work have recently been quoted, without correction, by Hickson, it may not
be superfluous to point out that the English edition of this work (I have not been able to consult the German
edition) abounds in typographical and other errata. Thus, on p. 216, the date of Stimpson's paper is given as 1837
(Stimpson was born in 1S32) instead of between 18.56 and 1859. On p. 217 it is stated that the present species
was "discovered in the Pacific Ocean by Dana in the course of his great voyage under tho command of AVilkes."'
As a matter of fact, the Bpecimeus were collected (no doubt by Stimpson himself) during the U.S. iS'orth Pacific
Surveying Expedition under Capt. .John liodgers. The incidental references to the carrying of the i/ouitr/ in the
brood-pouch (p. 217) and to the course of the respiratory current (p. 219) are our only authority for^believing that
I/cqialocarcinus differs in these respects from the majority of the lirachyurn. In the explanation of tig. 65 (p. 218)
Sideropora hystrix should probably read Seriatopora hystrhr.
t Ann. Mag. Nat. Hist, ((i) vi. 1890, p. 364.
X Bull. Liverpool Mus. i. nos. 3 & 4, pp. 81-82, plate.
§ Prof. F. Jeffrey Bell, who has kindly e.\amined my specimens, informs me that their fragmentary nature
renders an exact determination ditKcult, but that they seem to approach most closely to S. clegans, M.-E., though
differing from it in certain rliaraeters.
BRACK YUEA FKO:\r T0R11E8 STRAITS. 47
forms one side of tlic cavity in which tlio jmrasite was lodged, the other side being formed
by an expansion of the opposed j^art of the main l)i-anch, at the sides of which lobate
projections are beginning to dcAelop. A considerably more advanced gall is shown in
fig. 39. Here the gall is terminal in position and is roughly lenticular in form, the two
digitate lobes which compose it being perforated by fissures and only touching each other
here and there at the edges. A still older gall (fig. lOj, occupying a lateral position on
a branch, is closed except for two or three very small apertures at or near the margin.
These apertures are not placed regularly opposite to each other as Semper states. On the
outer surface of the gall the calicles are rather smaller than those on the normal branches
and are not arranged like them in series, but appear to be otherwise well developed. On
the inner surface of the gall the calicles, as Semper states, are small, shallow, and have
the septa only feebly developed. They are also in some parts distorted and drawn out to
an elliptical outline, but this distortion does not appear to l)e definitely related to the
marginal apertures of the gall as described by Semper, who attributes it to the action of
the current of water caused by the crab. Semper also found on the inner surface " very
distinct scars, whicii are evidently produced by continual scratching in (me spot," and he
concludes that the crab usually remains in one position within the gall. Such scars are
not visible in our specimens. In the older galls the outer surface rises into rounded,
irregularly placed swellings and short branches, as if the coral were about to resume the
normal habit of growth disturbed by the intrusion of the 2)arasite.
From Semper s earlier note we gather the not uninteresting detail that the polypes
on the mner surface of the gall are colourless.
As regards the further habits and life-history of Kapalocarcmus we have no informa-
tion. The fact that each gall is inhabited by a solitary female, while the male is as yet
unknown, would seem to iiidicate that both sexes are at first free-living, and that it is
only after impregnation that the female becomes imprisoned in a gall. The fact that
the youngest gall observed is of ample size to contain a full-grown Rapalocarciniis tends
to confirm this suggestion.
As regards the systematic position of Iliqxifocarcinm, we have to note in the tirst place
its close affinity with the CryptocJiirus (^oralliodijtets of Heller (" Beitr. z. Crust. Famia d.
roth. Meeres," SB. Akad. Wien, xliii. (1) 1861, p. 3(56, pi. ii. figs. 33-39). As Semper
has shown ('Animal Life,' pp. 217, 221-223)*, Cri/jitoc/uriis, like HapalocarciiiKs, is
parasitic on living corals. In this case, however, no closed " galls " are formed, the
crab living in massive corals (ex. Ooniastuea,) at the bottom of a funnel-shaped
depression, due to an arrest of the upward growth of the coral. The affinity between the
two genera is most clearly shown by the third maxillipeds, Avhich in both cases are
peculiar in having the merus-joint very narrow and the exopod rudimentary. The
structure of the facial region is somewhat similar in both, the antennules not being
retractile into fossettes, while the antemaae are very small and the orbits ill-defined.
The abdomen of the female in both genera is much enlarged, but in Cryptochirm it lies
» Simpers figure of Cn/idochinis differs considerably from those given !.y Tleller. He states, however, that the
Philippine form " ajjpears to l)e in no respect specifically different '' from that found in the Red Sea (o/;. (it. ]>. 2.S1).
48 DE. W. T. CALMAN ON A COLLECTION OF
wholly hehind the carapace, and is bent npon itself so as to form a deep pouch, open
only in front, while in Eapalocarcinus it is flexed nndcr the body in the manner usual
among the Brachyura. Among other differences the orbits of Cryptochirus are defined
externally by a strong tooth, and the basal joint of the antennules presents a serrate edge
anteriorly where that of Ilapalocarchms has only a stout dentiform lobe.
In describing Hctpalocarcimis, Stimpson noted its resemblance to Pimwtheres in the
large size of the abdomen and the softness of the integument, and he stated that its-
systematic position was probably between Pinnotheres and Hymenosoma. Apart from
the two points mentioned, there seems to be little in the chai-acters of the species as now
described to suggest affinity with the Pinnotheridre, while the third maxillipeds are
widely different in type from aiiytbing found in that group.
Heller expressed no opinion as to the systematic place of his Cryptochirus.
A. Milne-Edwards, however, has described under the name Llthoscaptus paradoxus (in
Maillard's ' Notes sur I'lle de la Reunion,' 2™^' ed., 1863, ii. Annexe F, pp. 10-12), a
form which, as Paulson has already pointed out, is in all probability identical with, or
closely allied to, Heller's species. Tiiis genus was regarded by Milne-Edwards as
representing a new family, " Lithoscaptes," among the " Bracliyures anormaux."
He writes, " par sa region cephalothoracique le Lithoscapte se rapproche des Ptanines
plus que tout autre groupe de Decapodes." Special resemblances to the Paninidie are
said to exist in the structure of the antennal region and in tlie shape of the thoracic
sternal region, which is broad in front but much contracted between the bases of the
last two pairs of legs. It is not clear, either from Milne-Edwards's or from Heller's
accounts (supposing the two genera to be identical), in what way the antennal
region resembles that of the Raninidse, while the thoracic sternum is not contracted
posteriorly, at all events in the female Hapalocar chins. Heller describes the sternum
of CryptocJiirm as " ziemlich breit, langlich oval." The third maxillipeds are said by
Milne-Edwards to resemble a little those of Reniipes, but this resemblance appears
to consist merely in the absence of a conspicuoiis exopod. Milne-Edwards describes
the abdominal appendages as uniramous and as existing on the first four somites.
According to Heller there are only three pairs, and this agrees with our examination
of Hapalocarcinus.
Paulson refers Llthoscaptus ( = Cryptochirus) to the Pinnotheridae, establishing for its
reception a new subfamily, which he designates Crypochirinai and defines as follows : —
" Cephalothorax convex, almost twice as long as broad. Inner antenna? without
fossettes and lying longitudinally. Basal portion of the outer antennae free. Third joint
of the outer maxillipeds considerably shorter than the second. Openings of the female
sexual organs on the sternum " (' Crust. Bed Sea,' Kiev, 1875, p. 72).
While the characters of Hapalocarcinns, as now described, show clearly that it must
stand alongside Cryptochirus, they give little help towards settling the place of the two
genera in the system. The position of the female genital apertiu'es shows that they
must be placed among the true Brachyura, although there are some curiou.s resemblances
to individual genera of the Anomura. Thus the endopod of the third maxillipeds
BRACHTUEA FROM TORRES STRAITS. 49
resembles somewhat that of Porcellana, while the rudimentary exopod suggests a com-
parison -with the Tllpindea. The number and uniramous condition of the abdominal
appendages also agree, except for the absence of the sixth pair, witli tlie last-named
group. These resemblances, however, are balanced by numerous important differences,
so that even were we to set aside the evidence of the genital openings, it would lie
impossible to place the genera in any of the groups of Anomura. On tlie other hand,
the characters of the third maxillipeds and of the abdominal appendages and the greatly
enlarged buccal area are unlike anything found among the Brachyura. The resemblance
to the Pinnotheridae ajDpears to be quite superlicial.
The characters usually relied upon to distinguish the various divisions of the Brachyura
have been in this case apparently so prof(Huidly modified by the pai'asitic habit of life,
that we can only regard these two genera as forming a family for the present incerke
sedis, for which the name Hapalocarcinidae will have to replace Mihie-Edwards's
" Lithoscaptes," the latter being based on a synonym of Cryptochlriis.
LocaUtij. " Torres Straits."
Distvlhution. Hapalucarcinus is recorded by name from Hawaii {Stimpson, Verrlll),
the Philippines, and (?) Bourbon {Semper). Coral-galls, possibly due to this species, arc
known from the Red Sea [Ehrenberg, Klunzlngev), Ceylon {Ven-iU), and the China
Sea {Bassett-Sndtlt).
EXPLANATION OF THE PLATES.
Plate 1,
Fig. 1. Filuiiinus cristipes, ii. sp., $ (enlarged).
2. ,, „ ,, Cephalic region iroin below.
3. „ ,, „ Right chela.
4. Cri/fjiucnemtis Huddon'i, u. sp., $ . Dorsal view, x G.
5. ,, „ ,, Ventral view.
C. ,, „ ,, Lateral view.
7. ,, „ ,, 'I'hinl maxilli[)c(l, inner face.
8. ,, ,, ,, Chela.
9. Palicus Jiikesii (White), cj, x 5.
10. „ ,, ,, Cephalic region from helow.
11. ,, „ „ Abdomen.
12. ,, „ ,, Second walking-leg from l)elow.
13. ,, „ ,, First abdominal appendage.
SECOMJ SERIES. — ZOOLOGY, VOL. VIII.
50 ON BEACHTUEA FROM TORRES STRAITS.
Plate 2.
Fig. 14. Palicus Whitei (Miers), ? , x 3i.
15. ,, „ „ Cephalic region from below.
16. ,, „ „ Abdomen, $ .
17. „ „ „ Abdomen, S-
18. „ „ „ Second walking-leg from below.
19. „ „ „ Fir.st abdominal appendage, ^ .
20. Palicus serripes (Ale. & And.), $ , x 4.
21. „ „ „ Cephalic region from below.
22. „ „ ,, Second walking-leg from below.
23. Hydstenus verrucosipes (Ad. & Wh.). Dorsal view (setre omitted), x 5.
24. „ „ „ Cephalic region from below.
25. Pseudomicippa varians, Miers, x 4.
26. „ „ „ Cephalic region from below (setse omitted).
Plate 3.
Fig. 27. Laiiibrus confragosns, n. sp., x 2^.
28. „ „ Outline from side.
29. Hapalocarcinm marsupialis, Stimpson^ ? . Dorsal view, x 13.
30. „ „ „ Ventral view.
31. „ „ „ Ventral view of cephalic region.
32. „ ,, „ Antennre and antennules.
33. ,, ,, „ First maxilliped.
34. „ ,, „ Second maxilliped.
35. „ „ „ Third maxilliped.
36. „ „ „ Chela.
37. ,, „ „ Dactylus of last walking-leg.
38. Branch of Seriatopora sp., showing at * beginning of " gall" formed by Hapalocarcinus.
39. A more advanced " gall," still widely open at edges.
40. A " gall," closed all round except for a few small apertures marked *.
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VII. Part I. 1896 0 10 0
Part II. 1897 0 12 0
Part III. 1897 0 6 0 . . .
Part IV. 1898 0 10 0 . . . .
Part V. 1898 0 18 0 . . . .
Part VI. 1898 0 13 0
Part VII. 1899 0 18 0
Part VIII. 1899. . . . 0 12 0
Part IX. 1899 1 0 0 . . . .
Part X. 1900 0 6 0
Part XI. 1900 0 2 9
VIII. Part I. 1900 0 10 0 . . . .
0 7 6
2nd Ser. ZOOLOGY.]
[VOL. VIII. PART 2.
THE
rl^
PR A NS ACTIONS
OP
THE LINNEAN SOCIETY OF LONDON.
REPORT ON A COLLECTION MADE BY Messrs. F. V. McCONNELL
AND .L -L aCELCH AT MOUNT RORAIMA IN BRITISH GUIANA.
{('omniiniirntrd by I'rof. E. Ray Lankester, D.C.L., F.R.S., Director Nat. Hist. Mus.)
\ , I-
Ji L O N P O N :
PRINTKl) FOR TIIK I.INNE.\N SOCIETY
BY 1 AVl.OK AMI> FKANl-IS. RKI) I. ION COURT. FI.F.KT STKEKT.
SOLD A'l' THE -society's APARTMENTS, BURLINGTON-HOVSE. J'TCCADILLY, W.,
AND nv LONGMANS. GREEN, AND CO., PATERNO.STER-ROW.
September 1900.
[ 51 ]
J I. Report on a Collection made by Messrs. F. V. McConnell and J. J. Quelch at
Mount Roraima in British Guiana. {Communicated by Professor E. Ray Lankester,
D.C.L., F.M.S., Director of the Natural History Museum.)
(Plates 4-6.)
Read 1st February, 1900.
r OR some years j)ast the British Museum has received many interesting collections
made by Mr. P. V. McConnell and Mr. J. J. Quelch during their various expeditions
into tlie interior of British Guiana. On the last occasion the explorers succeeded in
reaching the snmmit of Mount Roraima, and I have much pleasure in forwarding the
accomjjanying descrijjtions of the new species obtained during the expedition, all of
vi'hich have been worked out by members of the staff of the Zoological Department
of the British Museum, excepting the Crustacea, which have been dealt with by
Dr. de Man.
Mr. McConnell has kindly sent me the following note : — " The specimens were
obtained by Mi\ J. J. Quelch and myself on our second exjiedition to Roraima in August,
September, and October, 1S98. The route taken on tliis occa.sion Avas by the Mazaruni
and Curubung rivers to the Palls of Macrobah, a boat-journey of twenty days, and
thence by land to a point on the U})pei' Mazaruni where that river flows at a height of
1300 feet. Small bark-canoes were here obtained, and after three days' journey ujj the
Cako and Aruparu we arrived at the commencement of the ti'ail to Roraima. With the
exception of the last twenty miles, the whole journey, which occupied forty days, lay
through thick forest.
" Mount Roraima (8700 feet) is formed by a sloping base, surmounted by a rectangular
mass, fifty-four square miles in area, with perpendicular walls 2000 feet in height. On
the south-west, part of the wall has slipped, and lies diagonally across the face of the
upper part of the mountain. By following the ledge so formed the summit can be
reached without serious difiiculty." — E. Ray Lankester.
SECOND SERIES. — ZOOLOGY, VOL. VUI.
52 REPORT ON A COLLECTION FROM
MAMMALIA.
By W. E. DE WiNTON, E.Z.S.
Order RODE NT I A.
Ehipidomys Macconnelli, sp. n.
The general colour of the entire upper surface rich golden brown ; the fur is very soft,
from 11 to 12 millimetres in length, dull black for the greater part of its length with
bright red golden tips ; the slightly longer straight hairs have black tips ; the lower
surface is dirty white or drab, the fur being dull black with whitish tips, with no line of
demarcation between the colours of the upper and lower surfaces. The ears are large,
round, and naked, almost black in colour. The hands and feet dirty white, rather darker
on the upper surfaces, almost naked. Whiskers long, reaching beyond the shoulders.
The tail is brown, only very slightly paler beneath, practically naked, the very minute
hairs in no way hiding the scales ; at the extreme tip there is a long pencil of hairs about
10 millimetres long, but no l)ushy hairs on the sides.
Upper incisors dull orange, rather darker than those of the lower jaw.
Measurements taken from the specimen in alcohol: — Head and body 95 millim., tail 1J.7,
hind foot 2i<5, ear 17.
Skull — greatest length 26'5 ; greatest breadth ll-'S, across brain-case 12-7 ; narrowest
interorbital constriction 16 ; length of nasals 9"1 ; basal length 22'6 ; back of incisors to
])ack of palate 11"5 ; incisive foramina 5"9x2'5 ; diastema 7; length of molar series 5;
width outside first molar 5-5, outside last molar 59; mandible, tip of incisors to
coronoid process 12, to condylar process 17'8, to angle 173.
The type and only specimen was found on the summit of Rorairaa, Demerara, and has
been presented to the Museum by Messrs. McConnell and Quelch.
The skull is fragile and transparent, with large, smooth brain-case ; the zygomata are
very thin and weak, expanding very little beyond the sides of the head in the squamosal
region only. The molars are set in almost parallel rows ; the auditory bullae are very
small ; the foramen magnum is very large, the surrounding Ijones very thin, especially the
basioccipital condyles.
This new JR,kipiclomy.s is of much the same size as li. vdci'otis, Thos., from Colombia,
but the colour is distinct, being much darker above, while the underparts are greyish
with dark bases to all the fur instead of being pure white ; then the ears of the new
species are large, while those of the Colombian species are very small. Tlie tip of the
tail may or may not be distinct, for the end of the tail of the type of 11. microtis
appears to me to be wanting.
The broken state of the skull of R. microtis exckides the possil)ility of a fair com-
parison being made, but in the dentition the two species seem very similar.
MOUNT KOEAIMA IN BKITISH GUIANA. 53
BIRDS.
By R. BowDLEK Sharpe, LL.D., F.L.S.
(Plate 4.)
From the summit of lloraima Mr. McConnell brought several specimens of a
Zonotrichia quite distinct from the ordinary Z. pileata which is found over the greater
part of Central and South America. It is a larger and darker bird, much greyer,
and with the rump and flanks dark grey instead of brown. I propose to call it after
Mr. McConnell, who has done so much for the collections of the British Museum.
Zonotrichia Macconnelli, sp. n.
Similis Z. fileatce, sed major, obscurior, uropygio et hypochondriis saturate griseis,
nee brunneis distinguenda. Long. tot. frS poll., culm. 0-6, alae t2-8, caudse 2-35,
tarsi 0-9.
Hah. Summit of Mount Roraima.
EXPLANATION OF PLATE 4.
Fig. 1. Zonotrichia Macconnelli.
2. „ pileata.
REPTILES.
By G. A. BouLENGER, F.ll.S.
(Plate 5. figs. 1-2.)
1. AisOLis fusco-auratus, D'Orb.
This and most of the following Reptiles and Batrachians were obtained at the base
of the mountain, at an altitude of about 3500 feet. Prmiodactylus leucostictu,^ and
Oreophrynellu Q/(elchiia,ve fi-om the summit (8600 feet).
2. Centropyx calcaratus, Spix.
3. Centropy.x striatus, Daud.
4. Neusticurus rudis, sp. n. (Plate 5. figs. 1-1 c.)
Snout rather short, pointed ; canthus rostralis strong ; ear-opening as large a« the eye-
opening ; transparent disk of lower eyelid composed of five pieces. Rostral rather large,
the portion visible from above nearly half as long as tlie frontonasal, which is longer
than broad ; a pair of praef rentals ; a single frontal ; a pair of frontoparietcils, followed
by five shields in a row ; occipital region covered with small irregular shields ; three
S*
54 EEPOKT ON A COLLECTION FROM !. ;
large supraoculars, preceded by a small fourth and two or three granules ; nostril piei'ced
in a single nasal, which, like the loreals and suboculars, is in contact with the upper
labials ; latter, seven in number, fourth largest ; four lower labials ; chin-shields, one
anterior and two pairs. Gular scales smooth, smallest on a zone connecting the ears ;
collar bordered by six plates. Nape and back covered with small keeled scales inter-
mixed with large, oval, sharply keeled tubercles, disposed very irregularly, but avoiding
the vertebral line ; smaller tubercles on the sides. Ventral plates rounded posteriorly,
imbricate, in 8 or 10 longitudinal and 28 transverse series ; the plates of the four
median rows subequal, about as long as broad. Three slightly enlarged praeanal plates,
forming a triangle. 18-20 femoral pores. Tail feebly compressed, Avith the pair of
dorsal keels, formed by enlarged tubercles, feel)ly developed. Uniform blackish brown
above, whitish beneath.
Totallength 183 millim. From end of snout to vent ... 59 millini.
Head 14 „ Fore limb 19 „
Widthofhead 8-5 „ Hind limb 27 „
From end of snout to fore limb... 2.3 „ Tail 124 ,,
A single specimen ( 6 ) from the foot of Mt. Roraima, 3500 feet.
5. PmoNODACTYLUS LEUCosTiCTUS, sp. n. (Plate 5. figs. 2-2 c.)
Snout short, obtusely pointed; eai'-opening a little smaller than the eye-opening.
Frontonasal single, a little broader than long ; prsefrontals well developed, forming a
median suture ; interparietal large, hexagonal, larger than the parietals ; two pairs of
occipitals ; four supraoculars ; nostril pierced in a single nasal ; no loreal ; six upper
and five lower labials ; chin-shields very large, one anterior and four pairs, the first two
forming a suture, the two others separated on the median line by granules ; two
longitudinal rows of large transverse gular shields ; five collar-shields. Dorsal scales
hexagonal, strongly keeled ; lateral scales small, roundish, smooth ; 26 scales, including
the ventrals, round the middle of the body; 29 scales from occiput to base of tail.
Ventrals large, in 8 longitudinal and 19 transverse series. Four large praeanal shields,
forming a cross. 6 femoral pores on each side ( S ). Tail above with hexagonal
keeled scales, beneath with tetragonal smooth scales. Black above and beneath; each
scale or shield with one to three white dots, these dots larger on the ventral shields ;
chin white.
Totallength 127 millim. From end of snout to veut ... 50 millim.
Head 10 „ Fore limb 13 „
Widthofhead 6 „ Hind limb 19 „
From end of snout to fore limb... 19 „ Tail 77 „
A single specimen ( $ ) from the summit of Mt. Roraima, 8600 feet,
o. Lachesis lanckolatus, Lacep.
MOUNT KOKAIMA IN BRITISH (il'IANA. 55
BATRACHIANS.
By G. A. BouLENGER, F.R.S.
(Plate 5. fi-s. 3-6.)
1. Oreophhynella Quelchii, Blgr. (Plate 5. fig. 3.)
This tiny toad was described by me as the tyi)e ol' a new genus in 189") (Ann. & Mag.
N. H. [6] xvi. pp. 125 & 522) from specimens collected on the; suiimiit of Mt.
Roraima by Messrs. Quelch and McCoimell. Numerous specimens liave since been
obtained at the same altitude (8500-8000 feet).
I have now the pleasiu-e of describing a second species of the same genus, from the
base of the mountain.
2. Oreophrynella Macconnelli, sp. n. (Plate 5. fig. 4.)
Closely allied to 0. Qaelchii, but distinguished by the more prominent snout, projecting
much beyond the mouth, and by the greater distal expansion of the digits, which end
in distinct truncated disks. Interoi-lntal space broader than the upper eyelid. First
toe mucli longer than second, as long as fourth ; no distinct subarticular or metatarsal
tubercles ; the first and second toes appear to be opposable to the I'ourth and fifth, both
fascicles being bound by the thick integument, and the third toe, which is the shortest,
being free between them. The tarso- metatarsal articulation reaches the; eye. Upper
parts covered with small, smooth, feebly pronainent warts ; lower parts with flat
granules. Olive-brown above, with lighter marblings, and a series of small yellowisli
spots, forming a line on each side of the back from the eye to the groin, continued
obliquely across the upper surface of the femur ; upper lip yellowish, with two dark
brown bars below the eye ; lower parts whitish.
From snout to vent 22 millim.
A single specimen from the base of ^It. iloraima, 3500 feet.
Otophryne, g. n. Eii(jy,sto7uutidarum.
Pupil round. Tongue large, oval, truncate and free behind ; no teeth on the palate ;
two denticulated dermal ridges in front of the pharynx. Tympanum very distinct.
Fingers free, toes webbed at the base, the tips merely swollen. Outer metatarsals
united by the integument. Precoracoid jjresent, weak ; sternum cartilaginous. Sacral
diapopliyses rather strongly dilated *.
3. Otophryne robusta, sp. n. (Plate 5. figs. 5, 5 a.)
Habit stout; head rather small. Snout short, pointed, obliquely tnmcated at the end
and projecting beyond the mouth, which is inferior, well within the lines of tiie canthi
rostrales ; nostril equally distant from the eye and the end of the snout ; eye rather
small, little prominent ; interorbital space nearly twice as broad as the upper eyeUd ;
tympanum very large, close to the eye and once and a half its diameter. Fingers short,
• Osteological characters ascertained by means of a sciagraph kindly prepared by Messrs. Gardiner and Green.
5(5 KEPOHT ON A COLLECTION FROM
swollen at tbc end, first nearly as long as but not reaching so far as second. Toes very
short, depressed, swollen at the end, webbed at the base, the web continued as a fringe
alono- the sides ; no distinct subarticular or metatarsal tubercles. The tarso-metatarsal
articulation reaches the tympanum ; tibia as long as the foot, two-fifths length of head
and body. Skin thick, perfectly smooth, shiny on the upper parts ; a strong fold from
the eye to the insertion of the thigh. Blackish brown above ; groin and binder side of
thighs orange, spotted with black ; throat and breast dark brown ; belly and lower
surface of limits whitish with some brown dots.
Prom snout to vent 53 millim.
A single specimen from the foot of Mt. E-oraima, 3600 feet.
4. Hylodes marmoratus, sp. n. (Plate 5. fig. 6.)
Tongue circular, entire ; vomerine teeth in two very small oblique groups behind the
level of the choana;. Head a little longer than broad ; snout rounded, not prominent,
as long as the diameter of the orbit, with obtuse canthus rostralis and concave loreal
region ; nostril much nearer the tip of the snout than the eye ; interorbital space nearly
as broad as the upper eyelid ; tympanum distinct, one-fourth the diameter of the eye.
Digits moderately elongate, with well-developed disks, which are as large as the
tympanum, and strong subarticular tubercles ; first finger sliorter than second ; toes
quite free ; a small, oval, inner metatarsal tubercle. Tlie tibio-tarsal articulation reaches
l^etween the eye and the nostril ; tibia half the length of head and body. Skin smooth
above, with feeble oblique glandular ridges on the occiput ; belly granular. Grey-brown
above, with brown, dark-edged marblings on the head and body and cross-bars on the
limbs; dark bars radiating from the eye; grey-brown beneath.
Prom snout to vent 19 millim.
A single specimen from the foot of Mt. Roraima, 3500 feet.
5. Hyla albomauginata, Spix.
EXPLANATION OF PLATE 5.
MOUNT ROKAULA [.\ BRITISH GL'[A\A. 57
CRUSTACEA.
By Dr. J. G. de Max.
(Plate 6.)
PaLtEMON (MaCROBUACHIUM) QtiELCUI, .sj). II.
Thirty-seven specimens, only five or six of uiiich are full-grown, were collected in the
Upper Mazaruni river at an altitude of 2500 feet, and one young specimen was captured
at an altitude of 3500 feet on the Mt. Pvorainia range. Amongst the lormer is ])ut one
ova-bearing female, the rest are both males and young females, the full-grown specimens
being all males.
This pretty species, that I have the pleasure to name after Mr. .). J. Quelch, is
apparently closely allied to Fal. potluna, P. Miiller, from the Itajahy river. State
of Santa Catliarina, and to Fal. Iheringi, Ortm., from tlie State of Sao Paulo, both in
the south of Brazil ; but it is no doubt different, the second legs presenting characters
intermediate between those of the two quoted species. Palcemon Qmlchi is evidently
their representative in British Guiana. One full-grown specimen only is still provided
with both legs of the second ])air, in tlie others one of tliem is lost. The ova-bearing
female has also lost these legs, and in the numerous young individuals one leg of this
pair or even both are often wanting.
Falasmon Quelchl belongs to the species of sum// size, the adult individuals mcasurino-
only 55 millim. from tip of rostrum to the extremity of the telson. Examined under a
rather strong lens the cephalothorax presents a tine and rare puuctation, on wliich one
observes a short pubescence, for the rest it appears smooth. The rostrum (PI. 6. figs, l-i),
vertically moderately deep, is rather short, reaching only the end of the antennulary
peduncles or even only the middle of their terminal joint, so that it does not extend to
the end of the antennal scales. The iipper margin, usually very slightly convex above
the eyes, gradually descends downwards and carries sereii, eiyht, or nine low, rather
equidistant teeth, the first two of which commonly stand ou the cephalothorax, l)ut
often only one tooth stands on it, the second being placed above the orbital maru:in.
The lower margin is usually armed with two teeth, often, liowever, witli one only. The
formulae for 34 specimens are the following : —
o specimens ., ; 4 specimens j ; 9 specimens r, ;
* . - ' . r [ .
1 specimen !^.
The hepatic spine is small and placed below and posterior to the somewhat larger
antennal one. The apex of the telson, as usual shorter than the lateral appendages and
the flattened upper surface of which bears the two ordinary pairs of small spinules, is
triangular with a quite short median spine ; the inner spinules are somewhat longer
than the median point and considerably longer than the outer ones.
The free end of the antennal scales is obtusely angulated internallv and reaches a little
68 EEPORT ON A COLLECTION FROM
further forward than the short spine at the extremity of the external margin. The
shortest of the three antennular flagella is distinctly serrate and exceeds the free end of
the antennal scales hy its whole length. The external maxillipedes project with their
terminal joint beyond the peduncles of the outer antennoe.
The iirst pair of legs exceed, in the full-grown male, the antennal scales hy two fifth
parts of their carpus ; the latter is once and two-thirds as long as the hand, the fingers
very slightly longer than the palm.
The second legs are considerably stouter and longer than the first and somewhat
unequal. In the largest male, which is 54 niillira. long, both legs (PL 6. figs. 5 & 6) are
slightly longer than the body and both exceed the antennal scales by the whole length
of the carpus. The cylindrical merus widens slightly towards its distal end. The
carpus of both legs appears at first sight just as long as the merus, but measured exactly
it appears always very slightly longer than it. The carpus, quite narrow at base and
here much narrower than the distal end of the preceding joint, regularly widens towards
its distal extremity, so that it has a conical shape and its diameter at the distal end is a
little broader than that of the merus. The carpus appears, therefore, tioo and a half to
three times as loncj as thick at its distal extremity. The chela is two and a half times as
long as the oarpiis, and in both legs the pahn measures almost two-thirds the length of the
whole hand. The palm of the larger chela is distinctly broader than the widened distal
end of the carpus, being a little more than once and a half as broad ; the palm is about
three times as long as broad, and its width measures almost one-fourth the length of the
whole hand. The palmar portion of the hand appears .^lightly broader than thick, the
proportion being as 6 : 5 ; it is everywhere rounded both on tlie upper and lower surface
and on the sides. When the cliela is looked at from above, the outer margin of the palm
appears straight, but the inner slightly convex, and the inner border of the chela is a
little concave at the base of the fingers. The j^ointed fingers leave, when closed, a
narrow interspace between them, in the middle about as broad as the fingers thenaselves ;
the latter are almost cylindrical. The immobile finger is nearly straight and tapers but
very slightly towards the tij) ; the dactylus, however, is somewhat curved and tapers
more regularly. Each finger is armed with a strong conical tooth ; that of the index is
2)iaced just in the middle of tlie finger, that of tlie dactylus a little beyond it; three
much smaller obtuse teeth are observed between each conical tooth and the articulation,
and the third of these small teeth is double. On each finger a sharp cutting-edge runs
between the conical tooth and the tip.
The smaller chela (fig. 6) bears a close resemblance to the other, but the diS'erence
between its width and its height or thickness is still smaller, so that the palm appears
almost cylindrical and but slightly broader than the carpus. The fingers are regularly
tapering, the dactylus is less curved, and the interspace l>etween both is small, only half
as broad in the middle as the fingers. The toothing is about the same, but the dactylus
bears six small obtuse teeth between the large conical tooth and the articulation.
In the younger individuals the fingers are comparatively longer, so in a young male,
long. 36 mm., the palm is 4^ mm , the fingers 4 mm. long ; the former. If mm. broad,
is three times l)roader than long and 1 mm. thick.
MOUNT EOEAIMA IN BRITISH GUIANA. 59
Mg. 7 represents the second leg of a female, long. 42 mm., dcA^oid of eggs. The merus
measures 5 m.m., the carpus 5J, the hand lly mm., of which the palm occupies 6 mm.
The palm is If mm. broad, the carpus at its distal extremity 1| mm. The toothing of
the fingers, figu.red fig. 7 a, appears in this young individual still very feeble, the
dactylus showing only three teeth, the index also, but those are less prominent, more
rounded.
The second legs of these young individuals bear a close resemblance to Pal. potiuHu,
h\ Miill. {vide Ortmann, ' Os Camaroes da agua doce da America do Sul,' S. Paulo, 18i)7,
est. i. tig. D), but there can be no doubt that this species is a different one, for this
rescm1:)lance is only exhibited hij quite ijouiig individiiah.
The second legs are on all their joints roughened by small thorny points, that arc
crowded and numerous on their outer margin, less numerous on the rest of their surface,
and those of the lower surface and of the inner margin are distinctly somewhat longer ;
these legs are glabrous, devoid of hair, except a rare short pubescence, only perceptible
under a lens.
The ambulatory legs of the third pair project with a third of their pro^iodites beyond
the antennal scales, their carpopodites reaching as far forward as the peduncles of the
outer antenuu3 ; the legs of the fifth pair finally extend as far forward as the external
maxillipeds, but do not reach the free end of the antennal scales. The ambulatory legs
are rather slender. So are the meropodites of the third pair of tlic largest male 8 mm.
long, 1'25 mm. thick, the propodites l'\) mm. long and O'Si mm. thick, so that tlie
foi'mer are little more than six, the latter nine to ten times as long as broad ; for the
meropodites of the fifth legs (PL 6. fig. 8) these numbers are 7'5 mm. and 1 mm., for
the propodites 79 mm. and 0'7 mm., so that the meropodites are seven to eight, the
propodites eleven times as long as broad. The dactylopodites are short, measuring about
one-foui'th the length of the j)ropodites. The posterior margin of the propodites bears
two rows of spinules, so that in the third legs there are nine or ten spinules in the outer
and six or seven in the inner row. The ambulatory legs are a little hairy, but for the
rest quite smooth : the hairs are very short and fine, and arranged partly two and two in
longitudinal rows ; so that one row runs along the posterior margin of tlie meroj)oditcs.
The eggs are few hi number hut larye, having a diameter of 2^ mm. Concerning the
single female carrying these eggs, which is 38 mm. long from tip of rostrum to the
extremity of the telson, the following may be remarked : — The rostrum (tig. 2) reaches to
the middle of the terminal joint of the anteunulary peduncles ; the upper margin that
descends obliquely downward bears seven teeth, the second of which is placed above the
orbital margin ; the lower border is armed with two teeth, the interspaces are as usual
ciliated. Tlie external maxillipeds exceed the antennal jieduncle only by half their
leruinial joint. The tirst legs project only with the hands beyond the free end of the
antennal scales ; the hands measure just two-thirds the length of the carpus. The legs
of the third pair reach to the end of the antennal scales, those of the fifth to the end of
the antennal peditncles. The meropodites of the third pair are ij mm. long and f mm.
broad ; the propodites are 1< mm. long and ^ mm. broad.
Falcctaoii, potiumi, P. Miiller, dilFers at tirst sight by the chelae of the second legs
SECOND SKUIKS. — ZOOLOGY, VOL. VIII. 9
60
REPOET ON A COLLECTION FROM
having the fingers as long or even, according to Ortmann's figure, slightly longer than the
palm. PalcBnion Uierlngi, Ortm. {I. c. p. 211, est. i. figs. 7 e, 8) is apparently also
different. The carpus of the second legs, indeed, does not gradually and regularly widen
towards its distal end, but suddenly, so that the form is different.
Measurements in millimetres.
No. 1.
Length from tip of rostrum to extremity of abdomen
„ of second legs
„ of merus
„ of carpus
Width of the carpus at the distal end
Length of chela
„ of palm
Breadth „
Height „
54
Left.
58
10|
10|
4
25
15|
Right.
61
101
11
4
28i
18
64
No. 0. ' No. 6.
45
32
H
6
oi
12|
7
^
39
7
n
2|
18
10
■n
No. 6 is a detached leg.
EXPLANATION OF PLATE 6.
Figs. 1-4. Palmmon Quelchi, sp. u. Anterior portion of carapace in four examples, X 3 : Fig. 1 of the
largest male, long. 54 mm. ; Fig. 2 of the ova-beariug female, long. 38 mm. ; Fig. 3 of another
male, long. 52 mm. ; Fig. 4 of a young male, long. 35 mm.
Fig. 5, right, and Fig. 6, left leg of the second pair of the largest male, long. 54 mm., x 2.
Fig. 7, One of the legs of the second pair of a female without eggs, long. 42 mm., X 2 ; 7 a, toothing of
botli fingers of this specimen, x 25.
Fig. 8. Fifth leg of the largest male, long. 54 mm., X 5.
List of the known Species of the Genus Palsemon, Fabr. s. s., May 1900.
[The species printed in italics inhabit America and the West Coast of Africa. The locality-
indicated as the habitat is in every case taken from the first published description of the species.
Of those marked with an asterisk the descriptions were not accessible to me when preparing this list.]
1. acanthosoma, sp. n. (?) Nob. Katau, New Guinea.
2. acaiithurus, Wgm. Coast of Brazil.
3. acutirostris, Dana. Sandwich Islands.
4. africanus, Bate. Tambo river. — According to von Martens, 1869, = Gaudichaudii, M.-E. The
Tambo river, mentioned by Spence Bate as the habitat of his species, would, according to
von ^lartens, be situated in Peru! Confer also: Miers, ' On a Collection of Crustacea from
South America,' 1877.
5. africanus, Kingsl. West Coast of Africa. — Thalhvitz, 1891, supposes this species to be identical
with Pal, macrob)-ucMo7i, Herkl.
MOU>'T R01{AIMA IN BKITISH GUIANA. 01
6. Alphonsianus, IlfFm. Reuuion. — This species is identical with Pal. lUspar, Marts. Confer :
de Man, ' Crustacea collected by Max Weber/ 1892, p. 437.
7. altifrons, Hend. Delhi; River Jumna; Lahore.
8. amasonicus, Hell. Amazon river.
9. americanus, Bate. Lake of Amatitlan, Guatemala. — According to von Martens, 1869, = brachy-
dactylus, Wgm., and according to Miers, 1888, =jamair.ensis, Hbst.
10. Apjmni, Marts. Porto Cabello, Venezuela.
„ var. (Equutorialis , Ortm. Ecuador.
11. asper, Stps. In fresh water and in the river near Canton, China. — This species is identical with
nipponensis, de Haan.
12. asperulus, ^larts. Shanghai.
13. Audouini, Hell. Red Sea.
14. Audouini, Bate. Off New Zealand.
15. australis, Ortm. = sp., de M., 1887 ? Queensland. — Ortmann, ' Decapoden-Krebse des Strass-
burger Museums,' p. 708.
16. aztecus, Sauss. Gulf of Mexico.
17. bariensis, de M. Fresh water, Flores.
18. boninensis, Stps. Bonin Islands, in hill-streams.
19. Borcllii, Nob. San Lorenzo (Jujuy) j San Luis.
20. bruchydactylus, Wgm. East coast of .Mexico.
21. brasUiensis , Hell. Camaroes, Brazil, fresh water. — According to Ortmann a locality of this name
docs not exist in Brazil. Camaroes would be the Spanish name of these prawns ! {' Decapodcn-
Krebsc des Strassburger ]\Iusenms,' p. 711.)
22. brevicarpus, de Haan. Japan. — Confer: de Man, in Mas Weber's ' Crustacea," 1892, p. 418.
23. brevimanus, Fabr. India.
24. c(Pmentarius, Poepp. Mouth of the River Aconcagua. — This species is identical with Bithyn'is
loiKj'imana, Phil. Confer : ' Zoologischer Anzeiger,' 1894, p. 266 ; von Martens, ' Ueber
einige ostasiatische Siisswasserthierc,' 1868, p. 65 ; and Miers, /. c. 1877, p. 662. According
to Miers it is a variety of Pal. Gaudichaudii, M.-E.
25. callirrhoe, de M. Mandai river, Ketoengau river (Borneo).
26. carcinus, Fabr. India f.
27. consubrinus, Sauss. Gulf of Mexico, off Vera Cruz.
28. coromandelianus, Fabr. India.
29. Dana;, Hell. Sydney.
30. Dayanus, Hend. Orissa, Calcutta, Lahore.
31. dasydudylus, Streets. Tide-water of the Coatzacoalcos river, Isthmus of Tehuantepec—
According to Ortmann (/. c.) = mexicanus, Sauss.
32. Desamuri, Hell. New Gi-anada.
33. dispar. Marts. Isle of Adenare.
34. dolichodactylus, Hilgd. Mozambique.
35. dulcis, n. sp. ?, Thallw. North Celebes.
36. elegans, de M. Siuagar, Buitenzorg, Java.
37. endehensis, de M. • Flores.
38. ensiculus, S. Sm. Para.
39. equidcns, Dana. In the sea near Singapore.
40. escukntus, Thallw. • North Celebes.
+ Fabricin^ indicates the rivers of America as the habitat of this species— of course, wrongly.
9*
62 EEPOET ON A COLLECTION FROM
4L eurvriiyiiclms, Ortm., = latimaims. Marts. Fiji Islands. — Confer: de Man, iu Max Weber's
'Crustacea/ 1892, p. 482.
42. faustinus, Sauss. Antilles.
43. fluvialis, Streets. Coatzacoalcos river, among the Cordilleras.
44. forceps, M.-E. Rio de Jaueiro. — Aecording to von Martens, 1869, = acanthurus, Wgm.
45. formosensis. Bate. Eiver Tamsuy, Formosa.
46. gangeticum. Bate. Patna, India.
47. Gaudic/iaudii, M.-E. Chili.— Confer : von Martens, 'Ueber eiuige ostasiatische Siisswasserthiere/
1868, p. 65.
48. ffracilinianus, Rand. Sandwich Islands.
*49. gracilirostris, Miers. Upolu, Samoa Islands.
50. grandimanus, Rand. Sandwich Islands. — Confer : von Martens, /. c. 1868, p. 45.
51. heterochirus, Wgm. East coast of Mexico.
52. Hildebrandti, Hilgd. Madagascar.
53. Hilgendorfi, Cont. East coast of Madagascar, region of large forests.
*54. hirtimanus, Oliv.
*55. hispidus, Oliv.— According to Heller, ' Synopsis der im rothen Meere vorkommenden Crustaceeu,'
1861, this species occurs in the Red Sea.
56. Horstii, de M. Celebes, fresli water.
57. Idae, Hell. Borneo.
„ var. idella, Hilgd. Pond near Matomondo, Unguu ; Usaramo (German East Africa).
„ var. mammillodactylus, uov. var. ?, Thallw. North Celebes, Luzon.
„ var. subinermis. Nob. St. Joseph river, Innawi (British New Guinea).
38. Iheringi, Ortm. State of Sao Paulo, Brazil (fresh water).
59. jamaicensis, Hbst. Rivers of Jamaica.
60. japonicus, de Haan. Japan.
<jl. javaniciis, Hell. Java.
62. Jelskii, Miers. Guiana (Oyapok).
63. Lamarrei, M.-E. Coast of Bengal.
64. laminatus, Gollm., = jamaicensis, juv. ? Caracas. — Confer : von Martens, /. c. 1869, p. 24.
65. larapropus, de M. Celebes, Timor (fresh water).
66. lanceifrons, Dana. Luzon.
Q7. lar, Fabr. India.
68. latidactylus, Thallw. North Celebes.
69. latimanus. Marts. Philippines (Isle of Samar).
70. lepidaetyloides, de M. Flores (fresh water). — Confer: de Man, in 'Notes from the Leyden
Museum,' vol. xv. p. 308. — According to Coutiere = lepidactylus, Hilgd.
71. lepidactylus, Hilgd. Mozambique (Quellimane, Tette).
72. longidigitum. Bate. Habitat unknown.
73. longimanus, Fabr. East India.
74. longimanus, Htfm., = ornatus, Oliv. Reunion. — Confer: de Man, in * Notes from the Leyden
Museum,' vol. i. p. 172.
*75. lovgimmms, Phil., = rtnnentarius, Poepp. Chili (La Ligua river). — Confer : Philippi, in
' Zoologischer Auzeiger,' 1894, p. 266.
76. longipes, de Ilaan. Japan.
77. macrobrachion, Herkl. West Coast of Afiica (Boutry, near Dixcove).
78. madagascariensis, Hffm. Nossy-Faly.
*79. Malcolmsonii, M.-E. Nagpore. — Confer: Henderson, ' A Contribution to Indian Carcinology,'
1893, p. 444.
MOUNT EOEAIMA IN BKITISH GUIANA. 63
80. Malliardi, Rchtrs. Mauritius (Creole river. Black river).
81. Marite, Cout. River Ivaloiua, near Tamatave (Madagascar).
82. mayotteusis, liffm. Mayotte, Nossy-Faly. — Confer : de Man, in ' Notes from the Lcyden
Museum,' vol. i. 1879, p. 173, where it is proved to be a local variety of ornatus, Oliv.
83. mexicanus, Sauss. Coast of Mexico.
84. modestus, de M. Floras, fresh water.
85. MontezmncB, Sauss. Gulf of Mexico, off Vera Cruz.
86. Mooi-ei, Caiman. Lake Tanganyika.
87. mossambicus, Hilgd. Mozambique (Quellimane). — According to Coutiere = dispar, Marts.
88. multidens, Cout. River Kotofotsy, arm of the Onilahy, Madagascar.
89. Nuttercri, Hell. Brazil (Rio Negro).
*90. niloticus, Roux. Nile. — Confer : von Martens, /. c. 1868, p. 66.
91. nipponensis, de Haau. Japan.
*92. oliionis, S. Sm. Ohio, Mississippi.
93. 0/fersii, Wgm. Coast of Brazil. — Confer : Greeff, in ' Sitzuugsber. Gesells. z. Befdrdcrung der
gesammten Naturw. zu Marburg,' 1882, p. 30.
*94. ornatus, Oliv., = lar, Fabr.
95. parvus, Hffm. Nossy-Faly.
96. Patsa, Cout. River Mahauara (east coast of Madagascar) ; arm of the River Ouilahy (west
coast of the same island).
97. paucidens, Hilgd. Togo Country.
98. Petersii, Hilgd. Mozambique (Tette).
99. pilimanus, de M. Sumatra.
,, var. leptodactylus, de M. Java (Buitenzorg).
100. placidulus, de M. Saleyer, Celebes, Flores, Timor, fresh water. — Confer : de Mau, in ' Notes
from the Leyden Museum,' vol. xv.
101. placidu.s, de M. Sumatra.
102. potiporanga, F. Miill. Itajahy river.
103. potiuna, F. Miill. Itajahy river.
104. puncfatifs, Rand., — jamaicensis, Hbst. East Indies ? — Confer : Kingsley, in ' Bull. Essex
Institute,' vol. xiv. 1883, and Miers, in E. Whymper, Supplementary Appendix to 'Travels
amongst the Great Andes of the Equator,' 1888.
105. Quelchi, de M. Upper IMazaruni river. Mount Roraima (British Guiana).
106. reunionncnsis, Hfi'm. Rikmion. — Confer : de Man, in Max Weber's ' Crustacea,' 1892, p. 454.
107. Ritsema;, de M. Atjeh.
108. Rosenbergii, de M. Andai, New Guinea.
109. ruber, Hess., = ornatus, Oliv. Fiji Islands. — Confer: Ortmann, ' Decapoden-Krebse des Strass-
burgcr Museums,' p. 705.
110. rudis. Hell. Ceylon.
111. Savignyi, Bate. Bermuda Islands.
112. scabriculus. Hell. Ceylon.
113. sexdeututus, Streets. Tide-water of the Coatzacoalcos river, Isthmus o£ Tehuantepec. — According
to Ortmann (' Decapoden-Krebse des Strassburger Museums,' p. 711) this species is identical
with mexicanus, Sauss.
114. sinensis. Hell., = nipponensis, de Haan. Shanghai. — Confer: de Man, in * Notes from the Leyden
Museum,' vol. i.
115. singalangensis, Nob. Aier Manteior, near Mount Singalang (Sumatra).
116. sintangensis, de M. Sintang, Borneo.
6Ji EEPOKT ON A COLLECTION PEOM
117. spectabilis. Hell., = lar, Fabr. Tahiti. — Confer: de Mau, iu Max Weber's 'Crustacea/
p. 415.
118. spinimamis , M.-E. Antilles and coast of Brazil.— According to von Martens, I. c. 1869, = 01fersii,
Wgm.
119. sp., de Man, iu Zool. Jahrbiiclier, ii. 1887. Sydney.
120. sp., de Man, in Arcliiv fiir Naturg. 1888, p. 557. Amboina.
121. sp. (Macrobrachium ?), de Man, in Max "Weber's 'Crustacea,' 1892, p. 488. Celebes, fresh
water.
122. sp., Miers, in Ann. & Mag. Nat. Hist. ser. 5, v. p. 384 (1880). Java.
123. sp., Thallwitz, ' Decapoden-Studien,' 1891, p. 19. North Celebes.
124'. sundaicus, Hell. Java.
, var. bataviana, de M. Batavia.
, var. bracliydactyla. Nob. Amboina.
, var. de Mani, Nob. Amboina. — According to Nobili the last-named variety is identical
with that described by de Man in Zoolog. Jahrbiiclier, ix. Abth. f. System, p. 783, fig. 72.
125. superbus. Hell. Shanghai.
126. tenellus, S. Sm. Polvou, Occidental Department of Nicaragua.
127. tranquebaricus, Fabr. East India.
128. Trompii, de M. Ketoengau river, Mandai river, Sintang (Borneo).
129. ustnlatus. Nob. Rigo, British New Guinea.
130. vagns. Hell., = lar, Fabr. Amboina. — Confer : de ]\lan, in ' Notes from the Leyden Museum/
vol. i.
131. Vollenhovenii, Herkl. Coast of Guinea. — Confer : de Man, in ' Notes from the Leyden Museum,'
vol. i. 1879.
132. Weberi, de M. Celebes, fresh water.
MYRIOPODA AND ARACHNIDA.
By R. I. PococK.
Class DIFLOPOBA. (Millipedes.)
Pamily POLYDESMID/E.
Genus Odontopeltis, Pocock.
Odontopeltis Macconnelli, sp. D.
6 . Colour black or very dark blackish brown, with the external half of the keel bright
or dull red or yellowish brown, and, at least on the anterior terga, a median transverse
yellowish or red patch along the posterior border ; caudal process not pale ; autennge
blackish ; legs blackish brown or deep reddish brown, sterna brownish yellow. Dorsal
integimient smooth and shining or finely coriaceous ; keels horizontal, with smooth
edges, the posterior margin transverse and in the same straight line as the posterior
border of the tergum, as far back as the eleventh or twelfth somite ; the posterior angle
not spinate, mostly acutely angled, rarely approaching a right angle ; the anterior angle
widely rounded and obtuse. Caudal i^rocess triangular, narrowly truncate posteriorly.
MOUNT Et)llAIMA IN BRITISH GUIANA.
65
Anal sternite triangularly pointed ; sternum of eighteenth somite wider behind than the
length of the coxae of the last pair of legs. Copulatory limbs as in fig. 1, the basal
segment armed externally with a large downwardly-directed conical process; second
segment furnished with numerous thickly-set short hairs on the inner side at the base
and externally with many long bristles ; giving off distally two long processes, an upper
and a lower, directed obliqiiely forwards and downwards ; the upper process runs forwards
Fia;. 1.
Odontopeltis Macconnelli, sp. n.
(I. Lower view of left copulatory leg. h. Lateral view of external surface of right copulatory leg.
and downwards with a slight curve, then turns sharply externally, and ends in a sharp
tip curving upwards and forwards ; the inferior process is laminate, but narrower at base
and distally than in middle, with a slight sigmoid flexure when seen from the side ; seen
from below its inner edge is directed straight forwards in its basal third, then obliquely
forwards and outwards, the outer edge being convex ; distally the process ends in two
sharp prongs — an inner straighter and an outer semicircularly curved forwards.
2 . Stouter than male, with smaller keels.
rf . Total length 4.3 mm. ; width 6.
S . » 43 „ ; „ 7.
Loc. Summit of Roraima, 8600 feet alt.
Genus Euryurus, C. Koch.
EURYURUS ATRATUS, Sp. n.
2 . Colour : dorsal area a uniform black, ventral area a little paler. Head with a
smooth, oval, pale-coloured prominence in the middle line just between and below the
antennae ; a curved row of six setal pores above the labrum. Dorsal area smooth and
polished. Keel of second somite laterally emarginate, leaving the anterior imd posterior
angle acute and subdentate ; lateral margin of keel of third and fourth somites with a notch
behind the anterior angle, which is thus subdentate; remaining keels with posterior
margin finely serrulate, concave ; angle acute and becoming more and more acute and
produced in the posterior region of the body ; lateral margin of keels even, slightly
66
REPOET ON A COLLECTION FEOM
subsiniiate in front of the pores ; anterior angle couvexly rounded and on the anterior
somites slightly prominent. Caudal process narrowly oval posteriorly. Anal sternite
distinctly bituberculate.
Total length 41 mm. ; width 6-7.
Log. Base of Mount Pi.oraima (3500 feet).
Resembling in colour E. fumigatus, Peters, from Bogota (MB. Ak. Berlin, 1864,
p. 624), but differing from Colombian specimens in the British Museum that I have
rel'erred to fumigatus in having the lateral border of the keels of the second tergite
emarginate, the keels larger, and the caudal process narrower towards the extremity.
Class ABACENIDA.
Order ABANE^. (Spiders.)
Family BARYCHELID^.
Genus Cyrtogkammomma, Poc.
Cyrtogrammomma monticola, Poc.
Cyrtogrammomma monticola, Poc. Ami. & Mag. Nat. Hist. (6) xvi, pp. 139-140 (1895).
Loc. Summit of Mt. Boraima (8600 feet).
This genus and species were based upon a single female specimen. Additional
examples containing adults of both sexes enable me to supjilement the original
Fig. 2.
«. Extremity oi palpus of malo of Cyrlogrammomvni monticola. h. Eyes of ditto, c. Vulva of Anyphaina QuelcMi.
(I. Tibial spur of palpus of same. e. Tibial spur of pnlpus of Trcchahct MacconnelU.
MOUNT EOEAIMA IX BRITISH GUIANA. 07
description by stating the cliaraeters of the male and some additional features of the
female.
<^ . Colour, as in $ , a deep blackish brown ; hairs mostly olive-black, those on the
carapace, especially laterally, shining golden red, extremities of protarsi on upperside
also paler. Carapace flatter than in 2 , its length equal to patella -f three-fourtlis of
the tibia of the first and fourth leg, barely equal to protarsus of fourth. Labium (in s
and $ ) armed with a row of cusps varying in number from al)out four to nine ; maxilh«
also internally cuspidate. Palpus in <3 with the tibia almost as in 2 , but armed on
the inner side with six or more long strong spines, tliosc at the distal end protecting the
palpal organ when lying at rest backwards beneath the tilna ; tarsus long, like that
of the 9- , but scopulate only in its distal half beneath, excavated posteriorly ; spine
of palpal organ nearly straight, long, almost as broad at the base as the bulb and taperiu"-
aAvay to a fine point. Tibia of first leg without apical spur ; tibia of all the leo-s armed
with many long and strong spines : protarsus of first leg armed with two inferior spines
at base of scopula, which covers less than the ajiical half of the segment ; protarsus of
second armed with three spines, scopula short and scanty as in first leg ; protarsi of third
and fourth armed with many long and strong spines. (In the $ the anterior legs and
palpi are armed only with spiaiform setie and the scopuhe extend practically to the base
of the protarsi.)
Measurements in millimetres. — 6 . Total length 12 ; length of carapace G-5, of 1st leg
23, of 4th leg 26.
Family AXYPH^XID^.
Genus Anyph-«na, Sund.
ANYPH.EXA QUELCHII (Poc).
Aysha Quelchii, Pocock, Auu. & Mag. Nat. Hist. (6) xvi. pp. 140-142, fig. (1895).
Loc. P^oraima (summit, 8600 feet). Adult males and females.
On his previous expedition to Roraima, Mr. Quelch procured the only two specimens
of this species, an adult male and an immature female. I take the opportunity of
figuring the vulva (fig. 2 c) of the adult female and the tibial spur of palpus of the
male (fig. 2d).
Family PISAURID/E.
Genus Trechalea, Thorell.
(= Trichiria, V,. Koch.)
Trechalea Maccoxxelli, sp. n.
Colour. Carapace blackish, with pale lateral border and narrow radiating pale stripes,
the intervals between the radiating lines also ornamented with short flavous lines.
Clypeus pallid, with a broad dark baud on each side, running upwards and inwards from
the lateral angle; mandible black, clothed with loug yellow hairs, the yellow field
SECOND SERIES. — ZOOLOGY, VOL. VIII. \{)
G8 EEPOET ON A COLLECTION FEOM
interrupted by dark lines. Legs and palpi blackish banded with pale ; femora with
about four flavous patches, sometimes fusing together or connected with flavous stripes ;
patella distally flavous ; tibia with median and apical flavous stripe ; protarsus with broad
but indistinct distal flavous stripe ; femora reddish below at the base ; sternum and coxae
olive-yellow ; upperside of abdomen covered with greyish hairs intermixed with black
bristles ; black behind, with two bright yellow patches above the spinners ; lower side
orange-yellow.
Palp with tibial spur as in fig. 2 e.
Measurements in millimetres. — Total length 16, carapace 8'5 ; 1st leg 64, 4th leg 73;
tibia of 1st leg 17, of 4th 17.
Log. Mount Roraima (base, 3500 feet).
This species may be readily recognized by the form of the tibial spur of the palp.
Family AUGIOPIDJ^.
Genus Acrosoma, Perty.
ACROSOMA SCHRIEBERSII, Pcrty.
Acrosoma Schriebersii, Perty, Delect. Anim. Artie, p. 19i, tab. 38. fig. 9 (1830-34).
Loc. Base of Roraima, 3500 feet.
Genus Araneus, Linn.
Araneus audax, Blackwall.
Ejmra audax, Blackwall, Ann. Mag. Nat. Hist. (3) xi. p. 29 (1863).
Loc. Upper Mazaruni River.
Order SCORPIONES.
Eamily CHACTID^.
Genus Broteochactas, Poc.
Brotvochactas and Hadrurocliaclas, Pocock, Ann. Mag. Nat. Hist. (G) xii. pp. 77-78 (1893) ; id. Joui'u.
Linn. Soc, Zool. xxiv. p. 399 (1893) ; Kraepelin, MT. Mus. Hamb. xi. pp. 175, 178; id. Das
Tierr., Scorp. etc. p. 172 (1899).
Four species referable to this genus have hitherto been discovered, namely :—
1. Broteochactas Gollmeri, Karsch, Mitth. Miinch. cut. Ver. 1879, p. 133 (Chactas): nitidus,
Pocock, Jouru. Linn. Soc, Zool. xxiv. p. 399, pi. xxix. figs. 7,7 a (1893) {Broteochactas) .
Loc. Trinidad and Venezuela.
2. Broteochactas delicatus, Karsch, Mitth. Miinch. ent. Ver. 1879, p. 134, ? : opacus, Karsch,
op. cit. p. 134, (j yChactas) ; panamensis, Thorell, Bull. Soc. Ent. Ital. xxv. p. 27 (1894)
{Broteiis) .
Loc. British Guiana, Colombia, Panama.
MOUNT EOEAIMA IN BKITISH GUIAXA. 69
3. Broteochactas Sclateri, Pocock, Ann. Mag. Nat. Hist, (fi) xii. p. 80 (1893) [Hadrurochactas) :
? Schaumii, Karscli, Z. Natiirw. liii. p. JOfi (1880); ? quinquedentutus, id. op. cit. ]>. lO.'i
Loc. British Guiana [W. L. Sclater).
[Kraepclin (MT. Mus. Hamb. xi. p. 178, 1894, and Das Tierr., Scorp. etc. p. 173, 1899) states that
B. Sclateri is identical with B. Schaumii of Karsch, and possibly with quinquedentatus of this latter author,
both of which were recorded, though no doubt erroneously, from India. But the presence of only
five pectinal teeth in tlie hatter forbids, to my mind, such an opinion. B. Schaumii, judging from
Kraepelin's description, is closely allied to B. Sclateri, but the fact that the third caudal segment is higher
than wide la Schaumii, and wider than high in the two known examples of Sclateri, makes the synonymy
doubtful.]
4. Broteochactas parvulus, Pocoek, Ann. Mag. Nat. Hist. (G) xix. p. 3G4 (1897); Kraepelin, Das
Tierr., Scorp. etc. p. 174 (1899).
Loc. Amazons, Sautarem [F. O. P. Cambridge).
The following -well-marked new species were discovered by Messrs. McConnell and
Quelch : —
Bkoteochactas grakosus, sp. n.
Colour. Carapace, terga, and tail almost black ; legs, palpi, and vesicle of tail deep
reddisli brown, the legs with paler line and spots ; fingers black. Carapace and ferga
entirely covered with line close-set granulation ; ocular tubercle coarsely punctured ;
carapace a little longer than the first and second caudal segments, a little shorter than the
fifth. Coxce and sterna finely punctured, the last sternite weakly and closely granuhu"
laterally. Tail more than four times the length of the carapace, narrowed jjosteriorly,
the segments nearly parallel-sided : the first wider than long ; second slightly longer than
wide ; fourth not twice as long as wdde ; fifth a little more than twice as long as -wide ;
intercarinal spaces finely granular ; inferior and infero-lateral keels obsolete on segments
1 and 2, scarcely traceable on segment 3, represented on segment -l by iri-egularly
arranged larger granules ; stxperior and supero-lateral keels weak, weakly granular ;
inferior surface of fifth coarsely granular between the keels; vesicle granular, narrower
than third segment, -wider than high.
Chelce finely granular; upperside of humerus and Ijrachium with coarser granules
between the keels ; upperside of hand covered -with a reticulation of fine granules; inner
surface similarly granular ; back of hand coriaceous ; fingers granular at base. Width of
liand equal to length of external keel of hand-back, less than length of movable digit ;
digits longish, movable as long as the carapace, immovable about twice as long as its
basal width. Femora and tibiae of ^^^* granular, the granules intermixed with punctures
on the tibia, following segments closely punctured ; tarsi short, not twice as long as
high, convex above, incrassate distally, armed with long sette arranged more or l(!ss
regularly in two rows.
Measurements in millimetres. — Total length 46 ; length of carapace 6"5, of tail 29'.j ;
width of 1st segment 3-5, of 5th 2'8 ; width of hand ou ; length of hand-back .5,
of movable finger 8-5.
Loc. Base of Mt. Roraima (3500 feet alt.).
10*
70 EEPOKT ON A COLLECTION FROM
. BrOTBOCHACTAS POROSUS, sp. 11.
Colour. Trunk and tail blackish brown ; fourth and fifth segments of tail and vesicle
paler, reddish; legs also reddish broun, cheloe with humerus and fingers blackish,
hand and brachium redder.
Carapace, except on the normally smooth tracts, very distinctly punctured, especially
the area around and between the eyes, with very tine close-set granulation on its lateral
slope. Terga similarly jiunctured, witli a few very fine granules in front and on the
sides, the granules and punctures scarcely distinguishable with a hand-lens ; the tergum
of seventh abdominal somite much more distinctly granular, with larger granules along
the lateral border and two series forming indistinct crests on each side. Sterna punc-
tured, the fourth and fifth mure closely than the others, the third with a distinct smooth
patch in the middle of its posterior half.
Tail barely four times as long as the carajmce ; carapace as long as first, second, and half
the third segments, and about as long as the fifth ; scarcely narrowed posteriorly ; all the
segments wide, the second nearly twice as wide as long, the fourth about as wide as long ;
fifth oue-tliird longer than wide, abruptly narrowed behind; segments punctured through-
out ; the superior, supero-lateral, and infero-lateral keels strong and grantilar ; the inferior
median crests almost obsolete on segment 1, more evident on segments 2 and 3, but
represented by irregularly arranged granules ; on segment 4? the granules assume a more
definite, but still incomplete arrangement in two parallel rows ; median lateral keel
present on segment 1, represented by a few granules on segments 2 and 3, absent on
segment 4 ; fifth segment with its upper edges granular and sharp, a distinct median
lateral keel in the anterior half of the side, and three distinct and granular inferior keels,
the area between them also serially granular. Vesicle narrower than the tail, wider than
high, punctured but not granular below.
CheUe punctured even to the tips of the fingers; humerus with upper keels granular;
brachium with upper anterior keel obsoletely granular ; hand wide, distinctly though
not strongly carinate, not granular, except slightly so on the inner surface ; width of
hand about equal to length of hand-back; fingers long and slender, in contact, the
movable as long as the carapace, nearly twice the length of the hand-back.
Legs punctured, not granular; tarsi furnished beneath with long, close-set, irregularly
arranged bristles ; the fourth tarsus long, lightly convex above, about three times as long
as high.
Pectinal teeth 10 ( d 2 ); the teeth longer in d .
Measurements in millimetres. — Total length 24 ; length of carapace 3'5, tail 15.
Luc. Summit of Mt. Eoraima (8600 feet).
Some of the distinguishing features of these two species of BroteocJiactas are set forth
in the following table : —
a. Infero-lateral crests on all tlie caudal segments strong, as strong as the supero-
lateral; inferior medians present and granular; the integument punctured
throughout ; terga in male mostly weakly granular ; vesicle smooth ; hand
with two finger-keels, smooth, more globular; fingers long and slender,
movable as long as carapace. (Of small size, 25 mm. in length.) porosus, sp. n.
MOUM EORAIMA IN BEITISH GUIANA. 71
//. Infero-lateral crests on anterior three caudal segments obsolete ; inferior medians
absent; dorsal integument not noticeably punctured ; terga closely granular
throughout in male ; vesicle granular ; hand granular, less globular, with
compressed inner edge, without finger-keels.
a'. Carapace {^) entirely covered with close-set granules; sterna minutely and
closely punctured througliout ; lower surface of tail finely and closely
granular and punctured ; legs also densely and closely punctured ; fingers
longer, the movable as long as carapace, immovable more than twice as long
as its basal width. (Of large size, 46 mm. in length.) ffranosus, sp. u.
b\ At least the upper portion of the carapace smooth ; sterna and lower surface of
anterior segments of tail smooth aud polished, not punctured or granular j
legs mostly smooth aud polished ; femora at most weakly granular ; fingers
shorter, movable shorter than carapace, immovable not twice as long as
its basal width.
«'. Tail very thick, width of first caudal segment considerably exceeding the
width of the hand, and equal to length of first and second caudal segments
taken together ; hand very smooth, rounder, its inner edge less compressed,
with keel of underhand obsolete; tarsi longer, more thickly covered below
with long hair Scluteri, miiii.
b". Tail much thinner, width of first segment generally much less tlian, rarely
equal, to width of hand, aud less than sum of lengdi of first and second caudal
segments ; hand at least with its inner edge granular and subcom pressed;
tarsi shorter and more scantily clothed with shorter hairs . . Golliacri, delicatus, parvulus.
(For tabulation of the characters of the last three species, see my paper in Ann. Mag.
Nat. Hist. (6) xix. pp. 365-3G6, 1897 ; and Kraepelin, Das Tierr., Scorp. etc. p. 173,
1899.)
HYMENOPTERA, HEMIPTERA HETEROPTERA, HOMOPTERA,
NEUROPTERA, AND ORTHOPTERA.
By W. F. KiEBY, F.L.S., F.E.S.
(Plate 6. figs. A, B.)
The few specimens of these Orders which have been submitted to nae for identification
consist almost exclusively of common and well-known South American species. A few
species, mostly immature, are hardly in a condition to be determined with certainty.
Tliese are one Forficulide, three Blattidoe, and one Pentatomide. One Hemipteron
I have described as new.
The two species noted from the greatest elevation {Folistes annularis and Sympetrmu
gilvum) are southern representatives of forms found in North America.
72 llEPOET ON A COLLECTION FEOM
HYMENOPTERA ACULEATA.
APIDJE.
MeLIPONA INTERHrPTA
Melipona interrupta, Latr., Humb. &: Bonpl. Yoy. i. p. 2^J\, pi. 20. tig. 3(1811).
Melipona fasciculata, Smith, Cat. Hyiii. Ins. Brit. Was. ii. ]). 40G, ii. 25 (1854).
Horaima, 3500 feet.
VESPID^.
POLISTES ANNULARIS.
Vespa annularis, Johansson, Amoen. Acad. vi. p. 413. n. 93 (17G3).
Roraima, 8600 feet.
MUTILLID.E.
MUTILLA LARVATA.
Mutilla larruta, King, Nova Acta Acad. Leop. x. p. 310, pi. 22. fig. 0 (1821).
Roraima, 3500 feet.
Mutilla quadrum.
Mutilla quadrum, King, Nova Acta Acad. Leop. x. p. 320, pi. 23. fig. 8 (1821).
Roraima, 3500 feet.
EORMICID.E.
DOLICHODERUS BISPINGSUS.
Myrmica bispinosa, Oliv. Enc. Meth. vi. p. 502. no. GO (1791).
Roraima, 3500 feet.
A considerable number of specimens.
HEMIPTERA HETEROPTERA.
COREID^.
ACANTHOCEPHALA SURATA.
Diactor surafvs, Burm. Handb. Ent. ii. (1) p. 334. n. 2 (1835).
Roraima, 3500 feet.
REDUVIIDiE.
AcROCORis PERARMATA, sji. n. (PI. 6. fig. A ; B, profile of head.)
Long. Corp. 27 millim.
Male. Ru.fo-testaceous, clothed with a fine grey pile, all the spines tipped with reddisli.
Head long, the part behind the eyes slightly longer than that before ; two long pointed
spines near together just behind the antennie ; antcnnte with the first joint very Ions:,
MOUNT KORAniA IN BRITISH GUIANA. 73
blackish, with two whitish bands near each extremity, second joint about \ as long as
the first, whitish, with the extremity blackish ; third joint about as long as first, blackish
towards the l)ase, and yellowish beyond, blackish again at its junction with the fourth
joint, which is yellowish and about as long as tlie second. Front lobe of thoi-ax with two
strong erect spines at the back ; middle lobe with four, two in the middle, and two lateral,
all at about equal distances apart. Clavus brown. First joint of rostrum yellowish, the
remainder deep black, the tip extending just beyond the base of the head. Legs
unarmed, thickly pubescent, more or less blackish on the outer side ; tarsi black.
A])dorninal segments with four small lateral spines on the basal half, and three long,
triangular, whitish, lateral spots on each side beyond the middle. Scutellum with a
very slight terminal spine, if any.
Roraima, 3500 feet.
This curious insect does not seem to have much resemblance to any described species
HEMIPTERA HOROPTER A.
FULGORIDyE.
acriephia perspicillata.
Cicada perspicillata, Fabr. Spec. lus. ii. p. 3.22. u. 1 (1781).
Roraima, 3500 feet.
NEUROPTERA ODONATA.
LIBELLULID.E.
Sympetrum gilvum.
Dip/ax illotum, var. gilva, De Selys, Ami. Soc. Eat. Belg. xsviii. p. 43 (ISSi).
Roraima, SGOO feet.
O R T H 0 P T E R A.
LOCUSTID.E.
Chromacris speciosa.
Gryllus speciosus, Tluaib. IMtjm. Petersb. ix. p. 40, pi. 11. fig. 1 (1821).
Roraima, 3500 feet.
EXPLANATION OF PLATE 6,
Fig. A. Acrocoris perarniata, sp. n.
Fig. B. „ „ profile of head.
74 EEPOET ON A COLLECTION FEOM
COLEOPTERA.
By C. O. Waterhouse, V.P.E.S.
CARABID^.
OxYCREPis LEUCOCERA, Lacord.
A single example.
DYTISClDiE.
Rhantus elegans, "Waterli.
Several specimens found at an elevation of 8600 feet. A single example only was
obtained on a former occasion.
LUCANIDiE.
Charagmophorus lineatus, Waterh.
This genus and species were described from a single male example. Other males and
two females have now been found at 8600 feet.
The female has the elytra as in the male, with lines of very small grey scales ; but
tlie head and thorax are shining. The mandibles are short. The head is sparsely
punctured posteriorly, rather strongly transversely impressed in front and strongly and
closely punctured. The thorax is rather more convex than in the male, a little narrowed
anteriorly, ol)tusely angular at the sides at a short distance from the base, moderately
finely ptmctvired, the punctures rather unequal, not very sharply defined, separated
from each other by two to three diameters of a puncture ; the margins are impressed ;
the disk is longitudinally impressed. The front tibia? have the five teeth rather stronger
and more approximate than in the male ; the posterior tibiae have a small acute tooth at
the middle.
DASCILLIDiE.
ExAGONTUS, gen. nov.
Mentum a little broader than long, slightly narrowed anteriorly, corneous ; ligula
broad and transverse, acuminate at the sides ; labial paljii three-jointed, the basal joint
elongate, the second a little shorter, pear-shaped, the third somewhat the same shape but
inverted, acuminate at the apex. Maxilloe with two delicate subequal lobes ; the galea
consists of two portions, the basal part parallel, the apical portion shorter, clothed with
stiff hair ; the lacinia terminates in a curved acute tooth, which is surrounded by curved
stiff bristles. Maxillary palpi rather long, robust, hairy ; the basal joint rather short,
narrowed at its base ; the second joint stouter, at least twice as long as broad, gradually
and not very much narrowed towards the base ; the third joint similar in shape but a
little shorter ; the fourth a little longer than the second, club-shaped, acuminate at the
apex. Mandibles strong, curved, concave below, very acute at the apex, with a small
MOUNT EOEAIMA IN BRITISH GUIANA. 75
tooth about the middle. Labrum rather large, siibquadrate, rounded in front. Head
convex, deflexed, but in no way covered by the pronotum, parallel behind the eyes,
narrowed in front. Eyes rather prominent, coarsely facetted. Antennae placed a little
in front of the eyes, widely separated at their base, eleven-jointed, of moderate length,
of nearly equal thickness throughout, except the slender third joint; composed of
cylindrical joints, clothed with stiff pubescence ; the second joint globoss, the third joint
very narrow at the base, the following joints gradually narrower towards their bases.
Thorax strongly transverse, the side with a strong tooth-like prominence about the
middle. Scutellum triangular. Elytra broader than the broadest part of the thorax,
one-third broader than long, flattened dorsally; tlie surface uneven, irregularly
punctured.
Prosternum much reduced, with a diamond-shaped process between tlie coxae, the
coxal cavity completely open posteriorly.
Mesosternum slightly inclined, with a slight, sharply margined concavity in which the
prosternal process rests.
Metasternum rather short ; the episterna broad, a little narrowed posteriorly.
Abdomen composed of five visible segments below. Anterior coxae strongly transverse ;
intermediate coxae globose ; posterior coxae very narrow externally, very Avide internally.
Tibial spurs small but distinct. Tarsi five-jointed ; the basal joint nearly as long as
the two following taken together ; the fourth the broadest, concave above, so that it has
a tendency to be bilobed, clothed with soft pubescence below ; fifth jomt not very long,
with divaricating claws.
1 have some doubt as to the affinities of this genus. The structure of the antennae
points to affinity with the Ptinidae, near Hedobla ; but the broader, flatter form,
the freely exposed head, the transverse anterior coxae, and internally dilated postei'ior
coxae would place it in the Dascillidte, where I now joropose to place it. No doubt the
Ptinidae and Dascillidae should be placed nearer together than is usual in collections.
EXAGONTUS DENTICOLLIS, Sp. n.
Elongato-oblongus, parum convexus, fusco-castaneus, sat nitidus, brevissime griseo-
pubescens ; capita nigrescente, crebre punctato ; thorace utrinque dente valido
instructo ; elytris crebre pmictatis, pube gisea A'ariegatis, impressionibus nonnuUis
notatis.
Long. 3i, lat. 2 mill.
The antennae are modei'ately robust, the third joint more slender, the fourth to tenth
joints a little longer and broad, cylindrical, united to each other by their centres ; the
eleventh joint a little longer, elongate-oval. Apical joint of the maxillary paljji black.
The elytra are brown, closely punctured, with numerous lines of greyish-yellow
pubescence giving a mottled appearance. Each elytron has a large transverse imj^ression
below the scutellum, another about the middle, one below the shouldei' ; the surface of
the apical portion is uneven, and in certain positions three slight interrupted costae may
be traced.
SECOND SERIES. — ZOOLOGY, VOL. VIII. 11
76 ON A COLLECTION FEOM MOUNT KORAIMA.
TENEBRIONIDvE.
Cyrtosoma montanum, sp. n.
Oblonguni, nitidum ; capite, thorace femoribusque fere nigris ; elytris seneis, teauiter
striatis ; ore, autennis, tibiis tarsisque piceo-flavis.
Long. 11, lat. 5| mill.
Head finely and rather closely punctured. Antennae with the six terminal joints
gradually wider. Thorax finely but distinctly and rather closely punctured, with a very
slight, transverse impx-ession above the anterior angles and above the posterior angles ; the
sides with tAvo obtuse not very prominent angles. Scutellum pitchy. Elytra rather wider
than the thorax, one-quarter longer than broad, rather straight at the sides, obliquely
narrowed at the apex, finely striated, the striae indistinctly punctured, the interstices
scarcely convex on the back, but at the apex and at the sides they are slightly angularly
raised in the middle ; the fourth and fifth strise unite posteriorly about one-third from
the apex, the third and fifth unite nearer the apex, and the second and sixth within the
apical angle. The underside of the insect is for the most part pitchy red, shaded with
black on the metasternum and abdomen.
LAMIIDiE.
Alcidion sexnotattjm, sp. n.
Elongatum, angustum, fuscum, pube grisea vestitum ; thorace basi constricto ; ely tris
fasciis punctisque numerosis fuscis ornatis, singulo elytro ad apicem oblique
truncate, angulo saturali obtuso, angulo externo rectangulare ; antennis gracilibus,
corpore multo longioribus. d .
Long. 11, lat. 4 mill.
Compared with the majority of the species of this genus, this is very elongate and
narrow. The antennae are very long ; the basal joint is much narrowed at the base,
then quite straight and parallel to the apex ; the joints are tipped with black and have
some black pubescence, which forms a slight tuft at the apex of the third joint. The
thorax is short and transverse, lightly impressed on the disk, arcuately narrowed
anteriorly, slightly constricted at the base ; the middle of the disk and three or four
small spots at the sides are brown. The elytra have scarcely any trace of costae ; the
basal crest is well marked but not acute ; the basal area is brown, with two vague
oblique vittae of ashy pubescence marked with brown punctures ; at the middle there is
an irregular ashy fascia (descending at the suture) marked with brown punctures ;
behind this is a fascia formed by five elongate spots placed side by side, the sutural one
lower down than the others ; in the apical area there are six brown spots, three of which
are placed so as to make V I on the left elytron.
The following species, not of very special interest, were found at an elevation of
3500 feet :—
Passalus transversm, Dalm. ; Antichira dichroa, Mannerheim ; Pelidnota Icevissima,
Burm. ; .Pyrojihorus uoctiluca, Linn. ; Strongylium luemorrhoidalis, Eabr. ; HeiUpus
carinirostris, Schonh. ; Cratosomus suhangulatus, Sclionh. ; Sphenophorus hemipterus,
Linn. ; Trachyderes interruptus, Dup. ; and Jamesiu globifem, Eabr.
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[VOL. VIII. PAKT 3.
THE
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THE LLNNEAN SOCIETY OF LONDON.
ON THE STKUCTUEE ANH AFFINITIES OF ECHIDKOS UNICINCTOS.
13Y
A L I C E L. E M B L E T O N, B. Sc.
(Communicated by Piot". G. B. Howks, Sec. Linn. Soc.)
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[ 77 ]
III. Oh the Structure and Affinities of Echiurus unicinetus.
By Alice L. Embleton, B.Sc. {Communicated by JProf. G. B. Howes, Sec. L.S.)
(Plates 7 10.)
Read Ttb June, 19(i0.
J HE observations recorded in this paper were made by me in the Zoological Laboratory
of the Royal College of Science, London, during January, Pebruary, and March 1900,
under the direction of Professor Howes and Mr. M. E. "Woodward, to both of whom I
owe a debt of gratitude for their generous help and advice.
The material was obtained by Professor Howes from Professor Mitsukuri, of Tokyo,
through the kindness of Mr. H. Lyster Jameson, who had intended to rejiort upon it in
his recently published paper in the Naples ' Mittheilungen,' but for want of time had been
prevented from so doing.
The specimens, numbering between 90 and 100, were in alcohol, and had been
preserved in corrosive sublimate. On the whole they were in very good condition,
though much contracted, resiilting in great variability of shape both as regards the
entire body and the different organs ; in several individuals, parts of the alimentary canal
were forced out through bursts in the body-wall, presumably caixsed by sudden and
violent contraction. To a large extent these post-mortem changes obscured many
observations which, on fresh or uncontracted material, could have been made with
greater ease and certainty.
Doubtless this is the species mentioned — though not named specifically — by Willemoes-
Suhtn * as occurring on the Japanese coast, where apparently it lives in great abundance,
for he says : " Ein Echiurid der den Eischern als Koder dient und wohl in Schlamm
dicht am Ufer vorkommt. Der S-Jt Zoll lange Wurm stimmt ganz mit den Merkmalen
der Gattung Echiurns iiberein, hat aber hinten nicht zwei Hakenkriiuze, sondern nur
einen."
The average length of the body is about 7 oi' 8 cm. ; all the specimens are narrowest
at the posterior end, widening out gradually, as a veritable sac, between this and the
])rol)oscis. The term " proboscis " is used, though that organ is, in all, represented
merely by a bluntly-pointed prte-oral lobe of triangular outline (PI. 7. figs. 1-2, pi).) ;
it cannot J^e doubted, however, that this is due to its state of contraction, for in cutting
a series of microscopic sections of the anterior end of the body there was nothing
to suggest that the proboscis was missing, as is so often the case with the Bintish
iovm, Echiurus Fallasid {Guevin), called "E. vulgaris" hy Eorbes, Eorbes & Goodsir,
Sars, O. Schmidt, and Metzger.
* " Von der Vhallenr/er Expedition, lirieff von K. v. Willemoes-Suhm an C. Th. v. Siebold, vii.,'" Zeitschr. wiss.
Zool. Band xxvii 1876, p. cii.
SECOND SERIES. — ZOOLOGY, VOL. VIII. 12
78 ALICE L. EMBLETON ON THE STKUCTUEE AND
The only previous description of this species is that given by Drasche *, who had but
two specimens to work on, with the result that his observations are superficial and
limited to three imges. Of his specimens be records : " Beide Exemplaren fehlte der
Kopflappen (Riissel)." It may be that, having so little material, he was misled by the
highly retracted state of this organ, wliich, possessing as it does a very powerful
musculature, is very liable to extreme contraction. Even allowing for the contracted
condition, this proboscis is a much shorter organ than in Echlurus Pullasli, or Thalassema
neptuni, and never assumes the almost ribbon-like form it presents in the latter
species.
At the base of the proboscis-lobe is the mouth, placed ventrally (Pi. 7. figs. 1, 2, m.).
The whole outer surface of the body is densely papillated (PL 1. figs. 1, 2, 4), agreeing
in this respect with other species of Echiurus; the papillae show no very definite
arrangement, though they tend to run in transverse rows round the body, in wliich
direction also the individual papillae are elongated (PL 7. fig. J.).
As in allied forms, anteriorly on the ventral surface are two recurved setae (PL 7.
fig. 1, s.a.), pointing outwards and backwards ; posteriorly are the characteristic peri-anal
hooks (fig. 1, s.p.), which in this species form but a single circlet. Drasche gives
this as one of the three distinguishing features of the species — hence its name
" unicinciiis" or "one girdle." In speaking of this circlet of setse, however, he says
it is made up of 11 setaj ; but on examining my material the number seems to be by no
means constant, varying indifferently from 10 to 13 (PL 7. figs. 1, 5). The space
between the two setae situated most ventrally is always greater than that between any
other tw^o, giving it the appearance of an incomplete ring — though Drasche has observed
that the ring is complete ventrally.
Just behind the two anterior hooks, on the ventral side, it is possible to see with a
hand-lens two pairs of minute pores among the papilke ; these are the external apertures
of the segmental organs or nephridia. The reproductive prodvicts are passed to the
exterior through these pores.
Referring to the papilhe, Drasche says : " Nur an einer Stelle etwa 5 mm. hinter den
Bauchborsten, findet sich eiu 5h mm. breiter Giirtel von 13 Papillenkrauzen. Diese
Papillen sind etwas grosser als die iibrigen und habeu die Porm von auf die schmale Seite
ijestellten Ziegeln."
So far as my specimens are concerned no such tract is discernible, though on
submitting the body-wall to microscopic excxmination it is found that in the region of
the segmental organs the ordinary unicellular glands occur along with immense
compound glands (which will be more fully dealt with below). This particularly
glandular belt may correspond to the 13 rows of large papillte mentioned by Drasche
(PL 7. figs. 6, 7, £r.).
Probably the aoimal secretes a substance with which it lines the tubes or burrows in
\\hich it lives, using the two recurved anterior setae in climbing and its posterior circlet
* "Ueber ciue ueue Echiurus-A-ti aus Japan," Verhandlungen des zoologisch-botanischen Vereins in Wien,
Band sxx. 1880.
AFFINITIES OF ECHIURUS UNICINXTrS. , 79.
to liold itself in its tunnel. The setae have a significantly abundant supply of powerful
muscles (fig. 12), and the wall of the posterior end of the body is mucli thicker than
elsewhere, on account of the comparatively disproportionate development of the layer of
longitudinal muscles (PL 7. fig. 9j.
Setce. — Tliese agree on the whole with Spengel's description of those in Echinrns
Pallasii. TJiere is, however, some slight difference observable between the setsc of the
anterior pair (PI. 7. fig. 3) and of tlie posterior circlet ; the former ai-e large and very
markedlv recurved, while the latter are smaller and liner, with but a slisrht curvature.
The anterior seta? project into tlie body-cavity vei-y conspicuously, and possess numerous
strong radiating muscles (PL 7. figs. 5, 12; PL 10. fig. l-O). In minute structure each
seta presents a pointed, somewhat flattened free end ; about a third of its length back
there is a circular constriction, after wliich the seta widens out again to its blunt
extremity, which lies embedded in muscle and a protoplasmic mass from which it
originated, and in which young setae can 1)3 semi. The chitin of whicli tlic liook is
composed exhibits a transverse striation or annulation, less marked at the pointed end.
GreeflP says there is a fine canal in the suljstance of the seta at the free end running
a sliort way back ; this does not appear to be a canal but to be due to striation, as is so
often seen in the ordinary setae of Chajtopoda.
Bodij-waJl. — The body-wall varies in thickness in ditfereut regions, being thinnest
where the body is widest, and thickest at the tapering posterior extremity. It is not
unlikely that this is in part caused by differences in the state of conti'action, but this
alone could not account for the very striking divergence in the ])roportionate depths of
the respective muscle-slieatlis [cf. PL 7. figs. 6, 8, 9, 10).
Microscopic sections of the liody-wall serve to show tliat, in general histological
structure, it is almost identical with that of Echiurtts Pallasii as described by Greeff and
Spengel. The ejndermis is coA(ned by a well-defined cuticle, which is typical of the
whole group; in microsco2)ic preparations this appears as a sharp dark line running
along the outer border of the section, pierced by the skin-glands; and Ijeneath tliis
clearly marked band, and abutting on tlie epidermal cells, a vacuolated layer presents
itself, similar to that figured by Jameson * in Thalassema neptuni (PL 7. figs. 6, 7,
8, 9, 10).
The epidermis is composed of a single layer of columnar cells, but it is difficult to
distinguish them as such, for they are much distorted by the pressure of the numerous
unicellular glands and clear cells ; the epidermal cells give the appearance of a clear
matrix in Avhich these darkly staining bodies S(;em to be embedded. The nuclei of the
epidermal and gland-cells lie at their inner ends, and, as they stain almost as strikingly
as tlie glandular secretion, they form a dark border in section (PL 7. figs. 0, 7, 8, 9, 10).
Occasionally the unicellular glands assume giant proportions, become filled willi
secretion, and might well be mistaken for compound glands (PL 7. fig. G, (j.u.). In the
depressions between the papillae there are neither gland- nor " trigger-cells '' ; these
latter are seen scattered among the glandular tissue, being most noticcjible on the
proboscis (PI. 7. fig. 11).
* "Contributions to the Anitomy and Histologj- of Thalassema nepiani, Gaertner," /ool. Jahrb. lid. xii. 1899.
12*
80 ALICE L. EMBLETON ON THE STEUCTURE AND
Though on the whole, as is seen, the structure of the body-wall in this Japanese
species is largely identical with the descriptions given for allied types, yet in one
important point this EcJdurus differs from others in the group, for it possesses not only
unicellular glands but also large compound glands. These occur in immense numbers
but are strictly local in their distribution, being restricted to a wide belt in the anterior
half of the animal, coincident with the position of the segmental organs. They are very
conspicuous, being the most significant structures in sections of the body-wall, whether
they be taken in a transverse or longitudinal direction. As PI. 7. figs. 6, 7 ig.) show,
these glands are flask-shaped bodies, built up by radially grouped gland-cells {g.c) with
delicate walls ; the secretion (*.) from each component cell is poured into the main duct,'
which is lined by the cuticle. The nuclei (iic.) of the gland-cells collect at the basal
ends, and, being large, they make a sharp outline of the flask-like character of the whole
,organ (PL 7. figs. 6, 7). These glands are large and sink far down into the cutis (or
connective-tissue strata) {t.c), occasionally coming into contact Avith the outer circular
muscle-sheath (m.c). Invariably the ducts of these glands discharge on the summit of
a papilla ; and, seeing they are found in no other portion of the skin except near the
■segmental organs, it is not improbable that their secretion has some function to perform
in connection with the ripe eggs, which pass out by the segmental organs. Or, again,
the secretion may be used to form a lining to tlie burrows and tunnels in which the
creatures pass their lives. This can only be decided by carefully watching living
specimens. Sections of the body-wall of E. PallasU, specimens of which came from
St. Andrews, showed however, in the region of the segmental organs, where the
compound glands are located in E. unicinctus, only unicellular glands. Tliey were
similar to those in other parts of the body, being much modified, as Spengel noted, and
having each a long tapering neck, attached to an expanded body embedded in the cutis ;
on the proboscis they were larger than elsewhere, but in no region were there any
compound glands such as occvir in this Japanese species.
The cutis is beneath the epidermis ; it consists of connective-tissue brandling cells,
with nuclei scattered irregularly throughout. As in all the other layers of the body-
wall, no blood-capillaries can be detected (PI. 7. figs. 6, 7, 8, 9, 10, i.e.).
The muscle-sheath comes next ; the outermost layer is made up of circularly arranged
fibres; below this is a band of longitudinal muscles, followed on the inner side by
another layer of circular muscles, showing, however, a slight obliquity as compared with
the outer circvilar layer {m.c).
Sections cut in different parts of the body show all these parts in very varying
proportion. Median sections exhibit but feeble papillation, and the unicellular glands
are more scattered, while the large compound glands are absent altogether ; the cutis
and muscle are also proportionately reduced (PL 7. fig. 8). A great difference is
observable in sections taken through the thickened wall of the jwsterior end (figs. 9, 10),
the immense increase in depth being mainly due to a development of longitudinal
muscles ; the papillae are striking, but possess no compound glands.
It is difficult to discover how far the epidermis of the body-wall extends at the
posterior extremity of this worm, for there is no definite line of demarcation showing
AFFINITIES OF ECHIURUS UNICINCTUS. 81
where the epithelium lining the alimentary canal takes the place of the hndy integument.
A peculiar tissue is present within the anal spliinctor, continuing forward some distance
beyond the openings of the anal vesicles ; it is suggestive of l)eing the glandular
epidermis, modified as a result of its morphologically internal position. Its prol)al)le
signiticance and relations will be considered more fully in connection with the anal
vesicles.
In sections of the bodj'^-wall, and more particularly in the region of the proboscis,
there are present in the epidermis groups of sensory cells, similar to those figured by
Jameson in his paper on Thalassema neptuni. Each cell possesses a projecting process
or hair, which, piercing the cuticle, forms a trigger-like organ (PL 7. fig. 11, c.s.) ; I was
unable to trace any connection between these cells and nerve-branches from the lateral
trunks in the proboscis. The occurrence of such structui-es as these sensory cells would
appear to be exceptional from what Spengel says in the account he gives of his investiga-
tions on S?//«o'«s Pa//«m: "Beim Echiunis habe ich weder bewegliche Cilien noeh
Starre Haare zu Erkennen vermocht." The only organs of a sensory nature Avhich he
finds are those to which he applies the term " becherformige Sinnesorgane," comparable
with those mentioned by Eisig * as occurring in the CapitelUdeu. In Sipunculus
AndresBf found between the gland-cells occasional " Nervendorgane," particularly on
the proboscis, but also irregularly distributed over the body. Eisig concludes that what
Andreae calls " Seitenorgane " are in reality the same thing as his " Beeherorgane."
Alimentary Canal. — The alimentary canal is relatively very long, about 60 cm.; it
forms many coils, which are arranged for the most part quite irregularly ; yet I find
that some of the individual loops are constant in occurrence and disposition. In all
cases the mouth leads into a straight buccal cavity, which is fixed to the body-wall by
radial and dorsal mesenteric strands. Histologically this tract shows a ciliated glandular
epithelium which is thrown into folds, continuous with those on the prostomium. At
first the outer layer of longitudinal muscle is very inconspicuous (PI. 8. figs. 13, \Za),
but it gradually becomes more pronounced (PL 8. figs. 14, 14 a, m.l.). Beyond the
region of radial mesenteries the tube becomes narrower and takes an uninterrupted
backward course for about an inch ; in Ech'mrus Fallasll and Thalassema neptuni this
region is thrown into a double loop. As before, its lining is glandular, but the waUs
have become much more muscular ; at this jioint there is always a little knot of <1 to 5
close coils (PL 7. fig. 5, cr.), in which the muscular elements are greatly reduced and
the glandular nature of the epithelium is intensified, the foldings at the same time
becoming deeper. Prom the nature of this coiled portion I judge it to be distensible, so
it may function as a crop (PL 8. fig. 15). Beyond it, the alimentary canal decreases
still more in calibre, and runs almost straight to the posterior end of the body-cavity ;
its walls are immensely thick, owing to the remarkable development of circular muscle,
* H. Eisig : " Die Seitenorgane und becherfcirmigen Orgaue tier Capitcllidcn,"' MifctheiluDgen aus der Zoolo".
Station zu Neapel, Bd. i. p. 278.
t J. Andrea;: "Eeitrage zur Anatoiaie u. Histologic des Hiimnculus nadux, L.," in Zeitscli. wiss. Zool. Ed. sxxvi.
pp. 201-258, tabb. 12 u. lU (ISSl).
§2 ALICE L. EJJBLETOX ON THE STRUCTURE AND
while tlie longiiudinal fibres are almost wanting; the epithelium is correspondingly
leduced anl is devoid of glands (PI. 8. fig. 16). This narrow, muscular, non-glandular
part is probably to be looked upon as a gizzard, following as it does immediately upon
the crop-like region. Jameson describes the " crop " in Thalassema neptuni as coming
behind the gizzard ; though it is difiicult to see what its use could be in such a position,
for the crop is essentially a glandular distensible part in which food collects before it
can enter the narrow muscular gizzard, where it is crushed previous to its passage into
the delicate intestine. I find the cilia in somB specimens extend back as far as the
beginning of the intestine, though this is apparently an individual variation.
At the posterior end of the body this narrow gizzard, which extends for nearly 5 cm.,
widens out into the thin-wall^d intestine, and at the same time it turns forwaixls,
becoming a veritable tangle of complicated coils and loops ; mesenteric strands reappear
at this point which attach the coils, irregularly, to the body-wall. The beginning of the
intestine proper is marked by the appearance of the ciliated groove on the ventral side,
Avhich 25 cm. further on gives place to the collateral intestine (or siphon), at a point
about 12-5 cm. from the mouth. The intestine continues as far as the rectum (or large
intestine) ; its walls are very delicate, possessing very few fibrous elements ; its epi-
thelium is glandular and is raised into long slender villi (PI. 8. figs. 17, 17 a) ; above
the ciliated groove and collateral intestine runs a band of longitudinal muscle (PI. 8.
figs. 17, 21</, 21 b, m.L), which is continued on the rectum (Pi. 8. fig. 18, m.l). In all
the specimens 1 examined, the origin and end of the collateral intestine are clear and
unmistakable, though Drasche says he was unable to find the openings. At its origin
(fig. 19), the transition from the ciliated groove to the accessory intestine is very abrupt,
the latter standing up from the alimentary canal with a somewhat dilated and swollen
end. Posteriorly it passes over gradually into the ciliated groove, ending without a
dilatation, about 10 cm. from the anus (fig. 20). On opening tlie alimentary canal at
these points, and pinning back the walls so as to expose the interior, the aperture leading
from the collateral intestine into the alimentary canal is jilainly visible under a
dissecting microscope (PL 8. fig. 21, o). Beneath the collateral intestine there is a
shallow groove Avhich Jameson has called the "secondary ciliated groove" (fig. 21).
The collateral intestine has received various names : it is sometimes referred to as the
siphon, or accessory intestine; Spengel calls it the '• Nebeudarm," while Greeff mistook
it for a blood-vessel and speaks of it as the "Darmvene."
The rectum (PI 7. fig. 5; PI. 10. fig. 39, r.) is a straight, wide, thick-walled tube,
proceeding from the anus along the left side of the nerve-cord as far as the segmental
organs; it is held in place by a row of mesenteric filaments inserted exactly opposite the
lone-itudinal muscle, i. e. inserted on the dorsal side. At its anterior limit, where it
passes into the intestine, there is a marked constriction, though no sphincter muscle can
be seen on microscopic investigation. Tiiere are two endoparasites in this rectal tract —
a probably new species of the Infusorian Trichodma, and a Copepod, which is evidently
an entirely unknown form *.
* Tlds was confiriued by Dr. G. S. Brady, F.R.S., to \\liom some specimens were submitted.
AFFINITIES OF ECIIIUKUS UNICINCTUS. 83
On comparing the alimeatary canal of allied animals, as described by various authors,
Avith this species, I find the part anterior to the intestine has been divided up into
numerous distinct regions, llietseli calls it simply " intestin buccal," but Spengel says
it comprises a pharynx, oesophagus, and crop. Jameson finds the oesophagus of Echiuras
equals the oesophagus and gizzard of Thalassenia. Greefi" does not mention tiie crop, but
in other respects his description agrees with that of Spengel. In this Japanese form I
can divide the alimentary canal only into those areas mentioned already, i. e. : —
i. The huccal sac (or pharynx), on the first portion of which arc; the radial mesenteries.
ii. The coiled part (or crop).
iii. The narrow straight part (or gizzard).
iv. The intestine, bearing the ciliated groove and collateral intestine.
V. The rectum or large intestine, fixed to the left side of the nerve-cord and ending
at the anus.
Figs. 13-18, Plate 8, are diagrammatic representations of transverse sections through
these various regions, showing the comparative development of tlie muscular layers and
glandular epithelium.
Blood-Vascular Sijstem. — All the species of the genus Echiarm hitherto investigated
possess a very characteristic blood-vascular system. E. Fallasii, according to Greetf, has
two main vessels — a dorsal and a ventral — and " beide Gefasse verliinsen sich bis in
den Eiissel, das Riickenfass einfach und in der lliickenwandung des Ptiissels des
Russelarterie der mittleren Laugsrichtuug folgend das Bauchgefiiss in zwei Aesten, die in
den Seitenriiudern uuter Entsendung zalilreicher verlaufeu." In the proboscis he finds
an open communication between tlie blood system and the body-cavity ; anteriorly the
dorsal vessel swells out as a contractile heart, while the ventral vessel runs to the hinder
end of the hody, bound to the ventral surface of the alimentary canal, as the " Darmvene."
Spengel, in working on the same species, gives a very similar account of its vascular
system, though he denies the existence of a communication between the vessels and the
body-cavity, and does not give any support to the statement that a pulsatile heart is
present dorsally. The vessel which Greelf refers to as the "Darmvene" is, as Spengel
shows, the " Nehendarm," or collateral intestine. E. Pallasii, then, according to Spengel
and E-ietsch *, has a ventral vessel running the whole length of the body above the nerve-
cord, connected anteriorly by a loop with a short vessel on the dorsal surface of the
pharynx and oesophagus : these vessels are found in the proboscis. Jameson gives a
similar arrangement for Thalassenia neptiini; and Shipley f says : "A closed vascular
system exists in Echiurids, consisting of a contractile dorsal vessel running along the
dorsal surface of the anterior end of the alimentary canal, and continued along the
axis of the proboscis. At the tip of the prohoscis it bifurcates and each branch descends
along the edge until it reaches the base, where, having encircled the oesophagus, the
two unite, and are continued as the ventral vessel which runs along the dorsal surface
of the nerve-cord, and eventually ends blindly. There is also a vessel which passes from
* M. llietsch : •• Etude sur lea Gephyrieus aimos ou Echiuriens," in Eec. Zool. Suisse, vul. iii. IS8G.
t Shipley : Gqjhijrea aud Phoroitls. The Cambridge Xatural History, vol. Li. p. 450.
84 ALICE L. EMBLETON ON THE STEUCTUKE AND
tlie ventral vessel and encircles the intestine, opening into the posterior end of the
dorsal vessel. In Echiurus this same vessel encircles a stout muscle which runs from
the base of one of the ventral bristles to the other."
On examining Echiurus unicinctus, however, no trace of a closed system of blood-
vessels could be found. Many individuals were very carefully opened, but always with
the same negative result, there being no dorsal nor ventral vessel with their connecting
loop or " muscle-ring" (P). 7. tigs. 5, 12). Microscopic jiveparations served but to verify
this oliservation ; no vessel exists in relation to the nerve-cord, as is seen in sections
of the entire worm and in those of the nerve-cord alone (PI. 9. figs. 22, 23, 24, 25).
If the loop connecting the dorsal and ventral vessels were present, encircling as it does
in other species the muscle running between the basal ends of the two anterior hooks, it
w^ould ajjpear in the section of the. entire worm in that region (PI. 9. fig. 22, m.b.),
drawn with the " camera lucida," In similar sections taken through specimens of
Echmriis Pallasii and Thalassema nejituni the blood-vessels were unmistakably present,
cut across at various angles. Fearing my methods might be at fault, I applied them
to preserved material of Echiurus Pallasii and Thalassema neptuni ; both in dissections
and microscopic preparations the vessels occur as Spengel and Jameson describe, being
quite clear and distinct. In sections of tlie proboscis in these two allied forms, the
median and latei-al vessels are seen at once ; but sections in this region of Echiurus
nnicinctus are very ditFerent : there are no definite vessels present, but irregular sinuses,
which appear to be in two series — conceivably an efferent and an afferent system.
The ventral sinuses are elongated transversely (PI. 9. fig. 26, s.v.), and are rather
undefined, with thin walls w^hich are scarcely discernible. The upper sinuses, elongated
in the opposite direction, are less irregular, with more definite walls (tig. 26, s.d.).
In tracing the series of sections with great care from the anterior extremity of tlie
proboscis to its union with the body, it is seen that gradually the sinuses widen out,
forming large cavities (PL 9. fig. 24), which eventually run together in the mouth-region
and then merge into the body-cavity. Ova and coelomic corpuscles are found in all these
sinuses {o,c.), even at the tip of the j)roboscis. This is conclusive evidence, establishing
the fact that these sinuses are but forward extensions oi the coelom.
These sinuses are in all probability largely connected with the protrusion of the
proboscis, functioning much the same way as the blood-sinuses in the foot of the
Lamellibranchia, where the muscular foot is protruded by an injection of blood into its
substance.
Drasche, in speaking of this species, says : " Trotz sorgfiiltiger Untersuchung beider
Exemplare gelang es mir leider nicht, die Gefiissschiiuge aufzufiuden."
The fact that Drasche failed to detect the vascular loop in his tw'o specimens sujiports
my observations, which have the advantage of having been made on many individuals.
Nervous System. — The nervous system is arranged in a manner essentially characteristic
of all Ecliiurids. There is a single ventral nerve-cord, extending from mouth to anus
(PL 7. figs. 5, 12, n.) ; it is fixed to the ventral surface of the body-w^all by its lateral
branches, which for a short distance are quite free, then they peneti'ate into the tissue of
the body-wall, running parallel Avith the circular muscles (Pi. 9. fig. 27). These lateral
AFFINITIES OF ECIIIURUS UNICIXCTUS. 85
nerves do not originate accui'ately opposite one anotliev ; they spring from the ventral
region of the main cord, and are composed only of nerve-fibres. Spengel found in
Echinrus Pal/asii that tliese lateral nerves form definite rings or commissures in the body-
wall, as nerves from opposite sides meet in the middle dorsal line — their course being,
according to him, directly under the roAvs of papillae. In Echinrus uaicinctus the nerve-
cord is held in place by its side branches, there being no connective-tissue attachment, such
as is found in other forms, Rietsch, in describing the nervous system of JBonelUa minor,
remarks: " Le systeme nerveux se compose du cordon ventral depourvu de renflements
ganglionaires, et fixe aux teguments par un mesentere." Though it does not agree with
this form of attachment, yet it is not seated directly on the body-wall as in Thalassema
nepkmi, of which Eietsch observes : " Ce tronc nerveux n'est pas, comme chez la
Bonellie, I'elie aux teguments par un mesentere ; il repose directement sur la muscula-
ture." The cord is, in Echiuriis unicinctus, free from th(» body-wall, except in so far as
its fine lateral branches fix it in position, by becoming embedded in the musculature.
0\vin<r to the contraction of the bodv-wall. the central nerve-cord in these preserved
specimens is thrown into wavy folds (PL 7. figs. 5, 12 ; PI. 9. fig. 27, u.) ; this condition
makes it easy to see that the cord is free of the body-wall but for the attachment of
the nerve-branches.
Anteriorly the cord bifurcates (PI. 7. fig. 12 ; PL 9. figs. 23, 24, 27) ; the two limbs,
passing round the buccal cavity, eater the substance of the proboscis, at the tip
of which they meet, thus forming a true circum-oesophageal ring. In I'orms such as
E. Fallasii, or BoneUia, where the proboscis is a very extensive organ, this nerve-loop
is necessarily much elongated. But in the species under consideration, the proboscis
being scarcely more than a prostomial lobe, the nerve-ring is almost as compact as in the
Earthworm (PL 9. fig. 27).
Similarly, at the posterior end the cord splits into equal halves ; and these enter into
close relation with the wall of the rectum. It is not easy to follow these two branches,
but they apparently divide up into finer ramifications which supply the muscles of the
anal circlet of sctoe.
There are no ganglia in the A"entral cord.
The outline of the cord in transverse section (PL 9. fig. 28) is seen to be rather flattened
dorso-ventrally, and is not circular as in Echiurns Pallasii {vide Greeft', Spengel). The
cord has a coating of nerve-cells {c), most marked dorso-laterally — ventrally the cellular
elements are fcAver and more scattered. These nerve-cells vary much in size; they
enclose a central mass, made up of ramifying nerve-fibres (/.//.), the majority of which
run longitudinally as is seen in longitudinal sections (PL 9. lig. 30, f.n.). Many of
the fibres constituting this central fibrous mass are not nervous, but are inward
extensions of the protective connective-tissue sheath which covers the cord. The nuclei
which lie scattered throughout this fibrous area may belong to these connective-tissue
elements, or to the nerve-fibres, but from the preserved material it is impossible to iorm
a conclusive opinion on this point. In this outer sheath, the longitudinal muscles are
much less conspicuous than in similar sections of the nerve-cord of Echiurm Pallasii,
or of Lumhricus (PL 9. figs, 31, 32, sh.).
SECOND SERIES.— ZOOLOGY, VOL. VIII. 13
86 ALICE L. EMBLETON ON THE STRUCTURE AND
lluuniug through the dorsal region of the cord, emhedded m the cellular coat, is a
structure which Greeff refers to as the " central canal," in spite of its excentric
position (PI. 9. figs. 28, 32, c.ii.). Spengel calls it the '' Neuralkaniile," and says : " Ein
Gebilde problematischer Natur namlich eiuen Langskanal, der dicht unter der dorsalen
Mittellinie verlauft," but how far it runs he is not able to say. Greeff, however, observes
of it : " Dieser Kanal aber communicirt mit der Leibeshohle, ist ein Theil derselben, der
zuni'ichst aus dem, vvie wir oben gesehen hal)en, mit dem Blutgefass-system in directer
Verbiudung stelienden Kanalsysteme des Riissels hervorzugehen scheint."
In all my preparations, however, this " canal " has a similar structure and configura-
.tion to the so-called " giant fibres " of the Earthworm. I have in PI. 9. figs. 31, 32, made
comparative drawings of the dorsal region of the nerve-cord of Lmnbricus and of Echiuriis
imicinctus, both preserved and stained in the same way (Ehrlich's luematoxylin and eosin
or orange gave the most satisfactory results). Though Liimbriciis has three such
structures and Echiurus but one, the identity is very striliing. The same reagents, used
on the nerve-cord of Echim-us Pallasii, gave results as in the two former cases. In the
Earthworm, however, it was possible to trace the connection between the ganglion-cells
and the " giant fibres,"* being much more difficult to find with the central " fibre" than
with the lateral ones. Partly I could trace the connection in Echlurus uiiicinctus, though
not in such a way as to render the fact beyond dispute.
Anal J'esicles. — Situated at the posterior end of the body-cavity is a pair of very
. characteristic organs, generally known as the " anal vesicles," though they are occasionally
referred to as the "posterior nephridia."
In general form and histological structure, these vesicles show almost complete
agreement with those of Echiurus Pallasii as described by Spengel. The observations
were carried out with difficulty owing to the highly-contracted condition of the vesicles.
J)rasche evidently met with tlie same obstacle, for the only mention he makes of these
-organs is that " in beiden Exemplaren von brauner Farbe, liessen wegen ihres
macerirten Zustandes nur schlecht die Wimpertrichter erkeunen." However, in my
specimens the ciliated funnels are quite recognizable, even in a piece of a contracted
vesicle, Avhich was very slightly stained with alum-carmine and then clarified in oil of
cloves. Under the microscope this preparation had the appeai-ance of a mass of
diverticula, composed of very delicate tissue; the ciliated funnels (PI. 10. fig. 33,/.),
which took the stain rather more deeply than the surrounding tissue, were plainly observ-
able among the diverticula (rf.). Under a higher magnification the component cells could
be made out : those forming the diverticula jwssess very little protoplasmic contents, the
nucleus lying apparently in a vacuole (fig. 34, d. & nu.) ; but the cells forming the funnels
contain much granular protoplasm [pit-] very slightly vacuolated, and nuclei which
stain darkly {c.c. & /.).
Among the numerous specimens I examined, occasionally there occurred vesicles not
wholly contracted ; and such a piece of uncontracted material, treated with the same
reagents as before, shows the vesicle-wall smooth and unfolded, as in fig. 35. The wall
* Cy. Friedliinder : " Bcitriige zur Kenntnis des Cciitraluervensvstems von Lumhricas,'' Zeitsclir. f. wiss. Zool.
M. xlvii. 188«.
AFFINITIES OF ECHIUEUS UNICINCTIJS. 87
is very delicate, consisting of an epithelium of flattened cells {rp.), through which run
interlacing muscle-fibres {f.m.), forming an irregular network. The nuclei of these cells
are large and conspicuous, while the cell boundaries are quite distinct, though Spengel
says in Eckiurus Falhisii they are usually indiscernible. He also denies the existence of
a special system of blood-vessels which, according to Greeff, rim in the vesicle-wall —
in my preparations I see nothing to suggest that such a system of canals is present.
Greeff also says : " die Innenflache der Analkiemen ist mit wimpernden Cilien besetzt " ;
this is certaiuly not true of Echiunts imicinctus.
Standing out from the vesicle-wall are many ciliated funnels (PL 10. fig. 35, /.),
on the structure of which I have nothing new to add.
The vesicles are held in position by fine muscular fibres which attach them at various
points to the body-wall, but apart from this they float freely in the body-cavity beyond
their pomt of union with the rectum. I'l. 10. fig. 36 shows a transverse section of the
body at a point where the vesicles open — ventro-laterally — into the rectum, each through
a strikingly small aperture {a^, a'^) ; but according to some authors it is doubtful if this
should be looked upon as being a true part of the alimentary canal, or if it is not rather
an invaginated portion of the epidermis. There is, beneath the anal epithelium in this
region, a belt of peculiar tissue wiiich, in this species, extends forwards beyond the
openings of the anal vesicles; it has been thought to be modified glandular tissue
belonging to the epidermis, in which case the vesicles in Ee/ii/inis Kuic/i/cfiis would open
on the external surface which has become? secondarily internal. This special tissue has
been described by Spengel as " glandular " ; in all my preparations, however, it appears
to be a highly vacuolated connective tissue, differing from the ordinary connective tissue
only in possessing fewer fibrous elements and fewer nuclei (PL 10. fig. 37, t.p. ; La.) ; the
striking feature connected with this tissue is the abruptness with which it passes over
into the typical connective tissue {(m.), as is seen in fig. 37, where both types occur side
by side in one fold of the rectum-wall. The epithelium covering these different tissues
is the same, possessing, in both cases, an abundant supply of unicellular glands, whose
nuclei are situated at the basal ends of the cells ; many of these gland-cells sink down
into the tissue beneath, where they are seen cut across in all positions. Apart from
these glandular elements belonging to the epithelium, this special tissue gives no evidence
in support of the view that it is " glandular." What is its real significance is a
physiological problem which must he worked out on fresh or living material before it can
be settled satisfactorily. It is, however, probable that this animal performs some kind of
anal respiration, and that this area of vacuolated tissue is connected with the intake
of fresh sea-water, l)ecoming alternately turgid and flaccid. The significantly large
supply of radiating muscles on the rectum near the anus also suggests a suction action,
such as takes place in the rectum of a Holothurian, where there occurs a similar series
of muscles. In the straight posterior or rectal portion of the alimentary canal of
Echiurus unicinctiis (PL 7. fig. 5, r.) one finds two parasites — Trichodina and a Copepod,
and of these I hope later on to write a separate account. In all probability they
enter " per anum," for had they, or their eggs or spores, been taken in orally, they
would undoubtedly have been digested before reaching that part of the alimentary canal
13*
88 ALICE L. EMBLETON ON THE STRUCTUKE AND
in wliicli they live ; it therefore seems likely that they are admitted to the rectum in a
stream of water, having a respiratory significance, and taken in through the anal
sphincter. The most satisfactory preparations of this tissue were those treated in
bulk with alnm-carmine, having first cleared the tissue of corrosive-sublimate crystals
by immersing it for an hour in iodine solution. I found a sort of precipitate was
thrown down in preparations treated in bulk with Ehrlich's ha^matoxylin, followed
by a stain for a few minutes on the slide of a slightly acid solution of Griibler's
Orange G in 70 per cent, alcohol; but otherwise these formed very excellent contrast
stains.
Spengel says of Echiiirns Pallasii that the vesicles open exactly on the line of demarc-
ation between what he calls the "driisenlosen" and the " driisenreichen " areas; so that
regarding the " glandular " tissue as ejiidermal, the vesicles in this case open neither
internally nor externally, but on the boundary. According also to Eietsch : " L'orifice des
glandes anales serait exactement a la liiuite enti'e les teguments et I'intestin." The study
of the development of the Gephyrea, according to Korschelt and Heider*, proves that these
organs "do not arise, as was supposed, from the intestine, but are formed in the somatic
layer of the mesoderm" ; and further that " their entire mode of origin proclaims the anal
vesicles to be nephridia which only secondarily entered into connection with the hind
gut." If this is so, their function must be excretory ; and the observations of many
zoologists lend support to this theory. For instance, Shipley says : " The function of
these structures may be excretory ; or they may control the amount of liquid."
Danielssen and Korenf, speaking of the anal vesicles of Uaminf/ia arctica, assign to them
the function of nephridia, denying the possibility of their having a resj)iratory
significance, for they do not think the sea-water penetrates into the vesicles.
Rietsch observes : " Quant aux fonctions de ces organs, je crois, avec H. de Lacaze,
qu'elles sont avant tout glandiilaires, excretoires. Les courants determines j)ar les cils
desentonnoirs, de la glande elle-meme et de la portion terminale de I'intestin, ue peuvent
donner naissance qu'a un courant vers rexterieiu\ Les glandes, en se distendant, ne
determinent elles pas, malgre co courant, la penetration, a I'interieur de la glande, de
I'eau de mer qui servirait ainsi ;i la respiration du liquide de la cavite generale ? 'A
priori,' la chose n'est peut-etre jjas impossible, mais I'observation directe pourrait seule
prouver qu'elle a rcellement lien. II ne senible pas, en tout cas, que cette eau puisse
par les entonnoirs aller se meter au liqviide perivisceral, surtout chez YEcliiure et la
Thalusseme. En somme les fonctions respiratories des glandes anales demeurent
douteuses."
Huxley, Gegenbaur, Glaus, and Hatschek consider the anal vesicles as homologues of
the segmental organs of worms.
Other investigators, however, have looked upon them as being undeniably respiratory
in function. Greeff, in his account of these organs in the Ecliiurids, begins by saying :
" Als llespiration-organe mtissen in erster Linie die beiden Analsehiauche betrachtet
* Text-book of Embryolos;}- — Invertebrates, vol. i. p. 309.
t " Fra deu norcke Nordhavs Expedition (Gephyreer)," Nyt Mag. f. Nat. Vid. vol. xxvi. pp. 44-G6.
AFFINITIES OF ECHIUKUS UNICINCTUS. 89
werdeu, Kiemen im vollen Siuno der Wortes, volliiv den aualoi? soa^enannten ' Wasser-
lungen ' der Rolotlminen.'" Sclimarda's * opinion coincides with this, and Forbes and
Goodsir'sf went the length of calling the anal vesicles " Atheinsacke."
There may be rectal respiration, but it is extremely doubtful if the anal vesicles act
as lungs, for though there be a current of water in and out of the anus, it cannot
possibly flow freely through the vesicles, their structure rendering this impossible. In
the first place, the apertures leading into the vesicles from the rectum are too small to
allow of any appreciable amount of water entering them ; but allowing that the water
does penetrate to the vesicles, it cannot get further, for the ends of the ciliated canals
(described minutely by Spengel in IJc/ii/inm 'Fallasii) Avould act as valves, completely
preventing the jmssage of any liquid into the body-cavity; lastly, the cilia in the canals
are directed inwards, as Spengel and Greeff both remark, which is conclusive evidence
tbat fluid may be carried from the body out at the anus but not in the opposite
direction.
It, therefore, appears probable that the function of these organs is excretory ; the
granule-containing cells in the walls of the vesicles may function as carriers which
collect w'aste matter, and throw it and themselves away. The vesicles are full of bodv-
cavity fluid, which may be the product of the ordinary cells forming the peritoneal
lining of the ccelom ; for in no part of the animal is there any glandular tissue like the
chloragogenous tissue in Lumbricus Avhich secretes the coelomic fluid in that worm. It
has been suggested that the anal vesicles secrete the coelomic liquid, though, so far as I
can judge from my observations in Echiuriis umciuctun, tliere is no evidence to support
such a view.
Segmental Organs. — Posterior to the two anterior ventral setae are two pairs of
nephridia, or segmental organs (PL 7. flgs. 5, 12, o.s.), opening on the one hand into the
body-cavity, and on the other to the exterior. The lips of the inner opening are
produced out right and left into long tapering arms (PI. 7. fig. 12, l.s.), which are spirally
grooved and densely ciliated, but w^hich contain no canal. Drasche says there are 20-30
coils in each s])iral arm ; but in taking the average among many of my specimens, I
find it is less than this, there being not more than 12-20 coils. At the base of these
arms, and exactly between them, is the aperture leading into the vesicle from the body-
cavity (PI. 9. fig. 25 ; PI. 10. fig. 38, ap:-) ; the contents of the vesicle are discharged
externally through a minute pore in the body-wall {(ip.^). PI. 9. fig. 25 represents a
section through the entire body of the worm, and in it are seen the two openings of the
vesicle {up}, ap.-), the spiral arms {l.s.), and the vesicles containing ova.
In some individuals the vesicles, being highly muscular, an^ enormously distended by
the mass of reproductive products contained in them ; they stretch back some way
among tlie coils of the alimentary canal, when they are in this inflated condition —
this is also the case in Bonellia (Greeff).
Each vesicle possesses but one chamber, whereas in Bonellia it is divided into two by
* Zur Naturgcsch. tier Adria : 1. Bonellia viridis, pis. 4-7, in Memoirs of the Acad, of Vienna, 1852, vol. ii.
+ "On the Naturiil History and Anatomy of Thahisseiini and Echiurus" Edin. Now Phil. Journ. 1841, vol. xxx.
j)l. vii.
90 ALICE L. EMBLETON ON THE STRUCTURE AND
a partition. The inner surface of the vesicle is not ciliated, for the cilia on the spiral
arms do not continue in beyond the canal leading into the vesicle — in this canal the cilia
are directed inwards towards the interior of the vesicle. As Drasche has already pointed
out, there are two bands of longitudinal muscles attaching the segmental organs to the
body-wall.
These nephindia have been looked upon by some authors as reproductive organs, and
Pallas* named them " vesiculte genitales " : this error must have arisen from the fact that
they function as gonodncts, though they serve only as temporary receptacles for the
reproductive products. They are typical nephridia ; and Rietsch says : " Les poches
genitales des Echiurens sont evidemmeut homologues des organes segmentaires des
Annelides." They have miich systematic significance ; and differences in the excretory
oro-ans form important distinguishing features among the various species and genera of
the Gephyrea. Drasche gives as one of the characteristic marks of Echiunis niiicinctus
" die mit Spiralrinnen versehenen Trichter der Segmentalorgane."
Beprodiictke Organs. — In mature individuals the ova or spermatozoa are found
floating freely in the coelomic fluid : there are no special gonoducts, but the segmental
oro^ans perform the function of transmitting the ripe sexual products from the coelom to
the exterior. The sexes are separate in this species, though there is ajiparently no sexual
dimoi-phism, such as occurs, for example, so strikingly in Bonellia virkUs.
The ova and spermatozoa are the product of the peritoneal cells lining the body-cavity,
in one localized area — i. e., along the ventral surface, around the ventral nerve-cord,
chiefly at the posterior end of the body-cavity ; they are formed by a proliferation of
the peritoneal cells in this region. Wiien the sexual cells are ripe they become budded
off into the coelom, where they float about in the perivisceral fluid until they escape to
the exterior through the segmental organs. In microscoiiic sections of the proboscis,
ova are present in the sinuses almost at the tip of that organ, showing beyond question
that these sinuses are merely forward extensions of the ca-lom, and are not blood-vessels.
Greeff pictures the genital cells as budding off from a stem or rhachis ; Spengel .says
this axis is formed from the posterior extremity of the ventral blood-vessel : " Die Stiele
scheinen rundum vom einem gemeinschaftlichen Strang, einer gemeinschaftlichen
Rliachis, hervorzusprossen." Greeff and Spengel call this an " ovary."
The ova are, comparatively speaking, large ; they are dark and granular, with tlie egg
in the centre as a clear spot or vacuole ; there is an egg-membrane or shell.
The spermatozoa — of typical form with a globular head, and a flagellum-like tail —
float in the coelom in little balls or lieaps.
Mesenteries. — Several members of the class Echiuroidea possess definite mesenteries ;
and many zoologists have attached great importance to these structures, claiming
for them a systematic significance, inasmuch as they are supposed to be indicative of
primitive seomentation, of which almost all trace is wanting in the adult. This is the
view put forward by Spengel in his observations on EcJdunis Pallasii. Referring to the
septum-like membrane in the region of the pharynx, he says : — " Dieses Diaphragma ist
eine diinne, durchsichtige, von feinen Muskelfiiden durchzogene Membran, die sich vor
* Miscellanea Zoologiea. Hagw Comitum, 1766.
AFFINITIES OF ECHIUKUS UNICINCTUS.
91
den Baucliborsten senkreelit durch die Leibeshohle ausspauut und uur (lui-ehbrochen ist
von einem etwa ihr Centrum einnehmenden Loclie, das sich in einen bis avif die ventrale
Leibesvvand hinal)reicbendeu senkrechten Scblitz fortsetzt. Der linke Rand des Schlitzes
liefei-t die ventrale Bei;-i-enzung- des Locbes, indem er sicb nach rechts verbini-ert um
sicb an das Kreuznui;' der beiden Riinder stattbndet. Von vorn her aber steheu mit den
Eandern die beiden Blatter des Pbarynx-Mesenteriums in solcber Verbiudung, dass die
seitlichen Hiilften des Diai)bragmas als Fortsetzungen dieses Mesenteriums erscheinen."
" Icb habe das Verhalten dieses Mesenterinms so eingehend besebrieben, weil niicb die
Bildung des Diapbragmas aufs lebhafteste an die Dissepimente der Anneliden erinnert
bat und mir jede Andeutung von einer Segmentirnng l)ei den Ecliiuriden der
Beacbtung in hochsten Masze wertb scbeint. Icb werde es naturlicli unternehmen,
scbon jetzt die Bedeutung des Bef undes zu diskutiren ; icb wiinsche nuv die Aus-
fuhrlichkeit der Schilderung durcb diesen Hinweis zu recbtfertigen. Ueber eine f unktion-
elle Bedeutung dieses Diapbragmas linden sicb in dem iiber die Borsten bandelnden
Abscbnitte einige Bemerkungen."
Rietsch, describing the pharyngeal region of BoneUia minor, says : " Je n'ai pas
remarque ici de diaphragme analogue :i celui deerit par Spengel cbez VEchiure:'
According to Sluiter the general arrangement of the alimentary canal in Thalassema
erythrogmtninon is analogous with that of Echlnrus, ])ut there is no diaphrao-m.
Echinrus uiuciiuiu''.
Dissection of the anterior end of the body, showing the radial folds of peritoneum on the buccal sac. The dorsal
mesenteries have been cut through, ij., ventral nerve-cord; ph., pharynx; s.a., anterior seta; s.ni., mesenteric
strands ; »■., body-wall.
Greeff apparently has not noticed the "diaphragm" in Echiurus Pallasii, for the
only reference he makes to mesenteries is as follows : " Ein Mesenterium von
Bindegewebsfjiden und Muskelfasern hillt die Schlingeu unter einander zusammen und
befestigt sie audi mehr oder minder an die innere Leibeswand."
I .'
92 ALICE L. EMBLETON ON THE STRUCTURE AND
Danielssen and Korcn, in Epithetosoma norvegicum, speak of a ventral mesentery ;
thongh Sedgwick *, referring to tlie Echiuroidea as a wliole, says " there is no special
mesentery, bat strands of tissue run from all parts of the body-wall across the body-cavity,
to be inserted into the walls of the alimentary canal."
Jameson, however, in his recent paper on Thalassema ueptimi, recognizes a dorsal
mesentery running along the whole course of the alimentary canal except on the gizzard,
and a ventral mesentery, " bearing the ventral vessel througliout its entire length, is
attached to the alimentary canal anteriorly and posteriorly." ... "At the hinder end of
the oesopliagus it suddenly leaves the digestive tube with a falciform free margin and
extends as a short free fold along the ventral surface, attached to the peritoneal sheath
of the nerve-cord and bearing the ventral vessel. Posteriorly it approaches the gut
again, and attaches itself to the csecum, on the walls of which the solid continuation of
the ventral vessel ends." ..." Owing to the presence of the web of ventral mesentery
which occupies the oesophageal loop, the gut, with the two mesenteries, shuts off a
2)ortion of the body-cavity forwards, forming a kind of false diaphragm. This anterior
division of the body-cavity opens out by a single small hole through which the ventral,
and at times the dorsal, blood-vessel passes forwards ; this aperture is bounded by a
small piece of the left ventral body-wall wliich lies between the insertion of the
dorsal mesentery and the nerve-cord." ..." Tiie radially arranged folds of peritoneum
around the mouth and anus are quite irregular and consequently call for no attention."
I dissected Thalassevia nephmi and Echinrus Pallasii in order to see the mesenteries,
and conclude that wjiat Jameson calls "a kind of false diaphragm" corresponds to what
Spengel calls the "diaphrgam" in Ecitmrus Pallasil. But in Echiurus unicinctus there
is no structure comparable with either of these membranes ; the only mesenteries are
irregular strands attaching the alimentary canal and anal vesicles to the body-wall, and
the radiating mesenteries on the rectum and buccal sac. PI. 10. fig. 39, is a drawing of
the worm as it appears when first opened with the alimentary canal not yet uncoiled ;
obviously the arrangement of the mesenteric strands is quite irregular and indefinite {s.ni.).
The arrangement of the peritoneal folds on the buccal sac is shown in fig. 1, p. 91, Avhere
many of the dorsal attachments are shown cut through. Figs. 4.0-43 are drawings of a
series of sections cut through the anterior end of the body, in wdiich the radial folds
appear in transverse section ; they are most numerous dorsally, fixing the pharyngeal
portion of the alimentary canal fivmly to the body-wall.
As regards mesenteries, therefore, it appears that Echiurus unicinctus differs widely
from Echiurus Pollasii and Thalassema neptuni. There is no indication of segmentation
that I can see, in the structure of the adult of this Japanese Echiurus ; as to its probable
relationship Avith the segmented worms, that is a question which can only be settled after
a careful study has been made of the development of this and allied species, and by
following the fate of the larval organs. It is known in the trochophore larva of Echiurus
that the pair of mesoblastic bands become segmented up into 15 pairs of mesoblastic
somites, though, so far as is known at present, no segmentation occurs in the larva of
Eonellia.
'■ * student's Text Book of Zoology, vol. i. 1898, p. 529.
AFFINITIES OF ECHIURUS UNICINOTUS. 93
Sedgwick regards the Echiuroidca " as Annelids in wliidi segnioatation is teel)le,
showing faintly in the young, but, except in the repetition of the nephridia, .... l)eing
absent in the adult."
Proboscis. — The proboscis is essentially characteristic of the Echiui'oidea, usually being
a long highly contractile organ, A^hich, wlien fully extended, assumes an almost ribl)ou-
like form. This prostomium is formed from the oral and pi-;e-oral parts of the larva.
According to Grceff, the proboscis functions as a lung, the blood in its vessels beino-
separated from the sea-water only l)y a very thin layer of tissue : it thus becomes
oxygenated and, as he thinks that a communication exists between the blood-systeni and
the ccElom at the tip of the proboscis, the body-cavity fiuid in this Avay l^ecomes aerat(Hl at
the same time. S2)engel, as I have elsewhere remarked, denies the existence of tiiis opcTi
communication, and Rietsch says he has been unable to tind it. ,\.s regards the |)ro-
boscis sewing as a respiratory organ in Bonellia r irk/is, L., ilolando * says : " es ist kein
Anzeichen da, dass er ihm zum Athem oder als Kieme diene." Schmarda ao-rees with
Greeff in thinking that the respiratory fnnctiou is carried on by the proboscis and the
anal vesicles. It seems doubtful, to my mind, whetlier tlH> prol)()seis functions as a luni:,
if only on account of the well-known fact that it is completely thrown off on the slio-htest
provocation: if it subserved such an important function as respiration, it seems scarcelv
probable that the animal could so readily part witli it; yet in the specimens of EchlirniN
Pallasii which I have examined (obtained from St. Andrews) the proboscides were
missing, leaving in eacli case a scar round the mouth.
Greeff also assigns to the proboscis the function of prehension of food material,
by definitely seizing the prey and rolling it into the mouth ; but Spengel thinks this is
done by the cilia on the ventral surface, apart from any action of the organ as a whole.
The proboscis of Echiurm uiiiciiictus differs widely from that of allied forms : it is verv
small, thick, and blunt, being scarcely more than a prye-oral lobe (tig. '1), and in this
species it is difficult to see how it can act as a lung or for the prehension of food.
Doubtless it is much contracted in these preserved specimens, but, even allowinii- for
that, it is an extremely insigniticant organ compared with that of Echiurns Fallasii,
Thalassema, and Bouellin. If its insignitlcance were due to great contraction, then
the lateral nerve-cords would be thrown into deep folds, as is tiie ventral cord ;
but on dissecting out the nerve-ring in the proboscis-tissue (Fl. 9. Mg. 27) it is seen to be
very slightly wavy, which points to the fact that the proboscis cannot be extended far
beyond its condition in my specimens (PI. 7. tigs. 1, 2). There is no proboscis in the
aberrant genus Saccosoma, described by Danielssen and Koren.
The ventral surface is entirely clothed with cilia, which are not restricted to a groove.
The sinuses, which are so conspicuous in the tissue of the proboscis, as seen in
transverse sections, I have already referred to when considering the " blood-vascular
system." There maj^ however, be some correlation between the develo^imeut of the
blood- vascular system and the proboscis. It is known that, in those larvie of tiie
Gephyrea whose development has been followed, the vascular system is formed late, and
* Isis von Oben, i. 1S23, Taf. v. figs. 1 5.
SECOND SERIES. — ZOOLOGY, VOL. VIII. It
94 ALICE L. EMBLETON n\ THE STRUCTUEE AND
its natiiro seems to be entirely different from tliat in other animals. In the adult it
functions less as a circulatory system than as a hydraulic apparatus for expanding;; the
proboscis in the Echiuroidea, and the tentacles in the Sipunculoidea, where it is entirely
absent from the following forms, which have no tentacles ; Fetalostoma, Onchnesomn,
and Tylosoma. Similarly liere, in Echiurus unicmcfus, tlie absence of the vascular system
may he in direct correlation with the extrenie reduction of tlie proboscis. This view
gains some support from the fact that in tlie Priapuloidea, an allied group, where there
are no head appendages (such as proboscis or tentacles), there is no vascular system.
Mr. Sliipley * has called my attention to the fact that Thalassetna vegrande, Lampert,
has no proboscis, and but for the hooks there is no external indication as to which is
the anterior end.
Distribution and Jffinities. — Up to the present this species has only been met with on
the Japanese coast ; this is evidently the worm Willemoes-Suhm found there in great
abundance, and Drasche's two examples were collected by Dr. A. von Roretz on the east
coast of S. Japan ; my specimens came from Tokyo.
The affinities of tlie Echiuroidea, as a whole, are doubtful, and different zoologists classify
them very ditferently. Korschelt and TIeider, after studying the development of the
group, say that, "as regards the position of the Echiuridge, we agree with Hatschek's f
view ; he sees in them a division of the AnneUda, and In'ings them in relation with the
Ghsetopoda. The form and internal organization of the larva, as well as the mode of
origin of the setae, seem fully to substantiate this view. Even though the segmentation
(metamerism) no longer exists in the adult animal, it was nevertheless established iu the
larva, just as in the Chsetopoda and Archiannelida. The loss of the segmentation and
the reduction of the setae, as well as the enormous extension of the prostomium, or so-
called proboscis, make the Echiuridae appear as somewhat moditied forms." This is
confirmed by Conn |, who, with Hatscbek, inclines to separate the Echiuroidea from
the Sipuncixloidea. Shipley, however, is of opinion that the resemblances between these
tAvo great classes outweigh the differences, and that the Echiuroidea are derived from
the Chaetopoda, the nearest ally lieing iStern(is2:i/s ; that the Sipunculoidea are also allied,
but have departed much further from the Annelid stock. My observations on this
Japanese Echiurid lead me to believe that the connection with the Sipunculoidea is slight,
and that the features which separate the two classes are I'undamental.
As regards tlie affinities of this species with the other members of the genus, there are
several very imjiortant points of dilTerence, w^hich may even justify its being placed in a
new genus by itself: — (1) Unlike all other known genera of the Echiuroidea — with the
exception of the aberrant Saccosoma — it has no extensible proboscis, that organ being
represented merely by a short blunt prostomial lobe (PI. 7. fig. 2) ; (2) according to Shipley,
a closed vascular system is characteristic of all Echiurids, yet it is totally absent here —
this may be indicative of a very primitive condition, but, apart from that, it is, I think, a
* " Report on the Gepliyrean Worms collected by Mr. ,T. Stanley Gardiner at Rotnma and Funafuti," Proc. Zool.
Soc. 1898, p. -)72.
t " Ueber Entwicklungsgeschichte von Kcldnnts, etc.," Arbeiten zool. Inst. Wien, lid. iii. 1881.
t " Life-history of Thalassema," Stud. Biol. Lab., Johns Hopkins University, Baltimore, vol. iii. 1886.
AFFINITIES OF ECHIURUS UNICINCTUS. 95
diameter of sjeneric importance ; (3) the presence of a very definitely marked belt of
compound 2,'lands in the skin is a feature peculiar to this species, (4) as is also tlie single
circlet of peri-anal hooks, and (5) the spiral arms of the inner funnels of the segmental
organs ; (6) there is also no special mesentery nor diaphragm such as occur in Echiurus
Palhisii and Tlialassema neptuni ; and the alimentary canal seems to be simpler, for I
cannot distinguish the various regions named by Spengel, Jameson, and others.
'lliese peculiar features seem to me to have more than specific significance, and may
make it advisable to place this Japanese worm in a separate new geuus. On making
dissections of Ecliiiiriis laiicinctuH and Fallasli, and Thalassema neptuni, the identity
between the two latter was very striking, the only fundamental difference being that
Thalnsscnta lias no peri-anal bi'istles, its blood-system, prolioscis, mesenteries, nephridia,
body-wall, and alimentary canal all showing close agreement with Echiunis Pallasii ;
whereas, in these numerous and important respects, the Japanese " species ' is utterly
unlike the British Echiurid. Drasche, however, says " dieser neue Echiurus verbindet
also Merkmale von Thalassema mit deueu von Echvurus und erweitert somit nicht allein
die bisherigen Anschauungen iiber der Bau der Segmentalorgane l)ei Echitirm, souderu
moditieirt auch durch seinen nur einfaclien Hakenkranz die Umgrenzung der Gattung
selbst, wie sie bislier augenommen werden." His examination of tiie species, however,
was limited to two specimens, and was necessarily not exhaustive ; but after the facts
which have now come to light it would be necessary to extend the limits of the genus
very considerably, if the species unicinctus is to be retained within it.
EXPLANATION OF THE PLATES.
Plate 7.
Fig. 1. Echiurus unicinctus (natural size).
2. Ventral view of the anterior extremity (enlarged) .
3. Anterior seta, side view.
4. Piece of the external surface (magnified), showing the papilla! elongated transversely.
I A = anterior, j P = posterior.
5. Lateral dissection ; alimentary canal displaced.
6. Longitudinal section through the body-w;ill in the region of the segmental organ.s.
7. Section through a compound gland (highly magnified).
8. Transverse section througli the body-wall in the middle of the worm.
9. „ „ „ „ at the posterior end of the worm.
10. Longitndinal muscle-fibres of fig. 9, more highly magnified.
11. Section through a papilla on the proboscis, showiug a group of sensory cells.
12. Dissection of the anterior end, showing the absence of blood-vessels aud of the "' diaphragm or
septum " which occurs in E. Pallasii-
96 ALICE L. EMBLETON ON THE STKUCTURE AND
Plate 8.
Fig. 13. Diagrammatic section through the mouth.
14. „ „ „ pharynx.
15. „ „ » crop.
16. „ „ ,. gizzard.
17. „ „ „ intestine.
18. „ „ „ rectum.
13 a. [Drawn with the camera lucida.] Detailed drawing of part of fig. 13.
14 a. ,. „ .. „ „ .. .- H- 1-^-
17 «. .. „ „ „ „ „ ,, ^g- I''-
18 «. ., „ „ „ „ „ „ fig. 18.
19. Alimentary canal where the collateral intestine arises.
20. ., .„ ., .. „ ends.
21. „ „ opened, showing the aperture leading into the origin of the collateral intestine.
21 a. Transverse section through the intestine and coUatet-al intestine.
21 b. ,, „ „ ciliated groove.
Plate 9.
Fig. 22. [Drawn with the camera lucida.] Transverse section of the body on a level with the basal
muscle of the setae.
23. [With camera lucida.] Transverse section of the body at the nerve bifurcation.
24. ,. ,. , ,, ,. ,. ,, showing the sinuses of the proboscis
opening out as the ccelomic cavity.
25. [Drawn with the camera lucida.] Transverse section of the body where the segmental organs
open interiorly and exteriorly.
2r). [Drawn with the camera lucida.] Transverse section through the prostomium.
27. Dissection of the buccal loop (" oesophageal loop ") of the nervous system.
28. Transverse section of the ventral nerve-cord.
29. Part of same, more highly magnified.
30. Longitudinal section of the nerve-cord.
31. Dorsal region of a trans, sect, through the ventral nerve-cord of Luinbricus.
32. „ „ „ „ „ •, „ E. unicinctus.
Plate 10.
Fig. 33. Piece of contracted anal vesicle.
34. Section through same.
35. Surface view of a contracted vesicle (carm. alum ; clarified in oil of cloves).
36. [Camera lucida drawing.] Trans, sect, of posterior end of the body, whei'c the anal vesicles
open into the rectum.
37. [Camera lucida drawing.] Section of one fold of wall of rectum, anterior to the opening of the
anal vesicles, showing the two kinds of tissue (stained in bulk with carm. alum).
38. Longitudinal section through a segmental organ with its openings.
39. E. unicinctus when first opened, with its viscera '• in situ," showing the distribution of the
mesenteric strands.
40-43. Series of transverse sections through the body, showii'.g the radial arrangement of the
mesenteric strands. The coelom gradually passes over into the sinuses of the proboscis.
AFFINITIES OF KCHIUKUS LTNICINCTUS.
97
Reference Letters.
m.h., basal muscle between the setoe.
m c, circular muscle.
m.l., longitudinal muscle.
in.o., oblique muscle.
n., ventral nerve-cord.
lie., nuclei.
?«./., lateral nerve.
nu., nucleus.
o., ova.
o. (fig. 21), aperture.
(I.S., segmental organ (vcsi<de).
p. (fig. 1), papilhe.
p., ])(.'ritoiicum.
ph., ])rohoscis.
})h., })liarynx.
pp., granular protoplasm.
v., rectum.
s., secretion.
s.a., anterior seta.
s.d., dorsal sinus.
s.ni., mesenteric strands.
.t.p., posterior seta.
S.V., ventral sinus.
t.a., ordinary tissue with fibres.
t.c., connective tissue.
t.p., so-called '' glandular " tissue.
v., cavity of vesicle.
v.a., anal vesicle.
V.C., ciliated ventral surface.
w., body-wall.
SECOND SERIES. — ZOOLUGY, VOL. VlII.
15
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2nd Ser. ZOOLOGY.]
[VOL. VIII. PART 4.
THE
THA NS ACTIO XS
OF
THE LINNEAN SOCIETY OF LONDON.
THE TERRESTRIAL ISOPODA OF NEW ZEALANU.
BY
CHARLES CHILTON, M.A., D.Sc. (N.Z.); ELS. Lond.
RESEARCH FKLLOW, t'NIVERSITV OF EDINCUROH.
J
L () N I) O X
I'RINTKI) Foil I'lIK l.rXXKAN SOCrKTV
ItV lAVI.OK AMI. KKANCIS, KKI) I.ION COL'KT. Ff.KKT STREKT.
SOLD AT TIIK SDCIETV's APARTMENTS. BITRIJNGTON-HOLISE. PICCABILLT, W
AND BY LONGMANS, GREEN, AND CO., FATERNOSTER-ROW.
Jlay 190].
99 ]
IV. The TeiTCstrial Isopoda of Now Zealand. By Charles Chilton, 31. A.,
D.Sc. {N.Z.); F.L.S. Loudon; Bt'searcli Fellow, Unicersity of Edinburgh.
(Plates 11 16.)
■"I^HE first Terrestrial Isopoda described from New Zealand were those given by Dana *
in 1853, in his account of the Crustacea collected by the United States Exploring
Expedition ; in it he described and figured in considerable detail 7 species (includino-
one doubtful one), all of them from the northern part of New Zealand. In 1865 one or
two species were added by Heller f in the report on the Crustacea of the Novara
Expedition. In 1870 Mr. E. J. Miers % comjiiled a Catalogue of the New Zealand
Crustacea, and in connection therewith described some new species that were in the
collections of the British Museum ; he added 5 species, and his catalogue contains
altogether 12 species and one considered doubtful. During subsequent years a few
species were added by JVIr. G. M. Thomson § and myself || , and all the species known
were included in our " Critical List of the Crustacea Malacostraca of New Zealand." ^
In 1885 Budde-Lvmd ** published his " Crustacea Isopoda Terrestria," and added two
new species and mentioned most of those previously described, but as he was unable to
examine specimens he could give no additional information on them, and was obliged to
leave several of them under the heading of "imcertain sjiecies." Four other new species
were described and figured by Eilhol in 1885 in his " Mission de File Campbell," ff in
which he also gave references to previously described species.
In the present paper I endeavour to give a complete list of aU the New Zealand
Terrestrial Isopoda at present known, with descriptions of the various species and figures
where necessary. The material at my disposal consists chiefly of collections that I have
accumulated since 1884. Besides sj)ecimens that I have collected myself, I have
many from Mr. R,. Helms, formerly of Greymouth, Mr. J. McMahon, of Kenepuru,
Mr. W. W. Smith, of Ashburton, Mr. S. H. Drew, of Waugauui, Mr. H, Suter and
Mr. R. M. Laing, of Christchurch, and Mr. L. Hames, of Takapuua. Mr. G. M.
Thomson has very kindly placed in my hands the whole of his collection, including
* J. DaiKi : U. S. Exploring Expedition, Crustacea, vol. ii. pp. 713 et seqq (1853).
t Cam. Heller : " Keise der Novara," Zoul. Bd. 2, pp. 134-130 (1865).
X E. J. Miers: Annals &. Mag. iN'at. Hist. (ser. 4) xvii. pp. 'J.2b-'2,27 (1870); and Catalogue New Zealand
Crustacea, pp. 94-102 (1870).
§ G. M. Thomson: Trans. N. Z. Inst. xi. p. 232 & p. 249 (1879); and Annals & Mag. Nat. Hist. {ser. 0) xii.
pp. 225-227 (1893).
II C. Chilton : Trans. N. Z. Inst. xv. p. 73 & p. 149 (1883), & xviii. p. 159 (188(5).
% Trans. N. Z. Inst, xviii. pp. 141-159 (1880).
** " Crustacea Isopoda Terrestria,"' Copenhagen (18t>5).
tt " Mission de Tile Campbell,'' llecueil Mem. Acad. 8ci. (Venus) iii. part 2, pp. 439-446 (1885).
SECOND SEKIES. — ZOOLOGY, VOL. VIII. 16
100 DE. C. CHILTON ON THE
the type specimens of some species described by him. Through the kindness of Professor
F. Jeffrey Bell and Mr. R. I. Pocock, I have been able to examine type specimens of
Miers' species in British Museum, and Monsieur Adrien Dollfus has sent me species
from Europe that have heen most useful for comparison. To all these gentlemen I
desire to record here my hearty thanks.
Some of the work involved in the j)reparation of this paper was done in the Natural
History Department of the University of Edinburgh, where Professor Cossar Ewart
kindly gave me the use of a table, but a large part has heen done during vacations in the
laboratory of the University College, Dundee, and I desire to record my thanks to
Professor D'Arcy W. Thompson, C.B., for the facilities placed at my disposal, and for
permission to make free use of the rich stores of Terrestrial Isopoda in the collections
under his care, while to Dr. W. T. Caiman I am greatly indebted for much kind assist-
ance during the progress of the work.
I have, as far as possible, followed the classification and nomenclature adopted by
Professor G. O. Sars in his fine work on the Crustacea of Norway, and have to thank
him for sending me the parts hearing on the Isopoda. I have tried to give cill the
references specially dealing with the New Zealand sjiecies, but in the case of species and
senera that are also known from elsewhere, I have onlv 2,'iven one or two of the most
important ; for the benefit of workers in New Zealand, where books of reference are few,
I have given diagnoses of all the genera, taking these in most cases from Sars' work,
and have also given short notes on the characters of the families.
It will be seen that the Terrestrial Isopodan faima of New Zealand is fairly rich and
varied, all the families but one being represented. I am able to give 27 species, belong-
ing to 13 genera, of which only 8 or 4- are " uncertain species " ; for the sake of
comparison it may be mentioned that in the last list of the Terrestrial Isopoda of the
British Isles, as given by Canon Norman *, there are 20 species, belonging to 11 genera.
Moreover, it is probable that the number of New Zealand species Avill hereafter be
increased, especially when the North Island has been thoroughly searched, for at
present the majority of my specimens are from the South Island, and only a few more
or less haphazard collections have been made in the North Island. Of the outlying
islands of New Zealand, I have only two species from Chatham Islands, and one from the
Auckland Island. Three species are at present known from single sj)ecimens only, and
two others have been found only in one locality and on a single occasion. In addition
to the species given, specimens from ants' nests, probably belonging to Platyarthrus, were
referred to by Mr. W. W. Smith, in a paper dealing with some New Zealand Ants, and
were stated to have been sent to Europe with other collections from ants' nests. I have
endeavoured to trace these specimens, but without success, and so far Mr. Smith has
not been able to procure fresh specimens for me.
The Terrestrial Isopoda are \i'ell worthy of study from the point of view of the
geographical distribution of animals, and the facts of their distribution will be of great
value for testing the correctness of the views as to the origin of the fauna of particular
* Annals & Mag. Nat. Hist. ser. 7, iii. pp. 70~7S.
TEEEESTEIAL ISOI'ODA OF KEW ZKA l;.\ M). 101
countries and places, for they are strictly terrestrial animals, and as their youny are
hatched in the incubatory pouch of the female, it seems unlikely that they could cross
even comparatively narrow tracts of ocean, exeejjt by rare accidents, while a continuous
range of high moimtains would also be a formidable liarrier. In the NeAV Zealand
Journal of Science, vol. ii. (1884) p. 155, I have already called attention to the question,
and have also jjointed out that their distribution in aoy given land-area may be to some
extent influenced by floods in the rivers carrying logs with the Isopods attached to great
distances, and have given the following instance where this a{)pears to have actually
taken place.
The sjiecies Aruiadillo regulosus {=Cub(n-i.s i-egnlostis, Miers) is common on logs and
under the bark of trees in the bush, but I had not found it on the open Canterbury
Plains except at one place, Eyreton, where I got numerous specimens under some logs
that had been carted for firewood from the river Waimakariri, after having been washed
down by the river for at least twenty miles, probably furtlier, from places Avhere the
species was abundant. It seems likely that the Isopod had been washed down with the
logs, for I found it only at that particular spot at Eyreton, and af tei- the logs had all
been used it was no longer seen in that district.
It would, therefore, be interesting if some facts could be given as to the distributiou
of our Xew Zealaud species, especially of any that may l)e found in other countries.
Unfortunately, however, so little is at present known of the Terrestrial Isopoda of
Aiistralia aiul other lands of the soiithern seas, that little can as yet be said Avith
certainty.
Of the species, by far the greater part (18) are known only from New Zealand; two
species, Porcellio scaher, Latr. and Armadillidlmn vulgare, Latr., are cosmopolitan, aud
have probably been introduced by artificial means ; another species, Philoscia pubesceiis^
Dana, appears to be identical with a species found at the Cape of Good Hope and at the
Seychelles ; Actcecia euehroa, Dana, is found in Tasmania as Avell as in New Zealand ;
while Lie/id norfc-zcalandia', Dana, and Onisciis piincf.atns, Thomson, are reioresentcd
in Tasmania l)y closely allied species, and Ti/los neozelaniaus is jn'obably equally closely
related to T. sjnnidosus, Dana, from Tierra del Fuego. In the genus Trlchoiilscus it is
rather difficult to make any comparison between the numerous species, but the genus
is a very widely distributed one, and species are known from Tristan d'Acunha and
Valparaiso *, and from the Straits of Magellan f . The genus Armadillo is represented
in New Zealand by at least six S2)ecies, the greater number of the species of the genus
occur in the tropical countries, and Budde-LundJ has pointed out that about half of
them are frotn the islands and shores of the Pacific.
Of the distribution and occurrence of the diff'erent species in New Zealand itself a
little more can be said. Six species (i. e. lji[//a novce-zealandice, Tylos neozelaniciis,
Scypliax oruatiis, Actcecia euehroa, Actcecia opihensis, and Scyphoniscus waitatensis) are
* DoUfus : " Isopodes terrestres du 'Challenger,"' Socii'ti- d'Etudes Sdentifiques de I'liris, xii'^ Aniiei; (ISDO),
p]). 5 & 0 (separate oopj').
t Stebbing : Proc. Zool. Soc. London, 1900, Part iii. p. SfiG.
% Isopoda Tcrrestria, p. 16.
16*
102 DR. C. CHILTON ON THE
littoral, being- found on or near the sea-beach, and probably Sctjphax (?) aiicklaiidUe
should also be added to this list. Of these, LUjia noviC-zealandice is found all round the
New Zealand coast, and is very abundant under stones or sea-weed, especially on rocky
portions of the shore; Scyphax ornatus and Acicecia eiicJiroa are found on sandy beaches
either on the surface or burying themselves a little in the sand about high water mark
or a little lower ; Scijphax oruatus is probably abundant on all such beaches in the
North Island, but in the South Island has, so fjir, been recorded from "VVestport only ;
Ackecia euchroa is known from the south as well as the north, and is also found in
Tasmania. The remaining littoral species have as yet been recorded each from one
locality only.
Of the more strictly terrestrial forms, leaving out of account the two cosmopolitan
species PoreeUio scaher and Armadillklimn vulgare, and also Philoscla pubescens, which
is found at the Cape of Good Hope and elsewhere, we have Onlscus ptmctatus, found in all
pnrts of New Zealand ; ArinadUlo ambitiosns from all parts of the North Island, and from
Kenepiu-u and Greymouth in the South Island, laut not known further south ; while, on
the contrary, Armadillo rugnlosus and the three species of TricJioniscits are Avidely
distribiited in the South Island, but as yet not recorded from the North, though in the
case of Trlchoidsciis this is no doubt partly owing to their small size. Of the remaining
species too little is known to justify any general remark.
It may perhaps be well to mention here a few of what seem to be the more important
points brought out in this paper. I have been able to settle, in Avliat I hope will be
considered a satisfactory manner, uncertainties that have long existed with regard to
several of Dana's descriptions, and in so doing to reduce to the rank of synonyms some
species subsequently described (see Lkjla novce-zealandiiC, ScypJiax ornatns, PhUoscia
piibescens. Armadillo speaiosus) : I establish a new family, Scyphacidce, corresponding
mainly with Dana's subfamily Scyphacinw, Avhich had been ignored by most subsequent
writers, and show that the imperfect development of the seventh pair of legs, which Dana
had considered a character of the genus Scyphax, is merely an immature character which
in this instance is retained till a later period of life than usual, and settle the question as
to the relationship of Scyphax ornatus to Ackecia euchroa by showing that the only
connection between them is that both have the same habit of living on sandy beaches.
In the case of some of th(^ commoner species, I have had numerous sj^ecimens from
many localities, and have thus been able to make some observations as to the variations
that may be met with in these species.
In most of the species there is to be found on the dactylus a specially long and
peculiar seta which has characteristic forms in some, at any rate, of the genera.
Schiodte figured this " dactylar seta " many years ago in Tltanethes * alhus, and Weber
mentioned its presence in some species of Trichoniscus f , but I cannot find that any
one has drawn special iittention to it, though in some cases it is rather noticeable, and
together with the form of the dactylus itself, may be of use in readily identifying
* Bidrag til den uuderjordiske Fauna (Copenhagen, 1849).
t " Anatomisches iiber Triclioniscidon," Archiv fiii- ^likroskop. Anatomic, Bd. xis. p. 5S2.
TEEEESTEIAL KSOl'ODA OP NEW ZEALAND. 103
he liemis. In Ligia the dactylar seta is luiln-cxnched aud slightly cluhhed at the end ; in
TricIioniscK.s it divides into two brandies, each further subdividing into line filaments ;
in S'ci/p/ioiiiscKS it divides similarly, though differing a little in detail ; in 2)/Ion it
is rather short, unbranched, and has the distal iialf thicker and stippled-looking ; in
Actiecia cuchroa it is somewhat similar ; while in Acta-cia (?) uplheus'ts it is larger and
very distinct, and the stippled appearance of the distal portion looks under a high power
as if it were caused by the distal portion, resembling a narrow circular brush with short
hairs projecting all round it. It is sometimes lost in specially old aud large specimens,
but with this exception is always to be found in the genera mentioned ; I cannot,
however, find it at all in ScypJiax, Oiiiscm, Phlloscla, Armadillidinm, and Armaditlo.
Beyond the suggestion that it is a tactile organ, I can give no iiiformation as to its
function. A seta, probably aLso of a sensory nature, is found similarly situated in
Asellus aquaticm and some other Isopods, and also in many Amphipods, but in these it
is less prominent, and does not take such A'aried forms.
In all the genera, and especially those Jiitherto imperfectly known, I have examined
the mouth-organs in some detail ; an accurate knowledge of these will, I think, in time
help us on to\vards a natural classitication of this group, for they seem to be much more
constant than characters taken from the general shape of the body, from the uropoda, or
even from the presence or absence of air-cavities in the pleopoda. It is true that we
may get sudden variations in some of the mouth-parts, such as that I have described in
the outer lobe of the first maxilla of Sci/phoniscus, or by Dollf us hi the inner lobe of the
same maxilla in 3IcsarmmliIlo ; but these, occurring as they do in groups in which the
mouth-parts are otherwise very constant, are probably to be looked upon as sudden
variations or " sports " that have comj)aratively little value from a classificatory point of
view.
I give here a Table, based mainly on the mouth-parts, showing briefly what appear
to be the more important characters of the various families : —
A. ^landibles witli well-developed molar tubercle; inner lobe of 1st maxilla with
three plumose bristles.
I. Uropoda not concealed nnder pleon.
a. Antenuffi with flagellum multiartieulate; eyes large; male organ double . . Ligiid.e.
/;. Antcnnie with flagellum not more than G-jointed ; eyes small ; male organ
single Tkicho.viscid.e.
II. Uropoda concealed under pleon.
a. Segments of pleon separate Tylid.e.
b. First five segments of pleon coalesced Helleiud.k.
B. Mandibles without distinct molar tubercle, its place being taken by a brush-like
group of setae ; inner lobe of first maxilla with only two plumose bristles.
I. Maxillipcdes with terminal joints of moderate size ; lamellar longer than
masticatory lobe ScvpnyEiD^.
II. Maxillipcdes with terminal joints small aud almost rudimentary, hardly longer
than masticatory lobe.
a, Uropoda more or less projecting ; animals not rolling into perfect ball . . Omscid.e.
104 UK- G. CHILTON OK THE
c. Uropoda not projecting beyoud terminal segment ; animals rolling into
perfect ball Akmadilliid^.
While this table does not profess to be any very near approach to a natural classifi-
cation of the Terrestrial Isopoda, it is probable that the two large divisions A and B do
represent distinct groups in which development has proceeded on similar lines, in each
case leading from animals living on the sea-shore within reach of the waves and
breathing only very moist air, to others of pure terrestrial habits capable of breathing
ordinary dry air, i. e. the Helleridae in the one case, and the Armadilliidte in the other.
It is interesting to notice, too, how the protection affoi'ded by the animal's power of
rolling itself up into a ball has been acquired in different groups that are certainly
of independent origin, e. g. in the Tylidae, Hellerida?, Armadillidie, and to a less perfect
degree in some of the Scyphacidse {e.g. Acttecia), and perhaps also in some of the
Oniscida), and how similar is the general appearance of the body in each of these groups,
though of course the detailed arrangements by which it is acquired vary. If we go
beyond the Oniscoidea, we find a similar power of rolling into a ball and a somewhat
similar external apj)earance in the Sphseromidte and, among the Myriapoda, in the
Glonerida3.
A tabular arrangement of the Oniscoidea, based on much the same characters as
I have used, was given many years ago by Ulianin * ; his table, however, goes into
greater detail and separates the genera, and imfortnnately it is in the Russian language f.
For the benefit of those who may wish to identify their specimens without going to
the trouble of dissecting out the mouth-parts, I give the following artificial key to the
New Zealand genera, and a similar key to the species under each genus represented by
more than one species : —
A. Body convex ; animal capable of rolling into a l)all.
I. Flagellum of antenna 2-joiuted.
1. Outer branch of uropoda large and terminal Armadillidium.
2. Outer branch of uropoda small, inserted on the inner side of the enlarged base. Armadillo.
II. Flagellum of antenna with more than two joints.
1. Uropoda quite concealed beneath terminal segment Tylos.
2. Uropoda extending beyoud terminal segment and visible in dorsal view . . Actteciu.
B. Body more or less flattened ; animal not capable of rolling into a ball.
I. Flagellum of autenns many -jointed Lygia.
II. Flagellum 2-jointed.
1. Side-plates of metasome large, cxpainled PorcelUo.
2. Side-plates of nietasome small, adpressed . Metoponort/ms.
III. Flagellum with three to six joints.
1. Eyes large, crescent-shaped, of many ocelli Scyphax.
2. Eyes small, not more than three ocelli.
a. Body with longitudinal ridges Huplophthalmus.
* ' Crustacea Turkestriniii',' >St. Petersburg & Moscow, 1S75 (see Budde-Luuil, /. c. p. i'J.).
t I liiive to thank Mr. H. A. Webster, Librarian, University of Edin))urgh, for translating a portion of Dlianin's
work for me.
TERRESTKIAL ISOPODA OF NEW ZEALAND. 105
b. Jiody without lougitucliual ridges Trichoniscus.
'.^. Eyes of moderate size, more than three ocelli.
a. Pleon with lutcral expaiisious Onlscus.
b. Pleon with lateral ex])ansions.
i. Flagelluni much shorter than last joint of |i('(l[inek' Scyphoniscus.
ii. Flagellum ahout as long as last joint of peduncle Philoscia.
In tlie following- list 27 species are mentioned, bnt of these tliei-e are 4 which I have
not seen, aud which must be considered as more or less uncertain, though one, Armadillo
spinosHS, Dana, is in all pro1)ability a good species and distinct from the others given,
and I have tliereforc included it in the artificial key to tlic species. I have not been able
to do this with the other three species.
List of Species.
OXISCOIDEA.
I. LlGIID^.
1. Ligia iwrte-zeahuidice, Dana.
II. Trichoniscid/E.
2. Trichoniscits phormianus, sp. nov.
3. „ otakensis, sp. nov.
4. „ Thomsoni, Chilton.
.5. Hnplophtluthims Helmsii, sp. nov.
III. Tyli».e.
6. Tylo.s iieozelanicus, sp. nov.
IV. SCYPHACID^.
7. Scyphiix urnalus, Dana.
8. „ (?) aucUandue, G. M. Thomson.
9. Scyphoniscus waitatensis, nov. gen. et sp.
10. Actcecia euchrou, Dana.
11. „ opihensh, sp. nov.
V. OnISCID/E.
12. Oiiiscits ijdiictatus, G. M. Thomson.
13. ,, Leiieptirensis, sp. nov.
II. ,, CooL'ii, Filhol {nut accn).
15. Pliilosciu ptibescens, Dana.
16. „ novce-zealanditey Filhol (^not seen).
17. Porcel/io sraher, Latreille.
18. y MctopoiKjrfhus pru'niosus, Brandt.
VI. Akmadilliid.k.
19. ArmadUHdhiiii vulgare, Latreille.
20. Armadillo amljitiosi/s, Budde-Lund.
21.
22.
23.
24.
.25.
26.
27.
Dante, Heller.
s-peciosus, Dana.
rmpilosus, Miers.
niunolinus, Dana {not seen)
Handltoni, sp. nov.
Marmahoni, sp. nov.
sp/nosHs, Dana (not seen).
Family I. LIGIID^.
In this family the antennae have the flagellum multiarticulate, i. c. with more than
six or seven joints, the mandible has a well-developed molar tubercle with triturjiting
surface, the inner lobe of the first maxilla bears three plumose bristles, the terminal
portion of the maxillipede is of moderate size and more or less distinctly divided into five
joints, and the external male organ is double.
The family contains scA^eral genera, the best known being Ligia, Liyidiuni, and
Titaaethes. The genus Geoligia, Dollfus, appears to be very near to Ligia, but the
only known species, G. Simoni, lives far away from the sea, while all the species of Ligia
are found on the sea coast.
106 DE. C. CHILTON ON THE
If Styloniscns maffcllanicus, Dana, belongs to Trichoniscus as Stehbing * tbinks, it is
evident tbat tbc distinction between the Ligiidoe and the Trichoniscidge as regards the
antennae breaks down, for in that species the antenna may have the tlagellum with as
many as ten joints. Dollfus, when describing this species, had previously stated that
Styloniscus, Dana, is very near to Ligidlmn, and differs from it only in the uropods, which
want the long hairs characteristic of that genus t ; in making this statement, however,
he may have had in his mind also the species Styloniscns (jracilis, Dana, in which the
uropoda do resemble those of Ligldmm as Stcbbing has also pointed out, but it is
doubtful whether this species is really congeneric with S. iiuujellaniciis.
In any case the differences betM'een the Ligiidoe and the Trichoniscidfe are not great,
and the existence of genera intermediate in characters is only what we may natiu'ally
expect.
Dana placed Styloniscus in his sub-family Scyphacina>, liut from Stebbing's description
of the month-parts of ^S*. magelkmicus it is evident that that species at any rate cannot
come under the family Scyphacida; as I have defined it further on.
Genus 1. Ligia, Fabricius, 1798.
Liy'ia, Bate & Westwood^ British Sessile-Eyed Crustacea, ii. p. 44.2 (1868).
Ligia, Budde-Lundj Crustacea Isopoda Terrestria, p. 258 (1885).
Liffia, Sars, Crustacea of Norway, ii., Isopoda, p. 155 (1899).
The generic characters are given by Sars as follows : —
" Body regularly oval, or oblong oval, moderately convex above, with the metasome
not abruptly contracted ; last segment rather broad, with distinct epimeral plates. Eyes
large and convex. Antennulse very small, with the last joint rudimentary, uodiform.
Antennae rather strong and elongated. Mandibles with a ciliated lappet and numerous
j)enicils behind the cutting part. Maxillipeds comparatively short and stout, with the
terminal part rather expanded, epignath rounded. Legs gradually increasing in length
posteriorly, dactylus distinctly bi-unguiculate. Opercular plate of pleopoda sub-
branchial. Uropoda more or less elongated, basal part not produced inside, rami
narrow, styliform, subequal, each with a single apical spine."
This is the only genus of the family that is represented in New Zealand, and the
single species, i. novce-zealamUcc, described below, agrees well with the characters of the
genus as just quoted from Sars. It differs, however, from the characters of the family
in that the two hairy bristles on the inner side of the second maxilla are wanting, and
the terminal part of the maxillipeds, though showing distinct evidence of five joints, has
the three joints preceding the terminal one united together into one plate with the
sutures only partially indicated. The external male organs are considerably different
from those of the typical species L. oceanica. In all these points L. cmstralicnsis, Dana,
from Australia, closely resembles L. iiovce-zealandUv, and as these pecixliarities are
probably shared by other species which like them are nevertheless true Ligice, it will be
well to slightly modify the characters of the family as laid down by Sars in order that
these species may be included.
*■ i'roc. Zool. !5uc. I'JOo, p. 5{)G. t Mission du Cape Horn, Crustuces, p. 72.
TERRESTRIAL TSOPOUA OF NEW ZEALAND, 107
1. LiGIA NOV/E-ZEALANDI/E. (I'l. 11. fig. 1.)
Lijgia novi-zealandi(B , Daua, U.S. Explor. Expcrl. xiv. Crust, part ii. p. 7o9, pi. xlix. fig. 2 (1853).
Liyia nov(B-seal undue, Miers, Cat. Crust, of New. Zealand, p. 103 (187(5).
Ligia quadrata (Hutton, MS. Cat. X. Z. Crust.), G. .M. Thomson, Tran.'*. N. Z. Inst. xi. ]). 23:2, pi. x .\,
figs. 4& 4a (1879).
Ligia noKC-zeulandixc, Buckle-Lund, (^ni.stacea Isopoda Terrestria, p. 271 (188')).
Ligia quadrata, Budde-Lund, /. c. p. 271 (1885).
Ligia novce-zealandice, Filliol, Mission do Tile Campbell, p. 415 (1885).
Ligia quadrata, Filliol, /. c. p. 415 (1885).
Ligia neo-zelanica, Thomson & Chilton, Trans. N. Z. Inst, xviii. p. 157 (188U).
Ligia quadrata, Thomson & Chilton, /. c. p. 157 (1880).
Specific description^. — Body elongate oval, about twice as long as broad, rather
convex; surface finely granular, sometimes with minute seta;, giving it a i)uuctate
appearance. Outer antennae slender, minutely setose, two-thirds the length of the body ;
fifth joint of peduncle as long as the third and fourth combined, flagellum fully twice as
long as the fifth joint with about twenty joints. Eyes large, subquadrate, distinctly
angled towards the middle line, facets very numerous and of small size. Vertex with a
transverse depression just posterior to the angle of the eyes, interrupted in the middle.
Posterior bordei- of the first and second segments of the mesosome transverse, not
produced backwards at the lateral angles ; lateral angles of the remaining segments
progressively more and more produced backward, those of the seventh segment reaching
as far as the angle of the third segment of metasome. Side-plates (" epimera") large,
distinctly marked off from the middle jiart of the segment by a longitudinal sulcus in
the second, third, and fourth segments, the sulcus very iodistiuct in the remaining
segments. Legs sj)inose, gi^adually increasing in length posteriorly, the first and second
in the male having the carpus broadened, and the propodos and dactylus impinging
against it to form a subchelate hand, the first being broader than the second ; in the
female all the legs simple ; in each leg the dactyhts has a secondary slender nail about half
the length of the terminal nail; at the base of the terminal nail arises on the outer side
a long seta slightly clubbed at the end and reaching as far as the end of the terminal nail.
Terminal segment of the metasome subquadrate, its lateral angles acute but not much
produced, posterior margin regularly convex in the middle. Uropods with the peduncle
subcylindrical, about half the length of metasome ; the two rami of nearly equal length,
the outer often rather the shorter and more slender, both tapering, minutely setose, and
with one or two apical setae.
Colour : yellowish, closely speckled with black, givuig a greyish or slaty effect.
Length about 12 mm., breadth about 5'5 mm.
Habitat. — Very abundant on all the coasts of New Zealand, generally found under
stones or seaweed about high-water mark, but sometimes extending a little further
inland. It runs with great rapidity when disturbed.
Remarks. — It is only after considerable hesitation that I have united Liyia qiuulrata,
Thomson, with Lijgia noci-zealandice, Dana. When Mr. Thomson described his species
he was acquainted with Dana's description, but found that it differed from his sjjecimens
SECOND SEKIES. — ZOOLOGY, VOL. VIII. 17
108 DK. 0. CHILTOX OX THE
in some points tliat appeared very definite and well marlved, and lie therefore established
for them the new species Ligia quadi-ata. All specimens subsequently examined, both
by Mr. Thomson and myself, were found to agree with the characters as laid down for
i. quadrata, and thus to differ from L. iiovi-zealandice, Dana, and hence in our " Critical
List of the Crustacea Malacostraca of New Zealand," * under the heading Ligla novi-
zealandUe, the remark is made " I do not know this species, G. M. T." The points in
which Dana's description differed from our specimens are : —
{(i) The surface of the thorax and abdomen " covered with very short hairs.'"
{b) Base of caudal stylets " nearly as long as the abdomen."
((?) Branches of caudal stylets " qaite unequal " and the longer " hardly as long as
the thorax."
In none of the specimens that I have examined could the dorsal siu'face be said to be
" covered with very short hairs," and Mr. Thomson tells me that no liairs are to
be found in living specimens, which he has recently re-examined at my request, as I
thought it just possible that the hairs might have got worn off in the spirit sj^ecimens
tlaat I brought from New Zealand with me. I have been anxious to get for comparison
specimens from the Bay of Islands, where Dana's tyjie specimens Avere obtained, and
though I have not been successful in this, I have in Mr. Thomsoa's collection specimens
from Waiwera, a locality nortii of Auckland and not very far remote from the Bay of
Islands, and I find that these differ a little from our South Island specimens, and thougli
I regard them as undoul)tedly the same species, they show some slight approach towards
Dana's description. Thus the antennae are slightly longer and more slender and
distinctly more hairy than in the typical specimens of Ligla quadrata, and the surface
of the body when viewed with a higher power shows, especially at the edges of the
segments, very minute little setfe which, though they scarcely project beyond the surface
and are not deserving of the name of "very short hairs," must, I think, have given the
appearance which Dana has thus described. In South Island specimens these minute
points are much less marked but can occasionally be made out. The uropoda in the
"Waiwera specimens are a little more slender than in South Island ones, but as in them
the base is only about half as long as the abdomen, and I have not seen any in which the
base is '* nearly as long as the abdomen," but it must be remembered that in young
specimens with which Dana perliaps liad to deal the uropoda are considerably longer in
proportion than in fully-grown specimens. The branches of the uroj)oda are again
usually of nearly the same length, though the outer one is generally a little the shorter,
and the variation in their relative lengths is pretty considerable, and specimens in which
the difference was more marked than usual may have led Dana to describe them as
" quite unequal." The longest branch is, however, always much shorter than the thorax,
and I must regard Dana's statement that it is "hardly as long as the thorax" as an
unintentional exaggeration or else a mistake for " hardly as long as the abdomen.'''
In his " Catalogue of the New Zealand Crustacea," Miers refers specimens in the
British Museum to Dana's species without any question beyond remarking that "the
rami of the caudal aj^pendages are equal except in one specimen, where they are slightly
* Trausiiotious New Zealand Institute, xvili. p. 157.
TEEEESTEIAL ISOPODA OF >;EW ZEALAND. 109
unequal." When visiting the British Museum I found, however, that the speciniens
are labelled " ? Ligia notce-zealandiai, Dana," and tliat, so far as can be seen in their
dried and imperfect condition, they resemble my Waiwera specimens very closely, and
thus differ from Dana's descriptions in the other characters that I have pointed out as
well as in that of the rami of the uropoda.
I have discussed this question at what will probably be tliought to be undue length,
but I think that full reasons should always be given before one species is regarded as
the synonym of another, and it is well to hesitate before venturing to dispute the
accuracy of Dana's descriptions.
As this species is the largest and one of the commonest of the Terrestrial Isopoda of
New Zealand, and is, moreover, of a more generalized type than the others, it is deserving
of close attention by any who wisli to study the group, and I therefore give here a fairly
full accoimt of its external anatomy. I do not propose to consider its internal anatomv,
tliough it would no doubt repay careful consideration; indeed, I do not know that the
intenial anatomy of any species of the genus has yet been worked out in detail, thoun-h
many yeai-s ago Lereboiillet published an excellent paper on a species of the closely
allied genus Ligid'mm *, and Max Weber has more recently given a more minute account
of the anatomy of some species of the family TrlchoitiscicUc which comes close to the
Ligiidce f .
Detailed Description of Ligia nov;e-zealandia^. (PI. 11.)
The size is naturally subject to some variatioii, but all the specimens that I have seen
are considerably smaller than fully-grown specimens of L. oceaiiica. The following
measurements may be taken as about the average : — length of body 12 mm. ; greatest
breadth 6 mm. ; length of mesosome 7'5 mm. ; of metasome J< mm ; of antennic 10 mm. ;
of uropoda 5 mm. (base 2 mm., rami 3 mm.).
The head is oval, about three times as broad as long, the anterior margin regularly
convex and without lateral lobes ; the eyes are large and occupy nearly the Avhole of the
lateral margins, their anterior and posterior sides meeting at a distinct angle; the facets
are small and very numerous.
The surface of the head shows a transverse depression, interrupted in the middle, just
posterior to this angle of the eyes.
The first segment of the mesosome is about as long as the head. Its epimeral portions
extend anteriorlv about to the middle of the lateral margins of the head, tlie suture
marking them off from the central portion being indistinctly marked in posterior pai't of
the segment only ; the posterior margin straight ; the second and third segments similar
but a little longer than the fii'st ; the fourth segment the widest, its posterior margin
slightly concave, and lateral angles a little produced backwards ; fifth, sixth, and seventh
segments gradually narrowing; lateral angles aciite and more and more produced
backwards, those of the seventh segment reaching nearly to the postero-lateral angles of
* "Memoire siir la Ligidie de Persoon {Li(j{dium Persooni, Brandt)," Ann. d. Sciences Nat., Seconde Sc^rie,
tome XX. pp. ] 03-141', PI. 4 & .5.
t " Anatomisches iiber Trichonisciden," Archiv f. Mikroskop. Anatomic, lid. xix. pp. 570-048, Tab. xxviii.-xxix.
17*
110 DK. C. CHILTON ON THE
the third segment of metasome. The sutures dividing the epimera from the central
portions are fairly evident in the second, third, and fourth segments, but are indistinct
in the fifth, sixth, and seventh ; in some specimens they are indistinctly marked in the
fifth segment also, and the distinctness of the sutures is, I think, a character that is
subject to considerable variation, though Dollfus has established a nevr genus Geoligia
differing from Ligia only in having the epimera not distinct for a species, G. Slmoni,
found in the forests of Venezuela at an altitude of 1200 metres *.
The metasome is considerably narrower than the mesosome, the first and second
segments small and withoiit distinct epimeral projections, third, fourth, and fifth
segments subequal with well-developed epimera, lateral angles acutely produced
backwards, those of the fifth segment reaching very nearly to the postero-lateral angle
of the sixth segment ; sixth segment with its posterior margin deeply hollowed on each
side for the base of the uropoda, its central part regularly convex.
Surface of whole body slightly granular and with a few irregulai-ities, showdng under
a high power, especially at the sides, very minute set;ie which scarcely project beyond the
surface.
The antennul(e consist of the usual three joints, the first much the broadest, second
nearly as long as the first but narrower, the third very small, rounded at the end ; a
few minute setfe are present, chiefly on the second joint, but no " sensory setye " were
observed.
The autemxp are repi'esented in PL 11. fig. 1 u.'- : the first three joints are subequal,
short, nearly as broad as long, the fourth joint shorten than the fifth but broader ;
flagellum about as long as the last three joints of tlie peduncle together ; in the fig. «.-,
taken from a specimen 12 mm. in length, the flagellum is composed of fifteen joints, but
it may contain a greater or less number, Thomson says "flagellum 16- to 23-jointed."
There are numerous short and rather stout set;e oa the last three joints of the peduncle,
and finer setfe on each joint of the flagellum. In the South Island specimens these
latter are usually shorter than the breadth of the joint from which they spring, but in
the Waiwera specimens they are fully as long as the joint is wide, or may even slightly
exceed this length.
The mouth-parts are w^ell developed and of a more generalized type tban in most other
Terrestrial Isopoda. The upper lip calls for no special remark ; it is rounded, wdth a very
shallow emargination at its extremity, and provided with numerous sliort seta? in the
usual manner. The mandibles are strong and of the same general shape as in Ligia
oceamca ; in the right mandible the outer cutting-edge is formed of four stout teeth ; the
accessory appendage is slender, bends abruptly about the middle, and on its inner side is
prolonged into a slender acutely-pointed process ; its terminal part is nearly transparent,
and its basal part appears very pale brown and is evidently much less highly chitinized
than the corresponding part in the left mandible ; Ijetvveen this accessory appendage and
the molar tubercle is a soft membranous lobe, rounded at the end and thickly covered
with seta}, tho.se along its inner margin being longest and plumose; molar tubercle strong,
curving iuAvards, its truncate extremity covered with closely-set rows of short, stout seta?.
* " Voyage de M. K Simon au A'enezuola," Ann. Societe entom. dc Franco, vol. Ixii. (IbO^), p. 3-13.
TEREESTRIAL ISOPODA OF NEW ZEALAND. Ill
The left mandible has the outer cutting-edge of four teeth much as in the right, but the
accessory appendage is much stouter, thick and dark brown similar to the outer cutting-
edge ; it ends in several stout teeth, of which tlie outer one is the longest and strongest,
and is followed by two short double teeth ; the membranous lobe and the molar tubercle
are similar to those of the right mandible.
The loiccr lip consists of two broad lobes somewhat widely separated, with the ex-
tremities broadly rounded and thickly covered with short setae, most of which are
directed inwards.
Thejirst maxillcB are practically the same in form as tliose of Ligia oceanica, the outer
lobe being stout, longer than the inner, and provided at its extremity with about eight
strongly-curved setae, those to the outer side being the longest and stoutest. The inner
lobe is more delicate, apparently membranous ; its extremity appears concave on its inner
side, and it bears the three characteristic plumed setae, the distal one being very short
and the proximal one the longest.
The second maxillcs are stoutly formed, oblong in shape, about two and a half times as
long as broad, the extremity irregularly rounded, its inner half and the distal portion of
the inner margin fringed with sette ; there is also an oblique row of setae on the surface
of the maxilla near the end ; the outer margin bears fine setae towards the base, the more
distal portion being apparently free. I can find no trace of the two plumose setae which
are found in Ligia oceanica towards the end of the inner margin, and the division into
two lobes, which is partially indicated in Ligia oceanica and other species, is not recogniz-
able at all in the present species.
The maxiUipedes also show rather more coalescence of the different parts than those
of Ligia oceanica ; the first joint {coxa) is short and very broad, and the exopodite arising
from it is short, subtriangular, rounded at the end, and its free margins fringed wdth
setae ; its articulation with the bases is oblique, extending further distally on the anterior
(upper) surface than on the posterior, the extremity of the coxa being strongly convex
on the anterior surface but straight on the posterior ; the next joint [hasos) is nearly
oblong, fully two-thirds as broad as long, its outer margin slightly convex and bearing a
fringe of fine setae ; the inner margin is straight, and is bent inwards (i. e. upicavds, in
the usual position of the mouth-parts) to form a piece at right angles to the outer surface
of the maxillipedes ; this is thickly covered m ith short tine setae, and narrows distally
where it extends on to the masticatory lobe, which is formed ])y a prolongation of the
inner part of the basos ; the masticatory lobe is truncate distally, and bears there two
stout teeth and many finer setae. The terminal portion of the eudopodite ('' palp 'j
shows indications of being formed of five segments, of which only the first and last are
completely separated from the others, the second, third, and fourth being coalesced into
a flat plate w ith the lines of sutvire visible towards the inner side only ; on the outer side
the extremity of each joint is marked by one or two stout sette, the rounded inner
margins of the last ibur segments are thickly covered with short setae. Fig. mxp* shows
the maxillipede from its anterior aspect, i. e. that next to the second maxilla, and from
this point of view the conuectioji of the masticatory lobe with the basos can be clearly
made out ; w hen seen from the posterior (fig. uoxp.) the junction of the basos with the
112 DR. C. CHILTON OX THE
succeeding joint extends right across to tlie inner margin and makes the masticatory lobe
appear separated from the hasos though it is directly continuous with it on the anterior
surface.
The Jirst fair of logs differ considerably in the two sexes. In the female (PI. 11.
fig. \f} $ ) the appendage is similar to the succeeding pairs, though rather shorter; the
basos is somewhat oblong, and bears a few stout seta3 on its upper or inner side at the
distal end ; the lower or outer surface has a slightly hollowed depression, into Avhich the
more distal joints of the limb rest when they are bent back upon the basos, as they are
in the usual position of the legs. Tlie shape of the other joints and the arrangement of
the seta? on them can be readily made out from the figure : the propodos is cylindrical,
much narrower than tlie carpus, and lias on its inner margin a regular row of about
six short setae ; tlie dactylus is somewhat slender, and has the basal portion covered,
especially on the outer side, with short fine setae and a few spiniform ones ; the terminal
portion forms a strong, curved nail with margins regularly curved and without setie; the
accessory nail is about half as long as the tei'minal one and much more slender ; at the
base of the terminal nail arises from the outer margin a long, well-marked seta about as
long as the terminal nail, but usually curved backwards and having a slight club-like
swelling towards its extremity. These points, with regard to the dactylus, are repre-
sented in PL 11. fig. 1 p? $ *, which shows the extremity of the seventh pair of legs,
but with very slight modification the figure and description apply to all the pairs.
In the male tlie first pair of legs is nuich stouter than in the female, tlie meros is larger
and more triangular, Avhile the carpus is ovoid, being much expanded on the inner side,
and against it the joropodos and dactylus closely impinge and form a powerful subchelate
band ; the propodos is stout and slightly curved, and the dactylus rather stouter and
shorter than in the female. Tiie general appearance of this appendage in the male is
very like that of one of the guathopoda of an amphipod, or like the first pair of legs in
Fhreaioicus, but in these the subchelate hand is formed by the dactylus impinging
against the enlarged and swollen propodos, while in the present species the jn-opodos and
dactylus together impinge against the enlarged carjius.
The second jjair of legs in the female is quite similar in form and size to the first. In
the inale it has the form of a subchelate hand like the first pair, but the carpus is much
narrower and its inner edge, which forms the palm, is not so convex.
The third pair of legs in the, female is quite similar to the j)receding pairs in form, but
is usually a trifle longer; in the mcde it may have the carpus very slightly expanded, as
in the first and second pairs, but more generally it has nothing of the gnathopod form
and is almost identical with the corresponding appendages of the female.
The succeeding pairs of legs in both sexes are gressorial and similar to one another in
"•eneral form, but there is a gradual increase in length and sleuderness as we pass to the
seventh pair. In all there is the smooth, slightly concave depression on the basos against
which the other joints impinge, and the dactylus always bears the characteristic clubbed
seta already described, though in spirit specimens this may sometimes be lost, moi-e
frequently so in older and larger forms. The seceiith leg is represented in PI. 11.
fig. 1 p." ? , and it is scai'cely necessary to give a detailed description of it.
TERRESTRIAL ISOPODA OF NEW ZEALAND. 113
The pleopoda present the usual features, and all consist of a short basal portion or
protopoclite, from ^vhich spring the endopodite and exopodite ; of these the endopodite
is entirely branchial and has its margins perfectly free from seta), while the exopodite;
appears to be mainly opercular and usually has its margins more or less fringed with
plumose sette. It will be convenient to descril)e the [jleopoda of the female first, aud
then to point out the special modifications in the male.
The first j)^coj)od has the protopodite short and Iji'oad, rouglily rectaiignkir l)ut
narrowing a little externally ; on the outer side it bears a small rounded appendage with
margins free from setae, which appears distinct from the rest of the protopodite though
not distinctly separated by any suture or articulation. This appendage, which is found
on the first and second pleopoda of both sexes, is perha})s to be looked upon as an " epi-
podite" ; it will, at any rate, be convenient to refer to it by this name. The exopodite
issubovalin shape and much larger than the endopodite ; its margin bears a few irregular
plumose setae.
The second ijleopod of the female closely resembles the first, but is slightly larger ;
from the centre of the sternal plate of the segment is a small subtriangular projection,
truncate at the extremity ; the epipodite is longer, more pointed at the end, and bears
numerous finely -plumose seta?.
The third, fourth, and ffth jilcojwda nrt' n\\ similar in form, but each a little larger than
the preceding one. PL 11. fig. 1 j^Z/j.'* c3' shows the third pleopod of a male specimen,
but will serve almost equally well for that of a female ; from the centre of the sternal
plate of the segment arises an oval projection, which is produced distally to a fine point ;
there is no trace of the ejiipodite, but on the inner side the protopodite is pi'oduced into
a triangular acutely -pointed process the margins of which bear several jdumose setae ;
the exopodite is much larger than the endopodite, and is distinctly opercular in structure
and has the margins regularly fringed with long plumose hairs ; the fourth and fifth
pleopods are similar, but as we proceed posteriorly the endopodites, being less covered
by succeeding appendages, become more strongly chitinized and more abundantly su2>plied
with stellate pigment cells, the fifth one naturally most so, as it is completely exposed.
In the male the first and second pleopoda are specially modified for the purjjosc of
coptilation. In the first pair the pleopod itself is not very different from that of the
female, though the exopodite is rather larger and the endopodite is more pointed at the
apex, but it is closely associated with the external male organ, which no douht spriu"'s
from the last segment of the mesosome but is adherent to the pi"Otoj)odite of the pleopod
and in dissection always comes away with it ; it forms a long, narrow process, slightlv
narrowed and curved outwards at the end ; this is grooved throughout its whole len"th
on the posterior side, aud during life is closely ])re'ised against the anterior side of the
long process formed by the endopodite of the second pleojjod, and with it forms a tube
for the passage of the semen.
In the second pleopod of the male the protopodite and the exopodite present little
modification, but the whole of the endopodite is specially modified ; it forms a 2-jointed
penial appendage, strongly chitinized throughout, much more so than the male oi'gan
proper already described ; the first joint is short, lies transversely, and is moved by
114 DR. C. CHILTON ON THE
powerful muscles ; the second is long, semicylindrical, narrowing and curving outwards
at the extremity, which hears numerous fine short setfe with points directed away from
the apex ; the anterior aspect shows a well-marked groove, from the sides of which near
the middle numerous seta) project inwards towards the groove and appear to be for the
purpose of holding the male organ against this appendage and keeping it firmly in its
place; they probably do so by interlocking with similar setie on the male organ itself,
though these cannot be well made out.
The nrojwda are of the usual form, the basal portion irregularly cylindrical and some-
what twisted so that when detached it is difficult to get it to lie in its natural position ;
the outer ramus slightly narrower than the inner, but usually nearly or quite as long; it
bears two long setae at the apex, shorter setae being usually present on the inner branch ;
surface of base and rami covered with fine short setiie, giving it a roughened appearance.
Family II. TKICHONISCID^.
This family was established by Sars for Trichoniscus and a few other genera that had
previously been classed under the Ligiidse. It is closely related to that family, l)ut may
be recognized from it by the fact that the flagellum of the antenna has only a few joints
(not more than six or seven) ; the eggs are small, and contain only a few ocelli (usually
three), and the external male organ is single. The animals are usually small and live in
damp situations, none of the pleopoda being jirovided with air-cavities.
Two genera of this family — i. e., Trichoniscus and Haplufihthalmus — are represented
in New Zealand.
Genus 1. Tkichoniscus, Brandt. (PI. 12. figs. 1 & 2, and PI. 13. fig. 1.)
Trichoniscus, Braudt, Conspectus Crust. Oniscodorum, p. 12 (Bull. Soc. Moscou, vi. p. 174) (1833).
Pkilouffria, Bate & Westwood, Brit. Sess.-eyed Crust, ii. p. 454 (1868).
TrichoniscHS, Budde-Lund, Isopoda Tcrrestria, p. 243 (1885).
Trichoniscus, Sars, Crustacea of Norway, ii. p. 160 (1898).
Trichoniscus, Stabbing, Proc. Zool. Soc. London, 1900, p. 565 (1900).
Generic Characters. — Body more or less oblong, attenuated behind. Cephalon rounded
in front, with small though distinct lateral lobes. Side-i^lates of the three posterior
segments of mesosome more prominent than those of the four preceding segments.
Metasome abruptly contracted, with the epimeral plates of the two anterior segments
not concealed ; last segment narrowly truncate at the tip and slightly emarginate on
each side. Eye small but distinct, consi.sting of only three visual elements imbedded in
a dark pigment. Antennulse Avith the first joint rather large and curved, last joint
generally longer than the second. Antennae everywhei'e clothed with small appressed
spikes ; flagellum much shorter than the peduncle and gradually tapering distally. Oral
parts considerably prolonged, giving the buccal mass a pronouncedly conical form. Left
mandible with two, right with only a single penicii l)ehind the cutting-part. Maxil-
lipeds with the distal joint of the basal part rather large, and forming at the end outside
a broad lamellar expansion finely ciliated at the edge ; terminal part lanceolate, with the
TEEEESTEIAL ISOPODA OF KEW ZEALAND. 115
outer four joints confluent; masticatory lohe nearly as large as the terminal part, and
terminating in a narrow, finely-ciliated lash ; epignath ol)long-linguiform, with a rounded
expansion at the hase. Legs of moderate size, slightly increasing in length posteriorly ;
outer joints extremely spinous. Inner plate of first pair of pleojwda in male greatly
produced, hiarticnlate ; that of the second pair of diiferent structure in the different
species. Uropoda with the basal part rather broad and flattened, both rami terminating
in a pencil of delicate hairs. [Sars, /. c. pp. 160-161.]
Key to Spraies-.
1. Dorsal surface and imtcnna; with distinct tliou^li irregular tuljorcles T. otakensis.
2. Dorsal surface smootli or nearly so.
a. Surface with scattered longisli set;e. Animal small (I mm.) 7'. phormianus.
h. Surface without scattered sette. Animal large (7 mm.) . . T. Thomsoni.
1. TlllCUONlSCUS PHORMIANUS, .sp. nov. (PI. 12. ttg. 1.)
Philougria rosea, Chilton, Trans. N. Z. lust. xv. p. 11!) & p. 7o (iu part) (188,3) \_aot of Koch].
Philougriu rosea, Pilhol, Mission de I'ile Campbell, p. 13!) (in part) (1885).
Philijgria rosea, Thomson & Chilton, Trans. N. Z. Inst. .wiii. p. l."J7 (in part) (188Gj.
Specific Description. — Male not differing markedly from the female iu the general
shape of the body. Body oblong-oval, about two and a half times as long as broad.
Dorsal surface not very convex, smooth, or with a few small granulations and irregu-
larities ; cephalou and each segment of the mesosoine with a few scattered, rather long,
stout setfB, which are irregularly arranged and extend more or less over the Avhole
surface, but are most readily seen at the sides, especially in the anterior segments; on the
metasome there are few or none ; these seta^ readily break off iu spirit specimens.
Cephalou transversely oval, lateral lobes small, front slightly convex. Segments of the
mesosome of the usual form, the last three Avith the posterior angles recurved aud
acuminate. Metasome about one-quarter the length of the body, ratlier narrow ; first
two segments short, epimeral plates of the next three small and appresscd ; last segment
with the terminal expansion rather broad, the posterior margin straight or slightly
convex, and bearing three or four small sette.
Antenmc a little less than one-third the length of the body, rather slender, with long
setai at the extremities of the second, third, and fourth joints and along the inner margin
of the fifth ; these may arise from slight prominences, but the inner margin of the fifth
joint does not bear the distinct tubercles found in the next species ; outer margin of the
joint straight, with short fine seta; ; fiagellum as long as the fifth joint, of four joints
(sometimes only three), pencil of hairs at extremity long.
Uropoda long, outer branch more than twice as long as the base, conical, narrowing to
apex ; inner branch nearly as long, but much narroAver throughout and tapering very
gradually to the apex ; both covered with small appresscd sctui and Avith long seta' at
apex.
SECOND SERIES. — ZOOLOGY, VOL. VIII. 18
116 DR. C. CHILTOX OX THE
Colour light broAA'n, with irregular marblings of a darker brown.
Size about 4 mm.
Hdbitat. — Very common all over Canterbury, frequently found on the dead decaying
leaves of the New Zealand flax (Fhormium), and always in damp situations. Also from
Dunedin, Kenepuru, Greymoixth.
Remarks. — The separation of the New Zealand species of Trichoniscm presents
considerable difficulty, and it is quite possible that some modification may have to be
made in the division I am here adopting, though it is the best I can make with the
material now at my command.
The present species was originally confused by me with T. otakensis, and both
referred to Philougria rosea, Koch. Furtlaer investigation has shown that I was
dealing with two species, and tliat though each presents considerable resemblances to
Philougria rosea, Koch, neither can be considered as identical with that species.
The species now under consideration appears to be distinguished from the next species,
T. otakensis, by the smoother surface of the body, the more slender and smoother
antennye, the presence of stout sette on the surface, and by the fact that the male and
female are aj^proximately of the same general shape.
The stout setae on the cephalon and mesosome are very characteristic, but they readily
fall off in sjiirit specimens, and confusion may thereby be introduced. Some of my
specimens are now so free from all trace of these setse that I have sometimes been
inclined to think that there must be a form destitute of setie. On the other hand, I have
specimens from Kenepuru collected by Mr. MacMahon in which the setse are still
present ; they are rather more numerous and shorter than in Canterbury specimens,
and the surface is more uneven and tuberculated ; it is possible that these specimens
will require a separate sjjecies to be established for their reception, but in the meantime
I prefer to regard tliem merely as a variety of T. j)hor)uiaiius.
The mouth-parts show such a close general resemblance to those of other species of
the genus, such as T. rosens, that I have not given figures of them. The mandibles and
first and second maxillse present the usual characters ; in the maxillipedes the articulation
between the coxa and basos is oblique from the external to the internal face like that
already described in Ligixi iiovce-zealandice ; the masticatory lobe into which the basos is
prolonged is shorter than the palp, and bears at the end a sepai'ate conical portion,
thickly covered with fine seta? arranged radially and produced distally into the short
terminal lash ; in these points this species appears to agree closely with Trichoniscus
Leydigii as figured and described by Max Weber *.
The seven pairs of legs present no feature of special importance, and I have not
observed that any of them are specially modified in the male. The dactylar seta is
long and extends fully to the end of the dactylus ; at about the middle of its length
it divides into two branches, the outer one the thicker, l)otli further subdividina: into
numerous very fine hairs.
In the female the first pleopod is very like that of T. ptisillus figured by Sars, but the
* L. c. p. (51(3, pi. xxviii. fig. 18.
TERRESTRIAL ISOPODA OF NEW ZEALAND. 117
enclopotlite is larger in comparison -witli the exopodite. In the second pleopod tlio
endopodite is narrow and projects consideral^ly l)eYond the exopodite. In l)oth pairs
there is a lateral expansion of the protopodite corresponding to tlie "epipodite"
described in Ligid iiovte-zeulandicB. The remaining pleopcjda are of th<! usual form.
In the male the first two pairs of pleopoda are specially niodiflcd, as in other spcci(!S,
for sexual purposes, but they differ considerably in detail. Tlu! first pleopod, together
with the sexual appendage, is shown in figure pip} cf . The sexual appendage is soft
and membranous, spatulate in form ; the endopodit(> is narrow, subtriangular, and ends
in a very long, narrow, chitinous, styliform process whicli tajjcrs gradually to the very
acute apex. In the second pleopod (fig. ^j//;.-) the endopodite is modified into a 3-jointed
penial appendage, strong and higidy chitinised; it is of nearly the same breadth
throughout except at the extremity, where it narrows abruptly and ends acutely.
2. Trichoniscus otakensis, sp. nov. (PI. 12. fig. 2.)
Philoitgria rosea, Cfiifton, Trans. N. Z. lust. xv. |). 14i), unci p. 7?, (in part) (18<S;3).
Philijyr'iu rosea, Thomson & Cliifton, Trans. N. Z. Inst, xviii. p. 157 (in part) (1886).
Philougria rosea, Filliol, Mission de file Campbell, p. 439 (in part) (188r>).
Specific description. — Male and female differing in the shape of the body.
Female. — Body oblong oval, 2\ times as long as broad; whole dorsal surface thickly
covered with irregular, densely crowded, roughish tubercles. Cephalon Avith the lateral
lobes fairly large ; margins with two or three setae, but hardly denticulate ; front slightly
convex. Segments of mesosonie slightly separated laterally ; first four segments with
the lateral angles rounded, the last three with the postero-lateral angles recurved and
acuminate. Metasome rather less than one-fourth the length of the body ; last segment.
with its posterior margin straight and bearing three or four small setse.
Antennae rather stout ; fourth joint of j^eduucle stout; fifth joint narrowed at base
and expanding slightly distally, its inner margin with four or five distinct prominences,
from wiiicli short stout setae may arise ; outer margin straight, fringed with fine setae ;
flagellum nearly as long as the last joint of peduncle, composed of four joints. Uropoda
rather short, stouter than in T. phormiamts ; outer ramus twice as long as the base.
Male. — Much narrower than the female, the greatest breadth less than one-third the
length ; none of the legs specially modified.
Colour light brown, with markings of darker brown.
Length about 4 mm.
Habitat. — Widely distributed throughout the South Island, N. Z., in damp situations.
Memarks. — This species closely resembles the preceding one in most respects, but can
be readily distinguished from it by the tuberculatcd surface, the stouter antennae and
uropoda, and, in the male, by the narrow form of the body.
The mouth-parts, legs, and pleopoda (including those specially modified in the male)
closely resemble those of T. phormianus and do not call for special description.
I have a few specimens from Greymouth, collected by Mr. R. Helms, that I refer to
this species with some hesitation. The specimens, which appear to be all females, are
of sHghtly larger size, and have the body broader and more compact than in the typical
18*
118 DK. C. CHILTON ON THK
forms; the tuhercles l)oth on the body and on the antenu;e are particularly well marked,
and, in some specimens at any rate, the flagellum of the auteunge contains five joints.
3. Trichonisctjs Thomsoni, Chilton. (PI. 13. fig. 1-)
Philygria Thomsoni, Chilton, Trans. N. Z. Tnst. xviii. p. 159, pi. v. figs. 1-6 (1886).
Specific description. — Oblong oval, greatest breadth fully half the length, fairly
convex, surface quite smooth. Cephalon short, transverse, more than twice as broad as
long ; lateral lobes small, not visible in dorsal view, front slightly convex, a slight
transverse depression a little anterior to the eyes, and an oblique depression starting
near the median line between the eyes and extending backwards and outwards. Epiraera
largely developed, those of the first segment of mesosome produced anteriorly into
rounded lobes enclosing fully one-half of the cephalon, those of the second and third
segments with the posterioi- angles rectangular, those of the fourth to seventh segments
recurved and acuminate in progressive degree, those of the seventh i^eaching as far back
as the posterior border of the fourth segment of metasome. Metasome much narrower
than last segment of mesosome ; third, fourth and fifth segments with fairly-developed
but closely-appressed epimera ; last segment with posterior border straight and bearing
three or foiu' small setse.
Anteunre slender, fourth joint of peduncle nearly as long as the fifth and slightly
broader, all covered with fine setae ; one or two longer ones at the extremities of the
second, third and fourth joints ; flagellum about as long as the last joint of peduncle,
of at least five joints ; articulations between the more distal joints very indistinct.
Legs rather long, increasing considerably in length posteriorly, very spiny. Dactylar
seta large and well developed, dividing into two branches, each of which subdivides in
many fine hairs. Uropoda rather long, about two-thirds the length of metasome ; outer
ramus much the stouter, elongate, conical in outline ; inner three-fourths the length of
the outer, cylindrical, tapering very gradually, i)oth eadiug in a few settle.
Colour a light brown, with the greater part of the body covered with markings of a
much darker brown, sometimes nearly black ; legs with irregular alternate markings of
light and dark brown.
Length about 7 mm.
Habitat. — Widely distributed over the whole of the South Island.
Memarks. — This species can usually be recognised by the wide body with greatly
developed epimera, by the smooth, almost shining appearance of the dorsal surface, and
by the five joints in the flagellum of the antenna. In smaller specimens, however, the
epimera are not so much expanded, and the articulation in the flagellum may be A^ery
indistinct and identification is the more difficult. Though a tru^e Trichoniscus in the
mouth-parts, metasome, &c., the general outline is more suggestive of an Oniscus or
Forcellis.
The mouth-parts closely resemble those of the preceding species. The pleopoda also
are very similar, except that in the second pleopod the endopodite is more elongate in
the female, and in the male the penial appendage formed by it is of a slightly different
shape.
TEREESTEIAL LSOPODA OF NEW ZEALAND. 11!)
Genus 2. Haplophthalmus, Scliobl. (1860).
Haplophthnhims, Sars, Crustacea of Norway, ii. p. 10(i (1899).
Goiicvlc characters.—^' Body ohlont;'. moderatoly convex, sciilptuvcrl dorsally Avitli
morp or loss distinct longitudinal r-ilts, Ceplialon with tlio front triangularly produced,
tlioiigh scarcely defined fi'om the ejiistome ; lateral lobes i-ather large. Side plates of
mesosome lamellarly expanded, discontiguous. Metasorae not abruptly contracted,
epimeral plates of the two anterior segments small, those of the three succeeding ones
well developed, laminar; last segment of a similar shape to that in the two preceding
genera [Trichoiiiscus and Trichoniscoides\. Eyes very small, simple, subdorsal.
AntennuUiR and antenna? much as in Trichou/sciis. Oral ])arts likev/ise rather similar,
except that the terminal part of the maxillipedes is obscurely 5-articulate, and the
epignath simple, lanceolate. Legs short and thick, scarcely at all increasing in length
posteriorly. First pair of pleopoda in female very small and rudimentary ; those in the
male well developed, with the inner ramus strongly produced, biarticulate, terminal
joint spiniform ; inner ramus of second pair in male likewise produced, triarticulate,
last joint narrow, styliform. Uropoda with the inner ramus originating inside a broad
expansion of the basal j^ai't, and terminating, as in the genus Trichonisco'ules, in a single
slender spine." [Sars, I. c. p. 166.]
The genus is represented in New Zealand by the following species only : —
1. Haplophthalmus Helmsii, sp. nov. (Plate 12. fig. 3.)
Specific description. — Oblong-oval, about twice as long as broad ; strongly convex,
the central portion being raised somewhat abruptly above the epimeral portions;
epimera well developed and somew^iat wddely separated. Cephalon with the lateral
lobes large ; on the dorsal surface between the eyes are two rather large, rounded,
roughened tubercles ; surface in front of tliese sloping, rough and uneven : front bhmtly
triangular. All the segments of the mesosome bear at the outer border of the central
poi'tion a I'aised rounded ridge ; posteriorly this becomes more marked, and on the
seventh segment the ridges end in two well-marked tubercles projecting backwards a
little over the metasome ; on the fourth anterior segments of the mesosome there is on
each side a smaller and less-marked ridge internal to the one already described and
parallel to it. Metasome rather small, not quite one- fourth the length of the body ;
first three segments short and without epimeral expansions, fourth and fifth segments
longer and with well-developed epimera ; last segment very short, more than twice as
broad as long, posterior border sti'aiijht. Siu-face of metasome rough like tliat of the
wliole body but without distinct rido-es or tubercles.
Antennse short, not quite one-fourth the length of the body, rather stout ; fourth
segment of peduncle a little expanded, shorter than the fifth, which is narrowed at base,
all with appressed scales and a few short setae, one or two longer setae on the fifth joint ;
ilagellum as long as the fourth joint of peduncle, of three joints, ending in a pencil of
long hairs.
Legs of tlie usual character, short and rather stout, not visible in dorsal view; dactylar
1^>
DB- C. X ox THE
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C
bv R. Helm?
ro
This fami^ asrees witli tise Uk
' im ijaTiiis a wdl-
:jie imier lobe oi ii>e ik^
-iiieia of tJie ^esegid to seTiaiiii se^sneiis c»f lie T»tsiai!3e az^ quiic di<rinct
T\ ' ^ ins tir -"-"-- J" ~ " ' - - ^rS iiaT- - .
As tliis is Ti- —V r^-:- - _:.s- le : . - -evf as Li"
: r^niilT.
L Itlcjs XTv-t?TT ^xm-'v ^ nflrr. Pl2ti.e 1-3. ~^- --
be sasoe chamirteTs
asd s!lfiriT]y "•^"-~-7^.-^'«^ pTC«eess widi lais -r. ibe oas&l J Dim of the anT^nng^ ^iitiiwg
iBi-C' at- rareen iMs *Tui the otHital norgirL AnicrioT Tr.^T-^Hm straight. >3iojitiy
- -1
iirsi - - - ' ■ ' ' - - _ . ' '
^1/^
ijexi tij-ee Sr-irrDtsus "sriii - _
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-js of ia>i lirree sesments simiiir in sbsjt, iilt iar^eT. and iaiereasing in aae
:1t: aH cieai-T markid of frcan ligir siinnr-xs_ Lasn siesiiiE32i of i3ie:: : _„
TEERESTKIAL ISOPODA OF NEA\^ ZEALA]!^©, 121
quadrangular; surface convex and a little uneven, Avith depressed line parallel to hinder
margin ; posterior margin slightly convex and fitting evenly into the space between the
rather small side-plates of the fifth segment.
Eyes rather large, convex, with about 40 ocelli. .Vntennula? apparently 1-jointed
and immobile. Antennte reaching to jjosterior border of the first segment of mesosome,
last joint of peduncle jibout twice as long as the fourth and as long as the fiagellum ;
fir,st joint of flagellum strongly geniculate with the peduncle, of the same length as the
second, thii-d rather longer, fom-th very small, almost rudimentary ; the whole antennae
covered with numerous short, bluntish sette.
First pair of legs with anterior margin of the basos produced near the distal end into
a triangular process, and with a shallow groove posterior to this for the reception of the
distal portion of the limb when bent back ; second pair with similar but less marked
structure; legs scarcely increasing in length posteriorly, all very setose, the setae on the
posterior pairs larger and stouter than on the anterior pairs.
Eifth pleopoda with the exopodites .strongly chitinised, large, trapezoidal, articulated
at the postero-lateral angles, and projecting inwards and forwards so as to meet in the
median line and cover a large portion of the anterior pleopoda. Uropoda triangular,
outer side convex and with a few setae, inner margin straight, fringed with fine setae,
terminal joint small, bearing a few fine setae.
Colour. — Whitish or light yellow, with scattered black spots and usually with opaque
white or silvery spots arranged more or less closely in patches; some of the specimens
darker, especially along median line.
Length. — Al)0ut 14 mm.
Ha'ntat. — Lyall's Bay, ATellington [11. M. Luiiuj), "Wellington, under tussocks near
the beach" {(}. 31. Thomson).
Remarks. — I have some hesitation in describing this as a new species, for all the
species of the genus appear to be very similar in general appearance and to be distin-
guished chiefly by diS'erences in the front of the cephalon and the pleopoda, which are
somewhat difficult to describe accurately. It is probably not very different from
Tylos spiiiulosus, Dana, from Tierra del Fuego, but appears to be less spiny and to differ
in the antenme, for Dana describes and draws the flagellum as " 3-jointed, the first joint
but little shorter than the preceding, and the second as long as the following."
Family IV. SCYPHACID^.
Scyphacince, Dana (iu part), U. S. Explor. Exped., Crust, ii. p. "16 (1853).
Mandibles without molar tubercle, its place being taken l)y a tuft of long stiff setae or
bristles ; inner lobe of first maxilla with two plumose bristles ; maxillipedes with the
terminal joints fairly well developed, lamellar, longer than the masticatory lobe;
external male organ single.
The fiimily, for which I propose the definition just given, corresponds in part with
Dana's subfamily Scyphacina?, for he rightly oliserved that in the maxilKpedes, Scyphu.r
differs considerably from the Oniscidce, though his description that they arc 2-jointed is,
122 DK. C. CHILTOX ON THE
perhaps, a little misleading. In making it he appears to have counted the basos as
one joint and all the terminal part as the second. He did not reckon in the coxa, which
is usually more or less distinct, and he included the ischium, which is also \isually
distinct, Avith the terminal portion which generally shows indications that it is composed
of three or four joints. In cases of this kind the actual number of joints is less
important than the comparative sizes of those that are represented, though of course it
is not easy to express this in brief language.
I include under this family the genera Sci/pl/ca; Dana, ActcBcia, Dana, and Scypho-
niscus, gen. nov., all of which are represented in JS^ew Zealand. It will, I think, also
include Scyphacella, S. I. Smith, and Actoniscns, Hayer. both of which are discussed in
another part of this paper, and Philomfria manna, Chilton, which Stebbing has rightly
said cannot remain under PhUougria, probably also belongs to this family, though as yet
I have not had time to examine it sufficiently to say whether it can be referred to any
of the genera mentioned or not.
It may be worth while to point out that Kinahan, in his excellent " Analysis of
Certain Allied Geuera of Terrestrial Isopoda," published in 1857, appears to have
recognised the fact that Seypl/ax and Act(ccia probaloly formed types of separate
families, though owing to the great difference between thein in general appeai-ance, he
evidently did not think of placing them both in the same family *. The three genera
that I have included in this family all agree pretty closely in the mouth-parts and
pleopoda, and I am inclined to attach comparatively little importance to the external
form of the body.
Genus 1. Scyphax, Dana.
Scyphax, Dana, U. S. Explor. Exped., Crust, ii. p. 733 (1853).
Smjphax, Miers, Cat. New Zealand Crust, p. 101 (1876).
Sci/jjJiax, Budde-LiuKl, Isopoda Terrestria, p. .231 (1885).
Generic descr'qjtion. — Body somewhat convex, not cajjable of rolling into ix ball;
epimera moderately developed. Metasome not abruptly contracted, last segment not
much produced. Eyes large, of very many ocelli, crescent-shaped, occupying the sides
of the cephalon. Antennse with the flagellum 3- or 4-jointed. Second maxilla with
the outer margin a little angularly produced near the base. Mandibles with few
penicils behind the cutting part. Legs increasing in length posteriorly. Opercular
plates of pleopoda without any air-cavities. Uropoda exposed, inner branch arising
only slightly in front of the outer.
Remarks. — I have ventured to give a new diagnosis for this genus which was
established many years ago by Dana for the single species S. ornatns. In 1876, Miers
added a new species, S. intermedius, but this, as shown below, proves to be the same as
S. ornatus.
Another species, S. setiger, from New Caledonia, was added in 1885 by Budde-Lund,
who gave a diagnosis of the genus based mainly on external characters, and considered
* Natural History Ecvicw, iv. Proceedings of Societies, pp. 274 & 275 (1857).
TERRESTKIAL ISOPODA OF NEW ZKALAND. 123
it merely a subgenus of Oniscus. In 1874 a ge'iuis, Sc!i})hacclla, was established by
S. I. Smith*, who says: — "This genus differs from ScijiiIklv most notably in the form
of the maxillipedcis, whicli in Sci/plutx have the terminal segment broad and serrately
lobed, while in ovir genus it is elongated, tapering, and lias entire margins. In Scyphux
also the posterior pair of legs are much smaller I ban the others, and weak ; tlie last
segment of tlie abdomen is truncated at the apex, and tlie articulations b(!t\Aeeu tlic;
segments of the terminal portion of the antennte are much more complete tlian in
our species. Tlie general form iind appearance of the genera, are the same, and the
known species agree remarkably in habits " Budde-Lund t gives Smith's species,
Sci/ijltacella ureiilcola, as nearly related to Triclioiuscioi (illddns ; and Sars^;, following
Budde-Lund, refers to the genus Scijpliaccllu as coming under his family Trichoniscida^.
It appears, however, from Smith's remarks that his genus is really nearer to Scypliax
even than he thought, for of the four points of difference which he gives, two are based
on errors in Dana's description, for tlie seventh pair of legs in Sci/phax are small and
weak only in immature forms and the terminal segment is not truncate, the mistake
here having arisen from the fact that the lateral margins of the terminal segment arc;
not shown in his figure. In the other two points of difference Scijphacella certainly
does ajjproach Trichouiscas, but they are, I think, only of comparatively little
importance, and the spiny antenote and whole general appearance of Scupluicella are
more like Scyphux than any Trichonisciis that I know of. It is, moreover, e\i(lent that
Scyphacella cannot come under the Trichoniscidae as defined by Sars, for (1) the
metasome is not much narrower than the mesosonie, and (2) the eyes, instead of being
" small or wholly wanting," are large and prominent. Of course the question could be
settled at once if we knew whether the mandibh; in ^cijpJiacella has a molar tubercle
or not, and Avhether the inner lobe of the maxilla has three or two plumose bristles.
Unfortunately, no special information is given on these points, either by Smith or by
Hayer, who afterwards examined the si^ecies. But the mandibles of Scypluix are figured
by Dana, and presumably these drawings would be noted by Smith, who evidently
examined those of Scijphacella, for he says "mandibles slender," and if these had
possessed a molar tubercle he would almost certaiuly have noticed it.
Until the question can be settled by the examination of specimens, I think we are
justified in including Hcypliacella under the Scyphacidiie as nearly allied to Scyphax if
not actually identical therewith.
The genus Scyphax is represented in New Zealand only by one species, though another
is occasionally classed under it.
1. Scyphax ornatus, Dana (1853). (Plate 14. fig. 2, and Plate 15. fig. 1.)
Scyphax ornatus, Dana, U. S. Explor. Exped., Crust, ii. ]>. 7.3 1-, pi. xlviii. fig. 5 (1853).
Scyphax ornatus, Miers, Cat. N. Z. Crust, p. 101 (187()).
Scyphax iutermedius, Miers, Annals & Mag. Nat. Hist. ser. 4, xvii. p. 227 (1870); Cat. N. Z. Crust,
p. 102, pi. ii. fig. 8 (187G).
* Rep. U. 8. Fisheries, pt. i. p. /i'iT (1874).
t L. c. p. 249.
t L. e. p. 160.
SECOND SERIES.— ZOOLOGY, VOL. VIII. 19
124 -UK. C. CHILTON Oy THE
Scyphax ornatus, Thomson & Chilton, Trans. N. Z. Inst, xviii. p. 158 (1886).
Scijpha.r ornatus, Budde-Lund, Isopoda Terrestria, p. 233 (1885).
Sct/pkiM' iiitermedim, Budde-Lund, Isopoda Terrestria, p. 233 (1885).
? Pki/oxcla v'wlacca, Filhol, Mission de Pile Campbell, Crust, p. 445, pi. liv. fig. 5 (1885).
Scypluix oruiitiis, Filhol, /. (". p. 443 (1885).
Scyphax ialiirmedins, Filhol, /. c. p. 444 (1885).
Scyphax intermedius, Thomson & Chilton, Trans. N. Z. Inst, xviii. p. 158 (]88()).
Specific desc7Hplton. — Body elliptical, fairly convex, breadth about half the length,
sui-face finely gramilar, in smaller specimens sometimes rough with minute sette.
Surface of cephalon flat, depressed. Metasome not abruptly narrower than mesosome,
epimerte of third to fifth segments of moderate size, last segment triangular, much
broader than long, sides concave, extremity bluntly pointed, bearing a few short set*
and with a slight depression on its upper surface.
Eyes very large, crescent-shaped, occupying the whole lateral margins of tlie cephalon
and nearly meeting in front ; ocelli very numerous, about 150 to 200, arranged in four
longitudinal rows. Antennse about half the length of the body, spiny in small
specimens, in large ones with granulations or small tubercles in addition to the
small spines ; flagellum as long as the fifth joint of peduncle, which is considerably
longer than the fourth, consisting of three joints, the third l)eing followed l)y a minute
terminal joint ending in a tuft of short seta?, first joint longer than the second and
slightly shorter than the third. Anterior pairs of legs shorter and stouter than tlie
posterior pairs, which are rather long, the seventh pair not fully developed till animal is
nearly adult. Uropoda with the base large, extending a little beyond the extremity
of the terminal segment, lateral border with a distinct keel, rami rather narrow,
cylindrical, spinose, the inner one very slightly shorter than tlie outer and not arising
much in front of it.
Colour variegated, irregularly spotted with yellowish red, grey, brownish red or black.
Length of largest specimens about 18 mm.
Habitat. — On sandy shores in the North Island and also from "VVestport. Not found
in the south of South Island.
Bemarks. — This species was described and figured by Dana in 1853, but does not
appear to have been recognised since. Miers descril)ed his Scyphax intermedius as a
separate species in 1876, being misled by Dana's figure of the whole animal wdiere the
margins of the terminal segment of the metasome are not marked and the segment
consequently appears much more broadly truncate than it really is. I have seen
Miers's type specimen in the British Museum and have no doubt that it is only a large
specimen of *S'. oniafus, Dana. Although Eilhol's description and figures are not
altogether satisfactory, I have little doubt that his I'liiloscia ciolacea also belongs to
this sj)ecies ; the large crescentic eyes clearly show that the species cannot be placed
under Fhiloscia. In 1885 Budde-Lund described a species, Scyphax sctiger, from New
Caledonia which probably will come near to S. ornatus, Dana, though the eyes seem to
contain fewer ocelli and the proportions of the joints of the tlagellum of the antennae
are difi^erent.
Although Dana was undoubtedly dealing with immature specimens when he described
TERKESTKIAL ISOrODA OF XEAV ZEALAND. 125
tlio seventh pair of logs as "much smaller than tlie others, weak," it is nevertheless true
that ill tliis species the development of these legs appears to he delayed longer than is
usually tlie case. In specimens of from 4 to 't nun. in length, which are running
actively on tlie heacli and not otherwise immature, the seventh segment of the mesosomc
is small and tlie seventh ])air of legs repi-esented either hy a small hud or hy a weak,
non-chitinised appendage, m ith the joints only faintly indicated and surface free from
seta?; in specimens a little larger ((i mm.) the seventh segment is more developed, hut
still smaller than the sixth, and the legs are of the usual shape hut smaller than the
sixth and less ahundantly supplied Avith setae. In specimens of 9 mm. in length I found
the seventh segment and appendages fully developed ; the male organs were also present,
and the specimens apparently fully adult.
Most of the more important points in the appendages of this species have heen
referred to in the discussion of the genus already given. I gi\e here a i'cw additional
notes.
The maudihles are of the type usual in the family. The outer cutting-edge in the
right contains three or four stout teeth, brown in colour and higlily chitinised ; the inner
cutting-edge is more transparent, slender, and ends in two large teeth and one or two
smaller ones ; it is followed by a membranous lappet, the sides and margin of which
are densely setose; between this and the dense tuft of stilF plumose bristles is a single
large plumose seta. The left mandible is very similar, but the inner cutting-edge is
much larger and stouter, and ends in three large teeth which are brown in colour and
as strongly chitinised as those of the outer cutting-edge, and there are two plumose
setre between the membranous lappjet and tlie tuft of seta? representing the molar
tubercle.
Tiie first maxilla is of the usual form; in the second the extei'nal lobe at the end is
very small, and the outer margin shows a proininenc!' near the base like that drawn and
described by Sars in Oiiiscus and some allied genera.
In the maxillipede the exopodite is about half as long as the bases, oblong with the
end rounded ; the outer margin of the basos is somewhat expanded, and is fringed with
line setfe towards the distal end ; the masticatory lobe is about half as long as the
terminal j^ortion of the nraxillijiede, and lias the end obliquely truncate and fringed with
setae ; the ischium is short, distinctly separated from, adjacent points ; on the outer aspect
of both the basos and ischium are three or four short sette near the distal margin ; the
four terminal joints are coalesced into a single laniclliforra plate, with four distinct lobes
on inner side representing the different joints of which it is composed.
The legs of the mesosome are sjjiny, the anterior pairs shorter and stouter than the
posterior; the dactylus is long, and has its basal jiart thickly covered with short setae,
the terminal claw long, narrow and slightly curved, secondary claw very narrow, almost
like an ordinary seta. There is no special " dactylar seta."
In the male the first pleopod is remarkable in having the exopodite very large and
operculiform ; it is articulated as usual to the lateral part of the prolopodite, and extends
anteriorly and posteriorly into two large rounded lobes, which show branching thiokea-
19*
120 DK. C. CHILTON ON T]1E
ing's apparently intended to strengthen the large flat plate thus formed ; the endopodite
consists of a single long slender appendage, gradually narrowing to the end ; the
epipodite is formed of an oblong lobe with, rounded end. The external male organ is
short, rather narrow and rounded at the end.
The second pleopod of the male has tlie exopodite of fair size, subquadrate with angles
rounded off, and of similar structure to that of the first pleopod ; the endopodite is
represented by a 2-jointed appendage, the first joint extending directly inwards like a
prolongation of the base, the second at right angles to this, as long as the exopodite,
gradually tapering to an acute apex.
The succeeding pleopoda have the exopodite much larger than the endopodite, and
ajjparently mainly ojiercular ; the endopodite is subtriangular, with the inner portion
thickened, and is branchial in function.
The uropoda have been already sufficiently described.
2. ScYPHAX (?) AUCKLANDiiE, G. M. Thomson. (Plate 15. fig. 2.)
Acttecia aucklandue, G. M. Thomson, Trans. N. Z. Inst. xi. p. .349 (ISTQ).
Ackecia cnit-kl(indi<i\ Bndde-Lund, Isopoda Tcrre.stria, p. 239 (1885).
Acttpcia avcklanclice, Filhol, Mission de I'lle Campbell, p. 443 (1S85).
? Oniscus nova-zealandixe, Filhol, /. c. p. 441, PI. liv. fig. 7 (1885).
Actacia aucklandue^ Thomson & Chilton, Trans. N. Z. Inst, xviii. p. 158 (188(5).
Specific description. — Body oblong-oval, length about twice the greatest breadth, sides
of mesosome j^arallel. Cephalon with lateral lobes large and broad ; front triangular,
depressed, somewhat excavate in the middle (in dorsal vein appearing neaily straight) ;
an oblique ridge on each side running backwards and outwards behind the eyes from
near the centre in front to the posterior margin of the cephalon ; surface between these
ridges roughly tubercular. Side-2)lates of mesosome well develojoed, especially those of
the first segment ; posterior margin of first segment slightly sinuous, posterior angles
subacute, posterior angles of succeeding segments becoming more and more acute.
On each segment is a small oblique ridge on the side-plate running backwai'ds and
outwards, and betAveen these ridges eacli segment bears numerous irregular tubercles,
some rounded, others more acute ; on the three anterior segments they are rather less
marked and irregularly arranged ; on the four posterior segments they form a regular
row of somcAvhat pointed tubercles projecting a little backward along the posterior
margin of the segment, with irregular granulations anterior to this row.
Metasome not abruptly narrower than mesosome, first two segments short, their sides
overlapped by the last segment of mesosome ; side-plates of third to fifth segments large,
produced acutely backwards ; surface of each of these segments with a transverse row of
small tubercles, and with others irregularly distributed ; terminal segment short, much
broader than long, triangular, sides concave, extremity rounded and depressed, fitting
closely on to the bases of the uropoda.
Eyes of moderate size, situated on a slight prominence above the lateral lobes, and
external to the oblique ridges. Antennules easily visible. Antennte short, reaching
backwards to the posterior border of the second segment of mesosome ; first three joints
short, subequal ; fourth longer, somewhat broadened ; fifth a little longer than the
TERRESTEIAL LSOPODA OF \EW ZEALAND. 127
fourth, slightly sinuous, narrowed at base ; flagellum as long as the fourth joint, stout,
comjiosed of four joints, the first two short, suhoqual, third longer, fourth very short,
articulations not very distinct ; whole antenna roughly granular, with very few setaj.
Legs rather short, not visible in dorsal view, and not increasing in length posteriorly.
Pleopoda apparently as in Ormciis, opercular plates with outer margin incurved but
not so abruptly notched as in Oiiisciis; no air-cavities. Uropoda with the base large
and meeting in the median line, lateral portion expanded, flattened and keeled externally,
ending acutely external to the outer ramus ; both rami exposed, inner ramus arising a
little anteriorly to the outer, but extending backwards to the same point, but with apex
rounded and with a very few minute setae.
Colour brown.
Length 20 mm., breadth 9 mm.
Habitat. — Auckland Island {Mr. Jeniiiricj.s).
RenKo-kx. — The above description is taken from a female, the only specimen I have
seen. According to Mr. Thomson the male has the whole surface of tlie body nearly
smootb.
I refer Pilhol's Oniscus novfe-zeuIaudicB to this species with considerable doulst, for
his description and figui"e are hardly sufficient to permit of certain identification.
However, some points in his description as to the antennae and uropoda, and especially
that of the tubercles — " les granulations de la rangec posterieure sont plus detachees et
leui'sommet \m pen aigu est dirige en arriere " — apply exactly to the species in question.
He states that his specimens were obtained near Wellington.
I have jilaced the species under ScijpJia.r only provisionally, for I have had only one
specimen and have not been able to examine all the mouth-parts, &c. It can hardly
come under Actcscia, in which it was placed by Mr. Thomson, and though it has con-
siderable resemblance to Oniscus, it differs markedly from that genus in the antenna}
and uropoda, and also in the maxillipedes, for these, as shown in fig. 2 m.ip., have the
terminal part well developed, much longer than the masticatory lobe, and with clear
indications of the joints of which it is composed. In this, and in the maxilhe which I
have also been able to examine, the species reseuables Scyphax, and I think it will
certainly come under the same family, but ifc differs greatly from -Scj/jV/ca' in the cephalon
and in the much smaller eyes. It prol)ably lives on the sea-shore like the rest of the
Scyphacida-.
Genus 2. Scyphoniscus (novum).
Generic Characters. — Body rather narrow, lateral parts not greatly developed.
Cephalon with large broad lateral lobes. Metasome abruptly contracted, first tuo
segments rather short, third to fifth with small epimera, last segment short, triangular.
Eyes of moderate size, sublateral. Antennulae of three joints, the last small, but bearing
two or three sensory setse. Antennae with the flagellum of three ill-defined joints.
Mandibles with a membranous hairy lappet behind the two dentate lamellaj, followed by
a long recurved brush-like seta ; molar process represented by a dense tuft of recurved
setae of unequal lengtli. Anterior maxilla; with the outer lobe rather weak, some dentate
128 DR. C. CHILTOX ON THE
sette on its inner margin near the apex, end witlv a dense row of simple hair-like setiie ;
inner lobe small, delicate, with tAvo hairy bristles. Second maxillae distinctly bilobed at
the extremity, outer edge not angularly produced near the base. Maxillipedes with the
basos rather narrow; masticatory lobe well developed, rounded at end, terminal portion
of fair size with the last four joints coalesced into a single triangular plate. Legs short,
not increasing much in length posteriorly. Pleopoda simple, opercular plates delicate,
and none of them with air-cavities. UropD'Ja rather jiroduced, base only i^artially
covered by terminal segment, rami not flattened.
Jiemarka. — In general appearance this genus at first recalls Trichoniscus. The most
important character by which it is separated from other genera appears to be the structure
of the first maxillse, the outer lobe of which is very peculiar, and quite different from
that of any other Terrestrial Isopod that I am acquainted Avith.* In the mandibles with
the den'?e tuft of stiff setae apparently representing the molar tubercle and in the
maxillipedes this genus seems to resemble somewhat closely Sci/pJuu- ornalus, and the
two species of Actcecia described in this paper. Tiie type species, which is the only one
at present known, was found on the sea-beach within reach of high tides, and I at first
thought that the Philougria marina described by me in 1SS5 f would belong to the same
genus, but I find that it has the first maxillae normal, though, as Stebbing has recently
j)ointed out, the species cannot for other reasons come under Trichoniscus:^, and probably
will be found to belong to the ScyphacidEe.
ScYPHOXiscrs AVAiTATEXSis, sp. nov. (Plate 14. fig. 1.)
Specific description. — Body narrow oval, rather more than twice as long as broad ;
surface of cephalon and mesosome Avith small rounded tubercles and other irregulaiities,
generally forming a somev\hat irregular transverse row near the hinder edge of each
segment. Cephalon with the broad lateral lobes not very clearly defined at their bases,
front triangularly produced. Posterior margins of the first four segments of mesosome
straight, those of last three with tlie posterior angles more and more recurved posteriorly.
Metasome with the first segment short, second longer, third to fifth with moderate
appressed epimera ; last segment short, about twice as broad as long, triangular, ending
in a slightly rounded angle, sides concave.
Antennae with the fifth joint of paduncle much longer than tlie fourth, flagellum as long
as the fourth joint, of three joints, the first short, second twice as long, third narrow,
about as long as the first, Avhole antennae covered with short stout setie. Legs short,
dactyla stout, with the secondary nail small or obsolete ; dactylar seta arising near the base
of dactylus, longer tlian dactylus, 2-branched, the outer the stouter and plumose,
inner one apparently simple.
Colour brown.
* Dollfus }ias flrawii attention to some abnormalities in the mouth-parts of McsaniHiiliUo Alluandi, Dollfus,
the most striking being that the inner lobe of the fir.st maxilke bears nine hairy bristles, instead of the two usually
])rescnt in Armnill/Udi,um (Annales de la Socic'te Entom. de France (1892), l.Ki. p. 38C).
t I'roe. Linn. Soe. N.S.W., ix. part ,S, p. 463.
+ Troc. /ool. .Sof. London, ] 900, j). 5G5.
TEEEESTEIAL ISOPODA OF Ni:W ZEALAND. 129
JLeiigth of largest specimen seen o"5 mm.
Ilahilat. — Blueskiii Bay, Otago, undei' seaweed, &c. at high-water mark.
Hcnuo-lH. — Of this species I have only a few small specimens collected at Bluoskin
Bay about 1887. I have never found it since, tliongli I Iiave frequently searched the
same locality and otlier similar situations, and I am unal)le to say whether it attains a
larger size or not. In order that its affinities may be made as clear as possible, I have
figured the moiitli-parts in some detail, and add further description of some of its
appendages.
The upper li[) is of tlie usual structure.
The loj't mandible has the outer cutting-edge formed of three well-marked teeth
strongly chitinized, the inner cutting-edge similar and of four teeth ; near its base
arises a hairy niemljranaceous lappet with the innermost seta longer than the others •
next, at a considerable distance, follows a single long-curved, feathered seta ; and ne\t
from a slight prominence, a brush-like tuft of stitf simple setie, the; outer ones the
shortest, the others gradually increasing in lengtii, the innermost ones bein^ very Ion"-.
The right mandible is similar, but the outer cutting-edge appears to Ijear four tc(;!li
and the inner cutting-edge is smalle)", less ehitinous, and ends in three or four small
sharp teeth, apparently arranged in a circle roiuid the extremity.
Lower lip formed of two rounded lobes with numerous sette directed inwards and
between them a narrow tongue-like process with its margins thickly fringed with setic.
Theji/'fil m((xiUce has the outer lobe oblong in shape, its inner margin friuo-ed in distal
half with tine setiu, and towards the end bearing aljout ten dentate sette of various sizes
and irregularly arranged ; the extremity and a little of the outer margin bears a compact
row of long, simple, hair-like setie, the outer ones being the longest, the inner ones
stouter and a little incurved. Inner lobe of normal form, the distal plumose bristle
much shorter than the other.
The second maxilla; form a delicate oblong plate, distinctly cleft at the aiiex, out(U'
lobe about half as large as the inner, both roumled and bearing delicate setie, the inner
one with a row of short stouter sette also.
Tlie nuu-illipedes have the epipodite narrow ol)loug, as long as the basos, truncate at
end, end and distal half of outer margin with delicate setie ; basos narrow, its outer
margin not expanded ; masticatory loljo large, more than half the length of the palp ;
palp with the ischium distinct, but the remaining joints combined into a triangular
plate witli its inner margin thickly fringed with setie, and with a faint indication of
lobes indicating the separate joints.
The first and seventh legs are shown in the plate, and do not require further descrip-
tion ; the dactylar seta is large and well marked ; its general ajjpearance will be best
learnt from the figure.
The pleopoda appear to be all of about the same structure ; in the first the endopodito
is of nearly the same size as the exopodite, but narrower; tliere is a small "epipodite ''
arising from the basal jiortion. In the third and succeeding pleopoda the eudopodite
arises considerably more proximally than the exopodite, and is smaller in proportion.
I have not been abb; to examine the pleopoda in the nuile. The uropoda have already
been sufficiently described.
130 DR. C. CHILTON ON THE
Genus 3. Act^cia, Dana, 1853.
AcUpcia, Dana, U. S. Explor. Exped., Crust, ii. p. 7134 (185.3).
Actmcta, G. M. Thomson, Trans. N. Z. Inst. xi. p. 249 (1879).
Cylloma, Budde-Lund, Isopoda Terrestria, p. j6 (188;j).
Actmcia, G. M. Thomson, Proc. Roj'. Soc. Tasmania, 1892, ji. 12 (st'j)arate copy).
Generic description. — Body convex, capable of rolling into a ball, surface spiny.
Metasomc not abruptly contracted, terminal segment very short. Elagellum of anteanf«
4-jointed. Eyes very large and promiDcnt, on oval elevations along the sides of the
head. Maxillipedes with the terminal portion large, lamellar. Legs rather short, not
increasing much in length posteriorly. None of the opercular plates of the pleopoda
with air-cavities. Uropoda short, not projecting much beyond the outline of the body ;
base broad and flattened, outer portion produced, outer ramus short, inserted at the end
of the base near the inner margin ; inner ramus slender.
Remarks. — I propose to retain the genus Actcecla, Dana, for the following species, and
give for it the ahove diagnosis. In many respects it appears to resembie Arnuul'dloimcus,
Ulianin, with which Budde-Lund thought it to be identical, but that genus differs
considerably in the form of the head and in possession of air-cavities in the first two
pleopoda, and as it presumably belongs to the Oniscidce, it probably differs also in the
terminal portion of the maxillipedes, but on this point I can get no definite information.
Cijlloina, Budde-Lund, agrees so well with the genus in question in the eyes, and
terminal uropoda and general shape, that T think it must be the same, though Budde-
Lund describes it as having air-cavities in all the opercular plates of the pleopoda ; this,
however, perhaps arises from the fact that he had only a single badly-preserved
(probably dried) specimen, and natui-ally thought that it belonged to the ArmadilUdce
and came near to Armadillo.
Hayer's genus Actoiiiscus is, as he points out, nearly related to Actcecia, and tlie
difference that he gives in the antennae is more apparent than real, for Actcecia has only
four distinct joints in the iiagellum, and some of the other differences are of subsidiary
importance. On the other hand, the cephalon in his species is more Hke that of Arma-
dilloniscus, but whether it also resembles that genus in the possession of air-cavities in
the first two pleopoda or not, I cannot say. In general ajspearance it certainly appears
to resemble Acta'cia much in the same way as Sci/phacella resembles Scyphax.
The two New Zealand species of Actcecia may be distinguished thus : —
Eggs large, crescentic, outer branch of uropoda large, dilated distally A. euchroa.
Eyes moderate, outer branch of uropoda minute, acute at apex A. opitrensis.
1. AcTiECiA EUCHROA, Dana. (Plate 15. fig. 3.)
Actcecia euchroa, Dana, U. S. Exploring Exped., Crust, ii. p. 734, Plate 48, fig. 6 (1853).
Actcecia euchroa, Miers, Cat. N. Z. Crust, p. 101 (1876).
Actcecia euchroa, G. M. Thomson, Trans. N. Z. Inst xi. p. 249 (1879).
Armadilloniscus euchroa, Budde-Lund, Isopoda Terrestria, p. 239 (1885).
Actcecia euchroa, Filhol, Mission de Pile Campbell, p. 443 (1885).
Actcecia euchroa, Thomson & Chilton, Trans. N. Z. Inst, xviii. p. 158 (1886).
Actcecia euchroa, G. M. Thomson, Proc. Roy. Soc. Tasmania, 1892, p. 12, Plate ii. figs. 1-8 (1892).
TEEEESTKIAL ISOPODA OF NEW ZEALA.Ni). 181
Specific characters. — Body convex, surface with short, l)lLiiit, scattered spines, especially
on the margins of the metasomc and on the appendages. Cejihalon witli the front
rounded, with raised frontal margin, a little depressed in middle, a deep groove on each
side interior to the prominences I)earing the eyes. Inferior mai-gin of first segment of
niesosome thickened, not incised. Posterior angles of the first four segments of mesosome
sidjquadrate, those of tiie last three produci^d backwards, subacute. Epimeral positions of
third to fifth segments of metasome large and contiguous, and containing the outline of
the metasome. Terminal segment very short, convex, slightly pointed between the bases
of the uropoda. Eyes very large, on two crescentic prominences, occupying whole lateral
border of the cephalon ; ocelli very numerous, arranged in four or five longitudinal rows.
Antenna; with all the joints covered Avith short blunt spines, fiagellum about as long as
preceding joint ; its terminal fourth joint minute. Uropoda projecting beyond the
terminal segment, and pretty accurately filling up the space between tlie epimera of the
fifth segment ; outer part of base produced into a rounded lobe, outer branch inserted
on the inner portion of the distal margin, spatulate, extending a little beyond tbe
produced portion of the base ; inner rami, arising from the under surface of the base,
far in front of the outer, slender, scabrous ; apex uith a long bristle, which is visible in
dorsal view.
Colour light grey, with irregular black markicgs; during life sometimes coloured
as in Hcypliux oniatus.
Length about 10 mm.
Habitat. — Sandy beaches in New Zealand.
BemarLs. — This species is frequently found on sandy beaches in company with
IScyphax urnatiis, and is very similar to tliat species in colour and habits, so tliat Dana
thought it was perhaps the young of Scyjihaa;. From what has been already said, it is
clear that this is not the case, and that the two are perfectly independent ; I h ive
mature males of both species. When pursued, Actcecia euchrou rolls itself into a ball,
and is then tilmost indistinguishable from a grain of speckled sand ; under similar
cii'cumstances Hcypliax oniatus crouches down closely on the sand, and is then equallv
diificult to peicei\e. I liave always found these two species on the open sands and
never under cover, and the large and well-developed eyes that they both possess have
probably been developed in connection with their exposed mode of life.
The extremities have the usual three joints, though the articulations are not always
easy to see ; the last joint is tapering and bears a few setae. The mouth-parts show a
close general resemblance to tliose of Scyphax, the mandibles and first maxiUaj beiu"'
very similar; the appendage figured by Thomson as the inner lobe of this maxiUa is
really the second maxilla, which has the outer margin angularly produced near the base,
and the outer lobe at the apex very small and indistinct. In the maxillipedes the
ischium is distinct, but the following joints are all fused into a triangular plate with
lobes on the inner margin indicating the separate joints ; the masticatory lobe is small
and obliquely truncate, about half as long as the terminal portion of the maxillipede.
The legs have the dactyla short and stumpy-looking; the dactylar seta with distal half
thickened and presenting a stippled appearance.
SECOND SERIES. — ZOOLOGY, VOL. VIII. 20
132 DK. C. CHILTON ON THE
The first pleopoda of the male are shown in fig. 3, pip} 6 ; the male organ is single,
broader in basal half, end with small notch ; endopodite long, broad at base, apex curving
a little outwards, exopodite small and oval. The endopodite of the second pleopod
2-jointed, the second forming a very long, acute process. The other pleopoda present
no special features. None of them possess air-cavities so far as I can make out.
2. AcT^ciA OPiHENSis, sp. nov. (PI. 15. fig. i & PI. 16. fig. 1.)
Specific description. — Body very convex, rather narrow, more than twice as long as
broad. Surface fairly smooth, but with nurnerous scattered short setoe, especially on the
metasome and on the margins of the mesosome. Cejihalon with the front broad, straight,
with a slio'htlv-raised transverse ridcre. Posterior margins of segments of mesosome
nearly straight, those of the last two a little produced backwards at the lateral angles ;
inferior margin of fii-st segment thickened. Metasome convex; side portions of third
to fifth segments large and recurved ; terminal segment short, much broader than long,
its posterior mai'gin regularly convex. Eyes of moderate size, round. Antennae very
setose ; second joint of peduncle longer than the third and nearly equal to the fourth,
fifth about as long as fourth; flagellum as long as the second joint, of four joints, the
first longest, second and third subequal, fourth very small. Legs setose, scarcely
increfising in length posteriorly ; dactylar seta large and well-marked, simple, its distal
two-thirds thickened and apparently resembling a narrow circular brush. Uropoda with
basal joint very large, extending beyond terminal joint, exj)anded and plate-like laterally;
outer margm subcrenate and bearing four or five stout seta? ; outer branch small, conical,
scarcely projecting, tipped with a few small seta3 and one or two longer ones ; inner
branch reacliing a little beyond end of terminal segment, scabrous, and ending in two
long setae.
Colovr yellowish, with numerous black stellate markings, some specimens nearly
black.
Size. — Length about 6 mm.
Habitat. — Timaru, luider seaweed at hiiih-Avater mark.
Bemarks. — I have placed this species under Actcccia with considerable hesitation, for
it dilfers from the preceding species very markedly in the structure of the eyes. In ex-
ternal appearance it is rather like Tylos, but the mouth-parts are of course very different
and are in fairly close general agreement with those of Actcecia euehroa. The terminal
portion of the maxillipede is less lobed, and the masticatory lobe has a small terminal
lash very like that in some species of Trichoniscus.
The pleopoda in the female are of usual form and apparently all similar ; in each the
exopodite is slightly larger than the endopodite and overlaps about half the succeeding
one. In the male the first pleopoda are rather short and very strong, the endopodites in
close apposition and apparently coalesced along the median line, the ends curving
outwards ; the second pleopod with basal portion of endopodite long, end joint not very
acute.
TEREESTRIAL ISOPODA OF NEW ZEALAND. 133
Family V. ONISCID.E.
In this family the molar tubercle oi' the maiulible is replaced by a tuft of stilf setie,
the inner lobe of the first maxilla bears only two plumose bristles ; the terminal portion
of the maxillipcdes is short, scarcely longer than the masticatory lobe. The uropoda
are more or less exposed, and the inner ramus arises anteriorly to the outer.
The family contjiins numerous genera, of Avhich four are rej»resented in NeAV Zealand.
Genus 1. Uniscus, Linne (1767).
Ouiscun, Liuuicus, Syst. Nat. ii. p. 1061 (17(»7).
Oniscus, Bute & Westwood, Brit. Sess.-eyed Crust, ii. p. 166 (1808).
Uniscus, Budde-Luud, Isopoda Terrestria, p. 202 (1885).
Oniscus, Sars, Crustacea of Norway, ii. p. 170 (1899).
Generic dcucription. — "Body broad and depressed, with, the lateral parts of the
segments lamellarly expanded. Cephalon with well-defined lateral lobes, front im-
perfectly defined from the epistome. Metasome not abruptly contracted, last segment
considerably produced. Eyes large, sublateral. Antennula? wdth the terminal joint well
developed. Antenuse slender and elongated, with the flagellum comjiosed of three
articulations. Mandibles with numerous penicils behind the cutting-part. Legs
moderately slender, gradually increasing posteriorly. Opercular plates of pleopoda
without any air-cavities, those of the two anterior pairs deeply bilobed. Uropoda rather
produced, with the inner ramus originating far in front of the outer." (Sars.)
Although there are uo actual air-cavities in the opercu.lar plates of the first and second
pleopoda in Oniscus, Stoller * has recently shown that in the outer portion of these plates
there is a structure which performs the same function of allowing the animal to breathe
ordinary dry air.
1. Oniscus puxctatus, G. M. Thomson (1879). (PL 16. hg. 2.)
07iiscus jjuuctattis, (j. M. Thomsou, Traus. N. Z. lust. xi. p. 232, pi. x a. fig. 3 (1879).
Oniscus punctutus, Budde-Luud, Isopoda Terre.stria, p. 200 (1885).
Oniscus jnmctatvs, Filhol, Missiou de I'ile Campbell, p. 440 (1885).
Oniscus jjUHctutus, Thomsou & Chilton, Traus. N. Z. lust, xviii. p. 158 (1880),
Specific description. — Body oblong-oval, rather more than twice as long as broad,
rather convex, whole surface covered witii short setae which in dried specimens give the
appearance of small scale-like markings. Cephalon with the front depressed, produced
slightly into an obtuse lobe ; lateral lobes small, ending subacutely. Mesosome with
the posterior margins of the first three segments straight and their posterior angles
rectangular ; lateral angles of last foiu- segments produced more and more backwards,
acute. Epimera of third to fifth segments of metasome well developed, narrow, ending
acutely ; terminal segment not much produced, triangular, much broader than long, the
rounded apex reaching as far as the end of the base of the uropoda and bearing a few
minute setae which scarcely project beyond its margin.
* Zoologioa. llcft XXV. (^ks9y).
20*
134 15R. C. CHILTON ON THE
Eyes of moderate size, with fifteen to twenty ocelli. Antennae as long as the head and
first three segments of mesosome minutely setose throughout; second and third joint
suhequal, fourth longer, fifth twice as long as the fourth and longer than the flagellum ;
joints of flagellum increasing in length distally, and the third followed hy a styliform
process or hristle, fully as long as the first joint and dividing at the end into a compact
pencil of setae. Legs fairly long and very spinous, inci-easing considerahly in length
posteriorly. Eirst two pairs of pleopoda with the opercular plates partially l)ilohed.
Uropoda wdth the outer joint much longer and stouter than the inner, which arises only
a little anterior to it and reaches to the middle of the outer, both setose and bearing one
or two longer setae at the end.
Colour brown, with wavy stripes of white on each side the median line and often
with two lateral rows of whitish patches, the number and size of the white markings
varying greatly.
Lemjth about 10 mm.
Habitat. — Very common throughout the whole of New Zealand.
Remarks. — I have had some little hesitation in referring it to the genus Oniscus as
now restricted, for it differs distinctly from the definitions of that genus as given both
by Budde-Lund and by Sars in that the mandibles do not bear so many " penicils "
behind the cutting-part ; I find only one on each mandible, though another one or
sometimes two are situated on the setose membi-anaccous lappet just internal to the
cutting-edge. According to Budde-Lund Oniscus should have four or five penicils.
Moreover, the anterior segments of the mesosome have the posterior margin straight
instead of deeply sinu.ate, and the last segment is broader and less produced than is usual
in Oniscus. Budde-Lund (p. 206) suggests that the species under consideration may
lielong to Philoscia, and it certainly agrees with that genus in the mandibles and in the
posterior margins of the segments of the mesosome, but it differs from that genus in
possessing well-marked though small lateral lobes on the cephalon, and in having the
epimera of the mesosome of fair size, and those of the third to fifth segments of metasome
well marked. On the whole I prefer to leave the species under Oniscus, for though
allied genera are plentiful enough I cannot at present find one that will suit it better.
The species is widely distributed throughout New Zealand, and presents considerable
variation in colour, breadth, and compactness of the body prominence of the front and
lateral lobes, &c. I have some specimens collected by Mr. W. W. Smith on limestone
rocks at Albuy, Canterbui'y, which in colour resemble the variety marnwratus of
Porcellio scaber, the general surface being very light yellow, marked with somevrhat
sparsely-scattered black dots and markings ; the small spines on the surface are more
marked than in typical specimens, the front is less produced, and the lateral lobes of the
cephalon are very small and inconspicuous, while the joints of the flagellum of the
antenna? are more equal in length. In other respects they resemble Oniscus punctatus
so closely that I prefer, for the present at any rate, to consider them merely as a variety
of that species, for which I propose the name marmoratus.
The specimens from Mount Wellington, Tasmania, referred to this species by
TERRESTRIAL ISOPODA OF NEW ZEALAND. 13')
Mr. Thomson *, differ in having the side-plates of the third to fifth segments of the
metasome much smaller, and thus approach still more closely to Fhiloscia, under wliicli
they sliould perhaps be placed.
2. OXISCUS KENEPTJRENSIS, Sp. UOV. (PI. 16. fig. 3.)
Specific (Jescrqitioii. — Body regularly ohlong-oval, broad, the length rather less tlian
twice the greatest breadth ; dorsal face but sliglitly convex, finely granular, not setose,
each segment bearuig on each side of the median line a sliglitly raised and wrinkled
patch, most marked on anterior segments. Cephalon small, deeply sunk into first
segment of pereion, frontal edge regularly convex ; lateral lobes very small, subacute at
apex. Side-plates of mesosome greatly prominent, lamellar, contiguous, projecting
almost laterally, those of the first segment extending forwards into subacute lolies, which
reach to the level of the anterior margin of the eyes ; posterior angle broadly rounded ;
posterior margins of the second and tliird segments slightly sinuous ; posterior angles
rectangular, slightly rounded. Posterior angles of the fourth to seventh segments progres-
sively produced more and more backAvard and ending acutely ; those of the seventh reach
as far posteriorly as the end of the epimcral portion of the third segment of metasome.
First two segments of metasome of fair length, but wholly embraced by the preceding
segment; epiraeral plates of the next three segments produced and recurved, ending
acutely, the last reaching slightly beyond the end of the terminal segment. Terminal
segment triangular, broader than long, sides slightly concave, end rounded and reaching
as far as the end of the base of the uropods, posterior portion slightly depressed and
concave.
Eyes of moderate size, about fifteen ocelli. Antennae very similar to those of Onisctis
pnnctatus, scabrous, the minute setae being less prominent than in tliat species. Legs
long, spinose, the posterior pairs somewhat elongated. Opercular plates of pleopoda not
bilobed. Uropoda wdth basal joint large ; outer ramus broad at base, tapering regularly
and equally on both sides to an acute point, scarcely setose, inner ramus reaching to the
middle of the outer, minixtely setose.
Colour. — Slate-coloured, with white markings on the wrinkled patches on the mesosome
and usually with a lateral roAV of white patches at junction of epimera with the central
portion.
Lenyth about 11 mm., breadth 6'5 ram.
Habitat. — Kenepuru [J. 3IcMahoi)).
Hemarks. — In fully-grown specimens this species can be readily distinguished fi"om
Oniscus punctattis by the flattened body, the greatly-developed epimera, and the
wrinklings on the dorsal surface ; in younger forms all of these points are less marked,
but the species can then be usually recognized by the large and acutely-pointed outer
rami of the uropods.
* Proc. Roy. Soc. Tasmania, ISiti!, p. 10 (separate copy).
136 l>li- C. CHILTOK ON THE
3. Oniscts Cookii, Fiiliol.
OniscuH Cookii, Filhol; Mission de File Campbell, 1885, p. 442, pi. 54. fig. 6.
The following is the description given of tliis species by Eilbol : —
" J'ai rccueilli cette espece sous les pierres siir la portion ouest cle I'ile du milieu de la
Nouvellc-Zelande. Elle ne mesure que 0™"008 de longueur et 0""004 de largeur. Le
corps est ovalaire et remarquablement bombe ; la tete est large, les antennes externes
sont tres fines et leur cinquieme article plus developpe a la longueur du liagellum. II
n'existe pas de polls ni sur les articles basilaires des antennes externes, ni sur le flagellutn.
La base des articles composant les antennes est brune, alors que le sommet est blanc.
Les anneaux du thorax sont assez developpes d'avant en arriere. lis sont converts, en
grande partie, de tres fines granulations d'une teinte noire. Les granulations font defaut
en difi^erents points des anneaux et, la oil elles nianquent, on oliserve des surfaces un
pen creusees, d'une teinte noisette. Ces surfaces denuees de granulations, et apparaissant
en creux a cause des saillies que font les granulations qui les limitent, sont de formes tres
variables. Tantot elles sont arrondies, tantot elles se divisent et figurant de grossieres
arborisations. Sur les anneaux de I'abdomeu on retrouve ces plaques, mais elles sont
alors granuleuses, comme le reste des anneaux qui les presentent. Les stylets externes
sont les plus developpes et leur bord externe est garni, ainsi que celui des stylets internes,
de soies tres-fines, courtes et tres serrees."
I cannot recognize this species unless, indeed, it be Outsells kenepurensis.
Genus 2. Philoscia, Latreille (18(M).
Fhi/oscia, Latreille, Hist, des Crust. &c. t. 7, p. 43 (1801).
Philoscia, Bute & Westwood, Brit. Sess.-eyed Crust, ii. p. 448 (18G8).
Philoscia, Budde-Lund, Isopudu Terrestria, p. 207 (1885).
Philoscia, Sars, Crustaeea of Norway, ii. p. 172 (189'J).
Generic Characters. — Body oval, sliglitJy convex, with rather thin integuments.
Cephalon rounded in front, without any projecting lateral lobes. Side-plates of niesosome
but slightly protnineut. Metasome abruptly contracted, with the epimeral plates small
and appressed ; last segment not mucu produced. Eyes well developed, lateral.
Antenuai very slender, with the fiagellum composed of three articulations. Mandibles
with only a single penicil behind the cutting-part. Legs very slender and greatly
lucreasiug in length posteriorly. Opercular plates of pieopoda without any air-c£ivities
and scarcely biiobed. Uropodu not much produced, with the inner ramus not attached
so far in front as usual. [Sars, I. c. p. 173.]
1. Philoscia pubescens, Dana. (PL 16. fig. 1.)
Oniscus puhcsc.ms, Daua, U. S. Explor. Exped., Crust, ii. p. 7;30, pi. 18. fig. 2 (1853).
Otiiscus pubcsceiis, Miers, Cat. N. Z. Crust, p. 99 (187(1).
Philoscia vdna, liudde-Luud, Isopoda Terrestria, p. 219 (1885).
Philoscia jjulicsccus, Budde-Luud, Isojjoda Ttrref.tria, |j. 223 (1885).
Omscus puOesceiis, I'llhol, Missiou de File Campbell, p. 440 (I885j.
Oniscus pubescens, Tliomsou & Cliiltou, Traus. New Zealaud lust, xviii. p. 158 (188G),
Philoscia iiiiiiUjDoWlus, Bull. 8oc. Zool. de I'rauee, xviii. p. 1;58 (1893).
TERRESTEIAL TSOPODA OF NEW ZEALAND. 1^7
Specific description. — Body narroAV ohlong-oval, surface smooth and shining in large
specimens, in small specimens bearing numerous short setae. Pirst and second segments
of mesosome with posterior margins straight, lateral angles rounded, posterior margin of
third slightly sinuate, posterior angles of last three segments only slightly producinl
backward, subacute. Metasome al)ruptly narrower than mesosome ; epimera of third to
fifth segments small, closely appressed. Terminal segment triangular, flat, sides straight
or a little incurved, apex subacute.
Antennge very long and slender, from one-half to two-thirds the length of tlie body,
very hirsute, especially towards the end and in small s])ecimens ; third and fourth joints
together eqiial in length to the fifth, which is as long as the flagellum, the three joints
of which are subequtil ; terminal stylet slender, about two-thirds the length of the last
joint. Legs long, greatly increasing in length posteriorly, A^ery spinous. Opercular plates
of the second and succeeding pleopoda with three or four setae projecting at right angles
to the surface. Basal joint of uropoda reaching Avell beyond the last segment ; inner
surface scabroiis and with a few setae ; outer side Avith a well-marked groove, l)ecoming
shalloAver towards the base; inner branch rather more than half as long as the outer,
arisino- oulv a little in front of it, Avitli numerous short setfe and two lonii-er ones at the
apex ; outer ramus much stouter and conical, scabrous, and with some small setae but
fewer than on the inner ramus.
Colon J- light brown, often Avhitish, with various markings of darker broAvn, arranged
roughly in a median and two lateral longitudinal bands, frequently with a roAV of Avhitish
patches at the bases of the epimera. Less yellowish white Avith broAvn markings,
especially on the l)asal joints.
Length about 10 mm., breadth 4 mm.
Kabitat. — Under rotten Avood in forests, Whykare River {Dana) ; Howick, Auckland
(G. M. Thomson); Takapuna (i. Hames) ; Kenepuru, Marlborough {J. Mc3Iahon).
Hemarks. — I have little doubt that the specimens Avhich I have described above are to
be considered as belonging to Oniscus puhescens, Dana. From his description and figures
it is eAident, as Budde-Lund has already inferred, that he was dealing with a species of
Philoscia. His figure show^s an Isopod less narrow than most of my specimens, Imt it
was taken from a specimen only 3 mm. long, and I have specimens of about tlie same
size that correspond very closely to his figure, and from the series at my command I am
able to I'ecord the fact that in young specimens the metasome may be only as long as it
is broad at the base, Avliile in larger specimens it may be fully twace as long as broad,
and that the mesosome shoAvs corresponding A^ariations ; in large forms, too, the antenn;B
become longer and more slender. Dana's specimens were obtained from the north of
Auckland, and most of my specimens are also from places not A^ery far removed where
the species appears to be fairly common, and I know of no other species from that
neighbourhood to which Dana's description could apply.
I have little doubt that the species described by Budde-Lund from the Ca))e of Good
Hope under the name J?hiloscia mina, and afterwards recorded by DoUfus from three
localities in the Seychelles, is the same as the New Zealand species. Budde-Lund's descrip-
tion applies Avell to my specimens, and the groove on tlie outer surface of the base of the
138 DE. C, CHILTON ON THE
viropoda ajipears very characteristic. Dollfus's figaire is very like that of my larger
specimens, though the closely-appressed epimera of the metasorae are not shown, and in
accordance with wliat I have said the raetasome is show^n narroAver than it is in young
specimens. Dollfus calls attention to the fact that while Budde-Lund descrihed the
surface as glabrous, his specimens " presentent an contraire des poils epars, qui parais-
sent, il est vrai, assez caducs." I am able to explain the inconsistency, for my specimens
show that while the smaller specimens (even sexually mature) usiially possess numerous
scattered setoe, the largest specimens have the surface nearly or quite glabrous.
I give figures of the first and second j^leopoda of the male, which are of much the
usual type and do not call for special description. Attached to the male organ in con-
nection with the first pleopod is the vas deferens and apparently a large portion of the
testes, which came away with it when I dissected off the pleopod.
2. PmLOsciA nov^-zealandi^, Filhol.
Philoscia novte-sealandia, Fifhol, Mission de file Campbell, 1885, p. l^i, pi. 54. fig. 2.
Filhol gives the following description of this species : —
" Cette espece, que je crois nouvelle, a le corps allonge, ovalaire, assez bomb6 dans la
portion mediane. Les antennes externes sont couvertes sur leurs bords anterieur et
posterieur et sur leur face externe de poils courts, tres fins, tres serres et ayant dans leur
fornie, leur disposition quclque chose qui rappelle de petites opines. Les segments du
thorax sont granuleux sur presque toute leur etendue et les granulations sont d'un brun
noiratre. La o\i elles font defaiit la carapace est d'une teinte jaune clair. Les parties
granuleuses sont disposees de telle mani^re qu'elles constituent tout le long du corps de
I'animal trois sortes de bandes noiratres ; I'une mediane, les deux autres laterales. Les
stylets caudaux externes sont un pen plus longs que les stylets caudaux internes, mais
ils sont tons converts de poils tres fins sur toute leur surface. II existc une ligne de
ponctuations noires le long du bord externe des stylets externes.
m. m.
Longueur 0-022 a (YOUG.
Largeur ()-008 a 0-009.
•■&
J'ai trouve cette espece aux environs de W^ellington dans File du Nord, aux environs
de Dunedin dans la province d'Otago et enfin dans I'ile Stewart."
I cannot identity this with any species known to me ; the figure shows the side-plates
of the metasome much larger than is usual in Philoscia, but does not help in deciding
what the species Eilhol had before him.
Genus 3. Pokcellio, Latr., 1804.
Porcellio, Spence Bate & Westwootl, British Scssile-cyc-d Crustacea, ii. p. 473 (1868).
Pijrct'Uiu, Budtle-Lund, Isopoda Terrestria, p. 82 (1885).
Force/ lio, Stebbing, History of the Crustacea, p. 42() (1893).
Forcc.lliu, Vs. 0. Sars, Crustacea of Norway, ii. p. 17(5 (1899).
TERRESTRIAL ISOPODA OF NEW ZEALAND. 139
The folio-wins: are the ijeneric characters as mven bv Sars : —
" Body oval, more or less depressed, with the latei-al parts lamellarly expanded.
Cephaloii partly flanked by the side-plates of tlie lirst segiiiciit of inesosome, latei'al lobes
well developed, frontal lobe more or less projecting, and distinctly defined from the
epistome. Metasome not abruptly contracted, epimcral plates of the third to fifth
segments prominent and recurved ; last segment couically produced. Eyes, as a rule,
well developed, subdorsal. Antennae moderately slender, witii the flagellum composed
of two articulations only. Oral parts normal. Legs gradually increasing in length
posteriorly, last pair in male sometimes slightly differing from that in female. (Opercular
plates of tlie two anterior pairs of jileojjoda, and so]netimcs of the three succeeding
pairs, provided with distinct air-cavities. Copulative organs of male of a similar structure
to that in Oniscus. Uropoda distinctly projecting, outer ramus lanceolate, inner much
smaller, lineai-, and originating far in front of the former."
PORCELLIO SCABER, Latr.
Porcellio .irahrr, Latrcillc, Hist. Nat. des Crustaces et dps Insecte^, vol. vii. p. 45.
PorceUio xcaber, S])eiice Bate & Westwood, British Sossile-cyed Crustacea, ii. p. 475 (1808).
Porcellio granir/er, White, List. Crust. Brit. Mus. p. yj (1847), sine descr.
Porcellio (jraniyer, Miers, Ann. Mag. Nat. Hist. (ser. 4) xvii. p. '426 (1876), and Cat. New Zealand
Crustacea, p. 99 (1876).
PorceUio (jraniijer , Budde-Lund, Isopoda Terrestria, p. 129 (1885).
Porcellio yranif/er, Budde-Luud, I.e. p. 149 (1885).
Porcellio f/ranif/er, Thomson & Chilton, Trans. N. Z. Inst xviii. |). 158 (1886).
Porcellio (jraniger, Haswell, Cat. Australian Crust, p. 280.
Porcellio (/raiii(/er, G. M. Thomson, Proc. Royal Soc. Tasninnia, 1892, p. 4.
Porcellio scaher, Stehbing, History of the Crustacea, p. 427 (189."^).
Porcellio scaber, G. O^ Sars, Crustacea of Norway, ii. p. 17() (1899).
Specijic Characters. — " Body oblong-oval, about twice as long as it is broad, dorsal
face slightly convex and very rough, owing to the preseaoe of numerous rounded
tubercles. Cephalon with the lateral lobes rather large and rounded, frontal lobe less
prominent, obtusely triangular. Side-plates of mesosome of moderate size, witli the
posterior corners acutely produced. Metasome occupying about one-quarter of the
length of the body ; cpimeral plates of the third to fifth segments strongly recurved ;
last segment rather produced, terminating in an acute point slightly grooved dorsally.
Antennae less slender, scarcely attaining half the length of the body ; flagellum about as
long as the last peduncular joint, and having its two articulations of nearly equal size.
Last pair of legs differing but little in tlie two sexes. Opercular plates of only the two
anterior pairs of pleopoda with air-cavities. Uropoda with the outer ramus broadly
lanceolate, and comparatively larger in male than in female. Colour of dorsal lace
generally of a uniformly greyish black; sometimes, however, lighter, and variegated
with irregular dark patclies, more rarely black, with the side plates light yellowish.
Length of adult female 14 mm." (Sars.)
SECOND SERIES. — ZOOLOGY, VOL. VIII. 21
140 DK. C. CHILTON ON THE
Habitat. — Found in great abundance throughout the whole of New Zealand,
especially around buildings, in greenhouses, &c. ; rarely in the native bush.
Remarks. — This species is practically cosmopolitan, being found all over Europe,
North Aniei'ica, at the Cape of Good Hope, and Kamtschatka. In Australia it has
been recorded from Melbourne and Tasnaania, and I have specioaens from Sydney also.
In New Zealand it has hitherto been known under the name JPorcellio graniger, Miers,
though Mr. G. M. Thomson suggested some years ago that it was probably a cosmopolitan
species, and pointed out how easily it might be spread by artificial means*. Eudde-
Lund, in his " Isopoda Terrestria," had previously suggested tliat the New Zealand
species was perhaps the same as P. keiils, Latr., but a comparison of specimens has
shown that it differs considerably from that cosmopolitan species, but is undoubtedly
the same as P. scaber, Latr.
The variety marmorata, in which the general dorsal surface is lightly coloured and
variegated with irregular dark patches, is pretty abundant in New Zealand, and the
variety mavghuda " black, with side-plates light yellowish," is also sometimes seen.
Some years ago Mr. W. W. Smith sent me a large series of specimens from Ashburton,
some of the usual form (variety immaculatu), others of the variety marmorata, and a
large number with the whole dorsal surface (in spirit) of a reddish-brown varying from
a light yellowish-brown to dark orange-brown, some of them having the side-plates
lighter in colour than the centre portion ; there is a similar specimen in Mr. Thomson's
collection, and I have occasionally seeu similar specimens from other localities ; in this
form the tubercles on the dorsal surface are hardly so well marked as in some of the
darker forms belonging to the variety immaculata, but there seems to be considerable
variation in the degree of tuberculation in all the varieties.
Many years ago Brandt described albino and partially albino forms of this species,
and gave figures showing all the stages between complete albinos and the ordinary dark-
coloru-ed form'^f .
The great vaiiability in colour of this species has been noted by Eilhol in specimens
collected from New Zealand.
Genus 4. Metoponorthus, Budde-Lund, 1879.
Metoponorthus, Budde-Lund, Isopoda Terrestria, p. 161 (1885).
Metoponorthus, Sars, Crustacea of Norway, ii. p. 183 (1899).
Generic Characters. — " Body oblong, subdeprcssed, with very thin integuments.
Cephalon with the lateral lobes very small, frontal lobe obsolete. Side-plates of meso-
some but very slightly promiaent. Metasome abruptly contracted, with the epimeral
plates of third to fifth segments sub-appressed ; last segment comparatively short,
triangular. Eyes well developed, lateral. Antennuke very small, with the last joint
quite short. Autennte slender and elongated, fiagellum biarticulate. Oral parts nearly
exactly as in Forceltio. Legs slender, and greatly inci'easing in length posteriorly.
* "• Proc. Royal Societ)' Tasmania," 1892, p. 4 (.separate copy),
t Hor;E Soc. entom. Ross. T. viii. (1871), pp. 167-176, 1872.
TEERESTEIAL ISOPODA OF NEW ZEALAND. 141
Opercular plates of the two anterior pairs of [)leopoda with air-cavities, more rarely
also those of third or of all pairs. Copulative organs of male nearly as in Porcellio.
Uropoda rather produced, and of a similar structur(^ to that in Porcellio."
1. ? Metoponorthus truinosus, Brandt.
Porcellio pruinosus, Brandt, Consp. nioiiogr. Crust. Isop. turrcstr. p. 19, fig. 21.
Porcellio zealandicus. White, List. Crust. Brit. Mas. p. 9'.), 181-7 {sine dexcrip.) ; Miers, Cat. N. Z. Crust.
p. 100 (1876).
Metoponorthus pruinosu.t. Buddc-Luml, Isopoda Terrestria, p. 109 (18S5).
Porcellio neo-zelanicus, Thomas & Chilton, Traus. N. Z. Inst, xviii. p. 158 (1886).
Metoponorthus pruinosus, Sars, Crustacea of Norway, ii. p. 181 (1899).
Budde-Lund gives Pot-cellio zealandicus. White, as a douhtful synonym of Metopo
northus pruinosus, Brandt. I have seen the type specimen in the British Museum ; it
is dried and not very well pi'eserved, but is undoubtedly a Metoponorthus, and apparently
very closely resembles M. jjruinosus. It w^ould certainly not be extraordinary if this
cosmopolitan species were found in New Zealand, but I have never met with it, though.
AVhite's specimen, if really from New Zealand, must have been collected there before
1847, and we might naturally have expected that the species would have become
abundant since then.
To make the account of this species complete, I quote here the description given of it
by Miers : —
"Elongate oblong, finely granulous, the granules seriate on the posterior margin of
each segment. Head small, ti-ansversely oblong, with tlie latero-anterior angles not
prominent. Segments of the thorax (the last excepted) with the posterior and infero-
lateral margins straight, the infero-posterior angles obtuse ; last segment of thorax broad,
with the posterior margin concave, the infero-lateral margin straight, the infero-posterior
angle acute. Segments of the abdomen consideralily narrower than those of the thorax,
short ; terminal segment equilaterally triangular, slightly concave above, sides straight.
Caudal appendages with the base shorter than the terminal segment, the longer
(exserted) ranms narrow, acute, projecting beyond the termii^.al segment to a distance
equal to its own length. External antenufe very long and hairy — length nearly one-
third inch. New Zealand (Coll. Brit. Mus.)."
Eamily VI. ABMADILLIID.^.
In this family the body is generally convex, and the animals capable of rolling up
into a ball ; the metasome is not abruptly narrower than the mesosome. There are air-
cavities in two or more of the outer branches of the pleopoda, and the uropoda are
usually short and not produced beyond the terminal segment. In other respects the
family resembles the Oniscida, with which it is so connected by some intermediate
genera that, as Sars has poiuted out, it is rather difficult to get points of difference
that will apply in all cases.
The family contains many genera, several of which have been established during
21 *
142 DR. C. CHILTON UN THE
recent years by M. Adrien DoUfus, but all the New Zealand species appear to be
referable to ArniacUlliclmm and Armadillo.
Genus 1. Armadillidium, Brandt, 1830.
Armadillidium, Budde-Lund, Isopoda Terrestria, p. 49 (1885).
ArmadillirUum, Dollfas, Feuille des Jeunes Naturalistes, iii" Serie, 1" Mai 1892, No, 259 (1892).
ArmadilUdhim, Sars, Crustacea of Norway, ii. p. 188 (1899).
Genen'f! Charactet^fi. — " Body oblong or elliptical in form, very convex, and capable of
being rolled up into a perfect ball. Cephalon with the front distinctly niarginate,
latei'al lobes rounded and sharply defined at the base. Epistome vertical, forming above
a triangular shield, advancing more or less beyond the frontal edge. Side-plates of
first segment of mesosome large, securiform, not incised behind. Metasome. semi-
circular, with the edges continuous throughout ; last segment lamellar, quadrangular or
triangular in form, not extending beyond the limits of the epimeral plati s of the
penultimate segment. Eyes distinct, lateral. Antennuhe with the terminal joint but
little produced. Antennae, as a rule, not attaining half the length of the body, penulti-
mate peduncular joint scarcely longer than the second ; fiagellum biarticulate. Opercular
plates of only the first two pairs of pleopoda with air-cavities. Uropoda very short,
with the basal part broad, lamellar, outer ramus spatulate, inner naiTow, cylindric."
(Sars.)
1. Armadillidium vulgare, Latr.
Armadillo vulgaris, Latreille, Hist. Crust, vol. vii. p. 48.
Armadillo vidguris, Bate & Westwood, Brit. Sess.-eyed Crust, ii. p. 492 (1868).
Armadillidium vulgare, Budde-Luod, Isopoda Terrestria, p. 66 (1885).
Armadillidium vulgare, DoUfus, Feuille des Jeuues Naturalistes, iii"" Serie, No. 259 (l" Mai 1892).
Armadillidium vulgare, Sars, Crustacea of Norway, ii. p. 189 (1899).
Specific Characters. — " Body oblong-oval, more than twice as long as it is broad,
side-contours sub-parallel, dorsal face strongly vaulted and ^Jerfectly smooth. Cephalon,
seen dorsally, broadly quadrangular, transversely truncated in front, lateral lobes
comparatively small, rounded. Side-plates of first segment of mesosome with the
posterior corner acute. Metasome broad, semicircular, scarcely occupying more than
one-fifth of the length of the body ; last segment much shorter than it is broad at the
base, and slightly tapering distally, tip transversely truncated. Antenute very short,
scarcely exceeding in length one-quarter of the body ; fiagellum about the length of the
last peduncular joint, and having its first articulation somewhat shorter than the second.
Last pair of legs with the ischial joint rather large, equalling in length the succeeding
part of the leg. Copulative appendages of the first pair of pleopoda in male with the
tips slightly divergent ; opercular plate of the second pair rather produced, but scarcely
curving outwards at the tip. Uropoda with the outer ramus much shorter than the
basal part, and very broad, its distal edge being continuous with the last segment.
Colour of distal face somewhat variable, sometimes uniformly dark grey or nearly black,
sometimes variegated with lighter patches, generally arranged on the mesosome in three
longitudinal rows, one median and two lateral; between them, moreover, on each
TERRESTRIAL ISOPODA OF NEW ZEALAND. 143
segment is a group of more or less distinct flexuous stripes. Leng'th attaining
14 mm." (Sars.)
Habitat.— '^chon f J. C. Gully), and Mount Egmont (S. H. Drew).
Remarks. — I have a few specimens from Nelson and one from Mount Egmont tliat
undoubtedly belong to this species, whicli is very -widely distributed througliout all
Europe, and the adjiieent parts of Asia and Africa. According to Budde-Luud it has
also been widely dispersed, probably by artificial means, and has lieen found at New
York, Monte Video, Melbourne, &c. It has not been previously recorded from New
Zealand, and it is a little strange that it should have been taken at Nelson and Mount
Egmont, when it has not yet been found at any of the chief ports or in other parts
of the Islands.
My specimens agree very closely witli the figures and descriptions given l)y Sars and
Dolll'us, and I have been able to comjjare them with specimens from England, and
can find no points of diflPerence between them. In the male the first pair of legs has
the carpus a little more swollen than is shewn in Sars' figure, and has the propodos
bent back ujion it so as to form ait imperfectly subchelate hand. I find, however, that
the degree to which this structure is developed varies in different individuals, and it is
perhaps fully developed only in the adult male, or perhaps only during the breeding
season. The long ischium of the seventh pair of legs is also a characteristic of the fully-
grown male ; in the females it is only of normal length ; in the male, too, the meros
and carpus of the first six pairs of legs, and especially of the foui-th, fifth and sixth,
are much more setose than in the female.
With regard to tlie coloui*, Budde-Lund distinguishes two varieties : first, immaculata,
" e plumbeo griseus," and second, cariegata, " annidorum marginibus albis serieque
dorsali triplici vel quadruplici macularum flavarum." Dollfus, who has given a detailed
account of this species in the work quoted, says : " Les 6 sont generalement d'un gris
uniforme, ou avec quelques taches safranees; les 2 d'un brun plus ou moins clair, avec
des taches et marbrures pales." The few specimens that I have from New Zealand
are all males, and are of a uniform dark slaty-grey colour.
Genus 2. Armadillo.
Armadillo, Eufkle-Liind, Isopoda Terrestria, p. 35 (1885).
(icneric Characters. — Ptesembling Armadillidiimi in most respects, but with the shield
on the epistome much less marked, and the grooves for the antennae consequently very
shallow ; the side-plates of the first segment of the mesosome usually, and those of
the second segment sometimes, incised or grooved ; the last segment of metasome
subtetragonal, broader at the base than apex, sides concave ; the outer branches of all
the pleopoda \i itb air-cavities: uropoda with the base large, flattened and produced so
as to fill up the space between the side-plates of the fifth segment and the terminal
segment; outer branch small, inserted on inner margin of the enlarged base ; inner
branch arising more anteriorly, quite concealed in dorsal view by the terminal segment.
This genus contains many species, a large proportion of which ai-e found, as Budde-
144 DE. C. CHILTON ON THE
Lund says, on the islands and shores of the Pacific. In New Zealand there are at least
six species which may he distinguished as follows : —
A. Surface of Ijody uearly smooth.
I. Inferior maroia of first segment of mesosome simple.
a. Outer branch of uropoda very small, rudimentary A. speciosus.
b. Outer branch of uropoda of moderate size A. ambitiosus.
II. Inferior margin of first segment of mesosome grooved along its whole length . A. Dan(E.
III. Margin of first segment of mesosome notched behind A. rugulosus.
B. Surface of body with crests or tubercles.
I. Surface with numerous thin crests or flange-like processes A. Hamiltoni.
II. Surface with setose tubercles.
a. Tubercles large^ about four on each segment of mesosome A. Macmahoni.
b. Tubercles small, acute, numerous A. spinosus.
1. Armadillo ambitiosus, Budde-Lund. (Plate 16. fig. 5.)
Armadillo ambitiosus, Budde-Lund, Prospectus Crust. Isop. terr. p. 7 (1879) ; Isopoda Terrestria,
p. 34 (1885).
Specific description. — Body oval, very convex, nearly smooth, minutely punctate.
Dorsal surface of cephalon marked off from the pre-epistome by a well-marked ridge,
which at the sides projects a little above the surface of the cephalon, centre usually a
little depressed ; dorsal surface smooth or a little vineven, pro-ejiistome smooth, fiat.
Inferior margin of first segment of mesosome thin, with a small tooth posteriorly on the
inner surface ; second segment with the inferior margin entire, thicker in front, the
thickened part ending abruptly in a small tvibercle on tiie inner surface at some little
distance from the margin. Posterior margins of the anterior four segments sinuate,
posterior angle of first produced backwards subacute, that of second a little produced.
Terminal segment of metasome a little longer than its breadth at base, narrowing
abruptly, the posterior portion with sides parallel or slightly divergent ; posterior margin
truncate, slightly rounded or squarely truncate, often with a very small emargination
in the centre.
Antennae minutely setose, flagellum as long as fourth joint of peduncle, and shorter
than the fifth, its second joint three times as long as the first, apex with styliform
appendage about as long as first joint. Eyes somewhat large, with about 20 ocelli.
Basal joint of uropoda with the portion exposed dorsally narrow, about twice as long
as broad, end sharply rounded; extei'i or ramus slender, arising from well-marked notch
on inner margin of base, not reaching quite to the end of l)ase ; inner branch a little
shorter than the terminal segment. Colour usually brownish, varying considerably in
depth of tint, usually with wavy markings of a lighter colour on each side of median
line.
Length up to 1.5 mm., breadth about 7 mm., height 3".5 mm.
Habitat. — Widely distributed in North Island ; also found at Greymouth and Kenepuru.
Remarks. — This is a widely-spread species, and appears to show considerable
variation. It is quite possible that I am including under it forms that others might
look upon as separate species, but my difficulty has been that if I divide it up at all, I
TEREESTKIAL ISOPODA OF NEW ZEALAND. 146
would require to establish at least four or five new species for its members, and I shrink
from such an undertaking.
In specimens that I look upon as typical forms tlie whole surface of the head and
body is smooth, the ridge along the front of the eephalon is not very prominent,
especially in the centre, the end of the last segment is nearly straight (corners rounded)
and with indication of a slight emargination in the centre, and the " tooth " on the
inner side of the lateral margin of the first segment, and the small tubercle on that of
the second, are fairly distinct. I have some specimens from Greymouth, where the
ridge along anterior border of head is much move pronounced, the "tooth" and
" tubercle " less evident, end of last segment slightly rounded with no indication of
emargination ; the surface of the head, moreover, presents some slight irregularities, and
there are a few indistinct wavy elevations on the sides of the median line of the bodv •
the central part of the last segment is a little raised, though hardly sufficient to be
called ridged, and the basal portion of the last segment, and the epimeral portions of the
third to fifth segments of the metasome, are rather broader than in the type.
Two specimens from Wauganui in Mr. Thomson's collection agree with these Grey-
mouth specimens in the ridge on the anterior margin of eephalon, and in the " tooth and
tubercle," but the epimeral portions of metasome and the basal portion of the uropoda
are fully as narrow as in the typical specimens ; the end of last segment is quite squarely
truncate, and the whole body is nearly smooth. In these two specimens the inner
branch of the uropoda is shorter than in any other I have seen.
Specimens from other localities show other combinations of these various characters,
and on the whole I think it wisest to consider them all as belonging to one species.
2. Akmadillo Dan^, Heller.
Splierillo dana, Holler, Reise der Novara, p. 134, pi. xii. fig. 4 (1865).
Armadillo inconspkuus, Miers, Ann. & Mag. Nat. Hist. ser. 4, xvii. p. 22a (187(i); Cat. N. Z. Crust.
p. 95, pi. ii. fig. 4 (1876).
SpheriUo dancs et Armadillu inconspicuus , Thomson & Chilton, Trans. N. Z. lust. .wiii. p. J5'J (1880).
Armadillo dame, Butlde-Lund, Isopoda Terrestria, p. 39 (1885).
Armadillo inconspicuus, Budde-Luud, /. c. p. 39 (1885).
Armadillo inconspicuus, Filhol, Mission de I'llc Campbell, p. 439 (1885).
SpheriUo dance, Filhol, /. c. p. 440 (1885).
Specific deso'ipHor/ . — In the eephalon surface and general appearance closely re-
sembling A. amhitiosus. Posterior border of first segment of metasome only very slightly
produced backwards, that of others straight. Lateral margin of first segment with a
narrow groove extending along its whole length, becoming shallower anteriorly, second
segment also with lateral margin deeply grooved. Segments of metasome short, epimeral
portions less elongated than in A. ambitioims, terminal segment a little broader at base
than at tiie extremity, end truncate, slightly rounded.
Basal [)ortion of uropoda M'ith exposed portion small, end rounded, outer ramus
minute, inner ramus reaching very nearly to the end of last segment.
Colour brownish, with lighter wavy markings on each side the median line.
Length about 10 mm.
146 DR. C. CHILTON ON THE
Habitat. — Takapuna, Auckland (L. Hames) ; Auckland (Heller), Bay of Islands (Dana).
Filhol says that he has found this species in great abundance in the North Island,
and that it becomes less and less abundant towards the south, tliough he gathered a
few examples of it on Stewart Island. The statement as to the greater abundance in
the north is also true of Armadillo inconspicims, which this species closely resembles,
and there is nothing to show definitely that Filhol distinguished the one from the other.
I have a single specimen of this species before me from Takapuna, Auckland, and
have no difficulty whatever in identifying it with A. inconspicuus, Miers. It also agrees
very well with Heller's description, if we remember that the part he speaks of as the
outer branch of the uropoda is the outer distal portion of the base, and that his " inner
branch " is really the outer, though, as usual in this genus, arising from the inner
margin of the produced portion of the base.
In colour and general appearance this species is very like A. ambit iosiis, Budde-
Lund, but it may be readily distinguished from that species by the groove on the
lower margin of the fii'st segment of the mesosome, and by the minute outer branch
of the uropoda.
3. Armadillo speciosus, Dana. (PL 16. fig. 6.)
Armadillo speciosus, Dana, U. S. Explor. Exped., Crust, ii. p. 718, pi. 47. fig. 2 (1853).
Armadillo speciosus, Miers, Cat. N. Z. Crust, p. 95 (1876).
Armadillo speciosus, Budde-Luud, Isopoda Terrestria, p. 39 (1885).
Armadiilo speciosus, Filhol, Mission de I'ile Campbell, p. 439 (1885).
Specific description. — Body very convex. Ce])halon with frontal margin raised, a
little interrupted in the centre. Segments of mesosome each with transverse row of
indistinct tubercles or granules ; inferior margin of first segment simple, curving a little
outwards. Segments of metasome in close contact, the last nearly as broad at apex as
at the base; sides concave, posterior margin straight. Uropoda with the outer joint
minute, rudimentary, inner branch slender, extending to the end of last segment.
Colour light reddish brown, with darker markings.
Length about 6 mm.
Habitat. — Bay of Islands {Dana)\ Chatham Islands {Hutton).
Remarks.- — I have three or four specimens (mostly imperfect) from Cliatham Islands
that I have little hesitation in assigning to Dana's species. He describes his genus
Armadillo as having the external ramus of the uropods obsolete, and figures his species
A- speciosus in accordance witii this view. It is true that in the specimens before me
the outer branch is present, but it is so minute that it would be impossible to show it in
a figure the same size as that which Dana gives of the whole animal, and it would
be diflicult to see it with the magnifying power likely to be used for making such
a drawing, while in the view of the uropoda from beneath, which is the one shown in
Dana's enlarged figure, it is quite concealed. In other respects it agrees so well with
Dana's descrijjtion and figure that I identify it without much hesitation with his
species.
It can be readily distinguished from A. dance. Heller, which it otherwise greatly
resembles, by the absence of a groove on the inferior margin of the first segment of the
TEREESTKIAL ISOPOUA OF XEW ZEAL.AJVU. 147
mesosome. The specimens from Nelson, Welliug-toii, &c., jji^eviously identified with this
species hy Mr. Thomson and myself *, do not bclont^' to this species, but to Armadillo
(imhitiosus, Bndde-Lnnd.
4. AiiAiADiLLO iiUGULOSTFS, Miers, 1870. (PI. 16. iig. 7.)
Cubaris rugulosus, Miers, Ann. & Mag. Nat. Hist. ser. 1, xvii. p. '.l'l~i (1876) ; Cat. N. Z. Crust, p. 9(5,
pi. ii. fi-. f) (187G).
Cubans ruijulusiis, Chilton, Trans. N. Z. Inst. xv. p. 73 (18.S2).
Annadillo riii/ii/nsns, Buddc-Lund, Isopoda Terrestria, p. 40 (188.J).
Cubaris rugulosus, Filhol, Mission de File Campbell, p. ilO (1885).
Cubaris rugulosus, Thomson & Chilton, Trans. N. Z. Inst, xviii. p. 158 (1888).
Specific description. — Body moderately convex, surface of segments uneven, faintly
rugose. Head hroad and transverse, front margin revolute, first segment of mesosome
with two shallow depressions diverging anteriorly on the anterior part of the uj)])er
surface ; posterior margin sinuous, angles produced backwards; lower posterior max'gin
with a notch for reception of succeeding segment, the notch not extending along the
inferior margin ; second segment similarly notched. Dorsal surface of segments of
mesosome often with a slight groove parallel to posterior margin ; the second, third, and
fourth narrowed at sides with inferior margins rounded ; fifth, sixth, and seventli broader,
with inferior margins truncate. Metasome with terminal segment broadest at base, sides
at first suddenly converging, then parallel or slightly divergent, extremity square
truncate. Antennae finely hirsute, flagelluin shorter than fifth joint of peduncle, second
joint three times as long as the first. Uropoda with outer branch small, not quite
reaching to the end of last segment; inner branch short, reaching halfway from its base
to end of last segment.
Colour light brown, Avith variegated markings of a rich reddish brown, some speci-
mens very dark.
Length about 6 mm.
Huhitut. — Very abundant in Sotith Island.
Bemarks. — This species is much smaller than either of the two preceding, and may
generally be readily distinguished from them by the difi'ereut character of the notches
in the; posterior lateral margins of the first and second segments of mesosome. I have,
however, some specimens from Kenepuru in which these notches are less marked, and
the inner branch of the uropoda is a little longer tlian usual, and the colour is rather
greyish. The specimens are not very well preserved, and I prefer for the meantime to
consider them merely as a variety of the species imder consideration.
The oblique depressions on the first segment of the metasome described by Miers are
generally present, but are more distinct in some specimens than in others ; they are also
]n-esent in some specimens of Armadillo anibitiosus, Budde-Lund, and are probably more
or less the necessary consequence of the head fitting in to the first segment when the
animal is rolled up, and are therefore probably of little classificatory value. In some
specimens the irregularities on the dorsal surface are more distinct than in others, and
* Trans. X. Z. Inst, xviii. p. 15y.
SECOND SERIES. — ZOOLOGY, VOL. VIII. ^2
148 DR. C. CHILTON ON THE
tliey may even give the appearance of a poorly-marked transverse row of small tubercles
on the segments of the metasome ; it is perhaps sijecimens of this kind that Heller
described under the name Spherillo monolinus, but as he says nothing about the notches
on the inferior margins of the first two segments of the metasome, I do not feel
justitied in definitely identifying our species with liis.
5. Armadillo monolinus, Dana, 1853.
Spherillo monolinus, Dana, U. S. Explor. Exped., Crust, ii. p. 719, pi. 47, fig. 3 (1853).
Spherillo monolinus, Heller, Voy. Novara, Crust, p. 135 (18G5).
Armadillo Aucklundicus, Budde-Luiid, Isopoda Terrestria, p. 40 (1885).
I have seen no specimens that I could refer to this species. Dana describes it as
follows : —
" Head arcuate in front. Segments of thorax transversely marked with a beaded
ridge and laterally truncate, anterior segment longest and marked with two beaded
ridges. Abdomen semicircular, third, fourth, and fifth segments laterally obtuse, tlie
last with a nearly subquadrate apex, and much broader at base. Caudal appendages
subtriangular, shorter than breadth at base, inner margin broadly excavate. Antennae
nearly naked, flagellum hardly shorter than preceding joint. Length 4 lines."
Dana's specimens were obtained at " Wykare River, near Bay of Islands."
Heller describes a specimen from Auckland, and at the end of bis description says:
" I denote this species as ^S*. monolinus" without making it clear whether he is referring
it to Dana's species or giving it as a new species, using by inadvertence the same name.
Budde-Lund supposes Heller's species to be new, and therefore changes the name to
Aucklandicus. I tliink, however, that it is more likely that Heller intended to refer his
specimen to Dana's species, and there is nothing in his description inconsistent with this
supposition ; Miers had evidently taken this to be Heller's intention, but curiously
enough he docs not give the reference to Heller's descrij)tion, though he quotes it
for the habitat when describing Spherillo monolinns, Dana. It is possible that this
species may be the same as A. I'/if/idosws, Miers.
6. Armadillo Hamiltoni, sp. nov.
" Remarkably sculptured Terrestrial Isopod." — G. M. Thompson, Annals & Mag. N. H. ser. 6, xii.
p. 225, pi. iv.
Specific description. — Oblong-oval, breadth rather more than half the length, epimeral
portions greatly developed and projecting downwards and outwards, central part greatly
raised above the epimera and richly supplied with spines and crests. Cephaloa with the
dorsal surface produced forwards into a thin jilate ))rojecting far over the bases of the
antcnnte ; from the dorsal surface of the ceplialon arise two transverse lateral crests or
flanges, which project horizontally forwards as far as the central prolongation of the
vertex ; the posterior margin of the cephalon is curved upwards, and a little exterior to
the median line is produced upwards into a conical tooth ; the posterior margins of
each segment of the mesosome is similarly produced upwards and somewhat backwards
into two teeth, those on the seventh segment being very large and prominent ; the first
TEREESTRIAL ISOPODA OF jVEW ZEALAND. lid
segment has two pairs, and each of the other six segments one jmir, of conical teeth or
spines arising at right angles to the dorsal surface and situated in the same longitudinal
lines as the teeth already described. More laterally each of the first six segments l)ears
on each side two thin crests arranged in two longitudinal lines, but the crests of each
segment widely separated from tliose of the coutiguous segments ; in the seventh segment
the outer pair of crests is represented only by small spines. External to the outer row of
crests each segment bears tAAO or three small tubercles or spines, wliich are concealed in
dorsal view by the outer row of crests. Epimera very largely developed and projecting
someAvbat horizontally, that of the first segment much larger than any of the others ;
first two segments of raetasome concealed in dorsal view by the projecting liinder margin
of the last segment of the mesosome ; each of the third to fifth segments has tbe binder
margin produced into two teeth similar to those in the mesosome but smaller ; these
teeth increase in size from tbe thii'd to fifth segment. From the centre of the terminal
segment arises a small keel projecting backwards, and ending in a sharp tooth ; the
epimera of the third to fifth segments are very long and narrow ; liinder margin of
terminal segment straight, not much narrower than the base ; sides concave.
Eyes of moderate size, convex, of about twenty facets. Antennul;« and antennas not
observed. Uropoda with the lateral portion of the joint long and narrow, end rounded,
outer branch narrow, fully three times as long as broad, arising from a well-marked
notch in the inner margin of the base, not reacbing to the end of tbe terminal segment,
ending in a small seta ; inner branch very short, scarcely reaching as far as the base of
the outer branch.
Colour a rich brown, with niimerous markings of a darker brown.
Length 6 mm.
Habitat. — Petane, near Napier [A. Hamiltorb).
Remarks. — In accordance with a wish expressed by Mr. G. M. Thomson, I gladly
name this species after its discoverer, Mr. A. Hamilton. I have only the dried specimen
originally described and figured, but not named, by Mr. Thomson, but it is, I think,
sufiicient to show that the species may, provisionally at any rate, be placed under
Armadillo. The wealth of crests cind spines or teeth on the dorsal surface is quite
extraordinary, and I fear tbat the description Avill convey a very inadequate idea of the
actual specimen ; a better idea may be got by consulting Mr. Thomson's figures.
Mr. Thomson describes tbe legs as " very feebly developed and, as far as 1 could make
out, appear to want the dactylos." There are no legs now attached to the specimen, but
in the tube I found two fragments possessing normal dactyla similar to those found in
other species of Armadillo.
7. Armadillo Macmahoni, sp. nov. (PI. 16. tig. B.)
Specific descrijdion. — Body convex, tuberculated and setose, especially on the tubercles ;
surface with depressed hexagonal markings. Cephalon with the dorsal surface roughened,
somewhat setose, front with well-marked transverse ridge, a little lower in the middle.
Eirst segment of mesosome with inferior margin revolute anteriorly, posterior portion
deeply notched, second segment also notched, but with the inner lip of the notch not
150 DK. C. CHILTON ON THE
reachiu^ downwai-cls so far as lower margin. The mesosome bears four longitudinal
rows of tubercles (/. e., from tubercles on each segment), the two inner rows a little
external to the median line, the other two more lateral ; on the anterior segments the
tubercles are not very prominent, but on the succeeding segments they become larger
and more prominent and project slightly backwards ; on the seventh segment the two
medicin tubercles are very large and project backwards over the metasorae, while the
lateral tubercles are pooi'ly marked and form only slight elevations at the outer side
of the base of the others. The tubercles are covered especially near the apex with
numerons stiff setae. Shorter setae are also found on the rest of the surface, and some of
them, especially along the posterior margins of the segments, are broad and scale-like.
Metasome witli an indistinct median ridge formed by a setose tubercle on each of
the last four segments, that on the fifth the largest ; first and second segments short,
the first almost concealed by the preceding segment, all the segments of mesosome
fitting closely together ; terminal segment much broader than long, sides concave,
posterior margin slightly convex.
Eyes rather small, of about ten ocelli. Antennae short.
Uropoda with base fitting closely into the space between the side-plates of the fifth
segment and the terminal segment ; outer branch very minute, not projecting beyond the
inner margin of the base, inner branch reaching about to end of last segment.
Colour brown.
Length about 6 mm.
Habitat. — Keuepuru, Marlborough, in the bush [MacIIahon).
'Remarks. — I take pleasure in naming this fine species after Mr. Joseph MacMahon, to
whom I am indebted for many Terrestrial Isopoda collected at Keuepuru.
8. Aemauillo spinosus, Dana, 1853.
Upherillo spinosus, Dana, U. S. Explon Exped., Crust, ii. p. 72.3, pi. 47, tig. G (1853).
SpherUlo spinosus, Miers, Cat. N. Z. Crust, p. 97 (1876).
Armadillo spinosus, Buddc-Luiid, Isopoda Terrestria, p. (1885).
Spherillo spinosus, Thomson & Chilton, Traus. N. Z. Inst, xviii. p. 159 (1885,
The folloAving is Dana's description of this S2)ecics : —
" Body bristled throughout with subacute spines, margin either side a little produced
and segments laterally truncate. Head nearly trapezial, arcuate in front, and a little
broader than behind. Eirst segment of thorax largest ; segments of abdomen laterally
obtuse, the last subquadrate, not broader at base, truncate at apex.
" New Zealand, near Bay of Islands (Coll. Dr. C. Pickering; Dana). Under bark
of pine-trees."
Remarks. — I have seen no specimens of this species, but it appears to come near
A. MacMuhoni, differing, however, in having the spines much more nvimerous and more
acute.
TERRESTRIAL LSOPODA OF XEW ZEALAND. 151
EXPLANATION OF THE PLATES.
Jleferoice Lcftcvs.
a' = aiiteniuila.
a" = antciniii.
c = rephaloii.
/.,«. = labrum superior.
/./. = labnim inferior.
M.dex. = right iiiaiidiblc
M.sin. = left uiaiulil)lc.
vur' & m.r- = 1st and 2iul maxillie.
iiKijj. = inaxillipede (outer, /'./'., posterior or iiiulrr aspect) .
m.rp* = maxillipcde (inner, i.e., anterior or uppei' aspect).
prn = pereion (or niesosoine).
y,'--' ■ ■ . ; = 1st, .'2nd, ;5rd .... 7tli lej;-.
jA', &C. = extremity of 1st leg, &e.
J// = pleon (or luetasonie).
plp^'^, &c. = 1st, ,2nd pleopod (anterior aspect).
/)///•""*, &c. = 1st pleopod, &c. (posterior aspect).
urp = uropod.
[Wliere necessary, the sex is indicated l)y the sign (^ or % jjlaceil after the letters as above.]
Plate XL
Fig. 1. Lhi'ia )i(iV(i'-:ciihnidi(P, Dana. Dorsal view of whole animal and details.
Platk XII.
Fig. 1. Trichoni.icus phormianus , sp. nov. Porsal view and details.
2. Trichoniscus otakeiisis, sp. nov. Dorsal and side views of female, dorsal view of male, and
enlarged view of head and antennic of female.
'3. Haplop/i//i(i/juus Hebii.ti'i, sp. nov. Dorsal view and details.
I'LATK XIII.
Fig. 1. Tricliuii'isnoi Tlioiiisuui, Chiltdn. Dorsal view and details.
2. Tylo.i iK'ozclan'icus, s|). nov. Siih; view and details.
Pl.ATK XIV.
Fig. 1. Scyplioiiiscus ivaitaieii.iis, nov. gen. et sp. Dorsal view and details.
2. Scypha.v ornatif.t, Dana. Dorsal view and details.
SECOND SERIES. — ZOOLOGY, VOL. VIII. -3
152 DH. C. CHILTON ON THE TERRESTRIAL LSOPODA OF NEW ZEALAND.
Plate XV.
Fio-. 1. Scyphux ornatus, Daua (coutiuued). Details.
2. Scyphax (?) aucklandite, G. M. Thomson. Dorsal view and details.
3. Actacia euchroa, Dana. Dorsal view and detail-s.
4. ActeBCJa opihensis, sp. nov. Dorsal view of pleon and details.
Plate XVI.
Fig. 1. ActtBcia opihensis, sp. nov. (continued). Details.
2. Oniscus punctatus, G. M. Thomson. Details.
3. Oniscus kenepurensis, sp. nov. Dorsal view and details.
4. Phiioscia pubescem, Dana. Details.
5. Armadillo ambitiosus, Budde-Lund. Antenna and terminal portion of pleon.
6. Armadillo speciosus, Dana. Terminal portion of pleon.
7. Armadillo rugulosus, Miers. View from below of lower margins of 1st and 2ud segments of
pereiou.
8. Armadillo Macmahoni, sp. nov. Dorsal and side views and dorsal view of pleou and details.
AUG 3;
^^j
[ 152* ]
TEKRESTKIAL ISOPODA OF NEW ZEALAND.
(Trans. Linn. Soc, '2nd scr. Zool. vol. viii. pt. 4.)
EKRATA.
Page 10;}, line 7 from top, delete the comma after " portion."
0 from bottom, insert a comma after " lamellar."
17 from tup, for " (ilo/icridie " read " Glo?»erid8e."
„ for " with " riiiid " without."
,. for " eggs " read " eyes.''
,, for " Hayer ' read " llarger."
,, for " Hayer " read " Harger."
„ for " occasionally " read •' provisionally."
,, /(/(■ " vein "' read " view."
„ for " but with apex " read " both with apex."
,, /"/• '• Hayer "' read " Harger."
„ /;/■ "eggs '■ read " eyes."
„ between lines 'A & 4 from bottom insert : —
" Cylloma oculattnn. Budde-Limd, Isopoda Terrestria. p. 46 (1885)."
131, line 11 from bottom, /<«• ••extremities" read " antenniilse."
138, „ 14 Ironi top, /()(■" p. 144" /■««/'■ ji. 444."
14.5, „ 14 ,. /or " last segment " j-('((rf " uropoda."
148, „ 2 ,, /oy "metasome '" >-«a(Z •' mesosome."
l.")!, „ 18 „ for "yj/yj '■<"'■", &e. " read '' pJp '■-'*, &c."
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THE
TRANSACTIONS
OP
THE LINNEAN SOCIETY OF LONDON.
ETUDE D'UIS'E ESPECE NOUVELLE 1)E LEPADIDES
{Scalpellum giganteum, n. sp.) et de Poecilasma carinatum, Hoek.
PAK
A. GRUVEL,
FACULTY DES SCIENCES, BORDEAUX.
(Communique par !\1. Ic Pnif. Ct. B. Howes, Sec. Linn. Soc.)
^L O N D O N :
PIUNTKl) FOR THE I.rXNKAN SOCIKTV
HV 1 WI.OK AND KKANCIS, KKII I, ION COrKT. KI.KKT SlUKKT.
SOLD .\T THE society's .VPARTMENIS, HUKUNGTOX-llOL SE. I'lCCAlMlLY, AV.,
ANO HV r.ON(iM\NS, ORKEX, ANT) CO., rATERXO.SrEK-l!0\V.
Jiiipisl 1.001.
AUG -^^ i<5n?
V. Etude d'taie Espcce nouvelle de Lepadidos (Scalpollum gjiganteum, n. sp.) et de
Poecilasma Cfu-inatum, Tloel;. Far Monsieur A. Gruvel, Charge d'un Conrs de
Zoologie a la Faaulte des Sciences de Bordeaux. [Communique par M. Ic Pi'ofosscur
Howes, Sec. Linn. Soc.)
(Planche'^17.)
Lu le '2.1 fovrier, IDOl.
J E clois a la haute bienveillance de Monsieur le Professeur Hay Lankestcr, Directcur
clu British Museum (Nat. Hist.), et a ramahilite de son collahorateur, Monsieur Jeffrey
Bell, de pouvoir deerire aujourd'hui la plus grande espece actuellement couuue
ajjpartenant au genre Scalpellitm. C'est a cause de ses dimensions relativement con-
siderables que je propose de lui donner le nom de Sc. gigantetmi.
Sur le pedoncule de I'un des echantillons, se trouvait fixe un Poecilasma que I'etude
m'a dcmontre etre P. carinatum, decrit par lloek, dans son " K-eport on the Cirripedia "
du ' Challenger.' La comj^araison que j'en ai pu faire avee les echantillons types de Hoek
au British Museum, ne m'ont laisse aucun doute a cet egard. Jc reviendrai du reste plus
loin sur quelques-uns des caracteres de cette espece.
ScALPELLUM GIGANTEUM, n. sp. (Blanche 17. fig. 1.)
Diagnose. — Quatorze plaques cajiitulaires, entiercment calcifiees, recouvertes, presque
completement, par une opaisse cuticule chitineuse, I'apex seul faisant saillie en dehors.
Limite exacte des plaqiies impossible a determiner sans enlever la cuticule ; ces plaques
sont largement separees les unes des autres. Ciipitulum aplati siirtout vers le sommet,
carene reguHerement courbe, umbo a I'apex. Umbo des plaques careno-laterales tr^s
rapproche de la base, recourbc en avant et n'atteignant pas le bord externe de la carene.
E-ostre ovale, entierement cache sous la cuticule.
Pedoncule aussi long que le capitulum, li pen pres regulierement cylindrique, orne de
six series longitudinales et alternes, d'ecailles allougoes transvcrsalement, non imbriquces,
chaque rangee longitudinale comprenant de 10 a 12 ecailles recouvertes par la cuticule.
Dimensions: — Longueur du capitulum, 15 mm.; largcur, 32 mm.
Longueur du pedoncule, 45 mm. ; largeni-, 15 mm.
Cette espece est voisine de Sc. regiinn, Hoek, et Sc. Darwinii, Hoek, mais elle s'en
distingue nettement par un certain nombre de caracteres et en particulier la forme des
plaques et des ecailles.
Je I'ai designee sous le nom de Sc. gigantemn, car c'est la plus grande espece
actuellement connue.
CapiMim. — Le capitulum est aplati, beaucoup plus vers le sommet que vers la Ijase.
Vu par le cote, il represente un triangle curviligne, presque isocele (le cote dorsal etant
cepeudant un pen plus long que le cote ventral), dont les deux cotes seraicnt convexes ct la
SECOND SERIES. — ZOOLOGV, VOL. YIII. 24
154 MOXS. A. GRUVEL— ETUDE DVIS'E,
base legcrement concave. Le borcl anterieur forme uno ligne ;\ pen pres rcgvilierement
eonvexe.
Les quatoi-7,e plac^iies qu'il porte sent largement separees les unes des autres, surtout les
principales, et noyees clans une epaisse cuticule chitiueuse jaunjitre sale, un pen comme
chez Sc. sqimmulifcrum, Weltner, mais la cuticule a ici une epaisseur bien plus consider-
able. II est impossible de se rendrc uu compte exact de la forme et de la disposition de
ces plaques si on n'enleve pas, au prealable, la plus grande partie de la cuticule qui les
recouvre et qui, se poui-suivant entre elles, sert a les unir les unes aux autres.
Les stries d'accroissement des plaques sont, du reste, parfaitenient nettes sur la
cuticule. Celle-ci est couverte de nombreux poils courts fortement barbelcs, de petites
vermes cbitineuses couvertes de poils tres courts et enfin par de nombreux tubercules
cliitincux, plus fortement colores que le i^este de la cuticule et dissemines d'une fagon
assez dense mais irregulierement. Le tout produit au doigt, quand on le passe sur le
capitulum, la sensation qu'on eprouve en passant le doigt sur un velours un pen raide.
La cuticule se poursuit, du reste, sur le pedoncule, sans modification appreciable dans
sa structure.
Tergtmi. — Forme irregulierement quadrangulaire. Bord anterieur eonvexe ; bord
scutal eonvexe dans sa partie inferieure, concave dans sa partie superieure et termine
anterieurement par nn tres court espace rectiligne. Bord carenal droit et bord careno-
apical legerement concave, ce qvii fait que I'apex qui est droit, semble, cependant,
legerement recourbe en arriere. Une arete nette unit I'apex a I'augle inforieur, les stries
d'accroissement sont nettement visibles. (PI. 17. fig. 2.)
Scutum. — A peu pres triangulaire, bord anterieur eonvexe, bord inferieur tres legere-
ment eonvexe aiissi, enfin bord latero-tergal eonvexe sur ses § inferieurs et concave sur
son ^ superieur, ce qui fait que I'apex qui est droit, semble aussi legerement recourbe
en arriere.
La longvieur du bord inferieur n'egale pas la h de celle du bord anterieur. Une arete
nette va de I'apex a Tangle infero-interne ; stries d'accroissement bien marquees. (PI. 17.
fig. 2.)
Carene. — Regulierement courbe ; umbo a Tapex qui ne fait jias saillie entre les terga.
Bord dorsal eonvexe, ainsi que les bords lateraux. Le bord basal est fortement eonvexe et
son angle inferieur, mousse, n'atteint j)as le sommet des pieces careno-laterales. Concave a
sa partie interne, elle va en s'elargissant rc%ulierement du sommet vers la base. (Pig. 2.)
Plaque Jaterale siqu'rieure. — Irregulierement quadrilatere ; bord tergal droit, bord
scutal fortement excave, ce qui fait saillir I'apex en avaiit ; les bords posterieur et inferieur
sont regulierement courbe et se coupeut a angle tres arrondi. (Pig. 2.)
JRostre. — Petit et ovale ; bords lateraux caches par le sommet et les bords internes
tres concaves des pieces rostro-laterales. II est, du reste, entierement masque par la
cuticule.
Plaque rostro-lateralc. — AUougee, etroite, plus large dans sa partie anterieure que
dans sa partie posterieure. Le bord rostral est concave et delimite, avec son congencre,
un espace ovale oil se place la partie centrale du rostre. Cette piece rostro-laterale
est inclinee en avant.
ESPECE XOUVELLE DE LKPADIDES. 155
Plaque mfra-medio laterale. — De forme triaugulaive ; le Ijord iiiferieuv est Ic plus lon^-,
l^uis le bord j^ostei'ieur et eufiu le bord anterictir. A})cx dirige vers le sommet du
capituhiui.
Plaque careuo-laterale. — Allongee, etroite, inclinee en arriere. Apex fortement I'ecourbe
en haut et en avant et n'atteignant pas le bord inferieur ni le bord externe de la carene.
Umbo tres rapproche de la base. Le bord inf(''ro-dorsal est arrondi et egal environ a
une fois et demi la longueur du bord superieur. Quant au bord carenal, il est tres
concave et sa longueur depasse uu pen la moitie de celle du bord superieur.
Les limites de la cliitine qui borde les plaques ne correspondent pas du tout comme
forme et comme disposition il celles des plaques elles-momes. Cette limite est, dureste,
pour les plaques inferieures, tres diflicile k determiner. Les descriptions que nous
venous de donner portent sur les pieces calcaires seulement, telles qu'on les aperyoit ajjres
avoir enleve la cbitine qui les recouvre (fig. 2). Dans la fig. 1 les animaux sont repre-
sentes tels qu'on les trouve, c'est a dire entierement reconverts par la chitine.
Longueur du capitulum, 45 mm. ; largeur, 32 mm. ; epaisseur max., IG mm.
Pedoncule. — Aussi long que le capitulum et presque aussi large. De I'orme a peupres
rcgulierement cylindrique, mais cependant legerement dilatee vers le capitulum et vers
la base. II est orne de six series longitudinales et alternes d'ecailles allongces transver-
salenient, non imbriquees et separces, d'une serie a I'autre, par un espace chitineux assez
large, cliaque serie longitudinale comptant de 10 a 12 ecailles en general entierement
recouvertes par la cuticule, mais quelques luies out leur bord lil)re erode et mis a nu.
L'ecaille apparait alors, tres blanche.
Longueur du pedoucide, 45 mm. ; largeur, 15 mm.
Bonche. — Le labre est fortement saillant en avant [lab, figs. 7 et 8). Vu de face, il a
un peu la i'orme d'une couj)c ; la partie anterieure est arrondie et se trouve scparee par
un retrecissement de la partie posterieur ecrasee ; le bord posterieur est echancre en son
milieu pour recevoir les extrcmites des autres pieces masticatrices. Ce bord ne porte,
du reste, aucune espece d'ornements.
Les PaJpes (fig. 3) sont aplatis en lames et ornes vers les extrcmites libres et les
parties superiem-e et inferieure de soles assez courtes et fortement barbelees. L'epitlieiium
est charge de nombreuses cellules j)igmentaires d'un bruii noir et disposees avec assez
de regularite.
Les MancUbnles (fig. 4) portent trois dents, dont la superieure est la plus longue et
la plus forte ; elle est un peu pkxs eloignee de la deuxieme que celle-ci de la troisit-me.
L'angle inferieur est saillant et garni de soies courtes et robustes ; les plus hmguos
sont situes fi I'extremite.
Les Mdchoires (fig. 5) presentent ime forte dent suj)erieure, suivie de nombreuses
dents, formant une dizaine de rangees de plus en plus courtes et faibles. Entre celles-ci
s'en trouvent d'autres fines et non rigides. II u'existe pas d'encoche sur le bord fibre,
qui est d'abord legerement concave dans sa moitie superieure, mis legerement convexe
dans sa moitie inferieure. Le bord superieur des pieces porte des soies longues et flexibles.
Aucune n'est barbclee.
24*
156 MONS. A. GRUVEL— ETUDE D'UNE
Enfin les Palpes de la levre inferieure (fig. 6) sont les i:)irces les plus larges de
raj)pai'eil masticateiiiv Leur bord libre pruseute une legere eucoclie en son milieu et
cette encoclie determine ainsi la formation d'une saillie superieure et d'une inferieure,
portant chacune une toufi'e de soies, barbelees d'une fa§on identique a celle des soies
des palpes de la levre superieure. Sur le bord superieur, il existe aussi des soies beaucoup
plus longues, plus fines, et egalement irregulieres et barbelees.
Cirrhes. — Les cirrbes sont, en general, assez longs et robustes.
l'"" paire : Kames inegales, I'anterieure avee 12 articles ; c'est le 1' a partir de la base
qui atteint le maximum de largeur. lis sont garnis de soies nombreuses et assez courtes
sur toute la surface interne et sur les bords anterieur et posterieur.
La rame posterieure porte 16 articles, et la region moyenne, qui est la plus large, a une
largeur egale, environ, a la moiti^ de celle du 4" article de la l"' rame. Les soies sont
disposes de la meme fa^on.
La 2" paire est legerement separee de la l''^ Les rames sont a pen pres ogales, cependant
la posterievire depasse I'anterieure des 4 derniers articles extremement courts. La rame
anterieure porte 31 articles.
Les 3% 4" et 5'' paires de cirrhes sont semblables a la 2''.
Quant a la 6'" paire : la rame posterieure, qui possede 48 articles et a 50 mm. de long.,
depasse I'anterieure des deux derniers. Les soies qui les garnissent anterieurenient sont
longues, robustes et terminees en pointe tres fine a I'extremite ; elles sont disposees en 4
doubles rangees vers les articles terminaux. Posterieurement, a la limite des articles,
se trouvent groupees 5 ou 6 soies courtes.
Appendices ca/ulaux. — Formes cliacun de 4 articles, dont le basilaire egale en
longueur les 3 autres. Les articles vont en s'elargissant vers I'extremite libre,
de sorte que le dernier est le plus large et sa partie distale est, elle-meme, plus large
que sa base, de plus elle est nettement tronqviee (fig. 8'). A la limite des articles, du
e6te interne et du cote externe, se trouvent des groupes de soies dont les plus longues
sont celles de I'article terminal, et leur longueur ne depasse pas celle de cet article
lui-meme.
Penis. — Retreci et recourbe a sa base, il reste a peu pres cylindrique sur la moitie de
sa longueur. A partir de la, il s'amincit progressivement pour se terminer en j^ointe
fine, mousse et ornee d'un petit bouquet de soies, soies que Ton retrouve dissemiiiees sur
sa moitie termimile, irri'gulierement disposees. La longueur du penis, apres sejour dans
I'alcool, est de 12i mm.
Sur les cinq echantillons que j'ai eus a ma disposition, il m'a ete imj)ossible de
decouvrir la moindre trace de males dits compUmentaires.
Cette superbe espece, resemble un peu exterieurement et seulement en ce qui concerne
le capitulum a So. squamulifermH, "Weltner, mais elle a des affinites bien plus grandes avec
So. reglum, Hoek, et Sc. Dartdnii, Hoek, cependant les seuls caracteres des jilaques
capitulaires et des ecailles pedoncvilaires sulfisent a les dilferencier nettement — ainsi que
j'ai pu m'en rendre compte par 1' etude des echantillons memes du ' Challenger.'
LocalUe. — Cotes de Cuba, par cinq cents brasses do fond. British Museum.
ESPECE NOUVELLE DE LEPADIDES. 157
PCECILASMA CARINATUM, Hoek.*
J'ai (lit, ail dcljiit dc cc travail, que j'ai rencontn'' sur Ic pedonculc dc Tun des eclian-
tillons de tScalpellnni gu/antenvi, A. Gruvel, un Poic'dasina (|ue j'ai recounu iHre P. cariiou-
tum, Hoek. Ceci m'a amenc a faire une etude complMe de cette especc que je vais
rapporter ici, afin de complotcr quolques points laisses un peu daus I'Dmbre par Hoek.
J'y cxjouterai quolques dessins, car il me jiarait ucccssaire de represeuter I'aspect des
animaux que I'ou etudie avec une scrupuleuse exactitude. Les descriptions, aussi bien
faites soient-elles, presentent toujours une elasticite d'interpretation que u'olfre pas le
dessin ; rcxactitude des unes et de I'autre constituent done une condition uecessaire et
indispensable pour permettre aux classiiicateurs, meme les plus scvupuleux, qui viennent
ensuite, de commettre des erreurs involontaires et de les exposer a nommer des es2oeces
une secoude fois.
I)k((jnose. — Cinq plaques capitulaires. Carrne considerablement elargie a sa partie
jnferieiu'e et tcrminee en un disque plein sans fourche differenciee. Dents umbonales
du scutum trcs peu fortes ; bord careno-tergal du scutum ecbancre vers son tiers iuferieur
pour recevoir les parties laterales du disque de la careue. Angle infcrieur des terga
tronquc et a bord carenal sensiblement parallMc au bord occluseiu". Pedoncule court,
atteignaut seulement le \ de la longueur du capitulum, sans ornements nets.
Mandilnxles non semblables, trois dents a gauche et quatre a droite. Appendices
caudaux uniarticules et de longueur egalant a peu pres le j ou le \ de la longueur du
pedicelle de la 6'' paire de cirrhes.
Dimensions du plus grand echantillon etudie: —
Longueur du capitulum, 14 ram. ; largeur, 4^ mm. ; maximum d'epaisseur, 4 mm.
Longueur du pedoncule, 2^ mm. ; largevir, 2 mm.
Sabitat. — Cotes de Cuba, jjar un fond de 500 brasses. Collection du British
Museum.
Capiiidum. — Le capitulum etant environ trois fois aussi long que large, il en resulte
qu'il presente une forme elancee bien plus accentuee que ne le represeute le dessin de
Hoek. II est comprime lateralement vers sa region superieure, tandis que le maximum
dcpaisseur se trouve vers le quart iuferieur environ. II est ornc de cinq plaques d'un
blanc-creme, ne laissant entre elles aucun espace libre. Elles sont recouvertes par
une trcs mince cuticule, sans autres ornements que quelques rares polls irregvilierement
dissemines. On apercoit sur les plaques des striations trcs fines mais cependant
nettes.
Terya. — Les terga sont relativement tres pevi developpes ; I'apex est termine en pointe,
le bord anterieur droit. Tangle basal tronque, le bord adjacent a la carene etant sensible-
ment parallele au bord occluseur, mais la saillie de Tangle infcrieur du tergum vers le
scutum est beaucoup moins marquee que ne le figure Hoek.
Scuta. — Tres developpes par rapport aux autres plaques, mais non divises en deux,
lis sont renfles vers le tiers iuferieur et surtout daus la region anterieurc. Lc bord
anterieux ou occluseur est fortement saillant en avant, particulierement vers son tiers
* Hoek, l{e])ort on the Cirripedia. Voyage of ' Challenger,' 1S83.
158 MONK. A. GRUVEL— ETUDE D'UKE
superieur, mais il est cependant, a pcu pres regulierement courbe. Lc bord tergal est
droit. Le bord basal trc-s etroit, a pen pn'-s comme le bord basal de la cart'ne qui lui est
adjacent et qiii forme avec le premier une ligne legerement concave inferieurement,
entouraut le sommet du pedoncule qui semble ainsi legerement detacho du ca2)itulum dans
cette partie proximale. La particularite la plus caracteristique c'est que le bord carenal
forme, dans son tiers inferieur, une concavitr destine a loger les bords lateraux du
disque inferieur de la carene.
Assez frequemment, on remarque, a la partie infero-interne des scuta, a une petite
distance du bord basal, une ligne circulaii-e, limitant avec celui-ci, une surface legere-
meut concave, un peu en saillie et portant des cannelures paralleles formant des denticu-
lations a leur contact avec le bord basal. (Fig. 15.)
A I'umbo de chaque scutum se trouve une dent interne tres peu accentuee, mais nette,
cependant.
Enfin la convcxitc des deux plaques scutales est ideutique.
Carene. — La carene est courte. Son apex depasse de peu le niveau du milieu du bord
dorsal du capitulum. La partie superieure est beaucoup plus etroite que Tinferieure, mais
le minimum de largeur se trouve un peu au-dessus du disque basal. L'arete dorsale est
saillante.
La carene se dilate inferieurement pour former deux expansions aplaties, a bord libra
arrondi et allant, cliacune, se placer dans une cavite correspondante des scuta.
Vu du cote interne, il faut se representer ce disque, comme une sorte de trapeze isocele,
plie en son milieu et se2)are du reste de la carene jjar un Ijord legerement concave et en
relief (fig. 11). Dans ses regions mediane et laterales, cette partie du disque presente
une surface ressemblant aussi a un trapeze isocele, a cotes a peu pres paralleles a ceux du
premier et entierement en saillie siir la jiremiere surface. En son milieu, elle se prolonge
inferieurement en une pointe mousse. Cette pointe, qui n'est pas figuree par Hoek,
semble cependant normale.
Exterieurement, de cbaque cote de l'arete dorsale, les lames du disque presentent
parfois des cannelures paralleles, formant sur le bord basal des denticulations. La
presence de ces cannelures concorde avec celle des memes ornements des scuta (fig. 16).
Pedoncule. — Le pedoncule ne depasse pas en longueur le \ de celle du capitulum.
II ne presente aucune trace d'annulation reguliere et la euticule qui le recouvre est
simplement ornee par quelques polls courts et irreguliers.
Bouclie. — Le Labre porte, sur son bord libre, quelques dents faibles et courtes.
Les Paljjes de la lecre superieure sont allonges, subcouiques, avec quelques soies
longues et fines sur leur bord dorsal et d'autres plus courtes et plus raides vers leur
extremit(^ libre.
Les Mandihules sont bien exactement celles decrites par Hoek, c'est Ji dire que la
mandibule ^m«c7ce ne presente que trois dents, la troisieme formant deux jiointes saillantes
entre lesquelles s'en trouvent de pins petites, tandis que la droite porte quatre dents, la
quairieme etaut egalement bilobec, avec des pointes intermediaires moins accentuees.
Ce ne serait done pas la une anomalie, comme Hoek etait en droit de le suj)poscr,
ESPECE XOUVELLE DE LEPAUIDES. 159
mais bien une atrophic acquise d(''tinitiveraent. Quant a cc qui concerne les Machoires,
je n'ai rien a ajouter ni a modifier a la description de Hock.
Enfiu les Palpes de la levre iiiferieure sont aplatis, a bords autero-superieurs arrondis
et portaut de longues soies greles et flexibles.
Cirt'hcs. — D'une faeon generale, les cirrhes sont courts et, cependaut, restent greles.
La premiere jiaire est situee a la base du mamelon buccal et se trouve legeremeut scparee
de la deuxieme.
V paire. Rames presque cgales ; I'anterieure jiossede 5) articles, la posterieure 10.
L'article basilaire est le plus developpe eu longueur et en largeur. Les soies sont
nombreuses, surtout aux extremitos et du cote interne, en avant et en arriere.
2'' pa'tro. Dans la majorite des cas, les rames sont semblables a droite et a gauche,
leur longueur cgalant a peu pres deux fois celle de la rame posterieure de la l"'
paire. Celles du meme c6te sont sensiblemcut egales avec 15 et 17 on 18 articles
allonges. Mais dans quelqvies cas, il se produit du cote gauche une atrophic
singuliere qui porte sur la rame posterieure.
Tandis, en efifet, que la rame anterieure est normale, la posterieure est tres grele et j)lus
courte meme que celles de la premiere j^aire de cirrhes.
Les aiitres paires de cirrhes sont a j^eu pres identiques a la deuxieme paire normale,
avec de 15 a 18 articles.
Appendices caudimx. — lis sont cylindriques, imiarticules et termines par un bouquet
de soies courtes. Leur longueur atteint environ le -]- ou le \ de celle du pedicelle de la
6'' paire de cirrhes.
JPciiis. — Le penis est long et grele ; son extremite atteint celle des cirrhes de la
6* paire apres sejour dans Talcool. II ne presente aucune trace d'annulation propre-
ment dite et la cuticule qui le recouvre est simplement ornee de quelques polls,
disseminc5s sur sa perij)hcrie et se reunissant a son extremite en une touife assez rio-ide.
Distribution (jeograplnriue. — Pendant la campagne du ' Challenger ' nous dit Hoek
cette cspece a ete trouvee deux fois. La premiere fois, ce fut un tout petit echantillon,
par une profondeur de 390 brasses a Colubra Island (West Indies) ; la seconde fois, a
rile de 1' Ascension (Ocean Atlantique), par 420 ])rasses de fond, cinq echautillous furent
recoltes fixes sur de coraux. L'echantillou trouve sur les pedoucules de Scalpellum
giganteum vient des c6tes de Cuba. Enfin il existe an British Museum quelques beaux
exemplaircs de cette meme espece.
Je desirerais, en terminant cette courte etude, dire un mot de la collection du ' Chal-
lenger' que j'ai pu examiner tout a loisir pendant mon sejour au British Museum aux
vacances dernieres.
Avant d'avancer plus loin dans la lougue etude du groupe des Cirrhipedes que
j'ai enterprise et que je voudrais tacher de mener a bien, j ai pense qu'un peleriuage au
lieu meme oil lillustrc Darwin fit les briUantes et patientes recherches qui Fainenc'rent
a ecrire ses beaux volumes de la ' Monographic des Cirrhipedes," s"imposait tout d'abord.
160 MONS. A. GErVEL— ETUDE D'UNE
Je tenais a voir de pres sa collection conservce aii British Museum, ainsi que celle de
Hoek provenant des campagnes du ' Challenger.'
Je pensais, avec raison, que si bien des t'ois a Taide d'une description et de dessins,
on pent arriver a se faire une idee nette dune espece, il se prescntait des cas, pour des
especes trcs voisines, par exemple, ou la comparaison des types memes s'impose et
comme ces types ne sortent pas, generalement, des collections qui ont I'honneur de les
posseder, une visite au British Museum devenait indispensable.
Cette yisite, je I'ai faite et je puis affirmer, que grace a la haute bieuveillance de
M. le Professeur Bay Lankester, Directeur, ainsi qu'a I'amabilite de tons les Professeurs
et Assistants avec lesquels j'ai eu des relations, en particulier avec M. J. Bell, que j'ai
derange bien des fois de son travail, je n'ai pas perdu mon temps. Tout a ete mis de
la fapon la plus liberale a ma disposition, et je ne saurais trop les en remercier ici.
J'ai pu me rendre compte que j'etais dans le vrai en pensant qu'il fallait voir les
echantillons cux-memes, avant de pouvoir decider en connaissance de cause.
Meme quand Vauteuvfaii lui-meme ses dessins, il pent conimettrc des erreurs, s'il ne
les commet pas lui-meme, elles proviennent du graveur ou meme du tirage. Voila pour-
quoi probablement, certains types de Hoek sont le plus generalement assez diificilement
reconnaissables si Ion s'en rapporte aux dessins qu'il en a donnes. Comtne on ne pent
pas toujours voir les types, et que les descriptions sont parfois un pen elastiques, meme
les mieux faites, il en resulte que Ion pent dans ces cas nommer deux fois une espece,
avec la meilleure foi du monde.
En ce qui concerne le genre Scalpellum, beaucoup d'especes, particulicrement celles a
plaques imparfaiteuient calcifiees, ne montrent pas, dans les figures, la limite nette des
plaques cependant tres visible sur les echantillons.
Cela n'est nuUement une critique, mais une simple constatation. Ce sont la surtout
des defauts d'edition, mais il n'en est pas moins vrai que cela pent avoir des consequences
assez serieuses et, toutes les fois que j'en aurai I'occasion, je tacherai de rectifier ces
points obscurs des dessins, en me contraignaut, pour etre plus exact, a dessiaer moi-
meme a la chambre claire quand cela sera possible ou d'apres des documents photo-
graphiques, dans les autres cas.
Je crois meme que, toutes les fois que la dimension des echantillons le permet, il est
bon de remplacer le dessin entierement j)ar la photographic, comme je I'avais, par
exemple, pour la figure 1, representant un groupie de Sca^yellmn giganteum., mais des
difl&cultes d'exccutions m'ont oblige a remplacer la photographic jmr de simples dessins.
Ce sont la, malheureusement, des dilficultes qui se presentent souvent dans I'executions
des planches et centre lesquelles la meilleure volonte des auteurs est impuissante !
C'est la \\n fait evident, mais infiniment regrettable, meme en se placant au point de vue
exclusivement scientifique !
ESPECE NOUVELLE DE LEPADIDES. lOl
EXPLICATION DE LA PLANCilE 17.
[Tous ces dcssins ont cte fails par I'autcur, soit d'aprcs des photographies, soit a la chambre clairc]
Pig. 1. ScALPELi-UM GiGANTEUM daiis (liffcfciites positions.
2. Le memo, apres qu'uue partie do la chitiiie su))erfici(;lle a etc cnlcvee pour laisser voir la forme
(Ics plaques capitulaircs. Le pedonculc est simplement represcnte par son eoiitour. /. ter-
gum ; sc, scutum; ca, car6ue ; ca.la., plaque eareno-lateralc ; r.la, plaque rostro-laterale ;
s./a, plaque laterale supcrieure; i.la, plaijue itifra-laterale ; j/e, piHlom-ule.
3. Palpe de la Icvre supcrieure (gauche).
4. Mandibule gauche.
5. Machoire droitc.
6. Palpc de la levrc inferieure (droite).
/. Mamelou buccal vu de pi'otil. lab, labrc : p. lab, palpe di; la Icvre supt^rleure ; t/id, mandibule •
ma, machoire.
8. Mamelou buccal vu par la partie supcrieure (memos lettres).
8'. Appendice caudal droit.
9. PciiCiLAsMA CARiNATUM, Hock, VU dc protil, (lesxin a la ciniiabre rlalre.
10. TjC memo, vu par la face dorsale.
11. Maudibulos, («) gauche, {b) droite.
12. Machoire gauche.
13. Vue de la partie iufero-anterieurc de I'dcil/i.snio rari/iatum.
11. Vue iiifi'ro-iiiterue de la earene.
15. Scutum gauche, vue iufero-interne : (//, dout umboiiale interne.
1(). Carene, vue iufero-exterieure moutrant le.-; eaunelures des parties iat^rales du distjue.
17. Cellules pigmentaires de I'epithelium de Sc. (/it/unteum.
SECOND SERIES. — ZOOLOGY, VOL. VIII. 25
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[VOL. VIII. PART 6.
THE
TRANSACTIONS
OF
THE LINNEAN SOCIETY OF LONDON.
ON THE AFFINITIES OF ^LUROPLTS MELANOLEUCTS, A. m.-ED wards.
BY
E. RAY LANKESTER, M.A., LL.I)., E.R.S., E.L.S.,
DiKECTOB OF THE NaTUKAL HlSTORr DePAETMENTS OF TUB BkIXISH MuSEUM.
^LONDON:
PRINTKl) FOK I'HE I.INNKAN SOCIETY
BY lAVr.oU ANIi KHAN( IM, UKl) I.ION COURT. KI.EKT STREET.
SOLD AT THE SOCIETY'S A PARI'.M KNI S. BUKIJNGTON-HODSE. i'lCCAmi.LY, W.,
AND BY LONGMANS, GItKEN, ANP CO., I'ATERNOSTEll-KOW.
September 1901.
AUG 30
[ 163 ]
\L. On Ihe Affinities of ^Elui'Ojius uielanoleucus, A. Milne-Edwards. By E. Ray
Lankester, M.A., LL.D., F.B.S., F.L.S., Director of the Natural Ristory
Departments of the British Museum.
(Plates 18-20.)
Read 21st Februiiry. 1901.
oIX years ago my friend the late Professor Alphouse Milne-Edwards, Director of the
Jardin des Plantes and Museum of Natural History in Paris, kindly presented to the
Oxford University Museum a cast of the skull of the reraarkabla bear-like animal
brought home from Tibet by Pere David, and named by him TJrsus ineluiioleucus.
M. Milne-Kdwards in 1875 published a description of this animal and very distinctly stated
the conclusion that it must occupy a position intermediate between the Bears {(Jrsinte)
and the Panda {^Itirus). He formed for it the genus JEluropus*. But subse-
quently the late Sir W. Flower and Mr. Lydc^kker in their book, ' Mammals, Living and
Extinct,' 1S91. whilst dividing the section Arctoidea of the Carnivora into three families,
* Milne-Kdwards ' wrote as follows : — " L'ensemble de faits que je vieiis de passer en rovue, prouvo (|ue TAilurope
ne peut-etre rapporte a aucun des types geueriquos preeedemmeiit conniis. 11 appartient indubitablement a la
famille des Cariiassiers arotoides, dont les Ours sont !es priiicipau.x representants, et il ressemble beaueoup a ces
animaux ; raais ii tient encore plus peut-etre des Pandas, et il prcsentc un singulier melange de earacteres osteolo-
giqucs. Ainsi, par le mode d'articulation de la maohoire infcTieure, renorme doveloppement des arcades zygoma-
tiques, il ressemble aux Felins les plus robustes, et quebpies naluralistes le eomparent a THyerje ; niais la couforma-
lion de ses dents machelieres indique que c'est en realile un animal moins carnivore que ne le eont les Ours ....
Par la disposition de la couronne, la penultieme molaire a beaueoup d'analogie avec les molaires de divers Pachyderines
fossiles, notamment des Chwropoiiimus pcirisiensis .... Neanmoins e'est entre les Ours el les Pandas que rAiluroj)e
doit prendre place dans nos classifications methodiques, et la division qui le renferme ne parait avoir une vaieur
zoologique plus considerable que celle de la plupart des genres dont se compose I'ordre des Carnassiers.''
From this it appears that Professor Milne-Edwards regarded ^luropus as intermediate between the Bears and the
Panda, and therefore probably the representative of a distinct family, although this is not definitely stated. His
mention of the Ungulate resemblances of the molar teeth is significant, seeing that this is a feature which has been
often noticed as characteristic of those of JUlurus. It may be added that the resemblance of the lower jaw of
^luropus to that oi'.l-jluras is s])ecially noticed by Professor ililno-Edwards.
Writing at a laU.T date, the late Sir W. II. Flower', who regarded »/i7((r".v as the representative of a family
connecting the Procyonidae with the Ursidic, assigned JSIuropus to the last-named family, from the other members of
which it differs by the absence of an alisphenoid canal ^ The genus is described as " an interesting anneclant
' Recherches Hist. Nat. .\Iamm. p. 335 (1808-7")).
■ .'Vrticle " Mammalia," Enci/clopcedut Britannica, !Hh ed. vol. xv. p. 441 (1883).
° On the page cited above, in the 8th line from the bottom of the first column, the word " An " should be substituted
for " No."
SECOND SERIES. — ZOOLOGY, VOL. VIII. 26
1G4 DK. E. EAY LAXKESTER ON TJIE
viz. Ursidse, Procyonidse, and Mastelidse, placed ^Elurojnis in the Ursidte and ^lurus
in the Procyonidre.
]\Iy observations oa the cast of the skull which was in my charge at Oxford led me to
the conclusion that too much weight had been assigned by Flower and Lydekker to the
presence in ^luropns of the thii-d lower molar, which is lost in jElurus and T?rocyon
whilst retained in the Bears. The form of the teeth, especially of the upper fourth pre-
molar (so-called " caruassial ") and of the true molars, resemble 1 that of the same teeth
in ^lurus and Frori/on rather than that of the same teeth in ZTrtms. The teeth in ques-
tion are unlike those of any other living Carnivore, recalling those of some of the extinct
Ungulata in breadth, squareness, and tubereulation. The general form of my cast of the
skull oW'Elnropus differed greatly from that of any Bear. In all Bears the skull is much
longer and narrower both in its facial and cranial regions than is the skull of JEluropus.
Similarly the lower jaw differed greatly ia shape from that of the Bears. On the other
hand, the relative shortness and breadth of the skull of ^^Eluropm are closely repeated in
jFJurus; whilst the shape of the loAver jaw in the two forms I found to be singularly
coincident, not only as to outline and cui'vature of the jaw and coronoid process, but
especially in regard to the form of the condyles.
When I entered upon my duties at the Natural History Museum in London, in 1898,
1 looked up the material relating to yEluropus in the Collection, and was much pleased to
find that, in addition to a fine skull and moiuited skin, the Museum possessed certain
of the limb-bones and incomplete feet of JEluropus, obtained in 1896.
On comparing these bones — viz., Humerus, Radius and Ulna, Carpus and Metacarpus,
Temur, Tibia, Fibula, Tarsus and Metatarsals — with those of TJrstis, yElurus, and Procyon,
I found that my previous impression was abundnntly confirmed, and that in very impor-
tant and distinctive points jEluropus agrees with .EUirm and Procyou (more closely
with the former) and differs widely from Ursus.
I found a strongly marked entepicondylca- i-idf/e and foramen in the humerus of
jpluropus (PI. 20. figs. 1, 1 a). This does not exist in the Ursidse, but occurs in ^Elurus
and Procyon (PL 20. figs. 2, 2 a, 3).
Even more remarkable than this were the enormous accessory or extra-carpal bones
of the carpus. The iilnar or postaxial extra-carpal (the pisiform of man) is of very
laro-e size — larger than in any Arctoid — and springs obliquely backwards and outwards
CPl. 20. fig. ^,pi.). But the really extraordinary bone is the preaxial extra-carpal or
" prsepoUex " of some w riters, which is articulated to the scapho-lunar and to the preaxial
metacarpal, and has the size and position of a sixth metacarpal bone (PI. 20. fig. 8, r.s.).
form, connecting the true Bears with ^Slants and with several extinct genera. Dentition i. jj, c. \, p. ^-, m. f ; total
40. Premolars increasing in size from first to last, and two-rooted except the first. First upper molar with quadrate
crown, broader than long. .Second larger than the first. Cranium with zygomatic arches and sagittal crest immensel)-
developed, and ascending ramus of mandible very high, giving greater spaces for attachments of temporal muscle than
in any other existing member of the order. Bony palate not extending behind the last molar tooth Feet
bear-like, but soles more hairy, and perhaps less completely plantigrade. Fur long and thick. Tail very short. One
extremely rare species, A<^. mehi noltucus It is of the size of u small brown bear, of a white colour, with ears,
spots round the eyes, and limbs black."'
AFFIXITIES OF .ELUROPUS MELANOLEUCUS. 165
This bone has uo jjai-allcl Tor size and shape iu the Carnivora. A radial extra-carpal of
large size has been described in some Rod(mtia and Insectivora, but in the Carnivora as
iiitlierto described it is always small, and especially small iu the lar^'er forms fsee
Bardeleben, Troc. Zool. Soc. 1889, p. 259).
When we compare the carpus of ^luropus with that of the Bears, we find that there
is only a very minute bone in UrsHs corresponding; to the large radial extra-carpal of
^Eiurojms. In JElnnts, on the other hand, a radial exti-a-carpal of relatively large size
— but not so large proportionately as that of ^Eluropns — is present. In Trocijon the
radial extra-carpal appears not to be developed or to be minute.
In regard to this bone, then, ^Inropus differs markedly from Ursiis and agrees, though
not closely, with jEUwus. We are not acquainted with the mechanical requirements in
the use and movements of the fore-foot which its jji-esence subserves.
A close comparison of the bones of the forearm of Mluropus with the corresponding
bones of Ursus, ^Muriis, and Proci/ou, as also of the bones of the thigh and leg of the same
genera, confirms in many minute particulars the proposition that Mluropus differs from
Ursus, awdi agrees with ^lurns awA Pi-ocijon, and, moreover, agrees more closely with the
former than witii the latter of these two (see PL 20. figs. 5, 6, 7).
The comparison of the femora is very convincing in this respect, details of the form and
curvature of articular faces and of the devcloj^ment of the trochanters showing an amount
of minute coincidence as between ^THuropm and ^Nuriis which can only be interpreted
as the indication of close genetic affinity.
Having satisfied myself on these points, I icquested ^Ir. Lydekker, who had not pre-
viously examined the limb-bones of ^Slnropiis, to go over the specimens and to give me
the result of his observations. He entirely accepted my conclusion that -Eluropns must
be removed from association with the Bears, and associated with ^Elnnis and Froci/o/i.
It has accordingly Ijeen removed to that position in the exhibition gallery of the
Museum, and is no longer to be spoken of as "the Parti-c(jloured Bear," but as "the
Great Panda."
It seems to me that ^-Elurojms is probably an earlier form than either ^'Eluriis or
Procyoii, and it would not be surprising were affinities between it and extinct forms
discovered. In the meanwhile, the division of the Arctoidea into three families, the
Ursidae, Procyonidae, and MustelidiB, may be maintained, Avith the assignment of
^Eluropus to the Procyonidse alongside of ^Eltu-m and Pmcyon.
One peculiarity of the l)ones of JEluropus should, I think, be placed on record. I refer
to their great specific gravity. Not only are the bones (such as the lower jaAV and skull-
bones) powerfully built and of great actual thickness, but their weight is extraordinary.
I do not know any l)ones of terrestrial Mammalia of similarly high specific gravity.
Mr. Lydekker has kindly draAvn up for me a minute descriptive accoiint of IJic
skull and limb-bones of ^Eluropus and the details of their comparison with those of Vvsm,
JElunis, and Procyon. This account follows.
The illustrations of the skulls and limb-bones given in the Plates have been prepared
under my supervision by ^Ir. Gron\^ald at the Natural History Museum.
26*
166
DR. E. BAY LANKESTER ON THE
Detailed Description of the Skull and Limb-bones.
By R. Lydekker, F.B.S.
From jElurus (in which the formula of the clieek-teeth is p. f , m. f , and the total
number of the dentition 38) JEluropus differs markedly, not only by the absence of an
alisphenoid canal, but likewise by the presence of a third lower true molar ; in the latter
respect resembling the Ursidse, and more especially the extinct genus Hyanarctm, in
which the upper molars have short and broad crowns, and the last lower molar is much
less elongated than in the genus Ursiis, being, indeed, considerably shorter than the tooth
in front of it. The carnassials of Hycenarctus likewise come uuich closer to those of
^■■Eluropus than to the corresponding teeth of TJrsiis ; the upper one of the former genus
having a trilobed blade, owing to the development of a style in front of the protocone.
Its tubercular portion also carries a small hypocoue, which, however, is coalesced with
-at-
^.ci
-^
B
D
cl
Left upper carnnssial teeth of Raccoon (A), Himala_yan Long-tailed Panda (B), Great Panda (C), Cave-Bear (D),
and Hij(TnarctHS (E). «, protocone ; 6, paracone ; c, metacone ; (/, hypocone ; c, anterior stylo.
the protocone (fig. E). In ^Bluropus, on the other hand, the upper carnassial (which,
contrary to the statement of Sir William Flower, lias an inner root) has a large and
distinct hypocone situated behind the protocone, from which it is completely separate ;
this being a character it possesses in common with ^Bluriis and the American Procyonidae,
but found in no other Caruivora. In the Raccoon and its American relatives the
anterior style of this tooth is, however, much reduced, and as the metacone is also
relatively small, the blade, although still trilobed, is mainly formed by the paracone.
The lower carnassials of ^Inropus and ^Blurus are likewise constructed ou the same
general plan, being remarkable for the large size of the metaconid and entoconid. In
this feature they are essentially raccoon-like *, tlie main difference being that the
entoconid is double — distinctly so in jElurus, but rather less clearly in jElnropus. In all
* These features are well shown in the series of sectorial teeth of Carnivora exhibited on one of the pillars of the
Lower Mammal Gallery of the Natural History Museum, from which figs. A tj E were taken.
AFFINITIES OF ^LUROPUS MELANOLEUCUS. 167
three forms the metaconid is placed immediately on the inner side of the protoconid,
instead of partly behind it.
We have, thenrCorc, tlie important fact that ^Elttrm and ^EluropHs resemhh' the
American Procyonidte, and thereby differ from all otlier Carnivora, in the presence of two
lobes, or cusps, on the inner or tubercular jjortion of the upper carnassial. And since
^Elurus has been referred by Mr. Blantord to tiie Proeyonida;, there is a pr/md facie
presumption that yEliiropiis, in spite of its reteiitiou of the third lower molar, is likewise
a member of the saint; family.
'rurniiiii' for cotilirinatory evidence first to the external features of the animal, it may
be noticed that the type of coloration is essentially unlike that obtjiining in the Ursidai,
although a superficial approximation is presented by some specinu>ns of Ursus pruinosux.
On the other hand, if tlie coloration of ^Eluropus be compared with tliat o[\EJ/nnift, there
are certain similarities in the general plan. In both, for example, the limbs are black;
and tlie l)lack ring round the eye of the former corresjjonds to iIh; chest nut streak runnin"
through the same organ in the latter. The fur, too, of Ixtth animals is likewise very
similar in character, being comparatively sleek and smooth, and unlike the shaggy coat
of so many of the Bvav tribe. Of more importance is the form and expression of the face,
which in both animals is short, broad, and cat-like, entirely lacking the prolonged snout
characteristic ot the Bears. The ears, too, are strikingly alike in both genera ; and the tail,
although still very sliort, is longer in ^Eltiropus tlian in any Bear. The points in which
the feet of the former differ from those of the latter arc referred to in the above extract
from Sir William Flower's description of the genus.
As regards the features in which the skull of ^Elaropus rcsemldes that of ^Elurm and
differs from those of the Bears, mention had been made by Professor ^lilne-Edwards, and
before him by Professor P. Gervais, of some of the most important of these.
Botli agree in their relative shortness, wide zygomatic arches, convex profile, and
curvature of the inferior margin of the lower jaw, as well as in the great elevation, for-
ward inclination, and hook-like shape of the coronoid jnocess of the latter. The en(jrmous
development of the sagittal crest in ^Elnropus causes the convexity of the profile to be
more marked than in ^Elurus. The ^Eliiropiin skull differs from that of ^Eliirun in the
absence of postorliital processes to the frontals, and likewise by tlie non-prolongation of
the palate behind tlie last molar. In the presence of such, processes and the jirolonga-
tion of the palate, ^Eluriis is the more bear-like of the two, although it has no post-
orbital process on the zygoma. On the other hand, ^Eluropus approximates to the
Raccoon in the absence of postorbitiil processes on the frontals, although tlu; Baccoon
retains the one on the zygoma. Although by no means so short and wide, the skull of the
Raccoon, especially in its profile, is more like those of yElurm and ^Elnmpus than is a
Bear's skull A point of resemblance between ^Eluropus and Vrsua is to be found in the
circumstance that the maxillary foramen opens on the side of the skull well in advance
of the zygomatic root, whereas in ^^(f«?-;w and l'roc?/on it perforates the zygoma itself.
As Professor Milne- Edwards has pointed out that in .Eiuropus a depression in the bone
indicates the position of the alisjihenoid canal, the absence of the canal itself cannot be a
feature of any great morjiliological importance.
108 I^K- E. RAT LANKESTER ON THE
In addition to the great pe(;uliai'ity of their general form, to ^vhich allusion has been
already made, tlie jaws of both ^Elurus and ^Fl/iro/j/fsdiSi^v from the Ursine mandible by
the very reraarkal)le conformation of the condyle.
In the condyle of Ursus the articular surface forms a regular semicylinder, with its
inner margin approximately at right angles to the hmg axis. On the other hand, in the
two genera under special consideration the articular surface is like a riband wound
obliquely on a cylinder of which the ends have been cut off along the margins of the
riband. Consequently the inner margin of tlie articular surface forms an angle of about
45° with the axis of the condyle. No marked ajiproximation to this very peculiar type of
condyle is exhibited by the Raccoon. In ^El /frits the mandibular " angle " is fairly well
developed and nearly continuous with the long axis of the jaw, but in ^Eluropus it is
very small and inflected.
As regards the appendicular skeleton, concerning which no description has hitherto
])een published, the scapula in the Museum specimen of ^Eluropus being wanting,
comparison may be commenced with the humerus, which has the proximal epiphysis still
separate from the shaft, thus indicating that the animal was not fully adult at the time
of its death. Compared with the corresponding bone of Ursas (PI. 20. fig. 4), as
well as with that of the extinct Hycenarctiis, tlie humerus of ^Eliwopus (PL 20. fig. 1) is
broadly distinguished by the presence of an entepicondylar foramen, in which respect it
resembles ^Eliirus and the American Procyonidie. But this is by no means the only point of
difference from the one type and aj)proximation to the other. To begin with, the head
of the humerus in ^Elwojms, ^Elurus, and Vrocyou is much larger in j)roportion to the
rest of the bone than is the case in Ursus. Moreover, in the two genera just named the
inner margin of the deltoid crest forms a nearly straight line, and the j^i'ofile of the front
surface of the bone makes a slight but regular curve.
In Ursus, on the other hand, the inner margin of the deltoid crest is sinuous, and the
profile of the bone at the inferior termination of that crest (which forms a marked
rugosity) is angulated. In these respects the Frocyon humerus is nearer to the ^Elurus
than to the Ursus type. Inferiorly tlie humeri of ^Elttropns, ^Eliiriis, and Procyon have
a marked general resemblance, especially shown by the flatness and expansion, the shal-
lowness and width of the glenoid fossa, and the large size and tliitiuess of the entepicondyle,
which is most expanded in ^Eluropits. In U'rs/is the whole extremity is much less
expanded, the entepicondyle mucli more thickened from back to front, and the glenoid
fossa much deeper.
As regards the second segment of the arm, the most striking feature in both ^ElurojJii'S
and ^Elurus is its extreme relative shortness, the ulna being considerably inferior in
length to the humerus, whereas in both the other two genera it is longer. As regards
form, the most noticeable feature in the ulna of ^Eluropus and ^Elurus is the much
greater elevation of the oleci-anon above the summit of the greater sigmoid notch when
compared with the corresponding bone of Ursus. Although the elevation in Frocyon is
less than in the two first-named genera, it is great(!r than in the third. The radius is,
however, perhaps a more important bone in respect to the differences separating Ursus
from the two genera specially under consideration. In the former (PI. 20. fig. 7), in
AFFINITIES OF .^LUROPUS MELANOLEUCUS. 169
addition to its supoi-ior relative length, tliis l)one is specially distinguished by the compai-a-
tive narrowness of the distal extremity and the shar]) point formed by the preaxial
malleolus. In ^Inropus, on the other hand, tlie distal extremity of the radius (W. 20.
tig. 8) is much more expanded above the articuh^r surface, and the preaxial malleolus
does not descend to nearly the same degree. Consequently the profile of this extremity is
quite different from that of the corresponding bone in Ursns ; while (as may be seen from
the figures) there arc equally important differences in the contour of the distal articular
surfaces of the two bones. In all the aliove respects the radius of ^Eliirus (PI. 20. fig. (5)
is es.sentially similar to tiiat oWEZ/iroii/is. Although differing to a considerable degree,
the radius of Proci/oii comes nearer to the type of the two latter genera.
Passing on to the manus (PI. 20. fig. 8 ), the most noticeable feature in ^l^luropm is
the development of a very large preaxial extra-carpal (r.s.) in addition to the ordinary ulnar
sesamoid, or pisiform [pi). The former is wedged in between the scaphoid process of the
scaplio-lunar and the first metacarpal, so as to articulate largely witli Ixjtli bones. A
precisely similar arrangement obtains in ^EIki'ks, alihough the radial extra-carpal is of
very considerably smaller dimensions in that genus. (_)n the other hand, in the carpus of
Vrsus the radial extra-carpal * is reduced to a small nodule of bone articulating exclu-
sively with the scaphn-lunar. In the Raccoons, if such a preaxial bone is developed at
all, it must likewise be very minute. The huge size of this bone in ^Ehiropus, which
simulates a sixth metacarpal, is unique.
Both in ^ISluropm and ^Elnrns the metacarpals are relatively short, this abbreviation
being carried to tbe greatest extent in the former. In regard to this feature Froci/on
departs further from the type of the two genera under consideration than does I'rsns.
Einally, it may be mentioned that the terminal phalanges of ^Elnropns (PI. 20. fig. 12)
and ^'Elnrus differ from one another only in point of size, but are quite unlike the corre-
sponding bones of both Proci/ou and Urs/is. Tlieir most striking common features are
their shortness, widtii, and thinness, coupled with the great relative development of the
basal protecting sheath.
The femur of ^Eluropus (PL 20. fig. !)) is characterized by its shortness and flatness, as
well as by the small antero-posterior extent and nearly symmetrical lorm of the trochlear
surface for the patella. On the posterior aspect (fig. !) a) the digital fossa is very deep, and
the area between tlu> head and the two trochanters remarkable for its width and flatness.
Owing to this, the two trochanters are widely separated from one another. Another
feature is the relatively small size of the lesser trochanter, which is situated on the pre-
axial bordei- of the shaft, so as to be largely visible in a front view. Although its shaft
is proportionally longer and more slender, the femur of ^Eltiriis presents tJie essential
features of the corresponding bone of ^Eluroptis ; and in the femur oiProci/on, owin"' to
relatively greater shortness and width, the resemblances to the latter are even more
strongly marked, although less is seen of the lesser trochanter in a front \iew. On the
other hand, the Ursine femur (PL 20. fig. 10) is totally unlike that of the genus undt^r
consideration. Not only is the shalt much longer and more nearly cylindrical, but the
ti'ochlear surface for the patella is more elongated, and has one lateral border C')nsideral)lv
* See Flower, 'Osteology of the Manniialia,' i>. 2^7, fig. 93 (1SS3).
170 T)R. E. KAT LANKESTER ON THE
longer tlian the other. Much more striking are the differences at the proximal end oC
the hinder surface. Here the area between the head and the two trochanters is narrow
and uneven ; the trochanters themselves are closely approximated and connected with one
another by an oblique ridge ; while the lesser trochanter is relatively larger and situated
on the posterior surface of the shaft considerubly behind its preaxial border, so as to be
almost entirely concealed w^hen the bone is viewed from in front.
In a distal view of the femora of ^J^luropiis, ^Elnrus, and Procyon the two condyles
have their posterior borders nearly in the same vertical plane, whereas in Vrsus the innc^r
condyle projects nearly half an inch behind the plane of the posterior border of the outer
one.
Tiie evidence of this bone is therefore very strongly in favour of an intimate connec-
tion between the three genei'a first named and their wide differences from the fourth.
The tibia and fibula of u3^lunis correspond with the femur in relative shortness and
stoutness. In the tibia the ligament by which the patella is attached springs from a
separate ossification of the proximal epiphysis, as is the case in Bears and many other
mammals. Perhaps, in addition to its general proportions, the most distinctive feature
of this bone is the extreme shallowness of the groove on its distal surface for articulation
with the astragalus. In this respect it closely accords with the larger and moi-e slender
tibia o^ ^-Elur^is, but differs widely from Vrsus. In Procyon, the tibia is to some extent
intermediate between these two types. As regards the fibula, it will suffice to say that
in both ^Eluropns and ^lElurits it is characterized by the great expansion of its tw'o
extremities.
In correlation with the shallow "rooves on the distal extremitv of the tibia, the
astragalus of ^Eluropns is characterized by the slight elevation of the ridges on its tibial
surface, while above that surface it lacks the proiection which is so conspicuous in the
corresponding bone of the Bear. Another characteristic feature of the tarsus of the genus
under consideration is theantero-posterior elongation of the cuboid — a feature also shared
by the tarsus of ^Elurus, but not by that of Ursns, in which the cuboid is transversely
elongated. To the inner side of the navicular is articiilated in both ^Eluropus and ^Slurus
a very large tibial sesamoid, which is of a pointed form and extends down by the side of
the entocuneiform. I cannot find evidence of the existence of this tibial sesamoid in
either Procyon or Vrsus. Both ^Elurus and Procyon difftr from ^Eluropus in the greater
length and slenderness of the metatarsals, this feature, which is doubtless a fimctional
one, being most developed in Procyon. It may be added that the cuboid of the latter
genus is of the elongated .^jEJluropus type.
Prom the loregoing survey it will be evident that, so far as its dentition and osteology
are concerned, ^Uluropus is very closely allied to ^Elurus, the resemblance of some
of the bones of the two genera being so close as to suggest specific rather than generic
distincticm. Prom Vrsus the differences presented by the bones of both genera are
very marked ituleed. As regards the American representatives of the Procyonidie, the
dental and osteological resemblances presented by the tw^o genera in question are very
noticeable, although in some, but by no means all, respects these are greater in ^Murtis
than in ^Eluropus. In its retention of the last lower molar the latter serves, indeed, to
connect the former with Vrsus, Uycenarcius. Amphicyon, etc.
AFFINITIES OF ^LUROPUS MBLANOLEUCUS.
171
In spite of this difference in their dental formula, it appears reasonahle that if ^FJurus
he included in the Procyonidae, ^Eluropiis should likewise find a place in the same family.
The two may indeed he regarded as the representatives of a subfamily group of the
Procyonidae — the JElurince. And the following characters Avill serve to differentiate botli
the family from other Carnivora, and tlu; Old World subfamily from its American
representatives : —
Procyonid.'E. — True molars 3, except in yEIuro/ms, where they are |; tubercuhar, or inner, portion of
upper carnassial formed of two tlistiuct cnsps, the protoconc and hypocouc. Humerus usually
with an entcpicondylar foramen. Alisphenoid canal wanting, except in /Elurus. Feet planti-
grade ; tail generally long and ringed.
a. P;-oryo;H'«<e. — Blade of upper carnassial mostly formed l)y the paracone. No alisphenoid canal.
Two lower molars. Tail long. Steno's fissure present.
b. jElurincE. — Blade of upper carnassial formed to a great extent by the anterior style and meta-
cone, which approximates in size to the paracone, and thus renders the blade completely
trilobed. Steno's fissure absent.
1. Tail long; two lower molars, an alisphenoid canal, palate prolonged behuid m.Z, a
postorbital process, radial extra-carpal moderate ^Elunts.
2. Tail rudimentary ; three lower molars, no alisphenoid canal, palate not prolonged beyond m. 3,
no postorbital process, radial extra-car| al very large yEluropus.
In the retention of the third lower molar JEluropi^s is more generalized than JElurus,
but in its rudimentary tail and the obliteration of the alisphenoid canal it displays greater
specialization. The retention in both genera of the entcpicondylar foramen to the
humerus indicates that they must have branched off from the ancestral stock at an earlier
date than Uyienarctus and Vrsus. These relations may be roughly indicated in the
accompanying diagram : —
Canis.
Procyon.
JElurtis.
^
JElurojms.
Amphicyon.
SECOND SERIES. — ZOOLOGY, VOL. VIII.
27
172 AFFINITIES OF ^LUEOPUS MELANOLEUCUS.
EXPLANATION OP THE PLATES.
Plate 18.
Fig. 1. Palatal aspect o{ CTanmxn o{ yElur Opus melanoleuctis ; nat. size.
Fig. 2. heit su\e ot palate oi ^lur us ful(/ens; nat. size.
Fig. 3. Left side of palate of Procyon cancrivorus ; nat. size.
Fig. 4. Palate of Ursus arctus; ^ nat. size.
2). 4, carnassial ; pal., hinder border of hard palate ; pt., pterygoid ; c/L, glenoid cavity ;
/, tympanic; eu., eustachian canal; /.o., foramen ovale; /./a, foramen lacerum posterius ;
cl.f., condylar foramen ; x, Steno's fissure, the aperture of Steno's duct.
Plate 19.
Fig. J. Upper surface of right half of cranium of /Ehiropus melanoleucus ; nat. size.
Fig. 2. Upper surface of right half of cranium of yElurus fulc/eus ; nat. size.
Fig. 3. Upper surface of right half of cranium of Ursus arctus ; al)out \ nat. size.
Fig. 4. Palatal aspect of right ramus of mandible of yS^wro/JMs 7He/a//o/e«CM* ; nat. size.
Fig. 4 a. Outer aspect of same ; about f nat. size.
Figs. 5, 5 a. Palatal and outer aspects of right ramus of mandible of ^lilurus fulgens ; nat. size.
Fig. 6. Palatal aspect of right ramus of mandible of P/-oc7/o;( cflHcr/worMS; nat. size.
Figs. 7,7a. Palatal and outer aspects of right ramus of mandible of Ursus arctus; \ and \ nat. size.
»«. 1, carnassial, na., nasal; p.o., postorbital process of frontal; cr., coronoid process of
mandible ; b.c, base of corouoid process ; c, condyle of do. ; d, angular process of do. ;
i.e., internal condylar process.
Plate 20.
Figs. 1, 1 a, 1 b. Palmar, preaxial, and distal aspects of right humerus of ^luropus melanoleucus ;
S nat. size.
Figs. 2, 2 a, 2 b. Corresponding aspects of right humerus oi ^^lurus /uh/eu.i ; nat. size.
Fig. 3. Distal aspect of right humerus of Procyon cancrivorus ; nat. size.
Figs. 4, 4 e, 4 b. Palmar, preaxial, and distal aspects of right humerus of Ursus arctus ; § nat. size.
Fig. 5. Dorsal aspect of right radius of /Eluropns melanoleucus ; f. uat. size.
Fig. 6. Dorsal aspect of right radius of yElurus fulgens ; nat. size.
Fig. 7. Dorsal aspect of right radius of Ursus arctus ; "q nat. size.
Fig. 8. Dorsal aspect of right carpus and metacarpus of JEluropus melanoleucus ; uat. size.
Figs. 9, 9 a. Dorsal and ventral aspects of right femur of ^luropus melanoleucus; nat. size.
Figs. 10, 10 tt. Corresponding aspects of right femur of Ursus arctus; nat. size.
Fig. 11. Dorsal aspect of right tarsus and metatarsus of ^luropus melanoleucus ; uat. size.
Fig. 12. A terminal phalangeal of zEluropus melanoleucus.
d, deltoid ridge ; ewi.c, eutepicoudyle ; e«^/., entepicondylar foramen; //•., trochlea ; «, tuberosity
of radius ; pi., pisiform; sc.l., scapho-lunar ; ///., trapezium ; ^</., trapezoid ; u., unciform ;
cu., cuneiform ; r.s., preaxial extra-carpal ; h, head of femur; g.t., great trochanter; l.t., lesser
trochanter; t.f., trochanteric fossa; cd., condyles of femur; c«., calcanenm ; ar., astragalus ;
t.s., tibial sesamoid; ent.cu., entocuneiform ; m.cu., mesocuneiform ; ect.cu., ectocuueiform ;
cb., cuboid ; na., navicular.
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AUG 30 1902
2nd Ser. ZOOLOGY.]
[VOL. VIII. PART 7.
THE
TRANSACTION'S
OF
THE LINNEAN SOCIETY OF LONDON.
ON THE INTESTINAL TRACT OF BIEDS; WITH REMARKS ON THE
VALUATION AND NOMENCLATURE OF ZOOLOGICAL CHARACTERS.
BY
P. CHALMERS MITCHELL, M.A., D.Sc. Oxon., E.L.S., F.Z.S.,
Lectceer on Biology at the London Hospital Medical College.
^LONDON:
PRINTED FOR THE l.INNEAN SOCIETY
BY TAYLOR AND KRANCIS, KKD I-ION COURT, Kl.EET STREET.
SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON-HOUSE. I'ICCADILLY, W.,
AND BY LONGMANS, GREEN, AND CO., PATERNOSTER-ROW.
Uctoljcr 1901.
AUG 30 1SU2
r 173 ]
VII. On the Intestinal Tract of Birds ; with Remarks on the Valuation and Nonien-
clature of Zoological Characters. ByV. Chalmers Mitchell, M.A., D.Hc, Oxon.,
F.L.S., F.Z.S., Lecturer on Bioloyij at the London Hospital Medical College.
(Plates 21-23.)
Kead 2U\ March, litdl.
COXTENTS. _
Page
Introduction 173 ;
The Tract in I'alamedea cornuta as an Archecentrie Type J7.5
Valuation and Nomenclaturo of Characters 1 7S
Systematic Desckiption 1 82
Eatitae 1.S2
CarinatiE 184
' Colynibiformes 18 J
.Siiheiiiseifortues 186
Procollarii formes 188
f C'lcoinii'ormes 1 89
\ Anserit'ormes 20]
[ Falcouiformes . 2().j
Summary of the Pelargo-Col}-nihomorphinc Brigade 2l:J
Tinaniifnrmes 21.")
G;dliformcs 210
CiruiforDies 222
Charadriil'ormes 230
Summary of the Aleetoromorphine Legion 240
(hieuliformes 242
Coraoiiformcs 245
{ Passeiiformes 2.52
Summary of the Coraciomorphine Legion 2515
Summary of the Systematic Description 2.")7
Autonomic Nervous System 258
Morphology of the Intestinal Tract 2fil
Meckel's Diverticulum 2C2
MeckeFs Tract 264
Supra-duodenal Loop 265
Colic Ca!ca 267
Characters and Classifieation 27i>
Bibliography 273
Explanation of the I'lates 275
Introductton.
In 1813, in one of a series of papers on the Solvent Glands and Gizzards of 13irds,
Sir E. Home (15) gave a description and some measurements of the intestines in the
Ostrich, Emu, Cassowary, and Rhea; and in 181 i (16) the same avtthor described the
course of the intestines and the form of the caeca in a number of birds, giving figures, in
which, however, the coils were represented as freed from their blood-vessels and mesen-
teries. In the second edition of the ' Le90ns ' Cuvier (5) described the arrangement of the
intestinal tract in some dozen birds and distinguished regions, as the " duodenal loop " ;
SECOND SERIES. — ZOOLOGY, VOL. VIII. 28
1;
174 DE. P. CHALMERS MITCHELL ON THE
the " median loop," exteuding from the duodenum to the unpaired caecum ; the " colic
loop," extending from the median loop to the insertion of the paired caeca ; and the
" rectum." Following Meckel, he identified the " unpaired caecum" as the rudiment of
the yolk-sac. Cuvier, however, had no great range of facts before him, and refrained
from any general conclusions. Owen, in Todd's ' Cyclopaedia of Anatomy and Physiology '
(31), added little to the investigations of Home and Cuvier; and Macgillivray (22) did
little more than to point out that the subject might yet afford useful facts for taxonomy.
Thereafter the svibject was apparently completely neglected until Dr. Gadow (10, 11, 12)
began his extensive and extremely interesting investigations. Gadow studied and
figured the Ioojds and blood-vessels in a large number of forms, distinguished the loops
of the intestine as being " open" or " closed " according to the width of the mesentery
between the limbs of the fold, named the folds '" right-handed " or " left-handed "
according to the position of the descending limb ; but^. above all, described at length
and based taxonomic distinctions on the mode in which the loops were folded within the
coelom.
In 1894, working in the Prosectorium of the Zoological Society, I examined the
intestines of a number of birds in the method from which Gadow obtained his notable
conclusions ; but I found not infrequently that there were individual variations in the
disposition of the loops, particulai'ly as regarded their " right "- and " left-handedness,"
no doubt in connection with the writhing movements of the gut during life, and
came to the conclusion that there was more to be leai'ued from investigation of the
relation of the loo2)s to the unfolded mesentery than from consideration of the mode in
which the folds were packed. The method of examination, which I have since found
to be an extension of Cuvier's method, I described in 1895 (25), and, at greater length,
in 1896 (26). It depends on the morphological nature of the intestinal tract of Verte-
brates, which is a tube suspended in the coelom by a fold of mesentery attached to the
dorsal wall. The tube, in its course, describes an arc between two fixed points, the
pyloric end of the stomach anteriorly, and the insertion of the rectum to the cloaca
posteriorly. A third fixed point is given in the embryonic stage by the attachment of
the yolk-sac nearly in the middle of the ventral edge of the arc, and this point is often
marked in the adult by Meckel's diverticulum (see infra, p. 175). The intestinal tract
increases considerably in length between the fixed points, and along with its mesentery,
which similarly increases, it is thrown into a series of loops which are folded on one
another in the various modes described by Gadow. When the intestines have been
removed from the bird by section at the pylorus and cloaca and by cutting tlie mesentery
along its dorsal attachment, the cut end of the duodenum and of the rectum may be
pinned on a board to the operator's right-hand, these two points and the cut edge of the
mesentery stretching between them being placed in their natural relative jjositions. Then,
with some trouble in the more complicated cases, the various loops may be unfolded
to the left and pinned out ; whereuj)on the mesentery appears as a flat sheet, in shape
roughly the segment of a circle, the cut dorsal edge of the mesentery being the sector,
and the arc, which carries the intestinal tract, being irregularly distorted. The sheet of
mesentery is of course double, and tin; Iduod-vessels and autonomic nervous system run
between the two sheets, being situated outside the crelom. The figures which illustrate
INTESTINAL TEACT OF 151RDS. 175
this memoir represent the intestinal tracts of various birds as seen in preparations of
the kind described, and tlierefore illustrate the morphological conditions of the tracts.
In 1896 (26) I bad already a considerable material, and described and figured some of
the chief modifications of the common tyjic displayed in the groups of birds. Since
then I have been able to add largely to my material. I liave now examined many
bimdreds of birds, including a number of rare forms, and representing nc^arly all the
important groups of birds. Taking the System given by Dr. (ladow in Bi'onn's 'Thier-
Reich,' vol. vi. part ii. pp. 290-301, as one specially familiar to anatomists, I find that my
material includes, so far of course as living forms go, all the groups of llatites, and, of
Carinates, all the Orders, all the Suborders except Mesitcs, and all the Pamilies except
Mesitidoe, Galbulidse, and some Passerine families. For materials I am indebted to the
authorities of the National Collection, who have allowed me to examine a number of
s])irit-specimens, to sevei'al private persons, iucluding in particular Mr. C. Hose of
Borneo through Mr. Beddard, but above all to the Zoological Society of London.
Isearly all the work was done at the Prosectorium on materials from the Gai'dens or
sent to the Prosector. To my friend the Prosector, Mr. Beddard, F.R.S., I am deeply
indebted both for materials and assistance. To Prof. Howes, E.ll.S., I am indebted for
many valuable references to literature, and I have had the advantage of discussing the
nom.enclature of charactei's with Pi-of. Ray Lankester, E.R.S.
The Tract in Falamkuka cohnvta as an Aucuecentkic Type.
It is convenient to describe the conditions of the intestinal tract in one form in some
detail so as to provide a standard for comptirison and to avoid unnecessary repetition.
In a former paper (26. p. 138, fig. 2) I took the condition in an embryo of an Argus
Pheasant, about thirty days old, as a starting-point ; here, for various reasons which will
appear later, I begin with the condition in the Horned Screamer, Palamedea corimta
(fig. 1). When unfolded, in the fashion already described, the intestinal tract is seen
to present three main portions between the pyloric cut end {p.) and the cloacal cut
extremity (cZ.). The first division from^j. to the point w^iere, ^i]).v., the cut portal vein
is represented consists of a long, narrow, and closed loop ; this is the duodenal loop of
Cuvier : it contains the pancreas between its ascending and descending limbs, and
receives the ducts of the pancreas and liver. It always in the undisturbed condition
lies bent backwards towards the cloaca, ventrad of all the other loops, and is therefore;
the first portion of the gut to be seen when the abdomen is opened from the ventral
surface in the usual fashion. Its ascendiu"- and descending limbs are held together bv
a narrow outgrowth of the mesentery. The mesentery, after leaving the duodenum,
expands into a large, nearly circular fold, at the circumference of which, from the
end of the duodenum to c.c, the point of insertion of the cjBca, is suspended the great
portion of the intestine. About the middle of this portion is situated Meckel's Diverti-
culum (m.), which is well known to be a relic of the embryonic stalk of the yolk-sac
{cf. 26. tig. 2). This portion I name Meckel's Tract, as it carries Meckel's diverticulum.
In PaZ«??2^rf^«itpi'esents a number of minor, simple corrugations, but in more specialized
forms the loops present extremely definite and well-marked types of divergence from
28*
176
DE. P. CHALMEKS MITCHELL ON THE
this simple condition. Cuvier (5) divides this median portion into an "Anse moyenne"
extending from the duodenum to Meckel's diverticulum, and an ''■ Anse coUque," from
the diverticulum to the insertion of the ciBca. The examination of a larger number of
types, however, shows that althougli the remains of the yolk-sac when |)resent give a
point of orientation, still there is not a natural pomt of division between the loops
at the insertion of the yolk rudiment, which, indeed, most frequently lies at the
summit of a minor loop. Moreover, the adjective "colic" rightly belongs to a lower
Fig-. 1.
Intestinal Tract of Palamedca cornuta. From^). to jm'. is the Duodenum ; from p.ti. to ex., the insertion of the cajca,
is Meckel's Tract ; from c.c. to d. is the Large Intestine, here convoluted in its anterior portion, G *.
/>.!'., portal vein ; n.m.v., anterior mesenteric or duodennl vein; ;«.»(. v., middle mesenteric vein"; p.m.r., posterior
mesenteric or rectal vein : .r, " bridging " factor of duodenal vein from caeca : ^, recurrent factor of middle
mesenteric vein; p., pylorus; m., Meckel's diverticulum; cZ., cloaca; n.n.. nerves entering mesentery; v.n.,
" visceral nerve," ganglionated chain of the autonomic nervous system.
portion of the gut, as the colon of mammalian anatomy, fi'om which the name was
taken, is not the portion on which Meckel's diverticulum may be found. It is worth
noting, however, that very often a change in the colour of the gut, denoting a phase in
the digestive processes, begins just distal to the diverticulum. Meckel's tract, in the
* Since the large intestine is, with few exceptions, short in birds and but rarely differentiated into anything
comparable to a colon, it will be simider to refer lo it in this Memoir as j-ecfum.
INTESTINAL TKACT OF BIRDS, 177
undisturbed condition, lies very irregularly folded, or rather cruinpled-up dorsad of
the duodenum and veutrad of the rec^tuni. Its posterior portion is nearlv strai'>ht,
an extremely common state. The rectum or third portion of the gut extends from r.c,
the end of Meckel's tract, to cl., where it enters the cloaca. It is suspended by a long
straight piece of mesentery continuous with the mesentery of Meckel's tract. In the
figure, as in some of the later cases, the rectum is represented as folded over to the ri"'ht
so that tlie cut edge of the mesentery from the duodenum to the cloaca is twisted on
itself at the point where the ca?ca are inserted. The large intestine in most birds is
extremely short; its condition in Palamedea, where it is <ronvoluted and mi"-ht be
divided into colon and rectum, is unusual. The calibre of the gut xaries : the condition
seen in Palamedea is very usual ; the duoilenum is wider than Meckel's tract, and the
rectum is wider than either that or the duodenum. The coeca are equal in size and of
moderate length, being intermediate between the short thick stumps of Passerines and
the extremely long, irregularly dilated pouches of the fowl. The l)lood-vessels, shown
in black in this and the subsequent figures, are the veins. At p.v. is represented the
cut edge of the main portal vein on its way to break up in the liver ; the large factor
from the stomach and spleen is not represented, but the three characteristic intestinal
factors are figured. These are the anterior mesenteric or duodenal., draiuino- the
duodenum, and at x receiving a small factor which runs forwards from the cteca and
posterior part of Meckel's tract. The draining of these parts of the intestine by a
tributary or tributaries of the duodenal vessel is seen here in a simple form, but, as will
be seen later, becomes a peculiar and important structural feature. The second or
middle mesenteric factor oi the portal drains Meckel's tract; its main axis runs from
Meckel's diverticulum, but a strong factor, wliich I call the " recurrent mesenteric."
curves round from backwards following the contour of the tract. T\\c posterior mesenteric
or rectal factor, drains the rectum; in many birds it arises from two factors which run
backw^ards along the ventral surface of the kidneys and join opposite the junction of
the rectum and cloaca, entering the rectal mesentery at that point. These tlirec
components of the portal correspond to the three main divisions of the intestines the
Duodenum, Meckel's tract, and the Rectum ; they meet nearly at the same point, but,
in different birds, the order of their joining to form the common portal differs ; and,
after giving considerable attention to the matter, as I cannot find significance in the
order of tlieir junction, I shall not refer to it in this memoir. The arteries of the
mesentery are small relatively to the veins; their minor branches follow the minor
branches of the veins fairly closely ; the main bi-anches arise from the dorsal aorta.
The dotted chain marked v.n. in the figure is the part of the " autonomic nervous
system " which follows the intestine. It is extremely plain in some birds, as in
Palamedea, where it consists of a ganglionated chain following the general contour of
the intestinal loops. The nature of my material has not allowed me to follow it out in
the majority of my specimens. To certain general points in relation to it I shall reciu-
later.
178 DR. P. CHALMERS MITCHELL ON THE
Valuation and Nomenclature of Characters.
In tlie description of the structure of an organ or anatomical part as it occurs in a
large series of different forms, it becomes necessary from simple convenience to attempt
some kind of valuation according to which tlie series of facts shall fall into definitely-
named groups. When there is attempted the difficult passage from descriptive anatomy
to morphology, it is necessary that the valuation and nomenclature should be in relation
to the theory of descent with modification. I assume that birds were monophyletic in
origin, and that the existing forms have branched out in diverging directions from the
ancestral group. The members of this ancestral group, at the stage when they first
might have been called birds, possessed an heritage of characters and tendencies, and these
characters and tendencies have undergone modifications different in amount and nature
in the different groups. The first business is to come to a decision as precise as possible
as to the ground-plan, or archetype, the most ancestral condition of the structures under
consideration. In the present case, I find that the condition of the gut in Falamedea
(fig. 1) may be taken (after allowance for its length, as it is a large bird and vegetarian)
as representing closely the ancestral type. The form of the gut is extremely simple ;
it is distinguished from the intestinal tract of reptiles chiefly by the fact that the three
divisions — the duodenum, Meckel's tract, and the rectum — are sharply marked off one
from the other. Meckel's diverticulum, the morpliological median point of Meckel's tract,
occurs nearly at the actual median point. The pair of caeca are of moderate length and
are functional. The arrangement of the veins is also extremely simple. It will be seen
in the systematic part of this memoir that it is not difficult to refer the more comjolicated
types of arrangement of the tract to the condition in Falamedea. I propose to call such
a condition " archecentric," implying that it represents a jirimitive, ancestrcil, or central
condition, from which the conditions to be found in the other cases have diverged. It
is obvious that the possession by two or more groups of birds of a character in its
archecentric form cannot be an indication in itself that these groups are more closely
related to one another than they are to groups possessing the character in another form ;
for if the diagnosis of archecentricity be correct, the condition has been present in all
birds, and may be retained by any. For exam23le, I have recently (28, 29) endeavoured
to show that the condition in the wing known as diastataxy is archecentric ; that is to
say, that in the ancestral wing there was a gap in the series of quills proximad of the
fourth secondary quill. If that be correct, the fact that two groups of birds jiossess
diastataxic wings is no reason for uniting the groujjs.
When the ancestral condition is modified, it may be regarded as having moved out-
wards along some radius from the archecentric position. Such modified conditions I
propose to call " apocentric." Again, it must be obvious that the mere apocentricity of
a character can be no guide to the affinities of its possessor. Por instance, in the work
on the wing of birds, to which I have already referred, I tried to show that the condition
of the wing known as eutaxic is apocentric ; that is to say, that it is a modification of
the archecentric condition, which in this matter is the condition termed diastataxic.
Before deciding as to the value of eutaxy in a natural classification, it would be necessary
INTESTINAL TRACT OF BIRDS. 179
to decide whether the modification of the archecentric condition were a simple chani^e
that we might expect to occur in independent cases, or if it involved intricate and
precisely combined anatomical changes that we could not expect to occur twice inde-
pendently. In fact, having come to the conclusion that a character is apocentric, we
must pass on to consideration of the problem whether or no the apocentricity be imiradial
or iiiultiradial. In the case of eutaxy I came to tlie conclusion that it was tlie result of
a simple closing of the quill series, which might have occurred repeatedly, and probably
did occur repeatedly, and that therefore it was an instance of what I call here " multi-
radial apocentricity." Similar and common multiradial apocentricities, from which no
direct argument of kinship is to be drawn, are to be found in conditions depending on the
degeneration of a structure. If, for example, the presence of basipterygoid processes
be archecentric in birds, their absence is a multiradial apocentricity from which no
direct argument as to affinity may be drawn.
These multiradial apocentricities lie at the root of many of the phenomena that have
been grouped under the designation " Convergence." Especially in the case of manifest
adaptations, organs belonging to creiitvires veiy far apart genealogically may be moulded
into conditions which are extremely alike. It is of course the business of science to
distinguish the dissimilarities of genetic material under the similarities which are the
results of adaptation to a common purjiose. Ray Lankester (20) long ago designated
such ])ai'allelism of modifications as Homoplasy ; and there seems little reason to doubt
that a careful discrimination and elimination of the homoplastic features in so-called
cases of convergence would leave a residuum plainly showing the genetic differences,
and dispelling many of the hazy ideas which have been grouped round the word
convergence. In the case of the alimentary canal, it is easy to set apart certain
modifications as directly adaptive, and as therefore of no value when the character of
an organ is being considered as an indication of the natural affinities of its jjossessor.
Eor the apocentric modifications in question have been produced in different mammals
as well as in different birds, and hence in birds these modifications must be multiradial
and no indication of relationship. I find in the intestinal tract of birds four j^laiji
homoplasies — that is to say, four kinds of adaptation which produce multiradial apo-
centricity ; three of these are well known, the fourtli, so far as I am a\)'are, has not yet
been pointed out. That aU four occur among mammals as well as among birds is a
fortunate circumstance that definitely proves their multiradial character.
First Homoplastic Modification. Lengthening of the gut in graminiferous and grazing
birds. — This feature has been well known for long, and is in obvious relation to the
circumstance that the nutrient constituents of the food of such creatures are in a form
difficult to digest and in intimate admixture with a large bulk of indigestible material.
Much time and much surface are i-equired for digestion and absorption, and these are
provided for by increase in length of the gut.
Second Homoplastic Modification. Lengthening of the gut with thickening of its
wall and relative decrease of its calibre in piscivorous birds. — The very long and narrow
gut with stout walls is known in birds and mammals. The small calibre and thick wall
are doubtless a protection against mechaniccil injury by sharp bones. The extreuin
180 DR- P- CHALMEES MITCHELL ON THE
length is at first surprising, as we are accustomed to regard fish as being readily digestible.
However, a bird does not eat fish carefully with a knife and fork, but bolts it whole.
Dr. Hutchison, a recent writer on foods (17), calls attention to the large amount of
waste matter in uncooked fish, amovinting to fully seventy per cent.
Third Homoplastic Modification. Shortening of the gut in frugivorous l)irds.— The
tendency of the gut in birds and mammals which live chiefly on fruit to be very short,
thin-walled, and wide is well known, and is in direct relation to two simple physiological
factors. Tlie nutritious substances in fruits are in a form which renders them capable
of rapid and fairly com])lete absorption, and the organic salts present stimulate osmosis.
The ease of absorption makes a relatively large surface unnecessary, and tlie large calibre
of the gut not only diminishes the outflow from the blood caused by the presence of
organic salts, but it decreases the danger of violent purging. Tlie vigour of peristalsis
in birds is remarkable ; when the duodenum of a pigeon contracts it becomes as hard
and tense as a piece of cartilage.
Fovrth Homoplastic 3£od ificatiou . Increase of length rather than of calibre in large
birds. — I am not awa,re that the circumstance has been noticed, but it is the case that
in large birds and large mammals the gut tends to be relatively longer than in small
birds and small mammals. The explanation, I think, is simple. la the course of
phylogenetic increase in size, the various organs increase in size correlatively with the
v\hole, but m a mode corresponding to their functions. The first business of the
intestine is to present to the food-absorbing surface sufficient to supply the needs of the
whole body. As the calibre of a tube increases, its capacity increases more rapidly than
its surface ; it follows that to preserve the same relation of intestinal surface to intestinal
capacity, the length of the gut must increase more than the calibre in the course of
pliylogeuetic increase of size. It is interesting to notice that the caeca and the rectum,
two portions of the gut of birds in which absorption is not so great, increase in large
birds almost as much in calibre as in length, so that very frequently large birds display
Cceca and rectum which appear to be much wider than the anterior portions of the
intestinal tract. It is plain, from what has been said, that no genetic significance is to
be attached to such conditions, which are merely a result of the homoplastic modifi-
cation due to large size of the whole creature.
There is no need to discuss here the difficult problems as to the mode of origin of
homoplastic resemblances. To some extent they may be freshly epigenetic in each
o-eneratiou ; and Gadow's (12) investigations into the lengths of the gut in chicks as
compared with adults would seem to show that there is much to be learned as to the
occurrence of changes of form and length in direct relation to changes of diet. They
may have come about by a slow selection of strains with genetic variations in the
direction uf increased length or of capacity to acquire increased length at the stimulation
of i'ood ; or, on neo-Lamarckian principles, they may be the summations of the effect of
stimulations in a series of generations. It is enough to state that these homoplastic
modifications must be allowed for or " corrected " before the drawing of conclusions as
to relationship. Before the condition of an intestinal tract can be taken as affording a
clue to the affinities of its possessor, it must, in imagination, be shortened, in the case
INTESTINAL TEACT OF BIRDS, 181
of a large bird, or a graminifcrous or piscivorous bird, or lengthened in tlie case of a
frugivorous bird. In more general terms : when we Jire satisfied that an apocontricity
is multiradial, as is certain!}^ the case when it is homoplastic, we must neglect it when
we are dealing with the one character as a guide to affinity (fig. 2).
Pig. 2.
v-/
MfTACEMrfif
MULTIRADIAI
APOCENTRIC
PSnJDOCENTRiC
\
ARCHECENTR/C ;
Diagram to explain Nomenclature of Characters.
A complex npocentric modification of a kind that we cannot well expect to be repeated
independently, and that may be designated as uniradial, must be the most certain guide
to affinity. It happens frequently that such a modification forms a new centre aroimd
which new diverging modifications are produced. Such a centre I propose to call a
" Metacentre," borrowing a convenient term from physics. It is obvious that the
condition of a character, archecentric so far as the whole group of birds is concerned, is
metacentric with regard to the common stock of birds and reptiles, and that the trans-
formation of an apocentric character into a metacentre is simply an event in the general
process of divergent evolution. I justify the nomenclature which I am proposing
largely because it brings the valuation and classification of characters into line with our
conception of the general process of evolution.
Finally, there remains to distinguish a form of apocentricity extremely common and
often perplexing. Such conditions are marked by an apparent simplicity that, however,
reveals its secondary nature by some small and apparently meaningless complexity.
Such a condition that mimics the archecentric condition but which can be distinguished
from it, I propose to call '" Pseudocentric."
I trust that the ideas underlying this attempt at the valuation and nomenclature of
characters, so far from being novel, are merely a codification of criteria in common
SECOND SERIES. — ZOOLOGY, VOL. VIII. 29
182 DE. p. CHALMEES MITCHELL ON THE
employment among naturalists. I find, however, that such a codification was necessary
when I tried to arrange systematically the modifications of the characters with which
this memoir deals. So far as I have used theiu in my own work, I have found them
illuminating, and I offer them in the conviction that the rigorous discipline which their
use entails would prove of general utility.
SYSTEMATIC DESCRIPTION.
The Classification according to which this part is arranged is that given by Gadow in
Bronn's 'Thier-Reich ' (12).
EATIT^.
Casuarii. — In the Cassowaries and Emus the gut is archecentric in character. The
three regions are weU marked ; the duodenum is wide in the Cassowary, but longer and
narrower in the Emu. In both, Meckel's tract is nearly circular in form, with well-
marked minor folds in the Emu, and, in both, with a distinct relic of Meckel's diverti-
culum about the middle of its course. The rectum is short and nearly straight, this
being the only divergence from the archecentric character and a divergence displayed by
the vast majority of bii'ds. The loops are drained by the characteristic factors of the
common portal vein. The caeca are of moderate length as in the type. I have already
given a figure of the gut in the Cassowary (26. fig. 3).
Struthiones. — The Ostrich (26. fig. i) is also markedly archecentric. The diver-
gences from the archecentric type are very slight : the duodenum is rather longer and
has a minor loop situated on its distal limb ; the rectum is enormously long, being in
fact at least equal in length to the first two portions together ; the caeca also are
extremely long. The factor of the duodenal vein from the caeca comes off rather further
from the main stem of the portal vein, and instead of running forwards within the
mesentery, it leaves the mesentery and in consequence must be separated when the
intestine is unfolded : in this condition it forms what I call a " bridging " vein.
Rhe^. — In Bhea americana (fig. 3) the gut is archecentric, and indeed can be distin-
guished from that of Falamedea only in two points. Meckel's tract is somewhat
elongated, Meckel's diverticulum lying at its apex, and the caeca are much longer
relatively. The region of the caeca is drained by two " bridging " factors of the duodenal
vein which leave the mesentery and have to be divided when the duodenum is unfolded
outwards. A slight peculiarity that I have noticed in no other bird is that a factor of
the splenic vein (fig. 3, z) drains the proximal portion of the rectum.
Aptektges. — The gut of the Apterjjx (fig. 4), while markedly archecentric, is the
most modified of those among Struthious birds. The duodenum and the rectum conform
to type, save that the rectum is relatively shorter and is straight. The greater portion of
INTESTINAL TKACT OF BIRDS.
Fig. 3.
183
Intestinal Tract of Uliea aunr'ic ma,
X, x\ cut ends of " bridging" factors of duodenal vein, draininu; the distal portion of ileckel's tract and one
caecum ; r, factor of the splenic vein. General description as in legend of figure 1.
Fit?. -1.
~>I
^ x>
Intestinal Tract of Apterij.v ManieUi.
For explanation, see legends of figs. 1 and 3. S.D.K, apex of supra-duodenal loop.
29*
1.84 DE. P. CHALMERS MITCHELL ON THE
Meckel's tract forms a large circular fold with Meckel's diverticulum at its central point,
opposite the origin of the middle mesenteric vein, as in the archecentric type. But the
posterior portion of the tract, that to which the very long caeca are attached, is specialized
into a distinct minor loop. This I call the supni-duodenal loop, because, in the natural
condition of the intestine, it lies directly above the duodenum and is drained by factors
of the duodenal vein. This loop in Apteryoc is the first appearance of a specialized
structure to which I shall have to direct repeated attention.
Wlien correction has been made for the obviously homojilastic modifications in the
intestinal tract of Ratites — that is to say, when a reduction has been made in the case of
the larger forms for size and in the case of the herbivorous forms, — it is plain that while
all are archecentric, the Casuarii are by far the least modified, and that in this as in
many other characters they deserve the position assigned them as extremely primitive
types. Struthio and Rhea are more modified, and the modification is in the same
direction in each. Apteryx stands somewhat apart from the others, the specialization of
the supra-duodenal loop being well marked in it- The general character in all, however,
is so plainly archecentric that no argument can be drawn from the intestinal tract for or
against the polyphyletic nature of the group. Such similarities as they present to one
another or to other groups are simply part of the common inheritance of all birds.
CARINAT^.
COLYMBIFORMES.
CoLYMBi. — I have examined the Great Northern and Red-throated Divers [Colymhus
glacialis and C. septentrionalis), and have already figured the intestinal tract (26. fig. 5).
The duodenal lo'^p is straight and normal; the rectum is straight and very short.
Meckel's tract is pulled out into a series of short, straiglit, symmetrically arranged loops
with the Meckel's diverticulum nearly at the middle of its course, and with a well
specialized supra-duodenal loop drained by a bridging factor of the duodenal vein. The
three main factors of the portal vein are typical, and the cseca are of moderate length.
It is clear that the Divers present a certain degree of apocentricity in the gut. The
fish-eating habits have lengthened the tract, but the increase of length takes place not
by a series of irregular loopiugs but in a definite fashion.
PoDiciPEDES. — In the Crested Grebe, Fodicipes cristatus (fig. 5) the duodenum is a
large loop somewhat wider distally. Meckel's tract is still more s^Jccialized than that of
the Colymbi. It presents first a long minor loop, then a short loop, then a long loop,
on which, from the arrangement of the blood-vessels, I suppose the yolk-sac to have lain,
although it is not represented by a Meckel's diverticulum ; then follow two short loops,
and then a very large supra-duodenal loop with characteristic bridging vein. The cseca
are relatively short, but wide and functional, and the rectum is short and wide.
In Tachyhaptes jiitviatilis, the Little Grebe (fig. 6), the apocentricity seen m the
Crested Grebe is carried further in two points, otherwise the character of the tract is
■: 0 . ■
INTESTINAL TRACT OF BIRDS.
isls
similar. Two of the minor loops of Meckel's tract are carried out to a great length, one
of them bearing Meckel's diverticulum at its summit ; the caica are relatively much
longer, and the supra-duodenal loop is still more highly specialized. The Podicipedes,
then, present an apocentricity rather more marked tlian that of the Divers.
Fis:. 5.
Intestinal Tract of Pafliclpes cri.ftatus.
X, bridging factor of the duodenal vein draining S.D.F., the supra-duodenal looj) ; ?, probable
position of Meckel's diverticulum.
When allowance has been made for the size and piscivorous habits of the Colymbiform
birds, it appears tliat Avhilc the character of the gut has moved outwards from the archc-
centric position chiefly in the direction of the formation of specialized straight loops
(according to Gadow^'s nomenclature, the folding is orthoccelous), the apocentricity is not
great, and is ccrtainlv not of such a uniradial character as to associate them closely
with any other group.
186
DE. P. CHALMEES MITCHELL ON THE
SPHENISCIFOEMES.
I have already described and figured the intestinal tract o^ Eudyptes (26. fig. 6). I
now am able to add descriptions of tbe condition in Sj^heniscMs demersus (fig. 7) and
Aptenodytes Pennanti (fig. 8). In all three the intestine is extremely long and of
Fig. (3.
X
Intestinal Tract of Tavhyhaptes fluviatilis. Lettering as before.
slender calibre with firm walls; in the fignres the actual length is somewhat under-
estimated, so that the " cori-ection " for piscivorous diet has been made partially. The
three portions of the tract are well marked and are each drained by a characteristic
factor of the portal vein. The duodenum is extremely long ; it is thrown into a
complicated set of minor loops resembling the condition in the fish-eating Eagles, in
Eudyj)tes and Spheniscus ; in Aptenodytes it forms a loosely rolled spiral, a mode of
packing increased length that is not at all uncommon among birds and that must
be regarded as a mrdtiradial apocentricity. Meckel's tract in all three is moderately
specialized ; Meckel's diverticulum lies nearly at the middle of its course, but there is a
strong tendency for the numerous minor loops to be extended in length or expanded
INTESTINAL TRACT OF BIRDS.
Fig. 7.
187
Intestinal Tract of S/JwiiiscKS demersus.
l; supra-csecal kink ; other lettering as before.
Fis. 8.
Intestinal Tract of Aptetwdytcs Pennant!. Lettering as before.
188 DR. p. CHALMERS MITCHELL OX THE
into minor systems. Of these, two are well-marked in all : a very large loop towards
the distal extremity of ^Meckel's tract and drained by a factor of the middle mesenteric
vein, and a long narrow supra-duodenal loop which is drained by a " bridging " factor of
the duodenal vein. In Sjifieaiscits the portion between the supra-duodenal loop and the
rectum, wliich is very long in all, is expanded into a minor fold, an arrangement
characteristic of the Eagles, and to which I give the name " supra-csecal kink." The
rectum in all is extremely short, and the caeca are vestigial in those that 1 have
examined, but Gadow mentions that they were relatively long in a Eiidyptes.
It is plain that the character of the tract in the Sphenisciformes is markedly
apocentric. TThen due allowance has been made for the piscivorous lengthening, the
form still remains apocentric : the short caeca, the tendency to foi-m minor loops, the
supra-duodenal loop, the tendency to form a supra-ceecal kink, and the extremely short
rectum make up a peculiar and distinct type.
peocellaeiifor:mes.
Of these I have had an opportunity of examining only the Xorthern Petrel and the
Albatross. In Ftdmarus glacialis (26. fig. 7) the duodenum is compound, and ileckel's
tract is expanded into six very long, narrow and straight minor loops, the fourth of
which bears Meckel's diverticulum, while the last is a distinct supra -duodenal loop
di-ained by a "bridging" vem. The terminal portion of Meckel's tract is long, thrown
into sxipra-Cfecal kinks as in Spheniscu.^ and the Eagles, and is drained by the posterior
mesenteric vein. The cjeca are vestigial, and the rectum is extremely short. In
the Albatross, Diomedea exulans (fig. 9), the duodenum is simple, and ^reckel's tract is
drawn out into a series of extremely long nan'ow loops (the length of these is under-
estimated in the figure). The form of the tract is much alike in the Albatross and the
Petrel ; the chief difterence is that the fii-st three minor loops of the Petrel are repre-
sented in the Albatross by two long loops with shorter loops between them. A very
long loop bearing the Meckel's divei-ticulum then follows in both ; the lower portion
of Meckel's tract, the supi*a-caecal kinks drained by the rectal vein, the vestigial caeca,
and the verv short rectum are alike in each. The onlv striking ditference is that
there does not appear to be a definite supra-duodenal loop with •• bridging " vein in
Diomedea. The specimen I examined had been preserved for long in spirit, and
possibly 1 may have overlooked a ••bridsins" vein. althoiis:h I examined the resrion
minutely for it.
It is plain that the ProceUaruform birds present a markedly apocentric type of intes-
tinal tract, the special charactei"S being the transformation of the middle loop into a
number of definitely -placed minor loops, the appearance of supra-c<ecal kinks on the
large posterior portion of Meckel's tract, drained by the rectal vein, the vestigicd caecei,
and the very shon rectum.
Gtidow ^12) vuiites the Colymbiformes, Sphenisciformes, Procellariiformes, and extinct
Ichthyornithes into a '"Legion" under the name Colymbomorphce. Concemin? the
intestinal tract of the extinct forms we have no information. The others have all moved
out from the archecentric condition. Their apocentricity is fii'st a relatively increased
INTESTINAL TRACT OF BIRDS.
189
length in association with the nature of their food, and certainly multiradial ; second,
Meckel's tract in all, wliile remaining nearly symmetrical, tends to be drawn out into a
series of long, narrow, and straight loops, a feature wliich may give some clue to affinity ;
Fis. '■'.
Intestinal Tract of Diomedea eu-idans. Lettering as before
third, the rectum in all is very short. The Colymbiformes are least modified ; the
Penguins and Petrels are more modified, the tw^o latter sho-\dng degeneration of the
cseca and certain peculiarities in the ])Osterior portion of Meckel's tract, peculiarities
repeated in Steganopodes and Falconiformes.
CICONIIFORMES.
S T E GA N" O P 0 D E S.
(1) Phaethontid^. — Phaethon (fig. 10) displays a simple form of alimentary tract.
The duodenal loop is simple ; Meckel's tract is thrown into a series of narrow, minor
loops, some of which are slightly complicated Ijy folding. Meckel's diverticulum occurs
on one of the minor loops rather more near the distal end of the tract. The posterior
portion has no special supra-duodenal loop, and ends in a portion drained by the rectal or
SECOND SERIES. — ZOOLOGY, VOL. VIII. 30
190
DE. P. CHALMERS MITCHELL ON THE
posterior mesenteric vein and exliibiting' a short supra-csecal kink. The caeca are
reduced, but not to the same extent as in other Steganopodes except the Pelicans. The
rectum is extremely short. The three factors of the portal vein are of diagrammatic
simplicity.
Fig. 10.
Intestinal Tract of Phaethon fla I'lrostris. Lettering ;is before.
(2) SilLiD^. — In Sula bassana (fig. 11) the apocentricity of Phaethon is carried
further. The general arrangement is similar, hut the duodenum is longer and is com-
pound ; certain of the minor loops of Meckel's tract are longer, especially the last, that
forming a characteristic supra-duodenal loop with bridging vein ; and the kink on the
posterior part of Meckel's tract, that drained by the rectal vein, is larger.
(3) Phalacrocoracid^. — In Phalacrocorax carlo (fig. 12) the duodenum is very long
and narrow ; Meckel's tract is extremely elongated, in which respect it resembles the
condition found in Platalea and Phatnicopterns, and is symmetrically disposed about the
middle mesenteric vein, which in the archecentric fashion runs from a large Meckel's
diverticulum. The supra-duodenal loop is long; there is a large supra-csecal kink
supjjlied by the rectal vein. The cseca are vestigial, and the rectum is relatively longer
that in Phaethon and Sula. In Plotus anhinga (fig. 13) the form of the tract presents
marked dhferences. The duodenum is similar. The anterior portion of Meckel's tract
is relatively much shorter, and consists of a single narrow loop, followed by one very
INTESTINAL TRACT OF BIRDS.
191
Fig. 11.
I ntostinal Tract of Sula hassana. Letti'rinjr as before.
Fig. Ii>.
Intestinal Tract of Pkalacrocorax ,who. Lettering as before.
30*
19:
DK. P. CHALMEB:> AJITCHELL OX THE
long loop, at the end of which lay what I took to have heen a Meckel's diverticulum.
Then follows a verv long posterior portion, relatively longer than in the diagram, and
thrown into a series of straight minor folds, the posterior of which is a supra-duodenal
loop with a " bridging '" vein. Just beyond this is the trace of a supra-csecal kink. The
rectum is long and wide, in fact is typically archecentric. Of the usual pair of cseca,
only one \^-as developed, and that in a vestigial form, but Beddard (2) states that indi-
viduals vivry. some possessing two vestiges. The minor loops of Meckel's tract show a
Fis. 13.
tendency to
Meckel's diveniei
p as i: riicans- It I were wrons in my plaemg of
-Liii. acd what I took ior it was the merest traoe. : '.he
''.'-•■;'-' " - ' - irisriv even; :._ rs
Or the evideno? of the iatestLC-
- be inclined to place
1 4'' PRse-jLTTDi — In ?w.7.rf,ii -Sg-. 14) ^ite form of the iiitt^>
: diSece~ees are that in J"-
is v,3y like
e tract is
INTESTLVAL TEACT OF BIRDS.
193
relatively somewhat shorter, the minor expansions of Meckel's tract are less numerous,
that bearing Meckel's diverticulum being longer ; there is a supra-duodenal loop ; the
cseca are less developed and the rectum is longer.
Fig. 14.
Intestinal Tract of a species of Fregata. Lettering as before.
(5) Pelecanid^. — In the Pelicans {Pelecanus rttfescens, fig. 15) the form of the
intestinal tract is simple. The duodenal loop is long, narrow, and some^'hat twisted ;
Meckel's tract is nearly symmetrical round the middle mesenteric vein which runs from
the position of a large Meckel's diverticulum. There is a well-formed supra-duodenal
loop, a suprarcsecal kink drained by the rectal vein, and the caeca are relatively longer
than in other Steganopodes. The minor loops of Meckel's tract tend to be bunched up
towards the mesenteric vein, a feature that cannot well be represented in a diagium
showing the unfolded condition.
The Steganopodes are typically piscivorous, although some of them also take any kind
of floating carrion, and in aU of them allowance must be made for the piscivorous length
of gut. Their apocentricity, apart from such homoplasy, consists, as in the Cohonbo-
morphge, of a general tendency for Meckel's tract to be expanded into a series of short
straight loops. In the Colymbomorphse, however, these loops frequently increase in
length and become reduced in number; in the Steganopodes the tendency is rather for
the minor loops to increase in number, and for Meckel's tract to be either bunched up
194
DE. P. CHALMEES MITCHELL ON THE
tightly or much elongated as a whole. These two forms of modification of the middle
loop point towards the condition found in many of the Ibididse sucli as Platalea, and in
Fie. 15.
Intestinal Tract of Pelecanus rvfescens. Letterins; as before.
the Flamingoes, where Meckel's tract, in the unfolded condition not unlike that of the
Pelican,l[is in life twisted irregularly into a spiral. I find, then, in the apocentricity of
the Steganopod Meckel's tract an underlying resemblance to that of the Colymbomorphae,
as if a metacentric position had been common to all these, but from this metacentre the
two sets have diverged in difi'erent directions. The reduction of the cseca, the shortening
of the rectum, and the formation of minor loops above the cseca, but drained by the
posterior mesenteric vein, are common, Intt not invariable, in the whole set.
INTESTINAL TEACT OF BIRDS.
195
(1) ScopiD^. — In Scojms umbretta (fig. 16) the duodonura is long and slightly twisted.
Meckel's tract is nearly symmetrical, a large Meckel's diverticulum being near the centre
of its periphery, and is expcinded into a number of wide, irregular, but in the main straight
minor loops, the last of these being longer and forming a typical supra-duodenal loop.
The rectum is straight, of moderate length, and the pair of caeca are reduced. The veins
are typical. A small area supplied by the rectal vein corresponds to the supra-caecal
kink.
Fi-. 16.
Intestinal Tract of Scopus umhrettii. Lettering as before.
(2) Ardeid/E. — In the Herons and Bitterns, of which I have examined a number of
species, the ground-form is like that in Scopus. The duodenum is a long narrow loop,
nearly always considerably twisted to the left, usually more so than appears in tlie two
figures {Nycticorax grisem, fig. 17, and Ardeu candklissima, fig. 18). Meckel's tract
may be rather elongated as in Nycticorax, or relatively shorter as in Ardea. It is
always drawn out into a large number of minor loops, many of which are irregularly
folded, and not infrequently complex in themselves ; the distal minor loops in Ardea
(fig. 18) show this in a relatively simple form. In the Little Bittern and some other
Kg. 17.
Intestinal Tract of Nycticorax griseus.
C.C., single colic caecum characteristic of Herons ; 1-. 1 and 7i-. 2, supra-c;ecal kinks. Other lettering as before.
Fig. IS.
m
Intestinal Tract of Ardea candidissima. Lettering as before.
THE INTESTINAL TEACT OF BII{DS. 197
Herons tlie complexity of the minor loops is very great. The penultimate loop of
Meckel's tract is always a typical supra-duodenal loop witli " bridging " vein, and then
follows a siipra-caecal kink. It is typical of llie Ardeida?, as is well known, that one of
the two colic caeca he absent, and I have found no exception to this, but Beddard has
recorded such a case. The rectum itself is straight, but not very short.
A considerable jiart of the apocentricity of the int(^stinal tract in the Ardese must be
attributed to the fact thiit tliey are relatively large birds with a diet consisting chiefly
of fish, in consequence of which the gut is very long and narrow. Underlying this is a
general resemblance to the ground-form of the Steganopod and Colymbomorph gut,
shown Iti the symmetrv of Meckel's tract aroimd tlic middle mesenteric vein, the
persistence of a large Meckel's diverticulum opposite tlie end of this vein, and the
formation of a kink immediately above the colic coeca but supplied by the rectal vein.
The special Ardeine features aie the elongation of the duodenum as a narrow fold cmrvmg
to the left ; the throwing out of Meckel's tract into minor loops, which, straight in the
simpler forms and always folded over simply (" orthocoely " of Gadow), tend to loecome
complicated in themselves. Tlie persistence of only one of the colic caica is a most
peculiar feature, normally absent only in Scopus. But the absence occurs not infre-
quently as an individual abnormality in som(> other birds, and T shall make further
reference to such cases.
Although Scopus, so far as the character of the intestinal tract takes us, is more
archecentric than the Herons and Bitterns, it plainly belongs to their group rather than
to the Storks and Ibises.
C 1 c o N I .E.
(1) CicoNiiD^. — The Storks, of which I have examined a considerable number, present
a most interesting series of modifications. Anastomus oscitans (see Plate 21), the Indian
Open-bill, displays so little difference from the ground-form of the intestinal tract among
the set of l)irds I have been discussing, that I do not think it necessary to figure it in a
separate block. The duodenum is short and straight : Meckel's tract is thrown into a
very large number of short straight loops nearly symmetrically disposed around
the middle mesenteric vein, the latter running backwards from a large Meckel's
diverticulum. There is a very large and complicated svipra-iluodenal loop, more compli-
cated than in any of the other Storks I have examined, but di'ained by the usual bridging
vein. Above the reduced cajca lies a short kink drained by the rectal vein, and the
rectum is straight and of moderate length. In Fseudotaiitalus ibis (fig. 19) the typical
Ciconiiue apocentricity begins to appear, and this is of the definite type that I call
uniradial and am inclined to regard as a sure sign of affinity. The duodenum is
enormously long, but the bending which appeared in Herons is here transformed to a
spiral twist, represented in the figure as partially uncoiled, with the result that the vein
is otit of the mesent(n'y. Tiu; first minor loop of Meckel's tract is very large; and is in
itself slightly twisted, and, in the unfolded state, partly rolled in the duodenal s])iral.
The remaining portion of Meckel's tract consists of a few simple folds, symmetrical
SECOND SERIES. — ZOOLOGY, VOL. VIII. 31
198
DE. P. CHALMEES MITCHELL ON THE
about the middle mesenteric vein, which runs from a Meckel's diverticulum. The distal
portion of Meckel's tract forms first a distinct but short supra-duodenal loop, and
then a double kink drained by the rectal vein. The caeca are reduced and the rectum
is straight and short. Dlssura episcopm, Leptoptilus crumeniferus, L. argala, and Cicoma
alba are practically identical witli this condition. In Ciconia nigra, which I have already
figured (26. fig. 9), the condition is similar, except that the duodenal loop and the first
Fig. 19.
Intestinal Tract of Pseiidotantalus ihis. Lettering as before.
loop of Meckel's tract form more perfect spirals, which in the unfolded condition are
rolled together. In Mycteria americana (tig. 20) a similar condition exists, but the
duodenal spiral and the spiral on the first loop of Meckel's tract are still longer and
more complex, and are more intimately rolled together, with the result that certain
factors of the duodenal vein run across draining the loop with which the duodenum is
associated. These are represented in the figure as divided and the spirals are partly
untwisted. Among the Storks, then, we see the interesting state of affairs that the
character of the gut, starting from a metacentric position common to a large group of
birds, develops along a special radius forming a unirudial line of apocentricity.
INTESTINAL TEACT OF BIKDS.
199
Intestinal Tract of Mycteria americana.
-, cut ends of veins from first loop of Meckel's Tract to duodenal vein.
(2) Ibidid^. — I have already figured the intestinal tract of Flatalea leucorodia
(26. fig. 8). It is more archecentric than that found in other Ciconiiform birds. The
duodenum is a long narrow loop bent round to the left as in the Herons, but showing
no trace of spiral formation. Meckel's tract consists of an elongated system of short
loops, symmetrically disposed round the middle mesenteric vein, which runs from a
Meckel's diverticulum. This region of the gut is slightly twisted into a spiral in the
unfolded condition. The last loop of Meckel's tract forms a simple supra-duodenal fold ;
the caeca are short, and the rectum is straight but considerably reduced. Flatalea is
certainly low down in the Ciconiiform scale, but none the less it displays the features of
the group in a simple form, and is quite different in the character of the gut from
Nimienius (Plate 22), a low Charadriiform bird with which it has been compared.
31*
200
DK. P. CHALMEKs MITCHELL ON THE
Phcenicopteri.
Phcenicopterii)^. — In Fhoenicopterus ruber (tig. 21) the duodenum is a simple loop,
but with a slight Heron-like twist. Meckel's tract is very much elongated as in
Platalea, and, again as in that bird, it consists of very many short loops symmetrical
about the middle mesenteric vein which runs towards a Meckel's diverticulum. As in
Plaialea tliis region is twisted into a rude spiral, and there is a well-formed sujjra-
duodenal loop. The caeca are long, and the rectum is of moderate length but straight.
The only Anserine feature in the intestinal tract of the Flamingo is the presence of long
caeca; and this, as we shall see in the further course of this memoir, is an obviously
multu'adial apocentricity. In all other respects the intestinal tract of the Flamingo is
that of a low Ciconiiform bird.
Fig. 2L
-->'X
Intestinal Tract of Phcenicoj>ieriis rulicr. Lettering as before.
It is unnecessary to do more than sum up in a few words the conditions of the gut
found among the Ciconiiform birds. The ground-form is closely similar to tliat of the
Colymbomorphie, pointing towards the existence of a common metacentre for all these
birds. From this metacentre the groups of Ciconiiformes have diverged in different
directions and to different amounts. The Steganopods sliow a tendency to the bunching
up of the loops of Meckel's tract on the axis given by the middle mesenteric vein. The
Herons display a tendency to the individual elaboration of the minor loops of Meckel's
tract. The Ciconiine birds show the most definitely uniradial apocentricity, 'consisting
of an elaboration and intimate connection between the duodenum and the first looji of
Meckel's tract, with a reduction of the posterior portion of Meckel's tract, except the
supra-duodenal loop. The Ibidid*, or at least Platalea, show a primitive elaboration of
Meckel's tract as a whole, and this tendency is carried further in the Flamingoes.
INTESTINAL TKACT OF BIRDS. 2^
ANSERIFORMES.
P A L A M E D E yE.
PALAMEDEiDiE. — I described the intestinal tract of Palamedea above (fig. 1). The
intestinal tracts of Chauna chavaria (26. fig. 10) and of C. derbiana do not differ from
tliat of Palamedea in any essential detail. I am following Dr. Gadow's classification for
convenience of reference, but it is obvious that from the character of the intestines the
Palamedeae have no claim to be placed in close association with the other Anseriform
birds, or indeed with any of the Carinate birds witli which I have as yet been dealing.
The Palamedese exhibit what I take to be the most primitive or archecentric type of
intestinal tract to be found among birds. In the gut they have a general resemblance to
the Ratites, and to the lowest members of all the other groups ; to such they bear more
resemblance tlian to the Ducks and Geese, which in tlie intestinal tract exhibit a definite
or uniradial apocoitricity, no trace of which occurs in any of tlie three Screamers. On
the character of tlie intestinal tract alone I would not remove the Screamers from other
birds, Init rather all other bii-ds from the Screamers, leaving them in centrjil and
primitive isolation.
A N S E R E S.
Ansemd.^. — I have shown that in the Cavinates I have up to this point discussed,
with the exception of the Palamedete, there is strong evidence for the existence of the
apocentric type of gut which forms a metacentre from which the various groups have
diverged further. The metacentric character consists chiefly in the transformation of
Meckel's tract into a definite series of narrow, straight loops, folded in tlie mode Gadow
terms orthocoelous, and ari-anged nearly symmetrically round the middle mesenteric vein,
which fornix an axial line running from a Meckel's diverticulum. The form of the gut
in the Anseridae can be referred easily to such a metacentric condition. In Aiiscraiias
melanoleuca (fig. 22) this metacentric condition is reproduced Avith almost diagrammatic
fidelity, an interesting circumstance in view of the primitive position among the Anseres
generally assigned to this bird. The duodenum is a long narrow loop ; Meckel's tract is
thrown into very symmeti-ically-arranged narrow minor loops, the most peripheral of
which bears a large Meckel's diverticulum. Between this and the duodenum there are
three minor loops, a number typical of this part of the gut in tlie Anseres. The last loop
of Meckel's tract is a typical suj)ra-duodenal loop with the usual " bridging" vein from
the duodenum, and with the very long ca;ca closely applied to it. The rectum is not
very short but is straiglit. In Cygnus atratus, the tract of whicli I have figured in a
former paper (26. fig. 11), certain modifications of the type in Anserauas are present.
The first three minor loops of Meckel's tract are more complicated, showing a tendency
to give off !~econdary loops. The axial loop, which bears Meckel's diverticulum, is greatly
elongated and usually somewhat irregularly folded, with the result that the main vein
leaves the mesentery, being shorter than the loop itself. The minor loop, just posterior
202
DR. P. CHALMEES MITCHELL ON THE
•29
1 ntestinal Tract of Anseranas melanoUum. Lettering as before.
Fig. 23.
7
Intestinal Tract of Mergus albellus. Lettering as before.
INTESTINAL TKACT OF BIRDS.
203
to this, is usually elongated and may be similarly twisted. The supra-duodenal loop,
the cteca, and the rectum are as in Anseranas. The vast majority of Ducks and Geese
that I have examined, however different their size and habits, faithfully reproduce this
type. The diagram given for Ci/gaus atratus, witli the most trifling alterations, might
serve for Anas, Anser, JEx, Chanlelasmns, Berniolu, DciKjrocijgna, Fuligtda, JSesonetta,
Tadorna, and doubtless, so constant is tlie type, for many others. Mergus alheUiis, the
Smew, presents an interesting variation (fig. 23). The duodenum is unusually wide ;
Meckel's tract is thrown into a large number of very short loops at the periphery of an
almost circular mesenteric fold, but Meckel's diverticulum lies at the apex of a somewhat
longer fold lying in the axis of the system. There is a supra-duodenal loop, but it is not
Fig. 24.
Intestinal Tract of Sjtatidn cliipeata. Lettering; as before.
drained by a " bridging " vein. The colic ci,eca are paired, but practically non-existent,
and Beddard (2. p. 459) mentions an instance where one of the two was absolutely non-
existent. At first sight this intestinal tract appears more archecentric than that of any
of the other Anseriform birds except the Palamedese, but I do not doubt that it is an
instance of what I term pseudocentric simplicity. The caeca are obviously degenerate,
and in other Mergansers they are longer ; and examination of the first part of Meckel's
tract shows that it might have been derived by a fusion of the three first minor loops,
these having been more complicated tlian in Cygmis, and more as they are in Spatula
(fig. 24). There is a short supra-csecal kink, but it is not supplied by the rectal vein.
Spatula clypeala (fig. 24) shows a form of gut which is simply a further elaboration of
204
DR. P. CHALMERS MITCHELL ON THE
the Cygnvs type. The modifications are confined to Meckel's tract. The three most
anterior minor loops of that region are complicated by an elaborate series of minor
folds; the axial loop bearing Meckel's diverticulum is as in Ci/gims, but the loop next
posterior to that is enormously long and folded on itself, the vein having left the
mesentery and running a much shorter course than the loop itself. In Nettopus
coromcmdeliamis (fig. 25) the gut is relatively shorter, and an apocentric simplification
Fig. 2b.
Intestinal Tract of Nettopus coromandelianus. Lettering as before.
has taken place. The three proximal loops of Meckel's tract are represented by one
short and then a very long loop, and the loop next distad of the axial loop is contorted.
In these modified loops the veins leave tlie mesentery, a condition comparatively rare
among birds, but very common in the Anseres.
Considering them from the point of vievr of the intestinal tract, it is plain that the
other Anseriformes must be removed from the Palamedese. The latter are archecentrici
The former start from a metacentric position common to the Colymbomorphoe and the
Ciconiiformes, but have diverged apocentrically from that position, forming a specially
Anserine metacentre (Plate 22).
INTESTINAL TRACT OF BIRDS,
205
FALCON IFORMES.
C A T H A R T J^.
CATHARTTn.i=:. — Of these I have been able to examine only Cathartes aura (fig. 26).
The duodenum is long and is coiled into an ii-regular spiral. McckeTs tract presents first
one very Avide loop and then a nearly circular expansion symmetrical about a Meckel's
diverticulum and thrown into short irregular folds. Then follows a very large but
Fig. 2fi.
f
llite.'itinal Tract of CatJinrlcs aura.
Lettpriiip; :is before.
typical supra-duodenal loop with "bridging" vein. The cieca are totally absent, but
immediately above their normal position lies the kink sup])lied by the rectal vein. I
have already shown that this peculiar little loop is a recurring f(;ature among the
Carinates we have been considering. It occurs without exception in all tlie J'alconiform
birds.
SECOND SERIES. — ZOOI.OGV, VOL. VHI.
32
206
DE. P. CHALMEES MITCHELL ON THE
ACCIPITRES.
Gtpogeranid,e. — In Serpentar'ms reiitili corns (fig. 27) the duodenum is simple.
Meckel's tract presents first three narrow minor folds, and then expands into a nearly
circular portion like that in Cathartes, the axis being given by the middle mesenteric
vein which runs from a Meckel's diverticulum. Then follows a long supra-duodenal
loop, drained partially by a " bridging " vein. Posterior to this is a typical supra-csecal
kink, and then a pair of reduced caeca. The rectum is of moderate length and straight.
Fig. 27.
lutestinal Tract of Serpenlarius repfUivurns. Lettering as before.
Vtilttjrid^. — Neophron percnopte^'us (fig. 28) and other Vultures which I have
examined are closely similar. The duodenum is an irregularly expanded loop. Meckel's
tract presents three distinct minor loops (numbered 1, 2, and 3 in the figure), which
reappear constantly in Falconiform birds ; tlien follows an axial loop bearing Meckel's
diverticulum, and then several long, rather irregular loops, the last of which is a typical
supra-duodenal loop with " bridging " vein. The caeca are vestigial, and above them lie
two supra-caical kinks drained by the rectal vein. The rectum is short and straight.
INTESTINAL TRACT OF BIRDS.
Fig. 28.
307
Intestinal Tract of Neophron percnopferug.
1 . 2, 3, three anterior loops of Meckel's Tract. Other lettering as before.
Fat.conid.t;.
(1) GypaelitKP. — Gypohicra.r angolensis {1\^. 29) displays an alimentary tract extremely
like that of tlio true Vultures. The duodenum is simpler, being; a long, narrow loop.
Meckel's tract begins witli tliree distinct minor loojjs (1, 2, and 3) ; then follows an axial
loop, with ^Meckel's diverticulum, and the remaining part of the tract, including the
supra-c;ecal kiuks and the vestigial cieca, is precisely as in the Vultures.
(2) Polybori)}(P. — Tji J'oJyhorm hrmilienals (fig. 30), the Brazilian Caracara, the
duodenum forms a long, narrow loop, which is wound into a spiral. Meckel's tract
consists of a set of small loojis arranged very symmetrically, the axial loop as usual
bearing a MeckeVs diverticulum. There is a well-formed but single supra-c;«cal kink in
the nornial position, and anterior to that a supra-duodenal loop. The caica are vestigial,
and the rectum is short and straight.
(3) Accip'Urincp. — In Virnis cineraceun (fi<j. 31) is to be found what may be regarded as
a central condition lor the Falconidae. The duodenum is a large, very wide loop.
32*
208
DR. P. CHALMERS MITCHELL ON THE
Meckel's tract exhibits first three distinct minor loops, and then is prolonged axially,
bearing at its extremity a Meckel's diverticulum. The posterior part of the tract has one
or two minor loops and a large supra-duodenal loop with two " bridging " veins. Then
comes a large supra-caecal kink. The cseca are vestigial, and the rectum is short and
straight. Another Circus (the species of whicli was not identified), Clrcaeius galUciis,
and Helotarsus ecaudatus all exhibited precisely the same conformation.
Fix. 29.
Intestinal Tract of Gijpoliierax angolennis. Lettering as before.
(4) AquiUnd;. — 1 have examined Aquila anda.r, A. chrysaetus, A. Vei'reaitivi, Morj^hnios
guianensls. Spizacfns coronatiis, and Saliaetns alLicilla and H. leucogaster. I iiave
already described the conformation of the gut in tlie White-tailed Sea-Eagle (26. lig. 12).
The duodenum is A^ery long, being thrown into a complicated system of secondary folds.
Meckel's tract, also, is much elongated and its greater part is composed of a series of
short irregular loops suspended at the periphery of an oval stretch of mesentery, the apex
INTESTINAL TRACT OF BIRDS.
209
Piff. 30.
Intestinal Tract of Pohjhorus brasiliensis. Lettering as before.
Fis. 31.
Intestinal Tract of Clrexs churaeeus. Lctterin;; as before.
210
1)K. P. CHALMERS MITCHELL ON THE
of the system can*ying a Meckel's diverticulum. There is a well-formed supra-duodenal
loop, flhich is spirally twisted, and a large supra-csecal kink has a similar arrangement-
I'lie cac^ca are vestigial, and the rectum is short and straight. Saliaetus leucogaster
differs from the foregoing only in that its duodenum is spirally twisted. These two
birds are large and chiefly piscivorous, and the homojilastic increase in gut-length has
obscured the peculiar cliaracters of the Falconiform gut, but the general symmetry, the
peculiar duodenums, and the well-formed supra-caecal kinks are sufficiently distinctive.
The other Aquilinae repeat exactly the pattern displayed by Circus, the only slight
n^.odification I have found being that the supra-duodenal looji in Spiza'etus, although
(iniined by a " bridging "' vein, is very small. The sujjra-caecal kink is very large in
most and distinct in all.
(5) Buteoitiitce. — Of these I have examined Astur ladiiro, Asturlna magnirostris,
Buieo efythronotiis, JB.ferox, B.jucal, Jlilvus gomnda, M. iciinus, M. migrans. In all
the pattern of Circus is reproduced with close fidelity. The duodenum is irregular,
sometimes long, narrow, and straight as in the genus Buteo, sometimes irregularly
expanded, or folded upon itself several times as in some species of Jlilvus. Meckel's
tract is always as in Circus, save that in a Buteo no trace of Meckel's diverticulum was
retained. The supra-duodenal fold is always pi'esent as also is a large supra-caecal kink.
The caL'ca are vestigial and the rectum is short and straight.
(6) Faiconince. — I have examined Falco concolor, F. Feldeggi, F. lanarius, F. mela-
iiogeiiys, F. jjeregriiius, and Microhierax melauoleucus. The Falconinaj certainly exhibit
Fig. 32.
Intestinal Tinct of Falco iiielanor/enyg.
?'.»!., vise-oral nervc--fhain. Lettering; as before.
the most specialized or apoccntric form of gut among the rakcmidse. The duodenum i.s
always a large irregular loop, sometimes with minor folds (fig. 32), sometimes bent on
it-self (fig. 33). Meckel's tract is always very much elongated in the axial line, and
INTESTINAL TEACT OF BIKDS.
211
invariably bears at its apex an unusually large Meokel's divcrticuluui. In many Falcons
the first portion of Meckel's tract displays the three minor loops which recur among
Ealconiform birds. Jn Falco melanogenys {^^. 32) and F. Feldeggi (fig. 33) two of these
have disappeai'ed, probably in connection with the very great elongation of the tract as
a whole. The apical portion of Meckel's tract may be irregularly twisted as in the
Fijr. ;j;}.
Intestinal Tiaet of Fau-n F,!i/iyi/i. L'.ttcrini; as bcfure.
Peregrine Falcon and in Falco melaiioge)ii/s (fig. 32), or it may be coiled into an
irregular spiral, an apocentric peculiarity found in many specialized types, for instance
in Pigeons and Passerines. There is always a supra-dtiodenal loop and a supra-ciecal
kink. The caeca are vestigial ; in a Peregrine Falcon 1 found only oik; present. The
rectum is short and straiijht.
PandioniD/E. — In the Osprey, Pandion haliaetas (fig. 3i), the gut is enormously long,
and is of very narrow calibre, a modification obviously in association with j)iscivorous
habit. The duodenum is long and narrow. Meckel's tract is thrown into a very large
number of narrow loops, arranged round a nearly circular mesenteric expanse, the
diverticulum being in the usual place at the central [)i)iur, of the curved system. There
is no supra-duodenal loop, but there is a small supra-ciccal kink. The cieca are vestigial
and the rectum is short and straiijht.
The Falconiformes are on the average rather large birds with diet in the main
carnivorous, with some exceptions which are piscivorous, and a few whieii live on
insects. Except in the piscivorous cases, there is little correction to be made for diet.
212
DB. P. CHALMERS MITCHELL ON THE
The carnivorous forms are on the average the larger, and the carnivorous shortening
of the :;ut is in consequence disguised by the relative increase in length associated with
size. The general features of the group are a tendency for the duodenum to be irregular,
eiilar<;ed. very long, or spirally twisted ; for Meckel's tract to exhibit thi-ee definite minor
loops anterior to a median loop bearing a Meckel's diverticulum, and. posteriorly to that,
first one or two irregular loops and then a supra-duodenal loop and at least one supra-
cu'cal kink drained by the rectal vein : the caeca are always vestigial and the rectum
short and straight. The departures from this common type are — first, the irregularities
in tlie fish-eaters ; second, a progressive tendency for the lengthening of ^Meckel's tract
Fig. 34.
Intestinal Trnot of P((iidwn haliaetvs. Letterins: as before.
in the axial line with consequent obliteration of one or more of the otlier minor loops,
and, in the most apocentric ca.ses, with a spiral folding of the tract; third, Serpentarim
shows distad of the first three loops of Meckel's tract a circular expansion of the mesentery
bearing a number of minor loops, and this condition leads naturally to the condition in
('ailiartcs, where the circular expansion involves the second and third of the definite
loops on the proximal side of Meckel's tract. Attempts have been made to show a more
intimate relation between the Cathartte and some of the Ciconiiform birds than between
those and other Falconiformes ; there is no ground for such a conclusion in the structure
of the intestinal tract. Still less ground is there for attempting to place ia intimate
INTESTINAL TRACT OF BIRDS. 213
relation any of the Falconiformes with Cariama. The latter bird, as will appear later,
is definitely a member of the Gruiform assemblage, and for relations between the
Gruiform birds and the Falconiform bii'ds it is necessary, so iar as the characters of
the intestinal tract take us, to go back to the archecentric type underlying all birds.
Summary of the Pelargo-Coltmbomorphine Brigade. (Plate 21.)
At this point it is convenient to attempt a resmne of the conclusions to which study
of the Intestinal Tract has so far led.
Taking the form in Falamedea as the archecentric type, it appears that the Struthious
birds are grouped indifferently around it, as they all display tlie archecentric character
in an unmodified or very slightly modified form. Of the Colymbomorjihaj, Cohjmbus
exhibits the simplest type, in fact a slight but definite modification of the Falamedea
form, consisting in the expansion of Meckel's tract into a set of straight, narrow loops,
one of which is axial and bears the diverticulum, the others being arranged nearly
symmetrically about the middle mesenteric vein. There is usually a supra-duodenal
loop ; the cseca are functional and tlie rectum is short and straight. This form is of
great importance, as it is a Pelargo-Colymbomorphine metacentre from which radiate the
type of intestinal tract disj)layed by Gadow's first Brigade of birds, including the Legion
Colymbomorphae with the Colymbiformes, the Sphenisciformes, and the Procellariiformes,
and the Legion Pelargomorphse with the Ciconiiformes, Anseriformes, and Palconiformes.
The Grebes ai"e more apocentric modifications of this metacentric type, the change being
chiefly a reduction of the numl)er of loops of Meckel's tract with a corresponding increase
in the length of the individual loops. A slight but still more apocentric modification of
the Pelargo-Colymbomorphine metacentre produces a new central position, which may
be called the Steganopod metacentre. The two chief additional characters of this are,
firstly, a tendency to complication and lengthening of the duodenum, a tendency which
in nearly every ease is actually fulfilled ; and secondlj^, the appearance immediately
above the caeca of a kink supplied by the rectal vessel, altliough belonging to the
di"ainage-area of the middle mesenteric vein. Meckel's tract does not differ funda-
mentally (fig. 35) from the condition in the Pelargo-Colymbomorphine metacentre, but
the caeca are reduced. The Sphenisciformes are modified from such a condition only by
the greater length of their gut and consequent increase in the number of minor loops
on Meckel's tract. The Procellariiformes are modified from it chiefly in the reduction
of the number of loops on Meckel's tract and the great increase in length of the
individual loops, the axial of which may be spirally twisted in the most apocentric
forms, e. g., the Oceauitida;. Of the Ciconiiformes, the Steganopods retain theu* meta-
centric position. The Ardeaj start from that position {Scopus), but in the Ardeidse the
minor loops of Meckel's tract become very complicated and one of the two ca;ca is lost.
The Ciconii start from the Steganopod metacentre with forms like Anastomiis, but they
rapidly reach a more apocentric condition, the chief peculiarity of which is the spiral
twisting not only of the duodenum but of the first minor loop of Meckel's tract, and
the twisting of these two spirals together, so that sometimes the blood-vessels are in
SECOND SERIES. — ZOOLOGY, VOL. VIII. 33
i
214
DE. P. CHALMEES MITCHELL ON THE
common. Platalea is less apocentric than the Steganopod metacentre. In it Meckel's
tract cliflFers from that in the Pelargo-Colyrahomorphine metacentre only in that it is
elongated, thrown into many short folds, and sjiirally twisted as a whole. Phcenicopterus
is like Platalea, but it is still more archecentric, inasmuch as its caeca are functional.
The Falconiformes have developed from the Steganopod metacentre so as to display a
central type of their own, characterized by an elongation of Meckel's tract, in the line
of the axial loop, the latter bearing the diverticulum, and the settling down of the gut
Falco
Fig. 35,
Ciconine Types Falconing Vulturid^
Polyborin^
Buteomnffi
Procelkriifornies
Sphenisci
Podicipedes
\
Colymjbi
Gypohierax CatJiarLe?
Serpentarius
Spatula
Pelargo-
Colymbimorphme
Metacentre
Mergus
Archecentric Types
Palamedea
Ratitas
Evolution of Intestinal Tract in the Pelargo-Colymbomorpliine Brigade.
(For Colymbimorphine read Colymbomorphine.)
between this and the duodenum into three definitely formed loops. From this the higher
Falconidse move apocentrically, the axial loop of Meckel's tract increasing in length at
the expense of the others and being twisted into a spiral. Gi/pohierax, with its irregular
minor loops on Meckel's tract and its two supra-caecal kinks, leads to the Vulturidae.
Serpentarius with its spherically expanded Meckel's tract arises somewhat outside the
Falconiform metacentre, and may well lead to the more apocentric Cathartoe, which have
INTESTINAL TRACT OF BIRDS.
215
completely lost the caeca. Paudion stands by itself; its i^ut is so extremely long that
none of the usual minor loops except the supra-caecal kink can be identified. Haliaiitus
is a simple modification of the Falconiform metacentre, tlie irregularity being due to
increase in length. For the Anseridae it is necessary to go back to the Pelargo-Colymbo-
morphine metacentre. Aiiseranas is practically in that position unmodified, save that
the ca;ca are still longer, a condition common to all the Anscrid.Te except some of the
Mergansers. The Cygnus type, with its three definite and contorted minor loops on
the anterior portion of Meckel's tract and its A'ery long axial loop with peculiar blood-
vessels, gives an Anserine metacentre from which Nettoptis and Spatula have diverged
still further. Mergus is probably a pseudocentric modification of the Spatula type.
TINAMIFORMES.
Crypturid^. — Of these I have been able to examine the intestinal tract of seA^ei*al
specimens of JUtyiichotus rnfescens and Nothiira maculosa. The conformation is prac-
tically identical in these two forms. The duodenum is a long narrow loop ; Meckel's
tract {Rhyuchotus rnfescens, fig. 36) is divided into two nearly equal parts, the large
Fig. 30.
Intestinal Tract of liliynclwtus rufescens.
c, compare loop similarly marked in Gruiforraes, figs. 41, 42, 43, &c. Other lettering as before.
Meckel's diverticulum lying between the two. The first portion is one very long narrow
loop ; the second portion is an equally long and narrow supra-duodenal loop drained by a
branch of the middle mesenteric vein as well as by the usual " bridging " factor from tlic
duodenal vein. Meckel's diverticulum was very large in two specimens of Mhynchotus,
small in a third, and very small in Nothura. The small minor looj) just distad of it
33*
216 DE, p. CHALMERS MITCHELL ON THE
represented in figure 36 was present in two specimens of Bliynchotus, absent in another,
and present in Notlmra. The cseca were long in all, and Beddard (2) has notified the
presence of long caeca in the other genera and species, and of very peculiar caeca in
Calodronias. The rectum is short and straight.
The figure may be taken as representing the morphological character of the Tract in
the Tinamus, as there is little correction to be made for habits, and as the conformation
is practically identical in relatively large forms such as Rhynchotus, and relatively small
forms such as Nothura. It is clear, then, that the type of the intestinal tract in the
Tinamiformes is markedly apocentric, and differs from the archecentric type of Pala-
medea and of the Stnithiom birds in that the rectum is straight, and more notably in
that Meckel's tract is expanded into two long, straight, and narrow loops, one anterior,
the other posterior to the rudiment of the yolk-sac, both being orthocoelous. Naturally,
as the Ratites have intestinal tracts of archecentric conformation, it is as possible that
the Tinamiform gut may be derived from the Ratite gut as from that of any other
archecentric form. But so far as the character of the intestinal tract goes, there is no
reason to associate the Tinamus specially with the Ratites. Nor is there any reason
to associate the Tinamus in this matter with the Galliformes, for in these the gut
shows a fundamental archecentricity underlying a tendency to apocentric development
in the direction of expansion of that part of Meckel's tract supplied by the recurrent
branch of the middle mesenteric vein, and there is no trace of this feature in the Tinamus.
The gut of the Tinamus bears a close resemblance to that of specialized Ralline forms
such as Otis (fig. 45). The gut of Otis, as I shall show later, can be regarded without
difficulty as an apocentric derivative of that of less specialized E-alline forms, and for
this reason the resemblance between it and the Tinamu may be merely superficial.
There is, however, another reason for not rejecting the Ralline affinities of the Tina-
miform gut too readily. As I shall explain later, I have only gone a little way in study
of the autonomic nervous system, and therefore cannot yet weigh the value of evidence
to be drawn from the modifications exhibited in different birds. There are, however, at
least two strongly contrasting types. In one of these the visceral nerve of Meckel's
tract forms a ganglionated chain which follows the contour of the gut. This type
occurs in Falamedea (fig. 1, v.n.), in the Struthious birds, and in the Galliformes. In
the other type there are a much smaller number of larger ganglia on the visceral nerve
{Otis, fig. i<5, v.n.). This type occurs in some of the Ralline birds and in at least
Nothwa. Another striking resemblance is that between the conformation of the gut
in the Tinamus and in Opisthocomus. To this I shall recur later. The conformation
of the gut, then, supports those who, like Gadow, isolate the Tinamus as Tinamiformes,
but at the same time suggests possible relationships with Gruiform birds and with
Opisthocomus.
GALLIFORMES.
Mesites.— I have no information regarding this group. Naturally knowledge of the
conformation of the alimentary tract is much to be desired in the case of a bird the
afiinities of which are doubtful.
INTESTINAL TEACT OF BIEDS.
217
TuRNiCES. — In the Tumices (I have-not seen Pedionomus) the duodenum {Turnix
JDussumieri, fig. 37) is a long, narrow loop ; Meckel's tract is suspended at the periphery
of a nearly circular mesenteric expansion and bears no trace oi the diverticulum.
There is no specially formed supra-duodenal loop, but the very large caeca (which are
dilated towards their extremities) are drained partly by a bridging vein from the
duodenal vein. The rectum is relatively long and very slightly expanded into folds.
Fig. 3'
f
Intestinal Tract of Tnrnix Dussumieri.
)•., recurrent factor of middle mesenteric vein. Other lettering as before.
This intestinal tract is markedly archecentric. The simple character and the arrange-
ment of the blood-vessels are closely similar to the condition of Palamedea. The chi&f
modifications are absence of the Meckel's diverticulum, the increased length of the caeca,
and the relatively shorter rectvim. So far as the character of the gut indicjites systematic
position, there is no reason either to retain Turnix among the Galliformes or to remove
it from that assemblage. It is simply an archecentric form.
218
DE. P. CHALMERS MITCHELL ON THE
G A L L I.
Megapodidje, — In Talegallus Lathami (fig. 38) the duodenum is a straight, narrow
loop of moderate length. Meckel's tract is swung at the periphery of an expanse of the
mesentery, and forms almost a complete circle of which the middle mesenteric vein,
running from the Meckel's diverticulum, forms a diameter ; for the greater part of its
Fig. 38.
Intestinal Tract of Talegallus LafJiaim. Lettering as before.
course this tract is thrown into very regular corrugations, but its distal portion is nearly
straight. The recurrent factor of the middle mesenteric vein supplies a considerable part
of Meckel's tract. The coeca are of moderate length and are drained partly by a
"bridging" vein, and the rectum is fairly long and very slightly thrown into folds. A
striking feature in Talegallus, as in other Galli, is the conspicuous visceral nerve (u.«.),
which in this bird is a ganglionated chain following the sweep of Meckel's tract.
Ckacid^. — In Crax Datibentoni (fig, 39) the only notable difference from Talegallus
in the formation of the gut relates to Meckel's tract. This area is much elongated
distally, with the result that tbe recurrent factor of the middle mesenteric vein (r.) which
supplies the distal portion of the tract is much enlarged, and appears to be the direct
continuation of the middle mesenteric. That rein, however, in the embryo runs out as
usual to the diverticulum as I figured in the case of Argus glgantews (26. fig. 2), and its
real termination in the adult is one of the minor branches of the main blood-channei.
INTESTINAL TRACT OF BIRDS.
219
In cxssociation with the growth of the i^osterior portion of Meckel's tract, the visceral
ganglionated chain forms a curious elongated loop, which, on comparison of figs. 38
and 39, will be seen to he obviously connected with the distal growth of the tract. In
two other species of Crax and in two species of Penelope the conformation of the gut
with its blood-vessels and nerve was identical in all main points with the condition just
described.
Intestinal Tract of t'rax Dimhentoni. Lettering; as before.
Galled^. — I have examined a considerable number of these, including species of
Argus, Caccabis, CalUpepla, Coturnix, FrancoUniis, Gallns, Lophophorus, Pavo,
Phasianus, Perdix, and Tetrao, and in aU the conformation closely resembles that
shown in the figure of Pavo cristatus (fig. 40). Meckel's tract is more elongated in
the axial line than in the Cracidse, and the middle mesenteric vein runs obviously from
the remains of the diverticulum. There is a similar distal prolongation of the tract
drained by a large recurrent vein, and to the straight distal portion of this the very
220
DE. P. CHALMERS MITCHELL ON THE
large cseca are closely applied, and are drained partly by the recurrent vein and partly
by a " bridging " vein. The rectum is relatively short and straight.
In the Galli generally the only homoplastic cause of apocentricity that has to be
allowed for is the increased length in the larger forms, and this is not sufficiently great
to distort the morphological pattern of the intestinal coils. In all, the general character
is markedly archecentric ; in the Megapodidaj the archecentricity is most definite. In
the Cracidaj there is an apocentric modification of a definite nature, although not great,
and consisting in the peculiar expansion of the distal region of Meckel's tract with
Fia;. 40.
Intestinal Tract of Favo cristatus. Lettering as before.
coincident alteration of the visceral nerve and of the factors of the middle mesenteric
vem. In the Gallid;i3 a similar change has occurred, but has affected the blood-vessels
and the visceral nerve (the course of the latter, not indicated in the figm'e, very closely
resembles that in Talegallus) to a much smaller amount. On the other hand, the caeca
in the Gallidse are much larger, and the small factor of the duodenal vein, which in Crax
runs from them within the mesentery, is enlarged in the Gallidae and has broken through
the mesentery. It is plain that, so far as the conformation of the intestine indicates, the
Galli are all closely allied, but there is a distinct basis for their division into Megapodidse,
Cracidse, and GaUidoe.
Opisthocomi.
I have already described and figured the intestinal tract of Opisthocomus cristatus
(27. fig. 1). The duodenal loop is short and wide. Meckel's tract is thrown into three
well-marked narrow loops, the second of which is rather wider and has a tendency to be
twisted into a very slight spiral. In a chick and in two adults I found no trace of
INTESTINAL TKACT OF BIEDS. 221
Meckel's divei-ticulum, but towards the extremity of this loop the place of attachment
of the yolk-sac was marked hy a strong remnant of a ventral mesentery. In a third
adult this mesentery ran to a very small vestige of Meckel's diverticulum. The most
distal loop of IMeckel's tract was wide, and closely applied to it were the pair of large
caeca, drained partly by a "bridging" factor of the duodenal vein. The rectum is
unusually long, retaining the archecentric condition of being thrown into minor folds.
It is plain that the condition of the intestinal tract in this group is markedly apocentric,
except with regard to the rectum. From this point of view the Ojjisthocomi do not find
a natural place among the Galliformes — first, because in these latter the general condition
of the gut is archecentric ; and second, and more important, the small degree of apocen-
tricity displayed among some of them consists of a peculiar expansion of tlie distal
portion of Meckel's tract, and there is no trace of this feature in Opisthocomus. The
apocentricity of Opisthocomus may be compared first with that of the Tinamus. If
the very small loop seen just distal to Meckel's diverticulum in H/ii/iichofiis {&g. 3G) were
])rolonged, carrying with it the diverticulum, the condition in Opisthocomus would be
reached. A second suggestion of relationship is, as I have already pointed out, given by
comparison with the form of the tract in Ftei-ocles and the Pigeons. In Pterocles
(26. fig. 18) tlie condition of Meckel's tract and of the long caeca is extremely like that in
Opisthocomus. the most notable diff'erence being that in Opisthocomus the axial loop bearing
the diverticulum is slightly twisted. In the Pigeons (26. fig. 19) the twnsting of the axial
loop may be carried much further and the caeca are reduced. AVhen I first made this
comparison of the gut in these three groups, I accejited the common view that eutaxy or
quintocubitulism was a primitive condition, and diastaxy or aquintocubitalism a derived
condition, and it was with hesitation that I jjvit Opisthocomus, an eutaxic form, between
Pterocles, a diastataxic form, and the Columbidfe, then believed to be diastataxic. Since
then I have shown that the eutaxic condition is probably a muhiradial apocentricity
derived independently from the diastataxic, archecentric condition of the wing. Moreover,
I have shown (28) that among the ColumbidEe eutaxic forms occur. There is therefore
no difficulty in the way of supposing that the gut forms in the I'igeons and in Opistho-
comus are derivatives of the condition in Fteroclcs. In the paper in which I described
the intestinal tract of Opisthocomus (27) I pointed out also that there were resemblances
between the gut of Opisthocomus and the gut ot the Cuculidui, thereby recjilling Garrod s
(13) suggested relationship between Fowls, Opisthocomus, and Cuculidae. It is true that
among the Cuculidae Meckel's tract is thrown into three loojis, but an examination of a
larger number of Cuculidae has shown me that important differences distinguish the three
loops in Opisthocomus and the loops in Cuculidae.
Taking the Galliformes as a whole, it appears that tlie form of the gut in the Turnices
is archecentric; in the Galli it is still archecentric, but with a tendency to a special mode
of apocentricity ; in the Opisthocomi it is markedly apocentric, and the apocentricity is
quite different in kind from that found among the Galli, but with marked resemblances
to the condition in Fterocles and the Pigeons.
SECOND SEBIES. — ZOOLOGY, VOL. VIII. 84
•)0')
DR. P. CHALMERS MITCHELL ON THE
GRUIFORMES.
Eallid^. — lu Ocydi'omus australls (tig. 11) what J. find to be the typical Ralliue
conformation of the intestinal tract is presented. The duodenvim is a straight, narrow
loop of moderate length. Meckel's tract is dra\\ n out into a definite number of Iooj)s,
all of wliich are fairly straight. The first of these, marked " a " in the figures, succeeds
the duodenum; the second (" lO " in the figures) is axial and bears on its distal side a
hirge Meckel's diverticulum. The third, marked " r," like the others is a narrow loop
belonging to the drainage of the middle mesenteric vein ; and the fourth is a well-marked
supra-duodenal looji drained by more than one bridging factor of the duodenal vein. Tiie
FiK. 41.
-X.2.
Intt'stinal Tract of Ocytlrumiis idistralis.
a, /), c, the three h)(ip.s of MeckeFs Tract characteristic of the Gruiibrrue.-;. Other lettering as before.
long cicca arc; closely attached to this last loop of Meckel's tract. The rectum is straight
and of moderate length. Tliis conformation is found in all the members of the Rallida^
that I have examined, e. g., Aramldes ypecaha, Crew pratcnsis, (jullinula phoenicvni,
fon<)rii.i.s mariinicus, Porpliyrio cceriile^is, and P. polioccphalus. I described this condition
of 11k^ gut correctly in a former paper, but in the drawing of Crex (26. fig. 13) the third
iiiinoi- l()(»ji has been omitted by an unfortunate mistake, so that the description does not
INTESTINAL TRACT OF HTKDS.
223
f.illy with tlie figure. The diflerenecs which occur among the RallidjE are insignificaTit ;
in some, there are two " hridgiug " veins, in some, one; tlie Meckel's diverticulum, which
is always large, may he extremely large, the size of this structure heing a characteristic
feature of tlu; group ; the tlurd subsidiary loop of Meckel's tract occasionally is relatively
smaller than the other loops.
(ilUMD.i:.
(1) Gruiiuc. — In Griis virgo (fig. 12) and other species of (Iras, and species of
Avlhropoidvm and Balcarica are practically idcMitical, the large size has biouglit witli
it a relatively great increase of length, hut the fundanumtal ideutity of the coiilorniation
of the gut with that e.\hil)ited by the Rails is obvious. The duodenum is similar,
i-'is. 42.
liitcstiiinl 'I'riu t of GrMn v'ii-(\i). Lclteririf; as in tifr. 41.
although, on account of its great length, it is partly bent. Meckel's tract displays the
same four loops : " a " being widened out ; " i," the axial loop Ijcing very loni;- hut carrying
a Meckel's diverticulum in 1 lie characteristic ])osition, on the distal linil) of the loop;
"r" being small but compound; and the supra-duodenal loop to which the huig cteca
are attached being much enlarged. It is plain that we have here a simple modification
of the Ralline pattern, the modification being due to the relatively greater length of gut
tliat occurs in larger birds.
(2) Armnuio:. — The very interesting form Ara)}i,ns scolopuceus ilig. I.')) displays a
conformatiou of the intestinal tract which differs only in minor respects from that in the
221;
DR. P. CHALMERS MITCHELL ON THE
Rails and Cranes. The duodenum is normal ; Meckel's tract displays the loops which I
have marked "a" and "6" pi-ecisely as in the Rails, and "6" the axial loop bears
a large diverticulum on its distal limb. The posterior portion of the tract differs : in
place of "c," tlie third loop, and of the normal supra-duodenal loop, there are a set
of irregular small loops. The caeca are short but wide and in the natural condition
contain faecal matter. The rectum is short, wide, and straight. I am disposed to think
that the type ia Aramiis is more archecentric than the types displayed by the Cranes and
Rails. Although the axial loop with the diverticulum on its lower limb is character-
istically Ralline, the general conformation of Meckel's tract is much more like the
Kg. 43.
Tn
Intestinal Tract of Aramus sMlopaceus. Lettering as in fig. 41.
archecentric condition, and this similarity is increased by the moderate length of the
caeca and the absence of a specialized supra-duodenal loop. I think it is more probable
that the long caeca of the Cranes and Rails are an apocentric modification than that the
relatively shorter caeca of Araimis are pseudocentric degenerations, for, as I hope to show
later, in the vast majority of cases where the caeca are obviously degenerate, the
"bridging" factor or factors from the duodenal vein, which originally drained them,
persist to drain a specialized supra-duodenal loop, and there is no trace of this in
Aramus.
A
INTESTINAL TEACT OF BlllDS.
225
(3) P&op/i?iti(P.— As the position of Psojjhia is one concerning which there has been
LO little difference of opinion, it is interesting to find that the conformation of the gut
in Fsophia crepitans (fig. 44) is typically Ralline. As will be seen, it conforms in every
•way to the character typical of the Eallidse. As it is probable that the differences
between the type in the Cranes and that in the Rails is due merely to the laro-er size of
the former, there is no reason to be drawn from the character of the gut for placing
Fsophia preferentially either with the Cranes or with the Hails. It displays the "round-
form common to both sets. Psophia obscura is simplified in the direction of Cariama
and the Bustard ; the loops on either side of the axial loop, i. e. " i
practically disappeared, so that the conformation in Otis is closely imitated
Fig. 44.
and " c," have
Intestinal Tract of Psophia crepitans. Lettering as in fig. 41.
DiCHOLOPHlD^. — I have already described and figured the intestinal tract in Cariama
cristata (26. fig. 14), and the condition in Burmeister's Cariama {Chungo Biirmeistcri) is
practically identical. The duodenum is a long narrow loop. Meckel's tract displays
the Balline loops " a " and " 6 " ; but the two are merged proximally, " b " displaying a
Meckel's div erticulum on the distal limb. There is no trace of loop " c " ; but there is a
large supra-duodenal loop drained partly by a " bridging " vein, and having the long
caeca closely applied to it. The rectum is straight but of fair length. It is plain
that the intestinal tract of the Dicholophidaj presents an apocentric character of a
definite nature, but which may be regarded as a simple derivative of the type shown in
226
DR. P. CHALMERS MITCHELL OX THE
the EallidaB and Gruidac, the loops of Meckel's tract being long and straight, the first
and second being partly fused and the third having disappeared.
Otidid-E. — I baAe had the opportunity of examining only Otis tarda, the Great
Bustard. In it (fig. 45) the duodenum is a long narrov\- loop. Meckel's tract is thrown
only into two loops, of which the second is a very large supra-duodenal loop drained by
a pair of large bridging vessels, and has long and peculiar caica closely applied to it.
The proximal loop 1 take to be the axial loop seen in the Rallidae ; it bears upon it, low
Fis. 45.
r\:o.
u n
SOF
Intestinal Tract of Otis tarda.
n.ti.. nerves entering mesentery; v.u.ic.. gangJion in d\:odenum from -which nerves pass to caeca and snpra-
duodenal loop; v.n.g., large ganglion of visceral nerve; v.n., branch of visceral nerve in rectum.
down on its distal limb, a Meckel's diverticuluni. A ] air of very small but quite
definite folds, marked " r? " and "c" in the figure, I take to represent the similarly
designated large loojjs in the Rallidtie and Gruidse.
The intestinal trac't of Otis is certainly markedly apocentric. The loops are all very
definite and the supra-duodenal loop in jiarticular is highly specialized, being closely
applied to tlie duodenum, over which it lies in the unfolded condition, and from Avhich
it recei\es not only a pair of bridging veins, but t\Ao branches of the autonomic visceral
TNTESTINAf, TRACT OF FilKDS.
227
nerve roniinf^ from a special gang-lion in tiie duodena] loop (lig. 45, r.n..r.). The peculiar
caeca wiiicli have becji described l)y J^eddard (2. p. ."332) are specialized not only in their
^rcat length l)nt in liieir internal structure. The general character of the gut, however,
especially if 1 am correct in my interpretation of the small folds marked '• « " and " c,'
is obviously Kalline. The resemblance betAveeu the guts of 0//s and of the Tinaraus is
certainly striking, the only notable difference being that in the Tinamu the ciecji are
less specialized and that Meckel's diverticulum lies distad of the great loop of Meckel's
tract rather tiian actually on it. I regard tlu; closeness of the resemblauce as not
definitely m()r[)h()logical.
Rui.\o(;iii;tid.k. — lu Rhinochetus jnbalua, the Kagu (lig. 46), the duodenum is
Fig. 40.
Intestinal Tract nf KluHucIirtns jnhatn.-:. I.ctteriiig as in fij^. 41.
a simple, short, and narrow loop. Meckel's tract slunvs tirst a pair of minor loops, tin;
second of wliich is double, then an axial loop uith a large diverticulum about the middle
of its distal liml), then a large .simple loo]) markinl " r," then ;\ long narrow supra-
duodenal loop with a bridging vein. The cteca are rather short, and the rectum is
straight, wide, ami of moderate length. The intestinal tract of this bird certainly dilfers
considerably from that of other members of th(> Gruiform assemblage. The most
important diffi^rences are the presence of an additional loop on Meckel's tract, the
circumstance that the whole of that tract, with the exception of the supra-duodenal
228
DE. P. CHALMEES MITCHELL ON THE
loop, displays in the unfolded condition a slightly spiral twist, and the shortening of the
caeca, a character shared with Eurypyga. The resemhlance to the Ralline type, how-
ever, is fairly strong. Meckel's tract is produced into a set of minor loops, of which
two represent " a," while " 6," with the diverticulum midway on the distal limb, and " c "
are normal, and the last is a supra-duodenal loop simplified in correspondence with the
reduction of the caeca. Bhinochetus has been compared with Scopus, but the character
of the tract does not support this comparison. In Bhinochefm there is no trace of the
twisted duodenum, the diverticulum is in the Gruiform position, and Meckel's tract ia
thrown into definite minor loops.
EviiY-BYGUiM.— Eurypyga helias (fig. 47) departs stUl more from the other Gruifornies.
Fig. 47.
Intestinal Tratit of Eiirijpyja helias. Lettering as before.
The duodenum resembles that of the other forms, but Meckel's tract is much more
archecentric, the greater part of it consisting of an irregularly folded gut swung at the
periphery of an oval expanse of mesentery. The large diverticulum, however, is not at
the apex of the tract, but in the typical Gruiform position on the distal limb of the axial
loop. The distal portion of Meckel's tract is a definite supra-duodenal loop with
bridging vein. The caeca are vestigial, and the rectum is long and nearly straight.
Hkliornithid^. — In Ilelioruis fulica (fig. 4S) the typical Ralline or (iruiforra
characters again appear. The duodenum is simple and narrow. Meckel's tract displays
INTESTINAL TRACT OF BIRDS.
229
the three loops "«," "6," and " c " very definitely drawu out. In II. surlnumciisis, the
coni'ormation is similar, although the loojis are not so indepcndoTit. In neither is thei'e
a diverticulum present. The jiosterior portion of Meckel's tract is a definite supra-
duodenal loop drained by two " bridging " vessels. The caeca are of modci-ate length, and
the rectum is as in the others.
It is plain that what I term a metacentral condition underlies the conformation of the
alimentary tract in the Gruiform asscmlilage. This mctacentre possesses a simple
duodenum ; Meckel's tract is produced into four straight loops, of which " h" the axial
Fig. 48.
Intestinal Tract of Heliornis fuliai . Lottoring as in fig. 41.
loop, carries the diverticulum on its distal liml), and of which the last is a large supra-
duodenal loop drained by bridging vessels and with the long caeca closely attached to it.
The Rallidaj and Gruidae display this metacentral cliaracter without modification of any
important kind. The Dicholophidte and the Otididae have moved apocentrically from it
by reduction of loops " a " and " c " and greater specialization of the supra-duodenal
loop. Helioniis is metacentral. llhhtochetus and Eurypy(ja are slightly modified, the
apparent archecentricity of the latter being possibly pseudocentric.
SECOND SERIES. — ZOOLO&Y, VOL. VIII. 35
230
DE. P. CHALMEES MITCHELL ON THE
CHAEADRIIFOEMES.
L I M I C O L iE.
CHAKADRiiDiK. — I bave already described and figured tbe conformation of tbe intes-
tinal tract in the Curlew {Numenms arquata, 26. fig. 15). Tbe duodenum is straight
and narrow. Meckel's tract is more archecentric than in the Gruiformes, inasmuch as
the minor loops arc not well separated from tbe general course of the gut. None the
less, such minor loops exist, and markedly recall the common Gruiform type, altbougli
Ibc condition is less apocentric. There are three main loops corresponding to " «,"' " b,"
and " c " of the Gruiform gut, and, as in that assemblage, " b " the axial loop bears on
its distal limb a very large Meckel's diverticulum. The distal part of tbe Tract consists
.Fix. 49.
Tntestinal Tract"- of Tringa alpina. Lettering as in fig. 41.
of a supra-duodenal loop not well separated from the general outline of tbe Tract, but
drained by a " bridging" vein, and having closely attached to it the pair of long caeca.
The rectum is rather short and straight. Tbe condition in Himantopus is similar to this
but still more archecentric — that is to say, the loops are still less marked off from the
general sweep of Meckel's tract. Vanellus milg aris and V. cayennensis are like Numenius.
In Tringa alpina (fig. 49) a simple modifica tion of the Ktmieniiis condition is presented.
Loop " b " of Meckel's tract has grown out axiallv, leaving the large diverticulum at its
INTESTINAL TRACT OP BIRDS.
231
base, and the supni-duodenal loop is better separated. la Beouri:/ rostra acocelta a still
more a})ocentric modification in the same direction is reached. Tlie axial loop lias
grown outwards to more than twice the length attained in Truiga, and is folded irregu-
larly as a flat ribbon, partly rolled on itself and partly spirally twisted. The c;T)ca
remain long, and the other parts of the gut arc as in Trim/a. A still greater apocentrit;
divergence of precisely the same kind occurs in Scolopax rusticolu (Plate 22). I have
already figured the very remarkable conformation assumed by its gut (26. tig. 16). The
Fig. 50.
Intestinal Tract of Chionis alha. Lettering as in fig. 41.
change is simply that the axial loop has grown still longer than in llecurcirostra, and
the rolling-up and spiral twist is still better marked. The caeca have degenerated, and
have left isolated a small supra-duodenal loop with a bridging vein.
QinoyiiD.v..— Chionis alba (fig. ,jO) shows a comparatively slight modification of the
Xumeniiis type, and is plainly much more archecentric than most of the Charadriida}.
Meckel's tract is relatively longer; the axial fold with its large Meckel's diverticulum is
35*
232
DE. P. CHALMERS MITCHELL OX THE
Fie:. 51
Intestinal Tract of Glareola ocularis. Lettering as in ti''. 41.
Intestinal Tract of Thinocorys rumicivorus. Lettering as in fig. 41.
INTESTINAL TKACT OF BIRDS.
233
practically identical with that oi Numenius; but the fold"«"is represented by two
loops, and loop "c" is partly fused with, or not separated from, the very large supra-
duodenal loop to which tlie long cneca are attached.
Glaheolid.i:. — In Glareola ocularis (hg. 51), and 6'. pralincola is practically identical
in this matter, another very simple modification of the Numcnlm type is displayed.
The duodenum is similar; Meckel's tract is relatively shorter, and is thrown into Ioojjs
" <(," " W" with a small Meckel's diverticulum at its base, and a large and wide supra-
duodenal loop, loop "c " not being formed. The ctcca are attached in the usual way to
the supra-duodenal loop, and the rectum is short, wide, and straight.
Fifr. 53.
XI.
Intestinal Tract (if Parri
•I'll jdcana.
Lctturinn; as iu tiir. 41.
Thixocokid.e. — In Tli'mocon/s ruuiicivorus (tig. 52) a condition essentially similar to
hat found in Glareola is displayed. The duodenum is similar ; Meckel's tract is
uispended at the end of an oval stretch of mesentery, and in it loop " b " wit li (he
liverticulum at its base is obvious, although in the drawing it is represented as turned
ip instead of running out in the axial line. Loops " a " and " c " are not differentiated,
nd the supra-duodenal loop to which the long caeca are attached is not sharply
eparated off except at its apex. The rectum is straight, wide, and of moderate length.
Q^]dicnemid_1]:. — In CEdicnemus scolopad-, the Stone-Curlew, there is displayed a con-
irmation closely similar to that found in Tkinocori/s and Glareola. The duodeninn
234
DE. P. CHALMERS MITCHELL ON THE
is straight and simple, although, rather long. Meckel's tract displays a small loop " «,'
then an axial loop "i " with Meckel's diverticulum on its distal limb ; loop " c" is not
differentiated from the supra-duodenal loop, to which the long caeca are attached. The
rectum is straight. It is plain that CEdicnemus is more archecentric than the ordinary
Curlews, and that its nearest allies are the Glareolidae, Parridse, and Thinocoridre.
Parrid-E. — In Parra jacana (fig. 53, p. 233) the same ground-form is obvious, althougl:
Intestinal Tract oi Farjoi)hila elmrnca. Lettering as before.
there is a good deal of individual specialization. The duodenum is similar. Meckel's
tract exhibits loops " a " and " b" the latter with the diverticulum at its base. There is
no trace of loop " c," the distal portion of Meckel's tract appearing as a single very
large supra- duodenal loop supplied by two " bridging " veins and with the proximal
portion of the gut unusually dilated. The ctcca are vestigicxl, and the rectum is short and
straight. Except for the vestigial caeca, Parra is very like CEdicnemun.
INTESTINAL TEACT OF BIRDS.
235
It is plain that in the Limicola? there is displayed a uround-form very close to what I
described as the Gruifomi metacentre, but rather more areliecentric than that. Prom
this condition the Charadriidte exhibit a striking set of progressively more apocentric
moditications, reaching a climax in tlie conformation of the gut in Scolopax. The otlicr
iM<?. ;)o.
"X
Intestinal Tract of Slercoriirii's crejiidiitus. Lettcriug as before.
families, the ChionidiX', Glareolidae, Thinocoridoe, ffidicnemidiX', and I'arridyu, all display
types closely allied to tliat of Xiimenins, but still simpler. In other words, a form of
srut like that in Numenius is a metacentre common to the Gruiformes and the Limicohu :
from this the Gruiformes have diverged in one direction, the specialized Charadriida' in
another, while the other families cluster about the metacentre.
236
DE. P. CHALMERS MITCHELL ON THE
L A K I.
Larid.e. — I have already described and figured the conformation of the gut in Lams
mart litis (26. fig. 17). Tlie duodenum is narrow and straiglit. Meckel's tract is thrown
into a series of irregular loops, the first of which prohahly represents " «," while " h "
appears as a number of closely-set short loops, proximad of the diverticulum. " c " and
the large supra-duodenal loop are both present. The caeca are vestigial, and the rectum
Fig. 56.
Intestinal Tract of Slerna Inrundo.
is rather long. Other specimens of Lcmis marinus were identical, except that Meckel's
diverticulum was absent, a condition frequent in the Gulls. Larus argentatus was
closely similar, and L. ridibundus differed only in that the loops of Meckel's tract were
more definite and were twisted into a common spiral. In Tmjopldla churnea (fig. 54,
p. 231') the general conformation is similar. Loop " c" and the supra-duodenal fold are both
INTESTINAL TBACT OF RIKDS.
237
larger. However, the most striking circumstance is the appearance above the ca;ca in
the posterior part of Meckel's tract of a set of kinks drained by the rectal vein. In the
drawing, the relative importance of these has lieeii slightly exaggerated ; but I was
anxious to call attention to them, as their presence suggests either an unlikely ailinity
between the Larida^ (and the Alcidae) and the Pelargo-Colyml)oraorphine assemblage, or,
more probably, that the existence of tiiis kink is a miiltiradial apocentricity to which too
much importance must not be attached. In Slercorarlm crepidatns (tig. 5o, p. 2:55) a
Fi-. .17.
rnte>tinal Tract of Frnlercula arclicii. Lettering as in tij;. -11.
more archecentric type of gut is displayed, the arrangement rather closely resembling
that found in Thinocor^is and Glareola. The duodenum is simple. Meckel's tract exhibits
two loops proximad of tlie diverticulum, while distad of it is a vei-y Avide area re])re-
senting a supra-duodeual fold and an undifferentiated loo]) " c." The cteca are long aiul
the rectum is straight. In Sfcrna hlrundo (fig. 50) the conformation is more apocenlric,
and the pattern reseml)les that Ju many of the specialized (Iruiform and Charadriiform
types. Meckel's tract is thrown into three minor loops, which, however, are better left
SECOND SERIES. — ZOOLOGY, VOL. VIII. 3C
238
DB. P. CHALMEES MITCHELL OX THE
unidentified, as the diverticulum is not present, and there is a highly specialized supra-
duodenal loop left free hy the degeneration of the caeca and closely rcsemhliiig the
duodenum in shape. This similar moulding of the duodenum and the supra-duodenal
loop is a condition found in many of the higher forms Avith relatively short guts.
Alcid^. — In Fratercida arctica (fig. 57) a conformation of the same general character
as that in many of the Gulls appears. The duodenum is of moderate size and not
twisted. The greater part of IMeckel's tract consists of a set of sliort loops, the axial
of which bears a large diverticulum in the Ralline position, and of a very large supra-
Fis. 58.
ln(os(in;il Tract of Loinriit troilc. Lettering as before.
duodenal loop from which there has not been separated loop " c." Above the vestigial
caeca is the curious supra-cjecal kink to which attention was di*awn in certain Gulls.
In Lomvia iro'de (fig. 58) a Gull-like pattern is displayed. There is no diverticulum to
jjulicatc the exact topography. There is a well-formed supra-duodenal loop and a large
supra-oa>cal kink. The caeca are vestigial, and the rectum is extremely short.
The Larida^ and Alcidte are families of birds typically lish-eating, and in which
lengthening and narrowing of the gut has to be allo^^ (>d for. That alloAvance having
INTESTINAL TRACT OF BIEDS. 239
been made, the various conformatioBs exhibited by the dififerent genera of both families
(from the point of view of conformation of the gut no distinction can be made between
the families) appear obviously to belong to the simpler RaUine and Charadriiform
patterns. The only other possible suggestion of affinity is provided by the existence of
the supra-caecal kinks in some Gulls and Hawks ; but, as the other portions of the gut
are so unlike the conformations found among the Pelargo-Colymbomorphine assemblage,
I think the presence of the kinks in these few forms must be taken as a multiradial
modification.
Pleroclo' Columbine Assemblage.
Pterocles. — I have already described and figured the gut in Ptentcles blcmstins (26.
fig. 19). The duodenum is straight and rather narrow. Meckel's tract is expanded to
form three definite loops. The first of these is long, expanded at its apex, and curiously
bent on itself. The second is an axial loop bearing Meckel's diverticulum at its apex.
The third is a deSnite supra-duodenal loop, drained by a '" bridging " vein and having
the long caeca closely applied to it. The rectum is short and straight. In Plerocles
alchata the condition is exactly similar, except that the caeca are relatively longer.
C 0 L r M B .£.
I have examined a large number of Columbidse, including the genera Calcenas,
Chalcopelia, Chalcophaps, Columba (many species and varieties!, Columbula, Geopelia,
Goura, Leucosarcia, Ocyphajjs, CEna, Fhops, Phlogcenas, Ptiloptis, Stanifjenas, Trei'on,
and Turtur. The conformation of the gut is best understood by taking it as a simple
but more apocentric derivative of the condition in Pterocles. The duodenum is a
simple loop. Meckel's tract is thrown into the same three folds. The first is simpler
than in Pterocles. The second or axial loop is usually very much longer, and it may be
twisted into an elaborate spiral (26. fig. 19j, and as in Pterocles bears the diverticulum,
or a very small vestige of it marked by a ventral mesentery, or no vestige at all.
The third loop is a definite supra-duodenal loop very closely modelled on the duodenum
and diained by a "bridging' vein. The caeca are always vestigial and not infrequently
are absent. Beddard (2. p. 308) mentions that in an example of Tympanistria btcolor
only one vestige was present. I have found similar variations not infrequently in the
varieties of the genus Columba. but apparently only as individual variations. The rectum
is alwavs verv short and straisht.
There is little difference in the conformation of the gut in the different genera and
species. Tlie larser forms in most cases have the length of gut relatively greatly
increased, and the increase is most noticeable in the spirally twisted axial loop. In
some small forms, such as (Ena, the gut is absolutely and relatively shorter, and there
is little trace of the spiral. Precisely a similar state of affairs exists among Passerines,
where size is one of the chief factors in determining the length of the whole gut, and the
consequent elaboration of the axial spiral. In Pliloptis and other fruit-eating Pigeons
the whole gut is extremely reduced in length and is very wide. In sucii cases the
relation of the pattern to that in ordinary Pigeons can only be guessed.
36*
■2i0 BR. P. CHALMERS MITCHELL ON THE
Ei'om the point of view of couforniation of the iutestiual tract, it does not a])peai' that
the Pteroclo-Columbae stand in close relation to other Charadriiforni birds. The only
character in common is the tendency for Meckel's tract to be drawn out into long minor
loops, and tlie particular apocentrieity which reaches its climax in the elaborate spiral of
the axial loop of Pigeons is different from that in the specialized Charadriid:e such as
llecurvh'Ofttra and Scolopax, the difference being made plain by the relative positions of
Meckel's diverticulum. The comparison with Opisfhocomus seems to give a more fertile
suggestion.
Summary op the Alectoeomorphine Legion. (Plate 22.)
Gadow unites the Tinamiformes, Galliformes, Gruiforines, and Charadriiformes into
the Alectoromorpha3, the first Legion of his second Brigade, and I 2)ropose now to review
shortly the characters of the intestinal tract j^resented in this large assemblage. It must
be clear that the conformations of the gut within this group do not follow many of the
more generally accepted modes of subdividing the group so readily and naturally as in
the case of the Pelargo-Colymbomorphine Brigade. Those, I tliiuk, who have paid most
attention to the classification of birds will he prepai-ed for this result, for the many and
different divisions proposed suggest that the assemblage is still in indifferent order.
Prom the pohit of view of the subject of this Memoir, it is in the first place clear that
the Turnices and Galli stand apart from the other gronj^s, inasmuch as their pattern of
intestinal tract has remained markedly archecentric, differing extremely little from that
in Palamedea. Next, the Tinamidse retain no suggestion of Struthious affinity ; they
have moved ftir apocentrically, while the Batites, like the Galli, have remained close to
the archecentric condition. The Tinamidaj present on the one hand most striking
resemblances witli the Otididue, an apocentric type of the Gritiformes. On tlie other
baud, the radius of apocentrieity on which they lie suggests that of Opisthocornus,
Pterocles, and the Columbse. Opisthocornus, one of the Galliformes of Gadow, certainly
stands far apart from the others, and, so far as the gut is concerned, an affinity with
Pterocles and the Columbae is most clearly indicated. The guts of Columba3 and of
Pterocles are extremely similar, and there is no indication of any alfinity Avith the
conformations exhibited in the other Charadriiformes or Gruiformes.
The Galli, Turnices, Oplsthocomm, and Pteroclo-Columba' having been disposed of,
there is less difficulty in arranging the confontiations of the gut in other members
of the Alectoromorphine Legion in coherent order. A conformation that is a simple
derivative of tlie archecentric form appears to underlie all of them, and this may be
called a Liraicoline-Gruiform metacentre. The duodenum is simple ; Meckel's tract is
slightly elongated, but the elongation is proximad of the diverticulum so that that
appears on the distal limb of an axial loop. The distal region of Meckel's tract shows
the beginning of separation into a definite supra-duodenal loop to which the long caeca
are attached, and into a loop between that and the diverticulum. The rectum is short
and straight. Tlie simpler Limicoline families, such as the ChionidtE, Glareolida',
Tliinocoridoe, and Parridse, are extremely close to this Limicoline-Gruiform metacentre;
and the Lari, which from this point of view present no clear distinction into Alcidai and
lA'TESTINAL TRACT OF BIRDS.
241
L:iri(la', are (lerivatives of the pattern only sli-litly niodiCied l)y increased length. A
nioditication of the Limicoline-Gruiform metaceutre is found in many of the simpler
Cliaradriidse such as Numenius, and consists in the gradual shaping of Meckel's tract
into three loops and a supra-duodenal looj). Of tlic three, one is axial and carries a
large diverticulum on its distal limh. From this Charadri-Gruiform metaceutre one line
of •progressively increasing apocentricity, consisting chiefly in a lengthening of the axial
loop, is attended hy degeneration of the caeca; it runs through Trhiga and liccKrviroslra
to Scolopax. The Gruiform metacentre is another ajjocentric derivative of the Charadri-
¥\v^. 50.
Olidid.i
'kirddriid*
Charadriidas
Rhinochetu
Eurypvg
Columbidae
Gruin#
Charadrndc
Gruiform
MeLdCentre
OpisLhocomus
Pterocles
Tinanu
Chioiiidat
La^
Lari
Araiiudae
Chat'ddri-Gruiform
MetacenU^
CEdicnemidcS
Glareolidas
inocondae
Parridae
imicohne-
Gruifonn
Metacentre
Palamedea
ArchecenLre
Mvohitioii (if the Intestinal Tract in the Alectoromorphine Legion. (For UelorornU read Beliornis.)
(iiuiform mttaccutie, and is reached hy the first three loops of Meckel's tmct, including
the axial loop, heing more clearly marked otf and lengthening equally. Most of the
I'allidte rtmiain in this condition. The Gruinae are apoccntric modifications of it in
which all tlie loops, except that immediately following the axial loop, increase in length
individually. The Psophiina; and Heliornithidtr are practically unmoditied from it.
Ilhhwchetii.s is modified only in that the first loop of Meckel's tract is represented hy
two loops, and that the ca'ca are reduced. Eurij\>ii<i(i is more modified, its apjiarent
simplicity heing almost certainly secondary. The Uicholophidae and the Otididae arc
very apocentric modifications of it, the apocentricity consisting chiefly in the reduction
212
DR. P. CHALMERS MITCHELL OX THE
(jf the first and third loops of Meckel's tract and the great elongation of the axial and
supra-duodenal loops. Aramus is more difficult to place. It appears to be certainly
more archecentric tha,n other Gruiformes, and is to be derived either from tlie Charadrii-
forin-Gruiform metaccntre or, even more archecentrically still, from the metacentre
common to tlie Limicoli and Gruiformes.
CUCULIFORMES.
CUCUL I.
CucuLiD.E. — I liave examined the intestiual tract of Cacoiiuuitls laiiceulalaa, Carpo-
coccyx racUalus, Centropus (species ?), Crotophaga ani, Gulra pirirUjua, and Scythrups
nov(B-hollandi(e, and find that it presents the same conformation in all. 'J'he duodenum
(fig. GO) is straight and narrow. Meckel's tract is not very definitely produced into
Fi-. 60.
Intestinal Tract of Carporocri/x radlnfvs.
loops, l)ut on its periphery there are two loops anterior to the diverticulum, and, posterior
to it, one very long supra-duodenal loop to which the long coeca are attached. The
rectum is alvi^ays straight, but differs in length, being extremely short in some Cuckoos.
IXTESTJNAL TRACT OF HIRUS. 2i3
MusoPHAGiD.E.— I Lave examined several specimens of Corrjlhai.r chlorochlainijH and
C. persa, and have already figured the intestinal tract (26. fiii;. 20). The whole gut is
short and wide, iu obvious association with frugivorous hahit. This circumstance and
the fact that all the specimens T have s(>en were affected wiih Mihcrcnlar noduhs winch
rrade the examination of the gut unsatisfactory, mak(> it impossible to say anything very
definite regarding these foims. The duodenum is straight and of moderate width.
Meckel's tract appears in two portions, the proximal of wliich is a short, nearly circular
expanse on which there was no trace of a diverticulum, and tlic distal a supra-duodenal
loop very closely moulded on the duodenum and supplied by a large bridging vein. The
rectum was short and Avide, ar.d I found no trace of caicji.
V S I T T A C I.
Trichoglossib.t: : Psittacid.e.— Of the first family 1 have examined .species of
Trichoglosstis and LoHus; of the second, species oi Ara, Cacniua, Connriis, Chrijuotis,
Eclectus, Palceoriiis, Psittacus, and Stringops; but, as the conformation of the gut does
not differ in any appreciable fashion according to the families and subfamilies, I shall
treat them together. The larger forms have the gut relatively much longer than the
smaller forms, but, under the diversity so caused, the common type is apparent. In all.
the gut is very long and of small calibre ; the walls are thin, and tlie loops are folded
upon each otlier, and twisted and doubled iu a com])licated fasliion. Tiic masses of
twisted gut are bound together by connective tissue usually loaded with fat, and
"bridging" connections between tiie blood-vtssels in adjoining loops are common. I
have already figured (26. fig. 21) the type as displayed in Ara araraima. The duodenum
is long, narrow, and usually curved. IMeckel's tract is drawn out into very long and
narrow loops, of which the last is always a long supra-duodenal loop with a " bridging "
vein. Of the others, three usually are apjmrent as in Trichoglossus, and any of these
three may become compound. Thus in Striiiyops (fig. 61), of the proximal loops
(marked " a " and " h " in the figure), " a " is cxj)anded and siiows traces of subdivision ;
"6" is subdivided into three; "c" apjjcars as two loops, one of which is further
subdivided, and ouly a comparison with other forms shows that the great length of the
tract has brought about this complexity. I am not confident about the position of
the diverticulum. In Ara there was present AAliat I took to be a small representative
of it at the apex of loop " h," and in one other Parrot I have noted a similarly placed
trace. But in the others, including several other specimens q^ Ara ararmmu , anA in
other species of Ara there was no trace. The cu'ca always are absent, and th(> straight
rectum varies in length.
The diet of the Cuculidie, consisting of insects, friut, and ilcsli, is iu)t of a nature to
have caused much homoplastic modification, and the conformation of the gut is such as
to make it possible to derive it from any fairly archecentric type. Garrod (13) and
others have suggested an affinity between the Cuculida- and the Galli, and, in so far as
the conformation of the gut in Galli is archecentric, there is nothing definitely against
such an origin of the Cuculine gut. On the other hand, there is nothing definitely in
its favour, for such apccentricity as is indicated in the gut of the Galli is ditferent from
2U
DR. P. CHALMERS MITCHELL ON THE
that fouud among the Cuculiclse. In these, as we have seen, the tendency is for tlie
anterior portion of Meckel's tract to be produced into two loops, and this mode of
apoceutricity is typical of Coraciiform birds. The reduced gut of the Musophagidje
might have come from any simple source. The Psittaci live largely on hard grains and
Fis. 01.
Intestinal Tract of Strinr/ops hahroplilus.
a, h, c and S.D.F. are the four loops of Meckel's Tract typical of the Psittaci ; a, h, and c bcin?;
here subdivided, while S.D.F. is a supra-diiodcnal loop.
seeds, and this habit, together with tlie large size of many of the forms, has no doubt
greatly added to the length of the gut and helped to disguise its morphological form.
That conformation is markedly apocentrie, as shown by the complicated folding of the
loops, the length of the loops, the individuality of the supra-duodenal loop, and the
absence of ca'ca. No doubt such a type might have been derived independently from
any of the more archecentric forms, from, for instance, some of the simpler Galli, as lias
Ijeen suggested, but it is equally easy and perhaps more natural to derive the Psittacine
type from that occurring in the simpler Coraciiform birds.
INTESTINAL TEACT OF BIRDS.
245
COEACIIFORMES.
C o R A c r ^.
CoRACiiD-E.— I liave examined the intestinal tract of Leplosoma (species ?) and of
Corocim gan-ulu (fig. 62). In both, which are practically identical, the duodenum is a
Fig. 62.
S.D.f
Intestinal Tract of Cordcias r/arrula.
short loop relatively rather \\ ido. Meckel's tract is llirowii into two rather wide loops
which are separated by the diverticulum, the latter being conspicuous in Lepiosoma,
and only faintly indicated by a trace of a ventral mesentery in Conicias. The distal
loop is a supra-duodenal loop to which the long c;eca are attached. The rectum is
relatively short, wide, and straiijht.
MoMOTiii-K.— I have examined Todus ciridifi and some other sj)ecies of the same genus.
The general conformation closely resembles that in the Coraciidjc, but the part of ^Meckel's
tract anterior to the diverticulum is divided into loops and the ca>cii are shorter.
Alcedixid.e. — I have examined a number of these. The smaller forms do not differ
in any important respect, save the absence of the c:eca, from the conformation in the
Momotidfe. In the larger, such as Dacelo (tig. 63), the first of the two loops into whieli
Meckel's tract is thrown is very long and is wound into a spiral form. In all tiie
Kingfishers the cseca are absent, but there is a supra-duodenal loop more or less separated
from the general sweep of Meckel's ti-iet, and drained by a " Inidging" vein. The cteca
have disappeared com])letely, and the rectum is verv wide and rather long.
SECOXl) SERIES.— ZO(U.OGY, VOL. VIII. 37
246
DR. P. CHALMERS MITCHELL OX THE
Fis:. (53.
Intestinal Tract of Uaclo gigantea.
Pit;. (U.
m
Intestinal Tract of a spceii'S of Merops.
IXTESTIXAI. TRACT OF BIRDS.
•21]
Meropid.k.— 1 have examined two specimens of JIerop.s, tlie species not liaviii- been
identified. In l.otli (fio, (U) tlie duodenum ^yas sliort and rather wide. .Meckel's tract
forms an expanded region, tlie proximal part of wliich showed trace of production into
two minor h)ops. wliilc the distal part was a long- supra-duodenal loop not well separated
Irom the general sweep of the Tract. The cieca were not long, but were wide and
contained f:pcal matter, 'i'iic rectum was short and wide. In the middle of Z^IeckeFs
tract there occurred a distinct diverticulum.
Lpupiu.i;.— In [pnpa cpop.s (lig. 05) tlie duodenum, as iu mauv Coraciiforui l)irds
SDF
In'p.^tiiial Tiac:t of Uj'ii/i<( I'pops.
Im, probable position of Mcck'.'rs divprticiilum.
\\as a wide loop. Meckel's tract displayed two dislinet wide loops on its proximal iiorlion,
while the distal form(>d a long supra-duodenal loop drained by a bridging vein. There
37"
248
DR. P. CHALMERS MITCHELL OX THE
were no caeca, and the rectum was long, wide, and partly convoluted. la Rhytldoceros
plicatus (26. fig. 23), and other Horabills resemble it, the duodenum is extremely wide.
Meckel's tract is thrown into three distinct long loops, of which the last is a well-formed
supra-duodenal loop with " bridging" vein. The two anterior loops possibly correspond
to the two loops of Bacelo and other Coracii ; but the second, which is much the longer,
bears a distinct diverticulum near the apex on its distal limb. There are no cieca, and
the rectum is rather wide and straight. It is clear that the character of the gut does
not unite the Hoopoes and Hornbills closely.
Stbiges.
(1) Strigin.^. — Of these I have examined only Strix flammea. The conformation of
the intestinal tract in this is remarkably archecentric (fig. QQ). The duodeuum is a large
Fig. 66.
rn
Intestinal Tract of Stri.r Jhinmnut.
but simple loop. Meckel's tract is fairly symmetrical about the middle mesenteric vein
A\hich runs out towards a diverticulum. It is produced into first one definite loop (marked
"«" in the figure), and tlien has a circular expanse with a ratlier narrow neck, uniting
it above to loop 'Wr," and distally to a definite supra-duodenal loop. This expanse is
partly produced into indelinite minor loops. 'I'here are two long cteca attached to tlie
supra-duodenal loop, and the rectum is straight and of moderate length.
(2) BuBONiN^. — Of these I have examined species of Biiho, Asia, Athene, Syrnmm,
Speoiyto, and Gymnoglaux. Of these Bubo nmxlmus, which I have already figured
(26. fig. 22), Aslo, and Speolt/to do not dilfer in any important respect from Str'uv: the
INTESTINAL TEACT OF BIRDS.
249
supra-duodenal loop possibly is rather better separated from the general sweep of Meckel's
tract. In the others, except Gyynnoglaux, a slightly more apocentric modification has
taken place. In Syniium aluco, for instance (fig. 67), the duodenum is as in the others.
Loop '• a " of Meckel's tract is as in Strix, but the region between this and the diver-
ticulum is much enlarged, so that the diverticulum is no longer at the apex of the circular
expansion. The supra-duodenal loop has increased very much in size, and is well separated
Fig. 67.
Intestinal Tract of Syrnium aluco.
from Meckel's tract, and drained by a series of "bridging" veins. In Bubo maculosus
and B. poensis the conditions are very similar to that in Syrnium. but the part of Meckel's
tract anterior to the diverticulum appears in two well-marked loops, a conformation
which recalls that in many of the Coraciae. In Gymnoglaax, possibly in association witli
its small size, a simj^lification of the gut has taken place, with the result that there exists
conformation closely resembling that in many of the Rollers and other simple short-
gutted Coraeiiform birds, ^iteckel's tract is thrown into two wide loops separated by
the diverticulum, the posterior loop being a wide supra-duodenal loop with bridging
veins.
250
DE. P. CHALMEES MITCHELL ON THE
C A P R I il U L G I.
Steatormthid^e. — I liave examined only a spirit-specimen of Steatoniis, and that in a
fragmentary condition after it had already passed through the hands of other anatomists.
It was easy, however, to recognize the general conformation of the gut and its resemhlance
to the form in other Coraciiform birds. The duodenum was short and rather wide.
Meckel's tract showed two loops anterior to the diverticulum and two, not so well
separated, distad of it, the second being a distinct supra-duodenal loop with the functional
caeca attached to it. The rectum was long and straight.
PoDARGiD^ and Capriaiclgid-E. — I have examined a number of these, several of
which, however, were not identified as to species. In all, the form of the gut was
essentially similar. The duodenum {Kyctidromus albicollis, fig. 68) is sliort and wide.
Fiff. 68.
1 m-'
Intestinal Tract of N>jcti'lronvi,!> alriicoUis.
Meckel's tract, which is relatively short, appears as a circular expanse, upon the middle
of which the diverticulum was present in most cases, but in some it was absent. The
part of the Tract proximad of the diverticulum showed traces of expansion into tlie two
loops so common in Coraciiform birds. The part posterior to it had the long caeca
closely applied to it, and these were drained partly by a bridging vein. The rectum was
relatively long and straight. This conformation found in the Caprimulgidte is obviously
remarkablv archecentric, and is at once compirable with the primitive type.
C y P s e L I.
CypsELiD.-E. — I have examined several species of Swifts, and in all found the confor-
mation similar to that in Cypielus apns (fig. 69). The duodenum is short and very wide.
IXTESTIXAL TBACT OF BIRDS.
251
Meckel's tract is short, and forms a circular expanse wit:. :^. diverticulum nearly at
the central point. The posterior region of the tract, however, shows traces of former
Fig. 6y.
^_l-_ X
m
l^-~^;
Intestinal Tract of Cypftlus apvi.
complexity, in that it is drained partly by a bridsing vein, a circumstance to which 1
attach importance as evidence that the apparent simplicity of the gut is pseudocentric.
The cceca are absent, and the rectum is straight and wide.
Trochilid.£. — I have examined a number of Humming-birds, and found that the
conformation of the gut was in every case practically identical ^v ' Cupselu^.
It is worth while pointing out that the Humming-birds ditfer clearij .-. . _ .^formation
of the gut from the Passerine Xectariniidge, although in both the small size hn? brought
about a remarkable simplicity. The Xectariniida? possess the Passerine nij - cteca,
and show clear indications of the spiral t\vist in Meckel's tract which is a striking
Passer ne character, and there is no trace of either of these peculiarities in the
Humming-birds.
COLII.
I have already described and figured tlie intestinal tract of Colius capemi^ [z6. tig. 21).
It is relatively shorter and wider than in any other bird that I have examined, and this
modification, no doubt due to small size and frugivorous habit, has obliterated practically
completely tlie underlying morphological form. The gut consists of a very wide
duodenum, a Meckel's tract bearing a small diverticulum, and consisting of a simple
loop similar to the duodenum in size and appearance, and a short, wide rectum. There
are no traces of cieca.
252 DE. P. CHALMERS MITCHELL OIS^ THE
Trogones.
In Trogon puella (fig. 70) the duodenum is a large loop expanded towards its apex,
Meckel's tract shows two loops anterior to the position where I infer the yolk-sac to
have heen, although there was no diverticulum left as a vestige of it. The distal portion
of the Tract has the long caeca adherent to it, and these are drained by a "bridging"
vein. The rectum is short and straight.
PiCi.
Galbulid.e. — I have not seen any of this family.
Capitonid^e. — I have examined Tetragonops Frantzil and Ilegalcema asiatica. In
these the conformation of the gut is ranch alike. The duodenum (fig 71) is a short, wide
loop. Meckel's tract exhibits two short loops anterior to the diverticulum, and, distad
of that, forms a distinct supra-duodenal loop with " bridging " vein. There is no trace
of caeca, and the rectum is straight and wide.
Iyngid.i^. — In Iijihv torqidlhi the conformation of the gut is practically identical with
that in Megahcma.
IIhamphastiu.e. — I have examined Andigr)ia Baillom, Fteroglossus Jl-iedi, Rliam-
phastos ariel, and B. viteUinus. In all the duodenum is very wide, and Meckel's tract
and the short rectum do not differ from the form found in Megalcemu.
PiciD^. — In Dendrocopus major, Geclnus vlridis, and Leuconerpes candidus the con-
foi'mation of the gut is in every important respect similar to that found in Megalcema.
The Coraciiform birds tend generally towards the production of an extremely short,
Avide, and simple gut. The small si/e and the frugivorous habit, so common among
them, are agencies that liave combined in producing this pseudocentric simplicity.
Certain forms, such as Nyctldromus, are readily comparable with the archecentric type,
and present a simple duodenum, a Meckel's tract wliieh is a circular expanse divided by
the diverticulum into an antei'ior portion and a distal portion to which the long caeca
are attached, and finally a straight, rather wide rectum. The changes from such a
condition depend, first, on a widening of the duodenum ; second, on a tendency for the
portion of Meckel's tract proximad of the diverticulum to appear in two loops; and,
third, for a supra -duodenal looji to become distinct in the distal portion of Meckel's
tract. Erom tliis condition, various more apocentric modifications of the conformation
are produced, chiefly by shortening and simplification of the tract and by reduction
(if the caeca.
PASSEKIFORMES.
I have examined a very large number of Passcres, belonging to the family Eurylaemidae
of the Sviliclan.atores ; to the families Tyrannidse, Pittidae, Pteroptochidae, Dendro-
colaptidap, Cotingidae, Eormicariidae, and Pipridie of the Clamatores ; of the Suboscines I
have exammcd 31e)inra, and of the Oscines meraliers of the families Alaudida^, Tinic-
liidae, Pycnonotidae, Muscicapidte, Turdidte, Troglodytidae, Hirundinidae, Campeiihagidae,
INTESTINAL TKACT OF BIRDS.
253
Intestinal Tract of Troijon puflla.
Intestinal Tract of Megalmna asUuica.
SECOKD SERIES. — ZOOLOGY, VOL. VIII.
3S
254 THE INTESTINAL TRACT OF BIRDS.
Dicruridse, Ampelidre, Artamidse, Laniidae, Vireonidse, Sittidse, Oriolida?, Paradiseidse,
Corvidge, Stumiidfe, Meliphagidte, Zostei'opid:e, Nectariniidse, Dicaeidae, Coerebidse,
Mniotiltidfe, Icterida?, and Pi-ingillida?. This considerable labour from one point of view
had a negative result, for I failed to find in tbe characters of the alimentary canal any
indication of the families and groups into which systematists have subdivided the
Passerines. The differences that exist among them traverse the group irrespectively of
the divisions of systematists. These differences depend almost entirely on the size and
habits of the birds. The larger forms have a longer gut absolutely and relatively ; the
fruit-eaters have the gut extremely short and wide ; grain- and seed-eaters have the
gut rather longer. But for these differences the conformation is remarkably similar
throughout the group. The duodenum in all is a simple loop, but usually rather wide.
The rectum is always short, and the caeca, although present in every case that I have
examined, are reduced to short rather nipple-like structures of varying shape, but always
with thick walls and only a very small central cavity. Meckel's tract presents the
greatest divergences. The diverticulum is missing in the vast majority of cases. As
an exception, it is extremely large in Menura, and in a few cases it is present. Relying
partly on my own observations and partly on the extended labours of Gadow (12), I do
not doubt that the diverticulum in Passeres is situated on tlie apical point of the large
portion of Meckel's tract extending from the duodenum to the beginning of the supra-
duodenal loop. In birds with a long intestinal tract, as for instance in Corvus capellanus
(fig. 72), this portion of Meckel's tract is rolled up into a tight spiral which in the figure
is represented as pai'tly drawn out. In ])irds with a shorter tract, as for instance in
Cyclorhis albiveiitris (fig. 73), and this may be taken as typical of the vast majority of
Passeres, the spiral is represented by two distinct loops, which, in the unfolded
condition, are applied very closely to one another, and the folded jjair of loops have a
slight spiral twist. In all Passeres there is a supra-duodenal loop very distinctly
separated from the rest of Meckel's tract, modelled in the most exact fasliion on the
duodenum, and drained by one or more bridging veins from the duodenal vein. The
conformation of the gut in the majority of Passeres, although from its short form it falls
into a very simple pattern, appears in reality to be a higiily sj)ecialized type, the apparent
simj)licity being pseudocentric. In many birds with short guts the intestinal tract is
folded in an irregular and almost capricious manner, and the separate portions bear no
exact relation to one another. In the Passeres the first two loops of Meckel's tract are
most closely applied and extremely similar in caliljre and shape, and their arrangement
suggests most strongly that they have been derived by reduction from the proximal
portion of what was originally a long spiral. It would apj)ear to be certain that the
small Passeres have descended from larger birds. In the same fashion, the supra-
duodenal loop is so exactly modelled on the duodenum to which it is applied, as to
suggest specialization rather than simplicity. In Passeres generally, the blood-vessels,
instead of passing through the centre of the mesenteric expanse, run extremely closely to
the intestinal coils, and contribute to the general impression given by the conformation
of the gut as a highly specialized type.
Intestinal Tract of Corvus cupeUanus ; type of long-gutted Passerine.
/■, bridging- factor of duodenal vein from S.D.F. the supra-duodenal loop ; m. position of Meekel's diverticulum
at the apex of the spiral ; ii.s., it.g., v.n.i/., ganglia of the antonomic nerve-chain.
FiR. 7X
-/--Xl
Intestinal Tract of Cydorliis albiventris ; tyi)o of short-gutted Passerine.
d, apes,of duodenum, which in the unfolded condition is applied to /<, the apex of the supra-duodenal loop ;
)/(, position of Meckers diverticulum, corresponding to apex of spiral in fig. 7-.
38*
256
DE. P. CHALMEKS MITCHELL ON THE
SUMMAKT OF THE CORACIOMOKPHINE LeGION. (Plate 23.)
Gadow (12) unites the Cuciiliformes, Coraciiformes, and Passeriformes into the
Coraciomorphae, the second legion of his second Brigade. The simplest and apparently
most archecentric type of intestinal tract to be found among these birds occurs in the
Caprimidgi. This, which I call the Coraciomorphine metacentre, is a simple derivative
of the archecentric type, and is changed from that practically only in the shortness and
straightness of the rectum, although the duodenum tends to be rather wider relatively.
Prom this metacentre the Passerine gut is a very apocentric derivative, the changes
Fig. 74.
Passeres
Psittaci Ak
Upupid^
cedmi
Meropidas
Cuculidae
CoraciforiD
Cuculiform
Metacentre
Caprimulgi
Coracimorphme
Metacentre
Palamedea
Archecentre
Evolution of tho Intestiual Tract of the C'oraciomor])hine Legion.
(For C'oracimorphiue read Coraciomorphine.)
being the specialization of Meckel's tract into a proximal portion which bears the
diverticulum, and which is wound into a long or short spiral, and a distal portion
which is an extremely specialized supra-duodenal loop. The cfeca have also been
reduced in size. The conformation of the Cypseli and Colii may also be an apocentric
derivative of the Caprimulgid form, the apocentricity in both consisting of an immense
reduction in the length of the whole gvit, with degeneration of the cseca so that no
vestige of them is left, and with obliteration of the loops in Meckel's tract ; there may
or may not be a supra-duodenal loop. The Coraciidse present a less apocentric moditi-
INTESTINAL TEACT OF BIRDS. 257
cation. Meckel's tract api)ears in two loops separated l)y the diverticulum, the posterior
loop having attached to it the long cteca and so forming an incii)i(mt supra-duodenal loop.
Underlying the conformations in the other types there is a central pattern which I call
the Coraciform-Cucviliform metacentre, and in wliicli the duodciniui is frequently but
not invariably wide ; Meckel's tract is produced into two loops anterior to the diverticulum,
and a single, ratlier wide snpra-duodenal loop to which the long cteca are attached. Thr
Cuculida! retain this metacentral form, only sliglitly altered by increase of length, due
no doubt to the relatively large size of these l)irds. It is possible to derive the Psittacine
type as a very apocentric modification of this metacentre, the two anterior loops having
become enormously long, and often complicated in themselves, the posterior portion of
Meckel's tract being produced also into at least two complex loops, the distal of which is
a well-specialized supra-duodenal loop, and the ctcca having completely disappeared.
However, as the position of the diverticulum is not certain, this derivation of the
Psittacine type can be regarded only as provisional. The Striges present a much less
apocentric modification of the metacentre. The two anterior loops of Meckel's tract are
present, but the second is not so well separated from the general sweep of tlie tract, and
appears to be part of a circular expansion of the whole tract. But in some of tin; species
of Bilbo the metacentric position is reproduced exactly, and it is easy to regard the
conformations in Sfi-Lv, Syrninm, and so forth as very slight alterations of the common
form. The Meropida;, the Momotidie, and the Trogoues all retain the metacentric
position with extremely little alteration. The LTpupidae, the Alcedinidae, and the Pici
are all apocentric modifications of it, the chief difference being the disappearance of
the ca^ca and the elaboration of a distinct supra-duodenal loop.
Summary of the Systematic Description.
The various conformations of the Intestinal Tract in birds may all be referred to an
archecentric form, and this form is well displayed in Valamedea. The Struthious birds
(with which the Tinamidse are not included) adhere closely to this archecentric typr.
The Turnices and the Galli adhere to it almost as closely, but among the Galli an
apocentricity, consisting in the expansion of the distal portion of .Meckel's tract, liegins
to appear. The Tinamidae, the Opisthocomidae, and the Pteroclo-Columba; present
apocentric conformations which are not easy to derive from those in any other groups,
although there are certain suggestions of affinity with the type disjjlayed by Gruiform
birds such as Otis. Eor the present, however, it is preferable to regard the forms in
these birds as seiiarate, although allied derivatives of the central type. The Passeres
again do not show any clear affinity with the types in other birds. The conformation
displayed must be regarded as having come separately from an archecentric type,
perhaps from that common to tlie other Coraciomorphine birds, but which has
progressed very far apocentrically along a radius of its own. The conformations in all
other birds may be referred without difficulty to three main metacentres, which are
all simple, but different modifications of the archecentric type. The metacentre of all
the birds in the Pelargo-Colymbomorphine Brigade is a conformation in wliich the
258
DR. P. CHALMEES MITCHELL ON THE
symmetry of Meckel's tract about the median mesenteric vein and the Meckel's
diverticulum is retained, but in which Meckel's tract tends to be produced into a
number of long and narrow loops, the most distal of whicli is a supra-duodenal loop
to which the long cteca are attached. The metacentre of the Limicoline-Gruiform
assemblages (which contains the Alectoromorphine Legion without the Tinamiformes,
Galliformes, and Pteroclo-Columboe) is a derivative of the archecentre in which Meckel's
tract is no longer symmetrical about the middle mesenteric vein and the diverticulum,
but in wliich the region anterior to the diverticulum is produced beyond it so as to form
an axial loop, on the distal limb of wliich the diverticulum lies and which may be greatly
Passeres
Coracimorphine
Metacentre
Pelargo-
Colymbimorphme
Metacentre
Limicoli-
Gruiform
Metacentre
Fteroclo - Colum b as
Opisthocomi
Tlnami
Archecentre
Palamedea
Turnices
Ratitas
Galli
Evolution of the Intestinal Tract in Aves.
{For Coracimorphine and Colymbimorphine read Coraeiomorphine and Colymbomorphine.)
prolonged as in Scolopax. The members of the Coraeiomorphine Legion, except the
Passeres and a few pseudocenti'ic types, may be derived from a metacentre in which
the duodenum tends to widen, and in which the part of Meckel's tract anterior to the
diverticulum is produced into two loops. In Plates 21, 22, and 23, and in figu.res 35, 59,
74, and 75, the evolution and affinities of the conformations of the intestinal tract are
represented, and by no means necessarily the pedigrees and affinities of the birds
containing these conformations.
Intestinal Portion of the Autonomic Nervous System.
It was only at a late stage of this investigation that my attention was directed to the
peculiar features in the Sympathetic, or, to use the modern term, the Autonomic Nervous
System of Birds. In certain birds (for instance the Megapodidoe and Cracidae) the
INTESTINAL TEACT OF BIHDS. 259
so-called intestinal nerve is unusnally lar!>e and visible, and it was in members of tbese
families that I first noticed it. In the majority of cases, however, it is not readily seen ;
moreover, much of ray material consisted of rather badly preserved spirit-specimens, or
of birds that bad died of disease in which tul)ercular and fatty dei?enerations of the
mesentery were a conspicuous feature, and I am able only to <nve a few almost casual
notes, wliich, liowever, may serve the purpose of redirecting the attention of anatomists
to a very peculiar and interesting set of structures. A good deal is known al)out the
paired ganglionated mesenteric chain {Grcnzslrang of the German writers), thanks to
tlie observations of many writers, of whom the chief are Wiedersheim, Gadow, Gaskell,
and Marage. In the cervical region [Wiedersheim (36. p. 350), Gadow (12. p. 394),
and Marage {22„posshn)'] it divides into a deep and a more superficial portion, and irregular
traces of this division persist in different birds in the posterior parts of the body. In all
cases it appears to have more autonomy ; that is to say, to be in less intimate connection
with the metameric spinal nerves than in most other vertebrate forms. In the lumbar
region there arises from this a very complex plexus with large ganglia on the edge of the
stomach, on the ovary, on the supra-renal capsule, and further back near the rectum. It
is from these ganglia that the so-called intestinal nerve of birds arises. Concerning this
peculiar nerve the literature is very scanty. Remak (32, 33) appears to have called
attention to it first, hut most later writers pass it over almost completely Thus Gadow,
Wiedersheim, Piirbringer, and Beddard, in their ornithological and anatomical treatises,
pass it l)y. Oppel, in his great work on the anatomical tract (30), is content with
discussion of the plexuses actually in the wall of the gut. Gegenbaur (14), in the most
recent edition of his ' Text-book,' mentions the existence of such a nerve in reptiles, and
states that it is best developed in birds ("Am meisten sind diese Nerven bei Vogeln
entwickelt. Ein den Mitteldarm begleitender Nervenstamm geht am Enddarme in
mehrere anselmliche Ganglien "), and adds that it is undeveloped in mammals. Marage
(23) has given the best account of it, and states that it differs in its arrangement in
different birds ; but his figures are rather difficult to follow, and he is incorrect in statins'
that it does not occur in rapacious birds, and he seems to have overlooked it in tlie
Ratites. As I have already stated, I do not pretend to have made anything that
approaches to a complete study of it, and the figures that I have been able to give must
he taken only as rough anatomical notes. Special study by special methods on material
in good condition is required *. I can say, however, that in every case since I was
aware of its existence, where the material was in sufficiently good condition, I found it
present, and I do not doubt that this pi*esence is invariable. It arises usually fi-om two
or three main nerves, which leave the ganglia corresponding to the solar plexus and the
ganglia over the ovary and supra-renal capsule, and enter the mesenteric expanse which
is tlie sujjport of Meckel's tract. These, or some of these, are represented in figures 1
and 45, and I have worked them out in several unfigurcd cases. In fig. 72 there is a
* Since writing tliis memoir, my attention lias bi;en directed to a bcautil'ul memoir by Thebault (34), which
contains, inter alia, an elaborate study of the modes of origin of the intestinal nerve in various types of birds. —
P. C. M.
260 DR. P. CHALMERS MITCHELL ON THE
large ganglion at entrauce of these to Meckel's tract, and I could make out only two
nerves forming the oi-igin of this ganglion. One of the three nerves frequently runs
independently to the duodenum, as in Otis (fig. 45) and in Andigena Bailloni ; but no
doubt there may have been a cross connection between this and the main intestinal
nerve, as certainly is the case in the Pigeon. In the duodenal-fold looj) usually at least
one distinct and large ganglion is present (fig. 45, v.ii.v.; fig. 72, d.g.). The main
intestinal nerve which supplies Meckel's tract starts in common with the duodenal
nerve as in Palamedea (fig. 1) and the Passeres (fig. 72). It courses round the mesen-
teric expanse of the tract, giving off a series of minute branches to the gut, and where the
tract passes into the rectum it frequently receives one of the entering nerves, or is in
connection with the ganglion at the entrance to the mesentery of Meckel's tract, and
then turns down parallel to the rectum, and comes again into connection with the
posterior portion of the Grenzstrang. On its course this main intestinal nerve may
follow Meckel's tract more or less closely. Thus in Palamedea {&g. 1), Pandiou (fig. 34),
Talegallus (fig. 38), it foi'ms a loop corresponding to, but not absolutely identical with,
the course of the tract. In Falco melanogenys (fig. 32) it follows the general course of the
tract much more closely, having a loop following the supra-duodenal loop. In Crax
Danbentoni (fig. 39) it has a curious elongated and narrow distal portion, which corre-
sponds exactly with the typical apocentricity of these birds, in which the tract itself is
enlarged on its distal portion. This intestinal nerve in many cases — Struthious birds,
Palamedea (fig. 1), Talegallus (fig. 38), Crad- (fig. 39), other Galli, Falco (fig. 32),
Pandion (fig. 34) — presents a very large numljer of small ganglia evenly distributed along
its course. In other birds, e. g., Otis (fig. 45), Columba, Andigena, and Corviis (fig. 72),
there are a smaller number of much larger ganglia. As to the systematic value of these
differences, it is impossible to say anything definite on so small a range of information ;
but the sixbject is very promising. It appears as if the chain with many ganglia were
more primitive than the nerve with a limited number of large ganglia ; and it is certainly
the case that such birds as Otis, Columba, the Toucan, and the Passeres, Avhere the
number of ganglia is very small, are more specialized forms than those with multi-
ganglionated chains. It has to be remembered, however, that although the nervous
systems of many invertebrates with a small number of large ganglia appear to have been
produced by the concentration of multi-ganglionated chains, we have no right to extend
such a princi])le to other forms. The origin of this intestinal nerve in birds requires to
be worked out, and in this connection the observations of Andersson (i), who found that
in the Urodele Amj)hibia the main symjiathetic chain was subdivided into a Grenzstrang
and a Collateralstrang , are worthy of attention.
It is to be noticed that this intestinal nervous system lies, as the blood-vessels lie,
between the two layers of the mesentery, and therefore outside the coelom. In larger
forms, such as the Emu, it is often comparatively easy to strip otf one layer with its
load of fat from the mesenterial expanse supporting Meckel's tract, upon which the
ganglionated nerve-chain come plainly into view\ However, just as it occasionally
happens that some of the blood-vessels pierce the mesentery and form *' bridging" vessels
traversing the portion of coelom between two loops which happen to lie in contact, so in
INTESTINAL TKACT OF BIRDS. 2G1
certain cases nerves may " bridge/' or rather bore tliroui-li, the m.'sentery and pass across
a narrow portion of coelomic sjiace. The most obvious case of tills occurrence is where^
as in Otis, nerves leave a ganglion in the duodenal loop and pass to tlie cjbcu or the
supra-duodenal loop. I am practically certain that this happens in many of the small
Passerines, Avhere tlie supra-duodenal loop is a structure of considerable imjjortance, but
I am not yet prepared to demonstiati- this.
MOKPHOLOGY OF THE L\T1£ST1.\.\L TkACT.
The chief writers on this subject witliin conii)aratively recent times have l)een Tohit
(35), who deals chiefly with the tract in Man; Klaatscli (18), who in two extremely
important memoirs dealt with the relation of the tract and its ancillary viscera to tin-
mesentery; andMathes (24). who folloa^Ml closely tln^ work of Klaitsch, but dealt chiefly
with the development of tiie mesenteries in tiie Amphil)ia. None of these writers has
])aid special attention to the conditions tliat exist among liii-ds, but tlieir work has been
of great assistance to me in interpreting and coordinating my own investio-ations. Jt is
plain that in all the higher vertebrates the hitistinal tract is thrown into three main
portions which are homologous throughout the series. Of these, the first is the
Duodenum, which in birds is ahvays a closed loop lying ventral to the rest of the tracf.
It arises extremely early in ontogeny, and while in the majority of eases it remains
simple, it may develop many minor complexities, sometimes simply becomiuo- wider,
sometimes being thrown into numerous minor folds, and sometimes being twisted intd
more or less regular spirals, the sj)iral duodenum being in some cases (Storks) wound
with the spirally twisted proximal loop of Meckel's tract. Concerning the relations oF
the duodenum to the supra-duodenal loop I shall preseatly have more to sav. The
distal extremity of the duodenum, however the course of tliat may have been complex,
always returns to the dorsal edge of the mesentery very close to the starting point of the
anterior liml). and there passes into the second j)ortion of the gut. 'I'his portion, Avhich 1
name Meckel's Tract, extends from the duodenum to the insertioii of the cicea. The
first important point about this large region of the gut is that it represents an outgrowtli
of only a very small section of the primitiA'c gut. Its proximal extremity approaches its
distal extreuiitv so closely in the line of the dorsal attachment of the mesenterv, that in
the majority of eases it would be possilde to remove the Mhole of Meckel's tract and
suture the cut edge of the duodenum to the cut proximal edge of tlie rectum, and
a. most without dislocation reconstruct a primitive straight intestinul canal. In actual
development Meckel's tract, in all the vertebrates in which it is developed, arises as a
simple narrow loop in the line of tlie principal mesenteric artery. Toldt's figures, and
others given by Kollman (19), show this beautifully in the case of immau embryos, and
general comparative anatomy from the Frog to Man makes the morjihological nature of
Meckel's tract extremely jjlaiu. There can be no doubt that this is the most recent
phylogenetic development of the A'ertebrate gut, and that it ecu-responds to not mon-
tlian two, or possibly three, of tlie primitive somites of the budy. When tlie development
and comparative anatomy of the intestinal nervous chain in 13irds has been worked out.
SKCONIi .SERIES. — ZOOLOGV, VOL. V1J[. 39
2r>2 DE. P. CHALMERS MITCHELL ON THE
it is to l)e expected that the origin of that system from not more tlian three enterins:
nerve-hranches will he given an importance which as yet can only he suspected. The
nature of Meckel's tract as a recent outgrowth of a portion of tlie gut corresponding
only to a very limited numher of segments has an important hearing on medical work,
as it is heing found that there is an organic synipathy through the nervoiis system
hotween certain superficial areas of the skin and certain visceral organs or portions of th(^
organs; the sympathy depending on a common relation to the primitive segmentation of
the body, and heing of practical value in the diagnosis of affections of internal organs.
In those creatures in which there is a yolk-sac or umbilical cord, tlie outgrowth of
Meckel's tract from the primitive straight gut is opposite the vessels runninii' to these
structures ; and it seems tempting to regard the origination of Meckel's tract as heiui;
dependent on this mode of embryonic nutrition. However, as Meckel's tract is equally
definite and equally metamerically limited in the Frog, we cannot lay much stress on
this possible mode of origin.
The large intestine, from the caeca to the cloaca or anus, is the third distinct portion of
the intestinal tract. It always lies in the primitive position of the straight gut, dorsad of
all tiie other portions, and phylogenetically it is the oldest portion and corresponds to the
greatest numher of somites. It appears to he homologous throughout the vertebrate
series ; hut this homology depends on the identification of the paired caeca of Birds witli
the unpaired caecum of Mammals and Lizards — a homology not apparent when a bird
with a short gut is taken, hut much more convincing when the comparison is made
bet\A een a form such as Palamedea (fig. 1) or a Struthious bird and a Mammalian or
Lacertilian intestinal tract. In the more primitive types the large intestine is very long,
and may show traces of division into colon and rectiun ; in the higher forms, partly in
correlation with the greater development of the thoracic portion of the viscera, the
duodenum and whole intestinal tract has shifted far distally, with the result that the
large intestine is extremely small, and is here referred to simply as rectum (see footnotes,
pp. 176 and 271).
Meckel's Diverticulum.
Examination of the embryo of any bird makes it plain that this structure is the
vestige of the yolk-sac, and its retention in adult birds has been described by a number
of anatomists. It has been termed by most writers who understood its nature the
Diverticulum ccecum vUelli; but as it is obviously homologous with the caecum described
by Meckel as an abnormality in Man, and as in human anatomy it ])ears the well-known,
name of Meckel's diverticulum, I have preferred to give it that name in birds. I have
not found it present in any adult reptile, and I can find no record of its occurrence
there. Nor have I found it in any mammal other than Man. In human anatomy its
frequency of occurrence has received recent attention, and Birmingham (4) states that it
is present in about 22 per cent., varying from half an inch to five inches in length. In
birds it is present much more frequently ; indeed, its presence throughout life is a
character of very many groups. Gladow (12), summing up his own very numerous
observations and those of earlier writers, states that it is retained throughout life by
INTESTINAI, TKACT OF BIRDS. 2«o
SwimminGf-birds and most Waders, but that it disappears very early in Birds of I'rey,
Parrots, Woodpeckers, and Singini,^-birds. He reu:ards it as a fiinctionless rudiment.
Liinnberg and Jagerslviold (21) examined a lari,^' numher of birds, chiefly Sea-birds and
Waders, for it, not contentin^• themselves with siiiii'l,> individuals, and found it alisent in
most Gulls and Terns, present in Waders. Ducks, llcroMs. and some others. As at an
Cfirly stage in my investigations I found it an important jjoint of morphological orien-
tation, I searched for it in each of the very lari;e numher of birds upon which this
communication is based, and found its presence much more frequent than lias been stated.
An interesting feature in connection with it is, that in a large number of cases it is sup-
ported by a vestige of the primitive ventral mesentery. Usually this appears only as a small
fold of tissue tying down the diverticulum to the ventral edge of tht; sjut, hut sometimes
a strong band runs from this towards the liver, the latter of course being develojx'd in
the primitive ventral mesentery. Occasionally when the diverticulum itself was absent,
its place was indicated l)y the presence of a mesenterial vestige, as in many Pigeons.
I have found the Divei'ticuknu present and large in all the llatita^ ; of the Colymhifornu's
absent only in Pw//Vr/M' ; present in the Spheniscifornu's ; present in the Pi-ocellnriifomies ;
of the Ciconiiformes, present and large in all the Steganopodes but small in J-'/ofiiN, and
present and larue in the Ardeidte, Scopidie, Ciconiie, and Phcenicopteri, in many of the
Ciconiiformes being very large ; present and large in all the Anseriformes ; present in all
the Falconiformes (except a Birteoj, and very large in tlie FaIconida% hut usually small
in the others ; present and large in the Tinamiformes ; present hnt usually very small in
the Galliformes (except Tuniia:); present and very kuire in all f he (Iruiformes, often
extremely large, but, as a solitary exception, absent in Ile/iunun ; present and very large
in all the Li mi cola? ; among Lari small and olten absent in the Laridae, small, but
usually present, in the Alcidae ; present in the Pterocletidte, but usually absent and
always extremely small in the Columbida\ a feature in which they stand in marked
contrast with the Charadriiformes generally ; of tlu; Cuculiformes, always present
although small in the Cuculida? ; absent in the Musophagid*, and probably always
absent in the Psittaci ; of the Coraciiformes, always small, but present ratlier more often
than not in the Cor;iciie, present and small in most of the Striges, very small but present
in most Caprimulgi ; always very small and absent more olten than present in the
Cypseli, Colli, Trogones, Pici ; of the Passeriformes, absent in the vast majority of cases,
when present extrenudy small except as a solitary case in Menura, where it Avas very
large. In making this review I liave exchuled those .specimens which were marked in
uiy notes as obviously chicks or quite young birds, and the list may be taken as repre-
senting with fair exactness the incidence of the diverticulum among the .A.vian groups.
In all the eases where I have mentioned, in the paragraph above, that Meckel's diverti-
culum Avas small, I think that it was a vestige in the true sense, that is to say a functiouless
rudiment of an embryonic structure. !Sometim(^s tJic bunen remained, and contained a
few fracments of material resembling volk, and doubtless remains of tlie yolk. Rather
more frequently the " small " diverticula had no lumen, ami were mere nodular
excrescences on the wall of the 'J'ract. Among the cases which 1 have noted as large there
are certainly some (Katites, Palamedeu, etc.) in the same category. 'I"he curious feature
39*
•264 DB. P. CHAL:MEHS MITCHELL 0>' THE
therefore exists that a functionless restige is retained universally in some srroups and
not in other groups — a circumstance to \\hich the attention of those naturalists may be
directed who would see a purpose or "selection-value"' in every systematic character.
Meckel's diverticulum presents another condition of great interest. Lonnbers and
Jagerskidld (21) drew attention to the iact that in certain cases where the diverticulum
is large, it lias a patent lumen and a ttiick wall, and is slightly constiicted from the gut
(21, cited in Oppel 30, p. bo9K These authors, while they sugirest that the vestiirial
organ has been transformed into a ^land, do not go further in their microscopical
investigation than to point out that the mucous membrane of the diverticulum in these
cases is thickened by a series of folds. I have examined the microscopical structure in
several Anatifijrmes, Gruiforraes, and Charadriiformes, and find that glands occur
frequently in the foldings of the mucous membrane, and that the diverticulum in such
cases corresponds, with its folded wall containins: glands and lymphoid nodules, very
closely to the structure of the caeca in Passerines and Columbae. I am inclined to think,
therefore, that in many cases, particularly in the groups that I have mentioned (and
possibly in the Falconinse), Meckel's diverticulum has acquired a new function. It is at
the least suggestive that where the diverticulum has become glandular, the paired caeca
are either rudimentarv and functionless. or thev are verv lariie. thin-walled, and full of
faecal matter. "\iVhere, on the other hand, the paired cseca are chiefly glandular, the
diverticulum is either a functionless vestige or has disappeared. This, however, is not a
complete account of the relations of the two organs ; for, in the first place, it is based only
on a relatively small number of observations of microscopical structure, and in the second
place there are instancf s, such as the Psittaci, where both diverticulum and paired cteca
are absent in the adult.
The SuBSLDiARr Loops of Meckel's Tract.
In the section dealing with systematic description, I have already said all or nearly all
that 1 have to say regarding the minor loops into which the Tract so frequently is
produced. I have tried to show that these display patterns which persist through
systematic groups, the pei-sistency referring to their position with regard to the diverti-
culum and to their number. The arrangements of these minor loops, in fact, are
mstances of what I term uniradial apocentricities. "When Meckel's tr.ict is elongating,
in a large bird or in a bird the hahits of which demand a great length of gut, the
elongation does not take place at random, but in special regions and in special modes.
The combination of position and complexity is of a kind not likely to be repeated
independently, but to have common origin, and so to prove of systematic value. The
explanation of how these diiFerent and complex combinations came into existence I have
not attempted. Eefore that could be done, there is necessary the great labour of
following out in every stage of individual development the relations of the growing folds
of the gut to the blood-vessels, regions of the coelom, liver, air-sacs, sternum, stomach,
and so forth. The beautifid work of Klaatsch and of Mathes, elaborate and prolonged as
it was, does little more than to open up the lines of such enquiries. Until something
is known in each individual case of the nature of the " srowth-forces " in contiguous
IMHsTlXAI, TRACT OF BlKDs. 2{;
).j
"org-ans, of their compromises and eo-ordiiiatioiis, in fact of their phices with regard to
one another and to the whole corporeal republic in every staije of the growing eiubrvo,
nothing more can be said bvit that such complex uniradial apocentricities, if not in them-
selves possessed of " selection-value," may stand in correlation with structures that have
such value.
ThK SlPUA-DlJODKNAI, LoOl'.
In its natural condition the duodenum lies folded ventrally under the other portions
of the gut, and comes in vci-y close relation witii the distal portion of Meckel's tract.
Cuvier drew attention to tlie fact that in birds as in mammals the duodenum comes int..
intimate relation witli a posterior portion of tlie gut, a relation so intimate tiiat the
mesenterial folds suspending the two portions nuiy fuse after the fashion explained bv
Klaatscli. This portion of the gut which comes into relation with the ca^ca I have
called the supra-duodenal loop, preferring not to call it •• colon" as was done by Cuvier,
since that term is applied in ]\tammalian anatomy to a portion of the gut posterior t(»
the cteca, and thereroro belonging to what I have been terming the rectum. Within
the group of liirds various stages in the evolution of tliis curious inter-relation
between the proximal and distal portions of the gut are displayed. Thus in Paldwcdrd
(fig. 1) there is no supra-duodenal loop, and the gut may be uiifolded witliout any
difficulty or cutting of blood-vessels. The same conditions obtain in a nundicr of tin-
archecentric types, and in the systematic portion I have referred to these. lOveu in
Palamedea, however, the earliest stage in the formation of the connection is apparent.
A short recurrent factor of the duodenal vein runs in the mesentery at the dorsal portion
of Meckel's tract across from the terminal portion of that tract and assists in draininu-
the caeca. As the ca'ca increase in size this vessel becomes larger and of more
importance, and, in many cases which I have referred to in the systematic portion, the
recurrent vein along with a second and sometimes a tliird accessory recurrent vein i-iui
to nearer the apex of the duodenum, having traversed the mesentery as " l)ridging " veins,
and arisen from the long ca-ca and the posterior portion of Meckel's tract. Tn sucli a
mode, an intimate relation is established between the duodenum and tlie distal portion of
Meckel's tract, and, as in Ofifi, this relation may involve* not only tlie veins but the
nerves. The portion of the Tract to which I he ca'ca are adhercMit is not distinctly
marked off from the more proximal portion of the Tract in very many of th(' less
apocentric types. But, as I have shown in the systematic \Kn-{un\, pari puxsu with the
establishment of the •' bridging" veins, the distal portion of the Tract becomes a distinct
loop clearly marked off from the general sweej) of the Tract, finally, in tiiose birds
where the eieca have degenerated either completely or have shortened to glandular
nipples, the supra-duodenal loop is retained with its separateness from the rest of the
Tract and with its '• bridging " veins. In sucli a form the supra-duodenal loop generally
hecomes very accurately moulded to the contour of the duodenum, and its presence and
completeness are important reasons for seeing in the apparent simplicity of the gut in
many of the higher forms, such as the Passeres, a pseudocentricity — a condition apparently
simple, bvit still retaining evidence of past complexity. I do not tliink it can be doubted
'2C)6 BR. P. CHALMERS MITCHELL OX THE
that wlierever ia short-gutted birds without large caeca the supra-duodenal loop is present
in its elaborate form, these birds at one time had a longer gut, and almost certainly
possessed longer ca?ca. Where a gvit is long, and where its irregularly folded loops lie
closely together in a narrow space, the establishment of an intimate connection between
any of the crowded loops would call for little remark. Where, as in the Passeres and
many other short-gutted forms, there appears to be abundant space, and yet the
duodenum and the supra-duodenal loop are accurately moulded, the one on the other,
and in intimate vascular, nervous, and mesenterial connection, \\v nuist seek for the
origin of the conii)lexity in the past history.
There can be no doubt, I think, that the presence of a siiecialized supra-duodenal
loop marks a liigh degree of apoceutricity in the intestinal tract possessing it, but it is
equally plain that this apoceutricity is multiradial and no guide to affinity. The
))resence of cteca of at least moderate length is a fundamental or archecentric character
of birds; and, if the supra-duodenal loop has arisen in the mode I have indicated, it is
clear that it may have arisen repeatedly, and in my systematic description I showed that
actually it does appear, repeatedly and apparently independently, in the different groups.
The probably multiradial nature of the structure is also supported by the occurrence
of similar formations among Mammals, these formations not even being on exactly
homologous parts of the gut. The mammalian structures with which the Supra-
duodenal Loop may be compared are the loop which in Man has the tr;insverse colon as
its apical portion, and the sigmoid flexure, which in embryonic Man reaches much
further towards the duodenum. Naturally, I do not propose to enter at present into the
various modihcations presented by these structures ; it is enough to say that the
connections in Mammals between the duodenum and jiosterior regions of the gut are
frequently present, but less frequently than in the case of Birds.
Dealing with its occuri"ence in Man, Toldt sees in these connections between the
proximal and distal portions of the gut the mere result of apposition. Klaatscli
criticises this view, and rightly points out that, although many of the other loops are in
equally close spatial relations, fusion does not necessarily occur among them. In Birds,
however, as I have shown, secondary fusions and " ])ridging " veins are not absolutely
confined to the duodenum and sujira-duodenal loop. Thus in Fsittaci they are frequent
among the numerous long and narrow loops into which Meckel's tract is thrown. In
Anatidse secondary fusions and "bridging'' veins are of frequent occurrence in the case
of different portions of the same loop, notably in the axial loo[), and in Spatula in the
long loop distal to Meckel's diverticulum (fig. 24). Tinally, in the Ciconite bridging
veins and secondary fusion of the mesentery occur between the duodenum and the
j)VOximal minor loop of Meckel's tract. These occurrences, however, are less imjjortant
and much less frequent than the supra-duodenal connections. Klaatsch accounts
for these in the case of Reptiles and Birds by the mode in which the ccelomic divisions
are broken up by the intruding blood-vessels and viscera. I think, however, that, in
birds, the formation may serve a useful purpose ; the supra-duodenal loop is maintained
and made even more elaborate after the degeneration of the cit'ca, although it arose in
relation to the cieca ; moreover, it is specially perfect in cases where there is no reason to
INTESTINAL TRACT OV lURDS. 267
suppose that want of space played a i)art in its ontogenetic appearance. JMainly. wlien
thepyloi-ic valve relaxes and the contents of the stoniacli are poured into the duodenum,
the shock will he transferred 1o the supra-duodenal loop wliicli lies closely applied to
the duodenum and sometimes in sjieeial nerve-connection with it. Aiul thus discharffc
of the contents of the posterior region of the gut into the rectum may he set about
without the necessity of peristaltic waves traversinij the wliole len<,'tii of Meckel's tract.
Thk Colic C.kca.
(iadow (i2), Fiirbringer (9), Beddard (2), and Oppel (30), in their respective treatises
have devoted so much attention to the voluminous literature concerning the colic vxai
of birds tliat I need not refer to older writers. The arcIuH-entric condition of these
organs in Itirds, a condition which is probably an heritage from Itrptilian ancestors, is the
existence of a pair of cu'ca growing from the point when; the distal end of Meckel's
tract passes into the rectum. Such primitive ca-cn proximally are applied more or less
closely to the posterior poi-tion of Meckel's tract. They are of moderate length ; their
walls are not specially thickened, their lumens are widely open to tlie gut, and their
contents consist of food-material in a state more akiu to that in the rectum tlian to the
state in Meckel's tract. When, as happens frequently, there is a difference in colour
apparent through the intestinal wall and marking the diiferent stages of metabolism, tlie
colour of the caeca approximates to that of the rectum. These primitive caecii probablv
had a digestive function of some sort, for the presence in their walls of glands, of
absorbing veins, and occasionally of villi show that tliey were not mere reservoirs of
hecal matter. From the primitive condition various apocentric moditi('ations have
arisen. The c;eca may increase very greatly in size, and may develop spirally arranged
septa protru(iin<; nu)re or less into tlie cavity and deeply marking the exterior, as in
L'haiina and some of the Ratites and Gruitbrmes, or the whole external surface may bi'
prolonged into a luunber of papilliform hollow outgrowths, as in Calodromda described
by Beddard (2). These enlarged c;eca appear to retain their digestive functions.
Secondly, the ctpca may become very much reduced, but in such apocentric reduction I
am convinced that there are two quite different conditions to be noted. In the one case,
the reduction may be nothing more than the degeneration of an organ that has l)ecame
f unctionless ; and almost any stage from the archecentric size to complete absence may
exist. In the Columbidoe, for instance, the ca'ca if present are always small aiul
frequently are thin-walled, irregular (I have noted many cAses of individual absence of
one cjecuittl, and sometimes pigmented. In the fruit-eatinii' Pigeons, and indeed in
many other Pigeons apart from the nature of their food, thr \estigrs have disappeared
completely. Pr(^eisely a similar series of events occurs among most Kalconiform birds.
The caeca are small in all, and when present appear to he functionless, thin-walled
vestiges, frequently unsymmetrical (here, again, absence on one side has been noted in
different birds by different observers). Sometimes they appear to be absent, hut inflation
of the wall of the gut reveals the presence of slight, thin-walled rudiments in their
place. In the Vultures they are ^onu'times totally absent, and a similar absence is \x
2G8 DE. P. CHALMKHS MITCHELL ON THE
character of the Cathartidae. However, tliere is yet another condition possible among
these reduced CKca. Is'umerous obserAers liaA^e shown that in the cteca of many birds,
whether these be long or short, there occur in different regions masses of lymphoidal
tissue. Such masses may occur at the apex of long caeca as in the Owls, or in scattered
))atches as in Ducks ami FoavIs, or in concentrated swellings as in Otis. Berry (3) lias
)-(>cently shown tliat the occurrence of these lymphoid masses, in concentrated regions
I'orming vermiform apjjendices, or in scattered masses, is frequent in Vertebrates, and he
suggests that the vermiform appendix of Man is a specialized and not a degenerate
structure. The reduced ca?ca of many birds {e. g. Passerines) are quite different from
the obviously vestigial caeca that I have mentioned. They are regularly formed, nijjple-
like structures, the lumen of \\ hich is grcatlv reduced bv closelv-set villi and folds. In
these, small glands and lymplioid tissue are alumdant ; and in the Sparrow I have noted
the presence of abundant dark granules which are ajjparently matter in process of
excretion. I distinguish therefore in the conditions of the cieca among birds the
following : —
1. Archccentric Character. — Ca^ca large, thiu-w ailed, containing food-material on
which some process of digestion is taking place, but which may have in addition
the beginning of secretory or excretory function.
2. Apocenfric Characters.
A. Caeca enlarged, partly digestive, and partly secretory or excretory.
B. Caeca functionless, A-estigial or absent.
C. Caeca r(!duced but tiansformed solely to glandular organs, secretory or
excretory or both.
•Palamedea. Caeca archecentrie. Chauna apoceutric A.
IIatiTjE. Ca^cae archecentrie or ajiocentx'ic A.
CoLYMBiFOKMKS. Archeccntric.
Spheniscifokaies. Apocentric B. but according to Gadow archecentrie in some.
Pkocellakiifoumes. Apocentric B, occasionally one only present.
CicoxiiFOitMES. Archecentrie in the Pelecanidte. apocentric B in ihe Phaethontidic,
Sulida^ Fregatidae, and Plialacrocoracidie (in I'lotns one may be absent);
apocentric B in the Scopidfe, Ciconiidie, and Ardeidae, in the latter one only as
the normal but jiot invariable condition. In the J'hoenicopteri archecentrie.
Ansekifokmes (excluding the Palamedeida'). Apocentric A, usually Avith large roAvs
of glands, hut in Jfergns apocentric B and one occasionally absL-nt.
Palconifokme.s. Apoccnti-ic B, vestigial with frequent irregularities and absent in
Cathaitidae.
TiNAMiFoitMES. A])oceritiic A, extreme specialization in some.
(lALLiFORMEs. Apocontric A in (ialli and Opisthocomi ; archecentrie in Turnices.
■(lUUiFOKMES. In the Kallidae apocentric A; of the (iruida' archecentrie in the
Araminte, apocentric A in the others. In the Dicholojihidte and Otididae
apocentric A (extremely so in Ot/\s) ; in the P.iiinochetidac archecentrie ; in the
Eurypygidte ajwcentric, probably C ; iu the lleliornithidic archecentrie.
INTESTINAL TRACT OF BIRDS. 269
Charadriiformes.
IdmicolcK. In the Glareolidse archecentrie ; in tlie Chionidae, Thinocoridse, and
Charadriidye apocentric A, but. in some of the latter family apocentric C. In
the Parridye, Alcidte, and most of the Larid;e apocentric B and sometimes
practically quite absent, but in Sierc07'ariit.s archecentrie.
PterocletidcB. Apocentric A.
Columbid(B. In the vast majority apocentric B, occasionally one and very often
both absent, but in Columba Jivia and Columba cenas traces of i^^landular
structure and so apparently apocentric C.
CUCULIFORMES.
Cuculi. Archecentrie to apocentric A in the Cuculidte ; in the Musophagidaj
apocentric B, and usually, if not always, quite absent. In the Psittaci
apocentric B.
CoRACliFORMES. In the Coraciidfe, Momotidae, and Meropidsc archecentrie ; in the
Alcedinidoe and Upupidye apocentric B.
SttHffes. Apocentric A.
Caprimulgi. Archecentrie.
Trocjones. Archecentrie.
Cypseli, Colii, and Pici are apocentric B.
Passerifor:mes. All that I have examined are apocentric C.
Gadow (i2. pp. 688-693) has made a very careful study of the relations between the
character of the c;eca and the nature of the diet, and I have gained further information
from Finn (8) and from Evans (7), and from the keepers at the Zoological Gardens as to
the diet of birds. There is very far from an exact correlation between the apocen-
tricities and particular diets, but there is enough to make it plain that many of the
modifications are homoplastic ; that is to say, that they are multiradial apoceotricities
on which no argument as to affinity can be based.
Archecentrie Type. — The diet in these forms is by no means uniform, but there is
a preponderance of feeding on insects and caterpillars, grubs, and so forth ;
in fact, of what may be taken as more typical rejjtilian diet.
Ajwcentric Type A. A very large number of birds displaying much enlarged caeca
live on vegetable diet in which cellulose bulks largely, and it is to be remembered
that the digestion of cellulose occurs in the large cseca of some mammalia.
Apocentric Type B. Carnivorous diet, piscivorous diet, and especially frugivorous
diet are specially associated with the degeneration of the caeca to iunctionless
vestiges, or even with the complete disajipearance of the cseca.
Apocentric Type C. AVhen it is remembered that the vast majority of the I'asseres
display this type, and that yet almost every possible form of diet is found among
them, it is at once clear that diet is no complete explanation of the character of
the ca3ca. I am more inclined to think that type C is to be associated with a
high development of the Avian structure, and that it is a peculiar secretory or
excretory organ.
SECOND SERIES. — ZOOLOGV, VOL. VIII. 40
270 DE. P. CHALMERS MITCHELL ON THE
Some parallels may make the complexity of the problem of the cjeca apparent. The
Passeres display all diets and one type.
The Owls and the Falconiformes have almost identical diet, the larger forms of both
being carnivorous, the smaller forms chiefly insectivorous ; the Owls all exhibit apocen-
tricity A, the Falconiformes apocentricity B.
Fowls and Pigeons (except the fruit-eating Pigeons) have a similar diet ; Fowls all
exhibit apocentricity A ; Pigeons apocentricity B or rarely C
The Gulls have a similar diet consisting of lisli and garbage ; most exhibit apocentricity
B, but Stercnrarkis is archecentric.
On the other hand, Phoenicopteriis is distinguished from other Ciconiiformes by a
nearly purely vegetable diet, and it has the longest cseca in the group. The Mergansers
are distinguished from the other AnatidjB by their typically piscivorous diet, and they
alone among the Anatidie have apocentricity B as in most other fish-eaters.
Characters and Classification.
In tlie systematic descriptive part, my task was to treat the characters of the patterns
displayed by different birds as nearly as possible as if the gut were tlie whole animal, and
the various phylogenetic figures and the three Plates display Avhat I take to be the
relations of the intestinal tracts, and not necessarily the relations of the possessors of
these tracts. I have been taking, in fact, the anatomical structure as the unit, and not
the individual or the species. In a sense, this is a return to the old Hunterian method;
but its purpose reflects on the new problem of the nature and evolution of varieties and
species of genera and families. Granting that the Plates attached to this paper represent
with approximate accuracy the pbylogeny of the intestinal tract in birds, we have yet to
learn the relation of the phylogenetic tree of this structure to the phylogenetic trees of
other structures, and the relation of all these to the phylogenetic trees of these im-
permanent combinations of characters that we call species. Although the coincidence of
such trees is frequently assumed, there is no a priori reason to support such a proposition ;
and there is much recent work on the nature of characters and of their inheritance to
throw doubt on the proposition. The nature of the anatomical structure in any organism
depends in the first place on the nature of cei'taiu material transmitted from the parents
in the fertilized ovum (naturally it does not matter to the argument whether the trans-
mitted substance be what we call "matter" or "state of matter" sucli as mode of
rhvthm). Among other writers de Vries (6) has recently brought strong experimental
evidence to show that at least in plants the hereditary material is composed of inde-
pendent units which may be sorted out and recombined in each sexual generation. Of
such independent variables underlying the fully developed anatomical structures of
animals, we know practically nothing as to their number, nature, or modes of sorting
out and recombination as they pass from generation to generation. Next, the anatomical
structures of animals depend on the environment in which the combination of transmitted
units come to maturity as actual adult organs ; that is to say, they depend on the
INTESTINAL TRACT OF BIRDS. 271
various correlations with other organs and witli external conditions established during
embryonic, youthful, and adult life. In the case of tiie alimentary canal we know just
enough, from the trausforaiing experiments of Hunter and others, to feel certain that
a large part of the tinal structiu-e is determined by the conditions in Avhich the hereditary
material is grown. The phylogenetic tree of an organ sucii as the alimentary tract may
be little more than the exhibition of a long series of experiments in growing similar or
identical " germs " in different culture media. And, finally, we know nothing as to how
far these liereditary masses, as they were marshalled into the actual branches of the
actual phylogeuetic tree of the organisms that contained them, carried within themselves
historical limitations or determinations towards further development in specialized
directions.
Such questions, however they may seem remote from an actual study in comparative
organogeny, in reality lie at the root of every attempt to use characters in classification:
if by classification there is implied more than the convenient disposal of particular
animals in ])articular pigeon-holes according to their possession of obvious anatomical
cliaracters. The study of characters by theniselves and for tliemselves inust precede
the attempt to use these characters in genealogical classification.
In tlie study of characters, the first proposition is whether they are primitive or
modified, archecentric or apocentric. So far as the subject-matter of this paper is
concerned, I have tried to sliow that for Aves the presence of a specialized duodenal
loop, of a Meckel's tract consisting of a nearly circular expanse of mesentery Avitii a
simply convoluted gut suspended at its periphery and symmetrical about a median vein
running towards a vestige of Meckel's diverticulum, of a pair of colic cfeca with free
lumen of moderate length and Avith walls partly glandular and partly absorbing, and of
a rectum relatively long, are archecentric characters. When birds possess many or even
all of these characters, we are tempted to say, but we cannot say definitely, that they are
closely akin. The retention by some descendants of a common stock of an ancestral
heritiige in the ancestral form does not imply that such members are niore closely akin
than other members that have dissipated or improved the common heritage. I liave
shown that in all the great accej)ted groups of birds there are some members with the
archecentric condition.
The second proposition that may have to be made regarding characters is that they
are modified from the archecentric condition, or apocentric, and in this modification
degrees of apocentricity may be exhibited. I liave shown that tlie lengthening and
twisting of the duodenum, the appearance of asymmetry in Meckel's tract and its produc-
tion into variously-formed and variously-placed loops, the special formation of the loops
described as supra-caecal kinks and supra-duodenal loops, the great lengthening of the
colic caeca or their reduction to vestiges or to short compact glands, and the shortening
and straightening of the rectum *, are apocentric characters. That two birds possess
* Since writing this memoir, 1 liave been interested by noticing that Professor Metschnikoff attribntes manv of the
digestive troubles of man to inhcritauee of a long rectum. The higher birds of every group apparently have passed
through a similar condition, but have succeeded in very greatly reducing the length of the rectum. — P. C. M.
40"
272 DR. P. CHALMERS MITCHELL ON THE
apocentricity in an organ is no evidence of their affinity. In the first place, the apocen-
tricity may he the mere result of growing the same inherited " germs " in similar
culture media. I call such apocentricities multiradial, implying that they are the result
of similar conditions on the same plastic material. The lengthening of the whole gut
and the spiral twisting of portions of it, and in particular the lengthening of the cseca,
ai*e plastic or multiradial effects and can have no direct bearing on affinity. The
extreme shortening and widening of the whole gut and the disappearance of the cteca,
or even their transformation to nipple-like excretory organs, are multiradial. The
production of a supra-duodenal loop and its retention after shortening of the long cseca
in connection with which it arose are multiradial apocentricities. These again give us
no clue to affinity. There is no reason to suppose that even in the actual phylogenetic
tree of birds, a branch the members of which now possess an organ with archecentric
character, may not have come from a branch the members of whicii now possess an
apocentric character in the same organ. For, in the first place, the apocentricity may
have arisen after the branching; and, in the second place, if the apocentricity be truly
plastic, the transmitted germs in another environment may grow only into the ancestral
form. It is probable, however, that apocentricities, even if multiradial, leave some
legacy of coraj)lexity in their simplified descendants, and such conditions of character I
have called pseiidoceutric. The conclusion to which consideration of miiltiradial apocen-
tricity leads is that even if the phylogenetic trees based on the study of the intestinal
tract be absolutely correct, and if they be compared with equally correct trees based on
the examination and valuation of other characters, these trees may not coincide.
Apocentricities, however, may be so definite;, so precise, or anatomically so complicated,
that they appear to imply a phylogenetic contraction of the range of variability in
respect- to the structure in question. Such a demarcation of modification along a single
and definite radius I have called uniradial apocentricity. When furtlier divergent
modifications occur on a single radius, there is formed what I have termed a metacentre,
and what seems to imply that the range of vai'iability has been limited or defined, with
the result that future modifications all retain an indication of their more limited or
defined starting-point. I take it that the archecoutre of the alimentary canal of birds
is a metacentre of the alimentary canal of the Sauropsida, and that, again, is a metacentre
of th(! alimentary canal of the whole vertebrate stock. I have tried to show that the
uniradial apocentricities of the intestinal tract of birds relate to the number and position
of the minor loops into which Meckel's tract is produced.
The obvious use of the study of characters as regards classification is, then, the
valuation of characters as archecentric and apocentric, and the discovery among the
apocentricities of the uniradial modifications with their metacentres. When these have
been determined and valued, the characters have yielded all the material that they aff"ord
for genealogical enquiry. When one set has been exhausted, recourse must be made to
another set. In other words, the work of the anatomist whose i^oal is the achievement
of natural classification, is the study of the definition and limitation of A'ariation within
groups. There is, however, another and most important side to this valuation of
INTESTINAL TRACT OF ETRDS. 273
zoological characters. "When the archecentricities and apocentricities of a number of
organs or structures liavc been mapped out, the questions ai*e opened as to bow far
arcbecentric conditions of different organs are combined in the same animal ; how far
apocentric conditions ai'e so combined ; how far arcbecentric conditions of one set of
structures are associated witli apocentric conditions of other sets. In general terms, the
problem is how far different organs become modified or remain unmodified independently
or in association in tbc same individuals. If we compare the l)og-fish and the Frog, we
find that, in relation to the general characters of the Iclitbyopsida, the Dog-fish is arche-
centric as regards its skeletal system, apocentric as regards its urinc-genilal system,
while tbc Frog is apocentric in its skeleton and arcbecentric in its ui"i no-genital system.
Naturally, in a ])a[)er dealing with part of one system, this problem of association has
not been raised, but I desire to add my conviction, based on a general study of birds and
a particular study of small groups of birds, that the larger the number of forms in a
^roup and the nari-ower the anatomical range within which the group is confined, the
more close is tlie association of apocentric modification of different organs.
BIBLIOGRAPHY.
(i) AxDERSsoN, O. \. — Zur Kcinitniss des sympathichsen Nerveusystems der Urodelen Amphibien.
Zool. Jaliib. vol. V. pp. 181-210.
(2) Beduard, F. E. — The Structure and Classificatiou of Birds. London, 1898.
(3) Berrv, R. J. A. — Tlie True Csecal Apex or the Vermiform Appendix ; its Minute and Comparative
Anatomy. Journ. of Anat. xxxv. pp. 83-100.
(4) Birmingh.\m, a. — Some Points in the Anatomy of the Digestive System. Journ. of Anat. xxxv.
pp. ;5;5-6(i.
(5) CuviER, Cir. — Lch;ous d'Anatomie oomparee. 2ud edition, 1835.
(b) WE Vries, H. — Tl^e Law of Disjunction of Hybrids. Coniptes Kendus, 1900, p. 8-lu.
(7) Evans, .\. II.— Birds. London, 1899.
(8) Finn, F. — A Guide to the Zoological Collections in the Bird (Jallcry of the Indian ^luscuni.
Calcutta, 19tK).
(6) FijRHKixoER, M. — Untcrsuchungcn znr Morphologic und SystematiU der \'ogcl. 1888.
(ic) Gadow, H. — \ ersuch einer verglcichenden Anatomic des Verdauungssystemcs der Vogel. Jenaische
Zeitschr. xiii. 1879, pp. 92-171 & 339-103.
(ii) lu. — On the Taxonomic \'aluc of the Intestinal Convolntions in Birds. I'roc. Zool. Soe 1889,
pp. 303-315.
(12) Id. — "Vogel " in Bronn's Klassen und Ordnungcn des Thicr-Reich. I89I.
(13) Garrod, .\. H. — On the Anatomy of the Hoatzm. Froc. Zool. Soc. 1879, p. 109.
(14) Gegenuauk, C. — Verglcichcnde Anatomic der Wirbclthicre. 18S9.
(15) Home, Sir E. — On the ditlerent structures and situations of the Solvent (jlands of Birds
according to the nature of their food and their ditlcrcnt modes of life. Phil. Trans. 1M12,
p. 39 k
(16) Id. — The course of the lutcsliue, with the Varieties ic the form of tiie Cieca in the Carni\ orous,
Piscivorous, aud Graminiferous Birds. Phil. Trans. 1814.
274 M{. P. CHALMEKS MITCHELL ON THE
(it) Hutchison, R. — Food and the Principles of Dietetics. London, 1900.
(i8) Klaatsch, H. — Zur Morphologic dcr Meseuterialbildungeu am Darmkaual der Wirbelthiere.
Morph. Jahrb. 1892, pp. 385-450, 609-716.
(iq) Kollmann, J. — Lehrbueh d. Entwickelungsgeschichte d. Menscliens. Jena, 1898.
(20) Lankestek, E. Ray. — On the Use of the term Homology in modern Zoology, and the
distinction between Homogenetic and Homoplastic agreements. Ana. Mag. Nat. Hist.
1870, vi. pp. 34-43.
(21) LoNNBERG, E., und Jagerskiold, L. — Ueber das Vorkommen elnes Darmdivertikels bei Vugeln.
Verhandl. d. biolog. Vereins zu Stockholm, vol. iii. 1890-91, pp. 32-36.
(22) Macgillivr.w, W. — Observations on the Digestive Organs of Birds. Magazine of Zool. and
Botany, 1837, pp. 125-136.
(23) Marage, R. — Anatomie Descriptive du Sympathique chez les Oiseaux. Ann. Sci. Nat., Zool. vii.
pp. 1-72.
(24) Mathes, p. — Zur Morphologie der Mesenterialbildungen bei Amphibien. Morph. Jahrb. 1895,
pp. 256-288.
(25) Mitchell, P. Chalmers. — On the Anatomy of Chauna chavaria. Proc. Zool. Soc. 1895,
pp. 350-358.
(26) Id.— On the Intestinal Tract of Birds. Proc. Zool. Soc. 1896, pp. 13G-159, figs. 1-25.
(27) Id. — A Contribution to the Anatomy of the Hoatzin [Opisthocomus cristattts). Proc. Zool. Soc.
1896, pp. 618-628, fig. 1.
(28) Id.- — On so-called " Quintociibitalism " in the "Wing of Birds, with special reference to the
Columbae. Journ. Linn. Soc, Zoology, vol. xxvii. pp. 210-236.
(29) Id. — On the Anatomy of the Kingfishers, with special reference to the conditions in the Wing
known as Eutaxy and Diastataxy. The Ibis, 1901, pp. 97-123.
(30) Oppel, a. — Lehrbueh der vergleichenden mikroskopischen Anatomic der Wirbelthiere. Vol. ii.
1897.
(31 ) Owen, Sir R., in Todd's Cyclopaedia of Anatomy and Physiology. 1836.
(32) Remak, in Muller's Archiv, 1843, p. 481.
(33) Id. — Ueber ein selijstandiges Darmnervensystem. Berlin, 1847.
(34) Thebault, V. — Etude des Rapports qui existent entre les Systemes pueumogastrique et
sympathique chez les Oiseaux. Ann. Sci. Naturelles, Zool., vol. vi. pp. 1-252.
(35) Toldt, C. — Ban und Wachstumsveninderungeu der Gekrose des menschlichen Darmcanales.
"Wiener Denkschr. 1879, pp. 1-56.
(36) Wiedersheim, R. — Lehrbueh der vergleichenden Anatomie der Wirbelthiere. 2ud edition. 1886.
INTESTINAL TKACT OF BIRDS. 275
EXPLANATION OF THE PLATES.
Plate 21.
Kv()l\itioii of tlie Intestinal Tract in tlie Pelargo-Colyiiiboinorphinc Brigade.
[For Colymbimorphine read Colymbomorpliiiie.)
Plate 22.
Kvolntion of the Intestinal Tract in the Alcctoromorphine Legion.
Plate 23.
Involution of the Intestinal Tract in the Coraciomorpliine Legion.
[For Coracimorphine read Coraciomorpliine.)
In these Plates the affinities of the conformations of the Intestinal Tracts and not of the birds are
indicated. The outline of the coils of the tract in the unfolded condition is given ; the line of attachment
of the mesentery to the body lies to the reader's right. The diverticulum is represented as a dai'k spot
on the l(>tt. The duodenuni lies towards the upper end of the Plates, the rectum towards the lower end.
\\n\\ the cu'ca, when present, as a pair of d-.irk lines.
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THE
TRANSACiIOi\S
OP
THE LINNEAN SOCIETY OF LONDON
SUE QUELQFES LEPADIDES NOUVEAUX DE LA COLLECTION
DU BEITLSH MUSEUM.
. ^ PAR
A. GRUVEL,
MAITRE.DE CONFERENCES A LA FACDLTE DBS SCIENCES, UNIVERSITE DE BORDEADX.
[Communique par M. le Prof. G. B. Howes, Sec. Liun. Soc.)
^L O N D O N :
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!77 J
VIII. S/n- quelqiies Lepadldes noiioeaux de In Collection da British Jfuseum.
Far A. Grt'vel, MuHre de Coiifereiices <) la Factilte des Sciences, Universite
de BordeauA-. {CotiDnxniqne par M. Ic Prof. G. B. Howes, Sec. Linn.
Soc.)
-J
(Planche 24 et 5 figs, dans le texte.)
Lu le (I mars, lOOL'.
UURANT le sejoui- que j'ai fait a Loiidres, pendant les jiraudes vacances de 1900,
j'avais mis de cote uu certain nombre d'(''cliantillons de Lepadides que je cousidcrais
comma devant representer des especes nouvelles. et que, la duree de mon sejour, mal-
heiireusement trop courte. ue me permettait pas d'etudier sui- place, d'une fa9on
suffisamment a2)profondie.
Avec rautorisation. de Femineut Dii-ecteur du British ^luseum, M. le Prof. Jeffrey
Bell eut I'ainahilite de me faire envoyev a Bordeaux les cchantillons que je lui av^ais
desiiynes, et c'est le resultat des etudes dont ils ont ete I'objet de ma part, que je viens
consigner ici.
Les especes nouvelles provenant de cet envoi se repartisseut en trois genres : le genre
Alepas, le g. Pweilasma et le g. Scalpel/iin/.
SOIS-OKIJKE DES rEDONClMS. (I'EDUXCULATA.)
Faniille des Aiiasjiide.s. (iVxASl'iDJ:.) *
Sous-famille des ^llrpdillnt's. (^Vlepadi.we.)
I. Genre Alepas, .Sander- Itang, 182i).
Ce genre est un de ceux, ])eut-etre celui qui presente le jilus d'uuiformite d'organ-
isation parini les especes qui le composent. l^a dilferenciation a etablir entre elles est
d'autant plus delicate que ces auimaux, etant, soit entieremeut priv('\s de plaques capitu-
laires. soit reduits simplement k des scuta comes, sous cutieulaires, et sans forme bien
definie, n'ofFrent souvent que des caracteres exti'iieurs pen nets et \ariables d'uu
* Bicn que le genre Anaspklcs ait ete cree par G. M. Thomson pour un type nouveau de Sehizopode (Trans.
L'lm. Soc, 2nd scr. Zool. vol. vi. part 3, August 1S94), le nom de Anas/ndc: me semble pouvoir rtre conserve pour
uue famille do t'irrbipedes. — A. G.
SECOND SEIUKS. — ZOOLOGY, VOL. VIII. 41
278 A. GEUVEL— SUR QUELQUES LEPADIDES NOUVEAUX
echantilloii u rautred'une meme espece. II fautdouc, dans la plupavt des cas, s'attaclier
plus encore aux caracteres fournis par ranimal lui-meme, que par ceux donnes par la
Ibniic du capitulum et du pcdoncule, et ces caracteres internes sont eux-memes, parfois
si pen nets, que la difTiculte pour caracteriser nettement les especes en est ainsi singu-
lierement augmcutee.
Ainsi par exemple, entre les deux especes que je vais decrire, A. Belli et A. micro-
stoma, il esiste des differences tresnettes, qui ni'ont permis de les separer, mais neanmoins
si I'on prend certains types de chacune d'elles, dont les caracteres ne sont pas absolument
nets, quelqu'un qui ne serait pas bien exerce, les confondrait a pen pres surement, tout
au moins par leur forme exterieure. II est bon de decrire comme type de I'espece, dans
ce cas, des etres absolument ]^intacts, c est a dire, aucunement dcformes, ni par pression
:\ I'etat vivant on dans les flacons d'alcool, ou par toute autre cause. On sait, en effet,
combieu, parfois, ces animaux revctvis d'une enveloppe niolle, se laissent deformer et
rendre mcconnaissables quand ils sont places vivants et en trop grand nombre dans un
flacon d'alcool.
On ne saurait trop recommander aux voyageurs de ne pas tasser ainsi ces etres
mous dans leur flacons, comme ils le font, belas ! trop souvent, pour gagner un peu
dc place.
Les memes precautions peuvent ne pas etre prises quand il s'agit de Scalpelhmi, par
exemple, car, du moins en general, les plaques qui recouvrent le capitulum et le pedoncule
de ces animaux leur donnent une rigidite suffisante pour eviter a peu pres toute
<leformation.
J'ai rencontre dans cet envoi du British Museum trois especes nouvelles CCAlepas,
aux quelles j'ai donne les noms d'^. Belli, A. miernstoma, et A. indica.
Ellcs sont, en general, representees par des echantillons d'assez grande taille, surtout
111 derniere qui mesure une longueur totale de sept centimetres.
1. Alepas Belli, nov. sp. (PI. 24. tigs. 1 A, 2, 3, 17, 18, 19, 28, & 29.)
Diagnose. — Capitulum a peu pres triangulaire, avec bord anterieur presque droit.
Pas de Crete dorsale veritable, mais legere saillie sur toute la longueur. Orifice externe
allonge, retreci a sa partie superieure, arrondi, au contraire, a sa partie iuferieure.
Pas de scuta.
Cuticule presque iisse, avec, seulemeut, quelques plis irreguliers. Surface dorsale
absolument Iisse, delimitee par un sillon.
Pedoncule de forme cylindrique, separe du capitulum par un leger retrecissement de
celui-ci.
Appendices caudaux avec quinze articles.
Piames internes des 5'' et (5'' paires de ciirhes atrophiees et portant, chacune, vingt-sept
articles.
llabitat. — Cotes de Cuba.
DE LA COLLECTION DU BRITISH BIUSEUM. 279
Cette espece, dediee an Professeuv J. Bell, dn British Museum, se rap[)r()che
d'A. Lankestcrl. A. (iruvel.
La forme generale de I'auimal est assez gracieuse, uii peu elancee et non massive. La
longueur du podoncule est bien proportionnee ;\ celle du caiiitulum.
Le capitnlum est ii peu pres triangulaire, avec son bord antericur presque droit et sou
bord dorsal a peu pres regulierement arrondi. Cependant il fait une legere saiilie vers
son tiers iuferieurs et se releve au-dessus de I'oriilcc du capituluni \v)\\.v former une
saiilie assez develoj)pee j)recedee d'une partie legerement ereuse. 11 ny a ])as dir crete
veritable, mais ce bord dorsal fait cependant sur toute sa longueur une legere saiilie
aplatie.
Les parties iaterales (luea])itulum sout nettement compriraees ce qui donne precis(''ment
a I'animal son elegance particuliere. La partie la plus large, antero-posterieure, du
capituluni est placee au niveau de I'insertion du muscle adducteur des scuta.
L'orifice externe est de forme allongee, retreci vers sa partie superieure, dilate eii rond
a sa partie inferieure oil vieunent se placer les cirrlies. Les levres qui le bordent
presentent, exterieurement, un bord legerement deprime et remjili de plissements
paralleles les uus aux autres et tons perpendiculaires au bord anterieur de l'orifice.
La cuticule qui recouvre le capituluni est presque lisse, marquee seulement de quelques
plis irreguliers, rares ; mais il existe dans la region dorsale et de chaque cote un sillon
assez profond qui delimite une surface allongee, allant de la dtqiression siis-buccale
presque jusqu'a la jiartie inferieure du capituluni. Cette surface est extremement lisse,
au contraire du reste de la surface capitulaire, mais elle est sur le meme plan qu(^
celle-ci— c'est a dire, ni en creux ni en relief par rapport a elle. Le sillon qui la
delimite est rectiligne de chaque cote, puis remonte vers la partie dorsale, anterieurement
et posterieurement, pour aller rejoindre son syraetrique.
La cuticule est epaisse resistante, colorec en jaune tout au nioins, apres son
sejour dans I'alcool, et si peu transparente qu'il est impossible d'apercevoir au
travers le corps de I'animal, comme cela a lieu pour A. Laiikestrri, A. Gruvel, par
exf^mplc.
La surface externe de cette cuticule est ornee de granulations chitineuses cxrrondies
dont quelques-unes portent des crochets. II existe aussi quelques soies sensitives, mais
rares cependant.
Les dimensions du capitulum du plus grand eehantillou sont : — Longueur, 25 mm. ;
largeur, 16 mm.
Le pikloncule continue le capitulum dont il est separe par un leger retrecissement, sans
que, cependant, cette separation soit bien nette. II est un peu plus long que le
capitulum, mais beaucoup plus etroit, de forme cylindi'ique, c'est a dire avec une diametre
a peu pres le meme aussi bien au sommet qii'a la base. La cuticule s'y continue avec
les memes caracteres que siu- ie pedoucule, mais avec une epaisseur un peu moins forte,
cependant, et des sillous paralleles et plus nettement accuses.
Longueur du pedoncule, ;>2 mm. ; largeur, Do mm.
-11*
:^,S0
^. GKUVEL— SUE QUELQUES LEPADIDES NOUVEAUX
JiHbifcd.—Les rchanti lions cVJ. Belli i)rovieiinent des cotes de Cuba.
Collection du British Museum.
Mi.
/ \ ^^snx
AlcjidS Belli, nov. sp.
Fiir. 1. Mamelon buccal, vu par la partie supc'i-ieure : Lai., It-vre sup. : PI.s., palpe labial sup. ;
iiid., mandibule ; in.i ., maehoire ; PJ.i., palpe labial infcrieur.
Fig. 2. Palpe labial superieur droit.
Fig. 3. Mandibule droite.
Fig. 4. Miiclioire droite.
Fig. 5. Palpe labial infrrieur droit.
" Ap'pareil buccal. — Le labre fait unc forte saillie en avaut, sa ])artie interne est foi'te-
nient cliitinisee, et presente suv son bovd liljre des uodules chitineux arrondis repre-
sentant les dents. Les palpes sont allonges, aplatis en ranie et converts de polls sur
les bords antrrieurs et inferieurs (fij,'. 2).
Les niaudibiiles resseniblent beauconp ;i eelles d'yi. J.inikesUrl, elles sont cependant
DE LA COLLECTfOX DU HHITISII MUSEUM. 281
plus allong-ees et plus fortement arinees de pointes cliitineuses sur li's piirties laterales
des dents. Ces dents sont au nonibre des trois, la (iuatri(>me formant la poiute inferieure
do la piece (fig. 3).
Les mdchoires diilcrent siui])leiucut de colics (HA. LoalcHlei'i, ni ce que la partic
infci-ieurc, scalai-ifornie, se termine en saillie a Tangle inferleur (jui est arrondi dans
I'espece in-cc6dente. L'encocbe est aussi un peu uioius profondo cliez A. Belli
(fig- iO-
Enfin les palpes de la lecre inferieure constituent les pieces les plus volumineuses.
Lour bord libre est nettement arrondi et convert de deux touffes de soies longues,
flcxibles, nou separoes par uue encoclie (lig. .")).
Cirrhes. — En ce qui eouccrne les cirrbes, j'ai rencontre dans cette espeee des anomalies
noml)reuses et tout il fait singulieres. L'atropbie des rames est un fait extrcmement
commun, et je crois ponvoir dire, d'apres les ecbantillons examines que cette atrophic
se rencontre plus souvent que la disposition normale. Je ne veux pas parler en fait
d'irregularites de l'atropbie qui se manifeste sur la i-ame interne des 7y et (V paires de
cirrhes. Celle-ci est, en etfet, normale ; mais j'ai trouve des rames atrojijliit'cs sur les
quatre autres paires — tantot la rame externe, tan tot Uinterne, tantot d'un cote, tantot de
I'autrc. Cette irregularite absolue indiquc nettement que c'est la un phenomene
accidentel et dont il est impossible de tenir compte dans la speciticatiou. Seuls les
caracteres permunents doivent etre signales. Si j'insiste siu- ce point, c'est pour montrer
I'inconvenient grave qui pent resulter de I'etude des cirrbes cliez un seul individu, et dont
le resultat peut-eti'c une crreur de determination.
!"■ paire (PL 24. tig. 28). Les deux rames sont inegales, la rame anterieure etaut
plus courte que la posterieure d'environ le (piart de la longueur de cette derniere. EUes
sont epaisses ct garnies de tres nombreuscs soies longues tines et legerement l)arbelees
sur lour extremitc libre et la raoitie de leur longueur.
•1' paire. Karnes a pen pros egales, I'externe depassant cependant legerement I'interne.
Leur longueur est environ la moitic en plus de la rame posterieure de la 1"' paire. Les
articles sont assez allonges et chacun d'eux porte, anterieurement, une seule paire de
soies longues, flnement barbelees a I'extremite et a sa base deux paires tres courtes,
glabres, I'anterieure depassant lui pen la posterieure.
'i' paire et ^' ptairc a peu pros identiques a la deuxiemo, mais leur longueur est un pen
plus considerable.
.■)'■ paire et G" paire. Les rames externes sont normalcs et semblables aux deux
precodentcs, mais les rames internes sont atrophiees et egales, cbacune d'elle portant
27 arlieles. Ijcur longueur atteint seulement environ le ^/cr.s' de cello de la rame externe.
Les soies qu'elles portent sont aussi extremement reduites, excepte \v dernier article qui
porte une toutle de soies plus longues.
AppeiHlices terminaux (Caudal appendages). — Us sont I'ormes de I.") articles seulement,
assez largos a la base, avec des soies nuUes on tres courtes ; les articles s'allongent vers
le sommet, ainsi que les soies qui, au nombre de trois ou quatre, couronnent le sommet
du dernier article.
282 A. fTRUYEL— SUE QUELQUES LKPADIDES NOUVEAUX
Api^endices fiJamcnteux (Filamontaiy appendages). — Une seule paire, courte, termiiiee
en pointe effilee et situee a la base de la 1™ paire de cirrhes.
JPchiis. — Le penis (PI. 24. fig. 29) est long (env. 8 mm.). Forme cylindrique, allant
en s'effilant regulierement jusqu'au sommet qui est en pointe mousse.
Annnlation nette suv la moitie terminale ; quelques rares poils dissemines a la surface
et une toufFe a son sommet.
AJJinifes. — -En remerciements de I'extreme bienveillance avec laquelle M. le Professeur
Jeffrey Bell a l)ien voulu in'accueillir an British Museum pendant mousrjour a Londres,
et aussi pour la grande courtoisie qui a toujoui's regno dans nos rapports reciproques, je
lui dedie cette espece nouvelle. Alejx'.s Belli est voisin d .^. Laiikrsferi.
2. Alefas microstoma, nov. sp. (PI. 24. figs. 1 B & B', 7 & 8.)
Diagjiosc. — Forme gendrale plus globuleuse que celle du precedent. Bord anterieur
du capitulum droit an niveau de Torifice externe, mais saillant et arrondi en dessous.
Orifice externe etroit, cordiforme avec une gouttiere dorsale. Le capitulum presente
une legere crete tout le long du bord dorsal, surtout developp;^e a la partie inferieure.
Toute la surface est striee de sillons profonds, nonibreux et irreguliers, excepte une
surface courbe dorsale, delimitee par un sillon qui en fait tout Ic tour.
Pas de scuta. Cuticule jilus mince que celle d'J. Bell i.
Pedoncule a pen pres regulierement cylindrique.
Appendices caudaux formes de 15 articles.
Rames internes des 5'' et 6'' paires de cirrlies atrophiees et iaegales (29 et 20 articles).
Habitat. — Madere.
J'ai appele cette espece A. microstoma a cause de la petitesse relative de son orifice
externe. Voisine de la j)recedente.
La forme generale de I'animal est plus globuleuse que celle du precedent. Le
capitulum an lieu d'etre comprime est plutot un peu reufie lateralement ; sa longueur
est moindre et sa largeur plus considerable, ce qui, par rapport a rensemble, lui doune un
aspect moins elegant et plus trapu que le premier.
La plupart des grands echantillons que j'ai eu entre les nioius etaieut plus ou moins
deformes et les caracteres semblaient moins nets que sur un plus petit, absolument
intact et que je prendrai par consequent conime type de cette esjiece, au point de vue,
tout aix moins, de la forme exterieure.
Le bord anterieur du capitulum est droit en face de I'orifice, mais, au-dessoiis, 0 fait
une saillie arrondie en avant et se retrecit a sa partie inferieure pour aller se continuer
avec le pedoncule. Le bord dorsal, regulierement arrondi sur toute sa longueur,
presente une tres petite crete qui s'accentue un pen plus a la partie inferieure.
L'orifice externe, sans etre saillant, forme une sorte de tube, uettement separe de la
partie inferieure du capitulum par un sillon transversal. Get orifice, assez petit, est
cordiforme, avec un tres petit sillon a sa partie superieure, qui s'elargit tout a coup tres
regulierement pour laisser passer les cii'rlies.
DE LA COLLECTION DU BHITISII MUSEUM. 283
Toiitc la surface du capitulum est sti'iee de sillons })i'ofonds, raais tivs nomln'ciix et
irreg'uliers, oxcepte une surface dorsale delimitee do cliaqiie cote par im sillon eourbe
qui, partant du sommet du capitulum, descend d'abord h peu pres parallelement u la
lisne dorsale, puis s'cn eloig'nc progressivement, pour veuir la rejoiudre hrusquemeiit
;i la partic inferieurc dvi capitulum.
I^a surface coiirbe ainsi delimitee est absolument lisse et un peu eu relief par rapport
au reste do la surface capitulaire.
Le capitulum est nettement separe du pedoncule par uu retrecissement brusque, mais
la cuticule se continue a sa surface avee les memes caracteres.
La cuticule est moins epaisse que chez A. Belli, de couleiu- blanchatre, sale. Ello
porte comme ornements des nodules avec ou sans crochets comme chez A. Lankeslevi,
A. Gruvel, ainsi que des soies sensitives assez nombreuses.
Les dimensions du plus grand echantillon sont : —
Longueur du capitulum, 23 mm. ; largeur, 17 mm.
Longueur du pedoncule, 21< mm. ; largeur, 9 mm.
Comme on le voit, le ])edoncule est seulement un peu plus long que le capitulum ; il
est a peu pres regu.lierement cylindrique, un peu plus large, cependant, aux deux
extremites qu'au centre.
Le manteau ne presente rien de pavticulier.
Habitat. — Les echantillons etudies proviennent de Maderc.
Collection du British Museum.
ApjHfreil buccal. — L'appareil buccal, dans son ensemble, est tcllement identiqvie a
celui de I'espece precedente, que je n'aurais, 2)our ainsi dire, qi\'a repeter mot pour mot
ce que j'en ai dit. Je le juge inutile.
Cirrhcs. — Je n'ai pas a repeter ici ce que je disais a propos d'^. Belli. Les deux
echantillons etudies presentaient tous deux une constitution normale, I'atrophie portant
seulement sur les rames internes des 5° et G" ptiires de cirrhes.
V i^aii'c. La rame posterieure depasse I'anterieure de ses 5 ou 6 derniers segments,
bien que le nombre des segments soit tres different : 32 a la grande, 17 a la petite, mais
cela tient a ce que chez la premiere ils sont beaucoup plus courts que chez la seconde.
Ces rames sont moins larges que chez A. Belli et cliaque article au lieu de porter
i;n grand nombre de soies, n'en presente qu'une vingtaine environ, qui sont surtout
tres courtes sur la rame jiosterieure, elles sont raides et tinement barbelees sur leur
moitie libre.
2' paire. Rames egales, chacune d'elle etant environ j ^In^ longue que la rame
posterieure de la 1'^'^^ paire. Elles sont formees par un grand nombre d'articles tres
courts, portant chacun deux grandcs soies longues raides et finement barbelees, deux
tres courtes en arrieres de celles-ci et deux semblaliles a ces dernieres et inserees
lateralement.
3' et V paires. Un peu i)lus longues que la deuxieme par les rames, dout I'interne est
plus courte de deux segments que I'externe. Meme disposition des soies.
281. A. GEUVEL— SUE QUELQUES LEPADIDES NOUVEAUX
5' jmirc. Eame interne atroiohiee, avec 29 articles, mais les soles soiit beaucoup
moins reduites que cliez A. Belli.
6' ]}aire. Rame interne egalemeut atvophiee, mais formee seulement de 26 articles.
liCS rames internes des 5' et (!' paires soiit done inegales, au contraire A' A. Belli oii
elles soot egales.
Appendiceit tervihianx (Caudal appendag'es). — Semblables a ceux de la precedente
espece. lis jjresentent egalement 15 articles, dont le terminal tres court, porte uu
bouquet de soies.
Ap2)en(licesJHamentenx (Filamentary appendages). — Une seule paire assez longue a la
base de la premiere paire de cirrlies.
Penis. — Court (env. 6 mm.), trapu, conique, termino en pointe mousse. Forme d'un
graaid nombrc d'anneaux tres courts, avec quelques tres rares soies u la surface et un
l)0uquet au sommet.
AJlfinites. — Cette espece se rapproclie beaucoup de la precedente, dont elle differe
cependant par la forme exterieure et un certain nombre de caracteres internes que nous
avons mis en relief.
Je lui ai donne le nom d'A. ndcrostoma, tire du caractrre anatomique de la bouche,
tres petite par rapport au volume de rauinial.
3. Alepas indica, nov. sp. (PI. 24. figs. 1 C, i & 1', 21 to 27.)
Diagnose. — Le corps entier de Tanimal semble etre tout d'une venue, sans separation
nette entre le capitulum et le pedoncule qui est extraordiuairement developpe. Le
capituluni est tres comprime lateralement, avec le bord anterieur droit et le bord dorsal
rcsulierement courbe ; ce bord porte une crete transparente, haute d'environ 1 mUli
metre, sur toute sa longueur. L'oritice externe a la forme d'un triangle curviligne sans
gouttiere dorsale, et borde de levres tres nettement frangees. La cuticle est mince,
transparente, et ornee de plissemeuts tres fins, assez difiicilement visibles a I'oeil nu.
Le pedoncule fait suite au capituluni sans transition. Sa longueur atteint environ
trois fois et demi, et sa largeur egaie a peu pres celle du capitulum.
Pas de scuta. Apjiendiccs caudaux formes de 12 articles. Rames internes des 5" et
G' paires de cirrlies atropliiees et inegales (25 et 21 articles).
Habitat. — Singajioure.
A cause de son origine, j'ai apjiele cette espece A. iuclica. Elle vient se placer,
quoique tres differente, quant a sa forme exterieure, entre A. cornitta, Darwin, et
A. iiiicrosloma, A. Gruvel.
Cette troisirme esjiece est, certainement, une des plus curieuses que je counaisse
appartenant a ce genre, a cause do la longueur vraiment liors de projiortion de son
pi'doncule relativement a celle du capitulum.
Bieii que les (''chantilions ne soicnt pas dans un remarquable etat de conservation, il
m'a ete possible d'eii faire letude com^jlete.
Le corps tout entier de I'aninial semble etre tout d'une venue a cause de la longueur
du pedoncule. II est coloir, aprrs scjour dans I'alcool, en inarron fonce tiraut uu
DE LA COLLECTION DU BRITISH MUSEUM. 285
peu sur la teintc lie de vin. La cuticule est prcsque aussi transparentc que cclle
d'A. Lankesleri.
Le cnpUulum est beaucoup plus comprimc lateralemeat que chez A. Belli. Sou bord
anterieui' est droit, Ic bord dorsal reguliercment courbe, porte une cretc transparentc,
haute de pres d'un millimi'tre, sur toute sa longueur.
Rien ue rapj^tolle la surfaee dorsale signalee chez les deux autres especes. La surface
toute entiere du capituluni est lisse, sans stries ni sillons, excepte dans la region dc
I'orifice.
Celui-ei a la forme d'un triangle curviligne, dont la partie infcricure est occupee par
les cirrlics, sans qu'il existe trace, ;i la partie superieure, d'une gouttiere quelconque,
comme dans I'esjjt'ce precedentc, jiar exemple. L'orifice est borde de levres trcs nettement
frangees, avec des sillons transversaux tres apparents, places sur les parties latt;rales.
Sans etre saillant, l'orifice dcpasse legerement le bord antero-inferieur du capitulum,
dont il est supare par unc legere depression au-dessus de I'insertion du muscle adducteur
des scuta.
Le capitulum se retrccit legerement a sa partie infcrieure pour former le pedoncule,
mais la limite nette entre les deux est assez difficile a saisir.
La cuticule mince, transparentc, est ornee de plissemcnts tres lins, assez difficilement
visibles a rccil uu. On trouve a sa surface des plages formees de tubercules chitineux
de differentes grosseur, les plus gros etant generalemeut au centre. On trouve dans
quelques-uncs de ces plages une sole sensitive, mais de position tout a fait variable.
Les plages elles-memes sont disposees tres irregulieiement.
Le pedoi.cale qui I'ait suite au cajiitulum est tout d'une venue. Sa longueur egalo
environ trois fois et demi celle du capitulum et sa largeur est a peine moins considerable.
La cuticule qui Ic reeouvre, continuation directe de celle du capitulum, est lisse ct
presente quelques ornements semblables a ceux deja decrits.
Les dimensions du plus grand ecliantillon sont : —
Longueur du capitulum, 21 mm. ; largeur, 18 mm.
Longueur du pedoncule, 70 mm. ; largeur, 13'5 mm.
llahitat. — Ces cchantillons proviennent dc Singapoure, dans I'lnde Anglaise. Offert par
J. Hussell, 1886.
British Museum Collection.
AppareU buccal. — Sauf quelques details, I'appareil buccal de cette espece est identique
a celui des deux precedentes.
Les dents du UiJtrc sont de simples eminences irregulieres de la cuticule, n'ayant que
vaguement la forme de dents. La partie interne de cette piece presente des ornements
qui sont sur les parties laterales de fortes soies et des formations pectinees robustes
(fig. 22) et dans la region moyenne des formations egalement pectinees, mais beaucoup
plus fines.
Les jjolpcs sont identiques (PI. 24. fig. 23). Les mandibules portent sur les parties
SECOND SlililES. — ZOOLOGY, VOL. VIII. 12
236 A. GKUVEL— SUE QUELQUES LEPADIDES NOUVEAUX
lateralcs, en arricre des dents, des formations pectinees tres robustes (fig. 21.). Les
mdchoires ot los palpes de la levre infericure ne presentent rien a signaler (PI. 24. figs. 25
et 20).
Cirrhes. — La syraotrie de toutes les rames est absolue, dans cettc espece et I'atropliie
ne se manifeste egalement que sur les rames internes des 5" et 6"' paires.
1" jmire. La rame posterieure depasse I'anterieure de ses 5 derniers articles. La rame
anterieure trt-s large a la base, est couverte d'un grande nombre de soies finement
barbelees.
2" iKtire. Les rames, egales en longueur, atteignent environ le double de la longueur
de la rame posterieure de la l"' paire. Les soies y sont disposees comme dans A. micro-
stoma, niais il existc en jjlus une paire de soies courtes en arriere, et la partie antero-
supt'rieure des segments est fortement saillaute.
3^ paire. Eames egales, depassant celles de la 2" paire d'cuviron \ de leur longueur.
4° imire. Rames egales, depassant celles de la 3"" paire d'^ environ de leur longueur.
h' paire. IJame externe normale. Rame interne atrophiee, formee de 25 articles,
avec soies atrophic'es. Le dernier article couronne par un bouquet de G soies.
6' pff?>(?. Eame externe normale. E-ame interne egalement atrophiee, mais formee
de 21 articles sculement.
Les rames internes des 5'" et 6'' paires sont done atrophiees et incgales comme cliez
A. microstoma.
C'est la un caraetere specitique, sur lequel je me suis appuye pour etablir le tableau
synoptique qui suit. II est suffisamment precis pour jiouvoir etre employe dans la classi-
fication. II est a remarquer, en effet, que, lorsque les rames atrophiees ne portent pas le
meme nombre d'artieles, il y a une difference d'au moins 3 ou 4, suffisante, par
consequent, pour iioTivoir etre facilement controlee.
A])pendices termi>/ava\ — Les appendices terininaux sont seulement formes de 12
segments ; ceux qui sont a la base sont larges et courts, les autres etroits et longs. Les
(^uatre derniers seuls presentent des soies nettcs et le dernier, court, se termiue par un
bouquet de soies.
Appendices fhiviruteux. — Une paire a la base de la 1" paire de cirrhe, autant que j'ai
pu le voir avec des animaux en aussi mauvais etat.
Penis. — Assez long et cylindrique assez peu nettement annele.
Affinites.—^\yAn\a\i nom cut etc bien porte par un animal, c'eut ete celui Ac peduncu-
laia, mais comme il appartient deja a une espece de Hoek, je designerai — tout simplement
ceiui-ci par son lieu d'origine — A. indica.
DE LA COLLECTION DU BRITISH MUSEUM.
287
Tableau sytioptiquc ties especes da (jenre Alepas, Saader-Rang.
1. Scuta
presttits.
f
Scuta presents
rnais reoou-
vorts par la
culiculc.
Orifice trcs coxirt, a Icvres
frangees, tril'S petite espoce,
Rames int. dcs 5'" j prosque toujours fixce sur
et 0" paires ( Cklaris -'• minula, Philippi.
iiornialcs.
Orifice non proeiuincnt, mais
nettcment tubulaire. I'ura-
site sur Me'duses A. pantslta ?, Sander-Ilan^
llames int. des 5' /
et G" paires I pormo carrt'e. Orifice nettemeut
atrophiees et \ tubulaire A. quadmia, Auriv.
, hu'gales. \
/Orifice petit, tubulaire, Icgcr.
preeminent. Pas de cretes
dorsales. Pcdoncaile asscz
Jq,^, A. peduncuhda, Huck.
Orifice tubuleux, proem incut et
^;tl.^,it A. tahuhmi ?, (Juoy ct Gainiard.
llames int. des 5'^
et 6° paires
normales.
Orifice leger. proePiincnt, ^ a -t
' ( cretes mediaues et dorsales,
, , . . pen developpees A-ja^^nniai, Auriv.
Uaiues atrophiees '
■1. Pas
de scuta.
C'jahs.
llames int. des
5'' et 0'' paires
atrophiees. \
llames atrophiees
iiierjales.
I Orifice legi-T. ouvert, pas de
\ Crete mediano, mais bord
dorsal leger. saillant de toute
sa lougueur I- -D'"', A. Uruvel.
/Orifice leger. saillant, trois
cretes dorsales assez developpees,
dont une au-dessus de I'orifice. A. coniuta, Darwin.
Orifice non saillant, allonge,
nou tubulaire, une Crete
saillante sur touto la partie
dorsale ^- ""''<-•«> -^- ^»'"^'^^-
Orifice etroit, cordiforme, une
legere Crete geueralc dorsale,
surtout dJveloppee a la base . A. microstoma, A. Giuv-d.
Orifice leger. tubulaire et sail-
lant, une tres legere crete
dorsale gc'nerale, cuticule ex-
trememcnt transparente .... A. Lanlcsliri, A. Gruvel.
42
)«
2S8 A. GRUVEL— SUE QUELQUES LEPADIDES NOUVEAUX
Famille des Pentaspides. (Pentaspid.^;.)
Sous-famille des Ll'padines. (Lepadik.e.)
II. Genre Pcecilasma, DarAvin, 1851.
Ce genre n'est representd, dans I'envoi qui ni'a ete fait, que par une seule espece et
meme un unique ecbautillon que, bien entenclu, j'ai pris ie plus grand soin de ne pas
deteriorer.
J'estime, en efFet, que toutes Ics fois qu'une espece est nouvelle, on qu'oa la suppose
telle pour de nombreuses raisons, il faut, quand elle n'est representee que par un seul
echantillon, se borner a decrire I'exterieur de celui-ci, afin d'en tirer les caracteres
speciliques. Mais lanimal doit etre conserver intact, comme type qui sera etudie plus
comjjlt'temeut s'il arrive qu'un ou plusieui's individus semblables soient trouves plus tard.
II se pent cependant, que, dans certains cas, les caracteres exterieurs seuls si ont
nbsoluuieut insuffisants poui* ctablir une diagnose precise. Dans ces conditions, I'etude
de I'animal lui-meme s'impo-e — mais alors, les plus grandes precautions doivent etre
prises pour le deteriorer le nioins possible et un dessin tres rigoureux doit etre fait avant
d'v touclier.
Dans le cas present, les caracteres exterieurs seuls, ni'ayant paru suffisants pour justifier
la creation d'une nouvelle espece, je n'ai pas cru devoir deteriorer le type et je ne
donnerai, par consequent, que la description de I'exterieur.
Get ecbautillon a ete trouve par moi, fixe sur le pedoucule de I'un des exemplaires
d'Alepas indica, A. Gravel, dont j'ai donne j^bis baut la description complete.
Pcecilasma minuta, nov. sp. (PI. 24. lig. 5.)
Diagnose. — La forme generale du corps est assez elegante et elancee. Le capitulum
est comprime lateralemcnt, surtout clans sa paitie superieure. Terga triangiilaires, a
apex pointu et saillant. Dord scutal avec une encocbe en face de laquelle vient se placer
le sommet du segment anterieur du scutum. Cette encocbe est suivie d'une dent qui se
place entre les deux segments de cette derniere plaque. Scuta tres developpes, formes
par deux segments, un anterieur allonge, un posterieur large.
Carene courte, uniformement citroite, avec une crete dorsale.
Pedoncule a peu pres regulierement cylinlrique ; atteint environ la moitit3 de la
longueur de capitulum.
Dimensions : —
Longueur du capitulum, 2'G mm. ; largeur, lo mm.
Longueur du pedoncule, 1-23 mm. ; largeur, 061 mm.
A cause de ses petites dimensions, j'ai donne a cette espece nom de P. minuta. Elle ne
se rapprocbe nettement d'aucune espece couuue. C'est cependant de P. lenticula, Audv.,
(ju'elle semble etre la plus voisiue.
Le corps tout entier de cet animal presentc une forme assez gracieuse et elegante.
Les plaques capitulaires sont minces et permettent d'apercevoir par transparence le
DE LA COLLECTION DU BEITISH MUSEUM. 289
corps de I'animal qu'elles recouvrent. Corame ce corps est colore en jaime-orangc,
cela donne au capitulum uiie teiute legere. La cuticule qui separe les plaques est jaunatre
et celle qui recouvre le pedoncule est un peu rouge-brique ; ces couleurs t'taut, bien
enteudu, cellos dc I'animal qui a scjourno longtemps dans I'alcool.
Le capitulum est cornprime lateralemeut surtout dans sa partie superioure. Les pieces
qui le recouvrent sont serrees les unes centre les autres et ne laissent aucun espace
purement membraneux,
Les terga sont triangulaires, a sommet pointu et saillant. L'anglo basal est tronque,
le bord basal presente de sa region anterieure une encoche, en face de laquelle vient se
placer le segment antcrieur du scutum correspondant. Cette encoche est suivie d'une
dent qui est exactement en face de la ligne de suture, entre les deux segments de ce
meme scutum. Le reste du bord est parfaitement droit.
Les scuta sont tres developpes et chacun est divise en deux segments, un anterieur
ot un lateral, par une scissure legerement convexe anterieurement qui va de Tumbo a
I'apex.
Le segment anterieur est tres saillant en avant. Le maximum etant environ au niveau
du tiers superieur. L'angle superieur est regulierement arrondi, Tangle basal tres aigu.
C'est lui qui porte I'insertion da muscle adducteur des scuta.
Le segment lateral est, de beaucoup, le plus developpe d('s deux. Le bord basal
droit foi'me une tres legere pointe a son point de contact avec Tangle basal du segment
anterieur. Le bord dorsal ou careaal est reguliercmeut courbe ; en fin le bord tergal est
absolument droit.
On aperQoit encore, a Tumbo des terga et des scuta, les plaques embryonaires. On ne
trou.ve de stries visibles ni sur Tune ni sur Tautre de ces pieces.
La carene est courte puisqu'elle depasse a peine la milieu du bord dorsal du capitulum.
Elle est uniformement etroite du sommet a la base, ou, cependant, elle s'elargit un peu.
EUe presente une crete dorsale nette, mais arrondie.
La base du capitulum se retrecit beaucoup pour donner insertion au pt'doncule qui,
aussit6t, se dilate et deborde legerement les 2:)ieces capitulaires formant cette l)ase.
La longueur du pedoncule ne depasse guere la moitie de celle du capitulum. II est u
peu pres regulierement cylindrique, un peu plus large cependant vers le sommet qu'i\
la base. La cuticule qui le recouvre, assez mince, jaune-rougeatre, presente de petites
granulations cbitineuses un peu plus colorees que le reste de la surface et disposees a
peu pres regulierement en series circulaires et paralleles.
Affinites. — En raison de Textreme petitesse de Tindividu unique qui a servi a cette
etude, j'ai donne a Tespece le nomdeP. miimta. Je ne veux pas dii"e, bien enteudu, que
cette espece est tres petite, il est possible qu'elle atteigne meme une certaine taille, mais je
crois qu'elle ne doit jamais depasser de beaucoup les dimensions que je viens de donner.
Par sa forme exterieure, cet animal n'est comparable a aucune espece connue. Celle
dont il semble le plus se rapproclier est P. lenticula, Aurivillius *.
* On trouvera daus mon Momoire sur los Cirrbiptdcs du ' Travailleur ' et du ' Talisman," le tableau synoptique
des cspeces du geure FosciUisiiia. — A. G.
200 A. GRUVEL— SUE QUELQUES LEPADIDES XOUVEAUX
Fcmiille cles Polyuspides. (Polyaspid.e.)
Suns-famille cles Pollicipines. (Pollicipinje.)
III. Genre Scalpellum, Leach, 1817.
Dans le genre Scalpellum deux especes seulement a signaler : Tune qui constitue une
cspecfi nouvelle et queje dedie au savant qui a etudie les Cirrliipedes du 'Challenger,'
le Dr. Hoek, petite espece, extremement curieuse et qui ne rappelle aucuue forme
actuolleoient conuue ; I'autre, que je crois devoir rattacher a So. sexcormitum, Pilsbry,
Ijiou qu'elle ditfere du type dccrit j)ar quelques caracteres, secondaires, il est vrai,
1. ScALPELLrM HoEKi, nov. sp. (PL 24. figs. G, 9 to IG, & 20.)
Dmfjnose. — Capitulum plutot un peu globuleux, foi-me de 14 plaques, serrees fortes, a
stries tres nettement marquees. Bord anterieur a peu pres regulierement coui'be, avec
cependant un angle a la limitc des scuta et des terga et une saillie anterieurc de Tumbo
des pieces rostro-laterales.
La cuticule qui recouvi'e les jdaques est tres mince et glabre.
Carcne presqiie droite a sa partie inferieure, fortement arquee a sa partie superieure.
Umbo voisin de I'apex. Pus d'aretes laterales. Surface dorsale arrondie. Terga a a^ex
legerement tourne en arrierc. Scuta a apex droit. Plaques iufra-laterales allongees,
etroitts ct dont la partie intVrieure est i'ortemcnt recourbce en avant. Umbo a la base.
Umbo des pieces careno-laterales a la base et ne depassant j^ar le bord externe de la
carene. Postie allonge en forme de triangle curviligue. Pas de sous-carene.
Pedoncule assez long par rapport au capitulum, cyiindro-conique, orne de buit series
longitudiiialcs et alternes d'ecailles tres allongees transversalement, libres, largement
separees les unes des autres, excepte dans la region capitulaire ; chaque serie porte 8 ou 9
ecailles.
CaintuUm) .—Y\vX(ii legerement globuleux, surtout dans sa region inferieur. Bord
anterieur, a p)eu pres regulierement courbe, mais formant une angle entre les terga et
les scuta et une saillie de I'umbo des plaques rostro-laterales. La largeur egale a peu pres
les f de sa hauteur. 11 est forme de 14 plaqu.es fortes, solides, serrees les unes centre les
autres, a stries tres nettement marquees, recouvertes d'une cuticule a'bsolument lisse.
Le bord basal est tres legerement concave et un peu incline de haut en has et d'avant en
arriere.
Scutum. — Porme presque trapezoide. Bord anterieur regulierement courbe ; apex a peu
pres droit a peine rentrani. Bord tergal tres legerement concave, bord supra-lateral
droit a peu pres egal au precedent. Bord basal convexe ct un peu plus long. Une
arete plate et un peu large \mit I'apex a Tangle latero-basal ; la plaque porte des stries
paralleles non saillantes, qid se coutinuent sur le reste de la surface.
Tercjum. — Irregulierement triangulaire, car le cote basal est casse en deux parties
faisant entre elles un angle tres ouvert. Le bord anterieur est regulierement et assez
fortement courbe ; I'apex tourne en arriere, le bord carenal presque droit. Une arete
peu marquee unit I'apex a Tangle basal.
DE LA COLLECTIOX DU BRITISH MUSEUM. 291
Carene. — Etroitc au sommet, elle s'ulargit rapidemeut jusqu'a la liasc ou elle est trrs
large et se termine eu pointe mousse qui s'avancc eutrc les jjieccs careno-laterales
juscju'au niveau du pedoncule. La partie dorsale est arrondie, Idgerement carenee et la
plaque nc prescntc pas trace d'arctes laterales. L'umbo est tres voisiii de I'apex. Dans
sa paitie superieure, cette plaque est fortement recourhee en avant, taudis qu'elle est
presque droite dans sa region moyenne et inferieure. Enfiii, entre le bord aatcrieur
arrondi et le bord careual des terga se trouve une petite surface jiarfaitcment calcitiee et
qui se trouve dans un ])lau iuferieur, plus profond par rapport au reste de la plaque.
Plaques supra-htlcrales. — Triangulaires, avec le bord tergal et le bord scutal droits et le
bord careno-basal irregulierement couvese. Cette plaque portc une arete plate longeant
le bord scutal tres fortement marquee de stiies paralleles qui se j)oursuivcnt sur toute la
surface. L'umbo est a I'apex qui est droit et en pointe mousse.
Plaques rostro-laterales. — Irregulierement quadrangulaires, avee le bord scutal presque
droit, un peu concave et tr^s obliquement dirige de haut en bas et d'arricrc en avant.
Le bord rosl ral tres court est concave, de sorte qu'entre les deux vient se placer le rostre.
Le bord basal, court, est droit et se continue par le bord lateral tres convexe s'appuyant
sur la plaque iufra-Iatt'-rale. Une arete par de l'umbo et va rejoindre Tangle lat^ro-basai,
delimitaut uue large surface la erale et uiie petite surface anterieurc triangulaire.
Hostre. — Le rostre est tres facilement visible a la surface. II a la forme d'un triangle
curviligne allonge. Ses cotes soat legerement reconverts par le bord rostral des plaques
rostro-laterales.
Plaques iajra-latcrales. — Tres caracteristiques, un peu en forme de corne d'abondanee.
La partie superieure est la plus large et la ])laque se retrrcit progressivement jusqu'a son
umbo qui est tout a fait a la l)ase. La plaque, aliongee a sa partie inferieure,
est fortement recourbee en avant. Elle est marquee de stries paralleles tres nettes.
Plaques careno-laterales. — Presque la lorme d'un triangle rectangle si le cote infra-
lateral etait droit, mais il est concave en avant. Le bord carenal est a peu pres droit et
tourne autour de la base de la carene en se dirigeant vers la region mediane et dorsale du
pedoncule oil il rejoint celui du cote oppose. Une arete plate, large au sommet, etroite a
la base, va de I'apex a la base en bordant le cote carenal de la plaque. Cette arete
presente des lignes paralleles tres nettes qui se continuent sur la plaque et viennent
aboutir au bord basal, oil elles ferment une sorte de dentelure.
Longueur du capitulum, 5'50 mm. ; largeur, 3 mm.
Pedoncule. — Le pedoncule est assez long, de forme cylindro-conique. II est orne de
8 series longitudinales et alternes de plaques tres allongees transversalement, serrees dans
la region capitulaire, mais partout aillcurs largement separees par un intervalle purement
chitineux ; leur bord libre presente une saillie mediane dirigee vers la partie inferieure. La
cuticule ne les recouvre que vers la base. Chaque s6rie presente 8 ou 9 ecailles a peu pres
semblables.
Longueur du pedoncule, 275 mm. ; largeur, 175 mm.
llahUat. — Ocean Pacifique, sans autre indication. Cinq echantillons fixes sur des
Bryozoaires.
292 A. GRUTEL— SUR QUELQUES LEPADIDES JSOUVEAUX
Observations et Jffiiiiles. — Cette espece ne ressemble a aucune des especes actuellement
connues, par la loniie dc ses plaques infra-laterales et careno-laterales. Mais par ses
autres caracteres elle vieut se placer a c6te de Sc. luridwn, Auriv., dont elle se distingue,
du reste, tres facilement, d'aprus les caracteres des pieces que je viens de signaler.
Etude de Vonivud proprement dit. — Labre allonge triangulaire, encadrant dans ses
deux branches laterales, toutes les autres pieces buccales. Son bord libre porte une
trentaiae de masses cliitineuses arrondies, petites, toutes semblables.
Les PaJpes qui s'attachent en arriere et en dedans sont allonges, triangulaires,
avec un bouquet de soies terminales un peu longues et qiielques soies dorsales tres
petites (tig. 13).
Les Mandihules portent trois dents fortes avec Tangle inferieur armc de trois pointcs
chitineuses fortes et sur les parties laterales, de soies courtes et robustes (PL 24. fig. 15).
Les Ildchoires portent sur leur bord libre une encoche peu profonde situee un peu
plus du cote dorsal que du cote ventral. La zone dorsale porte deux dents longues et
fortes suivies de deux autres rangees plus courtes et nioins robustes. Quant a la zone
inferieure les dents assez fortes semblent divisees en deux groupes, tres peu separcs du
reste. Les parties laterales presentent des soies courtes et assez fortes (tig. 11).
Les Falpes de la levre inferieure sont aplatis, larges, avec un bouquet de grandes soies
non barbelees a la pointe anterieure et d'autres sur le bord dorsal et sur le bord libre,
mais ici en moins grande abondance. II y en a aussi sur les parties laterales.
Girrhes. — Les cirrbes sans etre tres longs, le sont cependunt suffisaniment et ils sont
egalement, greles.
La V paire est assez eloignee de la 2'^. Les rames sont a peu pres egales, I'anterieure,
etant cependaut un peu plus courte que la posterieure qui semble formee de 8 articles,
mais les limites des deux articles basilaires sont peu nettes. Ces articles sont tres peu
saillants, amies de soies nombreuses, assez courtes et tres finement barbelees.
I;a 2' paire a les rames egales et atteignant environ 2 fois la longueur de celles de la
premiere. Chacune d'elles est formee de 10 articles, etroits, allonges, legerement saillants
en avant et portant, en moyenne, 6 series doubles de soies longues, surtout vers la limite
superieure des articles et finement barbelees.
Les autres paires de cirrhes sont a peu pres semblables a la 2% mais un peu plus
longues.
Ap)pendices terminaux. — Ces appendices sont formes de trois articles nets ; I'article
basal, le plus long, porte bien vers son milieu une sorte de ligne d'articulation, mais elle
ne comprend pas toutes la largeur de I'article et ne forme pas, par consequent, deux
segments distincts. Cliacun de ces trois articles porte des soies plus longues vers la
limite superieure des articles. Le segment terminal porte un bouquet de longues soies
non barbelees et d'autres plus courtes a la base.
Au milieu de cbaque article se trouve une soie ; deux sur I'article basal.
Penis. — Absent. Pas trace de testicules. J'ai trouve au-dessous du prosoma une
vingtaine d'ccufs gros, ovoides, et en voie de developpement.
DE LA COLLECTION DU BRITISH MUSEUM. 293
Sur les cinq cchantillons examines, trois portaient 2 mfilcs compleraentaires un sur
chaque cote, profondement eufonces clans la cavite intei-palleale, 2 n'cn portaient qviun,
mais commc on le voit, il y en avait snr tons.
Comuic uue etude complete de ccs petits etres sera publiee dans les Resultats des
Expeditions du 'Talisman ' ct du ' Travailleui',' la description du male de cette espece y
troiivera sa place, qui me semble beaucoup mieux justifiec dans iiu travail d'ensemble,
que dans une description isolce qui ue signifie pas grand'chosc.
2. ScALPELLUM SEXCORXUTUM, Pilsbry.
L' unique eeliantillon que j'ai sous les yeux a ete designe par Miers sous le nom de
Sc. verticilUitum, c'est du reste cclui quo porte encore I'etiquette du flacon qui le
renferme. Comme Miers n'a jamais donne ni diagnose, ni description daiicune sorte
pour cette espece, elle a ete de nouveau nommee par Pilsbry en 1897 *.
Je me bornerai, par consequent, tout simplement, a montrer les quelques points qui ne
me semblent par repondre exactement a la description ou a la figure de Pilsbry, mais qui,
a mon avis, ne justifient par la creation d'une espece nouvelle. II est bon, je crois,
cependant, d'indiquer ces particularites pour montrer les modifications de details qui
peuvent se rencontrer dans cette espece.
Forme generale du capitulura triangulaire, le cote rostral presque droit, le cote dorsal
convexe. 13 plaques parfaitement calcifiees. Ces plaqvies, surtout celles de la rangee
inferieure, sout separees par un cspace purement chitineux, parfois c>asid(''ral)le. La
cuticule qui les recouvre en grande partie est couverte de polls fins et courts surtout
dans les regions dorsale et ventrale.
Carene regulierement courbe, mais redressee, umbo tres rapproche de I'apex. Sa
surface dorsale est arrondie, sans trace d'aretes laterales et nettement carenee ; tres
etroite au sommet, elle s'elargit progressivement vers la base.
Tergum triangulaire avec les cotes a peu pres droits, le cute carcnal etant le plus long
et legerement convexe. Une legere saillie de la plaque va de I'apex, droit et pointu,
a Tangle basal. Stries d'accroissement assez nettes, en partie cacliees par la cuticule.
Scutum convexe, triangulaire, avec le bord occluseur legerement concave et le Ijord
tergal droit, mais le bord basal fortement convexe, en partie cache sans la cuticule.
Apex pointu et droit. Stries radiales non marquees, les stries d'accroissement seules
visibles. Une bordure finement chitineuse et couverte de polls courts, ayant environ
1 mm. dans sa largeur maximum va de I'apex du tergum un peu au-dessous du
milieu du bord occluseur du scutum.
Les plaques supra-lalcrales en forme de triangle curviligne. Umbo a I'apex, en pointe
mousse, legerement recourbee en avant. Bord tergal a peu pres droit, bord scutal forte-
ment concave egalant environ 2 fois la longueur du bord tergal, enfin, bord basal
fortement convexe et egalant a peu pres trois fois la longueur du bord tergal.
Rostre triangulaire, dout la largeur egale, au moins, la hauteur. Apex tourne en haut
* Pil^liiy, IE. A. "Description of a remarkable Japanese Cirriped (Scaljtdlmn scvcornutum)." American
Naturalist, xxxi. p. 723 (lS!t7).
SECOND SERIES. — ZOOLOGY, VOL. VIII. 43
29i A. GEUVEL — SUE QUELQUES LEPADIDES NOUVEAUX
et legerement saillant. Toute la partie inferieure de la plaque est recouverte par la
ciiticule.
Les plaques rostro-laternles, careno-laterales et subcarenales, au lieu de se recourber en
has corame dans la description de Pilsbry, ont leur apex dirige, au contraire, vers le haut
et eloignc du capitulum, ce qui les fait saillir considerablement en dehors. Toutes ces
])laques sent triangulaires, plus hautes que larges ct recouvertes en grande partie par la
cuticule.
Pas de plaque infra-medio-laterales ni de sous-carene.
Longueur du capitulum, IG mm. ; largeur, 9 mm.
Fedoncule. — Cylindro-conique, aussi large que le capitulum a sa partie superieure et
allant en diminuant progressivement. Quolques rangoes circulaires, a peu pres paralleles,
d'ecailles allongees en forme de poiiites et presque cacliees par la cuticule couverte de
poils tres courts et tres fins.
Longueur du pedoncule, 10 mm. ; largeur moyenne, G mm.
Habited. — Une seul exemplaire recueilli par 22 brasses de foad et par 31° 31' lat. N.
et 133 44' long. E. Inland Sea. Presente par le Dr. J. G. Jeffreys.
Observations et Affinites. — Cettc espce me semble aussi rapprochee de Sc. Peroni,
Gray, et de Sc. Scorpio, Auriv., que de Sc. villosum, Leach, ou Sc. trispinosmn, Hoek, car si
la carene est presque droite, comme dans ces dernieres especes, il n"en est pas moins
vrai qu'elle est recourbee en avant et qu'il existe une surface antero-superieure, en partie
cachee par la cuticule, mais qui fait que I'umbo au lieu d'etre exactement a I'apex se
trouve un peu en arriere de lui *.
EXPLICATION DE LA PLANCHE 24.
Fig. 1. Reproduction pliotographique de :
A. Alepas Belli, n. sp.
B et B'. Alepas microstoma, n. sj>.
C. Alepas indica, u. sp.
2. Alepas Belli, dessine d'apr^s nature.
3. Le meme, orifice externe du capitulum, vu de face.
4. Alepas indica, d'aprfes nature.
4'. Le meme. Orifice externe du capitulum, vu de face.
5. Poecilasma minuta, n. sp., d'apres nature.
6. Scalpellnm Hoeki, n. sp.
7. Alepas microstoma jeune.
8. Le meme. Orifice externe du capitulum, vu de face.
Meme remarque que dans la note (*), p. 289.
DE LA COLLECTION DU BRITISH MU8EU.AI. 295
Fig. 9. Scalpelhim Hoeli. Careuc et plaques careno-laterales, vues ])ar la face dorsalc.
Rostre, vu de face.
Ecailles pwloiiculaires.
Mamelon buccal, vu par la partie supi'i-ieure.
Palpe de la levre superieure.
Machoire gauche.
Maudibule droite.
Palpe de la levre inferieure droit.
Ornements de la cuticule.
18. ,, „ „
19
'.li). Sra/pe/lum Hueki. Ap^jeiidices caudaux (caudal ajjpciiadges).
21. Alepas indica. Dents chitineuses du bord superieur du labrc.
'l:i a et b. Ornements des parties laterales du labrc ; c, ornements de la region moyeiine ; d, orue-
ineuts de la cuticule eapitulairc, avec les tubercules et la soie sensitive.
23. Le meme. Palpe de la levre superieure.
24. „ Mandibule gauche.
25. ,, Machoire droite.
20. „ Palpe de la levre inferieure, droite.
27. „ Portion de ramc dc la deuxieme paire de cirrhes.
28. Alepas Belli. Uue soie antericure de la V paire de cirrhes.
29. „ Penis.
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Part VIII. 1892 0 8 0 . . . .
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Part XI. 1894 0 2 6 . . . .
VI. Part I. 1894 2 0 0 . . . .
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VIII. Part I. 1900 0 10 0 . . . .
Part II. 1900 0 10 0 . . . .
Part III. 1900 0 10 0
Part IV. 1901 0 14 0
Part V. 1901 0 5 0
Part VI. 1901 0 10 0
Part VII. 1901 1 8 0 . . . .
Part VIII. 1902 0 4 0 . . . .
Price to
Fellows.
£ s. d.
10 0
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110
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13 3
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1f03
2nd Ser. ZOOLOGY.]
[VOL. VIII. PAET 9.
U\^
THE
TRANSACTIONS
OF
THE LINNEAN SOCIETY OF LONDON.
OK THE ACTIKIAN BITNODEOFSIS GLOBULIFEHA, veeeill.
BY
J. E. DUEEDEN, Ph.D., A.ll.C.Sc.(Lon(l.),
Bkuce Fellow, Juuns Huekins U>'iveksitv.
(Communicated by Prof. G. B. Howes, F.R.S., Sec. Liuu. Soc.)
LONDON:
PRINTED FOR TUE LIXNKAN s;OCibl\
1)Y TAYI.OK ANU KKANCIS, KKl) LION COUKT, KLKKT .SIIIEET.
SOLU AT TEK SOCIKTY's APARTMENTS, BlKLINGTON-llOLSE. I'lCCADlLLV, AV.,
AND BY LONGMANS, GREEN, AND CO., PATKRNOSTtU-ROW.
October 1902.
AUV' '903
r 00*7
[ 297 1
IX. On the Actinkm Buuocleopsis globulifeva, Ven'ill. By J. E. Duerden', Pli.D.,
A.It.C.Sc.{Lond.)., Bruce Fellow, Johns Hopkins TJaiversity. {^Communicated by
Prof. G. B. HoAVES, F.B.S., Sec. Linn. Soc.)
(Plates 25 & 26.)
Read L'Oth rebruary, 1902.
In a preliminary paper on the Jamaica Actiniaria (1898) I briefly descril^ecl a new West
Indian Bunodeopsis, without assigning it any specific name. The year previous I had
given a fuller account of two other species of the same genus, and it was intended to
institute a comparison with these. In 1899, Prof. A. E. Verrill figured, without any
comment in the text, a Bermudas Anemone which he identified as the Viatrix (jlohuli-
fera of Duchassaing and Michelotti (1860). Upon the appearance of the paper I drew
Prof. Verrill's attention to the similarity between his drawing and the new Bunodeopsis
which I had obtained, and the year following appeared his description of the form as a
new species, Bunodeojisis globulifera. The description is limited, however, to an
account of the excernal characters, and gives no attention to the peculiarities of
anatomy and minute sti'ucture.
A study of the internal structure and histology of the two species of Bunodeopsis in
1897 had revealed the presence of a well-developed ectodermal muscle and nerve-layer
on the column- wall, along with several other exceptional features, Avhose significance
was not then realized. About this time Dr. Oskar Carlgren (1893, 1900) was directino-
attention to the importance in Actiuian phylogeny of the occurrence of an ectodermal
columnar musculature, and its usual association with the absence of gonidial grooves in
the stomodseum, absence of ciliated bands from the mesenterial filaments, the non-
development of the basilar muscles, and the occurrence generally of a weak internal
musculature. Every species of Bunodeopsis thus becomes of special interest in
connection with this latest jihase of the morphological study of the Actinians.
The genus Bunodeopsis has been thus defined by me (1897, p. 6) : — " Tissues very
delicate. Tentacles elongate, readily retractile. Column short, beset proximally with
pedunculate or sessile vesicles, the vesicular area much broader than the capitulum.
Sphincter muscle feebly developed. More than six pairs of perfect mesenteries."
The genus at present is included within the family Aliciidae, which in the same paper
(p. 2) is characterized as follows : — " Ilexactineoe with a large, flat, contractile base.
Tentacles simple, subulate, and entacmseous. Column with simple or compound
outgrowths or vesicles over more or less of its surface, arranged mostly in vertical rows.
No cinclides. Sphincter muscle endodermal and diffuse, variable in amount of
development. Perfect mesenteries few or numerous. No acontia." In addition to
SECOND SERIES. — ZOOLOGY, VOL. VIII. 4-J,
298 DE. J. E. DUEEDEN ON THE
Biinodeoims, the family contains tlie genera Alicia, Cystiactis, and Thaumactis, and in
1898 Carlgren included the genus Phymactis of Milne-Edwards and Haime.
As a result of the discovery of the ectodermal columnar musculature in Bunodeopsis,
and also in Thanmactls, Carlgren (1898, p. 17 ; 1900, p. 28) considers that the family Aliciidae
as at present constituted is made up of heterogeneous members, and indicates that it will
be necessary to remove the two genera mentioned and associate them with the Tribe
Protanthese. He further states that he is preparing a revision of the family. With the
many representatives at his command Carlgren is in a much better position for carrying
out such a revision than any other student of the Actinise, and it is desirable that
a fuller account of Verrili's species of Bunodeopsis should be available.
Bunodeopsis globulifera, Verrill.
Bunodeopsis, u. sp., Duerden, 1898, p. 456.
Viatrix globulifera, Verrill, 1899, p. 146, fig. 20.
Bunodeopsis globulifera, Verrill, 1900, p. 559, pi. 67. fig. 4.
The base is loosely adherent or free, broad, flattened or eucircliug marine plants,
approximately circular, and thin- walled. In pi-eserved specimens it is usually irregular
in form, the middle deeply concave, and the periphery irregularly sinuate ; concentric
and radiating grooves are also present.
The column is erect, thin-walled, partly transparent, contractile, broad below and
gradually narrowing above, passing directly into the tentacles. The limbus is circular
in living, but strongly crenulated and lobed in preserved sj)ecimens. The lower region
of the column, for about two-thirds of the total height, is nearly covered with small
sj)heroidal vesicles, the upper one-third is naked and smooth. The naked region is
much more limited in extent in preserved polyps than in living specimens, and Verrili's
drawings and description would seem to have been founded upon the former. The
apex of the column is devoid of acrorhagi. It is very rarely infolded, but where
this takes place the disc, tentacles, and naked part of the column become hidden,
and only the region bearing the vesicles remains exposed.
The disposition of the vesicles sometimes appears irregular, though as a ride they are
seen to be arranged in vertical rows of different heights, but no regularity in longer and
shorter cycles has been established. Towards the base two or three rows of minute
tubercles may alternate with a row of large outgrowths. In both his figures Verrill
represents the vesicles as irregularly distributed, with scarcely any variation in size,
though in the text he notes that larger and smaller ones are mingled together. The
species undoubtedly varies greatly in its outward appearance.
The evaginatious are simple, sessile or sub-pedunculate, hollow, smooth throughout,
opaque, and thick-walled in comparison with the column-wall generally. They are
variable in size, some appearing as mere rounded tubercles, while others are seen as
distinct vesicular outgrowths ; most are spheroidal, but the larger examples may be
reniform. Dilierent sizes are intermingled, but in general the organs increase in size
from below upwards, at the same time diminishing in numbers.
ACTINIAN BUNODEOPSIS GLOBULIFEEA. 299
The tentacles are margiuate, strongly entacmteous, and very variable in number and
size. Any number from twelve to forty-eight may be present, according to the size of the
polyp. Verrill records examples with 18, 20, 22, 24, and 26 tentacles. Three or four
cycles are represented, but no regular an'angemcut, hexameral or otherwise, has been
made out. The inner members are mostly very long and slender, tapering towards the
extremity, and are smooth, very delicate, and transparent, and sometimes appear
minutely spotted. Very often the tentacles are imperfectly developed at some particular
region of the periphery, several appearing much shorter than the others, or represented
only by mere papillae. As a rule one such imperfect area will occur, but there may be
two, separated by one or more fully developed tentacles (PI. 25. fig. 1 b).
The tentacles are generally overhanging in living polyps, and in preserved specimens
they remain nearly erect and tapering, and a little shortened. On one or two occasions
they have been found completely infolded, with nothing but the vesicular part of the
column showing. Though such a strongly infolded condition is rather exceptional in
the present species it is very frequent in B. autilliensis, Duerd., and is somewhat
remarkable considering the practical absence of a sphincter muscle on the column
(p. 302). So complete is the infolding that the tentacles may even pass into the
mesenterial chambers, as shown in fig. 11. A basal constriction often occurs at the origin
of the tentacle from the disc (fig. Ic), and occasionally a tentacle has become detached,
its former position at the margin of the disc being indicated by a circular aperture.
The tentacles are thickest just above the basal constriction.
The disc is smooth, circular, flat or slightly convex, and transparent the mesenteries
and stomodteum showing through iu the living condition. The central part as a rule is
elevated in preserved polyps. The mouth is slit-like in living polyps, but in preserved
specimens it is widely open and circular, and tlie lips are thickened and j)rotruduig.
The stomodseal walls are smooth, and no gonidial grooves are distinguishable.
The polypal wall thi'oughout is nearly transparent and only slightly coloured.
During partial contraction the lower part of the column is sometimes a bright, opaque,
yellowish green, with brown or black lines or spots. On fu^ll distention, the vesicles and
column are a clear, delicate, yello^vish brown, due to the yellow cells within the
endoderm. The tips of the larger vesicles may be a little darker than the rest of
the wall, probably owing to the greater concentration of the stinging-cells (fig. 5).
The diameter of the base of the largest specimens is 12 cm. ; the height varies from
0'5 cm. to 1'3 cm. The length of the inner tentacles is about 1 cm., or nearly the same
as the height of the column.
The polyps are essentially shallow-water forms, being usually met w'ith around
Jamaica at a depth of three or four feet, with their base encircling the long, narrow,
cylindrical stems of the marine weed Ruppia, or more rarely adherent to the flattened
fronds of Dictyota and leaves of Thcdassia. It occurs in some abundance at Port
Antonio, and I have also found it aroimd Kingston Harbour. In this latter area the
closely allied B. antilliensis is very abundant, but I have not met with it at the former
locality.
44*
300 DK- J. E. DUERDEJN" OX THE
The polyps are easily detached from the marine plants to which they are adherent ;
indeed, in transferring the leaves from the sea, or from one jar to another, the polyps
wUl often slide off. It is douhtful if an actnal adherence really takes place, any more
than in the case of such organisms as Nudihranchs and Turhellarians. The presence of
a basal ciliation, to be described below, would iudicate that the contact with foreign
bodies is not very close. The extremely weak parieto-basilar muscles and absence
of basilar muscles are no doubt the structural features correlated with the incapacity for
firm fixation. In the laboratory specimens often free themselves, and, suspended in the
water with the tentacles fully expanded, may move slowly around, gradually coming
to rest asrainst the sides of the vessel. Probably their movements are assisted by the
strong ciliation on the base and column-wall. I have never observed any decided
rhythmic movements in swimm.ing. When free the base may be upwards, with the
tentacles hanging gracefully downwards ; or it may be in its normal lower position,
the tentacles then directed upwards and partly overhanging.
Carlgren (1900) also mentions that the BoIocerohJes IIcMurrichi, obtained by
Dr. Stuhlmann from Zanzibar, was likewise found to be able to detach itself and carry
on swimming movements much like a Medusa. Like Buiiodeojjsis, the genus .Bolo-
cerokles is considered to belong to the primitive Protantliese, as is also Gonactinia,
another form capable of swimming. Probably such loosely adherent Actinians should
be regarded as in some ways intermediate between the free Anemones with a rounded
aboral extremity and foi'ms with a flattened firmly adherent base.
Anatomy axp Histology.
Base.
The base is thin-walled, all the three constituent layers being very narrow (PI. 25. fig. 2).
In sections it is usually -03 mm. across. The ectoderm is remarkable for the short,
coarse cilia with which it is covered, and which are recognizable in basal sections even
when unpreserved in other parts of the polyp. In most preserved polyps only traces of
the external ciliation on other parts of the body-wall persist, while the ectodermal
stomodseal ciliation is usually very obvious, and less so that of the mesenterial filaments
and of the endoderm of the coelomic cavity. Probably the basal ciliation assists in the
movements of the polyps over the leaves of the marine plants on which they settle ;
or it may help the polyp when it assumes the floating habit. Its presence is in-
compatible with any firm adherence of the polyps by their base.
The basal ectoderm is constituted mainly of supporting cells, narrow granular gland-
cells, and a few clear gland-cells. A small nematocyst, measuring -012 mm., occurs
very sparingly. A rudimentary musculature and nerve-layer probably occur, but their
presence could not be determined with absolute certainty.
The mesogiffia is narrow, and contains a few minute connective-tissue ceUs. Elsewhere
throughout the polyp the mesogloea preserves the same character, that of a narrow,
homogeneous layer, containing but a few connective-tissue cells.
The basal endoderm varies much in thickness in different places. Where very narrow.
ACTIMAN BUXODEOPSIS GLOBULIFEEA. 301
the cell- contents stain deeply, being highly protoplasmic, and Zooxanthellae are absent
or nearly so ; where the layer is broader the cells are more vacuolated, the free surface
is somewhat irregular, and the symbiotic alg;3e are found in large numl)ers. Throughout
the endoderm of the polyp similar broad and narroAV areas are represented, according as
Zooxanthella? occur inal)undance or ai*e practically absent. A weak circular endodermal
musculature can be detected on the base, and in some specimens a nerve-layer separated
from the muscle-band by a fibrillar layer. Tiie latter, however, was not recognizable
in the section from which fig. 2 was taken.
Column-wall.
The proximal and distal regions of the column-wall vary somewhat in structure. The
former is broad and irregular in outline, owing to the occurrence of the evaginatious.
while the naked area is verv thin and delicate, measurini? in sections onlv '03 mm.
ft ' o »
across (fig. 4).
The ectoderm of the column-wall is weakly ciliated throughout, and both transverse
and vertical sections of the upper part otten exhibit fine ridges and grooves as a result
of uneqvial shrinkage (fig. 3). These appearances, however, are more characteristic of
the tentaciilar ectoderm, as shown in the same figure. The constituents are mainly
supporting cells and clear gland-cells ; nematocysts occur very sparingly, except in the
evaginatious of which they are very essential elements. A layer of delicate ectodermal
muscle-fibres occurs throughout the extent of the column, and in some places the meso-
gloea IS very slightly plaited for its support. The muscle-fibres are arranged with the
long axis vertically, so that they are cut cross-ways in transverse sections of the polyp,
and constitute a delicate but clearly-marked layer, the fibrils being highly refractive.
Outside the band of muscle-fibres occurs a weak nerve-layer {iir.l), the two distinctly
separated by a narrow interval crossed by fine fibrillar strands (figs, 3. 5, 16).
The natu.re of the fine strands occurring between the band of muscle-fibres and the
nerve-layer, and continued in some cases beyond the latter, is not readily determined.
They occur practically throughout the polyp, on both the ectodermal and endodermal
surfaces, but are much more definite in some individvials than in others. McMurrich
(1890. p. 113) refers to a somewhat similar appearance in various parts of the body- wall
of Cerianthus americanus, and regards the processes as mesogloeal prolongations.
Carlgren (1803, p. 26) describes a nearly similar structure in ProtanUtea simplex, and
also considers it to be due to very fine continuations of the mesogloea, which here and
there bear strongly refractive bodies, resembling sections of muscle-fibrils. In Carlgren's
preparations the details A-ary greatly according to the manner of preservation of the
specimens, and McMurrich' s two specimens of Cerianthus likewise diflfered.
The specimens now under consideration were preserved in formol and then transferred
to alcohol, and the interval between the mesogloea and the ganglion laj'er varies greatly
in width in different polyps, and even in different regions of the same individual.
Usually it is more pronounced on the endodermal side than on the ectodermal, and on
302 DR. J- E, DUERDEN ON THE
the former the strands sometimes appear slightly reticular in character. Small nuclei
occvir among the fibrils, and are regarded as the nuclei of ganglion-cells.
The same fibrillar layer is still better developed in some polyps of B. antilliensls. It
may occur on both the ectodermal and endodermal faces of the mesoglcea, and what
seem to be the continuations among the ectodermal cells are clearly seen (1891, pi. i.
figs. 3, 4). In Protcmthea, Carlgren (1893, pi. iii. figs. 4, 5) represents the mesogloeal
continuations, both on the ectodermal and endodermal aspects, as constituting a nearly
compact zone, while in the preparations of Bunodeopsis they appear as very djstinct
coarser strands, rather widely separated one from another.
The nerve-layer in Bunodeopsis is, as a rule, very narrow in sections, and appears
made up of interlacing fibrils, some seen longitudinally and others in section. In Carl-
gren's figures of Protanthea the layer is represented by minute dots only, indicating the
cut ends of the fibrils. Such an appearance also occurs in the stomodseal ectoderm of
Bunodeopsis (fig. 7, nr.l.).
The mesogloea of the column-wall is thin throughout, and both its inner and outer
surfaces are smooth, or very slightly plaited to afford additional support for the ecto-
dermal and endodermal muscu.latures.
The endoderm is broad in the lower region of the polyp, where, as a rule, the cells
contain many Zooxanthellse ; but the layer becomes very narrow above, as the symbiotic
algse are either absent or very rare, and the cells are then more charged with proto-
plasmic contents. The endodermal free surface is everywhere feebly ciliated.
The circular endodermal muscle occurs throughout the column-wall, usually supported
on very delicate mesogloeal plaitings. A slight increased development takes place
towards the upper extremity, but never to such a degree as to give rise to w^hat may be
regarded as a sphincter muscle (fig. 3, end.m.). A nerve-layer is usually seen, situated at
some distance from the mesoglcBa, and a little better developed than in the ectoderm.
The evaginations (figs. 4 & 5, ev.) are simple, hollow, rounded outgrowths of the lower
part of the column-wall, and in transverse sections are seen to be both entocoelic and
exocoelic in position, though without any regular alternation. In sections the walls at
their thickest are about "065 mm. across. The cavity communicates by a large aperture
with that of the ccelenteron. Both the ectoderm and endoderm have undergone certain
histological modifications compared with the character of the layers in the column-wall
generally. The ectoderm (fig. 5) is broad, somewhat strongly ciliated, and a very
distinct zone of oval nuclei occurs just within the outer margin, w^hile smaller, rounded
nuclei occur here and there throughout the thickness of the layer. The characteristic
elements, how^ever, are the large, thick-walled nematocysts (neni.), which occur in some
abundance in the outer area of the evaginations. Seen in sections, the nematocysts
exhibit few or no structural details, but appear as smooth-walled, nearly homogeneous
bodies, irregularly distributed among the supporting cells. The deeper examples more
especially are arranged at varying angles witli the surface of the ectoderm, but the outer
are practically parallel with the supporting cells. The former are probably individuals
in process of development and are strongly curved, while when they assume their
ACTINIAN BUNODEOPSIS GLOBULIFERA. 303
mature position at right angles with the surface, they are nearly straight. In macera-
tions the nematocysts appear as represented in fig. 8 b. They possess a thickened Avail,
and exhibit a tubu.lar appearance within, representing the tliickened basal part of the
thread, but no trace of the finer portion of the thread can, as a rule, be made out.
The musculature and nerve-layer (fig. 5, nr.l.) on both sides of the mesogloea are
extremely delicate in the evaginations, but can be clearly recognized in favourable
sections. The endoderni is broader than that of the column-wall generally, and is more
strongly ciliated. Its cells are usually crowded with Zooxanthellce {zoox.), though in some
cases the alga? are very sparse or altogether wanting. The histological character, as a
whole, would indicate that the columnar evaginations are to be regarded as special
nematocyst-beariug organs.
The size, character, and distribution of the nematocysts possess a certain importance
in systematic studies of the Anthozoa, and may be fittingly referred to here as repre-
sented throughout the different tissues of the present species.
Carlgren (l!)0(), p. 3) believes that the nematocysts may be made of great assistance in
the identification of species of Actinia^ and even in the recognition of genera. For the
time being he roughly divides them into " thin-walled nematocysts," in which the internal
spiral thi'ead shows distinctly, and " thick-walled nematocysts," which appear quite
smooth. Very often the internal spu-al thread is not seen within the latter, while
in others it may be recognized, but not so clearly as in the thin-walled form. With
regard to their general distribution in the polyp, Carlgren finds the thin-walled variety
to occur mostly in the tentacles and disc, but in certain sj^jccies and genera in the column-
wall and stomodajum also. The thick-wtilled forms are found in all parts of the ectoderm
of the column-wall, and their length is fairly constant for the species. As a rule the
longest thick-walled nematocysts occur in the stomodaeum ; those in the tentacles are as
long or somewhat shorter ; while they are the shortest in the disc, column-wall, and base.
The longest nematocysts usually occur in the acrorhagi, or in any other special out-
growths of the column-wall which may be develoi)ed.
Usually it is only in the living polyps that the stinging-cells can be examined in
their fully extruded state and their complete characteristics determined, including the
swollen basal part and the simple or complex evaginated thread. But certain distinc-
tions can be made out in macerations of preserved polyps, or even in sections. In the
species now under investigation the occurrence and character of the stinging-cells
are briefly as follows : — They are very sparingly I'epresented in the basal ectoderm by an
extremely small form of the thick-walled variety, measuring only '0] 2 mm. In the
column-wall they have been found only within the ectoderm of the evaginations. Here
they are all of one kind : large, thick-walled, slightly curved, and varying in length from
•038 mm. to '044 mm., with a diameter of about -008 mm.
The tentacular nematocysts include the two varieties. By far the majority are thin-
walled with the spiral thread showing very distinctly, and are broad at one end and
narrow at the other (fig. 8«). They range in length from -018 mm. to "028 mm. The
thick-walled form is numerous, and nearly as long as in the columnar ev'aginations, theu"
301- KR. J. E. DUEEDEN ON THE
length varying from -034 nun. to -Oi mm. They are invariably a little narrower than in
the evag'inations, measuring only -006 mm., and are nearly always perfectly straight.
In the disc stinging-cells are concentrated within the thickened lips, and are similar to
the thin-walled representatives in the tentacles.
The nematocysts within the stomodseal ectoderm include only one representative of the
curved columnar variety. They are very uniform in size, measuring "025 mm.
The mesenterial filaments are somewhat exceptional in being very sparingly supplied
with nematocysts, even in the lower region. They are all narrow and straight, and are
nearly of the same length as those in the stomodfeum, namely 025 mm.
jS^o uematoblasts occur anywhere within the endodermal epithelium.
It is thus clear that practically every region of the polyp possesses its own distinctive
nematocyst. Though generally distributed throughout, the thick-walled variety differs
slightly either in size or outline, or in botli, ia the different areas where it occurs.
Examples intermediate in type are to be found mingled with the others, so that it is
only when taken in nvimbers that the distinctive characteristics become of taxonomic
value.
Tentacles.
As the column-wall is rarely overfolded in preserved polyps, and the tentacles are
usually only slightly retracted, the latter are generally seen in their normal relationship
in both vertical and transverse sections (PI. 25. fig. -I & PI. 26. fig. 13). A transverse
section, such as is represented in fig. 13, demonstrates that the tentacles are outgrowths
of both the entoccelic and exocoelic mesenterial chamhers. In some instances, as at
ti and ^5, t., and t^, t^.j and ty,^, it appears as if two tentacles originated side by side from
a single mesenterial chamber, but on more close examination a pair of rudimentary
mesenteries (III.) can always be detected, so that one of the tentacles is eutocoelic and
the other exoccjelic. Such conditions prove that the appearance of a new mesenterial
pair is followed very closely by the outgrow^th of a tentacle from its entocoele. In the
polyp from which fig. 13 was taken no exotentacles had as yet appeared between the
entocoelic members ^5 and i,-, t^ and t-, ^n and iyj. For a short period, therefore, during
the s^rowth of new mesenteries, the number of tentacles mav be less than the number of
O t'
mesenteries, due to the lagging behind of the exotentacles ; but in the end the sum is
the same, a tentacle arising from each chamber, whether entocoelic or exoccelic.
The tentacular ectoderm and endoderm are both comparatively broad layers, but the
mesogloea is very narrow. In sections the wall is about '055 mm. across. The ectoderm
is usually thrown into minute rounded ridges separated hy deep narrow grooves, the
cells being elongated in the former and very short between (fig. 3). Nematocysts are
distributed in a peripheral zone with approximate uniformity throughout the length of
the tentacle, and are mainly of one kind : thin-walled, with the close spiral thread
showing distinctly, and broad at one end and narrow at the other (fig. 8 a). Numerous
examples of the larger, thick-walled variety, such as occurs in the evaginations, are also
present, but are slightly shorter and much narrower.
ACTINIAN BUNODEOPSIS GLOBULIFERA. 305
The longitudinal ectodermal muscle is supported upon the mesogla3a, which is usually-
thrown into slight folds to give it increased area, especially towards the proximal end.
The nerve-ganglion layer is also clearly developed.
The endodermal epithelium is rather broad in the partly retracted tentacles, and
exhibits an irregular free surface, while Zooxanthelhc are only sparsely distributed.
The endodermal muscle (end.m., fig. 3) is comparatively well developed, the mesogkea
presenting delicate plaits for its support, best seen just above the tentacular
septum (i.c.s.).
The tentacular cavity is separated from the general polypal cavity by a circular
transverse partition, perforated iu the middle, and situated a very short distance from the
origin of the tentacles (tigs. 3, 4, (5). The septum is covered on both sides by endoderm,
and the supporting mesogloea is narrow and lined by delicate muscle-libres on its
inner and outer surfaces, continuous with the circular musculature of the tentacles. Its
central free margin is thickened, and the mesoglcea becomes plaited {t.sph., tigs. 3 &6) in
a sub-dendroid manner to afford additional support for the muscle-tibres, which here
constitute a circular sphincter muscle. Various appearances of the partition in section,
showing its relation to the coeloraic and tentacular cavities, are indicated in figs. 4 and 13,
and the sphincter muscle {t.sph.), more highly magnified, iu PL 25. figs. 3aud 0. Where
in fig. 13 the section includes the middle of the partition the two free edges are seen to be
in the main directed downwards into the polypal cavity, so that the membrane must be
concave on its tentacular aspect and convex on its ccelomic side. In fig 13, tentacles
^1, ('2, and ^10 are partly constricted at their origin, and such a condition has been noted
among the external characters (fig. 1 c). No doubt the constriction is brought about by
the action of the sphincter. In fig. 3 the tentacular wall immediately below the
partition is seen to have become much thinned.
The presence of a tentaculo-coelomic septum and sphincter has already been recognized
in the genus Bolocera, including also the genus Boloceroides of Carlgren (1899, p. 43;
1900, p. 15). Carlgren (1893) descri!)es and figures the formation at the base of the
tentacles in Bolocera loiigiconiis, and the musculature is there not much better develojjed
than in Buiiodcopsis. In Bolocera the contraction of the sphincter often leads to the
detachment of the tentacles from the disc. In one or two of the Jamaica specimens of
Btmodeopsis glohuUfera circular apertures occur in the tentacular zone, indicating that
tentacles have disappeared therefrom, but I have never actually observed the j)rocess
taking place in the living polyp. In B. antllUensis, however, the tentacles were found
to be readily detached on handling the polyps, and in this species also a tentacular
septum and sphincter occur. Carlgren has pointed out that the partition comes off along
with the tentacles, and such would certainly be presumed from the great narrowing
which takes place immediately below the membrane, as represented in fig. 3.
McMurrich (1893, p. 154, pi. xxii. fig. 27) also describes and figures the tentacular
sphincter iu Bolocera occidua. In many of his sj)ecimens, mesenterial filaments were
lound protruding from the openings at the margin of the disc formed by the falling off
of the tentacles. The same author includes the deciduous character of the tentacles
SECOND SERIES. — ZOOLOGY, VOL. VIII. 45
306 DK. J. E. DUERDEN ON THE
within the definition of the family Boloceridse. Kwietniewski (1898, p. 394, pi. xxv.
fig. 10) has also described and figured the sphincter in Bolocera McMurricU { = Bolo-
ceroides McMurriclii (Kwietn.), Carlgren, 1899).
The occurrence of a perforated tentacnlo-coeloniic septum and sphincter would appear
to be somewhat restricted within the Actiniaria. So far, it is known only in the genera
Boloceroidcs, Bolocera, and Bunodeopsis. Its presence would seem to be associated
with deciduous tentacles, for the phenomenon is characteristic of all the forms above
mentioned.
According to Carlgren (1899), th.e genera Pohjstomidium and Polijopis, described by
R. Hertwig in the report on the ' Challenger ' Actiniaria as destitute of tentacles, are
really Boloccrce from which these organs have become detached. Haddon (1898) had
independently come to the same conclusion. Carlgren suggests that the detachment of
the tentacles may be a means of protection to the polyp.
McMurrich (1893) had already surmised, and later was able to confirm his suspicions
by an actual examination of the specimens at the British Museum, that the ' Challenger '
Actinian, lAponema multifommm, is also a Bolocera, from which the tentacles have been
detached.
Disc.
The oral disc {disc, PL 25. figs. 4 & 6) is thin-walled periphei-ally, and continues tlius
until the lips are reached, when it becomes swollen before terminating at the stomodteum
{st.). The discal ectoderm contains many clear gland-cells, and the radial muscle-
and nerve-layers are comparatively well developed. At the lips the ectodermal cells are
much longer, and thin-walled neraatocy.sts occur, similar to those in the tentacular
ectoderm, but stinging-cells appear to be absent from the remainder of the disc.
The discal mesogloea (fig. 6) is very narrow, and may be delicately plaited on both its
outer and inner siirfaces. At the lips it becomes much thickened and more strongly plaited,
aftording an increased area for the supjiort of the circular endodermal muscle. This
latter becomes so strongly developed as to constitute what must be regarded as a special
endodermal circumoral sphincter [or. sph., figs. 4 & 6). The action of the sphincter
muscle must have become relaxed during the narcotization of the polyps, permitting
the radial musculature to act, for in all the preserved specimens the oral aperture is
large and rounded.
A sphincter muscle in this position is very exceptional among the Actinite, but a like
concentration of muscle-fibres has been described by the brothers Dixon for B. rerrucosa
(1889, p. 322). It may be considered doul)tt'ul as to how far a special muscular develop-
ment would be represented when the oral aperture returns to its living, slit-like condition.
Other than in its concentration of muscle-fibres the discal endoderm presents no special
features. Comparatively few Zooxanthellte are present.
Stomodceum.
The stomodaeum is widely open in sections and usually somewhat irregular in outline
(figs. 9-15). Its walls are very delicate, measuring only -045 mm. across. It is
ACTINIAN BDNODEOPSIS GLOBULIFERA. 307
comjiaratively long, extending into the gastro-coelomic cavity for nearly one-half the
height of the polyp, its inner free mjirgin heing reflected (PL 25. fig. l). The ectoderm
is often thrown into vertical folds, which are without any regular disposition, not
necessarily corresponding with the mesenteries. The layer exhibits the same structure
all the way round, being in no way modified to form special gonidial grooves opposite
the directives. Prof. Verrill, in his description of the external characters of the Bermudas
specimens, mentions the occurreuce of gouidial grooves, but such are certainly not
indicated in the histology of the stomodseiim of the Jamaica representatives, and were
not recognizable in the living polyps. h\ other species of Actiniae where gonidial grooves
occur, the ectoderm lining them usually offers a n)arked contrast with that along the
lateral walls ; the cilia are more strongly developed, and fewer gland-cells and neuiato-
cysts occur than elsewhere.
The stomodseal ectoderm {ecL, PL 25. tig. 7) is strongly and uniformly ciliated all
rorxnd, and contains nematocysts [item.) and granular gland-cells. The former are sparse
towards the upjjer extremity, but as the lower free end is approached they become very
numerous, and are all of the thick-walled variety, such as occur in the columnar
evaginatious, but slightly smaller, measuring -0:^5 mm. in length. In the inner part of
the layer they are irregularly arranged at all angles with the free edge, but towards
the surface they become parallel with the other constituents. The ectodermal muscle
(ect.tn.) and ganglion-layer {/tr.l.} are only feebly developed.
The mesogloea is extremely delicate throughout, and remains smooth on both sides.
The endoderm closely resembles that lining the coelomic cavity generally, and contains a
few Zooxanthellge in some places, while in others they are absent, in which latter case the
epithelium is very narrow. The circular endodermal musculature is recognizable in
vertical sections.
Intermesenterially the stomodieum narrows gradually towards its free lower extremity
and becomes reflected, as on the left side of tig. i ; mesenterially the ectoderm appears
continuous with the iilament at the free edge of the perfect mesenteries. The stomodieal
waU on the right side of fig. J? terminates in a section of a mesenterial filament.
3Iesentei'ies.
The mesenteries are very varial)le in number in the many polyps examined, and
also in their cyclic relationships ; for a long time it seemed impossible to establish
any hexameral or other regular system of arrangement. The conditions encountered
will be best understood by reference to PL 26. figs. 9 to 15, representing outline
sections through the stomodaeal region of six different polyps.
In fig. 9 only three pairs of complete mesenteries occur, and two pairs which are
incomplete. The lower comjilete pair (d.) are directive mesenteries, having the
longitudinal muscle on the faces looking away from one another. The polyp from
which fig. 10 was taken possessed three pairs of complete mesenteries, including a
pair of directives and a single member of another complete pair. Within the wide
interval on the right side are nine, short, incomplete mesenteries, which present some
45*
308 DK. J. E. DUERDEN ON THE
indication of alternating large and small pairs. In this and the other figures the pairs
are numbered according to the order to which they belong, so far as this can be
determined with certainty. In the polyp whose mesenterial plan is represented by
PI. 26. fig. 11 a regularity of alternating complete and incomplete pairs is presented on the
leftside, but the right side is a region of irregularities. A mesentery also stretches across
from one part of tlie column-wall to the other, and encloses a rudimentary mesenterial
pair. In fig. 12 the left side j)ossesses perfect dicyclic hexameral symmetry, while
the development at the ujDper right-hand region is very irregular. As before, many
incomplete mesenteries occur at the upper right-hand corner.
Fig. 14 is tbe completed mesenterial plan of the polyp of w^hicii fig. 13 represents
a little more than half. As before, the first and second cycles on the left side are
regularly develojied, and in addition three I'udimeutary pairs of the third cycle occur,
a pair in each of the primary systems. The new pair in the dorso-lateral chamber
is situated on the ventral aspect of the second cycle pair, but on the dorsal aspect in the
middle and veutro-lateral chambers. On the right side the ventro-lateral system
corres^TOnds with that on the left, but the middle and dorsal systems are irregular.
Fig. 15 completes the series so far as I have been able to establish it. On the
left side of the directive axis all the pairs of the first, second, and third orders of a
hexameral polyp are represented. The second-order mesenteries (II. j, with the exception
of the dorso-lateral pair, have reached the stomodteum. One mesentery only is missing
on the right side from the number necessary to complete the hexameral plan, but at the
dorso-lateral region they are all incomplete and irregularly paired.
From the conditions thus presented there can be no doubt that normally the
mesenteries should be arranged in alternating hexameral cycles, with two pairs of
directives, and that members of the second order should also reach the stomodseum.
The increase in the number of the mesenteries, how-ever, does not proceed throughout
the polyp according to the usual order established for the Hexactiniae. According
to this, the six primary pairs of mesenteries appear successively in a bilateral manner ;
then the members of the second cycle arise successively in unilateral pairs within the
six primary exocceles, as a rule on each side of the polyp i'rom the dorsal to the ventral
aspect ; the third cycle pairs appear as single unilateral pairs within one of the two
exocoeles in each of the six primary systems, and then later in the remaining exoccele
in each primary system. In fig. 14. the sequence of the thu-d cycle pairs seems to
follow the rule that a pair is developed in each of the primary systems before a second
pair arises to complete the radial symmetry.
All the figures of BuHodeopsis indicate that on one side of the poly^D the increase
takes place in a fairly normal and regular manner, but that such is not the case for the
other side. In all the polyps a definite region of irregular growth occurs, both as
to the number of mesenteries which are present and also as to their paused relationships.
Evidently the hexameral cyclic regularity becomes more pronounced as the polyps grovi^
larger and the number of mesenteries increases, but I have met with no specimen
in which perfect regularity was attained. Probably some of the polyps obtained are as
ACTINIAN BUNODEOPSIS GLOBULIFERA. 309
yet immature, but no larger were to be met with, and, judging from the number
of tentacles and the dimensions, Verrill's Bermudas examples were in much the same
condition as the Jamaican.
Undoubtedly the imperfect development of the tentacles within a restricted area,
noticed among the external characters, is determined hy the imperfect growth of the
mesenteries. The two are associated phenomena; the rate of tentacular growth is
regulated by that of the mesenteries, and where the latter are irregular the tentacles
will be the same.
This regional irregularity of groAvth of the polyp would appear to be connected in
some Avay with an asexual method of reproduction. In the more common B. antilUensis
I have found that new polyps are produced by the process which Andres (1881) has
described as " Scissiparity." Fragments of varying size become gradually detached from
around the base of the column, attain an independent existence, and develop into new
polyps with tentacles and oral disc. Andres has followed the process of development of
the new^ individuals as it occurs in Aiptasia lacerata, and in this species the normal
hexameral disposition of the organs appears to be early assumed. I have not observed
scissii^arity in B. globulifera, but the other Jamaican species presents exactly the same
tentacular and mesenterial irregularities.
Irregularities would be much more likely to occur in polyjjs thus originating than in
individuals developed directly from the larva. The two species of Bunodeopsis occur
in large numbers wdthin restricted localities, and such a distribution is mo.st probably
the result of a like method of reproduction in each. The West Indian Aiptasia tagetes
(Duch. & Mich.) also reproduces in a most in-olific manner by scissiparity, and an
examination of its mesenteries and tentacles reveals many departures from the hexameral
regularity. McMurrich (1889), on the other hand, describes the Bahamas representa-
tives of this species as hcxamerous, without mentioning any asexual reproduction.
G. H. Parker (1899), who has made a detailed study of the effects of longitudinal fission
in Iletridium marf/iiiafum, finds the mesenteries are very irregularly arranged in what
may be assumed to be fission polyps.
Returning to the individual mesenteries, the members of the first order are found
to extend the whole length of the stomodseum, and some are continued as far as the
base of the polyp, across which they extend to the middle (PI. 25. fig. 4i). The complete
members of the second order may become free before reaching the stomoda^al termination.
At first the free edge of the mesenteries is straight, but soon it becomes greatly folded
and contorted, some mesenteries beconiing much more complex than others, so that they
may appear several times in the same section (fig. 4). The incomplete mesenteries
first appear in the uppermost part of the column-w^all, but all except the youngest
are better developed in the middle part of the polyp. Very often the mesenterial
mesogloea and that of the column-wall are seen to be interrupted, the continuity of the
circiJar endodermal muscle of the colvimn-wall being thereby preserved (PI. 26, fio-. 17).
"Within the stomodoeal region of most polyps the mesenteries are extremely narrow,
and, except at their origin in the column- wall, the musculature is so weak as to
310 DR. J. E. DUEEDEN ON THE
be discerned with difficulty. Near its insertion in the column-wall the mesentery
is usually a little broader, and the retractor muscle is represented as a smooth layer of
fibrils. In other polyps the mesenteries are thicker, and muscle plaits are very distinct
in the middle of their transverse length. Belon^ the stomodseal region all the
mesenteries become much broader, both the endoderm and mesoglcea increasing in
thickness, and the latter forms numerous plaitings for the support of the retractor
muscles {ret.m., PI. 26. figs. 16 & 17). The retractor muscle in fact assumes somewhat of
a circumscribed character, tnit the outline in transverse section is scarcely alike in any
tAvo mesenteries. Por a short distance from the column-wall the muscle-band is smooth ;
the plaitings beyond are at first minute, then gradually become larger and more
complicated, and terminate very abruptly, the mesentery afterwards being very narrow
and continued for some distance (fig. 10). At the central termination of the complicated
part of the muscle one or more of the mesogloeal plaitings may become greatly elongated
and minutely plaited, almost as if the mesentery w^ere about to branch (fig. 17, ret.m.).
The transverse musculature is comparatively strongly developed. In tangential
sections near the base the mesogloea of the mesentery is irregularly folded for its support
(fi-g. 2, trans.m.). When a complete mesentery is mounted as a microscopic object, the
vertical and transverse muscle-tiljrils are seen crossing one another at right angles.
The mesenterial endoderm is usually very narrow, and includes clear gland-cells and but
few Zooxanthellse. Traces of the nerve-layer, continuous with that on the column-wall,
can usually be followed (nr.l., fig. 16) for some distance beyond the insertion of the
mesenterv in the column-wall. In the lower region the mesenterial epithelium becomes
swollen just behind the mesenterial filaments [cf. figs. 16 & 17), and in some polyps this
region contains many clear spheroidal bodies, w^hich stain very deeply and stand out as
distinct objects from the rest of the tissue ; a few small particles are black and
irregularly shaped. Probably these foreign bodies are ingested nutrient particles, and
they are practically absent from some individuals. Where present in quantity they may
occur also in the endodermal epithelium of the lower part of the stomodseum, and
sparsely in the endoderm of the column-wall, and even in the tentacular endodermal
epithelium. The particles are easily distinguished by their homogeneous, deeply staining
character from the Zooxanthellse, which they greatly resemble in size and outline.
The swollen mesenterial endoderm immediately behind the mesenterial filament is
generally regarded as a special region for intracellular digestion (McMurrich, 1899,
p. 262), and in Btowdeopsis the greatest number of foreign particles occurs there ; but
their presence in other regions of the endoderm. would imply that ingestion is by no means
restricted to the region. The endodermal enlargements in the present species are often
somewhat irregulai- iu outline (fig. 17), not smoothly rounded as in most other forms.
At the origin of the mesenteries in the column-wall a delicate layer of muscle-fibrils,
arranged nearly vertically, occurs on the face bearing the transverse musculature. This
is the only trace of a parieto-basilar muscle ; in no instance does the mesoglcea become
plaited or afford any indication of the formation of a pennon, such as is characteristic
of Actinian species with a well-developed muscle.
ACTINIAN BUNODEOPSIS GLOBULIFERA. 311
A vertical section through a portion of the base, inchiding a tangential section of a
mesentery, is represented in PL 25. iig. 2. On the mesentery the mesoglojal folds wliich
support the transverse muscle {traiis.m.) are strongly developed, and tlie muscle-fibres
are seen in section. At the basal termination of the mesentery, however, no foldings are
indicated ; the muscle-layer is here perfectly smooth. Sucli an appearance can scarcely
be regarded as in any way indicating an incipient basilar muscle, for no corresponding
development takes place on the opposite face wliich bears the retractor muscle. In such
sections, the muscle-fibres of the latter are seen lengthways. Where present in other
species the basilar muscle is, as a rule, arranged on more or less strongly developed
mesogloeal plaitings on each side of the insertion of the mesentery on the base, the
muscle-fibres extending in a radial direction. The present species is therefore character-
ized by possessing a parieto-basilar muscle of the weakest character, while the basilar
muscle is altogether undeveloped.
The polyps are too small to permit of the presence or absence of mesenterial stomata
being determined by means of dissection. But in serial transverse sections iuterruiDtious
occur in all the perfect mesenteries close to the stomodseum and at its upjier extremity.
These represent the inner or perioral stomata, but no corresponding series has l)een found
near the column-wall, such as may represent the outer parietal stomata.
Mesenterial Filaments.
Mesenterial filaments occur at the free edge of all the mesenteries except the most
rudimentary. They appear on the complete mesenteries as these sever their connection
with the stomodgeum, while usually they are not developed on the incomplete mesenteries
until below the stomodoeal region. The filaments follow the contortions of the free edge
of the mesenteries, so that in the lower part of the polypal cavity the filament belonging
to one of the mesenteries may be seen cut through several times in the same section
(fig. i).
The filaments on the complete mesenteries are at first trilobed and exceptionally wide,
and both the glandular streak and ciliated bands are clearly represented (fig. 18). The
ciliated bands are far apart, and the tissue intermediate between them and the glandular
streak is irregularly thickened, so that the outline assumed by the filaments is somewhat
exceptional among the Actiniae, and varies greatly in different sections. A little within
the base of the filament the mesenterial mesogloea divides into three very narrow
branches, each of "which passes into a lobe, and there terminates in a clear expansion,
practically devoid of any cell-enclosures.
The middle portion of the trefoil is low, and the glandular streak at its apex
{g.s., fig. 18) is at first quite rudimentary in the degree of its development, merely capping
the middle of the filament as a more densely staining tissue. In addition to the
supporting cells, the glandular streak contains a few granular gland-cells and thick-
walled nematocysts.
On each side of the glandular streak, and sharply separated from it, occurs an accumu-
lation of undifferentiated endodermal-like cells, rarely regu.lar in its outline, recalling in
312 DR. J. E. DUERDEN ON THE
this respect the irregular endodermal thickenings on the mesenteries immediately behind
the simple filaments. The cells are much vacuolated, and here and there Zooxanthellae
may occur. To this tissue {/.s., fig. 18), which is partly developed on both the middle
and latei'al lobes of the filament, and separates the glandular streak from the ciliated
bands, I have already (1900, p. 14G) given the name of intermediate streak. It is
recognizable in most trilobed Actinian filaments, and McMurrich (1899, p. 259), in
describing the filaments of Zoanthus sociatus, has also independently applied the desig-
nation intermediate epithelium to what appears to be the corresponding tissue in the
Zoanthean filament. In the pi'esent species it often appears as if folding over or partly
covering the first portion of the ciliated bands, and is more trvily endodermal in character
than in most Actiniae. In some longitudinal sections through the filaments it presents
a distinctly banded appearance, with the deeply staining tissue of the ciliated bands at
the bottom of the depressions, as is so well displayed in McMurrich's figures oi Z. sociatus
(1899, p. 259).
The ciliated bands are strongly developed and nearly encircle the lateral mesogloeal
lobes. They are constituted altogether of ciliated supporting cells. On the front face of
the lateral mesogloeal axes they are continued in many transverse sections as a very
narrow band towards the middle ot the filament, diminisliing all the way, and overlaid
by the intermediate epithelium, as represented in fig. 18. In other sections the prolon-
gations are not recognizable. Posteriorly the ciliated bands pass into what I have
termed the reticular streak, which here closely resembles the mesenterial epithelium and
is continuous with it.
In the glandular, intermediate, and ciliated areas the trilobed mesenterial filament
thus presents three wholly distinct varieties of cellular tissue. That constituting the
glandular streak bears the closest resemblance to the stomodaeal ectoderm ; the inter-
mediate tissue is in no way distinguishable from the endodermal epithelium ; while that
of the ciliated bands, consisting wholly of long narrow supporting cells, is very specialized
and diff'erent from any other of the polypal tissues.
Prof. McMurrich in his study of the mesenterial filament of Z. sociatus has devoted
.si^ecial attention to the relationship of the ciliated bands and the intermediate region.
With its long, well-developed filaments, the species is specially fitted for such an investi-
gation. Discussing the endodermal origin or otherwise of the intermediate epithelium,
McMurrich affirms that it is certainly continuoiis with the stomodaeal ectoderm above,
and not with the endoderm, and from all the evidence before him is inclined to regard it
as being ectodermal like the rest of the ciliated baud epithelium. So far as histological
evidence can be dejjended upon, an endodermal origin would be assigned it in the
present species ; in character it is wholly distinct from the ciliated and glandular streaks
and stomodaeal ectoderm.
The trilobed condition of the mesenterial filaments in Bunodeopsis is continued only for
a short distance down the mesenteries, when it becomes replaced by the simple filament ;
but the species is not suitable for tracing the actual passage from the one to the other.
McMurrich (p. 2G1) states " that in Z. sociatus there is neither a histological continuity
ACTINIAN BUNODEOPSIS GLOBDLIFERA. 313
nor a histological identity of the ui^per part of the median streak of the filament with
tlie lower or gl andular streak proper." Embryologieal considerations also point to the
conclusion that a difierent origin must be assigned the apical median part of the
mesenterial filament in the uppermost region as compared with the whole filament
below. From the researches of McMurrich, H. V. Wilson (1888), Appellof (1900), and
myself (1899), there is good reason to suppose that the former is a downgrowth of the
stomodjeal (>ctoderm, according to my interpretation, to meet the upgrowing endodermal
filament.
In Bunodeoims the ciliated bauds quickly disappear along with the two lateral meso-
glceal lobes which supported them, and the simple filament remaining is altogether different
from the middle lobe of the more complex filament, although it corresponds with it
in position, representing as it does the termination of the free edge of the mesentery.
The unmodified endodermal tissue, which constituted the intermediate streak of the
complex filament, and covered the middle lobe laterally, is likewise no longer represented
in the simple lobe (PI. 26. tig. 16).
The cellular constituents along the front and sides of the simple filament, however,
most closely recall those of the glandular streak of the compound tilament, consisting
mainly of ciliated supporting cells and coarsely granular gland-cells, with the addition of
a few thick-walled nematocysts. The posterior part of the filament on botli sides is
constituted of much shorter cells, wliich seem to be all ciliated supporting cells, no
nematocysts nor gland-cells being recognizable. Histologically, and probably physio-
logically, the hinder region of the simple filament most nearly approaches the ciliated
bands, though no morphological identity seems possible.
At first the mesenterial mesogloea as it enters the posterior part of tha filament remains
swollen as in the trilobed filament, but in the lower part of the filament it becomes
flattened and rapidly thins out at each side. The anterior surface of the mesogloea
is provided with a layer of extremely delicate muscular fibrils.
In the lower region of the polyp the filaments and free edge of the mesenteries are
greatly folded and contorted, but they by no means crowd the gastro-ccelomic cavity
(PI. 25. fig. 4).
No gonads were developed within any of the jiolyps examined.
In its essential featiu'es, B. glohulifera closely resembles B. strumosa and B. antilliensis,
which were incompletely described in the paper (1897) "The Actiniarian Family
Aliciidae." An important external specific distinction is that the columnar evaginations
in the first are smooth and thick-walled throughout, while in the two older species they
are thin-walled in places, and the large nematocysts occur in restricted thickened bands
or tubercles. In sections this dift'erence is indicated by the practical uniformity in
thickness of the ectodermal evaginations of glohulifera (fig. 5), and their alternating
narrow and broad outline in strmnosa and antilliensis, according as nematocysts are
absent or present.
SECOND SERIES. — ZOOLOGY, VOL. VIII. 46
314 DR. J. E. DUERDEN ON THE
The three agree in such important characteristics as the presence of an ectodermal
columnar and stomoda;al muscle- and ganglion-layer, the absence of gonidial grooves, the
absence of a basilar muscle and of a well-developed parieto-basilar and endodermal
sphincter muscle, and the presence of ciliated bands on tlie mesenterial filaments.
In accordance with these later results, the definition of the genus Butiodeojjsis may be
emended as follows : —
Genus Bunodeopsis, Andres.
Actiniaria with ciliated flattened base, feebly adherent. Column-wall beset proximally
with simple or complex, sessile or pedunculated evaginations, bearing large thick- walled
nematocysts ; distal part of column smooth and delicate. Column with ectodermal
and endodermal muscle- and ganglion-layers ; sphincter muscle very feeble or absent.
Tentacles elongated, deciduous, separated from the polypal cavity by a circular perforated
septum provided with a sphincter muscle. Disc with thickened lips and circumoral
sphincter muscle. Stomodaeum with ectodermal muscle and ganglion-layers, without
gonidial grooves.
Mesenteries normally liexamerovis, two pairs of directives, more than six com^jlete
pairs, often irregularly developed. Parieto-basilar muscle very feeble. Basilar muscle
absent. Mesenterial filaments with ciliated bands. Without acontia and cinclides.
Prom the foregoing account it is clear that, while in many I'espects B. (jlobulifera
possesses primitive Actiuian characteristics, yet in others it is highly difi"erentiated. If
the conclusions of Carlgren (1900) as to the nature of the primitive Actiniae be correct,
amonir the former must be reckoned the occurrence of an ectodermal columnar and
stomodaeal muscle- and nerve-layer, the absence of gonidial grooves from the stomodaeum,
the absence of a basilar muscle, and the weakness of the internal musculature generally.
Among the latter will be included the division of the column into two distinct regions,
of which the upper is naked, and the lower bears liollow evaginations with the ectoderm
broadened and charged with thick-walled nematocysts ; the occurrence of a tentaculo-
coelomic septum provided with a sphincter muscle ; the circumoral ectodermal thickening
(lips) charged with thin-walled nematocysts ; the circumoral endodermal sphincter
muscle ; and tlie regional differentiation of growth of the tentacles and mesenteries.
The first-mentioned characters are those upon which Carlgren has founded the Tribe
Protanthete, which includes the lowest Actiniae, as contrasted with the Tribe Nynantheae,
under which are placed all the more differentiated Actinii3e. Carlgren regards as the
most important characteristic of the Protantheae the presence of the ectodermal longi-
tudinal muscle- and nerve-layer in the column- wall, and gives second place to the absence
of the basilar muscle. Wherever the first-mentioned structures occur, Carlgren would
separate their possessors from the forms witli whicla they have been associated, and jilace
them under the Protanthete, whatever the other characteristics of the sjjecies might be.
In a forthcoming report on the Porto Kican Actinians, to be published by the
U.S. Pish Commission, I have discussed these proposals somewhat, so that it is
ACTINIAN BUNODEOPSIS GLOBULIFEKA. 315
unnecessary to here enter into the subject in detail, except as concerns its bearing on the
present species. Suffice it to say that, while disposed to recognize the primary character
of the ectodermal columnar muscvilature in Actinia?, writers such as McMurrich, Iladdon,
and van Bcneden are not prepared to accord it the taxonomic importance which Carlgren
claims for it. Rather they would regard any species in which such a structure is
retained as among the lowest members of its own particular grovip. And this is the
view which seems most likely to result in a natural grouping of the Actiniaria, one which
will recognize both the primitive and the differentiated characteristics of a species.
Undoubtedly characteristics such as the presence of the ectodermal columnar muscu-
lature, the absence of a basilar muscle, and of ciliated bands from tlic mesenterial
filaments, are in themselves of vastly greater phylogenetic value than such features as
columnar modifications, the origin of one or more tentacles from a mesenterial chamber,
the nature of the sphincter muscle, the number and characteristics of the mesenteries, &c.
But these latter are the dilfei'entiations which alone are available for classificatory
purposes within the Nynanthete, and to wholly separate the lowest forms possessing them
would seem to take away the possibility of a phylogenetic grouping. The possessors of
the ectodermal columnar musculature do not appear to represent a homogeneous
assemblage.
Bunodeopsis is a good example of a genus in wliich, while certain primitive charac-
teristics are retained, imjiortant structux-al modifications have taken jilace in other
respects. To arrange it with others, on account of the former alone, would be to neglect
those differentiations upon which the classification of the Actiuise is mainly founded.
Memoirs and Papers cited in the Text.
1881. Andres, A. — " Iiitoruo alia Scissiparita ilelle Attiuie." Mitt. a. d. Zool. Stat, zu Neapel,
Bd. iii. Heft 1.
1900. Appellof, a.- — " Stiulien iiber Actiaien-Entwickluug." Bergens Museum Aarbog, 1900, no. 1.
1893. Carlhren, O. — " Studien iiber uordische Actinien. I." Kougl. Svenska Vet.-Akad. Handl.
Bd. x.w., no. 10.
1898. Id. — " Zoaiithaiien dor Hamburger !Magalhaensischen Sammelreise." Hamburg.
1899. Id. — " Ueber absclmiirbare Tentakel bei den Actiniarien." Zool. Anzeiger, Bd. x.\ii,
1900. Id. — " Ostafrikanisclie Actinien. Gesammelt von Herrn Dr. F. Stuhlmann 1888 uud 1889."
Mitt. a. d. Nat. Mus. Hamburg, Bd. xvii.
1889. DixoN, G. Y. & A. F. — " Notes on Bunodes thuUiu, Bunodes verrucosa, and Teal'ia crassicornis.^'
Proc. Roy. Dublin Soc., N.S. vol. vi.
1860. DucHASsAiNG, P., et Michelotti, J. — " Memoire sur les Coralliaires des Antilles." Mem. Reale
Accad. Sci. Torino, ser. 2, tom. xix.
1897. DuERDEN, J. E. — "Tbe Actiniarian Family Aliciidae." Ann. Mag. Nat. Hist. ser. 6, vol. xx.
1898. Id. — " Actiniaria around Jamaica." Journ. lustit. Jamaica, vol. ii., no. 5.
1899. Id. — " The Edwardsia-stage oi' Lebrunia, and the Formation of the Gastro-coelomic Cavity."
Jourc. Linn. Soc, Zool. vol. xxvii.
1900. Id. — " Jamaican Actiniaria." Part II. Stichodactyliuie and Zoantheae." Trans,. R, Dublin Soc.
vol. vii., ser. 2.
316 DE. J. E. DUERDEN ON THE
1898. Haddon, a. C. — "The Actiniaria of Torres Straits." Trans. R. Dublin Soc. vol. vi., ser. 2.
1898. KwiETNiEwsKi, C. R. — " Actiniaria von Ambon und Thursday Island." Semon, Zool. Forscli-
ungsreisen in Australicn, &c., Jena.
1889. McMuRRicH, J. P.— "The Actiniaria of the Bahama Islands, W. I " Journ. Morph. vol. iii.
1890. Id. — " Contributions on the Morphology of the Actinozoa. I. The Structure of Cerianthus
americanus." Journ. Morph. vol. iv., no. 2.
1893. Id. — " Report on the Actinire collected by the U.S. Fish Commission Steamer ' Albatross •" during
the winter of 1887-1888." Proc. U.S. Nat. Mus. vol. xvi.
1899. Id. — " Contributions on the Morphology of the Actinozoa. V. The Mesenterial Filaments in
Zoanthus sociatus (Ellis)." Zool. Bull. vol. ii., no. 6.
1899. Parker, G. H.—" Longitudinal Fission \u Metridium /Har^rma^MOT, Milne-Edwards." Bull. Mus.
Comp. Zool. Harvard, vol. xxxv., no. 3.
1899. Verrill, A. E. — " Descriptions of imperfectly-known and new Actinians, with Critical Notes on
other Species. III." Amer. Journ. Sci. vol. vii.
1900. Id. — ■" Additions to the Authozoa and Hydrozoa of the Bermudas." Trans. Conn. Acad.
Sci. vol. X.
1888. Wilson, H. V. — "On the Development of Manicina areo/ata." Journ. Morph. vol. ii.
EXPLANATION OF PLATES 25 & 26.
Reference letters.
I., II., III., = orders of mesenteries. c.s. = ciliated baud. col.w. = column-wall. d. = directive
mesenteries. disc = d\sc. ec/. = ectoderm. ec<.?«. = ectodermal muscle. end. ^endodcvm..
ewrf.wi. = endodermal muscle. ev. = columnar evagination. ^.a". = glandular streak. l.s.=-
intermediate streak. »!es. = m°soglcea. mes<.= mesentery. ?n.^/. = mesenterial filament,
wew. = nematocyst. w7"./. = nerve-layer. or.sph. :^ora.l sphincter. re<.»«.= retractor muscle.
s^.=stomodfeum. /.c.s.=:tentaculo-coelomic septum, t., <eH^ = tentacle. ^.s;;/j. = tentacular
sphincter. <>Yms.wi. = transverse muscle. »ooa-. = Zooxanthellae.
Fig. 1, a, b, c. Three different polyps, somewhat enlarged : a and b in the living expanded condition ;
c, a preserved polyp.
2. Vertical section through a portion of the basal disc, including a tangential section of a mesentery.
No basilar muscle occurs at the insertion of the mesentery on the basal wall, x 400.
3. Vertical section through the upper region of a polyp, including a part of one side of a tentacle,
the upper part of the column-wall, and one side of the tentaculo-coelomic partition with the
tentacular sphincter at its free extremity. The poly pal wall in all its three layers is here more
swollen than usual, x 400.
4. Median vertical section through an entire polyp, but including only the basal part of two
tentacles. The stomodseum is slightly protruding, and several mesenteries are cut obliquely.
X 70.
5. Vertical section through a columnar evagination. x 400.
ACTINIAN BUNODEOPSIS GLOBULIFERA. 317
(). Vertical section tlirougli the basal jjart of a tentacle with tlic tcntaculo-ccjelomic septum attached,
the disc, and upper portion of the stomodaeal wall. x 300.
7. Transverse section through a portion of the stomodteal wall, x 400.
8. Nematooysts : a, thin-walled form from the tentacular ectoderm; 6, thick- walled form from the
columnar evaginations ; c, from the stomodrcum ; «?, from the mesenterial filaments, x 1200.
!). Transverse section of a polyp with only three pairs of complete mesenteries, x 70.
10. Transverse section of a polyp with three pairs of complete mesenteries, one moiety of another
pair, and at one region numerous incomplete pairs. X 70.
1 1 . Transverse section of a polyp with four pairs of complete mesenteries, a moiety of two other
pairs, and a region with many incomplete pairs. A section of a tentacle whicli has intruded
within the polypal cavity is represented at the upper right-hand corner. x 70.
I'J. Transverse section of a polyp in which for the most part hexameral regularity prevails, but at
one region growth is proceeding irregularly, x 70.
13. Transverse section through part of the stomodseal region of a polyp, including the tentacular
zone. The relations of the tentacles to the mesenterial chambers and the teutaculo-coeloniic
septa are cleiirly shown. x 70.
If. Transverse section of the same polyp at a slightly lower level. Mesenterial pairs belonging to
the first, second, and third orders occur regularly throughout the greater part of the polyp,
but the sequence is irregular at the upper right-hand side. x 70.
15. Transverse section of a large polyp in which three orders of mesenteries are regularly
developed, except on the right upper side. Most of the pairs of the second order have become
inserted on the stomodseum. x 70.
Ki. Transverse section of a portion of the column-wall a little below the stomodseal region, with a
mesentery attached, bearing a simple mesenterial filament, x 400.
17. Transverse section of a mesentery and the portion of the column-wall to which it is attached.
X 300.
18. Transverse section through a trilobed mesenterial filament. x 400.
SIX'OXl) SElllES. — ZOOLOGY. VOL. VTII. i7
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^^^ 22 IJ03
2iid Ser. ZOOLOGY.
[VOL. VIII. PART 10.
lo^W'X
THE
TRANSACTTOi\S
OF
THE LINNEAi\ SOCIETY OE LONDON.
ON THE MORPHOLOGY OF THE BRAIN IN THE MAMMALIA,
WITH SPECL\L REFERENCE TO THAT OF THE LEMURS,
RECENT AND EXTINCT.
BY
G. ELLIOT SMITH, M.D. (Sydney),
FELwn- OF St. John's College, CAMnRiDOE, and Professor of Anatomy,
GOVEHXMEMT ScHOOL OF ilKDICI>-E, CilEO.
LONDON:
PRTNTKI) FOIl TIIK LIKNEAN SOCIETY
BY TAVl.OIi AND FRANCIS, RKT> l.ION COURT. FLEET STREET.
SOLD AT THE SOCIETY'S APARTMENTS, BURLINOTOK-TTOTSE. PICCADILLY, W.,
A-SV RY LON-GMANS, GREEX, AND CO., PATEENOSTER-KOW.
Februan/ 1003.
^UC 22 1903 [ 319 ]
X. On the 3forpholor/f/ of (hr Brain in the Mammalia, icith Special Reference to thai
of the Lemurs, Recent and Extinct. Bij G. Elliot Smith, M.D. {Sydney), Fellow
of St. John's CoUerfC, Cambridge, and Professor of Anatomy, Egypfiait Cocernment
School of 3Iedicine, Cairo. [Communicated by Prof. G. 13. Howes, D.Sc, LL.I) ,
F.R.S., Sec. Linn. Sac.)
(With C6 Illastrations in the Text.)
Itead ULh Mareli, VJ(r2.
Introductiox.
1 HIS investigation was undertaken pi-imarily to consider the possihility of homologizing
the sulci of the cerehral hcmis2)herc in different Orders of Mammals. A^'itli thi.s ol)jcct
in view, the brain was examined in almost every genus in the whole class of Mammalia ;
but as this yielded such an immense and unwieldy mass of data, it became necessary,
both for the sake of clearness and conciseness of exposition as well as to admit of the
adequate discussion of the signiticance of the facts, to limit the sco])e of the enquiry, or,
rather, to select the most instructive body of evidence which might tyjiify the
conclusions to be drawn from a study of the whole mass of data.
This process of curtailment has been accomplished in two ways. In the first place,
attention has been almost wholly concentrated on the consideration of the homologies of
tlic furrows called " calcarine " and " Sylvian." Tiiese are the only sulci (in addition, of
course, to the hippocampal and rhiual fissures) which are absolutely constant in all
Primates. At the outset of this investigation it became abundantly evident, firstly,
that if the calcarine sulcus is a distinctive feature of the Primates and is not represented
in the brain of other Mammalia, the search for homologues of the other, less stable,
furrow on the mesial surface of the hemisphere must a fortiori be utterly futile; and,
secondly, if the Sylvian flssiu-e of the Apes and Man is represented in other mammals by
no sulcus other than that inconstant and exceedingly unstal)le series of different *
furrows, which in various Carnivores, Ungulates, Edentates, Rodents, and Marsupials is
commonly called "Sylvian," the possibility of homologizing any sulcus on the lateral
aspect of the hemisphere of the Primates with a furrow on the surface of the brain ot
other mammals becomes utterly destroyed. That this not altogether obvious conclusion
must inevitably be drawn from the premiss just stated will be abundantly shown in the
following pages. "We shall, therefore, best arrive at some decision regarding the main
question, by devoting our attention chiefly to the Sylvian and calcarine furrows, without,
however, altogether neglecting the other sulci. In the second place, I have deemed it
desirable to further limit the dimensions of this work, by restricting the detailed account
* Xon-homologous.
SECOND SERIES. — ZOOLOGY, VOL. VIII. 48
320 PEOF. G. ELLIOT SMITH OX THE
of the sulci (and the organ in which they occur) to one circumscribed group of mammals ;
and, so as not to also narrow tbe scope of this discussion by so doing, I have freely
introduced comparative data, which serve to indicate the behaviour of each individual
sulcus in the whole Mammalian class.
Tlic brain of tlie Lemurs lias been selected as the aptest expression of these require-
ments. I can best explain this by anticipating some of tbe conclusions of this research.
Tlie features of the Prosimian brain become really intelligible only on the supposition
that the Lemurs have advanced a considerable distance in the main stream of the
evolution of the Primates and have then retrograded; among other manifestations of
this retrogressive process many interesting phases of the disintegration of the cerebral
sulci are exhibited, so that it becomes possible to recognize the constituent elements of
many compound sulci in the Primates, and so the more readily to compare them with the
lurrows found in other mammals.
My choice of the Lemurs was further confirmed by other considerations of a very
diiferent nature. The opportunity of examining a much more representative collection
of Prosimian brains ihan had previously fiilleu to the lot of any one observer, and the
imique facilities for studying the brain-form of the extinct Lemuroids, for which I am
indebted to the kindness of Dr. Forsyth jNIajor, were additional reasons for choosing the
brain of the Lemurs as the touchstone on which to test the hypotheses suggested by a
study of the brain <-.f other mammals. Moreover, the present state of our knowledge of
the mammalian brain is so unsatisfactory, that no apology would be necessary for
discussing its apparently obvious features in any Order, and least of all in the
enigmatical family of Lemurs ; for in no group of mammals is accurate knowledge
of the brain more urgently wanted than in the latter. The voluminous literature of the
Prosimian brain is jiarticularly barren so far as exact and authentic statements of fact
are concerned, and unduly rich in the grossest inaccuracies.
The foregoing statement of the object of these notes will explain their nature and
scope. They do not pretend to be a monograph on the Prosimian brain, but are rather
an account of the more variable features of the mammalian brain, as these are exhibited
in the Lemurs. I have deemed it advisable, if for no other purpose than to give some
idea of the " setting " of the j^attern of the cerebral convolutions, to give a brief account
of the outstanding features of the brain as a whole. The morphology of the mammalian
cerebellum is incidentally discussed also.
In attempting to settle such an essentially morphological problem as the search after
a fundamental plan underlying the apparently heterogeneous maze of cerebral sulci
found in the diflferent Orders, the observer is afforded an excellent opportunity of
appreciating at their true worth the factors which disguise the common plan. The
value of the evidence of the brain as an indication of the closeness of the bonds of
affinity between the Lemurs and other mammals hence falls not unnaturally within
the scope of this study.
/
MORPHOLOGY OF THE BEAIX IX TIIF. .MAMMALIA. 321
Tee Liteuature *.
Since Ticdomann f first described the brain of Lemur moiKjo:, a very considerable
numl)er of memoirs on tbc Prosimian braiu liave l)ceii published. Tiie interest of the
numerous early descriptions, Avhicli include those of Lcuret (1830), Schroeder A'an der
Kolk and Vrolik (IHiS 51), and van dor Iloeveu aud van Canipm- (185!)), is almost purely
historical. But the same period yielded an excellent memoir by Burmeister on the
braiu of Tarslns %, -which represents to the present day all the accurate information
we possess concerning- this ])rain. It is of further interest as the first really valuable
contribution to our knowledge of the Prosimian brain.
The most critically-careful descriptions of any Lemurs' brains are contained in the
two memoirs of W. H. Flower §, which include brief accounts of the brain in Lemur
fiilvus [_nigrifrons] and Nijctlcehus tardigradns, with notes on that of a Galacio.
In 1886 Richard Owen published the first account of the brain of CliiroiH.ys, in a
memoir 1| which lacks the clearness and definiteness of Elower's monographs.
In 1872 Paul Gcnwais gave an interesting account of the shape of the brain in the
A arious genera of LiMuurs from the data supplied by the study of cranial casts ^[ ; but as
lie made little or no attempt ** to check the results of his examination of the cranial
moulds by comparison with the actual braina, he draws many inferences concerning the
latter, and also with regard to tlie affinities of tlie Lemurs (see the title of his memoir I),
which are not justified by a study of the brain itself.
In 1875 A. Milne-Edwards gave an account of the brain in the Indrisinoe ft, whic-h
represents all tlie published data concerning the actual brains in the genera Lndris,
Fropitliecus, and Acti/iis. [Many recent \-\riters (siicli as Flatau and Jacobsohn, whose
acquaintance with the bibliography of their siibject is as slight as their knowledge oi'
the actual mammalian brain) attribute the knowledge of the brain in Lidrls to
Chudziuski, who borrowed from Milne-Edwards's monograph, and to Znckerkandl, A\hu
makes the merest reference to the brain of VropUhecua diadema + + .]
* Several of the titles, more especial!}- of the eailier memoirs, are not given in this list. A\ hen recently in
I'^ngland I read the whole literature of the Prosimian hrain, excepting only the memoir of Bischotf, which 1
unintentionally overlooked. But 1 neglected to make any notes on such memoirs as contained no information of
any value, and even neglected the titles of some. To an exile from the lands of lihraries, the gifts of reprints
such as 1 have to acknowledge from Dr. Forsyth ^lajor, Mr. Eeddard, and Professor IJuickhardt are especially
welcome.
i" ' Icones Cerebri .Simiarura et quorundam ilammalium rariorum,' Heidelberg, ISiii.
% ' Beitriige zur niiheren Kenntniss der Gattung Tarsitts,' Berlin, lS4(i.
§ " On the Posterior Lobes of the Cerebrum of the Quadrumaiui,'" I'liil. Trans. IS'iL' : and '• On the Brain of tlie
.lavan Loris (Steitops jttvdnkus, Hlig.)," Trans. Zool. Soc. 186(i.
]i "On the Aye-aye,"" Trans. Zool. Soc, vol. v. 180(5, pp. G8 et se'j.
•[ " Mcmoires sur les formes cerebrales propres a I'ordre des Lcmures,"" Journal do Zoologie, tome i.
^■* He refers to an earlier account of a Lemur"s brain in his Hist. d. Mainmiferes, which I have had no opportunity
of consulting.
tt In Grandidier's ' Histoirc Phj'sique, Xaturollc et Politique de JLidugascar," (2) tome vi. texte (Paiis.
1^7.")1.
Ji E. Zuckerkandl, ' Uebcr das Ileichcentrum," Stuttgart, 18S7, Taf. \i. fig. 2S (pp. 2'J, 45, & 70).
48*
:J22 PROF. G. ELLIOT SMITH ON THE
More recent numbers of the ' Histoire ' contain illustrations of tlie brain in the genera
Lemur, Sapalemur, and Lepidolemur.
The references to this will be found below in the accounts of these genera.
I have not seen Wernicke's short account * of the brain o^ Lemur f ulcus [inongoz], and
know of it merely from the passing i-eferences in three of the more recent memoir'^
(Itiotcd below.
In ISDO J. T. Oudemans gave us the best account we jiossess of the brain of the
Aye-aye f ; and in the same year W. Turner examined + in the light of comparative data
riower's above-quoted accounts of the Lemur's br.iin.
r. E. Beddard published an account of the brain of Sapalemur grlseus in 1891,
of those of an extensive series of A^arious species of Jjem.ur, Galago, Chirogale, Lor'n^,
Xyclicehus, and Ferodlct'icns in 1895, and those of Mapalemnr simns and another
ILapalemur grlseus this year (1901) §.
In 1895 the first of Theophile Chiidzinski's extraordinary memoirs 1| made its appear-
ance. It is a most peculiar document. The Avriter discusses the brains of Lemur,
Indris, and what he calls Loris gracilis. The data for the first two are apparently
borrowed from riowei-'s and Milne-Edwards's memoirs^; and the third — the only one
wliich the author himself describes — certainly belongs not to Loris gracilis, but to
ygclicehus tardigrudus. This error is the more singular because the author prefaces his
description with the remark that " it is easy, with a little practice, to recognize without
any difficulty the Order, Family, Genus, and often even the Species, to Avhich a particular
brain belongs " (p. 435). His second memoir, which deals with the brain of Chiromijs,
is so vaguely written and so poorly illustrated as to be almost valueless.
In the same year (1896) Theodor Zielien gave an account of the cerebral hemispheres
of a large number of Lemurs, the nucleus of Avhich consists of the collection in the
Pioyal College of Surgeons **.
In 1889 riatau and Jacobsohn gave a detailed account f f of the whole brain of a Lemur
macaco [or " macao,'' as they spell it throughout their book], and of a second bi-ain
* Archiv fin- rsycbmtrie, lid. vi. Taf. 4. fig. 10.
t " Beitriige zur Kcniituiss des Chiroinijs miuUii/dscai-ietinU, C'uv.," Xiituurk. Verb, der Koniiikl. Akadomie,
Amsterdam, Doel .xxvii.
T " The Convolutions of the Brain," Journal of .\natomy and Physiology, vol. xxv.
ii " Additional Notes upon llnpalemur griseas,'' Proe. Zool. Soc. ISUl, p. 456. " On the Brain in the Lemurs,''
I'joc. Zool. Soc. 18^5, pp. 142 ei scq. " Notes on the Broad-nosed Letnnr, Hfqxdi'mur siiiitis/' ProcZool. Soc. liiOJ,
1». 127. Compare also " On certain Points in the Anatomy of Callitlirir (ojv^iia^n,"' Novitates Zoologicaj, vol. viii.
Oct. 1901.
I " Sur les plis cerehraux des Lemuriens on general et du Loris grele en iiartieulier,"" Bull, de la Soc. d'Anthropul.
do Paris, 4'" serio, tome vi. 18U5, pp. 484-464. ■• Sur les ])lis cerebraux d'uii Aye-aye {C/wu-omi/s, MyslpithecHu,
Siugerat)," ihkl. tome vii. 1896, pp. 12 et sc-q.
*! I make this statement i'rom memory, as I am now unable to consult the original.
** " Ucber die Grosshirnfiinhung der Halbatlen uiid die Deutung einigcr Furobeu des mensohlichen Gehirns,"
Arch. f. Psychiatric, Bd. xxviii. 181J6, pp. 808 ct scq.
Tt 'Handbueh dor Anatomic uud vergleicheuden .\natouiic des Centralnervensystems der Siiugethiere,' 1 Teil.
litrlin, lt9!).
MORPHOLOGY OF THE BRAIN IN THE MAMMALIA. 323
Mliicli they call " Stenops gracilis.'" There can, however, be little doubt that tiie
ambitious authors of this extraordinarily inaccurate book have followed in the footsteps
of Chudzinski, and have mistaken S/enojis [Nycticebus] for Lor Is.
In the instructive memoir on the insula in the Carnivora which M. Holl * published
in 1809, thei-e is an interesting figure representing the lateral aspect of the right cei-ebral
hemisphere of a Lemur of unknown species.
In 1!)00 E. Zuckerkandlt gave a brief account of the structure Avhich he c:ills by llie
misl(;ading title " Balkenwindung " in four species of the genus Lemur and in Chiroin/js ;
;ind in the same year the most extraordinary contribution to the literatui'e relating to
the Prosimian brain was joublished +.
In addition to Tiu-ner, to whose memoir reference has already been made, several other
Avriters have discussed the anatomy of the Lemur's brain, without apparently adding any
new data to the common stock.
The writings of A. J. Parker § and Gegenbaur || belong to this categoi-y.
Dr. Porsyth Major has described the form of the brain in two sub-fossil Lemuroids f;
and Rudolph Burckhardt has published some most extraordinary criticisms of Porsytli
^Major's memoir, in the course of which he commits the blunders Avhich he erroneously
accuses tlie latter of perpetrating.
Max AVeber, and in a lesser degree Eugene Dubois and Theodor Ziehen, have
investigated the weight of the brain in the various Prosimian genera. Their results
are discussed in the body of this Memoir.
The nucleus of the material upon which this account is based consists of four brains of
Lemur fill c lis [labelled mougoz, ulbifrons, and two of tliem nlgrifro>is\ and one each of
Lemur varius. Lemur catta, Nycticebus tardiyradus, Loris gracilis, Perodicticus ijotta,
Galago Garneiti, Galago crassicaudata, and Microcebus Smithi, in the Collection of the
Royal College of Surgeons in London. These are the same brains upon which Ziehen's
memoir was based ; but as I found it necessary to remove the pia and arachnoid
membranes before I could pro2)erly see the surface of any of these specimens, the reader
will appreciate one of the reasons for any discrepancies which may bo found in the two
accounts of tlie same specimens.
Since Ziehen's visit to the College of Surgeons two brains of CJiiromys have been
placed in the Galleries. One of these was found in the College Stores and the other was
purchased from the Zoological Society, Mr. Eeddard kindly consenting to part with his
* " Ueber die Insel des CaruivorengcLirns," Arcb. f. Aimt. u. Phys., Anat. Abtli., Taf. 12. ti^;. Hi.
t " Beitriige /.ur Anatomic des lleicbcentrums," 8it/.UDgsbcritbtL'u d. kaiserl. Akud. der Wisseiisch. in Wini. Malb.-
naliirw. CI., Bd. cix. Abth. iii., .July 1000, \). 2.
J Ilonry C. Chapman, " Observations upon the .Vnatoray of Hijhlates Icuciscus and Chlromys imulnijusairiemt.i."
Proc. Acad. Xat. Sci. Philadelphia, May lOOO, p. 420.
§ " Morpholojiy "f the Cerebral Convolutions, with Special Reference to the Order of Primates," Journ. .\cad. Nat,
Sci. Philadelphia, 2iid series, vol. x. (ISUG).
II ' Vergleich. Anat. der Wirbelthiere,' i. Bd., Leipzig, 1898, pp. TGG & 7<)7.
i[ "On the Br;iius of two Sub-Fossil Malagasy Lemuroids," Proc. Itov. f^oc. vol. 62 (1897), pp. 46-50.
82 t TEOF. G. ELLIOT SMITH ON THE
specimen, so that one anatomist might have the chance of describing- and comparing the
two brains. Dr. Charles Hose lias from time to lime sent to Mr. W. L. H. Duckworth
in Cambridge a valuable series of excellently preserved brains from Borneo, which I
have had an opportunity of examining. Mr. Duckworth very generously placed at my
disposal two brains of Tursius, one of which had been fixed and preserved in situ by
means of i'ormalin injections, and the other had been extracted with all that care and
skill for which Mr. Hose is so well known among zoologists. This In-ain was preserved
for histological purjwses inMiiller's tiuid. Mr. Duckworth also permitted me to examine
many other specimens, including an excellent brain of Galago Garnetti. Captain Stanley
Flower has kindly placed at my disposal the fresh bodies of five Lemurs and SiNycticebus
which died in the Ghizch Zoological Gardens diuing the last ^qw montlis, and I have
thus been able to examine the brain of these animals in a jjei'fectly fresh condition. He
has also given me skulls of Lemur macaco and Nyciicchus tardigradus. From these I
liave been al)le to make cranial casts.
Dr. Forsyth Major, whose generous help has been afforded me in many ways during
the course of this investigation, gave me the brain of Microcebus Smithl and lent me
the skulls of various Lemurs. On his suggestion, the authorities in the Geological and
Zoological Departments of the British Museum kindly permitted me to examine their
large collections of crania of recent and extinct Lemurs. I have also examined all the
crania of Prosimije in the collection at the Royal College of Surgeons.
1 have thus been able to study the shape of the Ijrain and the arrangement of the
cerebral furrows not only in every genus and most species of recent Lemurs, but also
in the extinct Lemur Jullyi, Globileimir, and both an adult and a young Ilegaladapis.
,Vs Dr. Forsyth Major was engaged in investigating these A'aluable extinct forms, my
indebtedness to him is all the greater for allowing me to examine them.
It is commonly supposed that a mould of the cranial cavity can give little exact
information beyond indicating the shape of the brain. In most Apes this is to a very
great extent true. But in most mammals in which the pattern formed by the sulci is
not very complex (and also even in some, especially Carnivores, in A\liich a very rich
supply of sulci exists) a series of prominent ridges develop on the inner face of the
cranial wall so as to accurately map out the plan of the sulci. These ridges are especially
Avell developed in all Lemurs, excepting only the extinct form Globilemur, in which the
pattern becomes blurred as in most Apes.
It is, however, absolutely necessary, if one is to accurately interpret the plan of the
sulci from the arrangement of these bony ridges, to carefully study the actual braius of
several genera and the crania in whicli they were lodged, to apjjreciate several sources of
fallacy, which have been so unfortunately demonstrated in Gervais's imcontrolled study of
casts [oj). cit. supra). If this precaution be taken, one is able to map out the plan of
the sulci on the outer sitrface of the hemisphere in a plaster mould with the certain
conviction that he is accurately interpretiug the configuration of the brain itself.
From the specimens {Lemur, Kycllcebas, Jlicrocebus, and Tarsius) M'hich I have been
able to devote to histological purposes, I have been able to determine many interesting
features in the arrangement of the fibre-tracts, especially the optic and olfactory paths
MOEPIIOLOGT OF THE BKATX IX THE ^LA.MMALIA. 325
in the cerebral lioinisplieres and the anatomy of the pyramidal tracts. But as these
data have no direct hcarinf? upon the subjects with which this memoir is chiefly concerned,
and would demand a much larger numl)er of illustrations to make their arrangement
intelligible than I dare ask for at present, I sliall defer their publication until some
other occasion.
An investigation such as this, which aims at explaining th(! salient features of the
cerebrum in such a huge assemblage of animals as the Class Mammalia includes, must
entail many obligations.
The unrivalled collection of mammalian brains in the Galleries and Stores of the lloyal
College of Surgeons were freely placed at my disposal by Professor Charles Stewart. 'I'o
Dr. Henry "Woodward, Dr. Charles W. .Vndrews, and Mr. Oldficld Thomas, I am indebted
for the opportunity of examining the large collections of mammalian crania in tbe
British Mviseum. Mr. W. L. H. Duckworth, in Cambridge, also freely permitted me to
make use of the considerable collection of mammalian brains in his possession.
For valuable gifts of material for comparative purposes, I am under tlie greatest
obligations to Professor Howes, the late Mr. Martin "Woodward, Professors "Wilson and
Bahhvin Sjiencer and Mr. James P. Hill (Australia), Dr. Pobert Brown (South Africa),
Dr. Charles Hose (Borneo), and especially Captain Stanley Plower (Egypt), among many
others. "Without the rich (and I believe imprecedented) collection of representative
mammalian brains wliich I have thus been able to study, this communication would
have lost its only claim to consideration, l. e. that it truly sets forth the tendencies of
brain-evolution in the whole Mammalia, and not only in one small circumscribed group.
For the facilities which have been so generously afforded me in the accomplishment
of this work I must express my great sense of indebtedness to all the gentlemen who
have a.ssisted me.
The Cerebral HemlsphetvE ix the Genus Lemur.
It is not necessary to describe the general form of the brain in the various genera of
Lerauroidea, because the bulk of tbe literature quoted above deals with this subject, and
certain of them, such as that of Gervais, are concerned exclusively with the shape of
the brain. ^Moreover, all the important features of the brain-form will be discussed
incidentally in these notes along with those peculiarities which are causally related to
the modilications in shape.
When the dorsal surface of the brain of a Lemur is exposed by removing the roof of
the cranium, little else than the cerebral hemispheres can be seen. In front of the
pointed anterior pole of each hemisphere, the exposed area of the olfactoi-y bulb projects
very slightly more than one millimetre : and behind the broader, somewhat spatulate
caudal extremities of the hemisplieres in most cases only a very narrow band of the
cerebellum is exposed as a series of seven rounded knobs fringing the margins of the
hemispheres ((ig. 1). So that in order to study the relationships of the constituent
parts of the brain, it is necessary to turn to the examination of the ventral surface
(fig. 1). In front there is the broad flattened olfactory bulb, about 11 nun. long and
•til/*
PROF. G. ELLIOT SMITH ON THE
Fig. Z.
Fig. 1.- — Lemur macaco. The dorsal aspect of the brain. Nat. size.
Fig. 2. — Lemur fulms. The dorsal aspect of the brain. Nat. size.
Fig. 3. — Lemur varivs. The dorsal aspect of the brain. Nat. size.
NorF.. — The differences in the shape and the extent of cerebral overlap of the cerebellum in figs. 1, 2, and ;j are
probab!}' merely individual variations and have no specific value.
For the siguiBcance of the reference-letters see below ; the meaning of the letters d, c, and /' is explained in thy
text *.
* List of
JSulcu.-; ansatus — sa.
,, calcarinus — sc.
„ centralis — see.
,, coroualis. — seo.
,, cruciatus — scr.
diagoualis — sd.
ectosylvius anterior — sea.
„ „ posterior — seji.
,, genualis — sg.
,, intercalaris — si.
,, lateralis — si.
J, orbitalis — so.
,. paracalcarinus — spc.
,, postlateralis — sjd.
„ postsylvius — sj).
Refeeexce-lettees ts the Diagrams.
Sulcus pseudosylvius — S2is,
„ rectus- — sr.
„ retrocalcarinus — srs
„ suprasylvius — s«.
Fissura hippocampi — jh.
„ rhinalis — fr.
„ prima — fp.
„ secunda — /'«.
Bulbus olfactorius — bo.
Calcar avis — eal.
Cerebellum — c.
Claustrum — cl.
Commissura anterior — en.
Copula pyramidale — cop.
Corpus caUosum — cc.
MOHPHOLOGT OF THE BRAIN IN THE MAMMALIA.
327
Rg. 4.
Lemur fvlvus. — The \Tentral aspect of the hrain. X 2.
Corpus striatum — cs.
Corpora quadrigemina — cq.
Anterior pair vi corpora quadrigemina — cqa.
Posterior ,> „ — c(jik
Corpus geniexilatum ant. (latcrale) — cya.
„ „ post, (mcdiale) — cgp^
Fascia dentata— /tf.
Flocculus — -Jl.
Fornix — for.
Hippocampus — h.
Insula — inf.
I.obus flocculi — !f.
,, anterior — la.
„ medius — hn.
,, pyritormis — Ij'-
Nodulus — HO.
Nucleus amygdal;e — na.
Olive — 0.
Paraflocculus — jif.
Pons — p.
Pedunculus olfactorius — ^jo.
Psalterittm — ps.
Pyramid (bulbar) — jiif.
„ (cerebellar) — pyr,
Tractus olfactorius — tol.
„ opticus — to2}.
Trapezium — tr.
Tuberculum hippocampi — Ih.
„ olfactorium — to.
„ acusticum — (d.
Uvula — «!'.
Vallecula Sylvii — vS.
Oilier individual references are e.cplaiited in the te.vl.
The name " retrocalcarinc "' has been substituted for " postcalcarine "' throughout, the latter having been already
employed in another sense by other writers.
SECOND SERIES. — KOOLOGY, VOL. VIII. 49
328 PROF. G. ELLIOT SillTTI OX THE
about lialf as hroad, lying in contact witli the under surface of the eerebral hemisphere
except in the anterior lo mm. of its extent *.
Its posterior end is joined to a flattened peduncle Avhich is as broad as the bulb itself.
It is very short, for it pursues a course of only 3 or i mm. before it becomes attached to
the base of the cerebral hemisphere. In tliis respect it differs most decidedly from the
very long and attenuated olfactory peduncle of the Apes ; but, oa the other hand, a short
thick peduncle like that of the Lemur is found in the vast majority of mammals, and
must be regarded as the common mammalian type.
The lateral half of the peduncle is covered by a dense thick l)undle oO fibres which
constitutes the tractus olfactorius [lateralis]. Tliis white covering also extends over the
lateral border of the pediincle. The internal half of the ventral surface and the mesial
border present a greyish appearance, because the medullary fibres of the so-called mesial
tract are much less numerous and more scattered than those of the lateral tract. In the
elongated peduncle of all Monkeys the whole peduncle appears to consist of nerve-tibres,
so that its real constitution is not so obvious as it is in the Lemurs.
Just behind the insertioa of the olfactory peduncle into the base of the hemisphere
there is a transversely -placed, oval, puckered area of grey sul)stance, 4 mm. broad
and about 2-5 mm. in the sagittal direction. This is the tuberculum olfactorium —
a pecvdiar cortical formation coating the ventral surface of the head of the corpus
striatum. The features of this body are exceedingly constant thi'oughout the Mammalia,
l)ut become somewhat obscured in the adult In-ain of Man and the Anthropoid Apes.
The mesial surface of the olfactory peduncle extends on to the corresponding surface of
the hemisphere just in front of the tuberculum olfactorium.
The lateral wall of the olfactory peduncle is prolonged backward into continuity with
the pyriform lobe. The latter begins in front as a narrow horizontally placed grey band
3 mm. wide, the internal part of which is covered I)y the (external) olfactory tract.
Opposite the outer extremity of the tuberculum olfactorium it becomes bent at right
angles upon itself (ilg. G) and at the same time broadens to almost four times the width,
so that its posterior iTM-t forms a very prominent rounded boss projecting 4 mm. below
the level of the ventral surface of the anterior part. The transverse depression formed
by this sudden bend is the vallecula Sylvii — a furrow which is quite distinct from the
fossa and iissura Sylvii.
The vallecula Sylvii is much deeper and n\oie pronounced in the L^nuu- than in most
mammalian brains of other orders. In some of the smaller Xew- World A]jes, such as
the Hapalidie, the vallecula presents features exactly analogous to that just described,
the posterior pavt of the pyriform lol)e being bent at an angle of S)(K on the anterior
part ; but in most Apes the angle of bending hecomes much more acute, the posterior
part biilges forward over tlie anterior part of tlie pyriform lobe ; and the vallecula Sylvii
becomes a very deep cleft, which is commonly regarded as part of the stem of the true
Sylvian fissure.
* These measuremeuts refer to a fresh specimen of Lemur ftihiis, and may be regarded as a fair average of all
the specimens of the genus Ltinar which 1 have examined.
MORPHOLOfiY OF THE BRAIN L\ TUK .A!.\M MALTA. 329
At tlu> luitero-mesial aspect of tlic uatil'orin ciiiiiiencc (as the posterior part oi' the
pyriform lohe is often called) there is an elliptical area hounded laterally by a shallow
i;roove. Retzins has called liiis the " livrus luiiaris " and has sliown how remarkably
constant it is in 1h(> ^ra-nnialia *. I am not aware that anyone lias ever considered the
significance of tliis little body (which I liave studied not only in most Mammals, but
also in the Reptilia) ; its nature is at once revealed in the Lemur's bi-ain by a section at
rig'lit angles to its long- axis. It is clearly nothing else than the surface of the nucleus
amygdalte and is not therefore strictly a ])art of the pyriloi-m lobe.
At llie mesial border of the |)yrif()rni lobe we tiiul a peculiar depressed grey band,
whicli is erroneously labelled "gyrus uncinatus" by Tlatau and Jacobsohnf. It is
merely the low or end of the fascia dentata, whereas the uncinate gyrus is represented by
the pyriform lobe itself. These writers adopt a very umisual and utterly misleading
nomenclature Avhen they call the anterior part of the pyriform lobe by the name " gyrus
hi))p()canipi '" (p. 195).
In most Lemurs the rhinal fissure, which is the lateral boiuidaryof the pyriform lobe,
becomes abnost wholly obliterated. This also happens in such Chiroptcra as Pferopus
and Ci/i/onijc/('/-/s. I have, liowever, been fortunate enough to obtain the brain of a
Lemvrfiiicnft, in which the rliinal fissure is retained in a form as distinct as it is in most
mammals (figs. |. and (5). I hav(> also in my possession another brain of the same genus
in which the whole of the rhiiial llssure is visible, although it is very shallow ; and also
the cranial cast of yet another in which it is cpiite distinct. These three examples are,
however, exceptional.
The fissure is distinctly visible upon the lateral aspect of the first-mentioned brain
(fig. (5) pursuing a course (fx'om the cleft between the olfactory bulb and the hemisphere)
backward with a very slight upward inclination. Just behind the Vcillecula Sylvii it
suddenly bends downward at an obtuse angle and curves horizontally backward to a
point midway between the vallecula Sylvii and the posterior margin of the hemisphere.
Behind this point there is no line of demarcation betAveen the pyriform lobe and the
neopallium.
In Monkeys no part of the rhinal fissiu-e is visible upon the lateral aspect of the
hemisphere : the relatively greater size of the neopallium and the much smaller
absolute dimensions of the pyriform lobe shift it wholly on to the ventral, or even
mesial, surface.
In the brain of most members of the genus Lemur, all that remains to represent the
rhinal fissure is the angle and olten only the posterior limb of the angle (fig. 7). In
such cases, however, a careful examination enables one to distinguish the pyriform lobe
from the neopallium, and thus determine the situation from which the fissure has been
obliterated.
The olfactory tract (fig. 4) ends in tlie vallecula Sylvii in a distinct " tubercle of the
* Gu.slav Jictzius, " Windungon dcs Khincncephalous," Biolog. Unters., N. F. viii. p. 2 (1898). The condition which
he represents in the liodent DasijprocUi, Taf. 7. fig. 10, closely resembles that of the Lemue.
t Haudli. d. .Vriat. u. vergleich. .\nat., licrlin, IS'JO, Taf. o. fig. 2.
49*
330
PROF. G. ELLIOT SMITH ON THE
olfactory tract," wliich Retzius calls "gyrus intermediiis " [op. cU. »upra). To the
mesial side of this tuhercle there is a narrow transverse cleft — the locus perforatus —
between the tuberculum olfactorium and the large optic chiasma.
To rightly ap])reciate the nature of the exposed part of the fascia dentata (fig. 4), the
mesial surface of the hemisphere must be exposed by cutting through the cerebral
commissui-es and lamina terminalis and severing the attachment of the optic thtnlamiis
to the coi"pus sti'iatum (fig. 5).
Fig. 5.
Fig. Ga.
ss. d.
Fig. 5. — Lemur faJrus. The mesial aspect of the i-ight cerebral a.speet. x 2,
Fig. 6. — Lemur fulvua. Left lateral aspect of the brain, x 2.
Fig. 6 a.— Part of the same after removal of part of the caudal opercular lip of the sulcus pseudosylvius. x 2.
Fig. 6 h. — Part of the aame shaded iu order to roughly indicate the area of distribution of the claustrum (c/.). x 2.
Fig. 7. — Lemur macaco. The left lateral aspect of the brain. Natural size.
The hippocampal formation is thus partly exposed as a series of bands ascending
MORPHOLOGY OF THE BRAIN IX THE MA-M>rALIA. 331
approximately vertically from the patch of fascia deatata on the ventral surface to the
posterior end, or splenium, of the corpus callosum.
In the greater part of its extent tliis hippocampal formation conforms to tlio plan
which we may call characteristically mammalian (such as we find, for instance, in the
brain of Tamandua [vide tlie " Brain in the Edentata," op. ait. svpra, iiijs. 10 & 23,
p. 317)]. There is, as in Tamandua, a well-defined sub-splenial flexure, and the wrinkled
band of fascia dentata curves around on the under surface of the corpus callosum in the
characteristic manner and then tapers as it merges in the circumcallosal hippocampal
vestiges. A little hemispherical area of " hi])pocampus nudus " fills up the concavity of
the arc formed by the sub-splenial bending of the fascia dentata, just as I have represented
it in Tamandua [op. cU. p. 320, fig. 23). In all of these respects the hippocampal
formation conforms to the typical mammalian condition, which I have descril)ed in
detail in the above-quoted work.
In a recent memoir Zuckerkaudl has briefly referred to the "hippocampus nudus"
(his " BalkenAvindung ") in the Lemurs *. He describes it as being " zapfenformige "'
in Lemur catta, L. varius, L. monyoz, and L. coronatus, and " kurze zapfenformige " in
Chiromys.
The loAver extremity of the hippocamj)al formation presents a peculiar modification
not found in most mammals. Several years ago I called attention to the peculiar fact
that a part of the hipjjocamjjus is rolled in such a manner that it becomes completely
inverted: the surface which was originally dcej) is then superficial, and clce versa
(Jouru. of Anat. and Pliys. vol. xxxii. p. 28). A strip of this "inverted hippocampus"
separates the fascia dentata from the fimbria in the Lemur, as also happens in most
mammals; but this becomes greatly expanded at its lower extremity to form a "hippo-
campal tubercle," such as is found in Orycleropus (" Tiie Brain \xi the Edentata," op. cit.
fig. 23, p. 315). A hippocampal tubercle of similar constitution occurs in all the Apes,
and has been accurately described in the human brain by Giacomini and Hetzius
(compare Journ. Anat. and Phys. vol. xxxii. pp. 28 & 29).
In the Prosimian brain the tubercle vai-ies considerably in size and prominence, and in
the brain of Nycticebus practically vanishes ; it is never so j)rominent as it is in the
Apes, althovigh its constitution is the same. In some Insectivores, such as Macroscelides,
I have seen a condition closely resembling that of the Lemurs, and have also seen
instances of the same phenomenon in Orycteropus and in the brain of a Manatee. This
peculiar condition is apparently inherited by all the Primates, both Lemurs and Apes,
from some primitive Insectivore-like ancestor, which has also transmitted it to the
primitive Aard-vark and that peculiar Eocene stock which has persisted with little
change as the Sirenia.
The corpus callosum is very long and has a very plump caudal extremity or
splenium, and a very thick and fully-recurved genu. As a whole, it closely resembles
that of the Platyrhine Apes. The greatly attenuated psalterium, or hippocampal
commissure, resembles the common mammalian type, such as is found, for example, in
* E. Zuckerkandl, " Ecitr. /ur An;it. dcs Reichcentrums," Sit/.uiigsli. Akad. Wiss. Wicn, ilatb.-nat. CL, Ed. cix.
Abth. iii. I'JUO.
332 rUOF. G. ELLIOT SMITH OX THE
Tanulndiui, Fclis, or Ovis. It is, however, liardly so plump as in any of these mammals,
and yet it is not nearly so small and attenuated as that of the Apes. In the
Hapalida', for example, the psalterium becomes so extremely reduced {dele Jonrn. of
Anat. and Phys. vol. xxxii. fig. 23, p. 51) tliat Beevor failel to find it even by
histological means. In tlie Lemurs its size is more considerable, and the chief
accunuilation of its fibres is never wholly separated from the splenium by an interval
devoid of commissural fibres, as liappens in 3Iidas {op. cit.) and the other Hapalidse.
The septum lucidum, precommissural area, and anterior commissure present the usual
relations.
Immediately behind the hi^^pocampal tubercle there is a very deep indentation, which
may possibly represent the caudal i-emnant of the rhinal fissure (fig. 5, a). It begins
close to (but without actually joining) tlie hippocampal fissure and passes obliquely
downward and backward. On the base of the lu'ain (fig. 4) it has all the appearance of
ioining the hippocampal fissure. Its extent and dii'ection vary considerably in different
brains of even the same species. In many cases (as, for instance, fig. 5) its dorsal lip
projects downward as a prominent rounded knob overlapping the deepened notch.
Upon the mesial surface of the hemisphere there are a number of very stable and
constant sulci, and usually some otlier more variable furrovrs.
By far the most important of these is a group of three, which may lie called the
calcarine complex. This is a triradiate pattern of sulci situated behind the splenium of
the corpus callosum ; the ventral limb maybe distinguished as " calcarine," the caudal
as " retrocalcarine," and the dorsal as " paracalcarine."
Above and parallel to the corjius callosum there is a sulcus which will be called " inter-
calary " ; and in front of the genu there is generally a " genual sulcus " * of Krueg
(fig. 5).
There is always an oblique sulcus placed below the retrocalcarine sulcus, which for the
present may be referred to simply as b.
There is also a very oljlique sulcus (c) on the caudal margin of the hemisphere, midway
between the furrows a and h.
•That portion of the mesial surface of the cerebral liemis2:)liere which lies below and
behind the corpus callosum is adapted to tlie shape of the contiguous structures. A deep
depression receives the optic thalami, a much smaller and shallower depression below and
behind the splenium is produced by the qnadrigeminal l)odies, and the broad flattened
area behind it is so moulded by contact with the upper surface of the cerebellum.
The calcarine sulcus begins infei'iorly within the deep thalamic fossa just above the
sulcus a ; it ascends for about 3 mm. parallel to the hippocampal fissure and then
bending backward it crosses the rim of the thalamic fossa, traverses the mesencephalic
fossa, and ends in the cerebellar area, about 3 mm. behind the splenium. It attains a
depth of G mm. in its posterior part.
In every brain (with only one exception) belonging to the genus Lemur which I have
examined the calcarine sulcus was continuous Avitli the retrocalcarine sulcus, the two
* It is not altogelbcr certain whether it may not represent the " rostral" suleus.
MOKPHOLOGT OF THE BEAIX IX THE MAMMALIA. 333
rurrows unitiua: at an aiiLi'Ic of 1:^0 . The vetrocalcarine sulcus is coufined to the
cerebellar area. It passes backward wiili a very slight upward inclination for a distance
of S nun. The degree of ()bli(iuity of this sulcus shows considerable variability. In
some eases it is almost horizontal ; in otliers it is removed as much as 30" from that
plane. At its meeting place Avitb tlie ealcarine sulcus it is as deep as the latter.
The paraealcariue sulcus ]);isscs obliquely upward and forward from the angle of
junction of the otlier two sulci almost as far as (in rare cases even actually crossing) the
dorsal margin. In some cases its terminal piece is bent vertically upward. This sulcus
is generally shallower than the ealcarine, so that it appears to cut deeply into the upper
lips of (without actually joining) the sulcu.s formed by the junction of the ealcarine and
the retrocalcarine furrows. In one ease, however, I have seen the ealcarine and para-
ealcariue sulci joined, the retrocalcarine being the separate element. This is instructive,
because it shoAvsthat the reti-ocalcarine sulcus possesses a certain amount of individuality
and is not a mere mechanically-produced extension of the ealcarine sulcus. The sulcus
b varies considerably in length and depth in different brains. In the large brain of a
Lemur rariiis I have seen it represented by a veiy faintly marked depression less than
3 mm. long. In other cases it is a furrow 8 mm. long and 2 or even 3 mm. deejD. It
undoubtedly represents the most caudal element of that composite sulcus of the human
bi-ain known as the " collateral."
The sulcus c is even more variable. It may be a mere pit in the depression caused by
the roof of the bony capsule of the floccular appendage (so-called " petrosal " lobule) of
the cerebellum, or it may become a deep sulcus 5 or 6 mm. long.
On the external aspect of the neopallium the sulci are no less variable (tigs. G & 7).
The deepest and most constant furrow is that which is commonly called the Sylvian
fissure. It really consists of an apparent blending of two actually separate overlapjjing
sulci. One of these, which I shall call pseudosylvian, proceeds vei'tically upward (for a
distance of 4 mm.) from the bend (or more strictly from the posterior limb) of the rhinal
fissure. It ap2)e;u's to (but does not actually) join a second sulcus, which is obliquely
placed, so that the two sulci form an angle of abuut 150'. This upper oblique sulcus I
shall call the " suprasylviau."
If the lips of these sulci be separated, it will be found that the sulcus pseudosylvius
cuts deeply backward into the hemisphere, so that its posterior lip forms an operculum
(I mm. long and almost equally thick) which overlaps a dejjressed area (fig. 6 a). The
lower part of the sulcus suprasylvius overlaps the s. pseudosylvius (fig. (5), and the anterior
lip of this part of the furrow is also slightly operculated. Hence there is an area
(about 6 mm. long and 1 mm. high) submerged and hidden I'rom view by the anterior
opercular lip of the supi'asylvian and the posterior opercular lip of tiie pseudosylvian
sulci (fig. 6 a). The few observers who have in any way referred to this depressed area
regard it as the insula Keilii. IIoll {op. cit. supra) regards it as the submerged
homologue of Leuret's first arcuate gyrus of the Carnivora. The degree of accuracy of
these statements will be discussed below.
Between the upper end of the suprasylviau sulcus and the dorso-mesial edge of the
hemisphere there is a sagittal sulcus of very variable length, wliich may be distinguished
334 PEOF. G. ELLIOT SMITII OX THE
l)y the name "lateral." The auterior extremity of this sulcus always bends laterally, so
as to be approximately parallel to the upper part of the suprasylvian sulcus (fig. 1). Its
posterior extremity usually becomes bent in a mesial direction. It often happens,
hoAvever, that this mesially directed limb remains as a transverse sulcus widely
separated from the sulcus lateralis (fig. 3). It may be distinguished as the postlateral
sulcus.
Behind and parallel to the suprasj'-lvian and pseudosylvian sulci there is a sulcus of
varying length and depth, which may be distinguished as postsylvian.
The mode of termination of the supra- and postsylvian sulci is exceedingly variable
(compare figs. 1, 2, 3, G, and 7). In many cases the upper extremity of the supra-
svlvian sulcus bends backward so as to almost reach the postsylvian (fig. 6) : in
other cases (tig. 7) the latter curves forward to approach the suprasylvian ; and
in other cases again (fig. 3) the intercalated furrow is separated from both sulci.
We may distinguish it by the letter d, and defer the discussion of its significance until
later.
From a point about 5 mm. in front of the middle of the suprasylvian sulcus a furrow
(which may attain a length of 20 mm.) passes obliquely forward and inward to end upon
the anterior pole of the hemisphere. For reasons which will be discussed later, I shall
distinguish this sulcus by the not-obviously apjiropriate name " coronalis." In almost
all cases the posterior part of this furrow is bent backward and inward toward the lateral
sulcus (fig. 2). We may distinguish this little fragment from the rest of the sulcus by
the letter e.
Midway between this sulcus and the mesial border there is a very short furrow, the
direction of which is by no means constant. In most cases, hoAvever, it is directed
laterally and forward. For the present I shall call it the " sulcus y,"
Upon the orbital surface of the hemisphere there is a small sulcus of very variable
size, depth, and shape alongside the olfactory bulb. It is the orbital sulcus. In some
brains it is either quite absent or represented only by a very shallow dej)ression.
Immediately in front of the lower extremity of the suprasylvian sulcus (which is
generally exposed in the genus Lemur) there is often a small oblique sulcus in the
orbital margin. This I shall call " diagonal."
Near the floccular notch (in the caudal margin of the hemisphere) there is generally a
small oblique sulcus, which may or may not extend into the floccular depression and
overlap the sulcus c. It may be distinguished for the present by the letter g.
[In the foregoing enuniciation of theise sulci I have deliberately used term.s which no other writer has employed
in reference to the Lemur's brain, not only to avoid the hopeless confusion which the varied nomenclature of
Eeddard, Ziehen, Chudzinski, and Flatau and Jacobsohn wouki entail, but also for morphological reasons, which will
appear later and, I believe, justify this course of action.]
In different individuals of the genus Lemur the cerebral hemisphere exhibits slight
variations in size, shape, and the arrangement of its sulci.
The only writer, so far as I can remember *, who has even so much as hinted at the
* As most of the literature is not at present available, I am unable to consult the memoirs of Gervais in regard
to this point ; but, unless my memory belies mc, this statement is correct.
MOEPHOLOGT OF THE T5RAIN IX THE MAMMALIA. 335
possibility of these differences having any specific value is Beddard *. His remark that
"the bigger brains are on the whole more complex than the smaller" may seem a mere
truism; but I have found some noteworthy exceptions to it. Tlius one of the simplest
Lemur's brains I have seen Avas also l)y far the biggest : it belonged to a Lemur variiifi
and weighed 39 grammes, its cerebral hemispheres being each It) mm. long and 21-")
mm. broad ; it had no fronto-orbital sulci, very poorly developed orbital and collateral
sulci, and unusually short lateral, parallel, and .straight sulci (fig 3).
My examination of crania bears out llio statement of Beddard that the variety
nnjuanends has the simplest brain of all members of the genus. It is the only brain in
Avhich no trace of the sulcus / is found. But with this (possible) exception, all of
Beddard's remarks on supposed specific characters are the mere record of individual
variations, every one of which may occur in any species.
So far as my ob.servations go, it is quite impossible to assign any specific value to the
A-ariatious. But this may possibly be due to the fact that an insufficiently large number
of brains was available for examination. As the variations are practically AvhoUy confined
to the external aspect of the hemisphere, the investigation necessary to settle this
question can most readily be carried out on a series of skulls, from A\hich we can readily
de,termine the size, shape, and pattern of the sulci ia tlie cerebral hemisphere.
Concerning the exact size and shape — /. c chiefly the amount of cerebellum ovei-lapi)ed
— there are only very scanty data.
Max "Weber records the brain-Aveights of two specimens of Lemur variiis, Geoff. ; that
of a male 33 grms., being x^ of the body-Aveight, and that of a female 287 grms., being
tV of the body-AA^eight. A representative of this species died in the Ghizeh Zoologicjil
Gardens, and I found its brain-weight to be 39 grnis. and its body-Aveight 2900 grms.,
■i.e. the brain Avas J.f of the body-weight.
Max Weber also records the weight of the 1-raiu and its relation to the body-weio-ht
in two females of Z^W7?;- moiifjo:, L., as 28 (-^;) and 21-1 (,;iy) respectively. The only
other records of brain-Aveights by Ziehen, Flarau and Jacobsohn, are of little A^alue,
because they refer to s])ecimens the Aveights of Avliich were altered by the action of
reagents. I have found tliat the extent of tiie area of CL'rebellum uncoA-ered by the
cerebral hemisphere is subject to variation. It A\as largest m a brain of Lemur carlu^
(fig. 3), but, as I have found tlie extreme types of variation in the brains of two
specimens of Lemur fulrun (which died in the same Aveek and were examined under
exactly similar conditions), it does not seeni possible to attach any specific importance
to this fact. (Compare figures 6 and 7, also figures 1, 2, and 3.)
With regard to the variations in the uieasuiements of the hemisphei'es, my data arc
A'orv scantA".
Flatau and Jacobsohn give the following measurements for a specimen of Lemur
macaco {op. cit. p. 174) : —
Maximum length of cerebral hemisphere 18 mm.
Greatest breadth of cerebrum 38 mm.
Greatest height of hemisphere 28 in.n.
» " Oil the liriiin in the Lcnuiis," I'ruc. Zool. Soc. 1895, p. 143.
SECOND SERIKS. — ZOOLOGY, VOL. VII I. 50
;;:36 peop. g. Elliot smith ox the
These measurements are extraordinarily lari^e "when it is considered that the hrain
weighed only 26'3 grms., after heing in 10 per cent, formol solution for eight days.
The corresponding measurements in an adult female L. macaco I found to he
t3 mm., 37 mm., and 2C mm. respectively. These measurements (which were checked
hy comparison with the cranium) indicate a brain-shape very different to that of Elatau
and Jacohsohn's specimen. If the latter were swollen as the result of the formalin, the
value of the elaborate tables of measurements in their book is seriously discounted.
The hemispheres of a Lemur fiilvtis were 46 mm. long and 38 mm. broad ; and the
corresponding measurements in a Lemur varius were 49 mm. and 43 mm. respectively.
The size of the brain of the extinct Lemur Jullyi greatly exceeded that of any of the
recent Lemurs; for its cerebral hemispheres were 59 mm. long, 48 mm. broad, and
31 mm. deep.
Turning to the consideration of the cerebral sulci and fissures, a considerable amount
of variation is found.
The rhinal fissure usually becomes aborted, with the exception of a very small furrow
near the olfactory peduncle and the apparent recurved inferior extremity of the pseudo-
sylvian sulcus. In the specimen of Lemur fulvus described above, the greater part of
the fissure persisted. In another specimen of L. fulvus, and in one each of L. macaco
and L. varius, I have seen a faintly marked furrow representing the whole fissure.
The two brains of L. varius (one in the College of Surgeons and one from Ghizeli)
agree in the following respects : — The diagonal sulcus is absent ; the orbital sulcus is so
poorly developed as to be a mere insignificant depression ; the lateral sulcus is a simple
regular arc, widely separated from the postlateral and coronal sulci ; the sulcus d is not
joined to either the supra- or to the postsylvian sulcus ; the collateral sulcus is very small
and shallow ; and, apart from the small floccular sulcus, there is no sulcus behind the
postsylvian.
All of these features may be regarded as indicative of a tendency to simplicity, which
is all the more significant in that the brain in tliis species is appreciably larger than that
of most other members of the genus. It remains for future research to determine
Avhcther these features may be regarded as distinctive of the species varius.
I have examined two brains of Lemur macaco and several cranial casts. In addition
Platan and Jacobsohn [op. cit.) have given a full description of a brain of this species.
The diagonal and orbital sulci are constantly present, but the latter shows a tendency
to become irregular by the development of a lateral ranuis. Sometimes also its direction
varies. The mode of ending of the upper extremities of the supra- and postsylvian
sulci is variable. In most cases the upper extreinity of the suprasylvian sulcus is
simple ; but Platan and Jacobsohn represent (Taf. 3. fig. 1, -^j a small added horizontal
sulcus. Generally the upper end of the i)arallel sulcus curves forward, representing a
fusion with the sulcus d, which usually does not exist as a separate element in this
species. Unlike the condition found in L. varius, there is no distinct postlateral sulcus,
but there is a very ol)lique furrow, united at a very obtuse angle with the caudal end of
the lateral sulcus.
1 have never seen the sulcus described as " temporalis sccuiidus (oberer Teil) " by
MOKPllOLOGY OF TUE BRAIN IX THE MAMMALIA. 337
Platau and Jacobsolm in the brain of Lemur rariiis : nov have I ever found the sulcus
labelled "Nebenfurche " (Taf. 3. fig. 2, 2) in the l)raiii of any Leinur.
In the brain of Lemur fnlvus the diagonal sulcus is genei-ally well-developed and often
deeply notches the supraorbital margin.
The lateral and coronal sulci approach much more closely than is the case in the
other species, and in some instances they even seem to join. Tlius Beddard says (Proc.
Zool. Soc. 1895, p. 113) : — " In the brain of Lemur monyoz the angular [lateral ' and infero-
frontal [coronal] sulci are completely continuous.'' But even if this sometimes happens,
it is by no means the usual condition in L. fnlcus. Por, as the same writer remarks
(p. 113), this does not occur in L alblfrous, nor iu L. iiigrifroi/s nor L. (uijuaneiisis, all of
Avhicli may be included in the s-pecmsfulviis. ^Vithin this si)ecies all varieties of the upp(>r
ends of tlie supra- and postsylvian sulci are found. Sometimes a long arcuate sulcus d
is fused at one end to the suprasylvian sulcus and almost joins the postsylvian at the
other. Not infrequently the sulcus d is joined to the postsylvian and not to the supra-
sylvian ; but in some cases it remains independent of botli, ;is in tlie .-^pecies rari/is.
In most cases the caudal end of the lateral sulcus behaves as it does iu the species
macaco, and not as in rarius.
In the two specimens of Lemur ca/la which I examiiu'd thei'c was nothing distiuetivc
and they might well have belonged to the species fulrus.
The skull of Lemur rubrivetiter, Geoff., exhibits ridges corresponding to typical supra-
sylvian, postsylvian, lateral, postlateral, coronal, orbital, aud diagonal sulci, witii nothing
indicative of a distinct specific type.
Tlie large brain of the extinct Leinur Julli/i [Pahechirogaleus JuUtji, G. Grand.)
exhibits the tyj^ical Lemurid plan of sulci with diagrammatic clearness.
The suprasylvian sulcus with a long deejily-incised inferior exposed limb, the shallowei-
furrow indicatijig the pscudosylvian sulcus ; simple, long, linear orbital and diagonal
sulci; lateral and coronal sulci which almost (if not quite) meet as \\\ Lemur f ulcus;
and a .small sulcus f, all conform to the most typical Lemurid pattern.
The or])ital sulcus presents none of the irregularity which it has ao-quiivd iu most of
the recent Lemurs.
The Cerebr.\l Hemisphere in the Genus Hapalemuk.
An examination of tiie ridges on the interior of tlie ci-aiiial cavity of Ilapalemur
reveals a pattern of sulci which does not appreciably differ from that exliibited in tlic
genus Lemur. Tlie typical Sylvian fissure, a sliort lateral sulcus, a long coronal sulcus,
and a typical linear orbital sulcus are present.
Milne-Edwards has pu])lished illustrations of a brain of Ilapalemur simus * which
presents similar features. Beddard has described three brains belonging to this genus,
two of the species griseus and one of the species simus ■f. The hemispheres of the
smaller specimen of the former species were 33 mm. long and 2G mm. broad, the larger
'Histoire de Madiiga.scar,' (2) tome vi. pi. 122*. figs. 1. 2, '.i, 4, and .3.
Proc. Zool. Soc. lyul, pp. 127 & 12S ; also I'iol'. Zool. Soc. 18!)1, p. 4.J7.
50
*
338 PEOF. G. ELLIOT SMITH ON THE
being 35 mm. long and 2S mm. broad : the cori-esponding measurements •♦in the species
simits were 40 mm. and 31 mm. respectively. The brain of Sajjalemur therefore is
considerably smaller than the average size in Lemur.
In the braia of Hapalonur simus (Proc. Zool. Soc. 1901, p. 127, text-fig. 24) the upper
end of the suprasylvian sulcus is bifid; but it is not fused to a long sulcus d, as is the
ease in Beddard's first specimen of the genus, griseus {ibid. p. 128, text-fig. 25). There
is a typical coronal sulcus in both species, bent in exactly the same jnanner as so often
liappeus in the genus Lemur. Lateral, orbital, parallel, and even diagonal sulci and the
I'urrow _/ occur exactly as in the genus Lemur.
It is clear that the brain of Hapalemur is, to all intents, identical with that of Lemur ;
and the features which Beddard thinks may possilily amount to specific characters can
be nothing else than variations such as we find in almost every species of Lemur.
H. O. Forbes states* that "the brain [in the genus Hajxtlemnr] is narrower^ and
shallower than that of the genus Lemur, and presents no specially close resemblance to
the same organ in the Lndrisirue or the Lorisiiice." With the latter part of this quotation
I quite agree ; and if the former part is also true, it indicates, in comparison with
Lemur's brain, a slightly nearer approach to the pithecoid type.
The Cerebral Hemisphere in the Gends Lepidolemur.
The olfactory bulb appears to project beyond the hemisphere as it does in the Lorisinag,
the Galaginiie, and in Tarsius.
The pattern of sulci which a study of the cranial ridges in a series of skulls has
shown to exist in Lepidolemur differs in many respects from the arrangement which
is common to the genera Lemur and Kapalemur.
Fig. 8.
see. si.
Lepidolemur in usteliit.ua.
Lateral aspect of the cranial cast of the left cerebral hemisphere. Xat. si/.e.
In a skull of Lepidolemur musteliuus iu the British Museum there are ty^jical simple
linear Sylvian, lateral, orbital, and a very shallow postlateral sulcus, and another sulcus
apparently representing the conjoint elements e and /, such as often hap2)ens in Pro-
piihecus and Fcrodicticus. This specimen also closely resembles that of one of the
specimens of Avahis which I examined, the protruding olfactory bulbs coustitutino- a
small and relatively vmimjjortant difference.
A second skull of the same species exhibited a pattern of sulci exactly similar to this,
* '• Monkeys," vol. i. (Allen's Naturalist 'a Library, 1893), p, SI.
MOEPHOLOGT OF THE BRAIN IN THE MAMMALIA.
339
except that no trace of a ridge corresponding to tlie postsylvian sulcus was present.
There is neither a coronal nor a postlateral sulcus.
In a skull of Lepidolemiir leucopus in the British Museum there is no trace of eitlier
the orbital, coronal, or postsylvian sulci, or of the furrows e and /' in Lemur.
So far as I am aware, all that is known of the brain itself in this genus is represented
by a series of not altogether satisfactory drawings (without any description of any kind,
except the legend " brain of Lepidolemur ruficaudatus ") in Milne-Edwardss work *.
From an examination of the profile (fig. 1 b), one iniglit imagine that typical coronal
and lateral sulci of the Lemurid type were present ; but the appearance presented l)y
the sketch of the dorsal surface (fig. 1) is so different that it is difficult to believe that it
represents the same brain. As I had inferred from the examination of two of the three
skulls, there is no postsylvian sulcus.
These drawings unfortunately give us little information of any value.
So far as I am able to judge from the crania which I have examined, the brain must
present many features resembling those of the Indrisiiise and others (of less importance)
like those of the Lorisin;B (especially Ferodicticus) and Galagime.
F.g.9.
y. sp
Fig. 9. — Nycticehus tard'njradus. Dorsul aspect of a cranial cast. Nat. size.
Fig. 10. — Si/ctictbus tardiijradiuH. Left aspect of a brain. 2^at. size.
Fig. \\.—yijctictbus tardigradus. Lateral aspect of the left cerebral hemisphere of another b
rain. Nat. size.
The Brain ix the Lorisin.e,
It is convenient to consider Nycticehus first. The brain in this genus is not only
smaller than that of Lemur, but also presents some noteworthy difi'erences in shape.
The cerebral hemispheres of the only fresh specimen of yycticebus tardigradus m hich
I have seen are 34. mm. long, 28 mm. wide, 23 mm. high.
• Uistoire de Madagascar,' (2) tome vi. 43- fasc. 1S97, pi. 273. figs. 1, 1 a, 1 5, 1 c, and 1 d.
340 PKOF. G. ELLIOT SMITH OX THE
If a vertical line he drawn from the most prominent part of the " natifoi'm
eminence" of the pyriform lobe, it Avill he found that more than half of the hemisphere
lies behind the vertical line. This is not the case in Lemui-, but is so in all Apes.
There is a more pronounced caudal extension of the hemisphere, so that the cerebellum
is almost completely overlapped.
There is a much smaller " orbital area " in Nycticehus than in Lemur. In the latter
it extends in the caudal direction as far as the pseudosylviau sulcus, whereas in the
former it only extends to a point midway between the pseudo.sylviau sulcus and the
anterior jwle of the hemisphere.
In the genus Lemur the olfactory bulb is a horizontally flattened jdate, the bulk of
which is covered by the hemisphere. In the genus Ni/clicebiis the bulb is a vertical
sagittal plate, 5 mm. of the length (9 mm.) of which projects beyond the anterior pole
of the hemisphere. It is only 1-5 mm. broad (/. e. transverse thickness).
The anterior rhinal fi.ssure is not obliterated : of the posterior rhinal fissure only one
or two millia:ietres exist as a furrow, but if a i'resh brain be examined a very faint
depression is visible extending horizontally backward Avith a slight downward Ijend.
This can be traced on to the mesial surface.
The flexure of the ])yriform lobe is slightly more acute and pitliecoid than that (jf
Lemur, so that the vallecula Sylvii is more cleftdike. Tiie other parts of the rhiuen-
cephaloa closely resemble the corresponding regions in the genus Lemur, with the
exception of the lower end of the hipj)0campus. Eor in Nycticebus, iinlike every otliei-
Prosimian known to me, there is no well-defined hippocamiml tubercle. The inverted
hippocampus merely becomes slightly broader at its lower end, as it does in most
mammals (fig. 12).
6 re.
I.
fh.
Kyeiicehus tardigradns.
Mesial aspect of the right cerebral hemisj/here. Kat. size.
The corpus callosum is a very thin plate of fibres 12 mm. long; its anterior and
posterior parts are not so markedly thickened as they are in Lemur. There is an
exceptionally extensive arcuate genu.
The form and relations of the calcarine group of sulci are identical with those
described in Lemur. The paracalcarine sulcus is, however, much shorter. The
intercalary sulcus is relatively larger than it is in Lemur, and its posterior extremity
is slightly bifid : the upper limb being the deeper faintly foreshadows the upturned
posterior end of the calloso-margiiial sulcus of the Anthropoidea. There is neither
a genual nor a rostral sulcus.
.MORPHOLOGY OF THE BRAIX IX THE :MA:\r^rALIA. 341
The Sylvian fissure is composed of Iwo separate sulci, pseudosylvian and suprasylvian,
:\s in L('it//tr; ])ut in this g-cniis tlicy l)Otli pursue the same direction and overhip so
slightly that it requires the closest examination to detect the lowei" end of the supra-
sylvian sulcus. As the result there is no ohvious depressed area, such as is found in
Lemur, hut its homologue is found in tlie anterior lip of the furrow in the region of the
lower end of the suprasylvian sulcus. Tiie lower })art of the pseudosylvian sulcus
is very shallow.
Tlie up|i('r extremity of tlie supi-asylvian sulcus hecomes directly continuous
with the lateral sulcus, no suhmerged gyrus heing found to separate the two
elements.
The lateral sulcus hends hackward, and then rinally curves inward in a manner exactly
analogous to that of Lemur J'ulcu.s.
There is a small bracket-shaped sulcus J\ the concavity of wjiich faces the middle line.
There is a sulcus about 7 mm. lung in front of and parallel to the suprasylvian
sulcus. Its ujiper part is bent backAvard so as to become almost horizontal. Thus the
sulcus may be regarded as consi.>ting of two limbs x and y. There is always a small
sulcus just above the orbital margin which we may regard as the sulcus coronalis. In
my type specimen this is so small as to be little else than a mere pit.
There is a simple linear postsylvian sulcus. In my specimen (fig. 10) there are no
other sulci, nor even depressions to represent the sulci called a, b, c, and g in the brain
of Lemur.
In the specimen of Nyctlcehm turdigradus [Siciiops javcuiicus, Illig.] described by
Flower (Trans. Zool. Soc. vol. v. pp. 103 et seq.) there are some slight differences.
Thus tlu; sulcus y appears to be absent and x is joined to the coronal sulcus, the
resultant furrow being called " iufero-frontal," which is the name Flower gives to the
coronal sulcus of Lemur. The sulcus /is represented as a long sagittal furrow, which is
called " supero-frontal " (]). 101). The writer remarks that the Sylvian fissure " makes
a slight change in direction, continuing to extend backwards " (p. 101), but thinks that
the lateral sulcus of the Lemur is absent.
He represents a well-developed postlateral sidcus, such as is not found in my specimen.
Concerning this he says : — " A small sulcus running in a transverse direction on the
upper surface of the hemisphere, at the junction of the parietal and occipital lobes,
recalling the ' temporo-occipital ' (scisstire perpeJuUculaire exlerne) of the Quadrumana,
which iissure is, howevei-, Avanting in the smaller American Apes, as in Lemur " (p. 101).
We have seen above, however, that it sometimes occurs in Jjcmur, especially in the
species varius ; and even when absent as a separate element it is probably represented in
the " tail" of the lateral sulcus, both in Lemur and NijGlicebus.
h\ the specimen in the College of Surgeons Museimi (fig. 11) the postlateral sulcus
is a separate element on the left hemisphere, but is merged in the lateral sulcus on the
right side.
The upper end of the postsylvian sulcus bends forward, as it does in many Lemurs.
On the right hemispliere it presents an additional ramus parallel to the lateral sulcus.
In both hemispheres there is a little sulcus behind the upper end of the postsylvian
312 PEOF. G. ELLIOT SMITH OX THE
sulcus. In an Ape's hrain this would be called " infei-ior temporal." In this brain the
sulci X and ij appear to be joined to the coronal to form a triradiate pattern.
Tliere is also a small collateral sulcus (fig. 12, b), such as the brain of Lemur presents.
Beddard says that " there appear to be more differences between the brain of
[Ni/eficebits tardigradns] and that of its congener Nyctlcehus javanicus than loetween
the different species of Lemur .... the brain is more rounded in front, and the
cerebellum more fully exposed than is indicated in Sir W. Flower's figure of iS'.jaoanicus.
The angular fissure [Beddard thus refers to tlie sulcus designated / in my account and
mistakes it for the ' lateral '] is short and instead of l)eing straight it is crescentic . .
the concavities facing each other. . . Tlie infero-froutal [coronal] seems to be less
conspicuous tlian in the other species of the genus and on one side of the brain it
ran back to join the curved (anteriorly convex) presylvian fissure [tlie sulcus .r], the
homologue (?) of which latter in N. javanicus lias a totally different direction. The
])arieto-occipital ('Simian') fissures [postlateral sulci] are better developed than in
N. javanicus and reach the intercerebral sulcus" (Proc. Zool. Soc. 1895, p. 111).
The differences in shape which Beddard regards as specific features I have been
unable to find upon exaniining a series of crania, and the features of the sulci upon
which he lays stress are individual variations such as also occur in undoubted examples
of the species tarcUgradus. So far as one can be guided by the evidence of cerebral
anatomy, there is no reason for splitting up the species tardigradus to make another
species {javanicus).
There can be little doubt that the brain which Chudziuski described under the title
" Loris (jrocilis " (Bull. Soc. d'Anth. 1895, p. 43G) is really that of Nyciicehus tardi-
f/radus. It ])resents the arrangement of sulci typical of the latter ; and this is very-
different from that of Loris. The Sylvian fissure is joined to the lateral in the typical
manner, and the latter ends in a bifid extremity, which Chudziuski calls " portion externe
de la scissure occipitale " ; this is the postlateral sulcus.
The sulcus/, the postsylvian and the triradiate complex of the sulci .r, y, and coronal
resemble those described above in the specimen belonging to the College of Surgeons.
There is a small sulcus behind the upper end of the postsylvian.
Ziehen based his account of the bj-ain in this genus (Arch. f. Psych, xxviii.
pp. 902-906) on four specimens (one of Avhich is in the College of Surgeons, and another
in the British Museum). He calls the mesially-bent extremity of the lateral sulcus the
" Eiidiment der Fissura parieto-occipilalis lateralis."' [In respect to this sulcus the
condition represented in his drawing lesembles my type si)ecimen.]
The sulci which I have labelled/', x, and y he regards as representatives of the sulcus
centralis.
The sulcus behind the iipper end of the postsyl\i;ui (his "temporalis superior")
is called " temporalis inferior." Another small sulcus behind the loAver part of the
postsylvian, which I have not seen in any of my specimens, is regarded as the lower
part of the same sulcus.
He represents a small separate furrow above the postsylvian, such as is olten i)r( sent
in the Lemurs {d) ; and the connecting limb of the supra- and postsylvian sulci is also
]\I01?PH0L0GT OF THE BRAIX IX TITE .MAMMALIA. 343
shown. This lie hihels " Schoitelstiick zii 0 [postsylvian]." On the mesial surface
ho represents, in addition to the calcarine group and the intercalary, a small rosti-al and
two sulci labelled /3' and 7. Of these, j3' obviously represents the sh'ght depression
produced by the capsule of the floccular lobes, wliereas y represents tlie sulcus 1 liave
called " collateral " in Lemur.
Like Flower, lie denies the presence of an iiisuhi (p. DOG) ; l)ut this arises from a
misconception as to the nature of tlie region so-called. Both writers mean that
a definite subm(>rged area is not found in this genus.
Platau and Jaeobsohn repeat the curious error of Chudzinski and descril)e a brain,
which CAW be none other than that of Nycticebus tardigrddus, as being that of " Stenops
gracilis." For not only does it exhibit the plan of sulci distinctive of Ni/cticebus, but
its dimensions also are those of this genus and not those of Loris \_Stenoj)s]. If the
brain which they describe really belongs to the genus Loris, we should have expected
some comment from the authors as to its aberrant characters, w'hich a comparison of the
memoirs of Beddard and Ziehen ought to have demonstrated. [Moreover, a study of
their account (' Ilandbuch,' last few lines on p. 191- and p. 195) might lead one to suppose
that the authors do not clearly distinguish between the terms Loris gracilis, Stenops
gracilis, Stevops lardigradus, Stenops javanicus, and Nycticebus tardigradus.^]
The condition of their specimen differed from that of mine in possessing a separate
postlateral sulcus and an inferior temporal. In these respects, as w^ell as in the features
of the mesial surface, it resembles the specimen in the College of Surgeons.
The sulci which Ziehen called " central " the above authors regard as " precentral."
Otherwise they closely follow the teaching of the Utrecht psychologist.
They represent certain small furrows, which have not been mentioned above, such as
those labelled " Spur eines Sulcus fronto-orbital " and " Fissura sublimbica posterior."
In addition to these records, I have examined two crania of this genus, and from one
a plaster mould of the brain-cavity was made.
In one specimen (College of Surgeons, specimen 293, labelled "ior^ y«y««eCT<s")
I found characteristic snprasylvio-lateral, postsylvian, and orbital sulci, the triradiate
coronal -\-x-^y complex, and the sulcus/. In addition there was a separate postlateral.
There seemed to be a dingonal sulcus between the sulcus x and the Sylvian complex,
but of this I cannot be certain.
The other specimen, ^^ hieb was given me by Captain Stanley Flower, presented a pattern
which differed from the last in the absence of any trace of a diagonal sulcus ; it also
possessed modifications of the upper ends of the postsylvian sulci in the two hemispheres
exactly analogous to those found in the respective hemif^pheres of the brain in the
College of Surgeons. Thus the upper end of the sulcus was bent forward on the lel't
liemisphere ; but bifurcated, and its chief branch bent back on the right side. There is
also an inferior temporal sulcus.
It is interesting to note in these two casts that there is a definite postlateral sulcus in
all four hemispheres, and in none of them is the caudal extremity of the lateral sulcus
bent inward, as is invariably the case w^hen the separate postlateral sidcus is absent. A
comparison of the brain-case of my type specimen with the other cranium from Ghizeh
SECOND SERIES. — ZOOLOGY, VOL. VIII. 51
344 VROT. G. ELLIOT SMITH ON THE
shows that the separate postlateral sulcus occupies the same position in regard to the
skull as the mcsially l)ent end of the lateral sulcus of the other specimen.
The brain of PerodlcUcus presents some very interesting features.
The olfactory parts of the brain resemble the corresponding regions in Nyctlcehas.
The rhinal fissure resembles that already described in the latter genus.
ho- so- fr-
Perodictirits potto.
Left lateral aspect of the br<ihi. Xat. size.
The " Sylvian fissure " is composed of the pseudosylvian and suprasylvian sulci as in
Lemur, but the latter furrow is not joined to the lateral sulcus as it is in Nycticebns.
The lateral sulcus is a simple arc surrounding the upper ends of the suprasylvian and
postsylvian sulci. Tlie latter is bent forward at its upper end, as often happens in
Lemur. There is a typical well-developed orbital sulcus and also a well-defined coronal.
The most interesting feature of this brain, however, is an extensive transverse furrow^
placed between the coronal and lateral sulci in exactly the same maniier as the sulcus
centralis occurs in its most primitive form in the Apes. This sulcus is obviously
formed by a blending of the sulcus / of the Lemurs with a sulcus such as I called x in
Nijclicehus ; in the genus Lemur this sulcus is probably represented Ijy the bent piece c
of the sulcus coronalis. The mesial surface of this hemisphere resembles that of
Nyctlcebus.
]^eJdard described a brain of Perodiclicus (Proc. Zool. Soc. 1895, pp. 144 & 145) which
presented features essentially identical with that in the College of Surgeons. His account
is marred by the fact tliat he failed to recognize the lateral [liis "angular" svdcus] in
Nycticehus ; and most of his desci'iption of the brain of Ferodicticus, which is concerned
with this ])()int, is con.sequently valueless. He also quite needlessly confuses his account
by the misuse of tlie word " presylvian," which he applies to the sulcus which I have
tentatively suggested calling "central." The postsylvian sulcus bends forward as in tlie
last specimen.
Ziehen (Arch. F. Psych, xxviii. p. 901) describes tlie same brain of Ferodictlcus as
myself; but as I had to remove the arachnoid and })ia membranes before it was
possible to stiuly the arrangement of the sulci, his account can hardly be regarded as
an accurate description. His illustration (hg. 3) bears very little resemblance to the
specimen in the Iloyal College of Surgeons, and is obviously adapted from one of
Beddard's drawings {op. clt. tig. 2i5, p. 145). Ziehen quotes tlie writings of Beddard,
van der Hoeven, and van Camper in reference to the brain in Lerodicticus.
An examination of the ridges ujwn the inner walls of the cranium of a specimen of
MOEPHOLOOY OF THE BEAIX IX THE MAMMALIA. 345
I'crodictieus (Royal College of Surgeons, no. 2i)5) reveals a pattern identiciil with that
seen in the actual brain, which T have just described.
In the little brain of Loris (jracHis the olfactory parts closely resemble those of the
other two genera of Lorisinge. The upper ends of the supra- and postsylvian sulci
approach very near to one another on the left hemisphere ; on the right hemisphere the
former is prolonged into a hook surrounding the upper end of the postsylvian. There is
a simple arcuate lateral sulcus and a small postlateral. There is a small oblique orbital
sulcus ; and a shallow furrow placed far forward may represent the coronal sulcus.
so. -'P-
Litns r/r<icif<s.
Lateral aspect of the left cerebral hemispljerc. Nat. size.
There is also a small inferior temporal sulcus on the left hemisphere, as in the brain of
Nyeticehus. On the mesial surface the typical calcarine group and shallow intercalary
and collateral sulci are found.
Beddard examined the brain in two specimens of Lo7-is f/raci I is, and stated that "there
are only three fissures plainly visible — the Sylvian, the angular [lateral], and the antero-
temporal [postsylvian] " (Proc. Zool. Soc. 1895, p. 145). [His representation of the
brain (fig. 3) is, however, so studded with innumerable spider-like figures as to be
([uite unintelligible.] In one of his specimens there was a separate postlateral [his
" parieto-occipital "] sulcus on one hemisphere.
Gervais's illustration (Journ. de Zoologie, t. i. pi. 2. fig. 2) is useful as a demon-
stration of the actual size and shape of the brain in Zoi-is ; but Beddard's criticism
of his representation of the sulci is quite justified. Among the Lemurs it is possible, if
the investigator has studied the effect of brain-form on the cranial wall in some members
of the Order, to foi-m a very good idea of the configuration of the actual brain from an
examination of the brain-case. But Gervais has not exercised sufficient caution in the
representation of the plaster casts, and gives an altogether erroneous idea of the
configuration of the brain.
Ziehen's account of the brain in this genus was based upon the specimen in the Royal
College of Surgeons and another specimen of his own. He represents the lateral aspect
of the left hemisphere (Arch. f. Psychiat. xxviii. iig. G, p. yos). The Sylvian fissure,
orbital sulcus, a very short postsylvian sulcus and a " Scheitelstiick " above it, a lateral
and po.stlateral sulcus are all shown ; but in addition there is a transverse sulcus labelled e,
not present in Beddard's specimens nor in that in the College of Surgeons. The letter
e is used by Ziehen to designate the central sulcus in Pcrodiclicus, and both the sulcus
wiiich I call/ and that designated .r in Nijcllcehns.
51*
;34G
PEOF. G. ELLIOT SMITH ON THE
The Bhain in the Galagin.e.
The lirain of Gulago Guruetti may be regarded as the simplest and most generalized
among the Lemiivs, if Ave except that of Tarsiiis, which exhibits a peculiar combination
of primitive and pithecoid features imjierfectly disguised by its strangely-distorted form.
The features of the rliinencephalon in Galar/o resemble on the whole those of the
I-orisime ; but the olfactory bulb is jjlumper and the rhinal Assure is more complete than
it is in any other genus among the Primates.
The brain of this species is remarkable (among those of Lemurs of a corresponding size)
for the paucity of sulci. Only the most stable and constant sulci are present, and these
in a typical and clearly defined form (tig. 15).
Fig. 15.
(JaJago Oarnetti.
Lateral aspect of the left cerebral hcmi.sphere. Xat. .size.
The " Sylvian fissure " is constituted like that of the LorisinsB, no submerged area
being found between its two constituent sulci. The pseudosylvian sulcus is very
shallow.
Above the upj^er end of the suprasylvian (Sylvian) furrow there is a very short latei-al
sulcus, which is quite straight in one brain, but slightly arched in the other.
Ihere is a simple, straight, clean-cut orbital sulcus parallel to the anterior lateral
fissure.
Fig. 16.
spc-
Galago Garaetli.
Mesial aspect of the right cerebral hcmisiilierc. Xat. size.
In one of the two brains of Galago Garnetll there is a shallow depression far forward
on the hemisphere just above the orbital margin : it represents the coronal sulcus.
It resembles the brain of Lejndolenmr in having no well-defined postsylvian sulcus ;
but on the left hemisphere of one specimen there is a faint depression in the situation of
this furrow. This we have seen to happen sometimes in Lepkloleimir.
MORPHOLOGY OF THE BRAIX IX THE MAMMALIA. 347
The typical calcarine group of sulci closely resembles that already described ia Nucll-
cebus. There is a simple linear intercalary sulcus above the anterior two-thirds of the
corpus callosum. In one brain there is also a very small oblique rostral sulcus placed far
forward on the mesial surface of the hemisphere.
The hippocami)al formation presents the typical features, with a large hippocampal
tuliercle and distinct circumsplenial vestiges.
The cerebral hemisphere is 30-5 mm. long, and its posterior extremity is drawn out
into a distinct caudal (or occipital) process. The corpus callosum is 13-5 mm. long, and
its splenium is 11 mm. in front of the occipital pole of the hemisphere. The maximum
depth of the hemisphere is 195 mm.
Althougli the brain of Galago vrassicaudata is only a little larger tlian that of
Garnett's species, there is a noteworthy increase in the number and extent of the sulci.
This fact becomes more interesting and significant when it is observed that the pattern
mapped out by the new sulci is identical with that found in the genus Lemur.
The characters of the cerebral hemisphere of the Great Galago may thus be summed up
by saying that it is a brain of the same shape as (although slightly larger than) that of
Garnett's Galago, with the sulcal pattern of a Lemur.
The upper extremity of the Sylvian fissure bends slightly backward and approaches the
upper end of the postsylvian sulcus, as so often happens in the genus Lemur. The post-
sylvian sulcus is represented in the brain in the Royal College of Surgeons merely by a
G(iJ(ir/() crassicdidhitu.
I.aleiiil aspect of tbo left cerebral hemisphere. Xat. size.
short furrow corresponding to its upper end, whereas in Beddard's specimen* it is the lower
end of the sulcus which is deepest. In the skulls of this species which I examined
there was a low ridge representing a sulcus as extensive, though not so deep, as that of
Lemur. The postsylvian sulcus must be regarded as poorly developed in all the
Galaginae. Another instance of this has already been seen in the genus Lepklolemxr.
In Beddard's specimen the lateral sulcus (which he calls "angular") is represented as
a short linear furrow, like that of Garnett's Gtilago. But in the actual brain (and
also in a skull of this species labelled " Otogale Kirkil " in the British Museum) i have
found evidence of a much more extensive arcuate lateral sulcus surrounding the upper end
of the Sylvian fissure. In the brain in the Royal College of Surgeons the posterior end
of this sulcus is bent mesially ; and at a short distance behind it there are Iavo shallow
* Proc. Zool. Soc. 1895, p. 14G, fig. 4.
34S
PROF. G. ELLIOT SMITH ON THE
transverse furi'ows (in Bedclard's specimen there is only one, whieli is labelled " parieto-
occipital ") representing the postlateral (transverse occipital) sulcus. The anterior end
of the sulcus extends downward in front of the Sylvian fissure for a short distance, so as
almost to reach the caudal end of a sagittal sulcus (coronalis). The latter is bent in an
ol)tuse angle in the brain I have examined ; in Beddard's specimen it is apparently
represented by two separate sulci (labelled "infero-frontal ") in the same straight line ;
and in the skull of Kirk's Galago there is a ridge representing an extensive straight
sulcus directed obliquely forward and mesially, as in Lemur. Above the sulcus coronalis
tliere is a faint depression (/), such as is found in Lemur.
On the mesial surface the sulci are grouped as in Garnett's Galago. There is,
however, a sulcus a.
from an examination of the skull of GaJago 31onieiri (lloyal College of Surgeons,
specimen No. 285, Osteological Series), I can definitely state that the brain is of
the same shape as that of Garnett's Galago, and presents rhinal and Sylvian fissures and
an orbital sulcus of the same type. There must have been, however, an extensive,
tliough shallow, postsylvian sulcus, the upper end of which curves upward and forward
toward a lateral sulcus which was prolonged unusually far back. There was also
an extensive coronal sulcus, like that of Lemur.
In a skull of Galago Alleni the most noteworthy feature is the impression of a
crescentic or boomerang-shaped coronal sulcus.
Yig. 18.
Fig. IS.
Fig. 20
Kg. 21.
spc.
.TC .
Fig. IS. — Microcehiis SmitJu. Dorsal aspect of the bi-ain. Nnt. size.
Fig. 19. — Microcehvs SmitJii. Left lateral aspect of the brain. x 2.
YVr^. 20. — Microcchis ,Smithi. Lateral aspect of the loft cerebral hemisphere, dissectcfl to show the hippo-
campus and the calcarine eminence, x 2.
Fig. 21. — .Vicrocehiis Smith). Mesial aspect of the right cerebral hemisphere. x 2.
The cerebral hemisphere of the small brain of Jlicroeebus Smithi reseml)les that
of Galago in shape. It is 15-5 mm. long (without the olfactory bulb, which projects
almost 3 mm. in front of the hemisphere) ; its maximum depth is 10 mm., and its
greatest breadth 7-5 mm. (each hemisphere). The lateral caudo-ventral margin is
WOEPHOLOGT OF THE BRAIX IN THE MAMMALIA. 3 tO
13"5 iwm. long, and is so oblique that its lower end (measured in projection) is 10 mm.
further forward than its upper end. The natiform eminence of the pyriform lobe is
a very narrow conical projection on the base of the hemisphere. As in the genus
Lemvr, and unlike the condition found in the Lorisiate, the orbital excavation extends
back t(j the Sylvian fissure.
The thi]i, vertically-placed, leaf-like olfactory bulb resembles that of the Lorisinije.
The rhinal fissure is almost wholly obliterated.
The Sylvian fissure, which is constituted like that of Galago and the Lorisin;r,
extends obliquely upward and backward on to the dorsal surface, and divides the latter
into a broadly expanded posterior part (which extends as a thin leaf over the corpora
quadriyemina and the anterior part of the cerebellum) and a narrow tapering anterior
part.
There is a typically triradiate group of calcarine, retrocalcarine, and paracalcarine
sulci, the greater portions of which are situated in the mesencephalic fossa on the low er
surface of the caudal region of the hemis])here. There are no other sulci.
The notes on the brain in Ilicrocebus and Chirogale in the oft-quoted memoirs of
Gcrvais, Beddard, and Ziehen add little more. Beddard records the preseuce of a very
small lateral (his "angular") sulcus (Proc. Zool. Soc. 1895, p. 117).
The BiiAiN ix the Ixdrisix^e.
Practically all that is known of the structure of the brain in this subfamily is contained
in that part of the ' Ilistoire Naturelle des Mammiferes' of MM. Alph. Milne-Edwards
and Alfr. Grandidier which forms volume vi. of Alfr. Grandidier's great ' Histoire
de Madagascar.' The other memoirs which refer to the brain of any of the Indrisime —
viz., those of Zuckerkandl and Cluulzinski — either add nothing new or are borrowed from
Milne-Edwards's great work. Flatau and Jacobsohn derive their account of the brain of
Indris from Chudzinski, to whom thev accord all the credit for a research whicli is
either wholly or for the greater pait oljviously a reproduction of the account given Ijy
Milne-Edwards.
Gcrvais describes moulds of the cranial cavity iu the Indrisina?. lie dojs not seem,
however, to have exercised sufficient care iu the study of the impressions which correspond
to the cerebral sulci. Eor in the Lemurs, and especially in the genera Propitheciis and
Indris, it is possible to accurately map out all the sulci on the outer surface of the cerebral
hemisphere, if a critical examination be made of the prominent bony ridges on the inner
face of the cranium or of the furrows [)roJuced by these ridges on a mould of the cranial
cavity.
I have examined a considerable series of crania of the Indrisime in the British Museum
and in the Collection at the Royal College of Surgeons, and have thus been able to gain
a much more accurate conception of the jjlau of the sulci than that conveyed by the
lithographs which illustrate the ' Histoire de Madagascar.'
17r. Eorsyth Major lent me an excellent skull of an adult male Fropithectts diudona,
350
PEOF, G. ELLIOT SMITH ON THE
ill which the roof of the cranium had been removecl by a clean saw-cut. This I shall
adopt as a type.
The cerebral hemispheres were 52 mm. long, 40 mm. broad («. e. 20 mm. each), and
approximately 34 mm. deep.
The olfactory bulbs must have been almost, if not quite, completely overlapped
l)y the hemispheres ; and about as much of the cerebellum was exposed as is the case in
the genus Lemur.
The orbital excavation extends back as far as the lower end of the Sylvian fissure ; but
it does not extend upward to so gi*eat an extent as in Lemur.
Iig.22.
SCO.
sp
Fig. 22. — Propithecns diiidewa. Plan of the dorsal surface of the brain. Nat. size.
Fig. 23. — Propithenis diadema. Left lateral aspect of same.
Fig. 24. — Eight lateral aspect of same.
The rhinal fissures are much more extensive aud compk'te than they are in any other
subfamily 9i Lenuu's (excejiting the Galagos), as is well shown in Milne-Ed wards's and
Grandidier's illustrations (' Histoire de Madagascar,' (2) tome vi. pi. 86. figs. 2 & 3).
The Sylvian fissure does not differ from that of Lemur. In this, as in most of the
examples of Propithecns examined (compare also Milne-Edwards's pL 86. figs. 1 &, 3),
the upper extremity of the suprasylvian sulcus is strongly curved backward, being
obviously united to the small sulcvis (d) which is intercalated between the upper ends of
the Sylvian and postsylvian sulci in most Lemurs.
There is a typical postsylvian sulcus, the upper end of which is bent forward (in
varying degrees) in all cases. There is a clearly-defined orbital sulcus placed alongside
the olfactory bulb. It is subject to great variations in form. On the left hemisphere
of this and every other brain which I examined (although Milne-Edwards's case is an ex-
ception, op. cU. pi. 86. fig. 3) the simple primitive linear form of the sulcus is maintained;
MOEPHOLOGT OF THE BRAI?f IX THE MAMMALIA. 3.')1
\vhei"eas on tlie right side of this and two other specimens an irrei^idarity is introduced
by the development of a side-brancli. The diagonal sulcvxs seems to he more constant in
the genus Propithecus than in any other Lemur.
In only one case — Propithecus Coquereli (fig. 27) — was its presence doubtful; and, as
in that particular instance the lateral branch of the orbital sulcus is unusually long and
candally extended, the diagonal sulcus can hardly be regarded as absent even in that
case. The other sulci on the outer aspect of the hemisphere exhibit a very interesting
and most significant series of variations.
In the type specimen of Propithecus diadenia (figs. 22, 23, and 21') there are lateral
and coronal sulci not unlike those of Lemur fiiUnis, but, in addition, there is a small
transverse furrow intercalated between the lateral and coronal sulci, which recalls the
ventral extremity of the sulcus termed " central " in Perodicticus.
The posterior extremities of the lateral sulci on the two hemisjiheres curve mestally,
as so often happens in the genus iMmur when a well-defined postlateral sulcus is not
present. There can be little doubt that these mesially-curved extremities of the lateral
sulci may be explained in the same way, even though two short transverse furrows arc;
present in the right hemisphere behind the lateral sulci. In a specimen of Propithecus
Verreauxii (fig. 25), and also in the actual brain of Propithecus diadema represented in
ss..
Propitjifcus Veri-t'au.rii.
riuu of the sulci on the lateral aspect of the left cerebral hemisphere. Xat. size.
the 'Histoire de Madagascar,' (2) tome vi. pi. 80. fig. 1, the presence and Jiosition of a
definite postlateral sulcus, and the absence of an inwardly-bent tail to the lateral sulcus,
clearly complete the demimstration of the identity of these two structures. It is further
of interest to note the presence in the actual brain of Propithecus diadema of two
small transverse furrows such as we have seen in a cast of the same species : seeing that
these furrows exist behind a definite separate postlateral sulcus, they aff'onl a further
justification for the statements made in the above diseussion.
The most instructive features in the brain of Propithecus are tlie varying phases in
the group of furrows which we may term " central."
In the only representative of Propithecus dindeni'i (tigs. 22, 215, ami 21) tluu-e is a
snuxU transverse sulcus in the situation corresponding to that occupied by the little
furrow /' in Leiuur. On the left hemispliere there is a longer transverse sulcus (c)
SECOND SEIUE-S. — ZOOLOGY, VOL. VIII. 52
352
PEOF. G. ELLIOT SMITH ON THE
immediately external to this : it is interposed between the adjacent ends of the lateral
and coronal sulci. In Milne-Edwards's specimen of this species the correspondmg sulcus
is represented as being fused to the coronal sulcus (Hist. Madag. (2) tome vi. pi. 86.
fig. 2), just as the coronal sulcus is united to the furrow e in Lrmnr, and also in
ray specimens of Prop it heats Eihoardsi (figs. 28 and 29), F. Verreanxii (fig. 26), and
P. CoqvereU (fig. 27). On the right hemisphere of P. dladcma (figs. 22 and 24) the upper
sulcus (/) is longer, and the upper end of the lower transverse sulcus {e) is bent back
behind the end of the lateral.
Fig. 26.
SCO.
Fig. 2G. — Fnipitheciis Verrcaimi. Plan of the sulci on the flutcrior portion of another left
eerebrul hemisphere. Nat. size.
Fig. 27. — Pi-opitlients Cwiucrdi. Plan of the sulci on the anterior portion of the right
cerebral hemisphere.
Fig. 2S.- — Propithecus Edwardsi. Ditto.
Fig. '29. — Ditto. I,eft hemisphere.
In the specimen of Fropithecus Udwardsi (figs. 28 and 29) the arrangement found iu
Lemur is reproduced. The small furrow f is oblique, as it is in Lemur. On the left
hemisphere the upturned caudal apjieudage {e) of the coronal sulcus is apparently
separated from the main coronal element.
In Propithecus Coquereli there is no furrow f, and the sulcus e is unusually Jong
(fig. 27). It is joined to the coronal at a right angle, and overlaps the end of the lateral
on the caudal side. An exactly similar state of alfairs is found in one specimen of
Propithecus Verreaitxii (flg. 26) ; but in another specimen (fig. 25) of the same species
there is a long typical central sulcus, like that found in Perodicticus : it clearly
represents a fusion of elements c and /' and their complete emancipation from the
influence of the coronal and lateral sulci.
The arrangement of the bony ridges corresponding to the cerebral sulci in a cranivxm
MORPIIOLOGT OF THE BEAIX IX THE MAiMMALIA.
353
of Iiidris brcvicaudatns in the Museum of the Pioyal College of Surgeons (Osteological
Series, No. 253) is represented in fig. 30. The actual brain of a representative of this
species has been well represented by Milne-Edwards (Hist. Madag. (2) tome vi. pi. 87.
figs. 4, 4 ft, 4 h, and 4(?).
The draAvings published by Chudzinski (Bull. Soc. Anthrop. vi., vii), and reproduced
by Flatau and Jacobsohn (Handbuch, p. 203), appear to be borrowed from the figures 4
and 4 h of MiJne-Edwards's work.
In my specimen (fig. 30) the upper end of the suprasylvian sulcus is strongly
recurv^ed, as it is in Propitlwcns. There is an unusually long postlateral sulcus, as
there is also iu Milne-Edwards's specimen. In the skull the ridge for the lateral sulcus
was shallow and ill-defined, and seemed to be broken into three parts (/', /", and l'"), of
which only the anterior (/'") was promineat. It seemed to be joined to the coronal
Fig. 30.
Fig. 30. — latins hrfvramdvliis. Left lateral aspect of a cranial cast. Xat. .si/e.
Fig. 31. — li',(/u's lamfja-. Lateral aspect of the left cerebral hemisphere. Xat. size.
sulcus, which was much longer than in Milue-Edwards's specimen. Judging from the
condition represented by Milne-Edwards in the actual brain, I am inclined to regard
all three furrows, V, T, and V'\ in my specimen as representing the lateral sulcus. But
the relationship of the sulcus Z"' to the coronal seems to indicate that th(> lornici- sulcus
may also represent the furrow e of Propithecus (compare figs. 20, 27, 2.S, aud 2!J), which
may have fused with the lateral sulcus. This is a matter which can only be settled by
a careful study of a series of actual brains. This suggestion is put forwtird merely as
the solution which seems most likely to be correct.
Chudzinski regards the sulcus which is formed by the union of the furrows /" and /"
in Milne-Edwards's specimen as the " central sulcus." But there is iu both specimens a
small sagittal sulcus /, which represents the similarly-labelled furrow in I'mpWiecm
and Lemur.
The orbital sulcus is large aud asvmmctrieal in the two hemispheres.
52*
;5rj4 PEOF. G. ELLIOT SMITH ON THE
The rapsial surface of tlie hemisphere in the Indrisinte is known to me only in the
drawings (Hist. Madag-. (2) tome vi. pi. 86. fig. G, pi. 87. figs. 2 b and 4 c) of Milne-
JMwards's monograph. It closely resembles that of Chiromys, which will be described
later on in these notes.
Nothing is known of tlie smaller brain of Arahis except the scanty inforaiation
contained in Milne-Edward.s's memoir. The plan of the ridges on the interior of a
cranium oi Aral/ is laniger (in the Koyal College of Surgeons) is shown in fig. 31.
Tliere are well-defined representatives of the Sylvian, postsylviau, and orbital sulci,
shallow lateral and postlateral furrows, and also a deep sulcus x (in apparent continuity
Avith the lateral sulcus) rimning parallel to the Sylvian fissiu-e. Comparison with the
brain of Iiidris (fig. 30) seems to point to an homology with the element I'" in that genus.
Comparison with the plan found in Fropithecus (compare fig. 24) seems to suggest its
identity with the sulcus/of that genus. In Milne-Edwards's specimen it is not so distinctly
transverse in direction. In my specimen there is no trace of any ridge corresiwndiag to
the coronal sulcus. All this variability of the furrows around the adjacent ends of the
lateral and coronal sulci in the Indrisinae shows how unstable this region is in these
miimals. This instability is probably due to two distinct factors. In the first place, this
area is the seat of very pronounced growth-changes in all the Primates, and especially
the lowlier members of tlie Order, because the excitable area of the cortex is expanding
to a much greater extent than in other mammals. At the same time, there is in tlie
Ijemnrs, and especially the Indrisina?, another and a greater disturbing factor, which is
the pionouuced tendency to a disintegration of composite sulci : this must be the
expression of some retrogressive process of evolution. Tliis variability is perhaps best
exemplified by the behaviour of the sulcus /', wliich seems to be linked in different
individuals of the Indrisinse to the coronal sulcus, the sulcus c, and even to the lateral
sulcus.
The Cerebral Hemispheres in the Genus Datjbentonia
[more commonhj called Chiromys).
The brain of the Aye-aye {Baubentonia mcuhfgascariensis, Geoff., Chiromys madagas-
cariensis, Cuv.) is of extraordinary interest to the morphologist by reason of the
possession of what Chudzinski calls " contradictory characters." The same writer
lurther explains this remark by the statement that although the Aye-aye is no larger
than a small rabbit, or even not so big, it has a most voluminous brain, very rich
in cerebral folds*. He adds that "the form of the l)rain is ovoid, but its anterior
extremity is mai'kedly i-ounded. In this respect it resembles the l)raiu of certain
]lodentia (Castor, Ilijstrix) and even certain Marsiqiials {Pl/ascolomi/s)." Although
these comparisons are quite justified, rej^emhlances of a closer character are to be found
within the Prosimise themselves. Gervais describes the crucial cast of the Aye-aye as
being "plus arrondi et plus bouibe a sa face superieure que celui des Makis, du
Propitheque etde ITndri " f. He compares the appearance of its upper surface with that
of the Carnivore Ail uriis fid gens.
* Hull. Sw. Antlirop. svr. 4, vii. 1S9G, p. i:!. t Jourii dc Zool. i. 167l\ p. 23.
JIORPHOLOGT OF THE BRAIX IN THE MA:\IMALIA.
355
The olfactory liull) is a flattened cake-like mass 9 mm. wide and 11 mm. long, and
resembles that found iu the Indrisinte and in the genus Lemur. it is, howev'er,
completely overlajiped by the cerebral heraispliero, as Ouderaans and Chudzinski have
already pointed out ; Gervais's representation of it projecting' beyond the hemisphere
like that of Lemur is clearly erroneous.
About as much of the cerebellum is uncovered by the cerebrum as we have noted in
one specimen of Lemur /ulcus (fig. 3).
The arrangement of the cerebral sulci presents some most extraordinary features :
it is, moreover, especially noteworthy by reason of its extreme variability. The plan of
the fui'rows is so puzzling that no writer on the brain of Chlromys has ventured to
compare it in detail with that presented by other Lemurs. Thus, Owen makes use of
the nomenclature used in describing the brain in the Carnivora; Gervais compares it with
Fig 32.
Clt Iroinijs madarjascariensis.
The dorsal aspect of the braiu. Slightly enlarged.
that of Ailurus ; Oudemans likewise employs Krueg's nomenclature for the Carnivore-
brain; Chudzinski also is driven to a comparison with the cerebral pattern in the same
Order ; and, finally, A. J. Parker * quotes with approval the opinion of Pansch that the
Aye-aye belongs to the group of mammals which includes the Carnivora.
The chief reason for all this hesitancy in comparing the svdci of this brain with those
of the other Primates depends upon the fact that in Cldromys the suprasylvian sulcus
has become widely separated from the pseudosylviau sulcus, and the latter is exhibited
in different specimens iu various stages of disintegration. Thus there is no typical
Svlvian fissure ; and the failure to recognize that the latter consists (in the other
Primates) of an apparent blending of the suprasylvian and pseudosylviau sulci by the
submergence of the region between the overlapi)ing parts of these furrows {vide supra)
naturally gives rise to confusion w'hen the intervening area is not submerged, as is the
case in Chiromys. This separation of suprasylvian and pseudosylviau sulci also makes
* Journ. Acad. Philad. ser. 2, s. IS'JC.
550
PEOr. G. ELLIOT SMITH OX THE
the resemblance of the pattern to that found in tlie Carnivora much moi'e obvious than
it is in other Lemurs, and, to a certain extent, justifies the comparison Avliich all
previous writers have instituted.
It was, in fact, the search for an explanation for the extraordinarily puzzling arrange-
ment of the sulci in the brain of C/u'rowi/s-, which led me to conclude that the
sujirasylvian sulcus becomes merged in the " Sylvian fissure " in the Primates — an
hypothesis the accuracy of which has been so amply confirmed by later studies.
In one brain, ^vhich I shall distinguish as u, a deep f uitow begins near the bend of the
rhinal fissure, and passes upward with a very slight backward inclination for a distance
of about 11 mm. This is the pseudosylvian sulcus (figs. 33 and 3i).
Fig. 33.
sec
Fig- 34. si.
SCO.
S3.
Fig. 33. — C/iiromt/s mud<tiia.iiar'iciis':s. Lateral aspeit of the left cerebral hemisphere.
Fig. 34. — Lateral aspect of the right cerebral hemisphere.
An arched furrow formed by the union of the suprasylvian and postsylvian sulci,
after the manner of the pattern found in the Carniv^ora, surrounds the pseudosylvian
sulcus (figs. 32, 33, and 34). The suprasylvian sulcus begins at a distance of 5 mm.
in front of the upper part of the pseudosylvian, and arches obliquely upward and
backward to a point 15".j mm. from the mesial plane and 125 mm. in front of the
posterior margin of the hemisphere. At this jioint it joins the postsylvian sulcus,
and from the conjoint furrow a little ujiwardly-directed branch 2 mm. long is given
off. The postsylvian sulcus is 8'5 mm. behind the pseudosylvian sulcus. The two
hemispheres of the specimen a are practically symmetrical so far as these features are
concerned.
In a second specimen (figs. 35 and 36), which I shall distinguish as /3, the suprasylvian
sulcus presents similar features to those just described in k ; but the postsylvian and
pseudosylvian have become so greatly distorted and broken up that a pattern is formed
which is quite unlike that exhibited by the specimen a.
On the left hemispljere there is a much shorter and shallower pseudosylvian sulcus.'
The suprasylvian sulcus is separated from a very oblique sulcus (which is analogous to
the upper part of the postsylvian) by a narrow gyrus (fig. 35). And between this and
the small pseudosylvian there is a vertical sulcus (.r) \\hich is much longer and deeper
than either of the furrows between which it lies. It is clearly a furrow compensatory
to the fragmentary pseudosylvian and postsylvian sulci. Behind it there is a much
MORPIIOLOGT OF THE BRAIX IX THE MAl\r.M ALIA.
357
shorter furrow {y), placed below the displaced upper postsylvian element (/r). This
represents another fragment of the postsylvian sulcus.
In the rig'ht hemisphere the pseudosylvian sulcus is represented by a very shallow
depression, which is not worthy to be called a sulcus except in the uppermost 2 millimetres
of its extent. The oblique sulcus lo is here joined to the suprasylvian. The sulci x
and y are practically symmetrical on the two hemispheres. It seems probable that tlie
little ascending branch of the arcuate sulcus in the specimen a belongs to the postsylvian
sulcus. In a series of crania of Chlromys which I have examined, the condition of the
suprasylvian, pseudosylvian, and postsylvian sulci most nearly resembles that seen in
the specimen u in all cases.
Yig. 35.
.55. si.
Fig. 3.J. — Chiroiivjs riKidiiijascnriciisii'. Loft lateral aspect of another brain.
Fig. 30. — Chirumijs madagaicitrleusls. Lateral aspect of the right hemisphere.
In the two crania numbered 302 and 302 B in the Collection of the P^oyal College
of Surgeons, the bony ridge corresponding to the suprasylvian sulcus is much nearer to
the well-defined ridge representing the pseudosylvian sulcus (in all four hemispheres) than
the corresponding sulci are in the brain a. In the skvill numbered 302 the postsylvian
ridge seems not to be joined to the suprasylvian. But in the other cranium, as well as
in one of those in the British Museum, the suprasylvian and postsylvian ndges unite to
form a regular arc.
In Oudemans's sj^ecimen the arrangement of the sulci appears to have been even more
distorted than it is in my specimen /3. Tims on tlie left hemisphere {op. cit. pi. iii.
fig. 11) the suprasylvian sulcus (Avhich he labels " Sylvian") is Avidely removed from the
small upper fragment of the postsylvian (which he correctly labels " suprasylvia
postei'ior "). There is no sulcus corresponding to that which I labelled y (tig. 35), but
there is a Avell-defined sulcus x, which Oudemans regards as part of the postsylvian
(liis " suprasyl. post."). There are also two furrows radiating from the angle of the
rhinal fissure — one (which he does not label) passing to the representative of the svilcus x,
and the other (which he calls " Sylvian") passing vertically upward in front oft\\e lower
end of the suprasylvian. On the right hemisphere {op. cit. pi. iii. fig. 12) these two
sulci (suprasylvian and the last described) are united : the two sulci Avhich appear to
represent (this is merely a rough comparison, for no exact homology can exist) the
furrows ic and x in my specimen /3 are united to form a very sinuous sulcus.
358 PEOF. G. Er,LIOT SMITH OX THE
Oudemans explains this exceedingly puzzling condition in these words :— " Die Fissura
Sylvii zeig-te sich an der rechten Hemisphiire viel mehr ausgeprJigt als an der linken.
Nnr der obere Theil derselben war tief einschneidend, der nntere mehr superficiell,
besonders an der linken Ilemispliare, wo beide Tlieile selljst getrennt waren. Das wir
es hier jedocli gewiss mit der Fissura Sylvii zn tlinn haben, ergab sich durch Ver-
gleichung mit den Gehirnen einiger anderen Lemuriden." (Naturh. Yerh. Akad. Amst.
xxviii. p. 28.)
The closing phrase is true, but the writer misses the obvious inference, which must
follow from this important fact, viz. — that the Sylvian fissure of the Lemurs is
partly represented by the suprasylrian sulcus. In order to evade this conclusion, in
deference to the popular belief in the identity of the so-called " Sylvian fissure "
of Carnivora and the Prosimian Sylvian fissure, Oudemans is forced to call the lateral
sulcvis " suprasylvian." [For if the real suprasylvian is considered as representing
the "feline Sylvian," it is not illogical to call the lateral sulcus by the name
"suprasylvian," even if such an hypothesis makes it necessary to suppose that the
lateral sulcus has disappeared.] It was unfortunate for Oudemans that he did not
chance to see such a specimen as my example a ; for then there could have been
no doubt that the Lcmuroid Sylvian fissure is also in great part identical with tlie
suprasylvian sulcus.
Chudzinski, who happened to have a specimen Avhich (so far as I a)n able to judge
from his unsatisfactory account) resembled my specimen a, failed to recognize the
suprasylvian sulcus as the Lemuroid " Sylvian."
He regards the pseudosylviau sulcus as the Sylvian fissure, and says that it is joined
to the rhinal fissure (which he calls the fissure of the olfactory lobes) on the right side
but not on the left (Bull. Soc. Anthrop. ser. 4, vii. pp. 13 & 11). [This diff'ereuce between
the two hemispheres also seems to occur in my specimen «, but the condition of the brain
prevents me from speaking more decisively.] He is then so puzzled by the suprasylvian
and postsylvian sulci, that he does not attempt to compare them with the sulci of the
Primates, but, like all previous writers, seeks an analogy to the condition found in the
Carnivores {op. cil . p. 1()). His specimen seems to have been exceptionally interesting,
and it is therefore very disajipointing to find such imperfect drawings and such vague
descriptions, from which it is impossible to form a clear conception of the condition.
The postsylvian sulcus seems, if I read the account correctly, to be independent of the
suprasylvian on the right side ; and the latter appears to l)e so related to the pseudo.sylvian
as to be very slightly if at all difteient irom the normal Prosimian Sylvian fissure [but
this is by no means clear].
One cannot fail to be amazed that Chudzinski, who during the previous year had been
studying the brain in Lemur, Kycticcbns, and Indris, should have failed to recognize in
the suprasylvian sulcus of Chii'omys the homologue of the upper part of the Sylvian
fissure of these Lemurs.
It is difiicult to i'orm a clear conception of the condition of Owen's specimen from his
imperfect lilhographs and vague description ; and the interpretations of it by succeeding
writers (such as Pansch, Beddard, and Parker) only tend to still further confuse the issues.
:\l()KPIl()T.OGV OF TlIF, BRAIX 1\ THE .MAMMALIA. S-IO
Tlie remarks of (icrvais arc of intei'cst only in so far as tlicy call aifcnliou to the
presence of an additional arcuate sulcus [/. c. the scpiratiou of tli ; suprasylvian ami
the pseudosylvian sulci] without suggestiui;' any solution.
In my specimen a the lateral sulcus ])cgins ^posteriorly at a point 8 mm. from th(>
mesial plane and an equal distance from the posterior margin ; the anterior extremity is
11'5 mm. from the mesial plaiu> and 10 mm. from the; anterior extremity of the
hemisphere. The sulcus is 23 mm. long. In none of my specimens, nor iu that of
Oudemans, csn I find auy trace of a separate postlateral sulcus, hut in the left hemi-
sphere of the specimen /3 there is a characteristic mcsially-heut tailpiece such as we
have seen representing the joostlateral sulcus in many Lemurs.
The furrows which are situated in that region of the hemisphere wliicii lies in front of
the lateral and suprasylvian sulci are suhject to consideral)le variation. On the lelt
hemisphere of the specimen ^3 there is a typical orhital sulcus, and in a more dorsal
position a very obliquely placed sulcus, which a comparison Avith otiier Prosimian
hrains clearly shows to he the anterior part of the coronal sulcus. On the right
hemisphere of the same brain a shallow depression unites these two furrows. On the;
left hemisphere of specimen a, they are joined to form a single transversely-placed
arcuate furrow ; on the right hemisphere the orbital furrow has disappeared and there is
simply a transverse coronal element. In two of the crania (302 and 302 b) to which I
have refcri'ed, the two sulci are well developed and united to form figures, one like a
crescent, the other like a letter U placed on its side.
These sulci are exhil)ited in a typical manner in Oudeuiaus's memoir, and the names
" presylvian" and " prorean " which he applies to tliem we shall find in the subsequent
discussion to be not inappropriate.
Midway between the sulcus which I have called coronal and the anterior ends of the
lateral and suprasylvian sulci respectively, there ai-e two sulci in the left hemisphere of
the specimen 3. The upper of these may be compared Avith the sidcus distinguished by the
letter /in Lemur, and the lower one with that called c. These two sulci are united by a
shallow furrow to form a long transverse " sulcus centralis" (fig. 35).
On the rityht hemisphere of the same brain (fig. 36) the two furrows r and /are much
smaller and are Avidely separated. The sulcus / is joined to the anterior end of the
lateral sulcus, and e remains as a short independent furrow widely separated from all
the other sulci on the hemisphere.
On the left hemisphere of the specimen u (fig. 33) the two sulci ^ and /are united at
an obtuse angle, and the former has a horizontal ramus, which recalls the fact that it is
really the posterior fragment of the coronal sulcus. On the right hemisphere of this
brain the horizontal sulcus is independent of the moderately long central one.
On the right hemisphere of Oudemans's specimen there is a small triradiate sulcus/
(which he calls " coronal "), and a horizontal sulcus exactly like that exhibited by the
right hemisphere of my specimen a. This he calls " prorean," i. e. the same name
by which he denotes the anterior element which I call " coronal." This is of interest in so
far as he gives the same name (" prorean ") to these two widely-separated furrows, which
I also regard as two fragments of one sulcus ("coronal"). On thclcft hemisphere of
SECOND SERIES. — ZOOLOGY, VOL. YIII. ^^
300 PEOF. G. ELLIOT SMITH OX THE
liis specimen the / clement is a larger transverse sulcus and the e element is a small
Y-shaped furrow, which might possibly be explained by the condition seen in the left
hemisphere of my specimen a.
On both sides of three crania which I examined there is a long ridge representing a
fairly extensive *' central sulcus," such as might be produced by a deepening of the
connection between the furrows e and/ on the left side of my specimen /3.
If we summarize all this puzzling variety in the arrangement of the furrows on the
outer surface of the hemisphere of Chtromijs, the great outstanding fact is revealed that,
amidst all the variations which so modify as almost to disguise the identity of most of
tlie sulci, the suprasylvian and lateral sulci alone remain unchanged. They are the
deepest and most stable sulci on the outer surface of the hemisphere.
The other feature of interest is the strongly marked tendency toward the formation
of a transverse sulcus from the elements e and /, which is either the homologue of the
central sulcus or, at least, in the present state of our knowledge, fulfils all the known
criteria of a sulcus of llulando.
The peculiar variations which have been noted in the cerebral hemisphere of Chiromys
can only be explained on the supposition that a large and highly convoluted brain is
undergoing a process of retrograde metamorphosis which leads to the disintegration of
all except the most stable sulci. It is altogether inconceivable that the unstable group
of fragmentary sulci which rej^resent the pseudosylvian, postsylvian, and coronal sulci
can be stages in the evolution of a more complex pattern of furrows. All these facts
point to the conclusion that Cliii'omijs is not the lowly Carnivore-like primitive Lemur
which most other writers on cerebral anatomy suppose, but a highly-specialized Pi'imate
which has undergone most pronounced retrogressive changes, as the result of wdiich all
the more variable features of its brain stand revealed and, by contrast, show the underlying
common mammalian featiu'cs, which are the heritage of all ttie Meta- and Eutheria.
None of the writers who have described the brain of CItiromijs has placed on record any
information concerning the mesial surface of the hemisphere. I have examined this
aspect in four examples.
The corjDus callosum is 23 mm. long and very plump, Avith a considerably thickened
splenium. Its anterior part is bent downward so as to be boomerang-shaped (fig. 37).
This is unlike that of the other Lemurs, and may possibly be due to a retraction of the
anterior regions of the hemisphere, when the latter lost the more pointed form which it
must have had in the immediate ancestors of Chiromys. For the peculiar blunt anterior
pole of the hemisphere cannot be primitive, and when this process took place it probably
induenced the form of the corpus callosum on the mesial surface, just as it has modified
the coronal sulcus on the lateral aspect of the hemisphere.
The hippocamjjus is distinguished by the possession of an unusually large hippocampal
tubercle (fig. 38), and also possesses that peculiar furrow {a) which we have seen ia the
genus Lemur. I know of no brain other than tliose of the Primates in which these two
peculiarities are both found. The particular form they assume in Chiromys is distinctly
Prosimian.
Tlie most interesting featiire on tlie mesial surfiice is the grouping of the sulci.
MOKPHOLOGT OF THE BKAIX IN THE MA^r^IALIA.
8fil
The first cerebral hemisphere of the Aye-aye, which I cxamiiicd, is tliat represented
in fig. 38. At that time I had just coniph>ted an investigation into flie morphology of
the calcarinc sulcns, and had come to the conehision that tlie homologuo of the [anterior]
calcarine sulcus of tlie Primates was to be found in the retrosplenial part of the splenial
sulcus of Krueg in the brains of Carnivora, Ungulata, Ivlentata, Chiroptera, and
jMarsupialia (Proc. Anatomical Soc. of Gt. Britain and Ireland ; Journ. Anat. and Phys.,
189!)). In tliis particular hemisphere of Cliiroinijii there seemed, at a casual glance, to
be one great arcuate sulcus (such as we find in Tapinis, for example) rei3resenting a
fusion of the genual, intercalary, and calcarine sulci ; and so, occurring in a Lemur,
Pig. 38a.
Fig. 37. — Chiromi/s madar/ascni-ieiisls. Mesial aspect of the brain cut in sagittal section.
Fig. 38. — Chifomi/s madar/ascariensis. The mesial aspect of the right cerebral hemisphere.
Figs. 38 «, h. — Chironiiis. Modifications of the calcarine group of sulci, delineating the variations as if they
occurred in the /-ir/ht hemisphere, so as to be more directly comparable with figs. 37 and 38.
there seemed to be a striking confirmation of the hypothesis I then put forward.
However, upon separating the lips of the sulcus in the region immediately behind the
splenium, the great arcuate sulcus was found to be interrupted by a small narrow
submerged gyrus, so that the essential relations of the two furrows were thos(i
diaEcrammaticallv shown in fiij. 38. Examination of the other hemisphere showed the
existence of a broader bridge, and thus exj)0sed a state of affairs which is essentially
identical (fig. 38 b) with that found in other Lemurs. The subsequent examination of .i
second brain still further confirmed this view. In these four hemispheres of Cltiromyi
we find therefore every gradation of form between a condition which dilFers in the very
53*
302- PROF. G. ELLIOT SMITH ON THE
bliglitost degree from tliat common mammaliau form, such as the braiu of Taplrm
exhibits, to the characteristic Prosimian or Primate condition. The jiersistence of the
narrow g-vrus between the intercalary and pai'acalcarine sulci, when the mechanical
conditions clearly ])i-edispose to the fusion custoraaiy in non-Primates, is yet another
testimony to the view tliat Chiromys is a retrograde Primate.
The calcarine sulcus begins at a point S mm. lielow the splenium of the corjius
callosum and 2 mm. behind the hippocampal ti.ssare ; it jjroceeds obliquely upward and
backward to a ])oint 5 mm. behind and slightly below the level of the splenium : there
it bifurcates into retrocalcarine and paracalcarine branches. The former proceeds hori-
zontally backward and the latter vertically upward. In one instance the paracalcarine
is separated from the rest of the great cingular sulcus by a narrow submerged strip
(tig. 38) : in another case (fig. .38 a) there is also merely a very narrow strip separating
the sulci, l)nt the upper end of tiie paracalcarine is prolonged beyond the region of
approximation. This is the condition wdiicli Milne-Edwards (and Chudzinski after him)
represents as l)eing the usual form in the Indrisinse.
The Braix of the Extinct Lemuroib Globilemur.
In 1897 Dr. Eorsyth Major described * a })laster mould of the cranial cavity of the
large sub-fossil Lemuroid to which he had previously given the name Glohilcinur
Flaconfti.
I Avould not have deemed it necessary to attempt to add anythiiig to the clear and
sufficient (/. e. considering the state of the material) account given by this excellent
anatomist, if his memoir liad not been recently criticised by lludolf Burckhardt. "Whilst
I Avas actually engaged in studying the plaster mould and the cranium from Avhich it
Mas made in the British Museum, my attention was called to the brief report of Burck-
hardt's criticism, Avhich was then just published f . I therefore carefully re-examined in
the light of these criticisms the crania and the casts of Glohllemur [and also of 2Icga-
ladapis, to which I shall refer later in this memoir]. Professor Burckhardt has just been
ood enough to send me a co2:»y of the full memoir %, of Avhich the above-quoted note
may be regarded as an abstract.
The brain of Glohilcmur was considerably larger than that of any living Lemur. It
had large flattened olfactory bulbs projecting very slightly beyond the apex of the
hemisphere, as in the genus Lemur. In fact the whole of the exposed part of the
rhinencephalon closely resembles that of Lemur.
1'he cerebral hemispheres (fig. 39) cover the cerebellum to a slightly greater extent
than in most members of tlie genus Ljemnr ; the obliquity of the postero-inferior margin
of the hemisphere is about the same as in Lemur (fig. 40). In respect to both of these
points, Burekhardt's drawings are misleading [op. cU. tig. 1, p. 231).
So far as the shape of the hemispheres is concerned, this cast differs from that of any
of the larger Lemurs, such as Chiromys, Liidris, Froplthecus, and Lemur. It is least
» Proc. Royal Soc. 1S97, pp. 40 & 47, pi. 5. tigs. 1, 2, and ;i.
t 'TagoblaU dcs V. Litei'iuitionalfii Zdologcn-Cougresses,' ]!urlin, August liXIl, Nimimi'i- 4, 14lli August,
j ''Das Uehirn zweier subfussilcr Ricseulcmureii aus iladagasear,"' Aiiatomischer An/.eiger, Rd. xx. 1901,
pp. 2:29-232, fig. 1 , a & «.
s
:moupiiologt of the brain ix the mammalia.
3G3
uiilila' the latter, but the extreme broadeuiug' of the hemisphere ijosteriorly ami the
more pronounced tapering of its anterior part give the brain (as a whole) an oval form,
which is quite unlike that of Lemur. Porsyth Major compares its shape to that of
Jlicrocebits ; oa the whole I think that the dorsal contour of the brain api)roaches
nearest to that of the smaller Cercopithecidse.
Fig. 40.
Pig. ."39. — (•Johthmur Flarmirti. Tlie dorsal aspect of a cranial cast. x :\.
Fig. 40. — Gloliilemur F/acoiirti. The left lateral aspect of the same. x ,j.
The problem of mapping-out the pattern of the cerebral sulci in this brain is one of
singular and unexpected difficulty. In most Lemurs and in many of the Cebidse the
positions and extent of the various sulci can be determined with the greatest ease and
certainty. But in the crania of most Apes the ridges which corresjjond to the position
of the cerebral furroAvs become blurred and indistinct in the greater part of the surface,
excepting only the anterior region, which contains sharply-defined elevations representing
the sulci rectus, arcuatus, orbitalis, and fronto-orbitalis.
If a series of crania of Cercopithecidtie be examined, it will be found in many of i hem
that all the ridges behind that representing tlie sulcus arcuatus are so indistinct that it
would not be easy to map out the sulci, if we did not liappen to know the plan. It is
a i)eculiar iact that the ridge for the Sylvian fissure, which is always so distinct in the
Lemurs, becomes almost completely obliterated in the crania of many Ajjcs. And
the crests representing the intraparietal, the parallel, and, in a less marked degree, the
central sulci share a like fate.
A similar state of affairs is unfortunately jjreseut in Globllemur ; so that it is quite
impossible to map out the position of most of the sulci. As in the Apes, the ridges
representing the sulci on the anterior region of the hemisphere are by far the most
distinct.
Tlius there can be no doubt concerning the recognition of the orbital (t^) and coronal (Ij)
sulci (figs. 3!J & 40). There is a prominent bony ridge representing a furrow skirting the
fronto-orbital margin of tlie hemisphere, wiiich is probably the diagonal sulcus (e). But
the determination of the identity of the other furrows offers insuperable difficulties.
In both hemispheres there is a short vertical furrow (/), which both Forsyth ]\[ajor
and Burckhardt consider to be the representative of the Sylvian lissure or part of it.
^Oi PROF. G. ELLIOT SMITH OX THE
Tlie latter writer regards the furrow g to l)e the upper part of tlic Sylvian fissure, whicli
is separated from the lower part (/) by " eine kleiiie Untcrbrechung, die jedoch uur auf
llcehnung des Schadelaugusses zu setzeu ist " (Anat. Auz. 1002, p. 231). Natural
though sucli a suggestion is at a casual glance, its adoption would logically lead to many
difficulties in the interpretation of other furrows. Thus it is quite inconceivable that the
lateral (intraparietal or " interparietal " of German writers) sulcus, which we have learnt
to regard as one of the most stable sulci, not only in the Lemurs but also in the other
Mammalia, can be represented merely by the furrows d and n. The former of these {d) is
not even present on the right hemisphere, and ;/ is so shallow as to be of questionable
^aluc. Moreover, if n is ]mrt of any furrow, it is that marked g, and not d, to which it
nnist be linked. If Burckhardt had suggested that the lateral (intraparietal) and the
suprasylviaii (Sylvian) sulci had fused as in Nycticehns, Chrysothrix, and Nyctijjithecus,
ihere would have been less reason for criticising his suggestion on this jiarticular point.
But there are other reasons which forbid us accei:)ting the suggested homology as the true
interpretatioa. It is hardly i^robable * that a hemisphere whicli has extended backward
over the cerebellum to an extent at least as great as ever occurs in Lemur should possess
u Sylvian fissure which slopes backward so much more slightly and occupies such a
forward position. And if g is the Sylvian fissiu-e and h the superior temporal, as
Burckhardt suggests, it necessarily follows that half of the cerebral hemisphere lies on
the caudal side of the latter sulcus! This would be a state of affairs for which no
])arallcl exists elsewhere in the Primates.
I do not think that Burckhardt's suggestions are at ail conckisive or even probable.
But, on the other hand, I cannot offer any alternative scheme w^hich is not open to adverse
criticism. If the furrow / had not been present f I should have confidently believed
that this brain retained the true Lemurid pattern of sulci, g + n being the lateral,
h the Sylvian, and / + ,/ being the postsylviau ; but now" I cannot do more than
subscribe to the opinion of Forsyth Major that "it is never safe to attempt to make out
the exact homologies of the fissures in a cast of the brain-cavity" [op. cit. p. 17).
The BiiAiN ix Megaladapis.
One can speak with much more confidence concerning the features of the brain in
3Iegaladapis madugascariensis, Maj. ; but most that deserves saying concerniug this
brain has already been admirably said by Dr. Forsyth Major %.
The great contrast which the shape of this biain presents to that of Glohilenmr and
the extraordinary conformation of its anterior paits (more especially the elongation of
the olfactory peduncles and the relatively forward position of the optic chiasma)
have been clearly shown in the figures and descriptions of the work just quoted. The
*" In making this statoment I have not forgotten fliat the Sylvian fissure may occupy such an unusual position,
.i'«, for instance, in Tarsius.
t In the skull of a specimen of jS'i/clicehiis 1 have seen a bony ridge in a corresponding situation, which did not
represent any furrow on the brain itself.
* Proc. Ptoy. Soc. ISiJT, pp. 47-50.
MORPHOLOGY OF THE BEAIX IX THE ^LnrMALTA.
505
essential features are represented diasframmatically in the accompanying sketches
(figs. 11 & li>).
Fig. 41. — M'l/ahidiijiis tii(iJai/(i>:c(iriiii.sis. The dorsal aspect of ii (-Taiiial ca.^t. X ,-.
Fig. 42. — Mer/iiJadapis hiafhii/uscarunsis. Tile lateral aspect of the same. X ^.
The form of the brain (especially the full rounded ])lunt anterior ends of the hemi-
spheres and the absence of a fronto-orbital edge) resembles that of C/urom//s. Tlio
arrangement of the furrows approximates most nearly to that seen in the Indrisiuae and
especially l/ulris. They are, however, fewer in number and simpler t'aan they are in
tlie latter. There can be little doubt as to the identity of the suprasylvian, lateral,
coronal, and postsylvian sulci, and the furrow / represents the similarly-labelled sulcus
in other Lemurs. The appai'ent absence of the orbital sulcus is notewoi'tliy. Burck-
hardt's extraordinary suggestion that the furrow which I have called " lateral " repre-
sents the precentril sulcus is peculiar, bec;iuse no central sulcus is present; but it is
preposterous to represent as the precentral a furrow which, according to his diagram
ioj). ciL p. 233, fig. 2 a), is placed parallel to and on a lower plane than the frontal
sulcus.
But the most unwarrantable statements in Burckhardt's work are those accusing
Forsyth Major of mistaking the optic nerves for the olfactory bulbs. The latter
anatomist had carefully studied and described the skull of Ilegaladupis * long before
he attempted to describe the plaster mould of its cranial cavity. lie was therefoiH!
labouring under no misconception as to which lacunce in the cranial wall Avere the optic
foramina and which depressions were the olfactory fossee. So that when he came to
describe the cranial cast, there was no difficulty in recognizing as such the optic nerves
(even thovigh their position is so peculiar) and the olfactory peduncles (greatly elongated
though they are). In the account which Burckhardt criticises, the conformation of
the cranial region surrounding the disputed brain-area is carefully described (Proc.
* •' On Miyaladii^iis inath'ij'tscai-uusif, an extinct gigantic Lemuroii.1," Pliil. Trans. li. vol. 1S.5, 1804, pp. 2.5 & ■2(i.
300 Pi^UF. G. ELLIOT SMITH OX THE
Koy. Soc. 1897, pp. 48 & 49). And yet he would have us believe that so experienced
and careful an investigator as Dr. Forsyth Major had mistaken for the olfactory bulbs
structures Avhich are really the ojjtic nerves ; in other Avords, that he has been unable to
distinguish in the cranium between the olfactory fossa :md the o])tic foramen ! This
l)rcposterons charge was made at the Berlin Zoological Congress (Tageblatt, p. 4)
and repeated in the later memoir (Anat. Anzeiger, 1902, p. 234) in these words :—
" Die von Forsyth Major als Nervi optici gedeuteten Bildungen lialte ich fiir Bulhi
olfactorii " ; and concerning the true olfactory bulbs and their peduncles, he writes :—
" Diese Bildungen scheinen mir darauf zu deuten, dass hier nicht ein Tractus olfactorius
von enorraer Liitge vorliegt, sondern dass hier ein Tractionsdivertikel der Dura mater
gcbildet wurde, als deren Blutgefasse ich jeue Rauhigkeiten deuten mochte."
Such statements as these miglit be intelligilde if they had come from one who had
studied only the plaster moidd, but Burgkhardt visited the British Museum where he
had every opportunity for examining not only the original cast, but also the skull from
w liich it was made. That he failed to make use of these opportunities must be evident
to anyone wlio has seen the cranium ; for the merest glance is sufficient to show that
Dr. Forsyth Major has correctly identified tire optic foramina (or canals) and the
olfactory fossae (or rather the common olfactory fossa).
If further confirmation of his statements are wanted, there is in the British Museum
a recently acquired skull of a young Megaladapis wliich supplies the desired evidence.
This skull, moreover, amply confirms the pi-edictiou contained in tlie following quotation,
A\bich was made by Dr. Forsyth Major long before a skull of tl.e young 3Ie(jaladapis
was known : —
"When describing the skull of Mcyaladapis, I endeavoured to show that its peculiar low condition is
not primitive, but pseudo-primitive (Fiirbringer), that is to say, that it has been brought about by a
' retrogressive evolution/ or a i-etrograde metamorphosis, if the last term be preferred. If any further
proof were needed for this assertion, it would be furnished by the conformation of the brain, as described
at)ovc, i'or I trust that no anatomist will maintain that this was the primitive condition in Lemuroids.
It may fairly be predicted that, wlien we come to know the skulls of very young specimens of Mega-
ladapis, they will show a much closer approach to the ordinary Lemurid type in the conformation of the
l>iain-cavity and its walls, and the gap between the young and the adult in this respect will prove to be
wider than ])erhaps in any other known Mammal. However, in the Insectivora and most of all in
Centetes, we find also a very great difference between young and adult in the relative size and conforma-
tion of the brain (the brain being even absolutely smaller in the old), whilst tiie least divergence is to be
lound in Marsupials on the one side, in Man on the other, and this obviously for opposite reasons." —
Vroc. Roijal Soc. 181)7, p. 49.
I might add that the pattern formed by the sidci in the l)rain of this young Mecja-
Idilupls is like that of the adult.
The young Ileyaladapis possessed typical Sylvian and postsylvian sulci, and a faintly
marked lateral sulcus wiiich extended farther back than it does in the adult.
The plump olfactory bulbs do not project to so great an extent as in the adult ; and
they are, moreover, separated by a bony septum.
In these respects the young Meffaladapis more nearly resembles the average Prosimite
than the adult does.
-AIOIJI'IIOLOGV OK THE BKAIX IX Till; .Al A .\| .M.\ JJA.
;3fi7
Tilt; hrain of JI<'[ial<i(l<iit/.s must ho. i-egurded as the extreme Ibi-in resulting' from those
retroijjressive changes which aifect the brain in all the IVosimiu'. As one of the m;iny
indications of this, there is the fact, already emphasized by Forsyth Major, that the bi'.-iin
of (ilohilemur is distinctly larger and much richer in sulci tiian tliat of Jfcgaltidapis, in
spite of the fact that the brain of tlie latter should, iu accordance wilh the laws wiiich
regulate the siz(> of the hi-iin, be much the lari^er, Incause Jfrf/al/xhf/t/.s- is Iwice the size
of GlohilcmKr.
The general appearance of the latter is distinctly pitlu^coid, whereas the brain of
Megaladapi.s most nearly resembles those Lemurs, such as the ludrisina; and CItiruniijs,
the brains of which are least Ape-like.
TuE Cerebral Hemispheue ix Tarsius axj) the rROsiMi.E gkneralia'.
'J'lie hemispheres of Tdrshi.s assume a peculiar shape, quite unlike that of any other
mammal, being flattened in an extraordinary manner. A maximum depth of lO-') mm.
Fie. 44.
Fig. \'i.- — '['arsius s/mlriim. The duisal Mspect of (he liraiii. x 2.
Fig. 44. — Tarsius spectruyii. The ventral as])ect of the brain. X -.
Fig. 4.5. — Tarsius spectrum. The left lateral aspect of the hrain. X'at. si/.o.
in a hemisphere wliich is 16'5 mm. long and 95 mm. broad may seem to stultify this
statement ; but it is so deeply excavated in part of the region of greatest depth by an
vmusually extensive orbit and behind it by a deep fossa for the corpora (piadrigemina
and cerebellum, that it becomes converted into a thin irregular plate.
In a vieM' of the dorsal surface of the brain, the cerebrum hides all the rest of tin-
organ except the caudal margin of the cerebellum and its strongl}- projecting floccidar
lobes and olfactory bulbs (tig. 43).
The unusually extensive orbital excavation of the ventral surface of the licmis])hcres —
involving much more than the area usually called " orbital '' — gives to the basal aspect
of the brain an appearance spuriously resembling that of most birds, in which large eyes
SECOND SERIES. — ZOOLOGY, VOL. VIII. 51'
:](;9, PliOF. G. ELLIOT SMITH ON THE
nnd correspondingly enlarged orbits mould the In-ain in a manner not unlike tliat
of Tarsliis. This avian resemblance is still further enhanced by the slender, projecting,
olfactory bulbs. \\n view of the fact that several well-known writers on cerebral
anatomy constantly refer to topographically analogous regions {e. g. the occipital " lobe "')
in tlie hemisplieres of birds and mammals as being homologous, I cannot too strongly
insist that any such jjrej)osterous suggestions are not intended in the above comparisons.]
Gervais has already pointed out * that the brain of Tcwskis approaches nearest
to that of the Galagos in form. By reason of the fact that the smaller Galagina), such
as Microcebm, more nearly resemble Tarsias in size, it is perhaps not surprising to
find an even closer resemblance in the brain-form of the latter two genera. The brain
of Microcebus Smithi is slightly smaller than that of Tarsius f. Although the cerebral
hemisphere of 3Iicrocebus is 1 mm. shorter than that of Tarsias, it seems to overlap the
cerebellum to a slightly greater extent. The maximum depth of the hemisphere is less
than 1 mm. greater in Microcebus. Nevertheless the uatiform eminence (of tlie pyriform
lobe) apjiears to form a much more prominent boss in the latter, because it is narrower
and its anterior margin forms a right angle with the ventral margin of the anterior part
of the pyriform lobe. The smaller eyes and orbits in Microcebus lead to a much smaller
orbital excavation of tlie hemisphere, and consequently tlie shape of the brain is much
more like that of the majority of mammals. Each hemisphere of Tarsias is about
15 mm. broader (at its maximum diameter) than that of Microcebus.
The oil'aciory bulb is placed almost wholly in front of the hemisphere, and is laterally
compressed as in the Galaginse (especially 3Hcrocehus) and the LorisiutB. In both
I if these respects it differs markedly from the Lemurime, Indrisinne, and Chiromys, in
which the bulb is flattened dorso-ventrally and overlapped by the cerebral hemisphere.
In Tarsius the bulb presents a triangular outline in profile, its anterior extremity
forming an acute angle. Its size would be regarded as very small in any other Order of
mammals. A comparison of the size of the bulb with that of Microcebus (which in this
respect may be regarded as typical of the Prosimias) shows that the olfactory bulb is less
than half as large as it is in the Lemurs.
The olfactory peduncle is short and broad, as it is in all the Prosimiie and iu most
mammals excepting the Apes. Like that of the latter, however, and unlike that of most
mammals, the lumen of the peduncle and bulb is obliterated in Tarsius and other
Lemui"s. The short 2)eduncle of the olfactory bulb becomes continuous with a small
])yriform lobe, which is placed wholly on the ventral surface of the hemisphere, forming
approximately tiie mesial third of the great orbital depression. The rhinal fissure
is almost completely obliterated, as is the case iu most Lemurs and, among small
njammals, the Chiroptcra. But its situation is cle.irly recognizable by the difference in
the appearance of the surfaces of the pyriform lobe and the neopallium, and in one
liemisphere a shallow furrow remains to indicate its position. There is a small tuber-
culum olfactorium presenting features such as are common to all mammals.
In spite of the small size of the olfactory bulb, the hippocamjjus is relatively larger
* .Juuin. Ac Zoologic, tome i. p. -'■>. ^
t III Uiis >tatement 1 ii'f'er only to t!nj ^l^el■iIla•l]^ in my J>()s>^(.•^sioIl.
-AroKPiioT.onv of the brain in the .afammalia. 30!)
than in the Apes, Imt smaller than that of tlie othoi- Li'niurs. Although the hippociiuipii-^
is an appendage, so to speak, of llie apparatus of smell ami (Iwiiidles almost to nothing
in smell-less animals, like tlie Odontoceti, the size of the hippocainpal formation does nol
seem to vary directly (as that of the pyriforiu lobe does) with tlie degree of osniatism.
The hippocamjnis is distinctly larger in all Lemurs than it is in the Apes, and Tui-.siii.s
shares this peculiarity of the Lemurs, although in a lesser degree than the other Lemurs.
Thus, if we conijiave the hippoeampal swelling in the lateral ventricle of Tarslns with
that of Hapiilf (the Ape which most nearly ajiproaches it in size), we cannot fail to he
struck with the contrast between its plump columnar form in the Lemur and its slender
proportions in the Ape.
Tiie most that can be seen of the mammalian hippoeampal formation in a mesial view
of the hemisphere is a strip of fascia dentata and flmlnia, and a variable area of alveus-
covered "inverted hippoeam[)us *. In Tanuiis the fascia dentata and fimbria form two
hands of equal and uniform l)rea(ltli, not separated by any " inveited hippocampus."
These bands extend almost vertically upward, with only a very slight inclination forward,
and the fascia dentata passes directly to the splenium of the corpus callosum, without
the formation of any subsplenial flexure. Such a phenomenon does not occur elsewhere
among the Lemurs, but we find a parallel in tlie case of the Sloths f, in the brain of
Avhich there is, as in Tursiiis, an unusually short corpus callosum. 'J his is indicative of a
very primitive state.
The supra- and pre-callosal vestiges of the cephalic extension of the hippoeampal
formation call for no speci-il mention, because in Tarsius, as in all Primates, these
structures are extremely insignificant and difficult to demonstrate. 'Uhe hippoeampal
tubercle is relatively large in Tarsiun, seeing that it involves about one third of the
length of the hippoeampal formation.
The corpus callosum is remarkable (in an Order in which this commissure is dis-
tinguished for its great length) in being unusually short. It measures only 5-5 mm.
in a hemisphere 17 mm. long, whereas the corresponding proportions in a Lemur tiiongo-
are 17-5 : 39, in a Ni/cticchiis tardigradus 14 : 31, and in a Chironti/s 22 : 45. Its
thickness is remarkably uniform throughout, the splenial and genual extremities being
only very slightly thicker than the body, ^^'hereas in Lemur and most of the Prosimiaj
the splenium and genu are twice or even three times the thickness of l^he body. The
corpus callosum is also more obliquely placed than it is in Lemur. It exliibics a very
slight, regular, dorsally-directed convexity, and its slightly dependcmt anterior extremity
shows no sign of being bent backward to form a rostrum, such as we find in the xVpes.
In all these respects the brain of Tarsius approximates to the primitive condition
exhibited in the Insectivora.
In consequence of the relatively slight backward extension of the corpus callosum,
the psalterium or hippoeampal commissure is short and plump. In all other Trosimia',
as in most mammals, the great bulk of the psalterium is collected at the upper end
• Viilf .Tonni. .Vnat. and I'liyn. vol. xxxii. 1>!_)7. \'\k liO & 51.
t •• Jlraiu in llic Edentata, " Trans. Linn. Sue. ser. 2, Zool. vii. lSi)0.
54*
:]70
PKOF. G. ELLIOT S^flTH OX THE
of the lamina terminalis just above the anterior commissure, and is linked to the ventral
surface of the splenium of tiie corpus callosum by a thiu attenuated band, composed of
scattered commissural fibres. In the Hapalidae these scattered tiljres are \\aiiliny,
so that the psalterium is separated from the corpus callosum l)y a -wide gap ®. In
Tarsius there is little or no attenuation of the dorso-caudal part of the psalterium. A
])arallel for this is found in the brain of the Bradypodida^.
There is a septum lucidum composed of two thin sheets enclosing between tiiein
a cavum septi, which is freely open below in tlie Avide interval between the genu of the
Fit; 46.
Fig. 4ii. — Tdrsi'.i^ xjiic/i'iiiii. Surface exposed by a mesial sagittal section, x 4.
Fig. 47. — 'J'ar.iiii.^ s/ii'(iruiii. Diagram representing the mesial surface of the right cerebral hemi-!phere. x 4.
corpus callosum and the psalterium. In spite of a very generally-expressed belief to the
contrary, this is the condition whicli prevails in the vast majority (2)ractically in all) of
the Eutheria.
The anterior comniissure is slightly more than 1 mm. deep and just less than 1 mm.
thick (in tiie horizontal plane). For a Primate of such small ditneusions, these measure-
ments must be regarded as large, and es2:)ecially so when it is recalled that Tarsi m is the
most micvosmatic of Lemurs, and therefore the individual in which such a phenomenon
would a. priori be least expected. This means that the neopallial clement in the
* Vide Joiirii. Anat. and Thys. vul. xxxii. 1*07, fig. i?'i. p. ol.
MOIiPlIOLOGY OF TllH HHAIX 1\ TJIE \l A^IMATJA. :;71
commissure is relatively large. Aiul this is yet a further indication oi" the primitive state
of the braiu ol' Tarsim.
The anterior part of the mesial surface of the h-nuis[)here is quite smooth and flattened
against the correspond ing surface of the other heuiispliere. This flattened area. i)roloQg<'d
backward above the corpus callosum, extends for more than 6 mm. in the caudal
direction beyond the splenium of the corpus callosum as a baud less than 3 mm. Avide.
The whole of this flattened area is entirely devoid of sulci, no trace of the intercalary
(calloso-margiual), genual, or rostral sulci, which are found in some Lemurs, being
detected here. Ziehen says* IJiat in the region abcne tlie corpus callosum there is a
deep furrow on the mesial surface, ^\ liieii is probably nul a vascular groove, but is a true
fissure running parallel to the corpus callosum. In all of the four hemispheres of Tamlns
which I have examined this so-called sulcus a of Ziehen is nothiug else than the
impression of the anterior cerebral artery, as it oblitjuely crosses the surface of the
hemisphere, after having l)ent around the genu of tlie corpus callosum. A similar
impression may l>e fouml in any mammalian l)rain, as I have already pointed out in tlie
case of Oj-iiil]ioi-]ii/iicluiis\.
In a cerebral hemisjdiere 17 mm. long, 8 mm. (or almost half of its length) lies behind
the splenium of the corpus callosum. Such a marked caudal or (as it is commonly
called) occipital extension of the hemisphere is found nowhere else among the Lemurs,
and a parallel for it must be sought among the Apes, where such an exaggerated
backward prolongation is present in the Hapalidre and tiie smtdler Cebidae. It is a
si<Tniticant fact that this Pithecoid condition is found in a brain wbich in manv other
respects exhibits more primitive features than the majority of the Frosimite. We have
already seen that a narrow upper strip of this postsplenial area of tlie mesial surface is
flattened against tlie corresponding surfacs of the othei" hemisphere. The rest of this
region is deeply excavated in a \e\'\ irregular manner in adaptation to the shape of the
corpora quadrigemiua and cerebellum, wiiich are jjartly overlapped by the occijjital
prolongation as l)y a cap.
Immediately below (or to the lateral aspect of) the flat area there is a deep cup-shaped
depression about 3 mm. in diameter, placed about 2 mm. beliind the splenium. This
is produced by the anterior quadrigeminal body. In this depression there is a triradiate
arrangement of deep siilci diverging froLu its centre. The calcarine sulcus extends
downward and forward toward the hippocampal Assure ; the retrocalcarine extends
horizontally backward ; and the paracalcarine extends upward, but does not quite reach
the flat area. The other two sulci, however, just minage to cross the rim of the cup-
shaped depression.
By comparison with otlier Pi'osimian brains, it will be shown later on that tlie
sulci a and y prolxxbly re[)resent the true calcarine sulcus (/ c. that called " anterior
calcarine" by Cunninghaui), and that the sulcus /3 is the i-etrocalcarine (Cunningham's
"posterior calcarine"). All of these sulci are very deep and completely involute the
mesial wall of the very large pithecoid posterior cornu of the lateral ventricle. In other
* Archiv f. Psycli. P.d. xxviii. p. l.tcO.
t " Purtbcr Observations on the Braiu in thu ilonotromuUi," Journ. Auut. and I'll}--., vol. xxxiii.
372 PEOF. G. ELLIDT SMITH OX THE
Avords, there is formed a calcar avis of such i^reat iivoportions as to closely resemble in
shape and size that of the smaller Apes. This calcar is larger than that of the other
Prosimia', although the condition found in the Galaginae nearly approaches it.
It is noteworthy that in Ziehen's memoir on tlie Lemur's brain this calcarine sulcus —
the only true sulcus found in the brain— is entirely ignored, although reference is made
to a large number of spurious sulci. If this significant sulcus had really been absent, as
Ziehen's account would lead one to suppose, this would have placed the bi-ain of Tarsias
in a category quite apart from that of all the other Lemuroidea (and in fact all Primates),
instead of adding vet another convincing testimonv to tlie close Prosimian affinities
of Tarsiiis.
About midway between the calcarine sulcus and the ventral border of the hemisphere
there is a very deep irregular notch extending backward on to the caudal margin of the
liemisphere. When the brain is in situ in the skull, this deep depression is occupied by
the bony case of the strongly projecting fiocculai' lobe of the cerebellum.
Ziehen* represents two separate depressions in this situation: one on the mesial
surface, which he refers to as /3, and the other he calls " die Einbuchtung der untereii
Contours des TemiJoro-occipitallappeus." They are obviously both parts of the one
lloccular fossa. Although when describiua; the Tarsius-brain Ziehen inclined to the
\ie\\ that the sulcus /5 is merely a depression caused by a bony projection, yet in the
same memoir we find this writer discussing which of two sulci in the hemisphere of
Xijcticehus (p. 906) represents the "sulcus /3 in Tarsiiis" \ Then he adds, " Am
Avahi'scheinlichsten ist mir, dass /3 wenigstens z. Theil audi dem Yereinigungspunkt von
p, o, luul TT [the calcarine group of sulci] entspricht." If this suggestion is correct,
Ziehen's drawing (tig. 2, p. 900) is very misleading and erroneous, for the relationship of
his sulcus /3 to the floccular fossa (his " Einbuchtung") is very different to that of tlie
true calcarine sulcus.
Just below the retrocalcarine sulcus there is in one of my specimens a shallow
horizontal furrow, which crosses on to the caudal margin of the hemisphere. As it is
very shallow and lodges a branch of the posterior occipital artery, it is possible that it is
caused by the latter. In neither hemisj^here of ray second specimen is there any such
sulcus.
The narrow cleft between the optic chiasina and the tuberculum olfactorium, which
contains the representative of the locus j^t'iforatus of other mammalian brains, is
prolonged transversely outward across the pyriform lobe as a vallecula Sylvii. Opposite
the outer end of the latter a furrow extends outward (upward) and slightly forward
across the orbital excavation of the hemisphere, which it divides into two approximately
equal parts. It occujjies the situation of and probably represents the Lemurid Sylvian
fissure. It does not extend so far up as tlic dorsal lip of the orbital excavation, so that
it is not visible on the dorsal surface, although an indentation of the margin (tig. 13)
indicates its situation. A comparison with the condition found in Microcehm shows that
this must represent the Lemurid Sylvian or suprasylvian sulcus. When we remember
that the area which is usually called " orbital surface " lies wholly in front of the
* Arch. f. Psvchiat. xxviii. ISHG. p. OUU.
MOHPIIOLOGV OF THE BRAIN TX THE ^FAMMAMA. H?.''
Sylvian tissuve in othei- brains, it will be seen that tiic position of tliis " Sylvian ttssiire"
ill Tarsliis is very peculiar. The further discussion of the sipjniflcance of this furrow
w ill be more intelliij:ible if it bo postponed until the nature of the Sylvian lissiire of tlie
other Lemurs has been considered.
Apart from tiiis sulcus, the rest of tli(> cranial as|);»ct of tiu^ hemisphere is smooth ami
devoid of sulci.
There is a very slii^-ht llattenini;', unworthy of the u;un(> of furrow, just behind and
parallel to the caudal margia of the orbital excavation. It may possibly represent the
postsylvian sulcus.
,\n even slig'liter flattening is found parallel to the mesial plane in a situation wliei-e
the intraparietal (lateral) sulcus is found in other Lemurs.
\n view of the considerable dillerences whicii are shown to exist by comparing the
brains of various Lemurs, the resemblance between the brains of Tars/us and the smaller
Galaginie is a fact of considerable significance. The peculiar features of the calcarine
group of sulci so strikingly re2:)roduce those found in all the other Prosimiie, and in no
other mammals of a similar size (with the possible exception of the Edentate Mani-s),
that they alone constitute an important indication of the closeness of the bond of affinity
Avliich unites Tavsiits to all the other Lemurs.
It is also jieculiarly instructive to note that the chief diff'erences between the brain of
Tarsiits and the other Lemurs occur in the commissural region ; and in respect to this
Tarsins approximates much more nearly to the primitive nrammalian type than do the
other Prosimite.
As in Microcebus, Tarslus possesses only one sulcus other than the calcarine grou))
(and, of course, the hippocampal fissure). This other sulcus is the Sylvian or, perhaps,
its suprasylvian element. In these respects these small Frosimian brains resemble those
of the lowlier Apes.
In Galago small lateral and orbital sulci make their appearance on the outer aspect,
and intercalary and rostral sulci on the mesial surface. In the Great Galago the lateral
sulcus becomes longer, and a coronal sulcus makes its appearance ; the sulcus/ and th(;
postsylvian furrows also may become developed.
In tlu' larger Lemurs diagonal postlateral, collateral, and other furrows may develop.
TlU: BUAIN-SIKM.
By this term I mean those parts of the brain which remain after removing the
cerebral hemispheres and cerebellum.
In my Memoir on the Brain in the Edentata I pointed out that these parts are subject
to much less variation in the ditfereut mammalian Orders than are the cerebral hemi-
spheres and cerebellum. A careful study of these regions in T<u->iius, Lemur, Microcebius,
and, by macroscopic means only, in Chiromys, Galago, and several other Prosimiie, has
revealed no features which are not equally peculiar to all mammals, so tliat no lengthy
consideration of these regions is necessary.
There is practically nothing concerning these parts of the brain in the literature of
37 !• PROF. G. ELLIOT SMITH ON THE
the I'rosimiiX', for the feAv writers * who refer to any region other than the cerebral
hemispheres do not record anything worth recalling in reference to the hrain-stem.
There is nothing in the walls of the third ventricle, the pineal body, the ganglion
habcnnlae, the habennlar commissure (commissura superior), the hypophysis, corpora
mammillaria, the interpeduncular ganglion, the crura cerebri, pons Varolii, and corpus
trapezoidea to distiuguish them from tlie great majority of mammals and from the
smaller Apes. Many of the structiu'es eniuuerated in this list are found to be consider-
ably modified in Man and the liigher Apes in comparison with the majority of mammals.
Tims the corpus trapezoides becomes gradually covered by the pons Varolii ; the inter-
peduncular ganglion becomes deeply buried between the crura cerebri and pons, and is
unrecognizable as a distinct ganglionic mass; the corpora mammillaria become more
distinctly separated the one from the other. But these changes occur in the transition
from the smaller to the larger Apes, and hence it is not surprising that, like the former,
the Lemurs should preserve the common mammalian features.
There are only three features of the Prosimian l)iain-stem that call for special note.
Tliese are the relative size of the corpora quadrigeaiina ; the projiortions of the optic
thalamus and its external (anterior) geniculate liody ; and the olivary body in the
medtdla ol)longata.
Tlie general featitres of the corpora quadrigeraina are remarkably constant in all the
]\Ieta- and Eutheria. The features which are sttbject to most variation are the relative
proportions of the two pairs of collicttli or qtiadrigeminal bodies, and the size of the
ti-actus pedttncularis trausversus.
Gustav Retzius has recently published some excellent representations of this region
in the brains of Homo, Aiitliropopithecus, Sim.ia, Ursus, Lnlra, Phoca, Bos, Equus, Ovis,
S'lis, Lepus, and Macropns f ; Ziehen has described its features in the Marsupialia and
IMonotremata % ; and I have briefly described its appearance in the Edentata § and
Monotremata ||. Since then 1 have carefully examined this region in representatives
of every mammalian Order with the special purpose of comparing it with the Prosimian
mesencephalon. The most important result of this study is graphically demonstrated in
the acconijmnying series of diagrams, which represent the dorsal aspect of the thalamic
and mid-brain regions in Lemur, Tarsim, and a heterogeneous collection of mammals
including Cercopithecna, Fclis, Procavia, Lepus, Lasijpus, and Macroscelides, which
r(>present every phase of the changing jiroportions of these ])odies in the whole
Mammalia. All the diagrams are ma<?nihed in the same desTree.
The most striking feature of the diagram representing the condition in Lemur is the
relatively enormous size of the optic thalamus and the smalhiess of the corpora quadri-
gemina. As we descend the mammalian scale there is a progressive diminution in tlic
* Burmeister (2'a/-s;((4-), Owen ^C7(i/-ojn(/sX Milne-Kdwnrds ( liidrisiuie), Ondi-muns ((,'/(!/-07(iy6'), and Flatau and
Jacobsohn {Jjemxu- macaco).
t Biolog. Unters., N. F. Bd. viii. uo. 5, Taf. IG & 17.
J Jenaische Denkschrift, 1897.
§ Tran.i. Linnean Soc. ser. 2, Zool. vol. vii. ISfl'.
II Journ. Anat. and Phvs. vol. xxxiii.
MOEPHOLOGr OF THE BE A IX IN THE MAMMALIA.
0/ J
size, both absolute and relative, of the optic thalamus. In ihc lowly Metatheroid
Insectivore Ilacroscelides the dorsal area of the thalamus is little if anv biiraer than
the anterior quadrigeminal body.
Fig 48 '^'
(a)
cqp.
cqa.
cqp.
cgp.
cqp
C9P-
cqp.
A series of diugiiinis rcpie?eiitii)g tlio dorsal surface of the tbalamic and mesencephalic regions in {a) Lemur,
(h) Tarsias, (c) Cercopilheais, {d) Fdis, (<>) Frocavia, (/) Lepiis, (g) Dasypvs, and (h) Macrosc elides. All
inagnitifd 2 diameters.
In these respects Lemur distinctly conforms to the peculiar condition of the Primates,
and difter.s most markedly from all other mammals.
It is equally signiticant that in a large-eyed mammal like Lemur the anterior quadri-
geminal bodies are so small as they are, not only relatively, but absolutely. They aro
certainly much bigger in Tarsius, but in this creature the eyes attain such exceptional
dimensions.
SECOND SERIES. — ZOOLOGY, VOL. VIII. 55
376 PEOF. G. ELLIOT SMITH OX THE
In Levmr (and all the rrosimice) the lateral (anterior) geniculate body becomes much
larger and more distinct than it is in most mammals : and this feature it also shares with
the other Primates.
It might perhaps l^c imagined that the larger dimensions and more distinct separation
of the lateral geniculate body from the rest of the thalamus are due to the fact that the
optic tracts are unusually large in all the Lemurs. But this cannot be the whole
explanation, because in many other large-eyed mammals (see many llodents, Ungulates,
and others) there is no such distinctness of the external geniculate body as we find in all
Lemurs and other Primates. This phenomenon must be due to an increased perfection
of tlie cortical path for visual impulses, rather than to a mere enlargement of the lower
optic tracts.
The four quadrigerain-al bodies rise to the same level, and resemble in shape those of
the Myrmecophagidte more nearly than those of any otiier mammals (excepting the
other Primates). There is a well-defined tractus peduncularis transversus, but it calls
for no further mention.
If the region of the medulla oblongata in the Lemurs be compared with that of other
mammals of a corresponding size, the only noteworthy distinctive feature of the former
will be the relatively greater prominence of the olivary body. It is, however, somewhat
smaller and less prominent than it is in most Apes, the chief olivary nucleus being
smaller in proportion to the internal accessory olivary nucleus in the Lemurs than
it is in the Apes.
The Calcakixe Sulcus and the Occipital Prolongation' of the
Hemisphere.
The foregoing notes will have made it clear that a triradiate group of three sulci —
which I have called calcarine, retrocalcarine, and paracalcarine respectively — form an
iirrangement which presents identical feaiures in all the Prosimine. No one has ever
questioned the identity of the furrow labelled calcarine with the sulcus so-called in tlie
Apes. The sulcus which I have distinguished Ijy the name "paracalcarine" is regarded
l)y most writers as the representative of the parieto-occijiital sulcus in the Apes.
Plower was, I believe, the first to suggest this view ; but afterwards renounced it in
lavour of the hypothesis that the paracalcarine sulcus represents one oL" the limbs of
bifurcation which the calcarine sulcus presents in most Apes. When I began writing
this memoir I subscribed to the latter view, chiefly because the parieto-occipital sulcus
is absent in several of the lowlier Cel^idee and all of the Ilapalid^e ; and it seemed therefore
to be a furrow Avhich becomes evolved in the Apes : but in develo2:)ing the argument
which is here unfolded, I became convinced that there may be another and more probable
Avay of regarding this matter. None of the writers wlio call the paracalcarine sulcus
"parieto-occipital" seem to have considered the possibility of it being any other
furrow.
Pv-ecognizing the identity of the calcarine sulcus in all the Primates, we may pass on
to the consideration of its relationship to the lateral ventricle, with a view to determine
MOEPHOLOGY OF THE BRAIN IN THE MAMMALIA. 377
if there be any q\ialitativc diffcveace in Ihe nature of the occipital prolongation of the
hemisphere in Lemurs and Apes. For the extent of this region of the hemisphere, or,
rather, its relation to the cerebellum, is commonly supposed to be a feature of the
iitmost importance as an index of cerebral development, even by writers who have
devoted much labour to the study of this part of the brain.
This will lead not unnaturally to the enquiry as to whether the oxlcarine sulcus is
found in any mammals other than ihe Primates.
The most conflicting statements are found in the literature (quoted earlier in the
notes) regarding the posterior cornu of the lateral ventricle.
Burmeister (1846) described an extensive diverticulum of the ventricle in Tarsim,
and about the same time Vrolik denied its presence in }\ijcticebu,s \Slenops\. Both ol'
these statements are true.
In 18C2 riower thought that he had found a postx'rior cornu and calcar in Lemur
/ulcus {^nigrifrovs] and also in a Galago of unknown species ; but while his monograph
was being revised in proof he became sceptical and added in a footnote: "A further
examination of this specimen [a brain of dalago in tlie Museum of tiie Charing Cross
Hospital], and of the brains of some allied genera, leads me to doubt whether the above
described ' cavity' in the posterior lobe existed before dissection, the length of time that
it had been in spirit having greatly facilitated this process. If it did not, it will justify
the statement of the absence of the hippocampus minor by anatomists who have looked
at this structure only in its relation to the postei*ior cornu, but at the same time will
afford a further illustration of what I have endeavoured to show .... viz. : — that the
part of the brain to which this term [calcar] has been applied can exist indej)endently
of the ventricular cavity " (Phil. Trans. 1S02, p. 198, footnote). I have quoted
this very interesting and luminous note in full, not only by reason of its important
bearing upon tiie question at issue, but also to express surpri.se that its author shoukl have
suddenly stopped on the way to the full interpretation of tlie morphology of the calcar
and calcarine sulcus, and have categorically denied its existence in the Cat's bi-ain, after
thus removing all the obstacles to such an obvious inference as to the identity of the
so-called " spleuial '' sulcus in the Cat and the Prosimian calcarine. In the same year
he wrote a second memoir [which, however, was not published until 1800 (Trans.
Zool. Soc. vol. V.)], in which he makes the statement that it is imjiossible to determice
whether or not the posterior cornu exists in }\ijcticehus tardigradus [Stenops Javanicus]
(p. 100).
In the memoir on rhe brain of Chiromys, which was published in the same year
(Trans. Zool. Soc. vol. v.), Owen seems to represent a small posterior cornu (in fig. 5).
This fact is all the more significant if it is intentional, as the author was engaged at the
time his memoir was Avritten in a heated controversy, in which he maintained that the
posterior cornu and the calcar were human characteristics.
In Milne-Edwards's contribution to the History of Madagascar ((2) tome A-i. 1876,
p. 206) the author refers to Flower s demonstration of tiie existence of a calcar in Lemur
and Galago (quoting Piiil. Trans. 1862, p. 197), and states that he has confirmed this in
the case of the forms studied by Flower, but denies its (calcar's) existence in Indria,
378 PKOF. G. ELLIOT SMITH ON THE
Propithccits, and Acaliis. The writer lias apparently overlooked Plower's footnote
(quoted above) which clearly shows that a posterior cornu is not necessary to the
existence of the calcar. The latter must be present in tlie Indrisinse (because a calcarine
sidcus occurs in this subfamily as it does in the I,emurinffi) even though the posterior
cornu may be absent. For this is obviously what the author means to imply. His other
statements regarding the posterior cornu and calcar avis (on p. 205) are contradictory
and their meaning is not altogether clear.
The only other writers (so far as I am aware) to refer to the posterior cornu in the
Lemurs are Flatau and Jacobsohn (' Handbuch,' p. 189) ; and as they have obviously not
read any of the literature quoted above (excepting Burmeister's remark concerning
Tars/'iift) they dismiss this, the most crucial feature of the Prosimian brain, in that
casual manner, which is unfortunately the rule rather than the exception in their barren
and misleading work. Thus they dispose of the whole question in the case of Lemnr
macaco in the words : — " Der Seitenventrikel ist bei den Halbaffen von gleicher Eeschaf-
fenheit, wie bei den wahren Affen und audi von den gleiclien Gebilden begrenzt "
(p. 1S9), as though there were no problem to be settled. Moreover it will lie shown that
this observation is misleading, if not quite erroneous. The only other Prosimian brain
which the authors examined was one which they label " Stenops gracilis," but wliich
obviously belongs to " Nycticebus tardigracbi.sy Now although the interesting problem
as to the existence or absence of the posterior cornu in this genus had been definitely raised
and loft unsettled {vide supra) by Plower (whose numerous and important contributions to
the Comparative Anatomy of the mammalian brain these writers of a text-l)ook on this
su])ject almost wholly ignore), Platan and Jacobsohn merely state (in tlie paragraph
corresponding to that quoted above regarding Lemur) : — " Die iibrigen Vorhaltnisse am
Gehirn des Stenops sind so iihnlich denen am Gehirne des Maki, da^-s cine l)esondere
Eesprechung unnotig erscheint" (p. 199). But, as in the case of Lemur, they give
tables of measurements, among which figures " Abstand der vorderen Spitze des
Vorderhornes vom hinteren Pol des Hinterhornes " (pp. 199 and 189) *.
Ziehen, Beddard, and all other recent writers do not even so much as refer to the
subject.
After carefully examining the specimens concerning which the above-([uoted state-
ments of Plower's were made, I felt convinced that an undoubted patent posterior
cornu exists in the genus Lemur. I subsequently made dissections of the brains of
Nycticebus, Microcebiis, Tarsius, and two other specimens of Lemur, all of which had
been in preservative solutions for long periods ; and the results seemed to confirm my
earlier conviction. But recently I examined the fresh brain of a Lemur f ulcus, and, to
my great surprise, found mo trace of any patent posterior cornu. Since then I have been
unable to find any posterior cornu in three other fresh brains of various species of Lemur
and a fresli brain of Nycticebus. In view of these unquestionable facts, and of the
• This reference to a " Hinterborn " is, liowevor, meaningless, for tbey use the term in referring to the brain of
all mammals.
MOEPJIOLOGY OF TJIE BRAIN I\ TJIK MAMMALIA.
Of'
equally ticcisivc remarks of Milne-Edwards*, and Flower's hesitation in adopt iii;^' thc
view -which seemed so obvious, one naturally becomes sceptical of the existence of such
a ventricular diverticulum in any Prosimian brain.
If a horizontal section be made through tlie cerebral hemisphere of any J.emur so as
to pass just beloAV the level of the corpus callosum, the pericalcarine region will ]>res(mt
the appearance represented in the figures 1!) {Xt/c/.icebi(s) and ."iO {Lemur J'uIcks).
The hippocampus is prolonged mesially into continuity with a long pallial folium, which
extends almost as far as the mesial plane and then becomes sharply bent upon itself so
Fig. 49.
Fig '■-0
sc.
Fig. 41.1. — yi/ctii-eliii6- tardhifiiiliix. llciri/.outiil st'ction through the cerebral hemisphere
iinnicMliatfl}- bcknv the ecirpus callosum. .x -.
I'ig. SO. — Lemur faints. Part of a eorrespouding seetiou. x 4.
Fig. 51. — C'mro'pitJieciis snhti'iis. A. eorrespondiiig section. x -.
as to form the slightly oblique anterior wall of the deep calcariue sulcus. At the
bottom of the calcariue sulcus the pallium again becomes acutely fle.\ed and forms
the posterior lip of the calcariue furrow. It is of the utmost importance to recognize
at this stage that the anterioi- wall of the calcariue sulcus forms a part of the bouiulary
* Only so far as the Iiidrisiu;i! arc concerned ; for, in view of these results of my recent re-examination of the-
genus l.tmur, 1 fail to understand the significance of Milne-Ed wards's reference to this genus (v(V/« su/>i-(i).
380
PEOF. G. ELLIOT SMITH ON THE
of the lateral ventricle, its medullary layer bein"- part of the vcutricular liuing- ; whereas
the posterior wall of the sulcus is far removed from the ventricle, and its medullary layer
is not free but is fused to the general medullary mass of the hemisphere.
No part of the pallial lining of the retrocalcarine and paracalcarine sulci takes any
direct share in forming the walls of the lateral ventricle.
If we next examine a similar section in the cerebral hemisphere of an Ape, such as
Cercopilhecits (fig. 51), essentially the same state of affairs is revealed. The posterior
wall of the calcarine sulcus is far removed from the ventricle and its medullary layer is
fused to the general medullary mass.
As the result of the greater obliquity of the calcarine sulcus and of the smaller size
of the hippocamj)Us, a much larger area of the anterior (or lateral, as it has now become)
wall of the calcarine sulcus is exposed in the ventricle and forms the " calcar avis."
The only difference Ijetween the conditions in the Lemur and the Ape is a quantitative
and not a qualitative one. As the result of the larger dimensions of the neopalliiim, the
Fig-.52.
The left cerebral hemisphere of (a) Nycticelvs, (h) Microcdms, (c) Tarsina, and {iJ) Ihipalc, dissected
to show the hippocampus and the calcarine eminence (.<■) \not the true calcar in (/ and h'].
hemisphere has extended further backward and has produced a posterior diverticidum
of the lateral ventricle, in the mesial wall of which the calcar is found. In the Lemur
the homologue of the calcar is also found, even though there is no posterior cornu of the
ventricle, but it fiices the alveus of the hij^pocampus, so that no bulging calcar can be
seen Avhen the ventricle is opened.
If the lateral wall of the hemisphere be dissected away in the brain of a Lemur which
has been toughened in some preservative fluid, a line of cleaA'age readily extends
backward from the posterior angle of the ventricle between the dense medulla of the
calcarine cortex and the looser medullary matter in contact with it; in this way the
observer can readily be deceived into the belief that a jiosterior cornu exists and that
a rounded calcar (iig. 50, .;•) projects into it. The region .r, however, is not the
homologue of the true calcar, as a comjjarison of figs. 50 and 51 will at once show.
The appearance of the " calcar," so exposed in the brains of jS'i/cilcehtis, Jlicroccbiia,
MORPHOLOGY OF TUB BRAIX IN THE MAMMALIA. 381
Tarsius, and Uapale, is shown in the accompanying- drawings (fig. o2). In Tarsias and
JIapale the swelling exposed is partly and perhaps wholly a true calcar, like that of
Cercojnthecus.
In the hrain of the Apes, as also in Lemurs, tlie walls of the paracalcarine and
retrocalcarine sulci do not come into relationship Avith the ventricle.
If next we practice a corresponding section in the hrain of a Dog (fig. 53), a state of
affairs is exposed which is either identical witli (hat of tlic brain in th(? Primates
or presents the closest resemblance to it. Immediately to the caudal side; of the
liippocampus the neopalliiun behaves in tlie same manner as it does in the brain of
Lemurs and Apes ; but tlie deop, oblique sulcus is not usually called " calcarine," but
Fig. 53.
calr
— sc.
Can is film Uiaris.
A section analogous to these represented ia tigs. 50 aud 5L x i*.
by Krueg's title " splenial." A glance at this figure will at once show that, unless there
be some overwhelming argument to the contrary, we must regard this part of the splenial
sulcus of the Dog as the representative of the calcarine sulcus of the Primates. It is
commonly argued that there can be no calcarine sulcus in the Carnivora because there
is no posterior cornu ; Ijut the same writers do not deny the calcarine nature of the
analogously-placed furrow in the Lemtirs, even though tliere is no posterior cornu there.
We may therefore, at any rate as a working hypothesis, refer to this part of the Dog's
splenial sulcus by the name " calcarine." Now the conditions which are found in the
Dog also ^n-evail in every Carnivore, without exception. It is therefore very surprising
to find Plower speaking of "the absence of anything resembling the calcarine sulcus"
in the Cat's brain in the same memoir in which he made the important observation
(quoted above) that the existence of a patent posterior cornu is not a necessary condition
of the presence of a calcar, and therefore of a calcarine sulcus. Even if the calcarine
sulcus should ultimately prove to be not homologous with any part of the splenial
furrow, it is clearly erroneous to say that the latter does not " resemble " it. In the
case of certain Carnivores, such as the Seals, a definite posterior cornu of the ventricle
is found, and in some cases, c. g. Phoca, it reaches large dimensions : in these animals
the "splenial bulging" in the lateral ventricle so closely "simulates " the "calcar'' of
the Primates that there can be no reasonable doubt as to their identity. This fact was
well known early' in the last century, for Serres's observations, that " the hippocampus
minor [calcar] has hitherto been observed only in Man," which continue ". . . . I have,
382 PROF. G. ELLIOT SMITH OX THE
liowever, found it in the Apes and Seals [cliez les Singes et les Plioques] " *, gained a
wide currency in the text-Looks of the period t, and have even persisted in certain French
treatises np to tlie present day. And yet this important fact has been wholly ignored
in the discussion of the possible homologues of the calcarine sulcus in other mammals.
[In making this statement I am not unmindful of the fact that in the memoirs of
Murie on the brain in the Seals and in the Manatee an anthropocentric nomenclature is
adopted and a furrow on the surface of the hemisphere is called " calcarine " ; Murie
also describes a posterior cornu and a calcar in Otaria and Manatus7\
The calcar was rediscovered and described in the brain of the Seals by Fish in 1898 ;
but this writer so signally failed to appreciate the morphology of this region of the brain
as to entertain a doubt as to wlietlier the sulcus, which produced this calcar, ought to
l)e regarded as the calcarine or the parieto-occipital ! %. He makes the following
quotation from Wilder's " Anatomical Technology " : — " Between the ordinary Carnivora
and the Monkeys are two groups whose brains should be studied with especial care ;
the Seals liave a rudimentary postcornu and occipital lobe, and these parts are said
to be developed in the Lemurs which have aflQnities with both the Carnivora and the
Primates " (p. 80).
Although Fish quotes the statements of Tiedemann, who represents Fhoca as lacking
a posterior cornu, he appears to be ignorant of the above-quoted observation of Serres,
in spite of its Avide cui'rency in such works as those of Cuvier, Leuret and Gratiolet, and
Topinard. His suggestion that the splenial sulcus of the Piunipedia may possibly
represent the parieto-occipital sulcus becomes all the more amazing when he makes such
definite statements as: — " [^Phoca'] shows a jjostcornu relatively as large or larger than
in the primate brain, with a distinct calcar or hippocampus minor in which a portion of
the splenial appears as a total fissure " (p. 88).
I have confirmed all these statements of fact in the Seals and prepared a dissection
of this region in Phoca for the Musevim of the Eoyal College of Svu'geons (Si^ecimen
D. 377, Physiological Series).
Since Krueg carefully described and named the " splenial " sulcus in a large number
of Carnivores and Ungulates, no one has questioned the identity of the furrows so-named
in the two Orders. The different position of the furrow on the hemisphere in the
Ungulata modifies its obliquity so that it does not so closely resemble the calcarine
sulcus in the Primates as that of the Carnivora does. Xevertheless the relationship of
its two walls to the lateral ventricle and its depth (in comparison with neighbouring
sulci) would suflBice to show its identity, even if we w^ere not acquainted with its
* ' Anatomic Coiuparee du Cerveau,' Paris, 1826, t. ii. p. 470.
t Vide Leuret, " Aiiat. Comparec du Systeme Nerv." tomo i. 1839, p. 402, and a fuller account in Gratiolet "s
edition of the same work, tomo ii. 18.57, p. 74.
t P. A. Fish, " The Praia in the Fur-Seal, Callorhinus iirsiniis ; with a Comparative Description of those of
Zaiophus, Phoca, Ursas, and MonacJius," Journal of Comparative Neurology, vol. viii. nos. 1 and 2, July 189S, p. 79.
" The conditions .... might naturally suggest homology with the ape and human calcar and that the splenial
fissure, in this seal possessing a postcornu, might he homologized with the occipital [the contest shows that this
term undouhtedly refers to Uio parieto-occipital sulcus] or calcarine fissure in Man,"
MOEPHOLOGT OF THE BRAIX IX THE MAMMALIA. 3So
(leA-elopmontal history, wliicli thorouglily e.stablishes the homology. In some of the
larger Ungulata, for example Cameliis, tliere is a small posterior cornu into which
the " spleuial " eminence projects just as the calcar hulges into the Primate ventricle.
In the Cetacea the splenial sulcus is obviously identical with that of the Carnivora and
Ungulata, and its resemblance to the calcarine sulcus of the Primates is enhanced in
some genera, e. g. HijperoodoH, by its relation to a distinct posterior cornu of the
ventricle, wliicli has been described by Ziehen *.
During my investigations on tlie brain in the Edentata, the results of which were
presented to this Society almost four years ago, I was very much astonished to lind
a well-defined swelling on the mesial wall of the lateral ventricle of Orycteropus which
simulated the calcar avis. ]\Iy astonishment was due to the fact that at that time I was
not acquainted with any of the above-mentioned Avritiugs and knew only of the teaching
of such anatomists as Plovver, Turner, and Cunningham, who have so formally and
categorically denied the possibility of bomologizing the calcarine sulcus of the I'rimates
with the spleuial furrow in other mammals. Nevertheless the mass of evidence in
lavour of the identity of these two furrows, which a study of the Edentata and other
mammals yielded, seemed to me to be so overwhelming, that I adopted this view even
though it is in direct contradiction to the \iews of those anatomists, from Avhom I have
acquired much of my knowledge of the mammalian brain f.
Among the heterogeneous collection of Edentata, there are found all stages in the
form of the calcarine sulcus intermediate between the Carnivore condition on the one
hand, and the simpler form fouiul in the Chiroptera, Marsupialia, and Insectivora {i. e. if
GaleopitJiec/ts can be called an Insectivore). In no true Insectivores is there a calcarine,
or "splenial" (as most anatomists would call it) sulcus. But this is not necessarily
of any great significance, because in tlie smaller Chiroptera and (Polyprotodont)
Marsupials there is no such sulcus, although the larger members of these Orders
{Fteropus, Cyiioityderis, Thyhiciuns, and practically all the Diprotodontia) present
a typical one.
The most peculiar fact, so far as the distribution of this furrow is concerned, is
its absence in most of tlie Ilodentia. In only one or two of the largest members of the
Orders, such as JlydrudiaTiis, is any such furrow developed ; and even in these few
cases it is small and insignificant. In the Mouotremata the calcarine sulcus cannot
be recognized ; moreover, it is highly probable that none of the mesial sulci in the
hemisphere of Echidna represents the calcarine sulcus of the Metatheria and Eutheria.
The most primitive form of the calcarine sulcus is seen to best advantage in such
brains as those of ThyJaciniis, Trichosurus, F/nisco/arcfus, JIacropns, Cynoiiycteris,
Fteropus, and most of the Edentata. Of these I select for representation tlie hemi-
sphere of Pteropus poUocepjlialus, not that it diff'ers in any respect from any of the other
forms enumerated, but because the presence of the calcarine sulcus in this genus has
* W. Kiikeuflial and Th. Zielcf)i, " Ueber das Centraliierveusvstem der C'o'taeeeu. iiebst Untersuchungen iibc-r
die yergleioheude Anatoruie der I'laceiitalicr,"' Denkscbrifteii der mediciii.-naturwisseusch. GesellscUaft zu Jeca,
1893, pp. 89, 117, inter alia.
t Trans. Linn. Sue. ser. 2, ZooL vii. 1S99. pp. 32^ ;JL!3.
SECOND SERIES.- ZOOLOGV, VOL. VIXI, 56
3S4
PROF. G. ELLIOT SMITH ON THE
been categorically denied hy Ziehen *. He states that ia the brain of Pleropiis niedius
(which is identical with that of P. poUocepludus ■\) none of the sulci which he labels
TT, o, and p in the Lemurs (corresponding respectively to the paracalcarine, calcarine,
and retrocalcarine of my account) are present; but he believes the sulcus a (the
intercalary of these notes) to be prolonged backward around the splenium. It is quite
unnecessary (because the discrepancy is so obvious) to point out hoAv little accord there
is between this view and that held by the same writer with regard to the Carnivora
and Cetacea.
The comparison of the section th'-ough the hemisphere of Pteropus (tig. 54) wath
that of Lemur and C'anis is quite sufficient to show the identity of the conditions in the
three forms.
F12-.546.
Fig. 54 a. — Pleropus 2'>oUocephahts. Mesial aspect of the right cerebral hemisphere. X .3.
Fig. 54 6. — Pferopiis j)ol{ocfj)hah(s. A horizontal section through the posterior part of the
hemisphere in the plane y (fig. 51 a).
In order that the relationship of the calcarine sulcus to the retrocalcarine and para-
calcarine sulci may be rightly appreciated and its systematic value may be correctly
appraised, it is necessary to consider certain other aspects of this problem.
Tlie exaggerated importance which Owen "^ had attached to the calcar avis or so-called
* -Irch. f. rsychiat. ISyO.
t ^ide Gustav Eetzius, " Zur Morphologie der Fascia Dentata," &e., Biolog. Unters. X. F. viii. 3, Taf. 14. fig. 3.
j Froc. Linn. Soc. TS5S ; also Annals & Mag. Xat. Hist., June 1S51, Eede I/ecture, and elsewhere.
MOEPHOLOGT OF THE BRAIX IX' THE :\IAM.MAI,IA. 385
" liippocanipus minor," as a supposed distinctive feaiure of the human brain, led to
a very heated controversy in ISGl, during- the course of which many anatomists
subjected the "occipital " region of the brain in the Primates to a careful examination.
One of the most vahiabie results of this unseemly strife avhs the introduction of the
useful term " calcarine suhms," which Huxley conferred upon tlu; external furrow,
the obverse of which forms the calear avis in the lateral ventricle. Another result
was the deuionstratiou by Iluxley, Flower, and others, that the calear and its sulcus are
not distinctive of the human braiu, but are commoQ to all the Apes or even Primates,
according- to Flower*.
The calear was first described early in the eighteenth century by Morand f, who
likened the little projection on the mesial wall of the posterior eoruu of the lateral
ventricle (in the human brain) to a cock's spur and ctiUed it ''ergot,'' the Latin
equivalent of which is '' calear acis." Towards the latter part of the same century
Vicq-d'Azyr J called it " //ippocamjxiis minor," because it appeared to be continuous with
the larg-er swelling known as hippocampus \^major'\.
According- to Broca, Cruveilhier § was probably the first writer to detinitely refer to
the constancy of the sulcus which produces the calear, or " aufractiiosite de la caritc
dUjilalc " as he called it.
In 185 i Gratiolet wroug-ly believed || that the sulcus in question was continued into
the hippocam2)al fissure, and included both under the latter name. This misconception
heljoed to estal)lish the misleading name " hippocampus minor " which Vicq-d'Azyr had
introduced. The merit belongs to Huxley of pointing out the fallacy of Avhich these
names were the expression, and, by reinstating the name calear avis and introducing the
new term " calcarine sulcus,'' he rid Descriptive Anatomy of a serious source of confusion.
I have introduced these historical notes here, because it has unfortunately become
necessary to emphasize once more, firstly, the contention of Huxley that the term
" hippocampus minor " should be for ever banished with the misconception which it
symbolizes; and that the term "calear avis," or simply "calear," be substituted; and,
secondly, that the true and exact significance of the term "sulcus calcarinus" may not
be forgotten, as it is by most modern writers oia the anatomy of the brain. As the name
" calcarine" is derived from "calear," it necessarily follows, as Huxley showed, that the
essential part of the sulcus and tliat strictly deserolag the title " calcarine" is thefun'ow
which produces the calear avis. But Huxley himself pointed oiit that the sulcus " extends
beyond the calear and the jwsterior cornu"^f. And he also did much more than this.
For, profiting by the earlier and not wlioUy successful attempt of Gratiolet, he for the
first time clearly distinguished among the complex arrangement of mesial sulci the
distinct hipiiocampal, calcarine, collateral, and parieto-occipital elements. Huxley
* '■ Oh the Posterior Lobes of the Cerebrum of the Qiiadrumanu," Phil. Traus. ISG2, pp. 185 et seq.
t Morand, ' Hi.stoire de TAcademie,' 1744.
t Vieq-d'Azyr, ' Traite d'Aiiatomic et de I'hysiologie," tome i. J 7S6.
§ Cruveilliier, ' .\natomie Descriptive,' l'"^'"" Edition, 1H.i6, torn. iv. p. <>()3.
|l Modern neurological literature eoiitains iiiuiuueralde repetitions of this error.
% "The Jjrain oF Atcltx,'' Troe. Zool. Hoc. !S!Jl, p. 2oo.
56*
386 PROF. G. ELLIOT SMITH ON THE
thus clearly anticiiiated most of the recent work in this region, by distinguishing the
parieto-occipital from the calcarine sulcus and hy recognizing that the latter extends
backward, far beyond the calcar.
Within recent years Cunningham * has called attention to a differentiation of the
calcarine sulcus into anterior and posterior elements, which are to be distinguished the
one from the other not only by a difference in the time and mode of development, but
also, in many cases, by their features in the adult human brain. The " anterior calcarine "
sulcus is the first to develop and becomes an extremely deep sulcus, which is wholly
responsible for the bulging of the calcar in the ventricle. In other words, it is the true
" calcarine sulcus."
The sulcus which Cunningham calls " posterior calcarine " develops later and quite
independently of the anterior sulcus ; it never becomes as deep as the former ; as a rule
it does not share in the formation of the calcar, and in many cases it is separated from
the anterior or calcarine sulcus by a submerged fold of cortex. It is, to use
Cunningham's own words, " a secondary sulcus in every sense of the term " f. It
is therefore of a very different nature to the true calcarine sulcus, and, as it is convenient
to have a distinctive name, I shall call it " retrocalcariue," because it is placed on the
caudal side of the calcar.
In his great monograph on the human brain, RetziusJ objects to Cunningham's
teaching and says that he " does not believe that such a fundamental difference between
the fore acd hinder parts of the fissure, as Cunningham sees, can be recognized. The
front part grows at least as often without a hinder addition and itself forms the hinder
part." The close of this quotation is interesting, as being practically a concession of the
writer's position and also by reason of its bearing upon the condition found in the Apes,
which Cunningham regards (erroneously, I believe) as differing essentially from that
found in the human brain. For a study of Comparative Anatomy makes it almndantly
clear that in the case of two sulci developing in the same line, it often happens that
the minor sulcus seems to be formed merely by a j^i'olongation of the major sulcus
rather than as an independent element. The posterior part was called " calcarine " only
because it was supposed to be part of the true calcarine sulcus ; but as this is not strictly
so, the distinct name " retrocalcariue " has been introduced. This has been done, not for
pedantic reasons, hut because a separate name becomes absolutely necessary in
Comparative Anatomy, \\here the luudamental distinction between the two elements
becomes more pronounced.
Describing the development of the sulci in a Sheep's brain, Krueg says that, in a foetus
10 cm. long, two fuirows have appeared : one on tlie lateral aspect is the faintly marked
representative of the fossa Sylvii of the hiunan foetus, and the other is a more definitely
* D. J. CiiuniugUam, " Contributions to tlie 8urf;ico Anatomy of the Cerebral Hemispheres," Mem. Eo\-. Irish
Academy, July 1SU2.
t Op. cit. p. 49.
- 'Das Meu.schcnhirn," Jena, 1896.
The passage here quoted is taken from an article by Cunningham, Jourual of Anatomy and Thysiology, vol. xxxi.
p. 595. See also Ketzius, ' Biologischen Uutersuchungen.'
I^rOEPHOLOGY OF THE BRAIX 1\ THE .MAMMALIA. 387
marked sulcus on tlie mesial surface, which he calls " splen/al/'s " *, a name which has
])een vevy generally adopted. Ue adds: — "According to Eckerf the fissura parieto-
occipitalis J develops in the human brain at ahout the same time as or soon after the
fissura Sylvii : I am very much disposed to consider it as the homologue of my fissura
splenialis " (p. oOD). In a later memoir he describes the splenial sulcus in TJnguiculate
Mammals. He explains the [)eculiarities of the fissura liippocampi and the fissura rhinalis
as lines of demarcation between cortical areas which are histologically diS'erent. Both of
these fissures develop very early, and after their appearance there is a long pause before
any other sulcus presents itself. The first one to do so in the Cat's hemisphere is the
sulcus splenialis (in a i'lctus of 10+ i"5 cnr. long). In such a foetus it appears " as a
flatteucd arc surrounding the posterior end of the corpus callosum and the upper end of
the hippocampal fissure, midway between these structures and the upper and hinder
margins of the hemisphere " §.
From a comparison of these interesting facts with those In-ouglit to light ])y
Cunningham and Eetzius, the conclusion is forced upon us that, if any sulcus on the
mesial surface of the human hemisphere is to be regarded as the homologue of the
'• splenial" sulcus of the Sheep and Cat, it must be, not the mor])hologically unstable and
unimportant parieto-occipital sulcus which Krueg suggests, but the true " calcarine
sulcus."
That such an interpretation was not entertained by Krueg is obvious from his
quotation of Meynert's happy guess j|, when he called "a short ascending branch of the
f. splenialis" in the Bear together with a short accessory furrow under it the "sulcus
occipitalis," and the f. splenialis itself the "sulcus calcarinus " ^. On the same page
Krueg quotes Benedikt as calling " the f. splenialis in the Bear the fissura occipitalis
inferior . . . and the mesial part of the f. medilateralis the fissura calcarina " (p. 6i2)**'.
If, liowever, we adopt Krueg's idea, we must, iu the light of our fuller knowledge
of the developmental history of the human brain, regard the splenial sulcus as partly
homologous to the calcarine.
In Broca's Collected Works (p. 321, fig. 20) a short oblique furrow is represented
in the hemisphere of a lloebuck, Avhich obviously corresponds to the sulcus Krueg
calls " postsplenialis." Broca calls this sulcus " the analogue of the calcarine
fissure," whereas he regards the splenial sulcus as part of the "■ limbic fissure."' He
* Julius Krueg, " I Cber die Furchung der Urosshirnriiide der riigulateii," Zeitsch. f. wissenscli. Zool. Bd. xxxi.
p. 308, November ISTs.
t A. Eckcr, " Zur I'lutwickl. der Fiirchen und AVindungen dei- Grosshiruhemisph. im Fotua des Mensehen,"
Arch. f. Anthrojiol. iii. Bd., ISO.j.
± Ecker is ^vroug in Ibis ; for it is the calcarine, and not the parieto-occii)ital, sulcus which develops first.
§ Julius Ivrueg, " Ueber die Furehung der (irosshiruriiide der zonoplaceiitiileu Saugetiere," Zeitschr. f. wissensch.
Zool. Bd. xxxiii., isso.
II Meyncrt, ' Archiv f. P33cliiatrie,' Bd. vii., liill. The context shows tlial .Meynert's suggested homologies
were little else tliau sheer guesswork.
•[ Krueg, op. tit. p. 041.
*■* Benedikt, ' .Vnatomisehe Studicn an VerbrechergeLinien," Wien, 1879,
388 PliOF. G. ELLIOT SMITH ON THE
adds in explanation that lie " calls it s'dlon calcarhi, although the ergot (calcar)
from which its name is derived exists only in the Pi'imates " (p. 322). It is clear from
this that Broca believes that the postsplenial sulcus represents that sulcus of the Primate
brain which produces the calcar, i. e. the true calcarine and not the retrocalcarine sulcus.
In the brains of most Carnivora and Ungulata. as well as in many other mammals, a
series of inconstant and exceedingly variable sulci make their appearance behind the
calcarine [i. e. the retrosplenial part of the splenial) sulcus. In the smaller members
this so-called *' postsplenial" sulcus tends to run parallel to the " splenial " sulcus. But
in the larger Carnivores, especially the Bears and Seals, and the larger Ungulates, such
as the Cow, Horse, and Camel, there are usually a series of postsplenial sulci ; and it
usually happens that the deepest of these pursues a course which is not parallel to the
calcarine (splenial), but inclines obliquely upward aiul backward at an angle with
the latter. It not unfrequeutly happens that this "retrocalcarine" sulcus, as we may
not inaptly term it, joins the calcarine. The tendency of the retrocalcarine element to
pursue a course which is not parallel to the calcarine becomes more pronounced in such
mammals (the Seals, for example) as possess a distinct caudal (or occipital) prolongation
of the hemisphere. It is, therefore, not surprising to find that in those mammals — viz.,
the Primates— in which such a caudal extension of the hemisphere is most pronounced
(so that the mechanical factors which tend to induce the cortex to become folded in the
longitudinal direction are most j)otent), the retrocalcarine sulcus always assumes
the longitudinal, and never the vertical, direction *. It is still further determined
in this course l:iy the increasing obliquity of the calcarine sulcus, which is brought about
partly by the bcickward extension of the hemisphere itself, and partly by the caudal
elongation of the corpus callosum. In fact, the direction of the calcarine and retro-
calcarine sulci is brought, as the result of the operation of these various factors, so
accurately into the same line that in most Primates the less important retrocalcarine
element seems to develop merely as a caudal prolongation of the more precocious
and stable calcarine sulcus.
The ])rain of the Camel affords a striking illustration of these facts. The mesial
surface of its cerebral hemisphere exhibits a triradiate group of deep sulci behind
the splenium of the corpus callosum (fig. 55). Superficially tiiese furrows apj)ear to be
jdined to one another; but if their lijjs be separated, it will be found in some cases that
the deep cdcarine sulcus is confluent with neither the intercalary sulcus, as in most
mammals, nor with the retrocalcarine sulcus, as in most Primates, but is separated from
l)oth by deeply submerged gyri, as is frequently the case in the human brain. It
is, moreover, noteworthy that the bulging (calcar) which is pushed into the definite
jjosterior cornu of the ventricle in this brain is produced wholly by the calcarine, and not
by the retrocalcarine nor intercalary sulci.
* The varijibility of this sulcus clearly shows that it possesses little stability, aud cannot be regarded as a
deliuitely fixed landmark like the true calcarine. It probably arises in order to relieve the teasiun of the
expanding " visual " cortex, and its form and position are determined by conditions which are chiefly, if not wholly,
mechanical, la these respects it presents a most marked contrast, to the calcarine sulcus.
MORPHOLOGY OF THE BRAIN IN THE MAMMALIA.
389
It is obvious, tlicrefore, that it is tlic rctrocalcariac aud not tlio true calcarine sulcus
whicli Broca has called " sillon calearin " in tlie brains of the Roebuck and Ass. The
src.
Cant eJ us </roiH(_(/{H-itts.
Mesial aspect of the right ctn'hrul hemisphere. Xat. size.
true calcarine sulcus in these mammals is represented in a part of tlie great furrow
^Ahich he calls " linii)ie."
If my view of this matter be correct, most recent English writers have departed much
further from the true interpretation than even Ivrueg aud Eroca.
In his well-know^n memoir " On the Convolutions of the Brain," Sir William Turner *
gave expression to very decided opinions concerning the calcarine sulcus and posterior
cornu. Writing about the l)rain of Hapnle Jacchns, he states : — "There is no splenial
tissvire, but ojiposite the splenium a distinct calcarine fissure " (p. 141). This amounts
to a categorical denial of the homology of the calcarine and splenial sulci.
On the same page he makes the I'urther statement (in this case referring to the In'ain
of Nycticehiis 'iStenops\) that " the calcarine fissure . . . proves the jiresence of both
posterior cornu and calcar avis " f . The data upon which this statement is based are
borrowed from Flower (Phil. Trans. 1802), Avho says, in the memoir quoted, that he found
it impossible to determine in this particular brain "whether or not the posterior cornu
exists." Nevertheless Turner goes on to say that " if the surface of the hemisphere
be examined with the view of determining the presence of an occipital lobe by the
evidence of a fissure, the calcarine fissure is that which is to be regarded as of primary
importance" (p. ll'l). This means presumably that there is no "occipital lobe" in any
mammals other than the Primates, seeing that the " calcarine fissure" is supjjosed to be
absent. The splenial sulcus, which occupies the position of the calcarine sulcus, i=i
regarded by Turner as merely a part of the calloso-margiual sulcus (p. 111).
* Joarn. Anat. and Phjs. vol. xxv. 1890.
t I have shown in these notes that there is no posterior cornu in the hrain of Nyi-iirehus.
390 PROF. G. ELLIOT SMITH OX THE
Alongside these remarkable statements we roiglit place Flower's comment concerning
"the absence of anything reseinl)ling the calcarine fissure [in the Cat's brain]"
(Trans. Zool. Soc. vol. v. p. 109). Flower's failure to recognize the identity of the
calcarine and splenial sulci is perhaps the more imintelligible, because he admits [mde
.supra) that a patent posterior cormi is not a condition absolutely necessary for the
existence of a calcar and a calcarine sulcus.
In his earlier work on the Seal's T)rMin *, Sir W. Turner suggests homology between
the collateral sulcus of the Primates and splenial sulcus of " Q,uadrupeds " ; and, if
I read aright the memoir quoted above (Journ. Anat. and Pliys. a'oI. xxv.), the
splenial sidcus may be represented in the Primates by both the calloso-marginal and
collateral sulci. In comparison with the calcarine, these two furrows (calloso-marginal
and collateral) are of very minor importance f.
Many other writers might be quoted to show the conflicting views whicb are put
forward to exj^lain the morphology of the splenial sulcus on the one band and the
calloso-marginal, calcarine, and collateral on the other. It will, however, suffice for our
purpose if we consider cbiefly the views of two modern writers, whose views are
diametrically opposed, the one to the other. Professor Ziehen finds in the Primate brain
some representative of almost CA^ery sulcus of the common mammalian brains ; whereas
Professor Cunningham adopts the extremist view expressed in his statement that,
" excejit in the case of certain of the main furrows {e. ff. Sylvian J, hippocampal), it is
very questionable indeed if there is any homological corres2)ondence between the sulci
of a primate brain and the sulci of a quadrupedal ])raiu." These remarks, written
several years ago, have been confirmed as lecently as this year, at the meeting of the
British Association.
Ziehen lias discussed the representation of the calcarine sulcus among the general body
of mammals in several of his memoirs; but we shall chiefly consider the remarks in his
memoir on the Prosimiau brain § as being most pertinent to this discussion. In his
description of the " calcarine group " of furrows in Nycficehus, he says that the retro-
calcarine sulcus (whicii he labels o) is undoubtedly the calcai'ine fissure; and the calcarine
* ' Challenger ' Reports.
t Coiicerniug the latter sulcus Alex. Hill gives utterance to a eharacteristie remark : — " There is no doubt in m)^
mind as to the horaolog)' of the ectorhinal [rhinal] fissure of the Dog with the collateral lissure in Man. ... To use
any other term than collateral fissure appears to me pedantic" ("The Hippocampus," Phil. Trans. 1893, p. 408).
As the rhinal fi.ssure is one of the iew furrows which can certainly be recognized by the histological features of its
lip.s, we can with certainty state, even if we are thereby stigmatized as pedantic by Dr. Hill, that the rhiual fissure
is not the collateral sulcu.s.
One is surprised to find so cautious a writer as I^rofessor D. J. Cunningliam apparently subscribing to the same
view ; for he makes the statement that the " incisvra temporalis [i. e., a remnant of the rhinal fissure] . . . may be
regarded as a forward prolongation of the collateral fissure'' ('Manual of Practical Anatomy,' vol. ii. 189G, p. 501).
'i'his error is probably to bo explained by the fact that the jiart of the rhinal fissure which is on the caudal side of
(he incisura temporalis is commonly regarded by Human Anatomists (who disregard the distinction between the
jjyriform lobe and the neopallium in the so-called uncinate gyrus) as part of the collateral sulcus.
J The case of the >Sylvian is rather unfortunate, as will be seen later, for the sulcus called '• Sylvian '' in the
■■ (Quadrupeds '" is ceiiaiiihj nut the homologuo of the Primate " Sylvian fissure."
§ Archiv f. Psychiat. lid. x.xviii. 181J0, p. OOiJ.
MOEPHOLOGY OF THE BRAIN IN THE MA.M^fALIA. 301
(his /)) is (lie continuation of tlicretrocalcaiine and not of the paracalcarine (his tt). Th(!
furrow TT merely cuts into the upper lip of p + o. In reference to this statement, he calls
attention to Bischotfs memoir *. In his discussion of the lioraoloo-ies of the calcarine
group of svilci, Ziehen compares the furrows of tlie Prosimian bi-ain with those of the
Carnivora, and especially Fhoca hispida. Like Broca, he regards the posterior branch of
the splenial sulcus ("' ramus horizontalis posterior ") as the calcarine sulcus, and the
" occipito-temporal part of the splenial " as the " stem of the f. parieto-occipitalis and
fissura calcarina " (p. 919). The confusion in these statements arises from the failure to
recognize that it is the so-called " stem of the paiieto-occipital and calcarine furrows "
which alone deserves the title " calcarine "f, whereas the furrow he calls " calcarine " is
really the retrocalcarine. But if the calcarine sulcus of the Prosimiae is represented by
the end piece of the splenial svilcus in the Carnivora, why does he consider the obvious
homologue of the latter in Fteropus to be the sulcus a (calloso-marginal or intercalary),
and not the sulcus p (calcarine) ?
A similar confusion of the true calcarine sulcus and the posterior calcarine has been
made by the German Anatomical Nomenclature Commission, calling the former
" f. occipito-calcarina" and the latter " f. calcarina," in spite of the fact that Huxley
introduced the name '•' calcarine " to distinguish that sulcus which produces the calcar,
i.e. the so-called " occipito-calcarine." Burt Wilder, who has disagreed with the
conclusions of the German Committee on so many other points, seems to embodv
the same confusing use of terms in his system of nomenclature.
Many other writers, such as Benedikt and Meynert, apply the term calcarine to that
portion of the sulcus which can only by courtesy, as it were, be so called, because it is
really a separate sulcus, the retrocalcarine, which has secondarily become confluent with
the true calcarine. If it were not for this confusion of terms, the conclusions at vvliich
these writers have arrived might have been received as an approximately accurate
expression of the homologies existing between the furroAvs of the Carnivora and the
Primates; ])ut this can hardly be said of them in their present confusing form i.
Pansch's criticism of the teaciiing of Meynert is utterly futile in the light of our
present knowledge. The climax of his argument — "Wo ist bei einen Raubthier
ein Calcar avis?" — becomes meaningless, if we compare tigs. 51 and 53 of this
work §.
The works of the English anatomists Turner and Cunningham are in direct conflict
with all the other writings quoted above. I have already referred to Turner's views.
* Siteb. d. bayer. Akad. d. Wisscnsch. 187U, p. 47S.
t /. f., if the slightest attention be paid to the detinition given by Husley when he coiued the word calcarine.
The calcarine sulcus is the furrow wliieh produces the caJcar.
X Theodor Meynert, " Die Windungeu der couvexeu Oberfluche des Vorder-Hirnes bei llenschen, Affen nnd
IJaubthiereu," Arch, f. Psychiatrie, 1877.
Maurice Benedikt, " Nouveile contribution i rAnatomie compart'e du Oerveau," Bull, dc la Soc. d'Anthrop. de
Paris, 4e serie, t. vii. March 1890, p. 22(;.
§ Adolf Pansch, '• Bemerkungen iiber die Faltuugen des &rosshirns imd ibrc Beschreibung," Arch. f. Psychial.
Bd. viii. 1878, p. 1'38.
SECOND SERIES. — ZOOLOGY, VOL. VIII. 57
392 PROF. G. ELLIOT SMITH OX THE
Cunningham has said that " we arc not in a position at present to offer an opinion . . .
upon the statement that the calcariae fissure is also developed in brains below the
Primates " *. Pive years later he stated that, " except in the case of certain of the main
furrows {c g. Sylvian, hippocampal), it is very questionable indeed if there is any
homological correspondence between the sulci of a primate brain and the sulci of a
quadrupedal brain." In spite of all the research in this field since these quoted passages
were written, Cunningham has recently reafhrmed his unaltered belief in his view
in these words : — "We now know that the primate cerebrum was not only distinguished
from that of all lower mammals by the possession of a distinct occipital lobe, but also by
having imprinted on its surface a convolutionary design which, in all but a few
fundamental details, was different from that of any other order of mammals " f .
What is this occipital lobe, w hich is so distinctive of the Primates ? Cunningham
himself does not tell us what he means by the expression. If it is merely the caudal
prolongation of the hemisphere above the cerebellum, then many large Carnivoi*es have
an equal or perhaps better right to be regarded as Primates than some of the Lemurs ;
and if it is the posterior cornu which is the diagnostic sign, then the Lemurs cannot be
regarded as Primates, whereas the Seals, the Camel, and some Cetacea are to be classed
along with the Apes. If, however, it is the calcarine sulcus which, to quote Turner,
" determines the presence of an occipital lobe," how can we draw a line of demarcation
in respect to this feature between the Primates and other mammals? So far as I am
aware, these arc the only possible criteria of an " occipital lobe," and none of them,
either separately or in conjunction, can be regarded as exclusively distinctive of
the Primates.
If we refuse to admit the homology of the cilcarine sulcus of the Primates and
the retrosplenial part of the splenial sulcus of other mammals, we have a far more
ditiicult problem to solve. For an explanation is then needed of the reasons for
regarding as different two sidci (in different groups of mammals) presenting identical
relations to the lateral ventricle and to the neighbouring brain-regions, a similarly
precocious appearance in ontogeny, and a constancy under varying conditions in mammals
which have widely diverged from the primitive stock. Why should we suppose that the
splenial sulcus, which is the most constant and most precocious neopallial furrow in the
mesial wall of the hemisphere in the Marsupialia, Cliiroptera, Edentata, Carnivoiu,
Lngulata, and Cetacea, fails to develop in the Primates, and that another and a different
furrow develops in the Primates in a position, which exactlij eot'respoiids to that occupied
by the splenial in other mammals and at the same epoch in development ? Such a
phenomenon is utterly inconceivable. The calairine and the retrosplenial part of the
splenial sulci are certainly identical.
But, it may be argued, the calcarine sulcus is an independent furrow, whereas the
" splenial " is prolong(>d into a supracallosal course — the intercalary sulcus. Put in the
Myrmecophagidai, Bradypodidai, and Manida) the calcarine is not joined to the inter-
* " Complete Fissures of the Human Cerebrum," Journal of Anatomj' and Physiology, vol. xxiv. p. 343.
t Presidential Address, Section H, Anthropology, British .issoeiation, September 1901.
MOKPIIOLOGT OF THE BRAIN IX THE MA.M.MALIA. 893
calary sulcus, aud in some of tlic smaller Marsupials, such as Trlchosurm, there is no
intercalary sulcus, the simple calcarine furrow alone being present.
It may further he ohjected that a true calcarine sulcus can exist only w lien there is a
posterior cornu of the lateral ventricle. If so, the Lemurs cannot have a calcarine
sulcus, whei'cas the Seals and tlie Camel possess such a furrow.
It is quite impossible to frame any definition of a calcarine sulcus or of an "occipital
lobe" which can he raised as a harrier between the Primates and the other mamnials,
such as Cunningham would have us erect. The calcarine sulcus and the calcar avis arc
not the exclusive property of the Primates, but are the common heritage of all tlie
Metatheria and Eutheria.
In most mammals this sulcus becomes joined to the intercalary, a furrow of little
morphological importance; V)ut in the Primates the backward prolongation of the corpus
callosum so alters the direction of the calcarine that it forms an acute angle with the line
of the intercalary, so that for purely mechanical reasons the two furrows fail to unite,
and the calcarine becomes confluent with the retrocalcarine, which is another furrow of
quite secondary importance.
It is not without consifleiable significance that these phenomena occur in an equally
pronounced form in the Lemurs as in the Apes, in sjjite of the fact that the mechanical
conditions favouring such a .sundering of the calcarine-intercalary junction are at least
equally marked in many Carnivoi-es, in which the separation does not occur. It Avould,
however, be unwise to attach too great an importance to this iact, because we
liud the calcarine sulcus separated from the intercalary in the Anteaters, Sloths, and
Pangolins.
The fact that no trace of the parieto-occipital sidcus is found in the Hapalida; and
several of the smaller Cebidte seemed to suggest the possibility of that furrow being a
purely Simian feature, which had become evolved in the Apes only. I was therefore
inclined at first to follow Flower's teaching, and regard the paracalcarine sulcus of the
Lemurs as one of the limbs of the bifid extremity of the retrocalcarine sulcus of the Apes.
But the fact that it branches off from the point of union of the calcarine and retrocalcarine
sulci shows that it occupies a position analogous to that of the lower extremity of the
parieto-occipital sulcus in the higher A2:)es.
But are we justified in calling this furrow parieto-occipital ? Most wi'iters do so
without the .slightest hesitation.
In order to settle this point I have examined the brain in every genus of Monkeys,
and have found that in the higher Apes, where the true parieto-occipital sulcus of
Human Anatomy can alone be said to exist, this furrow is com[)o.sed of two and
frequently more elements. In the majority of cases, however, there are two sulci,
dorsal aud ventral, which overlap to a considerable extent, the intervening area of cortex
usually becoming submerged so as to obscure the dual natiu'c of the resultant furrow.
Tlic ventral furrow I have called "paracalcarine," Ijucause it pursues a course alongside
the calcarine sulcus in its most typical form, e. y. in Slmia. The dorsal sulcus is
obviously due to the bias a\ hich is given to the developing cortex near the dorsal edge by
the intraparietal sulcus. This furrow becomes so sharply bent that it forms an acute
394 PROF. G. ELLIOT SMITH OX THE
angle, Avhioh iwints toward the mesial surface, and so affects the neighbouring area that
a vertical sulcus usually develops in response to this mechanical bias.
The paracalcarine sulcus in the Lemurs obviously represents the ventral element ouly,
and not the whole of the parieto-occipital.
I have introduced the term " intercalary "' to distinguish a furrow which has hitherto
received no exclusive title. In most mammals it is joined to the calcarine, and the
comjilex is called "splenial": in the Primates it is separated from the calcarine and
joined to the genual sulcus to form the calioso-marginal sulcus. In many Carnivores ir,
is also joined to the crucial sulcus. It is therefore intercalated between the three
furrows — calcarine, genual, and crucial — to any or all of which it may be joined. lu
spite of its constancy in the Mammalia, the intercalary furrow is morphologically
unstable and readily becomes broken up into several fragments in the larger Ungulatcis
and Carnivores. A careful examination of the behaviour of this furrow in the whole
Mammalia leads me to regard the paracalcarine as being equivalent to the separated
posterior fragment of the intercalary sulcus, or as a new compensatory-calcarine furrow
developed only in the Primates. In the case of sulci which possess so little individuality
and morphological value as the intercalary and the parieto-occipital, it is of little use
attempting to establish any exact correspondence, because this can hardly be said to
exist even in two hemispheres of the same brain. All that can be said with certainty is
that the tension of the growing cortex, which v/ould be relieved in the Carnivora by the
conOuence of the intercalary and calcarine sulci, is instrumental in forming the para-
calcarine sulcus in the Lemurs. This does not, however, imjily the identity of the
two furrows. The instability of the paracalcarine element is shown in the Cebidai :
in CalUthrix it is either absent or very diminutive, in Chrysothrix it forms the typically
vertical ventral element of the parieto-occipital sulcus, and in Hf^yctipilhecus it is
horizontal. Of course the latter cannot be described as homologous with the typical
furroAV of ChrysothrLv, but merely as the expression of the same causal factors. There
can therefore be no strict homology between such an unstable furrow and any sulcus,
and especially such a variable element as the intercalary, in otlier mammals.
When we consider how tickle this sulcus is, the fact of its extreme constancy of form
and relations in all the Lemurs, not excepting even the aberrant Tarsius and Chlvomys,
becomes a valuable testimony to the closeness of the bond of affinity whicli unites all the
Prosimite. In fact, a study of the mesial aspect of the hemisphere affords convinciug
proof of this undoubted kinship.
In many Lemurs of various genera an oblique sulcus {b) appears on the ventral side
of th(! retrocalcarine sulcvis. A comparison of this with a series of Primate brains shows
it to be the posterior part of that composite and morphologically unimportant furrow
which is called " collateral " in the brain of Man and the Apes.
:\FORPlIOLOGr OF THE J5RAIX IX THE [MAMMALIA. :V.)o
TuK Morphology of the Sulci ox tiik Lateral and Dorsal
ASPECTS of the CeREBRAL HEiriSPIIEKK.
Iiinninerablo attempts liave been made, with results as vari(!d as they are numerous,
to solve the problem of determining to how great an extent, if at all, it is possible to
institute exact comparisons between the sulci of two brains.
If the two hemispheres of any highly convohitcd brain I)e cr)mpai-ed, a certain number
of furrows Avill be found in eacli, wliieh cannot l)e exactly liomolos-ized witii any sulcus
in the other hemisphere. And not unnaturally the number of such "incomparable"
elements increases when we compare the hemispheres of different individuals, species,
genera, families, and orders. The crucial question is thus reduced to the enquiry as to
the possibility of recognizing any sulci as the common property of several Oi'ders.
The most divergent and mutually contradictory views are held at the present time on
this subject. Thus there is the negative teaching, which is perhaps most forcibly and
dogmatically expressed in a posthumous memoir by A. J. Parker in these words: —
" Tiie mistake is ol'ten made .... of attempting to compare their [the Mammalia in
general] convolutional characters with those oi' the Primates, including Man .... We
cannot expect, nor do we find, any exact honiological relations between the convolutions
of this phyllum [sic] and those of other mammalian phylla [sic] " *. The same view is
expressed in a less jn'onounced form by D. J. Cunningham.
The view of Gegenbaiir may be summed up in his own words by the statement " dass
eine Homologie nur in sehr engen (Jreuzen besteht und bei sehr vieleu gar nicht durch-
fiihrbar ist " (Vergl. Anat. der Wirbelthiere, i. Band, 1898, p. 769).
Although in many of his earlier memoirs Turner had instituted comparisons between
the sulci in different mammalian Orders, yet in his kitest contribution he comes to the
conclusion that in each Order of Mammals a special pattern of sulci is evolved peculiar
to itself.
Most other writers who have in any way dealt with this subject have come to the
conclusion that it is possible to homologize the sulci of Primates to a greater or less
extent with those of other mammalian Orders, luit there is an infinite variety ol'
siiggestions as to tlie exact manner in which this is to be done.
Before avc consider a few of the suggested interpretations, it will conduce to clearness
if I first explain, by a reference to the Cat's brain, the nomsnclature employed in
describing the sulci in Orders other than the Primates.
The furrow commonly called " Sylvian fissure " is an offshoot of the rhinal fissure :
as the usual designation is undoubtedly erroneous, I shall call it the pseudosylvian sulcus
(fig. 56 i). The sulcus which is best known by the name "presylvian" {Owen), I have
termed "orbital" for reasons which will be apparent later. The furrow which is
commonly called " posterior suprasylvian " has been designated by its original name
" postsylvian " {Owen), not only because the latter is more appropriate, but chiefly that
* " llorpholog)' of the Cerebral Convolutions with Special Kei'erence to the Order of Primates," Journal ot the
Acad, of Xatural Science, Pliiladelphia, 2nd series, vol. x. 18!Mj, p. 27().
396
PEOF. G. ELLIOT SMITH OX THE
the name "suprasjlvian " may be reserved for the other sulcus, of such vastly greater
importance, to which it often happens to be linked in the Carnivora.
The other names are sufficiently explained by the accompanying diagrams (figs. 56 a
and h). One of the earliest attempts to compare this pattern of sulci— which we may
Fig. 56a
scr.
1 I
so.^dsca.
sps.
sep.
pi". ."6 a. — Fcli>: domestka. • Dorsal aspect uf bi'n,iii. Xat. size.
Pin-. 5(; /,. — FlVis ilomesiica. Lateral aspect of the left cerebral hemisphere. Nat. size.
regard as the common Mammalian plan — Avith the sulci of Man and the Apes was tliat of
Pansch *. He regarded the iipper of the three curved sulci on the lateral aspect of the
Doo-'s brain (/. e., presumably the conjoint lateral and coronal sulci) as the boundary
of the postcentral convolution {i. e., presumably the intraparietal sulcus of Turner,
the "interparietal" sulcus of most Continental writers). The middle and low^er curved
sulci (i. e., the suprasylvian, postsylviin, and ecto.sylvian) are not represented, according
to this writer, in the brain of the Primates.
In Owen's ' Anatomy of Vertebrates,' 1868, the orbital (presylvian) sulcus is regarded
as part of the Sylvian fissure.
Hitzig, as the result of a physiological investigation into the distribution of the motor
areas in the Dog and Ape, came to the conclusion that the central sulcus of the
latter is represented in the former by the ansate and the anterior part of the suprasylviau
sulci f.
In another memoir by Pansch the orbital sulcus of the Carnivora is regarded as the
representative of the precentral sulcus of Apes : the suprasylviau [iu his foi-mer paper it
was the corono-lateral] corresponds to the intraparietal, and the coronal sulcus to the
central (Rolando's) J.
Meynert, who committed the extraordinary error of thinking the crucial sulcus [ivhieh
* " Ueber d. t)-pische Aiiordung d. Purchen u. 'Wiiiduiigcn nuf deu Grosshirnhemisphiiren der Menscben u. der
Affen," Arch. f. Anthropologic, Bd. iii. p. 227 (1SG8).
t ' Untersuohungeii iiber das Gehirn.' Leipzig, 1S74.
X " Ueber gleichwerthige Regioiiou am Grosshirn der Cariiivoren uud der Primatcu," Ceiitralb. f. d. med.
Wissenscli. no. 31, p. 041 (1S75).
MORPHOLOGY OF THE BRAIX IN THE :VIAMMALIA. 397
he called central, l. c. Puolaiido's] absent in the Bear, consiilers that the central sulcus of
Man is represented in the Carnivora by the coronal sulcus *.
Bnjca regarded the orliital (presylvian) sulcus of Carnivora as the representative of
the human sulcus of Rolando f.
In 1878 Pacsch once more returned to the discussion of this problem, whicli he had
lieen considering for more than ten years. After criticising the views of other writers,
and more especially Meynert's, he maintains J the same views concerning the homologies
of the coronal and suprasylvian sulci which he advanced three years before {op. cit. supra).
On physiological grounds Munlc regards the crucial sulcus of Carnivores as the
liomologue of the central sulcus of Primates, and cites the recent work of Kiikenthal
ixnd Ziehen and of Eberstaller in support of this view. This enumeration of the different
views which have l)een enunciated by different writers might l)e considerably extended if
it would serve any useful purpose §.
It will be noticed that all of these writers (as well as tliose who arrive at negative
results) iittempt to institute direct comjiarisons between highly specialized repi'esentatives
of tlie Carnivora like the Dog and Cat and the higher Apes. No writer has taken the
obvious precaution of testing the accuracy of his suggestions by an appeal to the simplest
and most generalized types in the two Orders : nor, again, is it altogether clear why tlie
vast majority of writers invariably seek for homologues for the sulci of the Primates in
only one Order of Mammals (in most cases the Carnivora), unless it be because the
non-Primate type of sulci is seen in its most sj)ecialized form in these latter.
Every writer (including even those who, like Parker and Cunningham, deny the
homologies of the sulci in the Primates and other Orders) commits the fundamental
error of regarding the so-called Sylvian fissure of the Ciirnivora and other Mammals as
the representative of the Sylvian lissure of the Primates. But there cannot be the
slightest doubt that the two furrows are not homologous.
The only means of arriving at any reliable conclusions as to the possibility of homolo-
gizing the sulci of the Lemurs with other mammals is to critically study the behaviour of
these furrows in the Mammalia as a whole. And as this is the only means of truly
appreciating the relationship of the Prosimian brain to that of other mamooals, I j^ropose
to bricHy summarize here the data relating to the cerebral sulci in other Orders wJiich
have been stated more fully elsewhere ||.
Eirst of all, the behaviour of those sulci which I have represented in the Cat's brain
may be studied in the Carnivora. In the Primates, the two most stable furrows of the
neopallium, and those to make their appearance earliest in development, are the calcarine
* " Die Winduiigen der convexcn Oberfliicho des Vorder-Hirns bei Menschen, Aften uud Itaubthiereu,"' Arch. f.
Psychiatrie, Bd. vii. p. 250 (1877).
t ■' Anatomic coraparoe des circonvolution.s Co rub rales," Kev. d'Anthropologie, 1878.
X " Bemerkungen iiber die Faltuugeu des Grosshirns uud ihrc Beschreibung," Arch. f. Psychiatrie, Bd. viii.
1878.
§ Brief reviews of the Literature will be found iu memoirs by J. X. Langley ("The Structure of llie Dog's
Brain,'' Journal of Physiology, vol. iv. pp. 268-270) ; Jules Soury (" Systcmo Nerveux Central," tome ii.,
especially pp. 910 & !Jlo) ; and Flatau and Jacobsohu (•• Vergl. Anatomic," op. cit.)
II Catalogue of the lloyal College of Surgejus, Second Edition, vol. ii. 19U2.
398 TEOF. G. ELLIOT S^riTII ON THE
sulcus on tlie mesial surface (with wliicli at present we are not concerned) and tlie
Sylvian fissure on the outer surface. It might reasonahly be assumed therefore that,
if the sulcus usually called " Sylvian fissure" in the Carnivora were correctly so-called, it
would be the most constant and most precocious furrow^ on tlie outer aspect of the
neopallium. But this is not so. For if the developmental history of the neopallial sulci
be studied in any Carnivore *, it will be found that several sulci, such as the suprasylvian
and corona], make their appearance long before the pseudosylvian sulcus. Then, again,
if the brain be studied in the whole assemblage of Carnivores, it will be found that the
suprasylvian, coronal, and lateral sulci exhibit far greater stability and maintain their
characteristic features with far greater constancy than does the pseudosylvian sulcus.
And in the more generalized and primitive Viverridai the latter becomes very imperfect
and inconstant, and even at times disappears entirely. Occasionally the same
phenomenon occurs in some of the other Carnivores: I have seen the pseudosylvian
sulcus of the Cat reduced to very diminutive proportions, and the ectosylvian sulci
deepened to compensate. It is clear, therefore, that the so-called "Sylvian fissure" of
the Carnivora is not the morphologically stable and precocious furrow which we should
expect if it were the true representative of the similarly-named feature of the brain in
the Primates. Moreover, the nature of this false Sylvian furrow varies considerably
within the limits of the Carnivora. In the iEluroidea, as for example in the Cat's brain,
the lips of the pseudosylvian sulcus are formed by the first arcuate gyrus of Leuret, i. e.
the strij) of neopallium Avhicli is bounded peripherally by the ectosylvian sulci. In the
Arctoidea (as was long ago demonstrated by Sir William Turner, and recently confirmed
by Holl and the writer [Catalogue of Royal College of Surgeons t]), the w'hole of the first
arcuate gyrus is buried in the false Sylvian furrow, so that the lips of the latter are
formed not by the first (as in the Jilluroidea) but by the second arcuate gyrus of Leuret,
-/. e. by the area bounded peripherally by the suprasylvian and postsylvian furrows.
And in the Pinnipedia (as I have recently shown in the above-quoted Catalogue, p. 2S6)
the whole of the anterior limb of the second arcuate gyrus tends to become buried in the
great cleft-like so-called " Sylvian fissure," so that in many Seals the anterior lip of the
furrow is formed by the third arcuate gyrus of Leuret. And if a series of Carnivores be
examined, all intermediate stages will be found between these extreme types.
Moreover, it has been show'n by Holl %, who has so admirably demonstrated the real
nature of the so-called "Sylvian fissure" in Carnivores and Ungulates, that the
claustrum also extends beyond the area depressed in the pseudosylvian sulcus, even in
the Arctoidea ; so that, if anyone is inclined to attach importance to this structure as an
indication of the extent of the insula (or area submerged in the Sylvian fissure), it is
ol)vious that the pseudosylvian sulcus of Carnivores is in no sense homologous with the
true Sylvian fissure of the Primates.
By far the most precocious and most stable sulcus on the lateral aspect of the
neopallium in the Carnivora is the suprasylvian (/. e., the furrow which most writers
* I have examined embryonic brains of the Cat, Dog, and Bear, and have confirmed the teaching of Krueg
(Zeitsch. f. wissenseh. ZooL Bd. xxxiii.) iu this matter.
t VoL ii. (2nd ed. VM'2) p. 277, fig. 154, % Arch. f. Anat. u. Bhys., Auat. Abth. 1899 and 1900.
JIORPirOLOfiY OF THE RRAIX IN THE MAMMALIA. 399
call "suprasylvia anterior"). Its caudal extremity may or may not be joined to
the relatively unimportant postsylvicxu sulcus (/. c, the "suprasylvia posterior" auct.) ;
the latter is not only much more variable, hut develops much later than tlie true
suprasylvian sulcus. In most Carnivores tlie two sulci are united to form a regular a.rc
around the pseudosylvian sulcus. But it often happens in almost every genus, and
constantly in Uerpestes and its allies, as well as in many Seals, that the sliprasylvian is
)iot joined to the postsylvian sulcus. The morphological value and importance of tin;
two furrows is so vastly dilTereut, that 1 have thought it advisable to wliolly discard the
common and misleading title " suprasylvia posterior," and return to tlic original nam.;
" postsylvia " (Owen), which is both simpler and more accurate, as the further discussion
Avill show.
The ectosylvian sulci are features of little morphological importance, and are to b^
regarded as furrows compensatory to the pseudosylvian sulcus. In many gener.i botli
ectosylvian sulci are absent : in flie Ilya^nidfc the anterior ectosylvian sulcus is absent;
and in the Bears the two sulci are sul)merged. Even when present the sulci are
exceedingly variable.
Next in importance to the suprasylvian sulcus come tlie orbital (presylvian), coronal,
and lateral, and, after a wide interval, the crucial, postlateral, arcuate, and prorean sulci
in that order.
The presylvian (or, as I prefer to call it, "oi-bital ") sulcus makes its appearance at a
very early period, and is almost as constant as tlie suprasylvian. It usually becomes
considerably prolonged so as to join the rliiual fissure below and so as to extend near
to the dorso-mesial edge of the hemisphere above. But both these phenomena must
be regarded as secondary modifications, Avliich take ]Aiuie under the influence of purelv
mechanical factors, and are therefore devoid of great morphological imnortance. The
true fundamental orbital element is a small furrow jjlaced in the neopallium alono-sid**
the anterior end of the rhinal fissure : as such it is perhaps the most constant and one of
the most stable of all the neopallial sulci in the Mammalia. The coronal and lateral sulci
are second in importance only to the suprasylvian and orbital. The former is perliaps
veiy slightly the more precocious, but occasionally this sulcus is absent when the lateral
sulcus is present, as sometimes happens in Uerpeslcs. [This is also especially the ctise in
Rodents and in many Marsupials. On the other hand, the coronal sulcus is often
present when the lateral sulcus is absent in some Ungulates, and esjjeeially in FrocavuiA
In the Carnivora as a whole these two sulci are very constant, and in most cases they
are united to form one furrow.
The direction of the coronal sulcus is subject to considerable variation in the different
families. In tlie larger members, and especially the Ursida^, it becomes almost transverse ;
whereas in the simple, generalized Viverridse its direction is almost sagittal.
A postlateral sidcus is often united to the caudal end of the lateral, but it is exceedingly
variable and may be entirely absent. In its simplest form it often consists of a small
transverse furrow behind (and not joined to) the lateral sulcus.
The crucial sulcus, which is often regarded as a distinctively Carnivore feature, is not
constant. Its size is exceedingly variable. So large doi's it become in the Bear thai
SECOND SERIES. — ZOOLOGY, VOL. Vlil. 68
-100 PEOF. G. ELLIOT SMITH ON THE
Meynert refused to re:?arcl it as the "crucial" sulcus, and called it "centi-al." In the
Viverridae it is often absent, and its simplest form is exhibited in some of the members
of this family as a small depression on the dorsal surface of the hemisphere, a short
distance in front of the outwardly-bent anterior end of the lateral sivlcus.
The ansate sulcus is a branch of the lateral, which develops behind the crucial sulcus.
It is not constant, and when present is subject to great variation.
Much less constant than any of these furrows is the sulcus wliicli Krueg calls
"diagonalis." It is often present in the Felida- as a separate element: more often its
identity is lost by merging into tlie anterior ectosylviau sulcus or the suprasylvian.
Tf we represent graphically the inferences draAvn from the foregoing discussion as to the
features of the primitive Carnivore, a scheme such as the accomiianying diagram (fig. 57)
would be the result. This is practically the brain of Vicerra.
Fig. 57.
scr. ^f'- si.
SCO.
ss.
Jr. ."ps.
A selicmc representing the sulci in a liypotheticnl generalized Carnivore.
That this is really the primitive Carnivore type will be shown by an examination of
the brain-casts of the Eocene Carnivora.
Thus in a natural cast of the brain of Sfenoplesiclis Co//l/ari in the British Museum
(M. 1723) there is a conjoint corono-lateral sulcus (which is quite sagittal in direction)
and a well-defined suprasylvian sulcus, as in Viceira. There is r;o sign of crucial,
])seudosyIvian, or postsylvian sulci.
Fig. 58.
Steiinplesicl.is Cai/lii.ri. Dorsal aspect of a natural cast of the cranial cavity. Xat. size.
In the excellent natural cast of the cranial cavity of CijHoliijcBnodoii Cayluxi (British
Museum, M. 4199)— another Carnivore, which, like aSV^»o/j/(^'s/c//.s\ comes from the Upper
Eocene formations of France — w^e find similar lateral and suprasylvian sulci, and two verv
shallow furrows representing the pseudosylvian sulci [a similar phenomenon is often
MORPHOLOGY OF THE BKAIX IX THE :^IA^r.MALlA.
401
found in tlic Viverrida']. There are neither postsylviaii nor postUiteral sulci ; but
the suprasylvian and lateral sulci extend almost to the posterior extremity of the
hemisphere *.
Fig. 59a.
Fig. 59 6.
sL
sps.
Figs. 50 n & h. — CiiiwJiiiirnoiloii Cai/hi.ri. (it) dorsal .■iml (/>] lalorul aspects of a natural cast o
jf part of
tlic cranial cavil v. Nat. sii:e.
If we turn to the examination of tlie lateral surface of the hemisphere in tlie Ungulata
(leaving out of account certain sulci, such as the intercalary, which often leaves the
mesial and extends on to the dorsal surface f ) we shall find in Procai-ia, which is perhaps
the most generalized member of the Order, an arrangement of sulci whicli is obviouslv
identical with that of the hypothetical primitive Carnivore.
The suprasylvian and coronal sulci closely resemble those of Viverra, and especiallv
the Eocene Carnivores, both in position and extent. The lateral sulcus is usually less
well-developed, and is always separated from the coronal sulcus ; but there cannot be
the slightest doubt as to its identity with the similarly-named sulcus in the Carnivora.
In some cases the lateral sulcus is absent.
The i)se\ulosylvian sulcus is almost always absent, but in one brain (of eight examined)
I found a furrow resembling tliat sometimes seen in J'ieeri'u. Tliat this sulcus was no
mere vascular furrow was shown by the presence of a vertical postsylvian sulcus J in that
brain and not in others. Another instance of a similar phenomenon is shown in Turners
figure of the brain of ILjrax \_Procaijia'] (Journ. Anat. and Physiology, vol. xxv. 1900).
A small typical orbital sulcus is almost invariably present, although it has been
overlooked by both Krueg and Turner. There is no diagonal sulcus.
If the developmental history of the cerebral hemispheres be studied in a series
of Ungulates §, it will be found that the earliest sulci to make their appearance on the
outer surface of the hemisphere are those which from their relations and behavioiu" are
* There is not much resemblance between this cast and that tigured b)- Filhol. There is no doubt, however that.
even if this is not what it is represented to be, it is certainly an Eocene Carnivore.
t As a similar phenomenon is presented in the brains of certain Carnivores, such as lldictis, and, less decidedly
in Mcles, there can he no doubt as to the homologies.
+ This showed that there was a furrow of sufficient depth to lend a bias to the neighbouring cortex.
§ Such investigations have been made by Krueg (Zeitsch. f. wissensch. Zoologie, 13d. ixxi.), and more recently by
Paul Martin (" Zur Entwickelung der Gehirnfurchen bei Katze und Rind," Arch, fiir wissensch. u. prakt. Thierheilk.,
]Jd. xxi. Heft i., 189.5), and I have studied this problem in embryos of Ovis. Bos, Capra, Sus, and C'amelus.
Bradley has published a few data concerning the Horse's brain in the Journ. Anat. and Phys. vol. xxxiii. pp. 215
& 5«7.
58*
402 PEOF. G. ELLIOT SMITH ON THE
oln-iously the suprasylvian, orbital, and coronal sulci. In their subsequent developaaent
all of these sulci differ very cor.sidcrably from their homologaes in the Carnivora, so that it
is imjiortant to bear in mind that these sulci are regarded as homologous not only
because they are the most constant and similarly-placed farrows, but because their mode
of deA'elopment demonstrates their identity in the two Orders.
Holl has recently demonstrated that the sulcus commonly called " Sylvian fissure " in
tlie Uucrulata is not homolos'ous with anv one of the various tvpes of pseudosvlvian sulcus
Found in the Carnivora. The " Carnivore jiseudosylvian " sulcus is sometimes found in
the Ungulata, e. g. in the genus Bos, but it is not that Avhich is commonly called
"Sylvian fissure." The latter (which we may call the "Ungulate pseudosylvian "
sulcus) is formed by the meeting of two sulci, \Ahicli are analogous to the ectosylviaa
sulci of the Carnivora *.
After the sulci mentioned above, the most constant furrows are the lateral and diagonal
sulci. The former sulcus is not linked to the coronal sulcus as so often happens in the
Carnivora ; but it is sometimes joined to the suprasylvian sulcus. The diagonal sulcus
is much more constant than it is iu the Carnivora, and it is almost always placed just in
front of the anterior end of the suprasylvian sulcus.
It is doubtful whether true homologues of the postsylvian and postlateral sulci are
ever found in the Ungulata.
If we review the whole Ungulate Order so as to determine the constant sulci,
it will be found that those furrows only will be picked out Avliich, as the evidence of
embryology shows, represent those also found in the hypothetical primitive Carnivore.
If we next examine the Edentata the same fundamental plan of sulci is again found.
Thus in the genus Bradnpus we find pseudosylvian, suprasylvian, lateral, coronal,
orbital, and diagonal sulci arranged in a manner so like that seen in Fch's, that
one cannot question the homologies implied in the nomenclature just used. The
developmental notes publislied by Pouchet show that one is thoroughly justified in
maintaining this aIow. In the nearly allied, but larger, genus CIiolwpus the
pseudosylvian sulcus disappears ; it thus adds further testimony to the slight importance
of this sulcus.
In the Anteater {Tamandua) there is always a definite orbital sulcus and a conjoint
corono-lateral, but it is a peculiar fact that in some cases the suprasylvian sulcus may be
absent, although in others it is well-developed. There is no pseudosylvian nor diagonal
sulcus.
One of the most interesting and instructive brains in the whole mammalian series is
that oi IIijrmecoplKKjd Jalxda ; and all the more so because it is subject to peculiar
variations.
The normal or common type of brain exhibits a conjoint corono-lateral, a postlateral,
a postsylvian, an orbital, and a jirorean sulcus, as in the Caruivora, and in addition a
])seudosylviau sulcus of a peculiar nature. Moreover there are sometimes shallow
*" Holl, " UebtT die IujcI dcs UDgulatcngcliirnfb,"" AvlIi. f. AiKit. u. Thys. 1000. Compare also Catalogue of the
lioyal College of Surgeons, L'uJ Lditiou, vol. ii. figs. It' J {\>. \i\j'J) uud 107 (p. '•ii'd).
MORPHOLOGY OV THE BHAIX L\ THE MAMMALIA. 403
depressions representing diagonal*, posterior ectosylvian, and crucial sulci of ilie
Carnivore-type.
The feature of most interest, however, is the ])S('U(losylviau sulcus. Tliis usually
consists of a shallow depression above the bend of the riiiual iissure: there is a well-
defined posterior lip, but no anterior lip. Eroni the apex of the depression a deep
and well-defined sulcus begins and arclics backward close to the upper end of the post-
sylvian sulcus.
The reader will observe that so far no mention has been made of the suprasylviaii
sulcus, the most stable sulcus on the outer surface of the hemisphere in the Carnivora
and Ungulata. It might, perha])s, be supposed that, as this sulcus is sometimes absent
in Tanumdua, it might also be absent, as a rule, in Jlynnecophucja. JUit it is hardly
likely that a cerebral hemisphere which is provided with lateral, coronal, orl)ital, post-
lateral, postsylvian, pseudosylvian, proreau, and even diagonal, posterior ectosylvian and
crucial sulci, should lack just that particular sulcus which is the most constant and stable
sulcus in the Mammalia. Such a phenomenon is so highly improbable as to be almost
inconceivable. There can be no doubt whatever that the suprasylvian sulcus is present
as the deeply-incised, dorsal, arcuate appendage of the pseudosylvian sulcus.
Eortunately there is an anomalous brain in the Collection of the Royal College of
Surgeons [D. 281, Catalogue, p. 22o, fig. 1)5] which proves that this is so. On the left
liemisphere of this brain the condition described above is found ; whereas on the right
hemisphere the suprasylvian sulcus f is separated from the pseudosylvian sulcus and
joined to the jiostsylvian, as in most Carnivores.
In Orijcteropm the suprasylvian sulcus is only poorly developed, whereas the coronal,
lateral, orbital, and ])rorean sulci are exceedingly well-developed. There is no trace of a
pseudosylvian sulcus.
In Manls there are typical suprasylvian, orbital, corono-lateral, and pseudosylvian
sulci, as in Brudii])ux.
In the Armadillos the orbital sulcus is always, and the suprasylvian sometimes present.
In many of the Rodents there is an extraordinary j)aucity or even absence of sulci.
The lateral is perhaps the most constant sulcus, and the orbital and coronal sulci are
found in many Rodents. In Hi/drocha'nis aiul Lagodumus there is a su2)rasylviau
sulcus ; and in the former numerous other furrows of doubtful homology. The most
interesting brain among the Rodeutia is that of Dollcholls. In addition to the typical
orbital and sagittally-directed corono-lateral sulcus there is a suj)rasylvian sulcus like
that of LagostomH.s, l)ut it is joined to a pseudosylvian sulcus to form a conjoint Sylvian
furrow, such as is usually found in the Edentate M'jrmecophaga. These forms are of
particular interest in comparison with the condition found in the Lemurs, as will be
apparent later ;|:.
* The diagoual sulcus, both of the liradypodidoc and tlii' MynneropLagidii!, is always dircctod upward and forward
as in the Ungulata and Primates, and is not subject to the irregularity and changes iu direction which wc find in tha
Carvivora.
■^ .\ comparison of the two liemispheres shows that the supra.sylvian sulcus is certainly present in both.
J Sea Catalogue of the Royal College of Surgeons, 2nd Edition, vol. ii. p. 205, fig. 75.
4-04 PliOF. G. ELLIOT SMITH OX THE
111 the Cliiroptera the suprasylvian and lateral sulci are sometimes present in. the
genera Fteropits and Cijnonycteris, and resemble the corresponding furrows in the
Edentate Tamandua.
In the true Insectivora there is usually an orhUal sulcus only. This is particularly
well-developed in Gynimira. The little brain of the aberrant Galeopithecus is peculiarly-
rich in sulci. It possesses a very dee]} and unusually extensive suprasylvian sulcus of
Tinusual form, and in addition well-defined orbital and pseudosylvian sulci.
The smaller Polyprotodont Marsupials resemble the Insectivora in possessing no well-
defined sulcus except the orbital on the outer surface of the cerebral hemisphere. In
Tliylacinus and most of the Diprotodontia there are also suprasylvian, pseudosylvian, and
prorean sulci ; and in Phascolomys and many of the MacropoJidse there is a typical
lateral sulcus *.
If all of these scattered data be collated it will be possible to determine those features
which are the common property of all the Metatheria and of those Eutheria which do
not belong to the Order Primates. If we attempt to form a hypothetical type of cerebral
hemisphere embodying all these features, it must be macrosmatic and must present a
well-defined I'hinal fissure. It must have a deep and well-defined suprasylvian sulcus, a
small orbital sulcus which neither joins the riiinul fissure nor is prolonged far in the
mesial projection: coronal and lateral sulci which are nearer the sagittal than the
transverse direction and which are quite separate. In addition there mat be pseudo-
sylvian, postsylvian, postlateral, oblique, ansate, and crucial sulci, for none of these are
the exclusive property of one Order, but all, as the brief review given above shows, tend
to appear in all mammals. Moreover, there is to be noted a tendency in several Orders
(in Myrmecophaga among the Edentates, in Dolichotis among the Hodents, and in the
Seals among Carnivora) for the area which separates the overlapping parts of the supra-
sylvian and pseudosylvian sulci to become submerged or so reduced that the two furrows
form one sulcal complex (fig. 00).
If, after this review of the conditions which prevail among the Mammalia in general,
the plan formed by the sulci in the Lemurs be examined, it will be apparent that the
•cirrangement in the latter is unquestionably a slightly modified form of the hypothetical
common mammalian type. This exact reproduction of just those sulci which are most
stable in other mammals, and the A'ariability shown by just those furrow^s which are
less stable in other mammals, can be no mere fortuitous phenomena, but positive
^^vidence of morphological identity.
There can be no doubt that the Sylvian fissure of the Lemurs is formed by the peculiar
union of the suprasylvian sulcus with the less stable pseudosylvian of other mammals.
.1 ust as the suprasylvian element in this complex is the only sulcus of the lateral
surface which is ab.^olutely constant in all Lemurs, so it is the most stable sulcus in the
Mammalia generally. That it is really the suprasylvian sulcus which is found in the
Lemurs is abundantly shown by the peculiar condition found in C/uromys.
In order of constancy the orbital, lateral, and coronal sulci come next, just as also
* These homologies in the Marsupialia arc suggested by a comparison of the mode of development of ihe furrows
ill Ibc brain of J/wrojuti- and that exhiMtcd in other mammals. e-*peeially certain Ungulala.
MOT^PllOLOGY or Till' BRA IX IX THE MA^nfAlJA. 405
happens in the Mammalia as a \vh()l('. Next in order follows tlie post sylvian snlcns,
again as in other mammals. And iinally, Ili(> most variable I'ro^iniian tx'atnres — the
postlateral, diagonal, and cnicial lurrows— ar(> exactly tliose wliiuh coine next ia the
order of frequency imiong mammals in general.
The homologies thus suggested are so revolutionary and so opposed to all the current
teaching, that it is necessary to submit each sulcus seriatim to the closest scrutiny in the
light of all the comparative evidence available, either in the facts of direct observation
or in the writings of other anatomists. At the same time, it will be cjiivenient to extend
the range of comparative studies and Include the Apes and Man.
The Morphology of the Sylvian Fissure.
The Sylvian fissure is found in its complete and typical form only in tlie human brain.
The p)erij)heral lips of three sulci become opercular and appi-oach until they meet : the
peculiar pattern thus formed upon the surface by the meeting of these labia is commonly
called the Sylvian fissure.
The region which becomes overlapped by the oporcula is called the insula E/Cilii and
is bounded by three sulci — the inferior, superior, and anterior limiting sulci. If the
human brain be compared with a sufficiently complete series of brains of Primates, it
will become so obvious as to amount to an absolute demonstration that tlie inferior
(morjjhologically posterior) limiting sulcus represents the pseudosylvian, the superior
limiting sulcus the suprasylvian, and the anterior limiting sulcus the diagonal sulcus,
respectively, of the Lemurs and other mammals.
It has been clearly shown by Marchaud * and Cunningham f that the anterior limiting
sulcus of the island of Reil is the liomologue of the fronto-orbital sulcus of the
^Vnthropoid Apes. If anyone is disposed to dispute this contention and follow the lead
of Herve, Eberstaller, Waldeyer, and Giacomini (among others), a glance at the series of
brains of Anthropoid Apes, and especially those of the Gorilla, in the Museum of the
Royal College of Surgeons will convince him of liis error J. The anterior limiting sulcus
of the insula is cei'tainly the homologue of the fronto-orbital sulcus of the Apes. The
latter is best developed in the genera Aiithropopithemts, Simia, and Ili/lobates, but is
constantly found in a less well-developed condition in Scmuopilhecus, Papio, and
frequently in other genera of Old-World Apes and sometimes also in the larger members
of the family Cebidtc.
In the Hapalid:e and the smaller Cebidoe, and sometimes also in tlie smaller Cerco-
pithecidae, this sulcus is absent. It is therefore interesting to note its presence in many
individuals of the genus Lemur as well as in some of the Indrisintc. Undue importance
nmst not, however, be attached to tiiis fact, because the fronto-orbital sulcus of the
Lemurs and Ajies is almost certainly the homologue of the diagonal sulcus of the
Carnivora, Ungulata, and Edentata.
* "Die Morphologic des Stirnlappens uiid der Insel dor Aiitliropomorplien,"' Jena, 1^:);3, pp. 91 et sc'i (Arbeiteii
aus d. patholog. Inst, zu Marburg, Bd. ii.).
t " The Insular District in the Cerebrum of the Anthropoid Ape.s," Journ. of Anat, and Phys. vol. xxxi. pp. 1 cl sc'i.
t See especial!}- specimen D. G58, Coll. Surg. Catalogue, p. -io'J.
.JOG PKOF. G. ELLIOT SMITH OX THE
The only -n-riter t\ ho has attempted to seek for the common mammalian homologue of
this fronto-orbital sulcus of the Lemurs is Ziehen *. He came to the conclusion that it
represented the presylvian (orbital) sulcus of other mammals ; hut the fallacy of his
argument is quite patent and capable of easy refutation. Ziehen re2:ards the sulcus
which I have called " orbital " in the Lemurs as the representative of the " intraorbital "
sulcus of the Carnivora, and the diagonal (fronto-orbital) sulcus as the presylvian (which
I call orbital). Thus he attempts to homologize a sulcus which is one of the most stable
features of the Prosimian brain with an unimportant furrow like the intraorbital which
is found in only a few of the larger Carnivora and is morphologically of no importance ;
and, on the other hand, he suggests the identity of the exceedingly stable presylvian
(orbital) sulcus of the Carnivora with the fronto-orbital sulcus which is found only in the
o-enera Lemur, Propilheciis, and Indris of all the Prosimia?, and not constantly even in
these. He attempts to justify this by the statement that the presylvian sulcus may bo
absent in a Carnivore, and quotes the example of Eerpcstcs. I have, however, examined
ten cerebral hemispheres of Herpestes, and have never found the orbital (presylvian)
sulcus absent, though it is always very small f. But even if it were always absent not
only in one but in several genera of Carnivora, this would not justify the homology of
one of the most constant sulci of tiiis Order with perhaps the most unstable sulcus in
Lemurs.
In the literature of cerebral morphology the orl)ital (presylvian) sulcus of the
Carnivora has been represented as the homologue of many sulci liy different writers.
According to Owen, Meynert, and many others (even in Edinger's recent text-book,
1899), it represents part of the Sylvian fissure; according to Pansch it is the precentral
sulcus ; and according to Broca and Eberstaller (and also Soury in his great work ' Le
Systeme Nerveux Central,' 1899) it is the homologue of the central (PiOlando's) sulcus.
The evidence of Comparative Anatomy (and by this I do not mean the futile attempt
to compare the brain of one specialized Carnivore like the Cat with that of the highest
Primate) shows decisively that the presylvian sulcus of the Mammalia cannot possibly be
x-epresented in the Primates by any of these four sulci — Sylvian (Owen), precentral
(Pansch), central (Broca), or fronto-orbital (Ziehen). The orbital (presylvian) sulcus is
perhaps the most constant sulcus (in the neopallium) in the Mammalia. It is found in
the brain of almost every Carnivore f, Ungulate, Cetacean, Edentate (excepting Chlami/-
dophoi'us), ami in all Piodents, Insectivores, and Marsupials which have any cerebral sulci.
It is therefore highly improbable that such a stable sulcus should either be entirely absent
or be represented by an inconstant element in the Primates. Moreover, the brain of the
J'rimates possesses a sulcus in the same position in the hemisphere as the presylvian
occupies in other mammals ; and this sulcus is the most constant furrow in front of the
Sylvian fissure. This can be no other than the orbital sulcus (the " triradiate " sulcus of
'J'urner ; the " presylvian " sulcus of Owen). The correctness of this identity of presylvian
and orbital sulci becomes more conclusively demonstrated as the range of comj)arison
increases. If we compare, for example, the pi-esylvian sidcus of the Rodent JDoUcholis,
* Aroh. f. Psych. M. xxviii.
t It is actuallj- absent in the Ilctpcbtiue Crossai-c/ivf,
MORPHOLOGY OF THE BKAIX IX THE .M.\M>rALIA. 4.07
oi- of the Cai-nivoi-e TlceiTa ov the Ung-iilatc Procaria, with tin; orbital sulcus of such h
Prosimian bmiu as that of (iaUKjo or ^\q\\ Leinnr, cue cannot deny that the two sulci are
morphologically identical. If we examine a larg'c series of iunuan brains, it will be found
that the stable portion of the orbital sulcus consists of a deep oblique sulcus, the posterior
end of which approaches close to and often joins the outer limb of bifurcation of tlu^
olfactory sulcus, just as the presylvian sulcus of the Carnivore's joins the rliinal fissure ;
but the simjile linear form of the orbital sulcus of Man is usually disguised by a scries
of inconstant branches, so that it seems to lose the simplicity of form it possesses in most
Apes. If this fundamental part of the human orbital sulcus be compared with the pre-
sylvian sulcus of the larger members of the other Mammalirm Orders, such, for example,
as the Camel and the Seals, a demonstration of the identity of these two furrows will b(;
afforded which is quite as striking as the similarity of the presylvian sulcus of DolicholtH
to the orbital of Galago. Such instaucas might be multiplied without limit.
So that, if we take into consideration all the facts which a study of the brain in all
mammals yields, there is a mass of evidence a)uounting to absolute demonstration that
the sulcus variously called pi'esylvian, supraorbital, orbital, and triradiate in different
mammals is fundamentally one and the same furrow.
Between the orbital sulcus and the lower end of the Sylvian fissure we occasionally
find a furrow in the Lemurs and the New- World Ajies, and generally in the 01d-"\Vorld
i\j)PS : this is the frouto-orbital sulcus. AVith regard to this homology in the Lemurs,
I quite airree w ith Zi( hen. r>ut, on the other hand, the evidence which I have been al)le
to collect points conclusively to the truth of the suggestion of Eberstaller and Cunningham,
that the fronto-orhital sulcus of the Apes [and Lemurs] is the anterior limiting sulcus of
the insula of the human brain.
In most Ungulates, in many Carnivores, and in the Bradypodida: and .\fiin)tecupluMiii,
a small oblique sulcus makes its appearance in the position exactly corresponding to that
occupied by the fronto-orhital sulcus in the Primates : it is called the diagonal sulcus
(Krueg), and is placed between the orbital and the suprasylvian su1<m. It is highly
improbable that a small oblique sulcus such as the diagonal (especially in the Ungulates
and in Bradi/pus) should develop in the analogous position in three different Orders so
as to exactly reproduce the features of the fronto-orhital sulcus of the Lemurs and Apes
without being homologous. The probability that " diagonal," " fronto-orhital," and
" anterior limiting" are merely different names for the corresponding furrows in different
mammals is immeasurably greater than the reverse.
The evidence upon which Professor D. J. Cunningham chiefly relied to clinch the proof
of the identity of the fronto-orbital and anterior limiting sulci, was the distriljution of
the claustrum. This consists of the deeper part of the cortex which composes the island
of Beil. In the human brain the clausti-um becomes separated from the rest of the
cortex by a definite medulla ly layer in a region coextensive with the insula, /. e. so far
forward as the anterior limiting sxilcus. In the Chimpanzee [and the same holds also for
the Orang and Gorilla] the claustrum extends as far forward as the fronto-orbital sulcus.
In the Cercopithecidie and Cebid^c it is unfortunately the fact that we cannot vise such
evidence with the same degree of definiteness, because the anterior extremity of the
SECOND SERIES. — ZOOLOGY, VOL. Till. ^'J
408
PliOF. G. ELLIOT SMITH OX THE
claustrum becomes so diffuse that its extent caiiuot be accurately mapped out. But its
distributiou shows clearly that, tlie anterior part of the iusula is freely exposed on the
surface of the brain, as it is in the Anthropoid Apes. In the genus Lemur the claustrum
is an exceedingly plump and well-defined strueture. It extends back-ward as far as
the posterior [inferior] limiting- sulcus (which is the pseudosylvian sulcus), and extends
forward slightly beyond th^ fronto-orbital (diagonal) sulcus (fig. 00(0, ;it the same time
becomius: somewhat diffuse.
Fig. 60a
Fig. (!(). — A scliciuc representing the sulci common to most !Miimm;ili:i.
Fig. Wa. — Lemur fuhnts. Horizontal section through the bniin. x 1'.
This may perhaps seem to throw doubt upon the homology of this sulcus in the
Lemurs and the Anthropoid Apes. It is impossible to appreciate the true value of such
evidence until we know more than wc do at present of the significance of the claustrum *.
* This subject lias been studied by T. E. Clark (" Comparative Anatomy of the Insula," .Touru. Comp. Neurology,
vol. vi. June L'SiJO, pp. 5'.J-1((],;3.5 figures), and more recently by M. HoU (" Ueber die Insel des Carnivorengchirns,"'
Arch. f. Anat. n. Physiol. [.Vnat. .\bth.] '.'> u. 4, pp. 217 ct seq.. lS!j9; '• Ueber die Insel des Uugulatengehirns,'"
■ihidoii, ') u. 0, pp. l'!t.j i1 srq., KM II I J.
MOKPHOLOGY OF THE BEAIX IX THE :\[A:\r:\r.\LTA. 409
Among Clark's conclusions it is slated tliat " the ckuslrum may iiiclud'' parts of areas
other than the insula " (p. 91), /. e. it may exleiid beyond tlie situations of the limit int;-
sulci.
The most revolutionary and novel sugg'ostion in the foregoing discussion is the
homology of the suprasylvian sulcus of mammals in general partly with the superior
limiting sulcus and partly -with the ujjper (or posterior) part of the Sylvian li>sure of tho
Lemurs and Apes.
Such an interpi elation of the morphology of the cerebral sulci has never been
suggested hitherto; and yet I belies-e that, if the identity of these two sulci be (h-nied,
it is impossible to homologizc the majority of the sulci in the brain of the i'rimates witli
those of other mammals. For if the furrow which is deepest, most stable, and most
precocious in the Carnivora, Ungulata, and Edentata is not represented in the Primates,
and, moreover, l)y that i)articular sulcus which is also most constant and develops
earliest, then it is utterly futile to seek for the homologues of the other uu)re varial)l(!
sulci.
The only -writer (Ziehen) who has seriously attempted to homologizc the Prosimiau
sulci Avith those of other mammals [or, more strictly, with those of the Carnivora], found
representatives in the Primate brain for the lateral, coronal, an.sate, crucial, presylvitm,
ectolateral, medilateral, intraorbital, and postsylvian [his " fissura suprasylvia posterior'"]
of the Caruivora, and yet imagined that the most stable and deepest sulcus cf the;
Carnivora — the suprasylvian sulcus * — was missing when so many unimportant sulci are
reproduced. Such a suggestion is utterly inconceivable ; and, even if we knew nothing
of the instructive phenomena exhibited in Chimmijs,'n\ Myrmecophaga, and in Doltchoti.s,
the conclusion must be forced upon us that either the suprasylvian sulcus is merged in.
the Sylvian complex, or it is impossible to institute exact comparisons between the sulci
of the Primates and any other mammals.
It is quite unnecessary to point out the fallacy of the old teaching of Pansch and
Meynert, that the suprasylvian sulcus represents the intraparietal sulcus of the
I'rimates.
Several writers have ai)preciated the oljstruction in the way of a saiisfactory interpre-
tation of this region ; but I believe no one has hitherto suggested the solution which all
impartial observers must admit to be true.
Thirty years ago Gervais saw the diihculty, ^ihen he wrote :—" Lemurs never have
more than two convolntidns around the Sylvian fissure, whilst even the smallest
Carnivores have three " f ; but did not explain which of the Carnivore furrows disguised
its identity by merging in another sulcus.
Quite recently Holl recognized that the " Sylvian lissure " of Lemnr is composel of
two separate sulci ; but he suggested the ectosylvian, and not the suprasylvian, as the
dorsal element %.
* Ziehen (Arch. f. Psych. Bd. xxviii.) says :—'• Dor vonkre Eogcnsabschnht dcr F. suprasylvia [/. ^. the true
sulcus supr^svlvius] fehlt dem Halhaffrngi-hiru " (p. 021).
t Journ. a; Zoologie, tome i. p. 27. t Arch. f. Anat. u. Phys., A„at. Ahth. U.OO.
59*
410 PROF. G. ELLIOT .s^riTll OX THE
In the intevpretatioii of the morphology of sulci, one is not disposed to place any gi-eat
reliance upon physiological evidence ; i'or, even supposing that the functions of the
various regions were knowii, it does not necessarily follow that homologous areas are
homodynamic in ditferent Orders. Nevertheless it is of interest to note how fully
the results of physiological research support the view that the u})per part of the Sylvian
fissvu-e represents the suprasylvian sulcus.
Thus Terrier records * that stimulation of the dorsal lip of the suprasylvian sulcus in
the Cat causes retraction of the angle of the mouth and movement of the eyes to the
opposite side, Avhereas pricking or sudden retraction of the opposite ear is the result
of applying the electrodes to the lower lip of the suprasylvian sulcus. Stimulation of tlu'
corresponding tips of the upper part of the Sijti-ian fssiire in a moiikeii (Macacus)
produced ilie same results as that elicited by exoitatian of the lips of the suprasijloia ii
sulcus in the Cat (tig. 335). The results obtained by Ferrier in the case of Lepus (p. 73 i)
are in full accord.
In Edinger's summary of the results of physiological experiment on the cerebral cortex
he represents tv.'O diagrams-!, horrowed from Gustav Mann, showing the lateral aspect of
the brains of the Cat and Rabbit respectively : in these the suprasylvian sulcus is repre-
sented seimrating two areas labelled respectively "Oculomotor" [the dorsal] and " Auris "
[the ventral]. Edinger inserts a diagram of the human brain alongside these, and labels
the dorsal lip of the posterior end of the Sylvian lissure " Ocnlomot." and the ventral lip
" Auditus.''
Now, as it would be absurd to imagine that the results adopted by Eerrier, Schafer,
Mann, and Edinger Avere in any Avay biassed by such a (to tliem) rank heresy as the
belief in the identity of the Sylvian and supi'asylviau sulci, the evidence which they
unconsciously produce is of value as an indication that one who entertains such a belief
cannot he accused of flagrantly disregarding physiological data. Moreover, this wholly
unexpected support from ihe i^hysiologists is the last link in a long cliain of irrefutable
evidence (only a feAV links of which have been incorporated in these notcsj that the upper
[caudal] part of tlie Sylvian lissure of the Primates and the dorsal limiting sulcus of Reil
(the " opercular " sulcus of Marchand) represent the supi-asylvian sulcus of quadrupeds.
The stable suprasylvian sulcus, by its " union " with that variable suprarhinal kink
which I have called " pseudosylvian sulcus," gives a fixity to the latter Avhich it did not
possess l)efore ; and the result is the Sylvian fissure, or, to be strictly accurate, part of the
postericn- limb of the tissura Sylvii.
In the ijeneral review of the condition of the sulci in the Mammalia, it Avas noted that
in the Eodent Dolichotis and in the Edentate 3Ii/rmecophaga a peculiar union of the
suprasylvian and pseudosylvian sulci occurs. In the light of the foregoing discussion,
thc^'.e facts may be simply expressed by the statement that these two mammals have a
Prosiniian Sylvian tissiu'c.
If, alter cai-efully studying the arrangement of the sulci in the ^Eluroid and Cynoid
* E. .\. Scliiifpr, ' Text-ISook of Physiology," voL ii. liiUd, fig. :3;!3, p. 7;i4, fig. S.'So, p. 735.
-j- •• TUr Anatomy of the Central Xervous Sy^tcm," Aun'iican traiisUitioii, l^US.1, p. l?2.j.
MORPHOLOGY OF THE BRAI\ JX THE :\rAMMALTA. Ill
Carnivores, the student next examines the brain in tlie Arctoidoa, he will find that the
Avhole of the first arcuate gyrus of Leuret has heconic submerg-ed in a great cleft in
the situation of the pseudosylvian sulcus, and that there is a pronounced tendency for the
anterior limb of the second arcuate gyrus to become submergtul. This tendency, which
is seen in most Bears, becomes more marked in Me/c\s; and in most of the Seals the
Avhole of the anterior limb of the second arcuate gyrus and the gi-(;ater poi-t of tin;
suprasylvian sulcus become swept into the pseudosylvian clcff. There is therefore in
the Arctoid Carnivores a tendency to the production of a coiulition which is analogous
to, witliout however being altogether identical with, the state of affairs which has
already become evolved in the Primates. It is of interest to note tliat when the supra-
sylvian sulcus is swept into the pseudosylvian sulcus in Dtarla, (hlob(r»iis [Trichechus],
and Fhoca, it becomes separated from the postsylvian sulcus (to which it is united in
most Carnivores). "When the suprasylvio-pseudosylvian comjjlex is formed in M'/nw-
cophafia, the postsylvian sulcus becomes disconnected ; but in the anomalous case {ckle
supra) in which the suprasylvian sulcus is not joined to the pseudosylvian, the former
sulcus was joined to the postsylvian.
These peculiar phenomena are of great interest in the interpretation of the Prosimian
brain. It has frequently been noted in this memoir that in the Prosimiie, and especially
in the genus Lcmni; the mode of ending of the upper extremities of the suprasylviau
and pseudosylvian sulci is exceedingly variable, and indubitably points to a connection
between these two furrows, which has only recently been broken. In the peculiax-lv
interesting brain of Ch/roiin/.s, the suprasylvian and the variable pseudosylvian sulci are
widely separated, as in most quadrupeds; and, like the parallel case alforded by the
aberrant brain of Jff/nuecojj/nu/a, the suprasylvian sulcus tiien becomes joined (in most
cases) to the postsylvian sulcus.
The extreme variability of the postsylvian sulcus in €'///' roii/i/s is chiefly the result of
the peculiar inconstancy of the pseudosylvian sulcus ; for it will be obvious, from an
examination of the brains of Procacia and tlie Ungulata, that the characters of the
postsylvian (which is a morphologically unstable sulcus) are determined chielly by the
nature of the pseudosylvian sulcus. The irregularity of the jjostsylvian sulcus mav
also be partly due to the fact that {Jhiroinys has obviously been subject to marked
retrogressive changes.
In most Lemurs the postsylvian sulcus is a simple linear furrow, which obviousiy-
represents the parallel or superior temporal sulcus of the Apes and Man.
Although the pseudosylvian sulcus has a morphological importance and stabilitv
which are immeasurably less than that of the suprasylvian sulcus, it is obvious that it
is the former which is the chief determinant of the peculiar " Sylvian formation." That
this is so is shown by a study of the Carnivora, in many of which (as, for instance, in
many Canidpe, the larger Pelidte, and Hysenidae) a condition so spuriously like that of
the true Sylvian fissure is produced without any participation by the suprasylvian sulcus,
that the mistake of regarding the Carnivore 2)seudosylvian sulcus and tiie Primate
Sylvian fissure as homologues is not unnatural. The suprasylvian sulcus, however; adds
stabilitv to a region which in other mammals is notoriouslv variable and iickie. Tlie
\
\
\
1,12 PEOF. G. ELLIOT SMITH OX THE
upi^er [or caudal] part of the " Sylvian fissure " is purely suprasylviau, but the lower
[or anterior] part of the sulcus is formed by the meeting of the dorsal lip of the supra-
sylviau sulcus with the ventral lip of the pseudosylvian sulcus.
In nrany Prosimian brains, especially of the genera Lemur, Fropithecus, and Iiiclris
(l)ut also, to a less marked degree, in Galngo, Perodicticus, and Nycticehm), the lower
end of tlie suprasylviau sulcus may be seen emerging from the " Sylvian fissure " some
distance above the rhinal fissure. Several writers have noticed this. Ziehen, for instance,
Ciills it the " Vorderster Abschnitt der Fissura circularis externa (s. Reilii) " (Arch. f.
Psych. Ed. xxviii. p. 929).
These relations of the pseudosylvian and suprasylviau sulci are identical with those
foTuid in the smaller Cebidse and CercopithecidiB ; but in these Monkeys the lower end
of the suprasylviau sulcus is not exposed. It begins to emerge in the Cynocepludi and
Se^nnopUheci, and in ILjlohatcs and the Antliropoid Apes it becomes prolonged forward
(as the " opercular sulcus " of Marchand) so as to almost meet the fronto-orbital
[diagonal] sulcus. Even in the Simiidje the dorsal lip of this forward extension of the
suprasylvian sulcus and the anterior lip of the diagonal sulcus tend to become opercular,
and extend downward and backward until in m(«t hiunan brains they meet the
opercular caudal lip of the pseudosylvian sulcvis, and thus completely cover the anterior
insvilar area, wliich is always exposed in non-hiunan brains. In Man alone is tlie
Sylvian fissure complete. In the Apes the "stem" of tlie fissure is sometimes formed,
but the so-called Sylvian fissure of Monkeys really represents only a jmrt of the posterior
limb of the human Sylvian fissure. The fully ibrmed anterior limbs are never found,
except in the brain of Man.
The extent of submergctl area varies considerably in the different Prosimian genera.
In Lemur there is a vei-y considerable area, chiefly overlapped by the opercular lip of
the pseudosylvian sulcus: in Tar^vH*-, Ji/t•;"0(?t'Z//^s■, and Ayf^/Zcei^s there is practically no
submerged area, so that if the term " insida " be used as synonymous with " overlapped
neopallium," the opinion of Flower and Ziehen that there is an insula in Lemur but not
in Isiicticclus is justified.
Put it has Ijcen demonstrated that in the Apes a considerable part of the insula is not
submerged and extends forward as far as the frocto-orbital sidcus. If my identification
of the latter sulcus in the Lemurs is correct (and in this matter I am supported by the
opinion of Ziehen and Flatau and Jacobsohn), the insula must be regarded as being
exposed in a very considerable part of its extent. We have no certain criterion to inform
us which neopallial areas should be regarded as " insula." la the human brain the
insida is that area of neopallium (1) which is suliinerged in the Sylvian fissure, (2j is
circumscribed (almost completely) by limiting sulci, (o) is applied to the lateral aspect
of the nucleus lenticularis, and (4) is split to form a claustrum. If we adopt any of
these criteria in the Lemiu's, the region so mapped out will fidfil )ione of the other
three conditions.
The next group of sulci to be considei'ed includes the lateral, coronal, and crucial.
The sulcus ^^hich I have called "latcr;d" in the Lemurs is regarded by Ziehen as
the lepreseutative of the " Bogenfurche " of the Dng's brain, which is composed of the
/■
/
/
/'
/
/
/
MORPHOLOGY OF THE BHA]\ IX Till- .MAMMALIA. H :l
lateral, ansate, and coronal sulci*; the sulcus wliidi I distinqaiislied as "coronal" he
regards as the prorean ; and lie agrees with me in the rccogaitiou of liu^ sulcus/ as part
of the central sulcus. The latter sidcus is considered by Flatau and Jacobsohuf to
represent the superior precentral sulcus, because it occupies a positiou iu relitiou to the
intraparietal sulcus analogous to that whii-h is presented iu the ])raiii of Apes.
I shall discuss first of all the representatiou of these sulci in (he general body of
mammals, and then follow the evolution of (he corresjTOnding furrows in the Apes.
If we follow the example of Ziehen, aud limit our mammalian horizon to such
specialized Carnivores as Caii'is and Fhoca, neglecting all the generalized Carnivores and
the Ungulates, Edentates, and llodenls, we might perhaps arrive at the same conclusion
and regard the sulcus I have called " lateral" as the homologue of the conjoint corono-
lateral furrow of many Carnivores. If, however, w^e study the developmental tendencies
not only in one or two specialized forms, but in the whole Mammalia (in such a manuer
as I have roughly sketched out already), the whole evidence will point to the conclusion
that the sulci which I have called "lateral" and "coronal" respectively are in reality
the homologues of the sulci of the same names in rirerm, l^rocaria, and all mamniais.
In other words, the coronal sulcus, inst(-ul of being represented in the furrow which I
have called "lateral," as Ziehen would have us believe, really exists in the Lamurs as
the separate sulcus which I have called " coronal." The latter sulcus (Ziehen's r) is
regarded by the same writer as tlic houiologue of the prorean.
The prorean sulcus has a peculiar distril)ution in the mimnnlian series. It is seen in
both the Polyprotodont(r/////«e///».<.) and Diprotodout Marsupials (Macropodidae, P/i«s(?o-
lomys, and extinct Thylacoleo), in 3[iii-niecoph(i<i(i and Orncfcropiis alone of the Edentata,
and reaches the height of its development iu the Carnivora, and especially iu some of the
Arctoidea, such as Gulo. If we seek for this sulcus iu the more "■eueralized Carnivora
and especially tlie Viverrida:-, no such separate element as the prorean sulcus will be
found. In the Viverrida^ the coronal sulcus is prolonged forward so as to take the
place, functionally, if not morjihologically, of the prorean sulcus.
A similar phenomenon occurs in nrost Ungulates, ami in the generalized Hyrax
(Procaria) the coronal sulcus extends far forward on to the apex of the hemisjjhere.
Much light is thrown upon this perplexing problem by a comparison of the arrangemi^nt
of these sidci in certain llodents aud Edentates with that found in other mammals.
In the Carnivora the coronal is sejjai'ated from the ^^rorean sulcus by a mesial extension
of the orbital (presylvianj sidcus. This ])rolongation is certaivdy a purely mechanical
phenomenon; i.e., the tension of the growing cortex becomes relieved by the extension
of a pre-existing furrow, which in this case is the orbital. In Ori/cteroptis, and some-
times in 3Iyrmecophngfi, the mesial prolongation of the orbital is united not only with
the prorean but also with the anterior end of the coronal sulcus. In the l)rain of
Dolichotis (Rodentia) the orbital sulcus is not prolonged beyond its morphological limits
(if such a phrase is permissible), but the corono-proreau complex retains a form which
is curiously reminiscent of the united coronary and prorean sulci in Oi-yctei'opiis. This
* Arcli. f. I'sycliiat. Bd. xxviii. p. iil'l. t llaiiJbucli, p. ITS.
414 TROF. G. ELLIOT SMITH OX THE
uould seem to indicate that tlie "coronal" sulcus of the Viverridie, of the Ungulata,
and of the Lemurs oucfht properly to be regarded as prorean-coronal. For the constant
development of the prorean sulcus in those hrains (of such widely separated mammals
as the Marsupials, Edentates, and Carnivores) in whicli its union with the coronal sulcus
is prevented by a " mechanical " extension of the orl)ital seems to indicate that the
prorean siilcus possesses some definite individuality and mori)holog-ical stability.
In the brain of Chiroinys there is a peculiar modification of this region (which shows
unmistakable signs of retrogressive change). In some cases there is a definite typical
orl)ital sulcus and, in a more dorsal position, another sulcus which is obviously the
anterior part of a coronal (or prorean-coronal) sulcus which has become broken into two
fragments, like that of Isycticehus. In other cases the two sidci (/. e. the orbital and
the anterior fragment of the j)rorean-coronal complex) become united. Oudemans calls
this sulcus simply " prorean." 1 think that Oudemans's and Ziehen's use of the term
" prorean," without being absolutely erroneous, fails to express the whole truth.
There can be no difficulty in interpreting the sulci lateralis and coronalis in terms
-which are applicable to the Apes, because these two sulci (in the form seen in the genus
Lemur) are exactly reproduced in some of the smaller Cebidae, such as C'allithn\r.
The lateral sulcus becomes the intraparietal (or " interparietal," as the Germans call it)
sulcus ; and the coronal sulcus is identical with that variously termed "fruiitalis
j))'iitcq)al/s," '• rectus^' " rostmlis,'" among many other designations. It includes the
representatives of the two sulci called respectively "rectus" and " arciiahis" in the
Apes. The latter certainly x-epresents the inferior precentral sulcus of the human brain ;
whereas the homology of the former is the subject of great divergence of opinion.
Herve, El)erstaller, Waldeyer, Giacomiui, and Dubois * consider that the sulcus rectus
becomes the sulcus frontalis medius of Eberstaller in the human brain ; whereas
Cunningham and Marchand incline to the old view of Gratiolet, that it represents the
sulcus frontalis inferior. As I have shown in the above notes that the fronto-orbital
sulcus (whicli Waldeyer, Giacomiui, and their school regard as the homologue of the
inferior frontal sulcus) is certainly the anterior limiting sulcus of Reil, the evidence
seems to point to the view of Cunningham and Marchand as the true interpretation.
Tiie jjostlateral sulcus is as variable in the Lemurs as its rejiresentative in the
Carnivora and other mammals is unstable. Beddard, Ziehen, and Flatau and Jacobsohn
tall it parieto-occipital ; Chudzinski labels it " occipital," a term which is ap^iarently
applied in the same way as " parieto-occipital " used by the other writers. Beddard
further complicates matters by applying the term " Simian" [/. e. Altenspalte] to it in
JS'ijcticebns (Proc. Zool. Soc. 1895, ]). lit).
In the discussion of the mesial surface of the hemisjiherc it is shown how misleading
and erroneous it is to call this furrow "parieto-occipital."
The postlateral sulcus undoubtedly becomes the sulcus occipitalis transversus of
Ecker in the Apes and Man.
* '• PitheraiifhropHs erecti's" riocecdwgs of the Fourth Litcniational Congress of Zooh)g_v at Cambridge, Ic'JS
<^,ubli8hed ISOO), p. s3.
MOKPHOLOGT OF THE BRAIX IX THE MAMMALIA.
415
It is unnecessary to enter any furtlier into tlu^ discussion of the later evolution of
this sulcus ill the Apes, Ibrmy views on this subject have been far more aptly expressed
than I am capa1)le of stating them, in Ciumingham's monog-raph *. There still remains
I'or discussion the most difficult problem in the interpretation of the L'rosiniian sulci,
/. e. whether the central (Rolando's) sulcus is r(>prescuted in any form in the Lemurs.
The central sulcus is easily recognizable in all the Siiniidie and Cercopitlieciche.
Nor can there be much uncertainty concerning its identity in tiie larger Celiida".
If the smaller Cebidae be studied without comparison with the larger members of the
family, it may seem very doubtful whether the small central sulcus in such a form as
(lirnsothrix (for example) might not really represent the ramus postcentralis of th«!
intraparietal sulcus. But an examination of the brain in tlu; whole Family shows that
the postcentral sulcus first appears long after the central itself is fully formed. So that
in those Cebida) in which only one sulcus is present in the appropriate place, it may be
regarded as the central.
The simplest form of a well-defined central sulcus may be seen in the geiuis Pithcci((
as a simple transverse sulcus in the interval between the intraparietal and " coronal "'
sulci.
In the genus Calllfhrir there is sometimes no trace of a central sulcus ; in other cases
there is merely a shallow depression in front of the intraparietal sulcus, and tiie situation
% 61,
P'uhecia. — Dorsal aspect of bniiii. Xat. sizp.
where one would look i'or the upper end of the central sulcus ; in other cases again there
is a small transverse furrow in this situation.
Beddard represents this sulcus in its best-developed form in the brain of CuUUhrix
torquata f, and calls it precentral.
The hraiu of CoUithriv is of peculiar interest to tlie student of the Prosimian organ,
because the arrangement of the sulci on the outer surface of its cerebral hemisphere
exactly reproduces that found in the Lemurs, as Sir William Flower forty years ago
* D. .J. Ciinniiigluim, "Surface Anatomy of the Primato Cerebrum," ' C'uuniugbam Memoirs,' Xo. VIJ. Itoyal
Irish Academy, 1892, p. 221.
t "On certain points in the Anatomy of CciUithrir foi-qi'rdi," Xovitates Zoologio;e, vol. viii. October 19(»1,
pi. xi. tig. 5 (dorsal aspect, wrongly labelled " mesial ").
SECOND SERIES. — ZOOLOGY, VOL. VIII. 00
416 . PEOF. 0. ELLIOT S:\iriH ON THE
showed. Tlie Sylvian, " lateral," " coronal," postsylvian," and orbital are (|uite lemuvoid
in position and constitution. The presence of the small and variable sulcus ia front of
tiu" intraparietal j^lateral] sulcus completes tlie resemblance.
The possibility of recogiiizinij,- a central sulcus in the Lemurs has recently been
discussed by Beddard *, wlio believes that " the lemui-s are to be differentiated from
the higher primates by the fact that their T)rains do not exhibit any trace of a fissure of
Rolando [central sulcus], so constant a feature of all monkeys and apes." To justify
the latter statement he regards the anterior extremity of the intraparietal [his " supra-
angular "] sulcus as the representative of the central sulcus, chiefly because it " is in an
ideal i^osition for a fissure of Rolando"; i.e. it is midway between the anterior and
jiosterior poles of the hemispheres, which according to Beddard is the position of the
central sulcus in the genera Ateles, llycetes, Cehus, Bmchijurus, and Lagothr'ix. This
argument, however, is quite fallacious because it ignores tlie fact that a relatively
enormous increase in the extent of the pallial area in front of the Sylvian fissure occurs
within the family Cebidic, so that in the lowlier members of the Family the central
sulcus, if present, must be placed relatively nuidi further forward in the hemisphere
than it is in the lar^rer "'enera. An indication (jf such a teudencv is exhibited in the
brain of I'UIiccid.
In the earlier memoir of Ziehen a very different interpretation of the homologies of
these sulci is suggested ; and Beddard makes no attempt to refute these arguments.
Ziehen regards the sulcus wliicli is found in front of the lateral sulcus in Pcrodicficus
as tlie central; he also considers that in Ni/aticcbiis the central sulcus is represented by
the two sulci /'and .;■ + y. 'Ilie sulcus /'in the genus Lemur is considered by Ziehen to
represent the central sulcus or the upper portion of it, whereas Elatau and Jacobsohn t
anticipated Beddard in regarding it as " precentral." These writers then proceed to
criticise the argument of Ziehen that the furrow/ represents the central siilcus, because
it occupies the appropriate position in regard to the lateral (intraparietal) sulcus. They
argue that the sulcus in question might with equal justice be i-egarded as the sujierior
precentral by reason of its relationship to the coronal (their sulcus "' frontalis"), utterly
ignoring the fact that the precentral sulcus never occurs in the absence of the more
stable central sulcus.
They further lu'ge, in opposition to Ziehen's suggestion, the T- or Y-shape of the
sulci, their sagittal direction, and the fact that, according to Chudzinski's diagram, there
is a sulcus resembling the supposed central sulcus of Lenin}- in front of a definite central
sulcus in the l)rain of Iiulris. They summarize their criticism of Ziehen's conclusions
regarding the central sulcus in these words : — " Die Moglichkeit, dass der untere der
beiden qtieren, hinter dem Sulcus frontalis [coronalis] gelegeneu Furchen [i. e. tlie
sulcus ^ of i<?»i«/- and the sulcus x f ^ of Xyct'tcehus] homolog dem Sulcus centralis
der Affen ist, lasst sich niit absoluter Sicherheit nicht bestreiten, wenn auch die Gestalt
uiul Lage der Furche luehr dem Sulcus pra-eentralis (arcuatus) ents])richt ; die obere
[sulcusy] der beiden queren Furcheu ist aber sicker ein Homolog des Sulcus pra?con-
« •>'.jvitates' iUOK p. :3(j5. t •Haudbutli; p. ITS.
MOIil'JlOI.OdV OF Till- BK.\I\ J\ Till-: .M A.M.MA LIA. 1.17
trails superiov, da sie bei den lioclistcn ArlcMi dvv llalh.-iffcn Ljenau wie bei don waliren
Afleii zwisclien Sulcus frontalis (princi2)alis) und Sulcus centralis yelegen ist " *.
It will lie seen, thercFore, that there is little semblance of agreement with reaard to
the central sulcus in the three most recent memoirs on tiie Frosimian l)rain.
There is no necessity to ag-iin review all the data atForded by a study of the genera
Leiiiur, Niicticchits, Ferodicticns, I'roplthccnH, and ChiromijH. It is sufficient to recall
that a very pronounced tendency is exhibited in these varied forms for the small
sulcus/ to fuse Avith the caudal extremity (/') of the coronal sulcus, with the resulting-
formation of a areat transverse sulcus, wliich can be nothing- else than the central
sulcus of the Primates.
Ox iiiK Ai-i-iNiTiKs OF Tin: Le.wi'i;s.
It does not come within the scope of this work to consider in all its hearings the
vexed question of the relationships of tli(> Lemuroidea. It would, indeed, be presump-
tuous to attempt such a diOicuit task, whi(di has balllcxl many zoologists with the best
equipment for performing it. All that I am concerned with at present is the setting
I'ortli of such evidence as the brain affords, and the determination of its value as an
index of the aflinity of its possessor to other manimals.
The greatest divei-geuce of opinion exists among zoologists as to the relative closeness
of the bonds w hich link the Lemurs and the Apes.
Many writers, such as Gervais and Alilne-Ed wards, have relegated the former to an
Order quite distinct and separate from the Apes ; and Plower virtually, though iu)t
nominally, subscribes to the same opinion, for he retains the Lemurs among the Primates
chiefly in deference to "the traditional views of zoologists." On the othei- hand,
Linuteus, Lesson, Iluxley, Broca, and Mivart assign them a subordinate rank in the
I'rimates. Nevertheless ^livait, who is generally quoted as a champion of this view, has
stated " that ihe ditfereuces between the apes and lemurs are so many and so great,
that it cannot be considered otherwise than in the highest degree improbable that dm
the evolutionary hypothesis) they took origin from any cominon rcjot-form that was not
equally the progenitor of other mammalian Orders "f. So that ]\livart also virtually
places himself in the ranks of those who favour a separate ordinal rank for the Lenuirs.
Dr. Forsyth Major's careful studies of the skeleton and teeth of the recent Lemurs,
and of an interesting series of extinct Lemuroids, within the past few years, have uttei'ly
stultified ITower's statement that "as a more complete knowledge of their organization
has been gradually attained, the interval which separates them structurally from the
monkeys has become gradually more evident." Forsyth Major's conclusion is so
diametrically opposed to Flower's teaching that, to quote his own words, his "position ....
w ill have to be expressed in classification by giving up the two sepjiratc suborders of
Primates, thus going one step further than Mivart "" %.
* Flutau and .Tac-nb.si.liii. • IlaiKlbiirli,' ji. 178. t Arliclo •' Ape," F]iicyc,lop;r(liu lirilaiiiiira.
X •• Suiimniry ol I'lcsuul l\iiuwli-'d<;e of Fxtiiitt I'liiiudc* from Madiigascar," Goologicul Maga/.iiic. n. .s., Uccado 4,
vol. vii. Xuvimlioi- I'JuU, \i\>. VM-AM).
60*
418 PEOF. G. ELLIOT SMITH ON THE
It is liavdlv necessary to insist on the Aalue of cerehral cliaraeters as an index of tlic
aflBnities of the animal, when it is recalled that the mammalian brain is perhaps the
only organ whicli can be trnly said to represent in itself tlie whole organism. For even
thou"h tlie l)rain is responsive to changes of habit in the animal, and innumerable other
factors which in various ways influence the jDroportions of its various parts and modify
its shape, yet, in its essential plan, it is perhaps the most conservative organ in the
whole body. Thus the same fundamental plan of tlie brain is found throughout the
Eutheria, and, with the exception of the striking changes in the constitution of the
cerebral commissures, it is common also to all the Metatlieria. In the foregoing notes
I have attempted to show that underlying the exceedingly variable and apparently
haphazard disposition of the sulci of the cerebral hemisphere — which, being tlie most
recently-moditied, are consequently the most changeable regions of the brain — there is a
fundamental group of homologous furrows, which are the common property of the
Avhole Meta- and Eutheria rather than the exclusive distinction of one Order.
The fact of having thus discarded as obsolete and unjustifiable the belief that each
Order of mammals evolves for itself a number of furrows which are incapable of being
liomologized with those of other Orders, may seem to have broken down the barriers
Avhich separate the Lemurs from other mammals (for even those who have most
magnified the difierences between Apes and Lemurs have been compelled to adopt
from t!ie higher I'rimates their nomenclature for the Prosimian sulci). But this is
by no means so. Eor, having demonstrated the identity of the various elements
Avhich may unite in various manners to Ibrm the most heterogeneous series of patterns
in the difi'erent Orders, we are the l)etter able to appreciate the Ordinal value of
these varied groupings of the series of sulci, the morphological values of wiiich we
know, than we should if a new set of features of unknown importance were evolved in
each Order.
It surely does not lessen the great systematic importance of the fact that the calcarine
and sujji'asylvian sulci behave in a very peculiar and identical manner in the Lemurs
and Apes, to know that the direct homologues of these furrows are disposed in a
strikingly different manner in the various other mammalian Orders. On the contrary,
the fact that the different patterns are composed of such stable elements enhances the
value of the plan of the sulci as an index of relationship.
The peculiar constancy of the arrangement of the calcarine, retrocalcarino, and
paracalcarinc sulci is alone sufficient to indicate the close l)onds of affinity which unite
all the Lemurs, and even such difTerent members as Toi'si/i-s and C'h/roii/)/.s, and at the
same time separate them from all other mammals.
The confluence of the calcarine and retrocalcarino sulci is peculiarly distinctive of the
Pi'imates, and in this respect the Lemurs conform to the Ordinal type. In many of the
Jsew- World Apes the paracalcarinc sulcus is much less perfectly developed than it is in
the Lemurs; and in this respect the latter closely resemble the highest Apes, in which
the paracalcarinc sulcus forms the ventral clement in that peculiar combination of two
sulci which is commonly distinguished as " parieto-occipital." In making this state-
ment I am not unmindful of the probability that this type of paracalcarinc sulcus may
be meiely its most primitive Ibrm.
MOEPHOLOGT OF THE BEAIX ]\ THE r^EAMMALIA. 11 <)
re.
The an-angcment and beliaviour of the sulci ou tlie outer aspect of the heniisph(
closely follow those of the Ape, and especially of the lo\vlier Cehidie. There is the;
same peculiar form of suprasylviau sulcus merged ia the Sylvian fissure in both
Families. The lateral sulcus is closely similar, as in fact it is in all mammals, hut it
shows a peculiar tendency, which is exhibited in NycllcebuH in (he Lemurs and in
Chnjmlhrix, Xiictii)itheci(.'<, and JEycctes in the Cebidte, to become concurrent with \\\o
suprasylviau (Sylvio-intraparictal union), which is a striking instance of the Avidf;
separation of the Primates from all other mammals (in most of Ayhich the first stage of
this process, /. e. the pushing, as it were, of the suprasylviau sulcus on to the pseudo-
sylvian sulcus, has not begun, so that tlie further stage of pushing (he lateral sulcus into
tlie Sylvian complex is not even possible); at the same time it is a further testimony to
the affinity of the Lemurs and the Cel)idfG.
So far as the other sulci — the coronal, the orbital, the diagonal (fronto-orbital), the
central rudiment, the postsylvian, and the postlateral — are concerned, there is tin;
closest identity between all of these in the Lemurs and such lowly Cebidu; as Cairdh,-i.i\
the importance of which is again enhanced by the contrast affonU^d by the beJiaviour of
their respective representatives in other mammals.
There is no sulcus, perhaps, more distinctive of the Primates than the noi-mally-
constituted central (Rolando's) sulcus. Such a sulcus (as a unit) is not found in any
other Order of mammals ; but there is a widespread tendency in the Lemurs — in
Terodicticus of the Lorisina>, in certain cases in FropUhecn.s, and in Cldroinijs — to the
development of this characteristic sulcus.
The fronto-orbital sulcus, w'hich, like the paracalcarine, is absent in many New- World
Apes, is often as fully de\ eloped in the Lemurs {Lemur, PropUhecns, and Indris) a^ it is
in the larger Cercopithecidtc. And even if this furrow (as in the analogous case of the
paracalcarine) is represented among the other mammals (in the diagonal sulcus), it
it still important that a furrow which is not ju'esent in many New-World Apes j-hould
occasionally develop) in the Lemurs in the same situation and have the same form as tiie
fronto-orbital sulcus in the larger Apes.
The degree of aflinity of one mammal to another is a purely relative matter, which it
is difficiilt to express adequately. But if we take all the features of neopallial sulci into
consideration, it can confidently be stated that, judged by the standard of the otlier
mammalian subdivisions, the ditTerences between the brain in the Lemurs and such
Primates as the Cebidfe ara ctn'tainly not worthy of being granted either Ordinal or
even Subordinal imjiortance.
The differences between the brain in the Edentate Pamilies Myrmecophagidw, Dasy-
podid;e, and Bradyjiodida^ (which we recognize to be united by bonds of affinity) are
considerably greater than those m hich divide the Lemurs and Cebidte. Or, to adopt a
more certain or, at least, more genei-ally recognized scale of relationships, there is a
much closer resemblance between the brain in the Lemiu's and Apes than thei-e is
between those of the ^Eluroid and Arctoid Carnivores.
So far as the neopallial furrows arc concerned, the Lemurs m.iglit be regarded as
inseparable from the Cebidte, if it were not for the fact that the other parts of the brain
exhibit-verv definite distinctive features.
420 PEOF. G. ELLIOT SMITH OX THE
'J'lie brain of a Lemur is relatively smaller and more macrosinatic than that of an Ape.
In addition, there are differences in the so-called occipital region.
Tlic data for comparing tlie size of tlie brain in the Lemurs with other mammals are
unfortunately somewhat scanty, and (if we neglect the wholly unreliable figures yielded
by weighing organs which have been modified by the action of preservative agents) are
almost \vhf)lly the result of the careful investigations of Professor Max Wclier of
Anisterdani f.
Tlie \vei"-hts recorded bv Weber i are as follows : —
3
Brain- Propniiloii to
SpiiciKs. wi'hjht. Budy-ux'itjht.
Leiii II r ruriiis, G coff . , (^ 3."? 1:1 0:>
„ ? 28-7 1:75
* „ „ „ e 39 1:74-3
Lriii/ir iiio)ii/oz, lAnu., $ ^8 1:76
$ 21-1 1 :60
Nijctkcbiis lard'u/radtis, d\a\ , ^ 8'l.S 1 : (il
„ 6 "■"•- 1 :•"'!•
Fcro(/ic//ciis jju/to, IjCiSon !>'o 1 : 7()
1'5-^^ 1:--
'■>■■>'■> l:''i
('I//roiiii/.s i/nidiiija.scariciisis, Geoff.. ? 42''J3 1 : 37
The record of the weight of the brain in CMj-om^s is identical with that of Oudemaus
(Xatuurk Verb. Kon. Akad. Amst. xxvii. 1890, p. 27).
As tlie result of an elaborate series of investigations, Professor Eugene Dubois arrived
at the conclusion that " in the Lemuridse, ]S\i/cficeb!is and Tars/us occupy the sanu^
dea-ree on the scale of brain development as measured by its quantity," and further tliat
" this is about equal to that expressed by the lower indices in the Ungulata and
Camivora" §. In the same place, however, it is stated that "the lower OkMA'orld
apes ... do not distinctly exceed in this respect the Ungulata and'Carnivora."
In Ziehen's tables there are three records (all by Max AVeber) of brain-weights in the
Ilapalidaj, ranging from 9-8 grm. (1 : 20) to 12-H grm. (1 : 26) ; and three records in
the genus Filhecia (two by Plower and one by Weber), showing a variation of 22 grm.
(1 : 20) to 3G-2 grm. (1 : 15) in the brain-weight. These numliers are sufficient to show
how much more riclily the New-World Apes are endowed Avith brain-sul)stance Than the
Lemurs are.
Duliois's statement concerning Xycticebiia and Tarsi us presumably does not ap})ly to
t ' Vurst udii-ii iibcr das Ilirngcwioht der Siiugethierc," Leipsig, ISUli.
t)ther records have been published by Burmeister, liischott', FLd:iu and Jacobsohii. and Ziehen, Imt the weights
;ire not those of fresh brains and cannot l)e corrected with any accuracy.
X All these weights except that marked with the asterisk (") are nuuted from Hardeleben's ' Haiidbuch der
Aiiatoniie des Menschen,' Nervensystem (by Prof. Ziehen), p. y<iS.
^ •• On Piducaiithru2ii'.s ereiiiis," Proc. 4th Intern. Congr. Zoo!., Cambridge, l^li^, p. V\.
MORI'IIOLOOY OF THE J3RAIX IX THE MAMMALIA. il'l
llio Leiinirs as a wliolo. It' avo examine tlie few reconls which arc quoted al)ove, and
compare them Avith data ohtaiued from othin- mammals in exactly the same manner hy
the same investigator, it w ill then be seen from a comparison of the hody-weii^hts of
those non-Primate mammals which possess the same absolute brain-weights as various
Lemurs, that the proportion of brain to body is considerably hi^-her in th(« latter.
Compare the series of brain-weights in the genus Lemur, wiiieli vary from 21-1 grm.
to 39-1 grra. (representing the followinij proportions of brain to body : — 1 : GO, 1 : To,
1 : 7<'), 1 : 8f, and the exceptional 1 : 103), with the following measurements (jf
Carnivora : —
liniin- Pi-djiDrl'ioii to
Spi:(1i:s. irfir/lit. Iliiihj-u-niiht.
I'ircrru taii(/(il(iiii;a, VtViiy '.\() 2 1 : lo;}
Fclis cutKs, Liiui '.VJA\ ] : lO;,
1-yh domcslica, Uui :W-5 1 : [() [ '^"' '"-'"^-^^ "*'
l 7 niPU'suroiiKMits.
, ;52 1 : ]2S
('fiiii.<! /ar/oj/>is, Ijinn. .'51 \ :•>()'>)
„ .'iD-l 1 : S4. i
Pdiiuloi iiriis miixdiij/ii, (ir:iv 'I'l 1:111
Ciilict's vittdld, Sclnx-h 21-1 1 : !»I
HiTjiestfs all/'icdiiilii, Ciiv 2'.Vi 1 : 81
In all of these Carnivora the proportion of the brain-weight to th(^ body-weight is less
(and in many cases very considerably so) than it is in Lemurs with brains of a corr(>-
sponding size. And in other mammalian Orders the disproportion in the size of the body
per unit of brain-weight is even more pronounced. The Lemurs ;ill have relatively
large brains in comparison with other mammals, but are much uku-c; poorly endowed in
this res])ect than the Apes.
The data are too scanty to enable us to decide whether this difference in the brain-
weights of .Vpes and Lemurs is due merely to the failure of the latter to keep ])ace in
brain-growth with the former, or is the result of an actual reduction in size.
The most noteworthy distinction between the brain of the Lemurs aiul the A])es is the
relative size of the olfactory regions of tli<' brain.
The l)rain of the Ajjcs is distinctly uiicrosmatic. The olfactory bulb is very sniall,
and is borne on a long attenuated peduncle ; the anterior part of the i^yriforni lobe is
so reduced as to be recognizable with difliculty, and the posterior part is so small that it
is almost always confined to the ventro-mesial aspect, and rarely extends on to the
lateral as^iect of the hemisphere ; ami the liippocam])us is relatively very small.
In mammals the size of the olfactory parts i>f the brain is subject to great
A'ariations.
Thus in all aquatic mammals they b(!come greatly reduced in size [e. cj. in Orniiho-
rhijuchus, the Pinnipedia, the Sirenia, the Cetacea), or even entirely absorbed (in the
Odoutoceti). In mammals which lead an arboreal life they ai"e relatively smaliei' than
in those which lead a terrestrial existence, e. ij. the rhineacephalon is relatively much
122 PEOF. G. ELLIOT SMITH OX THE
isiualler in the arboreal Cycloturiis tlian in tlie terrestrial Ilyrmecopharia, two Edentates
of the same family; but it is the peculiarity of the Apes that they have become
extremely microsmatic for no apparent reason, the dirnimition of their olfactory organs
being much more pronounced than it is in any other arboreal animal. This phenomenon
is undoubtedly to be mainly attributed to the increasing efficacy of the neopallium as an
" organ of mind," and its consequent usurpation of the rule of guiding the creature,
i\ hicli elsewhere among mammals is fulfilled chiefly by the sense of smell.
It is a very significant fact that the Lemur which exhibits evidence of being the
most primitive — Tarsias — possesses the most extensive occipital prolongation, and is
more microsmatic than any other of the Prosimioe. Thus the olfactory bull) of Tarsius
is only half as large as that oi llicrocebus; and yet the former is appreciably greater
than that of Ilidas.
The importance \vhieh has been attached to the amount of cerebellum covered by the
cerebral hemisphere as an index of the degree of brain-development is altogether
disproportionate to its true significance. By this I do not moan to imply that the degree
of caudal prolongation of the hemisphere is not a very important indication of the size
of the cerebral hemisphere, but it must not be forgotten that it is merely one of several
indications of the degree of cerebral expansion. The failure of the cerebral hemispheres
to completely overlap the cerebellum is an obvious point of distinction between the
brain of the Lemurs and most Apes. In the Lemurs the extent of the backward
pi'olongation of tlie hemisphei'e varies considerably. In Nycticebus, for example, there
is only a very narrow strip of cerebellum uncovered. In the Galagiufie and in Tarsius
the extent of the caudal prolongation of the hemisphere is perhaps greater than in any
other Lemur; /. e., if its dimensions are estimated in relation to the position of the
corpus callosum and by the size of the calcar, rather than by the situation of the caudal
pole of the bemisjihere with relation to the cerebellimi. The relative greatness of
the occipital extension of the hemisj)here in these particular forms is all the more
significant, because Tarsius and Galago are the Lemurs which have probably become
least changed (so far as the brain is concerned) from the early Prosimian type. It is
hence probable that the immediate ancestors of the Lemurs possessed brains of a more
pithecoid form. There are many other suggestive facts which point in the same
direction.
In most mammals the calcarine and intercalary (? paracalcarine) sulci become
continuous (probably for purely mechanical reasons), and it is hardly conceivable that
tlie small caudal extension of the hemisphere in the recent Lemurs is sufficient to
so completely dissociate the calcarine from the intercalary sulcus, and to lead to its
confluence with the retrocalcarine sulcus, or in fact to supply the mechanical factors
which call the latter sulcus into being. It is difficult to conceive of the present relations
of the three sulci of the calcarine group having been produced except in a brain with a
mxich more pronounced extension of the hemisphere than at present exists.
It has been shown by Prof. A. Milne-Edwards, and emphasized by Dr. Eorsyth Mnjor*,
* ' Xovilates Zoologicie,' vol. i. lyjk p. 33,
MORniOLOGY OF THE BRAIN I\ THE MAMMALIA. 423
that in the young Pi-opitheexs Edwardsi tho caudal extonsion of tlic cerebral liemispliere
over tlic cerebellum is distinctly greater than it is iu the adult. Trom an e.^amiuatiou
of the cranium of a young (15 days old) PropithecuH corouatus, which Dr. Forsyth
Major lent me, I have been able to establish ibis also for aiiother species of a later age.
However, in the brain of a young (15 days old) Lemur fulvii.s; which Captain Stanley
Flower gave me, the cerebellum was exposed to as great an extent as in the adult.
In different specimens of adult Lemurs, even of the same species, I liave found
considerable variety as regards the amount of cerebellum uncovered.
If we consider the marked disproportion in the size of the brain iu the Lemurs and
the Apes respectively, the possibility presents itself that the former may at one time
have been much larger than it is now. If such were tho case, it is certain that th(?
process of reduction in size would be chiefly expressed in a curtailment of the caudal
(occipital) prolongation.
A study of the peculiar series of variations in the Indrisina?, Cliii-oniijs, and to a less
extent in all other Lemurs, clearly siiows tliat the hypothesis of a retrogressive change
can alone explain this extreme variability.
It may be urged that it is not the mere retraction, so to speak, of the caiulal pole of
the hemisphere whicli distinguishes the occipital region of the Lemurs from that of tlie
Apes, but also the absence of the posterior cornu in the former. This is a feature the
importance of which has been grossly exaggerated. It is merely one of the expressions
of a large neopallium, and the question of the presence or absence of the calcar and
calcarine sulcus is not in any way involved in the issue. In any large mammal (large
mammals being provided with actually, though not relatively, more extensive neopallia
than the small) the increase in size of the neopallium may lead to the formation of
a posterior cornu. We see this in the Carnivora, Ungulata, and Cetacea, as well as iu
the Primates. The Primates, being preeminent in neojmllial greatness, naturally present
in most cases a distinct posterior cornu.
The direct result of the "retraction" of the occipital extension of th(! hemisphere
would be the obliteration of the posterior cornu.
The brain of Tarshts, which in most respects is so eminently Propitliecoid, is dis-
tinguished from other Lemurs by the possession of a distinct ijosterior cornu indis-
tinguishable Irom that of the smallest New- World Apes.
When it is recalled that the Eocene Lemuroids jiossessed brains no smalhn- in size
than those of the recent lorms (Cope), it would indeed be surprising if the ])raiu should
have remained stationary in size through all the succeeding ages.
So far as the brain is concerned, the Galagimc are (with the exception of Tarsias) the
most generalized of all the Lemurs. And, in view of the foregoing suggestions, it is not
without significance that they also possess the most pronounced occipital prolongation.
The cerebral features of tlie members of the genera Lemur and Hapalemur are not
far removed from those of Galago. The peculiar variability in the extent of cerebelhuii
covered by the cerebrum is suggestive of a " retraction " of the latter.
Far more specialized than these genera are the Indrisina', Lcptdolemui- being in most
respects intermediate between the latter and Lciinir. In different modes the Indrisine
SKCOND SEllIKS. — ZOOLOGY, VOL. VIII. 01
42't PEOr. G. ELLIOT SMITH O^ THE
type of brain is ])eculi;u-ly specialized in the genus Chiroii/i/s and the extinct Lemuroid
JIcguhuhipLs.
It is impossible, with any degree of accuracy, to place the peculiarly pithecoid form
of brain which GloUlennir presents. So far as its characters can be determined, it
resembles the type of the genus Lemur and the Gulaginae.
The Loi'isinse again present peculiar cerebral featru'es. In many respects the brain
is peculiarly simple, after the same manner as tliat of tlie Galagime. In FerocUcticus,
and to a less extent in Ni/ctlcehas, there is a peculiar tendency toward the formation of
the typically Primate central sulcus.
Tarsius is obviously related, so far as the evidence of the brain is concerned, to the
Galaginee. This resemblance cannot be disguised even by the peculiar distortion of
the brain in the former. It is in every respect distinctly Prosimian. But it also
exhibits tAvo interesting series of features. So far as the corpus c:illosum, the hippo-
campal formation, and the cerebellum are concerned, the brain of Tarsius conforms
to the simplest Eutherian type, such as we find in the more generalized Insectivora —
for example, Erliiaceus. Yet its relative microsmatism (in comparison with other
Lemurs), the proportion of its hemispliere which is placed on the caudal side of the
corpus callosum, and the presence of a definite posterior cornu of the lateral ventricle,
all indicate a nearer approach to the Simian condition than occurs in the brain of
any other Lemur. Tai-sins possesses at once the most generalized and the most
pithecoid brain of all the Lemuroidea.
In this connection it is interesting to quote some remarks concerning Aintptoniorphus
which were made seventeen years ago by E. D. Cope: — " The brain and its hemis^jheres
are not at all smaller than those of the Tarsius, or of the typical lemurs of the present
period. This is important in view of the very small brains of the tlesh-eating and
luigulate Mammalia of the Eocene period as yet known. In conclusion, there is no
doubt but that the genus Anaptomorphns is the most Simian lemur yet discovered, and
probably represents the family J'rom Avhich the anthropoid monkeys and men were
derived" *.
Zittel also has remarked that the Eocene Lemuroids {Adapis, Mlcrocluei'iis) of Europe
" combine features of the existing Lemurs and true A])es " f.
It is not easy to draw any certain conclusions from the evidence of the brain as to the
relations of the Primates to other mammals.
The larger Carnivores show a tendency toward the development of a Sylvian complex
like that of the Primates. There is a singular al)sence of any such tendency in the
Ungulata. Like the latter, how^ever, the Primates always retain a coronal sulcus, which
in the main is nearer the horizontal than the vertical direction. It is a peculiar fact
(hat the separation of the calcarine and intercalary sulci, which is so marked a feature
of the Primates, should occur (practically) nowhere else in the Mammalia, except in the
three Edentate families Myrmecophagidic, Bradypodidse, and Manida^. In the case of
* "Tlie Lemuroidea and the Liscotivora of the Euoeiie Period of Xorth America," American Naturalist,
May l^s."), p. 407.
t Quoted l.y IL 0. Forbes, 'llonkeys,' vol. i. p. 111.
.A[O]{IMI0LOGY OF THE BKAT\ IX TliF. .M A.MAr.UJA. 12-")
the Anteatei's there is anotlier peculiar Primate feature^: the suprasylvian and pseudo-
sylvian sulci blend to form a Sylvian lissure. it is quite impossible to say how much
importance is to be attached to such ])henoinena.
After seeking for some explanation for ail the apparently conflicting features of the
Prosimian brain, the following tentative working hypothesis as to the ancestry of
the Lemurs shaped itself in my mind, and I ins(>rt it here merely as a slender bond
connecting certain facts scattered throngh these notes.
The brain of the Primates was derived from some lusectivore-like type, tiie cerebral
hemispheres of Avhich attained a precocious development and, as one of the expressions
of their greatness, bulged backward over the cerebellum. In consecpience of this great
extension of the " physical organ of the associative memory of visual, auditory, a-ivd
tactile sensations," the sense of smell lost the predominance which it exercised in th(>
piimitive mammal (and in all the Orders of recent mammals), and the olfactory parts of
the brain rapidly dwindled. This early Primate developed its distinctive type of
calcarine sulcus and " Sylvian lissure," the lateral, coronal, ajid orbital sulci, and
the characteristic central sulcus.
In the keen struggle for existence, the Lemurs ceased to keep pace with the other
Primates so far as the increase in the size of tlie brain is concei'ncd. Thev became
more specialized, and their brain prol)ably shrunk, thus leading to a retraction of
the occipital pole of the hemispheres.
AVith the diminution of the size of the neopallium the sense of smell comes to play
a more important part, and a secondary re-enlargement of the olfactory regions occurs.
The blotting-out of the rliinal fissure may be an indication of this jjhenomenon.
This somewhat crude and tentative scheme is put forward to give expression to
the view that the Lemurs are highly sj)ecialized Primates which share most of their
brain-features in common with those of the Apes, and that some at least of the
apparently primitive characters may be due to a secondary retrogressive modification of
a more highly-developed type of brain.
ADDENDUM.— The Cerebellum.
(Pteceived April 21, 1002.)
In my account of the brain in the Edentata*, I called attention to the fact (which
had previously been recognized independently by Stroud and Kuithan) that underlying
the apparently irreconcilable differences and seemingly divergent designs exhibited in
the cerebellar architecture of mammals, there is one common fundamental plan which
becomes variously elal)orated in its details in different animals.
In attempting to describe the cerebellum in the Edentata I found that the nomcnclatiuv!
and mode of subdivision usually adopted in works on Human Anatomy were so
ill-adapted to the needs of Comparative Anatomy, that it was quite impossible to describe
* '• Thf Liiiiiu in the Edentata," Trans. Linn. Soc. ser. 1', Zool. voL vii. IS'JO, pp. 3G0 et si'q.
61*
■i2C) PROF. G. ELLIOT S.AIlTIi OX THE
the cerebellum of many mammals in the current terminoloo-y ; and that even in the case
of those Avhich conld be so described, the account would be altogether unnatural and
hopelessly cumbrous. The chief reason for this is that the cerebellar fissure (fissura
horizontalis magna), ^^liich is regarded in Human Anatomy of such fundamental
importance as to be invariably taken as the starting-point in the primary subdivision, is
a most insignificant cleft of quite secondary importance in most mammals, and is even
absent jiltogether in others.
It therefore became necessary to seek some more stable line of demarcation between
the various cerebellar regions. Accordingly I proposed a tentative scheme of subdivision
based uj)on the recognition of the fissure called " preclival " as the primary natural line
of separation of the mesial parts of the organ. My reason for adopting this fissure was
the fact that it is present in all mammals, without exception, and is, moreover, the
deepest fissure crossing the mesial plane. Ample confirmation of the justice and
jiaturalness of this selection was afforded by the fact that two other investigators
had independently come to the same conclusion — Stroud, as the result of a study of the
development of the organ in the Cat and in Man, and Kuithan from embryological
studies on the Sheep.
Fig.ez.
cgp.
^'^P- p. Cr. py. Ca.
I'arsios s/jcclriim.
Left lateral aspect of the brain-stem and cerebellum, exposed bj- cutting through the thalmo-striate
junction and removing the cerebral liemisphere. x 4.
When I examined the cerebellum in Tarsias, JlicroceljHS, Galago, Lemur, JS^/fcdcebus,
and CJiii-Oiiii/s, it was not sur25risiag to find that it conformed to the plan wliicii has
])reviously been shown to prevail among such divergent groups of mammals as the
Ungulata, Edentata, and Carnivora. Yet I was not j)repared to find within the Primates
a form of cerebellum such as that of Tarsixs, which so nearly approaches the most
generalized mammalian type met with in the Insectivora, Marsupialia, the Dasypodidse,
and some Rodentia.
In the one Order we find therefore every gradation of cerebellar form, from the
simplest and most generalized mammalian type to the most complex, though not
the largest, example of cerebellar architecture. The undoubtedly close affinity of the
different Primate genera enables us to speak with greater certainty of the appareiit
MOKPHOLOGT OF TIIK HRAIX I\ Till-: >r.\ MM ALIA.
427
homologies of the various parts than was the case when I was discussing sucli widt'ly
separated mammals as the Edentates and Man.
Fig. 63.
fs.
uv.
no.
Lemur macaco.
Diiigrara representing- a mesial sagittal section of tlio cerebellum. x 2.
The accompanying series of drawings (figs.G4« to (/) representing mesial sagittal sections
through the cerebellum in Tursius, in the Insectivore Erluucens, in the Marsupial
Perameles (the pattern of which is identical with that of the Insectivore Ilacroscelldcs),
and in the Marsupial Notoryctes (the simplest cerebellum in the Mammalia), will serve
to indicate the dilfercnt manner in which essentially the same pattern becomes
elaborated in detail.
-':f3.
id ) fi.o.
Diagrams representing corresponding sections in (a) Tarsiii'!, X 4 ; (b) Ennaci'MS, x 4 ;
(c) Peniine/es, x 4 ; and (d) Xotori/ctes, x 8.
Such variations of the same simple plan might be adduced almost without limit front
among the small Insectivora, Marsupialia, Chiroptera, Rodentia, and Edentata, but a
comparison of Tarsliis with a small Insectivore and a Marsupial will suffice to explain
the general principle.
428
PROF. G. ELLIOT SMITH OX THE
It will l)e seen that, iu spite of its smaller size, the cerebellum of the Insectivore is
slightly richer in furrows than the Marsupial, and that, in order to liud in tlie former
Order a cerebellar pattern of the same degree of richness as Femmeles exhibits, we must
select some small form such as IlacrosceUdes or Oryzornctes.
It Avill be seen that the pattern found in Tarsius is much richer in fissnres than that
of either the Insectivora or the Marsupialia. And, if it be objected that this contrast
may be due to the smaller size of the organ in l^riuacem, the suggestion can be at once
i-efuted by showing that in the larger Insectivore Gymuura, in GaleopUJieeus, in many
Rodents like Lepus, and in the representatives of other Orders of a corresponding size,
there is a greater poverty of fissures than there is in Tarsius. This fact might be more
accurately stated by saying that the extent of the cerebellar cortex is determined, partly
bv the dimensions of the animal and partly by its zoological rank.
Fi^.65.
yp. la.
Fig. Co. — Ltmvr macaco. Dorsal as];ect of the cerebellum. x 2.
Fig. G6. — Gazdla dorcas. Left lateral aspect of t lie cerebellum. X 2.
The mesial part of the cerebellum is divided into three regions by means of the deep
lissura prima, y//'., and by tl'.e shallower fissnra secunda,/s. (fig. 6A). The region iu front
of the tissura jn'ima (which is homologous Avith the so-called "preclival" tissvu'e) may be
called the anterior lobe, that between the fissnra prima and the fiss. secunda the central,
or, better, middle lobe; the third region being the posterior lobe. The anterior and
middle lobes may be quite simple vmdivided folia, as in Notoryctes, but the posterior
lobe is ah\ays subdivided into two lobules, which correspond to the parts called
" nodulus " and " uvula "' respectively in Human Anatomy.
A comparisoai of a series of mammalian cerebella clearly sIioavs that the jDOsterior lobe
does not increase in size to so marked a degree as the anterior and especially the middle
lobe do ; and, conversely, the ultimate subdivision of this lobe is evident at a much
earlier phase in the phylogenetic history than that of the other two lobes.
The most noteworthy featiu-e of the mesial section of the cerebellum in Tars/'/ts is the
plumpness aud relative greatness {i. e. iu comparison witli tliat of other small mammals)
of the middle lobe of the cerebellum.
The lowermost part of tliis lobe is much more deeply cut off from the rest than it is
in Er'uiacetts and Fei-amclcs. The simple folium so formed represents the " pyramis "
of Human Anatomy. It is the dorsal lip of the tissura secunda, and the rest of the
:\10HPll()L0Gr OF THE RRAIX IX THE MAMMALIA. 42!)
middle lobe is mueli more deeply incised, /. e. has a much more extensive surface than
that of all other mammals of a corresponding' size.
Wherefore the cerebellum of Tarsius exhibits definite signs of superiority over that
of other mammals when it is studied in mesial section.
The resemblance of the surface of the cerebellum in TarshiH to that of other small
mammals is much closer than one might expect from a study of the mesial section.
The two lobules (nodule and uvula) of the posterior lobe extend outward for a short
distance only and then seem — to the naked eye — to stop abruptly. If, however, a series
of sections be examined microscopically, this lobe will be found to extend outwards as au
exceedingly attenuated baud of cortex which suddenly expands upou the lateral aspect
of the middle peduncle of the cerebelkun to form a sliort plump bilobulated mass — the
llocculus.
The pyramid (which is the ventro-caudal lobule of the middle lobe) is ])rolonged
outward into a broad band on the caudal aspect of tlie cerebellum. It arches outward
and downward parallel to the margin of the uvula, so that it reaches the lower margin
of the cerebellum. It then passes forward and expands, just above the lioccidus, into a
narrow, widely-projecting rod of cortex — the paradoceulus.
This parafiocculus is enclosed in a bony capsule ibrnied by the petrous temporal.
It is that iiart of the floccular lobe which is commonly called tlic " flocculus," but it is
not the flocculus of Human Anatomy.
The cortex on the upper side of the paraflocculus is divided into a series of five or six
bands by means of shallow notches. The under surface is smooth. The distal extremity
is bifid.
It is the distinctness of the broad cortical band joining the pyramid to the para-
flocculus A\liicli specially emphasizes the resemblance of the cerebellum in Tarsin.s and
that found in the generalized mammals. In most Primates this simple relationship
becomes so completely disguised as to be quite unrecognizable.
Por reasons Aviiich will appear later, it is desirable to have some term Avitli which
to distinguish the ])and of cortex linking the pyramid to the parafiocculus. I shall
therefore call it the " copula pyramidis." Exactly how much of the uniform band
(in Tui-sius) is pyramid and how much is copida, will be appreciated later when the
different fate of the two parts will be studied.
The copula pyramidis forms the lowermost lobule of the lat(n-al part of the middle lobe.
The rest of the latter is subdivided into three parts, which I distinguished by the
letters A, B, and C in rhe Edentate cerebellum [op. cit.).
The area A is a simple band which forms the posterior lip of the fissura prima
and extends outwards and forwards (on the anterior or cerebral surface) as far as tlir
parafiocculus.
The area B is a little rounded knob which forms the most laterally-projecting part
of the so-called " lateral lobe " of the cerebellum. It is subdivided into three folia l)y
means of two deep fissures. The area B is wedged in between the lateral extremities ol'
the areas A and C, and is connected to the mesial part of the middle lobe only by a very
narrow stalk, the limiting furrows of which are alwavs shallow and sometimes absent.
430 PKOF. G. ELLIOT SMITH OX THE
The area C is a little bifoliate iiodule placed iijion the caudal surface lietwcen the area B
and the copula pyramidis which may he regarded as an area D. Unlike the area B, C is
freely connected "with the mesial parts by a broad band.
The tissures which indent the mesial part, so-called "vermis,"' of the middle lobe
do not extend, into the lateral regions. The limiting tissures of the areas A, B, C, and D
arise laterally and do not extend, on to the mesial area in Tarsius.
The anterior lobe ditfers in appearance from the other two lobes. It is broadest
in the middle line and gradually tapers toward the region of the entrance of the middle
peduncle into the cerebellum. Its surface is incised by two deep and tliree shallow
fissures, the former of which extend, as far as the middle peduncle.
In the brain of Lemur, the pattern exhibited in a mesial sagittal section of the
cerebellum is much more complicated than that of Tarsim, but the plan is essentially
the same (fig. 65). It so closely resembles that of Orycteropm which I have described in
my memoir on the brain in the Edentata {op. cit. fig. 29, p. 362), that it is unnecessary
to repeat the account. In the accompanying figure I have inserted letters to represent
fissures homologous Avitli those similarly distioguished in Ori/cteropiis.
The general plan of tlu^ cerebellum closely conforms to the type found in such
mammals as the Dog, the Anteater, and the smaller Apes.
The floccular lobe is divided, as in all mammals, into a ventral part — the floecidus —
and a dorsal part — the ])araflocculus. Tlie flocculus con.sists of a little wedge-shaped
mass closely applied to the lateral aspect of the pons Varolii immediately in front
of the tuberculum acusticum and external to the trigeminal nerve. Its broad anterior
aspect is subdivided into folia by five or six horizontal fissures. The posterior extremity
of the wedge-shaped mass is connected by a long white baud with the nodulus.
The paraflocculus consists of an ii'regular mass of folia placed on the dorso-lateral
aspect of the flocculus, so that in a lateral vicAV of the brain the latter is almost
completely hidden by it. From its lowermost part (/. e. the region nearest the flocculus)
a narrow stalk emerges and swells out into a plump projecting mass of folia arranged in
a feather-like pattern. This projecting part of tlie paraflocculus is often called the
" petrosal lobule" and more commonly simjdy " flocculus." It presents a similar form
in all Apes except the Simiidse. In Hylobates, however, the typical Primate foi'm
of floccular lobe is fully developed.
The nearest approach to this type of floccular lol)c among other mammals is exhibited
in the Ungnlata, of which the accompanying drawing of the lateral aspect of the
Gazelle's cerebellum is typical (fig. QQ).
In Tarsius the paraflocculus has been described as being directly linked by a simple
cortical band to the pyramid. In Lemur a vastly different state of aflairs obtains.
Starting from the stalk of the so-called petrosal lobule, a series of folia (and not a
continuous undivided band) forms a worm-like coil which curves forward, then upward
and horizontally backward (compare the Gazelle's cerebellum, fig. 66). The series of
disconnected folia then proceeds backward below the area B to the lower end of the
area C, and it becomes impossible to accurately sepai-ate the series of folia belonging
to the parafloccular (or perhaps it wovdd be more accurate to say " copu'Iar ") group
MOEPIIOLOGY OF THE BliAIX IX THE jr.\M.M.\LIA. 431
from tbose of the area C. The connection of the copula pyrainidis with tlie pyrainiil
becomes either entirely destroyed or reduced to exceedingly insignificant proportions.
From comparative studies of the udult cerebellum it is quite imiiossible to decide the
fate of the copula pyramidis in the larger cerebella. In many brains it seems to dwindle
or completely disajipear. In others again, especially Siiiiia salynts, it seems to undergo
enormous expansion and form the greater ])art, if not the whole, of tiie body which
is called tonsilla (amygdala) in Human Anatomy. This is a problem which can only ])e
certainly settled, by a study of its developmental history in a series of mammals.
The relations of these parts are considerably disturbed in the genera S/iiiic, JiUhrop<i-
jnilu'cus, and Homo by the extreme dwindling of the paraflocculus and the consequent
disappearance of its •' petrosal " part. The parafloccular remains become the flocculi
secuudarii of ITenle,
Of the various subdivisions of the middle lobe of the cerebellum in Lemur, the area A
is subject to perhaps the least change. It becomes furrowed by numerous new fissures,
but they are approximately parallel to the fissura prima, and hence do not greatly
disturb the pattt-rn. The area A represents the lobus lunatus posterior, and is mesially
continuous Avith the region between the fissura prima and the fissura b, which is called
the clivus monticuli in Human Anatomy (tig. 05).
The area B is a very jdump rounded mass "ivhieli forms the lateral pole of the
cerebellum (fig. 65). It consists of twelve vertical folia radiating in a feather-like pattern
from a very narrow vertically-placed leaf-like stalk, the mesial part of which is deeply
buried between the areas A and C. This stalk does not reach the nu;!sial plane, for
it becomes fused to the posterior aspect of the area A. The area B is the lobus postero-
superior. There is no separate folium cacuminis.
The area C is a plump vertical worm-like band wedged in between the area B and the
mesial parts of the middle and jiosterior lobes on the caudal surface of tlu; cerebellum.
Its connection with the suprapyraniidal part of the " Aermis " is constricted and depressed
in a vertical furrow. In the Apes tliis vertical band rapidly increases in breadth, and
the direction of its folia becomes oblique.
The general features of the anterior lobe are remarkably constant in all Mammals.
The differences consist chiefly of an increase in the number of lissurcs in proportion
to the size of the cerebellum.
In all other Prosimi:e tlie structure of the cerebellum does not vary to any marked
extent from that of lAtiiur. In the smaller members the fissures become much fewer,
and in Mieror-ebus we find an orgt,n which differs very slightly from that of Tarsins. In
Chiromys tbe primitive connection between the pyiamid and tlie parallocculus, wiiicii
has been seen to become obscured in Lemur, is retained in a distinct form.
[Since the foregoing pages were in type, seme supplementary notes Iiave been received from tlie
author. In order not to delay the issue of this part, it is intended to publish the notes mentioned in an
early number of the ' Journal ' (Zoology), vol. 28.— Sec. Lin.n. Soc]
SECOND SEKIES. — ZOOLOGY, VOL. VIII. (j^
(32 3IORPII()L0GY OF THE BRAIN IX THE AI.AMMALIA.
Key to the Sectioxs of the trecedixg Memoir.
Introduction 'M\)
Ijiteratnre 3:21
The Coreliral Ilciiiisphcrc in f.cnnir '.V2'^
The siinie in Hapalemvr :>'57
The s;mK' in 1 .cpidolehuir .3-58
Tiie Brain in tlic Lorisina; 33!^)
Tlic same in the Galaginie 3 K!
The same in tlie Indrisinse 319
The Cerebral Hemispiiere in Danbenton'ni [Clurotnijs) 3")!
'I'he Brain of the Extinct Lemuroid Globilcmur 3(!;i
The same in MrgaJadapis 3G I-
The Cerebral Hemisphere in Tarsius and the Pi'osinii;e 'M'i7
The Braiu-stem 373
Tlie Calcarine Sulcus and the Oc(dpital Prolongation of the Hemisphere '\7Ct
The ^Morphology of the Sulci on the Lateral and Doi-sal Aspects of the Ceri'bi'ul
Hemisphere . . 3i).')
The Morphology of the Sylvian Fissure ■!().")
On the Affinities of the Lemurs 417
Addendum — The Cerebellum -12.")
lU'ference-lcttcrs emjiloyed in the Text-cuts, with Ex])l;iiiations 320
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THE
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XI. On the Ncio Zealand Phijllohranchiafe Crustacea-IIacrnra.
By George M. Thomson, F.L.S.
(Plates 27-29.)
Rend 17th April, lOOi'.
1 HE proposed publication of a V)iographical census oi' the New Zealand Tauna makes
it desirable to supervise many groups of animals, the memliers of whicli have been
imperfectly described and in many cases incorrectly named. No groups liave suffered
more in this respect than the Crustacea, and, as a small contribution to our knowledce of
this part of the fauna, I have lately been examining the Prawns and Shrimps of these
islands, and find that very consideral)le revision is required in orfl(>r to classify them
correctly.
In Miers's ' Catalogue of the New Zealand Crustacea,' published in 1876, a summary of
the then known species was given. Twelve species of Phyllobranchiate Macrura are
there briefly described, but some of these are doubtful or incorrect. Rhynchocinetes
iypus, a species stated to be found also on tlie coasts of Australia and Chile, is introduced
into the list of New Zealand species on the authority of Britisli ]\Iuseum specimens.
Haswell also introduces it into the list of Australian Crustacea, apparently taking his
description from Miers's Catalogue. No collector has, so far as I know, met with this
species in this Colony, nor does it occur in any of the various local museums. I think,
therefore, that its inclusion in our fauna is a mistake. I have recently received specimens
of this very distinct and beautiful species from Mr. David G. Stead, of Sydney.
Again, Heller, in the Crustacea of the ' Novara ' (p. 105), describes a Slirimp from
Auckland, without any reference to its habitat, as CarkUna cnrcirostrls. This turns out
to be the same as mj Leander jlitviatilis; but both names are equally incorrect. The
species is found only in fresh water, and belongs to the genus Xiphocarls.
The inclusion of Atya 2)llipes in the New Zealand fauna is due to an error, as first
suggested by Miers himself, since the species is found in the Mid-Pacific islands, but not in
New Zealand. Spence Bate (' Challenger ' Report, p. G9;3) also states that Atya arnuda
and Atya splnipes occur in New Zealand, but this is probably also an error.
Of tlie remaining species in Miers's Catalogue, Alpheus nuvce-zealandice was described
by him from specimens in the British Museum. This species does not appear to have
been met with since, but this only emphasizes the fact that so few examples of the
Crustacea occur in our local museums.
The 'Challenger' collections added twelve species, belonging to seven genera, to
SECOND SERIES.— ZOOLOGY, VOL. VIII. (33
43i MR. G. M. THOMSON OX THE NEW ZEALAND
the local fauna, but none of these are represented in our museums. They are all
deep-sea species.
The following list includes the names of all the species kuowji up to the present
time.
Tribe C R AX GO N I D E A..
Tamily CRANGONID^E.
Genus 1. Pontophilus, Leach, 1817.
1. P0NTOPHII.TJS AUSTRALis, G. M. Thomson. (Plate 27. figs. 1-5.)
1879. Crangon austraUs, Hutton, MS. ; G. M. Thomson, Trans. N.Z. lust. vol. xi. p. 231, pi. 10.
fig. A 1.
Carapace (fig. 1) about one-third of the length of the animal, the whole surface some-
■\vhat rugose with very short hairs, and bearing five irregular longitudinal rows of spines.
The rostrum is very short, blunt, and hollowed above, and does not extend beyond the
eye-stalks. The eye-sockets are semicircular, tlieir lower limit defined by a sharp spine.
The margin then inclines a little forward, and at its lower edge ends in a rather long-
acute spine ; the whole margin is rather setose. On the median line the carapace is
slightly ridged and carries two short spines ; on each side and behind the orbital spine
is a second row of four spines ; and on the lower lobe of the carapace are two acute spines,
the posterior one at a little distance from the margin. Tlie pleon is nearly smooth. Its
second segment is shorter on the dorsal line than the first, but its lower mai'gins
are produced both backwards and forwards into rounded broad lobes. The third
and fourth are subequal in length ; the lateral margins of the fifth are produced
posteriorly into a two-spined lobe. The sixth segment is two-ridged above, and ends on
tlie sides in a three-spined process. 'J'he telson reaches to the extremity of the tail-fan ;
it is narrow and tajjering, its margins are fringed with setae, and it ends in three minute
teeth and a number of plumose hairs.
The eye-stalks are short, stout, and bhmt ; the eyes are reniform.
The first antennae (fig. 2) have the broad plate on the outside of their base reacliing
nearly to the extremity of the first joint; the latter is pi'odnced on its outer distal
margin ; the second and third joints diminish in length ; the flagella are short.
The second antennre have the scaphocerite about twice as long as broad, produced
considerably beyond the peditncle, and ending in a strong tooth.
Th(^ third maxillipeds are mtich elongated, extending considerably beyond the i)latc of
the fir.st antennie. According to Spence Bate ('Challenger' MaeriuM, p. 182), the
dactylos and 2)ropodos are wanting in the members of this family ; Avhile Stebbing (South
African Crust, p. 17) says : " in the third maxillipeds it is clear that the geniculation
occurs between the fourth and fifth joints, and in all probability the si.xth, which is much
longer than the fifth, represents a coalescence of the sixth and seventh joints." In this
PHYLLOBRANCHIATE CRUSTACEA-MACRUEA. 435
species (fig. 3) the basal joint bears a sliovt appendage (l)asecpliysis) not reaching to the
extremity of the meros. The iscliium is very sliort and almost completely merged into
the next joint, the meros. The carpus is about half as long; wh'iU) the terminal joint is
again elongated, bearing a few spines on tlie lower margin among tiie numerous hairs,
and ending in two very short spines.
The first pair of pereiopoda have the meros (tig. 1) keeled on tlie upper margin and
produced into an acute spine, the lower margin being flattened and Avingcd to partially
receive the hand when bent inwards. The hand is about three times as long as broad,
and its inferior margin ends in a straight spine ; the palm is very oblique.
The second pair of pereiopoda are very short and feeble, only about half as long as the
succeeding pair ; both fingers of the slender chela; are subequal and fringed with liairs.
The third pair are long, very slender, and stiliform, and extend beyond the extremity
of the flagella of the first antennte.
The fourth and fifth pairs are strong, witli long aiid acute dactyla.
The posterior pair of pleopoda (fig. 5) are subequal with the telsou; the outer branch
has the outer margin terminating in a short subapical tootii ; the inner is narrower and
rather longer.
The colour is a uniform sandy grey ; occasionally the fifth segment of the pleon
is coloured dark brown or has patches of brown coloration on it.
The following are the dimensions of a large ovigerous female : —
mm.
Extreme lengtli 38
Breadth at second scgnicut of carapace 7
Lengtli of carapace 1
o
„ scajjliocerite of first antcnnrc fi
„ last segment of pleon .">
„ telson 7
ITahitat. This is a common species on the New Zealand coasts within the 20-fathom
line, as the following list of localities shows ; — Dusky Sovuid, Paterson Inlet (Stewart
Island), Otago Harbour, Blueskiu Bay, Akaroa, Lyttelton Harbour, and Cook Strait.
Also at Napier {Filliol).
Distribution. Confined to New Zealand.
2. PONTOPHILUS GRACILIS, Batc.
1888. Bate, 'Challenger' Macrura, p. 187, pi. 87.
Hahitat. Off the New Zealand coast, east of Cape Turnagain, in 1100 fathoms.
Dtstrihation. Off" Tristan da Cunha, near Torres Straits, and near the riiilippine
Islands.
63*
43G yili. G. :S[. THOMSON OX THE NEW ZEALAND
Tribe P O L Y C A R P I D E A.
Family PROCESSID.E.
Genus 1. Glyphocrangon, A. Milne-Edwards, 188i.
1. GlAPlIOCRANGON REGALIS, Bate.
1888. Bate, ' Challciiger' Macrura, p. 517, pi. 93. tigs. 3 & 4.
Habitat. North of the Kermadec Islauds, 600 fathoms.
Distribniion. Off Eiji and Banda Islauds, at de^jths of 200 and 315 fathoms.
Eamily ALPHEID.E.
Genus 1. Alpheus, Fabricius, 1798.
1. Alphei's socialis (Heller). (Plate 27. figs. 0-12.)
1865. Heller, Voy. ' Novaj-a,' Crust, p. 106, pi. 10. fig. 1.
1876. Miers, Cat. N.Z. Crust, p. 8.2.
The carapace is produced into a very short acute rostrum, the lower margin of which
is straight ; over each orbit it is produced into a rounded lobe which ends in a minute
spine ; the posterior margin is straight on the dorsal portion, but on the sides is slightly
produced backwards; in length it is barely one-third as long as the body.
The tirst antennae have the basal scale about as long as the first joint of the peduncle ;
it is broad and flat above, and the basal portion of its outer margin is finely serrated ;
the second joint of the peduncle is twice as long as the first. The outer fiagellum
is stouter than, but less than half the length of, the iimcr (which is about two-thirds as
long as the animal), and tends to divide into two at the apex.
The second antennae have the basal scale reaching to the extremity of the peduncle of
th(,^ first pair; the basal spine is about half as long and very acute ; the flagellum nearly
as long as the body.
The first pereiopoda are very unequal, the left being the largest ia all my specimens.
In the largest specimens the propodos of the left limb has the upper and lower margins
nearly parallel, the upper having a narrow groove or channel, "while the lower is
transversely rugose ; the dactylos is strongly ciu'ved and bent outwardly, while on the
inner side it is furnished witli a stout blunt tooth. In smaller ovigerous specimens tlu;
dactylos is rounded and less claw-like, as if it had lost its chelate function (fig. 9). At
the hinge of the dactylos and propodos the opposing surfaces are flat and circular. In
the right limb the dactylos is about half as long as the propodos, is Avell developed, and
slightly curved ; the inner faces of the claw are densely fringed with hairs.
The second pereiopoda are considerably longer than the succeeding pair. The three
I'lIVLLOBEAXCHIATE CRUSTACEA -MACRUllA. 437
following paii's have tlie propoda (fig. 10) fuvnished with numerous spines on the lower
margin, and the dactyla with a strong inner tooth.
The last pair of pleopoda have the inner branch evenly rounded ; the outer (fig. 12)
crossed near the extremity by a strong diairesis, the outer margin of Avhich carries a
strong sjiine.
The telson (fig. 11) bears two spines on its upp(>r surface near each margin ; its
truncate and slightly rounded apex is about half as wide as the base, and ends iu
a fringe of sette.
o
The following are the dimensions of a large male : —
mm.
*Eiitire leugtli 4g
Length of carapace l,j
Deptli of carapace K)
Length of pleou 21
,, 1st perciopod (left) 37
,, propodos of same 22
„ dactylos of .same 8
„ 1st pereiopod (riglit) .3-1,
„ propodos of same 18
„ dactylos 10
„ telson (J
Habitat. This species occurs fairly commonly in the Xorth Island, and as far south as
Cook Strait and Tasman Bay. I have specimens from tlie following localities : —
Waiwera, Auckland, Hauraki Gulf {L. F. Ai/soii), Moko Hinau {Sandacjer), Portland
Island {Eohsou), Wellington Harbour {Fat-qiihar), and Tasman Bay, Nelson. I have
never obtained any specimens from the east coast of the South Island, the waters of this
part being more or less cooled by an Antarctic current, and Alpheus being apparently a
genus which loves the warmer seas.
DlstrihiUion. Australia.
Mature specimens sent me from Port Phillip by ]\Ir. S. AV. Fulton, and which are not
more than an inch (25 mm.) long, agree in nearly all details with the above, only the
large left hand is like those of my smaller specimens in having the rounded dactylos, the
supraorbital spines are much more produced (tig. S), and the extremity of the telson is
somewhat more rounded.
In the 'Challenger' Macrura (p. 510), Spence Bate gives a list of species of Alphens,
and refers A. sociaJis to Australia, and ^L. chirayricus, M.-Edw., and A. vorcc-zealamluc,
Miers, to New Zealand. According to M. -Edwards (Hist. Nat. Crust, vol. ii. p. 351),
A. chirayi'icKs occurs in the s as of Asia, and I have seen no subsequent reference to its
occurrence in New Zealand. Pnnn thc^ brief description given by M. -Ed wards it is a
distinct enough species.
* 111 all the rostrum-bearing species, 1 have measured the leiigdi i'rom tlie point of thi; rostrum to the
s.\ws. <>i the telsou.
138 ME. G. M. THOMSON ON THE NEW ZEALAND
2. Alpueus xov.e-zealaxdi.e, Miers.
1S7G. Micrs, Ann. & 'Slag. Nat. Hist. ser. 4, xvii. p. ;224 ; Cat. N.Z. Crust, p. 82, pi. 2. fig. 2.
I do not know tbi.s species. Miers savs it " is distiugui-shed by the absence of spinules
on the upper orbital margin, the shorter basal spine of the external antennte, ^c," —
characters almost too indetemiinate to found a specific distinction upon.
Genus 2. Bet-EUS, Dana, 1852.
It is difficult to find a definite character on -SThich to base the generic distinction
between Betceus and Alphens. The two points emphasized by Dana are the absence of a
rostrum and the inverted position of the propodos of the first pereiopods. But Stimpson
gives as tlie principal character of the Australian B. trispinosus, " front with a long
needle-like ro.strum, and armed with two acute orbital teeth, which are half the length
of the rostrum." In our Xew Zealand species these orbital teeth are present, though
small, and are not on the margin of the carapace, but on the front of the eyes. The
t«i.sting of the first pereiopods, by which tlie dactylos is on the lower side of the propodos,
appears to me to be due to a slight twisting of all three joints — the meros, carpos, and
especially the propodos.
The ova, as pointed out by Sjjcnce Bate (Chall. Macr. p. 564), are much larger and
more oval than the small spherical ova of Alphens. The only New Zealand species
appears to be very distinct. Spence Bate, considering a mutilated specimen of an
Australian species, B. microslyhis, was inclined to assign it to B. (equimanus, but his
figures are conclusive against this identification.
1. Betjels iEQUiMANis, Daua. (Plate 28. figs. 1, 2.)
18i32. Dana, T'.S. Explor. Exped. xiii., Crust, part i. p. JGO, pi. 3.j. fig. 11.
1876. :\Iier.s, Cat. N.Z. Crust, p. 8."i.
The front of the carapace is notched about as deep as the eye-sockets (tig. 1), above
which it rises slightly, and is again slightly notched beneath tlsem ; its antero-inferior
angle is rounded. The posterior margin is somewhat convex and acutely notched on
each side of the dorsal surface, the postero-lateral portions being produced backwards
into a rounded lobe; which lies under the first segment of the pleou.
The second segment of the plcon has the sides greatly produced both forwards and
backwards.
The tclson (fig. 2) is wedge-shaped, its apex ending in a bkmt tooth on each side, with
a somewhat rounded portion between, bearing a row of plumose setse.
The ophthahuopods are short, depressed, and slightly projecting from under the front
of the cara2)ace ; the eyes are nearly circular, and occupy the greater portion of their
upper surface. In front they are produced into a small spinule.
The first antenna^ have the joints of the peduncle subequal in length and diameter, the
first being slightly the longest. The basal scale reaches to the extremity of the second
joint. 'J'he outer short nagellum shows the normal tendency to divide into two at the apex.
PIl YLLOBRAXCII lATE CRUSTACEA-MACRUKA. 439
The second autemuB have the pedimcle only slightly exceeding that of the first pair.
The basal joint is produced into a short triangular lobe on the lower inarijin ; tiie
scaphocerite reaches nearly to the end of the peduncle, and its outer spine is short. The
flagella arc thick and tapering, and though I have not been able to observe the
movement, I am inclined to think they help the animal to spring.
The third maxillipeds are slender, and do not reach to the extremity of the peduncle of
the first antennfe.
The first pereiopoda are not very strongly developed and are .subcqual in form ; the
triangular carpos is produced on its lower side into a rounded lobe ; the propodos is
about as long as the rest of the limb and slightly curved at its extremity ; the dactylos
is short and curved, and, as well as the opposed pollex, is somewhat hairy at the tii).
The second pereiopoda are normally long and slender ; the carpos is 5-jointed, the
three middle joints Ijeing together equal to the first and last in length.
The succeeding pereiopoda are very strong and stout, and are used for walking over
tlie rocks and among the T^ced in T\-hich this species is found; the tiiird pair are
the strongest.
The f>leopoda have short, broad, foliaceous branches, except the first pair, which have
one of the branches long and slender, and the other rudimentary.
The last pair, forming the tail-fan, are nearly twice as long as thetelsou, and the plates
are broad and spreading ; the outer is divided by a distinct diaeresis into two nearly equal
parts, the upper margin ending in a strong spine ; the inner is ovate in form, and ends
in two sti'oug spines.
The integument is somewhat rough vrhh harsh points, especially on the limbs, and
these are rather hard and brittle, as is so often tlie case with shore species.
The colour appears to be dependent on the surroundings. The specimens from Stewart
Island, taken under stones, were of a uniform brownish-red colour; those from Moeraki,
caught on the seaweed, were olive-green.
The normal mode of progression appears to be by walking, but when disturbed the
animal escapes by vigorous leaps of a foot or more in length.
The following are the dimensions of a large male
niiu.
Entire Icnjjth .'JO
Length of carapace 10
,, telsou ;}
Deptli of carapace 5-3
Length of 1st pereiopod 10
„ 1st antennse to end of flagelhun 11
Habitat. This species occurs most commonly under stones or among Aveed between
tide-marks, less rarely in rock-pools. It is probably common on the coast. I have it
from the Bay of Islands (where Dana's original specimens were collected), "Waiwera, Capsi
Campbell, Moeraki, Dunedin Harliour, Stewart Island, and Chatham Islands (U. B. Kirlc).
rilhol records it from Cook Strait.
I) ist I'll lit ion. Confined to Xew Zealand.
4J.0 ME. G. -AI. THOMSON ON THE NEW ZEALAND
FainUy HIPPOLYTID.E.
Genus 1. Alope, White, 1817.
1847. ■\Vliite, Proc. Zool. Soc. p. 123.
1848. White, Ann. & Mag. Nat. Hist. ser. 2, vol. i. p. 225.
1876. Miers, Cat. N.Z. Crust, p. 84.
Cai-apace smooth, with a supraorhital spine and a suborhital tooth on each side,
liostrum short, armed with teeth above, and springing from a deep groove.
Ophthalmopoda short, stout ; ocellus well developed.
Eirst antennae short, Avith two flagella.
Second antennae with a large scaj)hocerite and a very long flagellum.
Mandible with shortened cutting-tooth (psalistoma) and a three-jointed palp
(synaphijiod).
First luaxillae 2-branched ; second j)air 3-branclied, aa ith wide mastigobranchial j)late.
Eirst maxillipeds with a 2-lobed mastigobranchia.
Second maxillijieds with a short podobranchial plume.
Third maxilliiieds very long and pediform, without any trace of branchiae.
First pereiopoda strong ; chelae well developed.
Second pereiopoda very slender, long, and minutely chelate; carpos 7-articulate (or
i)-articulate) *.
Third to fifth pereiopoda slender, with bifid dactylos.
Telson long and narrow.
The branchial formula is as follows : —
h i 7c 7 m n o
Pleurobranchise .... — — 1 1 1 1 1
Poflobranchia 1 — — — • — — — ■
Mastigobrancliife ... — r r r r r r
The mastigobranchiae are extremely rudimentary, consisting of a minute tubercle with
a mere trace of a plume.
Miers placed this genus in the Alpheidtr, but its distinct rostrum, exposed ophthal-
mopods, mandibles with reduced j)salistoma, the symmetrical first pereiopods, and
narrow tapering telson lead me to include it among the Hippolytidae.
1. Alope palpalis. White. (Plate 28. figs. 3-12.)
1847. White, /. c, &c.
1874. Miers, Zool. 'Erebus' & 'Terror,' Crust, p. 4, pi. 4. fig. 1.
The carapace is broad, widening posteriorly ; in front it bears a 4-toothed slender
rostrum, two teeth being near the point of origin, and two on the distal portion ; the
lower margin is entire. On each side of the rostrum, and about half its length, stands a
* See specific description, p. 441.
PIIYLLOBRAXCIIIATE CRUSTACEA-MACRUEA. 411
strong- acute spine. Below the orbit the front margin is produced into u siiort tooth.
The posterior margin of the carapace is rather deejjly hollowed dorsally; the siiles
are deej).
The pleon is about twice as long as the carapace ; its second segmeut is not very niucli
produced on the sides.
The telson is long and tapering, with a deep groove along the centre ; its extrcMnitv
hears two long spines, with a short one at each side (fig. l).
The first antennoe (fig. 5) have the basal Joint broad and produced on the outer side
into a spine which reaches to the middle of the second joint ; the inner side l)(>ars a sniiiU
tubercle near its base; the apex of tin; joint bears a transversB comi) of small spinules
l)oth above and below. The second and third joints together barely equal the lirst
in length, and the latter bears a very short joint on its outer side, from wliicli sjiriugs tlie
short thick flagellum, the inner flagellum b(;ing long and slender.
The second antenn;e have the peduncle slightly exceeding in length the peduncle of the
upper jiair. The scaphocerite is long and rather narrow, and is produced into a spine on
its outer margin, -which is subcqual with tiie peduncle in length. The flagellum is lon"'er
than the whole body of the animal.
The mandibles (tig. G) have a cylindrical molar process, a short and much-reduced
cutting-tooth (psalistoina), and a 3-jointed palp (synaphipod).
The first masillas (tig. 7) are two-branched, the larger (outer) branch consistiu"- of an
obovate plate fringed with seta-.
The second maxilla (lig. S) is three-brandied ; tlie inner l)ranch small, verv short, and
broad; the second long and plate-like, bilid, and fringed with sette ; the third slender
and stiliform ; mastigobninchial plate wide, produced both back and front.
The first maxilliped (fig. i)) bears a bilobed niastigobrancliial plate, a flat broad basal
plate fringed with seta\ a long cylindrical branch, and on the inner side of the latter a
smaller more slender branch (ecphysis ?).
The second maxillipeds (first guathopoda of Spence Bate) (fig. 10) have a short broad
plate on the inner face of the coxa, and on its outer a short branchial plume ; the Ijasos
carries a long cylindrical branch fringed towards the extremity with setoe ; the ischium
and meros are short, broad, and curving round ; the carpos is broader tiian long, and
bears a very broad but short propodos fringed with hairs.
The third maxillipeds are long and pediform ; the basal joint is furnished with a
slender single-jointed branch, and there is no trace of branchi;e on the lin\b.
The first pereiopoda are well developed, and are rough on their outer surface and
edges with small spinules ; the long triangular meros aad the short carpos (which latter
ends above, or rather on the outer side, in two or three strong tec!th) have deep sockets at
their extremities to receive the l)ase of the succeeding joint.
The second pereiopoda (tig. 11) are hnig and slender ; the basos is very short, the
ischium lono- and divided into two unequal joints, the meros still longer, and also divided
into two joints, the carpos seven-jointed, and the propodos short, with a minute chelate
dactylos.
The relative value of the joints of this limb is rather difficult to estimate. If we count
SECOND SJilUES. — ZOOLOGY, VOL. VIII. 6Ji
442 ME. G. M. THOMSON OX THE NEW ZEALAND
the first four joints as normal, and then regard all those between the fourth and the
])ropodos as forming the carpos, then the latter is nine-jointed. But a glance at the
limb itself seems to suggest that this is a forced and not very natural mode of treating it.
The normal mode of folding the limb into three parts takes place at what I have assumed
to be tbe distal ends of the ischium and meros respectively. On the other assumption I
should describe the ischium and meros as straight, the latter about half as long as the
former, the carpos as nine-jointed, first joint about as long as the two preceding, second
half as long, remaining seven usually bent at a considerable angle and not quite so long
as the first two.
The third to the fifth pereiopoda are avoII developed and diminish in size posteriorly.
The dactyla (fig. 12) are strongly toothed below so as to l)e two-clawed.
A small tubercle projects from the sternum between the bases of the third pereiopoda,
and a larger keeled protuberance between the bases of the fourth pair. This latter sternal
segment is produced posteriorly into two small acute lobes (fig. 3).
The first pleopoda in the males have a stout basal joint longitudinally folded at right
angles ; the two foliaceous plates are slender and acute, and the outer projects
outwardly nearly at right angles to the inner, so as to stick out from the sides of
the animal like two minute fans.
The succeeding three pairs of pleopoda have the basal joints rather long and stout, and
the foliaceous plates subequal with them in length. Tiie fifth pair are shorter and
thicker.
The sixth pair have the foliaceous plates subequal, narrow and rounded at the end,
and somewhat exceeding the telson in length.
The dimensions of a male specimen were : —
min.
tMitire length 77
Leiigtli of caraj)ace to exti'eniity of rostrum 25
Depth of carapace 14
Length of rostrum from first dorsal tooth 11
„ i)lcoii 52
,, telsou 12
,, first antoniup 28
,, second anteiiuie 85
,, peduncle of same 12
,, external maxillipeds 5i
,, first pereiopoda 33
,, second pereiopoda 37
Uahitat. I have only met with this species in rock-pools from the following localities . —
Cape Cami)bell, Sumner, Moeraki, Otago Heads, and Stewart Island. Eilholalso records
it from Cook Straits.
Distribidioii. Australia.
PHYLLOBRANCIIIATE CRUSTACEA-^^IACEUEA. 41.3
Genus 2. IIippolytk, Lcacli, 1815.
The generic character, as limited by Speuce Bate, is i^ivcn at length in tlie 'Challenger'
Macrura, p. 587.
1. HippoLYTE BiFiDiPvOSTias, Miers. (Plate 28. ligs. 1,'] IG.)
1870. Virhhis bijidirostnn, Miers, Ann. & Mag. Nat. Hist. ser. 4, xvii. p. 2:.'l ; Cut. N.Z. Crust, p. 81,
pi. 11. tig. 1.
Carapace with a jorominent acute spine over the eje-socket, a second subocular spiiiuh;
on the front margin just between the bases of the two ])airs of antennjK, and a tliinl just
close to the margin behind the base of the second antenna; infero-anterior angle
sul)acute. llostrum subequal with or slightly longer than the carapace ; upper margin
with two teetli placed just above the eyes, and a minute notch at the apex ; lower
margin with five to seven teeth.
The 2)leon is sharply bent (in all spirit-specimens) at the extremity of the third
segment, the dorsal margin of which is slightly produced and hood-like. Tlie fourth and
fifth segments have their inferior margins produced considerably back into rounded and
subacute lobes respectively.
The telson is long, narrow, and slender, ending at its sliglitly rounded apex in a row
of small spinules.
The ophthalmopods (fig. V.\) are rather elongated, and scarcely dilated towards the
extremity ; the eyes only occupy abt)ut a third of tlieir length.
The first antennte (fig. It) have the first joint of the peduncle flattened, and broadened
on the inside into a narrow wing which ends in a spine ; the Imsal scale is acute and
nearly as long as the first joint of the peduncle. The outer flagellum is short and
imperfectly segmented, as seems to be usual in species of this genus.
Tiie second antenna- have the peduncle very shoi-t, and the scaphocerite oval and
reaching beyond the extremity of the rostrum ; the flagellum is very slender and often
as Ion 2: as the entire animal.
The third maxillipeds (fig. 15) are long and stout ; the tliird joint is subequal in length
to the iburth and fifth together ; the latter terminates in a number of minute spines.
The first pereiopoda (fig. 1(5) are very short and stout ; the carpos is produced on its
upper margin into a blunt spine; the propodos is subquadrate in form.
The second pereiopoda are nearly twice as long as the first pnir ; tlie iscliium, meros,
and carpos are about subequal in lengtli, the latter being three-jointed and having tlic
middle joint the shortest ; the propodos is short and straight, witli a stout straight
dactylos.
The succeeding pairs of pereiopoda are long and well developed ; th(> nropoda have a
few long spinules on their lower margins, while the long straight dactyla are pectinately
spined.
The pleopoda are well developed, and the inner branches of tlie females bear a well-
developed process (st\lamblys of Spence Bate).
(U»
Ml, MR G. :^r. THOMSON ox THE XEW ZEALAND
The sixth pair are narrow and rather longer tlian the telson; the oater ou(! with
a two-spioed outer margin and a well-developed diajresis.
Tlie dimensions of a mature female were : —
mm.
Entire length 41
Length of carapace 14
„ rostrum 7
„ sixtli seguicnt of jileon i
,, telson 6
,, first antcnnre 28
,, first perciupoda 5
,, second pereiopoda 7
Colour. Specimens taken in the trawl outside Otago llarhour were l)right green
when alive.
Habitat. Kenepuru {J. McMal/mi), Otago Heads, 20 fathoms (trawled) ; Paterson
Inlet, Stewart Island, 10 fathoms (dredged).
Distribution. Confined to New Zealand.
Genus o. Merhippoltte, Bate, 18S8.
' Clialleiiger ' iMaciura, p. (IIS.
1. Merhippolyte spinifrons, M. -Edwards.
]8;i7. Hippolijtc .sjj>iiifruii.s, M.-Edw. Hist. Nat. Crust, ii. p. 377.
18 13. Hipiwlyte s/Anifnrns, Wliite, Dieff. N. Z. ii. p. 2(i8.
1876. Hi/ipoli/te .spinifronft, jNIiers, Cat. N.Z. Crust, p. SO,
LSS."). Hippuliite sp'niifrons, Filhol, ]\H.ss. de I'lle Cainpbi'll, p. 131, pi. 53.
The following is the description given hy Filhol (/. c. p. 431) : — " The smooth carapace
hears on its anterior and median portion a very strong spine (rostrum), which reaches
lieyond the front of the eyes. This spine has on the anterior portion of the upper margin
two acute teeth directed forwards. Two much smaller spines are produced on the median
line immediately hehind the first one. These are reduced and are directed upwards and
forwards. Immediately outside of the frontal anterior spine are two other long spines
without teeth, which project as far as the space hctween the two little teeth on the
rosti'um. The ocular peduncle is completely hidden. The first antenme have the hasal
joint rather large, and they hear two flagella, the one long and slender, the other short
and strong, and ajjout half as long as the preceding one. The second antennae have only
one flagellum ; the outer portion of the upper margin of the hasal joint is two-toothed ;
the jjalp is moderately enlarged. The first two pairs of feet tire slightly developed and of
equal length ; each of them ends in a small didactyle hand. The second pair is shorter
than the tliird, and its carpos is inultiarticulate ; it ends in a very small didactyle hand.
The third pair of feet is the strongest. The upper margin of the third joint of the last
three pairs of feet ends in a small spine; the extremity of tlie fingers hears on its inferior
PHYLLOBKAXCHIATK CRUsTACF.A-.M VCkl-R.V. 1. 15
mavgiii three or four little tufts of hairs. The third maxillipeds are long, pcfliform, and
rather large; the terminal joint ends in an acute apL'x. The length from the apex of
tlie frontal spine to the extremity of the telson is 37 mm. The telson is remarkahle on
account of the triangular form of its apex ; it l)ears three pairs of small spines, of whieli
one is very much reduced and scureely visihhi.
" This species, which I collected at low water in the rock-pools, appears to me to he
rather ahundant. It occurs, without any indication of its hahitat in New Zealand, in
the collections of the Museum of Paris. The development of the suborhital spines
enables it to be recognized at the first glance."
Habitat. Lyall's Bay, Wellington {FUhol).
Didrihidion. Confined to New Zealand.
1 have not met with this species.
Spence Bate suggests that J/, oriculalis, one of the ' Challenger' species desci-il)od by
him, and which was taken off New Guinea in GUO fathoms, is the same species.
Genus 4. Nauticaris, Bate, 1888.
'Challenger' :\r;uTura, p. 602.
1. Naiticaris STEWAirrr, G. M. Thomson. (Plate 29. tig. 1.)
1888. Hippoh/te Sfm'nrti, G. ]\L Thomson, Ti-ans. N.Z. Inst. xkI. p. 2:,<.), pi. i:*.. fig. 1.
Cara]jace relatively deep, depth nearly equal to half its length ; suriace smooth ;
suhocular spine well developed ; infero-anterior margin produced into a small spine,
nostrum very acute, ahout as long as the rest of the carapace; upper margin with six
prominent teeth, lower produced deeply and with two acute teeth.
Pleon with the third to the fifth segments jjroduced l)ackwards on their lower margins
into acute teeth ; sixth segment also produced back into an acute tooth on either side
ahove the lower margin, which hears the characteristic movable spine.
Telson narrow-oval ; apex rounded, liearing tAvo spines and a number of slender
setfE.
Ophthalmopods short and thick ; eyes occupying the slightly dilated upper half.
Pirst antenme have the basal joint loiiger than the next two together; stylocerite
reaching to extremity of the second joint ; inner flagellum not nuich longer than the
peduncle, considerably stouter than the outer.
Second antenme with a slender peduncle, equalling that of the first pair in length ;
flagellum ? (missing); scaphoeerite twice as long as the peduncle, very narrow and
tapering, the spine reaching beyond the foliaceous plate.
The mandililes, maxilkie, and maxillipeds clost^y resemble tlic same organs in
y. mafioiiis as figured by Spence Bate ('Challenger' ^[acrura, pi. lO'^).
The pereiopoda are similar to those of 3'. marionis. In the second pair the carpos is
li-15-jointed. In the succeeding pairs the claws are all didaetyle.
IM) MR. G. M. THOMSOX OX THE XEW ZEALAND
The following arc the dimensions of the only specimen : —
mm.
Length of entire animal 28
„ carapace 10
J, rostrum 5
,, plcou 15
,, tliird segment of pleou 2"5
„ sixtli „ „ 2
„ tclson 3
Hahttal. A solitary specimen was taken by the dredge in Paterson Inlet, Stewart
Island, in 10 fathoms.
Distrihuiioii. Confined to New Zealand.
This species is so near N. mariouis, Bate, that I retain it with some hesitation. It
differs markedly iu the dentition of the rostrum ; and although this is a very conspicuous
cliaracter, yet I am inclined to think that it is a variable feature, and consequently of
little value as a specific distinction. Bate describes the telson as hearing three pairs
of spinules, rather an unusual feature in the family. My specimen has only the
usual two.
The ' Challenger ' specimens of 1^. marionis w"ere taken off Marion Island in G9 fathoms,
and Prince Edward Island in liO fathoms, both localities nearly due south of Africa, and
off the Falkland Islands in South America iu 12 fathoms.
Family PANDA LID./E.
Genus 1. Plesionika, Bate, 1888.
1 . Plesionika semil.i;vis. Bate.
1888. Bate. 'Challenger' Macrura, p. 014., pi. 113. fig. 3.
Habitat. Off the Kermadec Islands, 520 fathoms.
Distribution. Philip])ine Islands, Borneo, Fiji, and east coast of Australia, from 250 to
1200 fathoms.
Genus 2. Pandalus, Leach, 1817.
1. PANDALLiS MAGNOCrLUS, Bate.
1888. Bate, 'Ciialleuger' Maerara, p. Wir, pi. 115. fig. 1.
Habitat. Stations 16G and 1G7 in Tasman Sea, west of Xew Zealand.
Histributioit. Contiucd to New Zealand.
PHTLLOBEANCHIATE CRUSTACEA-MACRURA. U?
Tribe M O X O C A E, P I D E A.
Family ATYIDJE.
Genus 1. Xiphocaius, von Martens, 1872.
Xiphocar'is, von Martens, Arch. f. Natnrjj. xxxviii. pt. 1, p. 139 (1872).
" Rostrum slender, compressed, dentate, usually long. All the pereiopoda slender and
■witli exopodites ; carpal joints of first two pairs without a distal excavation, articulating
in a normal way with the proximal end of the propodus. Abdomen with sixth segment
elongate; telson slender, truncate at tip." (-l/'O-y J. linf/Jjii,/,' The I5rachyura and
Macrura of Porto Piico,' p. 118.)
1. XiPHoc.vRis cuRViROSTKis (Heller), G. M. Thomson, (Plate 29. figs. 2-13.)
1865. Caridina curvirostr/s, Heller, Voy. Novara, Crust, p. 10,").
1870. Ciiridina currirostris, Miers, Cat. N.Z. Crust, p. 78.
1879. Leander fluviatilis, G. ^M. Thomson, Trans. N.Z. Inst. vol. xi. p. ^.''i], pi. 10. lig. A -.1.
Carapace nearly twice as long as deep, the lower margin curved in towards the
body ; front furnished with a prominent spine above and another Ixdow the eye-
socket. Rostrum slender and slightly curved upwards, 11-12-toothed above, beginnin;^
as a slight crest on the carapace, with two or three teeth bi'liiiul the eyes, a central
group of four or live in front of the eyes, and three or four small teetii close to the
apex ; 4-()-toothed below.
Picon having the second segment greatly dilated in the females, extending forward
over the posterior margin of the carapace; fourth and fifth segments with their infero-
posterior margins produced back into angular projections, which are more or less fringed
with setae. Sixth segment nearly as long as the fourth and fifth together.
Telson (fig. 3) rather long, narrow, and slightly tapering ; on each margin it beai"s two
short spines on its distal half. The extremity is slightly round(!d and ends in two very
short marginal spines and about six slender short sette.
The ophthalmopoda are nearly pyriform, the upper half being occupied by the large
rounded eye; the peduncle has a minute rounded lobe just at its base on the inner side.
The first antennre (hg. 1) have the peduncle reaching almost as far as the extremity of
the rostrum. The first joint reaches beyond the ophthalmopod, and from the base of its
outer margin there projects forward a stout stylocerite, which reaches halfway along the
next joint ; its outer margin is also somewhat produced. The second joint is shorter
than the first, while the third is only about half as long as the second. The flagella are
both rather slender, the outer being imperfectly divided into two.
The second antenna? (fig. 5) have a slender peduncle reaching only to the middle of
the second joint of that of the first pair; the second joint is jjroduced on its outer margin
into a short spine. The scaphocerite is produced along its outer margin into a spine
which reaches the extremity of the ^teduucle of the first antcnnie, while the oval-pointed
418 MK. G. -M. THOMSOX OX THE NEW ZEALAND
foliaccons plate is produced considerably beyond it ; the flagellum is slender and nearly
as long as the body of the animal.
The mandibles (fla;. G) liave a very distinct curved molar process and a well-developed
four-toothed cutting-plate ; there is no palp jjresent.
The first maxillte (fig. 7) are small, feeble, and thi-ee-branched ; the outer branch is
short, ovate, and bears two tine terminal sette ; the middle one is the largest and broadens
distally, its edge being fringed Avith short spines; the inner is short and has a rounded
finely setose edge.
The second maxillae (fig. 8) consist of two distinct processes: an inner foinned by two
broad, straight-edged, overlapjjing plates with a thick fringe of seta3 on their edges, and
overlapped at their base by two smaller, rounded, fringed plates ; the outer process extends
forward into a slender, rounded, fringed plate, and backward into a long slender portion
ending in long settp, whicli lie in tlie l)ranchial cavity of the body. Between the two
main portions of the limb there is produced a very small, slender, and rudimentary one-
jointed process, only visible under the microscope.
The first maxillipeds (fig. t)) are two-lobed, the inner lobe being semicircular and
fringed on the straight inner edge with numerous setae, the outer shorter and thinner ;
between them, and apparently projecting from the basal joint, is a long, slender, slightly
fiatteued process.
The second maxillipeds (fig. 10) have a short podobranehia on the coxal joint, and a
long plumose branch (ecphysis) on the basos ; the ischium is short, the meros still
shorter, whik^ the carpos is dilated into a short fi-inged plate ; the propodos is Ijent
against the preceding joints in the form of a large fi'inged plate. I can find no trace of
a dactylos.
The third maxillipeds (fig. 11) are long and pediform, four-jointed ; the short basal
joint carries a long exojiodite ; the next three are long and subequal, the terminal one
bearing numerous spines.
The first pereiopoda (fig. 12) are comparatively sliort and stout ; the exopodite reaches
nearly to the propodos; the carpos is sLort, broadened at its deeply^ excavated apex; the
stout propodos articulates with it at its lower angle; the dactylos and poUex are tliick,
spoon-excavate on their inner faces, aird furnished ■\^■ith a thick tuft of hairs at their
extremities.
The second pereiopoda (fig. 13) arc considerably lonner and more slender than the first
pair; the exopodite reaches only to the extremity of the meros; the carpos is long,
excavate at the tip, and the propodos, which resenildes that of the first pair but is more
slender, is also attached to it by its lower angle.
The three posterior pairs are long and nearly imarnied, except for a few spinules on
the meros and propodos ; the dactyla are simple ; eacii liud) bears a well-developed
exopodite.
The pleopoda have strongly developed foliaceous plates, and each bears a stylamblys.
The sixth pair have the basal joint short, and ])rodueed on the outer margin into a tootli ;
the foliaceous plates are subcqual in length, and the outer is crossed by a well-defined
diaeresis.
I'll VLI.OBRAXCIILATE CKlISTACEA-ArACKUHA. 1 !9
The following- are the dimenHiou.s of a mature female: — •
iiiiii.
licnjitli of entire niiimal ;}<)
,, cara[)ac(' <)
,, rostiuin X
„ pleon o>)
,, second segment ol' pleon ."■>
" sixth „ „ -,
» telson (j
Ilahitat. Preshwater streams apparently throughout the Colony. T have it from the
neighlKJurhood of Dunedin, Christchureh, Nelson, and Greymouth, also from the Bull(>r
Hiver in the South Island; and from the Waikato and the neighbourhood of Auckland
in the North Island.
Distribution. Confined to New Zealand.
This species is distinct from the Australian X. compressa, the most (conspicuous
difference being in the number of spines on the rostrum. AVhile X. CHrL'irostri.s shows
only slight variation in this respect, the Australian form is rather variable. Ortmann
(Proc. Acad. Philadelphia, 189I<, p. iOO) states that it has from 20 to 24 teeth on
the upper, and from 2 to i on the lower margin. Put the following figures giv(> tlie
dentition of a number of specimens from different iocalities, the first figure representing
the number of teeth on the upper, and the second the number on the lower margin : —
From Victoria. — (1) 'K> — 2 ; (2) \A — '>, tliis one had the end broken.
From N. S. Wales.— ( 1 ) .'51—9 ; (2) 23—7 ; (3) 24—6; (4) 24—5 ; (fi) 20-4.
From Norfolk Island.— (1) 31—5 ; (2) 27—9; (3) 34—7; (4) 25— G : (:>) 22—7.
From Norfolk Island.— (1) 22—3; (2) 21—2 : (3) 17—3.
The last three belonged to a form smaller than those previously received from Norfolk
Island, and having a shorter rostrum. Pi-oni the above it appears that the dentition of
X. covipressa may be expressed as follows: — Ujiper margin with 17-31' teeth; lower
Avith 2-9.
It is interesting to notice that while one slightly variable species, .1'. compressa, should
range from Australia (including Norfolk Island) to -lapan, a quite distinct one, uhich is
also distinct from the American X. ehniijala, occurs in New" Zealand.
Pamily PAL.^MONID/E.
Genus 1. Acanthephyra, A. Milne-Edwards, 1881.
1. ACANTnEPHTEA SIC'A, Bate.
1888. Bate, 'Clialk'nger' Macrura, p. 7;i9, pi. 125. fij;. 1.
JIuhitat. Off Cape Turuagaiii, 1100 fathoms ; off ]:]:ist Cape, 700 fathoms; and off the
Kermadec Islands at 520 and 030 fathoms.
SECOND SERIES. — ZOOLOGY, VOL. VIIT. 05
,150 ^IH. G. M. THOMSO.V OX THE NEW ZEALAND
Didrihiilion. This species occurs both in the Atlantic aud Pacific Oceans, aud from
Japan in the nortli to New Zealand in the souih, at deptlis varying from 200 to 2(J7')
fathoms.
2. ACAXTHEPHYRA liRACHYTKLSONIS, Bate.
1888. Bate, 'Challenger' Macrura, p. 753, pi. 126. fig. 7".
Sabilnt. Off the Kermadec Islands at depths varying from 200 to 030 fathoms.
Distribution. Seas of Japan, Philippines, Bauda, and the Falkland Islands at depths of
200-2010 fathoms.
Genus 2. Pal^mox, Eabricius, 1798.
1. Pal.emon AFFiNis, M.-Edvvards.
1837. Pahemon affiiiis, M.-Edvv. Hist. Nat. Crust, ii. p. 391.
1852. Palamon offinis, Daua, U.S. Explor. Exjied., Crust, p. 581, pi. 38. tig. 5.
1888. Pahemon tiffinis, Bate, ' Challenger' Macrura, p. 782, pi. 128. tig. 5.
1837. Pahemon Quoiunns, M.-Edw. Hist. Nat. Crust, ii. p. 393.
1843. Palienion Qmianm, White, Dieffenb. A^oy. N. Z. ii. p. 208.
1876. Leander affinis, Miers, Cat. N.Z. Crust, p. 85.
Bate states (7. c. p. 782) that this species is only distinguished from P. squilla, Fabr.,
of Europe, l)y having- the apex of the rostrum bifid, and four teeth instead of three
on the ixnder margin. But, as Miers points out, " the number of teeth varies in a
large series of specimens, while the bifid appearance is caused by the greater or less
approximation of the anterior tooth of the upper series to the apex of the rostrum — also
a variable character." And on this ground he iinites P. Qiioiaiiiis, M.-Edw., with
P. ajfinis, an opinion I quite agree with. I have never, howcner, found a specimen
with only six teeth on the upper margin of the rostrum, and I liave examined an
immense number from various parts of the Colony. The numljer varies from seven on
the upper and three on the lower to nine and six respectively.
This is the commonest Shrimp in New Zealand, and is the only one I liave seen
offei-ed for sale in the fish-shops in l^unedin, Christcluirch, and Wellington.
Habitat. Stewart Island, dredged, 10 fathoms; east coast of Otago in rock-pools, &c.,
commonly between tide-marks ; Akaroa, Lyttelton, Wellington, Bay of Islands, and
Chatham Islands. Eilhol (Miss, de I'lle Campbell, p. 131) records it from Campbell Island.
Distribution. A widespread southern species, found at the Cape of Good Hope, coasts
of Australia, Tasmania, and in the Falkland Islands.
2. Palj:jion xatatok, ]M. -Edwards.
1837. PdUeiiwn iiatator, Al.-Eihv. Hist. Nat. Crnst. ii. p. 393.
1888. Pahemon ualalor, Bate, 'Challenger' A[acruva,-i>. 78 t, pi. 12S. figs. 6 & 7.
1819. Leander eiratira, Desmarest, Aiin. Soe. Entom. France, ser. 2, vii. p. 87.
1800. Leander natator, Stinipson, Proc. Acad. Nat. Se. Philad. p. 109.
1876. Leander natator, Miers, Cat. N.Z. Crust, p. 8(5.
Tliis species is included by Miers among the New Zealand Crust;ic(!:i on the authority
PJl VI.r-ORRA^•CllIATE CHUSTACEA-MACRUKA. 451
of specimens so labelled iu the British Museum. T have not met with it, nor does it
oceur, so far as I am aware, in any of the local museums in the Colony, hut, beiui;-
a pelagic species of very wide distribution, it is probably quite a correct reference. At
the same time, Tilhol (^liss. de Tile Campbell, p. 1.31) states that Huttou has collected
it iu the South Islaiul.
Genus 3. Bithynis, Philippi, ISGO.
1. Bithynis oknatus, Olivier.
1812. Paliemoii ornutns, Olivier, Eiiryci. viii. p. (UiO.
1887. Vahamuii nntutus, M.-Edvv. Hist. Nat. (.'rust. ii. p. .'59(i.
1876. PaUeiiioii umutus, Aliers, Cat. N.Z. Crust, p. 87.
This is another species of wide distril)ution in the seas of the .southern tropical regions,
and is introduced into the list of New Zealand Crustacea on the authority of Heller, wlio
obtained it at Auckland. All Heller's identilications of New Zealand Crustacea are,
however, doubtful, as the ' Novara ' collections appear to have got mixed up.
I have not heard of its being found at any other part of the Colony, nor does it seem
to occur in the museum collections as a local species.
Genus J;. Bit vciiycaui'I's, Bate, 188S.
1. Bkachycarpus Aidouini, Bate.
1888. Bate, ' Challeuger ' Macrura, p. 7'JS, pi. V2'.). fig. j.
This species is fully described by IJate from a single female specimen, but it is
apparently common in shallow waters round the coasts. The number of teeth in the
rostrum varies from eight to nine in the upper, and from four to six in the lower margin.
When alive, the body of the animal is nearly transparent, the hepatic region being green.
On placing the specimens m alcohol the green turns to a In-ight red, and numerous red
star-like spots also appear on the bases of the limbs, the colours dying out after a day
or two.
Habitat. Cook Strait, 10 fathoms (' Challengoir ' Exped.) ; Tasman and Blueskin Bays,
10 fathoms, trawled.
Bhtrlhntion. Coiitiued to New Zealand.
Family NE.MATOCARCINID.E.
Genus 1. Nematoo'ARCIXUS, A. Milne-Edwards, 188 1-.
1. Nematocahciniis undulatipes, Bate.
1888. Bate, ' Clialleuger ' Macrura, p. 801, i)l. MO.
Ihdjilat. North of the Kermadec Islands, 600 fathoms.
l)i^lf(but(un. rhili2)pines and Biinda Islands.
<tr)2 MYt. G. M. THOMSON OX THE NE"\V ZEALAND
2. Nkmatocarcinis gracilis, Bate.
1888. Hate, ' Clialk'iigcr ' Macrura, p. 815, pi. l;52. tig. 8.
Habitat. North of the Kermadcc Islands, 600 fathoms.
Disti-il)iitloii. Fiji Islands, 610 fathoms.
J}. NeMATOCAKCINLS SERKATU.S, Bate.
1888. Bate, ' Challenger' Macrura, p. 819, pi. 1.32. fig. 11.
Habitat. Off East Cape, 700 fathoms.
This species is described from a solitary specimen. Bate says {I.e. p. S20), " associated
with Acanthephyra imrimrea " ; hut this is evidently a slip instead of A. sica, as the
former species is found only in the Atlantic Ocean, Avhereas A. sica was traAvled at
tlie above localitj^
1- Nematocarcinus hiatus, Bate.
1888. Bate, ' Challenger' Macrura, p. 821, pi. 132. tig. 12.
This species Mas constituted on a fragmentary specimen.
Habitat. Olf East Cape, 700 fathoms.
Family STYLODACTYLID.E.
Genus 1. Stylodactylus, A. Milne-Edwards, 18S3.
1. Stylodactylus discissipes, Bate.
1888. Bate, 'Challenger' Macrura, p. 851, pi. 138. fig. 1.
Habitat. North of the Kermadec Islands, 600 fathoms. Described from two
specimens.
2. SXYLODACTYLUS ORIENTALIS, Bate.
1888. Bate, 'Challenger' Maerura, p. 85-1, pi. 138. fig. 2.
Habitat. North of the Kermadec Islands, 600 fathoms. Described from a solitary
specimen.
It is worth recording in this paper that, in 1894,. the Otago Acclimatisation Society
liberated specimens of Peno'us canalicntatus, Oliv. (obtained from Sydney), in Dnnedin
Harbour. I made a protest at the time, using as my strongest argument to a utilitarian
and unscientific body the waste of money involved in attempting to introduce a troj)ical
species into our cold waters, for the south-east coast of the South Island of New Zealand,
as far north as Banks Peninsula, is washed by a cold southerly current. But some
fifty specimens were brought over and set free.
Nothing more has been seen of these Prawns, and it is very improbable that any of
them have survived. But should this species hereafter be found in New Zealand waters,
the fact of tiieir having been introduced will have to be taken into account.
PHTLLOBRAXCHIATE CRUSTACEA-MACRURA.
453-
EXPLANATIOX OF THE PLA.TES.
i'^ig. 1. Float of carapace, from above
2. Base ot first antenna.
3. Third niaxillipcd.
Plate 27.
Figs. 1-5. Pontophilm aiistralis.
Fig. -i. First pcreiopod.
5. Telsou and tail-fan.
Figs. G-12. Alpheus socialis.
Fig. 0. Part of large male specimen.
7. Front of carapace of same.
■S. Front of carapace of a specimen from
Port Phillip (Australia).
Fig. 9. Extremity of first pcreiopod of small
male (from inside).
10. Extremity of third pcreiopod.
11. Telson and tail-fan.
12. Outer plate of last plcopod.
Plate 28.
Figs. 1-.2. Betceus (equimanus.
Fig. 1. Carapace, viewed from above. | Fig. 2. Telson.
Figs. 3-12. Alojje jici/jjalis.
Fig. .3. Portion of sternum. Fig. 8. Second maxilla.
4. Telson and sixth pair of pleopods. 9. First maxillipcd.
First antenna.
Mandible.
First maxilla.
Fig. 13.
U.
Ophthalmopod.
First antenna.
10. Second maxillipcd.
11. Second perciopod.
12. Dactylos of third pcreiopod.
Figs. 13-lG. Hijjpo/j/ie bijidirostr'is.
Fig. 1."). Third maxillipcd.
1(). First pcreiopod.
Fig. 2. Side view.
3. Telson.
4. First antenna.
5. Second antenna.
<;. Mandible.
7 . First maxilla.
Plate 29.
Fig. 1. Nniitlrar'is Stetcartl.
Figs. 2-13. Xi/i/iocaris curvirostris.
Fig. 8. Second maxilla.
9. First maxillipcd.
10. Second maxillipcd.
11. Third maxillipcd.
12. First pcreiopod.
13. Second pcreiopod.
SKCOND SERIES. — ZOOLOGY, VOL. Vlll.
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AUG 2%. ia03
2nd Ser. ZOOLOGY.]
[VOL. VIII. PART 12.
THE
TRANSACTIONS
OF
THE LINNEAN SOCIETY OE LONDON.
ON THE MORPHOLOGY OF THE CEREBRAL COMMISSURES
IN THE VERTEBRATA, WITH SPECIAL REFERENCE
TO AN ABERRANT COMMISSURE FOUND IN THE
EOREBRAIN OF CERTAIN REPTILES.
BY
G. ELLIOT SMITH, M.A., M.D.,
Fellow of St. John's College, Cambridse ; Professor of Anatomy,
Egyptian Government School of Medicine, Cairo.
[Communicated by Prof. G. B. Howes, D.Sc, LL.D., F.K.S., Sec. Liiiu. Soc.)
•^^"'"ft;,.;//^.
LONDON:
PRINTED FOR THE LINNEAN SOCIETY
BY TAVLOU AND FRANCIS, RED LION COURT. FLEET STREET.
SOLD AT THE SOCIETY'S APARTMENTS, BUKMNGTON-HOTJSE. PICCADILLY, W.,
AND BY LONGMANS, GREEN, AND CO., PATERNOSTER-ROW.
Juhj 1903.
AUG 2L 1903 r .r- -,
XII. On the Morphologn of the Cerebral Commissures in the Vertehrata, with Special
Reference to an Aberrant Commissure found in the Forebrain of certain Reptiles.
By G. Elliot Smith, M.A., JI.D., Fellow of St. John's College, Cambridge ;
Professor of Anatomy, Egyptian Government School of Medicine, Cairo. {Com-
municated by Prof. G. B. Howes, B.Sc, LL.D.. F.B.S., Sec. Linn. Soc)
(With 36 Illustmtions in the Text.)
Read nth. June, 1902.
In spite of the innumerable memoirs which have been published on the homologies of
the cerebral commissures in the Sauropsida and Ichthyopsida, the essential problem has
practically remained untouched, and the subject still awaits a full discussion. I do not
propose at this time to attempt the complete solution of this matter, but merely wish to
point out the natm-e of the question at issue, and to bring forward some facts which seem
to suggest in which direction we must seek for the full explanation.
To make clear the exact scope of this communication, I may be permitted to explain
the circumstances under which it was written.
About live years ago the late Mr. Martin Woodward gave me the head of an adult
Sphenodon (preserved in alcohol), from which I was able to obtain a brain in a sufficiently
good state of preservation to l)e cut iu a series of coronal sections. When these had been
stained with lithium carmine, I was greatly surprised to find a band of commissural
fibres crossing the epithelial roof of the third ventricle midway between the " dorsal "
(hippocampal) and "superior" (habenular) commissures, such as I had hitherto seen
oulv in the Lacertilia. [In the following account I shall call this commissiu'c b^s' the
name ^' aberrans," because all the other terms in current use are highly objectionable.]
Shortly afterwards I was enabled to confirm this fact, and study the relations of the
coinmissura aberrans in a series of sagittal sections, wliich were cut from a specimen
kindly given to me by Dr. Hans Gadow of Cambridge.
Although the only contributions * to our knowledge of the l)rain of Sphenodon w^hieh
had been published at that time contained no reference to the presence of the pecuhar
commissure, I withheld the publication of ray incomplete studies, in the hope of
obtaining material which might be examined after treatment by the staining methods of
Weigert and Golgi.
In 1899 Professor Arthur Deudy, for the first time, placed on record the existence ol'
this peculiar commissural tract in Sphenodon f. Dendy's memoir was written in Xew
* Bald-nTii Spencer, " On the Presence and Structure of the Pineal Eye in Lacertilia," Quart. Journ. Micr. Sci.,
vol. xxvii. p. 165; G. Osawa, "Beitriige zur Anatomie der Hattevia puiictata." Archiv mikr. Anat., Bd. li. p. 587;
R. Wiedersheim, Grundriss d. Vergl. Anatomie. 1893. p. 2til.
t A. Deudj-, '• The Parietal Eye and Adjacent Organs in HjAenodon," Quart. Journ. Micr. Sci., vol. xlii. pp. 121
and 143.
SECOND SERIES. — ZOOLOGY, VOL. VIII. 67
456 PKOr. G. ELLIOT SMITH ON THE MOEPHOLOGT OF THE
Zealand, where it was impossible to consult all the recent literature relating to this
region of the brain, and hence he used the term " commissura fornicis," which was the
original name applied to this tract of fibres by Eabl-Riickhard. I might add, in passing,
that this original and now discarded term is unquestionably the least objectionable of the
names hitherto ajiplied to this conmiissure.
In addition to devoting himself to the study of the early developmental history of the
Tuatara, Professor Deudy also collected a valuable series of embiyos, which were sent
to Professor Howes in order that the histogenesis of the skeleton might be fully
investigated. After a number of beautiful series of coronal and sagittal sections of
embryos of various ages had been made by Mr. Swinnerton for this work *, Professor
Howes kindly suggested to Professor Dendy that I should study the developmental
history of the nervous system in the sjiecimens iu whicli he was investigating the skeletal
structures. Professor Dendy kindly gave his consent to this suggestion, and in addition
offered to obtain specimens of properly j)reserved adult brains. During the year 1899 I
devoted most of my vacations to the examination of this valuable material in Professor
HoAves's laboratory at the Royal College of Science, where these notes were written. As
I left England shortly afterward, I have had no chance of seeing the material again, and
as there seems little prospect of doing so in the immediate future, I am publishing in this
incomplete form the most interesting observations of which I had made a record. In
order to enhance the value of this, which being so imperfect is in itself slight, I have
introduced into the account a series of comparative notes, which 1 believe will shed some
light on one of the darkest and least-understood corners of the brain, and help towards
systematizing our knowledge for the future.
The roof of the forebrain of Sphenodon presents an exceedingly irregular form (tig. 1).
It consists of a thin layer of epithelium which, attached in front to the dorsal extremity
of the lamina terminalis, ends posteriorly at the situation of the habenular (" superior '")
commissure by becoming continiious with the thicker epithelium of the parietal stalk.
Its irregular course between these two points is represented on a greatly-enlarged scale
in figvire 2. From the upper end of the lamina terminalis (a) the epithelial layer bends
downward in the third ventricle and, after being thrown into many folds, ascends again
and becomes continuous with a long, narrow, irregular evagination of the roof, which is
commonly known as the paraphijsls. The irregular fold (/3), which is invaginated into
the ventricle between the lamina terminalis and the paraphysis, is pi'olonged laterally on
each side into a complicated fold which extends through the foramen of Monro into the
lateral ventricle. As these lateral processes are kno"mi as the choroid plexuses, the fold
/3 may be distinguished as the lamina chorioklea (generally spelt "choroidea" in English
works). The paraphysis is separated from a much longer and broader outgrowth of the
roof (y) by a simple transverse lamina, which is commonly called the " velum." The
irregular fold y is known as the " dorsal sac."
The varied nomenclature which is applied to these structures has been discussed in
* Vuh G. B. Howe.s and H. H. iSwiniiorton, '■ On the Dovelojjinent of the Skeleton ot the Tuatara," Trans. Zool.
>Soc., vol. svi. (1001) part 1, for an account of the material.
\
CEREBRAL COMMISSURES IN THE VERTEBRATA.
457
Sorenson's critical review * of the enormous mass of literature which is devoted to this
Fig. 1.
region of the brain
pUcIV.
hyp.
A mesial sagittal section of tlie brain of an embryo [Deudy's stage S] of Hi>lur,wdon puitcbttion. Magnified almost
10 diameters. Outline drawn with camera lucida.
The cerebral hemisphere and the anterior quadrigemiual body, which, of course, are not seen in a mesial section,
are also represented ; for explanation of abbreviations see page 500.
,p.s.
^"^'m^^i^^^^
Semi-schematic representation of the Roof of the Forebraiu in Sphemdon. Considerably enlarged from figure 1.
Camera lucida drawing. The commissura aherrans is immediately in front of the letter c.
The structures just enumerated are foimd in almost all Vertebrates. But in the brain
of Spheiwdon there is another structure in close relationshij) with the roof of the third
* A. D. Sorenson, " Comparative Study of the Epiphysis and Roof of the Diencephalon," Journ. Comp. Neurology,
vol. iv. (April 1894) pp. 12-72.
67*
458 PROF. 6. ELLIOT SMITH ON THE MORPHOLOGY OF THE
ventricle. It is the rouuded buudle of nerve-fibres forming the aberrant commissure
lying' upon the epithelivim of the velum (c).
If a series of coronal sections of the brain of Sphenodoii be examined, it will be found
that 01 the mesial structures only the lamina terminalis and that i:)ortion of the epithelial
roof -nhich lies in front of the commissiira aherrans stand in direct connection with the
cerebral hemis2)here. That portion of the roof -which is placed on the caudal side of the
commissiira aherrans, and which forms the dorsal sac (the so-called " Zii'belpolster " of
Edinger), is connected laterally with the dorsal lips of the optic thalami. In other words,
it forms the roof of the third ventricle, and is not directly connected with any part of the
cereb)'al hemisphere. This simple relationship of the epithelial dorsal sac to the optic
thalaini is shown in the accompanying diagram (fig. 3).
kM
A simple scheme to illustrate the relationships bf the epithelial roof (•' dorsal sac '") behind the
commissura aberriDis.
But the lateral connections of the forward continuation of this epithelial roof undergo
sudden and most significant changes immediately in front — i.e., on the cephalic side —
of the^ commissura aherrans. This fact (which, so far as I am aware, has never hitherto
been definitely formulated) might be diff'erently stated by saying that the commissura
ABEKRANS INDICATES IN THE MESIAL PLANE THE CAUDAL LIMIT OF THE ATTACHMENT OP
THE CEREBRAL HEMISPHERE to the rest of the neural tube.
In endeavoiiring to arrive at some explanation of the meaning of tliis ambiguous
cojmmissure, as I shall attempt to do later on in this memoir, its limbic situation will be
fc»und to be perhaps its most significant feature.
, Before investigating the relationships of the cephalic portion of the roof of the median
ventricle, it is expedient to study the relations of the lamina terminalis and the cerebral
liemisphere.
The examination of a series of horizontal sections through the brain of any Reptile or
Amphibian will reveal the lamina terminalis as a narrow band connecting two large
vertical masses of grey substance, which I propose to call the ^' paraterminal bodies"
Each corpus paraterminale forms the inner wall of the anterior portion of the lateral
ventricle, and extends forward from the lamina terminalis as far as the olfactory bulb or,
in those cases in which the bulb is pedunculated, to the olfactory peduncle. These two
Varge masses of neural substance are linked together by the lamina terminalis, which
CEREBRAL COMMISSURES IN THE VERTEBEATA.
459
extends across the mesial plane as a transverse band of an ejjithelial or nenroglial natnre
which is devoid of nerve-cells. In most cases, however, the commissures which grow
across the mesial plane in contact with the lamina terminalis cany with them some grey
substance (derived from the corpora paraterminalia) which forms the "commissure-bed" —
a secondary thickeninsr of tlie lamina terminalis.
Fig. 4.
A scheme to indicate the fundamental relationsliips of ecrtain portions of the cerebral hemispheres to the lamina
terminalis and optic thalanii ; being the plan of an ideal horizontal section.
If a coronal section be made througli the cerebral hemisphere of Sphenocloa, imme-
diately in front of the lamina terminalis, the relationship of the corpus ixiraterminale to
the suri'ounding areas will be more readily appreciated.
A coronal section through the two cerebral hemispheres of ISphfuodon (Dendy's foetal stage R) immediately in front
of the lamina terminalis.
Each paraterminal body will then appear as the thick plump mesial Avail of the
hemisphere, which is continuous Aentrally around the ventricle with the corpus striatum
on the lateral aspect of the ventricle.
The upper edge of the thick paratermiual body is fused with a relatively thin portion
of the wall of the ventricle, in Avhich the nerve-cells are collected into an extremely
460 PEOF. G. ELLIOT SMITH ON THE MORPHOLOGY OF THE
densely imcked and regular layer, instead of being scattered irregularly in the thickness
of the wall, as is the case in the paraterminal body. In the subsequent discussion it will
be seen that there is little reason for hesitation in homologising the mesial portion of this
tbin roof the hemisphere, /. c. the part adjoining the paraterminal body, with the hippo-
campus or cornu Ammonis of the Mammalian brain. Hence I may be permitted in
anticipation to call this region by the name "hippocampus." In the roof of the
hemisi^here the hippocampal structure passes into continuity with another cortical area
which, for reasons which will appear later, I shall call " neopallium." Upon the mesial
surface of the hemisphere a shallow furrow, which I shall call " suleus limitans"
indicates the dorsal limit of the paraterminal body and the line of junction of the latter
with the hippocampus.
Tlie mesial surface of the cerebral hemisphere in front of the lamina terminalis may
now be mapped out into areas corresponding to the hippocampus, paraterminal body, and
a small and ill-defined mesial portion of the corpus striatum. The accompanying sketch
Fig. C.
b.o. U.
The right cerebral hemisphere oi Uphcnodon. — Semi-diagrammatic representation of its mesial aspect. Enlarged.
of the mesial surface of the cerebral hemisphere in Sphenoclon serves to indicate in a
rough manner the approximate distribution of these several areas.
In the description of the mesial structures, it has already been noted that towards its
upper part the lamina terminalis apj^ears to expand into a fusiform enlargement which
serves as a matrix for the dorsal and ventral commissures. If a coronal section through
the brain be made in this situation, it will be seen that the two cerebral hemispheres are
joined the one to the other by this matrix, or thickened lamina terminahs, which forms
a bridge between the two hemisjiheres.
A comparison of the accompanying figure 7 (which passes transversely through the
commissures and lamina terminalis) with the previous illustration, will at once reveal the
identity of most of the parts of the brain exposed. Thus the relations of the " hippo-
campus " and " neopallium " to the paraterminal body and the corpus striatum,
respectively, are unchanged. In the thickened lamina terminalis will be noted in
addition the two cerebral commissures. The con/mlssura dorsaUs is a great U - shaped
CEEEBRAL COMMISSURES IN THE VEETEBRATA.
461
strand, each limb of wliicli ascends through the paratcrminal body and ultimately enters
the hippocampus : in tlic hippocampus its fibres are found scattering in the interval
between the regular column of cells and the lining epithelium of the lateral ventricle.
The eommisstira ventralis, on the other hand, presents the aiij^earance of a transverse
horizontal baud extending laterally into the corpora striata.
But in addition to all these structures the section passes through the recossus oj)(icus,
and hence opens up the third ventricle and the structures in relationship with its walls.
The walls of this portion of the third ventricle consist of the most cephalic portion of the
thalamic region. It may be noted here that in the region of the lamina terminalis
the anterior extremity of the thalamic region fuses not only with the corpus striatum,
but also with the corpus paraterminale.
p<3r<3.T
cv.
Coronal sectiou. — Brain of Icttal Sjilx'awhjn (Dendy's stage R).
A section (tig. 8) which passes immediately behind the lamina terminalis presents
many interesting points of difference when contrasted with the last figure.
As in the last figure, the relations of " hippocampus " and the " neopallium " to the
coi'pus paraterminale and corpus striatum, respectively, are unchanged. Bat the
ventral relationships of the paraterminal bodies have undergone a notable change. The
third ventricle appears to have extended dorsally, and to have given off a lateral
diverticulum on each side. This lateral diverticulum joins the lateral ventricle, and
thus (in this section) completely separates the paraterminal body from the optic thalamus.
This dorsal portion of the third ventricle, or centricnlKS communis, may be distino-uished
not inappropriately as the aula, a name introduced by Burt Wilder. The aula com-
municates on each side with the lateral ventricle by means of the foramen of Monro.
The roof of the aula consists of an irregularly folded jilate of cubical epithelium, and it
therefore diflPers very much from the relatively thick neuroglial plate which forms tlie
lamina terminalis. This roof, or tela, is attached on each side to the ventral lips of the
462
PEOr. G. ELLIOT SMITH OX THE MORPHOLOGY OF THE
corpora paraterminalia. The typical relationship existing between the thalamus and
the corpus striatum in this section calls for no description.
ne.
A coronal section through the brain of SpJunodon passing iuimediately Ijehind the lamina terminalis
(Dendv's foetal stage E).
If another coronal section be made midway between the upper extremity of the lamina
terminalis and the commissura aherrans, the epithelial roof of the cmla Avill be found to
present a very differetst appearance to the simple fold which was shown in the last
figure.
Upon each side an elongated and irregular fold of the epithelial roof is bulged through
the foramen of Monro (or porta, as Wilder haj^pily calls it) into the lateral ventricle,
where it forms the lateral choroid plexus, as the accompanying figure 9 shows. A
comparison of this illustration with the preceding one at once shows that this choroid
Fig. 9.
Portion of a coronal section through the brain of S/Aenodou (Dend.y"s stage E), midway between the upper
extremity of the lamina terminalis and the cuiiimissura aherrans.
plexus is not a portion of the wall of the hemisphere which has retained its embryonic
character, but a part of the roof of the neural tulje, which, as His has 2)ointed out, never
becomes anything else than neuroglia or simple epithelium.
The careful study of the disposition of the lateral chorioid plexus in almost any Vertebrate,
or better in a series of Vertebrates, proves beyond a doubt that the choroid plexus
CEREBRAL COMMISSURES IX THE VERTEBEATA.
463
{at least in the region of the foramen of Monro) never forms part of the cerebral lieiui-
spbere, but is merely an appendage of the roof of the primitive forebrain. The mesial
portion of the epithelial plate from which the choroid jilexuses (lateral) appear to spring
may be distinguished as the lamina chorioidoa. Tlie exact situation of this lamina is well
shown in figure 2 (/3). From that figure it is seen that the lamina chorioidea is separated
from the commissura aherrans h\ the paraphysis, and this explains the presence of the
paraphysis in the coronal section which has just been imder consideration. It consists
of a very irregular median diverticulum of the lamina chorioidea, which extends upwards
between the two cerebral hemispheres.
A comparison of the last three figures will siiow that the paraterminal Iwdy diminishes
in bulk as it is followed in the caudal direction. At tlie level of the coinmisura aherrans
(fig. 10) it has assumed a triangular shape, and has estal)lished relations which are
Portion of a coroaal section through the brain o{ Splnuod'ni ( Deiidy's stage It): passing through the
coiumisSKra aherrans.
extremely peculiar and perplexing. Thus tlie mesial l)orders of the bodies in question
have extended mesially, and fused the one with tlie other upon the roof of the third
ventricle. In addition, the ventral surface of the paraterminal body is fused to the dorsal
surface of the optic thalamus, producing on the whole an appearance which (especially
to the student of the Mammalian brain) is in the highest degree bizarre and astonishing.
The ventrolateral angle of the paraterminal body is prolonged downward, and tapers to
become continuous with a plain epithelial lamina («), which completes the gap in the
mesial wall of the hemisphere which would otherwise exist in the interval between the
corpus striatum and the paraterminal body.
The commissura aherrans consists of a strand of nerce-fihres which crosses ahove the roof
of the thii-d ventricle in the hridije of grey matter formed by the fusion of the caudal
extremities of the tioo corpora paraterminalia : after passing through the paraterminal
SECOND SERIES.— ZOOLOGl', VOL. VIII. 68
464
PEOF. G. ELLIOT SMITH ON THE MOBPHOLOGT OF THE
body the fibres end in the hippocampus in a manner exactly analogous to that tchich
characterizes the cornmissura dorsalis in the most cephalic region of the hijjpocampm.
In 1S99 I called attention * for the first time to the fact that in the brain of Ornitho-
rhynchus the roof {tela) of the forebrain extends forward to be attached to the upper
extremity of the lamina terminalis in front of the dorsal commissure. As a result of this
an-angement, a small diverticulum — recessus siqjerior — of the third ventricle is formed
upon the upper surface of the cornmissura dorsalis. The significance of this arrange-
ment may possibly be not very great, but the relations of the parts which stirround this
little recess are most instructive when compared with those which are found in certain
Reptiles, and especially the Saiirians.
The surface of the brain of Ch-nithorhynchus, exposed by a mesial sagittal section. ilagniSed '2 diameters.
I have introduced here a figure from the memoir to which I have jost referred, in
order to make the general relations of this peculiar recessus superior clear, and to
explain the appearances presented by sections which I intend to discuss. For a full
account of the figure the reader is referred to the original description.
If a section be made through the brain of Ornithorhynchus (in the plane x-y), the cut
surface will present an appearance which is reproduced upon a somewhat lai'ger scale in
the next illustration (fig. 12). The general features of this section will be familiar to
those who are acquainted with the writings of Alexander Hill (Philosophical Transactions,
1893), Symington (Journal of Anatomy and Physiology, 1S92), or of the present writer
(Journal of Anatomy and Physiology, 1896). I need merely call attention to the two
large commissural bands — the cornmissura dorsalis, connecting the typical hippocampal
formations which are placed upon its upper surface: and the coniniissura centralis,
deriving its fibres from the whole of the other cortical areas of the brain.
* '•Further Observations ou the Anatomv of the Brain in the ilonotremata,"' Journal of Anatomy and
Physiology, toL xxxiii. p. 315.
CEREBRAL COMMISSURES IX THE YERTEBRATA.
465
In this figure, however, there are certain features which have not hitherto been
represented. The most imjiortant of these is the little epithelial pocket which constitutes
the recessus superior. It is placed upon the upper surface of the dorsal commissure
between the mesial extremities of the fasciie deutatae. It is unnecessary for me to
explain how the fascia dentata reaches this position, which to the human anatomist is so
bizarre; because in several earlier communications I have explained this peculiar
phenomenon (more particidarly in Journal of Anatomy and Physiology, vol. xxxii. —
especially figure 3, page 30, which represents the analogous arrangement in
Fhascolarctos).
■ ' Fis. 12.
/■<^-
cat.
CM pans.. L.p.
Coronal section of the cerebral hemispheres of Omitliorliyadn's. x about 21. The pointer from the letters c.d.
actually ends in the alveus hippocampi, from which the cominissiira dorsalis is derived.
If the recessus superior be examined with a higher degree of magnification, its exact
relations will be more readily appreciated. But certain features are more clearly
demonstrated in a section Avhich passes immediately in front of the dorsal commissui-e
and, while missing the crossing fibres, passes through the grey matrix of this commissure.
The next figure represents the superior recess from such a section (fig. 13).
In tliis section a solid mass of grey matter may be seen resting upon the dorsal aspect
of the ventral commissitre in the mesial plane, and extending upward on each side to
fuse wdth the ventral margin of the hippocampal formation and its specialized fringe of
fascia dentata. This mass of grey substance represents the posterior extremities of the
corpora paraterminalia Avhicli have fused the one with the other in front of the lamina
terminalis. The thickening of the lamina terminalis, which is thus produced at the
expense of the paraterminal bodies, corresponds to the bridge of grey substance whicli
forms the matrix for the two cerebral commissures.
In an earlier contribution I called this the " commissure-bed." Into this " commissure-
bed " the fibres coming from the hippocampus — in other w^ords the fornix — pass, and
68*
466
PEOr. G. ELLIOT SMITH OX THE MOEPHOLOGT OF THE
many of them assume a longitudinal direction and associate themselves either with the
post-commissural group (which forms the columnar for nicis) or the precommissural group
of fihres \vkle Journal of Anatomy and Physiology, vol. xxxii. pp. 36 et seqJ].
Fig. 13.
fta.sc.
Part of a coronal section through the cerehral hemispheres of Ornithorhynclnis, to show the relation of the
parncommissural hod}- to the hippocampus.
The recesstis superior in this section rests upon the upper surface of the " commissure-
bed," and presents an oblong shape. It is lined with simple cubical epithelium: its roof
consists of a simple epithelial layer, and its characters resemble those of the rest of the'
roof {tela) of the third ventricle of which it forms the cephalic extremity ; it is
invaginated into the cavity of the superior recess to form the anterior extremity of the
mesial choroid plexus — the " dlaplexus " of Biirt Wilder : the lateral walls of the recess
are thickened by a small quantity of grey substance derived from the paraterminal bodies,
and in this grey substance longitudinal libres belonging to a peculiar set of fornix-tibres
which are found at the ventral margin of the fascia deutata are found. These fibres
belong, in all probability exclusively, to the group which I have called "fasciculus
marginaUs" (Journ. of Anat. and Phys., vol. xxxii. p. 38). In the next figiu*e these
features are shown upon a much larger scale (fig. 14).
The recessus superior exists and presents similar features in the brain of Echidna, and
therefore it will be legitimate to institute comparisons between the adult Oriiithorhynchus
and certain foetal stages of Echidna, Avith a view to the explanation of certain earlier
fcEial stages of Ornithorhyiichus. Such comparative studies are necessary, because it has
been impossible to obtain a comjDlete series of foetal stages of Platypus.
If sections be made through the brain of a foetal Echidna (at the age when the spines
are just making their appearance in the skin), it will be found that a section passing
through the two cerebral commissures will present features (fig. 15) which, on the
whole, so closely resemble those of the adult Ornithorhynchus just considered, that it is
CEREBRAL COMMISSURES IX THE VERTEBRATA.
467
unnecessary to describe them in detail. But there arc certain features in the region of
the recessus superior which reproduce tlie condition found in the Sauropsida more nearly
than does that of the adult Monotreme : in addition, the hippocampal formation is very
Kk. 14.
recs.
C.a/r-m l—
CM
Greatly enlarged representation of jiart of a coronal section of the brain of OrjiitJiorhijnchus, to show the recessus
superior and the adjoining structtires. The letters utarr/. refer to the fasciculus marginalis, labelled /««. in the
other figures. The adjoining diagram is an enlarged part of fig. 11, to explain the jilane (x-if) in which the
section was cut.
mucli simpler than it is in the adult, and its exact relations to the j'araterminal body
less ambiguous. Such a section is roughly represented in figure 15, and the regions
Fig. 1.5.
rec s
fd.
c.d vM. c.v -^ p<a.r<a..
A scheme of a coronal section of the brain of a very young h'cJiiiliKi, greatly enlarged.
surrounding the recessus superior have been reproduced again upon a larger scale in
another figure (16). The former illustration needs no further description, and I may
turn at once to the consideration of figure 16.
In many minor points the condition in the foetal Echidna ditfers from that found in
the adult Ornithorhy minis (and Ecliidnu). Instead of the lateral waUs of the superior
recess being slightly thickened witli grey matte)- derived from the paraterminal body,
468
PEOF. Q. ELLIOT SMITH ON THE MOEPHOLOGY OF THE
the little recess will be fovind to be walled on each side by the whole mass of the para-
terminal body, a large part of which (05) intervenes between the recessus superior and
the ventral margin of the hippocampal formation. [I nse the term " hippocampal
formation" here in preference to "fascia dentata," because the latter body is obviously
in the process of formation as a specialization of the lower extremity of the hippo-
campus.]
At the dorso-mesial angle of the hemisphere the neopallium will be found in a state of
transition. The scattered nerve-cells are becoming collected into a compact column,
which is readily recognized ixpon the mesial wall (after comparison with the adult brain)
as the layer of pyramidal cells of the hippocampus. This layer of cells is already under-
going a lateral bending, but it is a significant fact that only the very slightest furrow
Fig. 10.
CM
Scheme drawn from a coronal section through the brain of a young ErJiidnu, which had been
stained with lithium carmine.
has yet made its appearance upon the mesial wall (o). It is hence clear (as Hochstetter
has shown) that the supposed furrow called " Bogenfurchc " by Schmidt and " Ammons-
furclie " by Mihalkovics, which makes its appearance at a much earlier stage of
development, cannot have anything to do with the true hippocampal fissure, which at
this late stage is only just beginning to appear. In fact the slight furrow c, Avliich is the
first representative of the hippocampal fissure, is really due to the peculiar inrolling of
the hippocampal formation which takes place pari passu with the develojjment of the
fascia dentata. The latter arises (see fig. 16) as a rapid proliferation of the ventral
extremity of the column of cells which elsew'here develops into the layer of pyramidal
cells. As these cells increase in number at a rapid rate, the surface area corresponding
to them grows much more rapidly than the deeper regions, and becomes bent up into
the peculiar pouch-like shape which the fully developed fascia dentata always presents
in transverse section in the adult brain. This rapid growth in superficial extent seems
CEREBEAL COMMISSURES IX THE ^TIRTEBRATA. 469
to determine to a larg:e extent the peculiar shape of the hippocampal formation and the
presence of the hippocampal fissure.
If it were pertinent to the subject of this communication, I might introduce here a large mass of data
in support of these statements regarding the morphogenesis of the hippocampal formation [vide "The
Fascia dentata," Anatomischer Anzeiger, 189(J], and in support of Hochstetter's contention that the
hippocampal fissure is a much later feature to make its a})pearance than the supposed " Bogenfurche/'
and that the latter has a purely post-mortem existence. 1 have, howevci, referred to this question only
to point out that the fascia dcntata makes its appeatance comparatively late in the development of the
brain *, and is derived from the hippocampal formation by a specialization of its ventral edge.
Before leaving this section I wish to call attention to a furrow {(3) which separates the
paraterminal body (a) from the hippocampal formation, and which I may call " sulcus
limitans [hipj^ocamjn]" Most writers on the Reptilian brain regard the sulcus fB as the
homologue of the Bogenfurche (8) : this drawing (fig. 16) shows how erroneous such a
contention is.
Fig. 17.
pdra..
Scheme of a section through the rccessus superior of a foetal Ormthorhijnclius.
A section thi'ough the recessiis superior of a foetal Oj'nitJiorhijuchus (fig. 17), at a much
earlier stage than the Uchidna just considered, shows the "neopallium" (ne.) joining the
hippocampus (hip.) at the dorso-mesial angle of the hemisphere just as it does in Echkhia ;
but there is no definite fascia dentata, although a slightly exuberant mass of cells [f-d.)
represents the earliest jjhase of its development. [The other features of this section do
not concern us here and have been described elsewhere. " The Brain of a FcEtal
Oniit/ior/ii/iicf/us," Quart. Journ. Micr. Sci. vol. xxxix. p. 181.]
In the light of the facts wliich this brief examination of the relations of the java-
terminal body and hippocampus has revealed, it is interesting to examine the condition
of the analogous parts of the brain in Reptiles. For this purpose I have deliberately
chosen the brain of a Saurian, not because I suspect any close phyletic relationship
between the Saurians and the Monotreinata, but because the arrangement of the homo-
logous regions in the two types of the brain is so clearly identical, that the comparison
between the two becomes an extremely simple matter. The question whether this
* The foetus under consideration had in other respects attained to almost the adult type, and yet the fascia
deutata is in the primitive condition shown in the figure.
470
PROF. G. ELLIOT SMITH ON THE MORPHOLOGY OF THE
undoubted identity of arrangement is to be attributed to a close genetic relationship,
or to an evolutionary convergence, does not come within the scope of this enquiry : it is
sufficient that the homology is so clear as to be indisputable.
Of a large number of Sauropsidan brains which I have examined, I have chosen that
of a common Australian Hydrosauriis, because it lends itself so admirably to exact
comparison with the Monotreme brain.
If a section be made through the brain of this Monitor in a plane corresponding to
that which was made through the Prototherian brains which have just been considered,
a state of affairs will be found which it is easy to interpret after comparison with the
foetal and adult Monotreme arrangement.
In this section (fig. 18) two commissural bands are found dis2)osed according to a plan
Coronal section of cerebral hemis])heres of Hiidrusiniriis.
which is analogous to that of the Monotreme. The question whether these dorsal and
ventral commissures are not merely analogous but also homologous to the corresponding
commissures in the Monotreme will l)e discussed subsequently, but I may for the present
call attention to the similarity of their disposition.
I may remark, m passing, that no writer who has treated of this question, even though
the conclusions at which he arrives are diametrically opposed to my own, has refused to
admit the fact of the correspondence in position of these commissui*al bands in the Reptile
and Monotreme. [As an example, I might refer to the writings of Osborn, ' Morpholo-
gisches Jahrbuch,' Bd. xii. p. 223, and numerous other instances might be quoted.]
Upon the upper surface of the ventral commissure, in the section under consideration,
two bulky masses of grey substance are found joined across the middle line by bridges of
grey substance. These grey masses are the posterior extremities of the paraterminal
bodies, which are relatively much larger than the corresj)onding bodies in the Mono-
tremata. In the next figure they are represented upon a larger scale. Upon the upper
surface of the paraterminal body a recessus superior is found resembling in all its features
that of the adult OrnUhorlvjnclms. Thus it presents an epithelial roof, invaginated to
form a mesial choroid plexus — " dlaplexxis " of Wilder. Its lateral walls are thickened
CEEEBEAL COMMISSUEES IX THE YERTEBEATA.
471
with grey substance derived from the paraterminal body, as in the Mouotreme ; and in
this grey mass a number of longitudinal nei've-fibres are found which constitute the
mesial elements of a larger series of fibres which are found iu that part of the para-
terminal body which clearly corresponds to the region distinguished as a in the foetal
Echidna. There can be no valid reason for refusing to regard these fibres as the
homologues of the '■'fasciculus maryinalis " of the Monotrcme. The fact that a foramen
or " intercommissural recess " is found in the " commissure-bed " in the Reptile and not
in the Monotreme, need cause no difficulty, because a sagittal section of the lamina
terminalis at once shows the unimportant nature of this feature : on the other hand, the
Fig. lii.
Portion of a coronal section through the brain of a Monitor (Hijdrosaiinis).
hip.
<a.L\'.
yoara
Scheme of lamina terminalis to show the plane of the section ,r-_i/ above.
presence of such a recess between the columns of the fornix — the vecessus triangularis of
Scbwalbe — is a well-known fact in the case of the brains of the Meta- and Eutheria.
The dorsal edge of the paraterminal body is fused with the lower extremity of a thin
portion of the wall of the hemisphere, which presents a peculiarly distinctive histological
structm'e. It is characterized by the presence of a very regular closely-packed column
of cells, which is placed somewhat nearer the ventricle than the surface of the hemi-
sphere. The component cells of this column are moderately large pyramids, the apices
of wdiich are directed toward the surface ; from the basal aspect of each cell an axis-
cylinder process passes off and contril)utes to the formation of a layer of nerve-fibres
SECOND SERIES. — ZOOLOGY, VOL. VIII. H!)
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CEREBRAL COMMISSURES IN THE VERTEBRATA.
473
suggestions utterly nugatory. In addition, the fact that Spitzka called the commissure of his " hipjjo-
campi " "corpus ca//osiun " * seems to indicate a distrust of his own suggestions.
In 1888 I'^dinger definitely homologized the medio-dorsal cortical formation in a number of Reptiles
as the " Ammo mr hide," or " Ammonst'ormation," and in a series of excellent figures of coronal sections of
the brain of a Saurian he leaves no doubt as to the meaning of ids written statements ; for he labels the
medio-dorsal cortex in these illustrations " Cortex Ammonis " \. It is instructive to quote his reasons
for this conclusion, because they are logical and well-founded, and have a considerable interest in view
of the fact that their author was led into a singular error five years later and gave up his earlier and
correct belief for another view. In 1888 he wrote {up. cit. p. 116): — "Das betrefFende Rindeastiick
[medio-dorsal cortex] muss man als orstes Auftrcten der Ammonsrinde [hippocampus] in der Tierreihe
ansehen. Die Ammonsformation wird characterisirt durch ihrc Lage iiber dem Hemisphiirenrand,
durch den ilir anlagerndcn Fornix uud durch den auf ihn folgenden Plexus choroides. Sic enthlilt also
bei ihrem ersten auftrcten alle die Elemeute welche spater, wenn auch durch Windungen und
dergleichen verwischt, das charactcrische Ammonshorn bilden."
Fis. 20.
Part of a transverse section through the cerebrum oi Hydvosnurus (immediately in front of the commissures),
showing the decussating fornix-fibres (a).
This is not the first representative of the cornu Ammonis in the animal series, for it can be distinctly
recognized in a large number of the lehthyopsida ; nor is it accurate to speak of part of the paraterminal
body as the " fornix." Rut in spite of these inaccuracies, Edinger^s argument is perfectly legitimate ;
for the position [at least of the caudal part] of the medio-dorsal cortex in the mesial wall of the
* According to Kdiuger, Abhandl. d. Senckeuberg. (icsellschaft, 1^96, p. 314.
t L. Edinger, " Untersuchungen iiber die vergleichende Aiiatomie des (Jehirns— I. Das Vorderhirn," Abhand-
lungen der Senckenbergischen naturforschendea Gesellschaft, 1888, Taf. iii. figs. 21, 22, 23, and 24.
G9*
474 PROF. G. ELLIOT SMITH ON THE MOEPHOLOGY OF THE
hemisphere and its relation to the choroid plexus are valid reasons for regarding the cortex in question
as the representative of the coruu Ammouis, or hippocampus, of the Mammalian brain.
Having homologized this caudal portion of the medio-dorsal cortex with the hippocampus, Ediuger,
with good reason, did not hesitate to regard the cephalic extension of the same histological formation as
also part of the hippocampus. But the amazing feature of Edinger's memoir is that, after having
recognized this medio-dorsal cortex as the hippocampus^ the author does not hesitate to call the
commissure which is derived from it by the name " corpus callosum." And this name has clearly not
been employed in the wider sense as including the hippocampal fibres, because upon page 117 this
statement is found : — " Ueber der Commissura anterior liegt (fig. 22) die grosse von Osliorn mit
Recht als Balken bezeichnete Mantel-Commissur. Ihre Fasern stammen aus dem Gebiet der grossen
Zellen in den dorsalen Wand uach aussen der Ammonsrinde."
The figure 22, to which Edinger refers, represents all these commissural fibres springing not " aus dem
Gebiet nach aussen der Ammonsrinde," but from the hippocampus itself, and, so far as ray own
observations go, this figure is perfectly accurate. In other words, the great majority, if not all, of the
fibres of the dorsal commissure in the Reptile spring from the hippocampus, and therefore cannot be
called " corpus callosum."
It is perfectly incomprehensible why Edinger, who definitely recognized the hippocampus and the
fibres of the dorsal commissure springing from it (as his figm'es clearly show), should have called the
commissure " corpus callosum." He seems to have deliberately gone beyond his facts in order to make
a spurious pretence of agreement with Osboru. NowOsborn (Morphologisches Jahrbuch, 1887, Bd. xii.
cit.) did not for a moment deny the origin of the fibres of his so-called "corpus callosum" from the
mesial cortex in the Reptile. The fallacy of his argument was his failure to recognize this mesial cortex
as hippocampus. Edinger clearly recognized the latter fact, but failed to see the logical outcome of his
suggestion. He seems to have utterly ignored the facts which he had himself clearly demonstrated, for
the sake of a pretended confirmation of the conclusions of Osborn.
Two years after the publication of this memoir of Edinger's, Brill published a brief communication *,
with a view to make known the work of Spitzka, and to record a series of original observations upon the
brain in the Lacertilia, Ophidia, Crocodilia, and Chelouia. He argued that the position of the medio-
dorsal cortex in the hemisphere and its histological structure showed its homology to the fascia dentata.
He regarded the rest of the dorsal cortex as the hippocampus and subiculum cornu Ammonis. And in
support of this hypothesis he propounded (upon wholly inadequate grounds) the statement that " the
fascia dentata is more primitive than the hippocampus," and therefore not unnaturally " forms the greater
part of the Reptilian hemisphere." In the introduction to this discussion I pointed out that the fascia
dentata is a specialization of the ventral edge of the pre-existing hippocampus, which makes its
appearance relatively late in the ontogenetic history of the Mammalian brain. Moreover, it consists of
a peculiar modification of the surface (only) of the hippocampus, as I have shown elsewhere ("The
Fascia dentata," Anatomischer Anzeiger, Bd. xii. 1896, p. 124.). So that it cannot exist apart from the
hippocampus.
In its typical form the fascia dentata is found only in the Mammalia, whereas the hippocampus can
be certainly recognized in all the Sauropsida, Amjjhibia, and Dipnoi, and even in some still lowlier
vertebrates. It is therefore preposterous to argue that " the fascia dentata is more primitive than the
hipuocampus," as Brill does.
In 1890 C. L. Herrick began the publication of a series of contributions to the morphology of the
brain. The first of these memoirs which deals with the question at issue appeared in the first number
of 'The Journal of Comparative Neurology," which this prolific writer founded in 1891. It deals
* N. E. lirill, " The True Homology of the Mesal Portion of the Hemispheric Vesicle in the Psauropsida [sicV'
Medical Hecord, March 29, 1890, pp. 343-345.
CEREBRAL COMMISSURES IN THE VERTEBRATA. 475
mainly with tlic brain u\' a Lizard {Sccloporiis), a Snake wiiicli lie does uot name, and a 'i'nrtle (As/ndo-
vectes). In the Lizard, he says (p. 15), tliere is in the occipital lobe "an undoubted horaologue of the
hip])ocampus." Li the Snake he says that " the portion liomologous to the hij)poeainpus is relatively
highly ditl'crentiated " ; and that even "the portions corresi)onding in cellular structure to the fornicate
and uncinate i^-yri may be distinguished " ! By these statements, which are nonsense as they stand, the
author really means to record a would-be important fact. In a later communication upon the brain of
DideljJiys (" The Cerebrum and Olfactories of the Opossum," Journ. of Comp. Neurology, vol. ii. 1892)
the author calls the fascia dentata the "gyrus uncinatus," the hippocampus the " gyrus fornicatus,"
and the hip])ocami)al fissure the " splenialis fissure." The monstrous nature of such use of well-known
terminology does not seem to have been a])preciated by their author, for when Dr. Hill called his
attention to the unusual application of these terms he naively remarked that "obscurity was needlessly
introduced by using the terms t/i/rns fornicatus and uncinutus for parts which are more
often named cornu Aminonis and fft/n/s dentutus respectively, without discussion of homologies
concerned" (Journal of Comp. Neurology, vol. iii. (1893) ! In the light of these remarks it is possible
to understand his statement of 1891, which I have already cpioted above. According to this inter-
pretation, it is possible to distinguish in the hippocampus of the Black Snake a marginal region
corresponding to the fascia dentata and another region corresponding to the hippocampus (sensu
stricto).
In this (1891) memoir, Merrick gives a good description of the cephalic portion of the real hippo-
campus, which he calls " fronto-median lobe " (p. 18), since he does not recognize its hii)[)ocampal
nature. This is very surprising, because he records the fact that the so-called " occipital cortex " (which
he regards as the whole of the hippocampus) and his " fronto-median lobe " are not only continuous,
but pi-csont similar histological features. He does not hint at the fact that the " fronto-median lobe "
may also be hippocampus : and his apparent ignorance of the earlier memoirs of Spitzka, Edinger, and
Brill exiilains the lack of the suggestion from outside sources, which his own iiistological studies should
have supplied.
The reasons which dictate this distinction between the caudal and the cephalic regions of one uniform
and indivisible histological formation are probably to be found in the fact that the caudal portion
corresponds in its topographical position to the hippocampus in most Mammals. Yet, strange to relate,
neither the study of Didelpliys nor the knowledge of the work of Edinger and others appear to have had
any influence upon Herrick's interpretation; for, in spite of the fact that the hippocampus extends
forward in the ^larsupial, and in spite of the suggestions of Edinger regarding the hippocampus in
Reptiles, he, in his later works, still clings to his original view that it is confined to the caudal part of
the hemisphci'e. And for this reason it is not surprising to find that, after deujnng in a positive and
formal manner the hippocampal nature of his "fronto-median lobe," he regards its commissure as the
true corpus callosum — for this is the natural and logical inference to be drawn from his erroneous
interpretation.
In 1893 Edinger returned to the discussion of the problem of the hippocampus in the Keptilia*, and
practically renounced the view (undoubtedly correct) which he had published in 1888. In the later
work he attempted to extend the region of the hippocampus to the lateral aspect of the hemisphere in
the Chelonia, being apparently deceived by the spurious resemblance of an inverted cortical area in this
situation to the Mammalian hippocampus. Not only did he commit this extraordinary error, but also
the less excusable one of locating the fascia dentata upon the lateral aspect of the hemisphere external to
the Idjjpocampus (cornu Ammonis). It is only just to add tJiat he subsequently appreciated the ground-
lessness of these suggestions, and amply acknowledged his error (Abhandl. d. Senckenberg. Gescll.
l89G,p. 326).
* L. Edinger, " Eeichapparat und .immonshorn," Auat. Anzeiger, Bd. viii.
476 PEOr. G. ELLIOT SMITH ON THE MOEPHOLOGT OF THE
I may pass over the consideration of the works of the brothers Ramon y Cajal, of Koppen, Schulgiu,
Botazzi, Maracino, and Milia, because they contribute nothing fresh to the morphological problem under
discussion, and as they have been reviewed by Edinger (Abhandl. d. Seuckeuberg. Gesell. 1896, pp. 318-
319).
In 1892 a most important contribution to the discussion of the problem made its appearance, its
author l)eing Adolf Meyer (" Ueber das Yorderhirn eiuiger Reptilien," Zeitsch. f. wiss. Zoologie, Bd. Iv.
p. 63).
Meyer attempted to institute comparisons between the Reptile and the foetal, rather than the adult,
Mammal. He called the paraterminal body the "septum lucidum," but this loose application of a term,
which is strictly associated with a special modification of the dorso-caudal extremity of the paraterminal
body, and which is found only in the more highly organized j\Iammalia, can hardly be regarded as happy,
and at the outset prejudices the chance of an exact comparison. He described a furrow in certain
Reptiles, which marks the line of demarcation between the paraterminal body and the cortical area,
which I have referred to as hippocampus. This furrow is the fissura limitans hippoccmipi, and Meyer
homologized it with the furrow which separates the so-called " Randbogeu" into outer and inner arcs.
While this comparison with the somewhat problematical " Randbogen " of the Mammalian foetus is
unfortunate, he further compared the cortical portion of the mesial wall of the Reptilian brain with the
" outer Randbogen " of the mammal, but he did not commit himself to any more precise homology
than this.
He declined to call the dorsal commissure " corpus callosum," as Osborn, Edinger, and most writers
before him liad done. The reason for this is not that the fibres arise from the region which he compared
with the " outer Randbogen," but becarise the commissure in question is situated in the lamina terminalis,
and not in a secondary fusion of the mesial walls of the hemisphere. This argument is fallacious, as it
is based upon a misconception as to the real position of the primordial corpus callosum.
In 1896 Edinger once more reconsidered the whole question of the morphology of the mesial wall of
the liemisphcre in Reptiles ; and, as I have already mentioned, he then corrected the errors which he
committed in 1893. He approached the study of this subject again, not only with the much wider
knowledge which his own researches has yielded, but with the benefit of the results of the work of
Spitzka, Brill, Herrick, and especially Meyer, on the Reptilian brain ; and of Symington, Hill, and
myself, upon the lirain of the Proto- and Metatheria. Von Kolliker also, in the same year, added the
weight of his authority to the controversy, but apparently merely as the interpreter of the data collected
by the writers quoted above. Indeed, the view with which the names of Edinger and Kolliker became
associated in 1896 is so plausible and so insidiously deceptive that it is necessary to state it at some
length, with the objections to it. In 1894 Kolliker stated that some of the uncrossed fibres of the
formx in the Mannnalian brain are derived from the gyrus foruicatus — that is, from a part of the
neopallium beyond the limits of the hippocampus *. In reviewing this work shortly afterwards,
Edinger t tised the term 7?aw6?!t7»«?;«J9 as a synonym fo;- "gyrus foruicatus"; and elsewhere he subse-
quently used the term "gyrus Ibnbicus" in the same sense. I mention this specifically, because both
of these terms had been previously used as synonyms for the bidusiuin grisemn, which is not a part of
the gyrus Ibrnicatus, but is composed mainly of the vestigial supracallosal portions of the hippocampus
{vide Schiifer, ' Quain's Anatomy,' ed. x. vol. iii. pt. 1, 1893, p. 159). In 1896 Edinger spoke of the
gyrus limbiciis in nuimmals as the frontal continuation of the " Ammonsrhtdr,'^ " welche nicht mehr
eingerollt ist und uber dcm Ventrikel liegt." From the reference to the inroUing, it is clear that the
author cannot be using the term " Ammonsrinde" in the literal sense of " gyrus hippocampi" (which is
* " Ueber don Fornix lougus von Forel,'' Verhandl. d. anatom. Gesellschaft in Strassburg (Anat. Anz. Bd. is.
p. 45, Erganzung).
t " Leistungen auf dem Gebiete der Anatomie des Centralnervensystems," Schmidt's Jahrbiieher, Bd. ccxlvi.
CEREBEAL CO.MMISSUBES IX THE VEKTEBRATA. 477
rc'uUy in coiitimiity with the gvnis I'oniicatus * ), but as a synonym for tlie more usual name " Ammons-
horn," i. e. coruu Ammouis or hippocampus. He further added : " Ich halte es fiir wahrscheinlich, dass
in dem Rindeugebicte, welches bei den Reptilien als Aramonsrinde bezeichnete wurde, die Elementc des
Gyrus limi)iciis ^</(c/ der Ammonsriude gegcbeii sind " (" I'ntersuchungeu iibcr d. vergl. Anatomic d.
Gehirns. 3. Neuc Studien iibcr das \'orderliirn der Reptilien/' in Abhandl. d. Senckcnberg. Gescllsch.,
1896). Thus, while he was clearlj' using the term " gyrus limbicus" in the sense of the " gyrus fornieatus "
he spoke of this structure as the forward continuation of the hippocampus, as if the latter were identical
with the pallial area confusedly known as the " hijjpocampal gyrus." The first quotation is so vaguely
expressed that one might readily imagine the author to ])e using the term "gyrus linibicus" (or " Raud-
windung") in the sense of the inJusiiiin or vestigial hippocampus, as Schiifer does in ' Quaiu's Aiiatomv.'
In this ease tlie statement would be quite intelligible; but this interpretation the author never intended,
for while in the second cpiotation {siipi^a) he referred to the "Gyrus limbicus iinil der Aramonsrinde,"
in his ' \'orlesungen,' which were published contemporaneously, he called the imlusium {strire
lom/i/inlinales Lancisii) t\w forward continuation of what he somewhat rashly terms the " schon im
.\mmonshorne atrophisehen Windung, des Gyrus dentatus." (L. Edinger, ' Vorlcsungen iiber den ]5an
der NerviJsen Centralorgane des Menschen und der Tliiere,' 5th ed., Leipsig, 18'JG.)
It is clear from the foregoing quotations that Edinger, in renouncing his erroneous views of 1893, did
not revert to his original belief of 1888. On the contrary, he stated distinctly that the region which he
had previously called cornu Ammonis, and of which he now speaks as the " Ammonsrinde," contains
the elements not only of the " Ammonsrinde" or hippocampus, but also of the (ji/rus limbicus (or gyrus
fornieatus). Some inkling of the meaning of these extraordinary views is obtained from Ediuger's
definition of the fornix lum/us as that "welcher aus demjonigen Theile der Randwinduug, welcher
nicht zum Ammonshorne sicli einrolltc, aus dem Gyrus limbicus" (Vorlesungen, p. 210). In this
statement he was evidently attempting to harmonize the morphology of the cerebral cortex with the
supposed grouping of the fibres springing from it. In other words, he argued that the gyrus fornieatus
is the anterior continuation of the hippocampus, because tlie fornix is derived posteriorly from the latter
and anteriorly from the gyrus (according to the belief of KiiUiker [vide supra] and Edinger). Again, he
stated {op. cit. p. 168) that in the higher Mammals, as a consequence of the development of the corpus
callosum, the fibres, of the fornix coming from the hippocampus become separated from those which
come from the more anteriorly placed part of the " Randwinduug" (meaning the gyrus fornieatus).
By thus introducing the idea of a " Randwindung " or gyrus marginalis, which includes both gyrus
fornieatus and hippocampus, Edinger obviously intends to establish a morphological justification for
Kijlliker's belief in the origin of the fornix in part fi-om the gyrus fornieatus. \Vhether any fibres of
the fornix ever have any such origin is extremely doubtful {ride "The Fornix Superior," Journal of
Anatomy and Physiology, vol. xxxi. p. 90).
Kolliker in his ' Gewebelehre ' {op. cit. 189(5) ap)n'oved of Ediuger's interpretation, and translated
the vague and somewhat cryptic language of the latter into a clear statement, free from all ambiguity.
He thus distinctly, though unintentionally, reduced the argument to a reductio ad ubsurdum !
Thus Kolliker, whilst speaking of the Re])tilian brain (' Gewebelehre des ^lenschcn,' Bd. ii., zweitcr
Hiilfte, 1896, p. 825), writes : — "Edinger hat schon cine Andcutung, ob dieser Fornix der Reptilien
nicht dem Fornix hmgus der Siiugcr cntspreche, die ich voU unterstiitzen mochte. Die mediale Waud
des Yorderhirnes der Reptilien cntsjiricht ja nicht nur Animonsfalte, sondern audi dem Gyrus forni-
eatus der Sanger, aus welchem der Fornix longus entspringt. Ein Fornix inferior kann bei den Reptilien
kaum vorhandcu scin, da auch das Ammonshorn, in weleheni derselbe entspringt, nicht gebildet ist."
This means that Edinger and Kiilliker regarded the cortex of the mesial wall of the llcptiliau
* I mention this point specitically because elsewhere Ediuger has on several occasions confused the hippocampus
and gyrus hippocampi.
478 PKOF. G. ELLIOT SMITH OX THE MOKPHOLOGT OF THE
hemisphere as partly liomologous with the gyrus fornicatus, i. e. with a cortical area of the mammalian
brain which is not hippocampal. And yet both of these writers spoke with complete approval of
the view that the dorsal commissure, which springs from this mesial wall, is the psalterium or
hippocampal commissure. But if the mesial wall be partly gyrus fornicatus, its commissure cannot
be the psalterium, but must be the corpus callosum. There can be no doubt that the homology of
the mesial wall of the Reptilian hemisphere with the hippocampus is correct ; and the comparison
with the Monotreme brain, which I instituted in an earlier ])art of this memoir, shows conclusively that
no part of the mesial wall of the Reptilian hemisphere can be regarded as the homologue of the gyrus
fornicatus.
It is strange that when Edinger held the mesial cortex of the Reptilian hemisphere to be
hippocampus, he called its commissure not " psalterium " but " corpus callosum " (oy;. cit. 1888); and
that when, eight years later, he changed his view and regarded it as including not only the
hippocampus but a uon-hippocampal pallial area, he called it not " corpus callosum " but " psalterium,"
with a singular disregard for consistency.
A comparison of the Reptilian cerebral hemisphere with that of the foetal and adult Monotreme
conclusively demonstrates that the whole of the mesial wall of the former which is ])laced above
the paraterminal body, together with a considerable part of the roof of the hemisphere, represents
unquestionably the mammalian hippocampus, and nuthing else.
Although the Reptilian hippocampus is not so highly differentiated as its Mammalian homologue,
in that its margin is not yet definitely specialized to form a fascia dentata, I utterly disagree with
Kolliker in the statement that in the Reptilia " das Ammonshorn nicht gebiklet ist." ^
The arguments for the recognition of the medio-dorsal coitex in the Sauropsida as the hippocampus
may be concisely summed up in the following manner. A comparison of the brain of the Eutheria with
the Meta- and Protothcria shows that in both of the latter lowly orders of Mammals — widely as they
are undoubtedly separated the one from the other — the hippocampus has a much more extensive
distribution than in the formei'. Instead of being confined to the caudal portion of the hemisphere, it
extends forward as far as its morphologically anterior pole ; and as a search in the analogous region of
the Eutherian hemisphere has revealed vestiges of this cephalic extension of the hippocamjius, it is
natural to conclude tliat the ancestors of the Mam.malia possessed a hippocampus extending
FORWARD upon THE JIESIALWALL OF THE HEMISPHERE TO ITS CEPHALIC POLE, «. C. TO THE NEIOHBOUR-
HOOD OF THE OLiACTonY PEDUNCLE. The presence of a peculiarly distinctive histological formation in
the brain of the Reptile exhibiting not only the identical relations of the Prototherian cornu Ammonis,
but also possessing a structure which corresponds to one phase through which the Mammalian hippo-
campus passes in its ontogenetic development, can be rightly interpreted in only one way. These
reasons for thus homologiziiig the mesial cortex in the Reptile have hitherto never been categorically
formulated, and they constitute the only legitimate ground upon which such an interpretation can be
based. The early suggestions of Spitzka and Brill, and Edinger's extraordinary memoir of 1893, are
little more than guesswork. The value of the legitimate argument of Edinger in 1888 Avas utterly
destroyed by his later contributions in 1893 and 1896. And the sound work of Adolf Meyer left
the question in too undecided a state to be regarded as in any way settling the question.
A hippocampus presenting the general features already briefly indicated exists in all classes of Reptilia,
but the details of its structure vary somewhat in the different families. lu all Reptiles the hippocampus
forms not only the whole of the mesial wall of the hemisphere above the paraterminal body, but also
part of the dorsal surface of the hemisphere. In many of the Chelonia, such, for example, as Emys
* Much of Hie foregoing discussion has already been published in a short memoir entitled " Further Observations
upon the Fornix, with 8j)ecial Reference to the Brain of Si/dophilus," Journal of Anatomy and Physiology, vol. xxxii.
1898. It is repeated here in order to make the argument complete and intelligible.
CEREBRAL COMMISSURES IN THE VERTEBRATA. 479
europma, the cells of the liippoeampus arc luucli more diffusely scattered than in Hydrosaurus and, to a
lesser degree, in Sphenodon also. This more diffuse scatteriug of cells points to the more primitive
condition of the hijjpocampus exhibited in the Amphibian brain. Osboru has given such an exeelleut
representation of it in Kmys, that it is quite unnecessary for me to do this here (Osl)orn, Morph. Jahrb.,
Bd. xii. Taf. xiii. tig. I(i). In Chelone, hoHcver, the cells are much more compactly arranged, so as to
form a regular column which presents a close resemblance to that found in Sphenodon, which I have
already represented *.
In the Chelonia, Crocodilia, and Rhyuchocephalia the characters of the distinctive column of cells of
the hippocampus arc iJi'actieally uniform throughout the whole length of the coluuni ; w hcrcas in the
Laccrtilia, and especially in the Ophidia, the cells become much more numerous and considerably smaller
in size in the neighbourhood of the paraterminal body. This fact is now well-recognized, and has been
successively noted by Herrick, Edinger, Meyer, and myself among many others ; and the condition has
been adinii:i!)ly rcjiresentcd in the excellent figures wiiicli accompany Edinger's memoir of 1888
(he. fit.) and Meyer's of 189;} [hc.cit.].
In the Chelonia, no such alteration in the characters of the ventral extremity of the hippocampal
column of cells can be detected, nor am I aijle to detect any trace of such a change in the brain of
Sphenodo/i.
This peculiar feature of the hippocampus in the Saurians is significant. For it points to a more
rapid proliferation, resulting in the production of a large mass of cells of small size, in a position
analogous to that in which the fascia dcntata has been found to develop, by a similar process of rapid
proliferation and small-cell formation, in the Mammal. But it may l)e argued that the fascia dentata
in the Mammal is a group of cells set quite a[)art from the hippocampal column of pyramidal cells.
The developmental history of the fascia dentata, at which I have briefly hinted in the Monotreme,
and fliuch follows a similar course in the placental Mammal, shows that the fascia dentata, or rather its
"stratum grauulosum," is originally the ventral (or inner) extremity of the column of pyramids which
becomes separate at a later period of development. This process of separation can be actually seen in
the adult brain of any Monotreme or ■Marsupial — for in these Orders the fascia dentata becomes reduced
to very insignificant proportions at the cephalic extremity of the hippocampus; and as its cells are thus
brought into line with the stratum of pyramidal cells of the hippocampus proper, the picture of the
Saurian eonditi(m is complete {vide "The Fascia dentata," Anat. Anzeiger, 1896). It is not surprising
to find that this ditt'erentiation of the hippocampal column of cells is not found in all Reptiles, when we
recall the fact that in the Mammal the fascia dentata is late in making its appearance. And it is of
interest to note (although it is not quite pertinent to this discussion;, that, just as the fascia dentata is
the last part of the hippocampal formation to make its appearance, so it is the first part to dwindle
and disappear in the retrogressive changes which involve the anterior portitms (supra- and precommissural)
of the great hippocampal arc in the Eutheria, leaving a hippocampal vestige devoid of fascia dentata
{vide supra; also Trans. Linn. Soc. ser. 2, Zool., vol. iii. 1897, pp. 1-7-69; also Journal of Anatomy
and Physiology, vol. xxxii. 1897).
The problem of the meaning of this peculiar change resulting in tiie development of the fascia dentata
presents itself for solution. It is an extremely difficult question, which does not peruiit of being definitely
settled in the present state of our knowledge. But the following suggestions may form a tentative
hypothesis until such a fuller knowledge is forthcoming.
In the Reptile large numbers of nerve-fibres stream from the bulbus olfactorius into the neighbouring
area of hippocampus. In the Monotreme and Marsupial the corresponding fibres become collected
mainly at the ventral margin of the hippocampal formation (see Journal of Anatomy and Physiology,
* Good figures of the brain of Chelone mydas will be found in Edinger's memoir in the Abhandl. d. Seiickenberg.
natnrf. (iescUscb., ItSltd, Taf. 2. figs. 5 & G.
SECOKl) SERIES.^ZOOLOGV, VOL. VIII. 70
480 PEOP. G. ELLIOT SMITH ON THE MOEPHOLOGT OF THE
vol. xxxii.), and form the fasciculus marginalis. Olfactory impulses coming from the olfactory bulb are
thus poured into the ventral fringe of the hippocampus, and this fringe becomes specially modified to
form a large mass of " receptive cells " — the fascia dentata. In other regions of the brain modifications
of superficial cells are found in regions in which sensory paths impinge upon the cerebral cortex. This
is very slightly shown in the general cortex of higher Mammals because the sensory fibres are widely
scattered; but it is most pronounced upon the surface of the lobxis jjyriforniis {vide supra) : here a very
rich supply of incoming olfactory fibres covers the surface of the lobe, and the superficial cells become
specially modified to form the layer of " double pyramids," which is such a characteristic feature of this
region. In a similar manner a peculiar cortex is developed in the tuberculum olfactorium {vide supra).
It would seem, therefore, not unreasonable to suppose that as the fibres coming from the olfactory
bulb began to collect at the ventral margin of the hippocampus, the latter region became specially
modified to form as the fascia deutata a receptive organ for olfactory impulses.
In the Saurian brain a similar tendency is shown, and hence it follows that the small-celled ventral
portion of the Saurian hippocampus should be regarded at least as the physiological, if not the strict,
homologue of the Mammalian fascia dentata. In all the Saurians studied, the hippocampal formation
comes to an abrupt termination upon the dorsal surface of the hemisphere ; and the characteristic
column of cells ceases, and its lateral extremity is separated from the rest of the cortical region of the
brain by a pronounced hiatus. There is no reason for regarding the regions which are placed beyond
this hiatus as constituting part of the hippocampus. The mesial half of the roof of the hemisphere is
considered to be hippocampus because it is part of the one circumscribed histological formation,
concerning the homology of the mesial part of which there can be no question ; and there is no valid
reason for not extending this interpretation to the whole formation. But when this typical structure
comes to an end, it is only logical to infer that a region beyond the hippocampus has been attained.
Now, by comparison with the I\Iammalian condition, it is clear that this cortical area beyond the hiatus
must represent the neopallium, to which I have already referred in the Mammalian brain.
In the Mammal, however, the transition from hippocampus to neopallium is usually a more gradual
process than this abrupt method which characterizes the Saurian ; and if an analogy for this be desired,
it will be found in the Chelonia and Rhynchocephalia, where the regular hippocampal column (jrudualhj
gives place in the roof of the hemisphere to the scattered arrangement which is characteristic of the
neopallium. In the earlier figures illustrating coronal sections through the hemispheres of Sphenodon,
this gradual transition from hippocampus to neopallium has been shown.
The fibres which arise from or terminate in this hippocampus must be regarded as the fornix (in the
sense in which I defined this term : " The Relation of the Fornix to the Margin of the Cerebral
Cortex," Journal of Anatomy and Physiology, vol. xxxii. 1897) : the Reptilian fornix, like its
^Mammalian representative, consists of sl fornix transversus (Forel) or hippocampal commissure [psalterium,
lijra,fornia.'-commissurc), and a form u- longus (Forel) composed of descending post-commissural fibres
[columna fornicis) and " pre-commissural fibres" of Huxley.
The uncrossed fibres (or fornix longus) need not be further considered at present, but the crossing
fibres {fornix transversus) are of such great morphological importance in the discussion of the arrange-
ment of the cortex, that they cannot be dismissed in this summary fashion.
The fibres which spring from the hippocampus and proceed across the mesial plane to enter the
hippocampus of the other hemisphere cannot be regarded otherwise than as the homologues of the
commissure variously known in mammals as psalteriwn, lyra, cmmnissura fornicis, commissura hippo-
campi, commissura cornu Amnionis, fornix transversus, as well as by other names. This commissure is
regarded as " hippocampal," not by reason of any particular relation it may present to the lamina
terminalis [compare Meyer's reasoning], to the recessus superior [comijare the writings of Osborn,
Herrick, and many American and German writers], to the roof of the forebrain or to the foramen of Monro,
but because its jib res are derived from the hippocampus. Stated thus directly, this truism may seem too
CEEEBEAL COMMISSUEES TX THE YEETEBRATA. 481
obvious to ucecl formal presentation ; but a glaucc at tlic past history of tliis discussion woukl appear to
show that the inference does not naturally suggest itself to most writers ; and that in deciding the
question of homology, the great majority of investigators have made use of criteria which may readily be
shown to be either utterly false or of limited applicability.
In the foregoing descriptions it has been demonstrated that a large strand of fibres coming from the
hippocampus crosses to the corresponding region of the other hemisphere in the lamina terminalis and
forms the dorsal commissure. The fact that the morphological relations of this commissure are identical
with those of the dorsal commissure in the INIonotreme, which is known to be of purely hippocampal
origin, undoubtedly lends support to the contention that the dorsal commissure in the Reptilian
brain is also a commissure of the hippocampi, i.e. t\\c psa/terium. But such an argument can only be of
subsidiary value. The only essential reason for regarding the dorsal commissure as being hippocampal,
and not a true " corpus callosum," must be the demonstration of its origin from the hippocampus. So
far as 1 am awaie, no investigator, with the solitary exception of Ramon y Cajal, has ever seriously
attempted to claim an origin of any fibres of the dorsal commissure from the ])allial area beyond the
hippocampus. Edinger, it is true, appeared to do so in his first contribution to this discussion (op. cit.
1888), but, as I have already pointed out, his statements are not supported by his figures. And the
internal evidence of his memoir seems to point to the statement merely as an attempt to harmonize his
conclusions with the plausible and sweeping generalization of Osborn.
When Osborn himself argued in favour of the existence of a " corpus callosum " in the Reptilia, he did
not trace any fibres of the dorsal commissure beyond the medio-dorsal cortical jjlate. He was led to the
erroneous belief in the presence of the corpus callosum, because he did not recognize in the medio-
dorsal plate the true hippocampus (H. F. Osborn, "The Origin of the Corpus Callosum," Morpholog.
Jahrblich, Bd. xii.). The same remarks apply to the earlier observations of Stieda (" Studien iiber das
Centralnerveusystem der Reptilieu," Zcitschr. f. wiss. Zoologie, Bd. xxv. 1875).
Rabl-Riickhard, however, would not homologize the dorsal commissure in the Alligator with the
corpus callosum, and gave it the neutral name " coiumissui-a pal/ii anterior"*. His reason for this
hesitancy was that he regarded it as corresponding neither in position with the corjjus callosum, nor
with what one should regard as the primitive stage of the great Mammalian commissure. He somewhat
doubtfully suggested its homology with the commissure of the fornix, but gave no valid reasons for his
belief.
In his numerous contributions, to which reference has already been made, Herrick regai'ded the dorsal
commissiu'c as a true " corpus callosum," for the reason that he refused to admit the homology of the
medio-dorsal plate of cortex with the hippocampus.
Then, again, it was the relation of the dorsal commissure to the lamina terminalis, rather than its
origin from the region which he compared to the " outer Randbogen," which led Adolf Meyer to refuse
the title "corpus callosum" to the dorsal commissure in 1893 {(Jp. cit.). In order to justify his
])osition ^leyer, in 1895, ])rought forward some rather arbiti'ary criteria of a " corpus callosum " (which
1 shall discuss later), but they added little to the force of his original argument.
It is somewhat surprising to find that in 1896 Edinger {loc. cit.) regarded the dorsal commissure
(which he now calls the " commissura pallii anterior" after Rabl-RUckhard) as a, psalter ium, although
he regarded the origin of this commissure as being homologous not ouly with the hippocampus but also
with the gyrus fornicatns of Mammals. He attributed his change of belief to the work of Adolf Meyer
{op. cit.) and the writer (" The Cerebral Commissures in the Mammalia/' Proceedings of the Liunean
Soe. of N.S.W. 2nd series, vol. ix.), and very properly argued that, as the commissural link between the
hippocampi, it must represent the psalterium (Edinger, op. cit. 1896, p, 371). But in this ])assage he
seemed to forget that elsewhere iu the same memoir (p. 38 1 inter alia) he refused to admit that the
* " Ueber d;is Centralnervcmsystem der Alligators," Zeitschrift fiir wiss. Zoologie, BJ. xxs. 1S7S.
70*
^82 PROF. (I. ELLIOT SMITH OX THE MORPHOLOGY OF THE
cortical area in question was hippocampus only, and concluded by regarding it as also representing the
gyrus t'ornicatus.
His lack of consistency is almost as pronounced as it was eight years earlier, when he called the
medio-dorsal cortex " hijipocampus " and its commissure " coi'pus callosum." In 1896 he almost
reversed tlie process by calling the commissure " psalterium," but declining to admit that its area of
origin is simply "hippocampus."
I have dwelt at great length and with, 1 tear, much repetition upon the views of Edinger, because his
opinion has been widely accepted from time to time by writers of all nationalities as authoritative, and
hence, in all recent memoirs and text-books which treat of the comparative anatomy of the brain, the
doi'sal commissure in Reptiles and Amphibians is called " psalterium." It only needs a writer to come
forward (as Flower did in 1865, in the controversy regarding the commissures in the Monotremes and
Marsupials) and indicate the weakness of the basis for the general belief, to cause a general revulsion to
the conviction of the presence of a corpus callosum in all Vertebrates. For so history might repeat
itself. But although Edinger's argument laclvs cogency and consistency, the co)iclusion at which he
arrived regarding the dorsal commissure is undoubtedly correct. For the fibres of the dorsal commissure
must be hippoeampal, i. e. psalterium, because they spring from the hippocampus.
In this discussion I have not yet referred to the possibility of the fibres of the dorsal commissure
arising h'oni a wider cortical area than the hippocampus. The only author who, so far as I am aware,
directly admitted this possibility is Pedro Ramon y Cajal (' Investigacioncs niicrograficas en el Encephalo
de los Batraceos y Reptiles,' Zaragoza, 1894). I am only acquainted with this work at second hand
from the excellent figure which Edinger reproduces [op. cit. 1896, p. 319). In this figure a group of
fibres is represented passing upward from the dorsal commissure (which he calls " corpus callosum ")
through the paraterminal body and hippocampus to end in the pallia! area beyond the latter. These
fibres are labelled " Mehr lateral aufgezweigte Balkeufasern fiir die laterale Rindenplatte.'' In no
specimens which I have examined have I been able to satisfy myself that any such fibres came from the
dorsal commissure and extend beyond the hippocampus. It may be that they merely consist of fibres
which associate the hippocampus and the lateral pallial area, and have no connection with the com-
missural fibres. Sucii fibres are found in the Mammalian brain. But Ramon y Cajal distinctly
represented the fibres in question as coming from the dorsal commissure.
If these fibres really exist as he represents them, and if the suggestion which I have already made to
the effect that the area beyond the hippocampus represents the neopallium of the mammalian brain is
correct, it must of necessity follow that the fibres in question are strictly homologous with the mammalian
corpus callosum. That such is the case, however, I am very loth to admit, not merely for the reason
that I have never been able to detect such commissural fibres passing to the lateral pallial ai-ea, but for
wider morphological reasons. There can be no doubt that no such corpus callosum exists in eitlur the
Monotremata or Marsupialia, and that therefore, presumably, the ancestors of the Mammalia had no
true corpus callosum, /. e. they possessed a dorsal commissure which was pttrehj luppocampul. ^Nlore-
over, a study of the conditions prevailing in the Mammalian lirain when the corpus callosum makes its
first appearance, seems to point clearly to the conclusion that the true corpus callosum develops in
response to the demand of the rapidly growing neopallium for a shorter route for its commissural fibres.
Such a demand can never exist in the Reptilia, in which the neopallium is of such insignificant area ; and,
« priori, one woiild be extremely surprised to find a representative of the true corpus callosum in the
Reptilia. Yet all these strung a priori objections would not be of sufficient iceight to counterbalance the
positive evidence for a true corpus callosum which the substantiation of Ramon y Cajal's evidence would
afford. The arguments of Meyer concerning its relation to the lamina terminalis, or of other writers
concerning its relation to the recessus superior and roof of the third ventricle, would be of no avail if this
crucial fact were established. In the meantime the question must remain in abeyance, for Ramon v
Cajal's conclusions need confirmation. His observations were made and recorded without anv due
CEREBRAL COMMISSURES TX THE VERTEBRATA. 483
sense of the morphological importance attaching to them, because he regarded the dorsal commissure as
tlie " corpus callosnm," not having recognized the liippoeampus as such. Hence the fact that the fibres
have a slightly wider area of origin would mean little.
The possibility of the existence of a corpus callosuni in tlie Rcptiiia in this sense has not hitherto been
suggested, so far as I am aware ; but it is patent that no other forui of true corj)t>s cfillosiim ran exist in
the ReptiUa thnn this, as yet, hypotlietical commissure of the lateral palliunt.
I shall have occasion to return ineidcnrally to this point later on. At the same time 1 eainiot too
strongly insist upon the fact that, even granting the existence of these true homologucs of the corpus
callosuni iu the reptilian In-aiu, the vast majority of tiie fibres of tlie dorsal commissure are unquestion-
ably derived from the liippoeampus, and therefore cannot l)c regarded as part of the true corpus
callosuni.
In the year 1887 Belloiiei |)nblishcd a nieinoir on the cerebral etniiniissures in the Amphibia and
Reptilia *, in which he stated that in his " tratto superiore della couunissirra anteriore " decussational fibres
were interiuingled with true commissural fibres. The commissure to which he thus refers is the
commissuru ilorsalis or liippoeampal commissure, which he regai'ded as part of the anterior couimissure —
an utterly untenable view. The existence of the decussational fibres which pass obliqueh' from the
descending hippoeampal fibres of one side to the opposite side of the brain has been since observed by
many investigators, and may be readily demonstrated by the method of Weigert, especially in the
Saurians. These fibres are represented in the accompanying drawing (fig. 20, a, p. 473), which has been
made, with the aid of the camera lucida, from a coronal section of the brain of Jlydrosaurus just in front
of the commissuru ventralis. I am utterly unable to offer any explanation of this extraordinary course for
fibres obviously belonging to the fornix system: nor am I able to find iu the Mammalia any fibres which
can be compared with these decussating bundles. No other writer, so far as I am aware, has offered any
feasible explanation of this arrangement. Ramon y Cajal represented the fibres in his figure to which
I have referred above; and it may be possible — although improbable — that they may represent the
continuation of the fibres which he indicates as coming from the neopallium beyond the hippocampus.
In other words, the;/ may be pallial projection fifjres. I do not, however, place any eonfidence in this
hypothesis, and can only mention this vague possibility in lieu of the utter lack of suggestion concerning
these decussating fibres. A knoAvlcdge of the alleged existence of such fibres should impose upon
investigators great caution in attempting to interpret the reptilian fornix.
It is unnecessary to write at any length regarding the commissures in the cerebral hemispheres of the
Bird. A study of the process of development of the chick's brain and a comparison of the histological
structure of the mesial wall of the hemisphere in the adult show that iu birds a hippocampus of the
chelonian type makes its appearance but never attains to large dimensions. From it a small hippoeampal
commissure springs, as in the reptile.
To speak of a " corpus eallosum " in the bird, such as Bryce aud Warrington have done even so
recently as 1898 (Report of International Physiological Congress, Cambridge, 1898), is a gratuitous
assumption ; because the dorsal commissure is unquestionably hippoeampal. The same arguments
concerning this homology which 1 have applied to the reptilian condition might be equally appropriately
employed in this case also.
* "SuUe Comniissure cerebrali auteriori degli Anfibia e doi Eettili," 'Slcm. del. Real, -iccad. del. Sci. dell' Istituto
di Bologna, Ser. iv. Tom. viii. IS)"?".
484 PEOF. G. ELLIOT SMITH ON THE MOEPHOLOGT OF THE
TJie Com2Jarative Morphology of the Commissura aherrans.
The problem of the interpretation of the cerebral commissures is, however, by no
means so simple as the above account may lead one to suppose. For there is iu
certain Reptiles the additional commissure, which I have distingviished by the name
" aberrans.'' So far as I am aware, E-abl-Riickhard * was the first writer to call attention
to it, in the brain of Pscmimqsaurns. He regarded it as a rudiment of the fornix or
" eine Kommissur fiir die Ammonshorner (Forel)." His reason for this was the fact
that its fibres spread out in the ventro-caudal region of the mesial wall of the hemisphere,
which he regarded as hippocampus, presumably because it occupies a position which is
topograi^hically analogous to that of the Eutherian hippocampus. Osborn wrongly
condemned this interpretation, and confused the commissura aherrans of the Reptilia
with the commisstira habenularum in the Frog, to which he had recently given the name
" coimuissuru superior' i. It is, however, only just to add tliat Osborn made this
comment before he had examined any brain 2:)ossessiug the aberrant commissure %.
According to BrUl §, the first observation of the aberrant commissure must be
attributed to Spitzka, who recorded its presence in Iguana and called it '• corpus
callosum " the year before E,abl-E;uckliard described it in Psammosaiirus.
Edinger confirmed Rabl-Eiickhard's observation, and adopted his suggestion concerning
its homology with the "■ commisura fornicis'' \. In this memoir he calls the commissura
dorsaUs by the name " corpus callosum," although, like the aberrant commissure, it
springs from the area which he correctly labels " Ammonsliorn.''
Two years later, Honegger ^ objected to the suggested liomology of the commissura.
aherrans with the fornix, and, if I correctly understand his somewhat involved argument,
he regarded it as the representative of the commissural fibres of the taenia semicircularis
(vel teriuinalis).
* Kabl-lliickbard, " Uebcr (las Vorkommen eiues Fornixrudimeuts bei Eeptilien," Zoolog. Anz. 1881, p. 281.
t H. F. Osborn, "The Origin of the Corpus CaUosum," Morph. Jahrb., Bd. xii. ]>. 243.
+ [En pf'ssant I must refer to a matter not altogether pertinent to the sul)jeot of discussion. I do So in order to
prevent a confusion which constantly occurs in the literature of this part of the brain. Since the introduction of the
term " commissura superior" b}' Osborn for the habenular commissure, it has attained a very wide and general
recognition among comparative anatomists. The names of Osborn, Bellonci, Burckhardt, Shijaley, Edinger, Hitter,
Studnicka, Prenant, and Sorenson, to mention only a few of those who have employed this term, indicate how
general and cosmopolitan the use of this term is. And yet, since this almost universal adoption of the term, a
prolific writer on comparative neurology, who claims to be familiar with the literature relating to the reptilian
brain, applies this name to the dorsal or hippocampal commissure in the Monotreraata and ilarsupialia, viz.,
Theodor Ziehen, in Jenaisch, Denkschr., 1897.
While referring to a case (that of Osborii) in which the commissura superior had been mistaken for part of the
hippocampal commissure (c. aherrans), I wish to protest against the singular inappropriateness of applying this same
name "commissura superior" to tlie hippocampal commissure itself, and thereby deliberately introducing an elemeut
of confusion.]
§ N. E. Brill, 'Medical Record,' March 29, ISiio, pp. 343-345.
II Ludwig Edinger, op. cit. su^jra, 1888.
% J. J. Honegger, " Uebcr den Fornix, &o.,'' Reeueil Zool. Suisse, tom. v., 1890.
CEREBRAL COMMISSURES IX THE VERTEBRATA. 485
In 1892, Adolf Meyer described the commissure in Iguana and called it " commlssura
pallii posterior" *.
In a series of memoirs published in 1892 and 1893, Herrick f records the presence of
the aberrant commissure in a series of Lacertiliau and, if I understand him aright +,
Ophidian brains, and adopts the same view as Edingcr.
In a later memoir Rabl-lliickhard ("Einiger liber das Gehirn der Riesenschlange,"
Zeitschr. f. wiss. Zoologie, Bd. Iviii.) adopted Meyer's title " commissura pallii posterior "
and agreed with him as to its presence in Saurians only. Thus he found it in
Psammosaurus, Lacerta, Iguana, Podineniu, and Chameleo, but nol in any representative
of the Chelonia, Crocodilia, or Ophidia. Herriek, on the contrary, somewhat vaguely
intimated that it is present in the Oi)hidian brain (" Topography and Histology of the
Brain of certain Reptiles," Journal of Comparative Neurology, vol. iii. 1893). He
referred to the commissurn aherram as the " hippocampal commissure," but when he
added that "this does not ditfer in any morphological respect from the corresponding
body in mammals" {op. cit. p. 89), it becomes clear that he had utterly failed to
a])preciate the extraordinarily peculiar features of this commissure.
In his latest account (1896) of the commlssura aberrans, Edinger, Avho adopted the
term " commissura pallii posterior" committed a number of errors, which are A^ery
surprising when it is remembered that he had then made a series of accurate obser-
vations on this region of the forebrain for eight years. Thus in referring to this (with
the other commissures) he made the general statement " in der Lamina terminalis
verlaufen die Commissuren " [op. cit. 1896, p. 339), without any qualification or any
remark to indicate that the commissura aberrans does )iot lie in the lamina terminalis,
but upon the epithelial roof of the third ventricle. This statement, moreover, was not
due to a mere looseness of expression or lapsus calami, because in his illustration of the
mesial surface of the brain of Varauus (tig. 1, p. 337) he represented the doubtful
commissure as being attached to the upper extremity of the lamina terminalis. [This is
shown even more clearly in the reproduction of the same figure in his ' Vorlesungen,'
1896, fig. 7G, p. 120.]
In the text of his memoir upon the forebrain of Reptiles {op. cit. 1896) he makes a
statement concerning the position of the commissura aberrans to the effect that '• die
caudale Mantelcommissur [commissura pallii posterior] liegt direkt von der Stelle, wo
die Hirnwand sieh zum Plexus verdiinnt, wie ich das schon in meinen ersten Mitteilungen
gezeichnet babe" (p. 372). This statement is in accord with the above-quoted remarks
concerning its relation to the lamina terminalis and is unquestionably erroneous, because
the commissure is placed behind, and not in front of, the situation of the mesial attach-
ment of the latenxl choroid plexus. This relationship is one of its most significant
features.
* A. Meyer, Zeitschr. f. wiss. Zool. BA. Iv. (1892) p. Go.
t C. L. Herrick, "Topograph}- and Histology of the Braiu of certaiu Reptiles,'' Jouni. Comp. Neurology, lby;3.
j Jouni. Comp. Neur., Ib92, p. 179.
480 PEOF. G-. ELLIOT SMITH ON THE IMOKPHOLOGY OF THE
This peculiar commissure has a very restricted distribution in the Vertebrate series.
It is not found in the Chelonia nor in the Crocodilia, but seems to occur in all the
Lacertilia (and perhaps in some Ophidia, although further investigations are needed to
harmonize the conflicting statements of Edinger, Herrick, and Rabl-Riickhard upon this
point *). There is no corresponding commissure in the Mammalia, nor is there any
exactly analogous arrangement elsewhere in the Vertebrate series. As the term
" commissura })allii posterior " i^ in many respects misleading and inapj)ropriate, as I
shall have occasion to point out later on, I have introduced the name commissura aberrcois
in reference to its peculiarly aberrant nature and limited distribution.
In Dendy's otherwise excellent figure representing this commissure (fig. 15) the
latter has been erroneously represented as having a complete epithelial sheath. I
have represented its actual relation to the roof of the forebrain in fig. 2 (p. 457)
of tills memoir.
No stronger testimony of the failure to understand the meaning of this commissure
coidd be adduced than the fact that Edinger, who has contributed a not insignificant
share to our knowledge of the commissura aherrans, should make such fundamental
errors regarding its position as I have mentioned above. But such statements are typical
of the obscurity which invests this peculiar feature of the Saurian brain ; and it becomes
necessary therefore, in consideration of the foregoing analytical paragraphs, to reconsider
the whole question of the possible significance of this commissure, because the literature
of the subject affords no sufficiently well-founded clue to its nature to be of any serious
value.
In the preliminary study of the roof of the forebrain in Splienodon^ which served as an
introduction to this account, it was clearly demonstrated (^cide figs. 1 and 2) that tlie
commissura aberraiis had no connection with the lamina terminalis, but is placed in a
fold of the epithelial roof which is separated from ihe lamina terminalis not only by the
Imnina chorioidea (the fold from which the lateral choroid plexuses are derived) but also
by the pai'aphysis. The commissura aherrans, moreover, is situated caudad of the foramen
of Monro, and it marks (as I have already showni in the introduction) the line of
demarcation between the strictly thalamic portion of the roof and the more cephalic
region which is closely related to the cerebral hemisphere. It is therefore situated at a
most important morphological site, the exact location of which may be more readily
appreciated from a schematic plan of the forebrain such as the accompanying diagram
(fig. 21). In this scheme the mesial cavity or third ventricle will be seen to extend
forward as far as the lamina terminalis, and then to communicate by means of the
foramen of Monro on each side with the lateral ventricle. The lateral walls of the third
ventricle in this diagram are formed by the optic thalami, and the anterior extremity of
each thalamus is joined by an attenuated band to the tliiu free cortical fold which forms
he lahiiiDi. cundaJe of the cerebral hemisphere. This cortical fold is the hippocampus.
Erom it fornix-lihres arise and proceed towards the dorsal surface of the cephalic
* Edinger auJ Eabl-Eiickhard are inclined to regard it as being absent in ISnakcs. I Lave been unable to find
am' trace of il in the Australian Suake Pseudechis.
CEREBEAL COMMISSUEES IN THE VERTEBRATA.
487
extremity of tlie optic thalamus by the most direct route, which happens to be situated
in the posterior wall of the foramen of Monro. These fornix-fibres then proceed across
the roof of the forebrain and form the commissura aberrans. A study of the accom-
panying' scheme will sliow that the commissure is placed at tlie exact situation in which
the narrow epithelial roof of the slit-like third veiitriclc passes into continuity with the
broad roof of the " aula," which expands in the lateral direction on each side to form the
choroid plexus. The latter passes through the foramen ol" Monro into the lateral
ventricle.
Fis. 21.
Schematic representation of a horizontal plan of the brain to iudicMto the situation of
the cominissnfa al'irrans.
In the introductory remarks I called attention to the fact that the pai"aterniinal
bodv is not limited by the lamina terminalis ; and I showed, by the lielp of drawings
of coronal sections, that a process or caudal prolongation of the paraterminal body
proceeds backward abov^e the foramen of ]\Ionro, and fuses with the dorso-cephalic
anii'le of the optic thalamus. lu order to )nake these dithcult and very complicated
relations intelligible, 1 have constructed a crude diagram (fig. 22, p. 488) of tlie
mesial surface of the forebrain (in part only) from wliich all epithelial folds Jiave been
removed.
This scheme will explain ho^- the paraterminal body extends backward alcove the
lamina terminalis and above the foramen of Monro, and how this caudal prolongation
fuses with the optic thalamus. At the savne time the paraterminal body alfords a
matrix or "commissure-bed" for tlie comiiilasiira aherraiis just as it does for the
commissnra dorsoUs and commissura centralis. It was to this caudal extension of tlie
paraterminal l)ody that Edinger gave the name •' Fornlxleiste.'" Other writers called it
the '• fornix-body " or even the " fornix." The latter term has been so used because
SECOND SEKIKS.— ZOOLOGY, VOL. VIII. 71
488
PROF. O. ELLIOT SMITH OX THE MOKPHOLOOT OF THE
this part of the paratermiual body presents a relationship to the hippocampus which
is analoo-ous (topographically) to the fimbria in the Mammalian brain.
That the commissiira aherrans arises from the hippocampus is an undoubted fact ; for,
as one of the coronal sections (fig. 10) in the introductory part of this memoir shows, its
relationship to the caudal portion of the hippocampus is exactly analogous to that of the
dorsal commissure to the same formation iurther forward. Therefore Rabl-Riickhard
had some justification for calling this structure the commissura fornicis, especially as it
springs from that portion of the hippocampus which persists in the Eutherian brain.
But this name is objectionable, in that the designation '\fornicis " is equally applicable
to the commissural fibres which spring h-om the cephalic ])ortion of the hippocampus
and from the commissimi dorsaUs. Movc-over, this commissure has a peculiarly
aberrant course in comparison with the usual course of the fornix — and hence it
deserves a special name. The term " commissura aberrcms \liippocampt\ '' seemed to
me to fulfil these requirements.
Fig. 22.
This is diagrammatic, the relations of the various parts oot being drawn to scale. The true proportions
are shown in figs. 1 and 2, and the seiies of conin^il sections of Splienndon.
The explanation of the peculiar caudal extension of the paratermiual l)ody is a matter
of extreme difficulty, and one which needs a much fullei- knowledge of its beliaviour in
certain other Vertel)rate types than I at present possess. The question is complicated by
the fact that this caudal extension seems in some way to be associated with the presence
of the commissura aherrans. So far as I am a^^■are. it occurs in all Reptilian^brains in
which the latter commissure is found, and is lacking in the Chelonia and other Reptiles
in which the aberrant commissxtre' is not developed. One might be led to believe that
the coiiiiit.is.sura aherrans is merely a part of the commissura dorsalis (for both are
hippocampal) which had in some way become separated from the parent commissui-e and
in its wandering caudalward had drawn back on each side a prolongation of its matrix
of paratermiual body. But the features of the region in the adult render such an
CEEEBRAL CO.MMISSURES IN THE VEKTEBKATA. 489
explanation improbable, nor does the liistory of its developfnent in Sphenodon lend any
support whatever to such an hypothesis. This explanation would jilso involve the
necessity of believing that its fusion with the dorso-anterior extremity of the optic
tlialamus is secondary and acquired, and not the persistence of a more primitive
condition. Certain other aspects of the in'oblcni will be discussed later.
But in the meantime tliere are certain facts which must be kept in mind in attempting
to explain this phenomenon.
The ])araterminal body is a ganglionic mass interposed between tlie bnlbus olfactorias
on the one hand and the hippocampus on the other. The only libre-tracts which
establish any connections with it are derived from these two sources. (For tlie sake of
simplicity 1 do not specifically mention the connection with tlie cortex of the
tuberculum olfactorium, which is merely an indirect path from the olfactory bvilb.)
The morpliological relations of the hipjiocampus and paraterminal body are thus most
intimate ; the latter not only affords a path for tlie hbres (fornix) of the former, but it
forms the bond of union of the liippocanipus with the olfactory region of the brain, of
Avhich physiologically the hippocampus is merely an api)endage. Not only so, but a
study of the relations of the jjrimordtiim hippocampi to the paraterminal body in the
Ichthyopsida lends support to the view that the hippocampus may be nun'ely the
specialized upper part of the primitive paraterminal body.
Whether this be so or not, there can be no doubt that the most intimate relationship
exists between the two bodies, and any system of subdividing the cerebral hemisphere
which separates hipjjocampus and paraterminal body commits a fundamental error which
must render it nugatory as a natural subdivision.
The evidence afforded by comparative anatomy in the Vertebrate series points to the
region upon the dorsal margin of the paraterminal body as the original site of the hippo-
campus, and it is conceivable that, as in the process of evolution the hippocampus grows
backward with the hemisphere it may, in certain cases, cany with it an extension of the
associated paraterminal body.
The attempt to offer an explanation of the significance of the commissiira ahcrruu>i will
involve the discussion of some of the most difficult problems in the whole field of cerebral
morphology. For, before it is jDossible to apjirecijite the factors which seem to have
determined the development of this commissure in the place where it is found, it will
be necessary to consider In-iefly the involved questions : (i) of the relationship of the
optic thalamus to the cerebral hemisphere ; and (ii) the nature of the choroid plexus of
the lateral ventricle.
I shall discuss the second problem first, and that in relation to the Monotreme brain,
because, while the latter presents all the simplicity of the Sauropsidian condition, the
neighbouring r(>gions have attained to the full degree of Mammalian difl'ereuliation,
wherefore it is possible to recognize them with certainty.
The epithelial roof of the forebrain extends in Echidna (as in Oi-nithorhynchiis, vide
siqyra) from the commissura superior posteriorly to the antero-superior lip of the
thickened upper extemity of the lamina terminalis (see fig. 23). The relations of this
roof in the region of the recessus superior have already been considered.
71*
490
PRO!'. G. ELLIOT SMITH ON THE MOKPHOLOGY OF THE
c.p
c.d. I y
f.M.
Semi-schematic representation of the roof of the forehrain of Echidna viewed in sagittal section.
The relations of that portion of the roof which intervenes
between the commissura superior and the line x-y (iig. 23) conform
to the usual plan, being attached on each side to the taenia thalami.
The accompanying crude scheme indicates this usxxal mode of
attachment (fig. 2i).
Behind the cominlssura dorsalis a large complex fold of the
epithelial roof extends on each side through the foramen of Monro
to form the choroid plexus of the lateral ventricle. If a coronal
section 1)e made through the situation of the foramen of Monro,
the exact arrangement of this fold will lie intelligible (fig. 25) *.
Fis. 24.
dors.
v.jn.
vM
Coronal section of the brain of a foetal Eclddnu, x '20, passing through the foramen of ilouro.
In the mesial j)lane it is thrown into a complicated series of folds, which form the walls
of the paraphysis, and on each side the epithelial fold, forming the roof of the foramen
of Monro, projects into the lateral ventricle, and is complexly plicated to form the
* 1 have represented a section of a foetal brain, because it is ■well-nigh impossible to keep these delicate epithelial
folds intact in the adult.
CEEEBRAL COMMISSURES IX THE VEKTEBKATA.
491
characteristic choroid plexus. The distal attachment of this fold is to the ventral lip
of the fascia dentata.
If, however, a section (fig. 26) he made Ijehind the situation marked x-y in the figure
representing the rooL' (Hg. 23), the typical thalamic roof is found to have no connection
with the lateral plexvis. The dorsal attachment of the latter is still to the margin of the
lascia dentata, while its ventral extremity is inserted into the " tcenia chorioidea " (His),
ter
Section piissing- buliiiid the foramen ol Monro.
i. e. the stria terminalis. In the whole of the caudal extent of the ventricle the choroid
plexus presents similar attachments to the margin of the fascia dentata and the stria
terminalis (fig. 2G).
Fig. i;7. Fig. 2?^.
Fig. 27 shows the ])rimitive relation oi'
the optic thalamus to the posterior
lip of the cerebral hcmisplicre.
The backwai-d extension of the
hemisphere will stretch the attach-
ment to the optic thalamus at *,
and reduce it to the condition of a
thin epithelial lamina as in the
next figure.
jPcy.
Schemata to explain how the posterior choroidal fold may arise.
Now, although in the whole of its extent the epithelial layer of the choroid plexus
presents uniform features, it is difficult to admit a common origin for the whole
492 PEOF. G. ELLIOT SMITH OX THE MORPHOLOGY OF THE
structure. With regard to that part of tlie plexus which is found in the region of the
foramen of Monro, there can be little doubt of its origin from the primitive roof of the
forebrain. Its developmental history clearly shows that this part of the choroid plexus
was never anything else than a simple epithelial layer, and a comparative study of its
behaviour in different Vertebrates indicates that it ought to be regarded as a derivative of
the primitive roof, or " Beckplatte" of the forebrain. To speak of it as a portion of the
cerebral hemisphere which has undergone a retrograde change, or has retained its
primitive epithelial simplicity, is to make a gratuitous assumption, which receives no
confirmation from the facts of embryology. But the case is very different with that
portion of the choroid plexus which is not directly connected with the roof of the fore-
brain, but is attached to the stria terminalis. There is no evidence to show that this
portion is derived from the roof, and all the facts of development point to the conclusion
that its proximal attachment to the optic thalamus is a primitive and not a secondarily
acquired relation. Such being the case, the caudal extension of the epithelial choroidal
fold in the Mammalian hemisphere would appear to be derived from a stretching of the
attachment of the labium caiidale of the cerebral hemisphere to the optic thalamus. As
a result of this the connecting band becomes reduced to an epithelial lamina, which
becomes invaginated and folded by an extension backward of the choroidal folding which
begins farther forward in the region of the foramen of Monro. This hypothetical
explanation of the derivation of the posterior choroidal fold is grapliically demonstrated
in the accompanying schemata (figs. 27 & 28, p. 491).
No full history of the mode of development of the lateral choroid plexus has ever been
given, so far as I am aware. Most recent writers admit the undoubted origin of its
cephalic part from the roof, and either ignore the more caudal portion or frankly admit
their ignorance of its morphogenesis. The hypothesis which I have suggested to
explain the possible mode of production of its caudal portion receives some su.pport from
comparative studies, and is advanced on the present occasion because the examination of
the plexus in Sphenodon seems to have an important bearing upon the question at issue.
In Sphenodon the two attenuated portions of the mesial wall of the hemisphere —
using this expression in its ixsual but inexact sense — are separated the one from the
other by the commissiira aberrans. At the situation of the latter the ventral margin of
the paraterminal body becomes linked to the corresponding part of the other hemisphere,
and thus a bridge is formed which effectvially limits the choroidal fold. Behind the
situation of this commissure the process of attenuation of the connecting band between
the hemisphere and thalamus may be readily recognized. It is not difficult to conceive
how, in a brain such as that of a Mammal in which there is no barrier in the shape of a
conimissitra aberrans, the epithelial invagination which begins in the region of the
"porta" (foramen of Monro) may extend in the caudal direction and involve the thin
portion of the mesial wall of the hemisphere, which has a different mode of origin. This
hypothesis might explain the jieculiar attachments of the post-portal part of the choroid
plexus.
The existence in the brain of Spheuodon of a thin mesial hemisjihere-wall, which has
luidoubtedly become attenuated secondarily and forms a part of the true wall of the
hemisphere-vesicle linking the caudal part of the hijjpocampus to the optic thalamus,
CEREBRAL COINIMISSURES IX THE VERTEBRATA.
493
naturally suggests the possil)ility that the caudal or post-portal part of tlic choroidal fold
in the Mammal, which also links the hippocampus to the optic thalamus (tig. 27), may
have arisen in a similar manner.
The relations of the optic thalamus to the cerebral hemisphere have Ikhmi elucidated
within recent years hy the researches of His, Lachi, and Hoclistetter. 1 wish briefly to
call attention here to certain features in tliis relationship which these writers have
not emphasized, and which afford the })ro]mble explanation of the presence of the
commisswa aberrant in such a peculiar locality.
From the accompanying rough sketch, which is slightly modified from one of Hiss
drawings *, it will be conceivable that the optic thalamus extending forward as the
Fio;. 29.
teL
Rough sketch of a mesial view of the forebraiii of a human embryo uf 10'2 mm. neck-length.
Mainly after His.
lateral wall of the third ventricle, becomes directly continuous on the ventral side with
the corpus striatum at /3, whereas its dorsal part becomes continuous at a, not with the
ganglionic mass, but with the mantle (using this term in the original sense of lleichert).
The corpus striatum in the floor of the hemisphere may be seen extending through the
widely ojien foramen of jMonro at /3 to fuse witli the lower part of the optic thalamus.
But this figure is introduced more especially to demonstrate the relationship which
exists between the dorsal portion of the optic thalamus and the mantle of the hemisphere
at «. Tlie significance of this relationship will be rmderstood when it is remembered tliat
it is this '• parathalamic " area of the mantle which becomes modified to form the caudal
portion of the liippocampus, au'd that it is the connecting band between the hippocampus
and the thalamus which ])ecomes attenuated (according to the hypothesis advanced earlier
in this account) and is then converted into the caudal part of the lateral choroid plexus.
Bearing in mind these primitive relations of the thalamus to the eerebi-al hemisphere,
which obtaiu at some epoch in the development of all Vertebrates and persist into adult
lile in the lehthyopsida, the course of the fil)res of the commissnra aOerrans becomes
intelligible. For it springs from the parathalamic mantle, wliich becomes the hippo-
* " Formentwick. des llenschl. Vorderhims." AbhaniU. d. koiiigl. Siiclis. Uesell. d. Wissensth., Bd. xv. 2\o. viii.
1889.
494
PROF. G. ELLIOT SMITH OX THE :\rOEPHOLOGT OF THE
campus at a time before the attemiation of the hippocampo-thalamic band has begim,
and when, therefore, there is a close connection between not only the hippocampus and
the thalamus, but also the roof of the forebraiu and both of these structures.
That commissural fibres of the hippocampus should avail themselves of this direct
path across the roof of the forebrain is what one would naturally expect. The puzzling
feature of the problem is that the corresponding fibres in the Chclonia and Crocodilia
do not make use of this route, but pursue the more circuitous path via the dorsal
commissure.
Fig. 30.
O.C.
Sketch of the dorsal aspect of the left half of the forebrain oi u human foetus, drawn from His"s tig. 7, Taf. i. {Joe.
eit.), with certain moditications suggested by Hochstetters researches (' Zur Entwick. d. Gehirns,' 18!)8).
Eoof of cerebral hemisphere dissected away.
Summing up the conclusions of the foregoing arguments, it seems that the commisstira
(iberrans is a bundle of fibres derived from the caiulal portion of the hippocampus, and
therefore homologous (in part) with the psalterium of the Mammalia. But its behaviour
presents a marked contrast to that of the Mammalia : for, instead of pursuing an
extensive forward course to cross over in the lamina terminaUs, it avails itself of the
primitive direct connection between the caudal lip of the cerebral hemisphere and the
optic thalamus, and in this way reaches the roof of the third ventricle dLrectly. This com-
missure is found only in Sphenodon and the Lacertilia (and possibly in certain Ophidia)
as a separate bundle. From examination of the brain in a number of Amphibia and
related forms, it seems certain that an analogous commissui'c is found there also, but
that instead of pursuing an independent coiu-sc it crosses the roof of the third ventricle
alongside the fibres of the commisstira superior of Osborn. This phenomenon is readily
understood when it is remembered that the thalamic region is very short in the
Ichthyopsida, and the sitiiations of the commissnra uberrans and coramissura s/qjer/or
would be closely approximated. The fact that the so-called commissuro superioi- of
Amphibia derives fibres from the caudal extremity of tlie cerebral hemisphere is well
known, and has been recorded by almost all writers upon the Amphibian brain, so that
it is unnecessary for me to adduce evidence to prove this matter of common knowledge.
CEEEBEAL COIMMISSUEES IX THE TERTEBKATA. 495
But in the light of the above recorded observations upon tlie contmissuru abcrrat/s, it
seems more than probable that these hemispheral fibres of the commissura superior in
the Tchthyopsida represent the aberrant hippocampal comiiiissurc of the Lacertilia and
Rhynchocephalia, and do not strictly belong to the habenular or superior commissure.
This interesting fact explains the otherwise inexplicable confusion of commissura
superior and coMmissura aherrans in the mind of Osl)orn (^lorph. Jahrb., Bd. xii. pt. i.).
Up to this point I have altogether refrained from speaking of the conditions which
prevail in the Ichthyoj)sida, because the crucial problems witli whicli this essay is more
especially concerned find their readiest solution in tlio study of the Amniota. For, once
a trustworthy Imsis for comparison of the Sauropsida and Mammalia is established, and
the factors which are at play in the specialization of the various types, which Reptiles,
Birds, and I\[ammals afford, are appreciated, the key to the solution of the problems of
Ichythopsidau morphology is provided. The comparison of a simple reptilian Ijrain, such
as the Cheioaian type with the Amphibian, is a much simpler and more obvious process
than the comparison of the reptilian brain with that, say, of a placental mammal.
In the Amphibia all gradations of brain-form are found, from a type which is
practically identical with that of Protopterus or Lepidosireii, and differs relatively
slightly from that of Ammocwfes, up to a form which readily admits of comparison
Avith the Chelonian type.
It is quite lumccessary to enter here into any long account o£ the features of the Ichthyopsidan types
of cerebral hemisphere. So mueli has been written upon this subject within recent years, and the
present state of knowledge has so recently been succinctly stated by Gaupp (' Auat. d. Frosches/ 1897)
and Johnston (Journ. Coinp. Neurol. 1900, 1901, and 1902) among many others, that the general facts
concerning these types are now matters of common knowledge. But the question of the interpretation
of this huge collection of facts is a very different matter, and the utmost confusion reigns in the field of
Ichthyopsidan neurology. It will be sufficient, therefore, if I indicate, in the light of the conclusions
drawn from the comparison of Sauropsida aud Mammalia, the main sources of fallacy in the current
interpretations of the Ichthyopsidan cerebral hemisphere, aud point out what seems to me the true
basis for accurate comparison with higher forms.
In the history of the growth of anatomical science, more especially as regards the Yertebrata, it so
happens that most of the names of the various organs and of the subsidiary parts of the body were
originally bestowed upon the human subject : later on these names have been applied with more or less
accuracy to those parts in other animals which correspond, or are supposed to correspond, to tiie
structures so-named in the human body. In the case of the brain, for example, comparisons were
naturally made with the corresponding organ in other Mammals, and applied with doubtful results to
Keptiles and Birds. In the case of the Amphibia, the data which were available for the accurate
application of the terms of Human Anatomy have of necessity filtered through the uncertain channels
of the lowlier Mammalian and Sauropsidan neurological knowledge, and in the process have become
tainted and vitiated with all the inaccuracies which I have already ex|)osed in speaking of the reptilian
brain. Added to all the inaccuracy which necessarily results from such a process, there is the further
fertile source of error which results from futile attempts at the comparison of the Reptile and Amphibian.
Under such circumstances, it is no matter for surprise to find the present state of knowledge concerning
the morphology of the brain in the Amphibia almost chaotic.
One example may suffice to show that this latter remark is no mere figure of speech iu regard to
recent work in this particular field of research.
SECOND SERIES.— ZOOLOGr, VOL. VIII. 72
496
PEOF. G. ELLIOT SMITH ON THE MOEPHOLOGT OF THE
One of the most recent writers upon the braiu in the Ampliibia (Pedro Ramon y Cajal, " L'Encephale
des Amphibiens," Bibliographie Anatomique, T. iv. 1896, p. 234.) described as basal ganglion the lower
half of the lateral (fig. 31, ^) and mesial (/5) walls together with the connecting band (7) which forms
the basal angle of the hemisphere in the Frog's brain (in a coronal section in front of the lamina
terminalis). The dorsal half of the mesial wall (S) he called " si-pUim," meaning to imply that it is the
homologue of the •' septum lucidum" of Human Anatomy.
Fiir. 31.
A still more recent writer (Ernst Gaupp, ' Ecker's uud Wiedersheim's Anatomic des Frosches/ 2ud
part, erste Halfte, 1897) described as •' basal ganglion " only that part of Eamon y CajaFs so-called
o-ano-lion which forms part of the lateral wall (a) ; and the mesial portion (/3) he called '•' septum.'' To
complicate matters further, Ramon's "septum" {h) Gaupp described as " A mmonshorn," i. e. hippo-
campus. Thus the only point upon which these two writers were in agreement is that the lower half of
the lateral wall (a.) is " basal ganglion." The brain of the Dipnoi is admittedly distinctly Amphibian,
and as regards the cerebral hemisphere the resemblance is very striking. It is interesting to note that
in Protopterus Rudolf Burckhardt (' Das Centralnervensystem von Protopterus annectens,' Berlin,
1892) regarded the upper part of the mesial wail of the hemisphere (8) (Gaupp's " hippocampus " and
Ramon's "septum") as the " basal ganglion" {vide Burckhardt, op. cit., Taf. 3. figs. 22 & 23). The
" Ammomhorn," or hippocampus, he located at the base of the hemisphere, where Ramon placed the
"basal ganglion " and Gaupp the " septum" and " basal ganglion" [vide Taf. 3. figs. 20 & 21). Ramon
(who in his most recent contribution [op. cit.) contradicted himself upon several occasions) added to
the intricacy of the present tangle by calling the mesial portion of his " basal ganglion," i. e. Gaupp's
"septum," the epistriatum (/S). The reason given by Ramon for this use of the latter term is that the
reo'iou /3 corresponds to the area so-named in reptiles by Ediuger. The enormity of the error involved
in this amazing statement will be appreciated, when I mention that the name " epistriatum " was applied
by Edinger to a portion of the lateral wall of the hemisphere in reptiles which is situated immediately
above the true basal ganglion. The structure iu question is the ventral part of the " pallium." It is
inconceivable how Ramon can see in the lower portion of the mesial wall, moreover, iu a region which
he himself calls " basal ganglion," a homologue of Ediuger''s epistriatum. In the same memoir Ramon y
Cajal describes as the homologue of the hippocampus the same region which he called " septum "
elsewhere.
It would be possible to multiply examples almost without limit to show that even at the present time
the problems of the morphology of the cerebral hemisphere in the Amphibia are iu the apparently
hopeless state of confusion, of which this single striking instance is unfortunately typical.
In view of the above statements it would clearly be mere waste of energy seriously to discuss further
CEREBRAL (O.MMI.sSLKES IX THE VERTEBRATA.
497
the views of Ramon y Cajal and ISurckliardt upon the morphological aspect of the Amphibian brain, for
their conclusions arc obviously a tissue of guesswork.
The account wliich Ganpp, profiting by the blunders of his predecessors, has been able to give is
clearly based upon the latest memoir of Ediuger {op. cif. 189(5) upon the forebrain ol' reptiles, which I
have discussed in an earlier jiart of this memoir. It is only natural under such circumstances that
Gaupp's conclusions regarding the Ani|)liii)ian lieinisphere are marred by the same errors, which I have
already e.\posed (ride supra) in Edinger's memoir. But in addition Gaupp commits some additional
mistakes which create great confusion. J might quote one example. The reptilian hemisphere presents
a basal notch which corresponds to the vallectdn Si//i:i/ of the mammalian brain. Edinsjer calls it the
fovea limbica. Gaupp adoj)ts this term, but applies it to the groove which separates the bulbus
olfactorius from the rest of the hemisphere !
A coroual section in the plane of tlie cerebral commissuves affords jievhaps the mo.st
satisfactory basis npon which to iustitnte comparisons witli other forms, and the Fro"-
may be adopted as a type, not because it is the most suitable Amphibian for comparison,
but simply because it is best known. The accompanying diaprram (fig. 32) represents
schematically the appearance of a coronal section througli the brain of the Erog in the
plane x-t/ of the explanatory diagram. There are two commissures — dorsal and ventral —
in the lamina terminalis as in the E-eptilia. The commissura dorsalis is disti'ibuted in the
dorso-mesial area of the section, whiclt consists of a mass of grey substance containing
diffusely scattered nerve-cells [hip.]. Tiie epithelial roof of the recessus superior is
attached on each side to a mass of grey substance (jx'ra.) through which the flln-es of
Fijr. 3:^.
Fifr. 33.
c.c/.
^pard..
recs.
vM
Transverse section throusL the brain of a Frog (Ihina) passing through the two cerebral commissures
in the plane .r-»/, fig. 33.
the commissura dorsaUs pass to reach the area la])elled hip. The relative disposition of
these various structures is so obviously identical with that which has been seen to
characterize the reptilian l)raiu tbat, in the absence of any reasons to the contrary, I
do not hesitate to call the dorsal area the homologue of the hippocampus, its commissure
the hippocampal commissure, and the mass labelled ^j«ra. the ^' paratermlnal body.''
Upon histological evidence alone one would never have suggested the homology of the
■•-)*
498 PEOF. G. ELLIOT SMITH ON THE MORPHOLOGY OF THE
dorsal mass to the hii^jDOcampus, because its cells are diffusely scattered ; but upon
morphological grounds the homology cannot be questioned, and the condition of the
hippocampus in Emys and certain other Chelonia indicates that a compact column of
cells is not a necessary feature in the j)rimitive hippocampus.
The morphological jilan of the mesial wall of the cerebral hemisphere is so constant in
the higher Vertebratn, in Reptiles, Birds, Monotremes, and Marsupials, and even in the
placental Mammals (though disguised by revolutionary changes associated with the
advent of the corpus callosum), that this plan must be regarded as the common property
of the Vertebrata. The relations of tiie paraterminal body to the lamina terminalis and
olfactory bulb places the recognition of this body beyond question ; and the mass which
rests upon its upper Ijorder and exhibits traces of a cortical structure (according
to Nakagawa, Journal of Morphology, vol. i.), unless the Amphibian tyj^e of brain
conforms to some plan utterly different from that which all the Amniota possess in
common, must be the hippocampus.
Osborn and all writers who have come after him have assimied as a postulate that
the dorsal commissure and the area from which it springs are the undoubted repre-
sentatives of the topographically analogous structures in the E-ejitile. There is no other
alternative which can for a moment be entertained, and therefore the amorphous dorsal
mass in the mesial wall of the Amphibian hemisphere must be regarded as unquestionably
the hippocampus. The question of interpretation once settled, the enormous mass of
data concerning this type of brain may be readily interpreted. Three schematic repre-
sentations of coronal sections in the precommissural region (figs. 34, 35, 36) in foetal
Ornithorhynehus (it might equally be any other Mammalian foetus — vide His, E-etzius,
Hochstetter), an adult Reptile and Amphibian, will make the general j)lan clear.
It is quite unnecessai-y, with this key to the interpretation of the Ichthyopsidan brain,
to give an account of all the variations of type which this heterogeneous grou^J of
Vertebrates includes. The reader who adopts this suggestion will find ample con-
firmation of the constancy of the plan, which the study of the Amniota has revealed,
throughout the Vertebrata. The type of brain in the Anura is fuUy described by
Gaupp, who adopts, with some relatively slight inaccuracies, the interpretation which
T have put forward hei'e {vide Gaupp, oji. cit.).
The writings of Osborn (Morph. Jahrb., Bd. xii., and Journal of Morphology, vol. ii.),
and a large number of memoirs by other American writers, of which Kingsbury's
account of the brain of Necturus (Journ. Comp. Neurology, 1895) may be mentioned,
as well as Burckhardt's work on Triton and Ichthyophis (Zeitsch. f. wiss. Zool. lii.
1891), clearly show the manner in which the type becomes simplified and more
fish-like in the Urodeles, producing a condition closely resembling that found in the
Dipnoi *.
Through the kindness of Mr. E. I. Bles, of Cambridge, I have been able to study the
condition of the brain in a great many genera of the Urodela, and compare them with the
* Vide FuUiquet, '• Recherches sur lo Cerveau du Protopterus,'' Eeciieil Zool. Suisse, vol, iii. 18SG : and
Burckhardt, ' Das Centralnervensystem von Protopterus,' Berlin, 1892).
CEREBRAL COMMISSURES I.V THE VERTEBRATA.
499
Anura and Lepkloslren, for the chance of examining which last T have to thank
Mr. Graham Kerr, It Avoiilcl be impossible adequately to consider these forms here
without long descriptions and a large number of figures, and it would add com-
paratively little to the force of the above remarks. The general plan is identical with
that briefly indicated in the remarks on tlie Frog's brain ; and if the fallacies to which I
Fit;. :J4.
ne
pam.
A coronal section of the cerebral hemisphere of a foetal OrnithorhyiiclMs
in the precommissural region.
Fis. 3.5.
pans.
A corresponding section — adult Hi/drosintnis.
Fig. 36.
pans..
A corresponding section — Rana.
have already called attention in the works of Ramon y Cajal, Burckhardt, and to a lesser
degree of Gaupp, be borne in mind, the reader can have no difficulty in interpreting
arigbt the above-mentioned accounts of Osborn, Kingsbury, Burckhardt, and Fulliquet,
and the innumerable other recent accounts of the Ichthyopsidan brain.
500
MORPHOLOGY OF CEKEBRAL COMMlSSTHES IN YEBTEBKATA.
The vexed questions of the morphology of the piscine types of brain hardly come
within the scope of this memoir. But the investigator who studies the brain of
Animocoetes or Pefroniyzon, in the light of the information which a comparison of the
Amniote and Urodele types of brain has yielded, cannot ha-\'e much difficulty in seeing
in the jjrevailing conditions in the Cyclostome an extremely simple and archaic form of
the common vertebrate type. This is well shown by the important memoirs of Studuicka
(" Beitrage zur Anatomic und Entwickelungsgescliichte des Vorderhirns der Cranioten,"
Sitzungsberichte d. konigl.-bohmisch. Gesells. d. AVissensch., Math.-j)hys. Classe, 1895)
and Johnston ("The Brain of Fetromyzon," Joiu*n. Comp. Neurology, March 1902.).
In conclusion, I must acknowledge with the deejoest gratitude the liberal help so
generously afforded me by Pi'ofessor Howes, who not only obtained for me the material
for the research, but also devoted a vast amount of time and energy in helping me to
accomplish this task.
J/ist of Abh'eviations used in the Diagrams.
U.S., aquaeductus Sylvii.
alv., alveus.
b.o., bulbus olfactorius.
c.a., commissura alx^rrans.
c.d., ,, dorsalis.
c.h., „ liabeuulfe (superior).
C.J)., „ posterior.
C.V., „ veutralis.
c.f., columna fornicis.
ce., cerebral hemisphere.
cbl., cerebellum.
dors., dorsal sac (part of epithelial roof of the
third ventricle).
ep., epiphysis.
fasc, fasciculus raarginalis.
f.d., fascia deutata.
/.//., fissura hippocampi.
f.M., forameu Mouroi.
hip., hippocampus.
hyp., hypophysis.
I.V-, lobus pyriformis.
l.t., lamiua tcrmiualis.
mam., corpus mammillare.
m.o., medulla oblougata.
ne., ucopallium.
M.O., nervus olfactorius.
n.IV., uervus trochlearis.
(l.t., tlial.nmus opticus.
opt., tractus opticus.
/;., jiallium.
pcn\, parapliysis.
para., corpus paraterminale.
p.o., peduuculus olfactorius.
p.s., parietal stalk.
p.c.f., posterior choroidal fold.
plx.I., plexus chorioideus lateralis.
plx.IL, lamiua chorioidea.
ph. III., plexus chorioideus ventric. tertii.
plx.IV., ,, „ „ quarti.
q., corpora quadrigemina.
red., recessus iutercommissuralis.
rec.s., recessus superior.
ret., retina.
S.I., sulcus limitaus.
s.gr., stratum grauulosum fasciae dentatse.
St., corpus striatum.
ter., stria tcrmiualis.
tel., tela, or epithelial roof.
t.o., tuberculum olfactorium.
tuli., tuber cinereum.
V.I., veutriculus lateralis.
v.III., „ tertius.
v.IV., „ quartus.
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])ee(?mljer 1903.
I N D E X.
[Note. — Synonyms and Native Names are printed in itaJics. A star is added to names which appear
to be used for tlie first time.]
Aard-vart, 331.
Acanthephyra, A. Mihie-Eilir.,
44'.t.
brachytelsonis. S/nnre Bate,
450.
purpurea, A. Mi/ne-Eihr.. men-
tioned, 45:^.
sica, Spence Bate, 4-1 i), 4." 2.
Acanthocephala surat;i, Burni., ~'2.
Acanthophrys, A. Milne-Edf., men-
tioned, 37.
ucuhdtiis, A. llilne-Edw., .38.
cristimauus, A. Milne-Edir.,
mentioned, 37.
verrucosipes, Miirs, mentioned,
30.
Aceipitrcs, 20(5.
Accipitrinsp, l'il7.
Aebiciis alllnis, .Vlfrs, 33.
Aelielous grauulatus, //. MUur-Eihr..
'11.
var. unispinosus, Miers,
•J\.
Aera-pliia perspicillata, Fuljr., 73.
Acrocoris perarmata*, W. F. Kirhij,
71', 73.
Acriisoma, I'erti/, GS.
Schriebersii, read Sehreiliersii,
I'ertt/, OS.
Aeta'a calculusa, //. Milne-Eilw., '2,
.3. S.
granulata, And., mentioned,
8, 9.
var. carcharias, White,
10; mentioned. 2, 9.
hvstri.v, Mlcrs, 1 0 ; mentioned,
2, 11.
nodulosa, Adams (|- White, men-
tioned, 11.
SECOND SERIES. — ZOOLOGT,
Aetaea Peronii, var. squamosa, IJeii-
der.^on, 10.
Iliippellii, fCrauss, 7 ; mention-
ed, 2, 8.
.Vetnecia, Dana, 130 ; mentioned, 122,
128.
aacldandiir, G. M. Thomson,
120.
encliroa, Dana, 130 ; mentioned,
101, 102, 103, 131, l.")2.
opihensis *, Chilton, K>2 : men-
tioned, 101, 103, 152.
Aotoniscus, Hiujer, mentioned, 122.
-Vctumnus, Dana, mentioned, 18.
pidclier, Ortmann, 15.
setifer, De Ilaau, 10.
Adapis, Cuv., 424.
-Eluroidea, 398, 410.
^Elnropus melauoleucus, A. Milne-
Edn'ardf, On the Affinities of,
by E. Kay Lankestcr, 1(13-172.
, Cuv., carnassial teeth of, Kiii.
^Elurus, Cuv., carnassial teeth of,
163, 16(5-172.
fulgens, Cuv., mentioned, 172,
354.
.Es, Boie, 203.
Affinities of the Lemurs, 417.
.\ilurus, Cav., 355.
fulgens, Cav., 172, 354.
-■Viptasia laeerata, Andn.'i, mentioned,
309.
tagotes, DiicJi. 4' Midi., men-
tioned, 309.
Alaudidaj, 252.
Albatross, 18<S.
Alcedinidtc, 245, 256, 257, 269.
Alcid.-e, 237, 238, 240, 263, 2(i9.
Alcidion sesnotatum *, ]Vatcrli., 7(!.
VOL. VIII.
Alectoromorphine Legion, summary
of, 173, 240.
Alepadinae, 277.
Alepas, Sander- Ranij, 277.
Belli *, Gruvel, 278 ; mentioned,
280, 282, 283, 284, 285, 287,
294.
cornuta, Daruln, mentioned,
2,s7.
indica *, Gruvd, 2^4 ; men-
tioned, 278, 287, 288, 294.
japonica. Auric, mentioned,
2S7.
Lankesteri, Gruvd, mentioned.
279, 280, 282, 283, 285, 287.
microstoma *, Gruvd, 282 :
mentioned, 278, 284, 286,
287, 294.
minuta, I'hilijtpi, mentioned,
2S7.
parasita ?. Sander-Ramj, men-
tioned, 287.
pedunculata, llotk, mentioned,
287.
quadrata, Auriv., mentioned,
2^7.
tubulosa ■?, Quay ij- Gaimard,
mentioned, 287.
■Vlicia, Johnson, mentioned, 298.
Alligator, brain of, 481.
Alope, White, 440.
])alpalis. White, 440 ; men-
tioned, 453.
.\.lpheida?, 43().
Alpheus, Fahr., 436.
chinigrious, A. Mdne-Kdw.,
mentioned, 437.
novaj-zealandia;, Miers, 438 ;
mentioned, 433, 437.
73
>02
INDEX.
Alpheus socialis, Ildler. V-W ; meu-
tioiied, 453.
Ammoeoetes, brain of, 4!)5, 500.
Ampelidas, 254.
Anipbicvoii, Lart., mentioned, 171.
Aiiipliitrite yladiatnr, L)e Haaii, 'J.\.
Anaptomorphus, Cope, 424.
Anas, Luiii., 203.
Anaspidre', 277.
.•Vnastomus, liunn., 213.
oscitans, intestinal tract of, 1'.I7.
Auatidffi, 270.
Andigeiia Baiiloni, Wagl., 252, 2G0.
Anolis fusco-auratus, d'Orb., ,53.
" Anse coliciue," 170.
" moyenue," 170.
Anseranas, Le.-s., 215.
melanolcuca. Less., intestinal
tract of, 201, 202, 203.
Anserida-!, 201, 215.
Anseriformcs 173, 201, 204, 213,
268.
Anteaters, 3'J3, 402, 425, 430.
" Anterior limiling sulcus,'' 4i)7.
mesenteric or duodenal tract,
177.
Anthropoides, Vieill., 223.
Anthropopithecus, Blaiiw., 374, 405 ;
cerebellum of, 431.
Antichira dichroa, Maiinerlwim, men-
tioned, 70.
Anyphoena, StiiuL, 07.
Quflchii, Pocock, 07.
Anypha>nida>, 07.
Apes, brain of, 396, 421 ; olfactory
peduncle of, 328.
Apidie, 72.
Apocentric type, 178.
Aptenodytes Pennanti, Gray, intes-
tinal tract of, 180, 187.
Apteryx, Sliaw, intestines in, 182,
183.
Mantelii, Bartlitt, intestinal
tract in, 183.
Aquila audax, Lath., 208.
chrysaetus, Linn., 208.
Verreauxi, Lesson, 208.
AquilinuD, 208.
Ara, Briss., 243.
ararauna, Liun., 243.
Araclniida and Myriopoda of Mount
Roraima in liritish (juiana, bv
U. I. Pocock, 04-71.
Araniidic, 241.
Aramides ypecaha, ViiUL, 222.
Araminic, 223, 208.
Araraus, VieUl, 224, 242.
scolopaceus, GmeJ., intestinal
tract of, 223, 224.
Aranea3, 00.
Araneus, Linn., 08.
audax, Blachw., 08.
Aroania gracilipes, Bell, 28.
Archeceutric type, Palamedea cornuta,
lAnn., intestinal tract in, as, 173,
175,178.
Arctoidea, 398, 411.
" Arcuate sulcus,'' 414.
Ardea candidissima, Gmel., intestinal
tract of, 195, I'JG.
Ardeae, 195, 213.
ArdeidEe, 195, 197, 213, 263, 208.
ArgiopidiB, 68.
Argus, Tanm., 219.
giganteus, Teinni., intestinal
tract of, 218.
Argus Pheasant, intestine of, 175.
Arraadillidium, Brandt, 142; men-
tioned, 103, ftnotel28.
vnlgare, Lntr.. 142; mentioned,
101, 102.
■ var. immaculata, Budde-
Lund, 143.
var. variegata, Budde-
Lrnnd, 143.
Armadilliidte, 141.
Armadillo, Budde-Liind, 143 ; men-
tioned, 10], 103, 149.
ambitiosus, Budde-Lnnd. 144 ;
mentioned, 102, 145, 147,
152.
AiicHandicus, Budde-Lund, 148.
l)an;E, Heller, 145 ; mentioned,
144, 140.
Haniiltoni*, Chilton, 148 ; men-
tioned, 144.
iHCOnspicuu.'i, Miers, 145, 140.
Macmahoni*, Chilton, 149; men-
tioned, 149, 150, 152.
monoliniis, Dana, 148; men-
tioned, 144.
regulosus, read lugulosus.
rugulosus, HJiers, 147 ; men-
tioned, 101, 102, 144, 148,
152.
speciosus, Dana, 140; mentioned,
102, 144, 152.
spinosus, Dana, 150; mentioned,
144.
vulgaris. Lair., 142.
Arniadillouiscus, Ulianin. mentioned,
13i_>.
A rmadilloniscus f »f A*-oa,Eudde-Lund,
130.
Armadillos, 403.
.Artaiuidae, 254.
Asellus aquaticus, Leach, mentioned,
10.1
Asio, Briss., 248.
Aspidouectes, Wagh, brain of, 475.
Ass, brain of, 389.
Astur taehiro, Daud., 210.
Asturina magnirostris, Gmel., 210.
Ateles, Geoffr., 410.
-Atergatis tioridus, Linn., 5 ; men-
tioned, 2.
Athene, Boie, 248.
Atya armata, A. Mihie-Edw., men-
tioneil, 433.
pilipes, Newp., mentioned, 433.
spinipes, Newp., mentioned, 433.
Atyidw, 447.
" Auditory area," 410.
Aula, 4(il, 402.
Aulacolambrus hoplonotus, Adams J,-
White, var.'i, 41.
" .Aural area," 410.
Autonomic Nervous System of Birds,
258.
Avabis, Jourd., 338, 378 ; brain of,
321.
laniger, Milne-Edir. 4- Grandi-
dier, brain of, 353, 354.
Aye-aye, brain of, 322, 354, 301.
Aysha (Jueh-hii, Pocock, 07.
Ealeariea, Bri.ts., 223.
liarjchelidoe, 00.
Batrachians of Mount Roraima in
British Guiana, by (i. -A. Boulenger,
55.
Bear, brain of, 387, 388, 397, 411.
Bears, carnassial teeth of, 100.
Betteus, Dana, 438.
aequimanus, Dana, 438 ; men-
tioned, 4.53.
microstylus, Spence Bote, men-
tioned, 438.
trispinosus, Stinips., mentioned,
438.
Birds, Intestinal Tract of, by Dr. P.
Chalmers Mitchell, 173 275.
Birds of ilount Roraima in British
Guiana, by R. Bowdler Sharpe, 53.
Bithynis, Philippi, 451.
longirnaua, Phd.. mentioned, 01.
oriiatus, Olivier, 451.
INDEX.
503
Bitterns, 195, 197.
Black Snake, brain of, 47.J.
'• Bogent'urchc," 412, 4(i8, 4(;9.
Bolocera longicoruis, Cai-hjnn, nu^n-
tioned, 3U5.
McMurrichi, Kirietn., mention<'(l,
306.
occidua, McMurrich, mcatioticd,
30.5.
Bolocerciidos JIp'Miirriclii. Kuiein.,
mentioned, .'iOO, .3(1."), ;!i»(i.
Bonellia minor, JIarion, nervous
System of, bo, 91.
viridis, liolamlo, ftnote S9 ;
dimorphism in, 90.
Bos, brain of, 374, 402.
Boulenger, G. A., Beptiles of Mount
Koraima, British Guiana, 53--5-').
Batraohians of Mount Boraiiua,
British Guiana, oo-oii.
Brachycarpus, Sptnce Bute, 451.
Audoiiini, Spence Bale, 451.
Brach^ura, Ldtr., On a Colleclion of,
from Torres Straits, liy W. T. Cai-
man, 1-50.
Brachyiirus, Fisrii.. 41(').
Br.idypodicke, ;i70, 392, 407. 419,
424.
Bradypus, Linn., 407 ; brain of, 402,
403.
Brain in the extinct Lemuroid Globi-
lemur, 362.
in the Galaginte, 34(i.
in the Indrisinse, 349.
in the Lorisinie, 339.
in Mammalia. Morphology of,
by Dr. G. Elliot Smith, 319-
432.
in Megaladapis, Major, 364.
Brain-stem of Mammalia, 373.
Brazilian Caracara, 207.
Broteochaetas, Fucovh, 6S.
delicalus, Karsch, 68, 71.
GoUmeri, Karsch, 08, 71.
nitidus, Pocock, 68.
opacus, Karsch, 68.
parvulus, Pocock, 69, 71.
porosus *, Pocock, 70.
Schaumii, Karsch, 69.
Sclateri, Pocock, 69, 71.
granosus *, Pocock, 69, 71.
Brvtcus panamcnsis, Thorell, 68.
Bubo, Ctti'., 248, 257.
maculosus, VieilL, 249.
maximus, Flemhuj, intestinal
tract of, 24b.
Bubo poensis, Fruser, 249.
Bubonina.', 218.
Bulbus olfactorius, 489.
Bunodeopsis, Andres, 314.
antilliensis, Duerden, mentioned,
299,302, 305-316.
globulifera, ]\rnll. On the
Actinian, by J. E. Duerden,
297-317.
glol)\ilifera, VerriU, 298: men-
tioned, 297 ; anatomy and
histology of, 3iiU, ;iti5-31ti.
struniosa, Andr., nitTitioned,
313.
Bunodes thallia, Di.con, mentioned,
315.
verrucosa, Di.con, mentioned.
306, 315.
Burmeistcr's Cariama, 225.
Buteo, Cuv., 203.
erythronotus, A7»;/. 210.
ferox, (inid., 210.
jacal, iJdud., 210.
Buteouiiiie, 21(_).
Cacatua, Briss., 24.'!.
Caccabis, Kauji. 210.
Cacomantis lanceolatus, i'^. MiiUcr,
242.
Calappa hepatica, Linn., 25.
" Calcar," 381.
" avis," 385.
Calcarine fissure, .319.
'• group " of furrows, 390.
" sulcus," 332, 381-391.
sulcus and the occipital pro-
longation of the hemisphere,
370.
Callipepla, Wa;/!., 219.
CaUithrix, Fnc'l., 394, 414, 419.
torquata, llojjinannseijij, brain
of, 415.
Callorhinus ursinus, Linn., ftnote
382.
Caiman, W. T., On a Collection of
Bracbyura from Torres Straits,
1-5U.
Calodromas, Sclater if- &ilcin, 210,
207.
Catenas, Gray, 239.
Camel, brain of, 388, 3S9, 392,
393.
Camelus, Linn., brain of, 383.
Campephagida;, 252.
I Camptonij.v rolandifinns, A. Milue-
j Edw., 20.
I Cancer calculosus, \\. Milne-Edw.,
b.
liinhalns, II. Milne-Kdw., 0.
j Canidaj, 411.
Canis, LiHii., mentioned, 171, ;>""!,
413.
Caphyra rolundifrons, A. Milne-Filw.,
2il.
Capitonidip, 252.
Caprimulgida;, 250.
Caprimulgus, Linn., 250, 256, 2(;:5,
269.
Carabida', 74.
Curdiiia cucvirostrij,; Heller, 447.
Cariama, Briss., 213
Caridina eurvirostris, IJclUr, men-
tioned, 433.
Carinata;, intestines in, 173, 175,
184, 201.
Carnivora, Cnv., 381, :5.s3, 3b8 -
426; brain in, 361, 391.
" Carnivore [iseudosylvian sulcus,"'
402.
Carpilius, Dcsni., mentioned, 4^
cinclitnaniis, Wliite, 4.
convexus, Forsk., 4 ; mentioned,
Carpilodes, Dana, mentioned, o.
cariosus, Alcuck, mentioned,
4.
cinclimanus, Mitrs, 4.
rugatus. Late, mentioned, 4.
sp., 2, 4.
Carpococcyx radiatus, Temm., intes-
tinal ti'act of, 242.
Cassowary, intestines of, 173, 1*2,
183.
Castor, Linn., 354.
Cat, brain of, 390, 395, 397,
398, 410, 426: cerebellum of.
426.
Catharlffi, 205.
Cathartes aura, /yt/ui., intestinal tract
of, 20.5, 212.
CathartidK, 205, 268.
Cave-Bear, upper carnassial teeth of.
100.
Cebidx, 303, 371, 376, 393, 405.
412, 4.15, 419.
Cebus, Er.vL, 410.
" Central sulcus," 343, 359, 400.
Ceiitro[ius, ///., 242.
Centropyx calcaratus, SjiLr, 53.
striatus, Baud., 53.
501
INDEX.
( 'cnitopla X ciliata, Slinipsov, men-
tioned, 25.
sp., 25.
Cercopithecidse, 363, 407, 412. 415,
419.
Ceicopitheciis, Er.xL, 380, 381 ; brain
of, 374, 376.
sabfeus, Linn., 379.
(Jerebellum of Mammalia, 425.
Cerebral hemisphere, morphology of
the sulci ou the lateral and dorsal
aspects of (Dr. G. Elliot Smith),
395.
Cerebral hemispheres in the genus
Daubenlonia, Geoffr., 354.
in the genua Hapalemur,
Geoffr., 337.
in the genus Lemur, Linn.,
325.
in the genus Lepidolemur,
Peters, 33S.
in Tarsius, Starr, and the
Prosimi* generally, 3<)7.
Cerianthus amerieanus, L. Ji/as^.,
mentioned, 301, 316.
Cetacea, 392, 421, 423.
Chactas Gollmeri, Karsch, 68.
opacus, Karsch, 68.
<iuiniiuedci>tatus, Karsch, 09.
Scliaumii, Karsch, (>9.
Chactidse, 68.
Chalcopclia, Bonaparte. 2;i9.
('hamelco, Gray, brain of, 485.
C'haradri-Gruiforra metacentre, 241 .
CharadriidiB, 230, 231, 235, 240,
241, 269.
Charadriiformes, 173, 230, 240, 263,
264, 269.
f'haragmophoruslineatus,TF«to-7i.,74.
Chaulclasmus, Bonajmrte, 203.
Chauna, III.. 267.
chavaria, -Liiut., 201.
derbiana, Grai/,201.
Chelone, llitgen, brain of, 479.
Chelonia, A. Brongn., brain of, 474,
485, 486.
Chilton, Chas., The Terrestrial Iso-
poda of New Zealand, 99-152.
Chimpanzee, brain of, 407.
ChionidK, 231, 2.35, 240, 241. 209.
Chionis alba, intestinal tract of, 231.
Chirogule, Geoff roij, 322, 349.
Chirnmys, Citv., brain in, 321 42().
raadagascariensis, C'kw., brain of,
354, 355, 361 ; weight of
brain of, 420.
Chiroptera, Blum., 383, 392, 404;
brain in, 329,301,368.
Chlrtmydophorus, IJarl., 406.
Chlorinoides aculeatus, J/. Milne-
Edw., 38.
var. urmat.us, Miers, 38.
Coppingeri, HasweU, 38.
Chlorodim e.varatns, H. Milne-Edw.,
6.
fragifer, Adams t|- White, men-
tioned. 11.
niger, i^j)\s7i-., 11 ; mentioned, 2.
scitlj)tus, A. Milne-Edw., 12.
unguJatus, H. Milne-Edw., 11.
Chlorodopsis nielanodactylus, A.
Milne-Edw., 12.
spinipes, Heller, 12.
ChcBropotamus parisiensi.s, mentioned,
163.
Cholrepus, lllig., 402.
CJiorinus aculeatus, Milne-Edw., .38.
verrucosipes, Adams & "White,
36.
Chromacris speciosa, Tliunh.. 73..
Chrysothrix, Kaup, 364, 394,419;
brain of, 415.
Clirysotis, Swains., 243.
Chuuga Burmeisteri, 225.
Cicada perspicillata, Eabr., 73.
Cioonia alba, Linn., 198.
nigra, Bechst., 198.
Ciconite, 197, 263, 266.
Ciconii, 213.
Ciconiidie, 197, 268.
Ciooniiformes, 263, 268, 270 ;
intestines of, 173, 189, 204.
213.
Circaetus gallicus, Gmel., 208.
Circus, Lac., 208, 210.
cineraceus, Mont., intestinal
tract of, 207, 209.
Clamatores, 252.
Ca?rebid8e, 254.
Coleoptera of Mount Eoraima in
British Guiana, by C. A. Water-
house, 74-76.
Colic coeca of birds, 173, 207.
loop of birds, 1 74.
Colli, 251, 250, 203, 209.
Colius capensis, Layanl (nee Gniel.),
intestinal tract of, 281.
•' Collateral sulcus," 333, 394.
Coluraba, Linn., 239, 260.
livia, Bonnaierre, 269.
oenas, Linn., 269.
, Columha;, 239, 240, 264.
Columbida?, 221, 239, 241, 263, 2(i7,
269.
Columbula, Bonaparte, 239.
Colymbi, intestines of, 184.
Colymbiformes, 263, 268 ; intestines
of, 173, 184, 188, 213.
Colymbomorphse, 188, 193, 194, 197,
204, 213.
Colymbus, Linn., 213.
glacialis, Linn., 184.
septentrionalis, Linn., 184.
Commissura aberrans, 455-495 ;
comparative morphology of, 484-
500.
dorsalis, 400, 464, 472, 484,
4»7, 488, 490.
fornicis, 456, 484, 488.
]>alii posterior, 485.
superior, 484, 489, 490, 494,
495.
ventralis, 461, 464, 487.
Conurus, KuM, 243.
CoraciiB, 245, 249, 263.
Coracias garrula, Linn., inteslinul
tract of, 245.
Coracii, 248.
Coraciidae, 245, 256, 26'9.
Coraeiiformes, 173, 245, 250, 203,
209.
Coraciomorphfe, 256.
Coraciomorphino Legion, summary
of, 173, 256.
metacentre, 2.58.
CoreidiB, 72.
"Coronal sulcus,"' 334, ;{59, 413,
416.
Corpus paraterminale, 458, 459, 40:.'.
403.
" Cortex Ammonis," 473.
Corvidifi, 254.
Corvus, Linn., 260.
capellanus, Sdater, intestinal
tract of, 254, 255.
Corythaix chlorochlamys, Sluih-ii,
intestinal tract of, 243.
persa, Linn., intestinal tract of,
243.
('otingidse, 252.
C'oturnix, 219.
Cow, brain of, 388.
Cracida^ 218, 219, 220, 258.
Cranes, 224.
Cranijon nustralis, Hutton MS.. 434.
Crangonid®, 454.
Cratosomus subaugulatus, ■^'c7/u»/i.,
mentioned, 76.
INDEX.
r,o:
Crax, Linn., 219, 220, 2(;i).
Daubeiitoni, Grai/, intestinal
tract of, 218, 219, 260.
Crested Urebe, 184.
Crex pratensis, Bechst., 222.
Crocodilia, brain of, 474, 485, 486.
Crotophaga ani, Linn., 242.
•■ Crucial sulcus," 40ii.
Crustacea of Mount Roraima in
l^ritish Guiana, by Dr.J.U.de Man,
.■57-64.
Crustacea, On the Xew Zealand
Phyllobranchiate Crustacea- Ma -
crura, by George M. Thomson,
433-473.
Cryptochirus, Hdlcr, mentioned, 48,
49.
coralliodytes, Heller, mentioned,
47.
Cryptocnemus Grandidieri, .1. Mihie-
EduK, mentioned, 20.
Haddoni*, Calm., 25.
Huldsworthi, Miers, mentioned,
20.
obolus, Oi-tm., mentioned, 20.
pentagonua, Stinips., mentioned,
25, 26.
Crypturida;, 21.5.
Cnbaris regulosus, read rugulosus,
Miers, 147 ; mentioned, Ull.
Cuckoos, 242.
Cuculi, 242, 269.
CuculidiB, 221, 242 244, 250, 203,
209.
Cuculiformes, 17.3, 242, 25(;, 2(i3,
269.
Curlew, 230.
Cyelax suborbicularis, Stimps., 39.
(\i/ch)maia marejaritala, A. Milne-
Edw.. 3i).
Cyelorhis albiventris, Sdater, intes-
tinal traut oF, 254, 255.
Cycdoturus, Gra//, 422. I
Cygnus, Linn., 215.
atratus. Lath., intestinal tract
of, 201,203. I
Cylloma, Budde-Lund, 130.
oculatum, BTidile-Liiml, men- \
tioncd, 152*. [
Cyrao Andreossyi, And., 13. I
melanodactylus, De Haan, 13.
Cymoijulia, Eoux. 2!>.
carinipes, Paulson, 29 ; men-
tioned, 30. j
J'm/.-W;;, White, 29.
serripes, Alcock & Anders., 32. j
<_'i/mopoli(i Whitei, Miers, 31.
Cynocophali, 412.
CynohyaMiodou Caylu.xi, 400, 401.
Cynoidea, 410.
Cynonycteris, I'eterx, 383, 104 ; brain
of, 329.
Cypseli, 250, 250, 203 -2(;il.
Cypselida;, 250.
Cypselus apus, fJnn., 250, 251.
Cyrtogrammomraa, Poeock, 0().
nionticdla, Ptn-Dck, (i(i.
Cyrtosoma montanum *, Watrrh., 76.
Cystiaotis, Eilic, mentioned, 298.
Dacelo, Leadi, 248.
gigantea. Lath., intestinal tract
of, 245, 246.
Dascillida>, 74.
Dasypodid;e, 419, 420.
Dasyprocta, ftnote 329.
Dasypus, Linn., brain of. 374, 375.
Daubentonia, Oeoffr., cereliral hemi-
.spheres in, 354.
madagascaricnsis, Gmd., brain
of, 354.
" Deckplatte " of forebrain, 492.
Dendrocolaptidie, 252.
Dendrocopus major, Linn., 252.
Dendrocygna, Strain^., 203.
Diactor suratus, Burm., 72.
" Diagonal sulcus," 400, 407.
Diaplexus, 400, 470.
Dica;ida>, 253.
Dic.holophids, 225,241, 208.
Dicrurid;B, 254.
l)idel|ihys, L^inn., brain of, 475.
Diomedea exulans, Linn., intestines
of, 188, 189.
Diplax illutum, var. ijllva, De Helys,
73.
Diplopoda, 04.
Diprotodontia, 383, 404.
Dissura episcopus, Bodd., 198.
Dog, brain of, 381, ftnote 390, 390,
397, 412; cerebellum of, 4.30.
Dog-fish, 273.
Dolichoderus bispinosus, Olic, 72.
Dolichotis, Desm., 403-413.
Dorippe sima, //. Milne-Ediv., men-
tioned, 29.
Ducks, 201, 203, 2(53, 208.
Duerden, J. E., On the Actinian
Bunodeopsis globulifeni, I'errill,
297-317.
Duodenal loop of birds. 173, 175.
Duodenum of birds, 178.
Dytiscida-, 74.
Eagles, intestines of, 188.
Echidna, brain of, 383, 466-471,
489, 490.
Eehiurus Pallasii, Gaerin, mentioned,
77-95.
unicinctus, von Drasdie, On the
Struetnre and .\flinities of,
by Alice L. Embleton, 77-
97.
vulgaris, Forbes, mentioned,
77.
Eclectus, M'a>jl., 243.
Edentata, 383, 392, 402-414, 425,
426, 430 ; brain in, 301.
Emble ton, Alice L., On the Structure
and Affinities of Eehiurus unicinc-
tus, 77-97.
Emu, intestines of, 173, ] 82.
Emys, Brongn., brain of, 498.
europaea, Sehneider, brain (if,
478.
Epeira auda.r, Blackw., (iS.
Epithetosoma norsegicum, Daniehseii
6) Koren, ventral mcsonterv in,
92.
Equus, Linn., brain of, 374.
" Ergot," 385, 388.
Erinaceus, Linn., 424, 427, 428.
Etisodes electra, Ilerbst, 7 ; men-
tioned, 2.
frontalis, Dana, 7 ; men-
tioned, 2.
scidjitilis, Heller, 7.
Etisus lacvimanus. Hand., 7 ; men-
tioned, 2.
Eudyptes, Vieill., 180, 188.
KuryhemidK, 252.
Eurypyga, Illi;/., 228, 229, 241.
helias, I'allas, 228.
EurypygidjB, 228.
Euryurus, C. Koch, 65.
atratus *, Pococl; 65.
fiimigatus, PeUrs, mentioned,
fiii.
Eutheria, 38.3, 393, 404, 418.
Eti.vantltas tuberctdosux, Miers, 8,
9.
Exagontus *, Waterh., 74.
dcnticoUis*, \V<iUrh., 75.
506
INDEX.
I'aloo concolor, Temm., 210. I
FeldcKgi, Schkijd, intestinal
tractof, 210, 211. j
lariarius, Schle(/el, 210.
melanogenys, intestinal tract of,
210, 200.
peregrinus, l^inist., 210.
Fulconidir, 207, 203.
l'"alconiformes, 173, 205, 213, 214,
263, 268, 270.
Faleoninie, 210, 204.
Fascia dcntata, 330, 469.
Fasciculus margiualis, 4()0, 471.
Felifla?, 411.
" Feline sylvian sulcus," .3.58.
Felis, Linn., brain of, 332, 374, 375,
402.
catus, Linn., weigUt of brain of,
421.
doniestica, Gmel., weight of
brain of, 396, 421.
" Fissura sublimbica j)osterior," .343.
Flamingoes, 194.
" Flocculus," 430.
Foramen of Monro, 402, 463.
Formicariidie, 252.
Formicida;, 72.
" Fornixleiste," 487.
Francolinus, Stejih., 219.
Fratercula arctica, Linn., intestinal
tract of, 237, 238.
Fregata, Cnv., intestinal tract of,
102, 193.
Fregatid;e, 192, 208.
Fringillidse, 254.
Frog, 202, 273 ; brain of, 496,
497.
'• Frontal sulcus," 414, 410.
'• Fronto-orbital sulcus,'" 4o7.
Fulgoridfe, 73.
Fuligula, Stejih., 203.
Fulmaris glacialis, Linn., intestines
of, 188.
r.alagina>, 338, 339, 368, 372, 422,
' 424 ; brain in, 346.
Galago, Ciiv., 321, 322, 340, 348,
373, 377, 407, 412, 422, 423, 420.
Allenii, Wa/crJi.. 348.
crassicaudata, G'loJJ'r., 323;
brain of, 347.
Garnctti, Ot/ilh,^, 323, 324 :
brain of, 346.
Monteiri, Barthtt, brain of,
348.
Galbulidre, 175, 2.')2. I
(ialeopithecus, Pai/a.s-, 383, 404, 428. 1
Galictis vittata, Schnh., braiu-weight
of, 421. 1
Galli, 218, 221, 240-244, 257, 258,
208. j
Gallida?, 21 9, 220.
Galliforme.s, 173, 210, 221, 240, 203,
208.
Gallinula phoenicura, Penn., 222.
Gallus, Linn., 219.
Gazella dorcas, Livn., 428.
Gecinus viridis, Linn., 252.
Geese, 201, 203.
Gelasituus tctragonon, //< rh.st, 24 ;
mentioned, 3.
variatus, Hess, mentioned, 24.
" Genual sulcus," 332.
Geoligia, DoJlfus, mentioned. 105.
Simoni, Z'o///w«, mentioned, 105,
110.
Geopelia, Swains., 239.
Gizzards of birds, 173.
Glareola, Briss., 237.
ocularis, T'erreaii.r, intestinal
tract of, 232, 233.
pratincola, Linn., 233.
GlareolidK, 233, 234, 235, 240, 241,
209.
Globilemur, Major, 307 ; brain of,
324, 362, 303, 304, 424.
Flacourti, Major, biain of, 302,
363.
Glyphocrangon, A. Milne-lu/m., 436.
regalis, Spence Bate, 43(>.
Gonactinia, Sars, mentioned, 300.
Goniastraja, Milne-Edw. i|" JJaime,
mentioned, 47.
Gorilla, Uenffr., brain of, 405, 407.
Goura, Stcph., 239.
Great Bustard, 220.
Koitbern Diver, 184.
Gruida\ 223, 220, 268.
Gruiform metacentre, 241.
Grui formes, 173, 222, 228, 230, 235,
240-242, 263-208.
Gruinse, 241.
Grns virgo, Linn., intestinal tract of,
223.
Gruvel, A., Etude d'une Espece
nouvelle de Lepadides (Scalpellum
giganteum, n. sp.) et de Poecilasma
Ciirinatum, JJoek, 153-1 0(t.
8ur (pielques Lejiadides nou-
veaux de la Collection du
British Museum, 277-295.
GrifJliis .•'peciosus, Thiinb., 73.
Guira piririgua, Bnjfon, 242.
Gulls, 230. 238, 239, 263, 270.
Gulo, Storr, 413.
Gvmnoglaux, Cahanis, 248, 249.
Gyranura, Vig. cj- Mors/., 404,
428.
Gypaetina;, 207.
Gypogerunidie, 200.
Gypohierax, Biijij)., 214.
angolensis, Gmel., intestinal
tract of, 207, 208.
" Gyrus hippocampi," 329.
" intermedins," 330.
" lunaris,"' 329.
" unciuatus," 329.
Iladrxirorhaefas, Pocock, 68.
Sehiteri, Pocock, 69.
Haliaiitus, >Sav., 215.
albicilla, Linn., 208.
leurogaster, GmeL, 208, 210.
Hamingia arctica, A) (u'i;/.s-se'w i^- Kuren,
anal vebicles of, 88.
Hapale, /?/., brain of, 369, 380,
381.
jacchus, Linn., 389.
Hapalemur, Geoffr., brain of, 322,
337, 338, 423.
griseus, Link, brain of, 322.
simus, Schlez/el ^' Pollen, brain
of, 322, 337.
Hapalids, 328, 332, 371, 370, 393,
405.
Haiialooarcinus, Stim2^s., mentioned,
45-49.
marsupialis, Slim^ts., 43 ; men-
tioned, 2, 45, 50.
Eaplopbthahnus, Schobl, 114, 119.
Hclmsii *, ClriUon, 119; men-
tioned, 151.
Hawks, 239.
Hedobiii, Latr., mentioned, 75.
Heilipus earinirostris, ScItdnJi., men-
tioned, 76.
Helictis, Gray, ftnote 401.
Ilcliornis, Bonn., 241, 263.
fulica, Bo'ld., intestinal tract of,
22-<, 229.
surinamensis, Gmel., 229,
Heliornithida% 228, 241.
arehecentric, 268.
Hclotarsus ccaudatus. Baud., 208.
Hemiptera Heteroptera of Mount
Pioraima, by VV. F. Kirby, 71-73.
INDEX.
mi
Herons, 195, 197, 199, 263.
Herpestes, III, 399, 406.
albicauda, Vuv., brain-weight of,
421.
Himaiitoijus, JSriss., 230.
Hippidea, mentioned, 49.
Hippocampal fissure, 310.
"Hippocampus," 461.
" minor," 3S5.
" nudus," 331.
Hippolyte, Leach, 443.
bifidirostris, Miers, 443 : men-
tioned, 4.53.
spinifrons, A. llilne-Edw.,
444.
Stewarti, G. M. Thomson, 445.
Hippolytida?, 44(t.
Hirundinidaj, 252.
Homo, Linn., brain of, 374.
Homoplastic modifications, 179,
180.
Homoplasy, 179.
Homoptera of Mount lloraima, by
W. F. Kirby, 71-73.
Hoopoes, 248.
Horned Screamer, intestinal tract in,
175.
Horse, brain of, 388.
Huenia jjroteus, De Ihian, 34.
Humming-birds, 251.
Hyajnarctus, Gerv., upper carna.ssial
teeth of, 166.
Hyaenidoe, 399, 441.
Hyastcnus Brockii, dr Man, 36.
convesus, Miers, 35.
oryx, -1. MiJnt-Edvj., 35.
Sebce, White, mentioned, .35.
spinosus, ..-1. MUne-Edw., 3.5.
verrucosipes, Adams c$' White,
36, 50.
verrucusipes, A. Milnc-Edw..
36.
Hydrochoerus, Briss., 38:?, 403.
Hydrosaurus, Kiwp, brain of, 470,
471, 472, 473, 479, 4S3, 499.
Hyla albomarginata, Spix, 56.
Hylobates, 111, brain of, 405, 412,
430.
Hvlodes marmoratus *, Bouhiujcr,
56.
Hvmenoptera of Mount lloraima. by
"W. F. Kirby, 71-73.
Hymenosoma, Deam., mentioned, 48.
Hyperoodon, ],ac., 383.
Hyrax, Berm., 401, 413.
H^strix, Linn., 354.
IbididoB, 194, 199.
Ibises, VM.
Ichthyoiihis, Fitz., brain of, 498.
Ichthyopsida, 273; brain of, 493, 494.
Ichthyoruithes, 188.
Icteridoc, 254.
Iguana, Baud., brain of, 484, 485.
Inacbiniv, 4(1.
Inachiiif (Ojii'inopus) uranea. Do
Haan, 34.
Indian Open-bill, 197.
Indris, Oeoffr.. brain of, ;!21. :i22,
349, 354, 35S,i!(i2, 365, 377, 406.
412, 416, 419.
brevicaudatus, Geoffr., brain of,
3.53.
Indrisina", brain in, ;i2I, 1538, 349,
354, 368, 378, ftnote 379, 405,
423.
" Inferior temporal sulcus,"' 342.
Insectivora, 331, 383, 42'.).
"Insula." 343, 412.
"Intercalary sulcus," 332.
Intestinal Tract of Birds, by Dr. ?.
Chalmers Alitchell, 173-275.
Intestines of Cassowary, 173, 182.
of Emu, 173, 182.
of Ostrich, 173, 182.
of llhea americaua, VieiJJ., 173,
182.
" Intraorbital sulcus," 4(K5.
" Inverted hippocampus," 331.
lonornis martinious, Linn., 222.
Isopoda, The Terrestrial, of Xew
Zealand, by Chas. Chilton, 99-162.
lyngidffi, 252.
Ij'iix torquilla, Linn., 252.
.lanicsia globifern, Fahr., mcntldiied,
76.
Kagu, 227.
Kingfishers, 245.
Kirby, W. F., Hymcnoptera, Hemi-
pt.era Heteroptcra, Homoptera,
jS'europtera, and Orthopfera of
Mount lloraima in British (iuiana.
71 73.
Kraussia nitida, Stim/is., 24.
Labium caudale, 486, 492.
Lncerta, Linn., brain of, 4>5.
Lacertilia, brain of, 474.
Eachesis lanceolatus, Lacfp., 54.
Lagostomus, Brook, 403.
Lagothrix, Geojfr., 416.
Lahaina ovata, Dana, mentioned, 36.
Eambrus coufragosus *, Caiman, 50 ;
mentioned, 1.
hoploHotus, Adams & While,
41.
tarpeius, Adams ^ White, mon-
tioned, 42.
(Aulacolambrus) hoploiiolus,
Adams 4' While, var. ?, 41.
( ) , var. lougieollis,
Miers, mentioued, 41.
(Parthenolambrus) calappoides,
Adams 4- White, 41.
( ) confragosus *, Caiman,
42.
LamiidaB, 76.
Lamina chorioidea, 456, 463, 486.
LaniidsD, 254.
Lankestcr, E. Bay, On the Affinities
of iEluropus melanoleucus, A.
^Lil ne-Edwurd^, 163-1 72.
Beport on a Collection made by
I". V. McConnell and J.. .1.
Queleh at Mount lloraima in
British Guiana, 51-76.
Lari, 236, 240, 241, 263.
Laridie, 237, 241, 263, 269.
Larus argentatus, Gmel., 23(i.
marinus, Linn., intestinal tract
of, 236.
ridibuudus, Linn., 236.
"Lateral sulcus," 334, 412, 413,
416.
Leander affinis, Miers, 450.
erratica, Desmarest, 450.
Jlariatilis, G. M. Thomson, 447 ;
mentioned, 433.
natator, Miers, 450.
natator, Stimps., 450.
Lemur, Linn., mentioned, 331-348 :
brain of, 322, 324.
alhifrons, Geoffr., brain of, 32:?,
337.
anjuanensis, Geoffr., brain of.
335, 337.
catta, Linn., brain of, 323. 'Xi\,
337.
c ironatus, Gray, brain of, KHi.
fulvus, Geoffr., brain of, .3,'J5,
336, 337, 341, 355, 378, 379,
408, 423.
fulvus, Gioff'r. [mongo/,, Linn.\,
brain of, 322-330.
508
INDEX.
Lemur fulvus, Geoffr. [nigrifrons],
brain of, 821.
JuUyi, (rVrt)irf«Z(C)-, brain of, 324,
336.
inacaco,X/Hii., brain of, 322, 324,
326, 330, 335, 330, 378, 427,
428.
macao, read macaco, Linn., 322.
mongoz, Linn., braiu of, 321,
322, 323, 331, 335, 337;
■weight of brain of, 420.
nigrifrons, Milne-Edw. & Gi'an-
didier, bruin of, 321, 323,
337, 377.
rubriventer, Geoffr., 337.
varius, Geoffr., brain of, 323-
337 ; weight of, 420.
I.cmuriiiae, 3G8, 378.
J.emuroidea, 325, 372, 417, 424.
Lemurs, Morphology of the Brain of,
by Dr. G. Elliot Smith, 319-432;
affinities of, 417.
Lepadides, Sur quelques Lepadides
nuuveaux de la Collection du
liritish Museum, par A. Gruvel,
277-295.
Lepadides (Scalpellum giganteum,
n. sp.),Etude d'une E.spece nouvelle
de, et de Po?,cilasma carinatum,
IIoclc, par A. (iruvel, 153-169.
l.epadiiise, 288.
Lepidolemur, Peters, brain of, 322,
346,347,423; cerebral hemisphere
in, 338.
leucopus, Major, brain of, 339.
mustelinus, Geojfr., brain of,
338.
ruficaudatus, Grandidier, 339.
Lepidosiren, JSatterer, brain of, 495,
499.
Leptodius esaratus, //. Milne-Edw.,
6.
■ exiiraius, Alcock, 6.
Leptoptilus argala, Linn., 198.
crumeniferus, Cuv., 198.
Leptosoma, Bonaparte, 245.
Lepus, Linn., brain of, 374, 375,
410.
Leuconerpes candidus, Otto, 252.
Lcueosarcia, Goidd, 239.
Lcucosia Haswelli, Miers, 27.
longifron.s, var. pulcherrima,
Miers, 27.
lonqifrons. var. ^(MW(cc/-/»ia,
Alcock, 27.
■pidclierrlma, Miers, 27.
Leueosia splendida, Hasicell, 27.
Libellulida;, 73.
Ligia, Fahr., 106 ; mentioned, 1 10.
australiensis, Dana, mentioned,
106.
neo-zelanica, Thomson & Chilton,
107.
novK - zealandife, Dana, 107;
mentioned, 101, l(i2, lOC,
103, 109, 116, 117, 151.
novf-zeaJandiif, Miers, 107.
oceanica, Linn., meutioucd, 106,
109, no. 111.
qxiadrata, Thomson, 1U7. Ill8.
Ligidium, Brandt, mentioned, li»9.
Persooni, Brandt, mentioned,
ftnote 109.
" Limbic furrow," 389.
Limicolse, 230, 235, 263, 260.
Limicoli, 242.
Limicoline-Gruiform metacentre, 240,
241, 25S.
Liomera, Dana, mentioned, 4.
cinctimana, lF/)t(^,4; mentioned,
2,5.
lata, iJana, 4.
lata, A. Milne-Edw., 4.
Liponema multiformum, R. Hrrtiri<j,
mentioned, 306.
Lissocarcinus orbicularis, Dana, 20.
Lithoscaptus paradoxus, A. Milne-
Edw., Ai<.
Little Grebe, 184.
Looustida-, 73.
Lomvia truile, Linn., intestinal tract
of, 238.
Lophactsea granulosa, -Zi't^»^)., 6; men-
tioned, 2.
granulosa, A. Milne-Edw., 6.
Lophophorus, Temm., 219.
Lophopilumnus, Miers, 18.
Lophoxanthus frontalis, liatldjun,
mentioned, 19.
Lopho/.ozymus dodone, Herhst, 6 ;
mentioned, 2.
octodentatus, H. Miln,e-E<lw., 6 ;
mentioned, 2.
Loris, Oken, 322, 323, 343.
gracilis, Genff'r., 322, 323, 342,
343, 345.
Lorisina;, ."138, 339, 345, 34'.), 368,
419.
Lorius, Brissun, 243.
Lucanida% 74.
Lumbricus, Linn., mentioned, 85;
nerve-cord of, 96.
Lujia (jraniilata, H. Milne-Edw., 21.
pelagica, H. Milne-Edw., 21.
sanqninolenta, H. Milne-Edw.,
21.
Lutra, Lian., brain of, 374.
Macacus, Car., brain of, 410.
McConnell, F. V., and Quelch, J. .1.,
Eeport on a Collection made at
Mount Roraima in British Guiana,
51-76.
Macrobrachium Quelchi *, de Man,
57 ; mentioned, 60.
Macropodidit, 404, 413.
Macropus, Spiv, brain of, 374, 383,
ftnote 404.
Macroscelides, brain of, 331, 374,
375, 427, 428.
Maia (Huenia) proicws, De Haan,
34.
Mammalia of Mount Roraima in
British Guiana, by W. E. de
Winton, 52.
Man, Dr. J. G. de, Crustacea of
Mount Roraima, British Guiana,
57-60.
List of the known Species of
the Genus Palaemon, Fabr.
s. s., May 1900, 60-64.
Manatee, brain of, 331, 382.
Manatus, Bond., 382.
Manicina areola ta, Linn., mentioned,
316.
Manida;, 392, 424.
Manis, Linn., 373, 4()3.
Marsupialia, brain in, 361, 383, 392,
393, Itnote 404, 414, 426.
Meckers diverticulum in birds, 173-
272.
tract, 173-272.
tract in Palamedea cornuta,
Linn., 176.
Median looj) of intestines of birds,
174,
Megaladapis, Major, 324, 362, 364,
365, 366, 424.
madagascariensis, Jia/., brain of,
364, 365.
Megalasma, Gratj, 252.
asiatica, Lath., intestinal tract
of, 252, 253.
MegapodidiB, 218, 220, 258.
Meles, Linn., 411.
Meliphagidic, 253.
Mdtpoiia Jasciculaia, Smith, 72.
INDEX.
^Felipona intorrupfa, Latr., 12.
Menaisth'm.'i inujuslus, Dana, 'A\.
mouoceros, Latr., 34.
monoceros, Alcock, 34.
ilenura, Lutli., 252, 254, 263.
i[ergaiisers, 203, 215, 270.
Mergus, Linn., 2GS.
albellus, Linn., 203.
ilerhippolyte, Spifni-e Bate, 444.
orientalis, Sjyenee Bate, men-
tioned, 445.
spinifrous, A. Milne - Edw.,
444.
MeropidfB, 247, 2oC>, 257.
archecentrio, 269.
Merops, Linn., intestinal tract of,
246, 247.
Mesarraadillo, DoVf'ot, mentioned,
103.
AUuandi, renri .VUuaudi, Dollfun,
abnormalities of the mouth-
parts of, ftnote 128.
JFesentery of birds, 174, 175.
Mesites, Geoffr., 175, 2 Mi.
Mesitidte, 175.
Metatheria, calearine sulcus ol', 3.^3,
3!»3. 404, 41S.
lletopograpsus messor, Forsk-., 24.
var. gracilipes *, de M<iii,
24.
messor, Kingsley, 24.
Metopouorthus, Budde-I.und, 140.
pruinosus, Brandt, 141.
Metridium marginatum, Mihie-Ediv.,
mentioned, 309, 31(;.
Micippa, Leach, mentioned, 40.
niascarenica (Kossm.), Miers,
40.
philyra, I/erbst, 40.
superciliosa, Haswetl, mentioned.
41.
Microcebus, Gioffr., brain of, 324,
349, 368, 372, 373, 378, 380, 412,
422, 426, 431.
Smithi, Gray, brain of, 323, 324.
348, 308.
JficrochCErus, Wood, 424.
Miorohalimus, Haawdl, mentioned,
40.
deflesifrons, HasvM, mentioned,
40.
Microhierax melanoleucus, Blytli,
210.
Microphrys sly.r, A. Milne-Edw.,
37.
Midas, Geoffr., 332, 422.
SECOND SERIES. — ZOO LOG V,
Middle mesenteric factor, 177.
ililkpora, Linn., mentioned, 46.
Milvus govinda, ISyI.ex, 210.
ietinus, -Sav., 210.
migrans, Bodd., 210.
Mitchell, Dr. V. Chalmers, On the
Intestinal Tract of Birds ; with
liemarks on the Valuation and
Xomenclature of Zoological Cha-
racters, 173-275.
Mithrax asper, Milne-Edw., 39.
Mniotiltidie, 254.
Moniotidas 245, 256, 257, 2W\.
Monacli'.is, Kaup, ftnote 3S2.
Monitor, Cnv., brain of. 470, 471.
Monkeys, rliinal ti.-sure in, 329.
Morphnus guiancnsis, DuikL, 208.
Morphology of the lirain in the
ilammalia, with Special Reference
to that of the Lemurs, Keeent and
E.xtinct, by Dr. (J. Klli.it Smith,
319-432.
of the Cerebral Commissures in
the Vertcbrata, with Special
Reference to an .Vberriint Com-
missure found in the i'ore-
brain of certain Reptiles, by
Dr. G. Elhot Smith, 4.55-
500.
of the sulci on the lateral and
dorsal asjjccts of the cerebral
hemisphere, 393.
of the Sylvian fissure, 405.
Multiradial aiioceutricity, 179.
Jluscicapidin, 252.
Musophagid», 243, 244, 2(53, 269.
Mulilla larvata, Kluy, 72.
quadrum, Khty, 72.
Mutillidie, 72.
Mycetes, ///., 416. 419.
Mycteria americana, Linn., intestinal
tract of, 198, 199.
Myra australis, IJaswell, 27.
coalita, Hdyend., mentioned,
27.
dubia, Miers, mentioned, 27.
fugax, Fiihr., 26 ; mentioned,
27.
pentacantha ?, Alcock, 26 ;
mentioned, 27.
Myriopoda and Arachnida of Mount
Roraima in Rntisli (juiaiia, by
R. I. I'ocock, 64-7 1 .
Myrmeeophaga, Linn., 40.'), 404, 407,
409,410,411, 413, 422.
jubata, Linn., 402.
VOL. Vlll.
Myrmecophagidas 376, 392, ftnote
403, 419, 424.
Myrniica ii.ipinosa, Oliv., 72.
" Xatiform eminence " of pyrlforni
lobe, 340.
Xauticaris, Spence Bate, 445.
marionis, Spence Bate, men-
tioned, 445, 446.
Ste« arti, G. M. Thomson, 445 :
mentioned, 453.
A'(/.vjo ceraste.i, Ortm., 37.
serpulifera, Guerin, 37.
.lerpulifera, ililne-Edw., 37.
taurus, Pocock, 37.
'■ Xebenfurche," 337.
Nectariniida;, 251, 253.
Necturus, Raf., brain of, 498.
Is'ematocarcinidiC, 451.
Xematocarcinus, A. Milne-EdiK.,
451.
gracilis, Spence Bate, 452.
hiatus, Spence Bate, 452.
serratus, Sjience Bate, 452.
undulalipes, Spetice Bate, 451.
'• Neopallium," 461, 409.
Xcophrou porcnopterus, Linn., in-
testinal tract of, 206, 207.
Xeptunus arniatus, A. Miln--Edw.,
21.
pelagicus, Linn., 21.
peUujicas, De Haan, 21.
sanguinoleutus, JJerhst, 21.
sanyuinoleHluf, A. Milne-Kdw.,
21.
(.\chelouN) granulatus. If. Milne-
Edw., 21 ; mentioned, 22.
(-
-)
var. unispinosus,
Miers, 21.
( ) unispinosus, Miers, 21.
Nervous system, autonomic, of bird^,
2.58-261.
Xesonetta, Gray, 203.
.Vettopus, Brandt. 21-5.
eoromandelianus, Gmel., iutea-
tinal tract of, 204.
Xeuroptera of Slount Roraima in
Rrilish Guiana, by \V. E. Kirby,
71-73.
Xeusticurus rudis *, Bmdenyer, 'y.i,
of,.
Xe w Zealand, On the Phyllobranchiat «
Crustacea-Macrura, by George M.
Thomson, 433-473.
The Terrestrial Isopoda of, by
Chas. Chilton, 99-1.52.
74
510
INDEX.
'• Xoduliis ■' of cerebellum, 428.
Northern Petrel, 188.
Nothura, Wayl., 219.
maculosa, Temni., 215.
Notoryctes, Siirlimj, 427, 428.
Numeniiis, Linn., I'JO, 230, 231, 233,
235, 241.
arquata, Linn., intestinal tract
of, 230.
Nycticebus, Oeoffr., 322-347, 358,
3(54, 372-390, 412-426 ; brain of,
331.
javanicus, Geoffr., 342.
tardigradus, Gray, brain of, 321,
322, 323, 324*, 339-343, 360,
877-379 ; weight of brain of,
420.
Xycticorax griseus, J^iiin., intestinal
tract of, 195, 196.
Jsyctidromus, Gould, 252.
albicollis, Omel., intestinal tract
of, 250.
>'_vetipithecus, Spix, 3(>4, 394, 419.
" Occipital " region of brain, 3S5.
" Occipito-calcarine fissure," 391.
" Occipito-temporal fissure," 391.
( )eeanitid8e, 213.
'• (jculomotor area,'" 410.
Ocydromus australis, Sparrin., in-
testinal tract of, 222.
(-)cyphaps, Gould, 239.
Ocypoda ceratophthahiia, Pallas, 24.
I'dobienus, Linn., 411.
Odontoceti, 369, 421.
( )dontopeltis, Pocock, 64.
Jlacconnelli *, Pocock, 64.
(Kdicnemida;, 233, 235, 241.
(Kdicnemus, Temm., 234.
scolopax, Gmel., 233.
(Ena, Selby, 239.
Olfactory tract, 328, 329.
Oncinopus aranea, l)e Haan, 34.
neptunus, Adams & Wliite, .34.
Oniscidae, 121, 133.
Ouiscus, Unn., 133 ; luentioned, 103,
118.
Cookii, Filhol, 136.
kenepurensis *, Chilton, 135 :
mentioned, 136, 152.
noviK-zealandia', Filliol, 1 20,
127.
pubescens, Dana, 136, 137,
139.
Oniscus punctatus, Thomson, 133;
mentioned, 101, 102, 134, 135,
152.
" Opercular sulcus," 412.
Ophidia, £foi)r)n., brain of, 474,
485.
Opisthocomi, 220, 221, 25S, 268.
Opisthocomida;, 257.
Opisthocomus, III., 216, 221, 240,
241.
cristatus, Gmel., intestinal tract
of, 220.
Orang, brain of, 407.
" Orbital fissure," 395.
"sulcus," 399, 404, 40(i.
Oreophorus frontalis, 3Iiirs, 26.
Oreophrynella MacconneUi *, Bou-
lenger, 65 ; mentioned, 53, 56.
Quelehii, Bouhnger, 55, 56.
Ornithorliynohus, Blum., 371, 421 ;
brain of, 464-469, 489, 498, 499.
Oriolidic, 254.
Orthoptera of Mount Koraima in
British Guiana, by W. E. Kii'by,
71-73.
Orycteropus, Geoffr., 403, 413, 430 ;
brain of, 331, 383.
Oryzoryctes, Grand.. 428.
Oscines, 252.
Osprey, 211.
Ostrich, intestines of, 173, 182.
Otaria, Per., 382, 411.
OtididK, 226, 229, 240, 241, 268.
Otis, Linn., 216, 226-227, 257, 260,
261, 265, 268.
tarda, Uiin., intestinal tract of,
226.
Otogale Ivirkii, Gray, 347.
Otophryne *, Boulenger, bo.
robusta *, Boulenyer, 55, 56.
" Overlapped neopallium," 412.
Ovis, Linn., brain of, 374.
Owls, 268, 270.
Osycrepis leucocera, Lacord., 74.
Ozius guttatus, II. Milne-Edw., 13.
Pagophila cburnea, Pliipps, intes-
tinal tract of, 234, 236.
Pagurus coralliophilos, Ehrenh., men-
tioned, 45.
Palajcbirogaleus Jullyi, 6'. Grand.,
brain of, 337.
Palsemon, Fuhr., 450.
acanthosoma, sp. n. (?), XohlU,
mentioned, 6).
Palaemon acantburus, Wiegm., men-
tioned, 60.
acutirostris, Dana, mentioned,
60.
affinis, A. Milne-Edw., 450.
africanus, Kingsl., mentioned,
60.
africanus, Spence Bate, men-
tioned, 60.
Alphonsianus, Hoffm., men-
tioned, 61.
altifrons, Mend., mentioned,
61.
amazonicus, Hell., mentioned,
61.
americanus, Spence Bate, men-
tioned, 61.
Appuni, Marts., mentioned, (il.
var. £B(juatorialis, Ortm.,
mentioned, 61.
asper, Siitnjn., mentioned, 61.
asperulus. Marts., mentioned,
61.
Audouici, Hell., mentioned, 61.
Audouini, Spence Bate, men-
tioned, 61.
australis, Ortm., = sp., de Man,
mentioned, 61.
aztecus, Sauss., mentioned, 61.
bariensis, de Man, mentioned,
61.
boninensis, Stps., mentioned,
61.
Boreliii, Nolnli, mentioned, 61.
brachydactylus, Wiegm., men-
tioned, 61.
brasiliensis. Hell., mentioned,
61.
brevicarpus, De Haan, men-
tioned, 61.
brevimanus, Fahr., mentioncil,
61.
Ciementarius, Poepp., mentioned,
61, 62.
callirrhoe, de Man, mentioneil,
61.
carcinus, Fahr., mentioned, 61.
consobrinus, Sauss., mentioned,
61.
coromandelianus, Fahr., men-
tioned, 61.
Dante, Hell., mentioned, 61.
dasydactylus, Streets, mentioned,
61.
l)a}'anus, Hend., mentioned, 61.
Uosausuri, Hell., mentioned, 61.
INDEX.
511
Pala;moii dispar, ^fal■fx., mentioned,
61, 63.
dolichodactylus, Nilgend., men-
tioned, CI.
duleis, n. sp. ?, Tlialhv., men-
tioued, 01.
elegans, de Man, mentioned, 61.
endehensis, de Man, mentioned,
61.
ensiculus, .S". Sm., mentioned,
61.
equidens, Dana, mentioned, 61.
esculentus, Tludlw., mentioned,
61.
eunThynchus, Orlm., mentioned,
62. '
faustinus, Saijxs., mentioned, 62.
fluvialis, Streets, mentioned, 62.
forceps, Milne-Edw., mentioned,
62.
formosensis, Sjience Bate, men-
tioned, 62.
ganKcticum, Spence Bate, men-
tioned, 62.
Uaudichaudii, Milne-Ediv., 60,
61,62.
graeilimanus, Rand., mentioned,
62.
gracilirostris, Miers, mentioned,
62.
grandimanus. Rand., mentioned,
62.
heterochirua, Wier/m., men-
tioned, 62.
Hildebrandti, Jlihjend., men-
tioned, 62.
Hilgendorfi, Cont., mentioned,
62.
hirtimanus, Oliv., mentioned, 62.
hispidus, Oliv., mentioned, 62.
Horstii, de Man, mentioned, 62.
Ida?, Bell., mentioned, 62.
var. idella, Hih/end., men-
tioned, 62.
var. mammillodactylus, n.
var.?, Thallw., mentioned, 62.
var. subinermis, KohiJi.
mentioned, 62.
Iheringi, Ortm., mentioned, 57,
60, 62.
jamaicensis, Herbst, mentioned,
61, 62, 63.
japouicus, De Ilaan, mentioned,
62.
javauicus. Hell., mentioned, 62.
Jelskii, Mitrs, mentioned, 62. \
Palaemon Lamarrei, Milne-Ediv.
mentioned, 62.
larainatus, O'oUnt., mentioned,
62.
lamiiropus, de Man., mentioned,
(!2.
lanceifrons, Dana, mentioned,
62.
lar, Fahr., mentione<l, 62, 63,
64.
latidactylus, Thallu'., mentioned,
62.
latimanus. Marts., mentioned,
(>2.
lei)idactyIoides, de Man. men-
tioned, (i2.
lepidiictylns, JIdr/end., men-
tioned, 02.
longidigitum, Spence Bate, men-
tioned, 02.
longimanus, Fahr., mentioned,
62.
longiinanus, r!td., mentioned,
02.
longipes, De Ilaan, mentioned,
62.
maerobracliion, Herkl., men-
tioned, 60, 02.
niadagascariensis, Hoffni., men-
tioned, 62.
Maloolmsonii, Milne-Edw., men-
tioned, 62.
Malliardi, Richters, mentioned,
63.
Marias, Coxit., mentioned, 62.
mayottensis, Unffm., mentioned,
62.
mexieanus, Satiss., mentioned,
61. 63.
modestus, de Man, mentioned,
03.
Montezumjc, Sanss., mentioned,
63.
Moorei, Caiman, mentioned,
63.
mossambieus, Uilgend., men-
tioned, 63.
multidens, C'u»(., mentioned, 03.
natator, A. Milne-Fdic, 450.
Nattereri, Ifell., mentioned, 03.
nilotieus, Ron.v, mentioned, 03.
nipponensis, De Uaan, men-
tioned, 61, 63.
ohionis, S. Sm., mentioned, 63.
Olfersii, Wietjm., mentioned, 63,
64.
Pahvmon ornatiis, Oliv., = lar, Fahr.,
mentioned, (iij, 0.3, 451.
parvus, lloffin., mentioned, 03.
I'atsa, Coat., mentioned, 63.
liaucidens, Hilijend., mentioned,
63.
]'etersii,y/<7(/fH(/., mentioned, 63.
pilimanus, de Man, mentioned,
63.
var. leptodactylus, de Man,
mentioned, 63.
placidulus, de Man, mentioned.
Ii3.
]diicidus, de Man, mentioned,
0.3.
])otiporanga, F. Miiller, men-
tioned, (>3.
potiuna, /''. .Miiller, mentioned,
57, 5i), 03.
punctatus, Rand., mentioned.
03.
tiuelchi, de Man, mentioned,
6:5.
Quoianus, A. Milne-Edw., 4oO.
Ciuoianus, IV/iite, 450.
rcunionnensis, Hoffm., ■ men-
tioned, f53.
Ritsema^ de Man, mentioned,
63.
llosonbergii,(/c Man, mentioned,
63.
ruber, Hess., mentioned, 03.
rudis. Hell., mentioned, 03.
Savignyi,6^>e;!oe Bate, mentioned,
63.
scabrieulus, Uell., mentioned,
63.
sexdcntatus, Streets, mentioned,
63.
sinensis, Hell., mentioned, 03.
singalangen.si.s, Nohili, men-
tioned, 63.
eintangensis, ile Man, mentioned,
03.
sp., de Man, menticmed, 64.
spectabili.s, Uell., = lar, Fahr.,
mentioned, 64.
spinimauus, Milne-Edw., men-
tioned, 64.
sundaicus. Hell., mentioned, 0).
var. bataviana, de Man,
mentioned, 64.
var. brachydaetyla, Nohili,
mentioned, 64.
var. de Mani, yohili, men-
tioned, 64.
7L*
.-,12
l^DEX.
]'al;rmou superbus, Hell., 64.
Inu'lhis, S. Sm., 64.
(runqnebaricus, Fahr., 64.
Trompii, de Man, 64.
ustulatus, Nohili, 64.
vasus, Hell., — \aT, Fahr., 64.
Volleiihovenii, Herkl., 64.
Weberi, de Man, 64.
(Macrobracliium) Quelohi *,
de Man, .57; mentioned. GO.
I'ahumonidiB, 449.
rala^ornis, Vig., 24;!.
Palamcdea, Linn., 1M;3, 216, 217,
240, 257, 260, 262-26S ; intestinal
tract of, 201.
archecentre, 241, 256, 258.
cornnta, Linn., intestinal tract
in, as an arcliecentric type,
173, 175, 176.
I'alamedea", 201, 203.
Palamedeidae, 268.
Talicus, Phdij^in, 29.
.Tiikesii, White, 29. 49.
Jakesii, Bouvier, 29 ; mentioned,
31, 32.
serripes, Alcock cj' Anders., 32,
50.
serripes, Bouvier, ."52.
Whitei, Miers, 31 ; mentioned,
3, 50.
Whitei, Bouvier, 31 .
Panda, Great, upper carnassial teeth 1
of, 166.
Himalayan Long-tailed, KiO.
Pr.iidalidsB, 446. ,
Pandalus, Leach, 446.
inagnoculus, Spenee Bate, 44(!.
Pandion, Sav., 215, 260.
haliaetus, Linn., intestinal tract
of, 211, 212.
Pandioiiidoe, 211.
Pangolins, 393.
Papio, Er.vl., 405.
'• Paracalcarine sulcus," 3.32, 333.
Paradiseidae, 254.
Paradoxurus musanga, Gray, weight
of brain of, 421.
Paramithrax (Chlorinoides) aculeatus,
Milne-Edw., 38.
var. nrmatus, Miers, ,38.
Coppingeri, Haswell, 38.
Coppingeri, Miers, 38.
longispinus, De Haan, ftnote 38,
vemieosipes, Miers, 36.
I'araphysi.s, 456.
I'aratermiual bodies, 458, 497.
Paratymolus sexspinosus, Miers, 33. {
'• Parieto-occipital sulcus," 348, 376,
414, 418. I
Parra jacana, TAna., intestinal tract
of, 233, 234.
Parridte, 234, 2.35, 240, 241,
269.
Parrot, 243, 263.
Parthenolainbrus calappoides, Adams
cj- 117) ;^', 41.
coufragosus *, Caiman, 42.
Parthenope calappoides, Adams &
White, 41.
Passalus transrersus, Dalm., men-
tioned, 76.
Passeres, 2.52, 256-260, 270.
Passeriformes, 173, 252, 256, 263,
269.
Passerinae, 175, 264.
Passerine Nectarinlidse, 251.
Passerines, 211, 254, 268.
Pavo, Linn., 219.
cristatiis, Linn., intestinal tract
of, 219, 220.
Pedionomus, Gould, 217.
Pedunculata, 277.
Pelargo-Colvmbomorpbine Assem-
blage, 237, 239, 240.
Brigade, 257.
Brigade, evolution of the intes-
tinal tract in, 214.
, summary of, 173, 213.
metacentre, 258.
Pelargomorpha', 213.
Peleoanidaj, 193, 268.
Pelecanus rufescens, ZnM., intesliiial
tract of, 193, 194.
Pelicans, 190, 102, 193.
Pelidnota lajvissima, Burm., men-
tioned, 76.
Penaeus canaliculatus, Oliw, nieu-
tioued, 452.
Penelope, Linn., 219.
Penguins, 189.
Pentaspidai, 28S.
Perameles, Geoffr., 427, 428.
Pcrdix, Briss., 219.
Peregrine Falcon, 211.
Perodicticus, Benn., 322, .338, 339,
344-351, 412, 416-424.
potto, GmeL, 323.
Petrels, 189.
Petrorayzon, Linn., brain of, 500.
" Petrosal lobe " of cerebollam,
333.
" Petrosal lobule," 430.
Phaethon tlavirostris, Reichenb., in-
testinal tract of, 189, 190, 192.
Phaethontida3, 189, 268.
Phalacrocoracidse, 190, 192, 268.
Phalacroeorax carbo, Linn., intes-
tinal tract of, 190, 191.
Phaps, Selbii, 239.
Phascolarctos, Blaino., 383; brain
of, 465.
Phascolomys, Geoffr., 354, 404,
413.
Phasianus, Linn., 219.
Philoscia, LatreiUe, 136 ; mentioned,
103, 134, 13.5, 138.
mina, Budde-Lund, 136 ; men-
tioned, 137.
novae-zealandiae, Filhol, 138.
pubescens, Dana, 136 ; men-
tioned, 101, 102, 152.
violacea, Filhol, 124.
Philougria, Spenee Bate cj- Westw.,
mentioned, 122.
marina, Chilton, mentioned,
122, 128.
rosea, Koch, mentioned, 116.
rosea, Chilton, 115, 117.
rosea, Filhol, 115, 117.
Pliilijgria rosea, Thomson & Chilton,
115, 117.
Tliomsoni, Chilton, 118.
Phlogoeuas, 239.
Phoca, Linn., 411, 413; brain of,
374, 381, 382.
hi.spida, Er.vl., 391 ; brain of,
391.
Phoenioopteri, 263, 208.
Phoonieopteridae, 200.
Phcenicopterus, Linn., 190, 214. 26:?,
268, 270.
ruber, Linn., intestinal tract of,
200.
Phreatoicus, Chilton, mentioned,
112.
Phyllobranchiate Crustacea-Macrura,
On the New Zealand, by George
M. Thomson, 433-473.
Phymactis, Milne-Edw. Sf Jiaime,
mentioned, 298.
Phymodins monticulosus, Dana, 12.
sculptus, A. Milne-Edw., 12.
sculptus, Alcock, 12.
ungulatus, H. Milne-Edtv., II.
ungulatus, A. Milne-Edw., 11.
Pici, 252, 256, 257, 263, 269.
Picidae, 252.
Pigeons, 211, 221, 263, 267, 270.
INDEX.
513
J'ilodius spinijies, Heller, 12.
I'ihiinnus, Leach, menlioued, 1, 18.
cristipcs*, Caiman, 1" : men-
tioned, 1, 1!).
cursor, A. M'dne-Edw., 17.
dilatipes, Admitx <.\- White, men-
tioned, IS. I
Forskalii, Milne- FaIw., men-
tioned, 1 7.
lanatu.s (Latr.'?), Mier.f, 1 G.
longicornis, Hihjend., 1 fj.
nitidus, A. Milne-Edw., 14.
pulehcr, Miers, 15.
scabriusculus, Adams ij- iVJiile.
mentioned, 1 7.
serainudus, Jliers, If).
setifer, De Haan, li).
Shiiteri, de Man, mentioned, 17.
rinnipedia, 308, 421.
I'innotheres, Latr., mentioned, 48.
ripridse, 2.^2.
Pisa sf'ij-v, Milne-Edw., :}7.
I'isauridae, 67.
rithecia, Desm., 41G; brain of, 41.5;
weight of brain of, 420.
PittidjB, 252.
riatalea, Linn., 100, 194, 214.
leuoorodia, Linn., intestinal
tract of, 109. I
rieslonika, Spetice Bate, 446.
.semihfvis, Spence Bate, 446.
neuropil rions, A. MiJne-Edw., men-
tioned, 29. j
cristatipcs, A. Milae-Edif., men- '
tioned, 29.
spinipes, de Man, mentioned,
20.
riotus, Linn., 20.3, 268.
anhinga, Linn., intestines of,
190, 192. I
rooillopora brevicornis, Lam/.-., men-
tioned, 45.
ciEspitosa, Dana, mentioned, 45,
46.
elongata, Dana, mentioned, 45.
Poeock, 11. I., ilyriopoda and Araoh-
nida of Mount Eoraima in liritisli
Guiana, 64-71.
Podargidje, 250.
Podiceps, Lath., 263.
Podicipedes, intestines of, 184.
Podicipes cristatus, Linn., intestinal
tract of, 184, 185.
Podineraa, Wai/L, brain of, 485.
Pa'cilasma, Darivin, -bS; mentioned,
157.
Piceilasma carinatuin, //oe/.-. 157:
mentioned, 153, 161 .
lenticula, Auriu., mentioned,
288, 289.
minuta*, Gnmel, 288; men-
tioned, 289, 294.
Polistes annularis, J^Jiansson, 72 ;
mentioned, 71.
Pollicipina;, 290.
l'ulyaspid<T, 290.
PolyborinoB, 207.
Polyborus brasiliensis, (ImeL, intes-
tinal tract of, 207, '-'OO.
Pol_vdesmid;e, 64.
Polyopi.s, Hertwiij, mentioned, 30().
Polystomidium, Ihrtwiij, mentioned,
306.
Pontopbilus, Leach, 434.
australis, G. M. Thomson, 434 :
mentioned, 453.
gracilis, Spence Bate, 435.
Porcellana, LMm., mentioned, 45, 49.
Porcellio, Lair., 138: mentioned,
118. 140.
r/ranii/er, Wliite, 139.
laevis, lAitr., mentioned, 140.
neo-zelanicus, Thomas ij- Chilton,
141.
jjniinosns, Brandt, 141.
scaber, Lair.. 101, l(i2, 134,
139, 140.
var. imraacnlata, Budde-
Lund, mentioned, 140.
var. marginata, Brandt if-
liatzehurij, mentioned, 140.
Tar. marmorata, lirandt 4"
Jtatzehunj. mentioned, 134,
140.
Zealand ic as. White, 141.
Porcellis, read Porcellio, mentioned,
118.
Porphyrio ca'ruleus, VandcUi, 222.
poliocephalus. Lath., '2'2'2.
Porta, 462.
PortuHus {Thalamiia) (ircuatns. Do
Haan, 23.
" Posterior ealcarine fissure," 371.
" mesenteric or rectal factor,"
177.
" suprasylvian fissure," 39.3.
" Postsplenial sulcus," 387, 388.
" Postsylvian fissure," 395, 399.
"sulcus," 416.
" Precentral sulcus," 343, 416.
" Presylvian fissure," 395, 399.
" sulcus." 359, 406.
Primordium hippocam])i, 489.
Prionodactylus lencostictus •, Boa-
lene/er, 54 : mentioned, 53, 56.
Procavia, Ston; 309, 401, 407, 411,
413; brain of, 374, 375.
Procellariiformes, 2()3, 268; intes-
tines of, 173, 188, 213.
Processidae, 436.
Procyon, -Stoj-r, mentioned, 164-172.
cancrivorus, Cav., mentioned,
172.
Prodicticus potto, /.es.wn, weight of
lirain of, 420.
Propithecus, Benn., 338, 378, 40(;,
412, 417, 419 ; brain of, 321, 340,
350, 351, 352, 353, 354, 362.
Coquereli, A. MiJ.,e-Kdw., brain
of, 351, 3.52.
coronatus, A. Milne-Edn\, brain
of, 423.
diadema, Bennett, brain of, 321,
349, 350, 35] , 352.
Edwardsii, (Irandidier, brain
of, 352, 423.
Verreauxii, Grandidier, 352.
" Prorean sulcus," 359, 414.
Prosimia, BrLss., 425.
ProsiraiK, 354, .371, 3,72,391,406,
411, 422, 431 ; brain of, 367, 368,
374, 376.
Prosimian brain, 320, 321.
Protanthea simplex, Carlijren, men-
tioned, 301.
Protopterus, Owen, brain of, 495.
Psammosaurus, Filz., brain of, 484,
485.
Pseudocentric condition in intestinal
tracts of birds, 181.
Pseudomicippa, Hell., mentioned, 40.
nodosa. Hell., mentioned, 40.
tenuipes, A. Milne-Ediv., men-
tioned, 39.
variaus, J/i«/\s-, 39 ; mentioned.
40, 50.
Pscudophilyra tridentata, Mier.t, 26.
" Pseudosylvian sulcus," 410.
Pseudotantalus ibis, Linn., intestinal
tract of, 197, 198.
Pseudozius dispar, Dana, 14; men-
tioned, 16.
Psittaoi, 243,244,256,263, 265, 269.
Psittacid;e, 243.
Psittacus, Linn., 24.3.
Psophia, Liinn., 225, 241.
crepitans, Linn., intestinal tract
of, 225.
514
INDEX.
I'sopbia obscura, PeJ:., 225.
I'sophiinffi, 225, 241.
rterocles, Temm., 221, 239, 240,
241.
bieinctus, Teiwii., intestinal tract
of, 239.
Pterocletidie, 263, 269.
Pteroclo-Columbfe, 240, 257, 258.
Pteroclo-Columbine Assemblage, 239.
Pteroglossus Wiedi, Sturm, 252.
Pteroptochiflae, 252.
Pteropus, Briss., 383, 384, 391, 404 ;
braiu of, 329.
inedius, Temm., 384.
poliocephalus, Temm., brain of,
383, 384.
Ptilopus, StrickL, 239.
Pycnonotidoe, 252.
" Pyramis," 428.
Pyriform lobe, 329.
Pyrophorus noctiluca, JAn»., men
tioned, 70.
Quelcb, J. ,T., and McConnell, F. V.,
lleport on a Collection made at
Mount Iloraima in British Guiana,
51-76.
Rabbit, brain of, 410.
Raccoon, upper carnassial teeth of,
106.
Rails, 223, 224.
Kallida;, 2i'2, 223, 226, 241,
26S.
Ramphastos, see Rhampbastos.
Rana, Linn., brain of, 497, 499.
Ratita?, intestines in, 173, 175, 182,
216, 258, 263, 268.
Ratites, 201, 240, 263, 267.
Recessus opticus, 461.
superior of third ventricle, 464,
465, 466, 467, 468, 469, 470,
472, 489.
triangularis, 471 .
" Rectal sulcus," 414.
Rectum of birds, 174, 176, 178.
of Palmedea cornuta, 176.
Recurrent mesenteric factor. 1 77.
Recurvirostra, Lint)., 240.
avocetta, Linn., 231.
Red-throated Diver, 184.
Reduviidic, 72.
Reptiles of Mount Roraima in British
Uuiana, by U. A. lioulenger, 53.
Reptiles, Aberrant Commissure in
the Foreiirain of certain, 455-500.
" Retrocalcariue sulcus," 332, 333,
386, 390.
Rhamphastidae, 252.
Rhampbastos ariel, Vir/., 252.
vitellinus, Licht., 252.
Rhantus elegans, Wuterh., 74.
Rhea, Lath., intestines in, 173, 182.
Rhese, intestines in, 182, 183.
Rhinal fissure, 319.
RhinocbetidK, 227.
archecentric, 268.
Rhinochetus, Verr., 228, 241 .
Jubatus, Verr. S,- dcs Murs,
intestinal tract of, 227.
Rhipidomys, Wagn., mentioned, 52.
Macconnelli*, de Winton, 52.
microtis, Thos., mentioned, 52.
Rhynchocinetes typus, H. Milne-
Edw., mentioned, 433.
Rhynchotus, Spi.c, 216, 221.
rufescens, Temm., intestinal
tract of, 215.
Rbytidoceros plicatus, SJunr, in-
testinal tract of, 248.
Rodentia, 52, 403, 404, 413, 426;
brain in, 354.
Roebuck, brain of, 387, 389.
Roraima (Mount) : Report on a Col-
lection made by F. V. McConnell
and J. J. Quelch at Mount Roraima
in British Guiana, 51-76.
" Rostral sulcus," 414.
Saccosoma, Danielssen, mentioned,
94.
Sauropsida, 272.
Scalpellum, Leach, 290 ; mentioned,
153.
Darwini, Hoelc, mentioned, 153,
156.
giganteum*, Cr'no'e?, 153; men-
tioned, 157, 159, 160, 161.
Hoeki *, Gruvel, 290 ; men-
tioned, 294.
luridum, Auriv., mentioned.
292.
Peroni, Gray, mentioned, 294.
regium, Hoel-, mentioned, 153,
156.
Scorpio, Auriv., mentioned,
294.
sexcornutum. mishry, 293 ;
mentioned, 290.
ScalpeUum squamuliferum, WeUner,
mentioned, 154, 156.
trispinosum, Hoeh, mentioned,
294.
viliosum. Leach, mentioned, 294.
8celoporus, Wiegm., brain of, 475.
Schi/.ophrys aspera, Milne-Ediv., 39.
Scissure perpendiculaire esterne, 341 .
Scolopas, Linn., 235, 24(J, 258.
rusticola, Linn., intestinal tract
of, 231.
Scopidae, 195, 263, 268.
Scopus, Briss., 213, 228.
umbretta, Gmel., intestinal tract
of, 195.
Scorpiones, 68.
Screamers, 201.
Scyphacella, S. I. Smith, mentioned,
122, 123.
arenicola, S. L. Snrith, mentioned,
123.
Scypbacidffi, 121.
Sc>Ji>hacin(P, 121.
Scyphax, Dana, 122 ; mentioned,
102, 103.
Aucklandise, G. 2J. Thomson,
126 ; mentioned, 102, 152.
intermedins, Miers, 123 ; men-
tioned, 122, 124.
ornatus, Dana, 123; mentioned,
101, 102,122, 128,131,151.
setiger, Budde-Lund, mentioned,
122, 124.
Scyphoniscus*, Chilton, 127; men-
tioned, 103, 122.
waitatensis*, C7i(7/o)i, 128; men-
tioned, 101, 151.
Scythrops novfe-hoUandite, I^ath.,
242.
Sea-birds, 203.
Seals, brain of, 381, 388, 390, 392,
393, 399, 404, 411.
" Second temporal sulcus," 336.
Semnopitheci, 412.
Semnopithecus, Cuv., 405.
Seriatopbora, Lam., 50.
elegans, Mihie-Edw., mentioned,
ftnote 40.
hystrix, Dana, mentioned, ft-
note, 46.
imbricata, Bassett-Smilh, men-
tioned, 46.
spinosa, 21ihie-Edw. <.f- Ilaime,
mentioned. 45.
subulata, Lumh., mentioned. 4.j.
Serpentarius, Cuv., 212, 214.
IXDEX.
515
Serpentarius ro|iHlivoru3, Daud., in-
testinal tract of, 2(H].
f^harpe, K. Eowdlur, liirds of Mount
Roraima, British Guiana, 53.
Sheep, brain of, 386 ; cerebellum of,
4l'6.
Sideropora digitata, I'ldUis, men-
tioned, 4().
bystrix, Ihnid., mentioned, ft-
notc 4G.
palmata, de Biaitiv., mentioned,
4().
" Sillon calcarin," 388, 389.
Simla, Linn., brain of, 374, 393,
405.
satyrus, Linn., cerebellum of,
431.
SimiidfD, 415, 430.
Singing: birds, 263.
Sipunculus, Brandt, mentioned, 81.
nudus, Linn., mentioned, ftnote
SI.
Sirenia, 421 .
iSittidiB, 254.
Sloths, 369, 393.
Smew, 203.
Smith, Dr. G. Elliot. On the Mor-
phology of the Brain in the Mam-
malia, with Special Reference to
that of the Lemurs, Eecent and
Extinct, 319-432.
On the Morphology of the
Cerebral Commissures in the
Vertebrata, with Special Ee-
ference to an Aberrant Com-
missure found in the Forebrain
of certain Eeptilcs, 455-500.
Solvent glands of birds, 173.
Spatula, Bole, 215, 20(!.
clyjieata, Linn., intestinal tract
of, 203.
Spootyto, (.ihxjer, 248.
i<plurruzius dispar, Stirapsoii, 14.
Sphenisciformcs, intestines of, 173.
186, 188, 213, 263, 2(;s.
Sphenisous demersus, Linn., intes-
tinal tract of, 186, 187, 188.
Sphenodon, Lund, brain of, 455, 479,
480, 489, 492.
punctatum. Gray, brain of.
457.
Spheiiophorus hemipterus, Linn..
mentioned, 76.
SfJieriUo dan(F, Heller, 145.
inonolinn.'!, Dana, 148.
s^tiiiosw, Dana, 150.
Spizaetus coronatu>i, Linn., 208.
'• Splcnial sulcus," 381, 387.
'■ Spur eines Sulcus fronto-orbital,''
343.
Starnocnas, Bonaj)., 239.
Steatornis, I/um/i., 250.
Steatornithidtc, 250.
Steganopodes, 189, 19o, 193, 197,
263.
Stenocinopina-', 40.
Stenoplesictis Cayluxi, Filhol, brain
of, 400.
Stenops (Xycticebus), 323, 343,377,
389.
gracilis, Geoffr., 323, 343, 378.
javanicus, Illirj., 341, 343, 377.
tardigradus, Linn., 343.
Stcrcorarius, Brixs., 270.
arebecentric, 269.
crepidatus, Gmel., intestinal
tract of, 237.
Sterna hirundo, Linn., intestinal
tract of, 237.
Sternaspis, Otto, mentioned, 94.
Stone-Curlew, 233.
Storks, 197, 261.
Striges, 248, 256, 257, 263, 269.
Strigina>, 248.
Stringops, Gray, 243.
habroptilus. Gray, intestinal
tract of, 244.
Strix, Linn., 248, 249, 2.57.
Hammea, Linn., intestinal tract
of, 248.
Strongylium ha^morrhoidalis. Fair.,
mentioned, 76.
Struthio, Linn., 216.
Struthiones, intestines in, 182, 183.
SturnidiB, 254.
Stylodactylidoe, 452.
Stylodactylus, A. Mihie-Edir., 452.
discissipes, Spence Bate, 452.
orientalis, Spence Bate, 452.
Styloniscus, Dana, mentioned, 106.
gracilis, Dana, mentioned, 106.
magellanicus, Dana, mentioned,
106.
Subclamatorcs, 252.
Suboscines, 252.
Sula bassana, Linn., intestinal tract
of, 190, 191.
Sulcus limitans (hijipocarapi), 460,
: 469.
I Sulids, 190, 268.
I " Superior temporal sulcus,'' 342.
] Supra-tiecal kink, 188.
Supra-duodenal loop, 173, 184,
265.
" Suprasylvian fissure," 396.
" sulcus," 333, 358.
" , anterior," 399.
" , posterior," .399.
Sus, Linn., brain of, 374.
Swifts, 250.
Swimming-birds, 263.
"Sylvian fissure," 319, 344, 346,
356, a57, 358, 373, 395, 398, 402,
409, 412, 425.
Sylvian fissure, morjihology of, 405.
" Sylvian formation,'' 411.
Sympctrum gilvum, de Selys, 73;
mentioned, 71.
Syrnium, Sav., 248, 257.
aluco, Linn., intestinal tract of,
249.
Tachybaptes iluviatilis, Tunst., in-
testinal tract in, 184, 186.
Tadorna, Ftem., 203.
" Tsnia chorioidea," 491.
Talegallus, Less., 220, 260.
Lathami, Gray, intestinal tract
of, 218.
Tamandua, Seha, 332, 402-404 ;
brain of, 331 .
Tapirus, 7i/-MS., brain of, .'i(U,
362.
Tarsius, Siorr, 338, 346, 394, 412,
418-431; brain of, 321, 324, 367 -
381.
spectrum, Pallas, brain of, 367,
370.
Tealia crassicornis, Di.ron, mentioned,
315.
Teeth, left upper carnassial teetli
of: —
Cave-Bear, 166.
Great Panda, 166.
Himalayan Long-tailed I'aiida,
166.
Hya;narctu8, 166.
Eaccoon, 166.
Tela, 461, 464.
" Tcmiwro-occipital fissure," 341.
Tenebrionidae, 76.
Terns, 2&3.
Tetragonops Frantzii, Sdaler, •2'>2.
Tetralia glabcrrima, TIerhst, 20.
ylaberrima, Ortmann, 20.
nigrifrons, Dana, mentioned, 20.
Tetrao, Linn., 219.
516
INDEX.
Thalamita admete, fferhst, 23.
cteruleipcs, dc Man, mentioned,
22.
crassimana, i'ann, mentioned, 22.
Danaj, de Man, mentioned, 22.
prymna, Herhst, 22, 23.
Savigniji, A. Milne-Edw., 23.
sima, U. Milae-Edw., 23.
spinimaua, de Man, mentioned,
22.
Stimpsoni, A. Milne-Edw., men-
tioned, 22.
Thalassema erythrogrammon, Leuck-
arl if- R'tipijell, alimentary canal
in, 91.
neptuni, Gaertner, mentioned,
78-8.5 ; dorsal mesentery in,
92, 95.
vegrande, Lampert, mentioned,
94.
Tliaumactis, Fowler, mentioned, 298.
Thinocorida;, 233, 234, 235, 240,241,
269.
Tiiinocorys, E.fchsch., 237.
rumicivorus, Esdtsch., intestinal
tract of, 232, 233.
Thomson, George II., On the Xew
Zealand Pl]3llobranchiate Crus-
taoea-Macrura, 433-473.
Thylacinus, Temm., 383, 404, 413.
Thylacoleo, Owen, 413.
Tiarinia angusta, Dana, 41.
angvsta, de Man, 41.
spinirostris, Haswell, 41.
Timeliidie, 252.
Tinami, 241, 258.
Tinamidae, 240, 257.
Tinamiformes, 173, 215. 210. 25S,
203, 208.
Tinamus, Lath., 210, 227.
Titanethes albus, .SchimUe, mentioned,
102.
Todus viridis, Linn., 245.
Toucan, 200.
Trachyderes interruptus, Diq)., men-
tioned, 76.
Trapezia eymodoce, Herlst, 20.
ferrnginea, var. areolata, Dana,
19.
anohita, Ortmann, 19.
var. inormis, A. Mihw-
Edtv., mentioned, 19.
Treclialea, ThortU, 07.
JFacconnelli *, Fucock, 07.
Treron, Vieill., 239.
I'ricbechus, Linn., 411.
Trichodina, Ehrcnb., mentioned, 82.
Tricboglossida?, 243.
Tricboglossus, Vig. t|- Horsf., 243.
Trichoniseidse, 114.
Trichoniseoides,(S'rtj'«,mentioned, 11 9.
Tricboniscus, Brandt, 114 ; men-
tioned, 101-103, 118, 119, 120,
123, 128.
albidus, Budde-Lund , mentioned,
123.
Leydigii, J/a.i' Wchcr, mentioned,
110.
otakeusis *, Chilton, 117; men-
tioned, 115, 110, 151.
pbormianus *, Chilton, 115:
mentioned, 110, 117, 151.
pusillus, iJcrtHcZ^, mentioned, 116. ;
rosens, Koch, mentioned, 110. !
Tbomsoni, Chilton, 118; men-
tioned, 115, 151.
Trichosurus, Less., 383, 393.
Triclaria, C. Koch, 07.
Tringa, Linn., 231, 240.
alpina, Linn., intestinal tract of,
230.
" Triradiate sulcus," 400.
Triton, Laur., brain of, 498.
Trocbilida;, 251.
Troglodytidte, 252.
Trogon pueHa, Gould, intestinal tract
of, 252, 253.
Trogones, 252, 250, 257, 203, 200.
" Tubercle of the olfactory tract,"
329.
Tuberculum olfactorium, 489.
Turdidae, 252.
Turnices, 217, 240, 241, 257. 258,
208.
Turnix, Bonn, 203.
Dussumieri, Temm., intestinal
tract of, 217.
Turtur, iSelhii, 239.
Tylida), 120.
Tylocarcinus styx, Herhst. 37.
s(>j.r, Miers, 37.
Tylos, Latreille, 120.
neo/.elanicus *, Chilton. 101, 1 :.'0 :
mentioned, 151.
spinulosus, Dana, mentioned.
101, 121.
I'vmpanistria bicolor, Boudp., 239.
l'\ninnid;e, 252.
iaa ie'ragona, Ortmann. 24.
tetragonon, Herhst. l'4.
Cngulata,392, 398, 401-426; brain
in, 361, 382, 383.
" Ungulate pseudosylvian sulcus,"
402.
Uniradial apocentricity, 179.
Unpaired caecum of birds, 174.
L'pupa epops, Linn., intestinal tract
of, 247.
Upupidae, 247, 256, 257, 268.
Ursus, Linn., 163-172; brain of,
374, ftnote 382.
arctus, Linn., mentioned, 172.
melanoleucus, A. Milne-Edw.,
mentioned, 163.
pruinosus, iJ/y/A, mentioned, 107.
" Uvula" of cerebellum, 428.
Vallecula Sylvii, 328, 329.
Vanellus cayennensis, Gmel., 230.
vulgaris, Bechst., 230.
Yaranus, Merr., brain of, 485.
Yaruna litterata. Fair., 24.
litterata, Kingsley, 24.
" Yentriculus communis," 461.
'■ Vermis," 430.
Vertebrata, Morpbology of Cerebral
Commissures in (G. Elliot 8mitli),
455-500.
YeSjja annularis, Jobansson, 72.
Vespidie, 72.
Viutrix globulifera, Bucha.tsdim/ i.j'
Mifhdotii, mentioned, 297.
i/lobulifera, Verril], 29^.
Virbias hijidirostris, iliers, 443.
Vireonidai, 254.
Viverra, Linn., 400, 401, 407,
413.
tangalanga. Gray, weight of
brain of, 421.
Viverridai, 398, 401, 413, 414.
Vultures, 267.
Vulturida=, 206, 214.
Waders, 203.
"Waterhouse, C. A., Coleojjtera of
Mount lioraima in British Guiana.
74-70.
White-tailed Sea-Eagle, 208.
Winton, W. E. do, Mammals of
Mount Iloraima in British Guiana,
i
Wuodpeckers, 263.
INDEX.
r>v
Xii/i/ho oetixlcntalii'!, H. ^filiic-
Jichv., 0.
(l^eptoiliu.s) exaratus, //. Milae-
Eihi\, nu'iilionod, '2, (J.
( ) c.varalits, Alcock, 0.
Xdiitliodes (jraiiusumanus, Dana, 1 1 .
Lamarckii, H. Mihie-Edii'., 11 ;
inentioncd, 2.
Xc'iioearcinus tubcrculatus, 11 Jiili , '.]-i.
Xiphooaris, v.jii Murtcnn, 417. /aloiihus, G'lU, t'tiioto 382.
couipressa, /// //<(f(», lu'MiLioiied, Zi):int.hiH sociatiis, Ellis, mentioned,
449. 312, 316.
oiii-vh-ostris (IL^ller), <!. M. Zonotricbia, 5fH'/<(/(«., mentioned, 53.
Thomson, 447 : mi'iitioiied. MaRCOnnelli *, Botvdkr Sharpf,
449, 4.-.3. 53.
L'longata, Garrin., mcntionod, pileata, Bodd., mentioned, 53.
449. ZostoropidLL', 254.
ERKATA.
Page OS, linos 1." and 10 IVom top, for Acroso.ma Scukieiieksii, Pcrty, ivad Acroso.m\ Schreiueusii, I'crty.
Page 71, line 2 from bottom, for Siuiictmin ijilviuit read Sympiti-nin (/ih'ii a.
Page S9, line 3 from bottom, /')/• Memoirs of the Acad, of A'ienna, 1852, vol. ii., n'ad Denksohr. k. Akad. '^'iss.
Wien, Hd. iv. Abt. 2(LS52).
Page 101, line 10 from top, for Aniindilh) i-eijulosiis { = Cuhai-is jv'^iJo,s»a', Miers) read Armadillo riijidosus
( = Ciibari\' rui/iilosus, iliers).
Page l<il, line In from top. for Cuhnria riqiilosuf, Miers, read Cnharis rmjiilosHs, Miers.
Page ini, line 11 from bottom, for Li/i/ia read Lvjia.
Page 1 Is, line 5 from bottom, for Porccllis read I'orcelllo.
Page 12S, line .5 from bottom, for Mrsannadillo Alhiandl, Dollfus, read Mesarmadillo Allaaadi, Dollfus.
Piige 107, line 1(.( from bottom, for .Elaroiniif read .Elvras.
Page 1 07, line 9 from bottom, for absence read presence.
Page 168, lines 17 and 18 from top, omit mention of Hyceaarctus, the humerus of which is unknown.
I'age 184, line 2 from bottom, fiU' Tachiilidjilin iliiviatilh read Tachi/hiiplusjliivinlihs, Tun.st.
Pago 180, lino 11 from bottom, for Tarhiihaptcs Jlnvlotilh read Tach>jhiipli's jlm-i'dUi!', Tunst.
SECOND .SJ'IUICS. — ZOf)l.()(;V, VOL. VIII.
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Part VII. 1901 1 8 0
Part VIII. 1902 0 4 0
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Part X. 1903 1 0 0
Part XI. 1903 0 6 0
Part XII. 1903 0 10 0
Part XII I. Index. . . " 2 9
IX. Part I. 1903 0 9 0
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