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NOVITATES
ZOOLOGICAE
A JOURNAL OF ZOOLOGY
Founded by
LIONEL WALTER, LORD ROTHSCHILD
and continued by
THE TRUSTEES OF THE BRITISH MUSEUM
Vol. 42, 1940-1948
LONDON :
PRINTED BY ORDER OF THE TRUSTEES
BRITISH MUSEUM (NATURAL HISTORY)
CROMWELL ROM), IONDON, S.W.7
I94O I948
NOVITATES ZOOLOGICAE
Vol. 42, 1940-1948
Tin.' parts of Volume 4? were issued as follows : —
Part I, containing pages i-2i(> and 13 plates, issued 26 April. 1940.
Pari II, containing pages 217-396, issued 251 1 1 October, 1940,
Part III, containing pages 397-508 and 4 plates, issued 20th March, 1948,
ERRATUM.
Page 93. "METRACRINIA gen. nov." should read "METACRINIA gen. nov.
CONTENTS OF VOLUME 42 (i 940-1948)
PART I.
PAGJ -
The Australasian Frogs of the Family Leptodactylidae. By II. \V.
Parker. (Plate I) i— 106
Neotropical Neoempheria (Diptera, Mycetophilidae). By F. W. Edwards.
(Plates I-II) . . . . . . . . . 107—130
Sour- Sapromyzid«e (Diptera) from the Solomon Islands and New
1 aledonia. By J. R. Malum h. (Plate I) .... 131 — 146
Studies on Diopsidae (Diptera). By J. F. Shillito. (Plate I) . . 147 — 1(>4
A Revision oi the Osmylid Subfamilies Stenosmylinae and Kalosmylinae
(Neuroptera). By D. E. Kimmins. (Plates I VIII) . . . 165- 202
On Australian Pamborini, Ozaenini and Scaritini (Coleoptera, Carabid te).
By M. Banningek ......... 203—214
Cercaria imbricata Looss 1S96, nee 1893 \ Note on Nomenclature. By
Miriam Rothschild ......... 215- 216
PART II.
A Monographic Revision of the Mexican Water Beetles of the Family
Elmidae. By H. E. Hinton ....... '217 — 396
PARI III.
Entomological Results from the Swedish Expedition, 10.54. to Burma
and British India — Coleoptera: Curculionidae. By Sir G. A. K.
Marshall . . . . . . . . . -397—473
A Synopsis of the Genus Mecedanum Erichson (Coleoptera. I olydiidae).
By H. E. "Hinton i; i Is 1
Notes on the Asteroids in the British Museum (Natural llistoi\
I. The Species of Astropecten. By I). Dilwyn John. (Plates 1 IV) (.85 508
NOVITATES
ZOOLOGICAE
A JOURNAL OF ZOOLOGY
hounded by
LIONEL WALTER, LORD ROTHSCHILD
and continued by
THE TRUSTEES OF THE BRITISH MUSEUM
l*
^
\9*0
^ Vol. 42
PART I, pp. 1-2 16
WITH THIRTLtH PLATES
26 April, 1940
LONDON :
PRINTED BY ORDER OF THE TRUSTEES
BRITISH MUSEUM (NATURAL HISTORY)
CROMWlill ROAD, LONDON, S.W. 7
1940
Price Fifteen Shillings
NOTICE
The Journal, Novitates £oologicae, was founded in 1894 by the
late Lord Rothschild as the official publication of his Museum
at Tring, and Lord Rothschild himself was joint editor until his
death in 1937. In September, 1938, the Trustees of the British
Museum assumed control of the Tring Museum, which had been
bequeathed to them by Lord Rothschild. In the meantime the
Journal was carried on under the joint editorship of the Hon.
Miriam Rothschild and Dr. K. Jordan until the close of Vol. 41.
Beginning with this part, the first of Vol. 42, the Trustees of
the British Museum have undertaken the publication of Novitates
Zoologicae, and the Journal will be edited at the British Museum
(Natural History), Cromwell Road, London, S.W. 7.
C. FORSTER-COOPER,
Director.
British Museum (Natural History),
London, S.W. 7 ,
12 April, 1940.
*•****«
:>^°
NOVITATES ZOOLOGICAE
Vol. 42 APRIL, 1940 No. 1
THE AUSTRALASIAN FROGS OF THE FAMILY
LEPTODACTYLIDAE
By H. W. PARKER, M.A.,
Department of Zoology, British Museum (Natural History).
(With one plate and twenty text-figures.)
THE first frog of this family to be named and described was the Rana australiaca
of Shaw, which was renamed Rana spinipes by Schneider a few years later.
The first to be referred to a definite family was Crinia georgiana Tschudi, described
in 1838 and placed by its describer in the family Cystignathi. As additional
species were discovered in later years they were referred to an ever-increasing
number of families according to the views of the times, and it was not until 1865
and 1868 when Cope published his papers on the classification of the Anura that
our present system began to take shape. Cope's disposition of the genera known
to him (1865, 1866a and b) was : —
Bufonidae Chelydobatrachus (=Myobatrachus).
Pseudophryne
Asterophrynidae Cryptotis (=Adelotus)
Cystignathi Pseudes Mixophyes
Ceratophydes Chiroleptes (=Cyclorana)
Criniae Crinia
Hyperolia ( = Uperoleia)
Helioporus
Neobatrachus ( =He!ioporus)
Borborocaetes (inc. Limnodynastes)
Boulenger's classification in the Catalogue (1882) differed but slightly from
this ; the various subfamilies of Cystignathi were dropped and Cryptotis was
also included in this family, renamed Cystignathidae ; Notaden was added to the
Bufonidae and the newly described Batrachopsis ( = Lechriodus) was referred to
the family Pelobatidae. The propriety of retaining the toothless forms in a
family (Bufonidae) distinct from the toothed genera (Cystignathidae) was criticized
from time to time, but Boulenger's arrangement persisted until 1922 when Noble,
following up an earlier piece of work by Nicholls (1916), advocated the union of
the Cystignathidae and Bufonidae under the latter name and the transference of
Lechriodus to this family from the Pelobatidae. Later (iq.ji) the same author
modified this arrangement by subdividing the Bufonidae into a number of sub-
families, to one of which, the Criniinae, all the Australasian genera were referred.
NOVIT. ZOOL., 42, I I
-
2 H. \V. PARKER
It was admitted that this subfamily was but poorly defined, and could hardly be
satisfactorily differentiated from some of the Neotropical groups of the same
family.
Noble's conception of the " Bufonidae " has not met with universal approval,
and herpetologists in general have shown a tendency to subdivide it into two,
Bufonidae (corresponding to Noble's subfamily Bufoninae) and Leptodactylidae
(s. Cystignathidae or Ceratophryidae), containing his subfamilies Criniinae,
Heleophryninae, Rhinophryninae, Pseudinae (= Ceratophyinae Parker, 1935),
Elosiinae and Leptodactylinae. But the principle of brigading the Australasian
genera with others from the Neotropical region and from South Africa as one
large family has not been seriously criticized.
It has long been realized that the limits and mutual relationships of the
Australian genera were very inadequately understood, and the present work is
an attempt to remedy this deficiency in so far as it is at present possible. More
and larger collections are still required and much more detailed anatomical work
is still necessary. But a larger amount of material has been available to the
present author than to any other previous worker (nearly 1300 specimens, including
types of 59 names) , and its study has emphasized the necessity for a reconsideration
•of a number of previously accepted beliefs. As far as the family and subfamily
status is concerned, it has become clear that the Australasian genera fall into two
groups, not absolutely differentiated, but tending to grade into one another and
confined to that zoogeographical region. They have been given subfamily status
and are referred to in the following notes as the Cycloraninae and Myobatrachinae.
These names have been selected in accordance with the principles laid down in
Articles 4 and 5 of the International Rules of Zoological Nomenclature (cf.
Parker, 1934 : 15). The first genus of the former subfamily whose name was
made the basis of a family or subfamily name was Chiroleptes (Chiroleptina
Mivart, 1869) ; this generic name unfortunately has to be displaced by Cyclorana
and the subfamily name must be changed accordingly (Art. 5). The name
Myobatrachinae, from Myiobatrachina Bonaparte, 1850, long antedates the name
Criniinae used by Noble (from Criniae Cope, 1866).
The relationships of these two Australian subfamilies to the South African
and Neotropical members of the Leptodactylidae cannot be satisfactorily
discussed until a comparative survey of the whole family has been completed.
Their mutual relationships can best be considered after a brief survey of the
various anatomical and morphological characters by which they are characterized
Internal Cranial Morphology.— This has only been studied in a single species,
Crinia georgiana (du Toit, 1934). During recent years a good deal of attention
has been directed by morphologists and anatomists to the cranial morphology
of the Anura and attempts have been made to modify the existing scheme of
classification on their findings. But it must be stressed that far too few species
have yet been examined for any satisfactory generalizations to be possible and
the variability of many of the characters has not been investigated. Du Toit
gives the following summary of characters noticed in Crinia georgiana : —
(1) The alary forms the lateral support of the entire lateral wall of the
apertura externa and vestibulum.
(2) The crista subnasalis is absent.
(3) The anterior undivided part of the crista intermedia is much bigger
than in Rana.
(4) The posterior vestibular " wulst " is absent.
(5) The plica obliqua is suspended from the tectum nasi.
AUSTRALASIAN LEPTODACTYLIDAE 3
(6) The infundibulum possesses an extra antero-ventral extension.
(7) A well developed recessus sacciformis is present.
(8) The cavum medium, being more posteriorly situated than in tin-
European Rana, does not share in the formation of the recessus sacciformis.
(9) The ductus nasolacrimalis opens into the roof of the horizontal
part of the infundibulum.
(10) The recessus lateralis attains an enormous size.
(11) The plica isthmi is absent.
(12) The intermaxillary gland opens into the buccal cavity by two main
ducts only.
(13) The septomaxillary consists of an anterior unpaired portion which
has two posteriorly-directed processes.
(14) The connective tissue between the latter bone and the lamina
inferior tends to disappear.
(15) The vomer (prevomer) is divided, consisting of an anterior edentulous
and a posterior dentigerous portion.
(16) The os en ceinture is paired.
(17) The foramen for the IVth nerve is situated in front of the foramen
opticum.
(18) The arteria carotis cerebralis enters the skull through a separate
foramen.
(19) The foramen perilymphaticum superius does not communicate
with the jugular foramen.
(20) The fronto-parietal fontanelle is large.
(21) There is only one dorsal fenestra in the chondocranium.
(22) The annulus tympanicus is sickle shaped.
(23) The hyomandibular branch of the trigeminal nerve is ventrally
situated to the plectrum.
(24) The posterior part of the operculum is weakly perichondrally
ossified.
(25) The articular region differs from that of Rana.
(26) Crinia possesses a bursa angularis oris.
(27) The hyale is confluent with the otic capsule.
Many of these characters are of widespread occurrence throughout the Anura
and can have little or no taxonomic importance, but others again (e.g. 23) may
be peculiar to the species or genus. Those which have been investigated in
more than one species of the subfamilies under consideration will be considered
separately.
Premaxillae. — These show no great diversity of structure ; they are denti-
gerous in the Cycloraninae except Notaden, but are edentulous in Uperoleia
(part), Metacrinia, Glauertia(past), Pseudophryne and Myobatrachus of the other
subfamily. In males of Adelotus the teeth are somewhat enlarged.
Maxilla.^ -This bom- is usually deeper than normal in the Cycloraninae
where it is dentigerous except in Notaden ; in some species of Cyclorana, notably
( austraiis, its outer face is heavily sculptured. In the Myobatrachinae the bone
is relatively less deep and is toothless in those genera where the premaxilla is
also edentulous (q.v.),
Septomaxillary. — Apparently present in all genera of both subfamilies ; its
variation has not been investigated.
Prevomer. — In the Cycloraninae this bone is large and entire with a strong
posterior portion bearing teeth (text-figs. 4-9). In the Myobatrachinae, however,
H. W. PARKER
— pr.br.
-prces.
Text-fig. i. — Hyoid and larynx of Heleioporns albopunctatus (after Trewavas). ar.
= arytaenoid; ca.cv. = cartilage of vocal chord ; c.l.p. = posterior constrictor
laryngis ; cr. = cricoid ; g.hy. = genio-hyoid muscle ; hgl. = hyoglossus
muscle; mem.h.a. = hyo-arytaenoid membrane ; o.hy. = omohyoideus muscle ;
pet.hy.a. = anterior petrohyoid muscle ; pr.br. = bronchial process ; pr.oes. =
oesophageal process ; p.v. = pulvinar vocale ; st.hy. = sternohyoid muscle ;
tend.c.l.p. = tendon of posterior constrictor laryngis.
-pr.ces.
Text-fig. 2. — Hyoid and larynx of Limnodynastes peronii (?) (after Trewavas).
= cardiac process. Other abbreviations as in text-fig. I.
pr.card.
AUSTRALASIAN LEPT0DACTYL1DAE
it is greatly reduced in size, is often divided into two, with the posterior, primitively
dentigerous, portion disappearing and teeth vestigial or entirely absent (text-figs.
13-15). In this latter subfamily a transitional series can be traced from species
such as Crinia laevis, in which vomerine teeth are still present and the bone is
entire (text-fig. 14), to forms such as Pseudophryne, in which both the anterior and
posterior portions may be completely lost. But in no member of this subfamily
is the bone as large, or the teeth as numerous, as in any member of the Cycloraninae.
There is no absolute correlation between loss of maxillary and vomerine teeth.
In the Cycloraninae maxillary teeth are lost only by Notaden, where vomerine
^pr.ces.
Text-fig. 3'. — Hyoid and larynx of Crinia signifera J. Membranous wall of larynx
removed, but vocal chords left in place. (After Trewavas.) cv. = vocal
chord ; g.hy.med. = median geniohyoid muscle ; st.hy.d. = dorsal part of
sternohyoid muscle. Other abbreviations as in text-fig. 1.
teeth persist ; in the Myobatrachinae vomerine teeth are absent in Uperoleia
rugosa whilst maxillaries still persist. There does appear, however, to be a
correlation between the vomerine condition and the tongue, with which also the
hyoid and intermaxillary glands are related (q.v.). Teeth, in order to function,
need a firm surface to bite on. In the case of the vomers this surface is the
tongue, and consequently it is not surprising to find that where the tongue is
large and broad, as in the Cycloraninae, the vomers are well developed, whereas in
the Myobatrachinae, which all have a much smaller, narrow tongue, the vomerine
teeth are obsolescent. It has been noticed in the Microhylidae (Parker, 1934 :
6) that there is a similar correlation between the presence of a large broad tongue
and an unreduced prevomer.
Palatine. — This bone is present throughout both subfamilies and shows no
important variation.
Ethmoid. — This bone is entire in the Cycloraninae, but may be divided, as in
1 Text-figs. 1-3 reproduced by permission of the Royal Society of London.
b H. W. PARKER
the family Microhylidae, in some members of the Myobatrachinae, e. g. Uperoleia,
Pseudophryne and Crinia. Not infrequently it is continued forwards as a peri-
chondral ossification in the tectum, solum and septum nasi (cf. Cyclorana
australis, text-fig. 5).
Fronto-parietals. — In the majority of the genera of both subfamilies these
bones are very small, laterally disposed and widely separated from one another.
In Cyclorana, Lechriodus, Mixophyes and Adelotus of the one subfamily they form
a median suture, but in the Myobatrachinae the genus Uperoleia alone lacks a
fronto-parietal foramen.
The Lower Jaw. — Shows no important modifications except in the genus
Adelotus where a pair of very large, tusk-like odontoids are developed anteriorly
in the male sex. The function of these structures is not known, but it is
interesting to note that a similar development has occurred in certain African
Ranids (Dimorphognathus, Petropedetes and Phrynodon) and in some Asiatic species
of the genus Rana.
Hyoid and Larynx. — These structures have been fully investigated by
Trewavas (1933), but in only a limited number of species — Heleioporus albo-
punctatus, Limnodynastes peronii, Crinia signifera and Mixophyes fasciolatus.
When due allowance is made for the fact that the only specimen of the latter
species which was examined was a juvenile, it appears that these four species
fall into two clearly-definable groups, corresponding to the two subfamilies, and
characterized thus :
A. Cycloraninae (Heleioporus, Limnodynastes and Mixophyes).
(1) Alary process narrow proximally, but expanded distally (text-figs. 1
and 2).
(2) Cricoid complete.
(3) Oesophageal process of the cricoid more or less slender.
(4) M. omohyoideus present.
(5) Mm. sternohyoideus and petrohyoidei attached at the lateral edges
of the hyoid plate.
B. Myobatrachinae (Crinia).
(1) Alary process of the hyoid a wing-like expansion of almost the whole
lateral margin of the hyoid plate, without narrow stalk (text-fig. 3).
(2) Cricoid incomplete.
(3) Oesophageal process of the cricoid broad and short, or almost absent.
(4) M. omohyoideus absent.
(5) Mm. sternohyoideus and petrohyoideus anterior inserted on the
ventral surface of the hyoid, reaching the middle line in the posterior part of
this insertion.
Certain of these characters, notably the loss of the omohyoideus and the
division of the cricoid, are believed to have little taxonomic significance for they
occur many times in the Anura in completely unrelated forms, but the shape of
the alary processes and the attachment of the sternohyoideus and petrohyoidei
appear to be correlated with other characters and may have considerable taxonomic
value. Thus the broad wing-like, sessile alary process is found in all the Myo-
batrachinae (Uperoleia, Crinia, Metacrinia, Glauertia, Pseudophryne and Myo-
batrachus examined), whilst the pedunculate type is common to all the Cycloraninae
(Cyclorana, Lechriodus, Mixophyes, Limnodynastes, Heleioporus, Philoria, Adelotus
and Notaden examined).
AUSTRALASIAN LEPTODACTYLIDAE 7
All the genera appear to lack the apical cartilage which is a characteristic
feature of most Ranids and Rhacophorids.
Ear. — This organ is present and well developed in all the genera of both
subfamilies except Pseudophryne, where the tympanum, annulus tympanicus,
cavum tympani, columella (plectrum) and Eustachian tube are absent. This
condition of the auditory apparatus occurs in many unrelated genera of the
Anura so that too great reliance cannot be placed upon it as a guide to phylogeny.
The facts that this condition is normal in the Urodela and that the fully-developed
anuran ear passes through a similar stage in its ontogeny1 suggest that the condition
may be a primitive one, and it is possible that its sporadic reappearance in so
many unrelated Anuran genera is a manifestation of a neotenic tendency.
The external and middle ear show some variations in different genera. The
annulus tympanicus is generally incomplete dorsally ; in the highly modified
burrowing genera there is a tendency for the extra-plectal (columellar) cartilage
to increase in size, and in Myobatrachus it forms a large thickened pad, as large
as, and closely apposed to, the whole of the tympanum. The size and distinct-
ness externally (due to thinning of the overlying skin) of this latter organ are very
variable, the general tendency being towards smaller tympana, more deeply
seated (partly covered by the m. depressor max. inf.) and protected by thickened
. skin in cryptozoic forms.
Tongue. — This organ is either subcircular or very broadly oval in the Cyclo-
raninae, or relatively long and narrow, sometimes quite small, in the Myobatra-
chinae. These two types of tongue are associated with a different disposition of
the openings of the intermaxillary glands (q.v.), and also with those differences in
the hyoid and prevomer which have already been mentioned. It seems highly
probable that all these differences are intimately correlated and that they are
concerned with the method of feeding. There appear to be no essential differences
in the nature of the food taken by the two subfamilies ; in fact the most specialized
genera of each, Notaden and Myobatrachus respectively, parallel one another
closely, for both are essentially myrmecophagous and specially adapted for life
in termitaria. Yet Notaden has a broad, subcircular tongue, vomerine teeth, a
long series of ducts from the intermaxillary gland, long, pedunculate alary processes
of the hyoid and sternohyoideus and petrohyoideus muscles attached laterally
to the hyoid plate, whereas in Myobatrachus the tongue is a small, narrow organ,
and all the associated structures of the hyoid region and intermaxillary gland are
concentrated towards the middle line, and vomerine teeth are wanting. The
prey appears to be the same in both genera, yet the mechanism by which it is
captured is very different.
Intermaxillary Gland. — E. Miiller (1932) has given a general account of the
disposition of these glands and their ducts in the Anura and has classified them into
five main groups. Representatives of only three genera of the present subfamilies
were examined by him, Pseudophryne, Limnodynastes and Uperoleia, and lie notes
that these three fall into twogroups. In Uperoleia an.1 Pseudophryne the openings
of the gland are concentrated near the centre line into two ducts, whereas in
Limnodynastes the openings of the gland are numerous (14) and form an irregular,
transversely oval patch, occupying a space almost equal to the inter-choanal
width. A survey of the gland in almost every species of the two Australian
subfamilies lends no support to Miiller's classification into five main categories, but
shows that there is, as might have been expected, a distinct correlation between
1 (hi Toit and de Villiers, 1932, S. Afr. /. Si ('., 29 : 449-4115.
8 H. W. PARKER
the disposition of the ducts and the shape of the tongue. The function of the
gland being to furnish the sticky secretion which enables the tongue to pick up
food, a broad tongue will need a widely spread series of ducts for the even distri-
bution of the secretion over its tip, whilst with a narrow tongue a corresponding
concentration of the ducts towards the centre-line is essential for efficiency.
This is what obtains in fact. In the broad-tongued Cycloraninae there are
numerous ducts arranged in a more or less regular transverse series over a width
almost, or quite, equal to the interchoanal space. Sometimes they form a regular,
continuous linear series which may even open under a single long fold of the
mucous membrane ; more frequently, however, they are arranged in three groups
on a transverse line, but every gradation between the two extremes can be found.
In the Myobatrachinae there are never more than two ducts, opening close
together near the middle line, and not infrequently the two open under a common
fold which results in the opening appearing single ; both conditions may be
found within the one species.
Shoulder Girdle. — This structure is relatively stable in the arciferal families,
as compared with the firmisternal, where reduction or loss of the clavicle and
procoracoid has taken place on numerous occasions. In the Australian Lepto-
dactylidae very few important structural differences have been noticed. Clavicles,
procoracoids and coracoids are invariably present and well developed ; the
omosternum may be absent, but when present is always small and cartilaginous;
the sternum also is never ossified, though in old individuals of some of the larger
forms there may be some degree of calcification. Myobatrachus alone shows any
great divergence from the normal, and here there seems tohavebeen a strengthening
of the girdle against lateral compression by a broadening of the clavicles and the
procoracoid region, whilst the coracoids have assumed a more oblique position
and are scarcely larger or stouter than the clavicles (text-fig. 20) . The modification
recalls, though to a less extent, the conditions to be found in the African Ranid
HemisHS which is also a cryptozoic genus. A secondary result of the widening
of the mesial ends of the clavicles and procoracoids is that the epicoracoids meet
edge to edge for about a third of their length and so approach an arcifero-
firmisternal condition.
In Crinia, and especially in Crinia haswelli (text-fig. 15a), the procoracoids
extend anteriorly beyond the clavicles, and their form strongly suggests that
the omosternum is homologous with the anterior portion of these cartilages.
Anatomists cannot agree as to the homologies of this structure in the Anura (de
Vos, 1938 : 56), but it seems highly improbable that they are correct in
believing it to be of different origin in various species of the same genus.
Vertebral Column. — As in most of the arcifera the vertebrae are pro-coelous,
but a peculiar and primitive feature of both subfamilies is the incomplete fusion
of the intervertebral condyle with the vertebra and the partial persistence of
the notochord ; in Cyclorana and Lechriodus only is the notochord completely
lost and the condyle firmly ankylosed to the vertebra. The normal number of
8 presacral vertebrae is found throughout the Myobatrachinae, but in the Cyclora-
ninae fusion of the first and second vertebrae occurs in five genera — Limnodynastes,
Heleioporus, Philoria, Adelotus and Notaden. The sacral diapophyses are some-
what dilated in all the genera examined, except Mixophyes, and the urostyle
articulates by two condyles.
Thigh Muscles. — The disposition of the muscles of the thigh, and particularly
the relation of the distal tendon of the m. semitendinosus to the mm. graciles,
has been shown (Noble, 1922) to be of considerable assistance in elucidating
AUSTRALASIAN LEPTODACTYLIDAE
anuran relationships. In all the Procoela it has been claimed that the semi-
tendinosus is separate from the sartorius and that its distal tendon is either
ventral to, or very rarely pierces, the gracilis major and g. minor. It is within
the present subfamilies that the rarer condition has been reported. In Limno-
dynastes ornatus the tendon of the semitendinosus pierces the actual gracilis major
and minor muscles, and in Pseudophryne australis a further stage in the inward
migration of the tendon is found, where it perforates, not the muscles themselves,
but their ligamentous head. The examination of further material reveals that
the inward migration of the tendon of the semitendinosus progresses ever further,
and that in the final stages it has passed dorsal to the gracilis entirely, and so
attained the condition hitherto regarded as characteristic of the diplasiocoelous
nrmisternia. In the two subfamilies a complete gradation from one extreme to
the other can be traced, but the ventral, presumably more primitive, condition is
only found in the Cycloraninae, whilst the dorsal condition is confined to the
Myobatrachinae. The various conditions can be roughly subdivided and classed
into four groups thus :
II.
III.
IV.
1 Noble, 1922, classes this species under the following section, but the examples examined
by the author show it to have a thigh-muscle complex very similar to that of Vpero'eia.
10 H. W. PARKER
Pectoral Muscles. — Jones (1933) has investigated the pectoral musculature of
Cyclorana aus traits, Limnodynastes tasmaniensis, L. peronii and Uperoleia mar-
morata in a survey of the pectoral myology of the Salientia. He finds that the
supracoracoideus profundus, found in all these four species, is characteristic of
the arcifera. The episternohumeralis, a variable muscle in the firmisternia, is
also present in the four Australian species as well as in all the other arcifera except
the genera Bufo and Rhinophrynus. The sternoepicoracoideus, a new muscle,
is found in Cyclorana and Limnodynastes but is absent from Uperoleia. In
Cyclorana it arises from the antero-lateral edge of the sternum and is inserted by
a narrow tendon into the dorsal surface of the epicoracoid ; similar conditions
are reported in the Discoglossidae and South American Leptodactylidae (Pleuro-
dema). In the two species of Limnodynastes it arises partly from the first
myocomma of the rectus abdominis as well as the sternum and so approaches
the condition noted in the genus Hyla (H. arbor ea and H. rubra), where the
muscle arises from the myocomma only.
Pupil. — A good deal of confusion has been caused at various times through
incorrect descriptions of pupil-shape. This is usually to be accounted for by
preservation, the degree of contraction, or distortion having masked the true shape.
In the majority of genera of both subfamilies the shape appears to be \y, but
irregularities of contraction may result in either the ventral or lateral angles
becoming unduly emphasized. In Mixophyes alone is it truly a vertical ellipse.
Viscera. — Hoffman (1931) has described certain features of the viscera, e.g.
disposition of lungs, liver-lobes, relations of stomach to duodenum, and the arrange-
ment of the rugae of the lining of the intestinal canal in Crinia georgiana. An
attempt is made to utilize some of these characters in distinguishing the South
African genus Heleophryne from the Cystignathidae (Leptodactylidae), but too
little is at present known of the variability of these organs for it to be possible to
assess their taxonomic value.
Consideration of these facts makes it evident that the two subfamilies are
clearly differentiated by the tongue and the associated structures such as prevomers,
vomerine teeth, the hyoid apparatus and intermaxillary glands. Other associated
characters, such as the vertebral condition and the thigh muscles, also lend
support to the view that the two groups represent different evolutionary lines.
But in these latter characters and in others, there is a complete intergradation
between the two, suggesting a not very remote common ancestry. It is, of
course, conceivable that the Myobatrachinae are not a natural assemblage, but
are a group of forms derived from the Cycloraninae by the evolution of the same
type of feeding-mechanism on more than one occasion. If the feeding-mechanisms
of the two were intimately associated with different foods such a view might have
something to recommend it ; but in fact no such difference appears to exist and,
as has already been pointed out, representatives of each group (Notaden and
Myobatrachus) appear to live under very similar conditions. Unfortunately
there is no evidence other than the morphological which will assist in elucidating
the relationships of the two subfamilies to one another and to the rest of the Anura.
The only fossils known which might throw any light on the subject are the Eocene
frogs of the Intertrappean beds of Bombay which have been variously referred to
Rana, Oxyglossus and Indobatrachus. If Noble's interpretation (1930) be correct,
then Indobatrachus should be referred to the Myobatrachinae ; it appears to have
8 presacral vertebrae, with the notochord persistent, and vomerine teeth in very
small groups, arguing a reduced prevomer. Unfortunately, as in most fossil frogs,
AUSTRALASIAN I.EPTODACTYLIDAE II
the difficulties of correct interpretation are very great, so that the reference of
this fossil genus to the Myobatrachinae must be regarded as somewhat uncertain,
and it provides no clue to the relationship of this subfamily with the Cycloraninae.
If correctly referred it indicates that the subfamily persisted in the Oriental
region long after the Australian members of the same group had become isolated,
and provides additional evidence, if such be necessary, that the whole family
Leptodactylidae at one time had a more northerly distribution.
The descriptive methods and terminology used in the following pages conform
to standard practice. The dimensions given are for the most part maxima, since,
with animals which continue to grow after the attainment of sexual maturity, it
is not practicable to give the actual range of adult size with any accuracy. Only
when size has been invoked as a specific or subspecific criterion has an attempt
been made to give ranges and means. In these instances the ranges are based on
obvious adults, i.e. females with distended ovaries containing pigmented ova
and males with secondary sex-characters. Such a series will naturally contain
a greater proportion of old individuals than of those which have only just reached
maturity, and consequently both the range and mean will err on the large side.
Only very approximate geographical ranges are given, though no doubt much
interesting information would be forthcoming were the geographical ranges of
the various species to be plotted and compared with maps showing climatological,
botanical, physiographical, geological and other data. But such maps, to be of
value, must be accurate and reasonably complete — criteria which cannot at
present be fulfilled even approximately.
During the course of this work so many herpetologists have assisted with
advice, information or material that to thank them all individually would need
too great a space, whilst to select a few would be invidious. It is, however, essential
to proffer thanks to Professor G. E. Nicholls, to whom modern amphibian taxonomy
owes so much. The collections he made in West Australia and presented to the
British Museum provided the focal point for much of the work, and arguments and
discussions with him have given the author numerous pointers and saved him
from many egregious errors. The bulk of the material, other than that in the
British Museum, has been received from the Museums in Amsterdam, Harvard,
Leiden, Perth (W.A.), Stockholm and Sydney; to the authorities in these
institutions the author wishes to express his great indebtedness.
REFERENCES.
Boulenger. G. A. 1882 Cat. Batr. Sal. Br, I. Mus.. ed. 2.
Cope, E. D. 1865 Nat. Hist. Rev., n.s., 5 : 97 -120.
1866 (a) J. Acad. nat. Sci. Philadelphia, (2), 6 : 67-97.
1866 (6) torn. cit. : 189—206.
Hoffman, A. C, 1931 S. A fir. J. Sci., 28 : 399-407.
Jones, E. J. 1933 Ann. Mag. nat. Hist., (10), 12 : 403-420.
Mivart, St. G. 1869 Proc. zool. Soc. Lond. : 288-294.
Muller, E. 1932 Morph. Jb., 70 : 131-216.
Nicholls, G. E. 1916 Proc. Linn. Soc. Loud.. 128 art. 1. : 80-92.
Noble, G. K. 1922 Bull. Amer. Mus. nat. Hist., 46, art. 1 : 1-88.
1930 Amer. Mus. Novit., 401 : 1 3.
1931 Biology of the Amphibia. New York.
Parker, 11. W. 1934 Monogr. I-'am. Microhylidae. London.
1935 Proc. zool. Soc. Lond. : 511.
du Toit, C. A. 1934 Proc. zool. Soc. Lond. : 1 eg -141.
Trewavas, E. 1933 Phil. Trans, roy. Soc. Lond., 222, b : 401-527.
Tschudi, J. J. von 1838 Mem. Soc. neuchdtel. Sci. nat. 1839, 2 : i-99-
DeVos, C. M. 1938. Anal. Anz., 87 : y, iu
12 H. W. PARKER
Subfamily CYCLORANINAE.
Raniformes (part) Dumeril & Bibron, 1841, Erpe't. Gen., 8 : 317.
Ranae (part) Fitzinger, 1843, Syst. Rept. : 31.
Cvstignathidae (part) + Discoglossidae (part) + Alytidae (part) Giinther, 1858, Cat.
Batr. Sal. Brit. Mus. : 26, 34, 37.
Asterophrydidae (part) + Scaphiopodidae (part) + Cystignathidae (part) Cope, 1865,
Nat. Hist. Rev., n.s. 5 : 97-120.
Asterophrydidae (part) + Cystignathidae, Pseudes (part) + Cystignathidae, Cera-
tophrydes (part) + Cystignathidae, Criniae (part) Cope, 1866, J. Acad. nat. Sci.
Philad., (2), 6 : 67-97.
Ranidae (part) + Discoglossidae (part) + Alytidae (part) Steindachner, 1867, Reise
Novara, Zool., Amph. : 7-34.
Cystignathidae (part) Keferstein, 1867, Nachr. Ges. Wiss. Gbttingen, 18 : 343.
Cystignathidae (part) + Discoglossidae + Alytidae (part) Keferstein, 1868, Arch.
Naturgesch., 34 : 251-273.
Alytidae, Uperoliina (part) + Ranidae, Cystignathina (part) + Discoglossidae,
Chiroleptina (part) and Asterophrydina (part) Mivart, 1869, Proc. zool. Soc. Land. :
288-294.
Cystignathidae (part) + Bufonidae (part) + Pelobatidae (part) Boulenger, 1882, Cat.
Batr. Sal. Brit. Mus., ed. 2 : 183-432.
Bufonidae (part) Noble, 1922, Bull. Amer. Mus. nat. Hist., 46 : 1-87.
Ceratophriidae (part) + Bufonidae (part) Waite, 1929, Rept. Amph. S. Australia : 244-266.
Bufonidae, Criniinae (part) Noble, 1931, Biol. Amph. 1496.
Leptodactylidae (part) Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 8.
Tongue broadly oval or subcircular, large. Intermaxillary glands with
numerous ducts arranged either in a long, transverse, linear series or in a trans-
versely oval patch or in three groups on a transverse line. Prevomer always
present, entire and well developed, with its posterior process bearing a long series
of teeth (text-figs. 4-9). Alary processes of the hyoid pedunculate (text-figs. 1-2) ;
sternohyoid and petrohyoid muscles attached to the lateral edges of the hyoid
plate. Distal tendon of the m. semitendinosus usually passing ventral to the
mm. gracilis, more rarely perforating them. Sternoepicoracoideus present
{Cyclorana, Limnodynastes) . First and second vertebrae fused in genera where
the notochord is persistent, free where the notochord is lost.
Key to Genera.
Maxillary teeth present.
A. Pupil vertical ; no fronto-parietal foramen ; 8 presacral vertebrae ;
sacral diapophyses not dilated. Toes webbed, the membrane
penetrating between the outer metatarsals . . Mixophyes.
B. Pupil horizontal ; digital webbing not penetrating between the outer
metatarsals.
(1) No fronto-parietal foramen in adults.
(a) First finger opposed to the remainder, the second very
short ; vomerine teeth between the choanae ; 8 presacral
vertebrae Cyclorana.
(b) First finger not opposable to the remainder ; vomerine teeth
behind the level of the choanae.
(i) Toes with small terminal discs ; no dentary pseudo-teeth ;
8 presacral vertebrae .... Lechriodus.
(ii) Toes without terminal discs ; male with 2 large, fang-
like, dentary pseudo-teeth ; first and second vertebrae
fused Adelotus.
AUSTRALASIAN LEPTODACTYLIDAE 13
(2) A large fronto-parietal foramen ; 1st and 2nd vertebrae fused
(a) Vomerine teeth between the choanae . . Heleioporus.
(b) Vomerine teeth behind the level of the choanae
(i) Vomerine series moderately extensive, extending laterally
beyond the inner borders of the choanae . Limnodynastes .
(ii) Vomerine series short and oblique, not extending laterally
beyond the inner borders of the choanae . . Philoria.
II. Maxillary teeth absent ; a large fronto-parietal foramen ; 1st and 2nd
vertebrae fused ; vomerine teeth small .... Notaden.
MIXOPHYES Gunther.
Mixophyes Gunther, 1864, Proc. zool. Soc. Land. : 46 (Type species — Mixophyes fasciolatus);
idem, 1864, Ann. Mag. not. Hist., (3), 14 : 311 ; Cope, 1866, /. Acad. nat. Sci. Philad., (2),
6 : 89, 93 ; Steindachner, 1867, Reise Novara, Zool., Amph. : 10 ; Keferstein, 1868, Arch.
Naturgesch., 34 : 254 ; Boulenger, 1882, Cat. Balr. Sal. Brit. Mus., ed. 2 : 188 ; Cope, 1889,
Bull. U.S. nat. Mus., 34 : 311 ; Nieden, 1923, Das Tierreich, Anura I : 518 ; Noble,
1931, Biol. Amph. : 497.
Myxophyes Cope, 1863, Nat. Hist. Rev., n.s., 5 : m.
Hyla de Vis, 1884, Proc. roy. Soc. Queensland, 1 : 128.
Maxillary teeth present. Prevomer entire, bounding the choana anteriorly
and laterally with an anterior projection towards the maxilla and a mesially-
directed, dentigerous branch which does not reach the palatine ; fronto-parietals
large, forming a median suture. Ear fully developed. Vertebrae procoelous ;
8 presacrals ; sacral diapophyses cylindrical ; coccyx articulating by two condyles.
Text-kig. 4. — Anterior cranial elements of Mixophyes fasciolatus juv. X 5. (From beneath.)
Omosternum cartilaginous ; sternum cartilaginous, entire posteriorly. Terminal
phalanges simple.
Distal tendon of the m. semitendinosus passing ventral to the tendon of the
m. gracilis. The hyolaryngeal apparatus has only been examined in a juvenile
specimen and Trewavas (1933 : 512) considers such material unsuitable for
comparison ; nevertheless it is apparent that the apparatus is Heleioporus-,
rather than Crinia-kkc, with a complete cricoid, m. omohyoideus present, and
the mm. sternohyoideus and petrohyoidei attached at, or close to, the lateral
edges of the hyoid plate.
Pupil vertical. Tongue subcircular, entire or emarginate, and scarcely free
behind. Toes webbed, the membrane penetrating between the outer metatarsals.
Mixophyes fasciolatus fasciolatus Gunther.
Mixophyes fasciolatus Gunther, 1864, Proc. zool. Soc. Loud.: 46, pi. 7, 1 1 j-; . 1 (Type locality :
— Clarence River, N.S.W.) ; idem, 1864, Ann. Mag. nut. Hist., mi. 14 : ^12 ; Krettt. 1867,
Cat. Industr. Prod. N.S.W. , Add. : 107 ; Steindachner, 1867, Reise Novara, Zool., Amph.:
14 H. \V. PARKER
10 ; Giinther, 1868, Proc. zool. Soc. Lond. : 479; Keferstein, 1868, Arch. Nalurgesch., 34 :
255. pl- 5. fig- 6 I Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 188 ; idem, 1885,
Ann. Mag. nat. Hist.. (5), 16 : 387 ; Fletcher, 1889, Proc. Linn. Soc. N.S.W., (2), 4 :
372 ; idem, 1890, op. cit., (2), 5 : 669-671 ; idem, 1892, op. cit., (2), 7 : 18 ; idem, 1894,
op. cit., (2), 8 : 529 ; Lucas and le Souef, 1909, Anim. Austral. : 267, fig. ; Harrison,
1922, Aitst. Zool., 3, 1 .'34; Nieden, 1923, Das Tierreich, Anura 1:518, fig. 350;
Trewavas, 1933, Phil. Trans, roy Soc. London, 222, b : 438, fig. 28.
Myxophyes fasciolatus Krefft, 1865, Pap. Proc. roy Soc. Tasmania: 16.
Mixophyes fasciolatus fasciolatus Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 10.
Hyla fenestrata de Vis, 1884, Proc. roy. Soc. Queensland, 1 : 128 (Type locality: — Tweed
River).
Head broader than long. Snout rounded, not prominent, 1-4 to 1-5 times as
long as the eye with obtusely angular canthus rostralis and oblique, slightly
concave loreal region ; nostril equidistant from the eye and the end of the snout ;
interorbital space equal to, or a little broader than, the width of an upper eyelid ;
tympanum very distinct, vertically oval, its horizontal diameter A to § that of the
eye. Fingers slender, the first a trifle longer than the second, which is shorter
than the fourth ; subarticular tubercles well developed on the metacarpo-
phalangeal joints only, with a smaller supernumerary tubercle proximal to each ;
two metacarpal tubercles. Toes two-thirds webbed, the edge of the membrane
midway between the third and fourth toes being level with the distal subarticular
tubercle of the third ; three phalanges of the fourth toe free from web ; sub-
articular tubercles moderate ; an elongate, oval inner, but no outer, metatarsal
tubercle. Tibio-tarsal articulation reaching the nostril or beyond the tip of
the snout.
Skin smooth above and beneath ; a curved supratympanic fold ; anal region
feebly granular.
Brown or olive above, the dorsum usually with scattered, irregular polygonal
darker markings ; a curved dark line from the tip of the snout, through the nostril,
along the canthus rostralis and along the supratympanic fold, often spreading over
the tympanum ; a dark interorbital bar which may be prolonged backwards as a
triangular or T-shaped marking. Flanks dark-spotted. Limbs with numerous
narrow dark cross-bars which are most defined on the concealed surfaces ; hinder
side of the thighs dark-mottled. Lower surfaces white, the gular region of the
male dotted and stippled close to the lower jaw.
Length from snout to vent : <J 63 mm. ; $ 97 mm.
Male with a diffuse nuptial pad on the dorso-lateral surface of the first
finger and edge of the inner metacarpal tubercle ; a vocal sac. Embrace
axillary ; spawning apparently in spring or autumn in New South Wales
(embrace in April, Harrison ; tadpoles in advanced stage of development
December, Fletcher).
The species is diurnal, frequenting the banks of creeks in deep, shady gullies
and taking readily to the water when alarmed.
Distribution : New South Wales east of the dividing range and southern
Queensland.
Specimens Examined.
AUSTRALASIAN LEPTODACTYLIUAE 15
Mixophyes fasciolatus schevilli Loveridge.
Mixophves fasciolatus Andersson, 1916, K. Svenska VetenskAkad. Handl., 52, 9 : 7.
Mixophyes fasciolatus schevilli Loveridge, 1933, Occ. Pap. Boston Soc. nat. Hist., 8 : 55.
(Type localities :—Millaa Millaa and Lake Barrine, Atherton Tableland, and 4000 ft.
Bellenden Ker Range, Queensland) ; idem, 1935, Bull. Mus. comp. Zool. Han.,
78 : 11.
Similar in general characters to the typical form, but with the toes £ webbed,
(only two phalanges of the fourth and none of the fifth being free) and with the
bars on the limbs tending to coalesce to form fewer, broader, bands.
Length from snout to vent : o DI mm- ; ? 83 mm.
Distribution : Atherton Tableland, Bellenden Ker Range, and Malanda,
Queensland. Intermediates showing intergradation with the typical form are
said to occur in the area from the Richmond River, N.S.W., to the Bunya
Mountains of southern Queensland.
CYCLORANA1 Steindachner.
Alyies (>ray. 1842, Zool. Misc., 2 : 56.
Chiroleptes (non Kirby, 1831) Gunther, 1858, Cat. Batr. Sal. Brit. Mus. : 34 (Type species : —
Alytes australis Gray); Cope, 1865, Nat. Hist. Rev., n.s., 5 : 108; idem, 1866, /. Acad.
nat. Sci. Philad., (2), 6 : 89, 93 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 267 ,
Cope, 1889, Bull. U.S. nat. Mus., 34 : 312 ; Noble, 1931, Biol. Amph. : 497.
Cheiroleptes (emend.) Spencer, 1901, Proc. roy Soc. Victoria, (2), 13 : 176.
Cyclorana Steindachner, 1867, Reise Novara, Zool., Amph. : 29. (Type species : — Cyclorana
novaehollandiae) .
Phractops Peters, 1867, Mber. Akad. Berlin : 30 (Type species : — Phractops alutaceus) ; Ogilby,
1907, Proc. rov Soc. Queensland, 20 : 32 ; Nieden, 1923, Das Tierreich, Anura I : 520 ;
Waite, 1929, Rep. Amph. S. Austral. : 245.
Mitrolysis Cope, 1889, Bull. U.S. nat. Mus., 34 : 312 (Type species: — Chiroleptes alboguttatus
Gunther) ; Nieden, 1923, Das Tierreich, Anura I : 524; Noble, 1931, Biol. Amph. :
947-
Maxillary teeth present. Prevomer large, bounding the choanae anteriorly,
with an anterior process which may meet the premaxilla, and a short, posterior,
dentigerous portion between the choanae, not overlying the palatine. Fronto-
parietals forming a median suture, their lateral borders raised to form a distinct
ridge. Ear fully developed. In adult individuals the dorsal surfaces of the skull
are often somewhat rugose and in P. australis a considerable amount of secondary
rugose bone is present on the nasals, premaxillae, maxillae, fronto-parietals and
squamosals, the zygomatic process being greatly enlarged and forming a broad
arcade across to the maxilla. Palatines with more or less distinctly raised ridges
and prominences ventrally. Vertebrae procoelous, the condyle completely
ankylosed and the notochord not persistent ; sacral diapophyses slightly dilated ;
urostyle articulating by two condyles; 8 presacral vertebrae. Omosternum
well developed ; sternum large, cartilaginous, bifid posteriorly ; clavicles strongly
curved. Terminal phalanges simple.
1 As Ogilby (1907, loc. cit.) has pointed out, the name Chiroleptes, so frequently used i"i
this genus, is a homonym and must be dropped. Hut his selection of Phractops to replace
it cannot be maintained. This name was proposed by IVter.s at a sitting of the Prussian
Academy of Science on January 10, 1867, and was not published until later 111 that year ,
the January I left of the Monatsbericht was actually not received at the library oi Gottingen
1 'mversity until .May. Hut Steindachner's account of the Amphibia of the Novara 1 ollection,
in which appears the name Cyclorana, was actually published and laid on the table of the
Vienna Academy on the same day that Peters read his paper (January 10) (ef. Ant. Akad.
wiss. Wien, 4 : 11).
1 6 H. W. PARKER
Distal tendon of the m. semitendinosus passing ventral to the mm. graciles.
Alary processes of the hyoid expanded distally, but arising as narrow stalks
proximally, similar to those of Heleioporus.
Pupil horizontal but with a downwardly-directed ventral angle. Tongue
subcircular, partly free behind. Toes more or less webbed. Second finger short,
the first opposable to the remainder.
The species of this genus fall into two clearly-marked groups as regards their
general habitus. First there are slender species with thin digital webbing of the
type usually associated with " grass-frogs " or aquatic species. Of these inermis
shows very little trace of the opposable inner digit and has two metatarsal
tubercles ; dahlii has the digital characteristic of the genus well developed and
has lost the outer metatarsal tubercle whilst alboguttatus has the inner metatarsal
Text-fig. 5. — Anterior cranial elements of Cyclorana australis. x 2. (From beneath.)
tubercle strongly compressed and so approaches the second group of species,
which are all of more or less globose habitus, and are typical fossorial species
with a shovel-shaped metatarsal tubercle and webbing, when present, of a thick
fleshy nature.
Synopsis of the Species.
I. Inner metatarsal tubercle not shovel-shaped ; habitus slender.
A. Toes f webbed ; two metatarsal tubercles ... C. inermis.
B. Toes fully webbed ; a single metatarsal tubercle . . C. dahlii.
II. Inner metatarsal tubercle shovel-shaped.
A. A straight dorso-lateral fold on each side of the back
(1) Zygomatic process of the squamosal heavily sculptured and
forming a broad suture with the maxilla. Habitus stout
C. australis.
(2) Zygomatic process not sculptured and separated from the maxilla
or only very narrowly in contact with it. Habitus slender
C. alboguttatus.
B. No definite dorso-lateral fold.
(1) Toes less than half webbed.
(a) Web midway between the 3rd and 4th toes not extending
beyond the proximal subarticular tubercle of the fourth
toe ; metatarsal tubercle shorter than its distance from the
tip of the inner toe ; a strongly marked colour-pattern
C. brevipes.
AUSTRALASIAN LEPTODACTYLIDAE 17
{b) Web midway between the 3rd and 4th toes reaching nearly to
the level of the distal subarticular tubercle of the third ;
metatarsal tubercle as long as, or longer than its distance
from the tip of the inner toe ; colour pattern indistinct
C. cultripes.
(2) Toes § webbed, or more . . . . . C. platycephalus.
Cyclorana inermis (Peters).
Chiroleples inermis Peters, 1867, Mber. Akad. Berlin : 30 (Type locality: — Rockhampton) ;
Keferstein, 1868, Arch. Naturgesch., 34: 267 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus.,
ed. 2 : 271 ; Andersson, 1913, K. Svenska VetenshAkad. Hand!., 52, 4:16; idem,
1916, K. Svenska VetenshAkad. Handl., 52, 9:12.
Phractops inermis Nieden, 1923, Das Tierreich, Anura I : 524.
Habitus slender ; head as long as broad. Snout pointed, slightly prominent,
1-5 to 1-6 times as long as the eye, with rounded canthus rostralis and slightly
oblique, feebly concave loreal region ; nostril midway between the eye and the
end of the snout ; interorbital space as broad as, or a little narrower than the
upper eyelid; tympanum very distinct, f to § the diameter of the eye. Fingers
moderately long, free, the first distinctly longer than the second, but the latter is
better developed than in the other members of the genus, and the opposition
of the first digit to the remainder is not well marked ; subarticular tubercles very
prominent ; two large metacarpal tubercles and rows of smaller tubercles on the
palm. Toes f webbed, the membrane midway between the third and fourth
extending beyond the distal tubercle of the third but not to the middle tubercle
of the fourth ; subarticular tubercles prominent ; a prominent oval inner, and a
smaller rounded outer, metatarsal tubercle ; a dermal fold or slight fringe along
the inner side of the inner digit and tarsus. Tibio-tarsal articulation reaching
the centre of the eye or between this point and a little beyond the tip of the
snout.
Skin with scattered warts above ; belly and hinder side of thighs finely
granular ; throat and chest smooth.
Greyish brown above with scattered, indistinct darker mottlings ; edge of
the upper lip with white flecks ; hinder side of the thighs spotted or marbled
with dark brown and white ; tibiae cross-barred ; lower surfaces immaculate
white except the throat of the male, which is marbled with brown.
Male with a vocal sac and a diffuse nuptial pad on the inner side of the
first finger.
Length from snout to vent : $ 35 mm.
Distribution : Queensland (Rockhampton, Kimberley, Torrens Creek).
Specimens Examined.
Cyclorana dahlii (Houlenger).
Chiroleples dahlii Boulenger, i8gd.Proc.zool.Soc.Lond. : 867, pi. 40. fig. 2 (Type locality : —
Daly River, N. Australia) ; Fletcher, 1898, Proc. Linn. Soc. N.S.W., 22 : 682, 684.
Phractops dahlii Nieden, 1923, Das Tierreich, Anura I : 522 ; Loveridge. 1935, Bull. Mm.
comp. Zool. Ham., 78 : 11.
NOVIT. ZOOL., 42, I 2
l8 H. W. PARKER
Habitus slender ; head as long as broad. Snout rounded, not prominent,
i'5 times as long as the eye,1 with rounded canthus rostralis and very oblique,
slightly concave loreal region ; nostril equidistant from the eye and the end of
the snout ; interorbital space narrower than the upper eyelid ; tympanum very
distinct, its horizontal diameter £ that of the eye. Fingers moderate, the first
much longer than the second and opposed to the remainder ; subarticular
tubercles moderately distinct, but the two metacarpals not well defined. Toes
fully webbed, the membrane extending to some extent between the metatarsals ;
subarticular tubercles not well marked ; an oval inner, but no outer, metatarsal
tubercle ; a slight fold along the inner edge of the metatarsus. Tibio-tarsal
articulation reaching the loreal region.
Skin feebly shagreened above ; a distinct curved supratympanic fold and a
median nuchal furrow ; lower surfaces smooth, except the hinder side of the
thighs, which are feebly granular.
Brownish-olive above, with a very faint, lighter, dorsal streak ; a dark
canthal stripe ; flanks and hinder side of the thighs spotted and marbled with
white. Lower surfaces immaculate white, except the sides of the throat and the
limbs, which are dotted with dark brown.
Male with a vocal sac.
Length from snout to vent : <$ 70 mm. ; 9 67 mm.
Distribution : Northern Territory.
Specimen Examined.
B.M. 95. 1 1. 14. 24 $ Daly River, N.T. Dahl. Cotype.
Cyclorana australis (Gray).
Alytes australis Gray, 1842, Zool. Misc., 2 : 56 (Type locality: — North coast of Australia).
Chiroleptes australis Giinther, 1858, Cat. Batr. Sal. Brit. Mus. : 34 ; Krefft, 1865, Pap.
Proc. roy Soc. Tasmania: 17; idem, 1867, Cat. Industr. Prod. N.S.W., Add. : 107;
Giinther, 1867, Ann. Mag. nat. Hist., (3), 20 : 54 ; idem, 1868, Zool. Rec, 4 (1867) : 145 ;
Keferstein, iStS.Arch. Naturgesch., 34: 267; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus.,
ed. 2 : 269 ; Boettger, 1894, Denkschr. med.-naturw. Ges. Jena, 8 : 109 ; Fletcher, 1898,
Proc. Linn. Soc. N.S.W., 22 : 678 ; Andersson, 1913, K. Svenska VetenskAkad. Handl.,
52, 4 : 15 ; Harrison, 1922, Aust. Zool., 3, 1 : 34.
Phractops australis Fry, 1914, Rec. W. Aust. Mus., 1 : 204 ; Nieden, 1923, Das Tierreich,
Anura I : 523, fig. 355 ; Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 13.
Cyclorana novaehollandiae Steindachner, 1867, Reise Novara, Zool., Amph. : 29, pi. 2, figs.
7-10 (Type locality : — Rockhampton) ; Keferstein, 1868, Arch. Naturgesch., 34: 267.
Phractops alutaceus Peters, 1867, Mber. Akad. Berlin : 31 (Type locality : — Rockhampton).
Mitrolysis alboguttatus (part) Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 13.
Upper surfaces of the premaxillae, maxillae, fronto-parietals and squamosals
strongly sculptured, the zygomatic process of the latter forming a broad suture
with the maxilla. Habitus stout ; head depressed, as broad as long in juveniles,
but increasing in breadth with age so that in adults it is very much broader
than long, more so in females than in males. Snout depressed, not prominent,
175 (adults) to twice (juveniles) as long as the eye, with rounded canthus rostralis
and very oblique, concave loreal region ; nostril equidistant from the eye and
the end of the snout or, more usually, a little nearer the latter ; interorbital
space equal to, or a little narrower than, the upper eyelid ; tympanum distinct,
slightly more than half, rarely §, the diameter of the eye. Fingers moderate ;
1 Loveridge's (1935 : 13) contention that Boulenger measured the snout from the
nostril appears to be without foundation, anil his assertion that the first finger may, in this
species, be shorter than the second may indicate a traumatic condition or confusion of
species.
AUSTRALASIAN LEPTODACTYI.I DAE 19
the first much longer than the second and opposed to the remainder; subarticular
tubercles prominent ; palm with small circular tubercles ; two metacarpal
tubercles. Toes J webbed, the edge of the membrane midway between the third
and fourth normally reaching halfway between the distal subarticular tubercle
of the third toe and the proximal tubercle of the fourth ; rarely it may reach
almost to the former of these points or only just beyond the latter ; subarticular
tubercles distinct ; a very large, shovel-shaped inner metatarsal tubercle, but
no outer ; a slight fold along the inner edge of the tarsus. Tibio-tarsal articula-
tion reaching the eye, or, in larger specimens, the tympanic region.
Skin regularly shagreened above, often with larger scattered warts ; a
distinct median occipito-nuchal groove ; a strong, curved, supratympanic fold
from which a short, straight lateral fold may branch ; a distinct dorso-lateral
fold on each side of the back ; sometimes a distinct gland between the angle of
the mouth and the insertion of the fore-limb. Belly and hinder side of the thighs
finely granular ; throat almost smooth in females, but distinctly granular in
males.
Brown or grey above with some darker markings, of which the most constant
are a cantho-temporal streak, a vertical bar beneath the eye and darker smudges
along the sides of the dorso-lateral folds which are themselves white in juveniles.
Flanks and hinderside of the thighs dusky with lighter mottlings (not circular
spots as in alboguttatus) . Lower surfaces white, the chin and throat blotched
with brown in females and juveniles, uniformly infuscate in breeding males.
Male with a vocal sac, and, at the breeding season, diffuse nuptial asperities
on the dorso-lateral surface of the inner finger and a small, but distinct, button-
like prominence where the columella auris abuts against the tympanum. Breeding
in West Australia appears to take place about midsummer.
Length from snout to vent : c? 87 mm.1 ; $ 97 mm.
Juveniles at metamorphosis, 27 mm.
Distribution : Northern West Australia, Northern Territory and Queens-
land.
Specimens Examined.
Nicol Bay, W.A. du Boulay.
Roebuck Bay, W.A. Dahl.
Port Darwin. Buckland.
Port Essington, N.T. Gilbert. Cotypes.
Alexandria Sta., N.T. Stalker.
Queensland. Thomson.
Rockhampton. Darnel.
Westwood, nr. Rock- Wilkins.
hampton.
Cooktown, Q. Bellenden Ker.
Port Denison, Q. Krefit.
Port Denison. Darnel.
" Australia." Krefit.
Alexandria Sta.. N.T. Stalker.
Rockhampton, Q. (Godeffroy Mus.) {Co-
type oiPhractopsalu-
ten , us Peters.)
Queensland. (GodetTroy Mus.)
MowlaDown, Kimber- Soderberg.
ley Div., W.A.
1 I he " male" 27 mm. long reported by Andersson (1913) proves on re-examination
to be a juvenile just through metamorphosis, with an incompletely rcsorbed tail.
20 H. W. PARKER
Cyclorana alboguttatus (Giinther).
Chiroleples alboguttatus Giinther, 1867, Ann. Mag. nat. Hist., (3), 20 : 54 (Type locality : — ■
Port Denison, Cape York) ; idem, 1868, Proc. zool. Soc. Lond. : 480 ; Boulenger,
1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 270, pi. 18, fig. 1 ; Fletcher, 1898, Proc. Linn.
Soc. N.S.W., 22, 1897 : 678.
Chiroleples albopunctatus (? lapsus calami) Fletcher, 1894, Proc. Linn. Soc. N.S.W., 1893,
(2), 8 : 525, footnote.
Mitrolysis alboguttatus Cope, 1889, Bull. U.S. nat. Mus., 34 : 312; Nieden, 1923, Das
Tierreich, Anura I : 524, fig. 356; Loveridge, 1935, Bull. Mus. comp. Zool. Harv.,
78 : 13 (part).
Habitus slender ; head as long as broad or only slightly broader than long.
Snout conical, slightly prominent, 1-8 to 2-0 times as long as the eye, with rounded
canthus rostralis and very oblique loreal region ; nostril equidistant from the
eye and the end of the snout ; interorbital space as broad as, or a little narrower
than the upper eyelid ; tympanum distinct f to § the horizontal diameter of the
eye. Fingers slightly depressed, with lateral " seams," the first much longer
than the second and opposed to the remainder ; subarticular tubercles prominent;
palm with small circular tubercles and two moderately well-developed metacarpal
tubercles. Toes nearly half webbed, the edge of the membrane midway between
the 3rd and 4th toes, reaching nearly to the level of the distal tubercle of the 3rd ;
subarticular tubercles moderately prominent ; a moderately large compressed
inner, but no outer, metatarsal tubercle ; a slight tarsal fold. Tibio-tarsal
articulation reaching the eye or the nostril.
Skin with some scattered warts above ; a curved supratympanic fold and
a distinct straight dorso-lateral fold on each side of the back ; a more or less
distinct median occipito-nuchal furrow. Belly and hinder side of the thighs
finely granular ; throat and chest smooth.
Olive or brown above, with some obscure black mottlings ; a dark canthal
streak ; flanks and hinder side of the thighs with numerous, closely set, circular
white spots ; edge of the upper lip brown, dotted with white. Lower surfaces
white, the throat and chest usually marbled and freckled with brown.
Male with a vocal sac opening by a slit on each side of the tongue and a
diffuse nuptial pad on the dorso-lateral aspect of the first finger and inner meta-
carpal tubercle.
Length from snout to vent : o 59 mm- ; ? 65 mm.
Distribution : Northern Territory ?, Queensland, and Northern New South
Wales. Loveridge (1935) has recorded this species from Alexandria, N.T., but
another specimen from the same locality obtained at the same time by the same
collector is preserved as a skeleton in the British Museum ; it has the zygomatic
process of the squamosal heavily sculptured and forming a broad suture with
the maxilla as in P. australis, whereas in P. alboguttatus the two fail to meet, or
only just make contact at a narrow point, and the squamosal is not sculptured.
B.M.
Mus.
AUSTRALASIAN I.EPTODACTYUDAE 21
Cyclorana brevipes (Peters).
Chiroleptes brevipes Peters, 1871, Mber. Akad. Berlin : 648 (Type locality : — Fort Bowen,
Queensland) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 269.
Phractops brevipes Nieden, 1923, Das Tierreich, Anura I : 523.
Chiroleptes brevipalmatus Giinther, 1876, /. Mus. Godeffroy, 12 : 47 (Type localities : — Port
Denison, Gayndah and Peak Downs, Queensland); Boulenger, 1882, op. cit.: 269, pi. 17,
fig- 5-
Phractops brevipalmatus Fry, 1915, Proc. roy.Soc. Queensland. 27 : 70 (part) ; Nieden, 1923,
Das Tierreich, Anura I : 522.
Habitus moderate ; head a little broader than long. Snout rounded,
scarcely prominent, 1-25 to 1-5 times as long as the eye, with obtusely angular
canthus rostralis and oblique, scarcely concave, loreal region ; nostril midway
between the eye and the end of the snout ; interorbital space narrower than the
upper eyelid ; tympanum distinct, its horizontal diameter half, or very slightly
more than half, that of the eye. Fingers moderate, the first much longer than
the second, and opposed to the remainder ; subarticular tubercles well developed ;
palm tubercular ; two distinct metacarpal tubercles. Toes less than \ webbed,
the membrane midway between the third and fourth not extending beyond the
proximal subarticular tubercle of the latter ; subarticular tubercles prominent ;
a large shovel-shaped metatarsal tubercle which is, however, distinctly shorter
than its distance from the tip of the inner toe. Tibio-tarsal articulation reaching
the tympanum or the posterior corner of the eye.
Back somewhat warty , a distinct curved supratympanic fold ; belly and
lower surfaces of the thighs finely granular ; throat smooth in females, granular
in males.
Pale olive or brown above with dark brown markings arranged as follows :
a curved cantho-temporal streak ; edge of upper lip spotted ; a large, subtri-
angular interorbital blotch, the apex directed forwards ; back with sharply
defined vermiculations, which may have a general longitudinal arrangement.
Usually a fine white mid-dorsal line. Upper surfaces of the limbs olive-
brown, spotted irregularly with lighter ; hinder side of the thighs dark brown,
uniform or with a few white flecks. Lower surfaces dirty white or pale brown ;
the gular region of the female with or without brown spots, that of the male
infuscate.
Male with a vocal sac and diffuse nuptial asperities on the dorso-lateral
surface of the first finger and metacarpal tubercle.
Length from snout to vent : <J y] mm. ; $ 42 mm.
Distribution : Eastern Queensland.
It seems probable that Loveridge (1935 : 12) is correct in regarding brevi-
palmatus as a synonym of brevipes, but it also seems highly probable that records
of both species from West Australia, South Australia, and the west of Queensland
and New South Wales refer to a species distinct from either [C. cuUripes), differing
in its stouter habitus, shorter digits, more extensive webbing, larger metatarsal
tubercle, more coarsely granular skin and obscure colour-pattern.
Specimens Examined.
B.M. 64.10.27.41 <J Port Denison. Darnel] Cotypes of C.
°7-3-4-37 ? Peak Downs. ,, / brevipalmatus
76.3.4.55 $ Gayndah. ,, > Giinther.
1926. 2. 25. 1 $ ' - 11 irooman, nr. Rock- Wilkins.
hampton.
22 H. W. PARKER
Cyclorana cultripes sp. n.
Chiroleptes brevipalmatus (non Gunther) Spencer, 1896, Rep. Horn Exped. C. Austral., (2),
(Zoology) : 165 ; Fletcher, 1898, Proc. Linn. Soc. N.S.W., 22 : 678-682 ; Lucas & le
Souef, 1909, Anim. Australia : 277.
Phraclops brevipalmatus Fry, 1914. Rec. W. Aust. Mus., 1 : 200 ; idem, 1915, Proc. roy. Soc.
Queensland, 27 : 70 (part) ; Waite, 1929, Rep. Amph. S. Australia : 248.
Phractops brevipes (non Peters) Loveridge, 1935. Bull. Mus. comp. Zool. Harv, 78 : 12.
Habitus moderately stout ; head a little broader than long. Snout rounded,
scarcely prominent, 1-4 to 1-5 times as long as the eye, with obtusely angular
canthus rostralis and oblique, slightly concave loreal region ; nostril very slightly
nearer the tip of the snout than the eye ; interorbital space a little narrower
than the upper eyelid ; tympanum | to § the diameter of the eye. Fingers
rather short, somewhat depressed, the first longer than the second and opposed
to the remainder ; subarticular tubercles prominent ; palm slightly tubercular ;
two large flat metacarpal tubercles. Toes \ webbed, the edge of the membrane
midway between the third and fourth reaching nearly to the level of the distal
subarticular tubercle of the third ; subarticular tubercles moderate ; a very
large, shovel-shaped inner metatarsal tubercle, as long as, or longer than, its
distance from the tip of the inner toe ; no outer. Tibio-tarsal articulation
reaching the tympanum.
Skin distinctly warty above ; a slight occipito-nuchal furrow ; a strong,
curved, supratympanic fold. Belly and lower surfaces of the thighs coarsely
granular ; chin slightly granular in males, smooth in females.
Dull yellow- or grey-brown above with some indistinct darker markings in
the form of a cantho-temporal streak, some labial spotting, a dark blotch on the
top of the head from the level of the posterior corner of the eyes forwards, and a
transverse bar just behind the occiput ; the area between the latter and the
head-marking lighter. Remainder of the back with very indistinct darker spots
and a fine white vertebral line. Groins and hinder side of the thighs brown,
mottled with lighter. Lower surfaces white, the chin dappled with brown in
females and infuscate in males.
Male with a vocal sac and 2 diffuse nuptial pads, one on the dorso-lateral
surface of the inner finger and the other on the inner metacarpal tubercle. Inner
2 fingers of breeding females slightly spatulate.
Length from snout to vent : <$ 46 mm. ; ? 50 mm.
Distribution : Western New South Wales, Northern Territory and northern
West Australia ; probably northern South Australia also.
This form has hitherto been confused with P. brevipes (q.y.) ; the two are
closely allied and may ultimately prove to be only racially distinct.
Specimens Examined.
B.M. 1908.2.25.33 <J Alexandria, N.T. Stalker. Holotype.
1908. 2. 25. 31-32 6*9 ■ ■ " ■• I „
ion o 28 I ? Wilcannia, Darling Helms. Paratypes.
River, N.S.W. >
Cyclorana platycephalus (Gunther).
Chiroleptes platycephalus Gunther, 1873, Ann. Mag. nat. Hist., (4), 11 : 35° (TyPe locality : —
Fort Bourke) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 268, pi. 17, ng. 4 ,
Fletcher, 1890, Proc. Linn. Soc. N.S.W. , (2), 5 : 673, 675 ; idem, 1891 op. at (2) 6 :
265 269 271 ; idem, 1892, op. cit., (2), 7 : 12 ; Spencer, 1896, Rep. Horn. Exped. C.
Austral.,2, (Zool.) : 160, pi. 13, fig. 1, pi. 14, figs. 5"9 I Lucas & le Souef, 1909. A mm.
Austral. : 278, fig. 1 ; Harrison, 1922, Aust. Zool., 3, 1 : 34.
AUSTRALASIAN LEPTODACTYLIDAE 23
Cheiroleptes platycephalus Spencer & Gillen, 1912, Across Australia, 1 : 59, pi. i, fig. a.
Phractops platycephalus; Nieden, 1923, Das Tierreich, Anura I : 520, figs. 352-354 ; Waite,
1929, Rep. Aniph. S. Austral. : 246, fig. 176 ; Loveridge, 1935. Bull. Mus. camp. Zool.
Harv., 78 : 12.
Habitus stout. Head broader than long, proportionately broader in older
individuals, and generally broader in eastern than in western specimens. Snout
rounded in eastern districts, more acuminate and prominent in the west, i-6 to
2-0 times as long as the eye, with indistinct canthus rostralis and very oblique,
rounded loreal region ; nostril equidistant from the eye and the end of the snout
or a little nearer the former ; interorbital space as broad as, or a little broader
than, the upper eyelid ; tympanum distinct § to \ the diameter of the eye and
varying in its proximity to the latter. Fingers moderate, depressed, the first
much longer than the second and opposed to the remainder ; subarticular tubercles
small but prominent ; two metacarpal tubercles, the outer sometimes rather
indistinct. Toes fully webbed, the membrane rather fleshy ; subarticular
tubercles small ; a large, shovel-shaped metatarsal tubercle much longer than its
distance from the tip of the inner toe ; a slight fold along the inner edge of the
tarsus. Tibio-tarsal articulation reaching the shoulder or the tympanum.1
Skin smooth or, more usually, with scattered warts, the condition varying
with the degree of distension on account of stored fluids in the body ; a curved
supra-tympanic fold ; lower surfaces granular.
In spirit, or when alive during the dry season the colour is olive-grey, grey-
brown or yellowish, freckled with indistinct darker spots and blotches not arranged
after any definite pattern ; lower surfaces dirty white, the throat of the male
dotted with brown or blue-black. After leaving their burrows at the onset
of rain the colour of juveniles and half-growTi individuals changes to yellow with
bright green markings, older specimens being duller, with more diffuse green
patches. The iris is golden with dark flecks, tympanum yellowish-brown, the
sides of the body and limbs orange brown and the digital webbing often distinctly
pink.
Length from snout to vent : $ 56 mm. ; $ 68 mm.
Male with a vocal sac and a diffuse nuptial pad on the metacarpo-phalangeal
knuckle of the first finger and extending on to the inner metacarpal tubercle.
Distribution : Arid districts of the interior of Australia from Murchison
in the west to the western districts of New South Wales, including the southern
parts of the Northern Territory and northern South Australia.
This species is essentially cryptozoic, frequenting arid regions and aesti-
vating through the dry season in burrows about a foot deep on the banks of
creeks and near water-holes ; impervious clay soils appear to be preferred, and
during the period of aestivation the frog is greatly distended with water stored
in the urinary bladder, the lymph spaces and body cavity. Breeding takes place
on the advent of rain (January to February in Central Australia) and develop-
ment is very rapid, possibly not more than two weeks elapsing from the laying
of the eggs to metamorphosis. The tadpole reaches a length of 69 mm. and has
the following characters : Body 1-5 times as long as wide ami \ the length of
the tail, ovoid, with eyes and nostrils directed upwards, the nostril nearer the tip
of the snout than the centre of the eye : spiraculum sinistral . anus dextral, near
the edge of the lower caudal crest ; tail acutely pointed, three times as long as deep,
the lower crest deeper than the upper, but not as deep as the muscular portion
1 It is the tibio-tarsal articulation which reaches the tympanum in the type as originally
described, and not the tarso-metatarsal as suggested by Loveridge (1935 '. '-'■
24
H. W. PARKER
at its base. Mouth, except the median third anteriorly, surrounded by papillae
which form a double row and invade the oral cone at the corners ; horny mandibles
strong and serrated ; labial teeth in series }, the innermost both above and below
divided, the outermost shortest. Body and muscular part of the tail dull yellow,
with faint blotches of darker ; caudal crests with dark mottling.
Length from snout to vent at metamorphosis : 31 mm.
Specimens Examined.
B.M. 1937.7.22. 1-2
1908.5.28.62-
64
97. 1 .20.29-30
97. 1 .20.28
1905. 10.31 .46
92.9. 16. 11
73.4.30.18
o\ 2??
Larvae.
?
¥
?
?
Dalgaranger Sta., near
Yalgoo, W.A.
Central Australia.
Charlotte Waters.
100 miles east of Lake
Eyre.
Euroka, N.S.W.
Fort Bourke, N.S.W.
Nicholls.
(Gerrard.)
Horn.
Hii'lier.
Rose.
(Sydney Mus.) Type.
LECHRIODUS Boulenger.
Asterophrys (non Tschudi) Doria, 1875, Ann. Mus. Star. nat. Genova, 6 : 355 ; Peters &
Doria, 1878, Ann. Mus. Stor. nat. Genova, 13 : 417.
Batrachopsis (non Fitzinger) Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 439 (Type
species : — Asterophrys melanopvga Doria) ; van Kampen, 1923, Amph. Indo-Auslr.
Archip. : 16.
Lechriodus Boulenger, 1882, Cat. Batr. Grad. Brit. Mus. : 116 (substitute name for Batra-
chopsis Boulenger) ; Noble, 1922, Bull. Amer. Mus. nat. Hist., 46 : 73 ; Nieden, 1923,
Das Tierreich, Anura I : 49 ; Noble, 1924, Amer. Mus. Novit., 132: 11; idem,
1931, Biol. Amph. : 497 ; Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 22.
Phanerotis Boulenger, 1890, Proc. Linn. Soc. N.S.W., (2), 5 : 593 (Type species : — Phanerotis
fletcheri).
Ranaster (part) Nieden, 1923, Das Tierreich, Anura I : 535.
Maxillary teeth present. Prevomer present, entire, its post-choanal portion
large, overlying at least the mesial half of the palatine, dentigerous ; fronto-
parietals well developed, forming a median suture. Ear fully developed.
Text-fig. 6. Anterior cranial elements of Lechriodus platyceps. X 2. (From beneath/
Vertebrae procoelous with very long transverse processes ; 8 presacrals ; sacral
diapophyses strongly dilated ; coccyx articulating by two condyles. Omo-
sternum cartilaginous ; sternum bifid posteriorly, cartilaginous. Terminal
phalanges simple.
AUSTRALASIAN LEPTODACTYLIDAK 25
Distal tendon of the m. semitendinosus perforating the gracilis complex.
Alary processes of the hyoid expanded distally but with a narrow stalk proximally
as in Limnodynastes and Heleioporus.
Pupil horizontal. Tongue subcircular, slightly free behind. Toes with a
rudiment of web and very small terminal discs.
Synopsis of the Species.
I. Head not broader than long ; first finger as long as or longer than the
second ; small species of which the females have normal digits and
the nuptial pad of the male is composed of minute spines.
A. Snout more than once and a half as long as the eye ; first finger
longer than the second . . . . . L. melanopyga.
B. Snout less than once and a half as long as the eye ; first finger equal
to, or a little shorter than, the second
(i) A small /\-shaped fold on the scapular region . L. fletcheri.
(2) A small / \-shaped fold on the interorbit and a pair of curved,
convergent dorso-lateral folds or rows of plicae . L. papuanus.
II. Head much broader than long ; first finger shorter than the second ;
large species, of which the females have the first and second fingers
strongly fringed and males have nuptial pads composed of closely set,
but relatively large spines ...... L. platyceps.
Lechriodus melanopyga (Doria).
Asterophrys melanopyga Doria, 1875, Ann. Mus. Stor. nat. Genova, 6 (1874) : 355, pi. xii,
fig. k (Type locality : — Wokan, Aru Islands).
Batrachopsis melanopyga (part) Boulenger, 1882, Cat. Balr. Sal. Brit. Mus., ed. 2 : 439 ;
van Kampen, 1923, Amph. Indo-Austr. Archip. : 17.
Lechriodus melanopyga (part) Nieden, 1923, Das Tierreich, Anura I : 49 ; Noble, 1924,
Amer. Mus. Novit., 132 : figs. 5, 7 (myology).
Vomerine teeth in long, curved series extending laterally a little beyond the
choanae. Head longer than broad ; its width contained more than 2-5 times in
the total length ; snout rounded, not prominent, i-6 times as long as the eye ;
canthus rostralis angular ; loreal region oblique, not concave ; nostril nearly
twice as far distant from the eye as from the tip of the snout ; interorbital space
narrower than the upper eyelid ; tympanum distinct, vertically oval, its vertical
diameter | the horizontal diameter of the eye. Fingers free, slender, the first
a little longer than the second ; subarticular tubercles large and prominent ;
a smaller supernumerary tubercle proximal to the base of each finger ; a prominent
oval inner, and a rather indistinct outer, metacarpal tubercle. Toes slender,
with a rudiment of web and small terminal dilatations; subarticular tubercles
prominent ; an elongate oval inner, but no outer, metatarsal tubercle. Tibio-
tarsal articulation extending a little beyond the end of the snout.
Skin almost smooth above, though minute pustules are present murr especially
on the upper eyelid, about the ear and on the Hanks. Two folds arise from the
posterior corner of the upper eyelid, the one running above the tympanum to the
flanks, and the other, somewhat indistinct and interrupted, composed of a row
of pustules, curves towards its fellow on the middle of the back and then runs
parallel with it on to the coccygeal region. Lower surfaces smooth. A very
small papilla on the heel.
26 H. W. PARKER
Brownish grey above, a light line connects the upper eyelids and may extend
forwards to cover the whole of the upper surface of the snout ; a dark bar runs
obliquely forwards from beneath the eye to the edge of the upper lip and may be
broadened anteriorly to cover the whole loreal region ; a deep black marking
may border the supratympanic fold inferiorly, its lower margin irregular and
crossing the tympanum ; other black spots may be present on the flanks, and
sometimes there are large, indefinite lighter areas on the back. Hind limbs with
alternating broader and narrower cross-bars above ; concealed surfaces of the
thighs and tibiae, as well as the lower surfaces of the forearms, tarsi and feet
blackish. Lower surfaces uniform white.
Length from snout to vent : $ 50 mm.
Distribution : Aru Islands.
Although the type of melanopyga was afterwards (Peters and Doria, 1878 :
417) said to be a juvenile when larger specimens from New Guinea were obtained
for comparison, the single specimen collected in the Aru Islands by the " Chal-
lenger " Expedition is of approximately the same size, and is a sexually mature
female. It also differs from specimens from New Guinea in its much narrower
head, longer first finger and smoother skin, so that there seems to be every
probability that the Lechriodus of the Aru Islands is not conspecific with that
from the northern and western parts of New Guinea which has hitherto been
known as melanopyga, but which must now receive a new name. Records of
a small "melanopyga" from S.E. New Guinea in all probability refer to L.
fletcheri (q.v.).
Specimen Examined.
B.M. 82.7. 14.31 $ Aru Islands. "Challenger" Exped. Topo-
type.
Lechriodus fletcheri (Boul.).
Phanerotis fletcheri Boulenger, 1890, Proc. Linn. Soc. N.SAV., (2), 5 : 494 (Type locality : —
Dunoon, Richmond River, N.S.W.) ; Fletcher, 1890, Proc. Linn. Soc. N.S.W., (2) 5 :
669-675 ; idem, 1894, op. cit., (2), 8 : 530 ; Lucas & le Souef, 1909, A nim. Austral. : 274 ;
Andersson, 1913, Jb. nassau. Ver. Naturk., 66 : 75 ; Fry, 1915, Proc.roy. Soc. Queensland,
27 : 69, fig. 1, pi. i, fig. 2 ; Andersson, 1916, K. Svensha VetenskAhad. Handl., 52,
9 : 10, pi. i, fig. 3 ; van Kampen, 1923, Amph. Indo-Austr. Archip. : 18.
Ranaster fletcheri Nieden, 1923, Das Tierreich, Anura I : 536.
Lechriodus fletcheri Noble, 1931, Biol. Amph. : 497 ; Loveridge, 1935, Bull. Mus. comp.Zool.
Harv., 78 : 22.
? Batrachopsis melanopyga Boulenger, 1898, Ann. Mus. Stor. nat. Genova, 18 : 17.
Vomerine teeth in slightly curved series which do not extend laterally
beyond the choanae. Head as long as broad. Snout rounded, not prominent,
1-4 times as long as the eye, with angular canthus rostralis and oblique, slightly
concave loreal region ; nostril 1-3 times as far from the eye as from the tip of the
snout ; interorbital space as broad as the upper eyelid ; tympanum distinct,
vertically oval, its vertical diameter f the length of the eye. Fingers moderate,
the first equal to or a little shorter than the second ; subarticular tubercles large
and prominent ; a slight, fleshy, tubercle-like webbing between the fingers ;
a prominent oval inner, and a longitudinally cleft outer metatarsal tubercle.
Toes slightly dilated terminally and with a distinct rudiment of webbing ; sub-
articular tubercles prominent ; an oval inner, but no outer metatarsal tubercle.
Tibio-tarsal articulation reaching beyond the tip of the snout.
Skin almost smooth or finely granulate above, with some larger pustules on
AUSTRALASIAN LEPTODACTYLIDAE 27
the upper eyelid and about the ear ; a strong narrow fold from the posterior
corner of the eye above the ear to about the middle of the flanks ; a small
/\-shapcd fold in the middle of the back ; smooth beneath ; a small papilla on
the heel.
Pale brown above, with some very small darker spots ; a narrow, dark,
transverse bar from eye to eye ; loreal and temporal regions darker with obscure
markings in the form of oblique lighter (reddish) bars radiating from the posterior
half of the eye ; sometimes a dark brown patch on the canthus rostralis and
below the supratympanic fold ; dorsal /\-shaped fold outlined with darker ;
limbs with alternately broader and narrower dark cross-bars ; concealed surfaces
of the thighs and tibia, and lower surfaces of the forearm, tarsus and foot dark
brown. Lower surfaces white, the edge of the lower jaw brown.
Length from snout to vent : $ 42 mm. ; $ 50 mm.
Male with a vocal sac opening by a slit on each side of the tongue, and
nuptial pads of very small spines on the dorsal surface of the inner finger, except
the terminal joint, on the inner metacarpal tubercle and along the inner dorsal
side of the second finger ; the pad on the first finger must contain many hundreds
of spines. Mature females with the two inner fingers dilated as in Limnodynastes.
Distribution : New South Wales, Queensland, Southern New Guinea.
Specimens Examined.
B.M. 90.7.28.1 imm. $. Dunoon, Richmond Helms. Type.
River, N.S.W.
97. 12. 10. 164 o* Vikaiku, Brit. N. Loria.
Guinea.
Lechriodus papuanus (Roux).
Phanerotis fletcheri papuana Roux, 1927, Rev. Suisse Zool., 34, 4 : 122, fig. 1 (Type locality : —
Lake Sentani district, Dutch New Guinea).
Vomerine teeth in two long transverse series behind the choanae. Head as
long as broad, moderately high. Snout blunt, slightly longer than the diameter
of the eye ; nostril quite close to the tip of the snout ; canthus rostralis distinct ;
loreal region high, oblique, slightly concave ; interorbital space narrower than the
upper eyelid. Tympanum distinct, oval, a little higher than long, its height a
little greater than half the diameter of the eye. Fingers and toes slightly dilated
at their tips, the first finger equal to the second ; subarticular tubercles very
prominent. Toes distinctly webbed at the base ; an elongate, oval inner meta-
tarsal tubercle, about half as long as the inner toe. Tibio-tarsal articulation
reaching a little beyond the tip of the snout.
Skin smooth and beset with small, pointed pustules, larger and rounded in
places, for example on the upper eyelids ; a small papilla on the heel. Dorsum
with prominent, short, narrow folds ; some on the interorbit form a /~\, others
form a pair of curved dorso-lateral lines from behind the orbits, convergent
towards the middle of the back and then running parallel to the lumbar region,
where there are also some short, oblique, secondary crests. Another fold runs
from the eye above the tympanum to the middle of the flanks. Lower surfaces
smooth except the median posterior portions of the thighs, which are granular.
Light grey above with black or grey-black spots and patches, All the dorsal
folds are outlined witli darker and the orbito-tympanic fold is bordered beneath
with a dark patch which covers the upper half of the tympanum ; below the end
28
H. \V. PARKER
of this glandular line, behind the fore-limbs, one or two black spots. Several
broad, dark grey bars on the sides of the snout, one at the tip, one beneath the eye
and another, smaller, between the two. Lower surfaces immaculate whitish-
grey. Fore-limb grey with some little black dots on the anterior surface of the
arm ; fore-arm with two broad, dark grey annuli anteriorly and behind and on
the elbow with longitudinal, blackish-grey marking. Fingers annulate with
blackish grey, light grey beneath. Hind-limb grey ; thighs with two indistinct
darker bars above, blackish brown beneath and on the anal region, this colour
sharply defined above and beneath. Tibia with a faintly indicated, darker,
oblique bar. Knee, a line along the outer edge of the tibia, heel and lower surface
of the tarsus black ; lower surfaces of the foot and toes dark grey (after Roux).
Immature 9 : 23 mm.
Distribution : near Lake Sentani, Dutch New Guinea.
A single example of this form is the only one known and its describer thought
that it was a characteristic Papuan representative of L. fletcheri. But the only
Papuan specimen of fletcheri available to the present author exactly resembles
the type from New South Wales and entirely lacks the dorso-lateral folds described
in the type of papuanus. Accordingly it seems probable either that there are
two distinct species, or that further material will show a much greater range of
variation than is at present known ; there is no evidence in favour of the
recognition of subspecies.
Lechriodus platyceps sp. n.
Asterophrys melanopyga (non Doria, 1875) Peters & Doria, 1878, Ann. Mus. Stor. nat.
Genova, 13 : 417.
Batrachopsis melanopyga Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 439 (part) ;
Boettger, 1894, Denkschr. med. Naturw. Ges. Jena, 8: 112; Lucas, 1898, Proc. Linn. Soc.
N.S.W., 23 : 359 ; van Kampen, 1923, Amph. Indo-Austr. Archip. : 17 (part).
Lechriodus melanopyga van Kampen, 1906, Nova Guinea, 5 : 163, 178 ; Barbour, 1912, Mem.
Mus. comp. Zool. Harv., 44 : 177 ; Fry, 1913, Mem. Queensland Mus., 2 : 48 ; idem,
1915, Proc. roy. Soc. Queensland, 27, 4 : 73, pi. i, figs. 1, 2 ; Nieden, 1923, Das Tierreich,
Anura I : 49 (part), figs. 105, 106 ; Noble, 1931, Biol. Amphib. : 113, fig. 38.
Holotype a female, number 76.7.18.6 in the British Museum, from Arfak,
Dutch New Guinea ; collected by A. A. Bruijn.
Vomerine teeth in curved series which do not extend laterally beyond the
choanae. Head broad and flat, considerably broader than long, its width con-
tained less than 2-5 times in the length from snout to vent. Snout rounded, not
prominent, 1-6 times as long as the eye, with angular canthus rostralis and
oblique, very slightly concave loreal region ; nostril 1-2 times as far distant from
the eye as from the tip of the snout ; upper eyelid 1-5 times the width of the inter-
orbit ; tympanum distinct, vertically oval, its vertical diameter f the length of
the eye. Fingers moderately long, the first a little shorter than the second and
both with strong lateral fringes which do not continue proximally beyond the
subarticular tubercles ; the latter are well developed but the supernumerary
tubercles at the base of each finger are merely indicated ; a large, prominent
inner metacarpal tubercle, containing a cartilaginous prepollex, and a small
indistinct outer. Toes with small terminal discs and a rudiment of webbing ;
subarticular tubercles well developed ; a small oval inner, but no outer meta-
tarsal tubercle. Tibio-tarsal articulation reaching the tip of the snout.
Skin regularly and profusely beset with small round pustules which may
AUSTRALASIAN LEPTODACTYLIDAE
29
form bead-like chains especially on the flanks ; some larger papillae on the
upper eyelid, about the ear and one on each heel. A straight dermal fold from
the posterior corner of the eye above the tympanum to the flanks, and a narrow
linear fold from the upper eyelid convergent towards the middle line about the
middle of the back and then slightly divergent and forming a sinuous line to the
end of the coccyx ; a slight curved fold across the interorbit. Smooth beneath.
Pale brown above ; upper surface of the snout to the interorbital fold lighter,
the fold itself dark-edged. An irregular dark brown band beneath the canthus
rostralis and supratympanic fold ; a few dark spots beneath the eye and on the
flanks, especially in the axillae and groins ; a faint, light marking on each side
of the back in the angles where the dorso-lateral folds approach one another and
then diverge. Limbs with narrow transverse cross-bars. Concealed surfaces
of thighs and tibiae and lower surfaces of the fore-arms, tarsi and feet dark brown.
Lower surfaces whitish, the edge of the lower jaw and gular region freckled with
brown.
Length from snout to vent : 79 mm.
These specimens agree with the holotype in general morphological characters
and colour with the exception of one or two details. The pustules are usually
more numerous and prominent, often forming lines, especially on the legs, where
they outline the transverse bars of colour ; the tibio-tarsal articulation usually
extends a little beyond the tip of the snout, the prepollex may be ossified, and
the dark colour of the concealed surfaces of the hind-limbs is broken up into spots
and dots. The first and second fingers of females are spatulate as described but
those of the males are cylindrical, but with nuptial pads composed of relatively
large spines ; one such pad occupies the dorsal surface of the inner finger, except
the terminal joint, a smaller one is situated on the end of the inner metacarpal
tubercle and a third on the inner edge of the second finger at its base ; the largest
patch does not contain more than about 180 spines. Male with a vocal sac.
Largest $ 70 mm. ; $ 96 mm.
Fry (1915 : 74) draws attention to the fact that specimens from New
Guinea had.a much broader head than the type of melanopyga, and this difference,
together with some others already noted, suggests that the two are not conspecific.
The width of the head cannot be an age-character, for two of Fry's specimens
measuring only 45 and 47 mm. characterized by the broad head are no larger
than the narrow-headed examples of true melanopyga recorded from the Aru
Islands, and the juvenile specimen from the Tor River, measuring only 21-5 mm.
from snout to vent, has a relatively broader head than an adult melanopyga. The
species plalyceps has been recorded from the following localities in New Guinea :
Arfak Mts. ; south of Geelvinck Bay ; Mansinam ; Hatam ; Mamberamo River ;
Idenburg River ; Tor River ; St. Joseph's River ; Fife Bay.
30 H. W. PARKER
ADELOTUS Ogilby.
Cryptotis (nee Pomel, 1848, nee Dana, 1852) Gunther, 1863, Ann. Mag. nat. Hist., (3), 11 :
27 (Type species: — Cryptotis brevis) ; Cope, 1865, Nat. Hist. Rev., n.s., 5 : 107 ; idem,
1866, J. Acad nat. Set. Philad., (2), 6 : 80 ; Steindachner, 1867, Reise Novara, Amph. :
30 ; Keferstein, 1868, Areh. Naturgesch., 34 : 267 ; Boulenger, 1882, Cat. Batr. Sal.
Brit. Mus.,ed.2 : 262; Cope, 1889, Bull. U.S. nat. Mus., 34 : 295; Noble, 1931, Biol.
Amph. 498.
Adelotus (substitute name for Cryptotis Gunther) Ogilby, 1907, Proc. roy Soc. Queensland,
20 : 32 ; Nieden, 1923, Das Tierreich, Anura I : 538 ; Noble, 1931, Biol. Amph. : 498 ;
Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 23.
Skull elongate, the pars facialis of the maxilla forming a long edentulous
suture with the pre-maxilla (text-fig. 7). Maxillary teeth present. Prevomer
bounding the choana anteriorly and mesially, with a short, oblique process
directed backwards bearing a short row of teeth and partially overlying the mesial
end of the palatine. Dentary with a pseudo-tooth at its anterior end, small in
females, very large and tusk-like in adult males. Fronto-parietals forming a
Text-fig. 7. — Anterior cranial elements of A delotus brevis. x 3. (From beneath.)
median suture. Ear fully developed. Vertebrae procoelous, the condyle
incompletely ankylosed and the notochord persistent ; sacral diapophyses
slightly dilated ; urostyle articulating by 2 condyles ; 7 pre-sacral vertebrae
(1+2 fused). Omostemum well developed, cartilaginous ; sternum broad,
cartilaginous, notched but not bifid posteriorly ; procoracoids broad. Terminal
phalanges simple.
Distal tendon of the m. semitendinosus perforating the gracilis complex
distally. Alary processes of the hyoid expanded distally, narrower proximally.
Pupil horizontal with a ventral angle ; tongue oval or subcircular, partly
free behind. Toes not dilated distally, with a rudiment of web.
Adelotus brevis (Gunther).
Cryptotis brevis Gunther, 1863, Ann. Mag. nat. Hist., (3), 11 : 27, pi. 4, fig. B(Type locality: —
Clarence River, N.S.W.) ; KreSt, 1865, Pap. Proc. roy. Soc. Tasmania : 17 ; idem, 1867
Cat. Industr. Prod. N.S.W., Add. : 107; Steindachner, 1867, Reise Novara, Zool., Amph.
30 ; Gunther, 1868, Proc. zool. Soc.Lond. : 480 ; Keferstein, 1868, Arch. Naturgesch., 34
268 ; Fletcher, 1889, Proc. Linn. Soc. N.S.W. , (2), 4 : 386 ; idem, 1890, op. cit., (2), 5
669-671 ; idem, 1892, op. cit., (2), 7:8; idem, 1894, op. cit., (2), 8 : 525, 527, 529
Lucas & le Souef, 1909, Antm. Austral. : 275 ; Andersson, 1913, A'. Svenska Vetensk.
Akad. Handl., 52, 4 : 11.
Adelotus brevis Ogilby, 1907, Proc. roy. Soc. Queensland, 20 : 32 ; Nieden, 1923, Das Tierreich
Anura I : 538, figs. 371, 372 ; Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 23.
AUSTRALASIAN LEPTODACTYLIDAE 31
Head strongly depressed and enlarged in the male, subtriangular, broader
than the body ; not broader than the body and more rounded in females.
Vomerine teeth in two short, oblique series behind the level of the choanae.
Snout rounded, slightly prominent, i-8 to 2-0 and 1-4 to i-6 times as long as the
eye in males and females respectively ; nostril a little nearer the eye than the
end of the snout ; interorbital space as broad as, or a little broader than the upper
eyelid. Tympanum hidden, but when uncovered by dissection vertically oval,
its horizontal diameter about half that of the eye. Fingers moderately long,
with a faint rudiment of web ; in females the first and second are strongly spatu-
late with a thick fleshy fringe along their preaxial margins ; subarticular tubercles
prominent ; two prominent rounded metacarpal tubercles. Toes moderate,
with a slight rudiment of web ; subarticular tubercles prominent ; metatarsus
with rows of small pustules ; a small, slightly oval inner metatarsal tubercle and
a much smaller outer situated at the base of the fourth metatarsal. Tibio-tarsal
articulation reaching the tympanic region or the posterior corner of the eye.
Skin more or less warty above, particularly on the upper eyelids and hind
limbs ; lower surfaces smooth.
Pale brown to olive or grey above with scattered darker blotches and lighter
dots ; synciput lighter. The most constant dark markings are a bar from the
tip of the snout through the nostril to the eye, sometimes continued as a straight,
oblique line from the posterior corner of the eye to the angle of the mouth and
edged beneath with lighter, a vertical bar beneath the eye and a large sub-
quadrangular or triangular blotch whose anterior edge connects the centres of
the upper eyelids. Limbs regularly cross-barred above. Lower surfaces dark
brown or blackish with small white (?) flecks on the gular region and bold spots
on the belly and lower surfaces of the femur and tibia ; groin and concealed
surfaces of the tibia, metatarsus and foot with large pink or crimson spots.
Male with a vocal sac but apparently without nuptial pads. Premaxillary
and anterior maxillary teeth strongly enlarged.
Length from snout to vent : $ 44 mm. ; $ 37 mm.
Distribution : New South Wales (East of the Dividing Range), and
Southern Queensland.
Specimens Examined.
Krefft. Cotypes.
Heron.
Breeding habits unknown. The inner fingers of the female suggest a foam-
beating habit similar to that of Limnodynastes, but the apparent absence of
nuptial pads in the male suggests that pairing may take place on land.
HELEIOPORUS Gray.
Rana Shaw, 1795, Nat. Miscellany. 6 : pi. 200; Schneider, 1799, Hist. Amphib., 1 : 129.
'Heleiuporus Gray, 1841, Ann. Mag. not. Hist., 7 : 91 (Type species: — H. albopunctatus) ;
Gray, 1841, in Grey, Journ. Exped. Austral., 2 : pi. i; Gunther, 185S, Cat. Batr. Sal.
Brit. Mus. : 38 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 271.
'Heltoporus Gray, 1841, in Grey, Journ. Exped. Austral., 2 : 447 ; Cope, 1865, Nat. Hist. R
n.s., 5 : 108 ; idem. iS<><>, /. A, ad. nut. S, 1 . I'lulad., (21. (. :-■>, M K 1. 1 1. m 1S1.S
1 In the specific synonymies these alternative spellings are not recorded separately.
32 H. W. PARKER
Arch. Naturgesch., 34 : 269 ; Cope, 1889, Bull. U.S. nat. Mus., 34 : 312 ; Fry, 1914, Rec .
W. Aust. Mus., 1 : 208 ; Nieden, 1923, Das Tierreich, Anura I : 525 ; Waite, 1929,
Rep. Amph. S. Australia : 240; Noble, 1931, Biol. Amph. : 497.
Heleioforus (err.) Krefft, 1865, Mon. Not. roy. Soc. Tasmania : 17.
Perialia Gray, 1845, in Eyre, Journ. Exped. C. Austral., 1 : 407 (Type species: — Perialia
eyrei) ; Cope, 1889, Bull. U.S. nat. Mus., 34 : 312.
Neobatrachtis Peters, 1863, Mber. Akad. Berlin : 234 (Type species : — N . pictus) ; Cope, 1866,
/. Acad. Nat. Sci. Philad., (2), 6 : 94.
Philocryphus Fletcher, 1893, Proc. Linn. Soc. N.S.W., (2), 8: 233. (Type species: — P.
flavoguttatus) ; Fry, 1914, Rec. W. Aust. Mus., 1 : 208 ; Noble, 1931, Biol. Amph. : 497.
Maxillary teeth present. Pre vomer well developed with a posterior den-
tigerous process which does not quite reach the palatine (text-fig. 8) ; a very large
fronto-parietal foramen, ethmoid entire. Ear fully developed. Vertebrae pro-
coelous, but the condyle incompletely ankylosed ; notochord persistent ; sacral
diapophyses slightly dilated ; urostyle articulating by two condyles ; 7 presacral
vertebrae. Omosternum small, cartilaginous, sternum undivided or notched
posteriorly, cartilaginous or calcified mesially. Terminal phalanges simple.
Text-fig. 8. — Anterior cranial elements of Heleioporus albopunclatus. X 2'5.
(From beneath.)
Distal tendon of the m. semitendinosus passing ventral to the tendon of the
mm. graciles. Alary processes of the hyoid expanded distally, but with a narrow
stalk proximally ; cricoid cartilage complete ; oesophageal process of the cricoid
slender ; m. omohyoideus present ; mm. sternohyoideus and petrohyoidei
attached at the edges of the hyoid.
Pupil horizontal, but with a downwardly directed angle ventrally. Tongue
large, broadly oval, half free behind. Toes more or less webbed.
The status of Philocryphus with regard to Heleioporus has been in dispute
for some time. Fletcher himself (1897 : 679) was doubtful whether it could
be maintained, but Fry (1914 : 206) advanced reasons for continuing to retain
the two apart ; Fry's views have been accepted by the latest commentator
(Loveridge, 1935 : 17), but Noble (1931 : 497), though maintaining the two, can
only distinguish them by the degree of distinctness of the tympanum. This can
be so extremely variable and is so dependent on preservation that its value seems
very problematical. Actually albopunclatus and australiacus are so closely
allied that in their young stages they are almost indistinguishable, and the
characters suggested by Fry for their generic separation are almost all age-
characters ; his skeleton of albopunctatus was almost certainly that of a juvenile.
Fletcher himself at one time (1889 : 376) referred one of his co-types of flavo-
guttatus (=australiacus) to albopunctatus, and Boulenger (1882 : 272) was unable
AUSTRALASIAN LEPTODACTYI.IDAE 33
to distinguish juveniles of austmliacus from albopunctatus. Examination of all
the material now in the British Museum suggests that some 8 forms are valid,
which may be distinguished by the key given below. If any generic separation
should be possible, or desirable, the division should apparently be between
sections I and II. The first of these, containing small species with an undivided
sternum and with diffuse nuptial pads, would then be known as Neobatrachus
Peters, of which N. pictus is the genotype.
Synopsis of the Species.
I. Sternum always cartilaginous, not bifid posteriorly. Small species,
the males with a diffuse nuptial pad. Metatarsal tubercle usually
edged with black or dark brown.
A. Toes f to fully webbed.
(i) Eye not longer than its distance from the tip of the snout. A
colour pattern of dark insuliform spots or irregular marblings
on a light ground.
(a) Skin warty above (spinulose in breeding males) ; meta-
tarsal shovel completely black . . . . H. pictus.
(b) Skin smooth or with spinules in breeding males ; metatarsal
shovel edged with brown or not coloured . H. centralis.
(2) Eye longer than its distance from the tip of the snout. Colour
pattern consisting of light bifurcating lines on a dark ground.
Skin quite smooth, but thick and glandular ; metatarsal
shovel edged with brown .... H. wilsmorei.
B. Toes not more than J webbed, except in males at the breeding
season, when the web continues as a tapering fringe to the tip of
each toe ..... . . . H. pelobatoides.
II. Sternum of the adult calcified and bifid posteriorly. Large species,
the males with large, conical, black horny spines on the dorsal surfaces
of the inner fingers.
A. Tip of the fifth toe not reaching beyond the nostril ; at most a
single, conical tubercle at the anterior corner of the eye ; flanks
not more warty than the dorsum ; a pattern of regular circular
cream-coloured (white) spots on a dark ground colour
H. albopunctatus.
B. Tip of the fifth toe reaching the tip of the snout or a little beyond ;
a fimbriated flap at the anterior corner of the eye ; flanks more
granular than the dorsum.
(1) Size large (up to 80 mm.) ; tympanum of adult large (f-| the
eye) and subcircular ; dorsum uniform purple-brown ; flanks
111" rally yellow-spotted H. austialtacus.
(2) Size moderate (up to 59 mm.) ; tympanum of adult vertically
oval, rather indistinct, S-| the diameter of the eye ; dorsum
brown marbled with grey or yellowish irregular markings H. eyra.
111. Incertae scdis H.sudcll
NOVIT. ZOOI.., 42, I 3
34 H. W. PARKER
Heleioporus pictus (Peters).
Neobatrachus pictus Peters, 1863, Mber. Akad. Berlin: 235 (Type locality: — near Adelaide) ;
Keferstein, 1868, Arch. Naturgesch. : 262.
N.ficlus (err. t\<p.) Krefft, 1865, Pap. roy. Soc. Tasmania : 17.
Helioporus pictus Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 272 ; Lucas, 1892,
Proc. rov. Soc. Victoria, 4 : 61 ; Fletcher, 1890, Proc. Linn. Soc. N.S.W., (2), 5 : 672-675 ;
idem, 1891, op. cit., 6 : 271-273 ; idem, 1894, op. cit., 8 : 529, 531 ; idem, 1898, op. cit.,
22:679; Lucas & le Souef, 1909, Anim. Austral. : 281, fig, ; Fry, 1914, Rec. W.
Aust. Mus., 1 : 207, fig. 11 ; Nieden, 1923, Das Tierreick, Anura I : 527 (part) ; Waite,
1929, Rept. Amph. S. Austral. : 249 (part), fig. 177.
Vomerine teeth in transverse series between the choanae. Snout rounded,
as long as the eye ; canthus rostralis rounded ; loreal region oblique, and very
slightly concave ; nostril equidistant from the eye and the end of the snout,
or a little nearer the latter ; interorbital space § the width of the upper eyelid ;
tympanum indistinct, vertically oval, its maximum diameter about half the length
of the eye. Fingers moderate, the first extending beyond the second, which
extends as far as, or very slightly beyond, the fourth ; the length of the third,
along its mesial side, very slightly shorter than the eye. Toes \ or fully webbed,
the membrane extending to the tips of all of them. Subarticular tubercles
scarcely developed on the feet ; a strongly compressed, prominent metatarsal
tubercle about as long as its distance from the tip of the inner toe, covered with
a black horny sheath. Tarso-metatarsal articulation reaching the eye, the tip
of the outer toe reaching the tip of the snout.
Skin with irregular small warts above ; smooth beneath ; anal region feebly
granular.
Greyish or pale brown above with numerous insuliform dark brown spots or
marblings, of which the most constant are an oval canthal stripe, a triangular
mark between the eyelids and a short, oval mark behind the upper eyelid ; lower
surfaces white except the sides of the throat and edges of the lower jaw, which
may be blotched or washed with brown ; anal granules white-tipped. Usually
a narrow, vertebral line. In life (Fletcher, 1891 : 273) there is sometimes a
yellow wash about the thighs and upper arms. Iris silvery or golden, veined with
black and with a dusky streak from the anterior corner to the centre of the pupil.
Breeding male with a distended throat but without definite vocal sacs, with
a diffuse, brown, nuptial pad on the upper surfaces of the first and second fingers
and with small, horny brown spinules scattered over the dorsal surfaces and on
the outer edge of the tarsus and fifth toe.
Length from snout to vent : cJ 36 mm. ; $ 47 mm.
Distribution : New South Wales, west of the dividing range ; Victoria ;
S. Australia in the south and east ; ? S. Queensland ; possibly also in eastern
districts of West Australia.
The records of this species from Central and N.W. Australia appear to refer
to a distinct, but related, form (vide H. centralis). H. sudelli Lamb, from Warwick,
S. Queensland, is usually placed as a synonym of pictus, but several points in the
description, notably the approximation of the nostril to the eye, and the equally
long first and second fingers, make this disposition seem very doubtful.
Specimens Examined.
B.M. 74.4.29.1283 $ Sandhurst, Victoria. [Beddome.]
99.2. 14. 1 $ Melbourne. Degen.
6). 10. 27.46 juv. Ryalstone, N.S.W. Krefft.
92.9.16.5 <J Urana, N.S.W. Fletcher.
R.R. 1937.7.29. 10 (J skel. "Australia." Green.
AUSTRALASIAN LEPTOD.U TYL1DAE 35
Heleioporus centralis sp. n.
Heleioporus pictus (non Peters) Spencer, 1896, Rep. Horn Exp. C. Austral., 2, (Zool.) : 166,
pi. 13, fig. 2, pi. 14, figs. 10-13 I Spencer & Gillen, 1912, Across Australia, 1 : 59, pi. i.
fig. c ; ? Andersson, 1913, A*. Svenska VetenshAkad. Handl., 52, 4:16; Harrison, 1922,
Aust. Zool., 3, 1 : 33 ; Nieden, 1923. Das Tierreich, Anura I : 527, figs. 358-361 (part) ;
Waite, 1929, Rept. Amph. S. Austral. : 249 (part).
Holotype a male, number 1905. 10.31 .47, from 100 miles east of Lake Eyre,
collected by H. J. Hillier, Esq.
Vomerine teeth in slightly oblique series between the choanae. Snout
rounded, as long as the eye ; canthus rostralis rounded ; loreal region oblique,
but not concave ; nostril a little nearer the tip of the snout than the eye ; inter-
orbital space I the width of the upper eyelid ; tympanum very indistinct, verti-
cally oval, its maximum diameter about half the length of the eye. Fingers
rather short, the first extending a little beyond the second, which extends well
beyond the fourth ; the length of the third, along its mesial side nearly as long
as the eye. Toes fully webbed, the membrane extending to the tips of all of them ;
subarticular tubercles very feeble ; a compressed inner metatarsal tubercle as
long as its distance from the tip of the inner toe. Tarso-metatarsal articulation
reaching the tympanic region, the tip of the outer toe marking the anterior corner
of the eye.
Skin smooth above and below, except for the secondary sex characters, which
consist of a diffuse nuptial pad on the first and second fingers, and small papillae
with black horny spinules scattered over the dorsum and along the outer edge
of the tarsus and foot.
Pale grey above with very obscure dusky mottling and a narrow white
vertebral line. Lower surfaces white.
Length from snout to vent : 42 mm.
The paratypes are a male and female from Central Australia, probably
Charlotte Waters, collected by the Horn Expedition (B.M. 97.1.20.32-33) and
two females from " Central Australia " (B.M. 1908.5.28.66-67).
These four specimens scarcely differ from the type in their morphological
characters. The webbing of the toes may be somewhat more emarginate, giving
an appearance of | web only, and the hind limb may be slightly longer, so that the
metatarsal tubercle almost reaches the posterior corner of the eye and the tip
of the outer toe extends a little beyond the eye. Both specimens collected by
the Horn Expedition have the metatarsal tubercle narrowly black-edged (this
colouring is much less extensive than in pictits), and in all the specimens it is
longer than its distance from the tip of the inner toe.
The colour pattern is more distinct than in the type, but its intensity is very
variable. When fully developed it is similar to that in the coloured plate illus-
trating Spencer's account (1896 : pi. 13, fig. 2) of the species. The largest male
and female, both from Charlotte Waters, each measures 50 mm. from snout to
vent.
These Central Australian frogs differ from the eastern and southern spei ies,
pictus, in their rounder, shorter snouts, shorter legs, smooth skin and the absence
or feeble development of the pigmented sheath on the metatarsal tubercle. It
seems probable that the specimens from N.W. Australia (Mowla Downs, Kimber-
ley Division) recorded by Andersson (1913 : 16) as lacking the dark metatarsal
tubercle really belong to this species; Loveridge's suggestion (1935: 16) that
they might be juvenile albopunctatus is untenable, for Andersson described the
nuptial characters of the male, and these are quite unlike those ol albopunctatus.
36 H. W. PARKER
In addition to the above-mentioned specimens the British Museum has a
series of tadpoles which accord with Spencer's description (1896) ; the labial
teeth are in -J- series.
Heleioporus wilsmorei sp. n.
Holotype a female, number 1937.7.22.3 from Wirarga, N.E. of Yalgoo,
Murchison, W. Australia ; collected by G. E. Nicholls.
Vomerine teeth in transverse series on a level with the posterior borders of
the choanae. Head short and high. Snout rounded, J the length of the eye ;
nostril slightly nearer the tip of the snout than the eye ; canthus rostralis rounded ;
loreal region nearly vertical ; interorbital space § the width of the upper eyelid ;
tympanum quite indistinct, but when exposed by dissection, vertically oval, its
maximum diameter about J the length of the eye. Fingers moderate, the first
extending beyond the second, which extends well beyond the very short outer ;
third, measured along its mesial side, as long as the eye. Toes f webbed, the
membrane thick and fleshy leaving the tips of the third to fifth free ; no sub-
articular tubercles on the feet ; inner metatarsal tubercle compressed, as long as
its distance from the tip of the inner toe and black-edged. Tarso-metatarsal
articulation reaching the tympanic region, the tip of the outer toe marking the
centre of the eye.
Skin quite smooth, but very thick and glandular above ; smooth beneath ;
a deep furrow below the vent.
Chocolate brown above, with a series of light markings as follows : a narrow
mid-dorsal line ; a longitudinal row of spots down the centre of the upper eyelid
to the posterior corner where they are continued as light lines which run back-
wards to the vent, sending off two branches on the way ; the first of these arises
above the tympanic region and runs obliquely backwards to the anterior part
of the flanks, and the second arising vertically above the axilla runs obliquely
to the groin. Lower surfaces uniform white.
Length from snout to vent : 50 mm.
The paratype is a female, in bad condition, found dead near Lakeside,
Calgoorlie, October 18, 1921 (B.M. 1937.7.22.4). It agrees with the holotype
in every way. Professor Nicholls has seen a third specimen from the coastal
zone north of Geraldton.
This species, which appears to be confined to the eastern parts of West
Australia, east of the Darling Range, is one of the pictus group, but is distinguished
by its much deeper head, large eye, shorter outer digits and very distinctive
colour pattern.
Heleioporus pelobatoides Werner.
Heleioporus albopunctatusvar. pelobatoidesW'emer, 1914, Fauna S.W. Austral., 4, 10, Amph. :
418 (Type localities : — Beverley and Broome Hill, W. Australia).
Helioporus pelobatoides Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 15.
Helioporus eyrei (non Gray) Loveridge, 1935, loc. cit.
Vomerine teeth in transverse or slightly oblique series between the choanae.
Snout rounded, as long as the eye ; nostril midway between the eye and the tip
of the snout ; canthus rostralis rounded ; loreal region very oblique, not concave ;
interorbital space £ the width of the upper eyelid. Tympanum indistinct in
non-breeding individuals, but becoming very distinct in males at the breeding
season ; in females and juvenile males it is vertically oval, its maximum diameter
AUSTRALASIAN LEPTODACTYLIDAE 37
§ that of the eye, but in adult males it is subcircular, though its diameter is
similar. Fingers moderate, the first extending slightly beyond the second, which
extends very slightly beyond the fourth ; third, measured along its median side,
a little longer than the eye ; subarticular tubercles well developed and the palm
of the hand beset with numerous small rounded tubercles. Toes in non-breeding
adults about £ webbed, but in juveniles at metamorphosis the webbing is more
extensive and males in nuptial dress have it carried to the tips of the toes as a
fleshy, tapering fringe ; subarticular tubercles small ; a compressed inner meta-
tarsal tubercle as long as or longer than its distance from the tip of the inner
toe and edged with brown. Tarso-metatarsal articulation reaching the centre
or anterior border of the eye, the tip of the outer toe extending well beyond the
tip of the snout.
Skin very warty above, especially on the head ; smooth beneath ; anal region
feebly granular.
Pale grey to dark brown above with irregular darker blotches or marblings.
The most constant are a canthal blotch, a triangular interorbital mark and a
pair of very large dorso-lateral blotches on each side of the middle of the back.
The warts and the anal granules are frequently white-tipped and a light dorsal
line may be present. Lower surfaces white or yellowish, the edges of the lower
jaw and sides of the throat washed with brown and stippled with white.
Males at the breeding season, in addition to the extension of the web already
described, have a diffuse nuptial pad on the first and second fingers, and scattered
horny spinules over all the dorsal surfaces.
A nocturnal species, burrowing by day ; normally the burrow is about 2 ft.
deep, but in damp localities the frog may be found in shallow cavities beneath
logs. Eggs are laid in the burrow and the embrace is lumbar (fide Prof. G. E.
Nicholls in litt.).
This species appears to be confined to the wetter south-western parts of
West Australia and is the analogue of the eastern pictus, with which it agrees in
habitus, leg-length and skin texture. The species from the central, dry zone
is more globular in habitus, with a shorter, blunter snout, higher head, and
shorter limbs and digits ; in fact it appears to be more highly modified for a
cryptozoic existence.
Specimens Examined.
B.M. 1931.7.1.18-23 <J. 4 ?? Mt. Toolbrunup. Baldwin.
1931.7.1.13-17 2 $$, 3 juvs.
1931.7.1.5-8 4juvs.,
1 larva
M.C.Z. 19430 $ (breeding) Tambellup.
■8219 juv. Pindawa. Schevill.
The specimen from Pindawa is one of the series referred by Loveridge to
H. eyrei, which he considered to be the same as the species more generally known
as //. pictus. But eyrei and pictus are certainly not conspecific, and these West
Australian frogs, which are very immature, cannot be distinguished from the
juveniles from Mt. Toolbrunup, which are almost certainly pelobatoides. Love-
ridge was led to believe that two species were present in his materia] owing to
the fact that his single adult was a male in nuptial dress. With the exception
of the nuptial characters of digital pads and dermal asperities, this breeding male
only differs from the Mt. Toolbrunup male in the greater amount of digital webbing
and more distinct tympanum ; but a non-breeding male has a larger, rounder,
38 H. W. PARKER
more distinct tympanum than females of the same collection, so that this character
is obviously variable and may well be seasonal ; the condition of the webbing
is believed to be similarly variable.
Heleioporus albopunctatus Gray.
Heleioporus albopunctatus Gray, 1841 (April), Ann. Mag. nat. Hist., 7 : 91 (Type locality: —
Western Australia) ; Gray, 1841, in Grey, Journ. Exped. W. Austral., 2 : 447 ; Giinther,
1858, Cat. Batr. Sal. Brit. Mus. : 39 (part) ; Giinther, 1867, Ann. Mag. nat. Hist., (3).
20 : 55 ; Krefft, 1867, Cat. Industr. Prod. N.S.W., Add. : 107 ; Boulenger, 1882, Cat.
Batr. Sal. Brit. Mus., ed. 2 : 271 (part) ; ?? Lucas, 1892, Proc. roy. Soc. Victoria, 4 : 61 ;
Fletcher, 1893, Proc. Linn. Soc. N.S.W., (2), 8 : 232 ; idem, 1898, op. cit., 22 : 678 ;
Lucas & le Souef, 1909, Anim. Austral. : 281 ; Werner, 1914, Fauna SAV. Austral.,
4, 10, Amph. : 417 ; Fry, 1914, Rec. W. Aust. Mus., 1 : 204 ; Harrison, 1922, Aust.
Zool., 3, 1 : 32 ; Nieden, 1923, Das Tierreich, Anura I : 526 ; Glauert, 1929, /. roy. Soc.
W. Aust., 15 : 44 ; Trewavas, 1933, Phil. Trans, roy. Soc. London, 222, b : 433, figs.
23, 24 ; Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 14.
Heleioporus albogultatus (laps, cal.) Gray, 1841, in Grey, Journ. Exped. W. Austral., 2 :
pi. 1, fig. 2 ; Giinther, 1858, Cat. Batr. Sal. Brit. Mus. : 33 (footnote).
Heleioforus alleopunctatus (err. typ.) Krefft, 1865, Mon. Not. roy. Soc. Tasmania : 17.
Vomerine teeth in long transverse series between the choanae. Snout
rounded, as long as the eye ; canthus rostralis rounded ; loreal region oblique,
not concave ; nostril equidistant from the eye and the end of the snout ; inter-
orbital space about § the width of the upper eyelid ; tympanum usually indistinct,
vertically oval, its horizontal diameter J to A the length of the eye. Fingers
moderate, the first longer than the second, which is longer than the fourth ;
subarticular tubercles well developed, and the palm with many tubercles in
addition to an inner and group of three outer metacarpal tubercles ; one or two
tubercles between the first and second and second and third fingers. Toes with
a distinct rudiment of fleshy web ; third extending beyond the fifth ; a large,
compressed inner metatarsal tubercle ; metatarsal tubercle reaching the tympanic
region and the fifth toe the anterior corner of the eye or the nostril.
Skin regularly shagreened above and on the flanks ; smooth beneath ; anal
region granular. A large parotoid gland extending posteriorly to the middle of
the flanks, and a similar glandular complex on the antero-ventral surface of the
thigh close to the groin.
Dark purple- or chocolate-brown above, regularly beset with circular cream-
coloured spots. A light vertical bar may be present on the tip of the snout
and a light diagonal stripe from below the eye to the angle of the mouth. White
beneath except for the throat, which may be lightly washed with brown.
Male in the breeding season with a series of black, horny, conical spines.
The longest, situated on the metacarpo-phalangeal knuckle of the first finger has
a bony core, which is articulated on the distal end of the first metacarpal ; a
series of much smaller, horny conical spines, but apparently without bony cores,
are situated distal to the above-mentioned large one on the first finger, and a few
may also be present on the second finger. No vocal sac.
Length from snout to vent : $ 68 mm. ; $ 69 mm.
Distribution : West Australia east of the Darling Range and ? N. Territory.
This species appears to be confined to the west of Australia, and various
eastern records have been regarded (Fletcher, 1893 : 233) as instances of faulty
labelling. Some West Australian records of albopunctatus are probably based on
examples of eyrei, and the Victoria record (Lucas, 1892) is probably due to a
misidentification. None of the 13 adult examples of true albopunctatus (including
the type) which the author has examined show any preocular dermal flap.
AUSTRALASIAN LEPTODACTYLIDAE 39
Specimens Examined.
Heleioporus australiacus (Shaw).
Rana australiaca Shaw, 1795, Nat. Miscellany. 6 : pi. 200 (Type locality : — New Holland) ;
Andersson, 1913, A'. Si'enska VetenskAkad. Hand!., 52, 4 : 3.
Philocryphus australiacus Fry, 1915, Proc. toy. Soc. Queensland, 27, 4 : 70 ; Harrison, 1922,
Aust. Zool., 3, 1 : 32 ; Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 17.
Rana spinipes Schneider, 1799, Hist. Amph., 1 : 129-139 (Type locality : — Islands of the
ocean around Australia) ; Shaw, 1802, Gen. Zoology, 3 : 112.
Helioporus albopunctatus (non Gray), Fletcher, 1889, Proc. Linn. Soc. N.S.W., (2), 4 : 376;
idem, 1890, op. cit., (2), 5 : 671 ; Werner, 1914, Fauna S.-W. Austral., 4, 10, Amph. :
417 (part : — Lion Mill ?).
Philocryphus flavoguttatus Fletcher, 1893, Proc. Linn. Soc. N.S.W., (2), 8 : 233 (Type locali-
ties : — Mt. Victoria, Blue Mts. ; Thornleigh ; Dural, near Paramatta ; near Manly) ;
idem, 1894, torn. cit. : 530 ; Lucas & le Souef, 1909, Anim. Austral. : 282, fig. ; Steel,
1912, Aust. Nat., 2 : 135 ; Fry, 1914, Rec. II'. Aust. Mus., 1 : 205, figs. 80, 9.
Helioporus flavoguttatus Fletcher, 1898, Proc. Linn. Soc. N.S.W., 22:679; Nieden, 1923,
Das Tierreich, Anura I : 528 (part).
Vomerine teeth in two long, transverse or slightly oblique series between
the choanae. Sternum in juveniles a broad cartilaginous plate, notched pos-
teriorly, but becoming calcified, the calcified style and posterior edge becoming
distinctly bifurcate with increasing age. Head broad, depressed. Snout rounded,
as long as the eye, with rounded canthus rostralis and very oblique loreal region ;
nostril equidistant from the end of the snout and the eye, or a little nearer the
latter ; interorbital space slightly narrower than the upper eyelid ; anterior
corner of the eye with a fimbriated flap of skin, well-developed in adults, less so
in half-grown examples and absent in the very young. Tympanum subcircular,
its horizontal diameter f to f the length of the eye. Fingers well developed, the
first a little longer than the second, which is a little longer than the fourth ; sub-
articular tubercles large ; an oval inner, and a group of 3 outer metacarpal
tubercles, as well as numerous palmar tubercles, of which the most constant are
one each between the first and second and second and third fingers. Toes with
a distinct, though slightly variable, rudiment of rather fleshy web ; outer toe
variable in length, but always shorter than the third ; a very large compressed
inner metatarsal tubercle. Tarso-metatarsal articulation reaching the rye, the
tip of the outer toe extending to the tip of the snout or a little beyond
Skin regularly shagreened above, more coarsely granular on the flanks ; a
large parotoid glandular complex not easily distinguished externally, and a small
oval gland on the antero-ventral surface of the thigh close to the groin. Lower
surfaces smooth or wrinkled ; anal region granular.
40 H. \V. PARKER
Dark grey- or purplish-brown above ; frequently a vertical yellow bar on
the tip of the snout ; warts of the flanks and posterior half of the upper lip yellow.
Lower surfaces white except the throat of the male, which has a dusky wash or
reticulation.
Male at the breeding season with a series of horny, conical spines on the
dorsal surfaces of the first, and sometimes also the second and third fingers.
The largest is situated on the metacarpo-phalangeal knuckle of the pollex and
has a bony core. Both sexes are also more or less profusely covered with small
black spinules. No vocal sac.
Length from snout to vent : <$ 80 mm. ; § 77 mm.
Distribution : New South Wales (Cumberland County) and West Australia
(Darling Range to Kajonup in the region of moderate rainfall).
Specimens Examined.
1925.8.24. 1-2 <J°_ Hornsby, N.S.W. (Rosenberg.)
1937. 7. 22. 15-26 , <J$andjuvs. New South Wales. Harrison.
1937.7.22.27 <J St. Ronan's Well, York, Nicholls.
W.A.
Heleioporus eyrei (Gray).
Perialia eyrei Gray, 1845, in Eyre, Journ. Exped Centr. Austral., 1 : 407, pi. 2, fig. 3 (Type
locality : — " On the banks of the River Murray "').
Heleioporus albopunctatus (non Gray) Giinther, 1858, Cat. Balr. Sal. Brit. Mus. : 39 (part) ;
Keferstein, 1868, Arch. Naturgesch.. 34 : 269 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus.,
ed. 2 : 271 (part) ; Werner, 1914, Fauna S.-W. Austral., 4, 10, Amph. : 417 ; Dakin,
1920, Aust. Zool., 1, 8 : 241 ; Nieden, 1923, Das Tierreich, Anura I : 526, fig. 357 ;
Glauert, 1929, /. roy. Soc. W. Aust., 15 : 44.
Helioporus insularis Loveridge, 1933, Occ. Pap. Boston Soc. nat. Hist., 8 : 92 (Type locality : —
Rottnest Island) ; idem, 1935, Bull. Mus. comp. Zool. Harv., 78 : 15.
Vomerine teeth in two long, transverse series between the choanae. Sternum
calcified in the adult and notched posteriorly. Head broad, depressed. Snout
rounded, a little longer than the eye, with rounded canthus rostralis and oblique
loreal region ; nostril equidistant from the eye and the end of the snout ; inter-
orbital space a little narrower than the upper eyelid ; anterior corner of the eye
with or without some small papillae. Tympanum more or less distinct, vertically
oval, the annulus tympanicus with a vertical diameter | to f that of the eye.
Fingers well developed, the first a little longer than the second, which is distinctly
longer than the fourth ; subarticular tubercles prominent ; a prominent oval
inner metacarpal tubercle (covering a bony prepollex in both sexes) and a group
of smaller outer metacarpal tubercles ; numerous palmar tubercles and conical
ones between the first and second, and second and third fingers. Toes with a
rudiment of fleshy webbing, the outer shorter than the third ; a very large com-
pressed inner metatarsal tubercle. Tarso-metatarsal articulation reaching the
eye, the tip of the fifth toe extending to the tip of the snout or a little beyond.
1 It has been suggested that this is the West Australian River Murray and not the
better known river of the same name in South Australia. There is some evidence to support
this view. Although the specimens are said to have been collected by Eyre, there is no
proof that this is so, and Gray expressly states that " The British Museum has received
from the different travellers various other species " (op. cit. : 406). The types are
entered in the register of 1843 as having been purchased from Turner (a dealer) and the
locality is given as West Australia. Other frogs in the same collection are Limnodynastes
dorsalis dorsalis and Crinia signifera signifera, of which the former does not occur in South
Australia.
AUSTRALASIAN LEPTODACTYLIDAE 41
Skin shagreened above, more definitely tubercular on the flanks ; a parotoid
complex, not discernible externally, and a small oval gland on the antero-ventral
surface of the thigh, close to the groin. Lower surfaces smooth. Anal region
granular. Brown above, with irregular lighter (grey or yellowish) marblings, of
which the most constant are a pair of oval spots above the shoulder and a vertical
light bar on the tip of the snout ; flanks and anal region sparsely dotted with
yellow or white. Lower surfaces immaculate white except the throat of the
male, which is slightly infuscate.
Male without or with only a single spine on the metacarpo-phalangeal knuckle
of the inner finger ; no vocal sac.
Dimensions : cJ<J 46-56 mm. (av. of 9 specs. 52 mm.) ; $? 48-59 mm. (av.
of 8 specs. 53 mm.).
Distribution : Coastal Zone of West Australia, west of the Darling Range,
south to Albany county, but excluding the damp south-west corner. Rottnest
Island ; ? Northern West Australia.
The species, which has usually been confused with alboputictatus, is closely
allied to australiacus and may prove to be a neotenic form of that species. In
West Australia the three form a complex, of which eyrei occupies the dry zone
to the west of the Darling Range, australiacus the relatively damp areas on both
sides of the mountains and albopunctatus the dry zone to the east. The two latter
occur together in the neighbourhood of St. Ronan's Well, but eyrei and albo-
punctatus meet in the south at Toolbrunup {circa 50 miles north of Albany).
Eggs laid in a frothy mass of jelly underground in April. Development
proceeds within the egg until the external gills have been lost and the operculum
developed ; hatching is conditional upon the nest being flooded. A well-
developed tadpole measures 58 mm., of which the head and body account for 26.
Eyes and nostrils directed upwards, the latter much nearer a line connecting the
anterior borders of the eyes than the end of the snout. Mouth terminal, directed
downwards, lips with a single row of papillae, the upper widely interrupted in the
middle ; angles of the mouth with short, transverse rows bearing three or four
papillae within the lips ; mandibles powerful and finely serrated ; three or four
rows of labial teeth above, the inner two or three widely interrupted mesially ;
three below, the outer shortest and the innermost narrowly interrupted in the
middle. Spiraculum sinistral, situated towards the ventral side. Anus dextral,
slightly anterior to the base of the tail. Young at metamorphosis 23 mm. from
snout to vent.
Specimens Examined.
B.M.
1 Fletcher (1893 : 233) has suggested that this example was one oi a series collected in
1866 by Masters in West Australia ; but it was received in London in 1862, the same year
in which the Leiden Museum received the specimens labelled " King George's Sound."
Heleioporus sudelli Lamb.
Heleioporns sudelli Lamb, 1911, Ann. Queensland Mus., 10 : 26 (Type locality: — Warwick,
S.E. Queensland).
Vomerine teeth in two groups between the choanae. Habitus stout. Head
broader than long, snout rounded, without canthus rostralis ; nostril nearer the
eye than the tip of the snout ; interorbital space narrower than the upper eyelid ;
tympanum indistinct. Fingers blunt, first and second equal ; toes short, blunt,
half webbed ; subarticular tubercles, especially those of the fingers, well developed ;
a small tubercle between the first and second and second and third fingers ; inner
metatarsal tubercle long and low ; no outer metatarsal tubercle. The hind limb
being carried forwards along the body, the tip of the longest toe reaches the eye
or just beyond. Upper surface of body and limbs minutely tubercular ; belly
smooth. Brownish or olive above marbled with darker ; a dark cross-band
between the eyes to the outer edge of the upper eyelid ; a light vertebral line and
a light blotch on the upper part of the shoulders ; upper edge of lower eyelid
silvery white.
Length from snout to vent : 45 mm.
Distribution : Known only from the unique type from Warwick, S.E.
Queensland.
(Possibly based on a specimen of Limnodynastes sp.)
LIMNODYNASTES Fitzinger.
Cystignathus (part) Dumeril & Bibron, 1841, Erpet. Gen., 8 : 392.
Discoglossus Gray, 1842, Zocl. Miscellany : 56.
Limnodynastes Fitzinger, 1843, Syst. Rept. : 31 (Type species : — Cystignathus peronii Dum.
& Bibr.) ; Giinther, 1858, Cat. Batr. Sal. Brit. Mus. : 32 ; Steindachner, 1867, Reise
Novara, Zool., A mph. : 25 ; Keferstein, 1867, Nachr. Ges. Wiss. Gottingen, 18 : 343 ; idem,
1868, Arch. Naturgesch., 34 : 256 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 :
258 ; Nieden, 1923, Das Tierreich, Anura 1 : 528 ; Waite, 1929, Rep. Amph. S. Austral. :
250 ; Noble, 1931, Biol. Amph. : 497.
Lymnodynastes (emend.) Cope, 1866, nat. Hist. Rev. : 113 ; M'Coy, 1867, Ann. Mag. nal.
Hist., (3), 20 : 182 ; Alexander, 1922, /. Linn. Soc. {Zool.), 36 : 462.
Wagleria Girard, 1853, Proc. Acad. nat. Sci. Philad., 6 : 421 (Type species by selection : — Cysti-
gnathus peronii Dumeril & Bibron) ; idem, 1858, U.S. Explor. Exped., Herp. : 42.
Platyplectrum Giinther, 1863, Ann. Mag. nat. Hist., (3), 11 : 27 (Type species: — P. marmo-
ratum); Cope, 1866, /. Acad. nat. Sci. Philad., (2), 6 : 89, 94; Keferstein, 1867, Nachr.
Ges. Wiss. Gottingen, 18 : 343 ; idem, 1868, Arch. Naturgesch., 34 : 260.
Borborocaetes (part) Cope, 1866, J. Acad. nat. Sci. Philad., (2), 6 : 94 ; idem, 1889, Bull.
U.S. nat. Mus., 34 : 312.
Opisthodon Steindachner, 1867, Reise Novara, Zool., Amph. : 9 (Type species : — O.frauenfeldi).
Heltorana idem, 1867, torn. cit. : 32 (Type species : — H. grayi).
Ranaster Macleay, 1878, Proc. Linn. Soc. N.S.W., 2: 135 (Type species : — R. convexiusculus) ;
Nieden, 1923, Das Tierreich, Anura I : 535 (part) ; van Kampen, 1923, Amph. Indo-
Austral. Arc/tip. : 19 ; Noble, 1931, Biol. Amph. : 497.
Maxillary teeth present. Prevomer present, entire, its post-choanal portion
large, overlying at least the mesial half of the palatine, dentigerous (text-fig. 9) ;
AUSTRALASIAN LEPTODACTYLIDAE
43
a distinct fronto-parietal foramen. Ear fully developed. Vertebrae procoelus
but with the condyle incompletely ankylosed ; seven presacrals ; sacral diapo-
physes slightly dilated ; coccyx articulating by two condyles. Omosternum
cartilaginous ; sternum undivided, cartilaginous or calcified. Terminal phalanges
simple. Males with a prepollex (PL I).
Pupil horizontal, with a downwardly directed angle ventrally. Tongue
large, oval, entire and slightly free behind. Toes usually more or less webbed,
not dilated distally.
Text-fig. io.
Text-fig.
Text-fig. ii.
Text-fig. 12.
Text-fig. 9. — Anterior cranial elements of Limnodynastes dorsalis dorsalis. X
(From beneath.)
Text-fig. 10. — Hand of Limnodynastes iasmaniensis $. X 4.
Text-fig. i i . — Hand of Limnodynastes salmini ?. X 4.
Text-PIG. 12. — Hand of Limnodynastes peronii $>. X 4.
T )i>t;tl tendon of the m. smiitendinosus passing ventral to, or perforating
the gracilis (ornatus). Alary process of the hyoid narrow proximally, expanded
distally ; cricoid complete ; oesophageal process of the cricoid slender.
This genus shows a very peculiar trend. In two species, fletcheri and perorm
(text-fig. 12), and to a less extent in a third, salmini (text-fig. 11), the phalanges
of the inner finger are greatly reduced in size, or, in some specimens, reduced
to a single one, but as a compensation, so that the inner finger is still approxi-
mately as long as the second, the inner metacarpal bone is disproportionately
44 H. W. PARKER
long. This compensation belies the possibility of the reduction being a concomi-
tant of loss of function unless it is assumed that there has been some secondary
change necessitating a re-development of an inner digit which was originally
functionless and vestigial. There is no evidence to lend support to such an
assumption and it is possible that an explanation is to be found by comparison
with the genus Crinia, in which also two species (laevis, text-fig. 16, and dar-
lingtoni) have the phalanges of the inner finger similarly reduced, though without
any compensating elongation of the metacarpal. Now it may be significant
that none of the species of Limnodynastes in which the phalanges of the inner
finger are reduced in size have nuptial pads, whereas all the remaining species of
the genus have them very well developed. In Crinia laevis, and probably C.
darlingtoni also, nuptial pads are likewise absent. But the former species at
least, and in all probability the latter too, pairs on land, whereas all the species
of Limnodynastes, whose breeding habits are known, including peronii with the
reduced phalanges, pair in water. It is highly probable that nuptial excrescences
are not so essential to frogs pairing on land as to those where oviposition occurs
in water. Reduction of the phalanges of the inner digit appears to be accom-
panied by loss of nuptial pads, but in the genus Crinia, where pairing occurs on
land, this is no handicap. In the water-breeding Limnodynastes, however, the
loss would be gravely disadvantageous and has been overcome by the enlarge-
ment of the metacarpal which can function as a nuptial spine, as it does in the
Ranid genera Petropedetes and Babina. It seems possible, therefore, that the
reduction of the inner finger is an orthogenetic tendency which proceeds regardless
of the requirements of the animals, and that to counteract the handicap it imposes
L. fletcheri and L. peronii have secondarily hypertrophied another structure to
perform the same function in a different way.
Synopsis of the Species.
I. Inner metatarsal tubercle shovel-shaped.
A. A large gland on the tibia dorsalis subsp.
B. No tibial gland.
(i) Toes not more than \ webbed ..... ornatus.
(2) Toes at least A webbed spenceri.
II. Inner metatarsal tubercle not shovel-shaped.
A. Inner metacarpal equal to, or somewhat longer or shorter than, the
second ; the proximal phalanx of the first finger very much longer
than the distal.
(1) First finger shorter than the second ; usually two metatarsal
tubercles ; vomerine teeth not extending laterally beyond
the choanae tasmaniensis.
(2) First and second fingers equal ; a single metatarsal tubercle ;
vomerine teeth extending well beyond the lateral borders of the
choanae ........ convexiusculus.
(3) First finger extending beyond the second, the proximal phalanx
of the inner finger twice as long as the second ; toes nearly free
salmini.
B. Inner metacarpal very much longer than the second ; phalanges of
the first finger subequal, or (in peronii) sometimes reduced to one.
(1) Toes with a distinct basal webbing ; snout not prominent . fletcheri.
(2) Toes long, slender, free ; snout pointed and prominent . peronii.
AUSTRALASIAN LEPTODACTYLIDAE 45
Limnodynastes dorsalis dorsalis (Gray).
Cystignalhus dorsalis Gray, Ann. Mag. nat. Hist., 7 : 91 (Type locality: — Western Australia) ;
idem, 1841, in Grey, Journ. Exped. W. Australia, 2, App. : 446; idem, 1845, in Eyre,
Journ. Exped. Central Austral., 1, App. : pi. 1, fig. 2.
W. {agleria) dorsalis Girard, 1853, Proc. Acad. nat. Sci. Philad., 6 : 421.
Limiwdynastes dorsalis Giinther, 1858, Cat. Batr. Sal. Brit. Mus. : 33 ; Krefft, 1865,
Pap. roy. Soc. Tasmania : 16 (part) ; idem, 1868, Arch. Nat urge sell. ,34 : 260 ; Boulenger,
1882, Cat. Batr. Sal. Brit. Mus., ed. 2: 261 (part); Fletcher, 1898, Proc. Linn. Soc.
N.S.W., 22, 1897 : 675; Andersson, 1913, A'. Svenska VeienskAhad. Handl., 52, 4 :
9, 10 ; Werner, 1914, Fauna S.W. Austral., 4 : 406; Alexander, 1922, /. Linn. Soc.
(Zool.), 34 : 462 ; Nieden, 1923, Das Tierreich, Anura I : 532 (part) ; Glauert, 1929,
J. roy. Soc. \V . Aust., 15 : 44 ; Noble, 1931, Biol. Arnph. : 1 14.
Limnodynastes dorsalis var. typica Fry, 1913, Pec. Aust. Mus. 10 : 24, pi. 2, fig. 2;
idem, 1914, Rec. W. Aust. Mus., 1 : 202.
Limnodynastes dorsalis dorsalis (part) Loveridge, 1935, Bull. Mus. comp. Zool. Han.,
78": 21.
Vomerine teeth in moderately long curved series separated from the maxillae
by spaces equal to, or a little greater than, half the length of a single series.
Snout rounded, not prominent, 1-25 to 1-5 times as long as the eye, with rounded
canthus rostralis and oblique, convex, loreal region ; nostril midway between
the eye and the tip of the snout ; interorbital space about as broad as the upper
eyelid ; tympanum indistinct, its horizontal diameter about half that of the eye.
Fingers moderate, the first very slightly longer than the second, the first meta-
carpal also being slightly longer than the second ; subarticular tubercles well
developed ; one or two small tubercles between the first and second, and second
and third fingers ; a prominent inner metacarpal tubercle, with a bony prepollex
and a flat, longitudinally cleft outer. Toes with a rudiment of web, variable
in amount, but not more than \ ; inner metacarpal tubercle large, shovel-shaped,
longer than its distance from the tip of the inner toe. Tibio-tarsal articulation
reaching the tympanum or the posterior corner of the eye.
Skin smooth, feebly granular or with some flat warts above ; a prominent
glandular ridge from beneath the eye to the forearm and a very large, conspicuous
gland occupying almost the whole upper surface of the tarsus. Smooth beneath ;
anal region feebly granular, with two larger glandular spots one on each side of
the vent.
Pale brown or greenish above with insuliform, clearly-marked dark brown,
sometimes light-edged spots in regular series ; a median series down the middle
of the back, commencing with a triangular one connecting the upper eyelids, is
bisected by a light vertebral line which is constantly present. A dorso-lateral
series runs from behind the eye to the groin. Flanks spotted and splashed with
darker, the inguinal region often marbled with white (? red) and brown. A
curved bar from the tip of the snout through the nostril and eye to the fore-limb ;
sub-aural glandular fold white or yellow. Hind limbs with dark spots on a
lighter ground above ; hinder side of the thighs and concealed surface of the
tibiae dark brown marbled with white. Lower surfaces white, sometimes faintly
<lusted with darker. Throat of the male infuscate.
Male with a vocal sac opening by a slit on each side of the tongue and nuptial
asperities on the metacarpo-phalangeal knuckles of the first and second fingers.
Length from snout to vent : <J 68 mm.
Distribution : West Australia ; Rottnest Island.
Specimens Examined.
B.M. 41. (2). 212 ^ W.Australia. Gould. Type.
41.3.6.7-8 2 o"(J Port Essington (?). Fleming.
43.5.19.27 juv. W.Australia. [Turner.]
46 H. W. PARKER
44.7.9.34 (J Houtman's Abrolhos. Gould.
95.6.21.6-7 2 (J (J Chapman River, W. A. Saunders.
1931 .7. 1.46-48 3 juvs. Mt. Toolbrunup, W.A. Baldwin.
96.1.30.8 skel. Chapman River, W.A. Saunders.
Limnodynastes dorsalis terraereginae Fry.
Limnodynastes dorsalis Giinther, 1876, J. Mus. Godeffroy, 12 : 47 ; Boulenger, 1882,
Cat. Batr. Sal. Brit. Mus., ed. 2 : 261 (part) ; Garman, 1901. Bull. Mus. comp. Zool.
Harv., 39 : 14 ; Andersson, 1916, K. Svenska VetenshAkad. Hand!., 52, 9:8; Nieden,
1923, Das Tierreich, Anura I : 532 (part).
Limnodynastes dorsalis var. dumerili (non Peters) Fry, 1913, Rec. Aust. Mus., 10 : 26-28,
pi. 3, fig. 2.
Limnodynastes dorsalis var. terrae-reginae Fry, 1915, Proc. roy. Soc. Queensland, 27 : 67, fig.
2a (Type locality: — Somerset, Cape York).
L. dorsalis terraereginae Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 21.
This north-eastern race is similar in general morphological characters to the
typical western form, except that the vomerine teeth may be slightly more exten-
sive laterally, the first finger rather shorter (being only as long as, or but very
slightly longer than, the second), shorter hind limbs, the tibio-tarsal articulation
reaching the shoulder or the ear, and the sub-aural glandular fold expanded
behind the angle of the mouth.
The colour is, however, very different. In juveniles there may be a pattern
of spots similar to that of the typical subspecies, but the light mid-dorsal stripe
is narrower and often partly or completely obliterated. With increasing size the
dorsal markings break up into smaller spots and the ground-colour tends to become
progressively darker, so that in adults the dorsal colour may be almost uniform
dark olive ; usually, however, there is some persistent trace of the spots and also
there is often a light streak, with indefinite margins running forwards from the
groin, obliquely upwards in the direction of the eye. Chin and throat with brown
reticulations. Males with the same secondary sex-characters as in L. dorsalis
dorsalis and females with the first and second fingers spatulate.
Length from snout to vent : <$ 69 mm. ; $ 76 mm.
Distribution : Queensland as far south as the Burnett River Valley.
Limnodynastes dorsalis dumerili Peters.
Limnodynastes (Platyplectron) dumerili Peters, 1863, Mber. Ahad. Berlin : 235 (Type locality: —
near Adelaide) ; Giinther, 1867, Ann. Mag. nat. Hist., (3), 20 : 54.
Platyplectrum dumerili Keferstein, 1868, Arch. Naturgesch., 34 : 262.
Limnodynastes dumerili Giinther, 1868, Zool. Rec, 1867, 4 : 145 ; idem, 1868, Proc. zool.
Soc. Lond. : 478.
Limnodynastes dorsalis var. dumerili Fry, 1913, Rec. Aust. Mus., 10, 2 : 26 (part), pi. 2,
figs, i-ia ; Waite, 1929, Rept. Amph. S. Austral. : 252, figs. 181-183.
Limnodynastes dorsalis dumerili Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 21
(part).
Limnodynastes bibroni (nom. nud., in err. pro dumerili?) Krefft, 1865, Pap. roy. Soc.
Tasmania : 16 ; idem, 1867, Cat. Industr. Prod. N.SJV., Add. : 107.
L. (ymnodynastes) dorsalis McCoy, 1867, Ann. Mag. nat. Hist., (3), 20 : 182.
AUSTRALASIAN LEPTODACTYLIDAE 47
Limnodynastes dorsalis McCoy, 1880, Prodr. Zool. Victoria, 5 : 12, pi. 42, fig. 2 ; Lucas &
le Souef, 1909, Anim. Austral. : zji, fig. ; Nieden, 1923, Das Tierreich, Anura I : 532
(part).
Limnodynastes dorsalis var. interioris Fry, 1913, Rec. Aust. Mus., 10, 2 : ^, pi. 3,
figs, i-ic (Type locality :— Merool Creek, Riverina, N.S.W.).
As understood by Fry (1913) this race ranges from Southern Queensland
southwards to South Australia and Tasmania except for a small area in western
New South Wales where his interioris was discovered. Loveridge (1935) , however,
includes Tasmanian specimens under the western dorsalis dorsalis, but it seems
highly probable that there is a distinct and recognizable race in Tasmania.
The status of Fry's interioris, with its strongly webbed toes and heavily
pigmented lower surfaces, is uncertain, but in view of the fact that this author
records specimens with these two characters from the central table-land and the
mountains of New South Wales it seems possible that interioris ought to be
united with dumerili. The material before the author is very scanty, but, so far as
it goes, the race appears to be distinguished by a somewhat shorter snout, less than
1-25 times as long as the eye ; the first finger slightly but distinctly shorter than
the second ; toes -}, or even slightly more, webbed ; a very conspicuous wart
on each side of the vent and the lower surfaces heavily marbled with brown.
Distribution : South Australia, Victoria, southern and western New South
Wales, including the mountain region.
Specimens Examined.
64.10.27.32-33 <J, Cj Adelaide. Krefft.
04. 10. 27.59-60 ?, Hgr.
Mus. Leiden 2033 $ Melbourne. van Kaathoven,
Limnodynastes dorsalis insularis subsp. n.
Limnodynastes dorsalis English, 1910, Proc. zool. Soc. Land. : 629, pi. 51, fig. 3 ; Nieden,
1923, Das Tierreich, Anura I : 532 (part) ; Blanchard, 1929, Aust. Zool., 5, 4 : 326.
/. imnadvnastcs dorsalis var. dumerili Fry, [913, & , . lust M us., 10, 2 : 26 (part)
Limnodynastes dorsalis dorsalis (part) Loveridge, 1934, PaP- roy. Soc. Tasmania, 1933 : 58
idem, 1935, Bull. Mus. comp. Zool. Harv., 78 : 21.
As suggested above, the Tasmanian representative of Limnodynastes dorsalis
may be recognizable as a distinct race. It resembles L. dorsalis dumerili in having
the first finger slightly, but distinctly, shorter than the second and in the possession
of a pair of large, prominent anal warts ; the colour, too, is essentially similar
to that of specimens from New South Wales and South Australia, though the
lower surfaces are only slightly brown-mottled. There is also some resemblance
to terrae-reginae in the degree of dilatation of the sub-aural fold posteriorly. It
differs from dumerili, however, in having a snout 1-5 to 17 times as long as the
eye and in the toes being almost entirely devoid of webbing.
It is essentially a burrowing frog, frequenting dry country, and hibernates
from mid-April until late in August. The breeding season, according to English,
is from August to October, the call of the male being a very loud booming note ;
Blanchard, however, found the frogs breeding in January, the eggs being laid in
floating, frothy clumps, often in contact with grass or sedges.
Length from snout to vent : <J 58 mm. ; $ 57 mm.
Specimens Examined.
B.M, 1901.9.13.1 imm. (J Near Launceston. English. (Cotype.)
M.C.Z. Harvard 19238 $ Eagle Hawk Neck, Blanchard
Tasmania.
'933° <J Tasmania. English
48 H. W. PARKER
Limnodynastes dorsalis grayi (Steindachner).
Limnodynastes dorsalis Krefft, 1865, Pap. roy. Soc. Tasmania : 16 (part) ; idem, 1866,
Trans. Phil. Soc. N.S.W., 1862-5 : 32 ; idem. 1867, Cat. Industr. Prod. N.S.W., Add. :
107 ; Fletcher, 1889, Proc. Linn. Soc. N.S.W., (2), 4 : 374 ; idem, 1890, op. cit., (2),
5 : 670-675 ; idem, 1891, op. cit., (2), 6 : 271-273 ; idem, 1892, op. cit., (2), 7 : 7, 8, 13 ;
idem, 1894, o£. «/., (2), 8 : 526, 528, 530 ; Harrison, 1922, Aust. Zool., 3, I : 33 ; Nieden,
1923, Das Tierreich, Anura I : 532 (part), figs. 362, 363.
Heliorana grayi Steindachner, 1867, Reise Novara, Zool., Amph. : 32, pi. 2, figs, n-14
(Type locality: — New South Wales) ; Keferstein, 1868, Arch. Naturgesch., 34 : 266.
Platyplectrum superciliare Keferstein, 1867, Nachr. Ges. Wiss. Gbttingen, 18:346 (Type
locality : — Australia) .
Heliorana superciliaris Keferstein, 1868, Arch. Naturgesch., 34 : 267, pi. 5, fig. 7.
Limnodynastes dorsalis var. dumerili Fry, 1913, Rec. Aust. Mus., 10, 2 : 26 (part).
Limnodynastes dorsalis dumerili Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 21 (part).
Fry (1913 : 29), in his consideration of the races of Limnodynastes dorsalis,
has pointed out that dumerili shows a great amount of local variation. If geo-
graphical races are to be recognized at all it seems certain that dumerili, sensu
Fry, must be subdivided, and one of its constituents is the small coastal form
of New South Wales, for which the name Heliorana grayi seems to be available.
This race is generally similar to the typical form, but is distinguished by its
smaller size (it rarely exceeds 55 mm. in length), subequal first and second fingers,
a pair of prominent warts on either side of the vent, and the subaural glandular
fold strongly dilated posteriorly.
The coloration is similar to that of the northern terrae-reginae except that
the hinder sides of the thighs are brown, regularly dotted with white or red.
Lower surfaces white or faintly brown dusted.
Distribution : New South Wales east of the dividing range ; ? South-eastern
Queensland.
Breeding period probably extensive ; spawn in a mass of frothy jelly ; tad-
poles very large, 2-5-3 m- m length ; young at metamorphosis 21 mm.
(Fletcher).
Specimens Examined.
B.M. 88.7.3. 11 6* Sydney. Fletcher.
1925. 5. 25. 1 $ Nowra, N.S.W. Rodway.
1929. 7.9. 1-7 7 juvs.
Limnodynastes ornatus (Gray).
Discoglossus ornatus Gray, 1842, Zool. Miscellany : 56 (Type locality : — Port Essington) ;
idem, 1845, in Eyre, Journ. Exped. C. Austral. II : pi. 2, fig. 2.
Perialia ? ornata idem, 1845, torn. cit. : 407.
Limnodynastes ornatus Gunther, 1858, Cat. Batr. Sal. Brit. Mus.: 33 ; Krefft, 1865, Pap.
Proc. roy. Soc. Tasmania: 17 ; Gunther, 1867, Ann. Mag. nat. Hist., (3), 20 : 54 ; idem,
1868, Zool. Rec, 1867, 4 : 145 ; idem, 1876, Mus. Godeffrov, 12 : 47 ; Boulenger, 1882,
Cat. Batr. Sal. Brit. Mus., ed. 2 : 262 ; Fletcher, 1889, Proc. Linn. Soc. N.S.W. , (2),
4 : 386 ; idem, 1890, op. cit., (2), 5 : 672 ; idem, 1894, op. cit., (2), 8 : 524, 529, 530 ;
Boettger, 1894, Denkschr. med.-naturw. Ges. Jena, 8 : 109,110; Fletcher, 1898, Proc.
Linn. Soc. N.S.W., 22, 1897 : 676; Lucas & le Souef, 1909, Anim. Austral. : 273,
fig.; Andersson, 1913, K. Svenska VetenskAhad. Handl., 52, 4:9, 11; Fry, 1914,
Rec. W.Austr. Mus., 1 : 202; Andersson, 1916, K. Svenska VetenskAhad. Handl., 52,
9:8; Nieden, 1923, Das Tierreich, Anura I : 533 (part) ; Kinghorn, 1931, Rec. Aust.
Mus., 18, 3 : 89 ; Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 22
Platyplectrum ornatum Keferstein, 1867, Nachr. Ges. Wiss. Gottingen, 18 : 346 ; idem, 1868,
Arch. Naturgesch., 34 : 261.
Platyplectrum marmoratum Gunther, 1863, Ann. Mag. nat. Hist., (3), 11 : 27, pi. 4, fig. a
AUSTRALASIAN LEPTODACTYLIDAE 49
(Type locality : — Clarence River, N.S.W.) ; Krefft, [865, Pap. Proc. roy. Soc. Tasmania :
17 ; idem, 1867, Cat. Industr. Prod. N.S.W., Add. : 107 ; Keferstein, 1867, Nachr.
Ges. Wiss. GSltingen, 18 : 345 ; idem, 1868, Arch. Naturgesch., 34 : 261 ; Giinther, 1868,
Proc. zool. Soc. Lond. : 480.
? Platxplcctrum m, idaitalr (."ope. iSiiii. /. . lead. nat.Sci. Pliilad., (2), 6 : go (Type locality : — ■
West Australia).
Opisthodon frauenfcldi Steindachner, 1867, Reise Novara, Zool., Amph, : <i, pi. 1, figs. 1-4
(Type locality: — New South Wales).
O. frauenfeldti Cope, 1867, /. Acad. nal. Sci. Philad. (2), 6 : 201.
Vomerine teeth in transverse, slightly curved series behind the choanae,
separated from the maxillae by a space equal to about the length of one series.
Snout short, rounded, not prominent, as long as, or very slightly longer than the
eye ; canthus rostralis rounded, loreal region moderately oblique ; nostrils
directed upwards, halfway between the tip of the snout and the eyes or slightly
nearer the latter ; interorbital space less than the width of an upper eyelid ;
tympanum very indistinct, \ to § the diameter of the eye. Fingers short, not
depressed, the first being either slightly longer or shorter than the second ; sub-
articular tubercles well developed and prominent ; a small supernumerary
tubercle proximal to the basal subarticular of each finger and some fleshy web
which may be tubercular between the first and second and second and third
fingers ; a prominent inner metacarpal tubercle, with a bony prepollex in males,
and a longitudinally cleft outer, of which the outermost portion may be very
indistinct or absent. Toes slender, with distinct basal webbing which frequently,
in adults, extends along the toes as a distinct fringe ; in juveniles and immature
specimens the web midway between the third and fourth toes does not extend
as far as the proximal subarticular tubercle of the fourth toe, but in adults it
usually reaches to this level or very slightly beyond it ; subarticular tubercles
prominent ; a large, shovel-shaped inner, but no outer, metatarsal tubercle.
Tibio-tarsal articulation reaching the ear or the eye.
Skin usually with small, not very prominent, warts which may form longi-
tudinal series on the back and on the upper eyelids ; some tubercles at the corner
of the mouth. Lower surfaces smooth ; anal region granular.
Colour very variable ; usually pale brown or grey above with numerous
irregular darker markings, of which the most constant are : — a curved streak from
the end of the snout, through the nostril, along the canthus rostralis, through the
eye and expanding over the tympanic region ; a vertical dark bar below the
anterior half of the eye ; an interorbital bar interrupted In the middle ; an X-
or H-shaped figure on the anterior part of the back, of which the anterior extre-
mities arise from the posterior corners of the eyes. Frequently the area betw een
the interorbital bar and the anterior portion of the X or H is uniformly light in
colour, and one well-marked colour variant is uniformly brown or grey over the
whole of the upper surface of the body, the cantho-temporal streak and labial
markings alone persisting. Usually the whole of the dorsum is covered with
sinuous dark markings which often have less-dark lines running close to and
parallel with their outlines. Limbs with alternately broader and narrower dark
cross-bars. Lower surfaces uniform white except the sides of the throat which
are freckled with darker. Concealed surfaces of the hind limbs dark brown,
uniform or stippled with white.
Length from snout to vent : J 41 mm. : j 42 mm.
Male with a vocal sac opening by a slit on each side of the tongue and with
3 patches of nuptial asperities, one on the dorso-lateral surface oi the inner meta-
carpal tubercle, one on the dorsal and lateral surfaces of the first finger and the
novit. zool., 42, 1 4
50 H. W. PARKER
other similarly disposed on the second finger. Fingers of the female not depressed
or fringed.
Distribution : West Australia, Northern Territories, Groote Eylandt,
Queensland, Thursday Island and northern New South Wales.
Specimens Examined.
Limnodynastes spenceri sp. n.
Limnodynasies ornatus (non Gray) Spencer, 1896, Rep. Horn Exped. C, Austral., II, Zool. :
156, pi. 13, figs. 3, 4 ; pi. 15, figs. 18-25 I Spencer & Gillen, 1912, Across Australia,
I : 58, pi. 1, fig. b ; Harrison, 1922, Aust. Zool., 3, 1 : 34 ; Nieden, 1923, Das Tierreich,
Anura I : 533 (part), figs. 364-367 ; Waite, 1929, Rept. and Amph. S. Australia : 256 ;
Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 22 (part).
Limnodynastes sp. Loveridge, 1938, Trans, roy. Soc. S. Aust., 62, 2 : 189.
Holotype a male, number 97.10.27.69 in the British Museum, collected at
Alice Springs, C. Australia, by Professor W. B. Spencer.
Vomerine teeth in rather short curved series, widely separated from the
maxillae. Snout almost vertically truncate, short, as long as the eye, with
rounded canthus rostralis and slightly oblique loreal region ; nostrils directed
vertically upwards midway between the tip of the snout and the eye ; inter-
orbital space a little less than the width of the upper eyelid ; tympanum hidden.
Fingers short, the first and second depressed ; first a little longer than the second ;
subarticular tubercles large and prominent ; a distinct supernumerary tubercle
at the base of all except the outer fingers, and traces of other tubercles along the
metacarpals ; one or two distinct, conical tubercles between the first and second
and second and third fingers ; a prominent inner metacarpal tubercle, with a
bony prepollex, and a rather flat outer. Toes pointed and webbed to the tips,
the lowest point of the web between the third and fourth toes reaching well
AUSTRALASIAN LEPTODACTYLIDAE 51
beyond the level of the distal subarticular tubercle of the third toe ; subarticular
tubercles small but distinct, the basal one (metatarso-phalangeal articulation)
of the fourth toe wanting ; a very large, shovel-shaped inner metatarsal tubercle.
Tibio-tarsal articulation reaching the tympanic region.
Skin smooth above and below ; anal region granular ; a few indistinct warts
on the upper eyelid.
Pale brownish grey above with irregular dark markings ; a chevron-shaped
cross-bar connecting the upper eyelids and a dark stripe from the posterior corner
of each eyelid, curving towards, but not meeting its fellow on the scapular region ;
remainder of the dorsum with irregularly-shaped dark blotches and spots ; a
dark streak from the labial margin through the nostril and along the canthus
rostralis and a vertical bar running somewhat obliquely forwards from beneath
the eye. Lower surfaces uniform white. Limbs with dark cross-bars.
A vocal sac opening by a slit on each side of the tongue. Four distinct dark
brown patches of nuptial asperities on each hand, one on the lateral aspect of
the inner metacarpal tubercle, another covering the dorsal surface of the inner
finger (except for the terminal joint) a third on the dorso-lateral aspect of the
second finger and the fourth along the inner edge of the third finger. The whole
of the dorsum is more or less beset with minute dark, horny spinules.
Length from snout to vent : 39 mm.
The paratypes agree closely with the type in almost all morphological
characters. The principal variation is to be found in the degree of webbing of
the toes. Spencer has stated that the amount of webbing is independent of sex
and age and that it overlaps that described for omatus. But in this he seems to
be wrong, for no specimen from Central Australia has as little web as orinttits,
in which this character shows very little variation over a wide area (p. 49). In
these Central Australian frogs (which differ also from omatus in their somewhat
shorter truncate snouts, shorter hind limbs, absence of the proximal tubercle
of the fourth toe, and the presence of a nuptial pad on the third finger) the minimum
amount of webbing is in females and immature examples ; but even here the
lowest point of the web between the third and fourth toes reaches almost or quite
to the distal subarticular tubercle of the third toe, whereas in omatus it never
extends beyond half way between the proximal and distal tubercles. In adult
males the lowest point is always beyond the distal tubercle of the third toe.
Length from snout to vent : <J 44 mm. ; 9 49 mm.
Distribution : Apparently confined to the central, desert regions of the
Northern Territories and South Australia from Oodnadatta to the Macdonnel
Range west to Hermannsburg and Officer Creek.
The species frequents soft, sandy ground, generally in the beds of creeks ;
it burrows by day to a depth of about a foot and becomes active by night. Spawn-
ing takes place in the early part of the year (January or February) in waterholes.
The tadpole reaches a length of 56 mm., the body 25 mm. and the tail 31, the
latter being 11 mm. deep and rounded terminally with subequal ventral and
dorsal crests. Body once and a half to nearly twice as long as broad ; nostrils
directed upwards, nearer a line connecting the anterior borders of the eyes than
the end of the snout ; spiraculuin sinistral ; anus dextral. Mouth surrounded
by a fringe of papillae except its median half anteriorly ; the papillae forming .1
single row except at the corners of the mouth where they invade the oral cavity.
Mandibles very finely denticulate ; two long rows of labial teeth anteriorly, the
inner divided mesially ; three long rows posteriori}', the innermost divided, the
outermost shortest. Toes webbed as in adult females.
52 H. \V. PARKER
Specimens Examined.
B.M. 97. 10.27.68 £ Alice Springs, N.T. Australia. Spencer. Type.
97.10.27.60-67 8 cJ(J, 2 $9 ,, ,, ,, ,, \
1910.5.28.32-35 3 0*6*. ¥ Hermannsburg, ,, Hillier.
1932.3.7.1-2 2 juvs. Central Australia. Basedow. |-ara"
M.C.Z. 18530-18546 9<J<J, 11$? Hermannsburg, N.T. Schevill. types.
5. juvs.
B.M. 97.10.27.69 skel. Alice Springs, N.T. Spencer.
1907. 10.3.2 Larva. Hermannsburg. Hillier.
Limnodynastes tasmaniensis Giinther.
Limnodynastes tasmaniensis Giinther, 1858, Cat. Batr. Sal. Brit. Mus. : 33, 134, pi. 2, fig. B
(Type localities: — Tasmania and New Holland) ; Peters, 1863, Mber. Akad. Berlin :
235 ; Krefft, 1865, Pap. Proc. R. Soc. Tasmania : 16 ; idem, 1867, Cat. Industr. Prod.
N.S.W., Add. : 107 ; Steindachner, 1867, Reise Novara, Zool., Amph. : 26 ; Keferstein,
1868, Arch. Naturgesch., 34 : 258 ; McCoy, 1880, Prodrom. Zool. Victoria, 5 : 11, pi. 42,
fig. 1 ; Parker, 1881, Phil. Trans, roy. Soc. Land., 3 : 99, pi. 18, figs. 5-8 ; Boulenger,
1882, Cat. Batr. Sal. Brit. Mus., ed.~2 : 260 ; Fletcher, 1889, Proc. Linn. Soc. N.S.W.,
(2), 4 : 374 ; idem, 1890, op. cit., (2), 5 : 669-675 ; idem, 1891, op. cil., (2), 6 : 271, 272,
274 ; idem, 1892, op. cit., (2). 7:7; idem, 1894, op. cit., (2), 8 : 527, 528, 530 ; Lucas
& le Souef, 1909, Anim. Austral. : 269, figs. ; English, 1910, Proc. zool. Soc. Lond. :
628, pi. 51, figs. 1, 2 ; Fry, 1915, Proc. R. Soc. Queensland, 27, 4 : 62 ; Harrison, 1922,
Au'st. Zool., 3, 1 : 33 ; Nieden, 1923, Das Tierreich, Anura I : 531 ; Waite, 1929,
Rept. Amph. S. Austral. : 251, figs. 179, 180; Kinghom, 1932, Rec. Aust. Mus., 18 :
361 ; Loveridge, 1934, Pap. roy. Soc. Tasmania, 1933 : 59 ; idem, 1935, Bull. Mus.
comp. Zool. Harv., 78 : 19 (part).
Lvmnodynastes tasmanicus McCoy, 1867, Ann. Mag. nat. Hist., (3), 20 : 182.
Limnodynastes peronii var. tasmaniensis Keferstein, 1867, Nachr. Ges. Wiss. Gollingen, 18 :
344.
Limnodynastes peronii (part) Fitzinger, 1861, S.B. Akad. iviss. Wien, 42 : 414.
Limnodynastes affinis Giinther, 1863, Ann. Mag. nat. Hist., (3), 11 : 27 (Type locality: —
Clarence River, N.S.W.) ; Krefft, 1865, Pap. roy. Soc. Tasmania : 16 ; idem, 1867,
Cat. Industr. Prod. N.S.W. , Add. : 107 ; McCoy, 1867, Ann. Mag. nat. Hist., (3), 20 :
182 ; Giinther, 1868, Proc. zool. Soc. Lond. : 480.
Limnodynastes peronii var. rugulosus Keferstein, 1867, Nachr. Ges. Wiss. Gottingen, 18 : 344
(Type localitv : — Sydney).
Limnodynastes platvcephalns Giinther, 1867, Ann. Mag. nat. Hist., (3), 20 : 54 (Type locality :—
Adelaide) ; idem, 1868, Proc. zool. Soc. Lond. : 480 ; Boulenger, 1882, Cat. Batr. Sal.
Brit. Mus., ed. 2 : 260, pi. 17, fig. 3 ; Fry, 1915, Proc. roy. Soc. Queensland, 27, 4 : 63 ;
Nieden, 1923, Das Tierreich, Anura I : 532 ; Loveridge, 1935, Bull. Mus. comp. Zool.
Harv., 78 : 19.
Limnodynastes tasmaniensis var. platycephalus Waite, 1929. Amph. S. Australia : 250.
Vomerine teeth in nearly straight transverse series, almost in contact mesially
but separated from the maxillae laterally by a space nearly equal to the length
of one series. Snout rounded, slightly prominent, 1-3 to 1-5 times as long as
the eye, with rounded canthus rostralis and oblique loreal region ; nostril midway
between the tip of the snout and the eye or very slightly nearer the latter ; inter-
orbital space equal to, or slightly wider than, the width of the upper eyelid ;
tympanum indistinct, about half the length of the eye. Fingers moderate, the
first distinctly shorter than the second, and with a shorter metacarpal than the
second ; subarticular tubercles moderate ; a prominent, inner, metacarpal
tubercle, with a bony prepollex in males, and a flat, longitudinally cleft, outer ;
fingers all with lateral " seams." Toes with a slight indication of web at the
base and with lateral " seams " which rarely are developed to form well-developed
dermal fringes ; subarticular tubercles moderate ; a prominent oval inner, and
usually a very small outer, metatarsal tubercle. Tibio-tarsal articulation reach-
ing the posterior corner of the eye in juveniles and males, the tympanic region
in females.
AUSTRALASIAN LEPTODACTYLIDAE 53
Skin smooth or very slightly warty above ; smooth beneath ; a narrow,
pronounced glandular fold from beneath the eye, beneath the tympanum to the
insertion of the fore limb. Hinder side of the thighs below the vent slightly
granular.
Light brown, yellowish or infused with pink above, usually with regular,
large, dark brown or green spots, of which the most constant is a triangular,
interorbital marking. The number, disposition and size of the spots is variable
and they may be very indistinct. There is constantly a curved, dark brown
stripe from the tip of the snout along the canthus rostralis, through the eye and
tympanum to the fore-limb, above the sub-aural glandular fold, which is yellow,
and frequently a yellow, orange or reddish vertebral stripe from the tip of the
snout to the vent. Limbs with dark cross-bars. Lower surfaces uniform white
except the gular region of the male, which is uniformly dusky.
Male with a vocal sac opening by a slit on each side of the tongue and brown
nuptial asperities at the breeding season on the inner metacarpal tubercle, adjacent
parts of the inner digit and metacarpo-phalangeal knuckle of the second finger.
Breeding females with the first and second fingers strongly depressed and spatu-
late, with thick fleshy fringes more strongly developed on their pre-axial margins.
Length from snout to vent : <$ 39 mm. ; $ 42 mm.
Distribution : Tasmania, Victoria, S. Australia, New South Wales and
southern Queensland.
The species frequents relatively dry areas and breeds throughout the year,
usually after rain. The call of the species is given in two keys, a lower " cook-
kuk-kuk-cook " and a higher (? female) " Kuk-ku-kuk." Amplexus is axillary
and spawning takes place in any available water-supply, though in captivity
the presence of plants in the water appears to have a stimulating effect. The
eggs are small, about 1-3 mm. in diameter, and are enclosed in a floating gelatinous
mass which contains many air-bubbles. This foam " nest " is produced by the
female paddling with her fore-limbs at the surface of the water before and during
the act of oviposition. This action, rendered more efficient by the dilated fingers
of this sex, causes a stream of bubbles to pass backwards under the bodies of the
frogs ; these accumulate in the cloacal region, where they become entangled in
the mucus which precedes and accompanies the extrusion of the eggs. The egg
mass, measuring circa 60 x 35 mm., contains about 200 to 300 eggs which, at
a temperature of 22D C, hatch in 48 hours ; the newly emerged larvae measure
from 2-5 to 5 mm. and immediately attach themselves to water plants, debris,
etc. Growth under favourable conditions is rapid ; the larval stage may be
prolonged to 4 or 5 months and the mature tadpole measures about do mm.,
though lungs are developed and functional 5 days after hatching . the juvenile
frog at metamorphosis measures 15 mm. Sexual maturity is reached (in captivity)
in the following year, 7 months after metamorphosis.
Numerous observations, often contradictory, on the behaviour and breeding
habits of the species have been published, notably in the Blatter fur Aquarun-und
Terrarienkundc. The following references may be cited : Zernecke, 1926, 37 :
4(15; Rehacek, torn. cit. : 120 ; W'olterstortf, ton:, cit. : 4<>; ; Schreitmuller,
1927, 38: 104; Weingand, 1928, 39:30; Geyer, /<>/;;. cit. : 195. 224; Hesse,
1932, 43 : 315 ; Luther, 1933, 44 : 362.
I have been unable to find any characters whereby pkUycephalus may be
distinguished from tasmaniensis, and in view of this and the fact that Loveridge
(1935 : 20) refers two frogs from Kangaroo Island. S. Australia, one to tas-
maniensis and one to plaiycephalus, it seems probable that they cannot l» retained
54
H. W. PARKER
as distinct species. Limnodynastes olivaceus de Vis, on the other hand, appears
to represent a distinct northern species and is not, as Loveridge (op. cit.) believes,
a synonym of tasmaniensis. A few specimens of tasmaniensis in which the outer
metatarsal tubercle is wanting approach the condition of olivaceus (q.v.), but the
two may be distinguished by their inner fingers, vomerine teeth, subarticular
tubercles and gular coloration.
Specimens Examined.
Limnodynastes convexiusculus (Macleay).
Ranaster convexiusculus Macleay, 1828, Proc. Linn. Soc. N.S.W., 2 : 135 (Type locality: —
Katow, Brit. N. Guinea) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 444 ;
Barbour, 1912, Mem. Mus. Comp. Zool. Harvard, 44, 1 : 177 ; Fry, 1913, Mem. Queens-
land Mus., 2 : 47 ; Nieden, 1923, Das Tierreich, Anura I : 537, figs. 369, 370; van
Kampen, 1923, Amph. Indo-Austr. Archip. : 19, fig. 3.
Limnodynastes olivaceus de Vis, 1884, Proc. Linn. Soc. N.S.W., 9:66 (Type locality: — ■
Mackay, Queensland) ; Boulenger, 1885, Ann. Mag. nat. Hist., (5), 16 : 387 ; Fry,
1915, Proc. roy. Soc. Queensland, 27 : 65 ; Nieden, 1923, lorn. cit. : 530.
Phanerotis novae-guineae van Kampen, 1909, Nova Guinea, 9, 1 : 36, pi. ii, fig. 4 (Type
locality : — Merauke, Dutch N. Guinea).
Limnodynastes salmini (part) Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 19.
Vomerine teeth in very long, slightly curved series, almost in contact mesially,
and separated from the maxillae by a space much less than half the length of
one series. Snout pointed, scarcely prominent, 1-5 to 1-8 times as long as the
eye, with rounded canthus rostralis and very oblique loreal region ; nostril
midway between the tip of the snout and the eye or a little nearer the latter ;
interorbital space equal to, or a little broader than the upper eyelid ; tympanum
indistinct, about half the diameter of the eye. Fingers moderate, rounded, with-
out lateral seams ; first finger equal to the second ; first metacarpal as long as
the second ; a very prominent inner metacarpal tubercle, with a bony prepollex
AUSTRALASIAN LEPTODACTYLIDAE 55
in males, and a flat outer, fissured longitudinally ; subarticular tubercles very
large and prominent ; sometimes a supernumerary tubercle on the base of each
finger proximal to the basal tubercle and indications of a tubercle between the
first and second fingers. Toes cylindrical, without fringes or lateral seams, with
the merest indication of web at the base ; subarticular tubercles large and promi-
nent ; a prominent, oval, inner, but no outer, metatarsal tubercle. Tibio-tarsal
articulation reaching the posterior corner of the eye.
Skin with numerous glandular areas above, which may be raised to form
distinct plicae (often as a post-mortem condition when immersed in too con-
centrated a preservative) ; a glandular fold from beneath the eye to the fore-limb ;
lower surfaces smooth ; anal region granular.
Grey or brown above, with numerous insuliform spots irregularly arranged
in linear series ; the most constant are a quadrangular pair closely apposed
commencing one on each upper eyelid and running backwards on to the scapular
region ; a dark blotch commencing on the tip of the snout, running through the
eye and covering the tympanic region ; a dark labial spot below the eye ; sub-
aural glandular fold white. Limbs with irregular dark cross-bars ; hinder side
of the thighs brown, mottled and spotted with white. Lower surfaces white,
the gular region with brown or grey reticulations.
Male with a vocal sac opening by a slit on each side of the tongue and a
nuptial rugosity on the metacarpo-phalangeal knuckle of the first finger. Female
without or with only slightly spatulate fingers.
Length from snout to vent : <$ 44 mm. ; $ 51 mm.
Distribution : New Guinea, Queensland and Northern Territories.
This species is very closely allied to tasmaniensis and to salmini, but may be
distinguished from either by the characters given in the synopsis. Loveridge,
misled by de Vis' original statement that there are two metatarsal tubercles,
concluded that Fry's redescription (1915 : 65) was based on a different frog and
placed olivaceus as a synonym of tasmaniensis. But the type and other material
examined reveals constancy in the presence of only a single metatarsal tubercle
and the presumption is that Fry was correct.
B.M.
Mus.
New Guinea.
<J Merauke, New Guinea. Koch. (C o t y p e of
Phain- rot is novae
guineae van Kampcn.)
$ v. W'eel.
Limnodynastes salmini Steindachner.
Limnodynastes salmini Steindachner, 1807. Reise Novara, Zool., Ainph. : 27, pi. 4. tigs. 12-
15 (Type locality : — Australia) , Keferstein, 1868, Arch.Naturgesch.,34: 259; Boulenger,
1882, Cat. Batr. Sal. Brit. Mas., ed. 2 : 259 ; Fletcher, 1889, Proc. Linn. Soc. N.S.W.,
(2), 4 : 374 ; idem, 1890, op. cit.. (2), 5 : 669-675 ; idem. 1891, op. at.. (2), 6 : 265, 272 ;
idem, 1894, op. at., (2), 8 : 530 ; Boettger, 1894, Denkschr. med.-naturw. Ges. Jena, 8 :
109; Nieden, 1923, Das Tierreich, Anural : 530; Loveridge, 1935, Bull. .Wiir. comp
Zool. liarv., 78 : 19.
Limnodynastes krefftii (non Giinther, 1863) Giinther, 1867, Ann. Mag. nat. Hist., (3). 20 : 54.
56 H. \V. PARKER
Vomerine teeth in long, slightly curved series, almost in contact mesially and
separated from the maxillae laterally by a space much less than half the length
of one series. Snout bluntly rounded, not prominent, about once and a half
as long as the eye, with rounded canthus rostralis and very oblique loreal region ;
nostril midway between the tip of the snout and the eye or a little nearer the
latter ; interorbital space subequal to the width of the upper eyelid ; tympanum
more or less distinct, A to § the diameter of the eye. Fingers moderate, rounded,
without lateral " seams " ; first metacarpal distinctly longer than the second ;
first finger extending beyond the second, the proximal phalanx very much
longer than the distal ; a prominent inner metacarpal tubercle, with a bony
prepollex in males, and a flat outer, fissured longitudinally ; subarticular tubercles
very large and prominent ; a supernumerary tubercle at the base of each finger
and a distinct tubercle between the first and second fingers.
Toes cylindrical, without seams or fringes, with the merest trace of web at
the base ; subarticular tubercles large and prominent ; a prominent, elongate
inner, but no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the
tympanum in adults, the posterior corner of the eye in juveniles.
Skin smooth or with some scattered flat warts above ; smooth beneath ; a
strong glandular ridge from beneath the eye to the insertion of the fore-limb ;
sometimes a strong pectoral fold ; anal region feebly granular.
Brown or pinkish above with small, scattered, subcircular dark spots ; an
irregular dark blotch from the nostril, along the canthus rostralis, through the eye
and covering the tympanic region above the subaural glandular fold, which is
yellow or red ; a dark labial spot below the anterior half of the eye ; a red or
pink line runs from above the shoulder obliquely to the groin and there may be
traces of a similarly coloured vertebral stripe, especially in the coccygeal region.
Groins and concealed surfaces of the hind limbs dark brown with small yellow or
white spots. Lower surfaces dirty white ; gular region more or less heavily
dusted with brown.
Male with a vocal sac opening by a slit on each side of the tongue. First
and second fingers of the female with a fleshy fringe on their preaxial sides.
Length from snout to vent : $ 63 mm. ; 9 7° mm.
Distribution : New South Wales both east and west of the dividing range
as far south as Sydney (Castlereagh) ; E. Queensland.
This species, which is so closely allied to olivaceus that the two have probably
been confused, is recognizable by its larger size, longer first finger and metacarpal,
somewhat shorter legs and colour.
Specimens Examined.
Limnodynastes fletcheri Boulenger.
Limnodvnastes fletcheri Boulenger, 1888, Ann. Mag. nat. Hist., (6), 2 : 142 (Type locality : —
Gu'ntawang, near Mudgee, N.S.W.) ; Fletcher, 1890, Proc. Linn. Soc. N.S.W., (2), 5 :
672, 675 ; idem, 1894, op. cit., (2), 8 : 529 ; Fry, 1915, Proc. R. Soc. Queensland, 27, 4 :
65 ; Nieden, 1923, Das Tierreich, Anura I : 531.
Limnodynastes tasmaniensis (part) Fletcher, 1892, Proc. Linn. Soc. N.S.W. , (2), 7 : 8, 9, 16 ;
idem, 1894, op. cit., (2), 8 : 529.
Limnodvnastes marmoratus Lamb, 1911, Ann. Queensland Mus., 10: 28 (Type locality: —
Goondiwindi) ; Fry, 1912, Rec. Aust. Mus., 9 : 98, 106.
AUSTRALASIAN LEPTODACTYLIDAE 57
Vomerine teeth in relatively short transverse series, separated from the
maxillae by spaces about equal to the length of one series of teeth. Snout
depressed, rounded, 1-3 to 1-5 times as long as the eye, with very obtuse canthus
rostralis and oblique loreal region ; nostril midway between the tip of the snout
and the eye ; interorbital space a little less, or a little greater, than the width of
the upper eyelid; tympanum indistinct, about half the diameter of the eye.
Fingers depressed, with lateral "seams" ; first not extending quite as far as
the second, the proximal and distal phalanges small, subequal, but the first
metacarpal greatly elongate, much longer than the second ; subarticular tubercles
distinct but not prominent ; no supernumerary tubercles and no tubercles between
the fingers. A prominent inner metacarpal tubercle, with bony prepollex in
males, and a small longitudinally cleft outer. Toes pointed, with lateral seams
and some webbing ; in males the web midway between the third and fourth toes
extends distally beyond the proximal tubercle of the fourth toe ; in female> to
about the centre of this tubercle ; an oval inner, but no outer, metatarsal tubercle.1
Tibio-tarsal articulation reaching the tympanum or the posterior corner of the
eye.
Skin smooth or somewhat warty and glandular above ; smooth beneath ;
anal region feebly granular ; a glandular ridge from beneath the eye to the
insertion of the fore-limb.
Pale brown or grey above, with darker brown or grey spots and suffused with
carmine or pink, especially on the upper eyelids. The most constant markings
are a labial spot beneath the eye, a curved band along the canthus rostralis and
from the eye to the fore-limb, an interorbital blotch and a subquadrangular
mark (very irregular) from the scapular region almost to the sacrum. Sub-
aural fold lighter ; limbs with dark spots and cross-bars above, the concealed
surfaces pale, mottled and stippled with darker. Lower surfaces uniformly
white, except that the edge of the lower jaw and the gular region may be very
feebly stippled with darker.
Male with a vocal sac opening by a slit on each side of the tongue ; first finger
swollen but apparently without a spinose nuptial pad. Second finger of the
female somewhat spatulate.
Length from snout to vent : (J 47 mm. ; ? 41 mm.
Distribution : New South Wales west of the dividing range, and southern
Queensland ; probably extending into northern Victoria and South Australia.
Specimens Examined.
B.M. 88.7.3.7-8 2<J(J' Guntawang, N.S.W. Hamilton. Types.
Limnodynastes peronii (Dum. & Bibr.).
Cystignathus peronii lhnm'nl & Bibron, 1841, Erpit. Gin., 8:409 (Type locality: New
Holland). Limnodynastes peronii Fitzinger, 1843, Syst. Rept. : ji ; Gunther, 1858 -
Batr. Sal. Brit. Mus. : 134 ; Fitzinger. [86x, S.B Akad. wiss. Wien, 42: 114 (part) ;
Keferstein, 1S07, \ ,, ,, 1, 11, . Gdttingen, 18:343; i 58 Xrch Valurgesck.,
34 : 257, pi. 5, fig. 1 ; Houlcnger. 1882, Cat. Batr. Sal. Brit. Mus.. ed. 2 : 258 . idem, 1 88 ,.
1 The original description of an outer tubercle as present appears to be erroneous.
* Originally, but erroneously, described as <J and $.
58 H. W. PARKER
Ann. Mag. nat. Hist., (5), 16 : 386 ; Fletcher, 1889, Proc. Linn. Soc. N.S.W., (2) 4 : 373;
idem, 1890, op. cit., (2), 5 : 669-673; idem, 1891, op. cit., (2), 6 : 272; idem, 1892, op. cit..
(2), 7:8-13; idem, 1894, op. cit., (2), 8:524, 526, 527,529; Lucas & le Souef , 1909, Anim.
Austral. : 269, figs. ; Andersson, 1913, Svenska VetenshAkad. Handl., 52, 4:9;
Harrison, 1922, Aust. Zool., 3, 1 : 33 ; Nieden, 1923, Das Tierreich, Anura I : 529 ;
Trewavas, 1933, Phil. Trans, roy. Soc. Lond., 222, B : 435, fig. 25.
Wagleria peronii Girard, 1853, Proc. Acad. nat. Sci. Philad., 6 : 421 ; idem, 1858, U.S.
Explor. Exped., Herpet. : 42, pi. 3, figs. 29-33.
Limnodvnastes peromi peronii Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 17.
Limnodynastes krefftii Gunther, 1863, Ann. Mag. nat. Hist., (3), II : 26 (Type locality:—
near Sydney) ; Krefft, 1865, Pap. rov. Soc. Tasmania : 16; idem, 1867, Cat. Industr.
Prod. N.S.W., Add. : 107 ; Steindachner, 1867, Reise Novara, Zool., Amph. : 26 ;
Gunther, 1868, Proc. zool. Soc. Lond. : 480.
Limnodynastes peromi var. krefftii Keferstein, 1867, Nachr. Ges. Wiss. Gbttingen, 18 : 345 ;
idem, 1868, Arch. Naturgesch., 34 : 258.
Limnodynastes lineatus de Vis, 1884, Proc. Linn. Soc. N.S.W., 9 : 65 (Type locality:—
Mackay).
Limnodvnastes peronii lineatus Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 18.
Limnodynastes dorsalis dorsalis (part ; spec. 19371) Loveridge, 1934. PaP- roy- S°c-
Tasmania : 58 ; idem, 1935, Bull. Mus. Comp. Zool. Harv., 78 : 21.
Vomerine teeth in long curved series separated from the maxillae by a space
not greater than half the length of one series. Snout rounded, distinctly
prominent, i-6 to i-8 times as long as the eye, with rounded canthus rostralis
and moderately oblique loreal region ; nostril midway between the tip of the
snout and the eye or a little nearer the latter ; interorbital space subequal to
the width of the upper eyelid ; tympanum indistinct or hidden, § to § the diameter
of the eye (exposed by dissection). Fingers depressed, the two inner with more
or less distinct marginal seams ; first not extending quite as far as the second,
with two very small, subequal phalanges, or with but a single one (PL I, figs. E, f);
the first metacarpal greatly elongate and much longer than the second ; subarticular
tubercles distinct, except on the first finger ; no supernumerary tubercles or
papillae between the fingers ; a prominent inner metacarpal tubercle, with a
bony prepollex in males, and a longitudinally cleft outer. Toes long, slender,
rounded, without marginal seams and almost completely free from web ; an
oval inner, but no outer metatarsal tubercle. Tibio-tarsal articulation reaching
the eye.
Skin smooth with some flat glands which may form longitudinal series ;
a pronounced glandular ridge from beneath the eye to the fore-limb ; smooth
beneath ; anal region feebly granular.
Pale brown or grey above, with dark brown markings in the form of rows
of spots or longitudinal stripes arranged thus. A median dorsal stripe commencing
on the synciput, broadening between the eyes, narrowing and running almost to
the vent ; this marking is often bisected lengthwise by a white or yellow vertebral
line. A dorso-lateral stripe or series of spots commences behind the upper eyelid
and runs parallel with the dorsal stripe to the groin. Flanks with some scattered
spots ; a curved band from the tip of the snout, along the canthus rostralis,
through the eye to the fore-limb ; sub-aural fold white or yellow. Limbs with
insuliform spots. Lower surfaces white, stippled with pale brown.
In some northern specimens, described as lineatus by de Vis, the longitudinal
stripes are very regular and clearly marked, but the available material is insuffi-
cient to decide whether or not this is a general characteristic of all northern
frogs. Loveridge (1935 : 18) regards lineatus as a distinct race, with a rather
shorter leg than the southern typical form, but in the two cotypes of lineatus
examined by the present author the hind-limbs are quite as long as in any
specimens from the south.
AUSTRALASIAN LEPTODACTYLIDAE
59
Males have a vocal sac opening by a slit on each side of the tongue and in
the breeding season the first finger is much swollen, though apparently without
any spinose nuptial pad ; instead, the metacarpal appears to furnish the means
of gripping the female, for the phalanges are dislocated from its tip and lie at
right angles to the axis of the digit, very much as in the African Petropedetes
johnstoni. Some specimens seem to have indications that the bone may at
times perforate the skin, as in some Ranids, to form a nuptial spine. In breeding
females the first and second fingers are depressed and spatulate, with fleshy
lateral fringes, more developed on the pre- than post-axial edges.
Length from snout to vent : <$ 61 mm. : $ 58 mm.
Distribution : Queensland, New South Wales, Victoria, Tasmania.
The species breeds at any time of the year, following rain, and shallow,
grassy pools are preferred as the spawning site. The call is an explosive " toe."
The egg-mass is similar to that of L. tasmaniensis (g.v.) but larger, 4-6 in. in
diameter and is usually found entangled in the sedges at the margins of the pool.
Specimens Examined.
B.M. 70.6.26.46-47
70.6.26.55
84.7.4.1-2
63.6. 16.89-91
63.6.16.85
63.6.16.?
62.8.1.29
87.7.16.7
88.7.3.5-6
1925.9.19-1
88.7.3.4
45-5-2J-7
60.6.19.34-35
Mus. Leiden 4251
1729
1864
B.M. 58.11.25.57
M.C.Z. Harvard 193 71
Austr. Mus R.8490
(part)
6*9
6". $
3 juvs.
<S
$
?
juv.
2$?
Hgr.
d*
o*
<??
6"
cJ?
3 Hgr.
6"
?
juv.
Australia (PQueensland).
Queensland.
Mackay, Queensland.
Sydney.
Sydney.
Burrawang, N.S.W.
Nowra, N.S.W.
Warragul, Gippsland,
Victoria.
?
[Erromango.]
Queensland.
Sydney.
?
" Van Diemen's Land
Stanley, Tasmania.
Tweed River, N.S.W.
Krefft.
Ling Roth. (Cotypes
of L. lineatus de Vis.)
Krefft. I Cotypes of L.
,, I krefftii
I Giinther.
(Stevens.)
Fletcher.
Rodway.
Baker.
C. Darwin.
(Cuming.)
(Godeffroy Mus.)
(Sydney Mus.)
( .. „ )
Sir A. Smith.
Scott.
PHILORIA Spencer.
Philoria Spencer, 1901, Proc. ray. Soc. Victoria, (2), 13 : 176 (Type species : — Philoria frosti) ;
Nieden, 1923, Das Tierreich, Anura I : 537 ; Noble, 1931, Biol. Amph. : 498.
Maxillary teeth present. Prevomer present, entire, its post-choanal portion
overlying the mesial half of the palatine and dentigerous, the teeth forming
oblique series nearly in contact mesially, but not extending laterally beyond the
inner borders of the choanae ; a distinct fronto-parietal foramen. Ear fully
developed. Vertebrae procoelous, but with the condyle incompletely ankylosed
and thenotochord persistent ; seven presacrals (?)' ; sacral di.ipophyses moderately
1 [n the single specimen examined there are only 5 free pre-sacral vertebrae of which
the foremost represents 1 + 2 fused. But fused with the sacrum anteriorly are two other
centra, the transverse processes of which arc small but free from the sacral diapophyses.
It seems probable that this shortening of the vertebral column may be an individual anomaly,
but should it prove to be normal it provides a very sharp distinction from all other members
of the subfamily.
60 H. \V. PARKER
strongly dilated ; coccyx articulating by two condyles. Omostemum carti-
laginous. Terminal phalanges simple. Distal tendon of the m. semitendinosus
passing ventral to the head of the gracilis. Alary processes of the hyoid small
and antero-lateral, as in Limnodynastes.
Pupil horizontal with a ventral angle. Tongue heart-shaped, slightly free
behind. Toes quite free, not or but slightly dilated distally.
These characters are taken from a single specimen of what appears to be
a northern species of Philoria and not from the genotype, which is not available
to the author for examination. There is thus room for doubt as to whether
the two really are congeneric and whether the foregoing diagnosis is really
applicable to Philoria. If it is, however, the genus is very closely allied
to Limnodynastes and differs only in the absence of digital webbing and the
reduction of the digits, a character in which it resembles Crinia.
Philoria frosti Spencer.
Philoria frosti Spencer, 1901, Proc. roy. Soc. Victoria, (2), 13 : 176 (Type locality :— Mt. Baw
Baw, Victoria) ; Lucas & le Souef, 1909, Anim. Austral. : 283 ; Nieden, 1923, Das
Tierreich, Anura I : 537.
Habitus stout. Head broader than long ; snout rounded, slightly longer than
the orbital diameter ; nostril nearer to the eye than the tip of the snout ; canthus
rostralis not strongly marked. Interorbital space as broad as the length of
the orbit. Tympanum not visible. Fingers blunt and free ; first finger not
extending so far as the second. Toes short and blunt, no trace of webbing ;
subarticular tubercles present ; three metacarpal tubercles, the inner one strongly,
the outer feebly developed. The inner metatarsal tubercles small and blunt.
Hind limbs short, stout and strongly built ; carried forwards the tibio-tarsal
articulation barely reaches as far forwards as the shoulder. Upper eyelids,
tympanic region, the top of the head and dorsal surface of the body, the upper
surface of the arm and fore arm and the upper surface of the leg covered with
small warts arranged in roughly longitudinal rows along the back. A very large
and prominent triangular shaped parotoid gland is present on either side extending
backwards over the shoulder region ; the longest side of the triangle runs parallel
to the mid-dorsal line, and these two sides are separated from one another by
a space which is narrower than that between the orbits. From the posterior
angle a special row of warts somewhat larger than the rest runs backwards to
the groin and is continued forwards over the surface of the gland, which is other-
wise comparatively smooth. Upper surface of body and limbs a general
dark brown colour with here and there small irregular light patches ; groins and
under surface of body and limbs yellowish, mottled with brown.
Length from snout to vent : 44 mm.
Habitat : Mount Baw Baw, Victoria (Spencer).
Philoria loveridgei sp.n.
Philoria frosti Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 16.
Habitus moderately stout. Head as long as, or longer than, broad ; snout
subacuminate, 1-4 times as long as the eye ; nostril much nearer the eye than
the tip of the snout ; canthus rostralis sub-angular ; loreal region slightly oblique
and distinctly concave ; interorbital space 1-2 to 1-5 times as broad as the upper
AUSTRALASIAN LEPTODACTYLIDAE 6l
eyelid ; tympanum just discernible, or hidden, its horizontal diameter half that
of the eye. Fingers very short, the first much shorter than the second ; small
subarticular tubercles beneath the proximal joints ; two metacarpal tubercles,
the outer largest and notched distally. Toes rather short, quite free, their tips
slightly spatulate ; subarticular tubercles feebly developed ; a small inner, and
no outer, metatarsal tubercle. Tibio-tarsal articulation reaching the tympanic
region in males, or not quite so far in females.
Skin slightly warty or quite smooth above ; a supratympanic fold but no
distinct parotoid gland. Smooth beneath.
Brown above, almost uniform, but the edge of the upper lip spotted with
lighter. Loreal and tympanic regions darker, the edge of the supratympanic
fold lighter ; flanks with some large dark spots ; groins and axillae white (? red
in life). Lower surfaces white, the chin and throat profusely and the belly
sparsely dotted with dark brown.
Length from snout to vent : $ 25-5 mm. ; $ 33 mm.
Distribution- : Macpherson Range, 3-4000 ft., S. Queensland.
Material Examined.
B.M. 1933.4.8.6 $ Macpherson Range, 3-4000 ft. Darlington. Type.
This specimen is one of the series recorded as P. frosti by Loveridge. No
material of Spencer's species is available for comparison, but there are a number
of differences between the description of frosti and these northern frogs, which
suggest that the two are not conspecific. Both species appear to be montane
forms.
NOTADEN Gunther.
Notaden Gunther, 187^, .Inn. Mag. nat. Hist., (4), 11 : 349 (Type species: — Notaden ben-
netti) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mas., ed. 2 : 328 ; Cope, 1889, Bull. U.S.
nat. Mus., 34 : 260 ; Nieden, 1923, Das Tierreich, Anura I : 162 ; Noble, 1931, Biol.
Amph. : 498.
Maxillary teeth absent. Prevomer bordering the choana anteriorly and
sending a branch diagonally backwards towards its fellow ; this branch is
raised into a knob-like prominence at its extremity on a level with the mesial
ends of the palatines and bears a few teeth or tooth-like structures ; a large
fronto-parietal foramen ; ear fully developed, the extra-plectal cartilage large,
but not as large as the tympanum. Vertebrae procoelous with the condyle
Incompletely ankylosed and the notochord persistent ; 7 presacrals ; sacral
diapophyses moderately strongly dilated ; coccyx articulating by two condyles.
Omosternum small, cartilaginous ; sternum cartilaginous or calcified, undivided.
Terminal phalanges simple.
Distal tendon of the 111. semitendinosus passing ventral to the mm. graciles.
Alary processes of the hyoid narrow proximally, expanded distally.
Pupil horizontal. Tongue subcircular, adherent. Palate with a series of
two or three smooth transverse folds in front of the pharynx. Digits not dilated
Synopsis of the Species.
I. First linger shorter than the second ; inner metatarsal tubercle 1-2 to
2-0 times as long as its distance from the tip of the inner toe. Dorsal
markings forming a cruciform or X-shaped pattern . N.bennetti.
62 H. W. PARKER
II. First finger as long as, or longer than, the second ; inner metatarsal
tubercle as long as its distance from the tip of the inner toe. Dorsal
markings not forming a definite pattern . . . N. nichollsi.
Notaden bennetti Giinther.
Notaden bennetti Giinther, 1873, Ann. Mag. nat. Hist., (4), 11 : 350 (Type locality : — Castle-
reagh River) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 328, pi. 27, fig. 3 ;
Fletcher, 1889, Proc. Linn. Soc. N.S.W., (2), 4 : 360 ; idem, 1890, op. cit., (2), 5 : 672-
675 ; idem, 1891, op. cit., (2), 6 : 265, 271, 272 ; idem, 1892, op. cit., (2), 7 : 12 ; idem,
1894, op. cit., (2), 8 : 530, 531 ; Lucas & le Souef, 1909, Anim. Austral. : 286, fig. ;
Nieden, 1923, Das Tierreich, Anura I : 162, fig. 212 ; Loveridge, 1935, Bull. Mus.
comp. Zool. Harv., 78 : 37.
Habitus globose. Head broader than long, its width contained about
3-75 times in the length from snout to vent. Snout short and deep, about as
long as the eye, vertically truncate and not prominent ; canthus rostralis
rounded ; loreal region vertical ; nostrils subterminal ; interorbital space as
broad as, or broader than, the upper eyelid ; tympanum hidden. Fingers rather
short, depressed, the first shorter than the second, which is longer than the fourth ;
small subarticular tubercles beneath the metacarpo-phalangeal joints and indi-
cations of tubercles between the first and second and second and third fingers ;
two rather indistinct metacarpal tubercles. Toes short, depressed, with fleshy
lateral fringes and basal webbing which midway between the third and fourth
toes reaches the middle of the proximal phalanx of the third toe ; no subarticular
tubercles ; a large shovel-shaped inner metatarsal tubercle which is 1-5 to 2-0
times as long as its distance from the tip of the inner toe. Hind limb, measured
from the vent to the tip of the fourth toe with the leg extended at right angles
to the body, as long as the distance between the vent and the anterior corner of
the eye or the nostril.
Skin very thick and glandular with closely set, raised warts except for 5
patches which are almost smooth and disposed as follows : a cruciform patch
on the head, the transverse bar connecting the upper eyelids anteriorly and the
longitudinal member running backwards to the nape ; a latero-dorsal sub-
circular patch above each arm, and a similar, but larger pair above the middle
of each flank ; a linear zone, less defined than the remainder, along the coccyx.
Gular region and chest smooth ; belly feebly granular and wrinkled ; circum-
anal region with scattered small pimples.
Colour pattern following the arrangement of the warts, which are dark
brown or black often stippled with white, on a greenish background with isolated
patches of rusty- or orange red ; the smooth areas are lighter, bright or greenish-
yellow. Lower surfaces yellowish white, the throat and chest spotted with
brown.
Length from snout to vent ; $ 45 mm.
The species is cryptozoic, frequenting the dry areas of inland New South
Wales and southern Queensland ; it is myrmecophagous and seldom seen above
ground except after rain. The skin exudes a milky venom when the frog is
handled.
Distribution : Plains of New South Wales and southern Queensland
west of the dividing range.
AUSTRALASIAN LEPTODACTYLIDAE 63
Specimens Examined.
B.M.
Notaden nichollsi sp. n.
Notaden bennetti (non Giinther) Andersson, 1913, K. Svenska VetenshAhad. Hand!., 52, 4 :
20, pi. 1, figs. 7, 8.
Habitus globose. Head broader than long, its width contained 3-4 to 3-6
times in the length from snout to vent. Snout short and deep, not prominent,
obtusely rounded and truncate ; canthus rostralis rounded ; loreal region oblique ;
nostril midway between the eye and the end of the snout ; interorbital space
as broad as or a little broader than the upper eyelid ; tympanum hidden. Fingers
rather short ; in adults the first as long as or slightly longer than the second
which is longer than the fourth; in juveniles the first is shorter than the second;
small subarticular tubercles beneath the proximal joints of the fingers ; two
rather indistinct metacarpal tubercles. Toes short, depressed, with thick,
fleshy fringes and webbing which between the third and fourth reaches nearly
or quite to the level of the distal end of the proximal phalanx of the third ; no
subarticular tubercles ; a compressed inner metatarsal tubercle as long as or
very slightly longer than its distance from the tip of the inner toe in adults ;
distinctly shorter in the very young. Length of the hind-limb, measured from
the tip of the fourth toe to the vent with the leg extended at right angles to the
body, as long as the distance from vent to eye.
Skin very thick and glandular with larger and smaller warts uniformly
distributed ; sometimes there may be some larger warts forming a very indefinite
linear series dorso-laterally from the posterior corner of the eye. Lower surfaces
smooth or wrinkled ; anal region with scattered pimples.
Adults olive-grey or brown above, the larger warts brown or blackish ; there
is often no defined colour pattern but sometimes traces of a faint, interorbital
triangle, its apex directed backwards, may be discerned, and where the warts
have a linear arrangement interrupted irregular dark lines may be apparent or
groups of dark warts with a faintly cruciform arrangement. Warts on the
flanks tipped with yellow. Lower surfaces uniform yellowish white. Throat
of the male infuscate.
Juveniles at metamorphosis with a well-defined pattern. Ground-colour
silvery grey ; a curved dark-brown streak along the canthus rostralis and a
large triangular interorbital marking, the apex directed posteriorly ; a series
of three dark triangles on each side, apices directed ventrally and their bases
sub-continuous to form a dorso-lateral stripe ; the light median zone thus cut
off with two dark spots one about the middle of the back and one in front of
the sacrum. Scattered warts on the back and sides white-tipped. Limbs
silvcr-gny with dark cross bars.
Male with a vocal sac; diffuse nuptial pads on the inner surfaces <>f the first
and second fingers and a series of pustules capped with horny spinules beneath
the lower jaw.
Length from snout to vent : <£ 63 mm. ; $ 64 mm.
Juveniles at metamorphosis (tail incompletely resorbed) : 12 mm.
Distribution : Northern West Australia.
64
H. W. PARKER
Specimens Examined.
B.M. 96.7.2.19 (J
1937-7-22.36-40 3 6*d\ 2 ?9
Swedish Mus. 1569
1570
1568
(part)
$
$
6", 4 juvs.
Roebuck Bay, W.A.
Rabbit-Proof Fence No.
1 . Far North, W.A.
Noonkambah, Kimber-
ley Divn., W.A.
St. George Range, Kim-
berley Divn., W.A.
Mowla Down, Mt. Alexa-
ander, 70 miles south
of FitzRoy River,
Kimberley Divn., W.A.
Dahl. Holotype.
Nicholls. Paratypes.
Mjoberg. Paratypes.
Subfamily MYOBATRACHINAE.
Cystigxathi (part) Tschudi, 1838, Mem. Soc. neuchatel. Sci. nat., 2 : 37, 78.
Faniformes (part) Dumeril & Bibron, 1841, Erpet. Gen., 8 : 317.
Alytae (part) + Bombitatores (part) Fitzinger, 1843, Syst. Rept. : 32.
Myiobatrachidae, Myiobatrachina Bonaparte, 1850, Consp. Syst. Herp., Amph.
Myobatrachidae Schlegel, 1850, Proc. zool. Soc. Lond. : 10.
Myobatrachidae 4- Cystignathtdae (part) 4- Uperoliidae -4- Brachycephalidae (part)
4- Engystomatidae (part) Giinther, 1858, Cat. Batr. Sal. Brit. Mus. : 3, 26, 39, 45, 51.
Brachymeridae (part) 4- Bufonidae (part) 4- Scaphiopodidae (part) 4- Cystignathidae
(part) Cope, 1865, Nat. Hist. Rev., n.s., 5 : 97-120.
Cystigxathidae, Criniae (part) Cope, 1866, /. Acad. nat. Sci. Philad., (2), 6 : 67-97.
Bufonidae (part) Cope, 1866, torn. cit. : 189.
Alytidae (part) + Phryxiscidae (part) Steindachner, 1867, Reise Novara, Zool., Amph. :
7-34-
Cystigxathidae (part) + Uperoleiidae Keferstein, 1867, Nachr. Ges. Wiss. Gottingen, 18 :
343-349-
Cystigxathidae (part) + Alytidae (part) 4- Phryniscidae 4- Engystomidae (part)
Keferstein, 1868, Arch. Naturgesch. 34 : 251-273.
Phryniscidae, Phryniscina (part) 4- Engystomidae, Brevicipitina (part) 4- Alytidae,
Uperoliina (part) 4- Raxidae, Cystignathina Mivart, 1869, Proc. zool. Soc. Lond.:
288-294.
Cystignathidae (part) 4- Bufonidae (part) Boulenger, 1882, Cat. Batr. Sal. Brit. Mus.,
ed. 2 : 183-432.
Bufoxidae (part) Noble, 1922, Bull. Ainer. Mus. nat. Hist., 46 : 1-87.
Ceratophriidae (part) 4- Bufonidae (part) Waite, 1929, Rept. Amph. S. Australia : 244-
266.
Bufonidae, Criniinae (part) Noble, 1931, Biol. Amph. : 496.
Leptodactylidae (part) Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 8.
Tongue narrowly oval or small ; intermaxillary glands opening by not
more than two ducts close to the mid-line of the palate. Prevomer much reduced
or absent ; vomerine teeth vestigial or absent (text-figs. 13-15). Alary processes
of the hyoid wing-like expansions of the lateral border of the hyoid plate (text-
fig- 3. P- 5) '• sternohyoid and petrohyoid muscles attached to the ventral surface
of the hyoid, approaching the mid-line posteriorly. Distal tendon of the m.
semitendinosus perforating the mm. graciles or their distal tendon, or passing
dorsal to them. Sterno-epicoracoideus absent (Uperoleia). Presacral vertebrae
S, and the notochord always more or less persistent.
Key to the Genera.
I. Ear fully developed.
A. Maxillary teeth present.
(1) Toes webbed ; a large fronto-parietal foramen
(2) Toes free.
(a) Fronto-parietal foramen closed in the adult
(6) Fronto-parietal foramen persistent, large .
Glauertiu.
Uperoleia.
Crinia.
AUSTRALASIAN LEPTODACTYLIDAE
65
B. Maxillary teeth absent.
(i) Toes webbed ; a large fronto-parietal foramen . . Glauertia.
(2) Toes free.
(a) Fronto-parietal foramen closed in adults . . . Uperoleia.
(b) Fronto-parietal foramen large, persistent.
(i) Clavicles broad mesially ; epicoracoids meeting edge
to edge anteriorly (text-fig. 20) . . . Myobatrachus.
(ii) Clavicles not dilated ; epicoracoids overlapping . Metacrinia.
II. No tympanum, cavum tympani, annulus tympanicus, columella
(plectrum) or Eustachian tubes ; maxillary teeth absent Pseudophryne.
GLAUERTIA Loveridge.
Pseudophryne (part) Andersson, 1913, A'. Svenska VetenshAhad. Hand!., 52, 4 : 19 ; Nieden,
1923, Das Tierreich, Anura I : 150.
Glauertia Loveridge, 1933, Occ. Pap. Boston Soe. nat. Hist., 8 : 89 (Type species : — Glauertia
russelli).
Maxillary teeth present or absent. Prevomer absent ; fronto-parietal
foramen very large and persistent, the fronto-parietal bones being confined to
the lateral regions of the cranium. Ear fully developed. Vertebrae procoelous,1
but the condyle incompletely ankylosed ; sacral diapophyses moderately dilated ;
8 presacral vertebrae ; coccyx articulating by two condyles. Omosternum
rudimentary ; sternum large, cartilaginous. Terminal phalanges simple.
Distal tendon of the m. semitendinosus passing through the tendon of the
gracilis complex (G. orientates) or passing dorsal to it (G. russelli). Alary process
of the hyoid a wing-like expansion of almost the whole lateral margin of the
hyoid plate.
Pupil horizontal with a ventral angle. Tongue small, oval, \ free behind.
Tympanum hidden. Digits not dilated ; toes webbed.
The species of this genus appear to be closely allied and to be descended
from some Criniine stock. They may be distinguished from the true Crinias by
their webbed toes. This difference, in itself, seems trivial enough, but the fact
remains that these web-footed species represent one line of descent whereas the
unwebbed species, whether with or without teeth, are a separate evolutionary
branch. To include all of them in the same genus would mask their phylogeny
and this seems adequate justification for the retention of Glauertia as a distinct,
if not very easily diagnosable, genus.
Synopsis of the Species.
I. Maxillary teeth present ; a tarsal tubercle ; toes webbed at the base
G. mjobergi (Andersson).
II. Maxillary teeth absent ; no tarsal tubercle.
A. Toes half webbed ...... G. russeli Loveridge.
B. Toes \ to J webbed ...... G. orientalis sp. nov.
1 One specimen examined appears diplasiocoelous, but owing to the incomplete fusion
of the condyle this might be an artefact.
NOVIT. ZOOL., 42, I 5
66 H. W. PARKER
Glauertia mjobergi (Andersson).
Pseudophryne mjobergi Andersson, 1913, K. Svensha VetenskAkad. Handl., 52, 4 : ig, pi. 1,
figs. 5, 6 (Type locality : — Noonkambah, Kimberley Divn., W. Australia) ; Nieden,
1923, Das Tierreich, Anura I : 150, fig. 197 ; Harrison, iQ2j,Rec. Ausl. Mus., 15 : 284;
Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 9.
Head as long as broad ; snout truncate, very slightly longer than the eye ;
nostrils directed upwards, nearer the tip of the snout than the eye ; canthus
rostralis rounded ; loreal region slightly oblique ; interorbital space convex,
once and a third as wide as the upper eyelid ; tympanum hidden ; occiput swollen.
Fingers free, not dilated, the first much shorter than the second, which is a little
shorter than the fourth ; subarticular tubercles well developed ; palm with
rows of small tubercles ; two large metacarpal tubercles. Toes with a distinct
rudiment of web and fleshy lateral fringes ; fifth much shorter than the third ;
subarticular tubercles conical ; two large, shovel-shaped metatarsal tubercles,
the outer obliquely transverse ; a conical papilla on the lower surface of the
tarsus, close to the tibio-tarsal articulation. Tarso-metatarsal articulation
reaching the posterior corner of the eye.
Skin with scattered small warts above ; a large parotoid gland ; a similar
gland along the flanks to the groin and a smaller gland on each side of the coccyx.
Smooth beneath, except the hinder sides of the thighs which are granular.
Pale grey above with a few large insuliform spots of which the most prominent
are a kidney-shaped pair bordering the parotoid glands above ; the small warts
light-tipped. Lower surfaces white.
Length from snout to vent : 20 mm.
Distribution : Known only from the type locality, Noonkambah, Kimberley
Divn., W. Australia.
Specimen Examined.
Swedish Mus. 1567 $ Noonkambah Mjoberg. Cotype.
Glauertia russelli Loveridge.
Glauertia russelli Loveridge, 1933, Occ. Pap. Boston Soc. nat. Hist., 8 : 89 (Type locality : —
Gascoyne River, near Landor Station, W. Australia) ; idem, 1935, Bull. Mus. comp.
Zool. Harv., 78 : 37, pi. 1, figs. 1-3.
Head broader than long ; snout triangular, truncate distally, a little longer
than the diameter of the eye ; nostrils directed upwards, nearer the tip of the
snout than the eye ; canthus rostralis rounded ; loreal region nearly vertical,
slightly concave ; interorbital space flat, once and a half to twice as broad as
the upper eyelid ; tympanum hidden. Fingers free, not dilated ; first much
shorter than the second which does not extend as far as the fourth ; subarticular
tubercles well developed ; palm with rows of small tubercles ; two metacarpal
tubercles. Toes half webbed, the membrane midway between the third and
fourth reaching the level of the distal subarticular tubercle of the third ; fifth
much shorter than the third ; subarticular tubercles well developed ; two large
shovel-shaped metatarsal tubercles, the outer transversely disposed. Tarso-
metatarsal articulation reaching the anterior corner of the eye in juveniles or
the axilla in adults.
Skin warty above ; more or less granular on the abdomen and beneath
the thighs.
Brown above with indistinct darker spots and with or without a light
(pink in life) vertebral stripe ; a light blotch on each scapular region (reddish
AUSTRALASIAN LEPTODACTYLIDAE 67
orange in life) and a similar but smaller spot may be present on each side of the
coccyx ; the two may be more or less connected by an indefinite zone of the
same colour. Dirty yellow beneath, the throat sometimes freckled with brown.
Length : <$ 23-5 mm. ; $ 30 mm.
Distribution : Known only from the type locality.
Specimens Examined.
Stellenbosch Univ. 2 juvs. Landor Sta., W. Austr. Glauert. ParaTYPES.
Colin.
B.M. 1937.7.23.3-5 $?, Hgr.
Glauertia orientalis sp. nov.
Pseudophryne fimbrianus ? Parker, 1926, Ann. Mag. nat. Hist., (9), 17 : 670 (Groote Eylandt).
Holotype a male, number 1908.2.25.34 in the British Museum from
Alexandria Station (190 08' S., 1360 43' E.), Northern Territories, Australia,
collected by W. Stalker.
Head a little broader than long; snout short, vertically truncate, shorter
than the eye ; nostrils directed vertically upwards, close to the tip of the snout ;
canthus rostralis rounded ; loreal region slightly oblique ; interorbital space
once and a third the width of the upper eyelid ; tympanum hidden. Fingers
free, the first shorter than the second, which does not extend quite as far as the
fourth ; subarticular tubercles well developed ; palm with rows of small, round
tubercles ; two metacarpal tubercles. Toes a quarter webbed, the edge of the
membrane midway between the third and fourth not extending beyond the proximal
tubercle of the fourth toe ; edge of the membrane raised into a tubercle ; fifth
toe much shorter than the third. Two large, compressed metatarsal tubercles,
the outer disposed transversely. Tarso-metatarsal articulation reaching the
tympanic region.
Skin slightly warty above ; a moderately distinct parotoid gland and a
smaller, circular inguinal gland. Lower surfaces feebly granular except beneath
the thighs where the granulations are more pronounced.
Brown above, each wart lighter and ringed with darker ; faint indications
of a light, mid-dorsal line ; parotoid and inguinal glands rufous. Lower
surfaces dirty white, the gular region infuscate. Hinder side of thighs brown
with a light (non-glandular) spot behind the knee.
A vocal sac opening internally by a slit on each side close to the lower jaw ;
base of the first finger on its inner side thickened and glandular.
Length from snout to vent : 26 mm.
Hind limb : 29 mm.
Paratypes : Two males from the type locality, and a female from Groote
Eylandt.
These examples show little variation. The parotoid and inguinal glands
are not always clearly defined and the digital webbing of the toes is not absolutely
constant, varying from ] to .', ; in none of them is it as extensive as in (',. russeUi.
The female has a deeply fimbriated cloacal flap, as in the genus Uperoleia.
These eastern specimens appear to differ constantly from the western (i.
russeUi in their shorter digital webbing and slightly shorter more acuminate
snouts ; but it must be admitted that with reasonably long series, these differences
may not be found to hold good and that the two forms may not be tenable.
68
H. W. PARKER
Specimens Examined.
Alexandria Sta., N.
Territory.
Alexandria Sta., N.
Territory.
Groote Eylandt.
Stalker. Holotype.
,, Paratypes.
Wilkins. Paratype.
UPEROLEIA Gray.
Uperoleia1 Gray, 1841 (April), Ann. Mag. nat. Hist., 7 : 90 (Type species : — Uperoleia mar-
morata) ; Gray, 1841, in Grey, Journ. Exped. Austral., 2 : 448; Giinther, 1858, Cat.
Batr. Sal. Brit. Mus. : 39 ; Steindachner, 1867, Reise Novara, Zool., Amph. : 33 ; Kefer-
stein, 1867, Nachr. Ges. Wiss. Gbttingen, 18 : 349 ; Giinther, 1867, Ann. Mag. nat. Hist.,
(3), 20 : 55 ; Keferstein, 1868, Arch. Naturgesch., 34 : 270.
Uperoleja Gray, 1841, torn. cit. : 436.
Hyperolia1 (emend.) Cope, 1865, Nat. Hist. Rev., n.s., 5 : 108 ; idem, 1866, /. Acad. nat.
Sci. Philad., (2), 6 : 94 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 267 ; Cope,
1889, Bull. U.S. nat. Mus., 34 : 312 ; Nieden, 1923, Das Tierreich, Anura I : 535 ; Noble,
1931, Biol. Amph. : 498.
Pseudophryne (part) Andersson, 1916, K. Svenska VetenskAkad. Hand!., 52, 9 : 13 ; Parker,
1926, Ann. Mag. nat. Hist., (9), 17 : 669.
Maxillary teeth present or absent. Prevomer reduced to two fragments,
one bounding the choana and the other overlying the mesial end of the palatine
(U. rugosa, text-fig. 13) or completely absent (U. mannorata) ; fronto-parietal
Text-fig. 13. — Anterior cranial elements of Uperoleia rugosa. x 6-5. (From beneath.)
foramen present in juveniles, but completely roofed over in the adult. Ear
fully developed. Vertebrae procoelous, but the condyle incompletely ankylosed ;
eight pre-sacrals ; sacral diapophyses moderately dilated ; coccyx articulating
by two condyles. Omosternum rudimentary ; sternum small, cartilaginous.
Terminal phalanges simple. Distal tendon of the m. semitendinosus perforating
the gracilis complex.
Alary process of the hyoid a wing-like lateral expansion of the whole of
the border of the hyoid plate.
Pupil rhomboidal, often somewhat vertically compressed. Tongue oval,
entire and free behind. Tympanum hidden. Parotoid glands large and
prominent. Digits free, not dilated.
The relationships of this genus are discussed below, under Pseudophryne.
1 Under Art. 19 of the Rules of Nomenclature the original orthography of a name may
be emended when, inter alia, an error of transliteration " is evident." What is " evident "
is not capable of rigid definition, so that it seems advisable to consider names as arbitrary
combinations of letters except when their derivation is expressly stated in the original publi-
cation of the name. Gray does not state the etymological derivation of Uperoleia and so,
although Cope's emendation to Hyperolia is probably based on a perfectly correct assump-
tion, there is no sure proof. Consequently it seems preferable to continue the use of the
name as it was originally proposed, the more so since this course will not involve any change
of the name Hyperolius Rapp, 1842.
AUSTRALASIAN LEPTODACTYLIDAK 69
Despite Loveridge's (1935 : 31) contention that ntgosa Anderson is at most
racially distinct from marmorata, there appear to be sufficient well-marked diffe-
rences to indicate that they are truly specifically distinct. Harrison (1927 :
284) suggested " with some diffidence " that Pseudophryne mjobergi Andersson
might be a synonym of U. marmorata, but the species appears to be referable
to the genus Glauertia.
Synopsis of the Species.
I. Maxillary teeth present ; metatarsal tubercles not compressed ; snout
rounded [/. marmorata.
II. Maxillary teeth absent ; metatarsal tubercles distinctly shovel-like, the
outer disposed transversely ; snout truncate . . . . U. rugosa.
Uperoleia marmorata Gray.
Uperoleia marmorata Gray, 1841 (April), Ann. Mag. nat. Hist., 7 : 90 (Type locality: —
Western Australia); Gray, 1841, in Grey, Joum. Exp. Austral., 2:448; Giinther,
1858, Cat. Batr. Sal. Brit. Mus. : 39; Krefft, 1865, Pap. rov. Soc. Tasmania : 17 ;'
idem, 1867, Cat. Industr. Prod. N.S.W., Add. : 107 ; Steindachner, 1867, Reise Novara,
Zool., Amph. : ^ ; Giinther, 1867, Ann. Mag. nat. Hist., (3), 20 : 55 ; Keferstein, 1867,
Nachr. Ges. Wiss. Gbttingen, 18 : 349; idem, 1868, Arch. Naturgesch., 34 : 270, pi. 6,
fig. 14 ; Harrison, 1922, Aastr. Zool., 3, 1 : 31.
Hyperolius {Uperoleia) marmoratus Parker, 1881, Phil. Trans, roy. Soc. Land., 3 : 134, pi. 24,
figs. 6, 7.
Hyperolia marmorata Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 267; Fletcher,
1889, Proc. Linn. Soc. N.S.W., (2), 4:376; idem, 1890, op. cit., (2), 5:672-675;
idem, 1891, op. cit., (2), 6 : 272 ; idem, 1892, op. cit., (2), 7 : 7-9; idem, 1894, op. cit.,
(2), 8 : 526, 528, 530 ; Boettger, 1894, Denkschr. med.-naturw. Ges. Jena, 8 : 109 ;
Fletcher, 1898, Proc. Linn. Soc. N.S.W., 22 : 678; Lucas & le Souef, 1909, Anim.
Austral. : 277 ; Nieden, 1923, Das Tierreich, Anura I : 535, fig. 368.
Uperoleia marmorata var. laevigata Keferstein, 1867, Nachr. Ges. Wiss. Gbttingen, 18 : 349
(Type locality : — Raudewick, N.S.W.).
Snout rounded, a little longer than the eye ; canthus rostralis quite indistinct ;
loreal region oblique, not concave ; nostril a little nearer the tip of the snout
than the eye; the distance from nostril to tip of snout contained 13 to 15 times
in distance from nostril to eye ; interorbital space once and a quarter to once
and a half as broad as the upper eyelid. Fingers moderate, with prominent
subarticular tubercles; the first much shorter than the second, which is a little
shorter than the fourth ; carpus with rows of small tubercles ; two well-developed
metacarpal tubercles. Toes with indications of fleshy, lateral fringes and
prominent subarticular tubercles ; third much longer than the fifth ; two large,
but not compressed, metatarsal tubercles. Tarso-metatarsal articulation reaching
the eye.
Upper surfaces smooth or minutely tubercular; a large parotoid gland;
lower surfaces smooth or feebly granular ; hinder side of thighs granular ; anal
flap undivided in males, fimbriated in females.
Pale brown to dark olive above, with or without darker spot-, and marblings,
of which the most constant are an interorbital blotch and a curved stripe on
the upper borders of the parotoids. A large yellow spot in the groin and another
behind the thigh close to the knee ; these areas do not appear to be glandular.
Lower surfaces pale brown to dark olive, minutely Stippled with lighter or pale
brown mottled with darker; chin of the male infuscate
Male with a vocal sac opening by a slit on each side of the tongue and a
glandular thickening of the base and inner side of the first fingei
70
H. W. PARKER
Length from snout to vent : $ 28 mm. : $ 31 mm.
As in Pseudophryne bibroni, examples from the coastal region of New South
Wales are very dark in colour and may be recognizable as a distinct race.
Habits cryptozooic. Breeding in early spring (N.S.W.) ; eggs laid singly,
over 200 in a clutch. Call note an explosive " akh " (Fletcher, 1889 : 376,
and Harrison, 1922 : 31).
Distribution : Australia.
B.M. 41 (2) 215
64.7.9.14
87.7
87.7
87.7
88.7
1915
1926
16. 1
16.2
16.3
3-9-23
2. 16. 1
1932. 10. 2.147-
148
Mus. Leiden 4259
Austr. Mus. R. 7981
R- 7379
(part)
<$
?
6"
<J (cleared)
c?
?
?
$
$9
$
$
Specimens Examined.
W. Australia.
Sydney.
Burrawang, N.S.W.
Parramatta.
Huskisson, Jervis Bay,
N.S.W.
W. Australia.
Queensland.
Upper Colo, via Rich-
mond, N.S.W.
Burrawang, N.S.W.
Type.
Giinther.
Fletcher.
Schrader.
Rodway.
Reischek.
(Mus. Godeflfroy.)
Uperoleia rugosa (Andersson).
Hyperolia marmorata (part) Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 267 (Cape
York) ; Fletcher, 1890, Proc. Linn. Soc. N.S.W., (2), 5 : 673 (Dandaloo).
Pseudophryne rugosa Andersson, 1916, K. Svenska VetenskAkad. Handl., 52, 9 : 13, pi. 1,
fig. 4 (Type locality: — Colosseum, Queensland).
Uperoleia marmorata rugosa Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 31.
Pseudophryne fimbrianus Parker, 1926, Ann. Mag. nat. Hist., (9), 17 : 669, fig. 3 (Type
locality : — St. George District, Queensland).
Snout vertically truncate, a little longer than the diameter of the eye ; canthus
rostralis rounded ; loreal region nearly vertical, not concave ; distance from
the nostril to the tip of the snout contained nearly twice in the distance from
nostril to eye ; interorbital space once and a third as broad as the upper eyelid.
Fingers depressed, the first much shorter than the second, which is a little shorter
than the fourth ; subarticular and metacarpal tubercles well developed ; carpus
with rows of smaller tubercles. Toes with feeble indications of fleshy lateral
fringes, especially proximally, and prominent subarticular tubercles ; third
much longer than the fifth ; two large, metatarsal tubercles, the outer the larger,
usually distinctly transverse and shovel-like. Tarso-metatarsal articulation
reaching the tympanic region.
Upper surfaces regularly warty. A distinct parotoid gland and a more or
less distinct lateral gland extending from the posterior corner of the latter towards
the groin and a circular gland on each side of the coccyx posteriorly. Lower
surfaces feebly granular ; hinder side of thighs distinctly granular. Anal flap
fimbriated in females.
Brown above with more or less profuse darker spots of which the most
constant is a V-shaped interorbital bar. Posterior half of the parotoids often
with a light spot. A distinct white patch in the groin and another on the thigh
behind the knee ; these areas are not glandular. Lower surfaces white, dusted
or spotted with brown ; chin of male infuscate.
AUSTRALASIAN LEPTODACTYLIDAE
71
Male with a vocal sac opening by a slit on each side of the tongue, and with
a thickened glandular pad on the inner side of the first finger.
Length from snout to vent : $ 26 mm. : ? 28 mm.
Distribution : Queensland and New South Wales west of the dividing
range.
Specimens Examined.
CRINIA Tschudi.
Crinia Tschudi, 1883, Mem. Soc. neuchdlel. Sci. nat. ; 38, 78 (Type species : — Crinia geor-
giana) ; Fitzinger, 1843, Sysl. Rept. : 32 ; Girard, 1853, Proc. Acad. nat. Sci. Philad.,
6 : 420 ; Cope, 1865, Nat. Hist. Rev., n.s., 5 : 107 ; idem, 1866, /. Acad. nat. Sci.
Philad., (2), 6 : 05 ; Keferstein, 1867, Nachr. Ges. Wiss. Gbttingen, 18 : 347 ; idem, 1868,
Arch. Naturgesch., 34 : 262 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus.,ed. 2 : 263 ; Cope,
1889, Bull. U.S. nat. Mus., 34 : 312 ; Werner, 1914, Fauna S.W. Austral., 4 : 407 ;
Nieden, 1923, Das Tierreich, Anura I : 539 ; Waite, 1929, Rept. Amph. S. Austral. :
257; Noble, 1931, Biol. Amph. : 498.
Cystignathus (part) Dumeril & Bibron, 1841, Erpit. Gen., 8, 392 ; Gunther, 1858, Cat. Batr.
Sal. Brit. Mus. : 26.
Ranidella Girard, 1853, Proc. Acad. nat. Sci. Philad., 6:421 (Type species: — Ranidella
signifera) ; idem, 1858, U.S. Explor. Exped., Herp. : 44.
Pterophrynus Liitken, 1862, Vidensk. Medd. Naturh. Foren. Kbh. : 302 (Type species: —
Pterophrynus verrucosus) ; Steindachner, 1867, Reise Novara, Zool., Amph. : 30.
Pterophryne Gunther, 1867, Ann. Mag. nat. Hist., (3), 20 : 53.
Camariolius Peters, 1863, Mber. Ahad. Berlin : 236 (Type species : — Camariolius varius).
Maxillary teeth present. Prevomer variable. In its most primitive con-
dition (leai, rosea, laevis and darlingtoni) a small anterior portion, bounding the
choana antero-mcsially, is connected by a narrow strip with a small posterior
portion overlying the mesial end of the palatine which often bears vomerine
teeth. In acntirostris only the anterior portion bounding the choana persists,
but in georgiana, signifera, tasmaniensis and glauerti both the anterior portion
and a small fragment overlying the palatine mesially are present, but unconnected ;
the posterior portion may, or may not bear teeth. A very large fronto-panetal
foramen. Ear fully developed. Vertebrae procoelous, the condyle incompletely
ankylosed ; notochord persistent ; sacral diapophyses slightly dilated ; urostyle
articulating by two condyles ; eight presacral vertebrae. Omostrrnum small,
cartilaginous, not fully differentiated from the procoracoids (text -figs, id and 18) ;
sternum undivided, cartilaginous ; procoracoids broad. Terminal phalanges
simple.
Distal tendon of the in. semitendinosus passing between the m, gnu ilis major
and minor.
Alary process of the hyoid a wing-like expansion of almost the whole lateral
margin of the hyoid plate ; cricoid cartilage incomplete : oesophageal process of
the cricoid broad or almost absent ; m. omohyoideus absent ; mm. sternohyoi-
deus and petrohyoideus anterior inserted on the ventral surface of the hyoid,
reaching the middle line in the posterior part of this insertion.
72
H. W. PARKER
Pupil horizontal but with a downwardly directed angle ventrally. Tongue
narrowly oval, partly free behind. Toes free, not or but feebly dilated distally.
Text-fig
Text-fig
Text -fig. 16.
Text-fig. 14. — Anterior cranial elements of Crinia laevis laevis. x 6-5. (From beneath.)
Text-fig. 15. — Anterior cranial elements of Crinia tasmaniensis. X 6-5. (From beneath.)
Text-fig. 16. — Ventral elements of the shoulder girdle of Crinia hasivelli. (Cotype $.)
Synopsis of the Species.
I. Prevomer undivided or with its anterior and posterior portions in
contact. Vomerine teeth usually present.
A. Belly, and usually the upper surfaces also, quite smooth. Toes not
fringed.
(1) Inner finger and toe not reduced, the latter § the length of the
second.
(a) Toes with distinct subarticular tubercles and slight terminal
dilatations ........ C. leai.
(b) Toes without or with very indistinct subarticular tubercles
and not dilated terminally ..... C. rosea.
(2) Inner finger and toe much reduced, the latter not more than half
the length of the second and the inner finger with, at most, a
single very short phalanx.
(a) Toes pointed ; groins and hinder side of the legs marked with
red and brown,
(i) Belly with a few large blotches . . . C. laevis laevis.
(ii) Belly regularly dusted with brown . C. laevis victoriana.
(b) Toes slightly dilated terminally ; no red on the groins or limbs
C. darlingtoni.
B. Belly more or less granular. Toes fringed ... C. haswelli.
AUSTRALASIAN LEPTODACTYLIDAE 73
II. Prevomer divided and the two halves widely separated or the posterior
absent. Vomerine teeth reduced or absent.
A. Vomerine teeth usually present ; toes not fringed ; belly granular.
Hinder side of thighs red in alcohol.
(1) Snout short, rounded, 1-14 to 1-3 times the length of the eye.
Average size of adults, <$$ 26, ?? 27-3 mm. . C. georgiana.
(2) Snout longer, pointed, prominent, 1-35 times the length of the
eye. Average size of adults 16-8 mm. . . . C. glauerti.
B. Adults with more or less distinct dermal fringes on the toes. Vomerine
teeth vestigial or absent.
(1) Belly smooth or only slightly granular.
(a) Two metatarsal tubercles ; loreal region very oblique ; belly
blotched with pink C. tasmaniensis.
(b) A single, inner, metatarsal tubercle ; loreal region nearly
vertical ....... C. acutirostris.
(2) Belly granular, a tarsal fold, belly not pink.
(a) Inner metatarsal tubercle £ its distance from the tip of the
inner toe. Nostril equidistant from the eye and the end
of the snout C. signifera signifera.
(b) Inner metatarsal tubercle about half its distance from the tip
of the inner toe. Nostril usually a little nearer the eye
than the end of the snout,
(i) Digital fringes well-developed, thin. Tibio-tarsal articu-
lation reaching the tympanic region. Snout somewhat
pointed and slightly prominent . C. signifera englishi.
(ii) Digital fringes short and fleshy. Tibio-tarsal articulation
scarcely reaching beyond the shoulder. Snout bluntly
rounded, not prominent . . . C. signifera Montana.
Crinia leai Fletcher.
Crinia leai Fletcher, 1898, Proc. Linn. Soc. N.S.W., 22, 1897 : 677 (Tvpe localities :— Bridge-
town and Pipe Clay Creek, near Jarrahdale) ; Fry, 1914, Rec. W. Aust. Mas.. I : 203,
pi. 28, figs. 2, 2a; Nieden, 1923, Das Tierreich, Anura I : 543; Harrison, 1027, Rec.
Aust. Mus., 15, 4 : 277 ; Loveridge, 1935, />'"// Mus. ""up- Zool. Harv., 78 : jo.
< rinia michaelseni Werner, 1914, in Michaelsen & Hartmeyer, Fauna SAW Australiens, 4 :
416 (Type locality: — Donnybrook, W.A.) : Nieden, 1923, Inc. cit.
Vomerine teeth usually present in transverse series behind the level of the
choanae. Snout depressed, rounded, not prominent, 1-3 times as long as the eye ;
canthus rostralis rounded ; loreal region very oblique ; nostril midway between
the eye and the tip of the snout ; interorbitaJ spare once and a quarter to once
and a half as wide as the tipper eyelid ; tympanum hidden. Fingers moderately
long with subarticular tubercles ; palm with a few indistinct granules ; two
metacarpal tubercles. Toes long, distinctly spatulate distallv, without dermal
fringes' ; subarticular tubercles distinct but not prominent ; a single, inner
metatarsal tubercle ; no tarsal fold. I tbio tarsal articulation reaching the eye
Skin smooth above and below; a di^tinrt parotoid glandular thickening;
hinder side of the thighs, beneath the vent, granular.
1 Fletcher speaks of a " tendency to fringed toes " ; Fry of a " distinct fringe or devoid
of a fringe," and Wernei says " Zehen gesaumt." The specimens examined have no trace
of fringes, but in some of the cotypes which are somewhat shrivelled there are lateral seams
on the toes.
74 H. W. PARKER
Pinkish grey or brown above, with, in young specimens, a large rectangular,
dark spot commencing between the eyes and extending backwards almost to the
hind limbs. This marking may have obscure mottlings within it and is not
generally completely persistent in adults. Usually with increasing age a process
of emargination commences posteriorly and spreads forwards ; the lateral borders
are often more or less persistent, sometimes as lines, sometimes as rows of spots,
and in extreme cases only a triangular dark interorbital spot persists ; synciput
grey ; upper lip with one or two dark bars radiating from the eye ; a dark stripe
from the nostril through the eye, interrupted on the parotoid region and sometimes
continued on the anterior part of the flanks. Limbs cross-barred ; a triangular
dark spot enclosing the vent, not clearly outlined below ; tarsus and metatarsus
blackish towards their outer edges beneath. Lower surfaces white, more or less
brown freckled.
Male with a vocal sac opening by a slit on each side of the tongue and a
glandular, non-rugose nuptial pad.
Length from snout to vent : <J 19 mm. ; 9 25 mm.
Distribution : Coastal belt of S.W. Australia from Albany northwards
almost to Perth.
Cryptozooic in habits and frequenting damp areas.
Specimens Examined.
M.C.Z. 18465-75 <j> Augusta, W. Australia. Brooks.
(part)
Austr. Mus. R.8337 2 <J<J, 2 juvs. Bridgetown, W. Australia. Cotypes.
,, R. 10324 (J, 2 59, 2 juvs. Pipe Clay Creek,
,, ,, R. 10978 $, juv. Augusta, ,, Brooks.
,, ,, R. 10959 §, juv. Near the mouth of the
Denmark River, W.A.
Crinia rosea Harrison.
Crinia rosea Harrison, 1927, Rec. Aust. Mus., 15 : 279 (Type locality : — Pemberton, in the
Karri Country, 218 miles south of Perth) ; Loveridge, 1935, Bull. Mus. comp. Zool.
Haw., 78 : 30.
Vomerine teeth strongly developed. Snout slightly acuminate, not promi-
nent, depressed, once and a quarter as long as the eye ; canthus rostralis rounded ;
loreal region very oblique ; nostril equidistant from the tip of the snout and the
eye, or a little nearer the latter ; interorbital space once and a half the width of
the upper eyelid ; tympanum hidden. Fingers short, pointed, with scarcely
perceptible subarticular tubercles ; palm smooth ; metacarpal tubercles faintly
indicated only. Toes moderate, pointed, not fringed, with scarcely a trace of
subarticular tubercles ; a very indistinct inner metatarsal tubercle ; no tarsal
fold ; tibio-tarsal articulation reaching the shoulder.
Skin smooth above and below ; no sub-anal granular patch ; glandular
areas behind the angle of the mouth, in the parotoid region and behind the thighs ;
a slight fold across the chest.
Dark purplish grey above, uniform or obscurely mottled with darker, or with
a median dark zone commencing between the eyes, with parallel sides running
backwards to the pelvic region, more or less deeply emarginate behind. Limbs
cross-barred. Lower surfaces whitish (red in life) with some brown freckling.
Gular region of male blackish.
Length from snout to vent : <J 22 mm. ; $ 25 mm.
AUSTRALASIAN LEPTODACTYLIDAE
75
Distribution : Pemberton, W.A.
A cryptozooic frog found under logs in wet forest country ; possibly dis-
tributed throughout the hard-wood forests of S.W. Australia.
M.C.Z. 18419
Specimen Examined.
Pemberton, W.A.
Darlington.
Crinia laevis laevis (Giinther).
Pterophrynus laevis Giinther, 1864, Proc. zool. Soc. Lond. : 48, pi. 7, fig. 4 (Type locality : —
Tasmania) ; idem, 1864, Ann. Mag. nat. Hist., (3), 14 : 314 ; Krefft, 1865, Pap. roy.
Soc. Tasmania : 16.
Crinia laevis Giinther, 1868, Proc. zool. Soc. Lond. : 480; Keferstein, 1868, Arch.
Naturgesch., 34 : 265 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 266 ; Lucas
& le Souef, 1909, Anim. Austral. : lid ; English, 1910, Proc. zool. Soc. Lond. : 630,
pi. 51, fig. 6; Harrison, 1922, Aust. Zool., 3, 1 : 31 ; Nieden, 1923, Das Tierreich,
Anura I : 542 (part) ; Blanchard, 1929, Aust. Zool., 5, 4 : 327, pi. 35 ; Loveridge,
1934, Pap. roy. Soc. Tasmania, 1933 : 60 ; idem, 1935, Bull. Mus. comp. Zool. Harv.,
78 : 29.
Vomerine teeth present or vestigial. Snout rounded, depressed, not promi-
nent, 1-4 times as long as the eye ; canthus rostralis rounded ; loreal region very
oblique ; nostril a little nearer the eye than the tip of the snout ; interorbital
space once and three quarters as broad as the upper eyelid ; tympanum hidden.
Fingers short, with scarcely perceptible subarticular tubercles ; first finger very
short, with the phalanges reduced to a single, small, cartilaginous or bony nodule
Text-fig. 17.
Text-fig. 18.
Text-fig. 17. — Hand of Crinia laevis laevis. x 9.
Text-fig. 18. — Ventral elements of the shoulder girdle of Crinia laevis victorianus.
at the extremity of the metacarpal ; two fiat, indistinct metacarpal tubercles ;
palm smooth. Toes short, not fringed, the inner short, but variable in length,
with one or two (type) phalanges ; subarticular tubercles scarcely indicated ; a
very indistinct inner metatarsal tubercle, but no outer, and no tarsal fold. Tibio-
tarsal articulation reaching the vicinity of the shoulder.
Skin smooth above or with a few scattered tubercles ; smooth below ; a
parotoid glandular thickening.
Brown or slate-grey above with small, irregular, scattered, red spots more or
76 H. W. PARKER
less bordered by black. Belly and throat with moderately large dark brown spots,
the ground-colour white, suffused with bluish anteriorly and pink posteriorly.
Hind limbs marbled with black or brown and pink ; a deep pink spot in the
axilla and groin, outlined or marbled with black. Throat of the male less spotted
than that of the female and coloured bright yellow.
Male with a vocal sac opening by a slit on each side of the tongue.
Length from snout to vent : c? 31 mm. ; $ 33 mm.
The species is nocturnal and cryptozooic, frequenting damp localities. Breed-
ing in late summer (March). The song is a short, rasping note repeated several
times in a few seconds. Eggs to the number of about 120 in a clutch are laid in
cavities underground in low-lying places, or attached to the stems of grasses
and rushes a little above ground level, but always in spots liable to be flooded
during rain. The developmental period within the egg is probably long (at least
five weeks), and the tadpole is hatched in an advanced stage of development,
without any trace of external gills (English and Blanchard).
Distribution : Tasmania up to 3000 ft.
Specimens Examined.
Crinia laevis victoriana Boulenger.
Criniavictoriana Boulenger, 1888, Ann. Mag. nat. Hist., (6), 2 : 142 (Type locality :— Warra-
gul, Gippsland, Victoria) ; Lucas & le Souef, 1909, Anim. Austral. : 276 ; Nieden, 1923,
Das Tierreich, Anura I : 543.
Crinia froggatti Fletcher, 1891, Proc. Linn. Soc. N.S.W., (2), 6 : 274, 275 (Type localities :—
Buninyong and Gong Gong, near Ballarat) ; idem, 1894, op. cit., (2), 8 : 523 ; Lucas &
le Souef, 1909, Anim. Austral. : 276 ; Blanchard, 1929, Aust. Zool., 5, 4 : 328.
Crinia laevis froggatti Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 29.
? C(rinia) sp. Lucas, 1892, Proc. roy. Soc. Victoria, 4 : 61. _
Crinia laevis (part) Nieden, 1923, Das Tierreich, Anura I : 542.
This race is very similar to the typical form in morphological characters and
colour, but has a slightly longer hind limb, the tibio-tarsal articulation reaching
nearly to the angle of the mouth, and the ventral surfaces are beset with smaller
spots and some degree of dark stippling, giving a generally more dusky appear-
ance. Frequently there is a dark triangular, interorbital spot, apex directed
posteriorly, which may, or may not, be continuous with two dark, sinuous streaks
which run backwards on each side of the middle line towards the groins. The
voice of the male is also different, consisting of a series of low rasping notes
followed by a series of high-pitched " pips ", thus— crrrack, crrrack, crrrack,
pip, pip, pip, pip-pip-pip-pip-pip-pip-pip. (Blanchard.)
Distribution : Victoria, south of the Dividing Range.
AUSTRALASIAN LEPTODACTYLIDAE JJ
R.8331-6 ) 5 £<£, 18$$, ' .., , ... .
R.10332-4 1 11 juvs 1 Wairagul, Victona.
Crinia darlingtoni Loveridge.
Crinia darlingtoni Loveridge, 1933, Occ. Pap. Boston Soc. nat. Hist., 8 : 57 (Type locality : —
3000-4000 ft., Queensland National Park, MacPherson Range, Queensland) ; idem,
1935, Bull. Mus. comp. Zool. Harv., 78 : 29.
Vomerine teeth absent.1 Snout obtusely rounded, not prominent, 1-3 times
as long as the eye ; canthus rostralis rounded ; loreal region moderately oblique ;
nostril equidistant from the eye and the tip of the snout ; interorbital space
1-5 times as broad as the upper eyelid ; tympanum hidden. Fingers very short,
the first reduced to a mere stump (without phalanges) ; the remainder with
distinct subarticular tubercles ; palm smooth ; two large metacarpal tubercles.
Toes short, not fringed, the first rudimentary ; remainder slightly dilated ter-
minally, with indistinct subarticular tubercles ; a small inner, and sometimes
traces of an outer, metatarsal tubercle. Tibio-tarsal articulation reaching the
shoulder in adults, the eye in juveniles.
Skin perfectly smooth above and below.
Brown or pinkish brown above with traces of a Microhylid type of colour
pattern consisting of a dark median band commencing between the upper eyelids
and connecting them, narrowing in the scapular region, broadening in the sacral
region, narrowing and then finally bifurcating, the two branches running towards
the groins where they terminate in darker spots. A vertical dark bar beneath
the eye. A lateral dark band from the nostril, through the eye, above the arm
and along the flanks. Hind limbs with a single diagonal cross-bar each on the
femur, tibia, and tarsus ; a dark spot on the knee and a triangular black spot
enclosing the vent. Some or all of these markings may become obscure and the
dorsal and lateral bands are frequently broken up, the posterior portions being
lost before the anterior. Lower surfaces heavily mottled and stippled with
brown, except on the belly and thighs.
Length from snout to vent : $ 19 mm.
Distribution : 3000-4000 ft., Queensland National Park, MacPherson
Range.
Specimen Examined,
M.C.Z. 18392 juv. Type locality. Darlington. Paratype.
Crinia haswelli Fletcher.
Crinia haswelli Fletcher, 1894, Proc. Linn. Soc. N.S.W., 1893, (2), 8 : 522, 526, 530 (Type
locality : — near head of Jervis Bay) ; Nieden, 1923, Das Tierreich, Anura I : 540.
Epicoracoids very strongly overlapping and extending forwards beyond the
1 The original description states " mandible toothed," but this is presumed to be a
slip, " maxilla " being intended.
78 H. W. PARKER
clavicles (text-fig. 16). Vomer entire ; vomerine teeth present, in short trans-
verse series well behind the level of the internal nares. Snout rounded, i-a to
1-5 times as long as the eye, shorter in juveniles than adults, with rounded
canthus rostralis and very oblique, convex, loreal region ; nostril equidistant from
the eye and the end of the snout ; tympanum hidden or just perceptible ; inter-
orbital space i-i to 1-2 times as broad as the upper eyelid in females, 1-2 to 1-4
times in males. Fingers slender, free, with well-developed subarticular tubercles ;
palm smooth ; two metacarpal tubercles. Toes slender, fringed ; subarticular
tubercles well developed ; a small inner metatarsal tubercle, but the outer minute
or absent ; no tarsal fold. Tibio-tarsal articulation reaching the tympanic
region.
Skin smooth or with scattered tubercles above ; a fold from the posterior
corner of the eye above the arm to the middle of the flanks, with a well-marked
parotoid gland below it ; a circular glandular spot on the flank just in front of
the groin. Belly slightly granular, hinder side of the thighs more strongly so ;
a small pectoral spot close to the insertion of the fore-limb, and a minute papilla
on the heel.
Colour in spirit, pale brown, with scattered small darker spots. A dark
brown band from the nostril through the eye and below the supra-tympanic
fold to the middle of the flanks. Lower surfaces uniform dirty white in juveniles,
pale brown in adults with the abdomen covered with large, circular white spots.
Groins and concealed surfaces of the limbs dark brown with large white spots.
A fine, light, mid-dorsal line may be present. In life, the ground-colour dorsally
is chestnut brown, drab, or silver-grey, with an incomplete darker band down the
middle, commencing between the eyes ; the light spots of the groin and concealed
surfaces of the limbs carmine or orange red.
Male with a vocal sac opening by a slit on each side of the tongue and glan-
dular, non-spinous nuptial pads at the base of the two inner fingers and on the
inner surface of the forearm.
Length from snout to vent : <J 26 mm. ; $ 32 mm.
Distribution : Coastal districts of New South Wales.
Specimens Examined.
Austr. Mus. R. 10335 2 $$, $, Hgr. Jervis Bay. Cotypes.
R.4210, cJ, $, 8 juvs. Maroubra, near Sydney.
5288, 6132-5
This species is in many respects intermediate between the smooth-skinned
species of the laevis group and the granular bellied signifera-georgiana group.
The peculiar shoulder-girdle, in which the omosternum shows its double origin
from the epicoracoids, may be considered primitive, and it is possible that haswelli
is the most primitive member of the genus.
Crinia georgiana Tschudi.
Crinia georgiana Tschudi, 1838, Mem. Soc. neuchatel. Sci. nat., 2 : 38, 78 (Type locality : —
King George's Sound) ; Keferstein, 1867, Nachr. Ges. Wiss. Gbttingen, 18 : 347 (part) ;
idem, 1868, Arch. Naturgesch., 34 : 263 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed.
2 : 264 ; Fletcher, 1889, Proc. Linn. Soc. N.S.W., (2), 4 : 375, 387 ; idem, 1898, op. cit.,
22, 1897 : 676; Lucas & le Souef, 1909, Anim. Austral. : 276; Werner, 1914, Fauna
S.W. Austral., 4 : 408 ; Nieden, 1923, Das Tierreich, Anura I : 540 ; Hoffman, 1931,
S. Afr. J. Sci., 28 : 405 (? possibly C. signifera) ; Loveridge, 1935, Bull. Mas. comp.
Zool. Harv., 78 : 23.
AUSTRALASIAN LEPTODACTYLIDAF. 79
Cystignathus georgianus Dumeril & Bibron, 1841, Erpe't. Gen. ,8 : 416; Bell, 1843, Zool. Beagle,
R*pt- : 33. pi- l6. fig- 4 : Gunther, 1858, Cat. Batr. Sal. Brit. Mus. : 30 ; Krefft, 1865,
Pap. roy. Soc. Tasmania : 16 ; Krefft, 1867, Cat. Induslr. Prod. N.S.W., Add. : 107.
Pterophryne georgiana Gunther, 1867, Ann. Mag. nat. Hist., (3), 20 : 53.
Crinia ignita Cope, 1866, /. Acad. nat. Sci. Philad., (2), 6 : 95 (Type locality : — West Aus-
tralia).
Crinia signifera ignita Loveridge, 1935. Bull. Mas. comp. Zool. Harv., 78 : 25.
Pterophrynus affinis Gunther, 1864, Proc. zool. Soc. Lond. : 47, pi. 7, fig. 2 (Type locality : —
Western Australia) ; idem, 1864, Ann. Mag. nat. Hist., (3), 14 : 312 ; Krefft, 1865,
Pap. roy. Soc. Tasmania : 16.
Crinia affinis Gunther, 1868, Proc. zool. Soc. Lond. : 480.
Crinia georgiana var. affinis, Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 265.
Crinia affinis affinis Loveridge, 1935, Bull, Mus. comp. Zool. Harv., 78 : 26.
Crinia stolata Cope, 1867, /. Acad. nat. Sci. Philad., (2), 6 : 201 (Type locality : — West
Australia).
( niua georgiana var. stolata Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 264 ; Fry,
1914, Rec. IV. Aust. Mus., 1 : 203, pi. 28, fig. 1.
Vomerine teeth usually present in adults. Snout moderate, rounded, not
prominent, 1-14 to 1-35 times as long as the eye (mean of 29 specs. 1-25) ; canthus
rostralis rounded ; loreal region oblique ; nostril equidistant from the eye and the
end of the snout ; tympanum usually slightly distinct A to § the diameter of the
eye ; interorbital space about once and a quarter as broad as the upper eyelid.
Fingers moderately long, free, with well-developed subarticular tubercles ; palm
tubercular ; two metacarpal tubercles. Toes long, without dermal fringes ;
subarticular tubercles well developed ; two metatarsal tubercles and an oblique
tarsal fold from the inner. Tibio-tarsal articulation reaching the tympanic
region or the posterior corner of the eye.
Skin above from quite smooth to distinctly and irregularly rugose, specimens
from the same area showing all intermediate conditions. From a perfectly
smooth frog {affinis) the next stage is the progressive development of a dorso-
lateral fold from behind the upper eyelid (ignita or stolata) ; smaller secondary
folds maybe associated with these and, with the development of ridges and warts
elsewhere, the dorso-laterals tend to be broken up. There is always a parotoid-
like glandular swelling above and slightly in advance of the arm and a small
dermal papilla on the heel. Cloacal flap fimbriated. Lower surfaces coarsely
granular ; a triangular, finely granular patch on the hinder side of the thighs
behind the vent.
As in C. signifera, the dorsal colour pattern is associated with the rugosity
of the skin. Smooth specimens have the affinis colour, uniform, or nearly uniform
pale brown above, with a dark brown lateral stripe from the nostril through the
eye to the groin, partially interrupted in the parotoid region. Where the skin
is warty the warts are each picked out in dark brown, so that in the ignita or
stolata varieties there are two dark longitudinal dorsal stripes separated from each
other by a lighter area and from the lateral stripe by an area of grey ; a triangular
or V-shaped interorbital blotch is also developed and dark markings radiating
from the eye on the upper lip. With the breaking up of the dorso-lateral folds
and the increased development of warts and folds elsewhere the whole dorsum
has a complex, irregular pattern of dark marblings separated by lighter brown
or silvery grey. Limbs cross-barred. Armpits, groins and hinder aspect of the
knees brilliant carmine, this colour being " fast " in alcohol (traces still present
after nearly a century) but fugitive in formalin (destroyed in six months). Lower
surfaces of juveniles and half-grown specimens washed with pale brown except
for a white median line ; the tips of some of the warts and especially a larger,
pectoral spot on each side white or rose-tipped. With increasing age the brown
8o
H. W. PARKER
wash tends to become less and less distinct, but the pectoral glandular spots
remain a clearer white than the rest of the lower surfaces. Throat of the breeding
male infuscate. "*
Male with a vocal sac opening by a slit on each side of the tongue, but
without any definite nuptial pad.
Length from snout to vent : <$q I8'5_33'5 (mean of 14 exs. 26-0) ; $$ 22-6-
35-0 (mean of 12 exs. 27-3).
Distribution : West Australia.
The localities " Port Essington " and " Sydney " recorded by Boulenger
for this species, based on specimens in the British Museum, are very doubtful
indeed, and can be neglected unless confirmation is forthcoming — an unlikely
event after the lapse of more than half a century.
Specimens Examined.
Like Crinia signifera this species exhibits a good deal of variation in colour
pattern in correlation with variations in the degree of development of the dorsal
warts and folds. The two species parallel one another very closely in these
characters, and the variants of each, though forming a continuous series, can be
roughly subdivided into three main groups, the smooth, uniform, affinis type of
pattern, the striped stolata or ignita pattern with its concomitant longitudinal
folds and the irregularly marbled and warty typical form. It seems probable
that Loveridge in his recent (1935) consideration of the genus has assigned too
AUSTRALASIAN LEPTODACTYLIDAE
81
much importance to these characters, and rather neglected the other morpho-
logical characters such as vomerine teeth, digital fringes and degree of develop-
ment of the tympanum which the present author believes to provide a surer index
of relationship. As a result Loveridge recognizes no less than five separate forms
(out of a potential six), but applies specific and subspecific names to them on the
basis of colour and geography. The result is as follows :
This also leads to some entirely fictitious conclusions (p. 28) as to the develop-
ment of vomerine teeth in frogs from the south-west of Australia and their absence
in eastern " forms " of the same " species." Actually the presence or absence of
vomerine teeth appears to be a good specific character and the two species may
be distinguished thus :
C. georgiana.
Vomerine teeth usually present.
Digital fringes absent.
Tympanum larger and slightly dis-
tinct.
Flash colours of carmine strongly
developed and fast.
C. signifera.
Vomerine teeth always absent.
Digital fringes present in adults.
Tympanum smaller, not distinct.
Flash colours of orange or carmine
absent or, if present, fugitive.
Crinia glauerti Loveridge.
Crinia glauerti Loveridge, 1933, Occ. Papers Boston Soc. nat. Hist., 8 : 57 (Type locality : —
Mundaring Weir, 30 miles N.E. of Perth, West Australia) ; idem, 1935, Bull. Mus. comp.
Zool. Harv., 78 : 24.
The status of this species is doubtful. In the first place, although it is
described as a " miniature replica " of C. georgiana, digital fringes are said to be
present and vomerine teeth absent — both characters of signifera rather than
georgiana. However, in the two paratypes examined vomerine teeth are present,
though few in number, and there are no true digital fringes, but only the lateral
seams found in most frogs; they have the appearance of small examples of
georgiana, though having been fixed in formalin, it is not possible to say whether
the characteristic red on the legs of that species was also present. If the name
is applicable to these small, but sexually mature, examples which are so closely
allied to georgiana, the only differences to be found distinguishing the two are
size and the length of the snout. In signifera it is found that there is a great
range of variation in the size at which sexual maturity is reached, and that this
is not wholly connected with locality ; it may conceivably be dependent on the
climatic and food conditions prevailing during the larval and post-larval period ;
the same state of affairs might well be expected to occur in georgiana and, in that
event, glauerti would fall within the range of size of georgiana. There remains
NOVIT. ZOOL., 42, I 6
82 H. \V. PARKER
only the length of the snout. In the examples of glauerti examined it is 1-35 times
as long as the eye ; this is equal to the maximum length found (twice) in a series
of 29 examples of georgiana. Obviously it is necessary to know much more con-
cerning the range of variation of this character in glauerti before it can be used
with confidence diagnostically.
Vomerine teeth present. Snout pointed, slightly prominent, 1-35 times as
long as the eye, with somewhat marked canthus rostralis ; nostril equidistant
from the eye and the end of the snout ; tympanum slightly indicated ; inter-
orbital space 1-4 times the width of the upper eyelid. Fingers slender, free, with
well-developed subarticular tubercles ; palm tubercular ; two metacarpal
tubercles. Toes long, free, without fringes, but with well-developed subarticular
tubercles ; two metatarsal tubercles, the inner about \ its distance from the tip
of the inner toe, an oblique tarsal fold from the inner metatarsal tubercle. Tibio-
tarsal articulation reaching the tympanic region or the posterior corner of the eye.
Skin somewhat warty with a pair of curved folds, convex towards the mid-
line, on the scapular region. Granular beneath and on the hinder side of the
thighs ; a pectoral glandular spot close to the insertion of the arms ; a small
papilla on the heel.
Blackish above (formalin preservation) with traces of a darker pattern,
especially an interorbital mark ; popliteal region white (? red in life). Lower
surfaces dirty white, marbled with black in females ; throat of breeding male
infuscate.
Male with a vocal sac opening by a slit on each side of the tongue ; no nuptial
pad.
Length from snout to vent : <$$ 14-5-15-5 mm. ; $$ 20-5 mm.
Distribution : Known only from the type locality, near Perth, West
Specimens Examined.
Mundaring Weir, near Perth. Darlington.
(Paratype.)
,, ,, ,, ,. Darlington.
(Paratype.)
Mjoberg.
Crinia tasmaniensis (Giinther).
1 Camariolius varins (part) Peters, 1863, Mber. Akad. Berlin : 236.
Pterophrynus tasmaniensis Giinther, 1864, Proc. zool. Soc. Lond. : 48, pi. 7, fig. 3 (Type
locality: — van Diemen's Land) ; idem, 1864, Ann. Mag. nal. Hist., (3), 14 : 313 ;
Krefft, 1865, Pap. roy. Soc. Tasmania : 16.
Crinia tasmaniensis Keferstein, 1868, Arch. Naturgesch. : 265 ; Giinther, 1868, Proc. zool.
Soc. Lond. : 480 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 266 ; Lucas &
le Souef, 1909, Anim. Austral. : 276 ; Nieden, 1923, Das Tierreich, Anura I : 541, fig.
373 ; Blanchard, 1929, Aust. Zool., 5, 4 : 324 ; Loveridge, 1934, Pap. Proc. roy. Soc.
Tasmania, 1933 : 60 ; idem, 1935, Bull. Mus. comp. Zool. Harv., 78 : 28.
Camariolius (Pterophrynus) tasmaniensis (?) Parker, 1881, Phil. Trans, roy Soc. Lond., 3 :
102, pi. 19, figs. 1-5.
Vomerine teeth vestigial or, usually, absent. Snout depressed, not promi-
nent, 1-3 to 1-5 times as long as the eye ; canthus rostralis rounded ; loreal region
very oblique ; nostril a little nearer the eye than the end of the snout ; tympanum
indistinct ; interorbital space 1-2 to 1-4 times as broad as the upper eyelid.
Fingers moderate, free, with well-developed subarticular tubercles ; palm tuber-
cular ; two metacarpal tubercles. Toes long, with well-marked dermal fringes ;
1 See p. 89 below.
AUSTRALASIAN LEPTODACTYLIDAE 83
subarticular tubercles well developed ; two metatarsal tubercles ; no tarsal fold.
Tibio-tarsal articulation reaching the shoulder or tympanum in adult females
and from the latter point to the eye in males and juveniles.
Skin smooth or with more or less distinct plicae and small warts dorsallv ;
glandular but smooth or feebly granular ventrally except behind the thighs,
which are strongly granular. Indications of a minute papilla on the heel ; a
parotoid glandular thickening.
Dark brown, grey or olive above, with darker longitudinal stripes or spots.
Usually a lateral stripe from the nostril through the eye, interrupted above the
arm, and a pair of dorso-lateral stripes separated from the laterals by a light
brown, orange or even reddish stripe ; a dark, triangular or V-shaped inter-
orbital spot and a vertical dark bar beneath the eye. Lower surfaces brown or
blackish, marbled with white anteriorly and with white or bright red spots and
blotches posteriorly and on the limbs ; the extent of each colour is variable, so
that the belly may appear light with dark vermiculations ; a pair of pectoral
glandular spots also red. This red pigment is fast in alcohol (persistent after
80 years) but destroyed by formalin. Gular region of the male almost completely
black.
Male with a vocal sac opening by a slit on each side of the tongue and a
glandular, not rugose, nuptial pad at the base of the inner digit, not extending
appreciably on to the forearm.
Length from snout to vent : £ 24 mm. ; $ 30 mm. Juveniles at metamor-
phosis : 9-10 mm.
The species is apparently strictly aquatic. Breeding season extended in the
summer months ; sexual maturity reached at the end of the second season after
metamorphosis. Eggs 46 to 69 in a clutch (Blanchard, 1929).
Distribution : Tasmania at relatively high altitudes (2500-4000 ft.).
Specimens Examined.
B.M. 58.11.25.63 3 <?<J, 2 ??, Tasmania. Cotypes.
2 juvs.
1928.4.25. 1 $ National Park, Tasmania. Blanchard.
1936.12.2.3-7 1 (J, 4 ?$ Cradle Valley, Tasmania.
1036. 12.2.8 $ skel.
Austr. Mus. R. 4400-4401, 1 2 <J<J, juv. Summit of Mt. Wellington, Tas-
5087 1 mania.
R. 10337, 10349 3 6*6*.? The Spring, M t . Well ington ,
Tasmania.
R. 10336 5 Tasmania.
R.10339-10340 o", 3 ?$ Cradle Valley. Tasmania.
K. 10338 (J Ulverstone, Tasmania.
R.10341, 10342, I 4 JJ, $ National Park, c. 4000 ft., Tas-
10345, 10347 t mania.
R.10343. 1034 |. ■ 5 .; j. 2 $$, Lake Fenton, National Park, „
10346, 10348 juv. Tasmania.
K. 10074 6"? Mt. Wellington, Tasmania.
R. 10352 <J§ Between Port Arthur and Roger's
River, Tasmania.
Crinia acutirostris Andersson.
Crinia acutirostris Andersson, 1916, Svenska Vetensk. Ikad. Handl., 52, 9 : 8, pi. 1, li;;. 1 (Type
locality: -Malanda, Queensland); Loveridge, [935, Bull. > Zool llarv.,
78 : 30.
Vomerine teeth absent. Snout truncate, not projecting beyond the labial
margin, once and a quarter as long as the rye; canthus rostralis angular; loreal
84 H. W. PARKER
region nearly vertical ; nostril equidistant from the eye and the end of the snout,
interorbital space a little broader than the upper eyelid ; tympanum slightly
distinct, a little less than half the diameter of the eye. Fingers free, with well-
developed subarticular tubercles ; palm smooth ; two metacarpal tubercles.
Toes long, slightly dilated terminally, with well-marked dermal fringes ; subar-
ticular tubercles moderate ; a single, inner metatarsal tubercle ; a tarsal fold
present or absent. Tibio-tarsal articulation reaching the eye.
Skin quite smooth or granular and tuberculate with spinose warts in breeding
males, these spines forming series on each side of the vent and on the hind limbs,
with a pair on the back behind the eyes and others forming a triangle on the
posterior part of the back ; sometimes a narrow glandular fold from the eye to
the groin. Smooth beneath. Sometimes a papilla on the heel.
Grey-brown above, uniform or with a darker, triangular, interorbital marking
and some smaller dark spots on the posterior part of the back ; a dark lateral
band from the nostril to the groin, sometimes narrowly light-edged above ; two
light bars on the lip beneath the eye ; groins yellow or red ; limbs cross-barred ;
lower surfaces white (reddish-yellow in life), with a reticulate brown stippling ;
throat of the male infuscate.
Male with a vocal sac opening by a slit on each side of the tongue.
Length from snout to vent : <$ 23 mm. ; $ 30 mm.
Distribution : Queensland.
Specimens Examined.
M.C.Z. 18416 $ Mt. Spurgeon, Q. Darlington.
B.M. 1938. 7. 2. 1 (J Mundubbera, Q. Sherrin.
Crinia signifera Girard.
Crinia (Ranidella) signifera Girard, 1853, Proc. Acad. nat. Sci. Philad. : 422 (Type locality : —
New Holland) ; Cope, 1867, /. Acad. nat. Sci. Philad., (2), 6 : 203.
Ranidella signifera Girard, 1858, U.S. Explor. Exped.. Herp. : 44, pi. 3, figs. 39-43.
Crinia signifera Boulenger, 1882, Cat. Batr. Sal. Brit. Mus.. ed. 2 : 265 ; Fletcher, 1889,
Proc. Linn. Soc. N.S.W., (2), 4 : 375 ; idem, 1891, op. cit., (2), 6 : 272-274 ; idem, 1892,
op. cit., (2) 7 : 8, 9 ; Lucas, 1892, Proc. roy. Soc. Victoria, 4 : 61 ; Fletcher, 1894, Proc.
Linn. Soc. N.S.W., (2), 8 : 526, 527, 528, 530; idem, 1898, op. cit., 22, 1897 : 676;
Lucas & le Souef, 1909, Anim. Austral. : 275, fig. ; English, 1910, Proc. zool. Soc. Land. :
629, pi. 51, figs. 4, 5; Werner, 1914, Fauna S.W. Austral. : 411 ; Andersson, 1913, A'.
Svenska VetenshAhad. Handl., 52, 4 : 12, pi. 1, fig. 1; Roux, 1920, Rev. Suisse Zool.,
28 : 115 ; Nieden, 1923, Das Tierreich, Anura I : 541; van Kampen, 1923, Amph.
Indo-Austral. Archip.: 21; Blanchard, 1929, Aust. Zool., 5, 4:326; Waite, 1929,
Rept. Amph. S. Austral. : 257; Glauert, 1929, /. roy. Soc. W. Austral., 15:45;
Trewavas, 1933, Phil. Trans, roy. Soc. Lond., 222, b : 436, figs. 26, 27.
Crinia signifera signifera Loveridge, 1934, Pap. roy Soc. Tasmania, 1933 : 59 '• Oeser, 1934, Bl.
Aquar. -u, Terrarienk., 45 : 355 ; Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 24.
Pseudophryne australis (part) Fitzinger, 1861, S.B. Ahad. iviss. Wien, 42 : 415.
Pterophrynus verrucosus Lutken, 1862, Vidensk. Medd. naturh. Foren. Kbh. : 302, pi. 1, fig.
4 (Type locality : — Australia).
Crinia verrucosa, Giinther, 1868, Proc. zool. Soc. Lond. : 478.
? Camariolius varius Peters, 1863, Mber. Akad. Berlin : 236 (Type locality : — near Adelaide).
Pterophrynus varius Steindachner, 1867, Reise Novara, Zool., Amph. : 31, pi. 2, figs. 1-6 ;
Kre'fft, 1867, Cat. Industr. Prod. N.S.W., Add. : 107.
Crinia georgiana var. varia Keferstein, 1867, Nachr. Ges. Wiss. Gbttingen, 18 : 349.
Crinia varia Keferstein, 1868, Arch. Naturgesch., 34 : 264, pi. 6, figs. 11-13.
Crinia stictiventris Cope, 1867, /. Acad. nat. Sci. Philad., (2), 6 : 203 (Type locality : —
Australia).
Pterophrynus fascialus Steindachner, 1867, Reise Novara, Zool., Amph. : 31, pi. 5, figs. 3, 4
(Type locality : — New South Wales).
Pterophryne fasciata Giinther, 1867, Ann. Mag. nat. Hist., (3), 20 : 53.
Crinia fasciata Giinther, 1868, Proc. zool. Soc. Lond. : 478.
AUSTRALASIAN LEPTODACTYLIDAE
85
Crinia georgiana var. laevipes Keferstein, 1867, Nachr. Ges. Wiss. Gottingen. 18 : 348 (Type
locality : — Sydney).
Cystignathus sydneyensis Krefft {nom. nud.), Keferstein, 1867, loc. cit.
Crinia affinis haswelli (non Fletcher) Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 27.
Crinia georgiana (part) Loveridge, 1935, torn. cit. : 23.
Pseudophryne semimarmorata (non Lucas) Loveridge, 1935, torn. cit. : 34.
This species exhibits a great deal of variation both in morphological characters
and colour-pattern. The latter is believed to have little taxonomic significance,
but at least two apparently distinct geographical races appear to be recognizable,
in addition to the typical form, as indicated in the key. None of the existing
names are obviously applicable to either of these forms, and it will lead to less
confusion to give them new names, which may subsequently have to be placed in
synonymy, rather than to use, incorrectly, ones which already exist.
Crinia signifera signifera (Girard).
Vomerine teeth absent. Snout short, rounded, not prominent, i-i to 1-4
times as long as the eye ; canthus rostralis rounded ; loreal region oblique ;
nostril equidistant from the eye and the end of the snout, the internarial space
less than, or equal to the distance from eye to nostril ; tympanum hidden ; inter-
orbital space about 1-3 times the width of the upper eyelid. Fingers moderately
long, free, with well-developed subarticular tubercles ; palm tubercular ; two
Text-fig
Hand of Crinia signifera.
metacarpal tubercles. Toes long, almost always with well-developed dermal
fringes in adults, though these may be only feebly developed or absent in juveniles
(see table of dimensions below) ; subarticular tubercles well developed ; two
metatarsal tubercles, the outer not, or scarcely, more than \ its distance from the
tip of the inner toe and an oblique tarsal fold from the inner. Tibio-tarsal
articulation reaching the tympanic region or the eye in males and the shoulder
or tympanum in females.
Skin above quite smooth to distinctly and irregularly warty ; a complete
transition can be traced in specimens from the same region between these extremes,
86 H. W. PARKER
with an intermediate stage with two strong folds down the back commencing
behind the upper eyelids and slightly convergent on the scapular region ; with the
development of additional warts elsewhere these folds may be broken up, though
usually traces of them persist ; a more or less distinct glandular aggregation
above the insertion of the fore-limb. Belly and gular region coarsely granular ;
a definite area finely granular below the vent on the hinder side of the thighs.
A small papilla on the heel. Cloacal flap fimbriated.
The colour pattern is intimately connected with the degree of wartiness of
the skin. Smooth specimens are almost uniform brown above, with a dark
lateral stripe from the nostril through the eye, interrupted behind the tympanic
region and continued along the flanks to the groin ; lips with dark bars irregu-
larly radiating from the eye ; usually there is an indistinct triangular marking
between the eyes, and a fine white line from snout to vent. With the develop-
ment of warts, each tends to be light in the centre, edged with darker ; in con-
sequence, where there are two well-developed dorsal folds a striped pattern is
developed, the centre of each fold forming a narrow light line bordered on each
side by darker, these darker areas being in turn separated from each other in the
middle and from the lateral stripe by broader light zones, which may be marked
with very narrow dark streaks. Where the dorsum is irregularly warty all over,
a pattern of obscure irregular darker marblings ensues. Limbs with diagonal
cross-bars. Anal flap invariably with a light dot. Lower surfaces more or less
profusely brown spotted, these markings attaining their maximum development
in specimens from New South Wales which have the colours almost reversed,
i.e. are brown with large white spots. There is constantly a pair of small white,
glandular pectoral spots, one close to the insertion of each fore-limb. Gular
region of male plicate and infuscate, but the edge of the jaw lighter. In life the
colours are browns and greys above, but there may be suffusions of yellow,
orange or carmine on the concealed surfaces of the limbs and flanks, these colours
being fugitive in alcohol.
Male with a vocal sac opening by a slit on each side of the tongue, and in the
breeding season, a non-spinous glandular area on the base of the inner finger and
distal half of the inner surface of the forearm.
Length from snout to vent, maxima : <$ 25 mm. ; $ 30 mm. Juveniles at
metamorphosis : 6-7-10 mm.
There is some geographical variation in size with latitude, and in the size
at which maturity is attained, and digital fringes developed. This may be
summarized as follows :
On Kangaroo Island there may be a distinct race, recognizable by even larger
size and suppression of the digital fringes ; unfortunately the series examined is
somewhat shrivelled, so that the latter character cannot be satisfactorily deter-
mined. Females of 23 mm. do not appear to be fully mature and have only
rudimentary digital fringes.
AUSTRALASIAN LEPTODACT YI.IDAE
87
The species is largely aquatic, with a considerable tolerance of differing con-
ditions from mountain streams to shallow, muddy pools. Breeding occurs
throughout the year after rain in New South Wales and South Australia. The note
of the males is an insect-like chirrup, embrace lumbar ; eggs to the number of
about 150 are laid in masses attached to objects at the bottom of the water ;
individual eggs are about 1-3 mm. in diameter, with a jelly-capsule of 4 mm. in
diameter. Hatching takes place in about ten days after oviposition, and the
tadpole stage is prolonged (estimated as three months by Harrison, 1922 : 31).
Distribution : New South Wales, Victoria (?), South Australia, West
Australia, Queensland and Northern Territories ; New Guinea (Merauke).
Specimens Examined.
88
H. W. PARKER
Austr. Mus. R.7413
R.8636
— 6953
— 6965
A.2879
R.204
R.4109
R.4209
R.4250
R.4327-9
R-4352
R.4364
R.4370
R-4397-9
R.4628-9
R.4764
R.5210
R.5888-
5892
R. 6304-6
R.6490
R.6496-8
R.6500
R.7328
R-7378-
738o
R-7397
(part)
R-7977
R.8085
R.8242
R8303,
8321
R.8487
R.8490
R.8497
R.8502
R.8638
(part)
R.8725
R.8765
R.9835
R. 10352
7 6"<J, 3 9?
6". 2?$
$
?
imm. (J
2 6"6\9
3
5 99
4 99
10 0*6"
3
3
6*. 2$?
6 33. 2 99
3. 9. 8 Hgr.
2 99
2 33. 2 99
4 33. 9 99. 3
j u vs.
4 33. 5 99. 5
juvs.
2 6V. 9
4 99
d"9
3 33. 9
10??
2 33. 9
6"
9. juv.
4 6V. 4 99
8 S3. 4 99. 2
juvs.
4 Hgr.
n cJ6", 99 and
juvs.
3 99
3
9
2 6*6", 3 juvs.
juv.
3. 2 99
Waroo, Inglewood, 60
miles W. of Stanthorpe,
Q-
Isis Scrub.
New South Wales.
Sydney, N.S.W.
Smithfield, near Sydney
N.S.W.
Kangaroo Valley, N.S.W.
Lindfield, near Sydney,
N.S.W.
Maroubra, near Sydney
N.S.W.
Warrell Creek, Nambucca
River, N.S.W.
Lindfield, near Sydney
N.S.W.
Woodford, Blue Mts.,
N.S.W.
Lindfield, N.S.W.
Bundanoon, N.S.W.
Lindfield, N.S.W.
Warrell Creek, Nambucca
River, N.S.W.
Tamworth, N.S.W.
Megalong Valley, Blue
Mts., N.S.W.
Gurravembi, near Macks-
ville, Nambucca River,
N.S.W.
Ditto.
Tweed River, N.S.W.
Wilde's Meadow, Moss
Vale, N.S.W.
Avoca, near Gosford,
N.S.W.
Bungendore, 2290 ft.,
N.S.W.
Burrawang, 2000 ft.,
N.S.W.
Yass, N.S.W.
Upper Colo, via Rich-
mond, N.S.W.
Maroubra, Sydney,
N.S.W.
Longueville, near Sydney,
N.S.W.
Maroubra, near Sydney,
N.S.W.
Wilde's Meadow, Moss
Vale, N.S.W.
Tweed River, N.S.W.
Wilde's Meadow, Moss
Vale, N.S.W.
Avoca, N.S.W.
Bundanoon, N.S.W.
Northwood, near Sydney,
N.S.W.
Long Bay Rifle Range,
Sydney, N.S.W.
Rose Bay, Sydney,
N.S.W.
Mosman, Sydney, N.S.W.
AUSTRALASIAN LEPTODACTYLIDAE
89
The specimens referred to haswelli by Loveridge (1935 : 27) have no
vomerine teeth, and, to judge from the specimen examined (19586), no red on
the limbs ; they appear to be normal smooth-skinned signifera with which several
of them had previously been associated in the collections of the M.C.Z.
The colour pattern of signifera is extremely variable, and is paralleled by
that of its congener C. georgiana ; an almost exactly similar series of colour
variations in a single species is exhibited by the S. American Physalaemus cuvieri
Fitzinger. Camariolius Darius Peters 1863 is included in the synonymy of this
species, although, as Cope (1867 : 203) and Boulenger (1882 : 266) pointed out,
Peters may have had two species confounded ; in the description of varius based
on six specimens, five from Adelaide and one from an unknown source, the belly
is said to be granular or smooth. The specimens from Adelaide are almost
certain to be signifera, but the other, if it had a smooth venter, may be tas-
maniensis. Should this prove to be so, Cope's restriction of the name varius
for the smooth-bellied form must stand and the name would have priority over
tasmaniensis (1864).
Crinia signifera englishi subsp. n.
Vomerine teeth absent. Snout short, somewhat pointed to slightly promi-
nent, with obtuse canthus rostralis and very oblique lores, 1-2 to 1-5 times as
long as the eye ; nostril equidistant from the tip of the snout and the eye, or a
little nearer the latter ; internarial space as long as, or slightly longer than, the
distance from eye to nostril ; interorbital space 1-25 to 1-5 times as wide as the
upper eyelid. Fingers moderate, free, with well-developed subarticular tubercles ;
palm tubercular ; two metacarpal tubercles. Toes moderate, with well-developed
dermal fringes in adults ; subarticular tubercles well developed ; two metatarsal
tubercles, the outer approximately half as long as its distance from the tip of
the inner toe, and an oblique tarsal fold from the inner. Length of the foot
contained 1-84 to 2-23 times in the length from snout to vent (mean of 32
specimens 2-oi). Tibio-tarsal articulation reaching the tympanic region.
Skin with small scattered warts above and usually with a more or less con-
tinuous, dorso-latcral, linear row on each side of the back ; a glandular aggre-
gation in the parotoid region. Lower surfaces and hinder side of the thighs
granular. A small papilla on the heel, ("loacal flap fimbriated.
Brown or greyish above, almost uniform sometimes, but usually with more
or less distinct traces of some or all of the following darker brown markings.
H. W. PARKER
A triangular interorbital spot, the apex directed posteriorly ; a dark dorso-lateral
stripe on the dorsolateral line of warts ; a canthal stripe continued behind the
eye as a temporal spot and a lateral stripe from above the fore-limb to the groin.
Lower surfaces yellowish white, mottled with brown ; a white, glandular pectoral
spot close to the insertion of the fore-limb. Anal flap lighter. Throat of the
male infuscate. In life the ground-colour dorsally is dark brown or grey as a
rule, males tending to be darker than females, and sometimes almost black.
There may be markings of bright orange and the hinder side of the thighs may be
washed with blood-red, though this colour is fugitive ; tadpoles and juveniles
black.
Male with a vocal sac opening by a slit on each side of the tongue and a
glandular, non-spinous nuptial pad on the inner finger and distal half of the
forearm.
Length : $$ 18-8-24-5 mm. (mean of 18, 22-2 mm.) ; $$ 21-5-30-0 mm.
(mean of 13, 24-9 mm.).
Largely aquatic with a wide tolerance, but very active on land. Call of
the male a cricket-like chirp ; breeding season chiefly from May to August ; eggs
laid in masses spread over stones and weeds at the bottom of shallow water.
Apparently does not hibernate (English).
Distribution : Tasmania.
Crinia signifera montana subsp. n.
Vomerine teeth absent. Snout short, bluntly rounded, not prominent, with
obtuse canthus rostralis and very oblique lores, 1-2 to 1-5 times (mean of 31,
1-34) times the length of the eye ; nostril a little nearer the eye than the end of
the snout ; internarial space longer than the distance from eye to nostril ; inter-
orbital space 1-2 to 1-5 times the width of the upper eyelid. Fingers moderate,
free, with well-developed subarticular tubercles ; palm tubercular ; two meta-
carpal tubercles. Toes with short, fleshy fringes in adults ; subarticular tubercles
well developed ; two metatarsal tubercles, the inner about half as long as its
distance from the tip of the inner toe ; an oblique tarsal fold from the inner.
Length of foot contained 1-9-2-35 (mean of 32, 2-12) times in the length from
snout to vent. Tibio-tarsal articulation reaching the shoulder.
Skin very warty above, the warts tending to form longitudinal ridges, of
which the most constant are an irregular dorso-lateral pair ; a glandular parotoid
aggregation behind the tympanic region. Lower surfaces and hinder side of
thighs granular. A small papilla on the heel. Cloacal flap fimbriated.
Colour pattern and secondary sex characters essentially similar to those of
englishi.
Length : S3 ^9'5~2yy mm. (mean of 15, 21-8 mm.) ; $$ 20-5-28-5 mm.
(mean of 16, 24-5 mm.).
Distribution : Mt. Kosciusko.
AUSTRALASIAN LEPTODACTYLIDAE
91
Cotypes Examined.
9742
o"9
Mt. Kosciusko, 5000 ft.
5500 ft-
7000 ft.
Pretty Point, Mt. Kos-
ciusko.
Lake Cootapatamba, Mt.
Kosciusko, 6500 ft.
Rawson Pass, Mt. Kos-
ciusko, 8800 ft.
MYOBATRACHUS.
Breviceps (non Merrem) Gray, 1841, in Grey, Journ. Exped. W. Austral.. II : 448.
Myobatrachus Schlegel, 1850, Proc. zool. Soc. Land. : 9 (Type species : — M. paradoxus) ;
idem, 1851, Ann. Mag. nat. Hist., (2), 7 : 70; Gunther, 1858, Cat. Batr. Sal. Brit. Mus. :
3 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 328 ; Cope, 1889, Bull. U.S.
nat. Mus., 34 : 259 ; Nieden, 1923, Das Tierreich, Anura I : 167 ; Noble, 1931, Biol.
Amph. : 498.
Myiobatrachus Schlegel, 1858, Handleid. Dierk., 2 : 59, 545.
Chelydobatrachus Gunther, 1858, Cat. Batr. Sal. Brit. Mus. : 53 (Type species : — Breviceps
gouldii Gray) ; Cope, 1865, Nat. Hist. Rev., n.s., 5 : 102.
Maxillary teeth absent. No prevomer ; a large fronto-parietal foramen ;
quadrato-jugal short, not reaching the maxilla ; ear fully developed, with a
very large, thick, extraplectal cartilage, covering the whole of the tympanum.
Vertebrae procoelous, with the condyle incompletely ankylosed and notochord
Text-fig. 20. — Ventral elements of the shoulder girdle of Myobatrachus gouldii.
persistent ; eight presacrals ; sacral diapophyses broadly dilated ; coccyx articu-
lating by two condyles. No omosternum ; clavicles short and stout, widely
dilated mesially ; epicoracoids meeting edge to edge for a third of their length
anteriorly and so approaching an arcifrro-firmisternal condition ; coracoids directed
obliquely backwards ; sternum small, cartilaginous or calcified (text-fig. 20).
92 H. W. PARKER
Terminal phalanges simple. Distal tendon of the m. semitendinosus passing deep
(dorsal) to the m. gracilis. Alary processes of the hyoid, broad wing-like expan-
sions of almost the whole lateral margins of the hyoid plate.
Pupil horizontal with a ventral angle. Tongue small, oval, partly free
behind. Digits not dilated distally.
Myobatrachus gouldii (Gray).
Breviceps gonldi Gray, 1841 (April), Ann. Mag. nat. Hist., 7 : 89 (Type locality: — Western
Australia) ; Gray, 1841, in Grey, Journ. Exped. W. Austral. II : 436, 448, pi. 1, fig. 1.
Myobatrachus gouldii Gray, 1850, Proc. zool. Soc. Land. : 10; idem, 1851, Ann. Mag.
nat. Hist., (2), 7 : 70 ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 329 ; Fletcher,
1898, Proc. Linn. Soc. N.S.W., 22, 1897 : 680, 682 ; Lucas & le Souef, 1909, Anim.
Austral. : 288 ; Werner, 1914, Fauna SJV. Austral., 4 : 421 ; Fry, 1914, Rec. W. Aust.
Mus., 1 : 208 ; Alexander, 1922, /. Linn. Soc. Lond. {Zool.), 34 : 462 ; Nieden, 1923,
Das Tierreich, Anura I : 167, fig. 216; Harrison, 1927, Rec. Aust. Mus., 15 : 287 ;
Kinghorn, 1932, Rec. Aust. Mus., 18 : 361 ; Loveridge, 1935, Bull. Mus. comp. Zool.
Harv., 78 : 37.
Chelydobatrachus gouldii Giinther, 1858, Cat. Batr. Sal. Brit. Mus. : 53, 138 ; Krefft, 1865,
Pap. roy. Soc. Tasmania : 17.
Breviceps Heliogabali Gray, 1841, in Grey, Journ. Exped. IV. Austral., II : pi. 1, fig. 1, caption.
Myobatrachus paradoxus Schlegel, 1850, Proc. zool. Soc. Lond. : 9 (Type locality : — Swan
River, W.A.) ; Giinther, 1858, Cat. Batr. Sal. Brit. Mus. : 3, 128 ; Krefft, 1865, Pap.
roy. Soc. Tasmania : 16 ; Peters, 1867, Mber. Akad. Berlin : 37.
Myiobatrachus paradoxus Schlegel, 1858, Handleid. Dierk., 2 : 59, 545, pi. 4, fig. 76.
Habitus globose. Head small, as long as broad, its length contained 4-6
times in the length from snout to vent. Snout bluntly rounded, 17 times as
long as the eye ; nostrils terminal, directed forwards ; canthus rostralis rounded ;
loreal region oblique ; interorbital space twice as broad as an upper eyelid ;
crown of the head dome-shaped owing to the presence of a pad of fibro-glandular
thickened skin ; tympanum hidden. Limbs very short and stout. Fingers
short, stout, depressed, the first shorter than the second ; fourth very short (the
phalanges reduced to small nodules) but very stout, due to the skin being thickened
and somewhat cornified ; a tract of similarly thickened skin extends along the
outer side of the forearm for a short distance ; subarticular tubercles indicated
on the proximal joints of the first three fingers ; an indefinite outer, but no inner,
metacarpal tubercle. Toes very short, free, depressed, the outermost stoutest ;
traces of subarticular tubercles on the proximal joints only ; an extensive, but
indistinct, outer metatarsal tubercle, the inner small and only faintly indicated.
The total length of the hind limb (vent to tip of fourth toe) about § the length
from snout to vent.
Skin shagreened and porous above, but not glandular ; similar beneath or
slightly warty on the abdomen.
Dull brown above, sometimes with faint, ill-defined darker and lighter areas
and small pink dots ; the cornified skin of the crown of the head and on the outer
fingers and forearm yellow or rusty brown ; a fine light vertebral line may be
present. Dirty white beneath, uniform or dotted with dark brown.
Length from snout to vent : <$ 44 mm. ; 9 57 mm.
A cryptozoic species frequenting termites' nests. Male with a vocal sac and,
at the breeding season (?), with numerous small, horny spinules very numerous
on the dorsal surfaces, larger and more scattered beneath. Eggs very large,
ovarian follicles 4 to 4-5 mm. in diameter.
Distribution : West Australia.
AUSTRALASIAN LEPTODACTYLIDAE 93
Specimens Examined.
METRACRINIA gen. nov.
Type species : Pseudophryne nichollsi Harrison.
Pseudophryne (part) Harrison, 1927, Rec. Aust. Mus., 15 : 285 ; Barbour & Loveridge, 1929,
Copeia, 170 : 12 ; Loveridge, 1935, Bull. Mus. cotnp. Zool. Harv., 78 : 35.
Maxillary teeth absent. Prevomer represented by a minute narrow, non-
dentigerous bone bordering the antero-mesial edge of each choana ; a very
large fronto-parietal foramen. Ear fully developed. Vertebrae procoelous, the
condyle incompletely ankylosed ; notochord persistent ; sacral diapophyses
broadly dilated ; urostyle articulating by two condyles1 ; eight presacral vertebrae.
Omosternum rudimentary ; sternum undivided, cartilaginous ; procoracoids very
broad. Terminal phalanges simple.
Distal tendon of the m. semitendinosus passing dorsal to the m. gracilis.
Alary process of the hyoid a wing-like expansion of almost the whole lateral
border of the hyoid plate.
Pupil horizontal with a ventral angle. Tongue narrow, oval, half free behind.
Toes free and not dilated distally.
Metacrinia nichollsi (Harrison).
Pseudophryne nichollsi Harrison, 1927, Rec. Aust. Mus., 15 : 284 (Type locality : — Pem-
berton, W.A.) ; Loveridge, 1935, Bull. Mus. cotnp. Zool. Harv., 78 : 35.
Pseudophryne bibroni (non Steindachner) Barbour & Loveridge, 1929, Copeia, 170 : 12.
Snout very short, bluntly rounded, not prominent, a little longer or a little
shorter than the diameter of the eye ; canthus rostralis rounded ; loreal region
slightly oblique ; nostrils much nearer the tip of the snout than the eye ; tym-
panum hidden ; interorbital space once and a quarter as wide as the upper
eyelid. Fingers short, with flat subarticular tubercles ; palm slightly tubercular ;
two metacarpal tubercles. Toes short, free .without dermal fringes, not dilated
terminally, with feebly-developed subarticular tubercles ; two rather indistinct
metatarsal tubercles ; no tarsal fold. Tibio-tarsal articulation not quite reaching
the shoulder.
Skin uniformly and regularly warty above, with a pair of curved folds from
behind the upper eyelids, convergent towards the middle line. Ventral surfaces
regularly and coarsely granular.
Uniform dark brown or blackish above, sometimes with obscure darker
markings or with a silvery-grey wash and minute pink dots. Lower surfaces
dark grey, blue or black, marbled and spotted witli white ; a pair of bright yellow
pectoral glandular spot-^ and similarly coloured spots on the poMnioi part ol
the belly in front of the thighs (sometimes confluent), in the popliteal region, on
1 In the single specimen examined an additional vertebra appears t<> I"- intercalated
between the urostyle and the sacrum, fused with the former but arti( ulating with the lattei
by a single condyle and by zygapophyses ; probably this is an anomalous condition.
Q4 H. W. PARKER
thighs and tibiae, and on the upper, inner side of the metatarsus. Sometimes
a narrow white vertebral line is present in juveniles and may persist as a coccygeal
stripe in the adult ; a similar line may traverse the hinder side of the thighs above
the vent. Throat of the male darker.
Length from snout to vent : <J 23 mm. ; $ 25 mm.
The species is sluggish in disposition, crawling rather than hopping, and is
often found in association with the ant Myrmecia regularis Crawley, but not with
any other species ; the frogs are often to be found using the galleries constructed
by the ants under logs and stones in damp places. When disturbed the frog
habitually turns over on its back to exhibit the brilliantly coloured lower surfaces.
The eggs are large, about 5 mm. in diameter, and are laid under cover of some
kind, such as a log, on land close to water ; 25 to 30 eggs appear to constitute
a clutch, and development reaches an advanced stage within the egg. Breeding
period apparently during late summer.
Distribution : West Australia, west of the Darling Range and south of
Geographe Bay.
Specimens Examined.
B.M. 70.6.26.56 2 ? Krefft.
Pemberton, W. Australia. Schevill.
Yanmah, near Manjimup, Glauert.
W.A.
[927.8.30.1-2 (J? Augusta, W.A. Brooks.
PSEUDOPHRYNE Fitzinger.
Bombinator Gray, 1835, Proc. zool. Soc. Lond. : 57.
Phryniscus (part) Dumeril & Bibron, 1841, Erpet. Gen., 8 : 722.
Pseudophryne Fitzinger, 1843, Syst. Rept. : 32 (Type species : — Phryniscus australis
Dum. & Bibr. = P. semimarmorata Lucas1) ; Giinther, 1858, Cat. Batr. Sal. Brit. Mus. :
45 ; Cope, 1865, Nat. Hist. Rev., n.s., 5 : 103 ; Steindachner, 1867, Reise Novara, Zool.,
Amph. : 34 ; Keferstein, 1868, Arch. Naturgesch., 34 : 271 ; Boulenger, 1882, Cat. Batr.
Sal. Brit. Mus., ed. 2 : 277 ; Cope, 1889, Bull. U.S. nat. Mus., 34 : 259 ; Nieden, 1923,
Das Tierreich, Anura I : 147 (part) ; Harrison, 1927, Rec. Aust. Mus., 15 : 286 ; Waite,
1929, Rept. Amph. S. Austral. : 264 ; Noble, 1931, Biol. Amph. : 498.
? Bufonella Girard, 1853, Proc. Acad. nat. Sci. Philad. : 424 (Type species: — Bufonella
a ucigera).
Maxillary teeth absent. Prevomer reduced to a fragment bordering the
choana mesially, or entirely absent, never dentigerous. Fronto-parietals widely
separated.2 Ear reduced, without tympanum, annulus tympanicus, cavum.
tympani, columella auris or Eustachian tubes. Vertebrae procoelous, but with
the condyle incompletely ankylosed ; notochord persistent ; sacral diapophyses
moderately dilated ; urostyle articulating by two condyles ; 8 presacral
1 Phryniscus australis Dumeril & Bibron, 1841, Erpet. Gen., 8 : 725, does not appear to
have been conspecific with Bombinator australis Gray, 1835, but to have been a composite
of two other species. One of these, that figured on plate 100, fig. 2, was considered typical,
the other (pi. 100, fig. 4) being a " variety " ; the typical form is the species subsequently
named semimarmorata by Lucas, which consequently becomes genotype of Pseudophryne
Fitzinger.
- W. K. Parker, 1881, Phil. Trans, roy. Soc. Lond., 172: 230, pi. 42, figs. 1-7, describes and
figures a skull ascribed to Pseudophryne bibroni. It differs from any member of the genus
examined by the present author in having the fronto-parietals in contact mesially, a larger,
bifid prevomer, an annulus tympanicus, the shape of the basi-sphenoid and the length of the
post-orbital part of the skull. It seems certain that there must have been a misidentification,
and, since Parker is so insistent on a number of characters which are " quasi-juvenile " or
show " arrested metamorphosis " or " relapse," the probability is that the specimens were
really juveniles of something very different.
AUSTRALASIAN LEPTODACTYLIDAE 95
vertebrae. Omosternum vestigial ; sternum undivided ; cartilaginous. Terminal
phalanges simple.
Distal tendon of the m. semitendinosus perforating the distal tendon of the
mm. graciles. Alary process of the hyoid a broad, wing-like expansion of the
lateral border of the hyoid plate.
Pupil horizontal. Tongue small, narrow and oval, half free behind. Toes free.
In recent years it has been believed that Pseudophryne and Crinia, though
at one time referred to two different families, were very closely allied, and the
former has even been described as a Crinia without teeth. But there are some
other differences distinguishing the two, amongst which the most important
appears to be the condition of the ear. In Crinia this organ is fully developed,
with Eustachian tubes, a columella auris, cavum tympani, annulus tympanicus
and tympanum, but in the great majority of the species of Pseudophryne, includ-
ing the genotype, all these structures1 appear to be completely absent. It is
scarcely conceivable that, amongst a group of closely allied species, such a pro-
found modification should appear haphazard, and the presence or absence of the
middle ear must be regarded as a good index of affinity. The only species of
" Pseudophryne " in which the ear is known to be fully developed are P. rugosa
and P. mjobergi Andersson, and P. nichollsi Loveridge. These three species
must accordingly be removed from the genus and their generic relationships
involve a reconsideration of the value of maxillary teeth as a generic character.
The affinities of P. rugosa with Uperoleia have already been recognized, and Love-
ridge (1935 : 31) has recently reduced rugosa to the status of a subspecies of
U. marmorata. This view is not entirely acceptable, for the question of teeth,
which are absent in rugosa but present in marmorata, has been entirely neglected.
But there can be no doubt that rugosa is closely allied to marmorata. Both
species are characterized by the presence of large parotoid glands, and differ from
any species of any of the allied genera Pseudophryne, Crinia and Glauertia in the
closure of the fronto-parietal foramen. With " Pseudophryne " nichollsi, how ever,
the case is rather different. The presence of a fully developed ear suggests
affinity with Crinia, but the species differs from any member of that genus in its
stout habitus and short limbs. It may, perhaps, be regarded as a toothless
Crinia, but it has so little in common with the other species that it has obviously
diverged a long way, and its relationships can best be expressed by referring it
to a distinct new genus. " Pseudophryne " mjobergi possessing a fully developed
auditory apparatus and teeth is allied to Crinia, but its webbed feet suggest that
it may be more appropriately placed in the genus Glauertia.
The description of the type species of Bufonella Girard is so reminiscent of
the very characteristically marked Pseudophryne australis that it seems possible
that the presence of a " tympanum," said to distinguish Bufonella, may be due
to an error of observation or an artifact.
Synopsis of the Species.
I. Two large shovel-shaped metatarsal tubercles ; a large inguinal, but no
femoral, gland ; inner toe with only a single phalanx . P. guentheri.
II. Metatarsal tubercles not shovel-shaped ; no inguinal, but usually a post-
femoral, gland ; inner toe with two phalanges.
A. No gland behind the thigh .... P. occidentalis sp. nov.
1 A vestige of the Eustachian tube may persist as a diverticulum from the mouth.
9° H. W. PARKER
B. A gland on the distal half of the hinder side of the thighs.
(1) Snout pointed, prominent, the internarial space shorter than the
distance between the nostril and the tip of the snout.
(a) Nostrils directed vertically upwards. Belly and throat
smooth ; dorsal warts few ; lower surfaces of the limbs
brown, mottled and spotted with white . . P. major sp. nov.
(b) Nostrils dorso-lateral. Belly and throat, especially in males,
very coarsely granular ; snout of male with a distinct
digging edge ; dorsum very warty ; lower surfaces of the
limbs and the inner digits uniformly flesh-coloured
P. semimarmorata.
(2) Snout rounded, not prominent, the internarial space greater than
the distance from the nostril to the end of the snout.
(a) Tip of the fourth toe extending to the end of the snout or
beyond.
(i) Dorsum lighter than the flanks, the two colours sharply
defined ; lower surfaces dark brown with very large
yellow marblings ; a single, bold, orange bar across
the lower surfaces of femur, tibia and tarsus P. coriacea.
(ii) No sharp line of demarcation between the colours of the
dorsum and flanks,
(a) A bold red or orange marking on the top of the snout
to the level of the hinder borders of the eyes and
a coccygeal stripe of the same colour ; white spots
on the upper arm, in the groin and on the femoral
gland P. australis.
(|3) No red or orange mark embracing the synciput but, at
most, a light vertical bar on the tip of the snout ; a
faint yellow coccygeal stripe ; a red or yellow spot
on the upper arm . . . . .P. bibroni.
(b) Tip of the fourth toe extending to the shoulder or slightly
beyond the eye. General colour of the dorsal surfaces black
P. dendyi.
Pseudophryne guentheri Boulenger.
Pseudophryne bibroni (part) Giinther, 1858, Cat. Batr. Sal. Brit. Mus. : 46, 137.
Pseudophryne guentheri Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 279, pi. 17, fig.
2 (Type localities : — Swan River ; N.W. Australia ; Australia) ; Fletcher, 1898, Proc.
Linn. Soc. N.S.W., 22 : 680 ; Lucas & le Souef, 1909, Anim. Austral. : 285 ; Werner,
1914. Fauna S.W. Austral., 4, 10 : 419; Nieden, 1923, Das Tierreich, Anura I : 151,
fig. 198; Harrison, 1927, Rec. Aust. Mus. 15 : 281, figs. 1-5; Waite, 1929, Rept.
Amph. S. Austral. : 265 ; Barbour & Loveridge, 1929, Copeia, 180 : 130 ; Loveridge,
1935, Bull. Mus. comp. Zool. Harv., 78 : 36.
Pseudophryne brooksi Loveridge, 1933, Occ. Pap. Boston Soc. nat. Hist., 8 : 59 (Type locality : —
Manjimup, near Pemberton, W. Australia) ; idem, 1935, Bull. Mus. comp. Zool. Harv.,
78 : 35-
Snout bluntly rounded, short, as long as, or very slightly longer than the
eye ; canthus rostralis rounded ; loreal region slightly oblique ; distance from
nostril to tip of snout less than the internarial space, which is equal to the distance
from nostril to eye. Fingers moderate, with well-developed subarticular tubercles,
the first much shorter than the second, which is a little shorter than the fourth ;
third longer than the snout. Toes short, the third longer than the fifth, which is
rather stout and fleshy proximally ; subarticular tubercles well developed. Two
AUSTRALASIAN LEI'TODACTYI.IDAE 97
large transversely disposed metatarsal tubercles which, in adults, are compressed
and shovel-like. Tip of the fourth toe reaching between thr shoulder and the
nostril in adult females, the nostril or somewhat beyond the tip of the snout in
males and juvenile-,.
Skin with irregularly scattered small warts above, sometimes with a pair of
folds convergent from the upper eyelid towards the scapular region ; a large
parotoid and a similar inguinal gland ; no gland behind the thighs. Lower
surfaces smooth or faintly granular on the throat, posteriorly and laterally on
the abdomen and on the lower surfaces of the thighs.
Brown above, irregularly mottled with darker, and usually with more or
less distinct traces of the following large, light blotches : one covering the upper
surface of the snout from the anterior third of the upper eyelids, an oval one on
each scapular region, a A-shaped one on the middle of the back and a broad line
along the coccyx. These markings are more distinct in juveniles than in adults,
and in the former there is often a very thin red line from snout to vent and a
narrow light line along the hinder side of the thighs. Lower surfaces white,
more or less heavily spotted or marbled with brown.
Male with a vocal sac opening by a slit on each side of the tongue.
Length from snout to vent : <J 31 mm. ; 9 36 mm.
Distribution : West Australia.
The paratypes of P. brooksi Loveridge which have been examined all appear
to be young ones, and the differences in leg-length and the size of the metatarsal
tubercles which were believed to characterize that species seem to be due to their
immaturity.
Specimens Examined.
Pseudophryne occidentalis sp. nov.
Pseiutophryne australis (non Gray) Stirling A- Zietz, 1892, Trans, roy. Soc. S. .hist.. 16 : 170 .
Loveridge, 1935, Bull. Mus. comp. Zool. Ham., 78 : 31.
Holotype a male, number 1937.7.22.42 in the British Museum, collected .i
Bruce Rock. West Australia, by Prof. ('.. E. Nicholls in April, 1927.
novit. zool., 42, 1 7
g8 H. W. PARKER
Snout rounded, slightly prominent, a little longer than the eye ; canthus
rostralis rounded ; loreal region oblique ; internarial space a little greater than
the distance from eye to nostril and once and a half as long as the distance from
the nostril to the tip of the snout ; interorbital space once and two-thirds as broad
as the upper eyelid. Fingers rather short, bluntly rounded terminally, the first
shorter than the second which is shorter than the fourth ; third as long as the
snout. Toes similar to the fingers, the third longer than the fifth ; an oval inner
and a small, flat, rounded outer, metatarsal tubercle. Tip of the fourth toe
reaching the end of the snout when the hind limb is adpressed.
Skin regularly beset with small warts above, except on the snout ; lower
surfaces distinctly granular except on the pectoral region. A parotoid, but no
inguinal or femoral glands.
Dull brown above with obscure lighter markings covering the upper surface
of the snout, on the upper arm, the coccyx and the knee and heel. Lower surfaces
white with bold brown marblings on the belly ; infuscate on the throat and
limbs.
A vocal sac.
Length from snout to vent 26 mm. ; hind limb 26 mm.
Paratypes a male and a female (B.M. 1937.7.22.43-44) collected at the same
time and place as the type and a male and female (M.C.Z. 18301-18302) from
Burara, south of Kalgoorlie, also collected by Prof. Nicholls. These examples
agree well with the holotype except that the tip of the fourth toe may only reach
the centre of the eye. The gular region of the male is more markedly granular
than that of the female, there may be a pair of light, oval markings, one on each
side of the vertebral line about the middle of the vertebral column, and the throat
is marbled like the belly.
This is the West Australian species identified by Loveridge with P. australis
Gray. But, though the type of the latter was originally said to have come from
the Swan River, the specimen is undoubtedly conspecific with the eastern frogs
usually referred to by that name ; no name appears to be available for the
western frog.
Pseudophryne major sp. nov.
Pseudophryne bibroni (part) Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 278.
Holotype a female, number 67.3.4.56 in the British Museum, from Gayndah,
S.E. Queensland.
Snout conical, rather pointed and somewhat prominent, once and a half
as long as the eye ; canthus rostralis rounded ; loreal region oblique ; nostrils
directed vertically upwards ; internarial space equal to the distance between the
nostril and the tip of the snout, § the distance from nostril to eye. Interorbital
space once and a third as wide as the upper eyelid. Fingers moderate, with large
subarticular tubercles ; the first shorter than the second, which is shorter than
the fourth ; third a little longer than the snout. Toes moderate, with distinct
subarticular tubercles ; fifth shorter than the third ; a distinct inner, and a small
circular outer, metatarsal tubercle. Tip of the fourth toe reaching the tip of the
snout when the hind limb is adpressed.
Skin with a few linear warts above ; a series running from the upper eyelid
converges on the scapular region towards its fellow and then diverges again,
becoming indistinct posteriorly ; another short lateral series from behind the large
parotoid gland to the middle of the flanks. Smooth beneath except the lower
AUSTRALASIAN LEPTODACTYT IDAE 99
surfaces of the thighs which arc granular ; an oval gland on the hinder side of
the thighs close to the knee.
Brown above, some of the warts faintly darker ; loreal region darker ; brown
beneath with yellow spots, small beneath the limbs, large on the abdomen; a
faint light spot on the upper arm.
Length from snout to vent 36 mm. ; hind limb 36 mm.
The paratype is a male, bleached and soft, number 67.5. 13.31 in the British
Museum from Cape York. It agrees with the type except that the tip of the
fourth toe only reaches the loreal region. Vocal sacs, opening by a slit on each
side of the tongue, are present. Length from snout to vent 28 mm. ; hind limb
27 mm.
This species is obviously one of the bibroni-coriacea-semimarmorata group
but is distinguished by its much larger size, more pointed snout, more oblique
lores and colour.
Pseudophryne semimarmorata Lucas.
Phryniscus auslralis (part) Dumeril & Bibron, 1841, Erpet. Gen., 8 : 725, pi. 100, rig. 2.
Pseudophryne australis Fitzinger, 1843, Syst. Rept. : 32.
Pseudophryne bibroni [non Giinther) Krefft, 1865, Pap. roy. Soc. Tasmania : 17 ; Boulenger,
1882, Cat. Balr. Sal. Brit. Mus., ed. 2 : 278 (part) ; English, 1910, Proc. zool. Soc.
Lontl. : 631, pi. 51, fig. 7.
Pseudophryne semimarmorata Lucas, 1892, Proc. roy. Soc. Victoria, (2), 4 : 61, 63 (Type
localities : — Oakleigh ; Heidelberg ; Ringwood ; Narre Warren ; Waterloo ; Gram-
pians).
Pseudophryne bibroni var. semimarmorata Fletcher, 1898, Proc. Linn. Soc. N.S.W.,
22, 4 : 665.
Pseudophryne blanchardi Loveridge, 1933, Occ. Pap. Boston Soc. nat. Hist., 8 : 91 (Type
locality : — Millgrove, Victoria) ; idem, 1935, Bull. Mus. comp. Zool. Harv., 78 : 33.
Snout obtusely pointed, about once and a third as long as the eye, that of the
male strongly prominent, with a blunt horizontal ridge terminally, caused by the
development of a fibrous pad in front of the premaxillae ; canthus rostralis
obtuse ; loreal region oblique ; distance from nostril to tip of snout equal to ($$)
or greater than (<J(J) the internarial space, which is only very slightly less than the
distance from nostril to eye. Interorbital space once and a quarter the width
of the upper eyelid. Fingers rather short, with large, flat, subarticular tubercles,
the first much shorter than the second, which is a little shorter than the fourth ;
third as long as the snout ; outer metacarpal tubercle larger and more distinct
than the inner. Toes with well-developed subarticular tubercles, the outer
shorter than the third ; two Hat metatarsal tubercles, the outer the smaller. Tip
of the fourth toe of the adpressed hind limb reaching the eye in females or the
tip of the snout in males.
Skin thick and regularly beset with numerous warts above, of which some
of the largest may form a curved dorso-lateral series on each side from the pos-
terior corner of the eye. No clearly visible parotoid gland. Lower surfaces of the
female smooth except for the lower surfaces of the thighs proximally which are
markedly granular, and the posterior part of the abdomen and throat which are
more feebly so ; almost the whole of the lower surfaces of the body and the
proximal parts of the thighs of the male, coarsely granular. A large oval gland
occupies the distal half of the hinder side of the thighs.
Uniform brown above, or some of the waits somewhat darker. Lowei
surfaces yellow, heavily marbled and spotted with dark brown on the belly and
throat. The chin, lower surfaces of the femora, tibiae, tarsi, inner half of the
foot, lower surfaces of the arms and inner lingers always immaculate. This
100 H. W. PARKER
light colour may also persist over the whole of the gular region, on the upper
arm and over the backs of the thighs, including the femoral glands, but the anal
region is often pigmented. The glandular secretion is rusty brown or orange in
colour and marks the position of the femoral gland clearly. There may be a
light coccygeal stripe and, where the anal region is pigmented, a light line above
the vent.
In life the upper surfaces are olive-green, the sides blue-black ; the under-
surfaces of the limbs and throat are pale greenish yellow, greener near the axillae,
whilst the unpigmented areas of the feet and hands are flesh-coloured. The
belly is white with light olive-green marblings (Fletcher, 1892 : 63).
Breeding in March to May in Tasmania. Eggs laid singly under stones and
in similar places in depressions where temporary pools may form after rain.
Tadpole developing to an advanced stage in the egg in the absence of water, the
operculum being fully formed. Hatching very rapid when the mature eggs are
brought into contact with water. Tadpole with papillae at the sides of the mouth,
horny, denticulate mandibles, a single long series of upper labial teeth, and three
lower, of which the innermost is divided mesially and the lowermost the shortest.
Male with a vocal sac.
Length from snout to vent : <J 28 mm. ; $ 33 mm.
Distribution : Tasmania, Victoria and ? southern New South Wales.
Specimens Examined.
B.M.
This species, which for long was confused with bibroni, may be recognized
by its longer, more pointed snout, more warty dorsum, and the immaculate lower
surfaces of the limbs and inner digits. Loveridge was misled into describing P.
blanchardi by comparison with specimens from New South Wales which were
incorrectly determined as P. semimarmorata ; some of these (Loveridge, 1935 :
34) have been examined and found to be examples of Crinia sp., probably
C. signifera.
Pseudophryne coriacea Keferstein.
Pseudophryne (new species) Krefft, 1865, Pap. roy. Soc. Tasmania : 17.
Pseudophryne bibroni var. australis ? Krefft, 1867, Cat. Industr. Prod. N.S.W., Add. : 107.
Pseudophryne coriacea Keferstein, 1868, Arch. Naturgesch., 34: 272, pi. 6, fig. 15 (Type locality :
— Clarence River, N.S.W.) ; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 278 ;
Fletcher, 1890, Proc. Linn. Soc. N.S.W. , (2), S : 669, 675 ; idem, 1892, op. cit., (2), 7 :
8, 9 ; idem, 1894, op. cit., (2), 8, 1893 : 525, 529, 530 ; Nieden, 1923, Das Tierreich,
Anura I : 149 ; Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : 33.
Pseudophryne australis forma bibroni (part) Andersson, 1916, K. Svenska VetenshAkad.
Handl., 52, 9 : 13.
Snout rounded, not prominent, once to once and a quarter as long as the eye ;
canthus rostralis rounded ; loreal region oblique ; distance from nostril to the
end of the snout appreciably less than the intemarial space, which is very slightly
shorter than the distance from nostril to eye ; interorbital space a little greater
AUSTRALASIAN LEPTODACTYLIDAE IOI
than the width of the upper eyelid. Fingers rather short, with large subarticular
tubercles, the first shorter than the second which is shorter than the fourth ; a
large, round outer metatarsal tubercle, but the inner only faintly indicated.
Toes short, with well-marked subarticular tubercles, the fifth much shorter than
the third ; a rounded inner, and minute outer, metatarsal tubercle. Tip of the
fourth toe of the adpressed hind limb reaching the end of the snout in females
or well beyond this point in males.
Skin thick and glandular, shagreened, with numerous flat warts above ;
smooth below except the proximal parts of the thighs which arc granular. A flat
oval gland behind the thigh on its distal half.
Pale brown to cream above, this area sometimes spotted with darker or
lighter and clearly marked off from a dark lateral band which extends from the
loreal region to the groin and sometimes encloses a lighter area along the middle
of the flanks. Lower surfaces brown, with large white spots and vermiculations,
or white with a few large black spots ; sometimes a large light inguinal spot.
Limbs dark brown with a large light spot each on the posterior side of the upper
and lower arms, femur, lower surface of the tibia and tarsus and upper surface
of the foot.
Male with a vocal sac.
Length from snout to vent : <$ 28 mm. ; § 34 mm.
Distribution : Eastern New South Wales ; S. Queensland.
Specimens Examined.
B.M. 04. 7. 22. 23-24 ?. Hgr. Lake Macquarie. Krettt.
QO.7.28.2 <J Near Lismore, Richmond Fletcher.
River, N.S.W.
M.C.Z. 194 1 $ Clarence River, N.S.W. Cotype.
Mus. Leiden. 4255 j Queensland. (Godeffroy Mus.)
Pseudophryne australis (Gray).
Bombinator australis Gray, 1835, Proc.zool.SocL.ond., 3: 57 (Type locality : — Swan River,
Australia).
Phryniscus australis Gray, 1845, in Eyre, Jonrn. Exped. Centr. Austral., 1 ^07, pi. 2, fig. 1 ;
Giinther. 185S, Cat. Batr. Sal. Brit. Mus. : 45 ; Pseudophryne australis Fitzinger, 1861,
S.B. Akad. wiss. IVien, 42 1415; Krefft, 1865, Pap. roy. Soc. Tasmania : 17 ; idem,
1867, Cat. Indus tr. Prod. N.S.W., Add. : 107; Keferstein. 1868, Arch. Naturgesch., 34:
271 ; Boulenger, 1882. Cat. Batr. Sal. Brit. Mus., ed. 2 : 277 ; Fletcher, 1889, Proc.
Linn. Soc. N.S.W., (2), 4 : 364, 376 ; idem, 1890, op. cit., (2), 5 : 670, 671 ; idem, [894,
op. cit., (2), 8, 1893 : 530 ; idem, 1898, op. cit., 22 : 679 ; Lucas & le Souel 1909, hum.
Austral. : 284 ; Harrison, 1922, Aust. Zool., 3, 1 : 26, fig. ; Nieden, 1923, Das I ierreit h,
1 intra I : 148.
Phryniscus albifrons Dumeril & Bibron, 1854, ErpH. Gen., 9 : 413, pi. 100, fig. 3 (Type
locality not stated).
I'm udophrvne albifrons Loveridge, 1935, Bull. Mus. comp. Zool. Harv., 78 : },z.
? Bufonella crucigera Girard, 1853, Proc. Acad. not. Sci. Philad. : 424 (Type locality:
New Holland) ; Keferstein, 1868, Arch. Naturgesch., 34 : 272.
Snout rounded, scarcely prominent, once and a quarter as long as the eye ;
canthus rostralis rounded ; loreal region oblique ; Internaria] space greater than
the distance from the nostril to the tip of the snout and equal to the distance from
nostril in eye ; interorbital space once and a quarter the width of the upper eyelid.
Fingers rather short, with prominent subarticular tubercles, the first shorter than
the second which is a little shorter than the fourtii ; third as long as the snout ;
a large outer metacarpal tubercle, but the inner is inconspicuous. Toes with
large subarticular tubercles, the fifth much shorter than the third : a circular
inner, and a very small outer, metatarsal tuber le fipol the fourth toe reaching
102 H. W. PARKER
the end of the snout or, usually, beyond, when the hind limb is adpressed ; the
tip of the outer toe reaching the anterior corner of the eye or the nostril.
Skin smooth with a few rather inconspicuous warts above ; a glandular
thickening in the parotoid region. Smooth beneath, except the lower surfaces
of the thighs, posterior part of the belly and flanks which are granular ; a large
oval gland on the distal half of the hinder side of the thighs.
Dark brown or blackish above with a few irregular reddish or orange dots
tipping the warts and the following constant red or orange markings : a large
subtriangular marking on the top of the snout embracing nearly the whole of the
upper eyelids, and a coccygeal stripe. A band on the upper arm, a large inguinal
spot on the anterior aspect of the femur and a similar spot on the femoral gland
are white. Lower surfaces dark brown with a few very large white blotches on
the throat and abdomen, and similar but smaller spots on the inside of the tibia
and upper surfaces of the tarsus and foot. Hands and feet dark brown with the
tips of the digits white.
Male with a vocal sac.
Length from snout to vent : J 24 mm. ; $ 27 mm.
Distribution : New South Wales over the Hawkesbury sandstone beds
{fide Harrison, 1921 : 27).
Eggs, few in number (circa 20), laid throughout the year in an underground
nest close to the edges of streams. Development proceeds within the egg until
rudiments of hind limbs are developed ; when brought into contact with water,
through the overflowing of the nearby stream, hatching takes place rapidly, and
the remainder of the development to metamorphosis takes place in about four
weeks.
Specimens Examined.
An attempt has recently been made to transfer the name aastralis from this
well-known eastern species to the western form described in this paper as Occi-
dent alts on the ground that, since the type-locality was given as the " Swan
River," the specimen must have belonged to the western species. But the
original specimen, which agrees closely with Gray's description, is so obviously
conspecific with eastern examples that there can be little doubt that the locality
" Swan River " is either erroneous or the less known river of the same name in
New South Wales was intended.
Pseudophryne bibroni Giinther.
Phryniscus australis var. Dumeril & Bibron, 1841, Erpet. Gen., 8 : 725 (part), pi. 100, fig. 4 ;
idem, 1854, op. cit., 9:413 (part).
Pseudophryne bibroni Giinther, 1858, Cat. Batr. Sal. Brit. Mus. : 46 (part) (Type localities : —
Australia; van Dieraen's Land); Peters, 1863, Mber. Akad. Berlin: 235; Krefit.
1865, Pap. roy. Soc. Tasmania : 17 ; idem, 1867, Cat. Industr. Prod. N.S.W., Add. :
107 ; Steindachner, 1867, Reise Novara, Zool.. Amph. : 34, pi. 5, figs. 1,2; Giinther,
AUSTRALASIAN LEPTODACTYLIDAE I";
1867, Ann. Mag. nat. Hist., (3), 20 : s.t ; Keferstein, 1868, Arch. Nalurgesch., 34: 271 ;
Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 278 (part) . Fleto her, 1889, I't oc.
/.inn. Soc. N.S.W., (2), 4 : 365, 376 ; idem, [890, op. cit., (2), 5 : 669-673 ; idem, [89] .
op. cit., (2), 6 : 271-274 : idem, £892, op. cit., (2), 7 ■ - 8 . /,/ i» C894, "/>. ri/., (2), 8,
1893 : 526-528, 530; Lucas, 1892, Proc. roy. Soc. Victoria, 4 : 61 ; Boettger, 1894,
Denkschr. med.-naturw. Ges. Jena, 8 : no; Lucas & le Soucf, 1909, Anim. Austral, :
285 ; Andersson, 1913, K. Svenska VetenskAkad. Hand/., 52, 4 : 17. pi. 1, figs. 3, 4 ;
Harrison, 1922, .lust. Zool., 3 : 26; Nieden, 1023, Das Tierreich, Anura I : 14S, fi^.
195; Waite, 1929, Rept. Amph. S. Austral. : 265, fig. 102 ; Blanchard, 1929, Aust.
Zool., 5, 4 : 326; Loveridge, 1934, Pap. roy. Soc. Tasmania. 1933 : 60 ; idem, [935, Bull.
Mus. comp. Zool. Harv., 78 : 33.
Pseudophryne australis forma bibroni (part) Andersson, 1916, K. Svenska VetenskAkad.
Handl., 52 : 9, 12.
Snout rounded, not prominent, once and a quarter to once and a third as
long as the eye ; canthus rostralis obtuse ; loreal region oblique ; internarial
space longer than the distance from the tip of the snout to the nostril, but shorter
than the distance between the latter point and the eye ; interorbital space about
once and a quarter the width of the upper eyelid. Fingers with distinct sub-
articular tubercles, the first much shorter than the second which is a little shorter
than the fourth ; third as long as, or a little shorter than, the snout ; two distinct
metacarpal tubercles, the outer the larger. Toes with distinct subarticular
tubercles, the fifth shorter than the third ; two metatarsal tubercles, the inner
the larger. Tip of the fourth toe extending beyond the end of the snout, the tip
of the fifth reaching to between the eye and the nostril when the hind limb is
adpressed.
Skin with some small warts above which may be irregularly scattered, but,
more usually, some of them form regular series, of which the most constant is a
curved one on each side of the back from behind the upper eyelid, convergent
towards its fellow on the scapular region and then diverging again ; a second
series is less frequently present in a straight line from the posterior corner of the
eye along the flanks. Lower surfaces smooth except the throat which, in males,
may be feebly granular, and the proximal parts of the lower surfaces of the thighs
which are always distinctly granular.
Brown above with obscure darker markings of which the most constant are
a dark streak below the canthus rostralis, continued backwards behind the eye,
and a dark line following the curved line of dorsal warts. There may be a light
vertical line on the extreme tip of the snout and a rather obscure yellow line
along the urostyle, the latter joining a transverse light spot above the vent
which often extends on to the femoral glands. There is constantly a red or
yellow spot on the upper arm. Lower surfaces brown with white spots, which
in often so large on the throat and belly as almost to obliterate the dark ground-
colour.
Male with a vocal sac.
Length from snout to vent : <$ 26 mm. ; 9 27 nun.
Distribution : Southern Queensland, New South Wales, Vii toria, Tasmania
and South Australia.
Eggs laid in April or May, a hundred or more t<> the clutch, in irregular
masses close to permanent water. Developmenl proceeds up to the development
of the operculum within the eggs, but a long free larval stage of 5-6 months
follows hatching. The larva is characterized by three transparent lymph spaces
in the head. The mouth is bordered laterally by papillae, has horny mandibles,
two rows of labial teel li 111 the upper jaw, the inner divided, and three in the lower
jaw, all undivided and the outermost shortest.
The four cotypes of this species were not all conspecific. One of them (b)
was subsequently made a cotype of P. guentheri by Boulenger (1882 : 279,
spec, d), but the others appear to have been destroyed or lost before 1882 ; none
of them is listed under bibroni by Boulenger. What species they really belonged
to it is now impossible to say ; two of them were from Tasmania, in which island
semimarmorata occurs as well as the form usually known as bibroni. Dumeril
and Bibron's Phryniscus australis, which Giinther quotes as a synonym of bibroni,
also appears to have been a composite ; the specimen figured on plate 100, fig.
2, has the typical coloration of semimarmorata, whilst fig. 4 of the same plate,
described as a " var." of australis, appears to represent the form now usually
known as bibroni. The situation accordingly is as follows : there is only one
surviving cotype of bibroni ; if the name were to be fixed on that basis, the species
at present known as guentheri would in future have to be known as bibroni and
another name found for this latter, well-known species. What species may have
been represented by the other, lost cotypes we cannot tell ; they may have
belonged to two different species. Certainly the Phryniscus australis Dum. &
Bibr., considered by Giinther to be the same as his bibroni, was a composite of
two species and Boulenger's " bibroni " is also a mixture of the same two forms.
The name must obviously be applied to one of these two ; there is no method
whereby one or the other can be definitely excluded, for the original description
is not sufficiently detailed. But the obvious choice is the one which involves
least change, and accordingly the name bibroni is retained for the species figured
under that name by Steindachner (1869 : pi. 5, figs. 1 and 2) and semimarmorata
Lucas retained for the other.
Harrison (1927 : 286) has drawn attention to the existence of a very dark,
coastal race of this species, and suggested that P. dendyi Lucas may have been
based on it. But Loveridge (1935 : 34) has discovered a dark, montane form
on the borders of Victoria (from whence dendyi was described) and this ought,
AUSTRALASIAN LEPTODACTYLIDAE 105
probably, to bear the name. Until much more material is available it does not
seem advisable to use a trinomial for Harrison's melanic coastal race.
Pseudophryne dendyi Lucas.
1'f.cutlnpliiyiH1 t/i-iidyi l.wrns. iticji, Proc. roy. Soc. Victoria, (2) , 4 61, 62 (Type locality :
tipper Wellington River, N. Gippslandj ; Lucas & le Souef, 1909, .hum. Austral. : 285 ;
Nieden, IQ23, Das Tierreick, Anura I : 1411 ; Loveridge, 1935, Bull. Mus. cmiip. Zool.
Harv., 78 : 34.
Snout rounded, not prominent, once and a quarter to once and a third as
long as the eye ; canthus rostralis rounded ; loreal region oblique ; internarial
space greater than the distance from the tip of the snout to the nostril, about
equal to the distance from the latter to the eye ; interorbital space once and a
third as broad as the upper eyelid. Fingers with large subarticular tubercles ;
the first much shorter than the second, which is scarcely shorter than the fourth ;
third a little shorter than the snout ; a large outer, and a much smaller inner,
metacarpal tubercle. Toes with well-developed subarticular tubercles ; the fifth
much shorter than the third ; two small, subequal metatarsal tubercles. Tip
of the fourth toe of the adpressed hind limb reaching to the shoulder or beyond
the eye.
Skin quite smooth or feebly granular above ; a pair of curved folds from the
posterior corners of the upper eyelids faintly indicated. Smooth beneath except
the lower surfaces of the thighs and posterior part <>I the abdomen, which maybe
feebly granular. A feebly developed gland on the hinder side of the thighs
distally.
Black above with or without small white dots which may be aggregated to
form a supraciliary line and some patches behind and below the eye. A yellow
patch on the upper arm, and a yellow coccygeal stripe meeting a transverse line
of the same colour above the vent. Lower surfaces black with large white
blotches ; limbs black, white marbled below ; dorsal surface of the hand sometimes
white ; digits black-tipped and sometimes transversely banded with white.
Length from snout to vent ($) 32 mm.
Distribution : North Gippsland and the mountains of New South Wales.
Specimens Examined.
B.M. 1936.10.2.3 imm. <j> Hartley Vale, Blue Mts., Darlington.
N.S.W.
Austr. Mus. R.8973 $ Barrington Tops, N.S W.
Harrison (1927 : 268) has reported the type of this species to be lost.
MSS. reed. October 21, 1939.
NOVIT. ZOOL., 42, I
106 . H. W. PARKER
PLATE I.
Photographs showing reduction of the inner finger and compensating enlarge-
ment of the metacarpal in the genus Limnodynastes.
(a) Limnodynastes tasmaniensis <$. Digit of normal proportions and the inner
metacarpal shorter than the second.
(b) Limnodynastes salmini $.
(c) Limnodynastes fletcheri £.
(d) Limnodynastes peronii $.
(e) Limnodynastes peronii imm. $. An individual variant with the number
of phalanges reduced.
(f) Limnodynastes peronii, breeding $. Phalanges dislocated from the
metacarpal which pierces the skin.
NOVIT. ZOOL., 42, C94C
PARKER PLATE I.
/ >
j»%
Limnodynastes spp.
NEOTROPICAL NEOEMPHERIA IO7
NEOTROPICAL NEOEMPHERIA (DIPTERA,
MYCETOPHILIDAE)
By F. W. EDWARDS, F.R.S.,
Department of Entomology, British Museum (Natural History).
(With two plates and twelve text-figures.)
UNTIL recently comparatively little knowledge has been obtained regarding
the species of fungus-gnats (Mycetophilidae) occurring in South America.
The British Museum Expedition to the Patagonian Andes in 1926-7 resulted in
the acquisition of material of some 250 species of this family (including Sciarinae),
mostly undescribed. Since then efforts have been made to obtain further fungus-
gnat material for the Museum from the American Continent, because it was fell
that the value and interest of any report upon the Patagonian collection would
be greatly enhanced if comparison could be made with the fungus-gnat fauna of
other prarts of America. The Museum has been especially fortunate in obtain-
ing within the last few years extensive collections made by Dr. A. Dampf in
Mexico and Mr. F. Plaumann in Southern Brazil, including material of perhaps
500 species of fungus-gnats, few if any of which are the same as those represented
in the Andean collection. It therefore seems probable that the number of South
American species of this family will eventually be counted by thousands.
The genus Neoempheria, which forms the subject of this paper, does not
occur, so far as is known at present, in the Andean region, but is widely distri-
buted throughout the tropics. Numerous species occur in the Ethiopian and
Oriental regions, and a few in Europe and North America. Nearly all the species
have conspicuously patterned wings, these patterns exhibiting great diversity
but remaining constant in any one species. The synonymy of the genus is as
follows :
NEOEMPHERIA Osten-Sacken.
1863. Empheria Winnertz, Yerh. zoot.-bol. Ges. Wien, 13 : 783 (preoccupied by Emphtria
Hagen, 1856).
1878. Neoempheria Osten-Sacken, Smithson. misc. Coll., 16, art. 2.
1909. Myoomya subgenus Neoempheria Johannsen, in Wytsman's Genera Insectorum, 93
(Diptera, Mycetophilidae) : 45.
i'ii 1. Pleonazoneura Enderlein, Stettin, ent. Ztg., 72 : 156.
mi 1. Neurocompsa Enderlein, Stettin, ent. Ztg., 72 : 158.
1912. Neoempheria Johannsen, Hull. Maine Agric. Exp. Sta., 180 : 157.
Genotypes: Empheria and Neoempheria, Sciophila striata Meigen (Europe) ;
Pleonazoneura, P. johannseni Enderlein (S. Brazil) ; Neurocompsa, N. ornati-
pennis Enderlein (S. Brazil).
Only three neotropical species have hitherto been referred to this genus :
varipeknis F. Lynch Arribalzaga (Misiones Terr.), maculipennis Williston
(St. Vincent, W.I.), and apicalis Kert6sz (Peru). To these however must
be added Sciophila formosensis F. Lynch Arribalzaga (Chaco), neura
johannseni End. and its "variety" evanesceus Enderlein, and Neurocompsa
nrnatipennis Enderlein (Santa Catharina). Enderlein distinguished In- genera
Pleonazoneura and Neurocompsa from Neoempheria chiefly on account oi
tin- strong vein-like fold between K and M, and Neurocompsa further by
NOVIT. ZOOL., 42, I 9
ioS
F. W. EDWARDS
the possession of an extra cross-vein-like thickening between this fold and
the radius. Actually however, the vein-like fold is traceable with greater
or less distinctness in most if not all species of Neoempheria, and the additional
" cross- vein " is not even a constant specific character in N. ornatipennis. The
distinctions mentioned by Enderlein are therefore valueless. There are indeed
some other characters by which the group to which Enderlein's species belong
may be distinguished, but in my opinion these are hardly even of subgeneric
value.
I have now examined over thirty neotropical species of this genus, mostly
collected by Herr F. Plaumann at Nova Teutonia, Santa Catharina, Brazil.
Through the kindness of Dr. Kastner of the Stettin Museum I have been able to
re-examine Enderlein's type material collected by Liiderwaldt, and have ascer-
tained that not two but six species were represented among the nine specimens.
Figures and re-descriptions of all these are included in this paper.
Species of this genus, like those of the allied Mycomyia, are notable for the
complexity of the male genitalia and for the very great differences often
found in these organs in species which are obviously very closely allied,
though it may also occur that species which on external features seem obviously
distinct show only small differences in the genitalia. In examining genitalia of
Neoempheria in the dry state it should be borne in mind that the whole hypo-
pygium is usually twisted or inverted, so that dorsal structures may appear
lateral or ventral and vice versa ; it is however easy to decide which is the true
dorsal surface by the presence on it of the anal segment. In the accompanying
figures the dorsal aspect of half of the hypopygium is shown on the left, the
ventral aspect on the right. The following parts may be recognized in most
or all species : (i) the large ninth tergite (t), divided nearly to the base into two
halves and often with the tip of each half produced into a long process (pt) of
varied form ; (2) the anal segment (as), composed of paired cerci and an unpaired
but sometimes divided sternite, always pubescent ; (3) the ninth sternite (s),
divided to the base into two lobes which are almost always completely bare and
membranous and may be difficult to see in the mount unless the specimen has
been stained ; (4) a pair of inner styles (is) lying beneath the tergite but articu-
lated with the sternite, never very hairy but often bearing a few or rather numerous
black spines or variously toothed ; (5) a pair of outer styles (os) articulated with
the sternite ventro-laterally, usually clothed rather densely with longish hair ;
(6) one or two pairs of processes of the sternite (ps), usually bare or only finely
pubescent ; (7) a pair of large fleshy parameres (pm) lying within or between the
lobes of the sternite, always completely bare, usually pale in colour and appear-
ing as broad flat plates in side view in the dry specimen ; (8) some small structures
connected with the aedeagus, visible only in the mounted specimen. In addition
to the structures of the hypopygium proper, the eighth tergite (&) and eighth
sternite (8s) show noteworthy specific differences in shape and in the number
of hairs (if any) on their posterior margins. The females as well as the males
show very considerable differences in the genitalia, but I have not investigated
these.
Differential characters applicable to both sexes are to be found not only in
the markings of the thorax and wings, but also in venation, trichiation of the
wing-veins, and chaetotaxy of the head. All the neotropical species known to
me possess only two strong bristles on the scutellum, in this respect differing
from the European species (including the genotype), which have four or more
such bristles. The neotropical species may be classified in about eight groups,
NEOTROPICAL NEOEMPHERIA
109
based on the length of vein Sci, position of fCu (base of cubital fork), length of
ocellar bristles (the pair immediately behind the black ocellar spot), presence or
absence of minute setulae (macrotrichia) on Sc and on branches of media and
cubitus, especially M2, and nature of markings (if any) of thoracic pleurae.
(The characters of the groups as defined below are not repeated in the specific
descriptions which follow.)
Group A.
Sc reaching almost to the end of the small cell, which is only about half as
long as stem of median fork ; Sc2 transverse and well beyond base of Rs. Costa
produced only a little beyond tip of R5. fCu before base of Rs. Sc setose on
more than its distal half ; Mi, M2, Cu, Cui and Cu2 all setose for the whole of
their length. Costal and basal cells all dark. Ocellar bristles quite short.
Thorax unicolorous. Halteres yellow.
The species of this group approach nearer to the genotype (N. striata Mg.)
than do those of any of the other South American groups of the genus.
Neoempheria neivai sp. n.
<J. Head yellowish except for the black ocellar spot ; antennal flagellum
blackish ; palpi largely dark. Thorax light yellowish on sides, rather more
reddish-yellow above, with black hairs and bristles, but quite unmarked ; a
Text-fig. i. — Hypopygia of Neoempheria : a. neivai; b, bipectinata.
pair of bare stripes between acrostichal and dorso-central hairs. Abdomen
with tergites 1, 2, 4 and 7 yellow, .;, 5 and 6 black; venter yellowish. Legs
yellow, including whole of hind femora ; tibiae and tarsi darkened. Wings
(PI. I, fig. l)with tip and whole anterior margin, including basal cell, dark :
also with a large dark patch below tip of Cu2. Hypopygium (text-fig. 1, a)
110 F. W. EDWARDS
very different in structure from that of all the other South American species
examined ; a notable feature is the apparent absence of the eighth sternite,
unless it is represented by a narrow bare strip of chitin fused on to the hypopygium.
2. Differs from <J in colouring of abdomen and legs : tergites 2-6 are entirely
black, only 1 and 7 being yellow ; hind femora almost entirely blackish, con-
trasting with the others. Thorax and wings as in <J.
Wing-length 6-7 mm.
Brazil : Nova Teutonia (Plaumann), 1 cJ (type), 2 2. British Guiana :
Kutari Sources (G. A. Hudson), 1 <J.
At the suggestion of Father Borgmeier this fine species is named in honour
of Dr. Arthur Neiva.
TV. neivai corresponds in many respects with the description of Sciophila
formosensis F. Lynch Arribalzaga,1 and there seems to be little doubt from the
description that Lynch's species is actually a N eoempheria of this same group.
Lynch's definite statements as to the abdominal colouring of his type (a male from
Formosa, Argentine Chaco) seem however to show that it is a different species ;
it is said to have the abdomen fuscous with the last two and part of the fifth
tergites yellowish. Lynch also does not mention any dark area below the tip
of Cu2.
TV. apicalis Kertesz,2 described from one female from Callanga, Peru, must
also belong to this group of the genus, but according to the description differs
from both TV. neivai and TV. formosensis in having three darker stripes on the
mesonotum and in details of abdominal markings. Kertesz, like Lynch, does
not mention the presence of a dark area below Cu2.
Group B.
Sc reaching to or beyond middle of the small cell, which is somewhat shorter
than stem of median fork ; Sc2 transverse and placed at base of Rs. Costa
distinctly produced. fCu before base of Rs. Sc setose on distal half ; Mi, M2
and Cui setose except narrowly at base, Cu and Cu2 setose throughout. Wing-
markings conspicuous, but costal and basal cells clear. Ocellar bristles short.
Pleurae with longitudinal dark stripe. Halteres with dark knob.
Neoempheria bipectinata sp. n.
(J. Head mainly yellowish, darker behind, especially towards sides ; ocellar
spot black. Antennae brownish, scape lighter ; seta on second segment rather
long. Palpi entirely black. Thorax mainly brown above, without darker or
lighter stripes, sides of mesonotum and most of pleurae yellowish ; a dark brown
band across base of postnotum, continued along each side across base of pleuro-
tergite and upper margin of stemopleura to base of front coxa, leaving anepi-
sternite entirely pale. Abdomen with tergites 1 and 7 yellowish, 2-4 each
yellowish with a median dark stripe connected with a rather narrow dark posterior
band, 5 and 6 mainly dark. Legs mainly yellowish but hind femora darker.
Wings as figured (PI. I, fig. 2) ; basal cell and base of cubital fork clear. CuP
and An both strong. Hypopygium (text-fig. 1, b) remarkable for the pair of
comb-like structures at base of tergite and the form of the anal segment.
1 1892, Bol. Acad. Cient. Cordoba, 12 : 416.
2 1909, Ann. hist. nat. Mils. Hung., 1 : 140.
NEOTROPICAL NEOEMPHERIA
III
9- Resembles <$.
Wing-length 4-5-5-5 mm.
Brazil : Nova Teutonia (Plaumann)
1 6" (type), 1 ?.
Group C.
Sc ending above base of Rs ; Sc2 oblique and practically at tip of Sc. R4
above or just beyond base of median fork, the small cell long. Costa distinctly
produced. fCu before base of Rs. M2 turned slightly downwards at extreme
tip. Arrangement of setae on veins differing in the two sexes ; in 3 Mi, M2 and
Cui are setose except at base and Cu2 is entirely setose ; in $ Mi is setose only
at tip, M2 bare or with a few setae at tip, Cui bare, Cu2 setose only at base. In
both sexes Sc is setose on most or all of the distal half, Cu sparsely setose. Wings
highly adorned ; basal cell with a dark spot at or near base. Ocellar bristles
short. Pleurae with oblique dark stripe, extending from base wing across anepi-
sternite to base of front coxa. A pair of bare stripes between acrostichal and
dorso-central hairs. Halteres with dark knob.
Neoempheria spinosa sp. n.
cj. Head largely yellowish above, but dark brown behind and towards sides ;
also darkened between the black ocellar spot and base of antennae. Antennae
with scape yellowish, flagellum black. Palpi wholly black. Thorax largely
brown above, but with the bare stripes pale ; sides of mesonotum broadly yellow,
Text-fig. 2. — Hypopygium of Neoempheria spinosa.
also most of pleurae ; scutellum dark above ; postnotum broadly dark at base,
somewhat lighter posteriorly ; pleurotergites dark brown on posterior half, The
oblique dark pleural stripe occupies anterior parts of anepisternite and sterno-
pleurite and extends rather faintly over front coxa. .Middle and hind coxae
entirely yellowish; hind femora somewhat darker than the others, but not
conspicuously so. Abdomen with tergites 1 and 2 largely pale, but both with a
112 F. W. EDWARDS
median dark longitudinal stripe, sometimes reduced to spots ; 3 mainly dark,
with a moderately large pale spot towards each side at base, these spots leaving
posterior and most of lateral margins broadly dark but just reaching sides at
base ; 4 mainly pale, but dark in middle above, the dark area broad basally
but not or scarcely reaching posterior margin, lateral margin also dark ; 5 and 6
mainly dark ; 7 pale (but retracted). Wings as figured (PL I, fig. 7) ; notable
features in the markings are the clear costal cell with a small dark area at its
base, dark spot near but not quite at base of basal cell, and more intense darken-
ing on inner edge of dark area at tip ; dark area in cell Cui uniform in tint ;
pale area below Cu2 crossing CuP. Hypopygium (text-fig. 2) very large, with
each half of tergite split into two parts, an upper finger-like part and a larger
lower part with many strong black spines at tip.
?. Resembles $ except in trichiation of veins, as noted in group diagnosis.
Seventh tergite large and mainly pale, but with its posterior margin dark ; setae
on its posterior corners not specially numerous or strong. Hind margin of
seventh sternite with some longish black hairs. Eighth sternite large, with
uniform sparse pubescence.
Wing-length 4-5-5-5 mm.
Brazil : Nova Teutonia (Plaumann), 8 <J (including type), 2 $.
Neoempheria brevicauda sp. n.
$. Very similar to TV. spinosa, with which it agrees in having the post-
notum darker at the base and lighter distally, posterior coxae entirely yellow,
dark area in cell Cui uniform in tint, and pale area below Cu2 crossing CuP.
Trichiation of wings as in N. spinosa 9. Differs from N. spinosa chiefly as
follows : Wings (PL I, fig. 8) with the pale area at base of median fork smaller
and dark area below Cu2 differently shaped. Abdomen with the yellow spots
on tergite 3 larger. Tergite 7 shorter and more extensively dark. Sternite 7
with few or no long hairs on posterior margin. Sternite 8 shorter, with a loose
tuft of dark pubescence near base on each side. The lateral flaps below tergite
7 (of uncertain homology) are smaller and much less pubescent than in N. spinosa.
Brazil : Nova Teutonia (Plaumann) , I ?.
Neoempheria simplex sp. n.
<$. Closely resembles N. spinosa except in the following details : dark
markings of mesonotum more intense, therefore contrasting more with the pale
parts. Postnotum uniformly blackish. Wings (PL I, fig. 10) with the sub-
apical pale area smaller ; dark area in cell Cui rather more intense proximally ;
pale area below Cu2 inconspicuous and not crossing CuP. (Abdomen lost after
mounting hypopygium ; markings probably agreeing with 9 as described below.)
Hypopygium (text-fig. 3, a) very different in structure, the tergite narrower,
undivided, with a small tuft of hairs but no spines at tip.
9. Resembles <J in markings of thorax and wings. Abdomen with tergites
1 and 2 more extensively dark above than in spinosa, 3 and 7 nearly all dark.
Structure of ovipositor quite different from that of N. spinosa and brevicauda ;
tergite 7 with many long dark hairs on its posterior margin and with about 6
long stout bristles on the part that is folded under on each side. Trichiation of
wings as in N. spinosa 9-
Wing-length about 5 mm.
NEOTROPICAL NEOEMI'HERIA
113
Brazil : Nova Teutonia (Plaumann), 1 g (type), 1 $.
A second ? (of which the wing is figured, PI. I, fig. 9) differs from the first
in having the subapical pale area of the wings larger, thus more resembling N.
spinosa ; the structure of its ovipositor however is the same as that described
above.
Neoempheria bilobata sp. n.
<$. Very much resembles the last three species, but differs from all of them
in having a dark streak running almost whole length of hind coxa. Postnotum
all dark. Tergites 1 and 2 each with only a small dark area above, 3 extensively
pale at sides. Wings (PI. I, fig. 11) marked similarly to those of N. simplex,
Text-fig. 3. — Hypopygia of Neoempheria : A, simplex ; b, bilobata.
except that the subapical pale area is differently shaped. Hypopygium (text-Jig.
3, B) quite different from that of N. spinosa or simplex : tergite bilobed at tip,
with about three stout bristles on inner lobe.
Wing-length about 4-5 mm.
Brazil: Nova Teutonia (Plaumann), 1 J.
Neoempheria borgmeieri sp. n.
? (?). Head yellow ; ocellar spot, palpi and flagellum black. Ocellai bristles
reaching about half-way from their insertion to base of antennae. Thorax
mainly yellow; mesonotum with three narrow parallel dark brown stripes
running the whole Length and connected <>n front margin, our along the inter-
acrostichal space, the other two sublateral, no truer oi additional stripes along
the lines of dorso-central hairs ; lateral stripes a little wider than the medial one.
Pleural markings as in other species "l Group C ; the oblique stripe well marked,
114 F. W. EDWARDS
pleurotergites indefinitely darkened posteriorly. Posterior coxae all yellow.
Wings with the markings more suffused than in other species of this group ;
basal cell more than half dark ; dark area in middle extending distinctly into
base of median fork, which it does not in the other species. Mi setose on distal
third, Cui on distal half ; M2 bare ; Cu2 setose throughout. R4 inwardly
oblique instead of transverse as is normally the case in this genus.
Wing-length barely 4 mm.
Brazil : Bom Retire, S. Cath., Prade, 20.L29 {Borgmeier), 1 $ (?).
I have described this specimen in spite of its damaged condition because its
colouring is quite distinct from that of all other Neotropical species of the genus
known to me. The markings and trichiation of the wings will distinguish it from
any species of Groups D or E.
Group D.
Venation as in Group C. Trichiation of veins alike in the two sexes ; Sc
bare or almost so ; Cu setose except at base ; Mi, Cui and Cu2 setose throughout,
but M2 completely bare. Branches of median fork rather widely divergent.
M2 tending to be turned upwards at tip. Wing-markings conspicuous ; basal
cell dark at base. Two of the bristles immediately behind ocelli are long, extend-
ing forwards almost or quite as far as base of antennae ; a second pair of ocellar
bristles also somewhat longer than usual, but not nearly so long as the median
pair. Bare stripes on mesonotum as in Group C. Pleurae with an oblique
dark stripe from wing-base to base of front coxa ; other dark pleural markings
sometimes present but separate from this stripe ; pleurotergite either entirely
pale or with the anterior margin darkened instead of the posterior as in Group C.
Knob of halteres dark (except in TV. puncticoxa).
Neoempheria plaumanni sp. n.
cJ. Head light brownish, ocellar spot black as usual. Antennae with scape
yellowish, flagellum brown above, yellowish beneath on basal half or more.
Palpi entirely black. Thorax brown above, darker (but not conspicuously so)
along the stripes of acrostichal and dorso-central bristles and at the sides of the
dark area ; sides of mesonotum rather broadly pale ; pleurae with yellowish
ground, the oblique stripe broad and distinct ; a dark brown band at base of
postnotum extends on each side across the suture between pleurotergite and
anepisternite. Abdominal tergites largely dark above, yellowish at sides, fourth
more extensively so but with a considerable dark area. Pleural stripe extending
on to front coxa ; hind coxa also with a dark streak outwardly. Wings as
figured (PI. II, fig. 13) ; costal margin mainly yellow ; tip of marginal cell
broadly dark ; brown spot over tip of Sc small ; a brown spot fills basal third
of upper basal cell, and a larger dark brown area fills base of cell R5 ; a brown
patch connects R5 with Mi at about middle of fork. Halteres with dark knob.
Hypopygium (text-fig. 4, a) elongate but not very stout ; tergite ending in a
long black finger ; outer style dark, widened beyond middle ; inner style entirely
pale, bent but simple.
?. Resembles <J, but thorax usually darker above.
Wing-length, 5-6 mm. ; body-length 6-7 mm.
Brazil : Nova Teutonia (Plaumann), numerous specimens of both sexes,
including type <$.
NEOTROPICAL NEOEMPHERIA
"5
Neoempheria plaumanni var ?.
$. Resembles N. plaumanni in most respects, including the presence of a
dark streak on outer side of hind coxa, but differs from that species and the other
two described below in that the wings (PL IJ, fig. 14) have no dark areas in
base of cell R5 and immediately below fCu.
Brazil : Nova Teutonia (Plaumann) , 1 ?.
Neoempheria flavicoxa sp. n.
cJ. Very similar to N. plaumanni, except that the hind coxa is entirely
yellow ; the mesonotal markings might be described as five dark stripes separated
Text-fig. 4. — Hypopygia of Neoempheria : a, plaumanni ; B, flavicoxa
by four yellow ones, all the dark stripes connected in front and the middle three
convergent behind ; the stripes are rather more definite than in Ar. plaumanni.
Wings (PI. If, fig. 15) almost as in plaumanni, but dark area below Cu rather
differently shaped. Hypopygium (text-fig. 4, u) resembling that of N. plau-
manni, but differing in various small details, notably in having the inner style
blackened and truncate at the tip.
$. Resembles J.
Brazil : Nova Teutonia {Plaumann), 1 S (type). 2 $.
Neoempheria puncticoxa sp. n.
(J. Very similar to N. plaumanni and N. flavicoxa, but differs from both in
having the last palpal segment mainly yellow, the postnotum with a dark spot in
middle at base instead of a dark basal band, front coxa with a small sepai.n.
dark spot in middle in front, and knob ot halteres almosl entirely yellow. Hind
coxa all yellow as in N. flavicoxa. Mesonotum with five dark stripes as in the two
n6
F. W. EDWARDS
allied species, but the outer pair of stripes less sharply defined outwardly. Hypo-
pygium (text-fig. 5) resembling that of TV. plaumanni, but differing in having the
inner style forked and in some other details.
? ?. Front coxa as in <J> but postnotal dark band almost complete (palpi and
halteres broken).
Brazil : Nova Teutonia (Plaumann), 1 c? (type), 1 $.
Neoempheria mulleri sp. n.
c?. Head yellow above, dark behind ; palpi black as usual ; scape yellow,
flagellum blackish. Ocellar bristles practically reaching base of antennae ; a
longish bristle on second antennal segment. Thorax in darker specimens largely
blackish-brown above, with a wedge-shaped median yellow stripe which is divided
by an indistinctly darker line along the acrostichal hairs, and towards each side
with a small oval yellow spot ; in lighter specimens the sublateral pale spot
may be .longer. Scutellum pale yellowish. Sides of mesonotum yellowish but
less broadly so than in some species ; ground-colour of pleurae yellow, but dark
markings more extensive than in most species, the usual oblique stripe being less
definite on this account ; whole anepisternite dark, also a large part of sterno-
pleura, but upper part of latter pale. Abdomen with large yellow areas on sides
of tergites ; tergite 2 with its posterior margin rather broadly yellow ; tergite 4
narrowly yellow at base and broadly so posteriorly, leaving a rather narrow
dark band, which is widest in middle ; 5 and 6 with large yellow lateral triangles
which do not quite meet in middle of posterior margin. Front coxae mainly pale
except at base ; middle and hind coxae with or without small dark marks out-
wardly. Hind femur darkened on distal half or more, but pale at base. Wings
(PL II, fig. 16) with costal and marginal cells alternately yellow and brown, a wide
brown area before tip of Sc ; basal half or more of basal cell dark ; a dark cloud
adjoins Mi in middle of cell R5, but does not nearly reach up to R5 as it does in
N. plaumanni and related species. Sc bare. Hypopygium (text-fig. 6, c)
resembling that of TV. plaumanni in that the tergite has a blackened finger-like
process at the tip, and the outer style is broad and dark, but the sternite is more
developed, forming a pair of large pale bare lobes. Inner style slender, darkened
at tip, with a single small tooth near middle ; inner processes of sternite with
bifid tips.
$. Resembles J. Sc usually with some setae at tip.
Wing-length 4-5 mm.
Brazil : Nova Teutonia {Plaumann) , 4 cJ (including type), 3 $.
Dedicated to the pioneer Brazilian Dipterist Fritz Muller.
This species is closely related to TV. varipennis F. Lynch A. (Misiones Terr.)
and to TV. maculipennis Will. (St. Vincent, W.I.), all three having very similar
wing-markings, but differing in other details.
Neoempheria varipennis (F. Lynch Arribalzaga).
1892. Empheria varipennis F. Lynch Arribalzaga, Bol. Acad. Cienc. Cordoba, 12 : 423.
A series of specimens in the Plaumann collection in the British Museum
agree with the description of TV. varipennis, and show the following differences
from the new species described above : Mesonotum not so dark, and with the
sublateral pale areas forming short stripes rather than spots. Fourth abdominal
tergite more extensively dark, the dark area nearly reaching posterior margin
NEOTROPICAL NEOEMPHERIA
117
Text-fig. 5. — Hypopygium of Neoempheria puncticoxa.
Text-fig. 6. — Hypopygia of Neoempheria t, varipennis ; B, latui ; c, miUleri ; D.maculi-
pi inns ; k, maculipcnnis ssp. bradleyi.
Il8 F. W. EDWARDS
in middle. Front coxa largely dark. Hypopygium (text-fig. 6, a) with tip
of inner style expanded and blackened, and with some other small differences.
Neoempheria maculipennis Williston.
1896. Neoempheria maculipennis Williston, Trans. Enl. Soc. London, 1896 : 262.
Three males (type and paratypes) of N. maculipennis in the British Museum
show the following differences from N. miilleri sp. n. Thorax paler, striped as in
N. varipennis ; sternopleura not at all darkened. Hind femora entirely yellow.
Fifth and sixth abdominal tergites, as well as the fourth, with complete pale
apical bands. Wings (PL II, fig. 18) with a small pale area in base of median
fork, otherwise as in miilleri and varipennis. Hypopygium (text-fig. 6, d) with
finger-like process of tergite differently shaped ; inner process of sternite simple,
not bifid ; inner style with two small teeth at some distance apart near middle.
Neoempheria maculipennis subsp. bradleyi subsp. n.
c?. Mesonotum more shining than in the types of maculipennis and rather
differently marked ; a small dark area above each shoulder connected with the
usual sublateral dark stripes and with very narrow dorso-central dark stripes,
but no trace of acrostichal dark stripe, so that the resulting mesonotal pattern
might be described as three rather broad yellow stripes narrowly outlined with
dark. Tergite 5 narrowly pale-bordered, 6 dark. Hind femora rather broadly
dark-tipped, but hind coxa entirely yellow, as in maculipennis. Sternopleura
dark in middle. Wings as in maculipennis, with a small pale area in base of
median fork. Hypopygium (text-fig. 6, e) very much as in typical maculipennis,
the parameres having the same rather unusual form, but inner style rather
differently shaped and with only one tooth ; eighth tergite with a longer row of
setae.
Peru : La Sombra, Putumayo District (J. Chester Bradley), 1 J, 22.viii.1920.
Type in Cornell University Collection.
Neoempheria sp. n. ?
?. Resembles N. miilleri and varipennis in most respects, but differs from
them and resembles the types of maculipennis in having hind femora entirely
yellow ; differs from all three in that the yellow mesonotal stripes are more
distinct, all reaching back to the scutellum, and the dark area above Mi extends
narrowly to R5, somewhat as in N. plaumanni and related species, though this
dark area (and more obviously the pale area which precedes it in cell R5) is much
more oblique than in those species. Fourth tergite with large apical lateral
yellow spots, but dark area reaching posterior margin in middle ; 5 and 6 with
small apical lateral yellow spots.
Wing-length 5 mm.
Brazil : Nova Teutonia (Plaumann) , 1 $.
This specimen may be a variation of N. varipennis or miilleri, but is more
probably a distinct species.
Neoempheria lanei sp. n.
cj. Head as in the allied species, but thorax much darker. Mesonotum
mainly blackish brown, the usual three or four paler stripes barely distinguishable
and the dark colour extending to the sides, leaving only a small yellow mark on
NEOTROPICAL NEOEMPHERIA IIQ
the margin some distance in front of the wing-base. Anterior pronotal lobes
dark brown instead of yellow as in all the other species of this group. Sterno-
pleura almost entirely dark, and pleuro-tergite more extensively dark than usual.
Abdomen with complete though rather narrow pale apical bands on tergites 2,
4 and 5. Front coxae almost all blackish ; posterior coxae also largely dark.
Hind femur blackish on distal two-thirds ; middle femur with a dark band before
the tip. Wings (PI. II, fig. 17) with the dark markings on the same plan as
in the last three species, but less extensive ; dark cloud in middle not involving
Mi or Cui. Hypopygium (text-fig. 6, b) very like that of maculipennis, but
inner style blackened and truncate at tip, with one blunt prominence near base.
Wing-length 4 mm.
Brazil : Juquia, Sao Paulo (/. Lane), I rj (type), presented to British Museum
by the collector.
Group E {Neurocompsa Enderlein).
Venation and trichiation of wings as in Group 1), except that Sc is usually
setose for a short distance at tip, and Sci is usually longer. Ocellar bristles
long, as in Group D, though not always reaching to base of antennae. Bare
stripes on mesonotum as in Groups C and D. Pleural markings as in Group D,
but the oblique stripe sometimes faint. Wing-markings less conspicuous than
in Group D ; basal cell entirely clear except for the cloud over base of Rs. Halteres
with knob mainly or all dark (except in N. liiderwaldti).
Neoempheria ornatipennis (Enderlein).
1911. Neurocompsa ornatipennis Enderlein, Stettin, ent. Ztg., 72: 159.
In addition to Enderlein's type $ collected by Liiderwaldt, I have examined
two males of this species collected by Plaumann at Nova Teutonia, and another
collected by Lane, probably near Sao Paulo.
The species is well distinguished from all others of the genus by the abrupt
sinuosity of the fold between R5 and M, with a dark ridge or thickening connect-
ing on to R5 at the top of the bend. Apart from this peculiarity N. ornatipennis
shows no very striking differences from others placed with it in this group. The
ocellar bristles reach the base of the antennae. Flagellum dark almost to the
base. Mesonotum mainly light brown, with three darker brown stripes connected
by a narrow dark band on front margin ; the sublateral pair of stripes broader
than the median stripe, which is very narrow and fades out posteriorly. Oblique
stripe of pleurae distinct. A narrow transverse brown band at base ol po 1
notum and extending on each side over base ol pleurotergite and on posterior
edge of anepisternite. Abdomen with tergite 1 dark above, yellow at sides, 2
rather broadly pale at sides and on posterior margin, 3 yellow at sides at base
but broadly dark posteriorly, 4 yellow with a dark area at base in middle, 5 and
6 mainly dark. Wings (PI. II, fig. 19) with a rather large pale area in base of
median fork ; vein Mi somewhat concave above in middle. All coxae yellow ; hind
femora somewhat darkened. Hypopygium (text-fig. 7. A.) rather peculiai in
that the two halves of the tergite are connected at a point instead "i bj a trans-
verse band ; inner style with a few black spines at tip; terminal finger ol tergite
short ; outer style rather broad. Eighth tergite with about >i\ setae in middle
of distal margin.
120
F. W. EDWARDS
Text-fig. 7. — Hypopygia of Neoempheria : a, ornatipennis ; b, evanescens.
Text-fig. 8. — Hypopygia of Neoempheria : a, kcislneri ; B, subdavata.
NEOTROPICAL NEOEMPHERIA 121
Neoempheria evanescens (Enderlein).
1911. Pleonazoneura johannseni var. evanescens Enderlein, Stettin, ent. Ztg., 72: 158.
cJ. In most respects very similar to N. kcistneri and other species described
below, but is rather smaller and with fainter wing-markings ; pleural stripe
present as in other species of this group, and abdominal markings as in N. kcistneri.
Features which may distinguish it from other species of the group are the faintness
of the acrostichal as well as the dorso-central dark stripes of the mesonotum and
the complete absence of setae on vein Sc. Hypopygium (text-fig. 7, b) quite
distinctive ; tergite hairy at tip, without any definite process ; inner style of
unusual form ; outer style rather short ; lobes of sternite with a small finger-like
projection.
Brazil : Santa Catharina (Liiderwaldt) , 1 J (lectotype, in Stettin Museum).
Neoempheria kastneri sp. n.
<?. Colour and markings almost exactly as in N. omatipennis except that
the hind femora are not in the least darkened, but are yellow like the others ;
second abdominal tergite with the dark area above rather longer, nearly reaching
posterior margin ; pale area in base of median fork rather smaller ; wings with
fold between R5 and M straight, Mi also quite straight. Hypopygium (text-fig.
8, a) : chiefly distinguished by the form of the inner style, which has a greatly
enlarged tip (this collapses in drying and maceration is needed to restore its
true form) ; outer style broadish ; the slender terminal process of tergite almost
as long as tergite itself. Anal segment with long setae. Eighth tergite with four
setae.
Wing-length 4 mm.
Brazil : Santa Catharina (Liiderwaldt) , 1 <$ (type) in Stettin Museum,
formerly a paratype of PI. johannseni End. Nova Teutonia (Plaumann), 3 $ in
British Museum.
Dedicated to Dr. Kastner of Stettin town Museum, in appreciation of his
kind assistance during my visit there in 1933, and subsequently.
Neoempheria subclavata sp. n.
<J. Closely resembles N. kastneri in all external features , hypopygium
(text-fig. 8, b) also similar in type, having the tergite largely bare, with one or
two long bristles at side in middle and with long terminal process ; anal segment
large, with long setae and with the sternite notched. Differs from N. kastneri
in having the outer style much longer and more slender, and the inner style less
swollen at the tip and without any black spines. Eighth tergite with only two
setae.
Brazil : Nova Teutonia (Plaumann), 2 o (including type).
Neoempheria luderwaldti sp. n.
tJ. Closely resembles N. kastneri in all external features, except that the
halteres are almost entirely yellow; wings as in PI. II, fig. 20. Hypopygium
(text-fig. 9, a) quite different : terminal process of tergite with a wide enlargement
before tip ; inner style divided almost to base into two arms, inner arm slender
and pointed, outer arm ending in a tuft of black spines ; outer style slendei
Anal segment small, with short setae. Eighth tergite with two setae.
122
F. W. EDWARDS
Brazil : Santa Catarina (Luderwaldt) , i <J (type) in Stettin Museum, formerly
a paratype of PL johannseni var. evanescens End. Nova Teutonia (Plaumann) ,
I tj in British Museum.
Neoempheria enderleini sp. n.
(J. Closely resembles N. kdstneri in nearly all respects, but yellow areas on
sides of third tergite occupying less instead of more than half its length, and
Text-fig. 9. — Hypopygia of Neoempheria : A, liiderwaldli ; B, enderleini ; c, smarti.
cubital fork more parallel-sided, with Cui distinctly turned up at tip. Hypo-
pygium (text-fig. 9, b) quite different : tergite without strong bristles and with
the terminal process extremely long ; inner style not swollen at tip and with
one long black bristle in addition to some shorter ones ; outer style less bristly ;
anal segment small and with short setae ; aedeagus with a long median rod,
which is not present in kdstneri. Eighth tergite differently shaped and with
six setae.
Brazil : Santa Catharina (Luderwaldt) , 1 <$ (type) in Stettin Museum,
formerly a paratype of PL johannseni var. evanescens End. Nova Teutonia
(Plaumann), I $ in British Museum.
NEOTROPICAL NEOEMPHERIA 123
Neoempheria smarti sp. n.
<3\ Closely resembles N. kastneri, enderleini and related species described
above, but smaller than any of them. Flagellum dark. Ocellar bristles nearly
reaching base of antennae. Mesonotum with sublateral stripes distinct and
median stripe fairly obvious, but dorso-central stripes not developed. Pleural
stripe as usual ; dark band at base of postnotum extending on to base of pleuro-
tergite. Abdomen with tergite I dark above, 2 with lateral and posterior margins
broadly yellow, 3 and 6 yellow laterally at base, 4 and 7 all yellow, 5 all dark.
Coxae and femora entirely yellow. Wings with rather faint markings arranged
as in the related species ; a rather large pale area in base of median fork ; cloud
below Cu2 indefinite. Sc completely bare. M2 markedly bent upwards at tip.
Hypopygium (text-fig. 9, c) : tergite bare except for a few fine hairs on the long
and simple terminal process ; inner style broad at base and divided into two
nearly to the base, dorsal part forming a long spine, ventral part longer and
stouter, bearing 7-8 black spines at its tip ; outer style slender. Eighth tergite
with 3 setae in type (perhaps normally 2).
Wing-length scarcely over 2 mm.
British Guiana: Mazaruni, secondary forest, 16.viii.37 (0- W- Richards
and J. Smart), 1 <?.
The structure of the hypopygium of this species is as nearly as possible
intermediate between N. luderwaldti and enderleini, the inner style resembling
the former and the tergite the latter species.
Neoempheria unispinosa sp. n.
cJ. Closely resembles N. kastneri, but mesonotal markings somewhat different ;
the median stripe is faint and abbreviated posteriorly, and there is a pair of
faint dorso-central stripes which are more distinct behind and unite in front of
the scutellum. Hypopygium (text-fig. 10, a) comparatively small and of quite
different structure : halves of tergite regularly tapering and with a single strong
bristle outwardly near base ; inner style slender, with one black spine at base,
tip bare ; outer style slightly clubbed ; ventral process of sternite long. Eighth
tergite with two setae.
Brazil : Nova Teutonia (Plaumann), 1 S (tyPe)-
Neoempheria flavicornis sp. n.
cj. Closely resembles N. kastneri, but ocellar bristles shorter (though still
long) and pleurotergite entirely yellow, in these respects differing from all the
last eight species described. Flagellum with only the distal half dark, base
extensively yellowish. Wing-markings (PI. II, fig. 21) as in the allied species,
but cubital fork rather differently shaped. Second tergite rather broadly yellow
posteriorly, third with yellow lateral patches occupying half its length, fourth
with a small dark area above. Hypopygium (text-fig. 10, b) somewhat of the
type of N. kastneri, but process of tergite not so long, with a small projection
near tip ; inner style elbowed but not swollen, with about six black spines in a
row at tip ; outer style rather short. Eighth tergite with four setae.
Brazil : Nova Teutonia {Plaumann) , 1 <J (type).
novit. zool., 42, 1 10
124
F. W. EDWARDS
Neoempheria rostrata sp. n.
(J. Closely resembles N . flavicomis and pilosa, and like them with the pleuro-
tergite completely yellow ; small external differences which may not be constant
are that the dark dorsal area of tergite 2 extends almost to the posterior margin,
and vein Sci is noticeably longer than Sc2 instead of only about as long. Hypo-
pygium (text-fig. 10, c) quite different : tergite with the basal part almost bare
except for a group of stout lateral bristles, process long, slender and pointed,
no shoulder at its origin ; inner style blackened, widened in middle and with an
Text-fig. 10. — Hypopygia of Neoempheria : a, unispinosa ; B, flavicomis ; c, rostrata ;
D, pilosa.
irregularly double row of short black spines on distal margin of the wide part, a
single fine hair in addition to the spines, but the long beak-like tip bare ; para-
meres each ending in two points.
Brazil : Nova Teutonia (Plaumann) , 1 3 (type).
Neoempheria pilosa sp. n.
cJ. Closely resembles the last species in all external features, having the
oblique pleural stripe distinct as in other species of this group but the pleuro-
tergite entirely pale. Hypopygium (text-fig. 10, d) very distinctive owing to
the structure of the inner style, which is slightly clubbed, the club clothed with
a tuft of hair. Tergite with three long bristles on basal part, the long terminal
NEOTROPICAL NEOEMPHERIA 125
process arising abruptly from a shoulder. Outer style moderately long and very
hairy. Eighth tergite with two setae.
Brazil : Nova Teutonia (Plaumami), 2 J (including type).
Females of this group cannot at present be associated with their respective
males, and there are probably many other species in addition to those described
above. One such is represented in the British Museum by a damaged male
from Manaos (G. V. Vredenberg).
Group F (Pleonazoneura Enderlein).
Wings and mesonotum as in Group E, but ocellar bristles shorter, reaching
only a little beyond front edge of the black spot, and pleurae lacking the oblique
stripe, at most with a small dark area immediately in front of wing-root. Halteres
with dark knob.
Neoempheria lindneri sp. n.
(J. Head with the area between ocellar spot and base of antennae darker
than usual. Ocellar bristles longer than in other species of this group. Meso-
notal colouring as in N. ornatipennis and kdstneri, except that the dark band on
postnotum is sharply denned, broadly V-shaped and distinctly removed from
base of postnotum ; pleurotergites with a sharply defined dark patch at base,
which does not extend on to anepisternite. Immediately below and in front of
wing-root is a small and rather ill-defined dark area which does not cross the
anepisternite. All coxae yellow. Abdominal markings as in N . kdstneri, tergites
2 and 4 being broadly pale on posterior margin. Wings (PL I, fig. 6) with the
dark markings stronger than in the species of Group E, and peculiar in having no
darker area below Cu2, but a slight cloud in basal half of basal cell. Hypopygium
(text-fig. 11, a) : in many respects very similar to that of N. enderleini, but inner
style rather differently shaped and darkened, with two black bristles at middle
and only two or three black spines at tip ; eighth tergite otherwise.
Wing-length 4 mm.
Brazil : Nova Teutonia {Plaumami) , 1 $ (type).
Dedicated to my friend Dr. E. Lindner of Stuttgart, who made a collecting
expedition to the Chaco in 1925-26.
Neoempheria shannoni sp. n.
cJ. Head largely pale above, dark at sides and on space between ocellar
spot and base of antennae. First few antennal segments yellowish ; palpi black
as usual. Mesonotum with acrostichal and dorso-central dark stripes equally
distinct and obviously darker than the ground colour, but sublateral stripes rather
wider and darker as usual ; postnotum with an ill-defined and indefinitely darker
band at base, not or scarcely extending on to pleurotergites. Pleurae entirely
yellow, no trace of darkening in front of wing-root. Abdomen with a continuous
blackish median longitudinal stripe, not interrupted on posterior margins of
tergites 2, 4 or 7 ; yellow areas at --ides of tergites as usual, j and 5 with whole
posterior margin broadly dark. Wings (PI. II, tig. 21) with the dark areas of
more intensive tint than in N. kdstneri and related species, but similar in arrange-
ment ; a pale area completely tills base of median fork ; a dark area present
below Cu2 as usual. Hypopygium (text-fig. 11, c) : tergite with each half long
and narrow but somewhat truncate at tip, with several strong bristles mar base ;
126
F. W. EDWARDS
inner style of rather complex form, with a long outwardly directed tooth at
middle and about four small black teeth at tip. Eighth tergite square-ended,
with about 10 setae.
?. Resembles <J.
Wing-length 4 mm.
Brazil : Nova Teutonia (Plaumann), 1 S (type), 1 ?.
Dedicated to my friend Dr. R. C. Shannon.
This species seems to be rather nearly related to the North American N.
illustris Johannsen, but in a female of the latter which I collected in Washington
Text-fig. ii. — Hypopygia of Neoempheria : a, lindneri ; B, johannseni ; c, shannoni.
in 1928 the mesonotal markings are more sharply defined, and Johannsen's figure
shows differences in the hypopygium, notably in the form of the inner style.
Neoempheria johannseni (Enderlein).
191 1. Pleonazoneura johannseni Enderlein, Stettin, ent. Ztg., 72: 156.
The lectotype male in the Stettin Museum closely resembles the last species
in external features and the hypopygium (text-fig. n, b) is also similar, but
differs in detail : the tergite is much broader, and the inner style has a regular
comb of ten or more small black teeth at the tip. Eighth sternite more hairy.
Brazil : Santa Catharina {Liiderwaldt).
Neoempheria costa-limai sp. n.
cJ. Head largely pale above as usual ; antennae with first five or six segments
yellowish, rest dark ; palpi black. Mesonotum with sublateral and acrostichal
NEOTROPICAL NEOEMPHERIA 127
stripes connected on front margin as usual, but dorso-central stripes not indicated.
Postnotum with rather faint dark basal band indistinctly extending on to pleuro-
tergites ; pleurae otherwise entirely yellow. Abdominal tergites 2 and 4 almost
all yellow, with a small dark area at base above. Wings (PI. II, fig. 23) differing
from those of all other species of Groups E and F described above in having the
small cell shorter, with R4 only a little beyond level of middle of stem of median
fork. Dark areas of wing rather faint, but cloud at tip completely filling median
fork. Hypopygium (text-fig. 12, a) in several respects very different from all
the other species : tergite with twisted tip set with dense patches of setae ; inner
style bare and with bifid tip, parameres also with bifid tips ; sternite on each side
with a long bare pointed process, as long as the long and slender outer style.
Eighth tergite larger than usual, with a row of about 16 setae.
Wing-length 3-5 mm.
Brazil : Nova Teutonia (Plaumann), 3 $ (including type).
Dedicated to Dr. A. Costa Lima.
Neoempheria biflagellata sp. n.
cj. Very similar to the last described species, differing mainly if not solely
in structure of hypopygium (text-fig. 12, b). Like the last species this has the
parameres divided at the tips and the lobes of the sternite drawn out into long
bare whip-like processes as long as the outer styles, but the tergite has a com-
pletely different shape, as has the inner style, and other details also differ ; the
eighth tergite is smaller.
Wing-length barely 3 mm.
Brazil : Nova Teutonia (Plaumann), 1 <3 (type).
Group G.
Sc ending above base of Rs ; Sc2 at its tip and transverse. R4 near level
of middle of stem of median fork, which is longer than usual, the small cell short.
Costa strongly produced. fCu beyond base of Rs. Trichiation of veins alike
in the two sexes, Ml and Cu2 setose throughout, M2 completely bare, Cui setose
except at base ; Sc completely bare ; Cu bare except for a short distance before
the fork. Wings with dark markings at middle and tip, but costal and basal
cells clear except for the dark area over base of Rs. Ocellar bristles long, and
bristle on second antenna! segment longer than usual. Pleurae with dark longi-
tudinal stripe or with lower part more extensively darkened, but upper part pale.
No definite bare stripes between acrostichal and dorso-central hairs. Halteres
yellow.
Neoempheria vogeli sp. n.
cj- Head yellowish except for the ocellar spot ; antennae with first four or
five segments pale, rest dark ; palpi black. Mesonotum uniformly brown above,
without stripes, but with the side margins rather narrowly yellow. Pleurae
yellow, with a dark stripe crossing middle of pleurotergite and upper margin of
sternopleura. Abdomen with tergites 2, 3 and 4 similarly coloured, with a
continuous median dorsal dark stripe and with the posterior and lateral margins
of each narrowly dark ; 5 more extensively dark. Coxae and femora all yellow,
Wings (PI. I, fig. 3) slightly darkened at base, with the dark area in middle
leaving much of the small cell clear, but covering basal half of stem of median
fork. Apical dark area leaving base of fork clear but extending below base of
128
F. W. EDWARDS
M2. Vein An reduced to a short stump. Hypopygium (text-fig. 12, c) : tergite
broad at base, the terminal process expanded and blackened at tip and with a
pair of long stout bristles ; anal segment large, the sternite deeply bifid ; appa-
rently there is no inner style ; outer style forming a broad flat dark plate with
a finger-like termination. Eighth tergite truncate and bare.
Wing-length 3-5 mm.
Brazil : Petropolis (Vogel), 1 cJ (type) , presented to British Museum by Father
Borgmeier in 1931.
Neoempheria lutzi sp. n.
$. Differs from N. vogeli as follows : Pleurotergite wholly dark ; sterno-
pleura with a short dark stripe above. Abdomen with tergites 2 and 4 mainly
Text-fig. 12. — Hypopygia of Neoempheria : a, costa-limai ; b, biflagellata ; c, vogeli.
yellow, with the anterior and posterior margins narrowly dark ; 3 (as well as 5-7)
mainly dark. Hind coxa slightly darkened outwardly. Wings (PI. I, fig. 4)
not at all darkened at base, with the dark area in middle completely filling the
small cell. Vein An long, but pale.
Wing-length 4-5 mm.
Brazil : Nova Teutonia (Plaumann), 1 ? (type).
Dedicated to Dr. Adolpho Lutz.
Group H.
Sc ending a little beyond base of Rs ; Sc2 a short distance from its tip and
transverse. R4 before fM, stem of fork hardly longer than usual. Costa strongly
produced. fCu before base of Rs. Sc, M2 and Cui completely bare ; stem of
Cu bare except for a short distance before the fork ; Mi and Cu2 setose through-
out. Wings dark in middle and at tip, basal half clear. Ocellar bristles short.
Mesonotal bristles scantier than usual ; dorso-centrals irregularly uniserial ;
NEOTROPICAL NEOEMPHERIA
129
acrostichals few. Scutellum with two strong bristles as in other Neotropical
species of the genus. Pleurae dark above, pale below. Halteres dark.
Neoempheria pereirai sp. n.
$. Head mainly dark brownish ; scape brown, flagellum and palpi black.
Thorax uniformly dark brown above and somewhat shining. Pleurae mainly
blackish brown, but lower part of pleurotergite and whole of sternopleura whitish
yellow, the two colours abruptly separated. All coxae whitish yellow, with the tips
pale brownish yellow ; femora and tibiae slightly darker than coxae, tarsi dark.
Abdomen mainly blackish ; tergite 2 with a rather narrow yellow transverse
band beyond middle but with whole of posterior margin dark ; 4 with a yellow
spot on each side at about the middle ; 7 mainly yellowish. Wings (PI. I,
fig. 5) narrower at base than in any of the other species, with the dark areas in
middle and at tip broadly connected on hind margin, leaving base only of cell
Cui clear. Veins CuP and An both obsolete.
Wing-length 3 mm.
Brazil : Salobra, Matto Grosso, vii. 1939 (J. Lane), 1 $ (type), presented to
the British Museum by the collector.
Dedicated to Dr. Clemente Pereira, Director of the Clube Zoologico Brazileiro,
under the auspices of which the expedition to Salobra was organized.1
1 A description of this species has also been included in a report on the Mycetophilidae
collected by the Salobra Expedition ; this report is being sent (January, 1940) for publication
in Boletim Biologico, S. Paulo (n.s.).
(MSS. reed. Jan. 20, 1940.)
PLATE I.
Wings of Neoempheria spp
1. N. neivai sp. n.
2. N. bipectinata sp. n.
3. N. vogeli sp. n.
4. N. lutzi sp. a.
5. N. pereirai sp. n.
6. N. lindneri sp. n.
7. N. spinosa sp. n.
8. N. brevicauda sp. n.
9. N. simplex sp. n., $ var. ?
10. N. simplex sp. n., J type.
11. N . bilobata sp. n.
12. TV. borgmeieri sp. n.
PLATE II.
Wings of Neoempheria spp.
13. N. plaumanni sp. n.
14. N. plaumanni sp. n., var. ?
15. N . flavicoxa sp. n.
16. N. mulleri sp. n.
17. N. lanei sp. n.
18. ./V. maculipennis Will.
19. N. ornatipennis (End.
20. N . luderwaldti sp. n.
21. N . flavicomis sp. n.
22. N. shannoni sp. n.
23. N . costa-limai sp. n.
24. N. biflagellata sp. n.
NOYIT. ZOOI... 42, 1940.
EDWARDS PLATE I.
Wings of Neoempheria spp
>i n, Lid
Nl IV] I . ZOOL., 42, 1940.
EDWARDS PLATE II
17
Wings 1 'I \ eoempheria spp,
SAPROMYZIDAE FROM THE SOLOMON ISLANDS 131
SOME SAPROMYZIDAE (DIPTERA) FROM THE SOLOMON
ISLANDS AND NEW CALEDONIA
By JOHN R. MALLOCH.
(With one plate.)
THERE are two papers by Curran dealing with the Sapromyzidae of these
islands.1 Species recorded here which were described by him or listed from
either group, are referred to in those papers. Only three species were recorded
from New Caledonia, but from the Solomons Curran recorded 4 genera and
18 species, 10 of the latter new. Where I am able to do so, I have identified
his species and made suggestions as to their status.
The materials on which this paper is based were kindly submitted for identi-
fication by the Imperial Institute of Entomology. All the material will be placed
in the British Museum (Natural History).
STEGANOPSIS de Meijere.
1910, Steganopsis de Meijere, Tijdschr. Enl., 53 : 145.
In 19292 I published a complete key to the species of this genus. Curran
described two species from the Solomon Islands in the paper referred to above.
There are four species in the present collection and below I present a synoptic
key to them.
Key to the Species.
1. Face glossy fulvous or orange-yellow, with a few deep black glossy
marks ; wings slightly yellowish brown clouded, with no definite black
and hyaline markings, the apical edge from near apex of second to
apex of fourth vein whitish hyaline ; all femora and most of tibiae
fulvous yellow .......... 2.
Face black, with or without pale dust ; wings black, with two oblique
hyaline fasciae on apical portion beyond the outer cross vein, and
usually some less distinct hyaline or subhyaline spots or streaks on
basal half ; femora and tibiae black, tarsi partly yellow or white . 3.
2. Face with two dark marks on each side, the upper one glossy black, tin-
lower one smaller and dark brown, and no central dark mark ; fron:
merely shiny on the interfrontalia, and with a large velvety black mark
covering the ocelli and extending to at least midway between anterior
ocellus and anterior margin (New Caledonia) . melanogaster (Thomson).
Face with a large oval glossy black central mark that extends from
epistome to between antennae and covers half the width of face, and a
small round black-brown spot against each eye margin near middle :
frons entirely glossy except the narrow dull orbits and a central line,
and with a small blackish mark over the ocelli . tripunctifades, sp. a.
1 1929, Amer. Mus. Nov., 375. New Caledonia.
193°. Proc. Calif. Acad. Sci., (4), 22, no. 1 : 32-41. Solomon Islands.
' 1929, Proc. U.S. nat. Mus., 74, art. 6.
NOVIT. ZOOL., 42, I II
132 JOHN R. MALLOCH
3. Head, thorax and abdomen glossy black, the frons with a distinct
greenish-blue lustre ; fore tarsi white, apical three segments and a
narrow basal mark on the metatarsus deep black ; wing as fig. I.
varimana, sp. n.
Head and thorax black, with dense whitish-grey dusted markings, centre
of frons rather dull pale grey, with slight grey dusting ; fore tarsi
yellow, extreme base of the metatarsus and all of the apical two
segments black ....... solomensis Curran.
Steganopsis melanogaster (Thomson).
1868, Lauxania melanogaster Thomson, Eugenics Resa (Insecta) : 567.
This species occurs in Australia and New Guinea. Curran recorded it from
New Caledonia. I have some specimens before me that apparently belong to
the same material as his, having been collected by Prof. T. D. A. Cockerell.
Steganopsis tripunctifacies sp. n.
$. Head glossy fulvous yellow except the narrow orbits and a central
frontal line, third antennal segment almost entirely dark brown, palpi infuscated
at apices. Bristles black. Frons wider than long, glossy on its entire width
above the upper orbital bristles, both the bristles on the edge of the glossy portion
subequal in length, ocellars merely short hairs, divergent and proclinate. Basal
antennal segment longer than second, third about nine times as long as wide,
and as long as height of head ; longest hairs on the aristae not as long as width
of third antennal segment. Eye higher than long, tapered below, not three
times as high as the gena. Face quite prominent centrally and evenly convex in
profile.
Thorax in type-specimen discoloured, fulvous yellow in front, showing three
broad blackish-brown vittae on the mesonotum that are apparently fused behind,
pleura mostly blackened, scutellum glossy black. Bristles weak, only the
posterior one of the four pairs of dorsocentrals well developed, the acrostichal
hairs minute, biseriate as in melanogaster. The sternopleura and mesopleura
each with one bristle.
Legs fulvous yellow, bases and apices of fore tibiae fuscous, apical four seg-
ments of fore tarsi black. Preapical dorsal bristle on fore tibia slender, as long
as the metatarsus, the one on mid tibia stout, much shorter, that on the hind
tibia slender, but shorter than the metatarsus.
Wings (fig. 1) brownish yellow, more yellowish on basal half of the
costa. Second wing vein evenly arched, running parallel with the costa from
apex of first vein to its forward curve to connect with the latter. White apical
margin linear. Halteres with the knobs blackish brown.
Abdomen thick, broadly ovate, and glossy black.
Length : 3-5 mm.
Type : San Cristobal, Solomon Islands (R. A. Lever).
In melanogaster the mesonotum has a number of yellowish-brown vittae that
are more or less distinctly margined with black so that there may be eight or ten
partial black lines or vittae in evidence in some specimens.
SAPROMYZIDAE FROM THE SOLOMON ISANDS 133
Steganopsis varimana sp. n.
$. A glossy black species, with greenish-blue lustre on frons, the basal three
segments of all tarsi yellow or white, the other segments and narrow base of the
fore metatarsus black, and the halteres black. Wing as fig. 2.
Frons entirely glossy, wider than long, depressed on posterior lateral angles,
posterior ocelli on edge of vertex, orbitals very near eye-margins, quite closely
placed at middle, ocellars microscopic. Antennae fulvous yellow, tip of third
segment slightly infuscated, aristae much longer than entire antennae, the longest
hairs nearly twice as long as width of third antennal segment, the latter about
six times as long as wide, and not as long as height of head. Eye higher than
long, narrowed below, emarginate on lower half behind, and about eight times
as high as gena. Postocular orbits silvery white dusted. Palpi black. Face
slightly convex in profile at middle, depressed below convexity, the epistome
projecting slightly.
Thorax glossy black, with black hairs and bristles, undusted. Mesonotum
with four pairs of dorsocentrals and acrostichals, the anterior pairs of both small
and presutural, and a series of fine hairs between the acrostichals and dorso-
centrals on each side. Scutellum slightly convex on disc, obtusely rounded in
outline, the edge thin.
Legs black, tsrsi as described in the foregoing key to the species. Fore tibia
with a very short moderately strong preapical dorsal bristle, mid-tibia with
longer and stronger, and hind tibia with a very short such bristle.
Wings black, with hyaline markings (fig. 2). Second vein undulated,
not parallel with costa on its entire extent, the submarginal cell comparatively
narrower than in tripitncti fades. Halteres black.
Abdomen thick, broadly ovate, glossy black, with black hairs and bristles
and very slight greyish dusting.
Length : 4 mm.
Type: Solomon Islands, Kolombangara, 5000 feet, at summit, 11. x. 1933
(H. T. Pagden.)
Steganopsis solomensis Curran.
1936, Steganopsis solomensis Curran, Proc. Calif. Acad. Sci.. (4), 22 : jj.
This species is readily distinguished from the others in this paper by the
densely grey dusted frontal orbits, the slightly metallic grass-green, rugose,
interfrontal area, and the grey dusted and black dotted and vittate mesonotum.
The frontal bristles are much stronger and more evenly spaced than in the other
species, while the genae are not one-fourth as high as the eyes. The halteres are
yellow, and the wings though marked much as in varimana have more evident
subhyaline markings on the basal half.
Length : 3-4 mm.
Originally described from Santa Catalina Is. I have specimens from Tulagi,
12.viii.1933 (H. T. Pagden), 23. hi. 1934 (R. A. Lever).
Steganopsis ceres Curran.
[936, Steganopsis ceres Curran, Proi . ( alif. Acad. Sci., (4), 22 : 32.
A careful study of the description of this species discloses no characters that
<nable me to embody it in the key given above, except possibly the smooth
instead of transversely rugose interfrontal area of the head. A teneral specimen
of solomensis before me has no frontal rugae.
Described from a single specimen from Guadalcanal Is.
134 JOHN R. MALLOCH
DIOIDES Kertesz.
1915, Dioides Kertesz, Ann. hist.-nat. Mus. hung., 13 : 491.
This genus is very similar to Cestrotus Loew, differing in having the aristae
very short haired. The frons is humped up, but not armed with a tubercle in
the middle, the sternopleura has two bristles in the species before me, though
the genotype is described as having but one, and the costal vein is as in Homoneura,
the short dense black setulae being continued to the apex of the third vein, where
they cease abruptly. In the species of Cestrotus (Turriger Kertesz) known to me
the costal vein is as in Sapromyza.
Dioides fuscoapicata, sp. n.
$. Head testaceous yellow, distinctly shiny at the bases of the orbital
bristles only. Frons with brown tinge on the upper orbits, a blackish brown
elongate mark over the ocelli, and another, drop-like, one in front of it that does
not extend to anterior margin, a dark brown streak on centre of face that begins
just above the apex of the facial elevation and continues down to epistome, a
black line in the suture between the face and parafacial, and a dark brown mark
on each gena close against the eyes. Antennae brownish black, third segment
and aristae fulvous yellow ; palpi fulvous yellow. Head in profile as figured by
Kertesz, the eye, however, not emarginate below on hind margin. Ocelli in line
between the upper orbitals, the ocellar bristles rubbed off in the type.
Thorax shiny testaceous yellow, largely blackened, the pleura yellow only
on the sutures, and slightly whitish grey dusted, the mesonotum with six blackish
vittae, the central pair carried over disc of the scutellum, the others largely fused
and indistinctly defined ; postnotum blackened. Bristling as in typical Homo-
neura species, the mesonotum with 8-10 series of short hairs between the dorso-
centrals, and no acrostichals but the prescutellar pair, the propleural quite strong.
Wing as fig. 3. The general colour brownish hyaline, inner cross vein
narrowly, outer one more broadly, clouded with dark brown, especially above.
Halteres brownish yellow.
Legs testaceous yellow, all femora with a broad fuscous to black mark on
basal halves and a narrower black mark at apices, extreme bases of all tibiae
black and a dark-brown ring near middle. Fore femur without an apical antero-
ventral comb and with the posteroventral bristles confined to the apical half,
mid femur with one apical posterior bristle ; all tibiae with a preapical dorsal
bristle, the one on hind tibia the weakest ; fore tarsi much longer than their
tibiae ; rather slender.
Abdomen dull testaceous yellow, with a narrow blackish apical line on each
tergite, the hairs and short apical bristles on tergites dark brown.
Length : 6 mm.
Type : Solomon Islands : Velio Lavetta, Parasso, 22. v. 1934 (H. T. Pagden).
The genotype, pictipennis Kertesz, from Formosa, has the wings more
extensively spotted with black, some marks being on the basal half, and the
outer cross vein is in a hyaline and not a dark mark.
MONOCERA van der Wulp.
1898, Monocera van der Wulp, Termeszetr. Fiiz., 21 : 425.
This genus was erected for the reception of a species that has in the male a
long forwardly projecting and downwardly curved thorn on the anterior margin
SAPROMYZIDAE FROM THE SOLOMON ISLANDS 135
of the frons. Kertesz in 19151 presented a revision of the species, describing
two as new, and included one that has the process or thorn on the epistome
instead of on the frons. I have seen the genotype, monstruosa van der Wulp,
from New Guinea, and have before me now a pair of J 'areata Kertesz, the aberrant
species with the furcate epistomal process.
The general characters, including the type of costal armature, are as in
Homoneura, the thoracic bristling being identical with normal forms in that
genus, but the arista is very regularly short haired, the ocellar bristles are directed
straight outward, the head is broader, the eyes are more narrowed below, the
fore femur has a very fine anteroventral comb on the apical half, and there are
two short black curved anterior spurs on the apex of the hind tibia.
Monocera furcata Kertesz.
1915, Monocera furcata Kertesz, Ann. hist.-nat. Mus. hung., 13 : 505.
Like all the other known species this one is testaceous yellow in colour, with
dark spots on the wings. The frons has four dark marks in the male, in the
female but one or none, the gena has a brown mark, and the mesonotum has
four dark brown vittae, the central pair linear, the outer pair broader, all four
fused about one-third from posterior margin. The abdomen has dark brown
spots or streaks on sides of the tergites, and the legs are yellow. Wings in the
genotype and furcata with, in cornitta Hendel and rhinoceros de Meijere without,
dark spots on apical section of the fourth wing vein. In furcata there are two
such spots, one at the apex and the other at the middle of that section of the vein,
while in monstruosa there is but the apical spot present. In monstruosa the thorn
is on the anterior part of the frons, in furcata it is on the epistome, projecting
forward and ending in two equally long laterally directed branches (fig. 4).
Originally described from New Ireland. Solomon Islands : (J, Bougainville,
Teopasina, 28.1.1936 (R. A. Lever) ; $, Guadalcanal Is., Kovagoumbi, 1. v. 1936
(R. A. Lever).
HOMONEURA van der Wulp.
1891, Homoneura van der Wulp, Tijdschr. Ent., 34 : 213.
This genus contains more species than all the others that occur in the Old
World, and is almost cosmopolitan, occurring in all countries from which I have
seen Sapromyzidae except New Zealand and Patagonia.
Curran has recorded two species from New Caledonia and twelve from the
Solomon Islands.
In 19292 I erected several new subgenera in Homoneura, and in this paper
I erect another. There is a great probability that some student of the family
will later accept these as distinct genera.
Key to the Species.
1. Fourth wing vein slightly but quite noticeably curved forward at apex
so that the first posterior cell is narrowed at tip ; mesonotum with a
strong but short prealar bristle just in front of the strong and long
supra-alar, and a long strong posterior intra-alar (subgenus Solomonia
novum) ......... leveri, sp. n.
Fourth wing vein not appreciably curved forward at apex ... 2.
1 Ann. hist.-nat. Mus. hung., 13 : 502.
' Proc. U.S. nat. Mus., 74, art. 6 : 12-15.
I36 JOHN R. MALLOCH
2. Head black, with a conspicuous brownish-yellow anterior margin to the
frons ; thorax and abdomen glossy black ; knobs of halteres black ;
arista with the longest hairs as long as width of the third antennal
segment ........ signatifrons (Kertesz).
Head rarely entirely black, never marked as above, and the thorax and
abdomen never entirely glossy black . . . . . 3.
3. Thorax and scutellum black, densely covered with whitish-grey dust, the
mesonotum with two broad dark brown vittae laterad of the lines of
dorsocentral bristles, the outer one on the extreme lateral edge ;
coxae and femora black, tibiae and tarsi testaceous yellow, aristae
plumose ........ viatrix (de Meijere).
Thorax shiny or glossy fulvous yellow or orange-yellow, the mesonotum
not vittate as above ; legs yellow, or other characters not as above . 4.
4. Longest hairs on the aristae as long as or much longer than width of
the third antennal segment ; wings without dark markings except
sometimes a faint narrow infuscation on the outer cross vein . . 5.
Longest hairs on the aristae not nearly as long as the width of the third
antennal segment, or the wings with distinct brown markings, some-
times on costa and cross veins . . . . . . .8.
5. Face black, lightly grey dusted, but glossy ; frons brown or fuscous,
shiny, upper orbits brownish yellow ; outer cross vein not clouded ;
a short but strong prealar bristle present ; fore femoral comb con-
sisting of closely placed and moderately long setulae . atrifacies, sp. n.
Face entirely orange-yellow ; frons concolorous to brownish yellow . 6.
6. Prealar bristle present ; fore femoral comb consisting of rather irregular
setulae that are thick at bases and very fine and hair-like at apices ;
outer cross vein narrowly brownish clouded ; abdomen without paired
black spots on the tergites .... immacnliventris, sp. n.
Prealar bristle represented by a few short setulae ; fore femoral comb
consisting of very short regular stout spinules ; outer cross vein not or
very indistinctly brownish clouded ; abdomen with or without one or
two pairs of small black spots, usually one on the fifth and one on
the sixth tergite at the lateral curves ...... 7.
7. Antennae entirely yellow ; fifth and sixth abdominal tergites with a pair
of spots each on lateral curves, or the fifth with such spots ; longest
hairs on the aristae longer than the width of the third antennal
segment ; anterior pair of orbital bristles much shorter than the
posterior pair russelli, sp. n.
Antennae yellow, third segment blackened at apex ; abdomen without
paired black spots on dorsum ; longest hairs on aristae about as long
as the width of the third segment ; anterior pair of orbital bristles
about as long as the posterior pair .... suspensa, sp. n.
8. Wing with distinct dark markings in addition to any clouds over the
cross veins ........••• 9-
Wing with at most faint clouds on the cross veins . . . .11.
9. Anterior of the three pairs of dorsocentral bristles on the mesonotum
in front of the suture ; face very wide . . Trypaneoides confusa, sp. n.
Anterior pair of dorsocentral bristles slightly behind the mesonotal
suture .......•■•• I0-
SAPROMYZIDAE FROM THE SOLOMON ISLANDS IJ7
10. Wing with a brown suffusion over the costa that extends to third vein,
outer cross vein clouded narrowly with brown ; antennae and palpi
yellow ........ laticosta (Thomson).
Wing with brown marks on both cross veins and apices of veins 2 to 4
horvathi Kertesz.
11. Mesonotum with one or more pairs of well-developed acrostichal bristles
in front of the prescutellar pair ....... 12.
Mesonotum with no outstanding acrostichal bristles but the prescutellar
pair 15.
12. Mesonotum fulvous yellow, slightly shiny, with slight even grey dust on
entire surface, the scutellum similarly dusted ; hind tibia with the
preapical dorsal bristle short and strong, subequal to the slightly
curved apical anterior hind tibial spur ; seventh abdominal tergite
short, with a few fine black hairs (4-6) on centre near apex, eighth
with a few similar scattered surface hairs, cleft V-shaped at apex ;
fourth to sixth tergites with three deep black marks, the first two
with the marks elongate, extending the entire length of exposed
surface, those on sixth more spot-like .... subnuda, sp. n.
Mesonotum glossy fulvous yellow or orange-yellow, without a trace of
dust, the scutellum also undusted ; preapical dorsal bristle on the hind
tibia much longer than the apical anterior hind tibial spur ; seventh
and eighth abdominal tergites not as above, with much more numerous
fine hairs ; the black abdominal markings when distinct not as above 13.
13. Small species, about 3-5 mm. in length ; male with a ridge or elongate
brush of dense stiff dark brown hairs or setulae on the entire extent
of the anteroventral surface of the basal segment of the hind tarsus,
higher on the apical half ; abdomen with three deep black spots on the
fifth tergite, the sixth almost concealed in type so that it is impossible
to detect any black spots ; seventh and eighth tergites with many
erect fine hairs of moderate length ; basal segment of the hypopygium
with two short forwardly directed points at apex . notaliventris, sp. n.
Larger species, over 4 mm. in length ; male with no ridge or elongate
brush of hairs as described above ; other characters not as above . 14.
14. Basal segment of the hypopygium with the apical thorn black and beak-
like ........ acrostichalis (de Meijere).
Basal segment of the hypopygium with the apical thorn pale and less
acute at apex ....... crockeri Curran.
15. Head dull yellowish brown, the frontal orbits yellowish grey dusted,
the postocular orbits greyish white dusted ; thorax yellowish brown,
densely grey dusted, the humeri, postalar callosities and margin of the
scutellum brownish yellow; abdomen brownish yellow, the apices
of tergites paler, third and fourth tergites eaeli with a wide central
black fascia, the fifth, and sometimes the sixth with a pair of large,
and sometimes rather diffuse black spots ; femora usually browned or
infuscated, the tibiae and tarsi testaceous yellow; inner cm,, vein
of the wing darker than the outer but not margined with dark colour
hawaiiensis Malloch.
Head, thorax, and abdomen shiny to glossy fulvous yellow , w ithoul con-
spicuous grey dust, the abdominal tergites without distinct black
fasciae; legs yellow; inner cross vein of the wing not noticeably
darker than the outer one ........ 16.
I38 JOHN R. MALLOCH
16. Wing epaulets yellow ; longest hairs on aristae about as long as width of
third antennal segment ; apex of third antennal segment blackened
suspensa, sp. n.
Wing epaulets black ; longest hairs on aristae about half as long as width
of third antennal segment ; third antennal segment entirely orange-
yellow ........ diversa, sp. n.
Subgenus SOLOMONIA novum.
This subgenus differs from typical Homoneura in having the fourth wing
vein slightly curved forward at its apex, the posterior intra-alar very long and
strong, and the prealar bristle present though short and strong.
Type species : Homoneura {Solomonia) leveri, sp. n.
Homoneura (Solomonia) leveri, sp. n.
$ and $. Head black, the frons more brownish, face with distinct whitish-
grey dust, frons with faint greyish dust, the orbits and narrow triangle shiny,
remainder dull. Antennae yellowish brown, third segment blackened apically ;
palpi yellowish brown. Frons subquadrate, slightly narrowed in front, all the
bristles long and strong, the two pairs of orbitals subequal in length, surface
hairs microscopic. Antennae of moderate size, third segment about 25 times
as long as wide, rounded at apex, second with one bristle above and two or three
below at apex, the former and central one below longer than width of third
segment ; arista with the longest hairs distinctly longer than the width of third
antennal segment, about equally long, but more sparsely haired below. Palpi
normal. Face convex, slightly more protruded to epistome, the foveae distinct.
Hairs ascending to about middle of lateral suture. Eye higher than long, tapered
below, not emarginate behind, and about ten times as high as gena. A number
of long bristles on back of head below.
Thorax black, shiny, with even, but not very conspicuous, whitish-grey dust,
most distinct on scutellum when seen from in front against the light, the humeri
sometimes yellowish brown. Mesonotum with three pairs of dorsocentrals, two
supra-alars, the posterior intra-alar long and strong, and a short strong prealar,
the latter sometimes duplicated, the other bristles as in typical Homoneura,
the sternopleurals 2. Intradorsocentral hairs in at least twelve rows ; only the
prescutellar acrostichals developed. Scutellum slightly flattened on disc, broadly
rounded in outline, the basal bristles very near base. Prosternum and disc of
pteropleuron bare.
Legs black, extreme apices of femora, all of tibiae, and tarsi, testaceous
yellow. Fore femur with the anteroventral comb fine but evident, and a com-
plete series of long posteroventral bristles ; mid femur with a series of short
strong bristles on apical half of the anterior surface and two strong curved bristles
at apex on posterior surface ; hind femur with a series of bristles on more that
the apical half, all tibiae with preapical dorsal bristle, the one on mid pair the
longest and strongest, that on hind pair the weakest and shortest ; apical anterior
curved spur on hind tibia very short.
Wing brownish hyaline, veins brown, rather ovate in shape. Inner cross
vein a little beyond apex of first vein and at about two-fifths from base of discal
cell ; outer cross vein at a little over half its own length from apex of fifth vein.
Knobs of halteres brown to black.
SAPROMYZIDAE FROM THE SOLOMON ISLANDS 139
Abdomen glossy brownish black to deep black, without distinct dusting.
Each tergite with an apical transverse series of quite strong black bristles.
Length : 6-8 mm.
Type : $, Solomon Islands : Tulagi, 23. hi. 1934. Allotype : Santa Isabel,
iii.1933. Paratypes : Guadalcanal, Popanu, 1500 feet, 16.xii.1934, and San
Isabel, Fate, ii.1933. All taken by R. A . Lever, to whom the species is dedicated.
Homoneura (Homoneura) signatifrons (Kertesz).
1900, Sapromyza signatifrons Kertesz, Termeszetr. Fiiz., 23 : 264.
This apparently common and very widely distributed species is represented
by a series of specimens from the Solomons, many from Tulagi, and a single
specimen from Russell Island. There are also four from Admiralty Island
(Froggatt).
Originally described from New Guinea, this species has already been recorded
by Curran from the Solomon Islands.
Homoneura (Homoneura) viatrix (de Meijere).
1910, Lauxania viatrix de Meijere, Tijdschr. Ent., 53 : 123.
This very distinct species was originally described from the East Indies,
and has also been recorded from the Solomons by Curran.
Dne specimen : Tulagi {Lever).
I examined the type-specimen when I wrote my paper on the Oriental
species in 1929.
Homoneura (Homoneura) atrifacies, sp. n.
$. A large glossy fulvous yellow species with the head preponderantly black.
It runs down to section 38 in Kertesz's key to the species.1 It differs from the only
two species in that segregate as follows : from nigrifrons Kertesz in having the
legs entirely fulvous yellow, and in having the mesonotum with several pairs
of acrostichals that are as long and strong as the dorsocentrals ; and from limbata
in having no black spots on the scutellum.
Head shiny black, with slight whitish dust, most distinct on the parafacials.
the entire occiput glossy fulvous yellow ; antennae brownish yellow ; palpi
fuscous. Frons about 1-25 times as long as wide, all the bristles except the
ocellars long and strong, the latter about half as long as the postvertical pair ;
surface hairs microscopic and very few in number. Eye about 1-5 times as high
as long, much tapered below, the hind margin not emarginate, height about ten
times that of the blackened gena. Antennae inserted slightly above middle of
eye in profile, reaching about half the distance to epistome, third segment about
2-5 times as long as wide, rounded at apex ; longest hairs on aristae as long as
third antenna] segment, almost as long below as above. Face slightly convex,
almost vertical, the foveae distinct.
Thorax glossy fulvous yellow, the scutellum concolorous and rather duller
than the mesonotum. Bristling normal, no well-developed acrostichals except
the prescutellar pair, the intradorsocentral hairs in ten rather irregular series,
the intra-alar short, presternum with a few minute black hairs, pteropleura bare,
sternopleurals 2.
Legs fulvous yellow. Fore femur with a distinct comb of short regular
1 1900, Termeszetr. FUt., 23 : 255.
I40 JOHN K. MALLOCH
setulae on the apical half of the anteroventral surface and a series of long postero-
ventral bristles, mid-femur differing from that of leveri in having but one apical
posterior bristle, hind pair with one or two quite strong preapical anteroventral
bristles ; all tibiae with a preapical dorsal bristle, as in leveri.
Wing brownish hyaline, veins brown, unclouded. Venation as in leveri
except that the fourth vein is not bent forward at apex, and the outer cross
vein is at less than half its own length from apex of fifth vein. Halteres yellow.
Abdomen concolorous with thorax, one or two of the apical tergites with a
blackish central spot, the genital processes black and finely black haired. All
tergites with an apical transverse series of black bristles, longest on third tergite.
Length : 7 mm.
Type : Solomon Islands : San Isabel, 28. ii. 1934 (R. A. Lever).
Homoneura (Homoneura) immaculiventris, sp. n.
$. Very similar in general colour and features to atrifacies, differing mainly
in having the head entirely fulvous yellow.
Frons hardly longer than wide, the ocellar bristles as long and strong as the
postvertical pair, eye hardly 1-25 times as high as long, less tapered below than
in atrifacies, and about nine times as high as the yellow gena. Antennae and
palpi fulvous yellow. Arista long haired.
Thorax as in atrifacies, the intra-alar bristle minute.
Legs and wings as in atrifacies except that the inner cross vein is very
slightly proximad of the apex of first vein and the outer cross vein has a slight
but evident brownish suffusion. Halteres fulvous yellow.
Abdomen concolorous with the thorax, and without any black tergal spots.
Structure as in atrifacies.
Length : 7 mm.
Type: Solomon Islands: Tulagi, 5.301.1934, Hibiscus (R. A. Lever).
Paratype : same locality, 20. ii. 1935, same collector.
Homoneura (Homoneura) russelli, sp. n.
cT and ?. A smaller species than immaculiventris, differing from it as stated
in the foregoing key to the species, and in having the fifth and sixth, or at least
the fifth tergite with a pair of deep black spots.
This species will run down to signata van der Wulp or neosignata Malloch
in my key to the Oriental species of this genus, but it differs from the former
in having only two pairs of black spots on the abdomen, the outer pair on the
fifth and sixth tergites lacking, and from neosignata in having the fifth and sixth
instead of only the fifth tergite in the male with a pair of black spots. The outer
cross vein of the wing has a slight brown clouding. In Kertesz's key (1900) it
runs down to signata van der Wulp, and in Curran's (1936) to orientis Hendel.
Curran records the latter from the Solomons, but the species now before me is
not the one I accepted (1929) as bioculata (de Meijere) from an examination of
the type. I have not seen the latter from the Solomons.
Length : 4-5 mm. Hypopygium of male as fig. 5-
Type: <J, Solomon Islands : Russell Is., 18. ii. 1934. Allotype: Tulagi,
3.xii.i934. Paratypes : Russell Is., 22. ii. 1934, Tulagi, 25. xi. 1934, and
Guadalcanal Is., Lunga, 29.iii.1934 (R. A. Lever).
SAPROMYZIDAE FROM THE SOLOMON ISLANDS 141
Homoneura (Homoneura) suspensa, sp. n.
$. Another very similar species, but with no black spots on the abdomen,
and the third antennal segment blackened at the apex. The hairs on the aristae
are not as long as the basal width of the third antennal segment, though tin-
longest are about as long as the central width.
The face is noticeably white dusted, which it is not in the next preceding
species, the inner cross vein is just beyond the apex of the first vein and very
little basad of the middle of the discal cell, and the outer cross vein is slightly
brownish clouded. There are no hairs on the pteropleura, there are some on the
presternum, and the mesosternum has a number of quite strong erect black
hairs but no pair of stronger downwardly directed anterior bristles as in some
other species.
Length : 4-5 mm.
Type: Solomon Islands: Guadalcanal Is., Lunga, 9. vii. 1933 (R. A. Lever).
Except for the fact that Curran states that the species he distinguished as
distincta Kertesz has some pteropleural hairs, I might assume that this was that
species. I have not been able to find distincta amongst my material.
Homoneura (Homoneura) laticosta (Thomson).
1869, Geomyza laticosta Thomson, Eugenics Resa (Insecta) : 598.
Two specimens of this easily identified species from the Solomon Islands :
Guadalcanal Is., Lunga, 28. v. 1936 (R. A. Lever).
Curran records the species from the Solomons ; I have recorded it from the
East Indies, where it is widespread and common.
Homoneura (Homoneura) subnuda, sp. n.
(J. A smaller species than any of the others in the group, with anterior
acrostichals, and readily distinguished from nearly all the yellow species by the
even, though not dense, grey or yellowish-grey dust on the mesonotum and
scutellum. This dust may be easily removed by damp, so that other characters
should be used in segregations to insure accurate identification.
Head entirely orange-yellow except a greyish mark in centre of occiput low
down, just above neck. Frons about 1-5 times as long as wide ; ocellar and post-
vertical bristles shorter than the others, subequal in length. Eye higher than
long, tapered below, and about seven times as high as gena. Antennae inserted
slightly above middle of eye in profile, third segment not infuscated, about
2-25 times as long as wide, rounded at apex ; aristae slender, with the longest
hairs a little longer than the basal diameter ; palpi yellow, lace slightly white
dusted, the parafai ials more noticeably so, foveae distinct.
Thorax orange-yellow, the mesonotum and scutellum rather dull because of
the presence of a coating of greyish dust, no vittae present. Bristling normal,
one pair of rather strong acrostichals midwav between the prescutellar pair and
the suture ; intra-alar and prealar lacking ; sternopleurals -'. the anterior one
much the shorter, Prostemum with some microscopic hairs, no evidence oi hairs
on the pteropleura.
142 JOHN R. MALLOCH
Legs entirely yellow. Fore femur with the anteroventral comb on apical
half rather well developed, mid-femur with one apical posterior bristle, hind
femur with some very short fine anteroventral bristles ; all tibiae with a preapical
dorsal bristle, the one on the mid-pair the longest and strongest, that on hind
pair not longer nor stronger than the apical anterior curved black spur.
Wings greyish hyaline, veins brownish yellow, paler basally, the cross veins
not clouded. Inner cross vein almost directly below apex of first vein and at
about five-sixths from base of discal cell ; outer cross vein at about half its own
length from apex of fifth vein. Halteres yellow.
Abdomen glossy fulvous or orange-yellow, with hardly a trace of dust, fourth,
fifth and sixth tergites each with three black spots, those on the first two in
centre extending from base to apex, vittiform, the laterals on sixth below curve
and seen only from below. Other characters as noted in the key.
Length : 3-5 mm.
Type: New Caledonia: Bourail, 22. v. 1928 (W. P. Cockerell).
The only species of this genus recorded by Curran from New Caledonia in
the Cockerell collection are varia Kertesz and horvathi Kertesz. The former is
unknown to me but has the aristae long haired and only one abdominal tergite
with a pair of black spots (the fifth), so is quite distinct from the present species ;
for the latter see p. 144.
Homoneura (Homoneura) notativentris, sp. n.
<J. A species of the same size as the one just dealt with above, but readily
distinguished from it by the peculiarly haired basal segment of the hind tarsi of
the male, which has a ridge or elongate brush of dense dark brown stiff hairs
from base to apex on the anteroventral edge that is highest on apical half.
Head crushed in type, but apparently similar to that of subnuda, though the
ocellar bristles are finer, and shorter than the postvertical pair.
Thorax slightly discoloured in type, with two or three shorter pairs of acro-
stichals in front of the additional central postsutural pair, otherwise as subnuda.
Legs entirely yellow. In addition to the hind tarsal distinction from subnuda
the preapical dorsal bristle on the hind tibia is fine and about three times as long
as the short stout apical anterior spur.
Wing greyish hyaline, veins pale brown. Inner cross vein slightly proximad
of apex of first vein and at middle of discal cell. Halteres yellow.
Abdomen fulvous yellow, glossy, with three rather large black spots on the
apical margin of the fifth tergite and none visible on the other tergites though
the apex of the seventh, all of eighth and base of ninth tergites are infuscated.
Possibly this is abnormal. The apex of the abdomen is quite stout, the seventh
and eighth tergites are furnished with more and finer erect hairs than in the
related species, and the apex of the ninth tergite is blackened and has the tip
with a slight production of its anterior angle similar to that shown by Curran
in sikaiana. Curran.
Length : 3-5 mm.
Type : Solomon Islands : Russell Is., Kaylan, 8.XK1934 {R. A. Lever).
This may be sikaiana Curran, but it differs from Curran's description in
so many characters, besides belonging to a different group containing many
closely allied species, that I do not believe it wise to refer it to that species. The
type-locality of sikaiana is Stewart Islands (Sikaiana Is.).
SAPROMYZIDAE FROM THE SOLOMON ISLANDS 143
Homoneura (Homoneura) acrostichalis (de Meijere).
1915, Lauxania acrostichalis de Meijere, Tijdschr. Ent., 58 (suppl.) : 51.
I have seen no specimens of this species from the Solomon Islands, though
Curran has recorded if from Anuda Island and also from Stewart Islands
(Sikaiana Is.). I examined the type-specimen when working on my 1929 paper
and have seen the species from several localities in the Pacific Islands region
and East Indies, but not from the Solomons.
Homoneura (Homoneura) crockeri Curran.
1936, Homoneura crockeri Curran, Proc. Calif. Acad. Sci., 22 : 40.
This species, as I accept it, averages a little larger than acrostichalis, but,
like it, has considerable variation in the black spotting of the abdomen. Usually
there are three dark spots or marks on the fifth to seventh tergites, or these
spots may be confined to one tergite, or even two spots only may be visible.
Both species have usually one or two very minute black hairs on the upper part
of the pteropleura, and the hypopygia are very similar.
Originally described from a single male taken on Guadalcanal Island. I
have a series of specimens from the type-locality, Tulagi, Russell Is., Stewart
Is. (Sikaiana Is.), and Admiralty Is.
Homoneura (Homoneura) hawaiiensis Malloch.
1927, Homoneura hawaiiensis Malloch, Proc. Hawaii, ent. Soc, 6 : 383.
This is probably anuda Curran, agreeing with it in all essential characters.
Two specimens from the Solomon Islands : Tulagi, and Guadalcanal Is.
Homoneura (Homoneura) diversa, sp. n.
$. A small glossy fulvous yellow species, with no dust on the mesonotum, a
pair of small deep black spots on the fifth abdominal tergite, the sixth partly
concealed in the type, so that it is impossible to say whether there are spots
present or not, but they are not on the apex as on the fifth tergite. The deep
black epaulet of the wing is distinctive amongst the yellow species in this report.
Head much as in suspensa, the longest hairs on the aristae about half as
long as width of third antennal segment ; ocellar bristles shorter than the post-
vertical pair, the latter much shorter than the other frontal bristles. Eyes
about 1-25 times as high as long, not much tapered below.
Thorax as in suspensa, the intradorsocentral hairs in about ten irregular
series, the presutural pair only of the acrostichals well developed, a distinct
bristle mesad of the presutural, sternopleurals 2, no pteropleura! hairs evident.
Legs yellow. Fore femur with distinct anteroventral comb on apical half ;
all tibiae with a preapical dorsal bristle, the one on hind tibia short, but longer
than the apical spur.
Wings yellowish hyaline, veins pale brown, neither cross vein clouded.
Inner cross vein almost directly below apex of first vein and slightly proximad
of middle of the discfal cell, outer cross vein about three-fourths of its own length
from apex of fifth vein.
Abdomen glossy fulvous yellow, with a pair of small round black spots at
apex of fifth tergite ; genital processes dark brown, finely haired.
Length : 3 mm.
Type : Solomon Islands : Guadalcanal Is., 26. ii. 1935 (R. A. Lever).
144 JOHN R. MALLOCH
Homoneura (Homoneura) horvathi (Kertesz).
igoo, Sapromyza horvathi Kertesz, Termeszetr. Fiiz., 23 : 260. ,
This is the only species with five dark brown marks on the wing, i.e. one over
each of the cross veins, and one at apex of each vein from second to fourth
inclusive, the last being removed slightly from the tip of the vein. The apices
of third antennal segment and of the palpi are black, and the longest hairs on the
aristae are about as long as the width of the third antennal segment. The meso-
notum has six narrow brown vittae, the lateral marginals very indistinct.
This is probably the species listed by Curran from Catalina Island as puncti-
pennis de Meijere. It is possible that the two names apply to the same species.
Solomon Islands: Tulagi, 4.vii.ig33 (R. A. Lever).
Recorded from Mueo Island under this name by Curran.
MINETTIA Robineau-Desvoidy.
1830, Minettia Robineau-Desvoidy, Mem. pre's. acad. Sci. Inst. France, 2 : 646.
Curran (1936) recorded two species under this generic name. He also placed
in his key two species that he apparently considered later to belong to Homo-
neura. It is the type of costal armature and not the presence or absence of
the intra-alar bristle that is the criterion for the separation of these two genera.
Whether the two species he described and left in Minettia belong here I am unable
to say, as I have not seen his material and have no other from the Solomons
before me at present.
The two species are Minettia surda Curran, which is a small species, 3-5 mm.
in length, mainly shiny black in colour, with plumose arista ; and Minettia
ethdia Curran, slightly larger and mainly rusty reddish yellow in colour, with a
pair of broad brown vittae on the mesonotum that are mesad of the dorsocentral
lines of bristles and extend faintly on to the scutellum.
MAQUILINGIA Malloch.
1929, Maquilingia Malloch, Proc. U.S. nat. Mits., 74 (6) : 35.
There are no species of this genus in the material before me from the Solomons.
In 1936 Curran recorded two new species in the genus. They are M. malaita
Curran and M. matema Curran from the islands after which they are named.
TRYPANEOIDES Tonnoir and Malloch.
1926, Trypaneoides Tonnoir and Malloch, Rec. Canterbury [N.Z.] Mas., 3 : 20.
This genus was erected for the reception of a species from New Zealand and
afterwards I placed 11 additional East Indian species in it when I dealt with the
Oriental Sapromyzidae in 1929. Subsequently several other species have been
added, one or two from Australia. Also in 1929 I dealt with the Sapromyzidae
of Samoa1 and placed in this genus Sapromyza caniventris Bezzi and S. leucosticta
Bczzi, with two additional new species from Samoa. Bezzi's species were described
from Fiji. Now over 20 species are known to belong to the genus. Below I
describe one more.
1 Insects of Samoa, (British Museum (Nat. Hist.)), 6 : 201-213.
SAPROMYZIDAE FROM THE SOLOMON ISLANDS 145
Trypaneoides confusa, sp. n.
?. A black, densely pale grey dusted species, with two chocolate-brown
transverse bands on the face, some black marks on the dorsum of the thorax,
the most conspicuous being a broad band across the mesonotum distinctly behind
the suture, and one on apex of the scutellum between the bases of the apical
bristles.
Head brownish yellow, densely grey dusted, the genae and lower postocular
orbits silvery white dusted, the dust on face more yellowish, face with two
chocolate-brown bands, the upper one broadest and enclosing antennal insertions,
the lower one above epistome, the latter narrowly yellowish grey ; prelabrum
dark brown ; antennae broken except the basal two segments which are blackish,
with dense dark grey dust ; palpi brown. Head wider than thorax, frons over
one-third of the width, wider than long, the vertex rounded, ocelli on the rounded
vertex, very small and closely placed, the ocellar bristles closely placed and strong,
proclinate, postverticals below vertical margin, cruciate, and fully as long as the
ocellars ; anterior orbital moderately long, close to anterior margin. Face
widened below ; eye almost round, about five times as high as gena, the latter
with a series of fine rather long hairs.
Thorax black, densely dark grey dusted, the mesonotum with a curved mark
above each of the humeri, a broad cross band behind the suture, and a narrow
transverse stripe on centre of the posterior margin, black ; scutellum with a
black mark across apex between the apical pair of bristles ; pleura without
black marks. Mesonotum with four pairs of long dorsocentrals, the anterior
pair in front of suture, and four smaller pairs of acrostichals ; sternopleura with
two bristles, the anterior the shorter ; mesopleura with one strong posterior
marginal bristle, and two fine discal bristles below middle that are curved
downward.
Legs testaceous yellow, all coxae and femora infuscated, the legs shrunken
in the type, so that it is impossible to give minute details. Hind femur with the
usual long anteroventral bristle beyond middle ; no definite anteroventral comb
on fore femora ; all tibiae with a preapical dorsal bristle.
Wings whitish hyaline, with brownish-black markings as in fig. 6, a hyaline
mark in apex of first posterior cell. Inner cross vein at middle of discal cell ;
first posterior cell widened at apex. Halteres broken off in type.
Abdomen black, with dense grey dust, three series of irregular black spots,
and the apices of tergites yellowish.
Length : 3 mm.
Type: Solomon Islands: Guadalcanal Is., Papanu, 500 feet, 15.xii.1934
(A'. .-I. Lever).
CEPHALOCONUS Walker.
[86i, 1 ephaloconui Walker, I'm, . Linn. Soc. I. "ml., 5 : 299.
I recently published a paper on this genus, giving the synonymy known to
me.1 One of the two known species is amongst the Solomon Islands material
before me.
Cephaloconus cyprinus (de Meijere).
1.11 ;. Ichlhyomyia cyprinus <le Meijere, Nova Guinea, 9 : 581.
Tulagi, viii.1933 (A'. .1. Lever).
1 0131.1, .Inn. Mag. nut. Hist., (n), 3 : | |
146 JOHN R. MALLOCH
In addition to the foregoing list of species there are several damaged specimens
of the genus Homoneura of which I do not care to offer identifications.
One of these specimens that had come loose in transit and which has lost
the head, legs and wings, but can be placed amongst those with bipunctate
abdomen, has a parasite embedded in the membrane of the venter of the abdomen.
I cannot identify this parasite, and cannot remember any similar occurrence in
the family in my rather extensive examinations from any part of the world.
(MSS. reed. June 22, 1939.)
EXPLANATION OF PLATE I.
Fig. 1. — Steganopsis tripunctifacies, sp. n., wing.
Fig. 2. — Steganopsis varimana, sp.n., wing.
Fig. 3. — Dioides fuscoapicata, sp. n., wing.
Fig. 4. — Monocera furcata, Kertesz, head from above (rj).
Fig. 5. — Homoneura russelli, sp. n., hypopygium of male.
Fig. 6. — Trypaneoides confusa, sp. n., wing.
NOV FT. ZOOL.. 42, ro^o
.MAI. U nil PLATE I.
Solomon Islands Sapromyzidae.
STUDIES ON DIOPSIDAE 147
STUDIES ON DIOPSIDAE (DIPTERA)
By JAMES F. SHILLITO, B.Sc,
South-West Essex Technical College.
(With one plate and six text-figures.)
THE Stalk-eyed Flies have from time to time attracted the attention of many
systematists, but since no critical revision of the whole group has been
completed during the last century the literature is confused, and of the 160
species named many must be regarded as synonyms.
For present purposes it is regarded as most expedient to treat the Diopsidae
as a family and to recognize eight genera ; the following short formal definition
suffices to distinguish this family from others :
Miscidae-Acalypterae, in which the head is produced laterally in
both sexes ; not only the eyes but also the antennae are borne on these
processes (" eye-stalks "). The thorax bears two, sometimes three, pairs
of spines.
Full keys indicating the position of this family are to be found in the works
of Imms (1938), Curran (1934), Lindner (1925), Hendel (1922), etc. A more
detailed subdivision of the Acalypterae has been given by Frey (1921), basing
his system on the structure of the mouth-parts. In his scheme Diopsidae
were placed in the Sciomyzaeformes, close to the families Sepsidae and
Megamerinidae. This agreed with a system suggested by Hendel (1916), who
later (1922) grouped these three families together as Sepsoidea-Sepsaridae.
This paper is intended to serve as an introduction to a series of papers at
present in preparation, in which the species of Diopsidae will be systematically
revised and their nomenclature brought up to date. The first part reviews the
most important contributions to our knowledge of these insects and the determi-
nation of the genera, and in the second part the genus Cyrtodiopsis Frey is revised
and extended.
I. THE FAMILY DIOPSIDAE.
(i) Historical Survey of Taxonomy.
The genus Diopsis was established by Linnaeus (1775) for his single species,
D. ichneumonea ; he pointed out that despite its apparent affinities with various
Hymenoptera, it must belong to the Diptera. Shortly afterwards further species
were described by Dalman (1817) and others, and were placed in this genus.
Fabricius (1805) established a new genus of Stalk-eyed Flies, Achias, which
fell within the original definition of Diopsis Linn. — " capite bicomi, oculis
terminalibus " — but from which it can readily be separated by the position of
the antennae on the eye-stalks. The original distinction in the form of the
antennae given by Fabricius was not correct ; in his key {he. cit., p. 15) Achias
is placed in the section " Antennis parum articulatis deflexis " while Diopsis is
quite unjustifiably placed in the section " Antennis minutis uniarticulatis."
Say (1817) described as D. brevicornis an American species which was
apparently an intermediate form and which was not infrequently regarded as
NOVIT. ZOOL., 42, I 12
I48 JAMES F. SHILLITO
an Achias, but for which Say himself later (1828) erected, as a new genus, Sphyra-
cephala, adding to his definition, " to be placed between Diopsis Linn, and Achias
Fab."
One of the earliest attempts at forming natural groups of these genera was
by Billberg (1820) in the catalogue of insects in his collection ; he arranged a
group (" natio ") Diopsides consisting of four genera, Loxocera, Calobata,
Achias and Diopsis, placing it between similarly constituted groups, Muscaedes
and Conopsides.
The first comprehensive account of these insects was that of Westwood
(1837), in which he gave an excellent summary of both the historical and morpho-
logical aspects of the genus Diopsis, to which he added several new species,
making a total in his list of 31 species. On the status of Sphyracephala, how-
ever, he did not give a decided opinion, though later (1848) he adopted the
name, not however without some misgivings.
In his first paper, which preceded Westwood's memoir, Macquart (1835)
separated Achias from Diopsis, grouping the former with the Muscidae and the
latter with the Sepsideae. Later (1843) he pointed out that this grouping was
for convenience only and that he regarded the differences as sufficient to justify
the erection of a new tribe Diopsideae, though in later papers he did not carry
out his other projected modifications.
Walker (1856, etc.) in his catalogues described some species of Diopsis
and included them in a sub-family of his own creation, Diopsides, ignoring
Billberg's use of this name.
Rondani (1875) published a key to nine genera of his sub-family (" stirps ")
Diopsidinae in which he grouped together all the Stalk-eyed Flies— " charactere
communi capitis ad latera in pedunculos ocelliferos sat aut valde producti " —
and Bigot (1880) , in his second and more important paper on these insects, followed
this principle and added two further new genera. An interesting contrast is
afforded by a more modern interpretation of the affinities of the genera then
grouped together (Table I) :
Table I.— Classification of Stalk-eyed Flies included in the Diopsidinae of
Rondani and Bigot.
I. Genera still retained in Diopsidae :
Diopsis Linnaeus.
Sphyracephala Say.
Synonyms : Hexechopsis Rondani.
Zygocephala Rondani.
Teleopsis Rondani.
Diasemopsis Rondani.
II. Genera now assigned to other families :
Achias Fabricius and its synonym Plegiocephala Rondani in
Ortalidae.
Zygothrica Rondani in Drosophilidae.
Laglaisia Bigot in Ortalidae.
Anaeropsis Bigot in Micropezidae.
JV.B. — Achias (s. str.) excludes at least two species originally placed in that genus but
now grouped with different genera in Trypetidae, namely, Achias maculipennis Westwood
(now Themara) and A. ichneumonea Westwood (now Pelmatops). In addition to the genera
mentioned above, stalk-eyed flies have been described in the following genera of Ortalidae :
Paragorgopsis, Richardia, Megalathoraca and Asyntona.
STUDIES ON DIOPSID \l 149
Subsequent research having exposed the artificiality of Rondani's grouping,
it was discontinued and the true Stalk-eyed Flies have latterly been treated
either as a family, Diopsidae, or as a sub-family, Diopsinae, according to the
status given to the Muscidae-Acalypterae.
Although no attempts have been made to revise the whole family, several
important partial revisions have been published and new genera have been
added since Bigot's time. Loew (1873) had produced a short summary of the
small genus Sphyracephala and allocated to it a fossil species which he described
from Baltic amber ; his paper was the basis of Osten-Sacken's remarks (1882)
on this genus, which in turn led to further revisions by Bezzi (1922) and by
Curran (1928a) (African species only).
The South Asian species were catalogued by van der Wulp (1896) and this
list was extended by Brunetti (1907) ; a later revision of the genera Teleopsis
and Sphyracephala by this author was published posthumously (1928) by which
time it was already more out of date than its editors had realized, for Hendel
(1917) had recognized a fifth genus, Pseudodiopsis (for Diopsis cothamata Bigot
— till then grouped with Sphyracephala by Osten-Sacken, Brunetti, etc.).
Eggers (1915a) discussed the characters and relationships of the African
genera and later (1925) published an important paper, again restricted to the
African species, in which he summarized the literature relating to the region,
gave a catalogue of species and concluded with a general discussion of the problem
of " stalk-eyedness."
Frey (1928) gave a key in which he added the genera Cyrtodiopsis (for D.
(hdnianni Wiedemann) and Megalabops (for D. quadriguttata Walker) to the
five genera already established.
Curran (19286) established a genus Diopsina for a new species from Africa,
and he also (1928c) published a key to the five African genera ; in another paper
(1931) he repeated this key to the genera (with one modification), and also gave
a key to the species of Diasemopsis along with descriptions of fourteen new species.
Subsequently Curran (1934) extended his generic key, ostensibly to cover
all the known genera and also a new genus Microdiopsis, which he immediately
recognized as a synonym of Pseudodiopsis Hendel (loc. cit., p. 495 in corrigenda),
but he omitted Megalabops and Cyrtodiopsis.
At the present time the following eight genera may be regarded as well
established :
Diopsis Linnaeus, 1775.
Sphyracephala Say, 1828.
Synonyms : Hexechopsis Rondani, 1875.
Zygocephala Rondani, 1875.
Teleopsis Rondani, 1875.
Diasemopsis Rondani, 1875.
Pseudodiopsis Hendel, 1917.
Synonym : Microdiopsis Curran, 1934.
Cyrtodiopsis Frey, 1928.
Megalabops Frey, 1928.
Diopsina Curran, 1928.
The following suggestions have been made for subdividing these genera :
Westwood (1837) classified the species in his genus Diopsis into four groups
according to the wing pattern. Eggers (1925) extended this classification by
150 JAMES F. SHILLITO
employing additional characters to form eight groups, mainly as an aid to identi-
fication and without in any way labelling these groups as sub-genera.
Frey (1928) however introduced a sub-genus, Eurydiopsis, for species of
Diopsis (s. str.) having short, blunt teeth on the oral margin ; Brunetti (1928)
also suggested such a division, but hesitated to adopt the principle, since inter-
mediate forms had been described. Curran (1931) also pointed out the possibility
of dividing Diasemopsis in a similar manner.
(ii) Geographical Distribution.
Until fuller records, based on more accurate identification, are available, it
is not possible to give more than the following rough outline of the distribution
of the various genera :
Sphyracephala. — Both Old World (Ethiopian, Oriental and Palaearctic
Regions) and North America (Nearctic Region). Also recorded in
fossil form in amber (Oligocene-Palaearctic Region).
Diopsis, Teleopsis and Megalabops. — Oriental and Ethiopian Regions.
Diasemopsis and Diopsina. — Exclusively from the Ethiopian Region.
Cyrtodiopsis. — Oriental and Ethiopian Regions. (The only record from
Africa is a new species described below, see p. 160.)
Pseudodiopsis. — Malay Archipelago and Ceylon (Oriental Region) and also
Papua and New Guinea (Australian Region).
Our knowledge of the localities in which the last-mentioned genus is known
to exist has been extended recently ; Malloch (1938) has recorded a specimen
from Mt. Lamington in Papua, but the following records have not hitherto been
published :
1. Examination of the holotype of T. bipunctipennis Senior White (1922) —
type locality Ceylon — has shown that this species is undoubtedly congeneric
and possibly conspecific with Pseudodiopsis cothumata (Bigot).
2. A series of specimens of P. cothumata (now in the British Museum (Natural
History)) was taken by Miss Cheesman in the Mount Cyclops region of Dutch
New Guinea in 1932.
These records from the Australian region provide an interesting exception
(the only one known to the writer) to the rule propounded by Osten-Sacken
(1883) which relates to the distribution of the various stalk-eyed flies :
" New Guinea, with its dependencies is the home of the wonderful forms
of Achias, Anaeropsis and Laglaisia, while Diopsis, so abundantly represented
in South Eastern Asia (even in Celebes), has not yet been found in New
Guinea." (It is obvious that Osten-Sacken here means Diopsis (sensu lato).)
(iii) Structure and Habits.
Very little has been published on this aspect of the Diopsidae ; Eggers
(19156) gives a full account (in Russian with English translation) of histo-
logical studies on the eyes of Diopsidae, from which he concludes that the higher
differentiation of the anterior cells gives the insect binocular vision of moving
objects in front ; those cells which are so situated that they cannot see objects in
the field of view of the other eye are not so specialized. In this connection,
it may be recalled that Wiedemann (1830) gave these flies the name " Perspectiv-
fliegen." Eggers implies that sexual differences are not observed in the eyes
of Diopsidae.
STUDIES ON DIOPSIDAE I5I
Morphological studies of a more comparative nature include those of Peterson
(1916) on the head of Sphyracephala brevicomis Say, of Young (1921) on the
thorax and abdomen of the same species, and of Frey (1921) on the mouth-parts
of D. ichneumonea, while Querner (1924) has dealt with the external morphology
of the head in D. apicalis Dalmann.
Concerning the habits of Diopsidae, it is interesting to note that they are
often found on windows, while the species from Transcaucasia described under
the name of Sphyracephala bahadjanedesi by Zaitzev (1919) was collected at a
light ; their normal habitat is the vegetation adjacent to streams. Species of
Sphyracephala appear more especially to choose sunny rocks in streams, from
which they have often been taken in large numbers ; several observations
collected by Sen (1921) tend to emphasise the gregarious nature of these flies.
Of their feeding habits nothing is known ; Sen (loc. cit.) fed specimens on
sugar solution, but it has generally been suggested that they are predaceous —
a supposition based on the structure of the fore-femora but not confirmed by
observation in the field. The structure of the eye-stalks and their function in
providing binocular vision is discussed by Eggers (19156 and 1925) and is regarded
by him as additional support for the predacity theory. Specimens have also
been taken on human faeces by Gibbins in Uganda (MS. note with specimens —
1935), but it was not recorded if they were actually feeding on them.
The first record of breeding habits was that of Mally (1920), who
quoted several instances of specimens being bred from maize stalks but, in
answer to the suggestion that Diopsid larvae are parasitic on maize stem-borers
gave as his impression " ... the larvae live in decaying vegetable matter
such as is commonly left by floods on the banks of streams. The parent flies
are probably attracted to maize stalks by decay induced by the borer."
The details of the life-history of Sphyracephala hearseiana Westwood have
been worked out by Sen (1921), who succeeded in rearing this insect under
laboratory conditions. As his paper is rather inaccessible the following summary
of his method is given here.
Several adults were placed in a glass jar, on the bottom of which was a
layer of earth covered by a large quantity of grass — this was kept very wet ;
sugar solution was fed to the flies when necessary, the jar being kept in a
shady place.
Under these conditions the eggs were laid on the grass about the tenth
day and the remainder of the life-cycle took about three weeks ; the larvae
fed on the decaying vegetation and were found to be particularly susceptible
to drying out.
Sen also described the larval and pupal stages, paying special attention to
the tracheal system, which he regarded as specially adapted to semi-aquatic
conditions of life.
(iv) Determination of the Genera.
Generic distinctions in the Diopsidae are based on wing structure and on
the ornamentation of the thorax. There has been considerable divergence in
the nomenclature of spines and bristles, both in keys and in descriptions,
and the lateral thoracic spines which have been variously called hypopleural,
pteropleural, metathoracic, etc., are here classed as PLEi ROT] RGAL, following
Young (1921) (see text-fig. i, 6). Brunetti's term supra-alar for the paired
spines present only in species of Teleopsis and Megalabops is retained (see
t<xt-fig. 1, 5). Linnaeus (1775) had stated that such spines were not known
152
JAMES F. SHILLITO
Text-fig. i. — Teleopsis sp., showing taxonomic characters, i, Eye ; 2, antenna ;
3, inner orbital bristle ; 4, outer orbital bristle ; 5, supra-alar spine ; 6, pleuro-
tergal spine ; 7, scutellar spine ; 8, apical bristle ; 9, haltere ; 10, collar. i-v,
longitudinal veins.
Text-fig. 2. — Wings of Diopsidae to show venation (markings omitted).
A, Diasemopsis ; b, Pseudodiopsis ; c, Sphyracephala.
STUDIES ON DIOPSIDAE
153
Text-fig. 3. — Heads of Diopsidae. a, Cyrtodiopsis ; b, Diopsis (slightly turned to
show facial teeth) ; c, Sphyracephala ; d, Pseiuiodiopsis ; E, Diasemopsis ; F,
Megalabops ; g, Teleopsis.
Text-fig. 4. — Scutella of Diopsidae. \. Cyrtodiopsis; e, Diopsis; c, Sphyracephala,
n, I hops: 1111.
154 JAMES F. SHILLITO
in other Diptera, but it may be noted that scutellar spines occur in several
genera of the Stratiomyiidae, and that species belonging to the genera Ephippinm
and Negrilomyia of the sub-family Clitellariinae also have well-developed supra-alar
spines comparable with the spines characteristic of Teleopsis. The Asilid genera
Chrysopogon and Codula are characterized by a pair of pre-sutural spiny bristles.
Pleurotergal spines have, as far as can be ascertained, no counterpart in
other families of Diptera.
Names for the bristles cannot be so accurately decided, but it would seem
that the two pairs generally found are best described as notopleural and
intra-alar. In Diopsina there may also be a pair of dorsocentral bristles,
but the specimens of this genus now available for study are too badly damaged
to enable a final decision to be made at the present time.
The following key was originally adapted from Frey's paper (1928) by Dr.
John Smart of the British Museum (Natural History), and I am indebted to
him for permission to include it here in a slightly modified form.
Diopsidae — $$ and $? (see text-figs. 1, 2, 3 and 4).
1. Alula present; scutellar spines truncate and bearing a bristle which is
much longer than the spine .....-•• 2.
Alula absent ; scutellar spines long — their bristles, if present, short,
not longer than the spine (in Diopsina spine and bristle are subequal) 3.
2. Fifth vein extending beyond the discal cell (1st M2) ; both inner and
outer orbital bristles present Sphyracephala.
Fifth vein not extending beyond the discal cell ; only outer orbital
bristles present Pseudodiopsis.
3. Thorax with paired pleurotergal and supra-alar spines ... 4.
Thorax with paired pleurotergal but no supra-alar spines ... 5.
4. Inner orbital bristle set on a small tooth in the middle of the eye-
stalks ; supra-alar spines short ..... Megalabops.
Inner orbital bristle set on a truncate cone towards the base of the
eye-stalks ; supra-alar spines long .... leopsis.
5. Thorax bearing paired bristles on dorsum
Thorax without paired bristles on dorsum
6. Scutellum truncate with a pair of discal bristles
Scutellum normal, without discal bristles
7. Body covered with coarse hairs ; inner orbital bristle strong ; scutellar
spines markedly curved ...... Cyrtodiopsis.
Body with but few sparse hairs ; inner orbital bristle weak ; scutellar
spines not markedly curved .....-• Diopsis.
6.
7.
. Diopsina.
DlASEMOPSIS.
With the assistance of the figures illustrating characteristic features,
undamaged specimens should be readily determined by this key, but valuable
confirmatory evidence may be obtained by examining the fore-femora, either in
situ, or preferably after clearing and mounting (text-fig. 5a and 56). Various
authors have made reference to the ornamentation of the fore-femora without
apparently realizing the value of this character as a diagnostic feature in deter-
mining the genus — especially in differentiating between Diopsis (text-fig. 5a, a)
and Diasemopsis (text-fig. 5&, d).
In all the Diopsidae examined by the writer, the ventral surface of the
STUDIES ON DIOPSIDAE
155
fore-femora bears a double row of small peg-like teeth, which vary considerably
in size and position in the different species. In addition to these, species
belonging to Diasemopsis, Sphyracephala and Pseudodiopsis (text-fig. 56)
bear spiny bristles alongside the small teeth. Brunetti (1926) was therefore
correct in assuming that his Diopsis fuscivenis would be more correctly grouped
with Diasemopsis, while an examination of Bigot's cotypes of D. dubia has
shown that this species too must be regarded as a Diasemopsis (cf. Brunetti
(1928) and Curran (1931)).
Text-fig. 5a. — Fore-femora of Diopsidae. A, Diopsis ; B, Cyrtodiopsis ; c, Teleopsis.
Tkxt-fig. 56. — Fore-femora of Diopsidae (cont.). D, Diasemopsis ; E, Sphyracephala ,
f, Pseudodiopsis.
All species of Cyrtodiopsis and some species of Teleopsis have distinctly
hairy fore-femora, the hair being much coarser in the former than in the latter.
The extraordinary specialized structure (text-fig. 5a, b) found on the fore tibiae
and the corresponding modification of the femora of some males of Cyrtodiopsis
dirfmuiuii (Wiedemann) will be dealt with in a subsequent paper.
Owing to the damaged state of the type specimen of Diopsina ferruginea
Curran (19286) no description of the scutellar spines was given. Specimens
from the British Museum (Natural History) were sent to Dr. Curran, who very
kindly confirmed that they wire congeneric ; the drawing of the scutellum
(text-fig. 4, n) was made from one of these specimens and a fuller description
will be given later in this series.
156
JAMES F. SHILLITO
The accompanying table (Table II) summarizes the data relating to chaeto-
taxy, the nomenclature of spines and the bristles of the fore-femora given above :
Table II.
ia, intra-alar ; sa, supra-alar; np, noto- pleural ; DC, dorsocentral.
1 Except D. absens Brunetti, in which the outer bristle is stated to be absent.
2 Except D. aethiopica Rondani, in which the inner bristle is weak (fide Curran (1931)
II. THE GENUS CYRTODIOPSIS FREY.
(i) Taxonomy.
The only direct reference to this genus in the literature is its definition by
Frey (1928) in his paper on the Philippine Diopsidae ; the genus was defined
in the key and a short note added about the " peg and hollow " structures of
the fore-leg (see text-fig. 5, b) in certain males of C. dalmanni (Wiedemann),
the genotype.
A study of the literature and an examination of material in the British
Museum (Natural History) has brought to light the following important facts :
1. D. quinqueguttata Walker and D. villosa Bigot belong to this genus ; the
description of the former (Walker, 1856) is characteristically inadequate, but the
type is in good condition (in the British Museum) and shows the covering
of hairs that is characteristic of this genus, which is so clearly emphasized by
Bigot in his description of D. villosa — " Sparsim et undique villosa." The
description of D. villosa contains the statement " ore bispinosa," whereas in
fact Bigot's co-types (now in the collection of Mr. J. E. Collin) show no facial
teeth and there is therefore no justification for separating these two species.
2. Curran (1936) was apparently unaware of Frey's paper, and two species
mentioned in his paper, the one identified as D. ferraginea Roder (1893) and the
other which he named D. whitei as a new species, both recognized by him as
related to D. dalmanni Wied., must also belong to Cyrtodiopsis.
Specimens agreeing well with the published data of both these species have
been found mixed with C. dalmanni in the British Museum. However it is quite
obvious that the specimens referred by Curran (loc. cit.) to D. ferraginea Roder
were misidentified, since that species, of which the writer has, through the courtesy
STUDIES ON DIOPSIDAE 157
of Dr. W. Ludwig of the Zoologisches Institut, Halle, examined the type, belongs
to Megalabops (as Frey (1928) suggested) and is not related to dalmanni. This
species is re-described below as C. currant sp. n.
3. It is very surprising that Brunetti (1928) should have described a typical
specimen of C. dalmanni (Wiedemann) as a new species, Teleopsis truncata,
especially since he drew attention to that specialization in structure of the fore-
leg which had been figured in previous accounts of dalmanni.
4. Also, in a collection of Diopsidae made by the author in Uganda, a new
species has been found which must be attributed to this genus ; this represents
the first record of this genus from the African continent, the other species being
exclusively from the Oriental Region.
(ii) Generic Characters.
The species now added to the genus do not call for any modifications of the
generic characters given by Frey (1928). The following formal definition may
be given :
Cyrtodiopsis Frey.
Diopsidae characterized by a coating of comparatively coarse hairs on most
parts of the body. The wings are without an alula ; the thorax bears pleurotergal
but no supra-alar spines and it is free from bristles ; the scutellar spines are strong
curved, hairy, occasionally with a very weak bristle or hair terminally situated.
The inner orbital bristle is strong. The abdomen is markedly clavate.
Genotype : Diopsis dalmanni Wiedemann (1830).
In many respects this genus is intermediate between Diopsis, which has
straighter scutellar spines and very weak inner orbital bristles, and Teleopsis,
which has supra-alar spines. Certain species of Diasemopsis have been described
as hairy, but these all have spines on the ventral surface of the fore-femora and
bristles on the dorsum of the thorax.
The following five distinct species fall within the above definition of the
genus :
1. C. dalmanni (Wiedemann).
2. C. quinqueguttata (Walker).
3. C. whitei (Curran).
4. C. currant sp. n.
5. C. africana sp. n.
(iii) Key to Species of Cyrtodiopsis (see Plate I, figs. a-g).
1. Head and body mainly black ..... quinqueguttata Walker
Head and body brownish, never black ...... 2.
2. 1st posterior cell (R5) with no median pale spot . . . africana sp. n.
1st posterior cell (Rs) with median pale spot ..... 3.
3. This wing spot small, isolated, distinct ; the sub-apical brown band on
the wing greater than the apical pale bands in width . currant >\>. a.
This wing spot larger and part of the transverse pale band ; the dark
sub-apical band on the w ing not greater than the apical bands in width 4.
4. Dorsum brown pollinose ; inner orbital bristle on a strong tubercle
ltd Curran
Dorsum glossy, not pollinose ; inner orbital bristle on a weak tubercle
dalmanni Wiedemann
158 JAMES F. SHILLITO
(iv) Descriptive Notes on the Species of Cyrtodiopsis.
1. Cyrtodiopsis dalmanni (Wiedemann).
Diopsis dalmanni Wiedemann, 1830, Auss. Zweifl. Ins. 2 : 560.
Diopsis dalmanni Wiedemann, Westwood, 1837, Trans. Linn. Soc. London, 17 : 309, 547.
Diopsis attenuata Doleschall, 1856, Natuurk. Tijdschr. Ned.-Ind., 10 : 413.
Diopsis latimana Rondani, 1875, Ann. Mus. Stor. Nat. Genova, 7 : 444.
Diopsis lativola Rondani, 1875, Ann. Mus. Stor. Nat. Genova, 7 : 445.
Diopsis albimana Rondani (in error for latimana), 1875, Ann. Mus. Stor. Nat. Genova, 7 : 445.
Diopsis dalmanni Wied., van der Wulp, 1897, Tijdschr. Ent., 40 : 184.
Diopsis dalmanni Wied., de Meijere, 1908, Tijdschr. Ent., 51 : 115.
Teleopsis truncata Brunetti, 1928, Ann. Mag. Nat. Hist., (10), 2 : 277.
Cyrtodiopsis dalmanni (Wied.), Frey. 1928, Notul. ent. Helsingfors, 8 : 70.
Of the descriptions listed above, van der Wulp's (in Dutch) and de Meijere's
(in German) are so complete that it is not necessary to redescribe this species
here.
The species is very variable and this no doubt is responsible for the list of
synonyms given above. It is possible that it could be sub-divided into geo-
graphical races, but until much longer series are available, such sub-division is
not practicable. From the British Museum material, however, it is apparent
that there is considerable variation in the wing pattern. Specimens from Java
(the type locality) show two forms (Plate I, figs. A, b) with intermediates, while
specimens from the F.M.S.,Siam and Sarawak (Plate I, fig. e) are slightly different
from both — the sub-apical pale band, usually composed of three spots, is absent
and the transverse dark band is both darker and wider than usual, thus approach-
ing in form the pattern of C. currani sp. n. (Plate I, fig. f).
The variation of this species is being studied biometricalry, and series from
as many different localities as possible are required, but it is more especially
important to know if the species exists in Burma, and if so, in which form it is
found.
2. Cyrtodiopsis quinqueguttata (Walker).
Diopsis quinqueguttata Walker, 1857, /. Linn. Soc. London, 1 : 36.
Diopsis villosa Bigot, 1874, Ann. Soc. Ent. Fr., 5 : 4, 114.
This species is readily distinguished from others in the genus by its black
appearance — in which it bears a superficial resemblance to D. subnotata West-
wood, which is found in the same region but which is generically distinct.
Rondani (1875) pointed out that the species he was describing as D. lativola
sp. n. and which is now regarded as synonymous with C. dalmanni (Wiedemann),
was apparently related to D. quinqueguttata Walker, but he did not give his
reasons.
The type of quinqueguttata is in fair condition, but it has been thought
advisable to give the following re-description from fresh material, collected by
Hobby & Moore in Sarawak in 1932 and now preserved in the British Museum.
$ and $. Length : 8 mm. Eye-stalk : 5 mm. Wing : 5i mm.
Head : black, face glossy black, not markedly protuberant, as in dalmanni ;
epistomium orange, facial teeth absent. The eye-stalks are darker at the base,
considerably lighter at the tips except behind the eyes — sparsely hairy all over.
Antennae typical in form, brownish with black arista. Inner orbital bristle long,
erect, slightly curved ; outer orbital bristle strong and long.
Thorax : dull black with pale pollinose triangular patches just before the
wings on the dorsum, also before and behind the wings on the pleura. Collar
very pale pollinose. Scutellar spines strong and hairy.
STUDIES ON DIOPSIDAE 159
Wing : venation typical, though careful examination of the wing shows that
the fifth vein (M4 + Cut) and the discal cell (ist M2) differ slightly from the
standard form (see Plate I, fig. n). Wing pattern as follows : distal third dark
except for a pale apical spot, middle third not quite so dark with a light spot
at the foot of the discal cell and a similar spot on the anterior margin opposite.
Legs : fore coxa pale yellowish, femur yellowish basally, but darker
apically and not very incrassate. Tibia black and tarsus, except for ist segment,
light. Middle legs dark brownish except for the apical half of the femur, which
is light. Hind legs dark brown except for the basal tenth of the femur,
which is distinctly paler.
Abdomen : mainly black with a sparse covering of hairs and with markings
as follows : first two segments very pale pollinose, third segment with a pair of
clear silvery-white patches on sides, fourth segment pale brown pollinose on
most of the basal half and two silver-white patches on the sides. Sixth segment
pale pollinose.
3. Cyrtodiopsis whitei (Curran).
Diopsis whitei Curran, 1936. Antcr. Mus. Nov., 833 : 1.
A single specimen in the British Museum with the following data : " In
jungle, Assam-Ganhati 21.x. 1920 R. Senior White" agrees exactly with
the original description, but unfortunately it has lost its abdomen (Plate I, fig. c).
There is nothing to add to the complete description given by Curran.
4. Cyrtodiopsis currani sp. n.
D. ferruginea Curran nee Roder, 1936, Amer. Mus. Nov., 833 : 2.
This species is similar to C. dalmanni (Wiedemann), but is not so hairy ; the
wing pattern is slightly different, but the inner orbital bristle is its most dis-
tinguishing feature, its structure resembling that of species of Teleopsis.
Length : 5 mm. Eye-stalk : 6 mm. Wing : 4^ mm.
cJ and $. Head : face yellow-brown, darker above the dark transverse
suture and black behind the eye-stalk. Antennae pale brown, with pale pilosity,
arista long. Inner orbital bristle on short conical base, stout ; outer orbital
bristle short, erect.
Thorax : brown, darker posteriorly, slightly brown pollinose anteriorly ;
collar shining brown with dark sutures ; scutellum pollinose, scutellar spines
basally concolorous with scutellum but darker apically, hairy.
Wing : mainly dark, hyaline at base ; tip tinged with brown, a narrow band
of three spots separating this from the dark portion of the wing ; three median
pale spots distinctly separate ; a pale band nearer the base (Plate I, fig. f).
Legs : fore coxa and femur reddish brown, tibia much darker, first
tarsal segment dull; dark, the remaining segments much lighter. Middle ami
hind legs similar, femora darkened distally, tibiae very dark brown, tarsi dark
with black hairs.
Abdomen: first segment dark brown, second somewhat darker, mat! ; third
light brown basally, darker where it widens ; fourth dark brown, nearly black
at edges with white spots on either side ; fifth and sixth segments blackish brown.
Siam — Bang Kloi, 19. xi. 1933. Holotype, A*.
Siam — Non Guri, 21. xi. 1933. Allotype, ?.
Both collected by W. R. S. Ladell and now in the British Museum.
1 6c
JAMES F. SHILLITO
Cyrtodiopsis africana sp. n. (Text-fig. 6.)
This is the only African species of this genus known at present ; it can
readily be distinguished from the Oriental species by its much smaller size, and
by the form of the wing-markings (Plate I, fig. g), in which it closely resembles
Diopsina ferruginea Curran (1928a), though this latter species has (as shown
by a specimen in the British Museum) setigerous scutellar spines and is generically
distinct.
tj and ?. Dark reddish brown. Length : about 5 mm. Eye-stalks : 4 mm.
Wing : 4 mm.
Text-fig. 6. — Cyrtodiopsis africana sp. n.
Head : dark reddish brown and markedly hirsute ; upper parts of the face
prominent, roughened ; lower parts glossy, with two small but distinct teeth.
The black suture on the lower side of the eye-stalks extends nearly to the oral
margin. Eye-stalks strong, glossy yellow-brown in the middle but darker
apically, with a sparse covering of hair on all surfaces. Ocellar spot black,
prominent, ocelli showing light by contrast. Inner orbital bristle strong, arising
from a small black conical base ; outer orbital bristle much stronger but not so
long, and with a similar base. Antennae brownish, the third segment covered
with fine light-coloured pile.
Thorax : glossy brown, showing the sutures clearly, somewhat darkened ;
hairs long, black, arranged in rows on dorsum. Pleurotergal spine strong with
black tip. Scutellum distinctly convex, deeper than long (as in Diopsina, but
bearing hairs and not distinct bristles) ; scutellar spines strong, slightly curved
STUDIES ON DIOPSIDAE Idl
distally — proximal third light, median third dark, distal third lighter than the
middle but darker than the basal third; more than 15 long black hairs on each
spine, evenly distributed.
Wing : mainly brown, showing seven clear spots arranged as shown (Plate I,
fig. g) (cf. also the illustration of the wing of Diopsina ferruginea Curran (19286 :
183), i.e. a subapical row of three contiguous spots, two centrally placed spots,
one from the anterior margin crossing the second vein and extending to the third
vein ; the other on the posterior margin at the end of the discal cell. The other
two spots form a nearly complete white band across the basal third of the wing.
Legs : conspicuously hairy ; fore : coxa brown, slightly white pollinose
on the lower surface, which is somewhat flattened ; femur not very incrassate,
much darker brown, with black hairs on the ventral surface, which also bears
two rows each of five very small teeth on the distal fifth ; tibia very dark brown
uniform all over ; tarsus very dark on dorsal surface, but with a golden pubescence
on the underside. Middle legs resemble the hind legs, but have no pubescence
on the tarsi. Hind femur dark brown with dark spot apically and a terminal
spine ; tibia dark brown and tarsus dark on upper surface, golden pubescence on
lower.
Abdomen: strongly clavate with terminal segments reflexed under (in
dried specimens). First segment slightly pollinose and therefore appears lighter
than the body. Second segment brown, blacker towards the edges distally.
Third segment glossy brown except on the distal margin, which is dark-banded ;
this segment is considerably wider distally. Fourth segment appearing more or
less hemi-spherical from above, shining dark brown, rather more hirsute than the
third segment. Other segments not normally visible.
Uganda, W.P., Nyakasura, Toro. Holotype, <$, 29.1.1935.
Uganda, W.P., Mpanga Bridge, nr. Fort Portal. Allotype, ?, 5.^.1936.
Paratype, <J, same data as holotype.
All collected by the author and now in the British Museum.
In life this Diopsid has a strong resemblance to an ant, the eye-stalks giving
the illusion of strong antennae and the wings held flat over the abdomen serve to
exaggerate the petiolation — an illusion that is more marked in the case of D.
circularis Macquart, in which the two circular marks on the wings coincide when
the wings are held in their normal resting position.
Acknowledgments.
In conclusion, it is my privilege to express my thanks to the British Museum
(Natural History) for granting me facilities for carrying out this study, and more
(■specially to Dr. F. W. Edwards, F.R.S., and to Dr. John Smart for the interest
they have shown in the work and for the helpful advice they have so willingly
offered. Also to Mr. H. Oldroyd, M.A., of the British Museum, and to Dr. F.
van Emden, of the Imperial Institute of Entomology, for information relating to
other families of Diptera. Mr. G. H. E. Hopkins, M.A., Senior Entomologist,
Medical Dept., Uganda Protectorate, very kindly referred me to Mally's paper
(1920), which I should otherwise have overlooked. For the loan of type-. 1 am
very much indebted to Mr. J. E. Collin, who now has part oi the Bigot Collection,
and to Dr. \V. Ludwig of the Zoologisches Institut, Halle, who sent Roder's type
of D. ferruginea for examination.
l62 JAMES F. SHILLITO
BIBLIOGRAPHY.
Bezzi, M. 1922 Note sur la presence en Algerie du Sphyracephala Hearseiana Westw. de
l'lnde et sur la synonymie de ce Diptere. Bull. Mus. Hist. Nat. Paris. 28 : 69-72.
Bigot, J. M. 1874 Dipteres nouveaux ou peu connus. I. Sur le genre Diopsis, especes
nouvelles. Ann. Soc. Ent. Fr., (5), 4 : 107-115.
1880 Dipteres nouveaux ou peu connus. XIX. Diopsidae. Ann. Soc. Ent. Fr.,
(5), 10 : 90-94.
Billberg, G. J. 1820 Enumeratio insectorum in Museo Billberg. Holmiae.
Brunetti, E. 1907 Notes on Oriental Diptera. I. Note on Sphyracephala hearseyana
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7 : 163-166.
1926 New Species of Diopsidae from the Belgian Congo. Rev. Zool. Bot. afr.
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- 1928 Revisionary Notes on the Diopsidae. (Edited by C. H. Curran ; re-edited
by E. E. Austen.) Ann. Mag. Nat. Hist., (10). 2 : 266-273.
Curran, C. H. 19283 Review of the African Species of Sphyracephala. Ann. Mag. Nat.
Hist., (10), 2 : 274-275.
— 19286 New Stratiomyidae and Diopsidae from the Belgian Congo (Diptera). Amer.
Mus. Nov., 324.
1928c Diptera of the American Museum Congo Expedition. Bull. Amer. Mus.
Nat. Hist., 58 : 167-187.
1931 Descriptions of new species of Diasemopsis Rondani. .-J mer. Mus. Nov., 463.
1934 North American Diptera. New York.
1936 A new Diopsis from India, with notes on others (Diptera). Amer. Mus.
Nov., 833.
Dalman, J. W. 1817 Anmarkningar vid slagtet Diopsis jemte beskrifningar och teck-
ningarpa trenne dithorande nyaarter. K. Svenska VetenskAkad. Hand!., 38 : 211-219.
[The descriptions are repeated in the same author's Analecta Entomologica, Holmiae
(1823).]
Eggers, F. O. 1915a On some new and incompletely known species of the family
Diopsidae from British East Africa. [In] Dogiel and Sokolow : Sci. Results of Zool.
Exped. to B.E.A. and Uganda in 1914, 1, pt. 6.
1915k On the structure of the stalked eyes of Diopsidae. [In] Dogiel and Sokolow :
Sci. Results of Zool. Exped. to B.E.A. and Uganda in 1914, 1, pt. 7.
1925 Diopsiden aus Deutsch-Ostafrika. Mit einem Nachwort iiber die Stielaugen
der Diopsiden. Zool. Jahrb. Jena (Syst.), 49 : 469-500.
Fabricius, J. C. 1805 Sy sterna A ntliatorum. Brunsvigae.
Frey, R. 1921 Studien iiber den Bau des Mundes der niederen Diptera Schizophora.
Acta Soc. Fauna Flora Fenn., 48, no. 3.
1928 Philippinische Dipteren. V. Diopsidae. Notul. Ent. Helsingfors, 8 : 69-77.
Hendel, F. 1916 Beitrage zur Syst. der acalyptraten Musciden, etc. Ent. Mitt., 5 : 294-
299.
1917 Beitrage zur Kenntnis der acalyptraten Musciden. Dlsch. Ent. Z., 1917:
33-47-
— 1922 Die Palaarktischen Muscidae acalyptratae Girsch., etc. Konowia, Wien,
1 : 145-160.
Imms, A. D. 1938 Text-book of Entomology . 4th ed. London.
Lindner, E. 1925 Die Fliegen der Palaearktischen Region. Stuttgart.
Linnaeus, C. 1775 Dissertatio Entomologica Bigas Insectorum sistens, etc. Upsaliae.
Loew, H. 1873 Ueber die Arten der Gattung Sphyracephala Say. Z. ges. Naturwiss.,
42 : 103-105.
Macquart, J. 1835 Histoire Naturelle des Insectes. Dipteres, 2.
1843 Dipteres exoliques, 2.
Malloch, J. R. 1938 Papuan Diptera. I. Diopsidae. Proc. Linn. Soc. N.S.W., 43 :
437-8.
Mally, C. W. 1920 The Maize Stalk-Borer. Bull. Dept. Agric. S. Africa, 3 : 89.
Meijere, J. C. H. de. 1908 Stud, iiber Siidostasiatische Dipteren. II. Tijdschr. Ent.
Amsterdam, 51 : 103-330.
Osten-Sacken, C. R. 1882 Diptera from the Philippine Islands, brought home by Carl
Semper. Berl. ent. Z., 26 : 187-246.
- 1883 Enumeration of the Diptera of the Malay Archipelago collected by Prof. O.
Beccari. Supplement. Ann. Mus. Stor. nat. Genova, 18 : 10-20.
Peterson, A. 1916 The head-capsule and mouth parts of Diptera. Illinois Biol.
Monog., 3.
Querner, F. 1924 Zur Kenntnis des Kopfes und der Mundteile von Diopsis apicalis
Dalm. Denkschr. Akad. wiss. Wien, 99 : 283-288.
STUDIES ON DIOPSIDAE 163
Rodek, V. von. 1893 Enumeratio Dipterorum quae H. Fruhstorfer in parte meridionale
insulae Ceylon legit, lint. Nachr. 19 : 234-236.
Rondani, C. 1875 Muscaria Exotica Musei civici Januensis observata et distincta. Ann.
Mus. Stnr. nat. Genova, 7 : 421-464.
Say, T. 1817 Descriptions of several new species of North American Insects. /. Acad.
Nat. Sci. Philad., 1 : 19-23.
1828 American Entomology (Philadelphia).
[Reprinted in Say. T. 1869. A description of the Insects of North America, 1 : 116-117.]
Sex, S. K. 1921 Life Histories of Indian Insects. Diptera : Sphyracephala hearseiana
Wt-stw. Mem. Dept. Agric. India (Ent. Series), 7, no. 6 : 33-38.
Senior White, R. A. hi:j New species of Diptera from the Indian region. Mem.
Dep. Agric. India. Ent. 7 : 107-169.
Walker, F. 1856 Catalogue of Dipterous Insects collected at Singapore and Malacca.
J . Linn. Soc. London, 1 : 36.
Westwood, J. O. 1837 On Diopsis, a genus of Dipterous Insects. Trans. Linn. Soc.
London, 17 : 283-313, 543-50.
1848 The Cabinet of Oriental Entomology, etc. London.
Wiedemann, C, R. W. 1830 Auss. Zweifl. Ins., 2.
Wn.p, F. M. van der 1896 Catalogue of the Described Diptera from South Asia. The
Hague.
Young, B. P. 1921 On the attachment of the Abdomen to the Thorax in Diptera. Mem.
Cornell agric. Exp. Sta., 44.
Zaitzev, F. A. 1919 A new element of the tropical fauna in the Transcaucasia (Diptera,
Diopsidae). Bull. Mus. Caucase, Tiflis, 12 : 1-6.
(MSS. reed. Jan. 20, 1940.)
PLATE I.
Wing pattern of species of Cyrtodiopsis. (x 12.)
a, b. C. dalmanni (Wied.) from Java.
c. C. whitei (Curran).
l). C. quinquegutlata (Walk.).
E. C. dalmanni (Wied.) from Sarawak.
f. C. currant sp. n.
G. C. africana sp. n.
NOVIT. ZOOL., 42, I 13
NOVIT. ZOOL., 42, 1940.
SHII.LITO— PLATE I.
Wing pattei n oi ' yrtodiopsis spp.
Son, ttf.
SUBFAMILIES STENOSMYI.INAE AND KALOSMYLINAE 165
A REVISION OF THE OSMYLID SUBFAMILIES
STENOSMYLINAE AND KALOSMYLINAE
(NEUROPTERA)
By D. E. KIMMINS,
Department of Entomology, British Museum (Natural History).
(With eight plates and twenty-eight text-figures.)
THE Osmylidae, as a whole, have attracted the attention of many neurop-
terists, notably Hagen, McLachlan and Kriiger. Many of them are large-
winged insects, with well-marked pattern and densely reticulated wings. These
features have led to the description of many forms, but unfortunately little
attention has been paid to genital structure in either sex, and it is probable that
many synonyms have been created, especially in the Osmylinae and Spilos-
mylinae. Kriiger, in his revision of the family in 1913-1914, based his classifi-
cation almost entirely on venation and placed much reliance on rather minute
venational characters. In some species at least, further material has shown
these characters to be variable, even differing on opposite sides of the same
insect. In spite of this, Kriiger's work forms a very useful basis for our classifi-
cation. Nevertheless in the Osmylidae the venation, being complicated, is by
no means stable, and should be used with some reserve as a basis for determination.
The recent acquisition by the British Museum of the historic collection of
Neuroptera built up by McLachlan, and also the types and many of the specimens
of Neuroptera from the Tillyard collection, has enabled me to make a more
careful study of the members of the two subfamilies Stenosmylinae and Kalos-
mylinae. These subfamilies form a convenient unit for study, inasmuch as the
genera (with the exception of Isostenosmylus) are restricted in their distribution
to Australia (excluding N. Queensland), Tasmania, New Zealand and Chile.
The exception is also South American, but it has a more northerly range, covering
Ecuador, Southern Brazil and Peru. Further collecting in Chile and Patagonia
may well extend the range of Isostenosmylus southward and will probably discover
further species of these subfamilies. The major part of the Australian Osmylidae
belongs to these two subfamilies, the exceptions being Eidoporismus pulchellus
Esben- Petersen, Porismus strigatus (Burm.), and one or two Spilosmyline species
of Papuan origin which have reached North Queensland.
The structure of the male genitalia confirms the opinion that the Osmylidae
are among the more primitive members of the true Neuroptera. In most
Neuroptera tin- tenth sternite in the male has been retracted within the abdomen
and has become much modified from the normal U-shaped sternite to a saddle or
arch-like structure. In these subfamilies are found examples of a tenth sternite
which is still external and covered with hairs (Stenosmylus, Oedosmylus), as well
as intermediate forms leading to the more or less internal structure of Isostenos-
mylus. In many Osmylidae there is found in the male a pair of eversible, finger-
like scent-glands, opening on the dorsum between the eighth and ninth abdominal
tergites. These are present in the Kalosmylinae, but appear to be absent in the
Stenosmylinae, in which the eighth and ninth tergites are fused.
novit. zool., 42, 1 14
l66 D. E. KIMMINS
The value of the male genitalia as a means of specific identification varies in
different genera. In Stenosmylus, Oedosmylus and I sostenosmylus these structures
are very helpful, particularly in the latter genus, in which the wings are rather
uniform. In Kempynus and Euosmylus they are of little use ; in fact, the male
genitalia of Euosmylus are practically indistinguishable from those of some
species of Kempynus. In the female the form of the eighth sternite and the
reflexed lobe hinged to its apex offer good characters. Another female structure
which is of assistance is the form of the accessory glands. In the Stenosmylinae
they take the form of a pair of slender, serpentine tubes, each terminating in
a small oval sac. In the Kalosmylinae this sac is more elongate, generally
constricted about midway and differs in form in the various species.
To examine the genitalia properly it is necessary to remove the apex of the
abdomen and to clear it by boiling in caustic potash solution. After neutralizing
the alkali with acetic acid, the preparation is stained in carbol-fuchsin and
mounted in Canada balsam in the usual manner. Staining is necessary to reveal
the shape of the basal parts of the tenth sternite. These structures gradually
thin down to a very fine edge which, if unstained, becomes invisible in Canada
balsam. All the drawings in this paper have been made from cleared specimens
in clove oil, before mounting. The various parts, shown separately, are not
necessarily drawn to the same scale, as this would mean an undue reduction of
some drawings.
The photographs of the wings have been made primarily with the object
of showing the pattern and in some instances this has been slightly emphasized,
with the result that venation in some areas is a little obscured. It was found
that illuminating the wings from behind only, whilst giving an excellent rendering
of the venation, tended to lose the finer shades of the pattern. Pale markings
formed by areas in which the veins were whitish were lost entirely, the veins,
being in silhouette, reproducing as black. Frontal illumination was therefore
used, with a white background some distance behind the wings. The only
disadvantage of this method is the slight tendency to reflection of the illuminant
from certain areas, unless the wing be completely flattened between glasses.
STENOSMYLINAE Kriiger.
Stenosmylinae Kriiger, 1913, Stettin, ent. Ztg., 74 : 23, 105-115, 214.
Anterior wing with one cross- vein in subcostal area, medius forked far
distant from base. Posterior wing without a sinuous cross-vein from the base
of the medius to the radial sector. (In some genera, Stenosmylus, Oedosmylus,
there may occur a feeble or incomplete sinuous cross-vein between the medius
and the stem of the radius.) Cu2 long. In the male apparently no eversible
scent-glands opening between the eighth and ninth tergites.
Key to Genera.
1. Wings brownish or black, with conspicuous opaque, white, apical, patches
Euporismus Tillyard.
Wings without such apical patches . . . . . . .2.
2. In anterior wing, first cross-vein to M beyond the basal corneous spot
arises from the stem of Rs ; M forks near the margin of the wing . 4.
In anterior wing, first cross- vein to M beyond the basal corneous spot
arises from the first branch of Rs ; M forks nearer the centre of the wing
a few cells beyond the second corneous spot ..... 3.
SUBFAMILIES STENOSMYLINAE AND KALOSMYLINAE
167
3. In anterior wing of $, part of Cuj and Cu2 or IA thickened ; Australian,
Tasmanian species Oedosmylus Kriiger.
In anterior wing of $, these veins normal ; S. American species
Isostenosmylus Kriiger.
4. A2 in anterior wing extends to level of origin of Rs Stenosmylus McLachlan.
This vein extends to level of origin of first branch of Rs.
Stenolysmus gen. nov.
STENOSMYLUS McLachlan.
(PI. I.)
Stenosmylus McLachlan, 1867, J. Linn. Soc. Land. (Zool.), 9 : 267.
1868, Brauer, Verh. Zool.-Bot. Ges. IVien, 18 : 398 ; 1913, Kriiger, Stettin, ent. Ztg., 74 :
106-112.
Genotype (fixed by McLachlan, 1867) : Osmylus tenuis Walker, 1853.
In anterior wing, first cross-vein to the medius beyond the basal corneous
spot arises from the stem of Rs ; A2 extends to level of origin of Rs. <$. Tenth
sternite largely external, hairy, varying in shape according to species. $. Hinged
lobe of eighth sternite slender, bifid at apex ; valves of ninth segment not heavily
fringed.
Distribution : Australia.
Stenosmylus tenuis (Walker).
(PL I, fig. 2 ; text-figs. 1, 2.)
Osmylus tenuis Walker, 1853, List Neur. Ins. Brit. Mus., 2 : 234 ; Hagen, 1866, Stettin, ent.
Ztg., 27 : 455.
Stenosmylus tenuis (Walker), McLachlan, 1867, /. Linn. Soc. Lond. {Zool.), 9 : 267 ; Kriiger,
1914, Stettin, ent. Ztg., 75 : 57, 114-117.
Walker's description of the general appearance is adequate, and the only
amplification I propose togive is the description of the $ and ? genital appendages.
cy. Eighth and ninth tergites fused dorsally ; ninth sternite short. Anal
Text-fig. i. — Stenosmylus tenuis (Walker), $ type, a, apex of abdomen, lateral ;
parameres, lateral ; c, the same, dorsal.
i68
D. E. KIMMINS
plates from the side broadly rounded, with a large group of trichobothria. Tenth
sternite external, triangular from above, from the side projecting in a stout hairy
finger. Its lower basal margin extended downwards in a pair of thin chitinized
plates. Between these plates lie the parameres, slender, curved rods, flattened
and fused basally. The fused portion bears on its upper surface a pair of triangular
flaps. Apices of the parameres dilated.
$. Seventh sternite simple, apical margin not produced. Eighth tergite a
little shorter than its depth ; sternite broad, convex basally and concave apically.
Text-fig. 2. — Stenosmvlus tenuis (Walker), $ paratype. a, apex of abdomen, lateral ;
b, eighth sternite, ventral ; c, the same, lateral.
About mid-way the lateral margins are each produced downwards in a short,
triangular tooth, carrying a number of setae. To the apex of the sternite is
hinged a slender process, rather shorter than the sternite, directed basally. This
process, which has a somewhat dilated base, is from the side parallel-sided ;
from beneath the apex is expanded and has a small V-shaped excision at the
centre of its margin. Ninth tergite short dorsally, side lobes pyriform. Valves
of the ninth segment straight, lanceolate, apex with a minute style. Anal plates
rounded, group of trichobothria large and ovate.
From Walker's series of three I select as type a male from Van Diemen's
Land. The other specimen from this locality is a female, as also is the example
SUBFAMILIES STENOSMYLIN AE AND KAI.OSMVLINAE
i6g
from Adelaide. In addition to these localities I have also seen examples from
E. Australia and New South Wales.
Stenosmylus stenopterus McLachlan.
(PI. I, fig. i ; text-figs. 3, 4.)
Stenosmylus stenopterus McLachlan, 1867, /. Linn. Soc. Lond. (Zool.), 9 : 267 ; Kriiger,
1914, Stettin, ent. Ztg., 75 : 57, 114-117.
McLachlan's description of the female type is, as usual, very good and
requires modifying only in respect of his account of the genitalia. The
Ticxt-fig. 3.
-Stenosmylus stenopterus McL. $. a, apex of abdomen, lateral ; b,
tenth sternite, dorsal ; c, parameres, lateral.
" long straight spine dilated towards the apex " is at the apex of the seventh,
not the base of the sixth, segment. The eighth sternite has a slender process
arising from each lateral margin, fringed with hairs. The " deeply bifid semi-
transparent valve " arises from the eighth and not the seventh sternite ; it is
bifid at both ends. The " sabre-shaped flattened yellow borer," the valves of
the ninth segment, bear at their apices very small styles.
c?. Eighth and ninth tergites fused dorsally. Anal plates fused dorsally,
quadrate from the side, angles rounded, the group of trichobothria set low, rather
ovate. Tenth sternite large and broad, centre of apical margin globose, extreme
apex produced in a slender curved finger. On either side of the centre and at a
lower level is a rounded lobe, which from the side is seen extending basally as a
sinuous ridge. Lower margins produced downwards, plate-like, apices incurved
under the central process. Parameres resembling those of 5. tenuis, apices less
dilated, and no triangular expansions at the base.
170
D. E. KIMMINS
Of the three examples in the type series, one is now very defective, and
appears to have been repaired with an abdomen of a female S. tenuis. Another,
with the same register number, 58-124, also lacks an abdomen, and I have there-
fore selected as type the female from S. Australia (Bakewell), 59-24. It is from
Text-fig. 4.
-Stenosmylus stenopterus McL., $ type, a, apex of abdomen, lateral ; b,
eighth sternite, lateral ; c, the same, ventral.
this example that the figures of female genitalia have been drawn. The distri-
bution of the species, as exemplified by British Museum material, covers S.
Australia, Victoria and N.S. Wales.
Kriiger has placed this species in the synonymy of S. tenuis. I feel sure that
he cannot have seen a true tenuis, for apart from the genital differences, S.
stenopterus has a noticeably narrower anterior wing, its apex more acute. The
pattern of the 'wing is variable, but I have not seen a stenopterus without an
apical streak, nor a tenuis with anything approaching a definite apical streak.
SUBFAMILIES STENOSMYLINAE AND KALOSMYI.INAK
171
Stenosmylus turneri sp. n.
(PI. I, fig. 3; text-fig. 5.)
$. Head and palpi shining brownish, antennae dull luteous. Pronotum
elongate, about as wide as the distance between the eyes, shining brownish, the
margins fringed with strong setae. Many of these setae arise from strongly
elevated bases, giving the lateral margins a serrate appearance. This feature is
a
b
Text-fig. 5. — Stenosmylus turneri sp. n., $ type, a, apex of abdomen, lateral ; b, eighth
sternite, ventral ; c, the same, lateral.
to be seen in other species of the genus. Thorax and abdomen brownish, less
shining, with scattered yellowish hairs. Legs pale brownish, irregularly marked
with dark brown. Wings hyaline, practically immaculate (two or three brown
specks on the gradate series and one on the fork of Cuj in anterior wing), stigma
brownish, venation mainly brownish. Wings shaped much as in 5. tenuis,
but without the prominent hair basis on the cross-veins, which are a feature of
tenuis and stenopterus.
Genitalia of the pattern of stenopterus ; seventh sternite produced at centre
of its apical margin. Eighth sternite shorter and broader, the hinged, bifid
appendage also much shorter, the forks not more than half the length of the
172 D. E. KIMMINS
appendage. Lateral margins about mid- way with slender tooth-like projections.
Base not bifid, but widely excised. Ninth tergite with a blunt process at its
lower basal angle. Valves of ninth segment sabre-like, with a minute apical
style. Anal plate short, deep, rounded, group of trichobothria elongate-oval.
Length of anterior wing 18 mm., posterior wing 16 mm.
Type $, S.E. Queensland, Tambourine Mts., 2-9.iv.35 (R. E. Turner), in
the British Museum.
STENOLYSMUS gen. nov.
(PL I, fig. 4.)
Genotype : Nymphes extraneus Walker.
The species for which this genus has been erected had a somewhat restless
time during the first few years of its entomological life. Walker placed it in
Nymphes, which it indeed resembles in the form of its wings, but the more slender
antennae and the absence of any but a marginal branch to Cuj bar it from the
Nymphidae. Hagen, in 1866, placed it in the genus Osmylus, then a rather
composite genus. The following year McLachlan, when revising Walker's list
of Neuroptera in the British Museum, placed it doubtfully in the genus Myiodac-
tylus, but at the same time pointed out that it possessed " ventral valves and a
borer analogous to those described . . . under Stenosmylus stenopterus."
I suspect that it was its more robust form and rather pallid appearance which
deterred him from including it in Stenosmylus . Whatever his reasons, I think
he was entirely justified, as both male and female genitalia show good characters
to separate it from Stenosmylus, although it is undoubtedly allied to that genus.
The chief difference in wing venation is in the length of A2 in the anterior wing.
This vein is much longer in the present genus, about half as long as A1( and
extends to about the level of the first branch of Rs. In Stenosmylus A2 is only
about one-quarter as long as Av reaching only to the level of the origin of Rs.
cJ. Eighth and ninth tergites and anal plates partially fused, anal plates
also fused together dorsally. Tenth sternite large, more modified than in Stenos-
mylus, the projecting hairy lobe smaller. Parameres shorter and stouter.
?. Seventh sternite with its apical margin produced in a broad plate, with a
slightly excised apex. Process hinged to apex of eighth sternite broad, truncate,
with two apical pockets or depressions. Eighth tergite longer than deep. Group
of trichobothria on anal plate small and nearly circular. Valves of ninth segment
elongate, somewhat twisted along their axes ; apical margin fringed with stout
hooked setae ; apex of valve with a small but definite style.
It is perhaps scarcely necessary to point out that the name Stenolysmus is
an anagram of Stenosmylus, and does not imply any special relationship with
Lysmus Navas.
Distribution : Australia.
Stenolysmus extraneus (Walker).
(PI. I, fig. 4 ; text-figs. 6, 7.)
Nymphes extraneus Walker, 1853, List. Neur. Ins. Brit. Mus., 2 : 230.
Osmylus extraneus (Walker), Hagen, 1866, Stettin, ent. Zlg., 27 : 453.
; Myiodactylus extraneus (Walker), McLachlan, 1867, /. Linn. Soc. Land. {Zoo!.), 9 : 262.
Stenosmylus extraneus (Walker), Banks, 1913, Trans. Amer. ent. Soc., 39 : 215.
Stenosmylus australiensis Esben-Petersen, 191 7, Vidensk. Medd. naturh. Foren. Kbh., 68 : 1
(Syn. nov.).
SUBFAMILIES STENOSMYLINAE AND KAT.OSMYLINAE
173
Walker's type is a female without locality, but I have no doubt that a female
of S. australiensis E.-P. from the Tillyard collection is conspecific. The descrip-
tion of the male genitalia is from a specimen in McLachlan's collection, determined
by myself.
cJ. Eighth and ninth tergites and anal plates fused dorsally, the anal plates
forming a hood. The latter from the side are rounded, a little flattened apically,
Text-fig. 6. — Stenolysmus extraneus (Walker), <J- a, apex of abdomen, lateral ; b,
tenth sternite, lateral ; c. parameres, lateral ; d, the same, dorsal.
and with a small rounded lobe at the lower basal angle. Tenth sternite large,
arched, partly external. Dorsal margin produced in a small setigerous lobe,
rounded from above, its upper surface elevated in a pair of convergent ridges.
Lower margins of tenth sternite produced tailward, curving upward towards the
apical lobe in a pair of somewhat membranous, flattened lobes. Parameres
short, deep, fused at their extreme bases. Lower margins with thin keel-like
flanges. Apices membranous, directed downwards and inwards. Near the base
on tiie upper surface are two horizontal, triangular wings.
174
D. E. KIMMINS
?. Seventh sternite produced in a broad plate, apical margin with a wide
shallow excision. Eighth sternite long, lateral margins each with a small tri-
angular process. To the apical margin is hinged a broad plate directed basally.
There is a deep concavity on its ventral surface near the base, a smaller depres-
sion on the upper surface at the apex and two pockets in the apical surface.
The apex itself is rather widely excised. Ninth tergite short, deep, lower margin
not very dilated. Valves of the ninth segment elongate, somewhat twisted, outer
apical margin fringed with stout, hooked, golden setae. Style small, elongate.
Anal plates short, rather trapezoidal, trichobothria-group small, circular.
Text-fig. 7. — Slenolysmus extraneus (Walker), $. a, apex of abdomen, lateral ; b, apex
of lobe of eighth sternite, ventral ; c, the same, lateral.
Length of anterior wing : <$ 22-26 mm., § 23-28 mm.
posterior wing : <$ 20-24 mm-. ? 21-26 mm.
Distribution : Australia. This species appears to be scarce in collections,
and I have seen only five examples, of which only one has a locality more
definite than Australia, namely, New South Wales, Beecroft, nr. Sydney,
19.iv.17.
OEDOSMYLUS Kriiger.
(PL II.)
Oedosmylus Kriiger, 1913, Stettin, ent. Ztg., 74 : 106-112.
Genotype : Oed. tasmaniensis Kriiger.
Wings rather elongate, otherwise much as in Isostenosmylus. In the female
anterior wing the apical part of C^ and either part of Cu2 and M3+4, or else IA,
SUBFAMILIES STENOSMYLINAE AND KALOSMYLINAE
175
are thickened, as are also some of the associated cross-veins. Tenth sternite in
male large, hairy, external.
Distribution : Australia, Tasmania.
Oedosmylus pallidus (McLachlan).
(PI. II, fig. 1 ; text-figs. 8, 9.)
Osmylus (?) pallidus McLachlan, 1863, /. En!., 2 : 113, pi. 6, fig. 2 ; Hagen, 1866, Stettin.
ent. Ztg.. 27 : 455.
Stenosmylus pallidus (McLachlan), 1870, Ent. mon. Mag. , 6 : 195.
Oedosmylus pallidus (McLachlan), Kriiger, 1913, Stettin, ent. Ztg., 74 : 216.
Text-fig. 8. — Oedosmylus pallidus (McL.), <J. a, apex of abdomen, lateral ; b, tenth
sternite, ventral ; c, parameres, lateral.
McLachlan's type is a female, and to his description may be added that in
the anterior wing, the apical halves of Cuj and Cu2, the cross-veins between these
in the same area, the basal portion of M3+4 and some of the cross-veins between
M and Cux are dilated to about twice their normal width. The artist seems to
have tried to portray this in pi. vi, fig. 2, but the dilatation of Cu, has been
carried too far basally. The female genitalia follow the stenosmyline pattern.
Seventh sternite scarcely produced, eighth from the side with a slender, basally
directed finger arising from tin- centre of the lateral margins. Posteriorly the
sternite tapers to a rounded apex. Hinged lobe slender, apex bifurcate, from the
side with a pair of triangular projections on its upper surface about midway,
directed somewhat outwards. Ninth tergite very narrow dorsally, lower portion
much expanded and with a narrow excision on its basal margin. Valves long
and narrow, with a small apical style. Anal plates pyriform from the side,
broad end uppermost.
176
D. E. KIMMINS
Anterior coxa with a row of five or six finger-like teeth on its anterior
surface near apex, each tooth carrying a socketed seta.
With this female I have associated three males from Hornsby and Hampton,
N.S.W. They have the same pallid, almost immaculate wings, the anterior
pointed and slightly falcate, hind margin lightly bordered with brownish.
cT Genitalia : eighth and ninth tergites and anal plates fused, the latter
forming a hood with slightly excised apex. Ninth sternite produced, apical
margin rounded, centre flattened. Tenth sternite forming a broad hairy lobe,
Text-fig. 9. — Oedosmylus pallidus (McL.), 9 type, a, apex of abdomen, lateral ; b,
eighth sternite, ventral ; c, anterior coxa, lateral.
apex rounded, sides sinuous from beneath. Lower margins of the sternite
produced downwards to form plates with approximated, acute, hooked apices.
Parameres slender, fused basally, bases dilated in triangular wings, apices
rounded, not very dilated.
Distribution : " Australia " ; New South Wales, Hampton, i.18, Hornsby,
12.iii.18, R. J. Tillyard.
Oedosmylus tasmaniensis Kriiger.
(PI. II, fig. 2 ; text-figs. 10, 11.)
Oedosmylus tasmaniensis Kriiger, 1913, Stettin, ent. Ztg., 74 : 26, 27, 94 (not described), 106-
112, 216 ; 1914, 75 : 117-119 (full description).
Stenosmylus tasmaniensis (Kriiger), Tillyard, 1926, Ins. Austr. N.Z. : 320.
SUBFAMILIES STENOSM YLINAE AND KAI.OSMYI.INAE
I77
Anterior wings elongate, 3-3-3-4 times as long as broad, posterior margin
rounded, not falcate, apex not pointed. Anterior coxa of female with numerous
short, socketed setae scattered over the anterior surface, not set on a row of
prominent serrations as in 0. pallidus.
(J Genitalia : eighth and ninth tergites completely fused, anal plates forming
an excised hood, fused at its centre to the margin of ninth tergite. Ninth sternite
short, only slightly produced. Tenth sternite prominent, apex hairy, from the
side dilated, upper angle produced upward in a truncate horn, lower angle
rounded ; from beneath it is parabolic, with the horn projecting beyond the
Text-fig. 10.
-Oedosmylus lasmaniensis Kruger, <$. a, apex of abdomen, lateral ; 6,
tenth sternite, ventral ; c, parameres, lateral.
apex. Lower margins produced downward, forming plates with outcurved,
hooked apices. Parameres slender, fused basally, less curved than in pallidus.
Apical half with a membranous expansion on lower surface, apices rounded and
divergent.
$. Seventh sternite scarcely produced, eighth elongate, apical margin
truncate, process of lateral margin short and stout. Hinged lobe stouter than
in pallidus, lower apical margin terminating in a pair of slender fingers with a
deep U-shaped excision between them. Upper apical angles short, plate-like,
dilated from beneath. The stem of the lobe is concave beneath. Ninth tergite
narrow dorsally, much expanded in its lower portion, no excision on its basal
margin ; valves narrow and elongate. Anal plates rhomboidal, upper angle
truncate.
Distribution: Tasmania: Hobart, George's Bay, i.i.85; Windermere
(? label indistinct), 23.xii.87, 22.L88; Cygnet, 27.xii.16.
178
D. E. KIMMINS
Text-fig. ii. — Oedosmylus tasmaniensis Kriiger, $>. a, apex of abdomen, lateral; b,
eighth sternite, ventral ; c, anterior coxa, lateral.
Oedosmylus latipennis sp. n.
(PL II, fig. 3.)
This species may best be distinguished from tasmaniensis by the shape of
the anterior wings. In latipennis they are definitely broader (three times as long
as broad), and the posterior margin is angled, not evenly rounded. The posterior
wings are also relatively broader. The pattern is much the same, but in the
female, Cux and IA are less swollen. I have been unable to discover any real
difference in the male genital structure of the two species ; unfortunately, both
female paratypes of latipennis lack the abdomen. Should further material reveal
no difference in this sex also, it will perhaps be necessary to reduce latipennis to
the rank of a subspecies. For the present, bearing in mind the fact that the
distribution of latipennis encroaches on that of tasmaniensis, I prefer to keep them
as distinct species.
Length of anterior wing : <$ 19-20 mm., $ 19 mm.
Type cJ, paratype $, N.S. Wales : Mt. Kosciusko, 4500 ft., 21.L85 ; para-
type <$, Mt. Kosciusko, 5000 ft., 22.1.1930, R. J. Tillyard; paratype $, Mt.
Wilson, 19.xi.1921, R. J. Tillyard; paratype cj, Tasmania: Triabunna, 25.xii.
1915, G. H. Hardy, in British Museum.
SUBFAMILIES STENOSMYI.INAE AND KALOSMYLINAE
I79
Oedosmylus montanus sp. 11.
(PL II, fig. 4 ; text-figs. 12, 13.)
Allied to 0. tasmaniensis rather than 0. pallidas, but differing in the form
of the wings and the genitalia. Anterior wing more elongate than in 0. latipennis,
posterior margin angled. Dilated veins of $ (Cu2 and IA) much as in tasmaniensis.
Anterior coxa of female as in tasmaniensis.
Text-fig. 12.
-Oedosmylus montanus sp. n., <J type, a, apex of abdomen, lateral ; b,
ninth tergite and anal plates, dorsal.
cJ Genitalia : eighth and ninth tergites fused, eighth more convex from side,
ninth with a pair of raised hairy lobes on the apical margin, in dorsal aspect
partly overlapping the trichobothria of the anal plates. Anal plates hood-like,
excised, more rounded from the side than in tasmaniensis. Tenth sternite
stouter, upper angle more slender, curving slightly basad, apices of the plate-like
lower margins stouter. Parameres with eversible membranous fingers at the
lower apical angles. These fingers are very delicate and may possibly occur in
other members of the genus.
$ Genitalia : lower margin of eighth tergite near base with a deep rounded
excision. Eighth sternite with a reniform prominence in basal half, basal margin
excised. Hinged lobe rather stout, apex more roundly excised, arms shorter and
i8o
D. E. KIMMINS
stouter. Upper portion of apex forms a transverse ridge. Lower portion of
ninth tergite broader, valves more slender, slightly curved.
Length of anterior wing : c? 20 mm., $ 21 mm.
Queensland : National Park, 3-4000 ft.
Type <£, 18.xii.21, paratype J, 20.xii.21, paratype 9. 21.xii.21, in British
Museum.
Text-fig. 13. — Oedosmylus montanus sp. n., $ paratype. a, apex of abdomen, lateral ;
b, eighth sternite, ventral.
ISOSTENOSMYLUS Kriiger.
(Pis. Ill and IV.)
Isostenosmyhis Kriiger, 1913, Stettin, ent. Ztg., 74 : 23, 216 ; op. oil., 75 : 48, 120.
Genotype : OsmvUis pulveritlentus Gerstaecker.
= Ausirosmylus Banks, 1913, Trans. Amer. ent. Soc, 39 : 215. (Syn. nov.)
Genotype : Osmylus pulveritlentus Gerstaecker.
In anterior wing, one cross-vein in the subcostal area. First cross-vein to
M beyond the basal corneous spot arises from the first branch of Rs. No thicken-
ing of Cu! and Cu2 or IA in either sex. In the hind wing, no basal radio-medial
cross-vein ; Cu2 long.
tj. Eighth and ninth tergites fused, line of fusion often indistinct ; dorsum
SUBFAMILIES STENOSMYI.I N Ai: AND KALOSM YI.INAE
181
often membranous and impressed in a longitudinal trough. Anal plates fused
dorsally, differing in form in the various species. Tenth sternite largely internal,
U-shaped, with a branch on lower surface at base. Paramens fused basally,
with a pair of basal, and sometimes also apical, branches.
Distribution : S. America (Ecuador, Peru, S. Brazil).
Text-fig. 14. — Isosteitnsiuvlits fusiitips sp. n., ,J paratypc. a, apex of abdomen,
lateral ; b, tenth sternite, lateral ; c, apex of the same, dorsal ; d, parameres,
lateral ; e, the same, dorsal.
Isostenosmylus fusciceps sp. n.
(PI. Ill, fig. 1 ; text-fig. 14.)
<J. Vertex, Irons to below antennae, and genae fuscous, remainder fulvous,
palpi dark fulvous. Antennae with two basal, and under surface oJ some succeed-
ing segments dark, remainder light fulvous. Pronotum nearly twice as long as
wide, fuscous, with golden-brown setae. Meso and metanota fuscous with
darker markings. Legs pale yellowish, tibiae finely speckled with blackish and
with a narrow, external, apical band ol blackish. Abdominal segments 1-7
fuscous, remainder fulvous, Wings greyish hyaline, marked with brownish as
in PL III, fig. 1. Veins mainly brownish, with some whitish areas.
MATT. ZOOL. 4_>. I 15
182
D. E. KIMMINS
Anal plates small, apical margins produced in short, broad spatulae, slender
from the side ; lower angles slightly produced and rounded. Ninth sternite
rather large, rhomboidal from the side. Tenth sternite largely internal, with its
apex produced in a short quadrate plate, whose apical margin is fringed with
fine setae. Sides of the segment slender, terminating in ovate plates. From
the lower margin of each side arises a flattened branch, bent twice at right angles,
so that its apex approaches the apex of the segment, the area between the branch
and the segment membranous. Parameres with the basal branches (the attach-
ment to the tenth sternite) directed apically and approaching each other ; in
dorsal aspect they appear somewhat lyriform. Apices of the parameres thin,
somewhat dilated, from above approximating and then diverging. Shortly
before its apex each paramere bears on its upper surface an upwardly directed
branch.
Length of anterior wing 25 mm., of posterior wing 23 mm.
Type J, 1 paratype <J, Peru : Callanga, from the McLachlan collection, now
in the British Museum.
Text-fig. 15. — Isoslenosmylus pulverulenlus (Gerst.), <?. a, apex of abdomen,
lateral ; b, tenth sternite, lateral ; c, apex of the same, dorsal ; d, parameres,
lateral ; e, the same, dorsal.
SUBFAMILIES STENOSMYLINAE AND KALOSMYLINAE
183
Isostenosmylus pulverulentus (Gerstaecker).
(PL III, fig. 2 ; text-figs. 15, 16.)
Osmylus pulverulentus Gerstaecker, 1893, Mitt, naturw. Ver. Greifswald, 25 : 166.
Isostenosmylus pulverulentus (Gerst.), Kriiger, 1913, Stettin, ent. 7Ag., 74 : 26, 112—115, 216.
Austrosmylus pulverulentus (Gerst.), Banks, 1913, Trans. Amer. ent. Soc, 39 : 215.
Isostenosmylus pulverulentus (Gerst.), Kriiger, 1914, Stettin, ent. Ztg., 75 : 120-121.
There is little to add to the general descriptions of Gerstaecker and Kriiger.
I have seen a more heavily marked specimen from " Espirito Santo " in which
the cubital spot at the level of the apex of A1 is extended forward as a narrow
streak to the radius.
Text-fig. 16. — Isostenosmylus pulverulentus (Gerst.), ?.
b, eighth sternite, ventral.
a, apex of abdomen, lateral ;
cj Genitalia : line of fusion between eighth and ninth tergites marked by
an internal thickening ; ninth more chitinized than eighth, its apical margin
produced in a rounded lobe, which is divided to its base by a V-shaped, mem-
branous, apparent excision. Margin of ninth sternite, from beneath, projecting
in an obtuse triangle. Anal plates obliquely truncate, lower apical angles
rounded. Tenth sternite rather as in fusciceps, the apical margin rounded, its
centre shallowly excised. Sides of the sternite slender, not dilated nt the ends,
lower branches broader. Parameres without an upper branch near apex, which
is much deeper. Basal branches less apically directed, fused basal portion of
parameres longer and more acute.
?. Apical margin of seventh sternite produced in an elliptical, somewhat
truncated lobe. Eighth sternite about two-thirds of the length of tergite, with
novit zool., 42, 1 I5§
1 84
D. E. KIMMINS
a longitudinal ridge ; basal margin curved down at right angles to meet the pro-
duced apex of seventh stcrnite. There is a transverse ridge about mid-length of
the sternite and the anterior angles are slightly raised and hairy. Attached
anteriorly by membrane is a slender lobe with a dilated and excised apex, directed
basally. From the side this lobe is angled near its base and the apex is divided
into an upper and lower portion by a rounded excision. Ninth tergite short and
deep, valves rather broad, lanceolate, each with a small apical style and a small
pointed process on the lower margin near the base.
Length of anterior wing 21-25 m111-
The above descriptions are based on a pair from Rio Grande do Sul. In
view of the general resemblance in wing pattern between the various species,
I suspect that Gerstaecker's example from " Hoch-Peru " may prove to be not
conspecific with those from Southern Brazil.
Text-fig. 17. — 1 sostenosmylus fasciatus sp. n., <J type, a, apex of abdomen,
lateral ; b, tenth sternite, lateral ; c, apex of the same, dorsal ; d, parameres,
lateral ; e, the same, dorsal.
SUBFAMILIES STENOSMYLINAE AND KALOSMYLINAE 185
Isostenosmylus fasciatus sp. n.
(PI. IV, fig. 1 ; text-fig. 17.)
cJ. Head light fulvous, marked with brownish on the genae, centre of clypeus,
below antennae and on vertex. Palpi and two basal segments of antennae
brownish. Pronotum nearly twice as long as broad, dark brown, with a broad,
yellowish, median fascia. Meso- and metanota dark brown, with yellowish
markings. Legs as mfusciceps. Abdominal tergites 1-7 brownish, with a narrow,
median, yellowish line. Sternites and apex of abdomen yellowish. Wings
greyish hyaline, marked with brownish as in PI. IV, fig. 1. Venation brownish,
with pale areas forming four indefinite streaks across the anterior wing. Area
between first and second branches of Rs rather broader than in preceding species.
Apical margin of ninth tergite produced, sternite from beneath moderately
produced, margin forming a flattened cusp. Anal plates small, basal margin
elevated in a small triangular projection. Apical angles produced, the lower
rounded and hairy, the upper broad, its apex from above rounded, from the side
with a shallow excision. Tenth sternite with rounded apical margin, its centre
more produced. Sides slender, apices moderately dilated, forming clavate
plates.
Parameres more developed than in preceding species. The fused base
angled upwards, basal branches directed upward. Apices larger and with a
complex system of folds on their upper outer surfaces.
Length of anterior wing 23 mm., of posterior wing 21 mm.
Type cT, Peru: Callanga; paratype <$, Bolivia: "Chaico", in the British
Museum, both from the McLachlan collection.
Isostenosmylus nigrifrons sp. n.
(PL IV, fig. 2 ; text-fig. 18.)
<J. Head fulvous, frons below and between antennae, with a black trans-
verse band from eye to eye. In the centre this band extends downward in a
tongue, and on either side of it, between the eye and the tentorial pit, is a round
black spot. Genae and palpi shining black. Antennae (incomplete) yellowish,
two basal segments brownish. Prothorax a little more than twice as long as
broad, blackish, with a yellowish median fascia. Meso- and metanota yellowish,
with black markings. Legs as in other species of the genus. Abdomen (dis-
coloured) brownish above, fulvous beneath. Wings similar to fasciatus, more
lightly marked, but with the posterior margin of hind wing distinctly clouded
with brownish.
From the side the dorsal outline of the eighth and ninth tergites is distinctly
angled at the point of fusion. Apical margin of ninth sternite produced at its
centre in a short finger. Anal plates with the upper and lower apical angles
produced, the latter only slightly, rounded, and hairy on its inner surface ; the
upper is larger and hooked downwards. Excision between upper processes in
dorsal aspect with sinuous sides. Tenth sternite from above with its apical
margin parabolic, its apex flattened or slightly excised. Bases of the arch
dilated only on their lower sides ; lower branches in form of quadrate plates
attached by short, thick stems. Fused bases of the parameres with a V-shaped
i86
D. E. KIMMINS
excision in dorsal aspect ; basal branches shorter than in fasciatus, directed
upward and incurved apically. Apex of paramere broad, plate-like, apical
branch also plate-like and separated from apex by an excision.
Length of anterior wing 22 mm., of posterior wing 20 mm.
Type jf, Ecuador: Intaj, from the McLachlan collection, now in the British
Museum.
Text-fig. 18. — Isostenosmyhts nigrifrons sp. n., $ type, a, apex of abdomen,
lateral ; b, tenth sternite, lateral ; c, apex of the same, dorsal ; d, parameres,
lateral ; e, the same, dorsal.
Isostenosmylus morenoi (Navas).
Oedosmylus morenoi Navas, 1928, Bol. Soc. ent. Esp., 17 : 91.
Navas has described, under the name Oedosmylus morenoi, an Isostenos-
mylus from Ecuador. He says that it resembles /. pulverulentus Gerst.,
and his figure certainly is very like the variety of that species which I have
mentioned from Espirito Santo. The type lacks most of its abdomen and
it will probably prove difficult to decide whether any of the foregoing
species are conspecific with it.
SUBFAMILIES STKNOSMYLINAE AND KALOSMYI.INAE
187
EUPORISMUS Tillyard.
(PI. V, fig. 1 ; text-fig. 19.)
Euporismus Tillyard, 1916, Proc. Linn. Soc. N.S.W., 41 : 43.
Genotype : E. albatrox Tillyard.
Wings with large, opaque, white patches at apices, remainder of anterior and
apical half of posterior dark brown, mottled with white. In anterior wing, first
cross-vein to the medius beyond the basal corneous point arises from the first
TEXT-FIG. ig. — Euporismus albatrox Tillyard, $ type. a, apex of abdomen, lateral
b, tenth sternite, lateral ; c, parameres. lateral.
branch of the radial sector. Medius forks several cells beyond the second
corneous point. Cu2 in posterior wing long. Gradate veins form a very even
line ; outer branches of radial sector very numerous.
Tillyard placed this genus between Oedosmylus and Porismus, considering it
to be allied to Porismus. At the time of publication (1916) Tillyard was probably
unacquainted with Kriiger's revision of the Osmylidae. His description unfor-
tunately omits all reference to the chief character separating Porismus from the
other subfamilies, namely, the presence ot numerous cross-veins in the subcostal
area. The type of Eupotismui has only one cross-vein in this area (the figure
100 D. E. KIMMINS
accompanying his description is inaccurate in this respect), and this, coupled with
the absence of a basal cross-vein connecting the medius with the radial sector in
the posterior wing, and the distant furcation of the medius in the anterior wing
places this genus in the Stenosmylinae. Tillyard is undoubtedly correct in
comparing it with Oedosmylus, but until the female is known one cannot say
whether certain veins are swollen, as in Oedosmylus. Its striking wing pattern
will readily distinguish Euporismus from other members of the subfamily.
Euporismus albatrox Tillyard.
(PL V, fig. i ; text-fig. 19.)
Euporismus albatrox Tillyard, 1916, Proc. Linn. Soc. N.S.W ., 41 : 44, pi. 1, fig. 1 ; 1926, Ins.
Austr. N.Z. : 320, pi. 24, fig. 2.
The type, a male, is now in the British Museum. To Tillyard's description
the following notes on the genitalia may be added :
No apparent eversible scent-glands between the eighth and ninth tergites,
which are fused ; ninth fused dorsally to anal plates, free margin produced in a
small rounded lobe on each side of the fused part. Ninth sternite short, tri-
angularly produced from the side. Anal plates fused together dorsally at base,
apically separated by a deep incision ; from the side triangular, apex rounded.
Tenth sternite large, projecting slightly beyond anal plates, hairy, apex truncate,
angles gently upcurved. From beneath, lateral margins are broadly emarginate.
Basal angles of sternite produced in curved branches, shorter than sternite,
apices from beneath clavate and outspread. Parameres slender, fused basally,
apices dilated, lower apical margin extended downwards and outwards, support-
ing the usual membranous eversible finger.
Length of anterior wing 30 mm., posterior wing 28 mm.
Queensland: Killarney, head of Condamine R., i.1914 {E. J. Dumigan).
Of the three paratypes mentioned by Tillyard (sex not stated) one is in Dr.
Esben-Petersen's collection, one in Mr. Banks's collection, and the third is presum-
ably in the Australian Museum, Sydney, or in the Entomological Dept., Council
for Scientific and Industrial Research, Canberra.
KALOSMYLINAE Kruger.
Kalosmylinae Kriiger, 1913, Stettin, ent. Ztg., 74 : 40, 104-113.
In anterior wing, one cross-vein in subcostal area ; medius forking near its
base, but beyond the origin of the first branch of Rs. In posterior wing a basal
cross- vein between the medius and the radial sector, not very sinuous ; Cu, long.
In the male a pair of eversible scent-glands between the eighth and ninth tergites.
Key to Genera.
1. In posterior wing, branches of medius widely separated, cells between
them long and sinuous ; New Zealand, Australia, Chile Kempynus Navas.
In posterior wing, branches of medius not widely separated, cells between
them short, quadrate ......... 2.
2. In posterior wing, M3 separates from M4 at about the level of the apex of
Cuj ; small species, New Zealand .... Euosmylus Kruger.
M3 separates from M4 two or three cells before apex of Cux ; larger species,
Australia ........ Australysmus gen. n.
SUBFAMILIES STENOSMYLINAE AND KALOSMYUNAE 189
KEMPYNUS Navas.
(Pis. VI and VII.)
Kempynus Navas, 191 2, Mem. It. Acad. Barcelona, 10 (9) : 59.
Genotype : Osmylus excisus Navas (= Osmylus incisus McL.).
Banks, 1913, Trans. Amer. enl. Soc, 39 : 214.
Genotype : Stenosmylus incisus McLachlan.
= Kalosmylus Kriiger, 1913, Stettin, ent. Ztg., 74 : 23, 96-102.
Genotype : Osmylus incisus McLachlan. (Syn. nov.)
= Osmylinus Banks, 1913, Trans. Amer. ent. Soc, 39 : 214.
Genotype : Osmylus longipennis Walker. (Syn. nov.)
Kriiger proposed the suppression of Navas' name Kempynus on the grounds
that (a) the name was incorrectly spelt, being derived from Kempny, and should
have been Kempnyus ; (6) the genus was based on a non-existent species [excisus
McLachlan).
The first is no reason for rejection, nor is there even a case for emendation
of the name, as it is perfectly clear that Navas intended this spelling, for he used
it no less than six times in his publication. As regards the second, this is due
to a lapsus calami by Mr. W. F. Kirby in labelling McLachlan's type of 0. incisus,
which was copied without verification by Navas. Even so, Kempynus excisus
is not a non-existent species, as a very full description was given by Navas when
erecting the genus Kempynus. I therefore consider Kempynus to be a valid
genus with as genotype Kempynus excisus Navas, = Osmylus incisus McL.
Wings falcate or entire. Anterior with one cross-vein in subcostal area ;
medius forking near base, but beyond origin of first branch of radial sector.
Posterior wing : basal radio-medial cross- vein present, M1 + 2 and M3+1 widely
separated, cells between them long, narrow and sinuous ; M3 separates from M4
several cells before apex of Cu, ; often a basal cross-vein from fork of medius to
cubitus ; Cu2 long. Relatively large species, expanse 42-55 mm.
Distribution : New Zealand, Australia, Chile.
Kempynus incisus (McLachlan).
(PI. VI, fig. 1 ; text-figs. 20, 21.)
Osmylus (?) incisus McLachlan, 1863, /. Ent.. 2 : 112 ; Hagen, 1866, Stettin, ent. Ztg., 27 :
455-
Stenosmylus incisus (McLachlan), 1870, Ent. mon. Mag., 6 : 195 ; 1873, Ann. Mag. nut.
Hist.. (4), 12 : 38; 1894, Ent. mon. Mag., 30 : 241-242. Hudson, 1904. .V. /.. Neur. :
5°-
Kalosmylus incisus (McL.), Kriiger, 1913, Stettin, ent. Ztg., 74 : 26, 96-102, 212-213 ; 1914,
op. cit., 75 : 104-106.
Kempynus incisus (McL.), Banks, 1913, Trans. Amer. nit. Soc, 39 : 215. Tillyard, 1926,
Ins. Austr. N.Z. : 320.
Kempynus excisus Navas, 191 2, Mem. R. Acad. Barcelona, 10 (9) : 59.
This species varies somewhat in the intensity of the markings. One form
has the smaller mottlings of the anterior wing merged into the ground-colour,
only the large markings along the posterior margin distinct.
cT Genitalia : ninth tergite short and deep, stemite triangular from vide,
apical margin rounded, with a small, shallow excision at its centre. Anal plates
fused, hood-like, apical margin broadly excised ; from side triangular, upper and
apical angles with rounded hairy warts, apical the larger. Tenth stemite largely
withdrawn, arched, apex from above truncate, from side rounded. Prom basal
190
D. E. KIMMINS
angles arise large curved branches, whose apices converge, and project a little
beyond apex of sternite. Parameres short, fused basally ; on the dorsal surface
near the base are a pair of outspread, slender wings. Beyond, the parameres
are curved upward, plate-like, lower margins produced downward and outward,
probably covered with membrane and each with an eversible finger, as occurs
in citrinus.
Text-fig. 20. — Kempynus incisus (McL.), <J type, a, apex of abdomen, lateral ;
b, tenth sternite, lateral ; c, the same, dorsal ; d, parameres, lateral ; e,
parameres, dorsal.
$. Seventh sternite slightly produced, and bearing on its apical margin a
pair of small, slender processes, separated by about twice their length. Eighth
tergite large and deep, sternite short, from beneath broad and sinuous, concave
at its centre, raised sides with a few radiating setae. Hinged lobe set at some
distance from sternite, connected to it by membrane, in side view arising between
the sides of ninth tergite. From beneath, it is bifurcate, the two branches separated
by a wide rounded excision, apices dilated, obliquely truncate and turned abruptly
downward. Ninth tergite narrow, not much dilated at its lower angles ; valves
SUBFAMILIES STENOSMYLINAE AND KALOSMYLINAE
I9I
rather broad, of medium length, a short, stout style just before the apex. Anal
plates short, deep, fused dorsally to form a hood, bearing two transverse bands
of stout setae, one at apex and other towards base. Accessory glands with long
ducts, each terminating in an elongate sac, constricted midway, the apical part
being the larger.
From three examples marked " Type " by McLachlan, I have selected as
Text-fig. 21. — K. incisus (McL.), $. a, apex of abdomen, lateral ; b, eighth sternite,
ventral ; c, accessory glands.
holotype a male in the British Museum, labelled " Auckland, N. Zeal." and
" excisus M'L. type " in W. F. Kirby's handwriting. There is another
paratype (without abdomen) from the same locality, labelled " Kempynus excisus
McLach." by Navas, and two male paratypes " Otago, Oxley," from McLachlan's
collection.
In the British Museum are additional specimens from the following localities :
Wellington ; Wainuiomata ; Wilton's Bush ; Waitara ; Kaitoke, all collected
and presented by Mr. (.. V. Hudson.
IQ2 D. E. KIMMINS
Kempynus citrinus (McLachlan).
(PI. VI, figs. 2, 3 ; text-figs. 22, 23.)
Stenosmylus citrinus McLachlan, 1873, Ann. Mag. nat. Hist., (4), 12 : 38 ; id., 1894, Ent.
mon. Mag., 30 : 242 ; Hudson, 1904, N. Z. Neur. : 51.
Kalosmylus citrinus (McL.), Kriiger, 1913, Stettin, ent. Zlg., 74 : 26, 96-102, 213 ; id., 1914,
op. cit., 75 : 106-108.
Kempynus citrinus (McL.), Banks, 1913, Trans. Amer. ent. Soc, 39 : 215 ; Tillyard, 1926,
Ins. Austr. N.Z. : 320.
This appears to be a more variable species than K. incisus, the extreme form
that I have seen being very prettily mottled on both wings, the membrane being
greyish rather than yellowish. Another variety has a brownish streak between
the branches of the cubitus in the anterior wing.
cj. Genitalia scarcely distinguishable from incisus. Apical margin of tenth
sternite less truncate, basal branch larger. Parameres with an eversible finger at
lower apical angle, arising from membrane covering paramere. This membrane
is very delicate and liable to damage during clearing in caustic potash, and
probably occurs in other species also.
$. Similar to incisus. Apex of seventh sternite slightly produced, rounded,
margin with a pair of very small teeth, sometimes absent. Eighth sternite roughly
triangular from beneath, concave at its centre, with a deep apical incision, which
is narrower than in incisus. Apices of hinged lobe less dilated, incurved and not
turned down. Ninth tergite narrow, slightly dilated at lower angles, valves a
little broader than in incisus. Anal plates nearly as long as deep, hood-like,
setae arranged in two groups. Accessory gland more constricted, terminal sac
set at right angles to stem.
The type is a female, labelled " New Zealand." Other specimens in the British
Museum from the following localities : Wellington ; Waitara ; Wainuiomata ;
Wilton's Bush (collected by G. V. Hudson) and from Ohakune (T. R. Harris).
Kempynus latiusculus (McLachlan).
(PL VII, fig. 1 ; text-fig. 24.)
Stenosmylus latiusculus McLachlan, 1894, Ent. mon. Mag., 30 : 241 ; Hudson, 1904, N. Z.
Neur. : 54 ; Tillyard, 1926, Ins. Austr. N.Z. : 320, pi. 24, fig. 1.
Kalosmylus latiusculus (McL.) Kriiger, 1913, Stettin, ent. Ztg., 74 : 26, 96-102, 213 ; id.,
1914, op. cit., 75 : 108-109.
This species appears to vary little, most examples before me being less mottled
(tessellated) than the type.
<$. Apex of anal plates slightly truncate from side. Tenth sternite more
robust, apical margin more hairy, from above broader and more rounded than in
incisus and citrinus.
9. Differing only in a few details from incisus and citrinus. Seventh sternite
with a pair of small triangular processes on its apical margin. Eighth sternite
very lightly chitinized, hinged lobe from side with apex dilated, lower angle
right-angled, upper rounded. The apex is not abruptly bent down as in incisus.
From beneath the arms of the lobe are less widely separated, excision between
them oval, not circular. Terminal sac of accessory gland pyriform.
Type is a female, labelled " Otira Gorge, on window at light." Subsequent
specimens from the following localities : Wellington, Hutt Forks (Stella Hudson)
and Wellington, Mangaterua R. (G. V. Hudson).
SUBFAMILIES STENOSMYLINAE AND KALOSMYI.IN .U
193
Text-fig. zz. — A', citrinus (McL.), o". Tenth sternite, dorsal.
Text-fig. 23. — K. citrinus (Mi I..). $. a, apex of abdomen, lateral ; b, eighth sternite,
ventral ; < . apex of accessory gland
i94
D. E. KIMMINS
Text-fig. 24. — K. latiusculus (McL.), o-c, 0* ; d-f, $. a, anal plate, lateral ; b,
tenth sternite, lateral ; c, the same, dorsal ; d, lobe of eighth sternite, ventral ;
e, apex of same, lateral ; /, apex of accessory gland.
Kempynus longipennis (Walker).
(PI. VII, fig. 2.)
Osmvlus longipennis Walker, 1853, List. Neur. Ins. Brit. Mus., 2 : 235 ; Hagen, 1866,
Stettin, ent. Ztg., 27 : 455.
? Stenosmylus longipennis (Walk.), McLachlan, 1868, /. Linn. Soc. Land. (Zool.), 9 : 268 ;
id., 1870, Ent. mon. Mag. 8 : 195.
Kalosmylus longipennis (Walk.), Kriiger, 1913, Stettin, ent. Ztg., 74 : 26, 96-102, 213-214 ;
id., 1914, op. cit., 75 : 109-110.
Osmylinus longipennis (Walk.), Banks, 1913, Trans. Amer. ent. Soc, 39 : 214.
This species is still represented in the British Museum only by the type.
This now lacks part of the abdomen, all but two segments of an antenna, one
anterior leg and four tarsal segments of the other ; the margins, particularly of
the anterior wing, are damaged. I quote Walker's description, enclosing any
additional notes of my own within square brackets :
" OSMYLUS LONGIPENNIS.
" Piceus ; prothorax linearis; pedes testacei, fasciis tarsisque piceis ; alae
ciner ascent es, Iongissimae, anticae fusco guttatae.
SUBFAMILIES STENOSMYLINAE AND KALOSMYI.ISAE
195
' [Head testaceous, marked with brownish on face below antennae, dark
brown above antennal bases and two impressed dark brown areas on vertex.]
Subcostal veinlets much more numerous than in 0. chrysops, and more contiguous
towards the base ; veinlets of the sector of the second radius [branches of the
radial sector] also much more numerous ; gradate veinlets forming many more
series ; veins more oblique towards the tips ; marginal veinlets much more
numerous ; areolets in the disk much more numerous, more alike and regular
in form, generally almost square. Pitchy : prothorax linear [24 : 19], longer
than that of 0. chrysops, and not narrower in front as is the latter [anterior angles
rounded] : legs testaceous, with pitchy bands [three on each tibia, apical, median,
basal, one large median and small apical on each femur] : tarsi pitchy : wings
grayish, very long ; fore wings very thickly covered with [diffuse] brown dots,
many of which are confluent [general appearance is of a brownish wing, with
darker markings along radius and cubitus, margins with small white dots] ; veins
brown [with pale interruptions along radius and cubitus, marginal gradate series
pale]. Length of body 5 lines ; [expanse] of the wings 30 lines.
" a. New Holland."
The type (sex doubtful) now carries a printed label " Australia " and a
manuscript label " O. longipennis Australia " not in Walker's handwriting. It
resembles a large and very darkly marked example of K. latiusculus (McL.).
One pair of wings has been removed from the type and placed between glass
slides for figuring and as a measure of protection against further deterioration.
I I \ I FIG. 25.-
-K. falcatus Navas, 9 type, a, eighth sternite, lateral ; b, the same,
ventral ; c, apex of accessory gland.
ig6 D- E. KIMMINS
Kempynus falcatus Navas.
(PI. VII, fig. 3 ; text-fig. 25.)
Kempvnus falcatus Navas, 1912, Mem. R. Acad. Barcelona, 10, (9) : 60; id., 1936, Rev.
Chil. Hist, nat., 40 : 180.
? Kalosmylus falcatus (Nav.) Kriiger, 1913, Stettin, ent. Ztg., 74 : 214, 221 ; id., 1914. °P-
cit., 75 : iio-iii.
Stenosmylus sp. McLachlan, 1894, Ent. mon. Mag., 30 : 241.
Navas' description of general appearance and figure of wing pattern are
reasonably good. His type is a female and is from Mulchen, not Meelchen as
given by Navas. The specimen mentioned by McLachlan is paler, but is
undoubtedly conspecific.
$ Genitalia : seventh sternite scarcely produced, eighth short, broad, apical
margin with two deep rounded excisions, separated by a triangular, elevated
promontory ; from the side this appears as a deep triangular keel. Lobe short,
bifurcate, apices incurved from beneath, angled upward from side. Ninth tergite
narrow, slightly dilated at lower angles ; valves broad, apices bluntly rounded.
Anal plates short, deep, angles rounded. Terminal sac of accessory gland slightly
constricted and folded back upon itself, duct not very long.
Type ?, Chile : Mulchen, i.1902, H. J. Elwes.
1 $, " Araucania", ii.1888, from McLachlan collection.
EUOSMYLUS Kriiger.
(PI. VIII.)
Euosmylus Kriiger, Stettin, ent. Ztg., 74 : 23, 102-105.
Genotype : Stenosmylus stellae McLachlan.
Wings falcate, venation much as in Kempynus, but branches ot medius in
posterior wing less separated, cells between them broader, quadrate, not sinuous ;
M3 separates from M4 about the apex of Cu: ; often a basal cross-vein from fork
of medius to cubitus. Small species, expanse 25-32 mm. Genitalia very similar
to Kempynus.
Distribution : New Zealand.
Euosmylus stellae (McLachlan).
(PL VIII, text-fig. 26.)
Stenosmylus stellae McLachlan, 1899, Ent. mon. Mag., 35 : 259-260 ; Hudson, 1904, N. Z.
Neur. : 52-54.
Euosmylus stellae (McL.) Kriiger, 1913, Stettin, ent. Ztg., 74: 26, 102-105, 214; id., 1914,
op. cit., 75 : in— 113.
Kempynus stellae (McL.) Banks, 1913, Trans. Amer. ent. Soc, 39 : 215.
var. connexus McL., 1899, Ent. mon. Mag., 35 : 259.
var. obliteratus McL., 1899, ibid., 35 : 260.
This species varies considerably in size, markings and breadth of wings.
^ genitalia are scarcely distinguishable from incisus. The ninth tergite is
a little broader at its lower angles, and the apex of the anal plate a little more
prominent.
?. Seventh sternite slightly produced, margin with a pair of small, obtuse,
triangular projections. Eighth sternite short, convex, about half as long as
tergite, apex excised ; lobe short, from beneath with short, stout arms, separated
by a U-shaped excision. Apices obliquely truncate, somewhat excavate, inner
SUBFAM1I IKS sTKNosMYI 1NAK ANH KAI.OSM VI. I N AK
ig7
angles the longer. From side, the lobe is slightly downcurved, arms gradually
dilating to apices, a small triangular process on upper surface at base. Ninth
tergite slightly more dilated at lower angles than incisus, valves short and broad.
Anal plates not as long as deep, angles rounded. Accessory gland hour-glass-
shaped, terminal sac the larger ; duct much shorter than in Kempynus.
Text-fig. 26.-
-Euosmylus stellae (McL.), $ paratype. a, apex of abdomen, lateral ;
b, lobe of eighth sternite, ventral ; c, accessory gland.
Type female, 2<Ji ? paratypes from Wainuiomata, near Wellington, also
other specimens from the same locality.
Var. connexus McL.
Type cT. 1 <S paratype from Wainuiomata ; other specimens from the same
locality and from Nelson.
Var. obliteratus McL.
Type (J, 3 cJc? paratypes from Wainuiomata ; other specimens from the same
locality.
AUSTRALYSMUS gen. nov.
(PI. V, fig. 2.)
Wings not falcate. Anterior with one cross-vein in subcostal area ; medius
forking near base but beyond origin of first radial sector. In posterior wing,
basal radio-medial cross-vein present ; branches of medius not very widely
separated, cells quadrate, not sinuous ; Ms separates from M4 two or three cells
198
D. E. KIMMINS
before apex of Cux ; no basal cross-vein from fork of medius to cubitus,
long. Species large, 40-44 mm. expanse.
Genotype : Australysmns lacustris sp. n.
Distribution : Australia (New South Wales).
Cu,
Australysmus lacustris sp. n.
(PI. V, fig. 2 ; text-figs. 27, 28.)
Head luteous, obscured with brown, particularly on the vertex (which carries
two piceous spots), below the antennae and on the margins of labrum and clypeus.
Text-fig. 27.-
-Australysmus lacustris sp. n.
b, tenth sternite, lateral ;
(J type, a, apex of abdomen, lateral
c, parameres, dorsal.
Antennae brown, joints faintly annulated with yellowish ; palpi brownish. Pro-
notum about three-quarters as long as broad, anterior angles rounded ; luteous,
with five brown spots, one median, elongate, and a pair on each side, like exclama-
tion marks ; basal margin brown. Meso- and metanota brownish, with obscure
yellowish markings. Legs luteous, tibia with piceous spots exteriorly at base,
mid-way and apex, posterior femora brownish above, terminal segment of tarsus
piceous. Anterior coxae of female not modified. Abdomen brownish.
SUBFAMILIES STENOSMYLINAE AND KALOSMYI.INAE
1.(0
Wings elongate, apices rounded, not falcate, membrane hyaline, pale yellowish
in subcostal area. Anterior wing much speckled with brownish, venation mainly
brown. Posterior wing not speckled, venation mainly brown, except for some
pale areas in apical half of wing.
<J. Eversible scent-glands between eighth and ninth tergites, apparently
shorter than in Kempynus. Ninth tergite produced, on either side of the centre
Text-fig. 28. — A. lacuslris sp. n., ? paratype. a, apex of abdomen, lateral ; b, lobe
of eighth sternite, ventral.
still further produced in two small, hairy, rounded lobes. Ninth sternite tri-
angularly produced, apex with a broad, shallow notch. Anal plates large, hood-
like, apex obtusely triangular from above, from side upper angle slightly
produced and rounded. Tenth sternite slender, rib-like, hairy, from abovearched,
apex truncate, basal branch projecting beyond apex ; parameres short, deep,
lower margins produced outwards, apices with eversible membranous fingers.
$. Seventh sternite produced at centre in rounded lobe. Eighth sternite
short, quadrate, concave ventrally ; apical margin with two deep excisions,
separated by a slender acute tongue. Lobe large, bifurcate ; from the Mde, deep,
upper margin with a semicircular excision before apex, lower with a shallower
200 D. E. KIMMINS
excision separating a rounded lobe from the apex. The latter is dilated and
hammer-like, upper angle longer than lower. From beneath, this upper angle is
truncate, directed inward, lower more slender and directed outward. A slightly
curved ridge on the inner lower margin at the base of each fork of the lobe. Eighth
tergite with a downwardly directed finger on the lower margin at base. Ninth
tergite from the side slender, sinuous, lower basal angle strongly produced basally
in a rounded lobe, projecting on either side of lobe of eighth sternite. Valves of
medium length, with small apical styles, lower margin more curved than upper.
Anal plates short, upper apical angle obtuse, lower rounded. Accessory glands
each with a single short ovate sac at apex.
Length of anterior wing : <$ 20-22 mm., ? 23 mm.
posterior wing : $ 18-20 mm., ? 21 mm.
New South Wales: Mt. Kosciusko, Blue Lake and Club Lake, i.ii.30,
R. J. Tillyard.
Type $, paratype $ from Blue Lake, paratype $ from Club Lake, in British
Museum.
This species bears a superficial resemblance to Eidoporismns pulchellus
E.-P. The latter may be readily distinguished by the distant origin of the first
branch of the radial sector and the position of the fork of the medius in the anterior
wing, far beyond the middle.
(MSS. reed. Jan. 3, 1940.)
PLATE I.
Anterior wings of Stenosmylus and Stenolysmus.
1. Stenosmylus stenopterus McL.
2. S. tenuis (Walk.).
3. S. turneri sp. n., $ Type.
4. Stenolysmus exlraneus (Walk.).
PLATE II.
Anterior wings of Oedosmylus spp. $.
1. 0. pallidus (McL.), Type.
2. 0. tasmaniensis Kriiger.
3. 0. latipennis sp. n., paratype.
4. 0. montanus sp. n., paratype.
PLATE III.
Wings of Isostenosmylns spp. <J.
1. I . fusciceps sp. n., Type.
2. /. pulverulentus (Gerst.).
PLATE IV.
Wings of Isostenosmylns spp. <J.
1. I . fasciatus sp. n., Type.
2. I. nigrifrens sp. n., Type.
SUBFAMILIES STENOSMYLINAE AND KALOSMVLIN AK 201
PLATE V.
Wings of Euporismus and Australysmus.
i. Euporismus albatrox Tillyard, $ Type.
2. Australysmus lacustris sp. n., $ Paratype.
PLATE VI.
Wings of Kempynus spp.
i. K. incisus (McL.) $.
2. K citrinus (McL.) J.
3. K. citrinus (McL.) $ var.
PLATE VII.
Wings of Kempynus spp.
1. K. latiusculus (McL.) ?.
2. if. longipennis (Walk.) Type.
3. K. falcatus Navas ?.
PLATE VIII.
Wings of Euosmylus stellae (McL.) and varieties.
1. E. stellae (McL.).
2. E. stellae, var. connexus (McL.).
3. E. stellae, var. obliteratus (McL.).
NOVIT. ZOOI.., 42, I l6
NOVIT. ZOOL., 42, C940.
KIMMINS I 'I. ATE I.
9k
\ 1 1 1 . mi] wings oi Slenosmyltcs and Stenolystnus.
r HI h
NOVIT. ZOOJ... 42, 1940.
KIMMINS PLATE II.
i
Anterior wings oi [1 male Oedosmylus.
Son, //./.
NOVIT. ZOOL., 42, 1940.
KIMMINS— PLATE II t.
Wings of male Isostenosmylus.
Photo I) I K
NOV] f. ZOOL., 42, 1940.
KIMMINS- I'l ATE [V.
^>z
^
if T-
a "-!
1
Wings of male Isoslenosmylus.
Photo l>l K
1 ..
NOVIT. ZOOL., 42, 1940.
KIMMINS— PLATE V.
Willis 11I Euporismus and Australysmus.
NOVIT. ZOOL., 42, [940
KIMMINS PLATE VI
Wings of Kempynus.
Phi I I) I A
NOVIT. ZOOL., 42, 1940.
KIMM1NS- -I'l ATE \'II.
Wings of Kempynus.
Son, Ltd.
NOVIT. ZOO].., 42, 1940.
KIMMIXS -PLATE VIII.
Wings til Euosmyltts slellae (Mel .) and varieties
AJIard {■ Son, Ltd.
AUSTRALIAN PAMBORINI, OZAENINI AND SCARITINI 203
ON AUSTRALIAN PAMBORINI, OZAENINI AND SCARITINI
(COLEOPTERA, CARABIDAE)
By M. BANNINGER.
TG. SLOANE was greatly hindered in his numerous studies on Australian
• Carabidae by being unable to consult many of Castelnau's and Blackburn's
types. The former are mainly in the Genoa Civic Museum and the latter in the
British Museum. Through the courtesy of Mr. Gilbert J. Arrow and Prof. O.
de Beaux I have been able to examine these, for which I express my sincere
thanks. The study of the types has enabled me to give new keys to the genera
Scaraphites, Euryscaphus and Philoscaphas , and to add remarks on some hitherto
doubtful names or supposed synonymy. Descriptions are also given of two new
forms of Pamborus and Mystropomus, which were among the Carabinae sent me
by Mr. P. J. Darlington, jun., for naming. They had been collected by himself
during his visit to Australia a few years ago.
Abbreviations : B = Coll. Banninger ; BM = British Museum (Natural
History) ; G = Genoa Civic Museum ; ZM = Zoological Museum, Berlin.
(!) signifies that the identity has been established by examination of the
type. For references to Sloane's papers the year of publication is given.
Tribe PAMBORINI.
I give below a key to those species of Pamborus which are known to me.
Geographical distribution is given only as far as it is definitely known ; uncertain
localities are omitted.
1. Elytra, in anterior half at least, with 6-8 broad and strong intervals
(including the sutural one) on each, of which the 7th and 8th are some
times less marked and more or less disintegrated into small tubercles.
Last ventral segment of <J truncate ...... 2.
Elytra each with about 13-15 narrow carinate intervals at least in anterior
half, more or less disintegrated into tubercles at sides and behind. Pro-
thorax with lateral basal impression uniting in a concavity with
marginal channel. Neck constriction behind the eyes deep in middle of
head. At least the last ventral segment more or less roughly rugose
or punctate .......... 6.
2. Prothorax with lateral basal impression uniting with marginal channel in
a broad concavity, the convex space between them widely separated
from posterior margin or reaching it but incompletely and flattened
(sometimes in opacus) ......... 3.
Prothorax with lateral basal impression uniting with the marginal
channel at the very narrow basal margination ; the space separating
them very convex close to posterior margin. Last ventral segment
in anterior half not roughly rugose or punct.i 1 e .... 5.
3. Aedeagus dentate on inner side not far from apex. Neck constriction
behind eyes more or less deep in middle of head. Elytra with 7th
novit. zool. 42, 1 17
204 M- BANNINGER
interval catenulate by a varying number of interruptions, the 8th
generally a little shorter but strong and entire. Surface more or less
greenish generally, especially on prothorax, partly bluish, sometimes
the elytra of a bronze-copper colour ; such specimens of (? individually)
smaller size and posteriorly more narrowed prothorax are probably
morbillosus Boisduval). Length, 24-34 mm- Victoria to Brisbane
alternans Latreille.1
Aedeagus not dentate at inner side near apex. Neck constriction of
head less deep or obliterated at middle ...... 4.
4. Elytra with 7th and 8th intervals very strong, the 7th not catenulate, the
8th not or scarcely crenulate. Surface more or less greenish. Length,
25-33 mm- Southern Queensland to Canoblas Mts. and Coonabarabran,
N.S.W. (Sloane, 1905) ....... viridis Gory.
Elytra with external intervals much narrower and less elevated than
the inner ones, the 7th catenulate by several interruptions, the 8th
more or less crenulate or disintegrated into short tubercles (but in a
single $ from Millaa Millaa near Cairns, Mus. Comp. Z00L, as in
viridis). Elytra generally more opaque and less convex. Neck con-
striction more or less obliterated in middle. Length, 27-32 (-35) mm.
Cairns district, N. Queensland . . opacus Gehin2 (Sloane 1905).
5. Elytra with 3rd interval catenulate at least in posterior half, the 5th and
7th more or less catenulate also in anterior half ; no line (or only a
vestigial one) of elevated small tubercles in the striae. Length, 21-
25 mm. Clarence River, Macpherson Range . macleayi Castelnau (! G).
Elytra with 3rd, 5th and 7th intervals entire, sometimes partly inter-
rupted in posterior half ; the striae with a line of elevated small
tubercles. Length, 21-24 mm. Brisbane brisbanensis Castelnau (! G).
6. Elytra with reflexed border bearing a few widely placed raised serrations.
Hind angles of the rather broad prothorax short and broadly advanced.
Last segment of o truncate. Length, 15-19 mm. Sydney to
Brisbane guerini Gory.3
Elytra with edge of reflexed border smooth ..... 7.
7. Last ventral segment of $ truncate. Prothorax broader, with short and
broadly advanced hind angles. Elytra more convex, the 4th and
8th intervals not catenulate. Length, 24-25 mm. Herberton
district, N. Queensland ; 1 $ Mac Ivor River (B) . . elegans Sloane.
Last ventral segment of rj rounded as in the $. Hind angles of the
longer prothorax narrower and strongly lobate. Elytra less convex . 8.
8. Elytra with 4th, 8th and 12th intervals more or less catenulate, the 14th
disintegrated into small longitudinal tubercles. Larger, broader.
Length, 23-26 mm. ; breadth, 8-5-10 mm. Dorrigo Plateau, west
of Coffs Harbour pradieri Chaudoir.
1 The examples from the Macpherson Range and Upper Richmond River have the side
margin of the prothorax broader, more grooved and not so sharply upturned as in those
from the other localities, which I consider as typical.
2 My interpretation is evidently identical with Sloane's, who also named specimens from
the same region opacus Gehin from the description. But Gehin's specimen was said to come
from the Murray River in N.S.W. , which is probably wrong. I do not know any alternans
to which the description applies, whilst it agrees very well with the species in question.
3 The colour varies very much : deep black, or elytra with golden margin, or the whole
elytra with a coppery or brassy lustre. The var. chaudoiri Castelnau mentioned in Csiki's
Catalogue has not been described, and the name does not occur in Castelnau's collection in
Genoa.
AUSTRALIAN PAMBORINI, OZAKNINI AND SCARITINI 205
Elytra with intervals much more entire and more regular, the 4th not at
all or only once interrupted in anterior half, the 14th entire in anterior
half. Smaller and rather narrower. Length, 20-23 mm. ; breadth,
7-5-8-5 mm. Barrington Tops, N.S.W. (S. of Dorrigo Plateau)
subsp. n. darlingtoni Banninger.
Pamborus pradieri darlingtoni subsp. n.
Chaudoir gives the length of his pradieri as 23 mm., and says expressly that
the 4th, 8th and 12th intervals are catenulate and the 14th disintegrated into
small longitudinal tubercles. The exact locality was unknown. These characters
agree exactly with those of six specimens before me from Dorrigo Plateau, Wesl
of Coff's Harbour, N.S.W. (Mus. Comp. Zool., Cambridge, Mass., and my coll.).
Sloane knew the species from the nearby Bellinger River and I have a specimen
named by him from Comboyne, N.S.W. The catenulation of the 4th interval
varies somewhat ; in one example it is missing in the anterior half. All intervals
are disintegrated at least from the anterior margin of the 4th segment. Upper
side with a coppery or greenish lustre, more pronounced at margin of elytra.
Length, 23-26 mm. ; breadth of elytra, 8-5-10 mm.
Further south, on the Barrington Tops Plateau, 5000 ft., Mr. P. J. Darlington
found in great number (about 60 specimens, of which 55 are before me) a very
interesting subspecies which is named after its discoverer darlingtoni subsp. n. It
is smaller and rather narrower. The intervals are much more entire, more
regular and reach nearer the apex before being disintegrated into tubercles.
The 4th is not catenulate, but in five examples there is a pit on one side or on
both sides in the anterior half. The 14th is more or less entire at least in the
anterior half. Lustre of elytra more greenish except near margin.
Length, 20-23 mm. ; breadth, 75-85 mm. Type and cotypes in the
Museum of Comparative Zoology, Cambridge, Mass., and, through the courtesy
of Mr. P. J. Darlington, in my own collection. Cotypes will also be sent to the
Queensland Museum.
Tribe OZAENINI.
Mystropomus regularis sp. n.
In the two species hitherto described, viz. subcostatus and chaudoiri, each
elytron bears, including the sutural one, four shining ribs on the very dull ground.
In chaudoiri traces of an additional interval are present between each two ribs ;
it is indicated by close shining granules, which are often dispersed also over the
other parts of the elytra. In the new species, regularis, the alternate intervals
are not obliterated nor different. Each elytron shows seven scarcely elevated
stripes, produced by a stronger lustre and the denser shining granules. It is
impossible to call them ribs. The 7th interval is scarcely distinguishable as a
rule, whereas there are sometimes traces of the 8th interval, especially behind.
I In' length "I 1 ; Hi mm. agrees with thai ol big specimens ol subcostatus.
1 cJ, 1 ?, Millaa Millaa, Atherton Table, 2500 ft., and 1 (J, Lake Barrine,
Atherton Table, 2800 ft., N. Queensland, collected by Mr. P. J. Darlington, jun.,
of the Museum of Comparative Zoology, Cambridge, Mass. Type in that
Museum ; the cotype will be sent to the Queensland Museum ; for the gift oi
the specimen from Lake I '..mine 1 am very much indebted to Mr. Darlington.
The pronotum is wider in front in regularis and more narrowed behind than
in average specimens of subcostatus, but judging by the greater material ol the
NOVIT. ZOOL., 42, I I7§
2o6 M. BANNINGER
latter at hand the shape of the pronotum is liable to considerable variation. The
distinctness of subcostatus and chaudoiri is somewhat uncertain. I have seen
the type of the latter (G). Identical specimens (B) are before me from Parra-
matta, Richmond River and Killarney Plateau in the extreme south of Queens-
land (both Mus. Comp. Zool. Cambridge, Mass.). In addition to the described
formation of the intervals the small size of 9-11-5 mm. and the long basal part
of the pronotum, the sides of which diverge rather strongly behind, are charac-
teristic. Amongst 16 specimens of subcostatus at hand from Salisbury, N.S.W.
(near Barrington), as well as in 3 specimens from Gosford, near Sydney, definite
approximations in all these respects occur, whereas there are hardly any in
12 specimens from the Dorrigo, west of Coff's Harbour.
Tribe SCARITINI.
Genus SCARAPHITES Westwood.
Scaraphites Westwood, 1842, Arcana Ent., 1 : 157.
Palpi filiform, last joint not triangular nor securiform. Triangular projection
of clypeus at each side of labrum wanting or scarcely marked. Suborbital grooves
to receive the antennae single, not divided. Paragenae not separated from sub-
mentum by a sharp oblique groove beginning at hind angle of mentum. Base
of elytra without ocellate punctures. Elytra without a costa at sides, lateral
border visible from above in its whole length. Upperside of front tibiae apically
with three strong teeth, without additional denticulations above the upper tooth.
The bifurcation of the two lower teeth, seen from behind, of variable position
with regard to the insertion of the tarsi. Entirely black, without metallic lustre.
Genotype : Scarites silenus Westwood 1842.
Key to Species.
1. Elytra without widely separated strong punctures on lateral declivity,
within the umbilicate series, and on apical declivity. Border at
shoulder not dentate and not folded over. Prothorax very feebly
sinuate on each side posteriorly, without setigerous puncture in the
angle ; base more or less imperfectly margined, especially at each side of
peduncle. Genae strongly conically projecting below the eyes.
Bifurcation of the two lower teeth of front tibiae (seen from behind)
at level of insertion of tarsi ; the excavate posterior surface in the
vicinity of the two upper teeth coarsely rugose. Intermediate tibiae
apically with a strong acute tooth. The episterna are narrower than
the epipleurae at the same level in a transverse line starting from
the inner anterior angle of the metepisterna ..... 2.
Elytra with some widely separated strong punctures on lateral1 and
apical declivities, in lucidus and humeralis on apical half only. At the
level mentioned above the episterna are of about the same width as the
epipleurae {silenus more or less excepted) ..... 3.
2. Smaller, narrower. Umbilicate series behind the shoulder only a little
farther from outer margin than in the apical third. Length, 34-40
mm. North Australia ; Ord River in N.W. Australia (Sloane 1898)
laticollis Macleay 1866. 2
1 The series corresponds to the 7th stria.
1 S. gigas is hardly more than a subspecies. Two specimens with the doubtful locality
Swan River (B) of 34 and 37 mm. in length are evidently less typical laticollis than the c?
I have from Darwin, N. Territory.
AUSTRALIAN PAMBORINI, OZAENINI AND SCAKITINI 2Q-]
Larger, relatively shorter and broader. Elytra with the border more
explanate, especially near the shoulder, where the umbilicate series is
much more distant from it than in apical third. Prothorax more
deeply emarginate in front, especially near angles, the sides more
converging to base, more rounded towards the hind angles and the
latter scarcely marked. Length, 37-51 mm. West Australia :
Murchison District to King Sound . subsp. gigas Castelnau 1867 (! G).1
3. Elytra with border wide at humeral angles, not folded over inwards,
but with edge interrupted just behind shoulder to form a humeral
prominence, projecting outwards. Prothorax shortly and strongly
sinuate on each side posteriorly, angles sharply marked, subrectangular,
with a setigerous puncture, basal margin more or less entire. Bifur-
cation of the two lower teeth of the front tibia not beyond the insertion
of the tarsus ; the excavate posterior surface in the vicinity of the
two upper teeth very smooth. Intermediate tibiae with external apical
tooth acute. Hind tibiae a little dilated at apex .... 4.
Elytra with border thickened and folded over inwards at humeral
angles and forming a tooth (sometimes much less so in hirtipes).
Genae somewhat projecting conically beneath eyes. Prothorax not,
or very feebly, sinuate on each side posteriorly, basal angles rounded
or very little marked. Bifurcation of the two lower teeth of the front
tibia markedly beyond the insertion of the tarsus ; the excavate
posterior surface of the two upper teeth and vicinity, with rare indivi-
dual exceptions (? worn), more or less rugose . . . . 5.
Elytra with border continuous at humeral angles, not or only a little
thickened, not folded over and not dentate (see also hirtipes). The
excavate posterior surface of the front tibia very smooth in the
vicinity of the two upper teeth ....... 6.
4. Border of the elytra very sparsely granulate near umbilicate series, the
declivity opaque but not granulate behind. Border of prothorax very
narrow and sharply reflexed, the sinuation near base generally strong
and very short. Length : 22-31 mm. South West Australia
(Chaudoir's locality, Melbourne, is certainly wrong)
lucidus Chaudoir 1863 (Sloane 1905).
The entire border of the elytra (as far as the row of strong punctures)
and the apical declivity densely granulate and opaque. Border of
prothorax broader, especially behind, and the sinuation at base gener-
ally a little longer. Front tibia with apical tooth on posterior side, at
base of insertion of tarsus, longer than in lucidus. Length, 30-41 mm.
" West Australia," Rottnest Is . . humeralis Castelnau 1867 (! G).
5. Intermediate tibia with upper apical tooth acute. The undulate striae
of the head ± confined to the frontal furrows. Border of the pro-
thorax not or feebly crenulate. Elytra ± regularly oval, not strongly
narrowed to base. Length, 22-36 mm. New South Wales (Sydney),
Eastern Victoria, King Is., Tasmania. Synonyms : macleayi Westw.
1842 (Cast. 1867, SI. 1905) ; intermedins Macl. 1865 (! Sloane 1905) ;
insulanus Sloane 1888 (! Sloane 1905) . rotundipenms Dejean 1825.
Intermediate tibiae with upper apical tooth dilated and obtuse. The
1 In one from Roebuck Hay (BM) the rugosity of the elytra is extremely rough and deep.
and the striae very distinct.
208 M. BANNINGER
undulate striae of the head much more extended. Border of pro-
thorax ± crenulate. Elytra very strongly narrowed to base, greatest
width much behind middle. Length, 23-34 mm- Western Victoria,
South Australia. Synonyms : crenaticollis Macl. 1864 (! Sloane 1893) ;
assimilis Sloane 1893 (! SI. 1905) hirtipes Macleay 18641 (! Sloane 1893).
6. Front tibia with the bifurcation of the two lower teeth not or very little
beyond the insertion of the tarsus. Intermediate tibia with upper
apical tooth almost regularly narrowed, acute or somewhat rounded
at the apex. Genae when seen from above not or only a little higher
than the eyes, very sharply projecting below eyes seen from the front.
Base of prothorax rounded or widely and feebly truncate, hind angles
not dentate, sometimes a little marked, margin entire at base. Elytra
of cJ extremely short, little longer than broad, sides very strongly
rounded, of the $ a little longer and the sides a little less rounded.
Length, 28-43 mm. Southern West Australia. Synonyms : cJ
bacchus Westw. 1842 ; <$ heros Castelnau 1867 (! G) ; ? master si
Macleay 1869 (! Sloane 1905)2 . . . silenus West wood 1842 ($).
Front tibia with the bifurcation of the two lower teeth much beyond the
insertion of the tarsus. Intermediate tibia with the external apical
tooth flattened and very broad to the rounded, straightly or obliquely
truncate apex. Genae, seen from above, projecting conically beyond
the eyes ........... 7-
7. Hind tibia with apex strongly produced and prominent externally.
Elytra with lateral border narrow at humeral angles. Sides of pro-
thorax not, or only faintly, sinuate behind, hind angles not (or faintly)
marked, posterior margin more or less obliterated. Length, 25-35
mm. South West Australia : Eucla (Sloane), Swan River (B).
South Australia : Yorktown (ZM). . subsp. pacificus Sloane 1888.
Hind tibia less dentate at apex. Elytra with lateral border wide at
humeral angles, striae faint, 8th interstice not convex. Prothorax
with sides rather strongly sinuate behind and basal angles rather
strongly dentate, front margin emarginate, especially near the broad
angles, which are distinctly prominent. Length, 31-36 mm. " Aus-
tralia "(B) . . . . . . lenaeus Westwood 1842.
Differing from lenaeus by less sinuate sides of prothorax and not (or
scarcely) dentate angles ; front margin hardly emarginate and angles
not produced. Elytral striae rather distinct, 8th interstice distinctly
convex at middle. Length, 27-36 mm. South West Australia : King
George Sound (Macleay, BM), " King River, south West Australia "
(BM) ....... subsp. latipennis Macleay 1863. 3
According to Sloane's Checklist (1905) four species remained unknown to
him : bacchus Westwood, heros Castelnau, humeralis Castelnau and martini
Castelnau. There is a fifth, confusus Westwood, which is not mentioned in any
1 The form of the shoulder of hirtipes is sometimes intermediate between that of rclundi-
pennis and of pacificus. In accordance with Sloane's decision and the rules of nomenclature
(Article 28), but contrary to Csiki's Catalogue, I give hirtipes priority over crenaticollis, though
the latter is described a page before.
2 As Sloane suggested, having the type before him, there seems scarcely any doubt
that mastersi Macleay from Mt. Barker is described from a ? of silenus. A $ specimen from
the adjacent Albany is in the British Museum.
3 S. pacificus, lenaeus and latipennis are very closely allied and probably at most sub-
species or perhaps individual aberrations of one species.
AUSTRALIAN PAMBORINI, OZAENINI AND SCARITINI 200,
of his papers. The study of Castelnau's types at Genoa enabled me to elucidate
two of the above names: humeralis is closely allied to lucidus, and heros is a
synonym of silenus. On the remaining three I make the following remarks :
The types of S. bacchus and silenus Westwood are missing from the Hope
Department of Entomology, University Museum, Oxford. The description says
that in bacchus the genae are produced into an angle beyond the eyes, while tin-
is not so in silenus. But in strong males they often also project considerably
beyond the eyes. Both are described from the same locality, Swan Kiver, and
comparing the two figures and descriptions, there seems to me no doubt that
bacchus is the broader <J and silenus the more parallel 9 of the same species,
which, according to the rules of nomenclature, has to take the latter name, as
Sloane chose it in his revisions. There are specimens of both sexes from Swan
River named bacchus in Castelnau's collection, which supports the above con-
clusion, and I have seen a specimen from Perth (BM) labelled by Sloane as
" silenus Westwood which is a small form of bacchus Westwood."
The type of S. lenaeus Westwood is also missing from Oxford, but there is a
specimen with the label in Westwood's handwriting " lenaeus West., ? small
male." I have examined it, through the courtesy of Prof. G. D. H. Carpenter.
It is hirtipes, the teeth of the intermediate tibiae disagreeing with figure and
description. I quite agree with Sloane's interpretation of lenaeus, having in my
collection a previously unnamed 9 with the very old label " N. Holl." 5. lati-
pauiis Macleay is evidently very close to it and hardly more than a subspecies.
There is a o of the latter from the type locality, King George Sound, in tin-
British Museum.
The type of 5. martini Castelnau is not at Genoa. According to the author,
the entire body is smoother than in rotundipennis and the punctures on the
declivity are larger. This agrees very well with the example I consider to be
lenaeus, which synonymy Sloane conjectured in 1907. But Castelnau's descrip-
tion does not say anything about the very different shoulders and posterior
angles of the prothorax. It seems to me better to leave the decision to the future.
The description of S. confusus Westwood is very poor and gives no locality.
It is impossible to say what it may be.
Genus EURYSCAPHUS Macleay.
Euryscaphus Macleay. [865, Trans, ent. Sue. A'. .SMI'., 1 : 187.
Palpi with the last joint triangular or securiform. Lower edge of genae
prominent and usually projecting in front below the eyes. The suborbital
grooves to receive the antennae are single, not divided. Parageiiae with a sharp
oblique, often almost transverse groove, beginning at the hind angle of the
mentum and separating the paragenae from the submentum and gula. Clypeus
on each side of the labrum with a more or less triangular projection. Pronotum
with the base sharply bordered above peduncle. Elytra without a keel on sides,
lateral border visible from above in its whole length, base with - e ocellate
punctures. Epipleurae very broad anteriorly, twice as wide as the metepistema,
or more. Upperside of front tibia witli two strong teeth, theii bifurcation, when
viewed from behind, much above the insertion of the tarsus; a variable
number of much smaller denticulations above the upper tooth. Mesotibia
witli the apical tooth acute, often rather small. Elytra very broad and short,
in the £ generally nearly circular, more elongate in the j>. Entirely black,
without metallic Lustre.
210 M. BANNINGER
The affinities of some Enryscaphus with the genus Carenum are very great.
I have from the Boileau collection two very large ? specimens (length, 34-35
mm. ; breadth of elytra, 13 mm.) from " N.W. Australia " which were named E.
hopei, and for some time I also considered them to be a Euryscaphus near politus,
in spite of their shape, which is too oblong for the genus. But the paragenae
are not defined behind by the deep oblique groove which is so characteristic of
Euryscaphus ; the genae do not project below the eyes and the front tibiae are
strongly tridentate. In Sloane's classification of Carenum (1900) into groups
one comes near the macleayi and transversicollis groups, but my specimens do not
agree with any of the species in question. In the anthracinum group of Carenum
and the groups with the penultimate joint of the labial palpi short and swollen,
grooves like those in Euryscaphus separate the paragenae from the submentum
and the gula.
Genotype : Euryscaphus angulatus Macleay 1865.
Key to Species.
1. Posterior angles of pronotum rounded, the base more or less rounded
between them or more or less lobate above peduncle ... 2.
Posterior angles of pronotum upturned and more or less rectangular.
Elytra without a discoidal puncture on apical third ... 6.
2. Elytra with a discoidal puncture about one third from apex . . 3.
Elytra without a discoidal puncture on apical third .... 4-
3. Lateral border of elytra very narrow, except near shoulder, feebly rounded
to the shoulder-tooth which is very distant from the peduncle. Pro-
notum with lateral and basal borders rather narrow, basal border
normal on each side of peduncle, epipleurae not (or hardly) visible from
above, base not (or scarcely) lobate, the wide longitudinal impression
at a considerable distance from lateral border wanting or very shallow.
Length, 31-43 mm. South Australia, Western New South Wales,
Victoria. Synonyms : $ bipunctatus Macleay 1865 (Sloane 1893) ; 3
howitti Castelnau 1867 (! Sloane 1893) ; 9 tatei Blackburn 1887 (! BM) ;
ferox Sloane 1888 (! 1890) , <J chaudoiri Blackburn 1892 (! BM)
obesus Macleay 1863 (! Sloane 1893) ?.
To be separated from obesus by the very broad lateral border of the pro-
notum and the stronger wide impression at a considerable distance from
lateral border. Length, 32 mm. ; breadth, 13 mm. South Australia
(Lake Eyre) . . . subsp. sulcicollis Blackburn 1892 (! BM).
Elytra with lateral border broad to apex, anteriorly shortly and strongly
rounded to the shoulder-tooth (sometimes more or less obliterated),
which is much nearer to the peduncle than in obesus and sulcicollis.
Lateral border of pronotum very broad, base generally strongly lobate,
the border very narrow on each side of peduncle and the epipleurae
visible from above, generally with the same wide shallow impression
at a considerable distance from lateral border as in sulcicollis. Genae
more projecting below eyes than in the two preceding forms. Length,
31-44 mm. South-West Australia. . subsp. ebeninus Sloane 1890.1
4. Reflexed border of elytra ending at humeral angle and forming a very
thick upturned humeral projection, less so in the $. Pronotum with
base rounded and strongly produced backwards above peduncle, lateral
1 There can hardly be any doubt that the three forms are subspecies of a single species.
AUSTRALIAN PAMBOKINI, OZAENINI AND SCAKITINI 211
margin more or less crenulate, epipleurae more or less broadly visible
from above on each side of peduncle. Very large species, 9 more
elongate than <J, which has almost round elytra. Length, 35-51 mm. ;
breadth, 15-24 mm. Central and North Australia. West Australia
(Lake Darlott). Synonym : $ titanus Sloane 1889 (! Sloane 1893)
waterhoasei Macleay 1864.
" Size moderate, elytra narrow and narrowly rounded behind." " Pro-
bably founded on a small form of waterhousei ." Length, 31-33 mm. ;
breadth, 12-13 mm. Bourketown District, N. Queensland. Ex Sloane
1893, 1897 ........ atratus Sloane 1893.
Reflexed border of elytra extending more or less beyond humeral angle,
not forming a thickened upturned projection. Lateral border of pro-
notum not crenulate, epipleurae not or very narrowly visible from
above on each side of peduncle . . . . . . 5.
5. Elytra rounded, not distinctly angular, at shoulder. (Base of pronotum
rounded and more or less lobate above peduncle.) Length, 22-33 rnm. ;
breadth, 10-14-5 mm- S. Queensland, New South Wales, Victoria,
? Eastern South Australia, " Central Australia." Synonyms : minor
Macleay 1865 (! Sloane 1905) ; affinis Castelnau 1867 (! Sloane 1893 =
minor Macleay) ; 9 hopei Castlenau 1867 (! G) ; arenarius Sloane
1888 (! Sloane 1890 = minor Macleay)
dilatatus Macleay 1865 (! Sloane 1905).
Elytra very distinctly angular at shoulder. (Base of pronotum more
truncate, not or less lobate above peduncle.) Length, 25-34 nun. ;
breadth 11-15-5 mm- S. Queensland, South, Central, and West
Australia, as far North as Murchison River. Synonyms : $ politus
Sloane 1893 ; ^$ concolor Sloane 1893 ; $ terrenus Sloane 1894
carbonarius Castelnau 1867.
6. Supraorbital pore present. (Frontal sulci less deep behind than in sub-
sulcatus, and fading out in the generally somewhat depressed neck.)
Central carina of mentum reaching the end of the tooth ; its sides
normally bordered. Marginal border of pronotum, including posterior
angles, much more widely upturned, with an anterior marginal puncture
and a broad flat depression inside anterior angles. Shoulders much
more rounded off than in subsidcatus. Length, 33-36 (-42) mm.
" Queensland " (Sloane, Check List 1905) ; Peak Downs (ZM) ; 1 9
without locality (B) . . . . . angulatus Macleay 1865.
Supraorbital pore wanting. (Frontal sulci sharp and deep behind, no
neck constriction.) Central carina of mentum reaching the strongly
upturned sides of the tooth. Lateral border of pronotum strongly
but not very broadly upturned, without setigerous punctures, trans-
verse impression behind front margin hardly perceptible even near
anterior angles. Shoulders square, strongly marked. Length, 22-29
mm. South Australia (Ouldea) . subsulcatus Blackburn 1887 (! BM).
E. carbonarius Castelnau and dilatatus Macleay.
It is almost impossible to name specimens belonging to group 5 from the keys
and descriptions hitherto published. A specimen from Queensland (BM) named
dilatatus Macleay by Sloane gave a hint. Another specimen which I recently
saw in the British Museum collection simplifies the question considerably. It
212 M. BANNINGER
is from Norseman, W.A., and was obtained from Sloane himself in 1923, thus
many years after his latest publication on the genus. Perhaps he had again
examined Castlenau's type in the Howitt collection (see Sloane 1905, 1907). It
bears the label " E. carbonarius Cast." with politus Sloane, concolor Sloane and
terrenus Sloane as synonyms. According to the descriptions and the material
at hand the differences of the pronotum are evidently subject to some variation,
which greatly complicates interpretation of the descriptions. The shape of the
shoulder affords a better character and it agrees very well with the geographical
distribution. There is, however, some doubt, as the type of carbonarius is said
to come from Cooper's Creek, which would be the extreme eastern point of dis-
tribution, and E. affinis Castelnau is from the same locality and was identified
from the types by Sloane as minor Macleay = dilatatus Macleay. It seems to
be somewhat doubtful whether the synonymy of the nine names is quite correct.
Genus PHILOSCAPHUS Macleay.
Philoscaphus Macleay, 1871, Trans, ent. Soc. N.SAV., 2 : 96.
Palpi with the last joint triangular or securiform. Lower edge of genae not
dentate nor projecting over the eyes. Suborbital grooves to receive antennae,
single, not divided. Paragenae not separated from mentum and submentum by
an oblique groove (sometimes a little marked in bamardi). Clypeus on each side
of labrum with a triangular projection. Elytra with one or two sharp costae on
each side, which disappear near apex. Base with some ocellate punctures, often
more or less disappearing in the rough sculpture. Shoulder dentate. Epi-
pleurae broad. Upper side of front tibia more or less tridentate, with or without
further denticulations above the upper tooth. The bifurcation of the two lower
teeth, seen from behind, above the insertion of tarsus. Colour variable.
Genotype : Carenum tuberculatum Macleay 1863.
Key to Species.
1. Disc of the elytra perfectly flat and quite smooth and opaque on inner
side of costa. Lateral margin and base of thorax and lateral margin
of elytra, as well as an irregular patch inside shoulder, of a bright green
or red-golden colour ; ventral surface cyaneous. Costa following the
curvature of the elytra widely separated from the margin, which is
visible from above in its whole length. Length, 14-16-5 mm. South
Queensland ....... bamardi Macleay 1887.
Disc of elytra with a very rough sculpture on inner side of costa (tubercles,
elevations). Upperside black . . . . . . .2.
2. Disc of elytra fiat on inner side of costa, with a few longitudinal impres-
sions like striae and a number of transverse and irregular ones, giving
a very uneven appearance. Costa seen from above very distant from
the lateral margin, towards which it slopes almost vertically ; lateral
margin broadly visible from above in its whole length.1 Length, 16-5-
18-5 mm. S.E. New South Wales. (Macleay 1873, 3. 324)
carinatus Macleay 1864.
1 The shape of elytra and position of the costa are, except for the rough sculpture extend-
ing to the lateral margin and black colour, much as in bamardi.
AUSTRALIAN PAMBORINI, OZAENINI AND S( AKITINI 21}
Lateral margin of elytra very narrowly visible from above in anterior
half or somewhat hidden by the costa. Disc more or less convex on
inner side of costa, with longitudinal rows of broad elevations and
scattered tubercles in the intervening space. Epipleurae very broad
Head smooth or only with traces of rugae between the eyes and the
furrows. Thorax strongly lobate above peduncle, disc generally
more or less smooth or with moderately deep transverse rugae . . 3.
Lateral margin of elytra completely hidden in its whole length, except
just behind shoulder and for a certain distance at apex, by the strongly
overhanging costa ; the large elevations generally less numerous, the
sculpture more disintegrated into more numerous smaller tubercles.
Epipleurae considerably narrower. Thorax strongly lobate above
peduncle, disc with very strong transverse rugae .... 4.
3. Elytra with a single costa at side (generally with marked bifurcation at
level of 2nd or 3rd segment). Length, 21-32 mm. New South Wales,
Queensland (Kuranda, B), Eastern South Australia
tuberculatum Macleay 1NO3.
Elytra with two costae at side, at least for some extent, corresponding
to the 7th and 8th interstices, the 8th more or less obliterated an-
teriorly or fused with the 7th, and both (especially the 7th) often more
or less interrupted posteriorly. Length, 23-33 mm- West and South
Australia. Synonyms : lateralis Macleay 1873 (! Sloane 1905) ;
crassus Blackburn 1887 (Sloane 1905) ; tepperi Blackburn 18871
costalis Macleay 1873 (! Sloane 1905).
4. Disc of elytra with one or two rows, among the irregular tubercles, of
stronger or more or less distinct tubercles which correspond to the
3rd and 5th interstices, the latter beginning at about middle of upper
visible part of basal carina. Length, 22-30 mm. S.E. Queensland
to Kuranda. Synonym : duboidayi Blackburn 1892 (! BM)
masters! Macleay 1871.
Disc of elytra with a semi-interrupted costa of two-thirds of their length,
beginning at middle of upper part of basal carina. Elytra narrower.
with base decidedly narrow. Sculpture of lateral declivity more
tuberculate above lateral channel, with a row of narrow elevations,
forming the line of a broken costa. The presternum has the inter-
coxalpart lightly channelled, not deeply excavate at the base. Length,
22-5 mm. ; breadth, 8-3 mm. North Queensland. Ex Sloane
bicoslatus Sloane 11)05.
' Sloane (1905:114), from description, placed /'. tepperi Blackburn (Angebuckina
in the synonymy of tuber culatus, though the description speaks oi two costae .it middle oi
each elytron for an extent of 2 mm. I cannot find any othei difiereni e hot ween tuberculatus
and costalis except the presence ol one or two costae. P. tepperi seems to be founded on an
example connecting them. There is a specimen (BM) < allabonna identifie I a << pperi
I'iLh L! mi 11 by A. M. Lea which, as well as the localities, supports that suggi ition,
(MSS. reed. July 22, 1939.)
NOTE ON CERCARIA IMBRICATA I.OOSS 215
CERCARIA IMBRICATA LOOSS 1896, nee 1893— A NOTE
ON NOMENCLATURE
By MIRIAM ROTHSCHILD.
IN 1938 a paper dealing with Notocotylid cercariae was published in which
it was proposed to divide these larvae into three main groups according to
the structure of the excretory vesicle (Rothschild 1938). " Cercaria imbricata
Looss, 1896" was selected as the " type " of the second group, characterized by
the anterior portion of the vesicle looping between the lateral eye-spots and
passing anteriorly to the median eye-spot. It is felt that attention should be
drawn to a slight error in nomenclature, the larva in question being more correctly
designated as " Cercaria imbricata Looss 1896, nee 1893." The reasons for the
correction are given below.
In 1893 Looss published a paper where in a footnote (p. 20) he records a
Monostome cercaria from Paludina impura from Leipzig (Germany). To this
larva he gave the provisional name (" provisorischen Namen ") Cercaria imbricata.
He considered this form might eventually develop into Monostomum attenu-
atum Rud. Where cercariae are concerned the term " monostome," together with
the indication of the host, is insufficient to constitute a description and the name
must be regarded as a nomen nudum.
In 1896 Looss found a monostome cercaria in " Melania tuberculata Bourg."
in Egypt which he thought was the same species as that provisionally named
C. imbricata by him in 1893. On this occasion he gives a good description, and
the name imbricata becomes valid for this species. He puts forward the view,
based on circumstantial evidence, that the adult form is Monostomum vcrru-
cosum Froel., not Monostomum attenuation.
In 1935 U. Szidat1 found a cercaria in Bythinia tentaculata (=Palwlina
impura) which she identified as Cercaria imbricata Looss. Cysts of this larva
were fed to ducklings, and an undescribed Notocotylus was recovered from the
birds. This she named N. imbricatus. The description of this cercaria is inade-
quate, but the figures clearly reveal that it is not the species found by Looss in
Egypt. The anterior portion of the excretory vesicle passes posteriorly to the
median eye-spot, thus placing the cercaria in the Monostomi group.
It seems very probable that Looss' original surmise was correct, and that
the cercaria he discovered in Paludina impura was the larval form of a Notocotylus
identical with, or very closely related to, N. attenuatus, whereas the species from
Melanin tuberculata was the larval form of Catatropis verrucosa. These two types
of excretory vesicle may well prove to be characteristic of the genera Notocotylus
and Catatropis respectively.
1 The Szidats have frequently shown a curious, even reckless disregard for the laws oi
nomenclature when naming cercariae and linking larval with adult forms. Thus in 1933
they identified a cercaria from Planorbis corneus as i . ephemera Nitzsi h, but on rearing the
adult form, described it under the name " Noto< otylus thienemanni n.sp." ! Several equally
glaring examples could be cited.
NOV1T. ZOOL., 42, I l8
2l6 MIRIAM ROTHSCHILD
Pending further research into the morphology of both cercariae and adult
forms of the Notocotylidae, it is perhaps unwise to attempt to correct the unfor-
tunate number of nomenclatural errors which characterize much of the literature
on this group. It will, however, save further confusion if Looss' species is clearly
referred to as Cercaria imbricata Looss 1896, nee 1893.
REFERENCES.
Looss, A. 1893 Zur Frage nach der Natur des Korperparenchyms bei den Trematoden,
nebst Bemerkungen iiber einige andere, zur Zeit noch offene Fragen. Ber. sacks.
Ges. (Akad.) Wiss., 45 : 10-34.
1896. Recherches sur la faune parasitaire de l'Egvpte. Premiere partie. Mem.
Inst, e'gypt., 3 : 1-252, pis. 1-16.
Rothschild, M. 1938 Notes on the classification of Cercariae of the Superfamily Noto-
cotyloidea (Trematoda), with special reference to the excretory system. Novit. Zool.,
41 : 75-83. figs. 30-40-
Szidat, L. & U. 1933 Beitrage zur Kenntnis der Trematoden der Monostomidengattung
Notoeotylas Dies. Zbl. Bakt. (Orig.), 129 : 411-422, 12 figs.
Szidat, U. 1935 Weitere Beitrage zur Kenntnis der Trematoden der Monostomiden-
gattung Notocotylus Diesing. Zbl. Bakt. {Orig.), 133 : 265-270, 5 figs.
{MSS. reed. Jan. 20, 1940.)
2 2 JUN1940
PRESENTED
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CONTENTS
The Australasian Frogs of the Family Leptodacty-
lidae. By H. W. Parker, M.A i
Neotropical Neoempheria (Diptera, Mycetophilidae) .
By F. W. Edwards, F.R.S 107
Some Sapromyzidae (Diptera) from the Solomon
Islands and New Caledonia. By John R. Malloch 131
Studies on Diopsidae (Diptera). By James P. Shillito,
B.Sc 147
A Revision of the Osmylid Subfamilies Stenosmylinae
and Kalosmylinae (Neuroptera) . By D. E. Kimmins 165
On Australian Pamborini, Ozaenini and Scarlatini
(Coleoptera, Carabhjae). By M. Banninger . . 203
CERCARIA 1MBRICATA LOOSS 1896, TWC 1893 A NOTE ON
Nomenclature. By Miriam Rothschild . . .215
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NOVITATES ZOOLOGICAE
Vol. 42
OCTOBER, 1940
No. 2
MONOGRAPHIC REVISION OF THE MEXICAN WATER
BEETLES OF THE FAMILY ELMIDAE
By H. E. HINTON, Ph.D.,
Department of Entomology. British Museum (Natural History).
(With 401 text -figures.)
CONTENTS.
Introduction ......
General Remarks . . . . .
(1) The Use of Quantitative Methods in Taxonomy
(2) The Use of the Internal Anatomy in Classification
Method ......
Material ......
Acknowledgments ....
A Key to the Tribes of Elmidae
A Key to the Genera of Larini of the World
A Key to the Genera of Mexican Elmini
A Key to the Larvae of the Genera of Elmidae
The Mexican Larini ....
The Mexican Elmini ....
References .....
PAOE
217
2l8
2l8
220
222
223
224
224
225
227
229
231
254
396
Introduction.
THIS monograph owes its origin to an attempt to determine the Elmidae collected
on an expedition undertaken by Dr. R. L. Usinger and the writer to Central
Mexico in the summer of 1933. At the time great difficulty was experienced in
assigning names to the various forms, the existing descriptions being too brief, and
in most instances unaccompanied by illustrations of essential characters such as the
male genitalia. It was also found impossible to determine the small amount of
material available of the immature stages. In the summer of 1934 an opportunity
was taken to collect again in Mexico, and on this expedition particular attention was
paid to the immature stages. As a result it has been possible in nearly all instances
to assign those stages to genera if not to species.
In this paper all available stages of each species arc described and figured. Such
parts of the internal anatomy as the alimentary canal, male and female reproductive
organs and central nervous system are described and illustrated for all genera
except one. It is believed that this is the first time that these characters of the
novit. zool., 42, 2. 1940. 19
2l8 H. E. HINTON
internal anatomy have been used in conjunction with those of the external
anatomy for defining genera in any family of the Arthropoda.
Altogether 36 species and one subspecies belonging to eleven genera are
described. Of these, nine species, one subspecies and two genera are described as
new.
General Remarks.
The Use of Quantitative Methods in Taxonomy.
The application of quantitative methods to taxonomy has been much neglected.
The usual taxonomic procedure has been, with few exceptions, entirely qualitative,
and for the most part only those characters which are conspicuously discontinuous
have been dealt with. The recent progress in the application of quantitative methods
has been reviewed by Richards (1938). Only the influence of the environment on
linear measurements will be discussed here.
It has been pointed out by Heincke (vide Richards, 1938), Zarapkin (1934) and
Richards (1938) that most of the structures of two species differ significantly in the
means of their measurements. This has always been found to occur among those
species measured by the writer. From this fact it might be concluded that significant
differences between the means of linear measurements of the structures of two
populations are good specific characters. The writer has had experience with a
group of insects living in a relatively stable environment, and therefore unusually
suitable for such studies, and has found that in these quite a small difference in
environment (in altitude and presumably in temperature) has resulted in significant
differences between the means of most of the linear measurements of two populations
belonging to the same species. Such significant differences are even apparent among
populations of the same species from the same stream, associated with altitude
differences of less than 1000 ft. (in Mexico at about 190 N.).
It is possible that in terrestrial insects the means of those living, for example,
on the north side of a hill would differ significantly owing to a difference in tem-
perature from those living on the south side of the same hill. It is also possible that
for the same reason there would be a significant difference between the means of the
same species living on opposite sides of a tree trunk. Apart from differences in
absolute size that can be attributed to temperature, the mean size of populations
is influenced by the type and quantity of food available in a particular locality, etc.
Although significant differences between the means of two populations are not
in themselves always specific characters, it might be claimed that while small
differences are not, very large differences are, and that in a particular genus or family
it is only necessary to determine the degree of variation for each category in the
taxonomic hierarchy. If this w-ere true it could be said that two populations whose
means differed by, say, 0-06 mm. belonged to different species, while differences of
less than 0-06 mm. were of subspecific importance. It is true that, on the whole,
interspecific differences in size are greater than intraspecific, but even in the writer's
very limited experience cases have been found in which the means of different
populations of the same species differ as widely as those of different species.
For example, the difference between the means of the length of the prothorax of
Heterelmis obscura taken at 8000 ft. and at 3500-4000 ft. is 0-034 mm., while the
difference between the mean of H. obscura taken at 3500-4000 ft. and that of
H. acicula taken in the same locality is only 0-022 mm. When a test of homo-
geneity is applied to these, it is found that the difference between obscura and
acicula is significant to the same degree as that between the two populations of
MEXICAN ELMIDAK
219
H. obscura, i.e. P < io~3. Furthermore, if the general decrease in size with a
decrease in altitude which has already been observed in obscura (text-fig. A) is
H. Ic,
W. o tlJJ
3,i ooft
S,l<ioft
l^oolt
Text-fig. A. — The mean length of the prothorax of five species of Heterelmis is
plotted against the altitude at which the population occurred. The means were
taken from populations of 14 to 75 individuals.
continued below the 3500 ft. level, at some poinl below this level the mean of
the prothorax length will be equal to that of acicula at 3500-4000 ft.
If all aspects of the environment were identical for two populations and
220 H. E. HINTON
significant differences could be found between the means of their linear measure-
ments, such differences would be genetical. If these differences were not posi-
tively correlated with characters of known value, it would be left to the judgment
of the observer to decide whether they were of specific or only of subspecific
importance.
For ordinary taxonomic purposes quantitative work of the type dealt with
above has a very limited application. Even in those cases where the specific
means are very different, it should be noted that there is frequently an overlap
m the measurements (vide text-figs. 366-368). This means that the species
must be first segregated on qualitative characters before quantitative work can
begin ; and at that point, unfortunately, taxonomic work usually ends.
The genus Heterelmis (text-fig. A) has been studied more extensively than
any other in this paper, and reveals the following general points : considering
one species at a time, the mean length of the prothorax increases progressively
with increase of altitude for obscura and obesa obesa (this is also true for Micro-
cylloepiis inaequalis and Neoelmis longula, p. 313 and p. 326) ; while for longula
(occurring only at the highest altitudes) an increase of altitude is accompanied
by a decrease of prothorax length. It is not known how far this effect of altitude
is due to modification and how far to selection. It is quite possible that within
obscura, for instance, certain genotypes are larger than others (in this genus
length of prothorax is positively and highly correlated with absolute size, p. 379),
and that the former flourish better at higher altitudes. It is probable, however,
that a considerable part of this effect is due to the influence exerted during
ontogeny, by the environment at different altitudes, on essentially the same
genotypes. There is no information as to which factor in the environment at
different altitudes is responsible for modifying size, but temperature may be
suggested as the most likely possibility. As regards longula, it may be suggested
that above certain altitudes — in this genus probably above 8000 ft. in lat. 19° N.
— such factors of the environment as very low temperatures have a dwarfing
effect.
Considering the relationship between the different species of Heterelmis,
with regard to altitude and prothorax length, it appears from text-fig. A that
the greater the mean altitude of the species the greater is the length of the pro-
thorax (and therefore the greater the absolute size). The scanty data obtained
for obesa plana and acicula are in general accord with this statement if it is
remembered that obesa plana does not belong to the same section of the genus
as acicula and obscura. Furthermore, when two species, e.g. obscura and obesa
obesa, are compared in the same environment, at 5600-7500 ft. for instance, it is
found that obscura is still much smaller — the difference in size being genetical.
So it may be concluded that the greater the size, as determined by genetical factors,
the greater will be the altitude of the habitat to which the species will tend to be segre-
gated. Unpublished data on a number of genera in this and other families
support this general principle.
The Use of the Internal Anatomy in Classification.
The use of internal characters in dealing with groups of family or super-
family rank in the taxonomic hierarchy has been dealt with already.1 It is
only necessary here to point out that without the use of such characters it is
much more difficult, and in many instances impossible, to build a system of
1 r939. Trans. R. Ent. Soc. Land., 89 : 133-184. 1 pi., 105 figs.
MEXICAN ELMIDAE 2.2.I
classification which gives even an approximate picture of the phytogeny of the
groups involved.
In the classification of the categories of lower rank, such as genera and
species, the internal anatomy has very seldom been used in the Arthropoda
Practically without exception, genera are based entirely on the characters ol the
external morphology. In the writer's experience there appear to be almost as
many characters of the internal anatomy which can be considered to be of generic
importance. The present study of the internal anatomy of a number of genera
has in some cases modified the conception of generic limits.
If the species of Elmidae of which the characters of the internal anatomy
are known be divided into genera solely on these characters, the resulting genera
frequently correspond exactly to genera which taxonomists have previously
founded on external structures alone. However, in some groups, e.g. Elsianus,
where the application of generic criteria used in the past, such as the presence
or absence of sublateral carinae on the pronotum, would have resulted in two
and possibly three genera, the similarity of the internal anatomy has caused a
more careful search for characters of the external anatomy which could be
considered to be of generic importance. This has led to the discovery of the
generic importance of such characters as the venation of the hind wings and the
distribution of the tomentose areas on the body — characters almost entirely
neglected in previous descriptions of genera. In the particular case of Elsianus,
it might be mentioned that the extent of the genus first suspected from a study
of the internal anatomy is supported by a study of the larvae.
Until now the genus Cylloepus seemed to consist of a fairly homogeneous
assembly of species, but an examination of the internal anatomy shows that the
species fall into two very distinct groups. These groups differ from each other in
the number of egg tubes to each ovary, the place of entry of the spermathecal
duct into the bursa copulatrix, the form of the lateral accessory glands of the
male, the number of caeca on the anterior margin of the midgut, and the number
of Malpighian tubules. Experience of the family gained elsewhere shows that
all these characters are of generic importance, and from this it is concluded that
the two groups are really very different genera. The only character of the
external anatomy which has been found to be correlated with these differences
in the internal anatomy is the absence or presence of a transverse belt of tomentum
on the hypomera. The species having a transverse belt of tomentum on the
hypomera have been placed in a new genus, Hexacylloepus. Among the described
species of Hexacylloepus a considerable amount of variation in the extent of the
tomentose belt is already evident ; and it is possible that when more species are
known, some will be found in which the tomentose belt is not developed. If this
were to occur, it would be necessary to study the soft parts of the internal anatomy
before these species could be placed in their proper genus. It is believed that
sooner or later, among some groups of genera in the family, dissection of the sofl
parts of the internal anatomy will be necessary before the species ran be refern d
to their respective genera.
No one to-day would claim that all genera should be distinguished solely on
characters of the antennae. Almost everyone would agree that all characters
of the external anatomy of the adults should be used in defining genera and in
attempting to build natural systems of classification ; and many would insist
that the external anatomy of other stages be considered also. However, in
practice, if not in theory, most systematists still draw a very sharp line between
characters of the external and internal anatomy.
222 H. E. HINTON
Specific differences in the internal anatomy certainly occur, and are often
more conspicuous than external differences. In the writer's very limited
experience specific differences are not as numerous in the gross structure of the
soft parts of the internal anatomy as they are in the structure of the external
anatomy. No instance has yet been observed of two species differing only in
the structure of the soft parts of the internal anatomy. There is no theoretical
reason why species differing in a few genes should not have the effect of these
genes confined to these parts.
Method.
While describing the species the problem arose as to how far the descriptions
should be complete. To describe and illustrate in detail every structure on the
body of each species is for all practical purposes a generally recognized impossi-
bility. Any modification of this procedure rests upon the judgment of the
individual worker. Some solve this problem by describing only those characters
which will enable one to separate the species from those already described in the
same genus. In practice this means that each time a new species, which differs
in structural details from those previously described, is added to a genus it
becomes necessary to re-examine and redescribe all the species ; and if such a
standard is accepted, whenever species are added to a genus, all other specific
descriptions in that genus must be revised. Fortunately a slightly higher
standard generally prevails to-day. The systematist selects from the characters
that he sees those which by comparison with other species of the genus, or by
analogy with the species of related genera, appear to him to be important ; and
to these he adds a few characters which he thinks are likely to prove useful when
more species are known, but makes no attempt at anything approaching a
complete description.
It has already been pointed out that a really complete description is a
practical impossibility, and that anything short of this must, at least where
those genera to which many species will be added are concerned, involve a certain
amount of redescription. The amount of redescription necessary is governed
largely by the number of species later added to the genus and the accuracy and
completeness of the existing specific descriptions. Although all redescription
cannot be avoided, an attempt is made here to describe and figure each species in
such detail that redescription will be reduced to a minimum, or, in those genera
of which few species remain to be discovered, will not be necessary at all. How-
ever, if other criteria are in the future needed to distinguish satisfactorily the
species of Elmidae, e.g. the number of branches of the recurrent nerve or the
biochemical properties of some tissue, the descriptions given here will have to be
rewritten.
Previous workers have seldom given precise measurements of the various
structures, and statements such as " prothorax longer than broad " are the rule.
It has been found that this degree of accuracy is not usually sufficient, and in
the descriptions given here the exact measurements have been made with the
aid of an eyepiece micrometer. It has also been found necessary to describe
more accurately than has been the custom the distribution of punctures and
tubercles. Statements such as " punctures coarse," " tubercles moderately
dense," etc., have been found to be too subjective and have been avoided by
giving the exact diameters of the punctures, or else comparing them with struc-
tures already measured or found to be specifically stable, e.g. the facets of the
MEXICAN ELMIDAE
223
eyes. The density of the tubercles and punctures has been expressed in terms
of their own diameters.
The 13 illustrations of the adult beetles were done by Miss 0. F. Tassart.
All other illustrations were done by the writer with the aid of a camera lucida.
Lines next to figures refer to a length of 0-20 mm. unless otherwise indicated.
Material.
The descriptions of the internal anatomy are drawn from species collected
by the writer in Bolivia, Brazil, French Guiana, Trinidad (B.W.I.), and England.
The material which forms the basis of this monograph was collected by
Dr. R. L. Usinger and the wrriter in central Mexico during the summer of 1933,
and by the writer in the same region in the summer of 1934. The entire collec-
tions of Elmidae in the British Museum (Natural History) have been available,
as have also the Mexican and Central American Elmidae in the collections of
the U.S. National Museum, the Museum of Comparative Zoology at Harvard
College, the Deutsches Entomologisches Institut at Berlin-Dahlem, and the
Museum National d'Histoire Naturelle of Paris. Collections of Elmidae have
also been sent by a number of individuals.
The following is a list of all the specimens of the species known to occur
in Mexico examined by the writer :
224
C. proximus Hinton.
Adults . . Mexico
C. puncticollis (Hinton).
Adults
C. sculptipennis (Sharp).
Adults . . Guatemala
Mexico
C. sexualis Hinton.
Adults
C. spinipes Hinton.
Adults
Cylloepus sp. ?
Larvae
Heterelmis acicula, sp. n.
Adults
H. longula Sharp.
Adults
H. E. HINTON
130.
I.
11.
98.
94-
2.
32.
I67.
H. obesa obesa Sharp.
Adults . Guatemala
Mexico
H. obesa plana, subsp. n.
Adults
H. obscura Sharp.
Adults . . Guatemala
,, . . Costa Rica
Mexico
Texas
H. tarsalis, sp. n.
Adults . Mexico
Heterelmis.
Larvae of several ,,
spp.
Total specimens
9-
5003.
1194.
38-
2.
1607.
1.
154-
16,500.
Acknowledgments.
My best thanks are due to Dr. A. D. Imms, F.R.S., for his advice and
encouragement during the course of this work. To Dr. G. Salt and Dr. W. H.
Thorpe I am indebted for many helpful suggestions. Dr. K. G. Blair and
Dr. M. L. R. Pettersson have also given useful advice on special points.
For the gift or loan of material my best thanks are due to Mr. H. S. Barber,
Dr. H. Bollow, Dr. P. J. Darlington, Dr. Fritz van Emden, Dr. W. Horn, Dr. R.
Jeannel, Dr. F. Plaumann, Dr. P. N. Musgrave, Dr. M. W. Sanderson, Dr. H.
Scott and Mr. F. X. Williams.
I am grateful to the Managers of the Balfour Fund (University of Cambridge)
for a grant of £200 and to those of the Worts Fund for a grant of £40 which
enabled me to work on the biology of some of the South American Elmidae, and
to collect properly preserved material for my studies on the internal anatomy.
A Key to the Tribes of the Elmidae.
Adults terrestrial, rarely entering the water. Body never with tracts
of tomentum. Front coxae usually strongly transverse and with the
trochantin exposed. Abdomen frequently with six visible ventral
segments ......... Larini.
Adults aquatic, seldom or never leaving the water. Body with tracts
of tomentum. Front coxae usually round and with the trochantin
concealed. Abdomen never with more than five visible ventral
segments .......... Elmini.
The tribe Ancyronychini (1904, Ganglbauer, Die Kdfer von Mittelenropa, 4
(1) : 108) should be relegated to synonymy with the tribe Elmini. According
to Ganglbauer the Ancyronychini are to be separated from the Elmini by the
very short presternum in front of the anterior coxae with a consequent exposure
of the ventral surface of the head. The length of the presternum in front of
the anterior coxae is not always even a good generic character. Among the
species of Stenelmis may be found many which have the presternum in front of
the anterior coxae extremely long and the head fully retractile, while many, e.g.
S. pallidipes (Cart.), etc., have the presternum very short and the ventral surface
of the head even more exposed than in Ancyronyx Er.
MEXICAN EI.MIDAE
225
A detailed study of the relationships of the Larini to the Elmini has been
made before, and it is only necessary to repeat here that it was found that there
was only one character by means of which these two tribes could be infallibly
separated, i.e. the presence of tomentose tracts in all Elmini and their absence
among the Larini. It has been shown that this division into two tribes is not a
very natural one, as it is supported neither by the structure of the larvae nor by
that of the pupae, and furthermore one genus, Phanoceroides Hinton, is referred to
the Elmini, but is really much more closely related to some of the Larini than
to any known members of the Elmini.
A Key to the World Genera of Larini.
1. Presternum broadly truncate posteriorly or at most feebly rounded ;
mesosternum without a depression for the reception of the prosternal
process. Elytra with two accessory basal striae on each elytron
between sutural and second stria. Africa, Madagascar
Potamodytes Grouv. (1896).
Presternum acute posteriorly ; mesosternum with a depression for the
reception of the prosternal process. Elytra with one accessory stria
or none on each elytron between sutural and second stria . 2.
2. Elytra with one accessory stria on each elytron .... 3.
Elytra without accessory striae ....... 8.
3. Pronotum on each side with a longitudinal carina or sulcus on basal
two-fifths. New Zealand, Australia . . Hydora Broun (1882).
Pronotum without basal carinae or sulci . . . . . -4
4. Pronotum evenly convex. Africa, Madagascar Hydrethus Fairm. (1889).
Pronotum with an apical transverse impression which at middle is on
anterior third and at sides on anterior fifth or sbcth. This impression
is but little developed in some genera, e.g. Potamophilus . . . 5.
5. Hind wing with a radial cross vein (text-fig. 1). Western North
America Lara Lee. (1852).
Hind wing without a radial cross vein ...... 6.
6. Second anal with either branch one or branch two absent ; between
first anal and third branch of second anal (plus first branch of third
anal) with only one vein. East Indies, Africa
Potamophilinus Grouv. (1896).
Second anal with first and second branches present so that between
first anal and third branch of second anal plus first branch of third
anal there are two veins . . . . . . . 7.
7. First branch of third anal joined to second anal before the latter has
given rise to any branches, the point at which these two become
united being a distance greater than the diameter of the anal cell
proximal to the anal cell. Australia . . Stetfwlus ('. Ov Z. (11)29).
First branch of third anal joined to third branch of second anal (i.e.
joined to second anal after the latter has given rise to branches .
text-fig. 2). Europe, East Indies (?) . Potamophilus Germ. (181 1).
8. Pronotum on each side with a longitudinal carina or sulcus on basal
two-fifths 9.
Pronotum without a carina or sulcus on each side . . . .10.
9. Pronotum without transverse impressions ; presternum very broad
between coxae. Japan . . . Dryopomorphus Hinton (1936).
226
H. E. HINTON
10,
Pronotum with a transverse impression on each side at apical third,
this impression extending mesally and posteriorly to join the impres-
sion formed by the longitudinal sulcus. North and South America,
West Indies Phanocerus Sharp (1882).
Pronotum with a strong transverse impression on apical third at middle
and at apical fifth or sixth at sides. Central and South America
Hexanchorus Sharp (1882).
Pronotum without a transverse impression. Central and South
America Disersus Sharp (1882).
l*jt->A/
Text-figs. 1-4. — (1) Lara avara amplipennis Darl. (2) Potamophilus acuminatus (F.).
(3) Wing of other side of same specimen drawn from ventral side to facilitate a com-
parison with the normal type of venation. (4) Potamodytes tuberosus Hinton.
Venation after Forbes.
Potamophilops Grouvelle (1896) would run to Hexanchorus in this key. The
type has been examined in the Paris Museum and appears to be identical with
Hexanchorus. However, at that time the wing venation was not examined,
MEXICAN ELMIDAE 227
and it is just possible that Polamophilops is a valid genus. Betelmis Matsumura
(1916) was referred by its author to the Larini. As nearly as I am able to judge
from the description and figure, this genus belongs in the family Psephenidae,
where it is possibly synonymous with M alaeopsephenus Waterhouse.
The hind wings of the Coleoptera have seldom been used for separating
genera. Apart from the fact that other characters are usually more readily
accessible, in most groups of beetles the hind wings are frequently atrophied ;
and it is this tendency towards reduction which is the most serious objection to
their use as generic characters. In those groups in which reduction of the wings
is known to occur, the genera should never be distinguished solely by some wing
character, e.g. venation, which is known to be affected by reduction, for this
would often result in the difficult position of having some species represented
only by individuals with reduced wings, and these individuals could not be
assigned to their proper genera. But even in those groups in which reduction
of the size of the wings with consequent reduction of the venation is known to
occur, it is possible, and often even necessary, to use the venation as a supple-
mentary generic criterion. In the tribe Elmini, as will be shown in my study of
the Mexican genera, reduction of the wing venation is a frequent occurrence, and
yet a consideration of the venation of those individuals which have fully developed
wings is an aid to a fuller understanding of important generic relationships.
The Larini are a group which is not truly aquatic but is riparian. The adults
are among the most agile of flying beetles, and after studying their habits it
is difficult to imagine atrophy of the wings occurring except at very infrequent
intervals. No individual that has been examined — individuals of most of the
described species have been examined — has shown the slightest tendency towards
reduction of the wings. Other families of beetles possessing strictly riparian
tribes or groups of genera resemble the Larini in that the wings are seldom if
ever reduced. Darlington (1936), who has studied the hind wings of the Cara-
bidae, says (p. 157), "... almost without exception species which live closely
associated with water have strongly developed flying wings " (italics are his).
A Key to the Genera of Mexican Elmini.1
1. Maxillary palp 3-segmented. Mid-gut with prominent regenerative
crypts ; anterior margin without caeca. Europe Elmis Latr. (1798).
Maxillary palp 4-segmented. Mid-gut smooth, without regenerative
crypts projecting from the outer surface ; anterior margin with or
without caeca .......... 2.
2. Base of pronotum feebly rounded in front of scutellum. Claws toothed.
Mexico ........ Tolriolus, gen. n.
Base of pronotum arcuately emarginate in front of scutellum. Claws
without teeth .......... 3-
3. Each elytron with a short accessory stria at base between first and
second striae. Each testis composed of three sperm tubes. North
and South America .... Elsianus Sharp (1882).
Elytra without accessory striae. Each testis composed of one or two
sperm tubes . . . . . . . . . -4
1 In this key I have included two genera not found in Mexico, Limnius and Elmis, since
in the past a number of Mexican species have been referred to one or another of these.
228 H. E. HINTON
4. Epipleura with a longitudinal line of granules which on anterior two-
fifths is half way between dorsal and ventral margins, while on
posterior three-fifths it is close and parallel to ventral margin. Hind
wing with a well-developed anal lobe. Alimentary canal with five
caeca on the anterior margin of the mid-gut. Each testis composed of
only one sperm tube. Each ovary with only two egg tubes. Central
nervous system with only the second and third abdominal ganglia
discrete. Australia, Central and South America
Austrolimnius C. & Z. (1929).
Epipleura without a longitudinal line of granules. Hind wing without
a well-developed anal lobe. Alimentary canal never with five caeca
on anterior margin of mid-gut. Each testis composed of two sperm
tubes. Each ovary with more than two egg tubes. Central nervous
system with a different combination of discrete abdominal ganglia 5.
5. Mentum twice as long as and broader than submentum. Each elytron
with three carinae or longitudinal rows of granules. Alimentary
canal with four caeca on the anterior margin of the mid-gut. Europe,
North America ...... Limnius Er. (1847).
Mentum never distinctly broader and always shorter than submentum.
Each elytron with at most two sublateral carinae. Alimentary
canal never with four caeca on the anterior margin of the mid-gut . 6.
6. Pronotum entirely tomentose. Hypomera entirely tomentose. Pro-
notum without sublateral carinae. Hind wing without a cubito-
anal cross vein. Alimentary canal without caeca on the anterior
margin of the mid-gut. Central nervous system with only the
second abdominal ganglion discrete. Central and South America
Xenelmis Hinton (1936).
Pronotum not tomentose. Hypomera never completely tomentose. Pro-
notum with a sublateral carina on each side. Hind wing with a cubito-
anal cross vein which may be complete or incomplete. Alimentary
canal always with caeca on the anterior margin of the mid-gut . Central
nervous system with at least three discrete abdominal ganglia . . 7.
7. Hypomera never tomentose. Pronotum on apical two-fifths always with
at least a moderately deep transverse impression which may be com-
plete or confined to sides. Alimentary canal with two, three, or no
caeca on anterior margin of mid-gut. Each ovary with six or less
than six egg tubes ......... 8.
Hypomera usually at least partly tomentose. Pronotum occasionally
with transverse impressions but these are never on apical two-fifths.
Alimentary canal with more than three caeca on anterior margin of
mid-gut. Each ovary with more than six egg tubes 9.
8. Pronotum with a moderately deep, occasionally incomplete, transverse
impression, with a median longitudinal discal impression ; on basal
half on each side with an oblique impression. Each elytron with two
sublateral carinae, only very rarely with one. Epipleura only rarely
tomentose. Hind wing without a trace of an anal lobe ; third anal
joining second anal. Alimentary canal with two caeca on the anterior
margin of the mid-gut ; hind gut with six Malpighian tubules. Each
ovary with six egg tubes. Central nervous system with the first three
abdominal ganglia discrete. North and South America
Microcylloepus Hinton (1935).
MEXICAN ELMIDAE 22g
Pronotum always with a very deep and complete transverse impression ;
never with a median longitudinal impression on disk nor with an
oblique impression on basal half. Each elytron always with only one
sublateral carina. Epipleura always tomentose. Hind wing with a
feebly developed anal lobe ; third anal not joining second anal. Ali-
mentary canal with three, two, or no caeca on the anterior margin
of the mid-gut. Hind gut with only four Malpighian tubules. Each
ovary with four egg tubes. Central nervous system with the first
six abdominal ganglia discrete. North and South America, West
Indies ....... Neoelmis Musgr. (1935).
Hypomera with a complete transverse belt of tomentum. Hind gut
with only four Malpighian tubules. Male reproductive system with
the lateral accessory glands not lobed. Each ovary with seven egg
tubes. North and South America . . Hexacylloepus, gen. n.
Hypomera without a complete transverse belt of tomentum. Hind gut
with six Malpighian tubules. Male reproductive system with the
lateral accessory glands lobed. Each ovary with more than seven
egg tubes ........... 10.
Body sub parallel. Hypomera usually without tomentum but occa-
sionally with a very narrow belt on anterior three-fifths adjacent to
sterno-notal suture. Hind wing with the second branch of the third
anal present. Prosternal process moderately narrow. Alimentary
canal with eight caeca on the anterior margin of the mid-gut. Each
ovary with about 18 egg tubes. Spermathecal duct opening into base
of bursa copulatrix. Central nervous system with abdominal
ganglia two to five discrete. North and South America, West
Indies Cylloepas Er. (1847).
Body usually obovate. Hypomera always with a ventral belt of
tomentum which at broadest point is two-fifths as broad as hypo-
mera. Hind wing with the second branch of the third anal absent.
Prosternal process broad. Alimentary canal with six caeca on the
anterior margin of the mid-gut. Each ovary with 11 egg tubes.
Spermathecal duct opening into the apex of the bursa copulatrix.
Central nervous system with abdominal ganglia one to six discrete.
North and South America . . . Heterelmis Sharp (1882).
A Key to the Larvae of the Genera of Elmidae.
Body flattened and usually at least feebly onisciform ... 2.
Body cylindrical or subcylindrical ....... 5.
. Posterior margin of pronotum with a double row of tubercles. EUROPI
Dupo^hilus Muls. (1872).
Posterior margin of pronotum with a single row of tubercles . . 3.
. Propleura not divided into an anterior and a posterior part ; meso-
and metapleura divided into three parts on each side. Brazil
Phanoceroid.es Hinton (1938).
Propleura divided into an anterior and a posterior part on each side ;
meso- and metapleura divided into two parts on each side . . 4.
. Eighth abdominal segment with the pleura bounded by tergo- and
sterno-pleural sutures. North and South America, West Indies
Phanocerus Sharp (1882).
23° H. E. HINTON
Seventh and eighth abdominal segments without discrete pleura and
both segments forming complete sclerotized rings. Europe
Elmis Latr. (1798)
5. Propleura divided on each side into three parts .... 6
Propleura not divided, or if divided then only into two parts on
each side .......... 10
6. Pleura bounded by sutures present on eighth abdominal segment.
Western North America .... Lara Lee. (1852)
Eighth abdominal segment forming a complete sclerotized ring . . 7
7. Pleura bounded by sutures present on seventh abdominal segment . 8
Seventh abdominal segment forming a complete sclerotized ring . . 9
8. Head with one ocellus on each side. Dorsal surface evenly convex.
Europe, North America .... Limnius Er. (1847)
Head with five ocelli on each side. Dorsal surface with numerous
large gibbosities. Europe . . Potamophilus Germ. (1811)
9. Pleura bounded by sutures present on sixth abdominal segment.
Central and South America . . . Hexanchorus Sharp (1882)
Sixth abdominal segment forming a complete sclerotized ring ; fifth
with pleura bounded by sutures. Bolivia . Disersus Sharp (?) (1882)
10. Propleura not divided, meeting on middle line of body so that
sternum is suppressed. Meso- and metapleura with only the sterno-
pleural suture present. North and South America, West Indies
Cylloepus Er. (1847)
Propleura divided into an anterior and a posterior part, and only with the
anterior part meeting on the middle line of the body. Meso- and
metapleura bounded by tergo- and sterno-pleural sutures . .11
11. Meso- and metapleura divided on each side into two parts . . . 12
Meso- and metapleura divided on each side into three parts. North
and South America .... Heterelmis Sharp (1882)
12. Pleura bounded by sutures present only on the first two abdominal
segments ........... 13
Pleura bounded by sutures present on more than the first two abdominal
segments ........... 14
13. Dorsal surface of body evenly convex. Europe . Latelmis Reitt. (1883)
All abdominal segments except ninth with the posterior part trans-
versely and strongly gibbous. Europe . . Riolus Muls. (1872)
14. Pleura bounded by sutures present on the first six abdominal segments.
Europe Esolus Muls. (1872)
Pleura bounded by sutures present on the first seven abdominal
segments ........... 15
15. Dorsal surface with numerous strong gibbosities. Mexico
Tolriolus, gen. n
Dorsal surface except for ninth abdominal segment evenly convex . 16
16. Tubercles of terga of first eight abdominal segments arranged in parallel
rows. North and South America . Microcylloepus Hinton (1935)
Tubercles of terga of first eight abdominal segments not arranged in
parallel rows .......... 17
17. Anterior margin of head on each side with a large and conspicuous
tooth. North and South America . Elsiantts Sharp (1882)
Anterior margin of head on each side without a distinct tooth . . 18
MEXICAN ELMIDAI 231
18. Anterior margin of clypeus on each side with a moderately large acute
tooth. Hind gut with six Malpighian tubules. South America
Macrelmis Mots. (1859).
Anterior margin of clypeus not toothed. Hind gut with only four
Malpighian tubules. Nori h and South America
Neoelmis Musgr. (1935).
In this key I have included all the genera represented in my collection as
well as Dupophilus Muls., which is known to me only by the description of Ber-
trand (1936). The larvae of three other genera, Ancyronyx Er., Macronychus
Mull., and Stenelmis Duf., have been described and figured in the literature at
various times, but in none of these descriptions have the important generic
characters been mentioned, so that it is not possible to deal with them here.
No character has been found which will serve to separate the larvae of the
Larini from those of the Elmini. As may be seen from the key, some members
of the Larini, e.g. Phanocerus, are much more closely related to members of the
Elmini than they are to members of their own tribe. Most Larini are much
larger than the largest Elmini, so that the large size of the mature larvae (usually
more than 7-0 mm. long and 1-5 mm. broad) is a useful character for distinguish-
ing the larvae of the two tribes. This in combination with those forms deter-
mined by elimination and locality and by breeding has enabled me to identify
the larvae of all American genera of Larini with the doubtful exception of Disersus
Sharp.
The Mexican Larini.
Two species of Larini occur in Mexico, Hexanchorus gracilipes Sharp and
Phanocerus clavicornis Sharp. A redescription of these two species with a
description of their immature stages follows.
HEXANCHORUS Sharp
1882. Hexanchorus Sharp, Biol. Centr.-Amer. Col., 1 (2) : 127.
1896. Xexanchorus (err. lyp. ?) Grouvelle, Bull. Soc. ent. Fr., 1896 : 78.
Body elongate, subparallel ; clothed for the most part with dense, moderately
long, usually recumbent hairs. Head when seen from below not capable of being
retracted into prothorax beyond the basal portion of the submentum. Antenna
as figured (text-fig. 8), 11-segmented. Mandibles with one obtuse subapical
and two obtuse apical teeth ; prostheca large and entirely membranous. Maxil-
lary palp (text-fig. 11) 4-segmented and stipes with a well-developed palpifer ;
galea and lacinia separate and each densely spinose. Labial palp 3-segmented
and prementum with a palpiger. Mentum broader than and as long as sub-
mentum. Gula at apex as broad as first segment of antenna. Pronoium with a
complete and deep apical impression which at middle is about on apical third and
at sides on apical fifth ; without longitudinal carinae. Elvira striate and punc-
tate ; without accessory striae and without longitudinal carinae. Hind wing
(text-fig. 12) without a radial cross vein and without an anal cell ; with the
second branch of the second anal absent ; first anal present only apically ;
cubito-anal cross vein complete or incomplete and joining cubitus to first anal.
Prosternum (text-fig. 16) very short in front of anterior coxae ; process long and
acute. Mesosternum with a deep and narrow groove for the reception of the
232
H. E. HINTON
prosternal process, this groove being posteriorly very broad and nearly as broad
as long where its posterior wall is formed by the mesosternum. Metasternum
with a longitudinal impressed line. Abdomen with the sixth ventral segment
externally visible. Legs with the front coxae transverse ; with the externally
visible part of the trochantin large. Claws without teeth. Alimentary canal
with six Malpighian tubules. Male reproductive system with two long and narrow
sperm tubes to each testis. The details of the internal anatomy are based on a
single pupa in poor condition. The number of caeca on the anterior margin
of the mid-gut, if any, could not be determined.
The wing venation is quite unlike that of any other Larini known to me, but
perhaps, in the absence of an anal cell, it is nearest to Phanocerus, a genus with
which it also agrees in having no accessory striae on the elytra and in possessing
two sperm tubes to each testis. The structure of the larvae would place this
genus very far from Phanocerus and near those genera having an anal cell in
the hind wing.
All legs have at the ventral apex of the fourth tarsal segment a fine nearly
erect seta which is about half as long as the fifth segment. This seta occurs
in all species I have been able to examine, e.g. gracilipes Sharp, caraibus Coquer.,
thermarius Coquer., tarsalis Hinton and tibialis Hinton, and is probably a
good generic character. Certainly no other Larini I have seen have a similar
seta.
Genotype : Hexanchorus gracilipes Sharp.
The specific characters of most importance seem to be the following :
(i) General proportions, length and breadth. It is particularly important
to measure the length exactly, as this character is most useful in
associating species with descriptions, there being but little intra-
specific variation in size.
(2) Punctation of all parts of the dorsal surface. I have not considered
the punctation of the ventral surface, for it is generally concealed by
the pubescence and besides there is an abundance of good specific
characters.
(3) Outline of the pronotum and the depth and general proportions of the
impressions on its surface.
(4) Proportion of scutellum and if flat or convex.
(5) Form of elytral apices.
(6) Shape of presternum.
(7) Shape of apical margin of fifth and sixth abdominal segments.
(8) Structure of the abdominal spicule.
(9) Secondary sexual characters. These are among the most highly
specific characters I have been able to find.
(10) Structure of the male genitalia. Here are found the most important
specific characters, and no one should describe a new species without
illustrating this organ.
The following secondary sexual characters have been observed in the species
before me :
(1) Female with the inner apices of the elytra turned up vertically (graci-
lipes).
(2) Front tibia of the male slightly shorter and more curved than that of
the female (gracilipes).
(3) Male with the inner apex of the fifth segment of the front tarsi dilated
(tarsalis) .
MEXICAN ELMIDAE 233
(4) Male with a fine, short, and nearly longitudinal carina on the inner
apex of the middle tibia (gracilipes, caraibus).
(5) Male with a fine tubercle on the inner apex of the middle and hind
tibiae (tarsalis).
(6) Male with a short and obliquely longitudinal tubercle on the inner apex
of the hind tibiae (tibialis).
(7) The disc of the metasternum and first three abdominal sterna is strongly
concave in all males and convex in all females.
(8) Apical margins of the fifth and sixth abdominal sterna always
differently emarginate in the two sexes.
(9) The structure of the abdominal spicule is completely different in the
male and female.
This genus is widely distributed in Central and South America. One species
occurs on Martinique and Guadeloupe.
Hexanchorus gracilipes Sharp.
(Text-figs. 5-20.)
1882. Hexanchorus gracilipes Sharp, Bu>L < enti Imer. Col., 1 (2) : 12S. t. 4, f. 7.
1937. Hexanchorus gracilipes Hinton, Ent. mon. Mag., 73 : 96, 3 figs.
Male : Length, 3-5 mm. -4-0 mm. ; breadth (across elytral humeri), 1-32
mm.-i-55 mm. Elongate, subparallel, moderately convex. Dorsal surface
clothed with fine, dark, suberect to erect, moderately dense hairs which are
about 0'06 mm. long, and also densely pubescent with much finer and shorter
hairs which are recumbent and usually greyish, though these often have a greenish
iridescence. Ventral surface with both types of hairs longer, more like one
another, and distinctly paler ; tibiae of middle legs pubescent only at extreme
base. Cuticle black to rufo-piceous ; two basal segments of antenna, mouth-
parts, trochanter, base of femora, all of tibiae, and claws paler. Head without
distinct impressions. Anterior margin of clypeus truncate, with the angle on
each side broadly rounded. Anterior margin of labrum shallowly and arcuately
emarginate at middle, with the angle on each side broadly rounded. Eyes
purplish in colour, narrowed posteriorly, and protected by an arch of long black
setae which arise from the head near the dorsal and ventral sides of the eyes and
meet above the eye at middle. Antenna as figured (text-fig. 8). Surface with
two types of punctures as follows : coarse punctures (about 0-008 mm. in diameter)
round and confluent to separated by once their diameters ; and fine punctures
about a third as coarse and separated by one to three times their diameters
Pronotum, across broadest point, which is at basal two-fifths, broader than long
(i-io mm. : 0-85 mm.) and base broader than apex (1-07 mm. : 075 mm.). Pro-
notum with the impressions as figured (text-fig. 20) and with the surface punctate
as head. Sharp (loc. cit.) said the pronotum had no punctures, but he can
scarcely have looked at it very closely. Elytra more than three times as long as
pronotum (2-80 mm. : 0-85 mm.), and with the broadest point across humeri
being only slightly broader than broadest point at apical third. Inner side of
apex of each elytron rounded. Lateral margins smooth. Humeri moderatelj
gibbous. Sutural interval on posterior two-thirds feebly raised ; other intervals
flat ; surface of intervals with the punctures no larger than finest of head and
NOVIT. ZOOL., 42, 2. 1940. 20
234
H. E. HINTON
pronotum and separated by two to ten times their diameters. Strial punctures
on mid-discal region round to subquadrate, separated longitudinally by less than
to once their diameters, and usually a third to half as broad as intervals ; basally
on disk they become finer and the intervals broader so that a ratio of one-eighth
as broad here often prevails ; at sides the punctures become slightly finer;
towards apex they become much finer and the striae shallower, so that on apical
sixth striae and punctures have nearly completely disappeared. Scatelhim
Text-figs. 5-1 1. — Hexanchorus gracilipes Sharp. (5) Dorsal view of male genitalia.
(6) Lateral view of right side of male genitalia. (7) Female genitalia. (8) Antenna.
(9) Ventral view of fifth abdominal sternite of male. (10) Ventral view of fifth
abdominal sternite of female. (11) Maxillary palp.
slightly longer than broad (0-17 mm. : 0-15 mm.) and moderately convex, being
distinctly elevated above the adjacent elytral intervals. Prosternum (text-fig. 16)
with the process long. Metasternum concave, the concave portion broadening
out posteriorly so that at posterior margin it embraces most of the disk ; with an
acute but low median longitudinal carina which extends from base forwards for
about 0-12 mm. and from end of this carina a narrow, shallow, impressed line
extends forwards, becoming narrower and shallower anteriorly so that it scarcely
attains the anterior margin of the metasternum. Abdomen with the entire
disk of the first two segments and the anterior half of the third strongly concave ;
MEXICAN ELMIDAE
235
first segment with the discal carina extending to posterior fourth or fifth ; apical
margin of fifth and sixth as figured (text-figs. 9 and 15) ; spicule as figured (text-
fig. 19). Legs with the front tihiae feebly curved inwards near apical third.
Middle tibiae glabrous except for extreme base, and with a fine, short, nearly
longitudinal carina on inner apex. Genitalia as figured (text-figs. 5, 6).
Female : Externally similar to male except as follows : (1) The inner apex
of each elytron is turned upwards at right angles to the general surface ; (2)
front tibiae are slightly less curved than those of male ; (3) middle tibiae without
a carina on inner apex ; (4) disc of metasternum much more feebly and less
extensively concave ; (5) disc of first three ventral abdominal segments convex,
Text-figs. 12, 13. — Hexanchorus gracilipes Sharp. (12) Hind wing. Venation after
Forbes. (13) Microtrichia near inner apical margin of wing.
not concave ; (6) apical margin of fifth abdominal segment as figured (text-fig.
10) ; (7) apical margin of sixth abdominal segment as figured (text-fig. 14) ; (8)
spicule as figured (text-fig. 17).
The eggs are nearly round with a diameter of about 0-30 mm. The surface
is densely reticulate throughout. The pattern near the micropyle is shown in
text-fig. 18.
Type: $ in the collection of the British Museum (Nat. Hist.). Mexico:
Chinantla (Salle).
Specimens examined : Five with data as above. 784, Mexico : Dist. de
Temascaltepec, Temascaltepec, alt. about 5600 ft., vi-vii.1933 (H. E. Hinton,
R. L. Usinger) and vi-vii. [934 (//. E. Hinton) ; 160, in the same district but at
Tejupilco, 3500-4000 ft., vii.1934 (H. E. Hinton); 217, Mexico: Estado de
Morelos, Cuernavaca, alt. 4800 ft., vi. [934 (H. E. Hinton). 1. PANAMA : Bugaba
236
H. E. HINTON
(Champion) and I, Chargres Basin, Upper Pequini Riv. (A. H. Jennings). 2,
Costa Rica : San Jose, La Caja, ii-vii.1932 (H. Schmidt).
Text-figs. 14-10. — Hexanchorus gracilipes Sharp. (14) Ventral view of sixth abdo-
minal sternite of female. (15) Same of male. (16) Presternum. (17) Spicule of
female. (18) Pattern on surface of egg near microphyle. (19) Spicule of male.
20
Text-fig. 20. — Hexanchorus gracilipes Sharp illustrated from the type.
Variations : The variations in the large series before me are mainly confined
to size and slight differences in punctation. In many specimens the greenish
iridescence of the fine hairs on the dorsal surface is not present. The series from
MEXICAN KLMIDAE
237
Cuernavaca is with few exceptions greyer than those from other localities, and
the size appears to be very slightly (0-20 mm.) but constantly greater (this has
not been verified statistically). A few individuals have the third elytral interval
feebly convex in the discal region.
Com partitive notes : The secondary sexual characters plus the structure oi
the male genitalia will readily separate the male from all other described species.
This species is probably nearest to H. tarsalis Hinton, from which the female may
be distinguished at once by the elevated apices of the elytra and the deeper
emargination of the apex of the fifth abdominal sternite.
Biology : They are found in small to large streams (1-30 metres) on the
downstream sides of stones, where they are often present in large numbers above
the water level. The writer has netted over 300 at one time from the side of a
large stone. They appear to be feeding on the algae growing near the water
level on such stones, and with sudden rises in the water level they are occasionally
submerged for a few seconds, but this apparently does not alarm them, for they
remain clinging to the same spot on the stone. When submerged they are com-
pletely surrounded by a film of air.
When the larvae are mature they crawl up a stone and burrow under the
algal or moss matting, often more than three feet above the water level. Under
this matting they construct a small cell in which they pupate. While in the water
the larvae feed on decaying vegetable material, as may be seen from an exami-
nation of their gut contents.
Pupae.
A single specimen was taken by peeling off the moss matting which was
growing above the water level on a large stone in a torrential stream. The pupal
cell was in the earth surrounding the roots of the moss, but the ventral side of
the cell was formed by the surface of the stone. Since only a single pupa was
taken it was allowed to mature for several days, and when it appeared the adult
was about to emerge, it was preserved in spirit and determined beyond question
as that of H. gracilipes by dissecting out the sclerotized parts of the male genitalia.
Description of Pupa of H. gracilipes Sharp.
(Text-figs. 21, 22.)
Male : Length, 4-25 mm. ; breadth (across broadest point which is at base
of prothorax), 1-32 mm. Head completely concealed from above by the pro-
notum. At middle near anterior margin of frons with a feeble, transverse gib-
bosity which is 0-20 mm. long and has on each side an acute tubercle which is
0-075 mm. high. Surface sparsely pubescent with fine, erect, pale hairs which are
0-07 mm. to 0-12 mm. long and are moderately densely distributed on the epi-
cranium, genae and frons, but form a dense transverse row near anterior margin
of frons, clypeus and labium. Antennae extend posteriorly and downwards to
a point exactly opposite posterior margin of front coxae. Pronotum similar in
shape and various impressions to that of the adult : with two anterior and two
posterior long stout spines as figured (text-fig. 21) ; surface clothed with hairs
which are similar in length and density to those of the posterior region of the
head. Mesonotum with a large oval gibbosity representing the scutellum beneath
(text-fig. 21) on middle posterior margin, and from this gibbosity a line carina
extends along middle line to anterior margin. Mesothorax and abdominal
238
H. E. HINTON
segments as figured ; ninth at apex with a median dorsal and two lateral pro-
jections (text-figs. 21 and 22). Surface of these segments throughout pubescent
as pronotum but slightly more finely and sparsely so. Wings with the anterior
pair extending to ventral side, attaining middle of fourth abdominal segment,
and near apex separated from each other by a distance equal to their greatest
breadth ; posterior wings similarly extending to ventral side but contiguous
near apex and attaining middle of fifth abdominal segment. Legs with the front
pair extending to posterior portion of metathorax and tarsi separated by half
their own lengths from each other on the middle line of the body ; second pair
Text-figs. 21, 22. — Hexanchorus gracilipes Sharp. (21) Dorsal view of pupa.
Left side of the end of the abdomen of the same.
(22)
extending to middle of first abdominal segment and with their claws contiguous ;
hind pair with the coxae, trochanters, base of femora, apex of tibiae and base
of tarsi concealed by the hind wings ; apical three segments of tarsi contiguous ;
claws attaining middle of sixth abdominal segment. Abdominal spiracles placed
on dorso-lateral sides and opening at apices of small tubercles.
Specimen examined : Mexico : Dist. de Temascaltepec, Temascaltepec,
alt. about 5600 ft., vii.1934 (H. E. Hinton).
Larvae.
The larva of H. gracilipes has been definitely determined as such, for the
cast skin of the last instar larva was found in the same cell as the pupa. In 1937
three species of larvae which apparently belong to the genus Hexanchorus were
collected in South America. Of these, one can be reasonably determined by
MEXICAN ELMIDAE 239
elimination and locality as that of H. tibialis Hinton. Of the two remaining,
one is from French Guiana, a region from which until now no adults have been
recorded, and the other was collected in Brazil without adults. The latter,
which was collected in localities as far apart as 1000 miles, may be H . thermarius
Coquer. or a new species. No exact locality record is extant for thermarius, but
if it is ever collected in the Amazon basin it may be reasonably associated with
this larva, which seems to be the common and possibly only one here. In
addition to these I have been able to study the larvae of H. tarsalis Hinton
collected by Dr. Fritz Plaumann in the same locality as the adult type. A key
to the five species is given below, and the two undetermined species are included,
for names can be assigned to them with reasonable certainty as soon as the adults
are recorded from the same region. In no knowrn case does the distribution of
one species of Hexanchorus overlap that of another.
A Key to the Larvae of the Species of Hexanchorus
1. Head with one ocellus on each side . . . . . . .2.
Head with more than one ocellus on each side ..... 3.
2. Mesothoracic and abdominal spiracles opening at the apices of very long
and slender spines ; ninth abdominal tergite strongly emarginate at
apex ; tergites of mesothorax and first two abdominal segments
densely and evenly tuberculate. French Guiana (St. Laurent du
Maroni) . . . . . . . . . H . sp. ?
Mesothoracic and abdominal spiracles not opening on the apices of very
long and slender tubercles ; ninth abdominal tergite truncate at
apex ; mesothorax and first two abdominal segments with complete
and transverse belts free of tubercles on anterior and middle part of
each tergite. Brazil (Manaos, Belem) . . . . H. sp. ?
3. Head with six ocelli on each side. Mesothoracic and abdominal spiracles
opening at the apices of large and conspicuous tubercles. Central
America ........ H. gracilipes Sharp.
Head with five ocelli on each side . . . . . . -4
4. Mesothoracic and abdominal spiracles not opening on large tubercles.
Bolivia (Yungas Valley) H. tibialis Hinton.
Mesothoracic and abdominal spiracles opening at the apices of large
and conspicuous tubercles. Brazil (Santa Catharina)
H. tarsalis Hinton.
From a study of these five species a brief generic diagnosis may be made,
since it is now possible to distinguish between those characters that are of
specific and those that are of generic importance.
Generic Characters of Larvae of Hexanchoniv
Body parallel, cylindrical. Head when viewed dorsally exposed and not
concealed by the pronotum ; anterior margin between base of antenna and
clypeus toothed ; with one to six ocelli on each side. Antenna 3-segmented,
not retractile. Mandibles of both sides similar and with three obtuse apical
teeth ; prostheca long, slender, and densely spinose. Maxilla with the palp
4-segmented and stipes showing no differentiation into a palpifei ; galea and
lacinia separate and apex of each densely spinose. Labium with the post-mentum
24O H. E. HINTON
undivided and the submentomental muscles absent ; labial palp 2-segmented
and prementum with a palpiger. Gula well developed. Prothoracic pleura
divided into three parts on each side, while those of the meso- and metathorax
are divided into two parts. Presternum divided into an anterior and a posterior
sclerite ; other thoracic segments with the sternum not divided. First six
abdominal segments with pleura bounded by tergo- and sterno-pleural sutures ;
segments seven and eight forming complete sclerotized rings. Operculum with
two strongly sclerotized claws attached to its dorsal membrane. Apex of ninth
segment rounded to strongly emarginate. Spiracles present on mesothorax and
first eight abdominal segments, and they may open at the level of the cuticle
or on prominent spines or tubercles ; sometimes they are present a short distance
from the median line on each side and again in some species they are found only
on the lateral margins. Tracheae with a number of well-developed air sacs or
without any. Three tufts of retractile, anal, tracheal gills are present. Ali-
mentary canal (text-fig. 25) with the six Malpighian tubules ending freely in the
body cavity near the rectum. Central nervous system with three thoracic and
eight abdominal discrete ganglia.
Description of Mature Larva of H. gracilipes.
(Text-figs. 23-47.)
Length, 5-1 mm. ; breadth (across broadest point, which is at base of meso-
thorax), 1-5 mm. Elongate, subparallel, and cylindrical to subtriangular in
cross section. Cuticle brownish ; with the antennae, mouth-parts and legs
paler brown to testaceous. Head slightly broader than long (0-65 mm. : 0-55
mm.) ; posterior margin broadly emarginate at middle ; coronal suture short ;
frontal sutures beginning on posterior sixth of head and diverging forwards to
meet anterior margin opposite base of antenna on each side ; anterior margin
between base of antenna and clypeus narrowly projecting forwards as far as
the anterior margin of the clypeus. Cuticle sparsely pubescent with fine, erect
hairs which are about 0-12 mm. long ; finely and densely alutaceous throughout ;
anterior five-sixths set with round, convex tubercles which are about 0-04 mm.
broad and are usually separated by less than to one and a half times their diame-
ters ; each tubercle with a coarse hairy seta which is at most slightly longer than
the diameter of the tubercle from which it arises ; dorsally with the basal sixth
and ventrally with the basal fifth free of tubercles. Eyes of five large and one
smaller and slightly ventral ocellus. Antenna (text-fig. 27) not retractile.
Clypeus nearly as long as second segment of antenna ; anterior margin feebly
and broadly emarginate, nearly truncate, with the angle on each side broadly
rounded. Labrum as figured (text-fig. 30) ; epipharynx entirely membranous.
Mandible (text-fig. 28) with three, short, obtuse apical teeth ; outer lateral
side with a small hairy seta ; prostheca (p) long and densely spinose. Maxilla
(text-fig. 29) with the setae as figured. Labium (text-fig. 26) with the setae as
figured. Thoracic and abdominal tergites for the most part sculptured as frontal
region of head but with the tubercles slightly larger and also with two other
types of tubercles as follows : (1) Median and lateral well-defined areas on thoracic
and first six abdominal segments (text-fig. 34) with very dense tubercles (text-
fig. 31) which are about half normal size ; and (2) tubercles about three-fourths
the size of the latter, nearly as dense, and occurring on irregular areas near the
median line of the thoracic and first five abdominal segments. The sculpture of
MEXICAN ELMIDAE
24I
a section on the right side of the fifth abdominal segment is figured (text-fig. 36) ;
posterior margin with a single row of tubercles (text-fig. 39) similar to the pos-
terior row of all segments except ninth abdominal which has no special posterior
row ; lateral margins of thoracic segments with two close and complete parallel
rows of tubercles from which arise long, flat setae (text-fig. 40) ; lateral margin-,
of first eight abdominal segments tuberculate as lateral margins of thorax but
with the dorsal row of tubercles present only posteriorly ; ninth abdominal
segment without a distinct lateral margin and with the sides throughout tuber-
culate as on dorsum. Thoracic and abdominal tergites gibbous and depressed
Text-figs. 23 25 (23) Brain of Hexanchorus sp. > (24) 1 radical air sac ol Hexan-
1 horns sp. ? (25) Alimentary canal of Hexanchorus gracilipes Sharp, (br) brain ;
(fg) frontal ganglion of stomodeai nervous system ; (H) hind gut ; (M) mid-gut ;
(mp) Malpighian tubule ; (O) oesophagus ; (P) pharynx.
as shown in text-tig. 32 ; ninth abdominal segment with the apex rounded (text-
(igs. 32 and 43). Sternum of thorax and first abdominal segment sclerotized as
figured (text-fig. 41). First six abdominal segments with pleura bounded by
tergo- and sterno-pleural sutures ; segments seven and eight forming complete
sclerotized rings, though in some species there is a pale line on these two segments
which is apparently the trace of a sterno-pleural suture. Cuticle tuberculate
similarly to dorsum but with the tubercles slightly finer and without areas of
fine, dense tubercles. Operculum (text-fig. 42) with two toothed claws attached
to its dorsal membrane (text-fig. 47). Legs with the front pair shortest and the
hind pair longest ; posterior legs sculptured similarly to front legs (text-fig. 35).
242
H. E. HINTON
Spiracles opening on the apices of large tubercles which on the mesothorax are
lateral on anterior third of segment and on first seven abdominal segments are
laterally on posterior third, while on the eighth the tubercles are on the apices of
the median gibbosities ; spiracles all similar in structure to that of the last
abdominal segment (text-figs. 45 and 46).
Text-figs. 26-30. — Larva of Hexanchorus gracilipes Sharp. (26) Ventral view of
labium. Setae of apical portion of the prementum are only approximately correct.
(27) Antenna. (28) Ventral view of left mandible. (29) Ventral view of right
maxilla. Apical spines of galea and lacinia only approximately correct. (30)
Dorsal view of labrum with the setae drawn to give the general appearance.
Specimens examined : Mexico : 14, Dist. de Temascaltepec, Real de Arriba,
alt. 6000-7000 ft., Temascaltepec, alt. 5600 ft., and Tejupilco, alt. 3500-4000 ft.,
all in vi-vii.1934 {H. E. Hinton) ; 12, Estado de Morelos, Cuernavaca, alt. 4,
800 ft., vi.1934 (H. E. Hinton).
All the larvae before me are apparently mature.
MKXICAN ELMIDAE
243
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244
H. E. HINTON
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MEXICAN ELMIDAE 245
PHANOCERUS Sharp
1882. Phanocerus Sharp, Biol. Centr.-Amer. Col., 1 (2) : 128.
1896. Phanocerus Grouvelle, Bull. Soc. ent. l-'r. : 78.
Body elongate, subparallel ; clothed for the most part with moderately long
and dense, usually recumbent hairs. Head when seen from below not capable
of being retracted into the prothorax beyond the base of the prementum. An-
tennae as figured (text-fig. 54) with the last six segments forming a compact
club. Mandible with two acute apical teeth ; with a well-developed but entirely
membranous prostheca. Maxillary palp (text-fig. 53) 4-segmented and stipes
with a well-developed palpifer ; galea and lacinia separate and each densely
spinose. Labial palp (text-fig. 52) 3-segmented and prementum with a well-
developed palpiger. Mentum shorter and narrower than submentum ; sub-
mentum (text-fig. 55) strongly concave posteriorly and at middle forming a
slight depression which is partly under the anterior margin of the gula. Pro-
notum (text-fig. 56) with a broad longitudinal impression on each side, which
anteriorly turns outwards to meet the lateral margin at apical half. Elytra
punctate and striate. Without accessory striae and without longitudinal carinae.
Hind wing (text-fig. 57) without an anal cell or radial cross vein ; with the first
branch of the second anal absent ; first anal very short, present only distally
and not connected to cubito-anal cross vein ; cubito-anal cross vein joining
second anal to cubitus. Prosternum moderately short in front of the anterior
coxae ; process long and acute. Mesosternum anteriorly with a narrow groove
for the reception of the prosternal process ; this groove merges posteriorly into a
larger and deeper depression the posterior margin of which is formed by the
metasternum. Metasternum with a complete longitudinal impressed line.
Abdomen with the sixth ventral segment not externally visible. Legs with the
anterior trochantin visible externally only if coxae are rotated caudally. Claws
without teeth. Alimentary canal (text-fig. 58) with eight caeca on the anterior
margin of the mid-gut. Hind gut with six Malpighian tubules which extend
forwards, often as far as the prothorax, and end freely in the body cavity near
the rectum. Male reproductive system (text-fig. 60) with two sperm tubes to each
testis, and with the testes always in the abdomen, seldom being as far forwards
as the first segment ; vas deferens with the proximal ectodermal part swollen
to serve as vesiculae seminalis which are connected to the ejaculatory duct
proximally to the two pairs of accessory glands. Accessory glands with the
median pair dorsal and much shorter than the slightly ventral and lateral pair.
Female reproductive organs (text-fig. 59) with nine egg tubes to each ovary.
Spermathecal duct opening into bursa copulatrix near base. Central nervous
system (text-fig. 61) with the thoracic ganglia not fused and with the first abdominal
ganglion fused to the third thoracic, third to fourth discrete, and five to eight
partly fused together. Stomodeal nervous system with the occipital ganglion
unpaired (text-fig. 61).
Genotype : Phanocerus clavicornis Sharp.
The internal anatomy of two species was studied, a new one from Bolivia
and P. congener Grouvelle from Trinidad. The new species is illustrated. Both
agree in all essential details. The relative size of both reproductive systems,
though particularly that of the male, differs enormously according to the sexual
phase of the individual.
This genus is not close to any other in the I.anni, though this is not so evident
from the structure of the adults, which in many ways is close to llexanchorus,
246
H. E. HINTON
as from that of the larvae. The larvae are very different from all known larvae
of the tribe, but very close to those of Phanoceroides Hinton (Elmini).
The species are widely distributed in America, including the West Indies.
The most northern record is that of Phanocerus clavicomis found in Texas.
1882.
1911.
1937-
Phanocerus clavicomis Sharp.
(Text-figs. 48-61.)
Phanocerus clavicomis Sharp, Biol. Cenlr.-Amer. Col., 1 (2) : 129, t. 4, f. 8.
Phanocerus clavicomis Schaeffer, /. N.Y. ent. Soc, 19 : 118.
Phanocerus clavicomis Hinton, Ent. man. Mag., 73 : 95.
Male: Length, 2-0 mm.-j'O mm. ; breadth (across humeri which is the
broadest point), 0-85 mm.-i-i mm. Elongate, subparallel, moderately convex.
Text-figs. 48-55. — Phanocerus clavicomis Sharp.
view of sixth abdominal sternite of male. (50)
Right lateral view of same. (52) Labial palp.
(55) Submentum (sm) and gula (g).
(48) Spicule of male. (49) Ventral
Dorsal view of male genitalia. (51)
(53) Maxillary palp. (54) Antenna.
Dorsal surface clothed with partly erect brownish hairs which are about 0-05 mm.
long and are usually separated from each other by about a third of their lengths ;
also more densely clothed with hairs which are finer, about a third as long, and
frequently in certain lights golden and shining. Ventral surface for the most
part with only the longer hairs which are here recumbent and frequently so dense
that the cuticle beneath cannot be seen. Cuticle shining and rufo-piceous to
pale brown ; mouth-parts, basal segments of antennae, and legs paler. Head
without distinct impressions. Eyes narrowed posteriorly ; margins around eyes
MEXICAN ELMIDAF.
247
laterally and posteriorly with very long, stout, black setae which curve across
the eye and generally meet about middle over eye. Antenna as figured (text-fig.
54). Surface with punctures which are usually round, about o-oio mm. broad,
and are contiguous to separated by once their diameters. Clypeus with the
anterior margin truncate and the angle on each side broadly rounded. Labium
with the anterior margin feebly, broadly, arcuately emarginate, and with the
angle on each side broadly rounded ; anterior margin except at middle with a
dense fringe of long testaceous hairs and on each side just behind angle with a
tuft of longer and denser hairs. Gula and submentum as figured (text-fig. 55) ;
mentum with the anterior margin truncate or at most very feebly rounded.
Pronotum across broadest point, which is at basal two-fifths, broader than long
(0-92 mm. : 0-77 mm.) and base broader than apex (0-87 mm. : o-6o mm.). Pro-
notum as figured (text-fig. 56) ; surface similarly but more sparsely punctate
56
Text-fig. 56. — Phanocerus clavicornis Sharp.
than head. Elytra between humeri and apical two-fifths nearly parallel. Humeri
feebly gibbous. Apices conjointly, feebly rounded but the apex of each elytron
is itself feebly rounded. Lateral margins smooth. Intervals flat and surface
of each punctate as pronotal disk. Striae feebly impressed but narrower and
slightly deeper towards apex ; strial punctures on disc round to subquadrate,
separated longitudinally by about their diameters, and from a half to a third
as broad as intervals ; towards sides these punctures become a little finer, but
towards apex much finer so that on apical fourth they are seldom as much as
one-third as broad as an adjacent interval. Scutellum Hat, broader than long
(0-12 mm. : o-io mm), with the base broadly arcuate and posteriorly coming
to a rounded point. Presternum moderately short in front of anterior coxae.
Metastemum with an impressed median longitudinal line which suddenly broadens
out and becomes deeper near anterior margin ; anterior margin on each side
projecting slightly, but distinctly over depression formed by the median line.
Abdomen with the apical margin ot the fifth sternite broadly rounded. Sixth
sternite as figured (text-fig. 4q). Spicule as figured (text-fig. 48). Genitalia as
figured (text-tigs. 50, 51).
Female: Externally similar to male except as follows: (1) The posterior
248 H. E. HINTON
projection of the concealed sixth abdominal sternite is much longer ; (2) the
spicule of the male has no homologous sclerotized part in the female. Eggs are
oval to nearly round, and the surface is marked with a close network of cells
which are usually round.
Type: In the British Museum (Nat. Hist.). Guatemala : Vera Paz, Corban
(Champion).
Specimens examined: 6, with same data as above, 2, Mexico: Dist. de
Temascaltepec, Real de Arriba, alt. 6000-7000 ft., vii.1933 (H. E. Hinton, R. L.
Usinger) ; 2, Mexico: Estado de Morelos, Cuernavaca, vi.1934 (H. E. Hinton).
160, Brazil : Santa Catharina, Nova Teutonia, 1934-1935 (F. Plaumann).
Other records : This species has been recorded from Texas by Schaeffer
(1911).
I 'aruitions : The absolute length and breadth in the series before me vary
very much. The elytral strial punctures in some rare individuals may be only
two-thirds as large and dense as in the specimen described.
Text-fig. 57. — Wing of Phanocerus clavicomis Sharp. Venation after Forbes.
Comparative notes : This is the only genus known to me in the Larini in which
there are apparently no differences in the structure of the male genitalia of the
different species. My figures of the male genitalia of P. clavicomis will apply
equally well to seven species before me.
There have been two other species described from continental America, one
from Columbia and the other from Argentina, but both are known to me only
from descriptions which are so inadequate that I am unable to say in what
manner, if any, they differ from P. clavicomis.
Biology : This species is generally found feeding on algae growing above the
water level on stones and sticks either in the stream or lying along the margin.
It may enter the water to oviposit, but is not generally found below the water
level as is the allied Phanoceroides.
Larvae.
By elimination and according to locality the larvae of several species of
Phanocerus have been associated with their adults. Although no evidence as
conclusive as breeding is available, I regard the generic determination of these
larvae as fairly certain for the following reasons : (1) the distribution of the
larvae exactly coincides with that of the adults over North and South America ;
(2) in Central Mexico, the Yungas Valley of Bolivia, and Trinidad, B.W.I., where
MEXICAN ELM I DAE
24.1
I have madr very large collections of the Elmidae, the larvae of all other genera
(except in Mexico where the larvae of Xenelmis and Austrolimnius are still
unknown) have been determined (those of Xenelmis and Austrolimnius must be
too small to be confused with Phanocerus) ; (3) the mature larva is of a size
to produce an adult equal in size to Phanocerus ; and (4) in the streams about
Manaos, Brazil, Phanocerus seems to be replaced by the closely allied genus
Phanoceroid.es, and the larvae of the latter are very nearly related to the larvae
I consider to belong to Phanocerus.
Text-figs. 58-61. — Phanocerus clavicornis Sharp. (58) Alimentary canal. (59)
Female reproductive system. (i>o) Male reproductive system. (61) Central and
stomotleal nervous systems.
From a study of the larvae before mi
generic characterization of Phanocerus.
it lias been possible to draw up a brief
Generic Characters of Larvae of Phanocerus.
Flattened, subparallel. Head when viewed dorsally exposed and not concealed
from above by the pronotum. With one ocellus on each side. Antenna 3-
segmented, not retractile. Clypeus completely fused with the Irons. Mandibles
of both sides similar and with three subacute apical teeth ; prostheca long,
slender and densely hairy. Maxilla with the palp 4-segmented and stipes showing
no differentiation into a palpifer ; galea and lacinia separate and apex of each
densely spinose. Labium with the postmentum undivided ; palp 2-segmented,
prementum without a palpiger, Prothoracic pleura divided into two parts and
with the anterior part meeting on the middle line. Meso- and metathoracic pleura
novit. zool., 42, 2. 11140. 21
250
H. E. HINTON
divided into two parts on each side. First eight abdominal segments with pleura
bounded by tergo- and stemo-pleural sutures. Operculum with two stronglv
sclerotized claws attached to its dorsal membrane. Apex of ninth segment rounded
or strongly emarginate. Spiracles annular and biforous ; present on mesothorax
a-'""
63
64
68
Text-figs. 62-68. — Larva of Phanocerns clavicornis Sharp. (6a) Alimentary canal.
(63) Oesophageal ring of tubercles and spines. (64) Portion of oesophagus anterior
to ring to show shape and distribution of spines. (65) Left lateral view of brain.
(66) Tracheal air sac. (67) Oseophageal sclerite. (68) Dorsal view of central and
stomodeal nervous systems.
and first eight abdominal segments and opening at the cuticle level or on the
apices of small tubercles. Tracheae with a number of well-developed air sacs.
Three tufts of retractile, anal, tracheal gills are present. Alimentary canal with
a dorsal oesophageal sclerite and anterior to this with a complete ring of spines
and tubercles. With six Malpighian tubules which end freely in the body cavity
generally near the rectum. Central nervous system (text-fig. 68) with three
thoracic and eight abdominal discrete ganglia.
MEXICAN ELMIDAE
251
Description of Mature Larva of P. clavicornis.
(Text-figs. 62 83 i
Length, 5-0 mm. ; breadth (across broadest point which is near base of
metathorax), 1-5 mm. Cuticle brownish-testaceous to brown. Head rectangular,
longer than broad (0-50 mm. : 0-42 mm.) ; coronal suture straight and 022 mm.
long : frontal suture extending nearly in a straight line on each side to margin of
head opposite base of antenna. Cuticle on basal belt nearly as long as epicranial
Text-figs. 69—72. — Larva of Phanocerus clavicornis Sharp. (69) Ventral view of
labium. Setae of prementum are only approximate. (70) Ventral view of left
mandible. (71) Dorsal view of ri^lit antenna. (72) Ventral view of left maxilla.
Spines on apex of galea are only approximately correct.
(coronal) suture not punctate, elsewhere with round punctures which are about
0-013 mnl- lir«>;ul and are separated usually by less than to once their diameters ;
from these punctures arise fine setae which are only slightly longer than the
diameters of their respective punctures. With a single ocellus on each side.
Antenna as figured (text -fig. 71). Clypeus fused to the front oi the head and
fronto-elypeal suture not visible even when the head i- cleared and stained.
Labium transverse (0-16 mm. : o-io mm.) ; anterior margin feebly arcuate and
with the angle on each side broadly rounded ; epipharynx smooth. Mandible
(text-fig. 70) with three subacute apical teeth ; outer lateral side apparently
without setae ; with a long, narrow, slender, and hairy prostheca. Labium with
the setae as figured (text-fig. 69). Maxilla with the setae as figured (text-fig. 72).
I Iter, ia< and abdominal terga with proportions as shown in text-fig. 7.;. Surfai e
throughout punctate as frontal region of he. id but with the punctures slightly
25^
H. E. HINTON
larger and sparser ; anterior part of each segment except pronotum with a
transverse row of acute tubercles and large flat setae (text-figs. 79 and Si) ; a belt
anterior to this is always finely and transversely rugose (text-figs. 79 and 82) ; at
middle each row is interrupted by a space of about 0-07 mm. to o-io mm., and this
row is also interrupted laterally on meso- and metathorax (text-fig. 73) ; at the
posterior lateral apex of each segment is a very long and narrow seta (text-fig. 83) ;
lateral ventral margins of each segment have a single row of seta (text-fig. 80).
The posterior margin of each segment except ninth has a complete row of large,
Text-figs. 73-76. — -Larva of Phanocerus clavicornis Sharp. (73) Dorsal view of larva
to show general shape. (74) Ventral view of pro- and mesothorax to show sclerites.
(75) Ventral view of first two abdominal segments to show sclerotization. (76)
Operculum.
flat, partly overlapping setae (text-fig. 79). Sternum of pro- and mesothorax
sclerotized as figured (text-fig. 74) ; sternum of metathorax similar to that of
mesothorax. First two abdominal sterna as figured (text-fig. 75) ; sterna of
abdominal segments three to eight sclerotized similarly to that of second ; ninth
sternum with an apical operculum (text-fig. 76) which has attached to its dorsal
membrane two well sclerotized claws. Surface of sterna of thoracic and abdominal
segments finely and sparsely setose, but anterior third of abdominal pleura with
a transverse row of larger and denser setae. Legs all fairly close in size and
chaetotaxy to front leg (text-fig. 77). Spiracles opening laterally on mesothorax
and first eight abdominal segments ; all similar to that of fifth abdominal segment
(text-fig. 78) ; with the cuticle around each of the spiracles slightly elevated but
not forming well-developed tubercles. On each side of the gut there are seven
tracheal air sacs (text-fig. 66) which occupy a position from first to sixth abdominal
MEXICAN ELMIDAE
253
segments. Alimentary canal as figured (text-fig. 62). Central nervous system as
figured (text-fig. 68).
Immature stages: Before me arc larvae representing at least two earlier
instars, but these apparently differ only in size- from the mature larvae.
Mllfci •
79
82
tf
1 [t-figs. 77 83. -Larva of Phanocerus clavicornis Sharp. (77) Vnterior face of right
front leg. (78) Fifth abdominal spiracle. (79) Third abdominal tergiteon right side
oi middle line. (80) A seta of the ventral and lateral marginal row, (81) Flat
eta oi anterior row of third tergite (82) Section ol anterior rugose belt of third
tergite. (83) Seta of posterioi lati ral apex of third tergite.
Biology : The larvae are found in small to large streams (about 1 30 metres)
oi last and Clearwater. They seem to prefer the -witter parts ol the streams.
Examination oi the gut contents of a number of specimens seems to -how that
they are living on algae (diatoms, etc.).
Specimens examined: 50, Mexico: Estado de Morelos, Cuernavaca, alt.
4800 ft., vi.1934 (//. E. Hinton); ,',2, Mexico: Dist. de temascaltepec, alt.
254
H. E. HINTON
3500-7000 ft., vi-vii.1933 (H. E. Hinton, R. L. Usinger) and vi-vii.1934 (H. E.
Hinton) ; 7, Guatemala : Escuintla, El Salto, 1100 It., 1934 (F. X. Williams).
The Mexican Elmini.
TOLRIOLUS, gen. n.
Body subparallel. Non-tomentose anas clothed with sparse and short,
recumbent hairs ; fine scale-like or hairy tomentum confined to the following
areas : (1) genae ; (2) epipleura ; (3) hypomera ; (4) sides of presternum,
mesosternum, metasternum, and abdominal sternites ; and (5) most of femora.
Head when seen from below capable of being retracted so that none of the mouth-
parts is visible. Antennae n-segmented. Mandibles with three blunt apical
teeth ; prostheca entirely membranous and with numerous fine spines or hairs
apically. Maxilla with the palp 4-segmented (text-fig. 87) and stipes with a
well-developed palpifer ; galea and lacinia separate and apex of each spinose.
Labium with the palp 3-segmented and prementum without a distinct palpiger.
Mentum transverse and as broad as, but only half as long as, submentum. Gula
at anterior margin three-fifths as broad as submentum, with the sides converging
so that at posterior margin it is only half as broad as submentum, and about a
fifth longer. Pronotum with the anterior margin moderately arcuate at middle
and on each side behind eye before apical angle deeply and broadly sinuate ; base
broadly and moderately deeply sinuate on each side and feebly rounded in front
of scutellum. Pronotum with a sublateral carina on each side extending from
near base nearly to anterior margin ; disk without distinct impressions. Elytra
punctate and striate ; each elytron with a single prominent sublateral carina on
eighth interval. Hind wing (text-fig. 93) without a radial cross vein or an anal
cell ; first anal absent ; second anal with the second branch absent ; third anal with
the second branch absent ; fourth anal well-developed ; and cubito-anal cross vein
complete and joining cubitus to second anal. Prostermtm very long in front of
anterior coxae ; process as figured (text-fig. 91). Mesosternum with a broad and
moderately deep groove for the reception of the prosternal process. Metasternum
with a median longitudinal impressed line. Legs with the visible portion of the
front coxae rounded and the trochantin completely concealed by the hypomera
and sternum. Claws each with a large and acute sub-basal tooth (text-fig. 90).
Genotype : Limnius ungulatus Hinton.
No properly preserved specimens have been available for a study of the
internal anatomy.
T. ungulatus has recently (Hinton, 1936) been referred to the genus Macrony-
choides Champion (1923), but though Tolriolus is most closely related to this genus,
it may be distinguished as follows : (1) the genae are tomentose ; (2) the hypo-
mera are tomentose ; (3) the pronotum has a distinct sublateral carinae, whereas
in Macronychoides there is no trace of carinae here ; (4) each elytron has a
sublateral carina on eighth interval, whereas in Macronychoides the lateral
intervals of each elytron are flat ; (5) the first anal of the hind wing is absent,
whereas in Macronychoides it is present ; (6) the presternum in front of the
anterior coxae is long (equal to about one and a half times the greatest breadth of
the front coxae), whereas in Macronychoides it is here short (equal to about a
third of the greatest breadth of the front coxae) ; and (7) the mesosternum has
the groove for the reception of the prosternal process broad, while in Macrony-
choides it is narrow.
MEXICAN KI.MIDAK
255
Tolriolus ungulatus (Hinton).
(Text-figs. 84-93.)
iui4. I.imnius ungulatus Hinton. Her. lint., Rio de /., 4 (2.) : 200.
1936. Macronychoidis ungulatus (Hinton), Trans. K. Ent. Soc. loud., 85 (18) : 428, figs. 27-
29 ; pi. 1. fig. 4.
Male: Length, 2-12 mm. -2-37 mm. ; breadth, 0-95 mm. -1-12 mm. Cuticle
shining and black ; basal three to five segments of antennae, mouth-parts, and
legs rufo-piceous. Head without distinct impressions ; surface with round, low,
often indistinct granules which are slightly finer than facets of eyes or about
90
Text-figs. 84-91. — Tolriolus ungulatus Hinton. (84) Dorsal view of male genitalia.
(85) Left lateral view of same. (86) Antenna. (87) Maxillary palp. (So) Female
genitalia. (90) Tarsus of hind leg. (91) Presternum.
0-012 mm. broad and are separated by two to tour times their diameters ; surface
between granules sparsely to densely microscopically alutaceous. Clypeus with
the fronto-clypeal suture straight and indistinct ; anterior margin broadly,
arcuatelv and deeply emarginate and with the angle on each side obtusely rounded ;
surface sculptured as head. Labrum with the anterior margin nearly truncate
and the angle on each side broadly rounded ; surface alutaceous as clypeus,
without granules, at middle on anterior third with a few punctures which are
about as coarse as granules oi head, and on each side with a few fine, recumbent,
testaceous setae which are about 0-04 mm. long. I'ronotum across broadest
poinl , which is at basal two-fifths, slightly broader than long (072 mm. : 0-70 mm.)
and base broader than apex lo-o; mm. : 0-45 mm.). Sublateral carinae and
impressions as figured (text-tig. 92). Surface with the apical third between
sublateral carinae with fine (about o-io mm. broad), irregularly shaped, obscure
punctures which are usually separated bv two to four times their diameters and
256
H. E. HINTON
surface between these punctures only sparsely alutaceous ; surface on basal
two-thirds of disk with low, round granules which are about as coarse or slightly
coarser than facets of eyes and are separated by one to four times their diameters ;
surface between these punctures for the most part contiguously to confluently
punctate with very fine punctures (about 0-007 mm. broad) ; surface between
92
Text-figs. 92, 93. — Tolriolus ungulatus Hinton. (92) Adult to show general appear-
ance. (93) Hind wing. Venation after Forbes.
lateral margins and sublateral carinae sculptured as basal two-thirds of disk but
with the punctures slightly finer and sparser. Lateral margins nearly smooth.
Elytra slightly more than twice as long as pronotum (1-50 mm. : 0-70 mm.) and
at broadest point, which is across apical two-fifths, broader than broadest
point across humeri (1-07 mm. : 0-97 mm ). Lateral margins nearly smooth.
Apices feebly and broadly produced and conjointly rounded. Surface with
the striae feebly impressed on sides and apex but not impressed on disk ; discal
strial punctures round to subquadrate, about a third to a half as broad as
MEXICAN ELMIDAE 257
intervals, and separated longitudinally by once or, more rarely, twice their
diameters ; these punctures become finer towards apex and coarser, denser
and deeper towards sides. Intervals flat and with the surface moderately densely
to sparsely alutaceous, and with a tew obscure granules at base and sides which
are similar in size to those of sides of pronotum. ScuteUum flat, subparallel,
longer than broad (o-io mm. : 0-075 mm.), base very feebly rounded, and apex
broadly rounded ; surface sculptured as adjacent parts of elytra. Presternum
with the carinae short and present only on basal third (text-fig. 91) ; when
viewed laterally the anterior three-fourths (not including process) is gradually
but moderately strongly bent ventrally ; process shaped as figured (text-tig. <)i) ;
surface of base of presternum and all of that of process sculptured like thai oi
head except that surface of process is also feebly rugose ; surface of presternum
elsewhere, and also that of hypomera, with the granules similar in size to thus,- of
sides of pronotum but feebly oblong and with the surface between granules
alutaceous like that of head. Mesosternum with the groove for the reception of
the prosternal process broad and long, posterior two-fifths deeper and about a
third broader, and extending to posterior eighth of mesosternum ; surface
on each side of groove sculptured as head but with the granules not distinct.
Metasternum with the median longitudinal impression extending to anterior
sixth, basally as broad as scutellum and half as deep as broad, and apically
shallower and a third to a fourth as broad ; disk feebly depressed posteriorly
and on each side in front of posterior coxae with a deep, nearly oval impression
which is about 0-05 mm. broad ; carina on each side of disk extending posteriorly
and slightly outwards half way to hind coxa ; surface of disk with round and
shallow punctures which are about 0-02 mm. broad and are usually separated by
twice their diameters ; surface between punctures for the most part only very
sparsely microscopically alutaceous ; surface of sides of metasternum sculptured
somewhat similarly to basal portion of pronotal disk but with the granules
distinctly oblong instead of round. Abdomen with the carina on each side of the
middle of the first sternite extending posteriorly and outwards to posterior fourth
of segment ; surface of sternites at sides sculptured as sides of metasternum but
with the granules slightly finer and sparser ; middle region of basal four sternites.
and to a slight extent that of fifth sternite, with the granules sparser than at
sides, rounded, and the surface between the granules often smooth and polished
Genitalia as figured (text-figs. 84, 85).
Female: Externally similar to male.
Type: In the U.S. National Museum. Mexico: Dist.de remascaltepec,
Real de Arriba, alt. 6000-7 » It., v-vii.1933 (H . H. Hinton, R. /.. Usinger).
Specimens examined ; 154, with same data as type ; 1, collected in the same
district at Tejupilco, alt. about 4000 ft., ii-vi.1933 [H. E. Hinton, R. L. Usinger) :
433. Real de Arriba, v vii.1934 (H. E. Hinton) ; (62, in the same district, but
at Rio Verde, alt. about 8000 ft., 14. vi. 1934 (H. E. Hmton).
1 ariations : In this long series there appears to be very little variation, the
most notable variation being in the length oi the metasternal discal carinae which
in some specimens are confined to .interior fourth, while in some they extend
three-fifths of the way to hind coxae.
Larvae.
The larvae oi this genus were determined by elimination and according to
locality. A briel generic diagnosis follows.
258 H. E. HINTON
Generic Characters of Larvae of Tolriolus.
Body subparallel and cylindrical to subtriangular in cross section ; dorsal
surface with numerous gibbosities. Head when seen from above exposed and
not concealed by pronotum ; anterior margin toothed on each side between base
of antenna and clypeus. Clypeal suture distinct. With one ocellus on each side.
Antennae 3-segmented (text-fig. 95) and feebly retractile. Mandibles of both sides
similar and with three obtuse apical teeth ; prostheca long, slender, and densely
spinose. Maxilla (text-fig. 102) with the palp 4-segmented and stipes showing no
differentiation into a palpifer ; galea and lacinia separate and apex of each densely
spinose. Labium (text-fig. 101) with the postmentum undivided ; labial palp
2-segmented and prementum without a distinct palpiger. Gula well-developed.
Prothoracic pleura divided into two parts and anterior part meeting on middle
line of body so that the sternum is here completely suppressed. Meso- and
metapleura divided into two parts. Abdominal segments one to seven with the
pleura bounded bv tergo- and sternopleural sutures and these two sutures
converge and meet at apex of seventh segment ; segment eight forming a complete
sclerotized ring. Operculum with two strongly sclerotized claws attached to its
dorsal membrane. Apex of ninth segment deeply and acutely emarginate.
Spiracles present on mesothorax and first eight abdominal segments and opening
on the apices of small tubercles. Tracheae without air sacs. Alimentary canal
with an oesophageal sclerite on the dorsal posterior margin of the oesophagus.
Hind gut with six ? (my material is not sufficiently well preserved to be sure of
this number) Malpighian tubules. Central nervous system with three thoracic and
eight abdominal discrete ganglia.
The larvae of this genus are close to Elsianus, but may at once be distinguished
by the gibbosities on the dorsal surface. All species of Elsianus (about 15 have
been examined) have the dorsal surface evenly convex.
Description of Mature Larva of T. ungulatus.
(Text-figs. 94-104.)
1936. Macronychoides ungulatus (Hinton). Trans. R. lint. Soc. Lond., 85 (18) : 420. figs.
30-40.
Length 4 mm. (text-fig. 94). Body subtriangular to nearly cylindrical in cross
section. Pro-, meso-, and metanotum subequal in breadth; abdominal segments
feebly narrowed caudally. More heavily sclerotized parts of body fusco-testaceous ;
antennae, mouth-parts, legs, opercular claws, and inter-segmental membranes
pale testaceous. Dorsal and ventral sclerotized areas mostly tuberculate, each
tubercle (except fine tubercles at anterior margin of segments) with a short seta
near apex on posterior side (text-fig. 103 a) ; also with a few sparsely scattered,
extremely fine setae arising independently of tubercles. Head capsule abruptly
rounded and truncate behind, not constricted to form a neck. Anterior four-
fifths of dorsal surface and sides slightly coarsely tuberculate, with the tubercles
separated mostly bv about one to three times their diameters ; surface also with
sparse minute pale areas from which arise setae, on basal one-fifth with three
similar but much larger areas on each side and from each arises a single seta.
Ventral surface with only the area between the ventral sutures tuberculate ;
elsewhere not tuberculate. Frontal sutures well-developed and extending on
each side from near the base of the antenna to unite at middle just before posterior
VIEXK AX ELMIDAI
259
margin. Single' ocellus on each side in a large pigment spot. Labium transverse :
anterior margin feebly rounded, and anterior angles broadly rounded. Antennae a -
figured (text-fig. 95). Mandibles as figured (text-fig. 97), and maxilla and labium as
figured (text-figs. [01, ii)2). Pronotum about a third broader than long; narrowed
anteriorly ; angles rounded ; with two prominent longitudinal ridges parallel
near base and from apical half diverging moderately towards apex ; near base on
Text-figs. 94 too. — Larva of Tolriolus ungulatus Hint. en. (94) Mature larva. (95)
Antenna. (96) Inner view of tip of mandible. (97) Mandible. (98) Left inner view
el middle leg. (99) Ventral view of operculum, (too) Dorsal view ..1 opercular
1 tin
each side with a rounded and elevated area. Surface slightly coarsely tuber-
culate with tubercles which are mostly separated by once to twice their diameters ;
anteriorly with the tubercles slightly liner and denser ; anterioi and posterior
margins with a row of flat quadrate tubercles (text-fig. 1036) from each of which
arises a multi-branched seta. Mesonotum more than twice as broad as lung (0-875
mm. : 0-375 mm.) ; similar to pronotum but with the longitudinal ridges joining in
front and not attaining apex ; with the hind angles feebly produced ; ant. 1101
belt ol very fine tubercles with the tubercles separated mostly by less than their
diameters. Metanotum generally similar to mesonotum. Dorsal abdominal
segments similar to metanotum but more strongly transverse and gradual!)
narrowing caudally : caudal segments with tin' hind angles increasingly strong!}
26o
H. E. HINTON
produced posteriorly. Eighth abdominal segment with the channel between the
dorsal ridges very feeble. Ninth dorsal three-fifths as broad as long (0-375 mm.
0-625 mm.), triangular, subcorneal, with a single median longitudinal ridge, and
with the caudal end deeply and triangularly emarginate. Pleural areas mostly
finely tuberculate as anterior margins of dorsal segments ; posterior margins of
these also with a row of flat and subquadrate tubercles. Ventral surface tuber-
culate as dorsal ; with the anterior margin of the presternum and postrrior
104
Text-figs. 101-104. — Larva of Tolriolus ungulatus Hinton. (101) Ventral view of
labium. (102) Ventral view of right maxilla. (1030.) Dorsal tubercle. (103&)
Tubercle of posterior marginal row of mesothorax. (104) Mesothoracic spiracle.
margin of all other sterna except the ninth with a row of flat and subquadrate
tubercles ; metasternum with two small median longitudinal and carina-like
tubercles at anterior margin ; first abdominal sternite with a single similar
tubercle at anterior margin. Operculum as figured (text-fig. 99) ; opercular
claws (text-fig. 100) very slender and strongly curved. Legs all similar to that
of mesothorax (text-fig. 98) but increasing in length posteriorly. Spiracles all
similar to that of mesothorax (text-fig. 104).
Specimens examined : 20, Mexico : Dist. de Temascaltepec, Real de Arriba
and Rio Verde, alt. 6000-8000 ft., vi-vii.1934 (H. E. Hinton).
The largest available larva is 5-0 mm. and the smallest 2-0 mm. long, but
MEXICAN ELMIDAE
z6i
these seem to differ only in size. All specimens were taken in company with the
adults clinging to roots of Salix and Alnus in cold and torrential mountain streams.
ELSIANUS Sharp.
1882. Elsianus Sharp, Hi<>l. Centr.-Amer. Col., 1 (2) : 131.
At the time of writing iS species have been described, and the genus is now-
known to occur from Texas to South Brazil. This genus will probably take
Text-figs. 105-108. — Elsianus tricolor Hinton. (105) Central nervous system (rum a
dorsal view. (io<>) Alimentary canal. (107) Male reproductive system. (108
Female reproductive system.
51 1 "ml place to Cylloepus Erichson in number of species. Though to-day only 18
species have been described there are over 20 new species in my collection from
South America. A redescription of the genus follows
Body elongate, subparallel. Dorsal surface clothed with short, sparse 01
dense, usually recumbent hairs. Tomentum confined to tin- following areas : (1)
genae : (2) epipleurae : (3) hypomera ; (4) sides of presternum, mesosternum,
and abdominal sternites, but in many species nearly the entire ventral surface is
clothed with scale-like or hairy tomentum ; and (4) all of legs except tarsi, tie >Ugh
in many species the scale-like tomentum is sparse or absent on the tibiae. Head
when seen from below capable of being retracted so that none of the mouth-parts
are visible. Antennae n-segmented. Mandibles with three apical acute teeth :
prostheca large and entire!} membranous with the apex spinose or hairy. Maxilla
262
H. E. HINTON
with the palp 4-segmented and stipes with a well-developed palpifer ; galea and
lacinia separate and apex of each densely spinose. Labium with the palp 3-
segmented and prementum with a well-developed palpiger. Mentum transverse
and about as broad as and about half again as long as submentum. Gula as long
as mentum, anteriorly more narrow or as broad as submentum, and always
narrowed posteriorly. Pronotum with the anterior margin moderately arcuate
at middle and on each side behind eye before apical angle deeply sinuate. Base
trisinuate, broadly and moderately deeply so on each side and more narrowly
and shallowly so in front of scutellum. With or without a sublateral carina ; if a
carina is present it may be complete or evident only near base ; in front of scutellum
sometimes with a gibbosity ; disk with or without a median longitudinal
Text-figs. 109, no.
—Ehiauus striatus Sharp. (109) Hind wing. Venation after Forbes,
(no) Microtrichia from inner apical margin.
impression. Elvira striate and punctate ; with an accessory stria on each
elytron on basal sixth or fifth between sutural and second striae ; without
sublateral carinae. Hind wings without a radial cross vein and without an anal
cell ; with the first branch of the second anal absent ; and with a cubito-anal
cross vein. Prosternum very long in front of anterior coxae ; prosternal process
long and posterior margin broadly rounded or acute. Mesosternum with a
broad and deep groove for the reception of the prosternal process. Metasternum
with a median longitudinal impressed line. Legs with the visible portion of
the front coxae globular and trochantin completely concealed by the hypomera.
Claws without teeth. Alimentary canal (text-fig. 106) with seven caeca on the
anterior margin of the mid-gut. Hind gut with six Malpighian tubules which
end near the rectum freely or embedded in the fatty tissue. Male reproductive
system (text-fig. 107) with the lateral accessory glands with several lobes. Each
testis with three sperm tubes. Female reproductive system (text-fig. 10S) with
eight egg tubes to each ovary. Spermathecal duct opening near base of bursa
MEXICAN ELM I DAE 263
copulatrix. Central nervous system (text-fig. 105) with three thoracic discrete
ganglia. First abdominal ganglion partly fused to third thoracic, two to four
free, and five to eight partly fused together though the limits of each are distin-
guishable.
Genotype : Elsianus striatus Sharp.
The internal anatomy of four species has been examined and found to agree
in essential details. E. bicnlor Hinton of Bolivia is figured.
Elsianus may be immediately distinguished from the other genera in the
tribe Elnrini (except Stenelmis Dufour) by the accessory stria on each elytron at
basal fifth or sixth between sutural and second stria. With the exception of
Stenelmis Dufour, it is the only genus known to me which has three sperm tubes
to each testis. From Stenelmis it may be distinguished by the densely pubescent
apices of the tibiae and the presence of caeca on the anterior margin of the mid-gut.
The specific characters of most importance in separating the species of
Elsianus seem to be the following :
(1) General proportions, length and breadth.
(2) Size and distribution of the punctures and tubercles on the various
sclerites.
(3) Condition of fronto-clypeal suture.
(4) Anterior margin of clypeus, whether rounded, truncate, emarginate,
or sinuate ; and also shape of angle on each side.
(5) Condition of anterior margin of labrum.
(<>) Outline of pronotum and extent and depth of the impression or
gibbosities on its surface. Shape and extent of sublateral carinae
if present.
(7) Shape of elytral apices.
(8) Condition of lateral margin of elytra, whether crenate or smooth.
(9) Shape of scutellum and if convex or flat.
(10) Shape of prosternal process.
(11) Shape of mesosternal groove and depth.
(12) Impressions on disk of metasternum.
(13) Condition of carinae of first abdominal sternite.
(14) Extent and depth of impression on middle of first abdominal sternite.
(15) Extent and depth of depressions on other sternites. A depression is
frequently present on apex of fifth sternite.
(16) Secondary sexual character-.
(17) Structure of male genitalia.
(iS) Relative size of the dorsal and ventral caeca on the anterior margin
of the mid-gut. In some species the ventral caeca are very much
-mailer than dorsal, while in other species all caeca aie the same
size.
(iq) Shape of median and lateral accessory glands of male reproductive
system.
The following secondary sexual characters have been observi d by examining
the 30 or so species in my collection :
(1) Male with the anterior portion of the labrum very densely clothed with
line, long, erect, pale hair-, while in the female the han- are at most
two-thirds as long and are confined to -ides (clypeatus, larsa/is).
(2) Metasternum of males I'll each side oi ili-k with an area clothed with
long, tine, testaceous, recumbent hairs (clypeatus, tarsa
264 H. E. HINTON
(3) Metasternal disk on each side with a large and deep depression in males
and with a broad, scarcely noticeable depression in females
{clypeatus).
(4) Metasternum with the median longitudinal line much more broadlv
impressed in male than in female (clypeatus).
(5) Middle of first abdominal sternite more depressed in male than in
female (scutellaris).
(6) Apical margin of fifth abdominal sternite differently formed in male
than in female (sp. n., Bolivia).
(7) Male with the shape of the depression on the apical abdominal sternite
different to that of the female (aeqnalis, clypeatus).
(8) Apical abdominal sternite of male with a broad, basal, non-granulate
and nearly impunctate area, while in the female this area is granulate
(chpeatus, scutellaris).
(9) Apical third of fifth abdominal sternite of male with an area on each
side which is densely clothed with very long, fine, golden hairs,
while in the female the same area is only clothed with very much
shorter and sparser hairs (clypeatus, tarsal is).
(10) Male with the apex of the hind tibiae broadened and on inner side
concave (sp. n., Brazil).
(11) Hind tibiae of male with the inner apical spur long and flexed inwards,
whereas in the female the corresponding spur is short and straight
(tarsal is).
(12) Hind tibia of male with the inner apical spur relatively broader than
corresponding spur of female (clypeatus).
(13) Males with long and fine hairs on ventral side of four basal segments
of all tarsi, whereas in the female there are only short and stout
hairs or spmes(aeqnalis).
(14) Males with long and fine hairs on ventral apices of four basal segments
of hind tarsi (sandersoni , striatoides).
A Key to the Mexican Species of Elsianus.
1. Pronotum with a prominent gibbosity near base in front of scutellum . 2.
Pronotum without a gibbosity ....... 3.
2. Species over 4-0 mm. long. Prosternal process with the apex acute;
mesosternum with the posterior part of the groove for the reception
of the prosternal process narrowed to an acute point. Mexico.
E. scutellaris Hinton (1934).
Species 4 -o mm. long or less. Prosternal process with the apex broadly
rounded ; mesosternum with the posterior part of the groove for the
reception of the prosternal process very broadly and feebly rounded.
Costa Rica, Mexico . .... E. graniger Sharp (1882).
3. Species usually about 6-0 mm. and never less than 5-5 mm. long. Mexico
E. grandis Hinton (1934).
Species never more than 5-0 mm. long ...... 4.
4. Male with no long and fine hairs on ventral apex of first four segments of
hind tarsi. Guatemala, Mexico. . . E. striatus Sharp (1882).
Males with numerous long (about equal to lengths of their respective
segments) and fine hairs on ventral apex of first four segments of
hind tarsi .......... 5.
MEXICAN ELMIDAE 265
5. Male genitalia with the median lobe extending much beyond apices of
lateral lobes. Mexico .... E. striatoides Hinton (1936).
Male genitalia with the median lobe not extending to apices of
lateral lobes. Mexico .... E.sandersoni Hinton (1936).
Elsianus scutellaris Hinton.
(Text-figs. 111-114.)
1934. Elsianus scutellaris Hinton, Rev. lint., Rio de /., 4 (21 : [96.
.l/(i/<' ; Length, 4-0 mm. ; breadth, i-q mm. Cuticle .shining and black,
with the antennae, mouth-parts, and tarsi rufo-piceous. Head without distinct
impressions ; surface with feebly oblong granules which are about a third coarser
than facets of eyes or 0-025 mm. broad and are separated usually by slightly less
than to once their diameters ; surface between the granules microscopically
alutaceous. Clypeus with the fronto-clypeal suture moderately deep and feebly
and arcuately emarginate for its entire breadth ; anterior margin feebly and
arcuately emarginate for its entire breadth, with the angle on each side very broac 11 v
rounded ; surface sculptured as head but with the granules slightly flatter and
sparser. Labrum with the anterior margin feebly rounded and with the angle on
each side broadly rounded ; surface with the basal and lateral parts extremely
densely and very finel}' alutaceous, middle apical region not alutaceous and with
punctures which are about a third as coarse as granules of head and are separated
by one to five times their diameters ; sides with a few long (0-075 mm.), fine,
testaceous hairs. Pronotum across broadest point, which is about at basal third,
broader than long (1-62 mm. : 1-50 mm.) and base broader than apex (1-40 mm. :
0-97 mm.). Sides arcuate, moderately strongly and shortly sinuate before basal
angles and very feebly and broadly sinuate at apical two-fifths. Lateral margins
feeebly and somewhat regularly crenate. Sublateral carina prominent, extending
from near base to apical seventh, and feebly sinuate and for short distance much
less prominent at about basal two-fifths ; on basal seventh in front of scutellum
with a longitudinal gibbosity which is o-io mm. broad, 0-22 mm. long, and about
0-07 mm. high ; disk with a very shallow moderately broad median longitudinal
impression which extends from apical two-fifths to near anterior part of gibbosity.
Surface of pronotum sculptured similarly to that of head but with the granules
round, not oblong, very slightly coarser, and usually separated by once to twice
their diameters ; extreme base and inner sides of sublateral carinae on basal
fourth with the alutaceous microsculpture much less dense than elsewhere.
Elytra more than twice as long as pronotum (3-4111111. : 1-50 mm.) and broadest
point, which is at apical third, slightlv broader than broadest point across humeri
i-q mm. : 1-8 mm.). Lateral margins finely and regularly crenate. Apices
moderately produced and each apex obliquely truncate so that the mure anterior
part of the truncation is mesal. Surface with the striae coarse and becoming
finer towards apex and slightly coarser towards sides ; discal stnal punctures
usually round, about a hall to a third as coarse as intervals, and separated longi-
tudinally by one to a little more than one diameter. Intervals more "i
Ks, tiat but with the third (or fourth it accessory interval is included) feebly
convex from basal tilth to eighth ; surface oi intervals sculptured as pronotum
but intervals at middle apical three-fourths with the granules much sparser.
Scutellum subovate, strongly convex, particularly posteriorly, longer than broad
(0*25 mm. : 0*22 mm.), base broadly and feebly rounded, and much narrowed
to the feebly rounded apex : surface sculptured as the adjacent elytra! intervals
MOVIT, ZOO! ., 4-1, J. [940. 2.-
266
H. E. HINTON
but with the granules slightly denser. Presternum with the process (text-fig. 112)
acute at apex and the margin opposite front coxae raised so that middle part of
process and basal part of presternum appear moderately depressed ; surface of
process coarsely and densely rugose and anteriorly transversely and more sparsely
so ; anterior portion of presternum granulate somewhat similarly to clypeus and
sides as well as hypomera granulate as pronotal disk. Mesosternum with the
groove for the reception of the prosternal process deep and broad and posteriorly
narrowed to an acute point ; surface at sides sculptured as middle apical region
Text-figs. 111-114. — Elsianus scutellaris Hinton. (ml Adult to show general
appearance. (112) Presternum. (113) Lateral view of male genitalia. (114)
Dorsal view of male genitalia.
of prosternal process. Metasternum with the median longitudinal impressed
line extending to anterior eighth ; disk with the posterior four-fifths deeply
depressed ; surface of disk with the granules about a third coarser than those of
pronotum, oblong and separated mostly by a third to once their length ; sides
of metasternum granulate as disk of pronotum. Abdomen with the first stemite
feebly depressed at middle and anterior and basal lateral margins moderately
strongly raised ; carinae of first stemite straight and complete ; surface of middle
of first sternite densely rugose and granulate, surface of other sternites granulate
somewhat similarly to pronotum ; basal middle of fifth abdominal sternite
without granules or punctures but only microscopically alutaceous ; apex of
fifth sternite with a large, tranversely oval, moderately deep depression.
Genitalia as figured (text-figs. 113, 114).
MEXICAN ELMIDAE 267
Female: Externally similar to male except as follows : (i) the mid. 11. oi
the first abdominal sternite is only very feebly depressed ; (2) there is no middle
basal impunctate and non-granulate area on the fifth abdominal sternite. ; and
(3) the apical depression of the fifth sternite is much shallow. 1 .
Type: £ in the British Museum (Nat. Hist.). Mexico: Dist. de Temas-
caltepec, Temascaltepec, alt. about 5600 ft., 2S.V.1933 (H. E. Hinton, R. L.
I 'singer).
Specimens examined : 1^. with same data as above but collected in 1934
(H. E. Hinton).
Variations : In some males the basal impunctate area of the tilth abdominal
sternite is confined to the extreme basal middle part, while in other specimens it
extends on middle as far as apical two-thirds.
Comparative notes : this species belongs to a small group which is charac-
terized by having a prominent gibbosity on the pronotum in front of the
scutellum. The only other representative of this group in North America is the
Guatemalan and Mexican E. graniger Sharp. From graniger it may be distin-
guished as follows : (1) Scutellaria is larger (4-6 mm. : 4-0 mm. — largest specimen
of graniger I have seen) ; (2) apices of elytra obliquely truncate with the most
anterior part of the truncation lateral, whereas in graniger the most anterior part
of the truncation is mesal ; (3) prosternal process with the apex acute instead of
broadly rounded ; and (4) mesosternum with the anterior part of the groove for
the reception of the prosternal process narrowed to an acute point instead of
broadly rounded. The male genitalia of the two species are also quite different
(cf. figures).
Elsianus graniger Sharp.
(Text-figs. 115-117.)
1882. Elsianus graniger Sharp, Biol. Centr.-Amer. Col., 1 (2) : 133.
Male : Length, 3-4 mm. -4-0 mm. ; breadth, 1-3 mm.-i-8 mm. Cuticle
shining and rufo-piceous to black ; antennae, mouth-part- and legs paler rufo-
piceous. Head without distinct impressions ; surface with round to feebly
oblong granules which are about 0-024 mm- broad or about a third coarser than
facets of eyes and are usually separated by once their diameters Or a little more ;
surface between granules microscopically alutaceous. Clypeus with the fronto-
clypeal suture moderately deep and so shaped that the anterior margin of the head
is feebly and arcuately emarginate for its entire breadth between antennae ;
anterior margin very feebly emarginate or nearly truncate for its entire breadth ;
and with the .mgle on each side very broadly rounded ; surface sculptured as
head but with the granules slightly Hatter. Labium with the anterior margin
broadly rounded and the angle on each side broadly rounded : surface a1 sides
and on basal middle sculptured as clypeus but more finely so ; -111 face on a middle
apical belt with punctures only, which are about two-thirds as coarse as granules
and are separated by one to four times their diameters ; each -id. with a few line,
long (about 0-05 nun.), recumbent, testaceous hairs. I'yonotum at broadest point,
which is at bas.d third, broader than long (1-35 mm. : 1-22 mm.) and base broader
than apex (l-20 mm. : 0-8o mm.). Sides arcuate, less strongly so at basal and
apical third but nowhere sinuate. Lateral margins feeblj and regularly crenate.
Sublateral carina.' prominent, extending from very near base to apical ninth, and
feebly sinuate and lor a short distance much less prominent at basal two-tilths ;
on basal seventh in front of scutellum with a gibbosity which is 0-12 mm. Ion-,
268
H. E. HIXTOX
0-07 mm. broad, and about 0-04 mm. high ; disk with a narrow, very shallow, and
indistinct median longitudinal impression extending from basal fourth to apical
fourth. Surface of pronotum sculptured similarly to that of head but with the
granules always round, often slightly coarser, and separated mostly by once to
twice their diameters ; surface along extreme base with the alutaceous micro-
sculpture much less dense. Elytra more than twice as long as pronotum (2-45 mm. :
1-22 mm.) and broadest point, which is at apical third, slightly broader than
Text-figs
115-
117
17. — Elsianus graniger Sharp.
(116) Left lateral view of same.
(115) Dorsal view of male
(117) Presternum.
renitalia.
broadest point across humeri (1-40 mm. : 1-30 mm.). Lateral margins finely
and regularly crenate. Apices moderately produced and each apex obliquely
truncate so that the most anterior part of the truncation is mesal and not as usual
lateral. Surface with the striae moderately coarse and becoming coarser towards
sides and finer towards apex ; discal strial punctures usually round, from a third
to two-fifths as coarse as intervals, and separated longitudinally by once their
diameters or a little more. Intervals all more or less flat but with the third
(not including extra interval formed by accessory stria) more obviously convex
from basal sixth to eighth ; surface of intervals sculptured as pronotum but on
discal part of apical half with the granules distinctly sparser. Scutellum sub-
ovate, strongly convex, broader than long (0-21 mm. : 0-20 mm.), base broadly
MEXICAN ELMIDAE 269
and feebly rounded, and slightly narrowed towards apex which is broadly
rounded; surface sculptured like adjacent intervals. Prosternum with the process
(text-fig. 117) broadly rounded at apex and margin near base feebly raised
so that area opposite this point appears feebly depressed ; surface of process
coarsely and densely rugose and also with the granules about a fourth coarser
than those of elytra and separated by two to four times their diameters ;
anterior portion of prosternum, sides, and hypomera sculptured somewhat like
pronotum. Mesosternum with the groove for the reception of the prosternal
process deep and broad and with the posterior margin broadly rounded ; surface
at sides sculptured as middle apical region of prosternal process but more finely
so. Metastemum with the median longitudinal impressed line broad and extending
to about anterior fifth ; surface of disk with the granules about a third to half
again as coarse as those of elytra and separated mostly by once their lengths,
though very occasionally they are contiguous ; sides of metastemum granulate as
base of elytra. Abdomen with the first sternite at middle only feebly and broadly
depressed and anterior and lateral basal margins moderately strongly raised ;
carinae of first sternite straight and complete. Surface of first sternite at middle
sculptured nearly as that of prosternal process, surface elsewhere and that of
other sternites granulate as pronotum but with the granules slightly flatter and
often feebly oblong. Apex of fifth sternite with a broad, transverse, and
-hallow impression. Genitalia as figured (text-figs. 115, 116).
I cm ale : Externally similar to male.
Type: In the British Museum (Nat. Hist.). Costa Rica: Cache [H.
Rogers).
Specimens examined : 52, Mexico : Dist. de Temascaltepec, Temascaltepec,
alt. about 5600 ft., vi-vii.1933 (H. E. Hinton, R. L. Usinger) ; 23, same locality
but vi-vii.1934 (H. E. Hinton) ; 3, -am.- data as above but at Tejupilco, alt.
about 4000 ft., \ii. ii)34 (H . E. Hinton) ; and 4, MEXICO : Estada de Morelos,
Cm rnavaca, vi.1034 (H. E. Hinton).
Variations : In some the sides of the pronotum are distinctly sinuate before
basal angles. The basal gibbosity of the pronotum varies from being distinctly
longer than broad to being quite round, and as regards prominence, from 0-05 mm.
to 0-015 mm- high.
Comparative notes : This specie-- can be compared only with the larger
E. Scutellaria Hinton, as these two are the only North American species possessing
a gibbosity in front of the scutellum on the pronotal base. I he differences
between these two are given under the d. scription of scutellaris.
Elsianus grandis Hinton.
1 i ext-figs. 1 18 1 jo.)
1113.). Elsianus grandis 1 1 mi mi, R, v. /:«/., Rio de J., 4 (2) : [95.
Male: Length, 57 nun. 6-0 mm.; breadth, 2-3 mm. 2-6 mm. Cuticle
shining and nrfo-piceous to black ; antennae, mouth-parts, and leg- paler rufo-
piceous. Head without distinct impressions ; surface with round, high granules
which are about 0-018 mm. broad or about .1- coarse a- facets oi eyes and are
usually separated by two to three time- their diameters; surface between the
granules densely and microscopically alulae, cm-. ( lypeus with the lrouto-
1 ij 1 eal suture deep and formed so that the ant. 1101 margin oi the head 1- very
f.ebly and arcuately emarginate for its entire breadth between antennae ; antei ioi
270
H. E. HIXTON
margin arcuately and very feebly emarginate for its entire breadth and with the
angle on each side very broadly rounded ; surface sculptured as that of head.
Labrum with the anterior margin feebly rounded and with the angle on each side
broadly rounded ; surface without distinct granules but densely and micro-
scopically alutaceous except for a narrow anterior belt which is broadest at middle
and which is only sparsely alutaceous ; sides with a few long (0-07 mm.), fine,
recumbent testaceous hairs. Pronotum across broadest point, which is at basal
I20
Text-figs, i 18-120. — Elsianus grandis Hinton. (11S) Dorsal view of male genitalia.
(119) Left lateral view of same. (120) Presternum.
two-fifths, broader than long (2-05 mm. : 1-62 mm.) and base broader than apex
(1-85 mm. : 1-17 mm.). Sides arcuate, nearly straight or scarcely noticeably
sinuate at apical fourth, and feebly sinuate before basal angles. Lateral margin^
finely and regularly crenate. Sublateral carinae prominent, extending from base
to apical fifth, and sinuate and for a short distance less prominent at basal two-
fifths ; disk without a distinct median impression ; with a moderately shallow and
broad depression beginning on inner side of sublateral carina at basal two-fifths
and extending obliquely backwards for a distance equal to a little more than the
length of the scutellum. Surface sculptured similarly to that of head but with
MEXICAN ELMIDAH 271
tin granules on disk not so high and separated by three to four times their
diameters, the surface between the granules much less densely alutaceous, and
the granules at sides of pronotum about a third coarser and slightly denser ; area
near inner side of sublateral carina near basal fourth nearly free (if granules.
Elytra more than two and a half times as long as pronotum (4-4 mm. : t-6 mm.)
and broadest point, which is at apical third, broader than broadest point across
humeri (2-45 mm. : 2-15 mm.). Lateral margins finely and regularly erenate.
Apices broadly and moderately produced so that each apex is obliquely truncate
and the most anterior part of the truncation is lateral. Surface with the striae
coarse and becoming finer towards apex and slightly coarser towards sides ; discal
strial punctures usually round, deep, about a third as broad as intervals, and
separated longitudinally by slightly more than their diameters. Intervals more
or less flat ; surface of intervals sculptured as sides of pronotum but with the
granules slightly sparser and on middle apical three-fifths and lateral half much
sparser and finer, here often being separated by as much as five times their
diameters. Scutellum strongly convex, nearly round but slightly longer than
broad (0-37 mm. : 0-32 mm.), and base and apex broadly rounded ; surface
sculptured as sutural interval. Prosternum witli the process (text-fig. 120)
broadly rounded at apex and the margin opposite middle coxae very thickly and
feebly raised ; surface of process sculptured as sides of elytra and also feebly
rugose ; surface of anterior portion of prosternum, sides and hypomera sculptured
as sides of pronotum. Mesosternum with the groove fur the reception ol the
prosternal process broad and deep but posteriorly much narrowed and much
deeper than elsewhere ; sides with the surface coarsely rugose. Metasternum
with the median longitudinal impressed line extending to anterior sixth ; disk
with the posterior four-fifths feebly depressed ; surface of disk with the granules
oblong, about 0-05 mm. long, and usually separated by once their lengths though
occasionally contiguous ; sides of metasternum granulate as hypomera. Abdomen
with the middle of the first sternite broadly and moderately deeply depressed on
most of the middle area ; first sternite without carinae ; surface of sternites
granulate as sides of metasternum but with the granules becoming finer and
slightly sparser towards apex ; middle apical portion of first sternite and middle
basal portion of second sternite with the surface between the granules nearly
free of the alutaceous microsculpture and highly polished. Genitalia as figured
(text-figs, us, 119).
Female : Externally similar to male.
Type: j in the British Museum (Xat. Hist.). Mexico: Dist. de Temas-
caltepec, Real de Arriba, alt. about 7000 ft., r3.vi.i933 (H. H. Hinton, R. I..
U singer).
Specimens examined : 3, with data as above; 5, with same data but taken
vi-vii.IO/34 {H. E. Hinton) ; and 2, in the same district at Rio Verde, alt. 71.1111-
Sooo ft., 14. vi. 1934.
Variations : The scutellum is in some specimens moderately narrowed in
apical half, whereas in others it is nearly round. The apical sixth of the scutellum
of one specimen is very abruptly flattened whereas in the remainder of the series
the scutellum is fairly evenly convex.
Comparative notes: This is the largest species <>i North American Rimini.
From its close allies E. striatus, /•. . striatoides and E. sandersoni it may at once be
distinguished by its much larger size (the Largesl specimen oi these time species
which I have seen is 5-0 mm., while the smallesl nt grandis is 5-7 mm.) and the
structure of the male genitalia (cf. figures).
272 H. E. HINTON
Elsianus striatus Sharp.
(Text-figs. 109, no, 121-124.)
1882. Elsiaims strialus Sharp, Biol. Centr.-Amer. Col., 1 (2) : 132.
Male : Length, y$ mm. -4-5 mm. ; breadth, 1-5 mm.-i-8 mm. Cuticle
shining and rufo-piceous to black ; antenna, mouth-parts and legs paler
rufo-piceous. Head without distinct impressions ; surface with round high
granules which are about 0-012 mm. broad or about as coarse as facets of eyes
and are usually separated by once to twice their diameters or occasionally
slightly more ; surface between the granules densely and microscopically
alutaceous. Clypeus with the fronto-clypeal suture moderately deep and so
shaped that the anterior margin of the head is arcuately and moderately feebly
emarginate for its entire breadth between antennae ; anterior margin of clypeus
moderately strongly, arcuately emarginate for its entire breadth and with the
angle on each side very broadly rounded ; surface sculptured as head. Labrum
with the anterior margin feebly rounded and with the angle on each side broadly
rounded ; surface without distinct granules and rather densely and microscopically
alutaceous except for a narrow anterior belt which is broader at middle and which
is without the alutaceous microsculpture ; the surface of this belt is frequently
with punctures which are about a fourth to two-thirds as coarse as granules of
clypeus and are contiguous to separated by five times their diameters ; surface
at middle of apex often impunctate ; at sides with a few fine, long (about 0-05 mm.)
recumbent, testaceous hairs. Pronation across broadest point, which is about at
basal third, broader than long (1-30 mm. : 1-15 mm.) and base broader than apex
(1-15 mm. : 0-82 mm.). Sides arcuate, broadly and distinctly sinuate before
basal angles, and very feebly scarcely noticeably sinuate at apical two-fifths.
Lateral margins feebly and regularly crenate. Sublateral carina prominent,
extending from very near base to apical seventh, and very feebly sinuate at
about basal two-fifths ; disk without impressions. Surface granulate as head
but with the granules on disk slightly finer, separated by two to three times
their diameters, and with the surface between them set with fine punctures which
are half as coarse as granules and are confluent to separated by once their diameters;
towards sides the granules become slightly larger and the punctures denser so
that finally at sublateral carina the surface is again microscopically alutaceous
as on head ; surface between sublateral carinae and lateral margins with the
granules about a third coarser and slightly denser than those of head. Elytra
nearly twice as long as pronotum (2-62 mm. : 1-15 mm.) and broadest point,
which is at apical third, slightly broader than broadest point across hum.ri
(I-55 mm- : 1 "45 mm.). Lateral margins finely and regularly crenate. Apices
moderately produced and each apex obliquely truncate so that the most
anterior part of the truncation is lateral. Surface with the striae coarse and
becoming slightly finer towards apex and slightly coarser towards sides ; discal
strial punctures usually round to feebly subquadrate, two-thirds as coarse as
intervals and separated longitudinally by once their lengths or a little more.
Intervals flat but near base discal intervals are very feebly convex ; surface of
intervals sculptured as sides of pronotum mesal to sublateral carina but with the
granules sparser and the surface between the granules not punctate and only
occasionally alutaceous ; discal region on apical half with the granules much
finer and sparser. Scutcllum subovate to nearly round, strongly convex, longer
MEXICAN ELMIDAE
273
than broad (0-22 mm. : 0-17 mm.), base broadly and feebly rounded,
apex
narrowed and feebly rounded ; surface sculptured as adjacent elytral intervals.
Prostemum with the process (text-fig. 122) broadly rounded at apex and the lateral
margins feebly and broadly raised so that the middle part of the process is mode-
rately concave; surface of process sculptured as ana of pronotum between
121
rEXT-Fics. 1 j 1 [24.- -Elsianus striatum Sharp. 1121) Adult to show general appear-
ance. (122) Presternum. 1125) Dorsal view <>t male genitalia. (124) Left la
view of|same.
lateral margins and sublateral carinae but with the granules about a fourth
coarser and slightly sparser; anterior ana of presternum granulate as ba al
discal pari of elytra ; sides and hypomera with the granules aboul as coarse as
those of anterior part of presternum but distinctly oblong and never round and
usually separated by oner their diameters. Mesostemum with the groovi foi
the reception "I the prosternal process deep and broad and narrowed posteriorly
2J4 H. E. HINTON
where it is much deeper than elsewhere ; surface at sides sculptured as prosternal
process. Metasternum with the median longitudinal impressed line extending to
anterior fourth or fifth ; disk nearly flat, posteriorly only very feebly depressed,
and surface with the granules about as coarse as those of prosternal process but
flatter and usually confluent to separated by nearly once their diameters so that
the whole surface has a subrugose appearance ; sides of metasternum with the
granules similar to those of sides of presternum. Abdomen with the middle of
the first sternite broadly and moderately strongly depressed ; first sternite
without carinae ; surface of basal middle of first sternite sculptured as prosternal
process ; surface elsewhere and that of other sternites granulate as sides of
metasternum but with the granules less oblong and more nearly round ; surface
of apical middle of first, all of middle of second, and basal middle of third sternite
highly polished between granules. Genitalia as figured (text-figs. 123, 124.)
Female : Externally similar to male.
Type: In the British Museum (Nat. Hist.). Guatemala : Rio Naranjo,
alt. 450 ft. (Champion).
Specimens examined: 3, Mexico: Dist. de Temascaltepec, Tejupilco, alt.
4000 ft., vii.1932 (H. E. Hinton) : 205, with same data as above but collected in
vi.1933 (H. E. H niton, R. L. Usinger); 11, as above but taken in vii.1934 (H. E.
Hinton); and 9, Mexico: Estado de Morelos, Cuernavaca, vi.1934 (H . E.
Hinton).
Variations : No variations worthy of mention have been noted.
Comparative notes : This is very closely related to E. striatoides Hinton and
E. sandersoni Hinton. From both of these only the males can be positively
distinguished by not having the ventral apical tufts of long and fine hairs on the
four basal segments of the hind tarsi and by the structure of the male genitalia.
The females of striatus are inseparable from those of striatoides, but from those of
sandersoni they may be distinguished by their smaller size — being never more than
4-5 mm., whereas 5-0 mm. appears to be the normal size of females of sandersoni.
Elsianus striatoides Hinton.
(Text-figs. 125-127.)
1936. Elsianus striatoides Hinton, Trans. R. Ent. Soc. Loud., 85 (18) : 422, figs. 15-17.
Male : Length, 4-5 mm. ; breadth, 1-9 mm. Identical to striatus except as
follows : (1) at ventral apex of each of the four basal segments of the hind tarsi
with about two to four very fine, partly erect testaceous hairs which are about as
long as their respective segments ; and (2) male genitalia with the apex of the
median lobe extending much further beyond apices of lateral lobes than is the
case in striatus (cf, figures).
Female : Externally similar to male except that the ventral apices of the four
basal segments of the hind tarsi have no long and fine hairs.
Type : J in the British Museum (Nat. Hist.). Mexico : Dist. de
Temascaltepec, Temascaltepec, alt. 5600 ft., 28. v. 1933 (H . E. Hinton, R. L.
Usinger).
Specimens examined : I, $ with data as above but collected on 5.V1.1933 ;
and 4, +; which may belong to this species collected in the same district but at
Real de Arriba, alt. about 7000 ft., 25. v. 1933.
MEXII AS I I. \llh U
275
Elsianus sandersoni Hinton.
( rext-figs. 128 130.)
1936. Elsianus sandersoni Hinton, Trans. R. Ent. Soc. Lond., 85 (18) : i-'o, figs. ii -14.
Male : Length, 4-5 mm.-jo mm. ; breadth, 1-87 mm.- 2-05 mm. Identical
to striatoides except that the genitalia have the apices of the lateral Lobes extending
I 1 1
127
130
Text-figs. 125-130. — (125) Dorsal view of male genitalia oi I '■■'• '
Hinton. (126) Left lateral view of same. (127) Presternum of samesni s. [i j.si
Dorsal view of male genitalia of E. Sanders mi Hinton. (129) Left lateral view ol
same (130) Presternum of same spei ies.
slightly beyond the apex oi the median lobe (if. figures). First segment oi front
and middle tarsi with long and fine hairs on the ventral apex which are similar to
those of first segment of hind tarsi. /•.. striatoides apparently has not fine long
hairs on the ventral apex of the middle and front tarsi, but this maybe due entirely
to the fact that my two male specimens are badly rubbed.
Female : Externally similar to male but without the long and fine hair- on
the tarsi.
Type: <J in the British Museum (Nat. Hist.). Mexico: Dist. deTemas-
caltepec, Tejupilco, alt. 4000 ft., vii.1934 (//. /•.'. Hinton).
276
H. E. H1NTON
Specimens examined: 1, J, Mexico: Estado de Morelos, Cuemavaca,
vi.1934 (H. E. Hinton) ; and 4 $$ with same data.
I have placed these females with this species because of their large size
(5-0 mm.) though, apart from size, I have no good evidence that they belong
to sander soni, as striatus has also been taken in this locality.
Pupae.
About 18 specimens of the pupa of Elsianus graniger Sharp were taken by
peeling off the moss matting which was growing above the water level on large
stones in a small torrential stream. The pupal cells were formed in the earth
held by the roots of the moss, and occasionally the ventral side of the cell was
formed by the surface of the stone. A few of the pupae were allowed to emerge,
and in this way the species was determined beyond doubt.
Description of Pupa of E. graniger Sharp.
(Text-fig. 131.)
Male : Length, 4-4 mni.-5-j mm. ; breadth (across metathorax and including
front wings), 17 mm. -1-9 mm. Head, concealed from above by pronotum so
131
Text-fig. 131. — Pupa of Elsianus graniger Shaxp.
that only the vertex is visible ; without distinct impressions ; surface densely
microscopically alutaceous and with fine, erect pale hairs which are about
0-10 mm. long or more and arise at intervals from less than to one and a half
their lengths. Antennae in normal position extending posteriorly and down-
wards to a point opposite anterior margin of front coxae. Clypeus with the
MEXICAN ELMIDAE ZJJ
fronto-clypeal suture deep and straight ; anterior margin nearly truncate and
the angle on each side broadly rounded ; surface as head but with a complete
transverse row of setae at middle which are similar to the longer ones nt head
Labrum with the anterior margin broadly, deeply, and arcuately emarginate
and with the angle on each side broadly rounded ; surface alutai eous as head but
with only a few tine, short, recumbent setae. Pronotum similar in shape to that
of the adult but with the sublateral carinae only feebly raised and the gibbosity
in front of the scutellum not evident. Each apical and basal angle with a long
(about 0-50 mm.), stout, curved seta ; apical setae curved outwards and usually
slightly shorter than basal ; surface similar to that of head but with the hairs
slightly denser. Mesonotum with a large and round gibbosity (text-fig. 131)
which represents the scutellum beneath. Thoracic and abdominal segments as
figured (text-fig. 131) ; ninth abdominal segment at apex with two Lateral
projections. Wings with the anterior pair extending to ventral side, attaining
middle of third abdominal segment, and near apex separated from each other by
a distance equal to more than their greatest breadth ; posterior wings similarly
extending to ventral side but much closer near apex and attaining apical margin
of third abdominal segment. Legs with the front pair extending to posterior third
of metathorax and tarsi separated from each other by about a third of their
lengths ; middle pair extending to posterior margin of metathorax and at apex
seldom separated by more than the length of the fifth tarsal segment ; hind pair
with the coxa, trochanter, basal three-fourths- of femur, apical half of tibia, and
base of tarsus concealed by hind wings and the apical segments of the tarsi are
contiguous along the middle line of the body and the claws extend to posterior
margin of fifth abdominal sternite. Abdominal spiracles placed on dorso-lateral
sides and opening on apices of small tubercles.
Female : Externally similar to male except that the middle apical margin
of the eighth abdominal sternite (this is the sternite which corresponds to the
sixth from base in the adults) is rounded and not moderately deeply emarginate
as in male. The sexes may also be distinguished by the developing external
genitalia.
Larvae.
The larvae of this genus have been definitely determined as such, for the
cast skin of a last instar larva was found with a pupa in a cell, and when the pupa
emerged it proved to be that of Elsianus graniger.
The following brief generic diagnosis is the result of a study of the larvae of
about 15 species in my collection.
Generic ( haracters of Larvae of Elsianus.
Body parallel, cylindrical. Head when viewed dorsally exposed and not
concealed by the pronotum; anterior margin on each side between base oi
antenna and clypeus toothed. With one ocellus on each side. Antennae ;
segmented and feebly retractile. Mandibles of both sides similar and with three
obtuse apical teeth ; prostheea long, slender, and densely spinose. Maxilla
with the palp 4-segmented and the stipes showing no differentiation into a
palpifer ; galea and lacinia separate and apex of each densely spinose. Labium
with the postmentum undivided ; labial palp 2-segmented and prementum without
or with only a very feebly developed palpiger. Gula well-developed. Prothoracit
pleura (text-fig. 132) divided into two parts and anterior part meeting on middle
278 H. E. HINTON
line of body so that the sternum is here completely suppressed. Meso- and meta-
pleurae divided into two parts. Abdominal segments one to seven with the
pleurae bounded by tergo- and sterno-pleural sutures ; segments two to seven with
a dorsal suture parallel to the suture usually considered to be the tergo-pleural ,
and if this dorsal suture is in fact the true tergo-pleural, then we must consider the
pleurae to be longitudinally divided on segments two to seven ; segment eight
forming a complete sclerotized ring. Operculum with two strongly sclerotized
claws attached to its dorsal membrane. Apex of ninth segment feebly to strongly
emarginate. Spiracles present on mesothorax and first eight abdominal segments
and usually opening on small tubercles. Tracheae without air sacs. Three
tufts of retractile, anal, tracheal gills are present. Alimentary canal with an
oesophageal sclerite on the posterior dorsal margin of the oesophagus. Hind gut
with six Malpighian tubules which end freely near the rectum. Central nervous
system with three thoracic and eight abdominal discrete ganglia.
This genus is close to no other known to me. The larvae of four Mexican
species have been studied and of these it has only been possible with any certainty
to give names to two. A table for the separation of these species follows :
A Key to the Larvae of the Mexican Species of Elsianus.
1. Metasternum and first abdominal sternite anteriorly at middle with a
longitudinal carina ...... E. graniger Sharp.
Metasternum and first abdominal sternite without longitudinal
carinae ........... 2.
2. First abdominal sternite with the transverse and anterior reticulate belt
extending to posterior fourth at middle of segment E. striatus Sharp.
First abdominal sternite with the transverse and anterior reticulate
belt never extending at middle as far as posterior two-thirds of
segment ........... 3-
3. Setae which arise from posterior row of all segments except ninth round
and densely and finely hairy ; anterior middle portion of prosternum
very strongly produced and anterior angles moderately strongly
produced . . . . . . . . . E. sp. ?.
Setae which arise from posterior row of tubercles of all segments except
ninth flat and apically branched ; anterior middle portion of basal
prosternal sclerite only moderately produced and anterior angles not
produced E. (?) sp. ?.
Description of mature larva of E. graniger.
(Text-figs. 132^146, 150.)
Length, 8-0 mm. ; breadth (across broadest point which is near base of
metathorax), 1-15 mm. Elongate, subparallel, and cylindrical to subtriangular
in cross section. Cuticle moderately shining and brownish ; antennae, mouth-
parts, and legs paler brown to testaceous. Head at broadest point which is at
basal third slightly broader than long (070 mm. : 0-57 mm.) ; coronal suture
0-07 mm. long, very broad at base and narrow anteriorly ; frontal suture on each
side extending in a nearly straight line to anterior margin of head near base of
antenna ; anterior margin between base of antenna and clypeus with a tooth-
like projection which extends slightly beyond anterior margin of clypeus ; surface
sparsely pubescent with fine and erect hairs which are about 0-04 mm. long ;
MEXICAN ELMIDAE
■279
8 = =
280
H. E. HIXTON
surface finely and densely alutaceous throughout ; surface on a basal belt as long
as epicranial suture only alutaceous and elsewhere set with round to feebly oblong
granules which are about 0-03 mm. broad and are usually separated by once
their diameters, each granule with a coarse seta which is as long or a little longer
than its respective granule. Thoracic and abdominal tergites for the most part
sculptured as head but with the granules slightly coarser and the alutaceous
microsculpture sparse or absent ; pronotum with a number of small oval or
irregular areas which are free of granules ; meso- and metanotum with these
Text-figs. 141-146. — Larva of Elsianus graniger Sharp. (141) Operculum. (142)
Anterior view of right front leg. (143) Posterior view of left middle leg. (144)
Posterior view of left hind leg. (145) A seta from the ventral margin of the oper-
culum. (146) Dorsal view' of right opercular claw.
areas less numerous ; and abdominal tergites one to seven with a single
anterior and lateral oval non-granulate area which is from a sixth to a fourth
as long as its segment ; tergites of segments eight and nine with the granules
largely replaced by punctures of the same size and density as the granules of
other segments. Posterior margin of all segments except ninth with a complete
ring of close tubercles from which arise flat and apically slightly branched setae
(text-fig. 150). All tergites, pleurites and sternites except those of pronotum
with an anterior transverse, short belt which is free of granules and is coarsely
and reticulately alutaceous ; sternites of abdominal segments two to five with a
complete row of close tubercles along posterior margin of this belt. Sternites and
pleurites for the most part sculptured as tergites ; first abdominal sternite with
the sculpture as figured (text-fig. 150). Metasternum and first abdominal sternite
MEXICAN F.I.MIDAE 28l
with a longitudinal carina on the middle of the posterior third to fourth (text-fig.
132). Apex of ninth abdominal segment (text-fig. 135) deeply and transversely
emarginate. Operculum (text-fig. 141) with the claws (text-fig. 146) not toothed.
Legs as figured (text-figs. 142-144). Spiracles opening on small tubercles and
all similar to those of mesothorax (text -fig. 140).
Specimens examined : 58, Mexico : Dist. de Temascaltepec, Temascaltepec,
alt. 5600 ft., vi-vii.1934 (H. E. Hinton) ; 4, with same data but collected at
Tejupilco, alt. about 4000 ft. ; and 13, Mexico : Estado de Morelos, Cuernavaca,
vi.1934 (H. E. Hinton).
Among my series there are several specimens representing at least one earlier
instar, but these differ in apparently no way but size from the mature larvae.
Comparative notes : This larva may be distinguished at once from all others
known by the presence of a longitudinal carina on the mesosternum and the first
abdominal stemite.
Description of Mature Larva of E. striatus.
(Text-figs. 147-149, 151.)
The larvae of this species were determined by elimination and according to
locality.
Length, 9-0 mm. ; breadth (across broadest point which is near base of meta-
thorax), 1-15 mm. Elongate, subparallel, and cylindrical to subtriangular in
cross section. Cuticle moderately shining and brownish ; antennae, mouth-parts
and legs paler brown to testaceous. Head across broadest point, which is at
basal fourth, slightly broader than long (0-75 mm. : 0-72 mm.) ; posterior
margin of head broadly and arcuately emarginate at middle ; epicranial suture
0-05 mm. long ; frontal suture extending on each side in a line which is
slightly curved outwards to anterior margin opposite base of antenna ; anterior
margin on each side between base of antenna and clypeus with a tooth-like
projection which extends slightly beyond anteror margin of clypeus and is
feebly curved inwards. Cuticle sparsely pubescent with fine, erect hairs which
are usually 0-075 mm. long and on each side mesal to and slightly behind eye with
an erect seta which is three times this length ; surface finely and sparsely or
densely alutaceous, though often free of this alutaceous microsculpture ; surface
on a basal belt as long as epicranial suture only densely alutaceous and elsewhere
with granules which are about 0-012 mm. broad and at sides of head are usually
separated by once to twice their diameters, while on middle they are frequently
replaced by equally small punctures which are similarly distributed ; from each
puncture or granule a coarse, short seta arises which is about a third longer than
its respective granule or puncture. Clypeus with the fronto-clypeal suture visible
but indistinct ; broader and shorter than labrum ; anterior margin truncate, with
the angle on each side rounded ; surface basally sculptured as middle region of
head while on anterior middle region it is only sparsely alutaceous. Labrum as
long as second antennal segment and slightly narrower than clypeus ; anterior
margin feebly rounded and the angle on each side broadly rounded ; surface
punctate and setose as middle region of head but anteriorly with finr punctures
from which arise fine, erect, golden hairs which are slightly larger than the usual
coarse setae. Mandibles similar to those of graniger but with the fine non-plumose
seta on the middle of the outer margin proportionally only about half as long.
Maxilla and labium fairly close to that figured for graniger (text-figs. 1 37, 138) but
with the stipes oi the first maxilla slightly more densely setose. Thoracic and
NOVIT. ZOOL., 42, 2. 1940. i,\
282
H. E. HINTON
abdominal tergites for the most part sculptured as head but usually only with
granules which are slightly coarser and on posterior segments have the apices acute
and produced ; pronotum with a number of irregularly oval areas which are free
of granules ; meso- and metanotum with a similar oval area on lateral fourth and
another on lateral two-fifths ; and first eight abdominal tergites with a similar
area on lateral fourth which may vary from being a sixth to a fourth as long as its
respective segment. Posterior margin of all segments except ninth with a
complete ring of close tubercles from which arise flat and apically slightly branched
Text-figs. 147-151. — (147) Ventral view of thorax of larva of Elsianus striatus Sharp
to show sclerotization. (148) Reticulate sculpture of anterior region of first abdo-
minal sternite of same. (149) Dorsal view of ninth abdominal tergite of same.
(150) Sternite of first abdominal segment of E. graniger Sharp. (151) Sternite of
first abdominal segment of E. striatus.
setae (text-fig. 151). All tergites, pleurites, and sternites except those of pronotum
with an anterior transverse belt which is usually free of granules and is coarsely
and reticulately alutaceous ; this belt is generally about a fourth to a third as
long as its respective segment but that of ninth is less than a sixth as long as that
segment ; all sternites except that of prosternum and ninth abdominal segment
with a complete and transverse heavily sclerotized belt or ridge along posterior
margin of reticulate belt. Sternites and pleurites with acute tubercles similar
to tubercles of tergites ; first abdominal sternite sculptured as figured (text-fig.
151). Postero-lateral angles of all abdominal segments strongly and-acutely
produced. Apex of ninth abdominal tergite (text-fig. 149) deeply and truncately
emarginate. Operculum with the claws not toothed and at base on dorsal surface
MEXICAN ELMIDAE
283
with a number of very long, fine, recumbent hairs. Legs similar in size and shape
to those figured for graniger but with the chaetotaxy slightly different. Spiracles
similar to those of graniger.
Specimens examined : 103, Mexico : Dist. de Temascaltepec, Tejupilco,
alt. about 4000 ft., vii. 1934 {H. E. Hinton) ; and 13, Mexico : Estado de Morelos,
Cuernavaca, vi.1934 (H. E. Hinton).
Included in this material there are specimens representing at least two
earlier instars, but these differ apparently only in size from the mature larvae.
Text-figs. 152-158. — Larva of Elsianus sp. ? (152) Dorsal view of ninth abdominal
segment. (153) Spiracle of first abdominal segment. (154) Presternum. (155)
Operculum. (156) One of the usual tubercles of the first abdominal sternite. (157)
A tubercle from the posterior marginal row of the fifth abdominal tergite. (158)
First abdominal sternite.
Comparative notes: This species may be distinguished from E. graniger in
not having a longitudinal carina on the metasternum and first abdominal sternite.
The sculpture of the first abdominal sternite (text-fig. 151) differs markedly from
all species known to me.
Description of Mature Larva of Elsianus sp. ?.
(Text-figs. 152-158.)
Length, q-o mm. ; breadth (across broadest point which is near base of
metathorax), r-12 mm. Resembles striatus but may be distinguished as follows:
(1) thoracic and abdominal tergites with the setae which arise from the tubercles
plumose and not smooth ; (2) posterior margin of all segments except ninth with
284 H. E. HINTON
a complete ring of close tubercles from each of which arises a long, round, and
densely hairy seta, instead of, as in striatus, a flat seta with a few apical branches ;
(3) middle of anterior margin of posterior prosternal sclerite very strongly instead
of moderately produced (cf. text-figs. 154 and 147) ; (4) anterior angles of posterior
prosternal sclerite (text-fig. 154) strongly produced and rounded, whereas in
striatus they are not produced and are angulate ; (5) sternite of first abdominal
segment with the anterior reticulate belt at middle confined to anterior fifth of
sternite whereas in striatus it extends to posterior fourth ; and (6) apex of ninth
abdominal segment (text-fig. 152) slightly arcuately instead of truncately
emarginate.
Specimen examined: 1, apparently mature larva. Mexico: Dist. de
Temascaltepec, Temascaltepec, alt. 5600 ft., vii.1934 (H. E. Hinton.)
Description of Mature Larva of Elsianus (?) sp. ?.
(Text-figs. 159-163.)
Length, 8-5 mm. ; breadth (across broadest point which is near base of meta-
thorax), 0-82 mm. Elongate, subparallel, and cylindrical to subtriangular in
cross section. Cuticle feebly shining and brownish-testaceous ; antennae,
mouth-parts, and legs paler testaceous. Head across broadest point which is at
basal fourth broader than long (0-75 mm. : 0-65 mm.) ; posterior margin of
head at middle very broadly and only feebly arcuately emarginate ; epicranial
suture 0-12 mm. long and narrow ; frontal suture on each side extending in a
moderately sinuate line to anterior margin opposite base of antenna ; anterior
margin on each side between base of clypeus and antenna with a large and straight
tooth which is nearly twice as long as clypeus. Cuticle sparsely pubescent with
fine, erect hairs which are usually 0-15 mm. long ; surface on a basal belt as long
as epicranial suture smooth to moderately densely alutaceous and without
distinct granules or punctures ; surface elsewhere occasionally moderately
densely alutaceous though usually smooth and with flat round granules which are
about 0-03 mm. broad and are separated mostly by once their diameters ; from
these granules arise setae which are about as long or a little longer than their
respective granules ; surface of head with the area enclosed by the frontal sutures,
except for a small antero-lateral area on each side, free of granules. Antennae
retractile up to apex of first segment. Clypeus with the fronto-clypeal suture
visible ; broader than labrum and as long as second segment of antenna ; anterior
margin truncate and with the angle on each side rounded ; surface, except at
sides where there are a few granules, free of granules and sculptured as frontal
region of head. Labrum narrower and slightly shorter than clypeus ; anterior
margin nearly truncate and with the angle on each side broadly rounded ; surface
similar to that of clypeus but near middle with a complete transverse row of close,
fine, erect hairs which are slightly shorter than usual ones of head. Mandibles
with the seta on middle of outer margin proportionally as long as that of graniger.
First and second maxilla in general form similar to that figured for graniger, but
with the apical region of the postmentum about a fourth broader than the basal.
Pronotum with the surface similar to that of frontal region of head but with the
erect setae generally about a fourth longer, and at anterior third there is a short
(about two granules wide) transverse belt of tubercles which extends from lateral
margin on each side of the middle third of pronotum ; lateral margins basally
and anteriorly with a few granules similar to those of head ; mcso- and metanotum
similar but with the transverse band at middle three times as long (four to six
MEXICAN EI.MIDAK
285
granules broad) and extending right across middle, though basally interrupted at
lateral third ; first three abdominal tergites with only a short posterior belt and
an oval lateral area, which is about a third as long as any segment, free of granules ;
tergites four to seven without a posterior belt free of tubercles but with the lateral,
oval, non-tuberculate area present ; tergites eight and nine without the lateral,
oval, non-tuberculate area. Anterior reticulate or coarsely alutaceous belt of
tergites, pleurites, and sternites present on all segments except pronotum, usually
about a fifth as long as any segment, and along posterior margin with a complete
transverse heavily sclerotized ridge which may or may not be tuberculate and is
present except on pleurites, though on eighth and ninth segments it forms a
complete ring ; on abdominal segments one to seven it is on the sternites arcuately
159
to/fff,
wSMm
*d
163
161
Text-figs. 159-163. — Larva of Elsianus (?) sp. ? (159) Operculum. (160) Dorsal
view of ninth abdominal segment. (161) First abdominal sternite. (162) Tubercle
from first abdominal sternite. (163) Tubercle from posterior marginal row of first
abdominal sternite.
curved so that at middle it is nearer anterior margin than elsewhere. Posterior
margin of all segments except ninth abdominal with a complete ring of close,
large tubercles from which arise fiat setae which are about twice as long as their
respective tubercles and are branched apically (text-figs. 161 and 163). First
abdominal sternite sculptured as figured (text-fig. 161). Anterior margin of
posterior prosternal sclerite similar to that of striatus (cf. figure). Posterior
margin of meso- and metastemum strongly and acutely produced. Pleurae of
abdominal segments two to seven longitudinally not divided and tergo-pleural
suture ventral. Postero-lateral angles of tergites rounded and not produced.
Apex of ninth abdominal segment (text-fig. 160) broadly, deeply, and arcuately
emarginate. Operculum (text-fig. 159) with the claws not toothed and at base
on dorsal surface with a number of very long, fine, recumbent hairs. Legs similar
in shape and size to those figured for E. graniger. Spiracles similar to those of
graniger and slriatits.
Specimens examined: 3, apparently mature larvae, Mexico: Dist de
Temascaltepec, Temascaltepec, alt. 5600 ft., vii.1934 (H. E. Hinton).
Comparative notes : This is the only species of about 15 I have examined
286 H. E. HINTON
belonging to this genus which has not the pleurae of abdominal segments two to
seven longitudinally divided. It may possibly belong to a genus as yet unrecorded
from Mexico.
AUSTROLIMNIUS Carter cS: Zeck.
1929. Austrolimnius Carter & Zeck, Ausl. Zool., 6 (1) : 61,
1929. Neosolus Carter & Zeck, Aust. Zool., 6 (1) : 68.
1932. Austrolimnius Carter & Zeck, Aust. Zool., 7 (3) : 204.
Carter and Zeck (1929) in their monograph of the Australian Dryopidae
erected the genus Austrolimnius to contain two species described by King (1865)
in the genus Elmis Latreille and two new species. In the same monograph they
erected the genus Neosolus to contain a single new species, N. tropicus. Neosolus
was later (1932) sunk by them as a synonym of Austrolimnius. Altogether ten
species and three varieties have been described from Australia. The following
North and South American species should be referred to A ustrolimnius : A .
chiloensis (Champion) (Elmis prothoracica Hinton) ; A. curtulus (Sharp) (Elmis) ;
A. formosus (Sharp) (Elmis) ; A. laevigatas (Grouvelle) (Elmis) ; A. pilulus
(Grouvelle) (Elmis) ; and A. sulcicollis (Sharp) (Elmis). Altogether eight species
are known from North and South America, and of these two have been collected
as far north as Mexico, A. formosus and A . sulcicollis.
A redescription of the genus follows :
Body obovate to subparallel. Dorsal surface glabrous or clothed with
sparse and short recumbent hairs. Tomentum confined to the following areas :
(1) genae ; (2) epipleura ; (3) occasionally (A. luridus C. and Z. and A. minutus
Hinton) sides of elytra between inner margin of epipleura and outer sublateral
carina ; (4) hypomera ; (5) sides of prosternum, mesosternum, metasternum,
and abdominal sternites, but in some species nearly the entire sternum of
the abdomen is clothed with fine tomentum ; (6) part or all of legs except
tibiae and tarsi. Head when seen from below capable of being retracted so
that none of the mouth-parts are visible. Antennae n-segmented. Mandibles
with three apical blunt teeth ; prostheca large and entirely membranous with
numerous fine spines or hairs apically. Maxilla with the palp 4-segmented
and stipes with a well-developed palpifer ; galea and lacinia separate and apex
of each densely spinose. Labium with the palp 3-segmented and prementum
with a well-developed palpiger. Mentum transverse and about as broad as and
two-thirds to three-fourths as long as submentum. Gula slightly narrower
than submentum, about a third longer, and with the sides nearly parallel.
Pronotum with the anterior margin moderately to strongly arcuate at middle and
on each side behind eye before apical angle shallowly to deeply sinuate. Base
trisinuate, broadly and moderately deeply so on each side and more narrowly and
shallowly so in front of scutellum. Pronotum with a sublateral carina on each
side which extends from base nearly to apical margin and may or may not be
prominent ; disk with or without a median longitudinal impression. Elytra
impunctate or with distinct seriate punctures but seldom with striae which if
present are only very feebly impressed ; each elytron with two prominent sub-
lateral carinae. Epipleura with a longitudinal line of granules which on anterior
two-fifths is halfway between dorsal and ventral margins, while on posterior
three-fifths it is close to and parallel to ventral margin. Hind wings with the
venation much reduced ; with a well-developed anal lobe (text-fig. 168) ; without a
radial cross vein and without an anal cell ; first anal absent ; second anal without
MEXICAN ELMIDAK
287
branches ; third anal only present basally and joined to second anal at extreme
base ; fourth anal short and indistinct ; and cubito-anal cross vein arising from
second anal at about the middle of the length of the latter but not complete to
cubitus. Presternum very long in front of anterior coxae ; prosternal process
long and very broad and posterior margin broadly rounded. Mesosternum with
a broad and deep groove for the reception of the prosternal process. Metasternum
with a median longitudinal impressed line. Legs with the visible portion of the
front coxae rounded and trochantin completely concealed by hypomera and
Text-figs. 164-166. — AustroUmnius tarsalis Hinton. (164) Male reproductive system.
(165) Central nervous system. (166) Female reproductive system.
sternum. Claws without teeth. Alimentary canal with five caeca on the
anterior margin of the mid-gut. Hind gut with six Malpighian tubules which
end near the rectum freely or embedded in fatty tissue. Male reproductive system
(text-fig. 164) with the lateral accessory glands with several lobes. Each testis
with only one sperm tube. Female reproductive system (text-fig. 166) with two
egg tubes to each ovary. Spermathecal duct opening posteriorly to bursa
copulatrix. Central nervous system (text-fig. 165) with three thoracic discrete
ganglia. First abdominal ganglion fused to third thoracic, two and three free, and
four to eight fused into a single large ganglion.
Genotype: : Elmis politus King.
The internal anatomy of only one species, A. tarsalis Hinton, has been
examined.
288
H. E. HINTON
This genus is close to Limnius Erichson, but may be distinguished as follows :
the middle region of the head is without tomentum ; Carter and Zeck (1929 :
61) claim that there is no tomentum on the head in Austrolimnius, but the genae
are clearly tomentose in their genotype and in seven other species I have been
able to examine for this character ; (2) there is no tomentum on the pronotum
Text-figs. 167, 168. — Austrolimnius sulcicollis Sharp. (167) Adult to show general
appearance. (168) Hind wing. Venation after Forbes.
between the lateral margin and the sublateral carina, whereas in Limnius
tomentum, at least anteriorly, is usually present here ; (3) the elytron between
the first and second sublateral carinae is never tomentose, whereas in Limnius it is
usually at least partly tomentose ; (4) the hind wing has a very well-developed
anal lobe, whereas in Limnius the anal lobe is at most only very feebly developed ;
(5) the second anal extends to inner margin of wing opposite junction of cubitus
and media, whereas in Limnius the second anal is present only basally ; (6) in
MEXICAN ELMIDAK 289
Austrolimnius there is an incomplete cubito-anal cross vein, while in Limnius
there is no trace of such a vein ; (7) the male reproductive system has only one
sperm tube to each testis, whereas in Limnius it has two sperm tubes ; and (8)
the central nervous system has only the second and third abdominal ganglia free,
whereas in Limnius, the second, third, and fourth are free.
The specific characters of most importance in separating the species of
Austrolimnius seem to be the following :
(1) General proportions, length and breadth.
(2) Colour. In some species, e.g. luridus, suffusus, etc., the elytra are
maculate.
(3) Density of the asperate microsculpture on the various sclerites.
(4) Size and distribution of punctures and tubercles on the various sclerites.
(5) Condition of fronto-clypeal suture.
(l>) Anterior margin of clypeus, whether rounded, truncate, emarginate,
or sinuate ; and also angle on each side .
(7) Condition of anterior margin of labrum and the angle on each side.
(8) Outline of pronotum. Shape and extent of sublateral carina. Extent
and depth of median longitudinal impression if one is present.
(9) Shape of elytral apices.
(10) Condition of lateral margin of elytra (inner margin of epipleura),
whether crenate or smooth,
(n) Number and extent, if any, of carinate intervals.
(12) Condition of epipleura. Form and extent of carina parallel to inner
margin if one is present.
(13) Shape of scutellum.
(14) Condition of prosternal carinae.
(15) Condition of anterior portion of presternum, whether straight or
lobed when viewed laterally.
(16) Extent and depth of mesosternal groove.
(17) Disk of metasternum, whether fiat or not and extent and depth of
the various impressions.
(18) Extent and prominence of carina on each side of metasternal disk.
(19) Length and prominence, if present, of carina of first abdominal sternite.
(20) Shape of fifth abdominal sternite.
(21) Secondary sexual characters.
(22) Structure of male genitalia. No case is known where these structure-,
are the same for two or more species.
The following are the secondary sexual characters that have been noted by
previous authors and those noted by the writer for the first time :
(1) Male with numerous very long, fine, recumbent, golden hairs on each
side of clypeus (formosus).
(2) Male with a dense tuft of long, fine, recumbent, golden hairs on anterior
side of metasternal disk (formosus).
(3) Male with the inner apical fifth of the front tibia swollen (montanus).
(4) Male with the front tarsi dilated (oblongus).
(5) Male with numerous very fine and long hairs on ventral surface of
four basal segments of front tarsi (tarsalis).
(6) Male with the inner apex of the middle tibia acutely produced
(oblongus).
(7) Male with the middle tibiae on inner apical two-fifths swollen and
here also with a few fine teeth (victoriensis, oblongus, sulcicullis).
2Q0
H. E. HINTON
(8) Male with a row of short and stout teeth on inner apical half of
middle tibia (diemenensis, oblongus).
(9) Male with the inner apical spur of middle tibia strongly curved inwards
(diemenensis).
(10) Male with a few large and acute teeth on inner margin of hind
trochanter (sulcicollis).
(n) Male with, on outer apical fourth of hind tibia, a row of long, fine
teeth and posterior to these with a few short and stout teeth
(diemenensis).
(12) Male with a long and acute tooth on inner apical two-fifths of hind
tibia (sulcicollis).
(13) Male with the inner part of apical two-fifths of hind tibia swollen
(tarsalis) .
(14) Male with the shape of the fifth abdominal sternite different to that
of female (formosits).
(15) Male with the surface of the fifth abdominal sternite less strongly
granulate then that of female (formosus).
(ib) Male with the sixth abdominal sternite differently shaped to that of
female (all species).
(17) Male with the last dorsal abdominal tergite differently shaped to
that of female (formosus).
A Key to the North American Species of Austrolimniiis.
1. Elytra without carinate discal intervals. Guatemala. Mexico.
A. formosus (Sharp) (1882).
Elytra with the third discal interval strongly carinate at base . . 2.
2. Pronotum with the median longitudinal channel about as broad at apex
as at base. Panama, Mexico . . . .-1. sulcicollis (Sharp) (1882).
Pronotum with the median longitudinal channel twice as broad on
apical three-fifths as at base (text-fig. 169). Panama
A. curtulus (Sharp) (1882).
169
Text-fig. 169. — Austrolimnius curtulus Sharp.
1 882
nij"
MEXICAN ELMIDAE
Austrolimnius formosus (Sharp).
(Text-figs. 170-180.)
Elmis formosus Sharp, Biol. Centr.-Amer. Col., 1 (2) : 140.
Austrolimnius formosus Hinton, Trans. R. Ent. Soc. Lond..
2QI
85 (18) : 42.1.
Male : Length, 1-3 mm. -1-5 mm. ; breadth, 0-62 111111.-0-75 mm. Cuticle
moderately strongly shining and black ; antennae, mouth-parts, and legs rufo-
piceous. Head without distinct impressions ; surface densely and finely asperate
and without distinct granules or punctures. Clypeus with the fronto-clypeal
suture straight and rather indistinct at middle ; anterior margin truncate to
very feebly rounded for its entire breadth and angle on each side very broadly
Text-fig. 170. — Austrolimnius formosus Sharp.
rounded ; surface asperate as on head and on each side with a dense tuft of fine,
golden-testaceous, recumbent hairs which are about o-io mm. long. Labium
with the anterior margin broadly, arcuately, and moderately deeply emarginate
at middle, and with the angle on each side prominent but obtusely rounded ;
surface slightly more sparsely asperate than that of clypeus ; at sides with
numerous fine hairs which are similar but generally shorter than those of clypeus.
Pronotum across broadest point, which is at basal third, not quite as broad as
long (0-53 mm. : 0-55 mm.) and base broader than apex (0-52 mm. : 0-32 mm.).
Pronotum with the carina and impressipns as figured (text-fig. 170). Surface
asperate as head but more evenly so. Elytra marly twice as long as pronotum
(ro2 mm. :o-55 mm.) and broadest point which is across middle (o-68 mm.) and
very slightly broader than broadest point across humeri. Lateral margins (i.e. inner
margins of epipleura) nearly smooth. Apices conjointly produced and rounded.
Surface without striae ; with three discal rows of seriate puncture's of which the
outer two extend from basal fourth to apical fifth and basally are round, about
292
H. E. HINTON
0-024 nlm- broad, and separated longitudinally by two to three times their
diameters ; towards apex these punctures become slightly denser and coarser ;
inner row of punctures similar to outer rows but not beginning until basal
two-fifths and extending slightly nearer to apex. Surface between punctures
more finely and much more sparsely asperate than that of pronotum. Inner
side of inner sublateral carina with a complete row of coarse, less distinct,
178
Text-figs. 171-180. — A ttstrolimnius formosus Sharp. (171) Dorsal view of male geni-
talia. (172) Left lateral view of same. (173) Female genitalia. (174) Antenna.
(175) Ventral view of fifth abdominal sternite of male. (176) Ventral view of fifth
abdominal sternite of female. (177) Maxillary palp. (178) Apical abdominal
tergite of male. (179) Apical abdominal tergite of female. (180) Sixth abdominal
sternite of male.
and sparser punctures ; between sublateral carinae and between inner margin
of epipleura and outer sublateral carina with punctures which are indistinct
about two to three times as coarse as those of disk, and are separated by once to
twice their diameters. Epipleura from humeri to apical sixth with a row of fine
(about as coarse as facets of eye) close granules parallel and close to inner margin
of epipleura. Scutellum flat, subovate, longer than broad (o-io mm. : 075 mm.),
base broadly rounded, and narrowed to apex and at apex feebly rounded ; surface
asperate as adjacent parts of elytra. Prosternum with the carinae prominent
MEXICAN ELMIDAE 293
and extending nearly to anterior margin ; when viewed laterally the anterior
half (not including process) is moderately but not sharply bent ventrally ; surface
of middle and of process asperate as pronotum, while the surface of the sides and
hypomera is asperate as that of head. Mesosternum with the groove for the
reception of the prosternal process broad and deep but not extending beyond
posterior third ; surface asperate on each side as that of prosternal process.
Metasternum with the median longitudinal line feebly impressed and extending
nearly to anterior fourth ; disk anteriorly nearly flat and with the posterior third
moderately strongly convex ; disk on each side with a prominent carina which
extends posteriorly and slightly outwards from middle coxa nearly to hind coxa ;
surface asperate as disk of elytra and on each side near anterior fourth of discal
carina with a dense tuft of fine, recumbent, golden testaceous hairs which are
about 0-05 mm. long ; sides of metasternum asperate as pronotal disk. Abdomen
with the first sternite without carinae ; surface of sternites one to four asperate
as metasternal disk ; fifth sternite (text-fig. 175) more densely asperate and also
with a few obscure granules. Genitalia as figured (text-figs. 171, 172).
Female : Externally similar to male except as follows : (1) there is no
dense tuft of fine and long hairs on each side of the clypeus ; (2) there is no dense
tuft of fine and long hairs on each side of the metasternal disk near anterior
fourth of discal carina ; (3) the fifth abdominal sternite is shaped differently
(cf. text-figs. 176 and 175) ; (4) the surface of the fifth abdominal sternite is more
densely and distinctly granulate than that of the male, the granules being here
about as coarse as facets of eyes and separated by two to three times their
diameters ; and (5) the last dorsal abdominal segment is differently shaped (cf.
text-figs. 179 and 178).
Type : In the British Museum (Nat. Hist.). Guatemala : Vera Paz, San
Joaquin (Champion).
Specimens examined: 1, with same data as type. 18, Mexico: Dist. de
Temascaltepec, Tejupilco, alt. about 4000 ft., vi.1933 (H. E. Hinton, R. L.
Usinger) ; 1, in the same district, Real de Arriba, alt. about 7000 ft., 28. v. 1933
(H. E. Hinton, R. L. Usinger) ; and 21, Mexico : Estado de Morelos, Cuernavaca,
vi.1934 (H. E. Hinton).
Variations : No variations, apart from absolute size, have been noted.
Comparative notes : This species may be easily distinguished from its only
Mexican congener and the only other North American species of the genus by
the absence of a basal carina on the third discal elytral interval.
Austrolimnius sulcicollis (Sharp).
(Text-figs. 167, 168, 181-188.)
1882. Elmis sulcicollis Sharp, Biol. Centr.-Amer. Col., I (2) : 139, t. 4, f. [6,
Male: Length, 1-3 mm. -1-4 mm. ; breadth, 070 mm. -077 mm. Cuticle
moderately shining and rufo-piceous to black ; antennae, mouth-parts, and legs
paler rufo-piceous. Head without distinct impivs>ions ; surface densely and
very finely asperate and without granules or punctures. Clypeus with the tumid
clypeal suture straight and rather indistinct at middle ; anterior margin truncate
and angle on each side broadly rounded ; surface asperate as head and on
side with a few fine, recumbent , golden-testaceous hairs which are about 0-05 mm.
long. Labrum with the anterior margin broadly and very feebly rounded and
with the angle an each side broadly rounded ; surface slightly more sparsely
2g4
H. E. HINTON
asperate that that of clypeus and on each side with a dense tuft of hairs each of
which is similar to those of clypeus. Pronation with the broadest point, which is
across base and equally across basal third, slightly broader than long (0-55 mm. :
0-50 mm.) and base broader than apex (0-55 mm. : 0-33 mm.). Pronotum with
the carinae and impressions as figured (text-fig. 167). Surface asperate as head
but, particularly on disk, more sparsely and evenly so. Elytra twice as long as
Text-figs. 181-188. — Austrolimnius sulcicoUis (Sharp). (181) Dorsal view of male
genitalia. (182) Left lateral view of same. (183) Last dorsal abdominal segment
of male. (184) Hind leg of male. (185) First abdominal sternite. (186) Fifth
abdominal sternite of male. (187) Middle tibia of male. (188) Antenna.
pronotum (i-oo mm. : 0-50 mm.) and broadest point which is across middle only
slightly broader than broadest point across humeri (070 mm. : 072 mm.).
Lateral margins nearly smooth. Apices broadly produced and conjointly
rounded. Surface without striae but with three discal rows of seriate punc-
tures of which the two outer begin on about basal third and extend to apical
sixth, while the inner row begins at about basal half and extends nearly to apex ;
on disk these punctures are two-thirds as coarse to a third coarser than facets of
eyes and are usually separated by two to three times their diameters ; anterior
MEXICAN ELMIDAE 295
half of inner row with the punctures distinctly finer than corresponding punctures
of outer rows ; surface between punctures finely, indistinctly, and much more
sparsely asperate than those of pronotal disk. Inner side of inner sublateral
carina with a complete row of coarse, less distinct, and more irregularly distributed
punctures ; area between sublateral carinae with punctures about two to three
times as coarse as discal ones, often indistinct, separated by once to twice their
diameters, and near apex partly arranged in two rows. Third discal interval
carinate on basal fifth. Epipleura from basal fourth to apical fifth with a feeble
ridge parallel and very close to inner margin. Scutellum flat, subovate, longer
than broad (o-io mm. : 0-07 mm.), base broadly rounded, and narrowed to and
feebly rounded at apex ; surface asperate as adjacent parts of elytra. Presternum
with the carinae prominent and extending nearly to anterior margin ; when
viewed laterally the anterior half (not including process) is only feebly and gradually
bent ventrally ; surface of middle and of process asperate as disk of elytra but
slightly more densely so, while the surface of sides and hypomera is about as
densely asperate as head. Mesosternum with the groove for the reception of the
prosternal process broad and very deep and not extending to posterior fourth ;
surface asperate as hypomera. Metasternum with the median longitudinal line
feebly impressed and extending nearly to anterior fourth ; disk only very feebly
convex at sides on posterior half ; disk on each side with a fine and feebly raised
carina which extends posteriorly and slightly outwards from middle coxa nearly
to posterior coxa ; surface of disk asperate as disk of elytra ; sides of metasternum
asperate as densely as sides of prosternum. Abdomen without carinae on the
first sternite ; sternites one to four asperate as metasternal disk ; fifth sternite
asperate as hypomera. Legs with middle tibia near apical fourth swollen on
inner side and with a row of fine short teeth (text-fig. 187). Hind trochanter with
an inner row of six stout, short, and acute teeth (text-fig. 184) ; hind tibia on inner
apical two-fifths with a large but short acute tooth. Genitalia as figured (text-figs.
181, 182).
Female : Externally similar to male except as follows : (1) the inner apical
fourth of the middle tibia is not swollen and there is here no row of short and stout
teeth ; (2) the hind trochanter is smooth and not toothed along its inner margin ;
and (3) there is no tooth on the inner apical two-fifths of the hind tibia.
Type : <J in the British Museum (Nat. Hist.). Panama : Volcan de
Chiriqui, alt. 2000-3000 ft. (Champion).
Specimens examined : 5, with same data as type but collected between 4000
and 6000 ft. ; I, Mexico : Dist. de Temascaltepec, Tejupilco, alt. about 4000 ft.,
15. vi. 1933 (H. E. Hinton, R. L. Usinger) ; and 8, at same locality as above but
collected in vii.1934 (H. E. Hinton).
Variations : No variations worthy of mention have been noted in the small
series before me.
Comparative notes : This species can be compared only with .1 . curtulus (Sharp)
of Panama but may easily be distinguished by having the pronotal channel as
broad at apex as at base, whereas in curtulus this channel is twice as broad at
apex as at base.
XENELMIS Hinton.
1936. Xenelmis Hinton, Trans. A'. Ent. Soc. Loud.. 85 (18) : 427.
The following species should be referred to Xenelmis : X. granata (Grouv.)
(Elmis) and A', micros (Grouv.) (Elmis). Five species of this genus are now
296 H. E. HINTON
known, one from Mexico and four from Brazil. A redescription of the genus
follows :
Body broadly oval. Non-tomentose areas clothed with sparse and short
recumbent hairs ; fine scale-like or hairy tomentum confined to the following
areas : (1) pronotum ; (2) epipleura ; (3) hypomera ; (4) sides of presternum,
mesosternum, metasternum, and abdominal sternites, but occasionally the entire
sternum of the abdomen is clothed with scale-like tomentum ; and (5) part of all
of legs except tarsi. Head when seen from below capable of being retracted so
that none of the mouth-parts are visible. Antennae 11-segmented. Mandibles
with three apical blunt teeth ; prostheca entirely membranous and with a few-
fine spines or hairs apically. Maxilla with the palp 4-segmented and stipes with
a well-developed palpifer ; galea and lacinia separate and apex of each with spines.
Labium with the palp 3-segmentedand prementum with a well-developed palpiger.
Mentum transverse and as broad as but only half as long as submentum. Gula
about as broad as submentum, nearly a fourth longer, and with the sides parallel.
Pronotum with the anterior margin strongly arcuate at middle and on each side
behind eye before apical angle deeply and broadly sinuate. Pronotum without
sublateral carinae and disk without a median longitudinal channel ; base
trisinuate, broadly and moderately deeply so on each side and more narrowly
and shallowly so in front of scutellum. Elytra punctate and striate ; each
elytron with two sublateral carinae each of which is formed by a row of close
granules. Hind wing (text-fig. 192) without a radial cross vein or an anal cell ;
first anal absent ; second anal without branches ; third anal only present basally
and at its apex joined to second anal at about apical two-fifths of the latter's
length ; fourth anal well-developed ; and cubito-anal cross vein absent.
Prostemum moderately short in front of anterior coxae ; prosternal process long
and very broad and with the posterior margin broadly rounded. Mesosternum
with a deep and very broad groove for the reception of the prosternal process.
Abdomen with the middle posterior margin of the first sternite very broad and
nearly truncate. Metasternum with a median longitudinal impressed line. Legs
with the visible portion of the front coxae rounded and trochantin completely
concealed by the hypomera and sternum. Claws without teeth. Alimentary
canal (text-fig. 191) without caeca on the anterior margin of the mid-gut. Hind
gut with six Malpighian tubules which end freely near the rectum. Male repro-
ductive system with the lateral accessory glands simple and not lobed. Each
testis with two sperm tubes. Female reproductive system (text-fig. 189) with
at least four egg tubes to each ovary. Spermathecal duct opening into apex
of bursa copulatrix (the duct then continuing in wall of bursa copulatrix so
that it appears actually to open near the uterus). Central nervous system with
three discrete thoracic ganglia. First abdominal ganglion partly fused at base
to third thoracic, second free, third partly fused at apex to terminal ganglion,
and four to eight fused into a single large terminal ganglion.
Genotype : Elmis bufo Sharp (1882).
Only one species, X. tarsalis Hinton, has been available for an examination
of the internal anatomy.
This genus seems to be most closely related to Limnius Erichson, but may be
distinguished as follows : (1) the head is without tomentum ; (2) the disk of the
pronotum is tomentose whereas in Limnius the pronotal tomentum is confined to
the sides between lateral margin and sublateral carina ; (3) the elytra are without
tomentum, whereas in Limnius they are often tomentose to inner sublateral carina ;
(4) there are no sublateral carinae on the pronotum ; (5) the hind wing has the
MEXICAN ELMIDAE 297
second anal extending to inner margin of hind wing opposite junction of cubitus
and media, whereas in Limnius the second anal is present only basally ; (6) the
alimentary canal has no caeca on anterior margin, whereas in Limnius caeca
arc here present ; and (7) the central nervous system has the fourth abdo-
minal ganglion fused to the terminal mass, whereas in Limnius the fourth is
free. It may be distinguished from Xenelmoides Hinton by its broadly oval
instead of subparallel body, the absence of tomentum on the genae, and the
much broader and nearly truncate middle posterior margin of first abdominal
sternite.
The specific characters of greatest importance in separating the species of
Xenelmis appear to be the following:
(1) General proportions, length and breadth.
(2) Colour. In some species the surface of the pronotum is a different
colour to that of the elytra.
(3) Density of the tubercles and punctures on the various sclerites.
(4) Distribution of the alutaceous microsculpture.
(5) Length and shape of carinae, if present, on head.
(6) Condition of fronto-clypeal suture.
(7) Anterior margin of clypeus, whether rounded, truncate, emarginate or
sinuate ; and also shape of angle on each side.
(8) Shape and distribution of rows of granules, if present, on pronotal
disk.
(9) Shape of anterior margin of labrum and that of the angle on each
side.
(10) Condition of lateral margin of elytra (inner margin of epipleura),
whether crenate or smooth.
(11) Number and length of carinate discal intervals, if any.
(12) Shape of scutellum.
(13) Condition of prosternal carinae.
{14) Condition of anterior portion of presternum whether straight or
lobed when viewed laterally.
(15) Shape of prosternal process.
(16) Extent and depth of mesosternal groove.
(17) Disk of metasternum, whether flat or not and the extent and depth
of the various impressions.
(18) Secondary sexual characters.
(19) Structure of the male genitalia. No case is known where these
structures are the same for two or more species.
The following are the secondary sexual characters that have been observed
in the species before me :
(1) Male with a prominent and short longitudinal carina on middle ol
first abdominal sternite near base (micros).
(2) Male with the inner apex of the front tibia with a short and
prominent carina (micros).
(3) Male with the inner apical fourth of the hind tibia moderately
strongly excavated and distal margin of this concave portion with
a small but prominent carina (micros).
(4) Male with numerous erect, fine, long, and pale hairs on ventral surface
of lour basal segments of middle and hind tarsi (micros).
(5) Male with the last segment of the hind tarsus strongly dilated and
ventral surface of dilated part densely pubescent (tarsal is).
novit. zool., 42, 2. 1940. 24
298
H. E. HINTON
Xenelmis bufo (Sharp).
(Text-figs. 189-198.)
1882. Elmis bufo Sharp, Biol. Centr.-Amer. Col., 1 (2) : 140.
1936. Xenelmis bufo Hinton, Ent. mon. Mag., 72 : 4, fig. 7.
1936. Xenelmis bufo Hinton, Trans. R. Ent. Soc. Land.. 85 (18) : 427 figs. 23 24 pi 1
fig- 3- '
Text-figs. 189-192.— (189) Female reproductive system of Xenelmis tarsalis Hinton.
(190) Adult of Xenelmis bufo (Sharp). (191) Alimentary canal of Xenelmis tarsalis
Hinton. (192) Hind wing of X. bufo (Sharp). Venation after Forbes.
Male : Length, 142 mm.-i-fa mm. ; breadth, 0-87 mm.-o-95 mm. Cuticle
feebly shining and rufo-piceous to black. Head on each side with a fine, low
carina which extends from a point which is about a third of the way across between
eyes, and is opposite posterior fourth of eyes, to posterior seventh of head ; these
carinae converge posteriorly but do not meet so that at posterior end they are
separated by a distance equal to half that of anterior end. Surface of head with
MEXICAN ELMIDAE
209
round, high granules which are about as coarse as facets of eyes or about o-oio mm .
broad and are usually separated by one to three times their diameters ; each of
these granules gives rise to a fine recumbent seta which is about twice as long as
its respective granule ; surface between granules densely and microscopically
alutaceous. Clypeus with the fronto-clypeal suture straight and deep ; anterior
margin very feebly rounded to nearly truncate and with the angle on each side
broadly rounded ; surface sculptured as head. Labrum with the anterior margin
truncate and with the angle on each side broadly rounded ; surface on basal half
smooth, on apical half with a narrow transverse belt of obscure, dense punctures,
and on each side with a few very fine, testaceous hairs which are often about
197
Text-figs. 193-198. — Xenelmis bufo (Sharp). (193) Dorsal view of male genitalia.
(194) Left lateral view of same. (195) Antenna. (196) Maxillary palp. (197)
Female genitalia. (198) Presternum.
0-05 mm. long. Pronotum across broadest point, which is at base, broader than
long (072 mm. : 0-55 mm.) and base broader than apex (072 mm. : 035 mm.) ;
evenly convex and surface with granules similar to those of head but usually
separated by two to three times their diameters ; area between granules much
more sparsely alutaceous ; at base in front of scutellum for a short distance
(about a seventh of the length of the pronotum) there are no granules. Elytra
slightly more than twice as long as pronotum (1-12 mm. : 0-55 mm.) and broadest
point, which is across middle, broader than broadest point across humeri (o-gomm. :
0-82 mm.). Lateral margins finely and regularly crenatc, these crenations being
due to laterally placed granules. Apices broadly conjointly produced and
rounded. Surface with the striae as broad as strial punctures and very feebly
impressed ; discal strial punctures round to subquadrate, about a third to two-
thirds as coarse as intervals, and separated longitudinally by one to one and a
half times their diameters ; these punctures become finer and denser towards
300
H. E. HINTON
apex and coarser and denser towards sides. Intervals flat and with the surface
sculptured as that of head except that here the granules are slightly coarser,
especially towards sides. Granules of sublateral carina about a fourth to a third
coarser than those elsewhere on elytra. Scutellum flat, subovate, broader than
long (0-15 mm. : o-io mm.), base very broadly and feebly rounded, and narrowed
to and acutely rounded at apex ; surface sculptured as adjacent parts of elytra.
Prosternum without distinct carinae ; when viewed laterally the anterior half
(not including process) is very feebly bent ventrally and the remainder of the
prosternum and its process is on the same plane ; process shaped as figured
(text-fig. 198) ; surface of all of prosternum and hypomera sculptured as disk of
elytra but with the granules slightly more distinct. Mesosternum with the groove
for the reception of the prosternal process a third broader posteriorly than anteriorly
and extending to posterior fifth of mesosternum ; surface at sides sculptured as
disk of elytra but without distinct granules. Metasternum with the median
longitudinal line feebly impressed and extending to anterior fifth or sixth ; disk
nearly flat but along anterior margin moderately depressed at middle and more
strongly so at sides ; surface of disk with irregular to round, large (0-03 mm.
broad), frequently confluent punctures and with only an occasional granule ;
between punctures and granules the surface is moderately densely microscopically
alutaceous ; sides of metasternum with the granules similar in size and distribution
to those of sides of elytra but otherwise sculptured as metasternal disk except
that the punctures are not as dense. Abdomen with the surface of sternites one
to four sculptured as disk of elytra and the surface of five similarly sculptured but
with the granules distinctly denser. Genitalia as figured (text-figs. 193, 194).
Female : Externally similar to male.
Type: In the British Museum (Nat. Hist.). Panama: Volcan de
Chiriqui, alt. 2600-3000 ft. (Champion).
Specimens examined: 1, Panama: San Miguel, Pearl Island (Champion).
30, Mexico : Dist. de Temascaltepec, Tejupilco, alt. about 4000 ft., vi.1933
(H. E. Hinton, R. L. Usinger) ; and 18, in the same locality in vii.1934 (H. E.
Hint on).
Variations : Except for the differences in absolute size as already noted, no
variations worthy of mention have been observed.
Comparative notes : This species is apparently most closely related to the
Brazilian X. tarsalis Hinton, but it may be distinguished from the latter by not
having a row of close granules on the basal half of the third discal elytral interval.
MICROCYLLOEPUS (Hinton).
1935. Microcylloepus Hinton, Stylops, 4 (8) : 178.
In addition to the species referred to this genus by Hinton (1935) , the following
belong here : M. thermarum (Darlington) (Elmis) and M. longipes (Grouvelle)
(Elmis). At the time of writing seven species and three varieties have been
described. In my collection there are 13 new species, and of these three are from
Mexico and will be described here. The species of this genus occur from Canada
to the south of Brazil and have been taken in the British West Indies on Trinidad
and Tobago. A redescription of the genus follows :
Body subparallel. Non-tomentose areas glabrous or clothed with sparse
and short recumbent hairs. Scale-like or hairy tomentum confined to the follow-
ing areas : (1) genae ; (2) epipleura — -many species have non-tomentose epi-
pleura ; (3) sides of prosternum, mesosternum, metasternum, and abdominal
MEXICAN EI.MIDAK
301
sternites.but in some species nearly the entire sternum of the abdomen is clothed
with fine tomentum ; and (4) part or all of legs except tarsi. Head when seen
from below capable of being retracted so that none of the mouth-parts is visible-
Antennae 11-segmented (text-fig. 218). Mandibles with three acute apical teeth;
prostheca long and entirely membranous with numerous fine spines or hairs
apically. Maxilla with the palp (text-fig. 219) 4-segmented and stipes with a
well-developed palpifer ; galea and lacinia separate and apex of each spinose or
hairy. Labium with the palp 3-segmented and prementum with a poorly developed
Text-figs. 199—201. — Microcylloepus carinatus Hinton.
accessory glands. (200) Male reproductive system.
system.
(199) Lateral view of lateral
(201) Female reproductive
palpiger. Mentum as broad and three-fifths as long as submentum. Gula
slightly longer than submentum, at anterior margin about two-thirds as broad as
submentum, and with the sides feebly converging posteriorly. Pronotum with the
anterior margin moderately arcuate at middle and on each side behind eye before
apical angle broadly and moderately deeply sinuate. Base trisinuate, broadly
and moderately deeply so on each side and more narrowly and shallowly SO in
front of scutellum. Pronotum on each side with a sublateral carina which
extends from base nearly to anterior margin ; at apical two-fifths with a broad,
moderately deep and complete or incomplete transverse impression which is
always distinct at sides but in some species it is not evident on discal region ;
disk with or without a broad and deep median longitudinal impression ;
beginning either at base or at basal third on each side in front of scutellum is a
302
H. E. HINTON
broad, deep or shallow impression which extends obliquely forwards, crosses
sublateral carina at basal two-fifths, and joins apical transverse impression near
lateral margin ; sometimes this oblique impression is confined to sides ; with or
without a median longitudinal, feebly convex to carinate line extending from base
in front of scutellum to posterior margin of longitudinal discal impression. Elytra
punctate and striate ; each elytron with a sublateral carina on eighth interval
and one on sixth, though in a few species the carina of the sixth interval is absent
or very indistinct. Hind wing without an anal lobe ; without a radial cross vein
Text-figs, 202
202
203. — MicYocylloepus carinatus Hinton. (202)
(203) Dorsal view of central nervous system.
203
Alimentary canal.
or an anal cell ; first anal absent ; second anal with the first branch present ;
second branch of second anal absent ; third anal with the second branch absent ;
fourth anal well-developed ; and cubito-anal cross vein present but incomplete
and joining cubitus to second anal. Prostemum very long in front of anterior
coxae ; prosternal process long, narrow to moderately narrow, and with the
posterior margin broadly rounded to nearly truncate. Mesosternum with a broad
and deep groove for the reception of the prosternal process. Metasternum with
a median longitudinal impressed line. Legs with the visible portion of the front
coxae rounded and trochantin completely concealed by the hypomera and sternum.
Claws without teeth. Alimentary canal (text-fig. 202) with two caeca on the
anterior margin of the mid-gut. Hind gut with six Malpighian tubules which
end freely near the rectum. Male reproductive system (text-fig. 200) with the
lateral accessory glands lobed. Each testis with two sperm tubes. Female
MEXICAN ELMIDAE 303
reproductive system (text-fig. 201) with six egg tubes to each ovary. Spermathecal
duct opening into apex of bursa copulatrix. Central nervous system (text-fig. 203)
with three thoracic discrete ganglia ; first three abdominal ganglia discrete, and
four to eight partly fused together, though the limits of each are distinguishable.
Genotype: Elmis pusilla Leconte (1852).
The internal anatomy of three species has been examined and found to agree
in essential details. That of .1/. carinatus Hinton is figured.
The species of Microcylloepus frequently bear a close resemblance to those of
Cylloepus Erichson (sensti stricto), but may be distinguished as follows : (1) the
pronotum has, at least at sides, a distinct apical transverse impression on apical
two-fifths, whereas in Cylloepus no such impression is present ; (2) the hind wing
has the second branch of the third anal absent, whereas in Cylloepus it is present ;
(3) the alimentary canal has two instead of eight caeca on the anterior margin of
the mid-gut; (4) each ovary has six instead of about 18 egg tubes; (5) the
spermathecal duct opens into the apex of the bursa copulatrix instead of into the
base ; and (6) the central nervous system has the first three abdominal ganglia
discrete, whereas in Cylloepus two to five are discrete.
A list of the characters which have been found useful in distinguishing the
species is as follows :
(1) General proportions, length and breadth.
(2) Colour.
(3) Size and distribution of the punctures and tubercles on the various
sclerites and the density and type of microsculpture between
them.
(4) Condition of the fronto-clypeal suture.
(5) Condition of anterior margin of clypeus, whether rounded, truncate,
emarginate, or sinuate ; and also the condition of the angle on each
side.
(6) Condition of the anterior margin of the labrum and the angle on each
side.
(7) Outline of pronotum and general proportions.
(8) Extent and depth of the various pronotal impressions.
(9) Shape of elytral apices.
(10) Absence or presence of tomentum on epipleura.
(11) Prominence and extent of the inner sublateral carina of the elytra.
(12) If third discal interval is carinate at base the prominence and extent
of the carina should be given.
(13) Condition of other elytral intervals, whether liat, gibbous, or carinate.
(14) Shape of scutellum and whether convex or flat.
(15) Condition of anterior portion of prosternum, whether straight or bent
ventrally when seen from the side.
(16) Extent of prosternal carinae.
(17) Shape of prosternal process.
(18) Extent and depth of mesosternal groove.
(19) Condition of disk of metasternum, whether flat or not, and the
extent and depth of the various impressions.
(20) Length, shape, and prominence of carina on each side of metasternal
disk.
(21) Length, shape, and prominence of carina on each side of middle of
first abdominal stemite.
(22) Secondary sexual characters.
304 H. E. HINTON"
(23) Structure of the male genitalia. This is one of the very few genera
in the family in which a number of species may have the struc-
ture of the male genitalia identical.
The following secondary sexual characters have been observed in the species
before me :
(1) Female with the sutural elytral interval strongly convex on apical third,
while that of the male is only feebly convex (carinatus).
(2) Female with the apex of each elytron produced to form an acute
tooth, while in the male the apices are broadly and conjointly
rounded (carinatus) .
(3) Male with the sides of the mesosternum between coxae densely clothed
with fine testaceous hairs, while in the female this part is glabrous or
very nearly so (spinipes).
(4) Male with the disk of the metasternum more deeply and extensively
concave than that of female (carinatus, obesus, spinipes).
(5) Female with the metasternal disk more densely punctate than that
of male (obesus).
(6) Male with the carina on each side of metasternal disk more prominent
that than of female (obesus).
(7) Female with the region near the apex of the fifth abdominal sternite
moderately gibbous (carinatus, spinipes).
(8) Male with the outer edge of the front and middle tibiae gibbous on
basal two-fifths (plaumani).
\ (9) Male with an acute tooth concealed in a tuft of erect, testaceous hairs
on inner apex of all tibiae (carinatus).
(10) Male with the ventral apex of the basal segment of front tarsi strongly
and acutely produced and at apex of produced portion with a long
and acute spine (spinipes).
(11) Male with the ventral mesal surface of the hind coxae with numerous
recumbent, testaceous hairs which are here much sparser or absent
in the female (spinipes).
(12) Male with the inner apical spur of the hind tibiae very long, slender,
and slightly curved forwards (spinipes).
A Key to the Mexican Species of Microcylloepus.
1. Scutellum strongly convex. Mexico . . . M. obesus, sp. n.
Scutellum flat, rarely very feebly convex . . . . . .2.
2. Humeri narrow, greatest breadth across humeri less or only very little
more than greatest breadth across pronotum. Elytra with the
punctures of the first and second striae on basal third broader than
intervals. Mexico ....... M. angustus, sp. n.
Humeri broad, always distinctly broader than greatest breadth across
pronotum. Elytra with the punctures of the first and second striae
on basal third seldom as broad as intervals ..... 3.
3. Small species with the mean length from apex of elytra to middle anterior
margin of pronotum 1772 mm. (altitude, 5600-7500 ft.). Elytra
with the third discal interval strongly convex to carinate on basal
fifth or sixth ; inner sublateral carinae usually as prominent as outer.
Guatemala, Mexico .... M. inaequalis (Sharp) (1882).
MEXICAN ELMIDAE 305
Large species with the mean length from apex of elytra to middle
anterior margin of pronotum 2-003 mm. (altitude, 5600-7500 ft.).
Elytra with the third discal interval at most only very feebly convex
on basal seventh ; inner sublateraJ carinae never as prominent as
outer and frequently so flat that they are scarcely noticeable.
Mexico ........ M. troihis, sp. n.
Microcylloepus obesus sp. n.
(Text-figs. 207-208.)
Male: Length, 2-215 mm. -2-415 mm.; breadth i-oo mm.-i-io mm.
Cuticle shining and rufo-piceous to black ; antennae, mouth-parts and legs paler
rufo-piceous. Head without distinct impressions ; surface with round granules
which are about 0-017 mm- broad and are usually separated by one to three times
their diameters ; surface between granules with round to irregularly shaped
microscopic punctures (about o-ooi mm. broad) which are confluent to separated
by twice their diameters — under low magnifications ( x 70) the surface does not
appear punctate but densely and microscopically alutaceous. Clypeus with the
fronto-clypeal suture well impressed and nearly straight ; anterior margin when
seen from in front shallowlyand arcuatelyemarginate for its entire breadth ; angle
on each side broadly rounded ; surface sculptured as head. Labrum with the
anterior margin feebly rounded and the angle on each side broadly rounded ;
surface without granules, basally and at sides with fine punctures similar in size
and distribution to those of clypeus, apical half at middle with the punctures
usually twice as large and half as dense and apical sides with numerous fine,
testaceous hairs which are about 0-05 mm. long. Pronotum across broadest point,
which is at basal third, as broad as long (0-70 mm.) and base broader than apex
(0-63 mm. : 0-48 mm.). Sides deeply and broadly sinuate opposite apical
transverse impression and very feebly sinuate to nearly straight before basal
angles. Sublateral carina extending from base very nearly to apex. Impressions
as figured for inaequalis (text-fig. 209) but without a longitudinal ridge from base
to posterior margin of longitudinal discal impression. Disk of pronotum very
densely, frequently confluently punctate with punctures which are from o-oo6mm.
to 0-02 mm. broad except as follows : on the middle portion of the anterior third
the punctures are usually separated by three to five times their diameters ; and
at sides near sublateral carinae on basal fourth the punctures are separated by
one to three times their diameters. Sides between sublateral carinae and lateral
margins sculptured as head. Elytra more than twice as long as pronotum (1-72
mm. : 0-70 mm.) and at broadest point, which is across apical third, broader than
broadest point across humeri (1-07 mm. : 1-05 mm.). Epipleura without
tomentum. Lateral margins moderately feebly and irregularly crenate. Apices
broadly and strongly produced and conjointly feebly rounded. Surface with the
striae moderately strongly impressed basally and feebly impressed on apical
third ; strial punctures on basal third of disk are round to feebly subquadrate,
deep, about as broad as intervals and separated longitudinally by less than to once
their diameters ; from basal third to apex these punctures gradually become
finer so that near apex they are half as coarse as at base, half to two-thirds as
broad as intervals, and separated longitudinally by two to three times their
diameters. Intervals flat except for third which is very feebly convex for a short
distance behind level of scutellum ; surface with an occasional microscopic
puncture and sparsely transversely alutaceous but on base at sides and on basal
3o6
H. E. HINTON
third between lateral margins and sublateral carinae with round granules similar
to those of sides of pronotum.and which on gibbous portion of humeri are separated
by about their diameters, while elsewhere they are sparser. Inner sublateral
carinae less prominent than outer. Sc iridium moderately strongly convex,
subovate, longer than broad (0-12 mm. : o-io mm.), base feebly rounded, and
apex more strongly and narrowly rounded ; surface sculptured as adjacent
elytral intervals. Prosternum when seen from the side with the anterior four-fifths
(not including process) gradually to moderately sharply and moderately strongly
2 04
206
207
Text-figs. 204-208. — (204) Dorsal view of male genitalia of Microcylloepus angtislus
Hinton. (205) Right lateral view of same. (206) Left paramere of same species.
(207) Dorsal view of male genitalia of M. obesus Hinton. (208) Left paramere of
same.
bent ventrally ; prosternal carinae prominent, present on basal fourth, feebly
diverging anteriorly, and posteriorly as broad as process at base ; process
moderately narrow, with the sides converging to apex and with the apex broadly
rounded ; surface for the most part densely asperate. Hypomera with the
surface densely asperate and with an occasional granule. Mesosternum
with the anterior two-fifths of the groove for the reception of the prosternal
process broad, deep, and smooth-bottomed ; on posterior three-fifths it is deeper
but only half as broad, extends nearly to metasternum, and the bottom is densely
asperate ; mesosternum between coxae moderately depressed and surface densely
microscopically punctate to asperate. Metasternum with the median longitudinal
line complete ; disk with the posterior four-fifths very broadly and deeply
depressed ; surface with punctures about 0-017 mm. broad which are separated
MEXICAN ELMIDAE 307
by one to four times their diameters, and at posterior third on each side near
discal carina with a broadly oval (0-07 mm.), moderately deep depression ;
discal carinae prominent, strongly diverging posteriorly, and extending to
posterior fifth of rnetasternum. Abdomen with the carina of the first sternite
prominent, feebly diverging posteriorly, and extending to posterior margin of the
segment ; middle area of first sternite between carinae only feebly depressed
anteriorly ; surface of middle of sternites one to four with punctures which are
usually separated by once to twice their diameters ; surface of sides of first four
sternites and all of anterior half of fifth very densely and microscopically punctate ;
apical half of fifth similarly punctate but also with round granules about 0-015 rnm.
broad which are separated by less than to twice or more times their diameters.
Genitalia as figured (text-figs. 207, 208).
Female : Externally similar to male except as follows : (1) the metasternal
disk is not as deeply and only about three-fifths as broadly depressed ; (2) the
metasternal disk has the punctures much denser, being seldom separated by more
than their diameters and frequently confluent ; and (3) the metasternal discal
carina are less prominent.
Type : $ in the British Museum (Nat. Hist.). Mexico : Dist. de Tema-
scaltepec, Rio Verde, alt. 8000 ft., 14. vi. 1934 (H. E. Hinton).
Paratypes : 3, with same data as type.
Variations : One of the two females before me has the metasternal discal
impression nearly as broad and deep as that of male but confined to posterior
half of disk, and the sides of disk near impression are feebly but distinctly convex,
whereas those of the female mentioned above are flat.
Comparative notes : This is the largest known species of Microcylloepus.
From all others it may be immediately distinguished by its moderately strongly
convex instead of flat scutellum and by the structure of the male genitalia.
Microcylloepus angustus, sp. n.
(Text-figs. 204-206.)
Male : Length, 1-42 mm. -1-67 mm. ; breadth, 0-55 mm. -0-67 mm. Cuticle
shining and rufo-piceous to black ; antennae, mouth-parts, and legs paler rufo-
piceous. Head without distinct impressions ; surface, particularly at sides, with
an occasional round granule which is about 0-012 mm. broad or slightly finer
than an eye facet ; surface everywhere densely microscopically alutaceous, this
microsculpture sometimes appearing asperate. Clypeus with the fronto-clypeal
suture deeply impressed and straight ; anterior margin when seen from in front
shallowly and arcuately emarginate for its entire breadth ; angle on each side
broadly rounded ; surface sculptured similarly to head and at sides also with an
occasional obscure granule. Labrum with the anterior margin feebly rounded
and the angle on each side broadly rounded ; surface without granules, only
sparsely and very finely alutaceous, with numerous very fine (about 0-004 nlm-
broad) punctures, and at anterior sides with numerous fine, testaceous hairs which
are about 0-05 mm. long. Pronotum across broadest point, which is at about
basal half, broader than long (o-6i mm. : 0-57 mm.) and base broader than apex
(0-50 mm. : 0-42 mm.). Sides moderately deeply and broadly sinuate opposite
apical transverse impression and scarcely noticeably sinuate before basal angles.
Sublateral carina extending from base very nearly to apex. The impressions are
similar to those figured for inaequalis (text-fig. 209). Disk of pronotum very
densely, frequently confidently, punctate with punctures which vary from o-oob
308 H. E. HINTON
mm. to 0-012 mm., except as follows : on the middle portion of anterior third the
punctures are often separated by as much as three times their diameters ; and on
basal fourth on each side of median ridge the punctures are separated by once to
twice their diameters. Sides between sublateral carinae and lateral margins as
densely punctate as middle of disk and also with round, obscure granules which
are as coarse as facets of eyes and are separated by one to three times their diameters.
Elytra twice as long as pronotum (i-tz mm. : 0-57 mm.) and at broadest point,
which is across apical two-fifths, broader than broadest point across humeri
(0-675 mm. : o-6o mm.). Epipleura without tomentum. Lateral margins
regularly and moderately strongly crenate. Apices broadly, strongly produced
and conjointly feebly rounded. Surface with the striae moderately impressed
basally and more feebly impressed towards apex ; strial punctures on basal third
of disk round to nearly subquadrate, moderately deep, usually about a third
broader than intervals, and separated longitudinally by less than their diameters ;
from about basal third to apex these punctures gradually become finer and sparser
so that at apex they are two-thirds to half as coarse and dense as at base.
Intervals flat except for third which is prominently carinate on basal sixth ;
surface microscopically punctate and sparsely, transversely alutaceous but on
base and on sides on basal third between lateral margins and inner sublateral
carinae with round granules similar to those of sides of pronotum but somewhat
sparser. Inner sublateral carina prominent and extending slightly beyond basal
half. Scutellum at most very feebly convex, subovate, longer than broad (0-08
mm. : 0-06 mm.), base very feebly rounded, and apex more strongly and much
more narrowly rounded ; surface sculptured like adjacent elytral intervals.
Prosternum when seen from the side with the anterior half (not including process)
gradually and moderately strongly bent ventrally ; prosternal carinae prominent,
present on basal three-fifths, diverging anteriorly, and posteriorly slightly broader
than process at base ; process moderately narrow with the sides nearly parallel
and the apex broadly rounded ; lateral margins of process strongly gibbous ;
surface as densely punctate as disk of pronotum, but on posterior sides and on
process frequently asperate. Hypomera with the surface densely asperate
and with occasional granules similar to those of sides of head. Mesostemum
with the anterior two-fifths of the groove for the reception of the prosternal
process broad, deep, and with the bottom smooth, while posterior three-fifths is
deeper, only a third as broad, extends nearly to metasternum, and the bottom is
asperate ; mesostemum between coxae moderately depressed and surface densely
asperate. Metasternum with the median longitudinal line complete ; disk on
anterior third moderately gibbous, but on posterior half broadly (0-07 mm.) and
deeply depressed on middle and with a deep pit at posterior margin ; on posterior
third on each side adjacent to discal carina is a broad (about 0-06 mm.), oval,
moderately shallow depression ; surface on middle of disk (except for bottom of
posterior depression which is microscopically asperate) with only an occasional
microscopic puncture and sparsely transversely alutaceous, while the surface
elsewhere is densely asperate as are sides of metasternum ; discal carinae
prominent, feebly diverging posteriorly, and extending to posterior third. Abdo-
men with the carinae of the first sternite prominent, moderately diverging
posteriorly, and extending to the posterior margin of the segment ; middle area
of first sternite between carinae scarcely depressed anteriorly ; surface of middle
of sternites one to four and extreme basal middle of fifth non-tomentose and with
microscopic punctures which are usually separated by two to four times their
diameters ; surface of sides of first four and most of fifth finely asperate when
MEXICAN ELMIDAE \(>q
the tomentum is rubbed off ; apical third of fifth sternite with round granules
about o-oio mm. broad which are separated by once to twice their diameters.
Genitalia as figured (text-figs. 204-206).
Female : Externally similar to male.
Type: rj in the British Museum (Nat. Hist.). Mexico: Dist. de Tema-
scaltepec, Temascaltepec, alt. 5600 ft., vii.i934(//. E. Hinton).
Paratypes : 16, with same data as type ; and I, also with same data but
collected on 15.vii.1933 (H. E. Hinton, R. L. Usinger).
Variations : In some specimens the middle raised portions of the pronotal
disk are not confluently punctate, the punctures here being separated by twice
their diameters. A few have the surface of the ridge, which extends from base
in front of scutellum to posterior margin of median longitudinal impression, with
a few obscure granules. In many specimens the strial punctures on basal third
of disk of elytra are occasionally longitudinally and laterally confluent. In some
specimens the frontal region of the head as well as the sides has a number of
round granules. In a few the middle portion of the metasternal disk has the
microscopic punctures separated by once to twice their diameters, but they are
here never dense enough to give the surface an asperate appearance.
Four specimens have been examined for the condition of the hind wings
and all found to have them greatly reduced, the apex barely extending to a point
opposite the posterior margin of the first abdominal sternite. As is the case with
most beetles having the hind wings reduced, the humeri of this species are greatly
narrowed.
Comparative notes : This species most nearly resembles M. thermarum (Darling-
ton) of Oregon, but may be distinguished by having the inner sublateral carinae of
the elytra moderately prominent and extending to apical half, whereas in thermarum
the inner sublateral carinae are absent or scarcely visible. Of the Mexican species it
can only be compared with .1/. inaequalis (Sharp) , and from this may be distinguished
by the structure of the male genitalia (cf. text-figs.), the narrow instead of broad
humeri and the discal strial punctures on basal third of elytra which in angustns
are generally broader than intervals, whereas in inaequalis they are seldom as
broad as intervals. .1/. angustns is also a much shorter species {vide Table I).
The means of the lengths of these two species were found to be significantly
M — M.,
different. Using the formula t = . — — = — ■ — = = .where M is the mean
8 •(S.E.M.,)2 4 (S.E.M..,)2
and S.E.M. is the standard error of the mean, t = 10. As may be seen from
Fisher's (1936) table of t, the probability, P, of these two samples being got from
the same populations is P < -oi. Or using Fisher's table of x, which is in effect a
continuation of t for numbers over 30, a better idea is got of the improbability of
the two populations being homogeneous for length, as the probability heir !->
P< io~9.
Table 1.
Lengths are ^ivcii in mm., and are taken from the apex oi the elytra to the middle anterior
margin of the pronotum. S.D. equals standard deviation ; S.K.I), equals standard error of
the deviation ; and S.E.M, equals standard error of the mean.
310 H. E. HINTON
Microcylloepus inaequalis (Sharp).
(Figs. 209-213.)
1882. Elmis inaequalis Sharp, Biol. Centr.-Amer. Col.. 1 (2) : 137.
1888. Elmis inaequalis Grouvelle, Ann. Soc. Ent. Fr.. 8 (6) : 408, t. 8, f. 11.
1934. Limnius mexicanus Hinton, Rev. Ent., Rio de ]., 4 (2) : 199 (syn. n.).
1935. Microcylloepus inaequalis Hinton, Sty lops, 4 (8) : 179.
1935. Microcylloepus mexicanus Hinton, Stylops, 4 (8) : 179.
Male : Length, 1-47 mm-i-a/) mm. ; breadth, 0-92 mm.-o-6o mm. Cuticle
shining and rufo-piceous to black ; antennae, mouth-parts and legs usually paler
rufo-piceous. Head without distinct impressions ; surface with an occasional
round granule which is about 0-012 mm. broad or slightly coarser than a facet of
an eye, these granules being more numerous at sides where they are separated
by twice their diameters ; surface between granules densely, microscopically
alutaceous, this microsculpture sometimes appearing asperate. Clypeus with
the fronto-clypeal suture deeply impressed and straight ; anterior margin when
seen from in front shallowly and arcuately emarginate for its entire breadth ;
angle on each side broadly rounded ; surface sculptured similarly to head but
even at middle with the granules separated by twice their diameters. Labrum
with the anterior margin feebly rounded and the angle on each side broadly
rounded ; surface moderately densely, microscopically alutaceous and also with
a number of microscopic punctures, and at anterior sides with numerous fine,
testaceous hairs which are about 0075 mm. long. Pronotum across broadest
point, which is at basal two-fifths, broader than long (o-6o mm. : 0-55 mm.) and
base broader than apex (0-55 mm. : 0-42 mm.). Sides broadly and moderately
deeply sinuate opposite apical transverse impression and scarcely noticeably
sinuate before basal angles. Sublateral carina extending from base very nearly
to apex. Impressions as figured (text-fig. 209). Disk of pronotum between
sublateral carinae with an asperate type of alutaceous microsculpture like that
of head on bottom of the impressions ; convex portions with microscopic (0-006
mm. broad) punctures which are usually separated by two to three times their
diameters ; surface of median longitudinal ridge which extends from base to
posterior margin of median impression very densely and microscopically punctate
so that it nearly appears asperate. Sides between sublateral carinae and lateral
margins sculptured as head but with the granules very slightly coarser and
separated by one to four- times their diameters. Elytra more than twice as long
as pronotum (1-20 mm. : 0-55 mm.) and at broadest point, which is across apical
third, broader than broadest point across humeri (0-82 mm. : 072 mm.). Epi-
pleura without tomentum. Lateral margins regularly, moderately strongly
crenate. Apices broadly, strongly produced and conjointly feebly rounded.
Surface with the striae moderately impressed basally and more feebly impressed
towards apex, so that on apical third of discal region the striae are practically
absent ; strial punctures on basal third of disk round, deep, usually about one-
half to two-thirds as broad as intervals, and separated longitudinally by one-half
to one times their diameters ; from basal two-fifths to apex these punctures become
finer and sparser so that near apex they are two-thirds as coarse and dense as
those near base. Inner sublateral carinae prominent and extending to apical
third of elytra. Intervals flat except for third which is strongly convex and
diverging outwards on basal seventh ; surface of intervals microscopically
punctate and sparsely, transversely alutaceous, but on base and on basal third
at sides between lateral margins and inner sublateral carinae with round granules
MEXICAN EI.MIDAE
311
which are similar to those of sides of pronotum but are usually slightly sparser.
Scutellum flat, subovate, longer than broad (o-io mm. : 007 mm.), base very
feebly rounded, and apex more strongly and much more narrowly rounded ;
surface sculptured as adjacent elytral intervals but with the granules slightly
denser. Prostemum when seen from the side with the anterior three-fourths
Text-1'igs. 209, 210. — Microcylloepus inaequalis (Sharp). (209) Adult to show general
appearance. (210) Hind wing. Venation after Forbes.
(not including process) gradually and moderately strongly flexed ventrally ;
prosternal carinae prominent, present on basal two-thirds, feebly diverging
anteriorly, and posteriorly slightly broader than process at base ; process
moderately narrow and with the moderately strongly gibbous sides feebly
converging to apex which is broadly rounded ; surface for the most part sculptured
similarly to that of head. Hypomera densely and distinctly asperate and with
a few obscure granules similar in size to those of head. Mesosternum with the
312
H. E. HINTON
groove for the reception of the prosternal process extending nearly to metasternum;
with the anterior three-fifths of this groove broad, deep, and smooth bottomed,
and with the posterior two-fifths only a third as broad, deeper, and with the bottom
densely asperate ; most of the mesosternum between coxae depressed and with the
surface densely asperate. Metasternum with the median longitudinal line complete ;
disk on anterior third moderately convex but on posterior two-thirds moderately
deeply and broadly depressed, and with a deep narrow pit on posterior margin ;
disk on posterior third or half adjacent to discal carina on each side with a broad
(about 0-075 mm.), oval, moderately deep depression ; surface on middle of disk
(except bottom of median longitudinal line which is densely alutaceous) sparsely,
transversely alutaceous and also with a few microscopic punctures ; surface of
211 212
Text-figs. 211-113. — Microcylloepus inaequalis (Sharp). (211) Dorsal view of male
genitalia. (212) Left lateral view of same. (213) Left lateral view of paramere.
disk elsewhere densely asperate as well as non-tomentose parts of sides of meta-
sternum ; discal carinae moderately prominent and diverging outwards and
extending to posterior third. Abdomen with the carinae of the first sternite
prominent, feebly diverging posteriorly, and extending to posterior margin of the
segment ; middle area of first sternite between carinae scarcely depressed
anteriorly ; surface of middle of sternites one to four and extreme basal middle
of fifth with punctures which are about o-oio mm. broad and are usually separated
by once to twice their diameters ; surface of sides of first four and most of fifth
tomentose but if the tomentum is rubbed off the surface is densely, microscopically
alutaceous ; apical third or fourth of fifth with round granules about 0-012 mm.
broad which are separated by less than to once their diameters. Genitalia as
figured (text-figs. 211-213).
Female : Externally similar to male.
Type : In the British Museum (Nat. Hist.). Guatemala : Guatemala City,
alt. 5000 ft. (G. C. Champion).
Specimens examined: 23, with same data as type ; and 10, Guatemala: San
Joaquin (G. C. Champion). The following 2926 specimens were collected in
Mexico : 4, Atlisco {Flohr, F. D. Godman) ; I, Vera Cruz (Koebele) ; 2071, Estado
MEXICAN klmihu
313
do Morelos, Cuernavaca, alt. 4800 ft., vi.1934 (H. E. Hinton) ; and 850, Dist. de
Temascaltepec, Tejupilco to Real de Arriba, alt. 3500-7500 ft., v-vii.1933 and
1034 (J933i H- E- Hinton, R. L. Usinger) (ig.',4, H. li. Hinton).
Variations : This species is the most variable member of the genus that has
been examined. The most striking variations are those concerning the sculpture
of various sclerites, particularly the pronotum and the disk of the elytra. The
convex parts of the pronotal disk are typically as described above, but in some
specimens all of the disk except anterior third, which is densely punctate, is con-
fluently punctate with punctures which are usually about 0-007 mm. broad, and
the intermediates between this extreme and the normal are numerous. In some
the strial punctures on basal third of disk of elytra are broader than intervals,
being in these individuals relatively as broad as in .1/. angustus. In many the
alutaceous microsculpture of the surface of the prosternal process is distinctly
asperate. A few specimens have the entire middle of abdominal sternites two
to five densely tomentose.
Samples selected at random were taken from localities at three different
altitudes and measured for length. The length was taken from the apex of the
elytra to the anterior margin of the pronotum, and the head was never included
Table II shows that there is an increase in size (the length of this species shows a
high positive correlation with breadth) with an increase in altitude, and, as may
lie seen from Table III, the differences between the means of the three samples
may be regarded as significant.
Table II.
Temascaltepec. 5600-7500 ft. 1-772
Cuernavaca, 4S00 ft. . . 1-726
Tejupilco, 3500-4000 ft. . 1-620
1 "25 1-575
1 -goo 1-425
1S50 1 400
0782
1 100
1152
00123
00142
° °I35
0-0179
00204
00194
20
30
38
Lengths are given in mm. S.D. equals standard deviation ; S.E.D. equals standard
error of lire deviation ; and S.E.M. equals standard error "f the mean.
Table III.
Temascaltepec and Tejupilco
Tejupilco and Cuernavaca
Cuernavaca and Temascaltepec
The significance (if tire difference between the means for length "f tin- samples from the
\1, M.
three different altitudes has been found by using the formula t ■.-
\/ (o. h...\l.l) " -f- (Sj.l-..M.;)
where M is the mean and S.E.M. the standard error of the mean. Since in all cases the sum
of the samples is over 30, the probabilities have also been given for values of x (Fisher, ro36),
as these probabilities of the homogeneity of the population are more accurate.
Comparative notes : Of those species already described, this ran only be con
fused with M . angustus, but may be readily distinguished as has been pointed out
above under the description of angustus.
NOVIT. /.(.)(>[.., 42, 2. 1940. 25
3i4
H. E. HINTON
Microcylloepus troilus sp. n.
(Text-figs. 214-220.)
Male : Length, 1-85 mm.-2-20i mm. ; breadth, 0-82 mm.-o-o,7 mm. Exter-
nally similar to inaeqiialis except as follows : (1) the elytra has the third discal
interval at most only very feebly convex on basal seventh, whereas in inaeqiialis
it is strongly convex to carinate on basal fifth or sixth ; (2) the inner sublateral
carinae of the elytra are never as prominent as the outer and are frequently so
Text-figs. 214-220. — Microcylloepus troilus Hinton. (214) Dorsal view of male geni-
talia. 1 2 15) Left lateral view of same. (216) Lateral view of left paramere of same.
(217) Female genitalia. (218) Antenna. (219) Maxillary palp. (220) Apex of left
coxite of female genitalia.
feebly convex that they are scarcely visible, whereas in inaeqiialis they are always
prominent and generally as prominent as outer carinae ; (3) troilus is nearly
always much larger and though there is some overlap the means for absolute
length are very different (vide Table VI) ; and (4) the male genitalia are different
(cf. text-figs. 214-216).
Female : Externally similar to male.
Type : c? in the British Museum (Nat. Hist.). Mexico : Dist. de Temascal-
tepec, Real de Arriba, alt. about 7000 ft., vi.1933 (H. E. Hinton, R. L. V singer).
Paratypes : 163, collected in the same district from Temascaltepec to Real
de Arriba, alt. 5600-7500 ft., v-vii.1933 (H. E. Hinton, R. L. V singer) ; 866, with
MEXICAN ELMIDAE
315
data as above but collected in 1934 (H. E. Hinlon) ; and 2(10, in the same district
at Rio Verde, alt. about 8000 ft., 14. vi. 1934 (H. E. Hinton).
Variations : There is very little variation in the density of the punctures on
the various sclerites. In no case are the inner sublateral carinae of the elytra as
prominent as the outer, being one-half to two-thirds as high in 64 per cent.,
while in about 36 per cent, they are so feebly convex that they are scarcely notice-
able. When viewed from the side the prosternum has the anterior three-fifths
(not including process) bent ventrally in about half the cases, while in the other
half the anterior three-fourths is bent. The prostemal carinae are usually present
on basal half of prosternum, in a few cases they are on basal three-fifths or two-
fifths, while in a very few they are confined to basal third. In a few case-, the
carinae of the first abdominal sternite extend only to posterior third of segment,
but in the great majority they are complete or very nearly so.
A sample was selected at random from each of two altitudes. The specimens
wen- measured for length from the apex of the elytra to the anterior middle margin
of the pronotum. Table IV shows a slight increase in size with an increase in
Mx-M2
altitude. But, using the formula t
, where M is the
\/(S.E.M.1)2+ (S.E.M.2):
mean and S.E.M. the standard error of the mean, t = o-6 and the probability of
the two samples representing populations homogeneous for length is 50 percent.,
so that the difference in size recorded in Table IV should be considered to be of
no significance.
Lengths are given in mm. S.D. equals standard deviation ; S.E.D. equals standard
error of the deviation ; and S.E.M. equals standard error of the mean.
Comparative notes : This species appears to be very close to .1/. obesus, but
may be distinguished by such qualitative characters as the flat instead of comix
scutellum and the differently formed male genitalia (cf. text-figs.). M. troilus is
a distinctly smaller species as may be seen from Table V, being Intermediate in
size between obesus and inaequalis.
Tabu-: V.
Lengths are given in mm. and are taken from the apex of the elytra to the anterior
middle margin of the pronotum. S.D. equals standard deviation ; S.l D equals standard
error of the deviation ; and S.E.M. equals the standard error of the mean. The two n
were compared using the formula x = — . \ 5-6sotha1 I' to
1 N (S.l M ,) ' (S E.M,.)«
and the difference between the means may be n gardi d as ifii ant.
3i6
H. E. HINTON
Species.
troilus, 5600-7500 ft.
inaequalis, 5600-7500 ft.
Table VI.
2 -0034
1-772
2- 125 I«25 OO656
1 925 1-575 00782
00070
00123
OOIOl
00174
43
20
Lengths are given in mm. and are taken from the apex of the elytra to the anterior middle
margin of the pronotum. S.D. equals standard deviation ; S.E.D. equals standard error of
the deviation ; and S.E.M. equals standard error of the mean. The two means were com-
pared, using the formula x = ' ~ \ 2 — . x = n and P<io-», so that the
^(b.L.M.,)- + (S.E.M.,)-,
difference between these two means may be regarded as being very significant.
Larvae.
The larvae of Microcylloepus have been determined by elimination and accord-
ing to locality, e.g. this is the only genus of the family found in both California
and Trinidad, B.W.I., and larvae have been collected by the writer in California
which are difficult to separate from those taken in Trinidad. Four species of
larvae are available, and from a study of these the following generic diagnosis
has been made.
Generic Characters of Larvae of Microcylloepus.
Body parallel, cylindrical, and evenly convex above. Tubercles of middle
region of first eight abdominal tergites for the most part arranged in parallel,
longitudinal rows. Head when viewed dorsally exposed and not at all concealed
by the pronotum ; anterior margin on each side between base of antenna and
clypeus without a distinct tooth. With one ocellus on each side. Antennae
(text-fig. 222) 3-segmented and not or only very feebly retractile. Clypeus not
separated from the front of the head by a visible suture. Mandible (text-fig. 230)
of both sides similar and with three obtuse apical teeth ; prostheca long, slender,
and densely spinose. Maxilla (text-fig. 221) with the palp 4-segmented and the
stipes showing no differentiation into a palpifer ; galea and lacinia separate and
apex of each densely spinose. Labium (text-fig. 223) with the palp 2-segmented
and prementum without a palpiger ; post-mentum undivided. Hypopharynx
with a sclerotized rod on each side. Gula not developed. Ventral sutures well-
developed. Prothoracic pleura (text-fig. 226) divided into two parts and anterior
part meeting on middle line of body so that sternum is here completely suppressed.
Meso- and metapleura (text-fig. 226) divided into two parts. Abdominal segments
one to seven with the pleura bounded by tergo- and sterno-pleural sutures and
these two sutures converge on seventh segment and meet at its apex ; segments
one to seven with a pale dorsal line or suture parallel to tergo-pleural suture ;
segment eight forming a complete sclerotized ring ; apex of ninth segment
shallowly and broadly, arcuately emarginate. Operculum with two strongly
sclerotized claws attached to its dorsal membrane. Spiracles present on meso-
thorax and first eight abdominal segments and opening at the apices of small
tubercles. Tracheae without air sacs ; with three tufts of retractile anal gills.
Alimentary canal with an oesophageal sclerite on posterior dorsal margin of the
oesophagus. Hind gut with six Malpighian tubules which end freely near the
rectum. Central nervous system with three thoracic and eight abdominal discrete
ganglia.
MEXICAN ELMIDAE
3*7
The larvae of this genus are very close to those of Elsianus and Neodmis, but
from the former may be distinguished by the absence of a fronto-clypeal suture
and the absence of a tooth on anterior margin of head between the base ol antenna
and clypeus. From Neodmis they may be distinguished by having six instead
oi four Malpighian tubules, and by having the tubercles on the dorsal surface
arranged in rows.
Description of Mature Larva of M. inaequalis (Sharp).
(Text-figs. 221-231.)
Length 4-0 mm. ; breadth (across broadest point which is near base of meta-
thorax), 0-55 mm. Elongate, subparallel, and cylindrical to subtriangular in
22 5
I 1 \i-fk,s. 22 1 225. — Larva of Microcylloepus inaequalis 1 Sharp). 1221 ) Ventral view
of right maxilla. (222) Dorsal view of left antenna. (223) Ventral view of labium.
Setae of prementum are only approximately correct. [22. p Dorsal view of right
opercular claw. (2231 Mesothoracic spiracle.
cross section. Cuticle moderately shining and moderately dark brown to brownish
testaceous. Head at broadest point, which is at basal third, about as broad as
long (0-30 mm.) ; posterior dorsal margin (text-fig. Z2(>) broadly, deeply, and
arcuately emarginate ; epicranial suture 0-012 mm. long ; frontal suture on each
side extending in a nearly straight line to anterior margin oi head near inner
base ol antenna ; anterior margin between base of antenna and clypeus without
a tooth ; surface sparsely pubescent with fine and erect hairs which are usually
about 0-03 mm. long ; surface on a basal belt which extends as fai as beginning
oi frontal sutures with round, microscopic (about 0-004 mm- broad) granules
3i8
H. E. HIXTON
which are usually separated by once their diameters ; surface elsewhere with
moderately convex, oval granules which are about o-or.2 mm. broad and are
usually separated by less than to one and a half times their diameters ; anterior
dorsal margin with a single row of these granules which are about a fourth coarser
and slightly denser ; all granules except fine ones of basal belt have a fine seta
which is about three-fourths as long as its respective granule on apical anterior
margin. Terga of thoracic and abdominal segments with the granules for the
most part similar to those of head but usually as coarse as those of anterior row.
Text-figs. 226-231. — Larva of Microcylloepus inaequalis (Sharp). (226) Ventral
view of thorax and first abdominal segment to show sclerotization. (227) A section
of the fifth abdominal tergite to show arrangement of tubercles. (228) Dorsal view
of ninth abdominal segment. (229) Operculum. (230) Mandible. (231) Inner
face of front leg.
Pronotum with the granules of posterior half partly arranged in a series of irregular,
parallel, longitudinal rows ; all of tergites of last two thoracic and first eight
abdominal segments with the granules arranged in similar rows (text-fig. 227) ;
posterior margin of all segments except ninth and anterior margin of pronotum
with a row of dense tubercles from which arise large, flat setae (text-fig. 227) ;
anterior seventh or eighth of all segments except pronotum with a belt of fine
(similar to those of basal belt of head) , flat-topped, subquadrate to round tubercles ;
ninth tergite with a median longitudinal, strongly convex ridge extending from
anterior eighth to posterior fifth and at middle about 0-07 mm. broad and 0-05
mm. high ; apex of this segment (text-fig. 228) broadly and shallowly, arcuately
emarginate. Sternites with the tubercles usually only two-thirds as coarse as
MEXICAN ELMIDAE 319
those nl tergites, separated by two to three times their diameters, and not arranged
in rows but otherwise similarly sculptured ; sternites of last four abdominal
segments with the tubercles coarser and usually separated by only twice their
diameters ; first abdominal sternite at middle anterior margin with a large
(0-03 mm. broad) and high tubercle. Operculum (text-fig. 229) with the claws
(text-fig. 224) not toothed. Pleurites tuberculate as their respective sternites
except for those of prothorax which are about twice as densely tuberculati-.
Legs all fairly similar in chaetotaxy to front leg (text-fig. 231) and with the front
pair shortest and the hind pair longest. Spiracles opening on apices of small
tubercles and all similar to those of mesothorax (text-fig. 225).
Specimens examined: 22, Mexico: Estado de Morelos, Cuernavaca, alt.
4800 ft., vi.1934 (H. E. Hintori).
Among the series before me are a few specimen-, representing at least one
earlier instar, but these apparently differ in no way but size from the mature
larvae.
NEOELMIS Musgrave.
1935. Neoelmis Musgrave, Proc. Ent. Soc. Wash., 37 (zj : 34.
[936. Neoelmis Hmton. /•.»/. Mon. Mag., 72 : 5.
[936. Elmis Darlington, Psyche, 43 : 79.
This genus was erected by Musgrave to contain a new species from Puerto Rico,
and to it he also referred Elmis apicalis Sharp, E. caesa Leconte, and E. minima
Darlington. Hinton (1936) added a few generic characters omitted from the
original description and referred Elmis simoni Grouvelle of Venezuela to the genus.
Darlington (1936) sunk this genus as a synonym of Elmis Latreille, a genus very
different from Neoelmis in the structure of the internal and external anatomy of
both larvae and adults. If Darlington is correct and Elmis and Neoelmis should be
united, then all the well-known genera of European Elmini, with the possible
exception of Stenelmis Dufour, must be sunk as synonyms of Elmis, for between
these and Elmis there is less difference than between Elmis and Neoelmis. Such
a course would invalidate the work of Erichson, Ganglbauer, Reitter, etc., on
the adults and the work of Bertrand on the larvae. Perhaps it is needless to
add that Darlington's conception of the generic limits of Elmis is followed by none
of the present authorities on the family.
At the time of writing there are 21 species known. These occur from southern
United States to south Brazil, and three species are known from the West Indies.
A it-description of the genus follows :
Body subparallel and usually long and narrow. Non-tomentose areas
glabrous or clothed with sparse and short recumbent hairs. Scale-like- or hairy
tomentum confined to the following areas : (1) genae ; (2) epipleura ; (.-;) sides
of presternum (in some species, e.g. maculata, there is no tomentum on the pro-
sternum) ; (4) sides of mesosternum, metasternum and abdominal sternites, but
in some species nearly the entire metasternum and tin- sternum of the abdomen
is clothed with tine tomentum ; and (5) part or all of legs except tarsi. Head
when seen from below capable of being retracted SO that none "1 the mouth-parts
is visible. Antennae 1 i-segmented (text-fig. 24')). Mandibles with three sub-
acute, apical teeth ; prostheca long and entirely membranous with numerous long
spines or hairs apically. Maxilla with the palp (text-fig. 245) 4-segmented and
stipes with a well-developed palpifer; galea and lacinia separate and apex "i
320
H. E. HINTON
each spinose or hairy. Labium with the palp 3-segmented and prementum with-
out a distinct palpiger. Mentum about as broad and three-fourths as long as
submentum. Gula slightly longer than submentum, at anterior margin about
tour-fifths as broad as submentum, and with the sides feebly converging, so that
Text-figs. 232, 233. — Neoelmis thoracica (Grouvelle). (1$!) Male reproductive
system. (233) Female reproductive system.
Text-fig. 234. — Hind wing of Neoelmis longula Hinton. Venation after Forbes.
at posterior margin it is only half as broad as submentum. Pronotum with the
anterior margin moderately arcuate at middle and on each side behind eye before
apical angle broadly and moderately deeply sinuate. Base trisinuate, broadly
and moderately so on each side and more narrowly and shallowly so in front of
scutellum. Pronotum on each side with a sublateral carina which extends from
base nearly to anterior margin ; at apical two-fifths with a deep, broad, and
MEXICAN ELMIDAE 32I
complete transverse impression ; disk on either side of transverse impression
evenly convex, though occasionally with a gibbosity in front of the scutellum.
Elytra punctate and striate ; each elytron with a sublateral carina on sixth
interval. Hind wing (text-fig. 234) with a very feebly developed anal lobe ;
without a radial cross vein or an anal cell ; first anal usually absent ; second
anal with the second branch absent ; third anal only present basally and not
joined to second anal ; fourth anal well developed : and cubito-anal cross vein
joining cubitus to second anal, though often incomplete. Presternum very long
in front of anterior coxae ; prosternal process long, narrow or broad, and posterior
margin of process nearly truncate to broadly rounded. Mesosternum with a
broad and deep groove for the reception of the prosternal process. Metasternum
with a median longitudinal impressed line. Legs with the visible portion of the
front coxae rounded and trochantin completely concealed by the hypomera and
sternum, flaws without teeth. Alimentary canal with three, two or no caeca
on the anterior margin of the mid-gut. Hind gut with four Malpighian tubules
which end freely near the rectum. Male reproductive system (text-fig. 232) with
the lateral accessory glands lobed. Each testis with two sperm tubes. Female
reproductive system (text-fig. 233) with four egg-tubes to each ovary. Sperma-
thecal duct opening into middle of bursa copulatrix. Central nervous system with
three thoracic discrete ganglia ; abdominal ganglia one to six discrete ; and
seven and eight fused, but with the limits of each distinguishable.
Genotype : Neoelmis gracilis Musgrave (1935).
The internal anatomy of three species has been examined and found to agree
in essential details, except that the number of caeca of the mid-gut varies. That
of N. thoracica Grouvelle is figured.
This genus appears to be nearer to Esolus Mulsant and Rey than to any other
described genus. From Esolus it may be distinguished as follows : (1) the hypo-
mera are not clothed with tomentum as they are in Esolus ; (2) the pronotum has
a deep and complete transverse impression on apical two-fifths, whereas in Esolus
it is more or less evenly convex ; (3) the hind wing has the second anal branched and
at least part of a cubito-anal cross vein is present, whereas in Eso/ns the second
anal is not branched and there is no cubito-anal cross vein ; (4) the anterior
margin of the mid-gut has three, two, or no caeca, whereas in Esolus no caeca
are here present ; and (5) the central nervous system has only the seventh and
eighth ganglia partly fused to form a terminal abdominal ganglion, whereas in
E'solus the terminal abdominal ganglion consists of a fusion of ganglia four to
eight.
A list of the specific characters which seem to be of most importance in
distinguishing the species of Neoelmis is as follows :
(1) General proportions, length and breadth.
(2) Colour. In some species, e.g. maculata, the elytra are maculate.
(3) Size and distribution of the punctures on the various sclerites and tin-
density and type of the microsculpture, if present, between the
punctures.
(4) Condition of fronto-clypeal suture.
(5) Anterior margin of clypeus, whether rounded, truncate, emarginate,
or sinuate : and also the condition of the angle on each side.
(()) Condition of anterior margin of labrum and the angle on each side.
(7) Outline of pronotum and its general proportions.
(8) Condition of disk of pronotum, whether with gibbosities or evenly
convex.
322 H. E. HINTOX
(9) Shape ot elytral apices.
(10) Shape of scutellum.
(n) Condition of anterior portion of presternum, whether straight or bent
ventrally when viewed from the side.
(12) Extent of tomentum, if any, on sides of presternum.
(13) Condition of prosternal carinae. In some species they are absent, but
when present may be parallel, diverging or converging anteriorly, or
contiguous anteriorly. If they are parallel their breadth apart
should be given.
(14) Breadth of prosternal process.
(15) Condition of sides of prosternal process. In some species they are
strongly gibbous, while in others they are not at all thickened.
(16) Extent and depth of mesosternal groove.
(17) Disk of metasternum, whether flat or not and the extent and depth of
the impressions.
(18) Length, shape, and prominence of carinae, if present, on middle of
first abdominal sternite.
(19) Secondary sexual characters.
(20) Structure of male genitalia. No case is known where these structures
are the same for two or more specie-,.
A list of the secondary sexual characters that have been observed in the
species before me is as follows :
(1) Male with a tooth on anterior margin of presternum (gigas, prosternalis,
simoni).
(2) Male with the disk of the metasternum more strongly and exten-
sively concave than that of the female {gigas, lobata, musgravei,
prosternalis).
(3) Each side of median impression of metasternal disk of male with
numerous testaceous hairs, and also densely punctate, while in female
this region is glabrous or nearly so and is also sparsely punctate
(musgravei).
(4) Male with carinae on the middle of the first abdominal sternite, while
in the female they are absent (prosternalis).
(5) Male with the carinae of the first abdominal sternite converging, while
in the female they are straight (gigas).
(6) Male with a tubercle on each side of middle of first abdominal sternite
(gigas, prosternalis).
(7) Male with the anterior middle of the second abdominal segment feebly
depressed (aspera, azteca, longula).
(8) Male with the inner apex of the front tibia gibbous (gigas).
(9) Male with a row of short and stout spines on inner apex of middle
tibiae (gigas, prosternalis).
(10) Male with a row of short and stout spines on inner ventral side
of apical three-fourths of middle and hind tibiae (apicalis,
aspera).
(11) Male with a row of small spines on ventral side of apical segment of
middle and hind tarsi (gigas, lobata, prosternalis).
In the following key for the separation of the Mexican species N. caesa Leconte
is included. This species is to-day recorded only from Texas, but will very likely
be found to occur in the north-western parts of Mexico which are a continuation
of the same Sonoran faunal region.
MEXICAN ELMIDAE 323
A Key to the North American' Species of Neoelmis.
I. Hypomera with the middle area entirely smooth and without an asperate
type of microsculpture. Mexico . . N. longula Hinton (1936).
Hypomera densely asperate throughout ...... 2.
3. Disk of metasternum without a depression on each side. Mexico
N. azteca, sp. n.
Disk of metasternum with an oval depression on each side ... 3.
3. Prosternal carinae parallel and separated by a distance equal to breadth oi
process near base ; presternum with a distinct ridge on each side
parallel to and between prosternal carina and sterno-notal suture.
Guatemala, Mexico N. apicalis (Sharp) (1882).
Prosternal carinae converging in front so that while at base they are
separated by a distance equal to the breadth of the process, at apex
they are separated by only half or less of this distance ; presternum
on each side without a distinct ridge between carina and sterno-
notal suture .......... 4.
4. Prosternal carinae at anterior end separated by a distance equal to about
half of the breadth of the process at base. Surface of flat parts of
metasternal disk without an asperate microsculpture between
anterior seventh and basal fifth. Mexico . . N. aspera, sp. n.
Prosternal carinae at anterior end separated by a distance equal to about
a third of the breadth of the process at base. Surface of flat part-- 1 if
metasternal disk nearly entirely asperate. Texas
N. caesa (Leconte) (1*74)
Neoelmis longula Hinton.
(Text-figs. 234-238.)
[936. Neoelmis longulus Hinton. Trims. R. lint. Soc. Lond., 85 (18) : J26, fig. ^1 . pi. 1.
fig. 2.
Male: Length, 1-97 mm. -2-20 mm. ; breadth, 0-82 mm. -0-90 mm. Cuticle
shining and rufo-piceous ; antennae, mouth-parts, and legs paler rufo-piceous.
Head without distinct impressions ; surface very densely and microscopically
alutaceous. Clypeus with the fronto-clypeal suture straight and very feebly
and indistinctly impressed; anterior margin when seen from in front broadly,
arcuately, and very feebly emarginate ; anterior angles broadly rounded ; surface
only sparsely alutaceous on most of the middle but at sides alutaceous as on head.
Labium with the anterior margin feebly and broadly rounded and the angle on
each side broadly rounded ; surface sparsely and transversely alutaceous basally,
and anteriorly with tine punctures which air about 0-010 mm. broad and are
separated by one to three times their diameters. Pronotum across broadest
point, which is at basal third, not quite as broad as long (0-575 mm. : 0-587 mm.)
and base broader than apex (0-52 mm. : 0-37 mm.). Sublateral carina extending
from near base to apical eighth. Impressions as figured (text-tig. 238). Lateral
margins nearly smooth. Surface on convex portions between sublateral carinae
with fine (about o-ooN mm. broad) punctures which are usually separated by three
to five times their diameters, and surface between punctures smooth : elsewhen
similarly sculptured except as follows: bottom of transverse impression with a
dense asperate type oi microsculpture, ami the middle of this belt extending
slightly posteriorly; the basal and anterior sixth of area between sublateral
3^4
H. E. HIXTON
carina and lateral margin is irregularly alutaceous to asperate ; and the outer
dorsal side of the basal three-fifths of sublateral carina is also microscopically
alutaceous. Elytra nearly twice as long as pronotum (1-47 mm. : 0-58 mm.) and
at broadest point, which is across apical third, broader than broadest point across
humeri (0-82 mm. : 0-76 mm.). Lateral margins nearly smooth (under magnifi-
cation of more than x 150 they appear very finely and irregularly crenate).
Apices moderately strongly, broadly produced and conjointly feebly rounded.
Text-figs. 2^5-240. — (235) Presternum of Neoelmis longula Hinton. (236) Dorsal
view of male genitalia of same species. (237) Right lateral view of same. (238)
Adult to show general appearance. (23a) Dorsal view of male genitalia of .V.
azteca Hinton. (240) Right lateral view of same.
Surface with the striae feebly impressed on basal two-thirds, but absent on apical
third ; discal strial punctures deep, round to subquadrate, two-thirds to slightly
broader than intervals, and separated longitudinally by a half to two thirds of
their diameters ; from apical half to apical third these punctures become rapidly
finer so that at apical third they are about a third as coarse, only half as broad as
intervals, and are separated longitudinally by twice their diameters ; from apical
third to apex they become slightly finer and sparser. Intervals flat ; surface of
intervals feebly, sparsely, very finely alutaceous and more finely punctate than
pronotal disk ; surface of intervals at extreme base and on basal fourth between
lateral margins and sublateral carinae with round, low granules which are about
MEXICAN ELMIDAE 325.
0-012 mm. broad ami are usually separated by oner to twice their diameters.
Scutellum flat, subovate, longer than broad (o-io mm. : 0-075 mm.), base feebly
rounded, and apex more strongly rounded ; surface sculptured as natural interval.
Prostemum when seen from the side with the anterior three-fifths (not including
process) moderately strongly but not sharply bent ventrallv ; prosternal carinae
present on posterior two-fifths, prominent, parallel, and separated by a distance
equal to about four-fifths of the breadth of the process between coxae ; between
carina and sterno-notal suture there is no distinct ridge ; process (text-fig. 235)
moderately narrow, parallel, with the margins moderately strongly gibbous, and
the posterior margin broadly rounded ; fine hairy tomentum present on most of
sides ; surface sculptured similarly to discal elytral intervals but antero-lateral
sides with a dense and asperate type of alutaceous microsculpture. Hypomera
smooth and highly polished except for basal fifth, apical sixth, margins near
sterno-notal suture, and bottom of depressions where it is mostly densely asperate.
Mesosternum with the anterior portion of the groove for the reception of the pro-
sternal process broad and deep, but at posterior half suddenly becoming twin ,1-
broad and slightly deeper so that all of the mesosternum between coxae except
lateral and posterior margins is depressed. Metasternum with the median longi-
tudinal line complete ; disk deeply and broadly depressed on middle posterior
three-fifths and the bottom of the depression is asperate ; surface of disk elsi
where with irregularly-shaped punctures which are about 0-007 mm. broad and
are separated by three to eight times their diameters, while the surface between
the punctures is smooth ; sides of metasternum densely, microscopically alutaceous
and this sculpture is sometimes asperate. Abdomen with the carinae of the
first sternite straight except posteriori}- where they are feebly curved inwards,
and extending to posterior fifth of sternite ; the entire middle area of the first
sternite between carinae is strongly depressed ; second abdominal sternite feebly
depressed on middle of anterior third ; surface of basal half of middle of first
sternite asperate, while apical half, middle region of sternites two to four, and
basal third of fifth punctate similarly to metasternal disk ; apical two-thirds of
fifth, as well as lateral part, with punctures which are about 0-OI2 mm. broad
and are separated by two to three times their diameters ; sides of sternites one
to four moderately densely, microscopically alutaceous. Genitalia as figured
(text-figs. 236, 237).
Female : Externally similar to male except that middle third of the second
abdominal sternite is not depressed.
Type: $ in the British Museum (Nat. Hist.). Mexico : Dist. de Temascal-
tepec, Real de Arriba, alt. 6000-7000 ft., vii.1933 (H. E. Minimi, R, L. Usinger).
Specimens examined : 10, with data as above but collected in v-vii ; 102. as
above but collected in v -vii.1934 (H. E. Hinton) ; and 7. in the same distrii 1 at
Rio Verde, alt. about 8000 ft., 14. vi. 1934 (H. E. Hinton). 13, Mexii 0 : Estado
de Morelos, Cuernavaca, alt. about 4800 ft., vi.1934 (H. E. Hinton).
Teratology: One female from Real de Arriba has a deep oval pit on the right
hand side of the metasternal disk.
Variations : In some specimens the carinae of the first abdominal sternite
only extend to posterior halt. There is an increase in si/.e with an increase in
altitude. This could not be tested on the material collected in the district of
Temascaltepec, as sufficiently accurate altitude data are not available. But the
series collected at Cuernavaca, which is at a lower altitude approximately within
the same latitude, iias a smaller mean value for absolute length (Table VII,
text-fig. 241). The significance of the difference between the means ot the series
326
H. E. HINTON
collected in Cuernavaca and that collected in the district of Temascaltepec was
M — M,
calculated from the formula t = . //cFM'\2 _l /q E M ^' WherC M iS the m6an
and S.E.M. is the standard error of the mean. In this case t = 5-5 and, as may
be seen from Fisher's (1936) table of t, the probability, P, of these two series
being got from the same population is considerably less than P<-oi. This
difference in absolute size of populations of the same species at different altitudes
is probably due to a difference in temperature during the larval life of the species.
From the district of Temascaltepec there are 13 specimens collected between
5600 and 7500 ft. These are so similar to longula that I have been unable to
separate them on any qualitative character. But, as may be seen from Table VII
or text-fig. 241, the absolute length of the longest is not even close to the absolute
length of the smallest of typical longula ; and they may be readily separated
on size even by the unaided eye. This series of small individuals (hereafter
called X) may be longula or another species.
Table VII.
The figures are given in mm. The lengths are taken from the apex of the elytra to the
middle anterior margin of the pronotum, and the head was not included as to do so would
have introduced a considerable error into the measurements, since the total length would
-vary in the order of 0025 mm. : 0100 mm. depending on how much the head happened
to be retracted into the prothorax. S.D. equals standard deviation ; S.E.D. equals standard
-error of the deviation ; and S.E.M. equals standard error of the mean.
If X is longula it is obvious that some factor is playing the part of a sieve
and sharply dividing the population into two size groups. This factor is not sex,
as males and females occur in the proportion of about 1 : 1 in both groups. The
population of longula may be infected by some protozoal or other disease the
presence of which would make the size of any individual much larger or smaller.
But even in this case lightly infected individuals intermediate in size would be
expected to occur.
It seems much more probable that X is, in point of fact, a new species. If
this is so, it should not be very surprising that these two species cannot be sepa-
rated when the crudity of the available qualitative taxonomic methods is taken
into account. X may differ from longula not only in one but in a great many
genes which control the histological and biochemical structure of many different
tissues. By the usual taxonomic methods applicable to dried specimens two or
more related species can only be distinguished if there happens to be a difference
in external characters. Even if the internal characters are alike, there may still
be many biochemical differences of specific importance.
Comparative notes : N. longula may be distinguished from all its North
American congeners by the smooth instead of asperate middle region of the
hypomera and by the structure of the male genitalia.
MEXICAN ELMIDAE 337
Neoelmis azteca, sp. n.
(Text-figs. 239, 240.)
Male: Length, 1-92 mm. ; breadth, 075 mm. Similar to longula except
as follows : (i) the round granules at the base and sides of elytra are much less
numerous and distinct ; (2) the prosternal carinae distinctly converge so that
anteriorly they are only half as far apart as at base ; (3) the asperate type of
alutaceous microsculpture completely covers the hypomera, and there are no
smooth and polished areas ; (4) the asperate microsculpture of the depressed
portion of the middle of the first abdominal sternite extends over the basal two-
thirds, whereas in longula it is confined to the basal half ; and (5) the male
genitalia are different (text-figs. 230, 240).
Female : Unknown.
Type: q in the British Museum (Nat. Hist.). Mexico: Dist. de Temascal-
tepec, Real de Arriba, alt. about 7000 ft., vii.1934 (H. E. Hinton).
Paratype : A male with the same data as above.
Neoelmis apicalis (Sharp).
(Text -figs. 242-245.)
1882. Elmis apicalis Sharp, Biol. Centr.-Amer. Cut.. 1 (2) : 136, t. 4, rig. 14.
1935. Neoelmis apicalis Musgrave, Proc. Ent. Soc. Wash., 37 (2) : 34, 35.
1936. Neoelmis apicalis Hinton, Trans. R. Ent. Sue. Loud., 85 (18) : 420. fig. 22.
Male : Length, 1-57 mm.-i-75 mm. ; breadth, 0-62 mm. -0-65 mm. Cuticle
shining and rufo-piceous ; antennae, mouth-parts, and legs paler rufo-piceous.
Head without distinct impressions ; surface very densely and microscopically,
asperately alutaceous. Clypeus with the fronto-clypeal suture nearly straight and
moderately feebly impressed ; anterior margin when seen from in front truncate
to very feebly rounded ; anterior angles broadly rounded ; surface at sides
sculptured as head but elsewhere only occasionally alutaceous and with micro-
scopic punctures (about 0-003 mm- broad) which are usually separated by two to
five times their diameters. Labrum with the anterior margin feebly and broadly
rounded and the angle on each side broadly rounded ; surface punctate as middle
region of clypeus but with the punctures denser and very slightly coarser. Pro-
notum across broadest point, which is at basal third, slightly broader than long
(0-48 mm. : 0-46 mm.) and base broader than apex (0-42 mm. : 0-33 mm.). Sub-
lateral carinae extending from base to very nearly apical margin. Impressions
as in longula. Lateral margins feebly and regularly crenate. Surface on convex
portions between sublateral carinae with fine (about 0*009 mm. broad) punctures
which are usually separated by three to five times their diameters, and the surface
between these punctures is smooth ; elsewhere similarly sculptured except as
follows : bottom of transverse impression with a very -.hurt belt (about a third as
long as that of longula) of a dense asperate type of mieroseulpture ; area between
sublateral carinae with the basal fifth, anterior fifth, all of apical impression, and
outer sides of sublateral carinae densely asperate. Elytra more than t\\ ice as long
as pronotum (l-l2 mm. : 0-46 mm.) and at broadest point, which 1- across apical
third, broader than broadest point across humeri (0-62 mm. : 0-57111111.). Lateral
margins nearly smooth Apices moderately strongly, broadly produced and con-
jointly feebly rounded. Surface with the striae feebly impressed on basal two-
thirds but absent on apical third ; discal stria! punctures deep, usually round, as
328
H. E. HINTON
3 —
a —
1 —
-\ — i — i — i — i — i — i — i — i — r~i — i i — i — i — i — i — i — r
jiii — 2i.sc— itlf— J7«— /«f_ i7SB— jfV™ igM— J«l« jiw- j?»'— jta»— J«J». J "■>— /*>>-»_ aaw— loir— jom-- mv
241
Text-fig. 241. — Frequency polygons of three populations of N. longula Hinton. The
polygons are not adjusted to equal areas. The measurements are from the apex of
the elytra to the anterior margin of the pronotum.
244
24B
Text-figs. 242-248. — (242) Presternum of Neoelmis apicalis (Sharp). (243) Dorsal
view of male genitalia. (244) Right lateral view of same. (245) Maxillary palp.
(246) Antenna. (247) Dorsal view of male genitalia of N. aspera Hinton. (248)
Right lateral view of same.
MEXICAN KI.MIDAE 329
broad to half again as broad as intervals, and separated longitudinally by two-
thirds to once their diameters ; at apical two-fifths the discal punctures become
only about a third as coarse and dense and from here to apex they gradually
become finer so that at apex they are no more than a fourth as coarse and dense
as those near base. Intervals flat, feebly, sparsely, microscopically alutaceous
and also with punctures which are similar to those of pronotal disk but are more
irregularly distributed ; surface of intervals at extreme base with an occasional
round, low granule which is about 0-012 mm. broad, and sides on basal fourth
between lateral margins and sublateral carinae with similar granules which are
frequently separated by no more than their diameters. Scutellum flat, subovate,
slightly longer than broad (0-075 mm. : 0-070 mm.), base feebly, broadly rounded,
and apex more strongly rounded. Prosternum when seen from the side with the
anterior three-fifths (not including process) moderately strongly and sharply bent
ventrally ; prosternal carinae present on posterior two-fifths, prominent, parallel,
and separated by a distance nearly equal to that of the process at base ; on each
side between carina and sterno-notal suture there is a prominent, thick ridge
about o-io mm. long ; process as figured (text-fig. 242) and with the lateral and
apical margins moderately strongly gibbous ; surface of prosternum and hypo-
mera throughout densely asperate except for middle in front of process which is
sculptured as disk of elytra. Mesosternum with the groove for the reception of
the prosternal process broad and deep on anterior two-fifths, where the bottom
of the groove is smooth, and suddenly becoming broader so that posterior three-
fifths is nearly twice as broad and has the bottom densely asperate and at middle
of this area there is a large (about 0-050 mm. long) oval, smooth-bottomed, deep
pit ; surface of sides of mesosternum asperate. Metasternum with the median
longitudinal line complete, on basal five-sixths 0-50 mm. broad and deep, and on
anterior sixth very narrow and feebly impressed ; disk on each side on basal
third with an irregularly round pit which is about 0-06 mm. broad ; surface of
disk and bottom of all impressions densely asperate except for a narrow smooth
belt along sides of median line ; sides of metasternum densely and obscurely
asperately alutaceous. Abdomen with the carinae of the first sternite straight,
prominent, and extending to posterior half of segment ; middle of first sternite
deeply depressed on anterior three-fourths ; surface of sternites sculptured as
follows : basal two-fifths of depressed portion of first sternite densely asperate ;
posterior three-fifths of middle of first sternite and all of middle of sternites two
to four only sparsely microscopically alutaceous ; and sides of first four sternites
and all of fifth sculptured as sides of metasternum. Legs with the inner ventral
side of apical four-fifths of middle and hind tibiae with a row of short and stout
spines. Genitalia as figured. (text-figs. 243, 244).
Female : Externally similar to male except as follows : the middle and hind
tibiae have no inner rows of short and stout spines.
Type: In the British Museum (Nat. Hist.). Guatemala: Vera Paz, San
Joaquin (Cham pi an).
Specimens examined : 1 , with same data as type ; and 2, in the same state
at Rio Naranjo, 450 ft. (Champion). 200, Mexico: Dist. de Temascaltepec,
Tejupilco, alt. about 4000 ft., vi.1933 (H. E. Hinton, R. L. Usinger) ; 69, with
data as preceding but collected in vii.1934 (H. E. Hinton) ; 2, Mexico : Estado
de Morelos, Cuernavaca, alt. about 4800 ft., vi.1934 (//. E. Hinton).
Variations : Apart from the variation in absolute size already recorded, none
worthy of mention have been observed.
Comparative notes : From N. longula this species may be distinguished by
NOVTT. ZOOL., 42, 2. I94O. 26
330 H. E. HINTON
its entirely instead of only partly asperate hypomera, from azteca by the presence
of a pit on each side of the metasternal disk, and from caesa by having the
prosternal carinae parallel instead of converging anteriorly.
Neoelmis aspera, sp. n.
(Figs. 247, 248.)
Male: Length, 1 -70 mm. ; breadth, 0-65 mm. Externally similar to apicalis
except as follows : (1) the fronto-clypeal suture is not distinctly visible ; (2) the
belt of asperate microsculpture on the bottom of the apical impression of the
pronotal disk is broader (about as broad as in longula) ; (3) the extreme base of
the pronotum is asperate as is apical part of disk ; (4) the prosternal carinae are
indistinct and converge in front so that anteriorly they are separated by a distance
equal to only half of breadth of process at base ; (5) presternum without a distinct
ridge on each side between carina and sterno-notal suture ; (6) nearly the entire
non-impressed part of the metasternal disk is smooth ; (7) the second abdominal
sternite is feebly depressed on middle anterior third ; and (8) the male genitalia
are different (text-figs. 247, 248).
Female : Externally similar to male except as follows : the middle and hind
tibiae have no short and stout teeth, and the basal third of middle of second
abdominal sternite is not depressed.
Type: <J in the British Museum (Nat. Hist.). Mexico: Estado de Morelos,
Cuernavaca, alt. 4800 ft., vi.1934 (H. E. Hinton).
Paratypes : 8, with data as above ; and I, Mexico : Dist. de Temascaltepec,
Real de Arriba, alt. about 7000 ft., vii. 1934 (H. E. Hinton).
Teratology : One male specimen from Cuernavaca has no pit on the left-hand
side of the metasternal disk, and the pit on the right-hand side is relatively only
three-fourths as long as is normal for this species.
Variations : In some specimens the carinae of the first abdominal sternite
extend to posterior fifth of segment, while in others it only extends to posterior
half, as is usual in apicalis.
Comparative notes : N. aspera is also very close to caesa, but differs in having
the prosternal carinae anteriorly separated by a distance equal to about half the
breadth of the process at base, whereas in caesa they are here separated by a
distance equal to only a third the breadth of the process at base ; and in having
the flat parts of the metasternal disk mostly smooth, whereas in caesa the disk
is here mostly asperate.
Larvae.
No larvae of this genus were collected in Mexico. By elimination and
according to locality the larvae of two Bolivian species of Neoelmis have been
determined, and the following generic diagnosis is drawn up from a study of these.
Generic Characters of Larvae of Neoelmis.
Body subparallel and cylindrical to subtriangular in cross section ; dorsal
surface evenly convex. Head when seen from above exposed and not concealed
by the pronotum ; anterior margin on each side between base of antenna and
clypeus with a short, indistinct, blunt tooth. Clypeus with the suture distinct.
With one ocellus on each side. Antennae 3-segmented and not or only very
feebly retractile. Mandibles of both sides similar and with three obtuse, apical
MEXICAN ELMIDAE 33I
teeth ; prostheca long, slender, and densely spinose. Maxilla with the palp
4-segmented and stipes showing no differentiation into a palpifer ; galea and
lacinia separate and apex of each densely spinose. Labium with the postmentum
undivided ; labial palp 2-segmented and prementum without a distinct palpiger.
Quia well developed. Prothoracic pleura divided into two parts and anterior
part meeting on middle line of body so that sternum is here completely sup-
pressed. Meso- and metapleura each divided into two parts. Abdominal
segments one to seven with pleura bounded by tergo- and sterno-pleural sutures
and these two sutures converging and meeting at apex of seventh segment ;
segment eight forming a complete sclerotized ring. Operculum with two strongly
sclerotized claws attached to its dorsal membrane. Apex of ninth segment
broadly and arcuately emarginate. Spiracles present on mesothorax and first
eight abdominal segments and opening at apices of small tubercles ; trachea
without air sacs ; with three tufts of anal, retractile, tracheal gills. Alimentary
canal with an oesophageal sclerite on the dorsal posterior margin of the oesophagus.
Hind gut with four Malpighian tubules which end freely near the rectum. Central
nervous system with three thoracic and eight abdominal discrete ganglia.
The larvae of this genus are very close to those of Elsianus, and may be
distinguished from some species of Elsianus by the absence of a dorso-lateral
suture parallel to tergo-pleural suture on segments two to seven. Some species
which appear to belong to the genus Elsianus have no dorso-lateral suture on the
abdomen, and from these Neoelmis can only be distinguished by the nearly tooth-
less anterior margin of the head and by counting the Malpighian tubules. The
adults of Neoelmis differ so greatly from those of Elsianus in the structure of both
their internal and external anatomy, that I consider the similarity of the larvae
of these two genera to be due to parallelism.
HEXACYLLOEPUS, gen n.
Body elongate, subparallel. Dorsal surface clothed with short, sparse or
dense, usually recumbent hairs. Hairy or scale-like tomentum confined to the
following areas : (1) genae ; (2) hypomera with a large area which begins opposite
middle of front coxae and extends transversely or obliquely upwards to lateral
margin of pronotum and forms next to this margin a belt which frequently
extends to anterior margin ; (3) epipleura ; (4) sides of presternum, mesosternum,
metasternum, and abdominal stcrnites, but in a few species nearly the entire
ventral surface may be clothed with tomentum ; and (5) all of legs except tarsi,
though in most species the tomentum is sparse or absent on many parts of the
tibiae and femora. Head when seen from below capable of being retracted so
that none of the mouth-parts are visible. Antenna n-segmented. Mandible
with three acute apical teeth ; prostheca entirely membranous and with the apex
spinose or hairy. Maxillary palp 4-segmented and stipes with a well-developed
palpifer ; galea and lacinia separate and apex of each densely spinose. Labial
palp 3-segmented and prementum with a palpiger. Mentum transverse and
about as broad and long as submentum. Gula anteriorly nearly as broad as sub-
mentum but posteriorly slightly narrowed. Pronotum with a sublateial longi-
tudinal carina on each side which generally extends from base to apex, though
in a few species it does not quite reach either base or apex ; with a median longi-
tudinal impression on disk. Elytra striate and punctate ; without accessory
striae ; with two sublateral carinae on each elytron. Hind wing (text-fig. 252)
332
H. E. HINTON
without a radial cross vein and without an anal cell ; with the first anal absent ;
second anal with the first and second branches present ; third anal with a second
branch ; cubito-anal cross vein complete and joining cubitus to second anal.
Prostemum very long in front of anterior coxae ; prosternal process long and
posterior margin usually broadly rounded. Mcsosternurri with a deep and
moderately narrow groove for the reception of the prosternal process. Meta-
sternum with a median longitudinal impressed line. Legs with the visible portion
of the front coxae rounded and trochantin completely concealed by the hypomera.
Claws without teeth. Alimentary canal with six caeca on the anterior margin of
249
250
Text-figs. 249, 250. — Hexacylloepus smithi (Grouvelle). (249) Male reproductive
system. (250) Female reproductive system.
the mid-gut ; with the outer surface of the mid-gut smooth. Hind gut with four
Malpighian tubules which end freely in the body cavity near the rectum. Male
reproductive system (text-fig. 249) with the lateral accessory glands not divided.
Each testis consisting of two sperm tubes. Female reproductive system (text-
fig. 250) with seven egg tubes to each ovary. Spermathecal duct opening into
apex of bursa copulatrix. Central nervous system with three thoracic discrete
ganglia. First abdominal ganglion partly fused to third thoracic, two to five
discrete, and six to eight partly fused together, though the limits of each are
distinguishable.
Genotype : Elmis smithi Grouvelle (1898).
H. smithi is the only member of this genus that has been available for an
examination of the internal anatomy. The majority of the species are so close
in external characters that they can be separated only on the structure of the
MEXICAN ELMIDAE 333
male genitalia which generally exhibit very striking specific differences. The
larvae are still unknown.
The following species should be referred to Hexacylloepus Hinton : H. abdo-
minalis (Hinton) (Cylloepus), H. abditus (Hinton) (Cylloepus), H. aciculus (Hinton)
(Cylloepus), H.fetruginea (Horn) (Cylloepus), H.flavipes (Grouvelle) {Cylloepus), H.
granosus (Grouvelle) (Cylloepus), H. granulosus (Sharp) (Elmis), H. grouvellei
(Grouvelle) (Cylloepus), H. horni (Hinton) (Cylloepus), H. indistinctus (Hinton)
(Cylloepus), H. plaumanni (Hinton) (Cylloepus), H. subsulcalus (Grouvelle)
(Cylloepus), and H. sulcatus (Grouvelle) (Cylloepus).
On external characters this genus can be separated from Cylloepus only by
having a complete transverse belt of tomentum on the hypomera. The internal
anatomy, however, is not at all close to that of Cylloepus, differing as follows : (1)
he anterior margin of the mid-gut has six instead of eight caeca ; (2) the hind
gut has only four instead of six Malpighian tubules ; (3) the lateral accessory
glands of the male are not divided into lobes ; (4) each ovary has seven instead
of about 18 egg tubes ; and (5) the spermathecal duct opens into the apex instead
of the base of the bursa copulatrix.
The species are confined to the Americas where they are found from the
southern United States to south Brazil. They also occur in the West Indies.
The specific characters of most importance in separating the species are
similar to those listed for Cylloepus. The following secondary sexual characters
have been observed in the species before me.
(1) Female with a broad depression on the apex of the fifth abdominal
sternite (abdominalis).
(2) Male with a long and acute tooth on the mesal margin of the hind coxa
(sp. n., not yet described, Brazil).
(3) Male with a row of close, stout, and short teeth on inner side of middle
tibiae (abditus).
A Key to the Mexican Species of Hexacylloepus.
1. Abdomen with an oval depression on the apex of the last sternite.
Mexico ........ H. apicalis, sp. n.
Abdomen without a depression on the apex of the last sternite . . 2.
2. Median longitudinal impression of pronotum not present on basal fourth,
or if present it is here very narrow and shallow so that it is scarcely
noticeable. Texas ..... H. jerruginea (Horn) (1870).
Median longitudinal impression of pronotum deep and broad on basal
fourth 3-
3. Male genitalia with the median lobe spatulate. Mexico H. scabrosus, sp. n.
Male genitalia with the median lobe evenly narrowed to the apex; with a
row of short and stout spines on ventral side of middle tibia . . 4
4. Male genitalia with the median lobe not twice as long as parameres.
Mexico H. abditus (Hinton) (1937).
Males without a row of short and stout spines on ventral side of middle
tibia. Male genitalia with the median lobe more than twice as long
as parameres. Mexico . . . . H. horni (Hinton) (11)37)
H . Jerruginea (text-fig. 259) is included in this key, as it is already known from
localities very near to the Mexican border and will undoubtedly be found in
Mexico later on.
334
H. E. HINTON
Hexacylloepus apicalis, sp. n.
(Text-figs. 252, 254-258.)
Male : Length, i-8 mm.-i-95 mm. ; breadth, 0-92 mm. Clothed with fine
short (about 0-025 mm- l°ng)> recumbent, testaceous hairs which arise mostly
at intervals equal to slightly less than their lengths ; antennae similarly clothed
253
Text-figs. 251-253. — (251) Hind wing of Cylloepus pimcticollis (Hinton). (252)
Hind wing of Hexacylloepus apicalis Hinton. Venation after Forbes. (253) Micro-
trichia of apex near inner margin of wing of C. puncticollis.
but with the hairs sparser, slightly longer, and less recumbent ; apical portion
of labrum clothed with equally fine but longer (0-050 mm. or more) and paler
hairs which are more erect, much denser, and usually confined to sides of labrum.
Cuticle feebly shining and for the most part finely alutaceous ; piceous to rufo-
piceous ; antennae, mouth-parts and legs somewhat rufo-testaceous ; tomentum
cinereous with golden reflections. Head with a feeble, broad, slightly oblique
impression on each side near anterior half of eyes extending to a line drawn across
MEXICAN KLMIDAE
335
head from anterior margin of antennal cavities. Antennae as figured (text-fig.
256). Clypeus with the fronto-clypeal suture well impressed and transverse ;
anterior margin broadly and very feebly arcuately emarginate, with the angle on
each side obtuse and bluntly rounded ; sides feebly arcuate. Labrum with the
anterior margin broadly and feebly rounded. Surface set with round to feebly
oblong and apparently flat-topped granules, which are about as coarse or slightly
coarser than facets of eyes and are mostly separated by less than a third to once
their diameters ; granules on clypeus finer ; labrum without granules and with
the very fine punctures separated by once to twice their diameters. Pronotum
at broadest point which is near basal half broader than long (075 mm. : 0-62 mm.)
and base broader than apex (0-65 mm. : 0-48 mm.). Apical margin as seen from
above strongly arcuate at middle and deeply sinuate on each side before apical
256
254
2 58
259
257
Text-figs. 234-258. — Hexacylloepus apicalis Hinton. (254) Dorsal view of male geni-
talia. (255) Left lateral view of same. (256) Antenna. (257) Presternum. (258)
Adult to show general appearance.
TEXT-FIG. 259. — Hexacylloepus ferruginea (Horn). Dorsal view of male genitalia.
angle behind eye ; apical angles acute, strongly produced forwards and slightly
inwards ; sides moderately arcuate, slightly more strongly so at basal half and
feebly sinuate before basal angles ; lateral margins finely and rather regularly
crenate, this crenation being due to granules placed on sides ; basal angles acute,
feebly produced outwards, inconspicuous ; base trisinuate, broadly and moderately
strongly so on each side and shortly and very shallowly so in front of scutellum.
Pronotum with the sublateral carina prominent, slightly converging towards
apex, moderately strongly sinuate at basal half, and becoming obsolete at apical
seventh ; median longitudinal impression extending from near base to very near
apex where it becomes obsolete, and broadest from basal fourth to apical half
where it is slightly broader than scutellum. Surface apparently microscopically,
confluently granulate throughout ; also set with much larger granules as follows
sides between lateral margins and sublateral carinae with the granules about a^
fine as those of head, usually round and separated mostly by two to three times
their diameters ; granules on outer side of carinae slightly coarser, often more
336 H. E. HINTON
oblong, and separated by about once their lengths ; sides of disk near sublateral
carinae granulate as area near lateral margins ; area on each side of median
impression with the granules slightly coarser and seldom separated by more than
once their diameters ; bottom of median impression without the coarser granules.
Base of pronotum without oblique impressions ; disk near sinuation of sublateral
carinae with a very shallow, very broad, and indefinitely bounded impression.
Elytra slightly more than twice as long as pronotum (1-40 mm. : 0-62 mm.) and
feebly broadening posteriorly to broadest point which is at apical third and here
broader than base of pronotum (0-92 mm. : 0-65 mm.). Apices feebly produced,
conjointly broadly and feebly rounded. Lateral margins finely and regularly
crenate, the crenation being due to fine lateral granules. Surface striate with
the striae becoming finer towards apex and obsolete at apical fifth ; discal strial
punctures round to feebly subquadrate, moderately deep, about a half to a third
as broad as intervals, and separated mostly by two to three times their diameters ;
these strial punctures become finer towards apex and are nearly obsolete at
apex. Discal intervals feebly convex and subequal in breadth, at base all except
sutural more strongly convex ; surface of intervals without the microscopic
granules of pronotum but with coarser granules similar to coarse ones of pro-
notum and generally slightly sparser ; sublateral carinae similarly but more
densely granulate. Scutellum flat, subovate, broader than sutural interval
(o-io mm. : 0-06 mm.), longer than broad (o-n mm. : o-io mm.), feebly and
broadly rounded basally, and slightly narrowed at apex ; surface granulate
similarly to adjacent portions of elytra. Prosternum with the process as figured
(text-fig. 257) ; middle area of prosternum moderately coarsely and very densely
granulate and subrugose ; sides more finely, more sparsely, and less rugosely
granulate. Mesosternum strongly depressed and with a large and deep median
pit ; surface more finely sculptured than surface of prosternum. Metasternum
moderately depressed posteriorly, with a fine median longitudinal impression
which is traceable nearly to anterior margin, sides with the granules finer and
separated by two to three times their diameters. First abdominal sternite with
the middle basal portion strongly depressed and the surface of the depression
slightly less coarsely and rugosely granulate than metasternum ; carinae of first
sternite scarcely developed, very indistinct. Surface of sides of first sternite
and all of other sternites with granules which are slightly finer than those of
pronotum and usually separated by two to three times their lengths. Apex of
last sternite with a large (about a third as long as segment), oval, moderately
deep impression which is free of granules and somewhat polished. Genitalia as
figured (text-figs. 254, 255).
Female : Externally similar to male.
Type : <J in the British Museum (Nat. Hist.). Mexico : Estado de Morelos,
Cuernavaca, vi.1934 (H. E. Hinton).
Paratypes : 176, collected at the same time as the type ; and 1, Mexico :
Dist. de Temascaltepec, Tejupilco, alt. about 4000 ft., vii.1934 (H. E. Hinton).
Variations : The series before me is very uniform in size and structure, but
there are slight differences in the density of the granules on the various sclerites,
and the apical pit on the last abdominal sternite is subject to a slight variation in
extent and depth.
Comparative notes : Both males and females of this species may be separated
from all other species of the subgenus except H. abdominalis (Hinton) by the
pit on the apex of the last abdominal sternite. The males of H. abdominalis
have no pit on the apex of the abdomen, and the females may be distinguished
MEXICAN ELMIDAE 337
from both males and females of apicalis as follows : (1) the median longitudinal
impression of the pronotum is not present on basal fourth in abduminalis , while
on this part of the pronotum it is very distinct in apicalis ; and (2) the sublateral
carinae of the elytra are much less prominent in abduminalis than they are in
apicalis. These two species are so widely separated geographically, one being
from southern Brazil and the other from central Mexico, that there is little
chance of their being confused in collections.
Hexacylloepus scabrosus, sp. n.
(Text-figs. 260-261.)
Male : Length, 2-0 mm. ; breadth, 0-95 mm. Clothed with fine, short
(about 0-025 mm- !°ng) recumbent, testaceous hairs which arise mostly at intervals
equal to slightly less than their lengths ; antennae similarly clothed but with the
hairs much sparser (the vestiture of the antennae cannot be more accurately
described, as all specimens before me are badly rubbed) ; apical portion of labrum
clothed with equally fine but longer (0-050 mm. or more) and paler hairs which
are more erect, much denser, and mostly confined to sides of labrum. Cuticle for
the most part finely alutaceous and shining ; piceous to rufo-piceous ; antennae,
mouth-parts, and legs paler rufo-piceous. Tomentum cinereous with golden
reflections. Head with a feeble, broad, slightly oblique impression on each side
near anterior half of eyes extending to slightly behind an imaginary line drawn
across head from anterior margin of antennal cavities. Clypeus with the fronto-
clypeal suture nearly straight and well impressed ; anterior margin very broadly
and very feebly arcuately emarginate ; angle on each side obtusely and bluntly
rounded ; sides feebly arcuate. Labrum with the anterior margin broadly
rounded in front, with the angle on each side broadly rounded. Surface set with
round to feebly oblong granules which are apparently flat-topped, as coarse as
facets of eyes, and separated mostly by less than a third to once their lengths ;
granules on clypeus slightly finer ; labrum without granules and punctate with
very fine, microscopic punctures which are separated mostly by once to twice
their diameters. Pronotum at broadest point near basal half broader than long
(0-82 mm. : 0-70 mm.) and base broader than apex. Apical margin as seen from
above moderately arcuate at middle and broadly and deeply sinuate on each side
behind eye before apical angles ; apical angles acute, strongly produced forwards
and slightly inwards ; sides moderately arcuate, slightly more strongly so at
basal half and scarcely noticeably sinuate before basal angles ; lateral margins
finely and rather regularly crenate, this crenation being due to granules placed
on sides ; basal angles feebly acute, nearly rectangular, and feebly produced
backwards ; base trisinuate, broadly and moderately strongly sinuate on each
side and shortly and very shallowly sinuate in front of scutellum. Pronotum
with the sublateral carina prominent, slightly converging towards apex, moderately
sinuate at basal half, and becoming obsolete at apical seventh ; median longi-
tudinal impression extending from near base to near apex where it becomes
obsolete, broadest from basal fourth to apical third where it is slightly broader
than scutellum ; base of pronotum without oblique impressions ; disk near
sinuation of sublateral carina with a very shallow and broad indefinitely bounded
impression. Surface of pronotum microscopically alutaceous so that it appears to
be minutely and conlluently granulate throughout ; also set with granules as
follows : side between lateral margins and sublateral carina about as fine as those
of head, usually round, and separated mostly by two to three times their diameters;
33«
H. E. HINTON
granules on outer sides of sublateral carina slightly coarser and usually separated
by about once their diameters ; sides of disk near sublateral carina granulate
similarly to but more densely than area near lateral margin ; area on each side
of median impression with granules about as coarse as those on sides of sublateral
carina and about equally dense. Elytra twice as long as pronotum (1-4 mm. :
0-70 mm.) and very feebly broadening posteriorly to broadest point which is at
apical third (here appearing scarcely broader than across humeri) which is broader
than base of pronotum (0-95 mm. : 0-71 mm.). Apices moderately produced,
conjointly broadly and feebly rounded, subtruncate. Lateral margins finely and
regularly crenate, the crenation being due to lateral granules. Surface striate
with the striae becoming finer towards apex and nearly obsolete beyond apical
fourth ; discal strial punctures round to feebly subquadrate, deep, about a third
to a half as broad as intervals, and separated longitudinally by two to three times
Text-figs. 260, 261.
r,o« 261
260
-Hexacylloepus scabrosus Hinton
genitalia.
(260) Dorsal view of male
(261) Left lateral view of same.
their diameters though occasionally they are more closely placed ; these punctures
become finer towards apex and are nearly obsolete at apex. Discal intervals
feebly convex and subequal in breadth, at base all except sutural more strongly
convex ; surface of intervals with the alutaceous microsculpture not similar to
that of pronotum ; also set with granules which are slightly coarser and sparser
but otherwise similar to those of pronotal disk ; sublateral carinae slightly more
densely granulate but with the granules here as elsewhere on elytra mostly
round. Scutellum flat, subovate, broader than sutural interval (o-io mm. : 0-06
mm.), longer than broad (o-n mm. : o-io mm.), feebly and broadly rounded
basally, and slightly narrowed at apex ; surface granulate similarly to adjacent
portion of elytra. Prosternum with the margins of the process somewhat raised
so that the median area appears slightly concave at middle ; middle area of
prosternum moderately coarsely and very densely rugosely granulate ; sides
more finely, more sparsely, and also less rugosely granulate. Mesostemum
strongly depressed and with the median pit deep and large ; surface slightly
more finely sculptured than middle area of prosternum. Metasternum moderately
feebly depressed posteriorly ; with a fine median longitudinal impression which
MEXICAN ELMIDAE
339
is traceable nearly to anterior margin ; disk sculptured as middle area of pro-
sternum ; sides with the granules slightly finer and separated by two to three
times or more than their diameters. First abdominal sternite with the middle
basal portion moderately depressed and here with the surface nearly as coarsely
and rugosely granulate as that of disk of metasternum ; carinae of first sternite
not evident ; sides of basal sternite and entire surface of other sternites with
granules which are slightly finer than discal pronotal ones and are usually separated
by two to three times their diameters. Genitalia as figured (text-figs. 260, 261).
Female : Externally similar to male.
Type : q in the British Museum (Nat. Hist.). Mexico : Estado de Morelos,
Cuernavaca, vi.1934 (H. E. Hinton).
Paratypes : 2 cJcJ and I 9 with same data as type.
Variations : In the small series before me no variations have been observed.
Comparative notes : This species is nearly identical in external characters to
H . apicalis, as may be seen from a comparison of the two descriptions. The only
external difference which can be relied on to distinguish the two is the apical
abdominal pit of both sexes of apicalis. The four specimens before me have
the prosternal process more strongly concave along the middle and the margins
of the process more thickly and strongly raised than is the case in apicalis,
but these slight differences may be bridged when a longer series is studied.
Hexacylloepus abditus (Hinton).
(Text-figs. 262-265.)
1937. Cylloepus abditus Hinton, Arb. morph. taxon. Enl. Berlin-Dahlem, 4 (2) : 106, figs.
13-16.
Male : Length, 17 mm. ; breadth, 077 mm. Clothed with fine, short
(about 0-025 mm- long), recumbent, testaceous hairs which arise mostly at
intervals equal to slightly less than their own lengths ; antennae similarly clothed
but with the hairs sparser and less recumbent ; apical portion of labrum clothed
with equally fine but longer (about 0-050 mm., but occasionally about 0-062 mm.)
and paler testaceous hairs which are more erect, much denser, and usually con-
fined to sides of apical portion. Cuticle for the most part finely alutaceous and
shining ; piceous to rufo-piceous ; antennae, mouth-parts and legs paler. Tomen-
tum cinereous with golden reflections. Head with a feeble, broad, slightly
oblique impression extending on each side near anterior half of eyes to a point
opposite anterior margins of antennal cavities. Clypeus with the fronto-clypeal
suture nearly straight and well impressed ; anterior margin broadly and feebly
arcuately emarginate, with the angle on each side broadly rounded and the sides
feebly arcuate. Labrum broadly and feebly rounded in front, with the angle
on each side broadly rounded. Surface set with round to feebly oblong, flat-
topped granules which are nearly as coarse as facets of eyes and are usually
separated by twice their diameters though often by much less ; granules on
clypeus finer and denser ; labrum without granules but punctate with fine,
microscopic punctures which are mostly separated by once to twice their dia-
meters. Pronotum at broadest point which is near basal half broader than long
(0-062 mm. : 0-52 mm.) and base broader than apex (0-57 mm. : 0-42 mm.).
Apical margin as seen from above moderately strongly arcuate at middle and
deeply sinuate on each side behind eye before apical angle ; apical angles
moderately acute, moderately strongly produced forwards and slightly inwards ;
340
H. E. HINTON
sides moderately arcuate but slightly more strongly so at basal half, scarcely
noticeably sinuate just before basal angles. Lateral margins feebly and rather
regularly crenate, this crenation being due to granules placed on sides ; basal
angles acute and very feebly produced backwards ; base trisinuate, broadly and
deeply so on each side and shortly and more shallowly sinuate in front of scutellum.
Pronotum with the sublateral carina prominent, slightly converging towards apex,
moderately strongly sinuate on basal half, and becoming obsolete at about
apical fourth ; median longitudinal impression extending from near base to near
apex where it becomes obsolete and broadest from basal fourth to apical third,
where it is slightly broader than scutellum ; base of pronotum without oblique
Text-figs. 262-267. — (262) Dorsal view of male genitalia of Hexacylloepus abdilus
(Hinton). (263) Left lateral view of same. (264) Antenna. (265) Prosternum.
(266) Dorsal view of male genitalia of Hexacylloepus hovni (Hinton). (267) Left
lateral view of same.
impressions ; disk near sinuation of sublateral carina with a shallow, broad,
indefinitely bounded impression. Surface microscopically alutaceous in such a
manner as to appear confluently granulate throughout ; also set with distinct
granules as follows : Sides between sublateral carinae and lateral margins set with
granules about as fine as those of head, usually round, and separated mostly by
two to three times their diameters ; granules on outer sides of sublateral carinae
slightly coarser and usually separated by once to twice their diameters ; sides of
disk near sublateral carina granulate as area near lateral margin ; area near
median impression with the granules slightly coarser and denser than those of
sides of disk. Elytra more than twice as long as prothorax (1-18 mm. : 0-52 mm.)
and feebly broadening posteriorly to broadest point, which is at apical third and
is broader than base of pronotum (0-77 mm. : 0-57 mm.). Lateral margins
finely and regularly crenate, the crenation being due to fine lateral granules.
Surface with the striae becoming finer towards apex and all except sutural obsolete
MEXICAN ELMIDAE 341
beyond apical sixth ; discal strial punctures round to feebly subquadrate,
moderately deep, about a third to a half as broad as intervals, and separated
longitudinally usually by two to three times their diameters ; these strial punc-
tures become finer towards apex and at apical fifth they arc shallow, sparse,
and only about a third to a sixth as coarse as discal intervals ; discal intervals
subequal in breadth and feebly convex, at base with the fourth interval more
strongly convex (there is much variation in this respect and sometimes none are
convex) ; surface of intervals at base alutaceous somewhat as on pronotum but
elsewhere on elytra the microsculpture does not appear granulate ; granules
mostly similar in size and density to those of pronotum ; granules on carinate
intervals slightly larger and denser but also round to feebly oblong as elsewhere
on elytra. Apices moderately produced, conjointly broadly and very feebly
rounded. Scutellum flat, subovate, broader than sutural interval (0-09 mm. :
0-06 mm.), longer than broad (o-io mm. : o-og mm.), feebly and broadly rounded
basally, and slightly narrowed to apex ; surface granulate similarly to adjacent
portion of elytra. Prosternum with the process as figured (text-fig. 265) and
feebly concave with the lateral margins somewhat raised ; middle area of pro-
sternum with the granules as coarse as those of elytra but with the surface also
densely rugose, at sides much less rugose and with the granules finer and sparser.
Mesosternum strongly depressed and with a median pit at the bottom of the
depressed area. Metasternum moderately strongly depressed posteriorly ; with a
fine, median longitudinal line which is traceable to nearly anterior margin ; disk
sculptured as middle area of prosternum but with the granules usually slightly
larger ; sides less rugose and with the granules sparser. Middle portion of first
abdominal sternitc moderately strongly depressed and with the surface of this
depression sculptured as sides of metasternum ; first sternitc without obvious
carinae ; sides of first sternite and all of surface of other sternites with granules
which are about as coarse as discal pronotal ones and are usually separated by
less than to twice their lengths. Middle tibiae with a row of small, short, stout,
and close teeth on ventral side. Genitalia as figured.
Female : Externally similar to male but without the row of spines on the
ventral side of middle tibiae.
Type : £ in the British Museum (Nat. Hist.). Mexico : Dist. de Temascal-
tepec, Tejupilco, alt. about 4000 ft., vi.1934 (H . E. Hinton).
Specimens examined : 56, with same data as type but collected on 16. vi. 1933
(H. E. Hinton, R. L. Usinger).
Variations : No variations worthy of mention have been noted in the series
available.
Comparative notes : The males of this species may be distinguished from those
of H. scabrosiis by the row of fine teeth on the middle tibiae and the differently
formed male genitalia, but tin- females, as far as I know, arc inseparable. H.
abditus and H. scabrosiis have not yet been found to overlap geographically.
Hexacylloepus horni (Hinton).
(Text-figs. 266, 267.)
!937- Cylloepus horni Hinton, Arb. morph. (axon. Ent. Beriin-DahUm, 4, (2) : 109, figs.
17, 18.
Male : Length, 17 mm. ; breadth, 0*8o mm. 1 have written a description
of nearly iooo words for this species, but a comparison of this description with
342
H. E. HINTON
that of abditus shows that the two are identical in nearly every respect, so that
to give the description of horni here would be a waste of space. For an under-
standing of the external characters of this species the student is referred to the
description of abditus. The males of the new species differ from those of abditus
by not having a row of short teeth on the ventral side of the middle tibiae and by
the structures of the male genitalia (cf. figs.). Both males and females of the two
species may be readily separated by the differences in the length of the legs. The
following table is drawn up from a male specimen of each equal in length and very
nearly equal in breadth :
Front femora.
Mnldle femora.
abditus 0-375 mm.
horni i 0-425 mm.
0425 mm.
0-487 mm.
0-40 mm.
0512 mm.
0437 mm.
0-587 mm.
•437 mm.
•537 mm.
0500 mm.
0650 mm.
Female : Externally similar to male.
Type : S in the British Museum (Nat. Hist.). Mexico : Dist. de Temascal-
tepec, Tejupilco, alt. 3500-4000 ft., vii.1934 {H. E. Hinton).
Specimens examined : 2, Mexico : Sierra de Durango, 1922 (C. Schaufuss).
Comparative notes : Apart from the characters of the male genitalia, this
species is inseparable from H. scabrosus. The latter has the length of the legs
intermediate between abditus and horni.
CYLLOEPUS Erichson.
1847. Cvlloepus Erichson, Naturges. Ins. Deutschl., 3 : 521.
1854. Cvlloepus Lacordaire, Gen. Col., 2 : 510.
1882. Cvlloepus Sharp, Biol. Centr.-Amer. Col., 1 (2) : 129.
This genus was erected by Erichson (1847) to contain Limnius araneolus
Miiller (1806) of Peru. Erichson had before him a male of C. araneolus, and a
good proportion of his brief generic diagnosis is devoted to describing the apical
segment of the labial palpus which in this, as well as in the males of many other
species of Cylloepus, is very much broadened at the apex.
The generic limits of this genus have been little understood. Grouvelle
correctly assigned five species to this genus and described many others as species
of Elmis Latreille and one as an Elsianus Sharp. Sharp (1882) in his work on the
Central American Elmidae correctly referred one species to Cylloepus and placed
a number of others in Elmis Latreille. Darlington (1927) described a number of
West Indian forms under the name of Elmis. These species were later correctly
placed in Cylloepus by Musgrave (1935), but were again assigned to Elmis by
Darlington (1936). A redescription of Cylloepus (sensu stricto) follows :
Body elongate, subparallel. Dorsal surface clothed with short, sparse or
dense, usually recumbent hairs. Tomentum confined to the following areas :
(1) genae ; (2) epipleurae ; (3) sides of prosternum, mesosternum, metasternum,
and abdominal sternites, but in a few species nearly the entire ventral surface
may be clothed with scale-like or hairy tomentum ; and (4) all of legs except
tarsi, though in most species the scale-like tomentum is sparse or absent on the
tibiae. The hypomera are nearly always without a trace of tomentum, and only
rarely is there a very narrow belt of tomentum along the anterior portion of the
sterno-pleural suture. Head when seen from below capable of being retracted,
so that none of the mouth-parts is visible. Antennae n-segmented. Mandible
MKXICAN ELMIDAE
343
344
H. E. HINTON
with three acute apical teeth ; prostheca large, entirely membranous and with the
apex spinose or hairy. Maxillary palp 4-segmented and stipes with a well-
developed palpifer ; galea and lacinia separate and apex of each densely spinose.
Labial palp 3-segmented and prementum with a palpiger. Mentum transverse
and about as broad and long as submentum. Gula anteriorly nearly as broad as
submentum and slightly narrowed posteriorly. Pronotum with a sublateral
longitudinal carina on each side generally extending from base to apex, though
in a few species this carina does not quite reach either base or apex ; with a
median longitudinal impression on the disk ; with a shallow impression which
Text-figs. 273,
Grouvelle.
2 74
274. — (273) Female reproductive system of Cylloepits consobnnus
(274) Female genitalia of Cylloepits punclicollis (Hinton).
extends obliquely outwards from base on each side of scutellum to sublateral
carina which it bisects at about basal third. Elytra striate and punctate ; without
accessory striae ; and with two or very rarely (scalptipennis Sharp) one sub-
lateral carina. Hind wing (text-fig. 251) without a radial cross vein and without
an anal cell ; with the first anal absent ; second anal with the first and second
branches present ; third anal with a second branch; cubito-anal cross vein com-
plete and joining cubitus to second anal. P roster num. (text-fig. 278) very long
in front of anterior coxae ; prosternal process long and posterior margin usually
broadly rounded. Mesosternum with a deep and moderately narrow groove for
the reception of the prosternal process. Metasternum with a median longitu-
dinal impressed line. Legs with the visible portion of the coxae rounded and
trochantin completely concealed by the hypomera. Claws without teeth. Ali-
mentary canal (text-fig. 268) with eight caeca on the anterior margin of the
MEXICAN ELMIDAE 345
mid-gut. Hind gut with six Malpighian tubules which end mar the rectum freely
or imbedded in fatty tissue. Male reproductive system (text-fig. 272) with the
lateral accessory glands divided into several lobes. Each testis of two sperm
tubes. Female reproductive system (text-rig. 275) with about 18 egg tubes to
each ovary. Spermathecal duct opening at base of bursa copulatrix. ( entral
nervous system with three discrete thoracic ganglia. First abdominal ganglion
partly fused to third thoracic ; two to five tree ; and six to eight partly fused
together, though the limits of each are distinguishable.
Genotype : Limnius araneolus Midler (1806).
The internal anatomy of three species has been examined and found to
agree in essential details. ('. consobrinus Grouvelle of Bolivia is figured.
In external appearance this genus is very close to Stenelmis 1 )ufour from which
it mav lie distinguished as follows : (1) There is no accessory stria on each elytron
at base between the first and second striae ; (2) the inner apex of each tibia is
densely pubescent, while in Stenelmis this part of the tibia is without a fringe of
hairs ; and (3) there are only two instead of three sperm tubes to each testis.
This genus is confined to the Americas, where it occurs from Southern United
States to Chili and is well represented in the West Indies.
The specific characters of greatest importance in separating the species oi
Cylloepus seem to be the following :
(1) General proportions, length and breadth.
(2) Si/e and density of the punctures of all the sclerites and the typi "1
microsculpture between the punctures.
(3) Extent and depth of the impressions on the head.
(4) Condition of fronto-clypeal suture.
(5) Anterior margin of clypeus, whether truncate, rounded, or emarginate,
and shape of angle on each side.
(6) Condition of anterior margin of labium.
(7) Colour. In many species the antenna is bicoloured.
(8) Outline of pronotum and the extent and depth of the various impre-
sions on its surface. Shape and extent of the sublateral carinae.
(9) Shape of elytral apices.
(10) Condition of lateral margin of elytra, whether crenate or smooth.
(11) Number of sublateral elytral carinae and number of carinate or convex
intervals.
(12) Depth and extent of striae and strial punctures.
(13) Shape of scutellum and if convex or flat.
(14) Shape of prostemum and prosternal process.
(15) Impressions on disk of metasternum.
(16) Condition of carinae on first abdominal sternite.
(17) Extent and depth of depression on middle of first abdominal sternite.
(18) If a depression is present on the second abdominal sternite its depth
and extent should be described carefully.
(iq) Depth and extent of depression sometimes present on apex oi fifth
abdominal sternite,
(20) Secondary sexual characters. These are, as in most genera "I 1 In
family, among the most highly specific characters.
(21) Structure oi the male genitalia. The structure of this organ is more
important than that oi any other in associating specimens with
specific descriptions. Unless a species is exceptionally distinct, it
should not be described without illustrating the genitalia.
novit. zool., 42, 2. 1940. -7
346 H. E. HINTON
Secondary sexual characters are abundant in Cylloepus, and the following
have been observed in the species before me.
(1) Male with the apex of the last segment of the labial palp very much
broadened (araneolus, barberi, consobrimis, palpalis, puncticollis,
opt at 11s).
(2) Male with the mesosternum on each side of the middle with a strong
gibbosity, at the apex of which is a group of erect hairs, while in the
female this portion is only slightly convex and without hairs (spinipes).
(3) Disk of metasternum not as strongly nor as broadly depressed in the
females as in the males (most species).
(4) Female with an acute tubercle on each side of metasternum in front
of mesal margin of hind coxa (ventralis).
(5) First abdominal sternite at middle more strongly depressed in male
than in female (most species).
(6) Male with the carinae of first abdominal sternite more convex than
those of female and curving inwards while in the female they are
straight (sculptipennis).
(7) Male with the carinae of the first abdominal sternite relatively longer
than those of the female (sculptipennis).
(8) Second abdominal sternite depressed in male and convex or flat in
female (sculptipennis, sexiialis).
(9) Male without a tooth-like tubercle on mesal margin of hind coxa
(spinipes).
(10) Male with three large teeth on ventral apical fourth of front tibia
(spinipes).
(11) Male with a carina-like swelling on inner side of front tibia near apex
(sexualis).
(12) Male with a row of close, stout, and short teeth on inner side of middle
tibiae (barberi, optatus, sexualis).
(13) Male with a large and flat tooth on inner half of hind tibia (spinipes).
(14) Male with the hind tibia at middle of its length more swollen and
curved than that of the female (optatus, sexualis).
(15) Male with numerous pale and erect hairs on the ventral surface of the
first four segments of the front tarsi (optatus).
A Key to the Mexican Species of Cylloepus.
1. Elytra on each side with only one distinct sublateral carina. Guatemala,
Mexico C. sculptipennis (Sharp) (1882).
Elytra with two distinct sublateral carina on each side . . . .2.
2. Base of pronotum in front of scutellum with an impression which is about
as broad as scutellum and half as deep. Elytra with the basal discal
intervals flat and with the surface of the intervals not granulate.
Guatemala, Costa Rica . . . . C. barberi Hinton (1934).
Base of pronotum in front of scutellum without an impression ; or if one
is present it is very shallow and scarcely noticeable. Elytra with the
basal discal intervals always granulate if flat ..... 3.
3. Elytra with none of the basal discal intervals strongly convex. Mexico
C. blairi Hinton (1936).
Elytra with one or more of the basal discal intervals strongly convex . 4.
4. Elytra with the fourth discal interval strongly convex at base . . 5.
Elytra with the fourth discal interval flat .... . . 7.
MEXICAN ELMIDAE 347
5. Elytra with the third disca] interval flat. Guatemala, Mexico
C. heterocerus (Sharp) (1882).
Elytra with the third discal interval near base nearly or more strongly
convex than fourth ......... 6.
6. Apical segments of antennae black, or if not black at least distinctly
darker than the two basal segments. Pronotum with the disk punctate
on either side of median longitudinal impression. Males with the last
segment of the labial palp very much broadened at apex ; metasternaJ
disk only feebly depressed ; abdomen with the first sternite depressed
at middle from base to apex ; front and hind tibiae without spines.
Mexico ...... C. puncticollis (Hinton) (1934).
Antennae unicolorous. Pronotum with the disk granulate on either side
of the median longitudinal impression. Males with the last segment
of the labial palpi not strongly broadened at apex ; metasternal disk
strongly depressed ; abdomen with the first sternite depressed at
middle only on basal three-fourths ; front tibiae on inner apical fourth
with three spines, and hind tibiae on inner side at basal two-fifths
with a large flat spine. Mexico . . . C. spinipes Hinton (1934).
7. Elytra with the fifth interval near base strongly convex for a short
distance. Males with a short and prominent carina-Uke swilling on
inner apical fourth of front tibiae. Mexico . C. sexnalis Hinton (1937).
Elytra with the fifth interval near base at most feebly convex. Males
without a carina on inner apical side of front tibia .... 8.
8. Antennae with the two basal segments pale rufo-piceous and the others
piceous to black. Males with the four basal segments of the front
tarsi not densely clothed with erect hairs on the ventral side ; abdomen
with the middle of the second sternite at most feebly depressed on
extreme base. Mexico. . . . . C. ■proxitnus Hinton (1937).
Antennae unicolorous. Males with the four basal segments of the front
tarsi clothed on ventral side with moderately long, erect, and pale
hairs ; abdomen with the middle of the second sternite moderately
strongly depressed on basal two-thirds. Panama, Guatemala,
Costa Rica C. optatus Sharp (1882).
Two species, C. barberi Hinton and C. optatus Sharp, which are widely dis-
tributed in Central America are included in the key given above, as they may be
found in Mexico when this country is more thoroughly explored.
Cylloepus sculptipennis (Sharp).
(Text-figs. 275-279.)
1882. Elmis sculptipennis Sharp, 81 <l, Centr.-Amer. Col., I (2) : 135.
Mule: Length, 2-8 mm.; breadth, 1 - 1 2 mm. Subparallel, moderately
convex. Clothed with fine, recumbent, brownish hairs which are about 0-05
mm. long and arise at intervals equal to more than once their lengths ; labrum
at sides with the hairs much longer and denser and paler. Cuticle shining and
rufo-piceous to nearly black ; densely and minutely alutaceous on head, pro-
notum, hypomera, prosternal process, meso- and metastemum, all of first
abdominal sternite, and sides "I sternites two to five. tU\id without distinct
impressions; surface densely and very minutely alutaceous so that under a
34«
H. E. HINTON
magnification of x 150 it appears to be finely granulate ; also set with low and
irregularly shaped granules which are slightly finer than facets of eyes and are
usually separated by less than to twice their diameters. Clypeus when viewed
from in front with the anterior margin feebly and arcuately emarginate for its
entire breadth, with the angle on each side broadly rounded ; clypeal suture
feeble and arcuate ; surface similar to that of head "but with only the base and
sides alutaceous. Labrum with the anterior margin truncate and the angle on
each side feebly rounded ; surface with punctures which are not more than half
as broad as granules of clypeus and are usually separated by less than to once
their diameters. Pronotum with the broadest point at basal two-fifths and here
broader than long (0-85 mm. : o-8o mm.) and base broader than apex (o-8o mm. :
0-54 mm.). General shape and form and extent of the various impressions as
278
9. — Cylloepus sculptipennis (Sharp)
Text-figs. 275-.
genitalia. (276) Right lateral view of same
(279) Adult to show general appearance.
(277)
(275) Dorsal view of male
Antenna. (278) Presternum.
figured (text-fig. 279). Surface sculptured as head except for area near inner
side of sublateral carinae on basal half, extreme base, and sides near lateral
margins which are only feebly or not at all alutaceous. Elytra more than twice as
long as prothorax (1-87 mm. : o-8o mm.) and feebly broadening posteriorly to
broadest point which is near apical third and which is distinctly broader than
broadest point across humeri (1-12 mm. : i-oo mm.). Lateral margins coarsely
and densely crenate. Surface coarsely striate, discal striae finer and shallower
as they approach apex but never altogether absent ; discal strial punctures
subquadrate to round, moderately deep, and at middle of disk as broad as intervals
to slightly narrower and separated longitudinally by slightly more than to slightly
less than their own diameters ; towards sides the punctures become coarser and
denser and towards apex finer and sparser. Discal intervals flat, the third at
base being only very slightly convex. Elytra with only one sublateral carina.
Surface of intervals with the granules slightly larger, more convex, and more
regularly round than those of pronotum and separated usually by one to four
MEXICAN ELMIDAE .549
times their diameters : sutural interval on basal two-thirds and first four inter-
vals on apical third with only an occasional granule ; surface between granules
with punctures similar to those of labrum but separated by one to five times
their diameters. Scutellum flat, subovate, broader than sutural interval near
base (o-i2 mm. : 0-05 mm.) and as broad as long ; surface sculptured as adjacent
elytral intervals. Prosternum with the anterior two-thirds (not including process)
moderately strongly but not sharply lobed ; surface granulate as elytral intervals
except for surface of process which is sculptured as head. Hypomera very
densely and regularly alutaceous, with only an occasional granule, and without
t omentum. Mesostemum sculptured as prosternal process. Metasternum with
the posterior third of disk only feebly depressed ; with a broad (about 0-03 mm.)
and deep median longitudinal impression which extends to apical fourth ; surface
of disk similarly but distinctly more coarsely sculptured than surface of pro-
sternal process ; sides of metasternum with the granules only showing through
the tomentum and separated by one to three times their diameters, the surface
here strongly resembling that of the legs. Abdomen with the carinae of the first
abdominal stemite curving slightly inwards, complete from base to apex, and
most prominent at apex ; first stemite at middle strongly depressed, this depres-
sion very shallowly but nevertheless distinctly encroaching to middle of basal
third of second stemite ; middle of first stemite on basal two-thirds densely and
minutely alutaceous and with deep round punctures which are about o-02 mm.
broad and are contiguous to separated by once their diameters ; apical third of
first sternite and all of middle up to apical four-fifths of fifth stemite not alu-
taceous and with the punctures similar to those of first stemite, but becoming
progressively finer as apex of abdomen is approached so that on basal fifth of
fifth sternite they are only half as coarse. Sides of abdominal stemites granulate
as sides of metasternum. Genitalia as figured (text-figs. 275, 276).
Female : Externally similar to male except as follows : (1) the carinae of
the first abdominal sternite straight and not curved inwards, absent on apical
sixth, and least instead of most prominent at apex ; and (2) the depression of the
middle of the first sternite does not extend on to second sternite.
Type : In the British Museum (Nat. Hist.). Guatemala : Rio Naranjo,
450 ft. (Champion).
Specimens examined : 8, Mexico : Estado de Morelos, Cuernavaca, vi.1934
(H. E. Hinton) ; 2, Mexico : Dist. de Temascaltepec, alt. 6000-7000 ft., vi.1934
(H. E. Hinton) ; and 1 with same data as above but at Tejupilco, alt. about
4000 ft., vii.1934.
1 ' dilations : Xo variations worthy of mention have been noted.
Comparative notes : This is the only species of Cylloepus known to-day which
has only one sublateral elytral carina, the usual outer one being completely
absent. In general appearance it is close to C. blairi Hinton. The mouth-parts
and wing venation are like those of typical members of the genus.
Cylloepus blairi Hinton.
(Text-figs. 280-283.)
1936, ( ylln, pus blairi Hinton, Ent. num. Mug., 72: 1, iij^s. 1—4.
Male: Length, 3-0 mm.; breadth, 1-2 mm. Subparallel, moderately
convex. Clothed with fine, short (about 0-037 mm- l°ng)' recumbent, brownish-
testaceous hairs which arise mostly at intervals equal to about their own lengths ;
350
H. E. HIXTOX
antennae (all specimens before me are badly rubbed) with a few equally fine but
more erect hairs at apical angles of segments ; apical portion of labrum with
equally fine but much longer (often o-o6 mm.) hairs which are apparently con-
fined to the sides ; beneath with the hairs generally finer and shorter but other-
wise similar. Cuticle moderately shining and for the most part finely alutaceous ;
piceous to rufo-piceous ; tomentum of body cinereous with golden reflections.
Head on each side near anterior one-half of eyes with a feeble, scarcely noticeable,
broad and oblique impression. Antennae as figured (text-fig. 282). Clypeal
suture well-marked ; anterior margin of clypeus very broadly and very feebly
282
283
Text-jigs. 280-283. — Cylloepus blairi Hinton. (280) Dorsal view of male genitalia.
(281) Right lateral view of same. (282) Antenna. (283) Presternum.
arcuately emarginate ; angle on each side obtusely rounded ; sides feebly arcuate.
Labrum broadly and feebly rounded in front, nearly truncate ; anterior angles
broadly rounded. Surface with the alutaceous microsculpture appearing con-
fluently granulate ; set with usually round granules which are about as coarse as
facets of eyes and are separated mostly by once to twice their diameters ; granules
on clypeus finer, middle apical margin nearly smooth and with a few fine punctures ;
labrum without granules, apical half punctate with fine punctures which are
separated by once to twice their diameters. Pronotum at broadest point near
basal half broader than long (0-925 mm. : 0-825 mm.) and base broader than
apex (0-90 mm. : o-6o mm.). Apical margin when seen from above moderately
arcuate and deeply sinuate on each side behind eye before apical angle ; apical
angles acute and moderately strongly produced forwards and slightly inwards ;
sides moderately arcuate, scarcely noticeably sinuate before basal angles, some-
what regularly, finely crenate ; basal angles nearly rectangular, scarcely
MEXICAN ELMIDAE 351
produced ; base moderately strongly trisinuate, broadly so on each side and very
shortly so in front of scuteUum. Pronotum with the sublateral carinae prominent,
slightly converging towards apex, feebly sinuate at apical and basal thirds,
complete from base to apical margin, and very broad from basal to apical third.
Surface sculptured as follows : a very broad and feebly raised portion extends
from base in front of scutellum to base of disk ; on each side of this raised portion
with a feeble, moderately broad, feebly curved, and oblique impression extending
to a broad, moderately feeble, indefinite impression near basal sinuation of sub-
lateral carina ; median longitudinal impression beginning at base and ending
at about apical one-half, throughout very shallow, broadest at basal two-fifths
where it is as broad as scutellum. Surface with the alutaceous microsculpture
similar to that of head ; disk with the granules slightly coarser than facets of
eyes but rather low [i.e. not strongly convex) and usually separated by two to
three times their diameters, though often, especially on apical portion of disk,
more sparsely placed ; granules on sides slightly denser but otherwise similar ;
granules on basal portion of sublateral carina slightly more convex and very
slightly denser. Elytra more than twice as long as prothorax (1-87 mm. : 0-82
mm.) and very feebly broadened posteriorly to broadest point at apical one-
third which is broader than base of pronotum (1-22 mm. : o-go mm.). Apices
moderately produced, conjointly broadly and feebly rounded. Lateral margins
moderately strongly and regularly crenate. Discal surface rather coarsely
striate basally, with the striae becoming finer towards apex, and only very
feebly impressed beyond apical third ; -discal strial punctures on basal half
nearly subquadrate, deep, about as broad as intervals, and usually separated
longitudinally by little more than their own diameters, these strial punctures
suddenly becoming nearly obsolete beyond apical half but at sides they are con-
tinued further towards apex. Discal intervals flat in apical half and (except
sutural) feebly convex on basal half and still more strongly convex at base, with
the third interval at base very feebly elevated ; surface of intervals, especially
at base, alutaceous somewhat similarly to pronotum ; set with granules which
are similar to pronotal ones but are more convex and on disk become much
sparser towards apex ; carinate lateral intervals with the granules similar but
slightly denser. Scutellum flat, subovate, broader than sutural interval (o-io
mm. : 0-087 mm.), longer than broad (0-12 mm. : o-io mm.), feebly and broadly
rounded at base, and slightly narrowed to apex ; surface sculptured similarly
to adjacent elytral intervals but with the granules slightly sparser. Beneath
with the surface alutaceous as pronotum. Prosternal process as figured (text-
fig. 283) ; middle area of prosternum feebly depressed, slightly rugose, and
obscurely granulate similarly to apical portion of pronotal disk ; sides more
evidently granulate ; hypomera sculptured similarly to middle area. Meso-
sternum with the surface sculptured similarly to middle area of prosternum but
more densely granulate. Metasternum feebly depressed posteriorly ; with a
broad (about 0-025 mm.), deep, median longitudinal impression which ends
abruptly at anterior fourth ; on each side of middle near base with a broad,
shallow, and subtriangular impression ; surface of disk with round, rather convex
granules which are distinctly coarser than facets oi eyes and are usually separated
by two times their diameters ; side similarly granulate hut with the granules
slightly less convex. Middle basal portion of first ventral segment moderately
depressed, at base rugose and obscurely granulate, and anterior portion with
the granules about hall .is coarse as those of metasternum and usually separated
by about five or more times their diameters: basal sides oi first and middle
352 H. E. HINTON
portion of second and third segments similarly granulate ; elsewhere on abdomen
with the granules separated by two to three times their diameters. Femora and
tibiae granulate similarly to abdominal segments ; tibiae with no rows of teeth.
Genitalia as figured (text-figs. 280, 281).
Female : Externally similar to male.
Type: g in the British Museum (Nat. Hist.). Mexico: Dist. de Tema-
scaltepec, Tejupilco, alt. about 4000 ft., vii.1934 (H. E. Hinton).
Paratypes : 18, with same data as type ; 2, collected at the same locality on
15. vi. 1934 (H. E. Hinton, R. L. Usinger) ; and 6, Mexico : Estado de Morelos,
Cuernavaca, vi.1934 (H. E. Hinton).
Variations : The scutellum in some specimens is distinctly more narrowly
obovate than that of the type. The depth of the pronotal depression near basal
sinuation of sublateral carina is slightly greater in some specimens, and the
median longitudinal impression of the pronotum appears in some specimens to
begin at basal fifth. These variations are, however, scarcely worthy of mention.
( omparative notes : This species can be compared only with C. heterocerus
(Sharp), which it resembles in general shape, but from which it maybe distinguished
by the following readily observable characters : (1) the antennae are slightly longer
and are entirely rufo-piceous, while in heterocerus the apical segments are dis-
tinctly darker than the two basal ; (2) the elytral intervals are subequally convex
at base, the third being only very slightly instead of very much more strongly
convex than the other discal intervals ; and (3) the basal discal strial punctur.s
are very coarse, being nearly as broad as intervals instead of seldom more than
one-third as broad as they are in heterocerus.
Cylloepus heterocerus (Sharp).
(Text-figs. 284-288.)
1882. Elmis heterocerus Sharp, Biol. Centr.-Amer. Col., 1 (2) : 135, t. 4. fig. 13.
Male: Length, 3-0 mm. ; breadth, 1-4 mm. Subparallel, moderately convex.
Clothed with fine, moderately short (about 0-05 mm.-o-o6 mm. long), moderately
recumbent, brownish-testaceous hairs which arise at intervals equal to half or
less of their lengths ; antennae (probably badly rubbed in specimens before me)
with a few fine and short hairs and on inner apex of eighth, ninth, and tenth
segments with a prominent brush or erect hairs ; labrum clothed as elsewhere
but with the lateral apical portions with denser and longer hairs ; beneath with
the hairs generally shorter. Cuticle moderately shining and for the most part
microscopically alutaceous ; piceous to feebly rufo-piceous ; basal two to five
segments of antennae, mouth-parts, and tarsi pale rufo-piceous. Tomentum
cinereous with golden reflections. Head between eyes with a very feeble and
broad impression which ends anteriorly on each side at base of clypeus and
extends posteriorly to slightly behind eye on each side and then transversely
across head. Antennae as figured (text-fig. 286), with the basal segments always
distinctly paler than the apical. Clypeal suture well-marked and nearly straight ;
clypeus when viewed from in front very broadly and feebly arcuately emarginate ;
angle on each side obtusely rounded, and sides feebly arcuate. Labrum broadly
and feebly rounded in front with the angle on each side broadly rounded. Surface
set with round granules which are slightly but distinctly finer than facets of eyes
and are usually separated by twice their diameters though often by less ; labrum
without granules, rather densely alutaceous, and punctate with fine punctures
MEXICAN ELMIDAjE
353
generally separated by once to twice their diameters. Pronotum at broadest
point near basal half broader than long (0-98 mm. : 0-90 mm.) and bast' broader
than apex (0-92 mm. : 0-72 mm.). Apical margin as seen from above moderately
arcuate at middle and deeply sinuate on each side behind eye before apical
angle ; apical angles moderately acute, rather broad, and moderately produced
forwards and slightly inwards ; sides feebly arcuate though slightly more strongly
so at basal half, feebly sinuate before basal angles, and scarcely sinuate at apical
fourth ; lateral margins finely crenate ; basal angles feebly acute, nearly rect-
angular, and scarcely produced ; base insinuate, broadlv and moderately deeply
287
Text-pigs. -'S4 -2SS. — Cylloepus heterocerus (Sharp). (2S4) Dorsal view of male geni-
talia. (285) Right lateral view of same. (286) Antenna, (287) Presternum. (288)
Adult to show genera] appearance.
sinuate on each side and shortly and more shallowly sinuate in front of scutellum.
Pronotum with the sublateral carinae prominent, most strongly raised on basal
half, very feebly curving towards apex, moderately strongly sinuate at basal
two-fifths, and feebly sinuate at apical one-third and becoming obsolete at about
apical eighth ; with impressions as follows : a very broad and feebly raised
portion extends from near base in front of scutellum to base of disk ; median
longitudinal navicular impression extending from basal third to apical third,
rather deep and well-defined, and broadest at about basal halt ot pronotum
where it is nearly as broad as scutellum ; at sides of disk near basal sinuation of
sublateral carina with a broad, deep, and slightly oblique impression which when
viewed laterally appears to feebly bisect sublateral carina and continue obliquely
to apical third ot sides near lateral margin where it again becomes as broad as on
disk ; at sides of disk near apical sinuation of sublateral carina there is a broad,
feeble, and transverse impression. Surface sculptured as follows: sides between
354 H- E- hixtox
sublateral carinae and lateral margins with granules which are round to obovate,
coarser than those of head, slightly coarser than facets of eyes, and occasionally
confluent but mostly separated by once to twice their diameters ; sides of disk near
apex with a few feeble granules and also with an occasional granule elsewhere ;
disk with the punctures fine, shallow, and separated by less than a third to once
their diameters so that the surface frequently appears to be eroded. Elytra more
than twice as long as pronotum (2-25 mm. : 0-90 mm.) and feebly broadened to
broadest point which is at apical third and here broader than base of pronotum
(1-40 mm. : 0-93 mm.). Apices feebly produced, conjointly broadly and feebly
rounded. Lateral margins very feebly crenate. Disk of elytra scarcely notice-
ably striate on basal portion and beyond apical two-thirds without a trace of
striae ; discal strial punctures about one-fifth to one-third as coarse as sutural
interval, rather shallow, usually round, and separated longitudinally by less than
to twice their diameters ; at sides between the two carinate intervals and
between outer carina and lateral margin with the punctures about half as coarse
as sutural interval, subquadrate, deep, and usually separated longitudinally by
less than their diameters ; these punctures become finer and nearly obsolete at
apex. Discal intervals flat, third strongly convex and slightly curved inwards
on basal sixth ; surface of discal intervals at base granulate similarly to sides of
pronotum but with the granules slightly finer ; granules on carinae coarser and
denser than discal basal ones. Scutellum flat, subovate, broader than sutural
interval (0-14 mm. : o-io mm.), longer than broad (0-17 mm. : 0-14 mm.), feebly
and broadly rounded basally, and more narrowly rounded at apex ; surface
impunctate, without granules, and strongly shining. Prosternum with the process
shaped as figured (text-fig. 287) ; middle area of prosternum feebly concave, set
with granules which are about as coarse as those at sides of pronotum and which
are seldom separated by as much as twice their diameters ; hypopleura similarly
punctate. Mesosternum with the groove for the reception of the prosternal
process very deep and broad ; surface granulate similarly to prosternum. Meta-
sternum very feebly depressed posteriorly; with a broad (about 0-037 mm.,
posteriorly), deep, median longitudinal line which is much narrowed anteriorly
but attains anterior margin ; on each side at base with a moderately large oval
and shallow impression ; surface of disk with the granules similar to those of
prosternum but separated by two to four times their diameters ; sides with the
granules denser, separated by once to twice their diameters. Middle portion of
first abdominal sternite feebly depressed ; all of middle of first sternite and
middle basal portion of second not granulate but finely alutaceous and with a
few sparse punctures ; sides of ventral segments with finer granules than meta-
sternum, which are usually separated by one to four times their diameters but are
even sparser on basal sides of first and second sternites and middle of third and
fourth. Femora and tibiae with the feebly obovate granules similar to those of
sides of metasternum. Genitalia as figured (text-figs. 2S4, 285).
Female : Externally similar to male.
Type: In the British Museum (Nat. Hist.). Guatemala: Vera Paz, San
Joaquim (Champion).
Specimens examined : 6, with same data as type. 3, Mexico : Dist. de
Temascaltepec, Temascaltepec, alt. 5000-6000 ft., vi.1934 (H. E. Hinton) ; and
one with same data as above but collected on 5.vi.ig33 (H. E. Hinton, R. L.
U singer).
Variations : No variations worthy of mention have been observed in the
small series before me.
MEXICAN ELMIDA] 355
( omparative notes : This species is nearest to C. blairi, and for the differences
between the two see the description of the latter. It might possibly be mistaken
for C. spinipes from which it may be distinguished as follows : (1) The antennae
are bicolorous instead of unicolorous ; (2) it is a shorter and proportionally broader
species ; (3) the pronotal disk is finely and irregularly punctate instead of densely
granulate ; (4) the elytral disk is nearly non-striate and moderately finely
punctate instead of noticeably striate and coarsely punctate ; and (5) the third
discal interval is only convex at base and the others are flat, while- in spinipes
all the discal intervals are moderately convex at base.
Cylloepus puncticollis (Hinton).
(Text-figs. 351, 297-302.)
1934 Slenelmis puncticollis Hinton, Rev. lint., RiodeJ., 4 : 198.
[937. ' ylloepus puncticollis Hinton, Arb. morph. taxon. lint. Berlin-Dahlem, 4 (2) : 106.
Male: Length, 2-8 mm-37 mm.; breadth, i-i mm.-i-j mm. Clothed
with fine, short (about 0-037 mm. long), recumbent, brownish-testaceous hairs
which arise mostly at intervals equal to distinctly less than their own lengths ;
antennae with the apical segments more densely clothed with slightly shorter
and more erect hairs ; beneath with the hairs generally shorter ; apical portion
of labrum clothed with equally fine but longer (about 0-062 mm.), paler and more
erect hairs which are rather dense at sides. Cuticle shining and for the most
part alutaceous ; black to dark rufo-piceous ; basal two or more segments of
antennae, mouth-parts, legs and surface beneath paler rufo-piceous. Tomentum
cinereous with feeble golden reflections. Head with a moderately shallow and
broad impression extending on each side near eyes to clypeal suture ; on vertex
with a moderately deep obovate impression which is nearly as broad as second
antennal segment. Antennae (text-fig. 299) bicolorous. Clypeal suture nearly
straight, strongly impressed ; anterior margin of clypeus when seen from in
front very broadly and feebly arcuately emarginate for its entire breadth, with
the angle on each side broadly rounded and the sides feebly arcuate Labrum
with the anterior margin broadly and feebly rounded, with the angle on each
side broadly rounded. Surface with the alutaceous microsculpture under a
magnification of x 150 appearing densely granulate ; set also with moderately
flat granules which are about as coarse as facets of eyes and are usually separated
by about once their diameters though at base and sides of head they are slightly
sparser ; clypeus similarly granulate ; labrum without granules, finely punctate
on a transverse middle band with punctures which are seldom separated 1>\ as
much as twice their diameters; extreme apex and base nearly impunctate. Pro-
notum at broadest point which is near basal half slightly broader than long
(1-12 mm. : 1-07 mm.) and base broader than apex (i-02 mm. : 0-85 mm.).
Apical margin moderately arcuate at middle and deeply sinuate on each side
behind eye before apical angle ; apical angles moderately produced forwards and
moderately acute ; sides feebly arcuate near basal half, nearly straight on api< al
half, very feebly sinuate at apical third, and feebly sinuate before basal angles ;
lateral margins moderately feebly crenate ; basal angles moderately acute,
scarcely produced ; base trisinuate, broadly and moderately strongly sinuate on
each side and more narrowly and shallowlv sinuate in front of scutellum. Pro-
notum with the sublateral carinae prominent, feebly converging towards apex,
moderately feebly sinuate at basal half, and becoming obsolete just before apical
35&
H. E. HINTON
289
291 293
292
295
Text-figs. 289-296. — (289) Dorsal view of male genitalia of Cylloepus optatus Sharp.
(290) Left lateral view of same. (2gi) Antenna. (292) Prosternum. (293) Dorsal
view of male genitalia of C. barben Hinton. (294) Right lateral view of same. (295)
Prosternum. (296) Antenna.
300
_ 30I
298
297 302
Iext-figs. 297-302. — Cylloepus puncticollis (Hinton). (297) Dorsal view of male
genitalia. (298) Left lateral view of same. (299) Antenna. (300) Labial palp
of male. (301) Labial palp of female. (302) Prosternum.
MEXICAN ELMIDAE 357
margin ; broadest portion of carina is from basal fourth to apical fourth. Pro-
notum with impressions as follows : a very broad and gradually widening raised
portion extends from base in front of scutellum to base of disk at about apical
two-thirds of pronotum ; on each side of this raised area a feeble impression
extends obliquely forwards from a puncture-like impression near base to join a
broad, moderately deep, oblique impression which is mostly on basal third but
which narrows and when viewed laterally extends moderately deeply across
sublateral carina at the basal impression where the carina is slightly narrowed ;
median longitudinal impression at base (here occupying most of basal raised
area) broad but becoming much narrowed at apical two-thirds, where it suddenly
broadens out to a navicular impression which becomes obsolete at about apical
sixth, this navicular impression being at its broadest point, which is at apical
two-fifths as broad as scutellum. Surface only feebly alutaceous it sides, not
alutaceous on disk ; disk with round to obovate punctures which van' from
being twice the diameter of a facet to being one-fourth this size, though most of
them are as coarse as facets of eyes ; these puncture-, are seldom separated by
more than their diameters, and near anterior margin the punctures become
denser and are often confluent so that here the surface appears to be somewhat
rugose ; sides of disk near sublateral carinae and bottom of depression mar basal
sinuation of sublateral carina somewhat rugose ; basal lateral portion of disk
finely and sparsely punctate ; surface of sublateral carinae so densely punctate
that it becomes rugose and in certain lights appears to be densely granulate ;
sides between sublateral carinae and lateral margins slightly more sparsely rugose
and granulate. Elytra more than twice as long as prothorax (2-40 mm. : 1-07 mm.)
and very feebly broadening posteriorly to broadest point which is near apical
third and is broader than base of prothorax (1-45 mm. : 1-02 mm.). Apices
moderately feebly and broadly produced and conjointly feebly rounded. Lateral
margins without distinct crenations. Surface moderately densely striate and discal
striae only slightly finer at apical tenth ; discal strial punctures round to sub-
quadrate, deep, and at middle of disk from a third to two-thirds as broad as
sutural interval and usually separated longitudinally by once to twice their
diameters ; discal punctures between basal convex intervals usually a little
finer and denser ; punctures on apical tenth about a fourth as coarse as those on
middle of disk. Discal intervals subequal in breadth and with the sutural
slightly broader than the others ; intervals nearly flat and at basal fifth with all
except sutural and fifth feebly convex, third distinctly more convex than second
or fourth and fourth slightly more convex than second and posteriorly not as
abruptly flattened as others but continued as a feebly convex interval to apical
third ; surface of intervals feebly alutaceous, on basal fifth with theconvex intervals
having the granules round, feebly convex, about as coarse as those on sides of
pronotum , and seldom separated by more than once their diameters ; towards apex
the granules of the intervals become much sparser; lateral carinae granulate
similarly to basal discal intervals ; sutural interval and intervals between strial
punctures much more sparsely granulate. Scutellum flat, subovate, broader than
sutural interval at base (0-12 mm. : o-IO mm.), longer than broad (0-17 mm. : 0-12
mm), broadly and feebly rounded basally, nearly truncate, and at apex moderately
acutely pointed ; surface without distinct granule-,. Prosternum with the process
as figured (text-tig. .502) ; prosternum with the .interior two-thirds (not including
process) strongly but not sharply lobed ; middle area coarsely but not distinctly
rugose and granulate, anterior marginal area and sides not rugose and more
finely granulate, and with round, feebly convex granules which are slightly
35§ H. E. H1NTON
coarser than facets of eyes and are usually separated by once to twice their
diameters ; hypomera with the granules finer, very feeble, and not distinct.
Mesosternum with the groove for the reception of the prosternal process feebly
triangular posteriorly and occupying most of the middle ; surface sculptured like
the middle of the prosternum. Metasternum very feebly depressed posteriorly
and on each side of middle at posterior margin with a large, moderately feeble
depression ; with a moderately narrow median longitudinal impression which is
distinct to anterior two-fifths and becomes obsolete shortly before anterior margin ;
disk with the usually round granules about half again as coarse as facets of eyes
and separated mostly by once to twice their diameters ; sides similarly granulate
but extreme sides with only an occasional very fine granule. Middle portion of
first ventral sternite of abdomen very strongly depressed ; lateral carina evident
only on basal half and even here very broad and but little raised ; anterior border
of depression rugose and remainder of surface with the granules from a half to a
third as coarse as those of metasternum and separated by three to four times their
diameters ; sides of first sternite for a short distance near depression granulate
nearly as coarsely as metasternum and sides elsewhere as well as those of second
and third segments with only an occasional granule ; middle portion of sides of four
apical stemites with the granules a third to two-thirds as coarse as those of meta-
sternum and separated by one to three times their diameters though slightly finer
and sparser on middle region of second, third, and fourth. Femora and tibiae with
the granules generally coarser than abdominal ones, about two times as coarse as
metastemal ones and similarly separated. First tarsal segment of all legs with
two fine and erect hairs at ventral apex. Genitalia as figured (text-fig. 297, 298).
Female : Externally similar to male but with the terminal segment of the
labial palpi about half as broad as that of male.
Type : $ in the British Museum (Nat. Hist.). Mexico : Tejupilco, Dist. de
Temascaltepec, alt. about 4000 ft., vi.1933 (H . E. Hinton, R. L. V 'singer).
Specimens examined : 1, with same data as above ; and 128, collected in the
same locality in vii.1934 (H. E. Hinton).
Variations : In addition to the usual slight variation in the size and density
of the punctures and granules on the various sclerites, this species exhibits a
moderate range of variation as follows :
(1) On the pronotum of some specimens there is no definite median longi-
tudinal impression from base to apical three-fourths. There is also
a slight amount of variation in the extent and depth of the median
navicular impression of the pronotum.
(2) The punctures of the pronotal disk are in some specimens mostly con-
tiguous to confluent so that the surface appears to be subrugose, while
in other specimens the punctures of this area are mostly separated by
half their diameters, and the surface never appears to be subrugose.
(3) A noticeable difference in the shape and breadth of the terminal
segment of the labial palpi of both males and females. In the
males the thickness of the apex may vary as 1 : 2.
(4) In a number of specimens the lateral margins of the prosternal process
are feebly raised, and occasionally the middle of the process is
feebly and longitudinally convex.
(5) The first ventral abdominal sternite varies from being moderately
depressed to being strongly depressed. This variation, except for
that in the absolute length of the beetle, is the most noticeable.
(6) A few specimens have the antennae practically uniformly coloured.
MEXICAN ELMIDAE ;v)
Comparative notes : The punctate instead of granulate pronotal disk will
separate this species from C. spinipes Hinton. From C. barberi Hinton it maybe dis-
tinguished as follows : (i) the surface of the pronotal carinae is granulate instead
of punctate ; (2) the basal discal elytral intervals are convex instead of flat ;
and (3) the depressed instead of flat middle portion of the first abdominal sternite.
The moderately convex instead of flat fourth basal interval of elytral disk will
serve to distinguish it from C. optatus Sharp.
Cylloepus spinipes Hinton.
(Text-figs. 303-309-)
1934. Cvlloepus spinipes Hinton, Rev. Ent., Rio de J., 4 (2) : 192.
Male: Length, 3-0 mm-3'4 mm.; breadth, i-i mm. -1-3 mm. Clothed
with fine, short (about 0-038 mm. long), recumbent, brownish-testaceous hairs
which arise mostly at intervals equal to slightly less than their lengths ; antennae
similarly but more sparsely clothed ; apical half of labrum clothed with equally
fine but much longer (as long as 0-087 mm- at sides) and paler hairs which are
more erect and much denser. Cuticle shining and for the most part densely
alutaceous ; piceous to dark rufo-piceous ; antennae, mouth-parts, and legs
paler rufo-piceous. Tomentum cinereous with moderate golden reflections.
Head with a scarcely noticeable, broad, feebly oblique impression which extends
shortly on each side near anterior half of eyes. Clypeal suture strongly impressed
and feebly arcuate ; anterior margin of clypeus very broadly and feebly arcuately
emarginate, with the angle on each side obtusely rounded and sides moderately
arcuate. Labrum broadly and feebly rounded in front, with the angle on each
side broadly rounded. Surface with the alutaceous microsculpture appearing
somewhat granulate ; with granules which are usually round, slightly coarser
than facets of eyes, and usually separated by less than to once their diameters ;
granules on clypeus similar ; labium without granules, apical half punctate with
very fine punctures which are usually separated by once to twice their diameters.
Pronotum at broadest point, which is at basal half, broader than long (1-05 mm. :
0-97 mm.) and base broader than apex (0-96 mm. : 0-72 mm.). Apical margin
as seen from above moderately feebly arcuate at middle and deeply sinuate on
each side behind eye before apical angle ; apical angles acute and strongly pro-
duced forwards and very slightly inwards ; sides feebly arcuate, more strongly
so at basal half, nearly straight at apical fourth and scarcely noticeably sinuate
just before basal angles; lateral margins feebly crenate ; basal angles feebly
acute, nearly rectangular and scarcely produced ; base trisinuate, broadly and
moderately strongly sinuate on each side and shortly and very shallowly sinuate
in front of scutellum. Pronotum with the sublateral carinae prominent, very
slightly converging towards apex, moderately sinuate at basal two-fifths, and
extending to apical margin, while the broadest portion of the carinae is from
basal fifth to apical fourth. Pronotum also as follows : a very broad and feebly
raised portion extends from base to base cil disk ; on each side ol this raised area
with a feeble, moderately broad, feebly curved and oblique impression extending
to broad, moderately shallow, indefinite impression near sinuation ol sublateral
carina which when viewed from a nearly lateral position seems to very shallowly
extend across sublateral carinae at about middle ; median longitudinal impres-
sion in some specimens extending from base to basal third as a very shallowly
360
H. E. HIXTON
impressed line and in others beginning only at basal third, but in all extending
from basal third to about apical third as a navicular, moderately deep impression
which is broadest at middle where it is not quite as broad as scutellum. Surface
with the alutaceous microsculpture similar but not as evident as that of head ;
set with round to feebly obovate granules which are nearly twice as coarse as
facets of eyes, low (only feebly convex), and are usually separated by slightly
less than their own diameters ; granules on sublateral carinae coarser and denser ;
303
Text-figs. 303-309. — Cylloepus spinipes Hinton. (303) Dorsal view of male genitalia.
(304) Left lateral view of same. (305) Antenna. (306) Inner view of hind tibia
of male. (307) Lateral view of hind tibia of male. (308) Inner lateral view of
middle tibia of male. (309) Inner lateral view of front tibia of male.
granules anteriorly on disk slightly finer and sparser. Elytra more than twice as
long as pronotum (2-30 mm. : 0-97 mm.) and feebly broadening to broadest
point, which is at apical third, and is here broader than base of pronotum (1-32
mm. : 0-96 mm.). Apices broadly and moderately feebly produced and con-
jointly broadly rounded. Lateral margins feebly but regularly crenate. Surface
striate with the discal striae moderately coarse at base, becoming finer towards
apex, and beyond apical fourth obsolete except for sutural ; discal strial punc-
tures round to subquadrate and at basal third a third to two-thirds as broad as
intervals, separated longitudinally by once to twice theirdiameters ; these punctures
are narrower basally and towards apex rapidly become fine and sparse so that at
apical eighth they are about a fourth as coarse as those at basal third. Discal in-
tervals nearly flat and subequal in breadth, at base all except sutural are slightly
MEXICAN I.I. Mil). \1 361
convex, and of the convex intervals the third is only slightly more so than the
others ; surface of intervals at base alutaceous somewhat as pronotum, elsewhere
much more sparsely and differently alutaceous ; on basal fourth set with granules
which are similar in size to those of pronotum and similarly distributed ; these
granules rapidly become sparser so that beyond basal half the disk is free of
granules ; granules of carinate intervals equal in size and density to those of
elytral base. Scutellum flat, subovate, broader than sutural interval (0-14 mm. :
o-io mm.), longer than broad (0-15 mm. : 0-14 mm.), very feebly and broadly
reunded basally, nearly truncate, and at apex narrowly rounded ; surface nearly
free of granules. Prosternum with the process rounded at apex ; presternum
and process evenly and moderately depressed, with the anterior two-thirds (not
including prosternal process) strongly and abruptly lobed ; surface granulate as
that of elytral base ; hypopleura slightly more sparsely granulate. Mesosternum
nearly entirely depressed for the reception of the prosternal process ; sides near
middle coxae very strongly and broadly raised and at apex with a dozen or less
slender, long (about 0-075 mm.), erect, brownish-testaceous hairs ; surface similar
to prosternum but more sparsely granulate. Metasternum with nearly the
entire discal region occupied by a deep, transverse, oval depression ; posterior
margin on edge of each side of middle with a moderately deep and large sub-
triangular impression ; with a broad (0-375 mm. broad in specimen 3-0 mm. long)
and shallow median longitudinal impression which ends rather abruptly at apical
fifth and basal seventh ; discal surface granulate similarly to pronotum ; sides
similarly sculptured but with the granules sparser and extreme sides with the
granules very flat. Middle basal portion of first abdominal sternite strongly
depressed and sculptured similarly to metasternal disk ; ventral sternites else-
where more sparsely granulate than disk of metasternum and especially sparsely
so at middle. Front tibiae with three tooth-like structures as figured (text-fig.
309). Middle tibiae with a ventral row of teeth (text-fig. 308). Hind tibiae
with a large, flat, knife-like tooth on inner ventral side at basal two-fifths (text-
figs. 306, 307). Surface of femora and tibiae granulate similarly to ventral
abdominal segments. Genitalia as figured (text-figs. 303, 304).
Female : Externally similar to male except as follows : (1) sides of meso-
sternum only moderately convex and without the group of long hairs ; (2) disk
of metasternum not as broadly nor as strongly depressed ; and (3) front and hind
tibiae without the tooth-like structures and middle tibiae without the row of
teeth.
Type: <$ in the U.S. National Museum. Mexico: Dist. de Temascaltepec ,
Real de Arriba, alt. 6000-7000 ft., v-vi-vii. 1933 (H. E. Hinton, R. L. Usinger).
Specimens examined : 68, with same data as type but a few of these also
collected at Temascaltepec, alt. about 5000 ft. ; 25, as above but collected in
vi.vii.1934 (H. E. Hinton).
Variations : The only notable variation is that in some specimens the median
longitudinal line extends from base to basal third as a very shallowly impressed
line, while in others it is absent on basal third and only present from basal third
to apical third.
Comparative notes : The males may be separated from all other species by
the secondary sexual characters of the front and hind tibiae. The females may
be distinguished from those of C. barberi by the granulate instead of punctate
pronotum, while from both C. puncticollis and C. sexualis they may be distin-
guished by (1) the strongly instead of feebly depressed metasternal disk and (2)
the densely granulate instead of punctate pronotal disk.
novit. zool., 42, 2. 1940. 28
362 H. E. HINTON
Cylloepus sexualis Hinton.
(Text-figs. 310-316.)
1937- Cylloepus sexualis Hinton, Arb. morph. taxon. Ent. Berlin-Dahlem, 4 (2) : 102, figs.
4-10.
Male: Length, 3-1 mm-37 mm. ; breadth, i-2 mm-1-3 mm. Subparallel
moderately convex. Clothed with fine, short (about 0-037 mm- l°ng). recumbent,
brownish-testaceous hairs which arise mostly at intervals equal to distinctly less
than their own lengths ; antennae with the apical segments more densely
clothed with slightly shorter hairs ; labrum clothed with equally fine but generally
much longer, more erect hairs which are most numerous at sides. Cuticle shining
and for the most part alutaceous ; colour black to dark rufo-piceous ; basal two
segments of antennae, mouth-parts, and legs paler rufo-piceous. Tomentum
cinereous with golden reflections. Head without distinct impressions, but ante-
riorly between eyes with a median longitudinal impression which is nearly as
long as basal segment of antennae and about two-fifths as broad as long. Antennae
as figured (text-fig. 312). Clypeus with the suture straight and strongly impressed ;
anterior margin of clypeus as usual when seen from in front very broadly and
arcuately emarginate, with the angle on each side obtusely and bluntly rounded ;
labrum very broadly and feebly rounded in front, with the angle on each side
broadly rounded. Surface densely and very minutely alutaceous in such a manner
that it appears minutely granulate ; also set with low granules which are about
as coarse as facets of eyes and are usually separated by once to twice their
diameters though sparser basally where the surface is also somewhat rugose ;
granules of clypeus slightly denser and more regularly distributed ; labrum
without granules, base and apex nearly impunctate, elsewhere with the punctures
fine (about two-thirds as coarse as facets of eyes), round, and usually separated
by once their diameters or less. Pronotum with the greatest breadth near basal
half not as great as length (0-97 mm. : 1-05 mm.) and base broader than apex
(0-87 mm. : 075 mm.). Apical margin as seen from above moderately arcuate
and deeply sinuate on each side behind eye before apical angle ; apical angles
moderately produced forwards and moderately acute ; sides moderately arcuate
on basal half, nearly straight on apical half, and moderately sinuate before basal
angles. Lateral margins finely crenate ; basal angles moderately acute and
feebly produced backwards ; base trisinuate, broadly and moderately deeply
sinuate on each side, narrowly and moderately deeply sinuate in front of scu-
tellum. Pronotum with the sublateral carina prominent, moderately feebly con-
verging towards apex, moderately sinuate at basal half, when viewed from above
slightly sinuate on apical third, and extending to apical margin ; broadest portion
of carina difficult to delimit but apparently extending from basal fourth to apical
third. Pronotum with the impressions as shown in text-fig. 318 for C. proximus.
Surface feebly alutaceous, more strongly so at bottom of impressions and at sides
and anterior margin but nowhere as strongly alutaceous as head ; disk punctate
with round to feebly obovate punctures which are moderately deep, usually a
half to two or more times as coarse as facets of eyes, often contiguous or confluent
and seldom separated by as much as half of their diameters ; anteriorly on the
disk with most of the punctures confluent and therefore appearing rugose ;
bottom of the various impressions nearly impunctate and therefore appearing
highly polished ; basal convex areas on each side of middle with a few punctures
similar to those of disk but slightly sparser ; surface of sublateral carina punctate
as on disk but so densely as to appear rugose ; sides near lateral margins much
MEXICAN hl.MIDAE
363
more sparsely punctate. Elytra more than twice as long as pronotum (2-42 mm. :
1-05 mm.) and feebly broadening posteriorly to broadest point which is at apical
third and here broader than base of pronotum (1-30 mm. : 0-87 mm.). Lateral
margins without distinct crenations. Surface rather coarsely striate, discal
striae slightly finer at apical tenth but not obsolete ; discal strial punctures
subquadrate to nearly round, rather deep, and at middle of disk from two-thirds
to as broad as intervals and separated longitudinally by slightly more than to
slightly less than their lengths ; towards sides the punctures become coarser and
Text-figs. 310-317. — (310) Dorsal view of male genitalia of Cylloepus sexualis Hinton.
(31 1) Left lateral view of same. (312) Antenna. (313) Lateral inner view of front
tibia of male. (314) Dorsal view of median lobe of male genitalia. (315) Labial
palp of male. (316) Prosternum. (317) Dorsal view of median lobe of male geni-
talia of C. proximus Hinton.
towards apex much finer. Discal intervals (five intervals) all nearly flat, second
feebly convex on basal fifth, third strongly convex but becoming flat at about
apical three-fourths, fourth flat (with punctures of the two striae almost con-
tiguous so that the fifth seems to be the fourth), and fifth moderately convex on
basal seventh ; surface of intervals at most only feebly alutaceous ; elevated
intervals with numerous round and low granules which are slightly coarser than
facets of eyes, and the surface here also somewhat rugose ; carinate intervals
similarly sculptured ; surface between punctures with only an occasional granule
and often feebly rugose. Scutellum flat, ovate, broader than sutural interval at
base (0-14 mm. : o-io mm.), longer than broad (0-17 mm. : 0-14 mm.), broadly
and feebly rounded basally, and at apex moderately acutely rounded ; surface
with a few coarse and shallow punctures. Prosternum with the anterior two-
thirds (not including process) strongly but not sharply lobed ; prosternal process
364 H. E. HINTON
as figured (text-fig. 316) ; surface of middle area densely, moderately coarsely
rugose and obscurely granulate, sides with the granules varying much in size but
usually about as coarse as facets of eyes or slightly finer and separated by one
to three or more times their diameters. Hypomera feebly, moderately sparsely
rugose and with a few fine, obscure granules. Mesosternum sculptured similarly
to middle area of presternum. Metasternum with all the middle of the disk
except anterior two-fifths moderately strongly depressed but with the area near
median line slightly less depressed so that it appears that there are two depressions
on disk, one on each side of median line ; area just before mesosternum strongly
declivous ; on each side of middle at posterior margin with a moderately large,
oval, deep depression ; with a broad (0-037 mm- basally), deep, median longi-
tudinal impression which extends broadly to apical third ; surface of disk granu-
late with round to obovate, flat-topped granules which are as coarse as facets of
eyes and are separated by once to twice their diameters ; sides similarly granulate
but extreme sides and pleura with only a few fine and obscure granules. First
abdominal sternite with the entire middle portion strongly depressed and pos-
teriorly at middle this depression seems to encroach upon middle of second segment
as far as apical two-fifths ; surface of the depression feebly rugose ; carinae of
first sternite prominent and extending from base to apex ; surface of sides of
first sternite and extreme sides of others with only an occasional granule ; surface
of segments elsewhere with rather flat, round granules which are half as coarse
as facets of eyes and are usually separated by two to three times their diameters.
Hind coxa with a moderately large and deep, usually oval depression. Femora
and tibiae very finely but otherwise similarly granulate to metastemal disk ; the
front tibia on inner apex has a toothed carina (text-fig. 313) ; inner ventral side
of middle tibiae with a row of teeth similar to that of front tibiae ; hind tibiae
with no rows of prominent teeth ; when viewed from inner dorsal side feebly
curved and moderately swollen at apical two-fifths for a short distance ; when
viewed ventrally this swollen portion is somewhat concave and the tomentum
is here particularly long and dense. Genitalia as figured (text-figs. 310, 311).
Female : Differs externally from male as follows : (1) the basal abdominal
depression does not encroach on to second sternite so that there is no trace of a
depression here ; (2) the front tibia has no carina or teeth as has been figured for
the male ; (3) the middle tibia has no row of teeth ; and (4) the hind tibia though
nearly as curved at apical two-fifths as in male is not as thick.
Type : 3 m tne British Museum (Nat. Hist.). Mexico : Dist. de Tema-
scaltepec, Tejupilco, alt. about 4000 ft., vii.1934 (H. E. Hinton).
Specimens examined : 8o, with same data as above ; 16, with same data but
collected 15-28 . vi . 1933 (H. E. Hinton, R. L. Usinger) ; 1, Dist. de Temascal-
tepec, Temascaltepec, alt. about 5000 ft. (H. E. Hinton, R. L. Usinger).
Variations : Besides slight variations in body size and density of granules
the following have been noted : (1) a noticeable variation in the depth of the
depression which parallels basal raised portion of pronotum at the point where it
joins the large impression near basal sinuation of sublateral carina, being some-
times absent at this point so that basal convex portion appears as part of the
disk ; and (2) a slightly noticeable difference in the proportions of length to
breadth of the prothorax.
Teratology : One female specimen is malformed as follows : the outer sub-
lateral carina (on eighth interval) is broadly interrupted on the left side at apical
two-fifths and less broadly interrupted on the right side at the same place, and
the inner sublateral carina is broadly interrupted at basal third on left side only.
MEXICAN ELMIDAE
365
Comparative notes : The punctate instead of granulate pronotal disk will
serve to distinguish it from C. spinipes. The bicoloured instead of uniformly
coloured antennae, the coarsely and densely instead of finely and moderately
sparsely punctate pronotal disk, the strongly convex instead of nearly flat third
basal interval of elytra, and the strongly depressed instead of nearly flat middle
of basal abdominal sternite will separate it from C. barberi. From C. optatus, to
which it is most closely related, it may be distinguished as follows :
C. sexualis.
Antennae with the two basal seg-
ments pale rufo-piceous and the
others piceous to black.
Elytra with the fifth interval at
base moderately strongly convex.
Male with the second abdominal
sternite depressed only at middle
of basal half.
Male with a short, prominent carina-
like swelling at inner apical fourth
of front tibia.
Front tarsi of males without nume-
rous erect hairs on ventral surface
C. optatus.
1. Antennae uniformly pale rufo-pi-
ceous ; sometimes with the two
basal segments slightly paler.
2. Elytra with the fifth interval at
base at most feebly convex.
3. Male with the second abdominal
sternite depressed at middle of
basal two-thirds.
4. Male without a carina on front
tibia.
5. Males with the four basal segments
of the front tarsi densely clothed
on ventral side with moderately
long, erect, pale-testaceous hairs.
C. sexualis superficially resembles C. puncticollis but may at once be separated
by the flat fourth and convex fifth, instead of flat fifth and convex fourth basal
discal elytral intervals. The terminal segment of the labial palp is normal in the
males of C. sexualis, while in those of C. puncticollis it is very much broadened at
apex.
Cylloepus proximus Hinton.
(Text-figs. 317, 318.)
1937. Cylloepus proximus Hinton, Arb. morph. taxon. Ent. Berlin- Dahlem. 4 (2) : 106, figs.
11, 12.
Male: Length, 3-3 mm. ; breadth, 1-38 mm. Similar to male of sexualis
except as follows : Elytra with the fifth interval very shortly and scarcely notice-
ably elevated, while that of sexualis is distinctly elevated for a short distance.
Abdomen with the depression of the first sternite not, or just barely, encroaching
on second sternite. Inner apex of front tibia without a toothed ridge, as is the
case in sexualis ; hind tibia with the apical half not as strongly curved or swollen
as that of sexualis. Male genitalia of both species very similar, the chief differ-
ences being found in the median lobes which have been disxrtedout and illustrated
(sexualis, text-fig. 314 ; proximus, text-fig. 317).
Type: o m the Deutsches Entomologisches Institut, Berlin-Dahlem.
Mexico : Puebla, Necaxa (Georg Heine).
Larvae.
The larvae of Cylloepus have been determined by elimination and locality.
Only one Mexican species is available, and it appears to be either C. puncticollis
(Hinton) orC. sexualis Hinton. Tin- larva of a species (C. consobrinus Grouvelle -)
366
H. E. HINTON
from the Yungas Valley of Bolivia is also available. The following generic
characterization is drawn from a study of these two species.
Generic Characters of Larvae of Cylloepus.
Cylindrical, parallel. Head when viewed from above exposed and not con-
cealed by the pronotum. With one ocellus on each side. Antennae 3-segmented,
feebly retractile. Clypeus with the suture distinct. Mandibles of both sides
similar and with three acute apical teeth ; prostheca long, slender, and densely
spinose. Maxilla with the palp 4-segmented and the stipes showing no differen-
tiation into a palpifer ; galea and lacinia separate and apex of each densely
spinose. Labium with the postmentum undivided ; palp 2-segmented and pre-
mentum without a palpiger. Gula well developed. Prothoracic pleurae not
Text-fig. 318. — Cylloepus proximus Hinton.
divided, meeting on middle line of body. Meso- and metathoracic pleurae with
only the sterno-pleural suture present. First abdominal segment with only the
sterno-pleural suture present ; second to eighth with discrete pleurae bounded by
tergo- and sterno-pleural sutures and these sutures are continued to basal half
of ninth segment ; these sutures do not converge where they terminate, as in
most Elmidae, but remain parallel. Anterior and posterior part of presternum
divided by a complete median longitudinal suture. Ninth abdominal segment
with the apex broadly and strongly emarginate ; operculum with two strongly
sclerotized claws attached to dorsal membrane. Spiracles annular and biforous,
present on antero-lateral part of mesothorax and first eight abdominal segments,
and opening at level of cuticle and not on well-developed tubercles ; tracheae
without air sacs ; with three tufts of retractile, anal, tracheal gills. Alimentary
canal with a small dorsal oesophageal sclerite on posterior margin of oesophagus.
Hind gut with six Malpighian tubules. Central nervous system with three thoracic
and eight abdominal discrete ganglia. Stomodeal nervous system with two
occipital ganglia and a single oesophageal ganglion.
In general appearance the larvae of this genus are very similar to those of
Stenelmis Dufour, but they may at once be distinguished from Stenelmis and all
MEXICAN ELMIDAE
367
other known genera by the complete absence of tergo-pleural thoracic sutures
and the presence of both tergo- and sterno-pleural sutures on the ninth segment.
Description of Mature Larva of Cylloepus ,s'/>.
(Text-figs. 319-329.)
Length, 8-0 mm. ; breadth (across broadest point which is near base of ftieta-
thorax), 0-70 mm. Cuticle brownish-testaceous to brown. Head rectangular,
slightly broader than long (0-52 mm. : 0-50 mm.). Epicranial suture nearly
322
l'i kt-fios 319-323. — -Larva of Cylloepus sp. (319) Ventral view of right maxilla,
(320) Ventral view of labium. Setae of prementum are only approximately correct.
(321) Antenna, (i^i) Mandible. (323) Mesothoracic spiracle.
straight and 0-07 mm. long ; frontal suture extending nearly in a straight curve
on each side to margin of head opposite base of antenna. Cuticle on a basal belt
which is nearly as long as epicranial suture finely and transversely alutaceous ;
elsewhere with round punctures which are about 0-02 mm. broad and are separated
by less than to nearly three times their diameters ; from these punctures arise
fine setae which are usually only slightly longer than the breadth of their respective
punctures though occasionally they are about five times this Length. Antenna
as figured (text-fig. 321) ; retractile to basal third of first segment. Frontal
margin of head with a tooth-like projection on each side of clypeus between
clypeus and base of antenna which is two-thirds as long as basal segment of
antenna. Clypeus with the fronto-clypeal suture distinct ; slightly longer and
broader than labrum ; and with the anterior margin truncate, with the angle on
368
H. E. HINTON
each side rounded. Labrum (text-fig. 329) with the epipharynx smooth. Man-
dibles, maxillae and labium as figured (text-figs. 322, 319, and 320). Thorax
and first abdominal segment with the proportions as figured (text-fig. 324).
Surface of thorax and abdomen for the most part punctate as frontal region of
head but with the punctures about a third larger ; posterior margin of each
segment except ninth with a dense and complete ring of tubercles (text-fig. 328)
near anterior margin of each segment with a dense and complete ring of tubercles
327
324
328
329
Text-figs. 324-329. — Larva of Cylloepus sp. (324) Ventral view of thorax and first
abdominal segment to show sclerotization. (325) Operculum. (326) Dorsal view
of ninth abdominal segment. (327) Posterior view of left front leg. (328) A section
of third abdominal tergite. (329) Dorsal view of labrum.
from which long flat setae arise (text-fig. 328). Legs all fairly close in size and
chaetotaxy to front leg (text-fig. 327).
Specimens examined : 2, apparently mature larvae taken in a small (5-30 ft.
across) and rapid flowing stream. Mexico : Dist. de Temascaltepec, Tejupilco,
alt. about 4000 ft., vii.1934 (H. E. Hinton).
HETERELMIS Sharp.
1882. Heterelmis Sharp, Biol. Centr.-Amer. Col., 1 (2) : 130.
At the time of writing 10 species and 1 subspecies of this genus have been
described. To these two new species and one new subspecies are added in this
paper. The species occur from southern United States to south Brazil, and are
MKXICAN ELMIDAE
369
also known from Trinidad and Tobago in the British West Indies. A redescrip-
tion of the genus follows :
Body subovate to subparallel. Non-tomentose areas glabrous or clothed
with sparse and short recumbent to suberect hairs. Scale-like or hairy tomentum
contined to the following areas : (1) genae ; (2) epipleura ; (3) hypomera from
posterior fourth to anterior half with a ventral belt which at broadest is two-
fifths of hypomera, and this belt extends to anterior margin as a narrow fringe
along sterno-notal suture ; (4) sides of presternum, mesosternum, and abdominal
sternites ; and (5) part or all of legs except tarsi. Head when seen from below
330
331
Text-figs. 330, 331. — Heterelmis longior (Grouvelle). (330) Alimentary canal.
Dorsal view of central nervous system.
(331)
capable of being retracted so that none of the mouth-parts are visible. Antenna
(text-fig. 337) n-segmented. Mandibles (text-fig. 335) with three subacute
apical teeth ; prostheca long and entirely membranous with numerous long spines
and hairs apically. Maxilla (text-fig. 334) with the palp 4-segmented and the
stipes with a well-developed palpifer ; galea and lacinia separate and apex of
each densely spinose or hairy. Labium with the palp (text-fig. 365) 3-segmented
and prementum without a distinct palpiger. Mentum as broad and three-fourths
as long as submentum. Gula about a fifth longer than submentum, at anterior
margin about three-fourths as broad as submentum, and with the sides feebly
converging so that at posterior margin it is only half as broad as submentum.
Pronotum with the anterior margin moderately arcuate at middle and on each
side behind eye before apical angle broadly and moderately deeply sinuate. Base
trisinuate, broadly amd moderately deeply so on each side and more narrowly
370
H. E. HINTON
and shallowly so in front of scutellum. Pronotum on each side with a sublateral
carina which extends from base to, or very nearly to, anterior margin ; without
or with a transverse impression at middle ; with or without a median longi-
tudinal discal impression ; and with or without an oblique impression on each
side on basal half. Elytra punctate and striate ; each elytron with a longi-
tudinal carina on sixth interval and one on eighth. Hind wing (text-fig. 341)
without an anal lobe ; without a radial cross vein or an anal cell ; first anal
absent ; second anal with the first and second branches present ; third anal
without a second branch ; fourth anal well-developed ; and cubito-anal cross
Text-figs. 332, 333. — Heterelmis longior (Grouvelle). (332) Male reproductive system.
(333) Female reproductive system.
vein complete and joining cubitus to second anal. Prostemum very long in front
of anterior coxae ; prosternal process long, very broad, and with the posterior
margin broadly rounded to nearly truncate. Mesosternum with a moderately
deep and very broad groove for the reception of the prosternal process. Meta-
sternum with a median longitudinal impressed line. Legs with the visible portion
of the front coxae rounded and trochantin completely concealed by the hypomera
and sternum. Claws without teeth. Alimentary canal (text-fig. 330) with six
caeca on the anterior margin of the mid-gut. Hind gut with six Malpighian
tubules which end freely near the rectum. Male reproductive system (text-fig.
332) with the lateral accessory glands lobed. Each testis with two sperm tubes.
Female reproductive system (text-fig. 333) with 11 egg tubes to each ovary.
Spermathecal duct opening into apex of bursa copulatrix. Central nervous
MEXICAN ELMIDAF.
371
system (text-fig. 331) with three thoracic discrete ganglia ; abdominal ganglia
one to six discrete and seven and eight only partly (used together.
Genotype : Heterelmis obscura Sharp (1882).
The internal anatomy of three species has been examined and found to
agree in essential details, and that of H. longior (Grouvelle) (Elmis) is figured.
This genus is close to no other so far described, and the fact that it is isolated
is also borne out by the structure of the larvae (vide infra). Occasionally the
smaller species superficially resemble members of the genus Hexacylloepus
339
Text-figs. 334-339. — Heterelmis longula Sharp. (33t) Maxilla. (335) Dorsal view
of left mandible. (336) Ventral view of same. (337) Antenna. (338) Outline of
labrum. (339) Outline of apical abdominal tergite.
Hinton, but may be distinguished from this genus as follows : (1) the hypo-
meron always has adjacent to sterno-notal suture a narrow belt of tomentum
which extends to anterior margin and there is never a complete transverse belt,
whereas in Hexacylloepus there is no anterior belt of tomentum adjacent to the
sterno-notal suture, and there is at about the middle a complete transverse belt ;
(2) the hind wing has the second branch of the third anal absent whereas in Hexa-
cylloepus it is present ; (3) the hind gut has six instead of only four Malpighian
tubules; (4) the lateral accessory glands of the male reproductive system are
lobed, whereas in Hexacylloepus they are not lobed ; (5) each ovary has 11 egg
tubes instead of only seven ; and (6) the central nervous system has the first
six abdominal ganglia discrete, whereas in Hexacylloepus the first is partly fused
to the third thoracic and only two to five are discrete.
372
H. E. HINTON
The most important characters for distinguishing the various species seem
to be the following :
(i) General proportions, length and breadth.
(2) Colour.
340
^.777777777
341
Text-figs. 340, 341. — Heierelmis longida Sharp. (340) Adult to show general appearance.
(341) Hind wing. Venation after Forbes.
(3) Size and distribution of the punctures and tubercles on the various
sclerites, and density and type of microsculpture, if present, between
tubercles and punctures.
(4) Condition of fronto-clypeal suture.
(5) Condition of the anterior margin of clypeus and labium, and also the
condition of the angle on each side.
(6) Outline of pronotum and its general proportions.
(7) Extent and depth of pronotal impressions.
MEXICAN ELMIDAE 373
(8) Shape of elytral apices.
(g) Shape of scutellum.
(10) Shape and extent of prosternal carinae.
(n) Breadth of prosternal process and shape of posterior margin.
(12) Extent and depth of the various impressions on disk of metasternum.
(13) Length and shape of carinae of first abdominal sternite.
(14) Secondary sexual characters.
(15) Structure of the male genitalia.
Only one of the species before me has secondary sexual characters, and since
only a male has been found, it is entirely by analogy with other genera that the
following are considered to be secondary sexual characters :
(1) Male with the disk of the metasternum strongly concave (tarsalis).
(2) Male with a large oval area of dense, recumbent, testaceous hairs on
inner side of posterior gibbosity of metasternal disk (tarsalis).
(3) Male with the apex of front tibiae moderately gibbous {tarsalis).
(4) Male with numerous pale, erect hairs on ventral surface of four basal
segments of front and middle tarsi (tarsalis).
A Key to the Mexican Species of Heterelmis.
1. Metasternal disk strongly, longitudinally concave for its entire length ;
tibiae of front legs clavate ; basal four segments of front and middle
tarsi of male with a conspicuous fringe of erect hairs ; male genitalia
with the median basal projection of median lobe triangularly
broadened. Mexico ...... H. tarsalis, sp. n.
Metasternal disk only feebly concave posteriorly ; tibiae of front legs not
clavate at apex ; basal four segments of front and middle tarsi of male
without a conspicuous ventral fringe of erect hairs ; male with the
median basal projection of median lobe aciculate, not triangularly
broadened ........... 2.
2. Basal segment of all tarsi with two close, stout, short spines on inner
apex ; parameres of male genitalia without an inner apical fringe of
hairs, though (when stained carefully) with an occasional very minute
hair 3.
Basal segment of tarsi without two stout spines on inner apex ; parameres
of male genitalia with an inner apical fringe of moderately long hairs 5.
3. Length of prothorax nearly always more than 0-925 mm. ; median lobe of
male genitalia with the basal median projection seen from the side
gradually sloping behind. Mexico . . H. longula Sharp (1887).
Length of prothorax nearly always less than 0-925 mm. ; median lobe
of male genitalia when seen from the side strongly declivous behind 4.
4. Pronotum with the disk not evenly convex, i.e. with impressions.
Mexico, Guatemala ..... H. obesa Sharp (1882).
Pronotum with the disk evenly convex. Mexico H. obesa plana, subsp. n.
5. Median lobe of male genitalia broad, opposite apices of parameres it is
0-050 mm. broad. Texas, Mexico, Guatemala, Costa Rica,
Panama, Brazil ...... H. obscura Sharp (1882).
Median lobe of male genitalia narrow ; opposite apices of parameres it is
0-170 mm. broad (it has not been possible to separate females of
obscura from those of acicula). Mexico . . H. acicula, sp. n.
374 H- E- HINTON
Heterelmis tarsalis, sp. n.
(Text-figs. 342-346.)
Male : Length, 2-2 mm. ; breadth, 1-02 mm. Obovate. Surface pubescent
as usual in the genus ; hairs on dorsal surface usually brownish-piceous. Cuticle
moderately shining and black ; antennae, mouth-parts and legs rufo-piceous.
Head without distinct impressions ; surface minutely, densely eroded and also
with the surface of the callosities punctate with fine (about 0-007 mm. broad)
punctures which are about half as coarse as facets of eyes and are separated by
less than to two or three times their diameters. Clypeus with the fronto-clypeal
suture moderately deep and nearly straight ; anterior margin feebly, arcuately
emarginate for its entire breadth, and with the angle on each side broadly rounded ;
surface more coarsely and sparsely eroded (or rugulose) than head ; the punctures
are larger and seldom appear to be on callosities. Labrum with the anterior
margin broadly, feebly arcuate and with the angle on each side broadly rounded ;
surface not eroded, minutely alutaceous, and with punctures which are finer
than those of head and slightly sparser. Pronotum at broadest point, which is
at basal third, broader than long (0-875 mm- : 0'66o mm.) and base broader than
apex (0-825 mm. : 0-500 mm.). Sides moderately strongly arcuate at basal third,
elsewhere more feebly arcuate but at middle opposite median transverse impres-
sion they are feebly sinuate. Median transverse impression broad, moderately
strongly impressed, and distinct to sublateral carinae ; at middle this impression
is crossed by a deep, oval impression which is about 0-075 mm. broad and 0-125
mm. long. Oblique impression begins on middle fourth of basal third of pro-
notum, is broader and not as deep as transverse impression, nearly meets trans-
verse impression at sublateral carinae, and extends obliquely forwards to meet
lateral margin at basal two-fifths. Base on each side of scutellum with a mode-
rately deep, oval impression which is only a little smaller than median discal
impression. Surface of pronotum between sublateral carinae with two sizes of
punctures as follows ; coarse punctures, which are about as large as facets of
eyes or occasionally slightly larger, are contiguous to separated by as much as
five times their diameters ; fine punctures usually about a fourth to a third as
coarse and separated by five or more times their diameters (as on apical discal
region), though often (as on apical sides) coalescing to form a densely and minutely
eroded area ; there are also numerous intergradations between these two sizes
of punctures. Sides of pronotum between sublateral carinae and lateral margins
minutely, densely eroded, and also with round granules which are usually about
a third coarser than facets of eyes and are mostly separated by once to twice
their diameters. Elytra twice as long as pronotum (1-3 mm. : o-66o mm.) and
broadest point is at about apical two-fifths. Striae very feebly impressed,
slightly more strongly impressed at sides and apex ; strial punctures deep and
usually round, on discal region from a fourth to more than a third as coarse as
discal intervals and separated longitudinally by once to twice their diameters.
Second and third intervals slightly broader than sutural ; surface of intervals on
a narrow basal belt and laterally beyond inner carinae with occasional granules
similar to those of sides of pronotum ; surface elsewhere with the punctures from
which the hairs arise as fine as finest of pronotum and separated by two to ten
times their diameters. Scutellum subovate and with only fine punctures similar to
those of elytral intervals. Prosternum which is unusually broad for this genus
has the ratio of breadth (at broadest point which is at basal fourth) to length
MEXICAN ELMIDAE
375
0350 mm. : 0-200 mm. ; the apical margin is broadly truncate with the angle on
each side broadly rounded ; the sides are reflexed upwards at an angle of about
30° with the result that the prosternal process appears strongly concave ; on
each side from base of process the fine, straight, i.e. not oblique, carina extends
to apical third of presternum. Metasternum with the disk strongly concave and
sloping downwards anteriorly, and at the bottom of this cavity with a fine,
median longitudinal line which extends from posterior margin to anterior fifth ;
on each side near posterior margin the disk is strongly and broadly gibbous, and
Text-figs. 342-346. — Heterelmis tarsatis Hinton. (342) Dorsal view of male genitalia.
(343) Right lateral view of same. (344) Dorsal view of paramere. (345) Lateral
view of median lobe. (346) Dorsal view of base of median lobe.
on the mesal side of each gibbosity is a large, oval patch of testaceous hairs which
are as coarse and half again to twice as long as discal hairs ; surface of disk at
middle with punctures which are but slightly coarser than those of elytral intervals
and are separated by two to five times their diameters ; at sides of disk and on
sides of metasternum the surface is often finely and densely eroded and also set
with punctures which are slightly coarser than coarse ones of disk of pronotum
but are similarly distributed ; extreme sides also with fine granules which are
slightly finer than coarse punctures. Abdominal stemites on middle between
tomentose portions with punctures a little coarser (base of basal segment with
punctures as much as a third coarser) than discal pronotal ones and separated
by less than to three times their diameters. Legs with the front tibiae gradually
becoming stouter towards apex so that at broadest point, near apical fifth, they
are o-ioo mm. in diameter. Ventral surface of basal four segments of front
376 H. E. HINTON
tarsi with about 25 erect hairs which are slightly longer than apical diameter (or
occasionally shorter) of the second segment. Ventral surface of basal four
segments of middle tarsi have about 60 erect hairs which are slightly longer than
apical diameter of second segment. Ventral surface of four basal segments of
hind tarsi with an occasional erect hair. Genitalia as figured (text-figs. 342-346) .
Female : Unknown.
Type: <J in the British Museum (Nat. Hist.). Mexico: Dist. de Temascal-
tepec, Tejupilco, alt. 3500 ft., vii.10.34 (H. E. Hinton).
Comparative notes : This is one of the most aberrant species of the genus.
It may be distinguished from all other described species of Heterelmis by the
following : (1) the straight instead of oblique prosternal carinae ; (2) the un-
usually broad and short prosternal process ; (3) the longitudinally concave disk
of the metasternum which slopes downwards towards anterior margin ; (4) the
broadly and strongly gibbous region on each side of posterior part of metasternal
disk ; (5) the dense patch of testaceous hairs on mesal side of each discal gib-
bosity ; (6) the clavate front tibiae ; (7) the fringe of erect hairs on the ventral
surface of the four basal segments of all tarsi ; and (8) the structure of the male
genitalia. It seems probable that of these characters only the first two will be
possessed by the female, the others probably being secondary sexual characters.
Heterelmis longula Sharp.
(Text-figs. 334-341.)
1887. Heterelmis longulus Sharp, Biol. Cenlr.-Amer. Co!., 1 (2) : 775.
Male : Length, 3-5 mm-40 mm. ; breadth, i-2 mm.-r6 mm. Elongate,
subparallel. Dorsal surface clothed with fine, recumbent to suberect, testaceous
setae which are usually about 0^075 mm. long and arise from minute punctures
which are separated by distances equal to or slightly less than the lengths of the
hairs ; on elytra these hairs are mainly confined to the surface of the intervals
so that in specimens which have not been rubbed they appear to be arranged in
longitudinal rows. Ventral surface — apart from tomentose areas — clothed with
finer, sparser, and generally slightly longer hairs. Cuticle moderately to strongly
shining and black to dark rufo-piceous ; antennae, mouth-parts, legs and often
middle portion of ventral surface paler rufo-piceous. Head without distinct
impressions ; surface minutely and densely eroded and punctate as follows :
throughout with numerous microscopic (about 0^005 mm. broad) punctures ; and
also with larger (o 020 mm. broad) punctures which are on the small, irregular,
flat callosities and are separated by much less than to twice their diameters.
Clypeus with the fronto-clypeal suture deeply impressed and nearly straight ;
anterior margin scarcely noticeably, arcuately emarginate for its entire breadth,
and with the angle on each side broadly rounded ; surface at sides sculptured
similarly to head but on middle region not distinctly eroded and with the punctures
slightly coarser and never on callosities. Labrum as figured (text-fig. 338) ;
surface punctate more finely but otherwise similarly to middle region of clypeus ;
basal region very finely, transversely alutaceous and without coarse punctures.
Pronotum (for dimensions see Table VIII) with the greatest breadth at about
basal third. Sides moderately strongly arcuate at basal third, elsewhere only
feebly arcuate ; feebly and broadly sinuate at middle half opposite transverse
impression. Transverse impression at middle broad, moderately strongly
impressed and distinct to sides ; at middle bisected by a deep, oval impression
MEXICAN KI.MIDAE
Ml
which is (in a specimen 360 mm. long) 0100 mm. broad and 0125 mm. long ;
near middle third on basal third of pronotum an oblique impression extends on each
side so as nearly to join transverse impression at sublateral carinae, this oblique
impression being half again as broad and equally as strongly inpressed as the
transverse impression. Near base on each side of scutellum in front with an oval
feebly impressed area about two-thirds as large as median oval impression.
Surface punctate as middle of clypeus, the punctures being of various sizes so
that the smallest are no more than a fourth as large as the largest and are usually
round but often irregular, and are confluent to separated by about (on apical
discal portion) twice their diameters ; sides between sublateral carinae and lateral
margins slightly more coarsely but otherwise similarly sculptured to middle
region of head though often with round granules which are a little larger than
punctures. Elytra longer than pronotum (ratio usually about 92 : 42). On
basal third feebly but very broadly impressed from first interval to inner carinae.
Apices broadly and conjointly rounded. Striae feebly impressed on sides and
apical fourth, elsewhere not distinctly impressed ; strial punctures deep, round
and only occasionally on lateral intervals subquadrate. Intervals with the second
and third slightly broader than sutural and with the others narrower ; sutural,
second, and third elytral intervals on basal third two and one-half times as broad
as their respective punctures ; surface of intervals occasionally smooth but
generally with fine transverse wrinkles and with the punctures at base occasionally
as coarse as coarse pronotal ones, though mostly about a fourth to a third as
coarse as coarse pronotal ones and separated by two to five times their diameters.
Scutellum subovate and surface with punctures which are finer and more sparsely
separated than usual punctures of elytral intervals. Prosternutn with the carinae
present on basal three-fourths (not including process), straight, as wide apart
as process at base, prominent on basal half and indistinct on apical fourth ; sides
of process strongly and broadly gibbous so that middle of process is concave ;
process as broad as in the great majority of the species and with the posterior
margin feebly rounded, nearly truncate. Metasternum with the median longi-
tudinal line attaining anterior fourth of disk, anteriorly fine but posteriorly,
particularly on posterior third, broader and deeper. Disk only feebly convex on
each side of median impression ; surface of disk with the punctures usually
round, about as coarse as coarsest of pronotum, and separated by less than to
twice their diameters ; surface between punctures smooth, though at sides, par-
ticularly anteriorly, very densely set with punctures which are about a fifth again
as coarse as usual ones and here also densely and finely eroded and with an occa-
sional granule which is about half as broad as coarse mesal punctures. Abdominal
sternites with the non-tomentose areas punctate as disk of metasternum except
for basal three-fifths of busal segment which is slightly more coarsely and densely
punctate. Genitalia of male as figured (text-figs. 347-352).
/■ cmale : Externally similar to male.
Type : In the British Museum (Nat. Hist.). Mexico : Salazar [Flokr).
Specimens examined : Mexico: 1, Salazar (Hdge) ; 1, Las Vigas (T'lohr) ; 4,
Fedral District, La Venta, 1933 (//. /■.'. Hinton, R. L. Usinger) ; and 160, Dist. de
TemascaJtepec, Temascaltepec to Las Cruzes, alt. 5600-9000 ft., vi vii. 1032-34
(1932, 1934, H. E. Hinton) (1933, H. E. Hinton, R. L. Usinger).
Variations : The punctation of the various sclerites may be slightly finer or
coarser than that ol the specimen described above, the transverse impressions of
the pronotum an' often deeper and broader or shallower and narrower, and the
median oval impression of the pronotal disk in exceptional cases is hall as broad
NOVIT. ZOOL., 42, 2. I94O. 29
378
H. E. HINTON
and slightly longer than in the specimen described above. The transverse impres-
sion on basal third of elytra is occasionally absent. In a few specimens there is
on the pronotal disk a narrow and very shallow impression extending from base
to discal oval impression.
The following remarks apply to the District of Temascaltepec. H. longula
occurred nowhere below 5600 ft. It was the only Elmid taken as high as 9000 ft.
From my data it is not possible to determine at which altitude this species was
most abundant. The majority of the specimens before me were taken at alti-
tudes above 7000 ft. At this altitude most of the other Elmids began to fall off
354
351
352
Text-figs. 347-354. — Heterelmis longula Sharp. (347) Dorsal view of male genitalia.
(348) Right lateral view of same. (349) Dorsal view of median lobe. (350) Right
lateral view of median lobe. (351) Dorsal view of paramere. (352) Abdominal
spicule of male. (353) First segment of front tarsus. (354) Female genitalia.
rapidly in numbers, so that even if the numbers of longula had remained constant
at the different altitudes the method used for collecting would yield the greater
number at the greater altitude. At 5600-7500 ft., for example, it was often not
possible to examine more than 10 to 20 netfuls a day, so great were the numbers
of Elmids in each netful, but at 8000 ft. more netfuls could be examined in the
same time. The material was divided into two groups, 58 collected between
5600 and 7500 ft. and 66 collected between 7500 and 9000 ft. Owing to insuffi-
cient accurate altitude data the two groups are divided at the 7500-ft. level very
inaccurately, and it is not unlikely that specimens placed in the first group in
reality belong to the second and vice versa. However, this difficulty in separating
the two samples at their adjacent levels does not materially affect what follows.
MEXICAN ELMIDAI
379
All specimens wen- measured for the length of the pronotum, breadth at
base, breadth at broadest point (which is at about basal third), and breadth
across apex. The results of a statistical study of these measurements are recorded
in Table VIII. The chief interest of this table is that it shows that the individuals
p < .01
Cft E A7£ "a 7
355
Text-fig. 355. — Heterelmis longula Sharp. Dot diagram of the correlation between
the length of the prothorax and the breadth at broadest point. Measurements are
given in mm.
100 1125 1 1 -jo Hi 5 >:oO 1 IIS 1150 I tts
356
Text-fig. 356. — Heterelmis longula Sharp. Dot diagram of the correlation between
the breadth at base of the prothorax and the breadth at broadest point. Measure-
ments are given in mm.
living at altitudes of 5600-7500 ft. are on the whole larger than those living above
7500 ft. The differences between the means of the two samples may be regarded
as significant for the four different measurements (Table IX). An increase in
one of the prothoracic measurements indicates an increase in absolute size, foi
the length of the prothorax was found to be highly and positively correlated with
the absolute length of the individual and the length ol the prothorax highly and
NOVIT. ZOOL., 42, 2. 11140. 2g§
38o
H. E. HINTON
positively correlated with its breadth. Using the formula r = ^ y' , where r
NDxDy
is the coefficient of correlation, S(xy) the sum of the products of x and y for each
individual — x and y being the differences between the individual measures and
their means, N is the number of individuals and Dx and Dy the standard deviation
of the two series, r = 0720 for length times breadth at broadest point (text-fig.
1 — r2
355). From the formula Dr . the standard deviation of the coefficient of
correlation is found to be 0-06. The formula t =
. \/N — 2 gives the
VI — r2
value of t as 837, and this value in Fisher's (1936) table Va shows that it is
extremely improbable that the value for r should have arisen by random sampling
from an uncorrelated population, the probability being very much less than •01.
For breadth at broadest point times breadth at base (text-fig. 356) r = o-8i ±0-07,
t = 11 and P< -oi.
Table VIII.
The altitude of sample A is 5600-7500 ft., while that of sample B is 7500-9000 ft.
Measurements are given in mm. S.D. equals standard deviation, S.E.D equals standard
error of the deviation, and S.E.M. equals standard error of the mean.
The significance of the difference between the means of the two samples of longula from
M, - M2
different altitudes has been calculated from the formula x
where M is the mean and S.E.M. the standard error of the mean.
vAS.E.M.,)* + (S.E.M.,,)1
Comparative notes : This is the largest North American species of Heterelmis.
It is close only to H. obesa from which it may be distinguished as follows : (1)
the parameres of the male genitalia are broader in longula ; and (2) the median
lobe when seen from the side is broader and more sinuate, and the basal median
projection is gradually sloping behind instead of strongly declivous as in obesa
(cf. text-figs.). Nearly all individuals may be separated from obesa by their
greater length, the pronotum nearly always being more than 0-925 mm. long.
It has not been possible to separate very small females from large females of
obesa.
MEXICAN ELMIDAE
381
357
Text-fig. 357. — Heterelmis obesa Sharp.
363
365'"
Text-figs. 358-365. — Hen 1 dims obesa Sharp. (358) Dorsal view of male genitalia.
(359) Left lateral view of same. (360) Dorsal view of median lobe of male genitalia.
(361) Right lateral view of median lobe. (362) Sublateral view of paramere.
Presternum. (364) Sixth abdominal sternite of male. (365) Labial palp.
382
H. E. HIXTON"
Heterelmis obesa Sharp.
(Text-figs. 357-365.)
1882. Heterelmis obesus Sharp, Biol. Centr.-Amer. Co!., 1 (2) : 131, t. 4, f. 10.
Male: Length, 2-5 mm.-2-3 mm. ; breadth, i-i mm.-1.-5 rnm. Similar to
longula except as follows : (1) length of prothorax nearly always less than 0-925
li, 27 2i 2.1 Jo
Ja. jj 3* is
1*0 tH ti {,3 <t<t
1= O OJJ'»™,
366
Text-fig. 366. — Frequency polygons of prothoracic dimensions of Heterelmis adjusted
to equal areas. Smooth line refers to length of prothorax, while broken line
refers to its breadth at base, (a and 1) H. obesa plana Hinton ; (b and 2) H. obesa
Sharp ; (c and 3) H. longula Sharp.
mm. ; (2) pronotum seldom with a median discal impression but in the rare
specimen where this impression is well-developed it is never as deep as that
of longula, though it is often proportionally as broad and long ; (3) elytra only
rarely with a visible transverse impression on basal third ; (4) median lobe of
male genitalia with the basal median projection, seen from the side, sharply
MEXICAN ELMIDAE
383
instead of gradually sloping behind ; and (5) the parameres arc narrower than
those of longula.
Female : Externally similar to male.
Type: In the British Museum (Nat. Hist.). GUATEMALA: San Joaquin
{(r. C. Champion).
Specimens examined: 8, with same data as tin- type. 5003 from Mexico
as follows : 7, Altisco (/"'. D. Hodman) ; District of Temascaltepec : 4708, Teinas-
caltepcc to Real de Arriba, alt. 5600-7500 ft., 5.VU.1932 (H. E. Hinton), 1933
(H. E. Hinton, K. L. Usinger), and 1934 (H . E. Hinton) ; 27S, Rio Verde de la
1 = 0 on' .
367
Text-FIG. 367. — Frequency polygons adjusted to equal areas of the breadth of
the prnthorax at broadest point, (a) Heterelmis obesa plana Hinton ; (b) H. obesa
Sharp . (c) //. longula Sharp.
Comunidad, alt. 8000 ft., 14.vi.i034 (H. E. Hinton) ; 1, Tejupilco, alt. 3500-4000
ft., vi-vii, 1932 (H. E. Hinton), 4, 1933 (H . E. Hinton, R. L. Usinger), and 1, 1934
(H. E. Hinton) ; 4, Estado de Morelos, Cuernavaca, alt. 4800 ft., vi. 1034 (#. &■
Hinton).
Variations: The punctation oi the various sclerites is similar to that of
longula, and the variations in density are much the same. In a few specimens
the transverse pronotal impression is barely noticeable, but in the great majority
it is as well developed as that of longula.
A statistical study was made of the prothoracic measurements of two samples
1 let ted at random to find out if there was an increase ol size with an increase
in altitude. 45 specimens were taken from 5600-7500 ft. and 75 from 8000 It.,
and the pronofmn ol all was measured for length, breadth al broadest point
3«4
H. E. HINTON
(usually at about basal third), breadth across base, and breadth across apex.
The results of this study are shown in Table X. Table XI shows that there are
no significant differences between the means of the various prothoracic measure-
ments, so that it appears that a difference in altitude of probably more than
500 ft. does not on the whole affect the absolute size of the individual.
1 - O OA&' ~~™,.
368
Text-fig. 368. — Frequency polygons adjusted to equal areas of the breadth of the
prothoraxat apex, (a) Heterelmis obesa plana Hinton ; (b) H. obesa Sharp ; (c) H.
longula Sharp.
Table X.
The sample, consisting of 75 specimens, was collected at 6000 ft., while the other was
collected between 5600 and 7500 ft. Lengths are given in mm. S.D. equals standard
deviation, S.E.D. equals standard error of the deviation, and S.E.M. equals standard error
of the mean.
MEXICAN ELMIDAE
385
Comparative notes : Its small size usually enables it to be easily distinguished
from longula, but, as may be seen from the frequency polygons (text-figs. 366-
368), the various pronotal measurements occasionally overlap. When obesa is the
same size as longula, the males can only be distinguished by an examination of
the male genitalia and the females cannot, as far as I know, be separated.
Table XI.
Length .
Breadth at broadest
Breadth at base
Breadth at apex
The difference between the means of the various pronotal measurements is significant in
no case. Even where the difference is greatest, i.e. between the means for length, it is not
twice the standard error of the mean.
Heterelmis obesa plana, subsp. n.
(Text-figs. 369, 370.)
Male: Length, 2-12 mm. -2-65 mm. ; breadth, i-oo mm. -1-25 mm. Similar
to obesa except as follows : (1) there is no distinct transverse impression on the
pronotum ; (2) it is a slightly smaller species (Table XII) ; and (3) the pronotum
is proportionally broader, the value for the mean of the length into that of the
breadth at broadest point is 1-30, while the value for obesa is 1-24.
Female : Externally similar to male.
Type : g in the British Museum (Nat. Hist.). Mexico : Dist. de Temascal-
tepec, Tejupilco, alt. 3500-4000 ft., vii.1934 (H . E. Hinton).
Paraiypes : Mexico : 7, with same data as type ; 1178, Estado de Morelos,
Cuernavaca, alt. 4800 ft., vi.1934 (H. E. Hinton) ; 7, Sierra de Durango (C.
Schaufuss) ; and 1, Cuautla (Hbge).
Variations : There is a little variation in the density of the punctures on
various sclerites, but no other variations worthy of mention have been observed.
The coefficient of correlation, r, for length times breadth at broadest point
is : r = 079 i 0-034, t = H'4 and P<-oi. For breadth at broadest point times
breadth at base, r = 0-87 ± 0-03, t = 19-4 and P <-oi.
Table XII.
Length
Broadest
I i.im'
Apex
0-0342
o • 04 1 3
00358
0-O222
o- 00250
o- 00265
0-00229
000142
Lengths given in mm. of various prou.ital measurements ul a sample selected at random
o[ //. obesa plana from Cuernavaca. S I >. equals standard deviation, S.K.I), equals tin-
standard error of the deviation, and S.E.M. equals the standard error of the mean.
386
H. E. H1NT0N
0130 0|13 0<iOO 0115 01SO 0 175 I boo icii 1050 . ili 1100
GRF A~EST aRCAJTH
369
Text-fig. 369. — Heterehnis obesa plana Hinton. Dot diagram of the correlation
between the breadth of the prothorax at base and the breadth at broadest point.
Measurements are given in mm.
o-tso
ous
0100
0725
I 0 750
0H5
0 100
0 lib
0 850
0 115
©Sao ojiS 0100 0U5 oiso 0115 ' °°o iol5
GR.CHTE&T 6ft£H3TM
370
Text-fig. 370. — Heterehnis obesa plana Hinton. Dot diagram of the correlation
between the breadth of the prothorax at broadest point and its length. Measure-
ments are given in mm.
MEXICAN' ELMIDAE
387
Heterelmis obscura Sharp.
(Text-figs. 371-376.)
1882. Heterelmis obscurus Sharp, Biol. < enlr.-Amer. < 61., 1 (2) : 130.
1888. Heterelmis obscura Grouvelle, Ann. Sue. lint. Fr.. 8 (6) : 400, t. 7, f. 8.
Male: Length, 1-9 mm. -2-2 mm. ; breadth, 0-9 mm.-i-i mm. Similar to
H. obesa except as follows : (1) the body is as a general rule proportionally more
elongate, seldom as broad ; (2) the impressions of the pronotum, particularly
376
Text-figs. 371—376. — Heterelmis obscura Sharp. (371) Dorsal view of male genitalia.
(372) Left lateral view of same. (373) Dorsal view of median lobe. (374) Right
lateral view of median lobe. (373) First segment of front tarsus. (5J(>) Dorsal view
of right paramere.
apical transverse impression, are deeper and more distinct, these impressions
being similar to those of longula but the median discal impression is as a rule not
so well developed ; (3) the ventral apex of the basal segment of all tarsi is as
figured (text-fig. 375), i.e. without two stout spines as in obesa and other members
of the longula group ; and (4) the parameres of the male genitalia (text-tigs. 371-
374, 376) have a conspicuous fringe of fine, long hairs on the inner apical half.
Female : Externally similar to male.
Type : In the British Museum (Nat. Hist). Guatemala: San Joaquin
[G. C. Champion).
Specimens examined: 34. with same data as type; and ;. GUATEMALA:
Guatemala City [G. C. Champion). Texas: i. Ft. Sam Houston. <'"si \ Ri< \ :
388
H. E. HINTON'
I, San Jose, alt. 1000-1200 m., 18. v. 1931 (F. Nevermann) ; and 1, Coronado,
alt. 1400-1500 m. 1607 from Mexico as follows : 249, Estado de Morelos,
Cuernavaca, alt. 4800 ft., vi.1934 (H. E. Hinton) ; Dist. de Temascaltepec : 21,
Tejupilco, alt. 3500-4000 ft., vi-1933 {H. E. Hinton, R. L. Usinger) ; 13, as pre-
ceding but in vii.1934 (H. E. Hinton) ; 1324, from Temascaltepec to Rio Verde
de la Comunidad, alt. 5600-8000 ft., v-vii (1932, 1934, H. E. Hinton) and 1933
{H. E. Hinton, R. L. Usinger).
Other locality records : Grouvelle (1888, loc. cit.) has recorded this species
from Theresopolis, Brazil, but since he did not compare the male genitalia
of his specimens with those of typical obscura from Central America, his locality
record cannot be considered as proof that this species occurs in southern Brazil.
Variations: As usual, the punctures on the different sclerites vary slightly
in density, and, apart from size, no other variations worthy of mention have been
observed.
Table XIII.
The sample of 28 specimens is from Rio Verde de la Comunidad, alt. Sooo ft. ; that of
38 is from Temascaltepec to Real de Arriba, alt. 5600-7500 ft. ; that of 73 is from Cuernavaca,
alt. 4800 ft. ; and that of 14 is from Tejupilco, alt. 3500-4000 ft. The measurements are
given in mm. S.D. equals the standard deviation, S.E.D. equals the standard error of the
deviation, and S.E.M. Equals the standard error of the mean.
A statistical study was made of the length of the pronotum and the breadth
at three different points. The samples were selected at random and each of the
four samples is from a different altitude. The results of this study are recorded
in Table XIII, and the significance of the difference of the means for all possible
pairs of altitude samples is given in Table XIV. A comparison of the sample
(73) collected at Cuernavaca (4800 ft.) with (14) that collected at Tejupilco
(3,500-4000 ft.) shows that for each pronotal measurement the mean value is
slightly greater for the former, but in no case is the difference significant. The
minimum difference in the altitude between these two samples is 800 ft. This
fact justifies the conclusion that in latitude 19° N. a difference in altitude of more
than 800 ft. is probably necessary before any significant difference in size can be
detected.
MEXICAN ELMIDAE
389
The altitudes of the four samples are given in Table XIII. The significance of the
difference between the means of the samples was calculated from the formula x =
— . „ „ ' " „ .. where M is the mean and S.E.M. the standard error of the mean.
V/(S.E.M.,)! + (S.E.M.,)»
The probabilities, P, for the various values of x were found in Fisher's (1936) table of x.
Comparative notes: No difficulty i> met with in distinguishing H. obscitra
from the species belonging to the H. longula group, all of which have the first
segment of the tarsi modified. In size and general appearance obscitra only
approaches — of those species I have already dealt with — obesa, and apart from the
differences mentioned above, the pronotal measurements of these two but rarely
overlap ; and in practice it has been possible to separate them on size with great
ease, only about 1 in 200, being mistaken.
Heterelmis acicula, sp. n.
(Text-figs. 377-381.)
Male : Length, 1-9 mm. -2-2 mm. ; breadth, ro mm.-ri mm. Similar to
male of obscura except as regards the structure of the male genitalia (cf. text-figs.)
which differ as follows : (1) the parameres are more slender and the apical fringe
of hairs is sparser ; (2) the median lobe is narrower (0-0170 mm. broad) opposite
apices of parameres, whereas the median lobe of obscura at the same point is
0-050 mm. broad ; and (3) the basal median projection of the median lobe is,
when seen from the side, rounded instead of pointed at the end, and immediately
behind is strongly instead of feebly sinuate.
Female : Unrecognized. There are probably a number of females before me,
but if so, I cannot distinguish them from the females of obscura. This means
that wherever obscura and acicula are found together, e.g. in Tejupilco, female
specimens cannot be assigned to their respective species.
Type : J in the British Museum (Nat. Hist.). Mexico : Dist. de Temascal-
tepec, Tejupilco, alt. 3500-4000 ft., vii.1934 (H. /•.'. Hinton).
Paratypes : 15, with data as above : and 16, collected in the same locality
in vi.1933 (H. E. Hinton, R. L. I 'singer).
I 'ariations : No variations worthy of mention have been observed.
Comparative notes: to male specimens were used in a statistical study to
determine the means oi four pronotal measurements and the significance of the
39°
H. E. HINTON
differences between these means and those of 14 male specimens of H. obscura also
collected in Tejupilco. From the results given in Table XV it is obvious that
acicula is a smaller species. In Table XVI the probabilities of the two species
being homogeneous for size are given. When acicula occurs at the same altitude
and latitude as obscura it is a smaller species. It is important to remember that
these size differences between the two species only apply in the same altitudes
and latitudes, e.g. if the great difference in the size of obscura taken above 7000 ft.
and below 4000 ft. is considered, it can be reasonably inferred that at lower
altitudes than Tejupilco we will get populations equal in size to those of acicula
at Tejupilco.
377
378
379
380
381
Text-figs. 377-381. — Heterehnis acicula Hinton. (377) Dorsal view of male genitalia.
(37S) Left lateral view of same. (379) Dorsal view of median lobe of male genitalia.
(380) Left lateral view of same. (381) Dorsal view of left paramere.
Larvae.
The larvae have not been associated with Heterehnis by rearing but by con-
stantly finding them associated with the adults over a number of years and in
many localities in Mexico, Peru, Bolivia, Brazil, French Guiana, and Trinidad
(B.W.I.). A distributional map of the larvae here considered to belong to
Heterehnis will fit only that of the adults of Heterehnis.
Generic Characters of Larvae of Heterelmis.
Body subparallel and cylindrical to subtriangular in cross section ; dorsal
surface (except for ninth abdominal tergite which has median longitudinal ridge)
MEXICAN ELMIDAE
391
evenly convex and with parallel, feebly oblique rows of closely placed and large
tubercles each of which bears a large, stout spine. Head when seen from above-
exposed and not concealed by the pronotum ; anterior margin on each side
between base of antenna and clypeus with a large and acute tooth. Clypeus
with the fronto-clypeal suture well developed. With one ocellus on each side.
Antenna (Text-fig. 388) 3-segmented and feebly retractile. Mandibles ot both
sides similar and with three obtuse, apical teeth ; prostheca long, slender, and
densely spinose. Maxilla (text-fig. 386) with the palp 4-segmented and the stipes
showing no differentiation into a palpifer ; galea and lacinia separate and apex
Measurements are given in mm. S.D. equals standard deviation, S.E.D. equals standard
error of the deviation, and S.E.M. equals standard error of the mean. Both samples are from
Tejupilco, alt. 3500-4000 ft.
Table XVI.
The significance of the difference of the means of four pronotal measurements of obst ura
\ I M
compared with acicula was calculated from the formula x = — r=
1 t/(S.E.M.,)" (S.E.M.*)"
where M is the mean and S.E.M. the standard error of the mean. The probabilities for
various values of x were found in Fisher's (1936) table of x. the samples of both spei ie
are from Tejupilco.
of each densely spinose. Labium with the postmentum undivided ; labial palp
(text-fig. -jSj) 2-segniented and prementum without a distinct palpiger. Gula
well developed. Prothoracic pleura (text-fig. 30.1) divided into two parts and
anterior part meeting on middle line of body so that the sternum is here com-
pletely suppressed. Meso- and metapleura divided into three parts on each side
(text-fig. .101). Abdominal segments one to seven with the pleura bounded by
tergo- and sterno-pleural sutures which in seventh segment (text-fig. 384) con-
verge and meet near posterior margin ; segmenl 8 forming a complete sclerotized
tint; ; apex of ninth very feebly and broadly, arcuately emarginate. Operculum
(text dig. 401) with two strongly sclerotized claws (text-tig. 398) attached to its
dorsal membrane. Spiracles present on mesothorax and first eight abdominal
segments and opening at apices of small tubercles; tracheae without air sacs ;
with three tufts of anal, retractile, tracheal gills. Alimentary canal (text-fig. 385)
392
H. E. H1NTON
with an oesophageal sclerite on the dorsal posterior margin of the oesophagus.
Hind gut with six Malpighian tubules which end freely near the rectum. Central
nervous system (text-figs. 382) with three thoracic and eight abdominal discrete
ganglia.
The larvae of this genus have a very characteristic appearance due to the
slightly oblique and parallel rows of close and large tubercles. Each tubercle
bears a spine so that the larvae appear to have a series of sharply carinate ridges
when viewed dorsally. The division of the meso- and metapleura into three parts
on each side will distinguish the genus from all genera except Phanoceroides Hinton,
h
Jl
1.
[
,,
382
385
Text-figs. 382-385. — Larva of Heterelmis languid Sharp. (382) Central nervous
system. (383) Dorsal view of head. (3S4) Ventral view of seventh abdominal
segment to show sclerotization. (385) Alimentary canal.
and from this it may be distinguished as follows : (1) its cylindrical instead of
flattened and onisciform body ; (2) the prothoracic pleura are divided into anterior
and posterior parts, whereas in Phanoceroides they are undivided ; and (3) by
the eighth abdominal segment which here forms a complete sclerotized ring,
whereas in Phanoceroides it has the pleura bounded by tergo- and sterno-pleural
sutures.
Description of Mature Larva of Heterelmis longula Sharp.
(Text-figs. 382-401.)
Length, 7-6 mm. ; breadth, at broadest point which is near apex of
abdomen i-o mm. Head with the frontal sutures complete (text-fig. 383) ; coronal
MEXICAN ELMIDA]
•>'i.;
suturi- short (ttxt-fi^. 383) ; head without other sutures. Parietals finely tuber-
culate on a basal belt which is as long as coronal suture : elsewhere for the most
part tuberculate as figured (tuberculate area of drawing) (text-fig. 383) and with
a number of fine setae ; four post-parietal setae (text-fig. 383) placed in an oblique
row. Frons with a tooth-like process on each side and with the surface tuber-
culate as the parietals. Clypeus about as broad as base of labrum but only
about a third as long. Labrum as figured (text-fig. 389). Ocellus on each side
composed of a single large pigmented area and hence not distinctly faceted.
fy* * 1 >ft ^tWX
389
390
Text-figs. 386-390. — Larva of Heterelmis longula Sharp. (386) Ventral view of right
maxilla. (387) Ventral view of labium. Setae of prementum are only placed to
Hive general appearance of this region. (388) Dorsal view of left antenna. (389)
Dorsal view of labrum. (390) Mesothoracic spiracle.
Antenna with the setae as figured (text-rig. 388). Mandibles of both sides similar ;
one-fourth longer than broad at broadest point which is at base ; and with a
seta-like prostheca which is about two-thirds as long as mandible and is itself
densely and very finely setose. Maxilla and labium with the setae as figured
(text-figs. 386, 387). Pronotum at broadest point, which is near base, slightly
broader than long (0-925 mm. : 0-850 mm.) ; anterior margin broadly and feebly
arcuate, apical angles obliterated, sides feebly arcuate, basal angles inconspicuous
and broadly rounded, and base feebly arcuate for its entire breadth ; surface very
evenly convex ; surface anteriorly with round to oval setose tubercles which are
about 0-025 mrn- broad and are separated by less than to twice their diameters ;
on posterior three-fifths of pronotum most of these tubercles are arranged in five
394
H. E. HINTON
rows on each side which are feebly oblique towards sides and become increasingly
well defined posteriorly ; posteriorly the tuberculation of this tergite resembles
that of the fifth abdominal segment (vide text-fig. 395) ; anteriorly this segment
has a row of flat setae which is similar to posterior row and both of these rows are
similar to posterior row of fifth tergite (text-fig. 395). Tergites of mesothorax,
metathorax, and first eight abdominal segments with the ratio of breadth to
length as follows : 0-950 mm. : 0-450 mm. ; 0-975 mm. : 0-450 mm. ; i-oo mm. :
0-400 mm. ; i-oo mm. : 0-400 mm. ; 0-975 mm. : 0-450 mm. ; 0-925 mm. : 0-425
39 5
Text-figs. 391-395. — Larva of Heterelmis longula Sharp. (391) Ventral view of thorax
and first abdominal segment to show sclerotization. (392) Dorsal view of ninth
abdominal segment. (393) Section to show tuberculation near anterior margin of
fifth abdominal tergite. (394) One of the tubercles forming the rows on fifth abdo-
minal tergite. (395) A section of fifth abdominal tergite to show various types of
tubercles.
mm. ; 0-850 mm. : 0-425 mm. ; o-8oo mm. : 0-450 mm. ; 0-700 mm. : 0-425 mm.
0-625 mm- : °'375 mnl- I aU 0I these tergites are alike in having the anterior and
posterior margins nearly truncate, the sides feebly arcuate, the apical and basal
angles broadly rounded, the surface evenly convex, and the tubercles and setae
as shown in text-fig. 395. Ninth abdominal tergite with the outline as shown in
text-fig. 392 ; with a median ridge extending from base to apex and on this ridge
are two nearly contiguous rows of tubercles, the rows diverging slightly basally ;
on each side with an extreme dorso-lateral row of setose tubercles extending from
base to apex, and also on each side with a ventro-lateral row of similar tubercles ;
areas between these rows of tubercles have themselves somewhat similar rows of
tubercles which are separated by less than to twice their diameters ; apical
MEXICAN ELMIDAE
395
margin with a conspicuous fringe of posteriorly directed setae which are about
0-03 mm. long. Ventral surface with the setose tubercles about two-thirds as
coarse as those of tergites and usually separated by one to three times their
diameters — two common types are figured from the fifth abdominal stemite
(text-figs. 396 and 400), and there are setae showing every intergradation between
them. Operculum with the shape as figured (text-fig. 401) and the claws as figured
(text-fig. 398). Legs all somewhat similar to front leg (text-fig. 397).
401
Text-figs. 396-401 . — Larva of Heterelmis longula Sharp. (396) One of the types of seta
found on the fifth abdominal sternite. (397) Inner face of right front leg. (398)
Dorsal face of right opercular claw. (399) Mature larva to show general appearance.
(400) One of the types of seta found on the fifth abdominal sternite. (401) Oper-
culum.
Specimens examined : 4, Mexico : Dist. de Temascaltepec, Las Crazes, alt.
9000 ft., vi.1934 (H. E. Hinton).
In addition there are 119 larvae collected in the same district from Tejupilco
alt. 3500 ft., to Rio Verde, alt. 8000 ft. ; and 31, Mexico: Estado de Morelos,
Cuernavaca, alt. 4800 ft., vi.1934 (H. E. Hinton).
From the distributional data available it is evident that there are at least
two species and one variety before me (longula, obesa, and obesa plana) and possibly
two other species (obscura and acicula). The larvae of longula were determined
as such by the altitude data, this being the only species which occurs in the Dist.
de Temascaltepec as high as 9000 ft. What is apparently the last stage larva of
396 H. E. HINTON
this species may be distinguished from the last stage larva taken at Cuernavaca
(where longula does not occur) as follows :
1 . Abdominal tergites of last stage larva without tubercles or only with an
occasional smaller tubercle between first and third rows of tubercles
H. obesa, H. obesa plana, and H. obscura.
Abdominal tergites of last instar larva with numerous large tubercles
between the first and third rows of tubercles (text-fig. 395) H. longula.
Larvae representing probably three earlier instars are before me. These are
all alike and similar to the mature larva except in the case of longula where the
mature larva has numerous tubercles between first and third rows of tubercles
of abdominal terga.
REFERENCES.
Bertrand, H. 1936 Captures et elevages de larves Coleopteres aquatiques. Ann. Soc.
Ent. Fr., 105 : 213-238, 1 pi., 44 figs.
Carter, H. J., and Zeck, E. H. 1929 A monograph of the Australian Dryopidae. Order:
Coleoptera. Aust. Zool., 6 (1) : 50-72, 7 pis.
1932 Four new species of Dryopidae, together with notes on the family.
(Order Coleoptera.) Aust. Zool., 7 (3) : 202-205, l P'-
Champion, G. C. 1923 Some Indian Coleoptera (11). Ent. man. Mag., 59: 165-179,
5 figs-
Darlington, P. J. 1936 A list of the West Indian Dryopidae (Coleoptera), with a new
genus and eight new species, including one from Colombia. Psvche, 43 (2-3) : 65-83,
1 pi.
1936 Variation and atrophy of flying wings of some Carabid beetles (Coleoptera).
Ann. Ent. Soc. Amer., 29 (1) : 136-179, 3 pis., 2 figs.
Dufour, L. 1835 Recherches anatomiques et considerations entomologiques sur les
insectes coleopteres des genres Macronique et Elmis. Ann. Sci. Nat. {Zoo!.), (2) 3 :
I5I-I74-
Erichson, W. F. 1847 Naturgeschichte der Insecten Deutschlands , 3, Berlin.
Fisher, R. A. 1936 Statistical methods for research workers. London.
Forbes, Wm. T. M. 1926 The wing folding patterns of the Coleoptera. /. N. Y. Ent. Soc,
34 : 42-68, 91-139.
Ganglbauer, L. 1904 Die Kafer von Mitteleuropa, 4 (1). Wien.
Grouvelle, A. 1888 Nouvelles especes d'Helmides. Ann. Soc. Ent. Fr., 8 (6) : 393-410,
pis. vii, viii.
1896 Note sur les subdivisions generiques des Potamophiliens. Bull. Soc. Ent. Fr. :
77-78.
Hinton, H. E. 1934 Miscellaneous studies in the Helminae (Dryopidae, Coleoptera).
Rev. Ent., Rio de J ., 4 (2) : 192-201.
1935 Notes on the Dryopoidea (Col.). Stvlops, 4 (8) : 169-179, 7 figs.
1936 A new genus and a new species of Elminae (Coleoptera, Dryopidae). Ent.
mon. Mag., 72 : 1-5, 7 figs.
1936 Descriptions of new genera and species of Dryopidae (Coleoptera). Trans.
R. Ent. Soc, Lond., 85 (18) : 415-434, 1 pi., 43 figs.
1937 Descriptions of new Brazilian Dryopidae and distributional records of others.
Ent. mon. Mag., 73 : 6-12, 4 figs.
1937 Additions to the Neotropical Dryopidae. (Coleoptera.) Arb. morph. taxon.
Ent. Berlin-Dahlem, 4 (2) : 93-1 11, 21 figs.
Musgrave, P. N. 1935 Two new Elmidae from Puerto Rico with description of new
genus (Coleoptera). Proc. Ent. Soc. Wash., 37 (2) : 32-35, 1 fig.
Richards, O. W. 1938 The formation of species. [In] Evolution ; essays . .
presented to E. S. Goodrich on his ■joth birthday. Edited by G. R. De Beer. : 95-110
Oxford.
Schaeffer, C. 1911 New Coleoptera and miscellaneous notes. /. .V. Y. Ent. Soc, 19
113-126.
Sharp, D. 1882 Heteroceridae, Parnidae, Georissidae. Biol. Centr.-Amer., Col., 1 (2)
116-141 ; 1887, Col. 1 (2) : 772-775.
Zarapkin, S. R. 1934 Zur Phanoanalyse von geographischen Rassen und Arten. Arch
Naturgesch., n.f., 3 : 161-186, 9 figs.
(MS. reed. Sept. 1, I939-) ,
THE ZOOLOGICAL RECORD
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CONTENTS
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A Monographic Revision of the Mexican Water
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NOVITATES ZOOLOGICAE
Vol. 42 1948 No. 3
ENTOMOLOGICAL RESULTS FROM THE SWEDISH
EXPEDITION 1934 TO BURMA AND BRITISH INDIA—
COLEOPTERA : CURCULIONIDAE
By Sir Guy A. K. Marshall, K.C.M.G., F.R.S.
HPHE fine collection of Curculionidae made by Dr. Rene Malaise in Burma
■*■ serves to illustrate once again the striking richness of that fauna. The dates
on the specimens range from April to November, but the great majority was
taken in only four months — April to July. In all, 343 different species were found,
but unfortunately a high proportion of these were represented by single specimens
that could not be determined.
Part of the collection was dealt with in a paper published in 1941 (Ann.
Mag. nat. Hist, (n) 8: 345-379) comprising the subfamilies Brachyderinae,
Otiorrhynchinae, Eremninae and Sitoninae. The species identified amount to
196, of which no less than 114 were new to science. If it be assumed that of the
147 unnamed species 85 per cent, are new (a conservative estimate) the new forms
would represent 70 per cent, of the species.
Although, as may be seen from the accompanying map, Dr. Malaise covered
a fairly wide area in his expedition, from Tenasserim in the south to Myitkyina
in the north, the great majority of the weevils were taken at Kambaiti, due east
of Myitkyina, as shown by the fact that of the 114 new species no less than 87 are
recorded from that locality only.
The only large collection of weevils from Burma that has previously been
worked out fully is that made by Leonardo Fea during four years' residence
(1885-89) for the Genoa Museum. This was dealt with by Faust (Ann. Mus.
Stor. nat. Genova, 34, 1894 [1895] : 153-370), who recognized about 450 species, of
which he identified 321, and of these 184 were new ; he estimated that in the
whole collection 60 per cent, (or 270 species) were probably new.
An interesting point is the great difference in the two collections. The
combined total of identified species is 483, yet of these only 34 are common to them
both. This is no doubt mainly due to differences in the localities investigated, for
Fea's captures came principally from farther south, especially around Bhamo
and in the Karen Hills ; but differences in methods of collecting seem also to have
affected the results.
NOV IT. ZOOL., 42, 3. 1948.
30
398
SIR GUY A. K. MARSHALL
0 50 100 150 200 Miles
0 ii5o 200 360 K.lometcrs
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 399
Except where otherwise stated, the types of all the new species are in the
Stockholm Museum and cotypes in the British Museum (Natural History).
In those cases in which no collector's name is given the specimens were
collected by Dr. Rene Malaise.
Subfamily Cleoninae.
Tetragonothorax macilentus Oliv. — Rangoon, 1.9, xii.
Nemoxenus zebra Chev. — 20 km. E. of Myitkyina, c. 200 m., vii.
Xanthochelus fannus Oliv. — N.E. Burma, Sadon, 1200 m., i<J, vi-vii. S. Shan
States, 40 km. E. of Taunggyi, 3$ 1$, ix-x.
Lixus vetula F. — S. Shan States, Taunggyi, 1500 m., 4(^3$, viii-ix.
Lixus prainae Fst. — 40 km. E. of Taunggyi, 1^, ix-x.
Lixus languidus Fst. — N.E. Burma, Sadon, 1^ 1$, vi-vii.
Gasteroclisus arcurostris Petri. — N.E. Burma, Sadon, 1200 m., 2$ 1$, vi-vii.
Subfamily Hylobiinae.
Peribleptus scalplus Boh. — Kambaiti, 7000 ft., 4^ 7$, v-vi. ; Sadon, 4000 ft.,
1 (j 1$, vi-vii ; Punkaung, between Sadon and Myitkyina, z£, vii. ; Washaung,
20 km. E. of Myitkyina, 2$, vii.
Peribleptus bardus Mshl. — Kambaiti, 6<J, v.
Peribleptus frenatus Mshl. — Kambaiti, i<J, v.
Peribleptus bisulcatus Fst. — S. Shan States, Taunggyi, 2<$ 2$, ix.
Peribleptus dealbatus Bdv. — Sadon, 4000 ft., 1$, vi-vii.
DYSCERUS Faust
Heller (Abh. Mus. Tierk. Volkerk. Dresden, 17 (3) 1929 : 12) sank Dyscerus Fst.
as a synonym of Pagiophloeus Fst., on the ground that the characters given by
Faust appeared to be unreliable ; but the assemblage of species thus formed seems
far from homogeneous and will probably need redivision, perhaps on other
characters. Moreover, I am not yet acquainted with the genotype of Pagiophloeus
(pacca F.), an examination of which will be necessary to establish the relationship
of the two genera. Meanwhile I provisionally refer to Dyscerus those species in
which the front tibiae curve inwards at the apex, however slightly, and to
Pagiophloeus those in which the upper edge of these tibiae is quite straight in the
apical two-thirds.
Dyscerus malaisei sp. n.
<J$. Derm dull black ; prothorax with a faint median stripe formed of two
or three rows of fulvous setae, some shorter setae along the basal margin, and a
patch above the front coxae ; elytra with a few inconspicuous . small spots of
fulvous setae, mostly forming a sinuous macular band at about one-third from the
base and across the top of the declivity ; underside with very sparse pale setae and a
narrow fulvous band across the base of the metepisterna.
Head longer than usual, subcorneal, bare, with rather coarse subconfluent
punctures that become smaller behind ; frons flat, two-thirds the width of the
base of the rostrum, with a deep median fovea. Rostrum elongate, rather slender,
gradually narrowing from base to middle, strongly dilated at apex, longer than
the pronotum, rather strongly curved, the dorsal outline continuous with that of
the head but sometimes separated by a small shallow impression in £ ', cS with
coarse confluent punctation from base to antennae and a single low median
carina, the punctures in $ smaller, non-confluent and confined to the basal half,
400 SIR GUY A. K. MARSHALL
the median carina obsolescent, without any trace of a sulcus above the scrobe in
either sex. Antennae piceous, the funicle with joints i and 2 equal, 3 a little
longer than 4, 4-6 beadlike, 7 slightly longer and broader. Prothorax nearly as
long as broad, variable in shape, being sometimes more or less rounded laterally
and widest near the base, or subconical with the sides almost straight and widest
at the base, the apical constriction also varying from deep to shallow ; the apical
margin dorsally truncate with a small sinuation in the middle, without any trace
of postocular lobes, the gular margin not sinuate ; dorsum flat longitudinally in
the median line, set with very large, more or less confluent, shallow foveae,
the intervals between them forming irregular variable narrow ridges, without any
trace of a median carina. Scutellum rather large, triangular, about as long as its
basal width, slightly raised, closely and finely punctate, setose. Elytra one-third
wider at the roundly rectangular shoulders than the base of the prothorax,
parallel for three-fourths of their length, with large obtuse posterior calli, the
apices very shortly and separately acuminate ; the shallow striae partly obscured
on the disk by more or less irregular large shallow foveae, which are often laterally
confluent and disappear entirely on the declivity ; the intervals narrow, irregular
and often partly obliterated, the higher parts usually with aggregations of small
flattened granules, intervals 3 and 5 being slightly more raised than the others.
Legs long and slender, black, with very sparse short white setae ; femora with
sparse fine punctures on the basal half, wrinkled apically ; anterior pairs of tibiae
incurved apically and hardly bisinuate on the inner edge.
Length 14-15 mm., breadth 5.5-6.0 mm.
N.E. Burma : Kambaiti, 7000 ft., 4^ 2$, iv-vi. 1934.
This interesting species presents three characters which have not previously
been noted in this genus, namely : the unusual prolongation of the head ; the
absence of any deep sinuation in the gular margin of the prosternum ; and
especially the absence of the punctate longitudinal sulcus on the rostrum just
above the scrobes.
Dyscerus posticus sp. n.
cJ. Derm red-brown, the apex of the rostrum, the basal half of the prothorax,
and a variable band across the top of the elytral declivity, blackish ; upper
surface almost bare, but the declivity of the elytra with rather numerous recum-
bent setae caked with yellowish powder.
Head with rugose confluent punctures ; frons half as wide as the base of the
rostrum, with a median fovea. Rostrum a little longer than the pronotum (9 : 8),
comparatively slender, almost parallel-sided from the base to the scrobes and
slightly widened apically ; dorsum with rather coarse confluent punctation and five
narrow low carinae, and a shallow sulcus above the scrobe. Antennae slender,
red-brown ; funicle with joint 1 longer than 2, 3-7 moniliform, subequal, 7 not
wider than 6 ; club more slender than usual. Prothorax very nearly as long as
broad, very gradually narrowing in a slight curve from the base to the broad
shallow apical constriction, the apical margin feebly arcuate dorsally, the gular
margin sinuate, without postocular lobes ; dorsum with the apical area coarsely
confluently punctate, the rest fairly closely set with small shiny convex granules,
each bearing a short recumbent seta, and a short median carina on the anterior
half. Scutellum subquadrate, sparsely setose. Elytra wider than the base of the
prothorax (13 : 9), parallel for two-thirds of their length, separately pointed at
the apex, the posterior calli prominent and obtusely angulate ; the rows of large
round punctures entirely regular, the rather sinuous intervals narrower than the
BURMA AND BRITISH INDIA— COLEOPTERA : CURCULIONIDAE 401
septa between the punctures, 3 being a little higher than the others and all with
an unevenly spaced row of small flattened granules, the setae, apart from those
on the declivity, sparse and minute. Legs red-brown, rugosely punctate and with
sparse yellow setae, all the femora with a round black apical spot on the inner face
and the hind pair with a larger one near the tooth ; tibiae incurved at apex and
very shallowly bisinuate on the lower edge, the front pair with a distinct tri-
angular mucro.
Length 9.5-10.0 mm., breadth 3.5 mm.
N.E. Burma : Kambaiti, 7000 ft., 2^, vi. 1934.
Dyscerus ordinal us sp. 11.
(J$. Derm dull black ; prothorax with sparse yellow setiform scales and a
broad ill-defined lateral stripe of these scales on each side ; elytra with small spots
of similar scales, some of them forming a common, deeply bisinuate, macular
band before the middle, a spot on intervals I and 2 before the middle, an indefinite
transverse macular band across the top of the declivity, and some irregular
markings on the declivity.
Head with small dense punctures and sparse setae ; frons two-thirds the
width of the base of the rostrum, with a small shallow fovea. Rostrum a little
longer than the pronotum, almost parallel-sided to the antennae, only slightly
widened at the apex, rather strongly curved and with a shallow depression
between the base and the frons ; dorsum in $ with rather rugose confluent
punctures and five low narrow carinae that do not unite at the base, there being
two shallow sulci laterally ; rostrum of $ more shiny, with the punctures less
rugose. Antennae dark red-brown ; funicle with joint 1 distinctly longer than 2,
3-7 moniliform, subequal, 7 scarcely broader. Prothorax slightly transverse
(8:9), rounded laterally, widest behind the middle, very slightly constricted at
apex, the apical margin gently arcuate dorsally, the gular margin deeply sinuate,
the postocular lobes feeble ; dorsum with more or less confluent small rugose
punctures and an abbreviated low rugose median costa. Scutellum shield-shaped,
rather densely setose. Elytra much broader at the shoulders than the base of the
prothorax (3:2), parallel for two-thirds of their length, with obtuse posterior
calli, the apices separately produced into a short sharp process ; the striae nanow
and quite regular throughout, containing small separated punctures ; the intervals
much broader, mostly flat but with low granular tubercles on which the scale
spots are situated, intervals 3 and 5 slightly higher than the others, especially near
the base where all the intervals bear some low shiny granules. Legs slender,
black, with sparse pale setae, the apices of the tarsi red-brown ; tibiae incurved
at the apex and shallowly sinuate beneath.
Length 9.5-11.0 mm., breadth 3.4-4.0 mm.
N.E. Burma : Kambaiti, 7000 ft., 6<$ 3$, v-vi. 1934.
Dyscerus direptus sp. n.
cJ$. Derm dull black, with markings formed of narrow fulvous scales ;
prothorax with a broad lateral fulvous stripe on each side, usually extending
downwards over the whole pleurae, but the pleurae may also be bare with a
single median spot and a narrow stripe just above the coxa ; elytra with small
fulvous spots arranged in the pattern which is highly characteristic of this genus :
three or four spots on interval 2, a curved macular band before middle on 4-9,
and a sinuous macular band across the top of the declivity.
402
SIR GUY A. K. MARSHALL
Head globular, bare, with dense small punctures, sometimes confluent ;
irons more than two-thirds the width of the rostrum at its base (5 : y), with a
deep median fovea. Rostrum a little longer than the pronotum, gradually
narrowed from base to middle, moderately dilated at apex, rather strongly curved,
with a shallow depression between the base and the head ; dorsum with irregular
shallow confluent punctures and three low narrow sinuous carinae that unite at
base, and a sulcus above the scrobes. Antennae piceous ; funicle with the two
basal joints equal, 3 longer than 4, 4-7 moniliform, 7 not broader than 6.
Prothorax a little longer than broad, parallel-sided from base to middle, then
narrowing to the deep collar-like apical constriction, the apical margin gently
arcuate dorsally, the gular margin deeply sinuate, the postocular lobes distinct ;
dorsum moderately convex longitudinally, highest at the middle, uneven, very
rugose, with unevenly separated granular elevations and a broad high rugose
median carina that does not reach base or apex. Scutellum almost semicircular,
smooth, bare, impunctate. Elytra wider than the prothorax (13 : 10), parallel
to beyond middle, with obtusely angulate posterior calli, the apices separately
acuminate ; the rows of large separated punctures mostly irregular on the disk
and sometimes transversely confluent here and there ; the intervals very irregular
and uneven, interval 1 with a single row of distant depressed granules, 3 with a
low subgranulate elongate elevation on the basal fourth (often interrupted), a
similar short elevation at the middle and another at the top of the declivity, 5
with several small distant granular tubercles, and some smaller ones on 7. Legs
as in bigutticollis, sp. n.
Length 10-12 mm., breadth 3.7-4.5 mm-
N.E. Burma : Kambaiti, 7000 ft., 2^2$, iv-vi. 1934 (type) ; Sadon, 4000 ft.,
1?, vi-vii. 1934.
Dyscerus asper sp. n.
<J$. Derm dull black, with sparse pale yellowish setae ; pronotum without
markings ; elytra with a small spot of denser setae (sometimes absent) at one-
third from base on interval 2, another slightly larger and a little further back on
6, and an indefinite patch on the declivity— these markings usually made more
distinct by a covering of yellow powder.
Head with rugose confluent punctures and sparse setae ; frons much narrower
than the base of the rostrum (3:5), with an indistinct shallow median fovea.
Rostrum a little longer than the pronotum (8:7), very slightly narrowed from
base to beyond middle, moderately dilated at apex, regularly curved, with a
shallow depression between the base and the head ; dorsum with a fine straight
median carina and a lower more sinuous one on each side, the interspaces with
irregular shallow confluent punctures, and a shallow sulcus above the scrobes.
A ntennae piceous ; funicle with the two basal joints equal, 3-7 equal and moniliform,
7 not wider than 6. Prothorax a little broader than long, almost parallel-sided
in the basal fourth, then narrowing rather rapidly with a gentle curve to the
apex, which is only shallowly constricted, the apical margin gently arcuate
dorsally, with feeble postocular lobes, the gular margin distinctly sinuate ;
dorsum slightly convex longitudinally, fairly closely set with small rounded
granules of unequal size and with an abbreviated narrow median carina.
Scutellum almost round, prominent, very convex, obsoletely punctate, with a
few fine setae. Elytra much wider than the prothorax (16 : n), parallel to beyond
middle, with obtuse posterior calli, the apices each with a short sharp process ;
the shallow striae with large unequal punctures that are somewhat irregular on
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 403
the disk ; the intervals not wider than the punctures, sinuous, uneven, with
irregular small low granules that often form small groups, intervals 3 and 5
partly higher than the others, 3 having a short granulate costa near the base.
Legs piceous, rugosely punctate throughout, with sparse narrow curved whitish
scales and fine recumbent yellow setae ; all the tibiae incurved at apex and
distinctly bisinuate on the lower edge.
Length 8.0 -10.5 mm., breadth 3-4 mm.
N.E. Burma : Kambaiti, 7000 ft., io^1 109, v-vi. 1934.
Dyscerus roratus sp. n.
$. Derm dull black to red-brown, unevenly mottled with groups of small
yellowish setiform scales.
Head with coarse shallow confluent punctures ; frons two-thirds the width
of the base of the rostrum, without any median fova. Rostrum longer than the
pronotum (11 : 8), almost parallel-sided from the base to the scrobes and rather
strongly widened apically, regularly curved, with a very shallow depression
between the base and the frons ; dorsum very finely punctate on the disk, rather
more strongly at the sides, with three narrow impunctate lines, the median one
becoming subcarinate at the base with a shallow depression on each side of it, the
lateral areas with a single shallow sulcus. Antennae piceous ; funicle with joint 1
distinctly longer than 2, 3-7 moniliform, 7 very slightly wider than 6. Prothorax
very nearly as long as broad, parallel-sided in the basal third or half, then narrow-
ing with a slight curve to the apex, which is very shallowly constricted, the
apical margin gently arcuate, the gular margin sinuate, without postocular
lobes ; dorsum flat longitudinally, unevenly set with low shiny granules, the
interstices being clothed with setiform scales, and with a low narrow sinuous
median carina that nearly reaches both base and apex. Scutellum ovate, flat,
impunctate, bare. Elytra wider than the base of the prothorax (3 : 2), parallel
for two-thirds of their length, the posterior calli much reduced and inconspicuous,
the apices jointly rounded and without any sharp processes ; the rows of large
subquadrate punctures mostly rather irregular, the narrow sinuous intervals all
of the same height and bearing a row of flattened granules, these being smaller,
more convex and more numerous near the base. Legs red-brown, with the knees
black, and with numerous yellow setae ; femora closely punctate except on the
thickened part ; anterior pairs of tibiae slightly incurved apically, and shallowly
bisinuate internally, the hind pair almost straight.
Length 10-11 mm., breadth 3.5-4.1 mm.
N.E. Burma : Kambaiti, 7000 ft., 2$, vi. 1934.
Dyscerus bigutticollis sp. n.
(J$. Derm rather dull black, with small spots formed of yellowish scales ;
pronotum with a spot on each side behind the middle ; elytra with three spots on
interval 2 (at one-third from base, behind middle and at top of declivity), a
curved oblique macular band before middle from stria 10 to stria 5 or 3, and
additional spots on the lateral intervals behind middle.
Head globular, bare, with small dense punctures ; frons more than half as
wide as the base of the rostrum (4:7), with a deep median fovea. Rostrum a
little shorter than the pronotum, parallel-sided from the base to the scrobes,
moderately dilated at apex, regularly curved, with a very shallow depression
between the base and the frons ; dorsum with small dense confluent punctures
and three low narrow carinae that unite at the base, and with two shallow
404
SIR GUY A. K. MARSHALL
punctate sulci laterally. Antennae black ; funicle with the two basal joints
equal, 3-7 moniliform, subequal, 7 not broader than 6. Prothorax as long as
broad, slightly rounded laterally, widest at or behind middle, deeply constricted
near apex, the apical margin feebly arcuate dorsally, the gular margin sinuate,
without postocular lobes ; dorsum almost flat longitudinally, with large confluent
punctures behind the subapical constriction and much smaller ones in front, the
narrow intervals unevenly raised and a short broad flattened median carina
about middle. Scutelkim subtriangular, bare, shiny, convex. Elytra wider than
the base of the prothorax (16 : 11), parallel for two-thirds of their length, with
large obtuse posterior calli, the apices separately rounded ; the rows of large
foveae often irregular and variable in shape, the intervals sinuous and as narrow
as the septa between the foveae, intervals 3 and 5 being partly higher than the
others. Legs slender, black, with very sparse fine whitish setae ; femora finely
punctate on the basal half, almost impunctate on the dilated part, and rugose at
the apex ; all the tibiae incurved at the apex and distinctly bisinuate on the
lower edge.
Length 9.5-11.5 mm., breadth 3.5-4.5 mm.
N.E. Burma : Kambaiti, 7000 ft., 8^ 29, iv-vi. 1934.
Dyscerus cognatus sp. n.
$$. Derm rather shiny black, with markings formed of pale yellow setiform
scales ; pronotum with a few sparse scales, without spots ; elytra with a deeply
bisinuate narrow ill-defined macular band at about one-third from base, a trans-
verse one across the top of the declivity as far as interval 5, and some indefinite
spots on the apical area".
Closely allied to D. bigutticollis sp. n., the description of which applies to it
except as regards the two prothoracic spots and the following characters : —
Head with the irons half as wide as the base of the rostrum. Rostrum slightly
widening at the immediate base ; the three fine dorsal carinae with two fairly
regular rows of much larger punctures between them, usually with minute
punctures among them. Prothorax parallel-sided in the basal half, then narrowing
with a slight curve to the broad shallow apical constriction ; the dorsal punctures
as large or larger but with their margins narrower and more raised, the median
carina narrower and extending from the middle almost to the apex. Legs with
the tibiae less incurved at the apex and much more shallowly bisinuate on the
lower edge.
Length 10-0-11-5 mm., breadth 3-7-4-4 mm.
N.E. Burma : Kambaiti, 7000 ft., 6<J i<j>, vi-viii. 1934.
Dyscerus caligatus sp. n.
(J$. Derm dull black, almost bare, with only very sparse short pale setae
and a few small spots formed by the aggregation of a few narrow pale scales,
being the remains of the typical pattern in this genus : prothorax with a small
sublateral spot on each side a little behind the middle and another in the middle
of the base ; elytra with a very small spot at one-third from the base on intervals
2 and 10, a larger one a little behind them on 6, small ones at the beginning of the
declivity on 2, 7 and 9, and a large one near the apex.
Head with rugose confluent punctures ; frons two-thirds as wide as the base
of the rostrum, with a shallow median fovea (obsolete in <$). Rostrum as long as
the pronotum in <J, a little longer in 9, parallel-sided from the base to the scrobes
and rather broadly widened at the apex, almost porrect at the base and curving
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 405
strongly downwards apically, the base separated from the frons ; dorsum closely
and coarsely punctate in <J without definite carinae, the punctures smaller in $
and with traces of fine narrow carinae, there being only one shallow lateral sulcus.
Antennae red-brown, comparatively stout ; funicle with joint 1 longer than 2,
3-7 moniliform, 5-7 shorter and transverse, 7 rather broader than 6. Prothorax
very nearly as long as broad, strongly rounded laterally, widest at a little behind
the middle, shallowly constricted at apex, the apical margin subtruncate dorsally,
the gular margin deeply sinuate, the postocular lobes feeble ; dorsum fairly
closely set with rugose uneven opaque granules and with a short narrow median
carina on the anterior half and a smooth irregular tubercle on each side of it.
Scutellnm shield-shaped, bare, opaque, with a shallow median impression. Elytra
at the shoulders only slightly wider than the broadest part of the prothorax,
subparallel to beyond the middle, with low obtuse posterior calli, the apices
separately rounded ; the large round punctures in fairly regular rows, the spaces
between them being mostly much greater than their diameters ; the intervals not
wider than the punctures, rather irregular, each with a row of very small shiny
granules that are more numerous on intervals 3 and 5, which are slightly higher
than the others. Legs rather stout, with sparse pale setae, red, with the knees,
front tibiae (except partly in the middle), and the tarsi (except joint 4), blackish ;
tibiae incurved at apex and distinctly bisinuate below, the front pair with a very
small mucro.
Length 5-7 mm., breadth 1-9-2-4 mm.
N.E Burma : Kambaiti, 7000 ft., 1^2^, v. 1934.
Dyscerus latipes sp. n.
$. Derm opaque, piceous, the apical collar of the prothorax red-brown, with
sparse yellowish setiform scales, some collected together in small groups, but the
only conspicuous spots being a round one on interval 6 of the elytra at a little
before the middle and a much larger oblong one behind the posteiior callus.
Head finely shagreened with some very minute sparse shiny granules ; frons
comparatively broad, five-sixths the width of the base of the rostrum, without
any media fovea. Rostrum a little longer than the pronotum, narrowing very
slightly from the base to the scrobes and widening at the apex, moderately
curved and forming an angle with the frons at the base ; dorsum with rather
large shallow confluent punctures, a narrow median carina near the base only,
and a distinct lateral sulcus. Antennae red-brown ; funicle with joint 1 longer
than 2, 3-7 moniliform and subequal, 7 slightly broader than 6. Prothorax
somewhat transverse (5:6), moderately rounded laterally, widest at about the
middle, shallowly constricted at the apex, the apical margin truncate dorsally.
the gular margin sinuate, the postocular lobes obsolescent ; dorsum closely set
with small shagreened granules of unequal height and a short narrow median
carina, the apical area sculptured like the head. Scutellum shield-shaped, some-
what raised, bare, uneven. Elytra at the shoulders broader than the prothorax
(4:3), parallel for two-thirds of their length, with prominent obtuse posterior
calli, the apices separately produced into a short sharp process ; dorsum somewhat
flattened on the disk, as far as interval 5 with regular rows of deep round
separated punctures ; the intervals rather uneven, 3, 5 and 7 a little higher than
the others and with numerous low shiny granules. Legs red-brown, with sparse
pale setae ; tibiae strongly compressed and laminately dilated, especially on the
406 SIR GUY A. K. MARSHALL
basal half, being there about as wade as the broadest part of the femora, the
dorsal margin highest near the base, rapidly narrowing and incurved at the apex,
the lower edge deeply bisinuate.
Length 5-5 mm., breadth 2 mm.
N.E. Burma : Kambaiti, 7000 ft., 1$, vi. 1934.
A very striking species on account of its ' small size and very abnormal
tibiae.
Dyscerus clathratus Pasc. — Kambaiti, 2^ 1$, iv-v. 1934.
Key to Burmese Dyscerus.
1 (22). Tibiae normal.
2 (3). Head elongate, rostrum without any lateral sulcus, apical margin
of presternum not sinuate ..... malaisei sp. n.
3 (2). Head globular, rostrum with a lateral sulcus, apical margin of
presternum sinuate ....
4 (5). Antennal club a little longer than the funicle . . longiclavis Mshl.
5 (4). Antennal club much shorter than the funicle.
6 (21). Elytra at the shoulders much wider than the broadest part of the
prothorax.
7 (8). Front tibiae with a distinct triangular mucro ; rows of punctures
on the elytra quite regular throughout . . . posticus sp. n.
8 (7). Front tibiae not mucronate.
9 (16) . Elytra each with a short sharp apical process.
10 (11). Elytra with perfectly regular rows of small punctures ; pronotum
rugosely punctate ...... ordinatus sp. n.
11 (10). Elytra with large, more or less irregular punctures ; pronotum
granulate.
12 (13). Funicle with joint 1 longer than 2 ; prothorax widest at middle,
not constricted at apex ..... clathratus Pasc.
13 (12). Funicle with joints 1 and 2 equal ; prothorax widest behind
middle, constricted at apex.
14 (15). Prothorax with coarse granular elevations and a broad rugosely-
punctate median carina, deeply constricted at apex ; elytral
spots formed of narrow scales .... direptus sp. n.
15 (14). Prothorax with small close shiny granules and a narrow
impunctate carina, shallowly constricted at apex ; elytra with a
general clothing of sparse yellow setae . . asper sp. n.
16 (9). Elytra without apical processes.
17 (18). Frons without any median fovea ; funicle with joint 1 longer than
2 ; elytra with moderately large punctures and a uniform
mottling of scales ; legs red-brown with numerous yellow
setae ........ roratus sp. n.
(17). Frons with a deep fovea ; funicle with the two basal joints equal ;
elytra with very large foveae and a few scale spots forming a
18 pattern ; legs black with very sparse setae.
19 (20). Rostrum dorsally with fine confluent punctation ; prothorax
deeply constricted at apex and with a spot of yellowish scales
on each side behind the middle . . . bigutticollis sp. n.
20 (19). Rostrum with a row of larger separated punctures on each side of
the median carina ; prothorax shallowly constricted at apex
and without any spots ..... cognatus sp. n.
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 407
21 (6). Elytra at the shoulders only slightly wider than the broadest part
of the prothorax ; legs red, with the knees, front tibiae and
tarsi blackish ; length 5-7 mm. .... caligatus sp. n.
22 (1). All the tibiae dilated and sublaminate, especially the hind pair ;
length 5-5 mm. ....... latipes sp. n.
In addition to the foregoing species Faust has also recorded from Burma
(Ann. Mus. Stor. nat. Genova, 60, 1895 : 230) D. notatus Pasc. (Java and Borneo)
and D. bispinulus Desbr., but in both cases the identification is open to doubt.
Moreover, Dr. Malaise also took single specimens of four other species of Dyscerus,
all found at Kambaiti, in May, 1934.
Kobuso binodosus sp. n.
cj§. Derm black, thinly clothed with brownish setiform scales.
Head with coarse shallow subconfluent punctation ; frons strongly convex
transversely, with a small median fovea. Rostrum of $ nearly straight, as long as
the pronotum, gradually widening at apex, with four dorsal rows of coarse,
longitudinally confluent punctures and a deep sulcus above the scrobes from base
to beyond middle ; rostrum of $ slightly longer, with the punctures much smaller
and not confluent. Antennae with joint 2 of the funicle a little longer than broad,
nearly as long as 1. Prothorax as long as broad, in <$• parallel-sided in the basal
third, then narrowing to apex, with a very shallow subapical constriction ; in $,
widest at base, gradually narrowing anteriorly, without any constriction ;
bisinuate at base, the dorsal apical margin arcuate ; dorsum slightly convex
longitudinally, with numerous small convex non-contiguous granules and a fine
median carina ; the granules somewhat larger and fewer in $. Scutellum semi-
circular, raised, smooth in $, with fine rugulose punctures in $. Elytra widest at
the slightly prominent shoulders, gradually narrowing behind, jointly rounded at
apex, the posterior calli forming a sharp conical tubercle in g, less developed in
$ ; the striae with large subquadrate punctures, those in striae 3-6 slightly
irregular ; the septa between the striae as broad as or broader than the intervals
and sometimes uniting laterally to form low transverse wrinkles ; the intervals
narrow, more or less undulate, with small sparse flattened granules, interval 3
slightly more raised in $. Legs shallowly rugulose, with sparse pale setae ; joint
2 of hind tarsi forming an equilateral triangle.
Length 10-11 mm., breadth 4-5 mm.
N.E. Burma : Kambaiti, 7000 ft., itf (type), 20. vi. 1934, i$, 4-8. vi. 1934.
The only known Indian species, K. crassus Mshl. 1936, is a much broader
insect ; the frons is less convex ; the antennae have joint 2 of the funicle transverse
and much shorter than 1 ; the elytra have the discal punctures quite regular, the
tubercle near the base of interval 3 is much lower, and the subapical calli are
normal and obtuse ; joint 2 of the hind tarsi is transverse and broadly truncate at
the base.
Niphades malaisei sp. n
<$$. Derm dull black ; pronotum with a narrow median line of grey scales ;
elytra with the whole of the posterior declivity and a very indefinite large patch
round each shoulder covered with dense brownish grey scales.
Head with dense coarse punctures and sparse brown setae ; frons a little
broader than the base of the rostrum, without any median puncture. Rostrum
shorter than the pronotum, only slightly curved, gradually dilated at apex, with
more or less confluent coarse punctures that are not in regular rows ; that of $ a
little longer and more shiny. Antennae with joint I of the funicle longer than 2,
408 SIR GUY A. K. MARSHALL
3-7 transverse. Prothorax as long as broad, moderately rounded laterally, widest
near or behind middle, not constricted at apex, feebly bisinuate at base, the apex
gently arcuate ; dorsum closely set throughout with large granules and with a fine
abbreviated median carina. Elytra broadly suboblong, parallel for three-fourths
of their length, nearly twice as broad at the roundly rectangular shoulders as the
base of the prothorax, shallowly trisinuate at the base, jointly rounded at the
apex ; dorsum moderately convex longitudinally, the shallow striae with punctures
that are comparatively small and regular near the suture and larger and more
irregular laterally, but all greatly diminishing apically ; interval 3 with five
rather large tubercles that are variable in shape and not always symmetrically
placed, and 5 with five or six smaller tubercles, the last being the largest and
forming the subapical callus ; all the tubercles with an apical tuft of very short
stout setae. Legs black, with fairly dense brown scaling and short suberect setae,
the femora with a broad pale ring ; corbels of hind tibiae only slightly longer than
the apical width of the tibia.
Length 8-9 mm., breadth 3-5-4-1 mm.
N.E. Burma : Kambaiti, 7000 ft., i<^3$, vi. 1934.
Nearly allied to N. granicollis Fst., which differs in having the rostrum much
more broadly dilated at the apex, with the punctures in regular rows ; antennae
with joint 2 of the funicle equal to 1 ; prothorax without any trace of a median
carina, with a longitudinal strip on each side devoid of granules, and the apical
margin strongly arcuate and overhanging the head ; the corbels of the hind tibiae
nearly twice as long as the apical width of the tibia.
Niphades debilis sp. n.
9- Allied to N. malaisei, but much smaller and with the tubercles and
colour pattern much less pronounced, the shoulder patches on the elytra being
very faint. The description of malaisei applies to it except in the following
particulars : —
Rostrum much more slender, not dilated at apex, the irregular punctures
finer. Elytra ovate, parallel for only half their length, much more narrowed
behind, the punctures in the lateral rows similar to those near the suture, the
tubercles on intervals 3 and 5 much smaller and flatter, 5 without any apical
tubercle, the preapical callus being almost obsolete ; all the setae (including those
on the pronotum) short and erect, whereas in malaisei they are all recumbent,
except those on the tubercles on the elytra.
Length 6-5 -7-0 mm., breadth 2-5-2-7 mm.
N.E. Burma : Kambaiti, 7000 ft., 2$, vi. 1934.
Niphades alni Mshl. — Kambaiti, 2$, v-vi.
AMPHIALODES gen. n.
Head with the frons at its narrowest much wider than the base of the rostrum ;
eyes broadly ovate, not more convex than the head, rather coarsely faceted,
of a shining silvery colour. Rostrum comparatively narrow, elongate ; scrobes
passing rapidly beneath the rostrum at about the middle ; mandibles with very
short blunt teeth. Antennae with the scape straight, slender and abruptly
clavate, not reaching the eye ; funicle a little shorter than the scape, joint 1 much
broader than 2, 7 transverse. Prothorax as long as broad, truncate at the base,
the apical margin arcuate dorsally, the postocular lobes moderate ; presternum
excavated in front of the coxae, the gular margin deeply sinuate. Scutellum
BURMA AND BRITISH INDIA — COLEOPTERA ! CURCULIONIDAE 409
concealed. Elytra convex, broadly ovate, more or less acuminate behind, jointly
truncate at base, without any humeral calli. Legs long, slender ; femora moder-
ately clavate, with a small sharp tooth ; tibiae straight, with a sharp uncus and
no mucro ; claws free, simple. Sternum with the front coxae as widely separated
as the middle pair, the prosternal process smooth and bare ; metasternum as long
as a middle coxa, the metepisternal suture entirely obliterated. Venter with the
intercoxal process about as wide as a hind coxa, which is transversely elongate ;
ventrite 2 much longer than 3 and 4.
Genotype: Amphialodes acuminatum, sp. n.
The only near ally of this genus is Amphialus Pasc, which at present is
known only from Ceylon. The latter differs in the following characters : the head
has the frons slightly narrower than the base of the rostrum, and the eyes are
finely faceted and black ; the prothorax is transverse ; scutellum distinct ; front
tibiae more or less bisinuate on the lower edge and the front coxae contiguous ;
metasternum much shorter than a middle coxa ; and ventrite 2 is not longer
than 3 and 4.
Amphialodes acuminatus sp. n.
<J$. Derm chocolate-brown, opaque with very sparse pale yellowish scales ;
prothorax with three very indefinite stripes of sparse scales ; elytra with an
irregular band of scales across the declivity and sparse groups of scales elsewhere.
Head with separated coarse punctures and sparse brown scales ; frons with a
shallow median impression. Rostrum of $ rather strongly curved, a little shorter
than the pronotum, parallel-sided and subcylindrical from base to antennae and
there irregularly punctate and with subrecumbent tranverse scales, the apical area
wider, bare, smooth and with very shallow punctures; rostrum of $ slightly longer,
more gradually widened at the apex, the strong punctures confined to the basal
half, the scales fewer and narrower. Antennae with joint 1 of the funicle about as
long as 2 but more than twice as wide, 2-7 widening distally, 3 as long as broad,
the rest more or less transverse. Prothorax moderately rounded laterally, widest
at middle, narrower at apex than at base but not constricted ; dorsum feebly
convex longitudinally, with strong separated punctures leaving an indefinite
impunctate median line on the basal two-thirds ; two small groups of erect brown
scales on the apical margin and a few similar scales on the disk. Elytra broadly
ovate (4 : 3), widest before the middle, rapidly acuminate behind and jointly
rounded at apex ; dorsal outline rising steeply from the base, then almost flat to
beyond middle, and less steeply declivous behind ; the impressed striae containing
rather large deep punctures which become smaller behind ; intervals broader than
the striae, somewhat convex, smooth and impunctate, interval 3 with an obtuse
elevation near the base and a larger one at the top of the declivity, 5 with two
much smaller elevations, one near the base and one at about the middle, all these
bearing suberect brown scales, and between the two tubercle on 3 are one or two
small tufts of similar scales. Legs with sparse recumbent narrow pale scales.
Length 4-0-4-5 mm., breadth 2-0-2-2 mm.
N.E. Burma : Kambaiti, 6600 ft., i£ 1$, vi. 1934.
Amphialodes graniger sp. n.
(J. Nearly allied to, though larger than, the preceding species, the description
of which applies to it except in the following particulars : —
The sparse scaling more generally distributed, without any definite band
across the declivity of the elytra.
410 SIR GUY A. K. MARSHALL
Rostrum more rugosely punctate. Antennae with joint I narrower and not
so globular. Prothorax more closely and somewhat rugosely punctate, with a low
smooth median carina on the apical half only. Elytra more narrowly ovate and
much less acuminate behind, the basal margin raised into an obtuse smooth
costa ; the dorsal outline rising much less steeply at the base and with the
apical declivity steeper ; the punctures much larger, as wide as the intervals,
and between the punctures a small shiny granule ; interval 3 with two similar
tubercles, but none on 5, without the prominent tufts of erect scales.
Length 5 mm., breadth 2-2 mm.
N.E. Burma : Kambaiti, 6600 ft., 1^. vi. 1934.
Euthycus sp. — Kambaiti, <J, v.
Euthycus sp. — Kambaiti, 1$, vi.
Genus ?. — Kambaiti, 1 ex., iv.
Genus ?. — Kambaiti, 1 ex., vi.
Subfamily Pissodin^
Carcilia granicollis sp. n.
$. Derm black to piceous, dull, thinly clothed with minute setae.
Head with fine dense confluent punctures. Rostrum rapidly narrowing from
the base to the scrobes, then gradually widening again to the apex, shiny, densely
punctate at the base, the punctures becoming much sparser anteriorly. Antennae
inserted behind the middle of the rostrum ($), red-brown, the club black ; funicle
with joint 1 a little longer than 2. Prothorax a little broader than long (9 : 11),
rounded laterally, widest behind the middle, not constricted at the apex, sub-
truncate at the base ; dorsum somewhat convex longitudinally, with close
reticulate punctures, the narrow intervals bearing small low granules, both
punctures and granules becoming larger laterally. Elytra only a little wider at the
roundly rectangular shoulders than the prothorax, very gradually widening to
beyond the middle, and broadly rounded behind ; the striae containing rather
large deep punctures varying in size and becoming much smaller near the apex,
where the striae are very deep ; the intervals flat varying in width, transversely
rugulose and becoming definitely granulate towards the base. Legs short, stout,
coarsely and rugosely punctate, black, with the tarsi red-brown ; middle tibiae
with the upper end of the corbel produced into a sharp triangular tooth. Underside
rather closely punctate and with bifid to quadrifid setae on the metasternum and
first ventrite ; ventrite 5 with the apical half shiny, almost impunctate, nearly
bare and depressed on each side.
Length 7-10 mm., breadth 2-7-4-0 mm.
N.E. Burma : Sadon, 4000 ft., i$, vi-vii. 1934 (Dr. Malaise). Assam : Naga
Hills, 1$ (Doherty type).
Type in the British Museum.
Closely allied to the Japanese genotype, C. strigicollis Roel., in which
however the pronotum has an obtuse elevation in the middle near the base and a
broad shallow constriction at the apex ; the striae on the elytra are narrower with
much smaller shallower punctures, and the intervals proportionately broader ;
the apical part of ventrite 5 in the female has a group of long erect setae in the
middle.
Subfamily Erirrhininae.
Echinocnemus adustus Chev. — Tenasserim, Mekane, 90 km. E. of Moulmein,
200 m., 7 $ 3$, xi. ; Siam, Medaw, Burma frontier, i<3\ xi.
BURMA AND BRITISH INDIA — COLEOPTERA ! CURCULIONIDAE 4II
Echinocnemus roelo/si Fst. — Moulmein, i$, xi.
Bagous sp. — S. Shan States, S. end of Inle Lake, 900 m., i^J, ix.
Bagous sp. — Washaung, ca. 200 m., 20 km. E. of Myitkyina, 1$, vii.
Smicronyx sp. — Myitkyina, 175 m., i^J, vii.
Subfamily Eugnominae.
Ixalma asperula Fst. — Kambaiti, 2$, v-vi.
Ixalma 3 spp. — Kambaiti, 3 specimens, v-vi.
Subfamily Nanophyinae.
Nanophyes nigritulus Boh. — Myitkyina, 175 m., 2$, iii.
Nanophyes sp. — Washaung, 20 m. E. of Myitkyina, 1$, vii.
Subfamily Apioninae.
Apion 2 spp. — Kambaiti, 2$, iv-vi.
Apion sp. — Punkaung, between Sadon and Myitkyina, i<$, vii.
Apion sp. — S. Shan States, Inle Lake, irf, ix.
Subfamily Attelabinae.
Euscelophilus burmanus sp. n.
<£$. Derm black (except the inflexed margins of the elytra which are dark
blue), not very shiny ; head and pronotum with sparse golden pubescence ;
elytra almost bare on the basal half, with a small indefinite golden spot on the
suture at one-fourth from the base, the apical half with a large common irregular
X-shaped mark of grey pubescence (silvery in some lights) ; underside with
rather sparse grey pubescence, which is denser in the middle of the metasternum,
the sides of which bear golden pubescence.
Head only slightly longer than its basal width, finely rugosely punctate, with
a small elevation in the middle bearing a narrow median carina and a shallow
median stria behind it, and a large depression behind each eye ; frons as broad as
the base of the rostrum, flat, with fan-like carinulae ; eyes moderately prominent.
Rostrum longer than broad, coarsely punctate, with the antennae inserted close
to the base. Antennae black ; the scape equal to joint 2 of the funicle, which is
longer than 1, the others progressively diminishing distally, 4 and 5 as long as
broad, 6 and 7 transverse, 7 the broadest. Prothorax transverse (8 : 11), moderately
rounded laterally, widest near the base, with a broad shallow constriction at base
and apex ; dorsum having in the middle of the disk a very large abruptly raised,
punctate tubercle divided longitudinally by a broad impunctate sulcus ; the areas
on each side of this tubercle with fine dense punctures and two shallow impressions,
the apical area more sparsely punctate, and the basal area transversely carinulate.
Scutellum transverse, trapezoidal, broadest at the base, rugose. Elytra sub-
quadrate, rather longer than their width at the obtusely projecting shoulders
(6:5), parallel from behind the shoulders nearly to the apex, the posterior
declivity vertical and appearing broadly subtruncate from above, the actual
apices separately rounded ; dorsum almost flat as far as interval 5, with a small
rounded shiny tubercle on each side of the scutellum ami irregular rows of rather
large shallow punctures, the intervals narrow and sinuous, 3 and 5 being slightly
more raised and 9 subcarinate. Legs short and stout, black, with sparse golden
setae ; femora with a small sharp blue tooth and also a small tooth on each side
412 SIR GUY A. K. MARSHALL
of the tibial socket ; front tibiae with two rows of denticles on the lower edge.
Pygidium smooth, with fine sparse punctures, erect pale setae and a median stria.
Length 4 mm., breadth 2-2 mm.
N.E. Burma : Kambaiti, 7000 ft., 3$ 1$, iv-v. 1934.
Judging by the description, E. gibbicollis Schilsky, from Peking, must be
nearly allied to the present species. But the former differs in having the antennae,
tibiae and tarsi reddish ; the rostrum is very short, broader than long ; the large
tubercle on the pronotum is not divided by a sulcus ; and the elytra have before
the apex three elevations, of which the middle one is the largest.
Trachelolabus burmanensis Voss. — Kambaiti, 7000 ft., io<J 9$, iv-vi.
Paramecolabus feae Fst. — S. Shan States, Taunggyi, 5000 ft., 1$, viii-ix.
Paramecolabus sp. — Kambaiti, 1 $, iv.
Euops championi Voss. — Kambaiti, 1$, hi.
Euops peguensis Voss. — Kambaiti, i(J, v.
Euops 2 spp.— Kambaiti, i<$ 1$, iv.
Euops nigricollis sp. n.
rj$. Derm shining blue or greenish blue above and below, except the disk of
the pronotum which is black.
Head with sparse irregular punctures, the eyes very narrowly separated,
longer than the temples and not exceeding them laterally. Rostrum almost
impunctate dorsally. Prothorax transverse, rounded laterally, widest at the base,
without any basal or apical constriction ; dorsum smooth and shiny, with sparse
minute punctures, the lateral punctures much coarser, the basal depression
punctate and transversely wrinkled. Scutellum transverse, suboblong, with the
hind margin gently arcuate, quite smooth and impunctate. Elytra subquadrate,
a little longer than broad, parallel from the roundly rectangular shoulders to well
behind the middle, broadly rounded behind ; the striae not impressed at the base,
fine behind the middle, broad and deep at the apex, the punctures in them
moderately strong and diminishing behind, mostly separated by their own
diameters or more in $, closer in $ ; the intervals broader than the striae, flat,
with a single row of fine punctures. Legs without a femoral tooth ; front tibiae
of {J simply curved, those of $ bisinuate on the lower edge.
Length 3-0-3-2 mm., breadth 1-7-1-8 mm.
N.E. Burma : Kambaiti, 7000 ft., i<J 1?, iv. 1934.
Closely resembles in general facies the Japanese phaedonius Sharp, which,
however, has the pronotum much more strongly punctate and with a transverse
impression ; the scutellum is quadrate ; the front tibiae are more slender, and less
deeply bisinuate in the female.
Hoplapoderus bistrispinosus Fst. — S. Shan States, Taunggyi, 1500 m., 1$, viii-ix.
Hoplapoderus hystrix F., var. echinatus, Gyll. — 40 km. E. of Taunggyi, i^J, ix-x ;
Sadon, 1200 m., i(J, vi-vii.
H. hystrix var. orientalis Voss. — Sadon, 2$ 2$, vi-vii.
H. hystrix var. caliginosus Fst. — Taunggyi, 2(j3?, viii-ix ; 40 km. E. of Taunggyi,
ig, ix-x.
Hoplapoderus breviceps Voss. — Kambaiti, 2000 m., 2^ 3$, v-vi.
Hoplapoderus gemmosus Jek. — Taunggyi, 1$, viii-ix ; Inle Lake, 900 m., 5^ 19,
ix (irf entirely black) ; Moulmein, 1$, xi.
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 413
Paroplapoderus malaisei sp. n.
$$. Derm black, with orange-yellow markings ; head and the basal half of
the rostrum black above, the remainder orange ; prothorax with a large irregular
lateral orange spot, a small triangular one in the middle of the base (sometimes
transverse), and a variable spot on the front margin of the pleurae (sometimes
absent) ; scutellum black ; elytra orange, with the following black markings : a
large patch covering the whole shoulder (but not reaching the lateral margin) and
curving inwards at one-fourth from the base as far as interval 3, a large oblong
lateral patch at about the middle between interval 5 and the margin, a transverse
spot immediately behind this, a broad sublunate band close to the apical margin,
a sutural spot at the base, and a spot covering the large median tubercle ; under-
side rarely entirely black, usually with an orange spot on the metasternum just
behind the middle coxae, the lateral margin of the venter broadly orange enclosing
a longitudinal row of three round black spots, the basal one large, the other two
smaller ; pygidium orange with two large black spots uniting at the base.
Head with a deep median sulcus from the frons to the base of the head and
a transverse stria behind the eyes ; frons rugose, with irregular longitudinal
striae and a few coarse punctures ; vertex more shiny, with sparse minute
punctures and faint striolae. Rostrum transverse, with a broad median sulcus on
the basal half, the apical half with strong separated punctures. Antennae stout,
orange ; funicle with joint 1 almost globular, 2 as long as broad but narrower than
any of the others, 3 and 4 longer than broad, 5-7 transverse ; club with the two
basal joints transverse. Prothorax transverse, almost semicircular, apart from the
very short apical collar which is constricted at the base ; dorsum coarsely and
unevenly plicate, with a shallow depression on each side of the disk behind the
middle and a narrow median sulcus which becomes much broader and deeper at
the base. Scutellum broadly sublunate, rugulose, with the apical margin sinuate
in the middle. Elytra with the sides sinuate in the basal half and broadly rounded
behind the middle, the shoulders projecting laterally in a small sharp angle ;
dorsum with somewhat irregular rows of large shallow punctures, the narrow
intervals rugulose on the disk, shiny and finely punctate laterally, each elytron
with an oblique row of three conical black tubercles from the shoulder to the
middle, the one above the shoulder the smallest and bluntest, the one near the
suture sharply pointed and much the largest. Legs yellow, the femora with a
broad brown subapical ring ; tibiae finely denticulate on the lower edge.
Length 5-5-7-0 mm., breadth 3-4 mm.
N.E. Burma : Kambaiti, 7000 ft., 6$ 2$, iv-vi. 1934.
The general colouring is very similar to that of the Chinese vitticeps Jek.,
but that the species may be readily distinguished by the tubercles on the elytra,
the humeral one being almost obsolete, and the large sutural one is broadly
rounded instead of sharply conical. Structurally malaisei is nearer to the Indian
lefroyi Mshl., which, however, has very different colouring, the elytra being
brown on the basal half and yellow apically, and the head has only a fine median
stria.
Tomapoderus cyclops Fst. — S. Shan States, 40 km. E. of Taunggyi, i<3\ ix-x.
Apoderus opacus sp. n.
(£$. Derm dull black, bare above ; mesepimera and metepisterna with sparse
pale setae, forming a dense whitish spot at both ends of the former and at the
apical end only of the latter.
novit. zool., 42, 3. 1948. 81
414 SIR GUY A. K. MARSHALL
Head of $ longer than the pronotum, subcorneal, deeply constricted at the
base, with the sides nearly straight, transversely wrinkled on the vertex with a
distinct median stria, the frons wider than the length of an eye, coarsely striolate
longitudinally, the eyes prominent ; head of 9 broader, more rounded laterally,
the vertex shiny with only obsolescent wrinkles and the median stria disappearing
in front, the eyes less prominent. Rostrum stout, as long as broad, with a deep
stria between the antennae. Antennae with joint 2 of the funicle shorter than 1
and longer than 3, 3-5 subequal and as long as broad, 6 and 7 transverse ; joint
2 of the club as long as broad. Prothorax subconical, a little broader than long,
moderately rounded laterally, the apical collar not constricted at its base, its
anterior margin shallowly sinuate, the dorsum rugulose. Scutellum trapezoidal,
transverse. Elytra very nearly as broad as long, parallel for three-fourths of their
length, the striae perceptible only with difficulty (except near the apex) owing to
the dense uniform rugulose punctation. Legs with a long sharp femoral tooth ;
tibiae with a strong dorsal carina between two sulci, the front pair curved inwards
at the apex in <J, straight in §.
Length 4-5 mm., breadth 2 mm.
N.E. Burma : Sadon, 4000 ft., i$, vi-vii. 1934 (type) ; Kambaiti, 7000 ft.,
1$, iv. 1934.
In Voss's key (Stettin, ent. Ztg. 88, 1927 : 2) this species runs down to the
Sumatran corporaali Voss ; the latter differs however in having no median stria on
the head ; the width of the frons is less than the length of an eye ; joint 2 of the
funicle is longer than 1 ; and the apical collar of the prothorax is deeply constricted.
Apoderus discalis sp. n.
(J$. Body testaceous, bare ; head and rostrum red-brown, turning to blackish
beneath ; prothorax with a broad black lateral stripe ; elytra dark red-brown,
turning to black at the base, with a large common testaceous band extending from
near the base to the middle and laterally as far as stria 5, its posterior margin
having a deep indentation on interval 3.
Head of $ longer than the pronotum (4 : 3), subconical, with the sides straight;
frons much wider than the length of an eye, impunctate, with two short divergent
striae ; vertex smooth, shiny, with only a faint indication of a median stria ; eyes
moderately convex ; head of $, a little shorter and slightly more convex. Rostrum
slightly longer than broad, almost impunctate above, with a broad shallow sulcus
between the antennae. Antennae testaceous ; joint 1 of the funicle bulbous, 4
usually slightly longer than any other, 7 slightly transverse ; club with joint 2
longer than broad. Prothorax about as long as broad, subconical, with the sides
straight, smooth and shiny above, with only a faint trace of a median stria.
Scutellum transverse, trapezoidal, with two small indentations in the middle of
the hind margin. Elytra rather longer than broad (6:5), shallowly sinuate
behind the shoulders, widest behind the middle, with a broad depression covering
the same area as the pale marking on the basal half and a small shiny black
tubercle on each side of the scutellum ; the rows of punctures moderately strong
on the basal half and becoming extremely fine posteriorly ; interval 5 costate
basally. Legs testaceous, with the apical half of the hind femora black.
Metasternum of $ with a deep median sulcus having a small sharp overhanging
granule on each side of it, the sulcus continued more narrowly between the
middle coxae.
Length 5-5-6-0 mm., breadth 2-3-2-5 mm.
N.E. Burma : Kambaiti, 7000 ft., 2<3I2$, iv-v. 1934.
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 415
Nearly allied to carbonicolor Mot. and praecellens Sharp, which have the
general body colour and the four anterior femora black ; the head is shorter and
more convex ; the elytra lack the tubercles on each side of the scutellum ; and on
the male metasternum there are two obtuse tubercles between the median coxae
(instead of the sharp granules behind them) .
Sharp {Trans, ent. Soc. Lond. 1889 : 46) states that these two tubercles
are on the mesosternum, which is erroneous. This draws attention to a character
in the Attelabinae that has not previously been noticed. In other Curculionidae,
when the middle coxae are separated, the space between them is occupied partly
by a backward process of the mesosternum and partly by a forward process from
the metasternum. These normally meet in the middle, but sometimes the
mesosternal process occupies the whole area and rarely may even extend behind
the coxae. But in the Attelabinae the intercoxal area is occupied entirely by the
metasternum. There is, however, one exception, namely, the more primitive
Tropical American genus Pilolabus Jekel, which in this respect resembles the
Khynchitinae ; moreover Voss has already pointed out {Stettin, ent. Ztg. 85, 1925 :
19) that in this genus the structure of the tibiae agrees with that found in the
same subfamily.
Sharp {I.e.) describes in praecellens some striking secondary male characters,
but these also occur in carbonicolor, with which he was not acquainted, and I can
find no reliable differences between the species ; praecellens must therefore sink
as a synonym.
Apoderus atronitens sp. n.
cJ9' Derm very shiny black, the underside of the head, the rostrum (except
the blackish mouthparts), the legs and antennae entirely, the sides and apex of
the venter, and the whole pygidium, yellow.
Head quite similar in the two sexes, only slightly longer than the width
across the eyes, which are moderately convex, the whole vertex and frons shiny
and impunctate, with only a faint median stria on the former, the frons much
wider than the length of an eye. Rostrum a little longer than broad, only slightly
narrowed at the base and gradually widening to the apex, with a broad sulcus
between the antennae, the apical part with fine sparse punctures. Antennae with
joint 1 of the funicle longer than any of the others, 2 shorter than 3 and equal to
4, 5-7 transverse ; club with joint 1 a little longer than broad, 2 transverse.
Prolhorax transverse (5 : 7), subcorneal with the sides gently rounded, the shiny
dorsum entirely impunctate, without definite impressions but with a fine shallow
median stria. Scutellum broadly lunate, shiny and impunctate, the apical margin
arcuate. Elytra nearly parallel-sided, with a small sharp angle at the shoulders ;
the discal punctures small and becoming much finer behind, the lateral punctures
much larger, the intervals broad, flat and impunctate. Legs with the tibiae
slender, straight and finely serrate on the lower edge.
Length 5-0-6-5 mm., breadth 2-5-3-5 mm-
N.E Burma : Kambaiti, 7000 ft., 28^ 14$, iv-vi. 1934.
Very closely allied to the genus Phvmatapoderus Voss, the only essential
structural difference being that in that genus all the species have a boss-like
tubercle in the middle of each elytron. Voss placed this genus in the tribe
Hoplapoderini, which is primarily distinguished from the Apoderini by the shape
of the head. On this character Phvmatapoderus certainly belongs to the Apoderini
and cann°t be regarded as more than a subgenus of Apoderus ; the five species at
present known are extremely closely related and might well prove eventually
to be only local races of a single species.
416 SIR GUY A. K. MARSHALL
Apoderus notatus F.— Myitkyina, 175 m., z<$, iii ; S. Shan States, Taunggyi,
1 (J 2$, viii-ix.
Apoderus blandus Fst. — Siam, Medaw, Burma frontier, i<J, xi.
Phymatapoderus elongatipes Voss.— Kambaiti, 4<J 3$, v-vi ; Sadon, 1200 m.,
3<J 2$, vi-vii ; S. Shan States, Taunggyi, 1500 m., 2$ 2$, viii-ix.
Centrocorynus rufulns Voss. — Sadon, 1$, vi-vii ; Taunggyi, 4$ 4$» viii-ix ;
Tenasserim, Sukli, 75 km. E. of Moulmein, 600 m., i$, x.
Centrocorynus scutellaris Gyll.— Kambaiti, i^, v ; Punkaung, between Sadon and
Myitkyina, i<J, vii.
Centrocorynus propinquus Voss. — Punkaung, i<J 1$, vii ; Sadon, 1$, vi-vii.
Cycnotrachelus flavonotatus Voss. — S. Shan States, Pekong, 900 m., 1^, x;
Sulki, 600 m., 75 km. E. of Moulmein, 1$, x.
Cycnotrachelus sp. — Kambaiti, i(J, iv.
Paracycnotrachelus nigrigenibus sp. n.
<$$. Derm red-brown ; head with the extreme base and a large transverse
frontal patch, the scutellum, and the apex of the femora, black ; the mesepimera
and metepimera with the usual pale pubescence.
Head of $ three times as long as the width across the eyes,* the cylindrical
basal part being less than one-third of the total length, faintly striolate trans-
versely on the posterior two-thirds ; frons broader than the length of an eye, the
two deep frontal sulci almost parallel ; eyes moderately convex ; head of § much
shorter and broader, 1-8 times as long as the width across the eyes, subcorneal,
without any distinct cylindrical portion at the base. Rostrum twice as long as
broad, with three sulci on the basal half. Antennae of <$ testaceous ; scape a little
shorter than joint 2 of the funicle, which is equal to or shorter than 3, 4 longer
than 3 ; club with joints 1 and 2 equal, 4 sharply pointed but without any curved
process ; of $, with the scape longer than 2 of the funicle, 2-4 subequal, club with
joint 1 longer than 2. Prothorax as long as broad, conical, with sides straight, the
apical margin only shallowly sinuate and without any subapical constriction, but
with a deep round fovea in the middle of the apical collar ; dorsum shiny and
impunctate, not rising up abruptly in front (as in cygneus F.) but almost on a
level with the elytra (as in insularis Fst.), the apical collar sunk below the level of
the disk, especially in $. Scutellum three times as broad as long, trapezoidal,
the truncate apex less than half as wide as the base. Elytra longer than broad,
parallel for a short distance, then widening to behind the middle ; the shallow
striae with strong punctures to beyond the middle and becoming much finer
apically, striae 5 and 6 somewhat irregular near the middle ; the intervals broad,
slightly convex, impunctate, 3 and 5 a little more raised near the base. Legs rather
slender ; front tibiae slightly curved inwards at apex in $, straight in <j>. Pygidium
black, with small separated punctures and very short erect pale setae.
Length 7-0-8-5 mm., breadth <$ 2-5-2-7, ?3-°-3'2 mm.
N.E. Burma : Kambaiti, 7000 ft., 6$ 4$, iii-vi. 1934.
In Voss's key to this genus (Stett. ent. Zeit. 90, 1929: p. 130) the male comes
nearest to insularis Fst., but that species lacks the black markings ; the antennal
club is fuscous and has a short curved apical process ; the eyes are much more
convex and slightly longer than the width of the frons ; striae 5 and 6 on the
elytra are quite regular ; the metepimera are much broader and ovate (linear in
nigrigenibus). _^_
• Voss appears to omit the cylindrical basal portion in measuring the length of the head in this
genus.
BURMA AND BRITISH INDIA — COLEOPTERA .' CURCULIONIDAE 417
The female more closely resembles that of montanus Jek., in which, however,
the apical collar of the prothorax is cylindrical, bears no median fovea and has the
front margin very deeply sinuate ; the scutellum is not trapezoidal, but lunate, the
hind margin being broadly arcuate ; the punctures on the elytra are much
smaller and striae 5 and 6 regular.
Paratrachelophorus erosus sp. n.
cJ$. Derm dark red-brown ; a broad frontal band on the head, the scutellum,
the apical part of the pygidium, the third tarsal joint and the apices of the other
joints, black ; sternum with the following markings of golden pubescence : a
small spot in the posterior lateral angle of the metasternum, a larger spot at the
apex of the metepimera, and a narrow stripe on the mesepimera.
(J. Head and rostrum nearly as long as the rest of the body ; the cylindrical
posterior part of the head twice as long as the subconical anterior part, the
former finely transversely striolate above and below, and a large deep fovea on
the hind margin of the black frontal band. Rostrum elongate (7 : 3), with two
shallow sulci from the base to the antennae, which are inserted near the apex.
Antennae elongate, red-brown, with the scape and the apex of the funicular
joints black ; funicle with the apex of joints 2-6 forming a knoblike projection
externally, 1 much the shortest, 2 much the longest being one and a half times as
long as 3, 7 much longer than 1 and clothed like the club. Prothorax twice as long
as its basal width, abruptly sloping upwards in front, transversely striolate above
and with transverse carinae beneath ; the apical collar not constricted, almost
parallel-sided, with a deep median fovea. Scutellum trapezoidal, the apical
margin shallowly sinuate. Elytra one and one third times as broad as long, only
slightly widened behind the middle ; dorsum with a broad shallow transverse
impression at one-fourth from the base and with rows of large subquadrate
foveolae, which do not diminish behind, the intervals very narrow and sinuous, no
wider than the septa between the foveolae, impunctate. Legs as in P. brachmanus
Voss. Sternum with the metepisterna impunctate, except for two or three
punctures near the upper margin anteriorly and a few in the lower inner angle.
Length 17-0-18-5 mm., breadth 3-5 mm.
9- Head about twice as long as the width across the eyes, without any
definite cylindrical part at the base, the sides almost straight. Rostrum not
quite twice as long as broad, the antennae inserted at the middle. Antennae much
shorter, with the funicular joints 2-7 distinctly clavate but not produced outwards,
2 only slightly longer than 3. Prothorax horizontal, as long as its basal width,
conical, with the sides quite straight, the dorsum more finely striolate, the
underside without any transverse carinae. Abdomen with an angular projection
on each side of the pygidium arising from near the lateral margin of the fifth
ventrite, the pygidium itself with a small tubercle in the middle of its apical
margin.
Length io-o-n-o mm., breadth 3-5-3-8 mm.
N.E. Burma : Kambaiti, 7000 ft., 4c?4$, iv-v. 1934.
Differs from all the four previously described species in its coarsely foveolate
elytra and in the tubercle on the pygidium of the female.
Paratrachelophorus sp. — Kambaiti, i^J, vi.
Paratrachelophorus brachmanus Voss.- — Kambaiti, <)<$ 7$, iv-vi ; Sadon, 4(J 5$,
vi-vii.
418 SIR GUY A. K. MARSHALL
Subfamily Rhynchitinae
Deporaus semiruber sp. n.
(J$. Black, with the apex of the rostrum reddish and the elytra entirely
bright red.
Head deeply constricted at the base, with dense strong punctures throughout
and fine subrecumbent white setae, the temples parallel and about as long as an
eye ; irons as wide as the base of the rostrum, with a short median sulcus, eyes
moderately convex. Rostrum about as long as the pronotum, with the apical
margin simple and a strongly punctate sulcus on each side from base to apex ; a
little shorter and stouter in $. Antennae of 9 black, short ; joint I of the funicle
nearly as long as the short scape, 2 longer, 3 a little shorter than 1, 4 and 5 a
little longer than broad, 6 as long as broad, 7 transverse ; club short and
comparatively stout, the two basal joints as long as broad ; antennae of $ with
the funicular joints a little shorter. Prothorax as long as broad, slightly rounded
laterally, shallowly constricted at the apex, widest behind the middle, the base
gently arcuate, the apex not marginate ; dorsum densely and evenly punctate in
(J, without any median stria, the punctures smaller and less dense in 9. the
subrecumbent fine whitish setae longer and denser in <§. Scutellum small,
subquadrate, black. Elytra with the sides straight and gradually diverging from
the shoulders to two-thirds, the rather deep striae with strong punctures, striae
9 and 10 uniting only at the apex ; the intervals somewhat convex and with a row
of distinct punctures, the subrecumbent pale setae much shorter than those on
the pronotum. Legs entirely black ; front and hind tibiae straight, the middle
pair somewhat curved ; joint I of the hind tarsi as long as 2 and 3.
Length 3-0 mm., breadth 1-3 mm.
N.E. Burma : Kambaiti, 7000 ft., i<$ i?, v-vi. 1934.
Apparently most nearly resembling the Chinese bicolor Voss, which has the
same colouring but differs, according to the very brief preliminary description
(Stettin, ent. Ztg. 99, 1938 : 106) available, in having the prothorax strongly
conical, the apex being about half the width of the base, whereas in semiruber it
is four-fifths the width.
Deporaus sp. — S. Shan States, Taunggyi, 5000 ft., 1$, viii-ix.
Deporaus sp. — Sadon, 4000 ft., 1$, vi-vii.
Deporaus spp. — Kambaiti, five specimens of five different species, vi.
Eugnamptus rufipennis sp. n.
$. Body entirely black, except the elytra which are red with a brown
marginal stripe extending indefinitely inwards to interval 9, but not reaching the
base or the apex.
Head with strong separated punctures which become obliquely confluent on
the irons, the temples parallel. Rostrum shorter than the pronotum, widening at
base and apex, dorsally with two continuous punctate sulci, the interspace
impunctate, broad at the apex and regularly narrowing to a sharp point at the
base. Antennae inserted at a little behind the middle of the rostrum, brown with
the articulations pale ; scape much longer than joint 1 of the funicle and equal
to 2, 2-6 subequal, 7 shorter. Prothorax longer than broad, gently rounded
laterally, widest beyond the middle, with a broad shallow apical constriction ;
dorsum with rather large subreticulate punctures and a very narrow abbreviated
median carina, set with numerous erect and suberect pale setae. Scutellum
triangular, red, sparsely setose. Elytra parallel to two-thirds, with a common
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE
419
transverse depression at one-fourth from the base ; the shallow striae with strong
punctures, striae 9 and 10 uniting close to the apex ; the intervals slightly convex,
with a row of small distant punctures and numerous erect and suberect pale
setae of various lengths. Legs black, with long erect setae.
Length 4-6-5-0 mm., breadth 1-5-1-7 mm.
N.E. Burma : Kambaiti, 7000 ft., 2$, vi. 1934.
The only Indo-Burmese species known as yet with uniform red elytra.
Eugnamptus pardalis sp. n.
£. Head, rostrum and prothorax bronze ; elytra pale yellowish brown with
numerous, more or less confluent, dark brown spots, except in the subbasal
depression, and a short dark stripe at about the middle of interval 4.
Head rugosely punctate, with suberect pale setae, the temples slightly
converging behind, the narrow frons with a very deep short sulcus, the eyes very
large. Rostrum shorter than the pronotum, almost parallel-sided at the base
and widening apically, with rugose confluent punctation dorsally and a narrow
median carina on the basal half. Antennae inserted at the middle of the rostrum,
brown, with the articulations pale ; scape longer than joint 1 of the funicle and
equal to 2, which is a little longer than 3, 3-6 equal, 7 slightly shorter. Prothorax
longer than broad, narrowing slightly from the base to one-fourth, then parallel
sided to the broad shallow apical constriction ; dorsum with fine subreticulate
punctation, which becomes transversely confluent at the apex, whereas towards
the base the punctures are less close, a short median stria and behind it a broad
shallow impression, and on each side just behind the apical constriction a small
low shiny tubercle ; the setae rather long, whitish and subrecumbent. Scutellum
bronze, trapezoidal, widest at the base and truncate at the apex. Elytra parallel
from the shoulders to beyond the middle, then widening slightly near the apex,
each with a broad shallow oblique depression at about one-fourth from the base ;
the striae with close small punctures and the intervals with dense irregular
punctures of about the same size as those in the striae, so that the latter appear
to be confused in parts, striae 9 and 10 coalescing near the apex. Legs brown, the
femora testaceous on the basal half and the tibiae with a broad testaceous band
in the middle.
Length 4-5 mm., breadth 1-5 mm.
N.E. Burma : Kambaiti, 4$, iv-v. 1934.
The colouring, combined with the unusual sculpture of the pronotum, will
distinguish this species.
Eugnamptus pannosus sp. n.
c??. Dark brown, with the head blue-black.
Head of $ with comparatively large separated punctures, the temples
parallel ; frons with either a very short shallow stria (type) or a deeper stria that
extends back to the top of the vertex ; head of $ with smaller punctures, much
larger eyes, and much narrower frons with a deep short sulcus. Rostrum of $ a
little shorter than the pronotum (6 : 7), widening at base and apex, the basal half
with two strongly punctate sulci dorsally, the anterior part with irregular
confluent punctures laterally, the impunctate median area narrowing to a point
at the base ; rostrum of <$ much shorter (4:7), parallel-sided at the base, the
anterior half with fine close punctures in the middle only. Antennae inserted at
the middle of the rostrum (<^) or a little behind it ($), brown to fuscous, the club
black with the last joint wholly or partly yellow ; scape much longer than joint 1
of the funicle and equal to 2, 3-6 subequal, 7 shorter. Prothorax longer than
420 SIR GUY A. K. MARSHALL
broad, parallel-sided in (J to the shallow apical constriction, very slightly widening
to two-thirds in $ ; dorsum with large close punctures and an abbreviated median
stria, with sparse erect pale setae. Scutellum trapezoidal, widest at the base, with
a few fine punctures. Elytra parallel to beyond middle and slightly widening
behind, with a broad shallow transverse depression at one-fourth from the base ;
the shallow striae with large close punctures, the intervals slightly convex with
a series of transverse impressions, especially on the apical half, with numerous
erect pale setae of varying lengths. Legs uniform dark brown ; <$ with the usual
mucro on the middle tibiae only.
Length 3-7-4-4 mm., breadth l*2— 1*5 mm-
N.E. Burma : Kambaiti, i^1 2$, v-vi. 1934.
Eugnamptus furvus sp. n.
$. Very closely allied to the preceding species, E. pannosus, sp.n., and
differing only as follows : —
Head with the punctures much finer and more sparse ; frons with or without
a fine short median stria. Rostrum more slender, with finer punctures that are not
confluent. Elytrawith the intervals flat, each with a row of small distant punctures
without any transverse impressions. Legs pale brown.
Length 4-0-4-5 mm., breadth 1-5 mm.
N.E. Burma : Kambaiti, 39, v. 1934.
Eugnamptus spp. — Kambaiti, seven specimens of seven different species, v-vi.
Rhynchites subplumbeus Voss. — Kambaiti, 4^ 1$, iv-v.
Rhynchites ? coarctus Voss. — Kambaiti, i(J 1$, iv-vi.
Rhynchites spp. — Kambaiti, three specimens of three different species, iv-vi.
Rhynchites sp. — Sadon, 1200 m., one specimen, vi-vii.
Rhynchites sp. — Inle Lake, one specimen, ix.
Rhynchites sp. — 40 km. E. of Taunggyi, one specimen, ix-x.
Subfamily Balanininae.
Balaninus pumilus sp. n.
$Q. Derm rather shiny black, the upperside with fuscous setae and markings
formed of rather sparse white setae ; prothorax with a broad white lateral stripe
enclosing two indefinite black spots (sometimes confluent longitudinally) and a few
white setae along the median line ; elytra with a narrow basal band reaching the
shiny humeral callus, another common bisinuate macular band behind the
middle reaching interval 8 when fully developed, these bands united by a narrow
sutural stripe (normally only two setae wide) and with scattered white setae
between them on the disk, interval 5 usually with a short stripe on the basal
third, and variable white setae around the apex ; underside with dense white
scales on the prosternum and along the sides of the rest of the body, the scales
elsewhere much sparser.
Head with the frons as wide as the base of the rostrum, with sparse white
setae. Rostrum strongly curved and about as long as the head and pronotum in
both sexes, with the scrobes lateral ; rostrum of £ (in lateral view) arising from
the upper half of the eye, with four punctate striae on the basal part ; that of $
arising from the middle of the eye, with two dorsal rows of punctures and a very
fine lateral stria. Antennae inserted at ($) or beyond (<$) the middle of the
rostrum, red-brown, with the club fuscous ; funicle with joints 1 and 2 equal, the
distal joints all slightly longer than broad. Prothorax somewhat transverse
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 421
(5:6), greatly rounded laterally, widest at about the middle, not constricted at
the apex, without postocular lobes ; dorsum with close reticulate punctation and
no median carina. Scutellum without any sulcus on each side of it, transverse,
rugulose, bare or with a few pale setae. Elytra with the lateral margins not
sinuate, separately rounded at the apex, the suture not depressed and without
raised interlocked setae apically. Legs comparatively short, black, with the tarsi
piceous, thinly clothed with white setae ; femora with a small sharp tooth, the
hind pair not exceeding the elytra ; tibiae almost straight, with a small mucro.
Pygidium of $ with a deep median sulcus.
Length 1-8-2-0 mm., breadth 10 mm.
N.E. Burma : Kambaiti, 7000 ft., 8<J 3$, v. 1934.
Very closely related to the European B. crux F., in which however the
white dorsal markings are formed of elongate truncate scales, the sutural stripe
on the elytra is much broader (six scales wide), the narrow lateral stripe on the
pronotum is widely separated from the pleural scaling, the prothorax is more
transverse and shallowly constricted at the apex, joint 1 of the funicle is longer
than 2 and the club smaller, and the male has no sulcus on the pygidium.
Balaninus spp. — Kambaiti, six specimens of six different species, v-vi.
Balaninus sp. — Sadon, 1200 m., 1$, vi-vii.
Balaninus sp. — Taunggyi, 1$, viii-ix.
Subfamily Anthonominae.
COPIDORRHINUS gen. n
Head elongate, subconical, temples about as long as the eyes, which are
large and closely approximated above. Rostrum short, stout, almost straight,
longer than broad, its depth about equal to its width, the dorsum in the basal
half strongly tectiform ; scrobes narrow, straight, subtransverse, passing down-
wards in front of the eyes. Antennae inserted at about the middle of the rostrum ;
scape clavate, much shorter than the funicle, exceeding the front margin of the
eye ; funicle 7-jointed, the club elongate, loose, with the joints separated.
Prothorax nearly as long as broad, with the disk raised into a high elevation, the
apical margin truncate and very oblique laterally, the basal margin fitting closely
to the elytra. Scutellum conspicuous. Elytra oblong, with obliquely rounded
shoulders, separately rounded at the apex, which entirely conceals the pygidium,
stria 10 disappearing behind the middle. Legs with the femora feebly clavate,
without any tooth ; tibiae straight, without any uncus or mucro, the hind corbels
dorsal in position ; front coxae narrowly separated, the median pair as widely
separated as the hind pair ; tarsal claws appendiculate. Sternum with the
presternum very short in front of the coxae ; mesosternum rather steeply
declivous ; metasternum somewhat tumid, much longer than a mid coxa, the
metepisterna rather wide, almost parallel-sided behind the basal dilatation and
broadly truncate at the apex, the hind coxae remote from the elytra. Venter with
the intercoxal process ogival, much narrower than a hind coxa, ventrite 2 not
soldered to 1 and hardly longer than 3, the intermediate ventrites angulate
laterally.
Genotype : Copidorrhinus bivittatus sp.n.
The nearest ally of this genus is Demimaea Pasc, which differs as follows : —
Head globose, rostrum not compressed dorsally, antennal club small and compact,
elytra much shorter and subtriangular, front coxae contiguous, tibiae uncinate,
femora with a small tooth, and the intermediate ventrites not angulate laterally.
422 SIR GUY A. K. MARSHALL
Copidorrhinus bivittatus sp. n.
$. Derm dark brown, with black setae and markings of grey or yellowish
setae ; head with a large patch of sparse pale setae behind the lower half of the
eyes ; prothorax with similar setae in the basal angles and on the pleurae ; elytra
with a common trisinuate pale band (interrupted by the striae) before the middle
and a similar common curved band across the top of the declivity, an elongate
spot on interval 5 being in front of the others.
Head with strong close punctures, the temples longer than an eye ; frons
elongate, only as wide as the narrowest part of the antennal scape, parallel-sided
for most of its length ; eyes large, broadly ovate, slightly projecting beyond the
temples. Rostrum stout, longer than broad (3:2), gradually widening from base
to apex ; dorsum with the compressed basal half with large shallow punctures
and rising into a sharp carina in the middle, the anterior part almost flat, shiny
and nearly impunctate. Antennae testaceous ; funicle with joint 1 much thicker
and somewhat longer than 2 ; 3-5 longer than broad, 6 and 7 moniliform, 6 as
long as broad, 7 transverse ; the large club as long as joints 2-7 of the funicle, its
two basal joints subequal and transverse. Prothorax transverse (7 : 8), narrowing
with a slight curve from the base to two-thirds and the sides there broadly and
deeply sinuate, then widening slightly to the apex, the base feebly bisinuate, with
the basal angles rounded ; dorsum raised into a large high obtuse elevation
beyond the middle, the elevation itself with dense small punctures, those on the
subapical part much larger, the basal third almost impunctate but opaque.
Scutellum much longer than broad, with dense pale setae. Elytra parallel-sided
from the shoulders to beyond the middle and broadly rounded behind, with an
indefinite shallow transverse impression at one-fourth from the base between
striae 1 and 4, and a deep common transverse depression on the declivity behind
the obtuse posterior calli ; the striae broad and deeply incised, with deep sub-
quadrate punctures ; the intervals about as broad as the striae, flat, finely rugulose,
the setae (both black and pale) rather dense and appressed. Legs dark brown,
with dense shallow punctures and recumbent setae, the tarsi testaceous.
Length 3-5-4-5 mm., breadth 1-5-2.0 mm.
N.E. Burma : Kambaiti 7000 ft., 2$, vi. 1934.
Demimaea sp. — Kambaiti, i$ 1$, iv-v. 1934. It is not certain whether
these are sexes of one species or belong to two different species.
APIONODES gen. n.
Head subglobose, frons a little narrower than the base of the rostrum, eyes
slightly more convex than the temples. Rostrum long and slender, with the
antennae inserted at not more than one-fourth from the apex in both sexes ;
scrobes oblique, rapidly passing beneath the rostrum. Antennae elongate,
slender ; scape longer than the funicle, abruptly clavate ; funicle with seven
joints ; club fusiform. Elytra subovate, much wider at the roundly rectangular
shoulders than the base of the prothorax, covering the pygidium at the apex,
with ten striae, stria 10 being complete, and without posterior calli. Legs long
and slender ; femora moderately clavate, with a small tooth ; tibiae almost
straight, not uncinate, with a minute mucro in $ but not in $ ; hind tarsi with
joint 1 longer than 2 and 3, 3 deeply bilobate, the claws appendiculate ; front
coxae subconical and contiguous. Sternum with the prosternum not excavated ;
mesosternum with the side-pieces not fused, the intercoxal process much narrower
BURMA AND BRITISH INDIA — COLEOPTERA I CURCULIONIDAE 423
than a coxa, truncate at apex ; mctasternum between the coxae not shorter than
a median coxa, with a transverse row of three foveae in the middle of the base.
Venter with the intercoxal process ogival, about as wide as a coxa ; ventrite 1
not fused with 2 in the middle.
Genotype : Apionodes longipes sp.n.
The Apion-like species on which this genus is based has affinities with
Anthonomus, but the latter differs inter alia in having the prosternum much
shorter in front of the coxae, the scrobes on the rostrum are entirely lateral in
position, and the tarsal claws are toothed and not appendiculate.
Apionodes longipes sp. n.
(J9- Derm rather shiny black to piceous, with sparse recumbent white
setae ; underside with much shorter and sparser setae, which at the sides of the
sternum are replaced by narrow white scales, these being densest along the
metepisterna.
Head with deep separated punctures and sparse recumbent setae ; frons
without any median fovea but with two erect setae beside each eye. Rostrum of
<§ a little longer than the head and pronotum (9 : 8), gently curved, narrowing
very slightly from the base to the antennae and somewhat broader in the apical
part ; dorsum opaque, with a smooth median carina and two narrower ones on
each side ; the lower margin of the scrobe finely crenulated and with a sparse
row of fine erect setae ; rostrum of $ a little longer, with the antennae inserted
slightly further from the apex, and without the crenulation and setae along the
scrobe. Antennae testaceous brown, the scape paler, with sparse short fine erect
setae ; funicle with joint 1 a little longer and thicker than 2, 3 slightly longer than
broad, the rest moniliform and subequal, not widening distally. Prothorax as
long as broad, parallel-sided to three-fourths from the base, then narrowing to the
apex, truncate at base and apex, the basal angles right angles ; dorsum feebly
convex longitudinally, with strong close punctures that are often transversely
confluent at the sides and an obtuse median carina that reaches neither base nor
apex, a transverse row of four pale erect setae along the apical margin and four
more set in a square in the middle of the disk. Elytra widest at the prominent
shoulders, which are smooth and shiny, and gradually narrowing behind to the
obtusely accuminate apex, with a broad shallow depression from stria 1 to 4 at
one-fourth from the base ; the rather deep striae with close punctures, the intervals
as broad as the punctures, rather convex, smooth and shiny, the alternate ones
with a row of four or five widely spaced stiff erect white setae. Legs flavous, with a
broad blackish band on the femora and sparse short recumbent white setae ;
femora with the tooth small and sharp, that on the front pair minute.
Length 27-2-9 mm., breadth i-o-l-l mm.
N.E. Burma : Kambaiti 7000 ft., 3c? 4?, iv-vi. 1934.
Sphinxis sp. — Kambaiti, one specimen, iv.
Rhynchaenus spp. — Kambaiti, two specimens of different species, iv.
< ienus ? — Kambaiti, two specimens of different genera, iv-v.
Subfamily Cioninae.
Cwnus obesus albopttnctatus Auriv. — Sadon, 1200 m., i<$, vi-vii.
424 SIR GUY A. K. MARSHALL
Subfamily Prionomerinae.
Ochyromera sericea sp. n.
$. Derm red-brown, rather thinly clothed with short recumbent fulvous
pubescence, through which the derm is visible, with scattered groups of paler
hairs, the pubescence lying in different directions, largely oblique or transverse
on the elytra.
Head subglobular, not constricted behind the eyes, with small dense strong
punctures and sparse pubescence ; frons three-fifths the width of the rostrum,
with a rather large shallow glabrous median fovea ; eyes gently convex, longer
than the temples. Rostrum long, slender, curved ; longer than the head and
prothorax (4:3), gradually widening from the antennae (inserted at five-eighths
from base) to the apex ; the basal part with coarse confluent punctures, a narrow
median carina and a short one on each side, the apical area with fine separated
punctures, and a short deep stria between the antennae. Antennae red-brown, the
club fuscous ; scape a little longer than the funicle, which has joint 1 much longer
than 2 (3:2), 4-7 subequal, as long as or slightly longer than broad. Prothorax
transverse (2:3), rounded laterally, widest at a little behind the middle, with a
broad constricted apical collar ; dorsum feebly convex longitudinally, finely
rugulose, with small low granules that are rather unevenly distributed and an
ill-defined smooth median line. Scutellum slightly raised, suboblong, with dense
grey pubescence. Elytra very slightly narrowed behind the broad roundly-
rectangular shoulders, thence parallel to beyond the middle, separately rounded
at the apex ; dorsum somewhat flattened on the disk in the basal half, with a
shallow common transverse impression as far as stria 4 at the basal fourth, with
rows of small deep punctures of varying sizes that do not diminish behind ; the
intervals broader than the punctures, somewhat uneven, with an irregular row of
small low shiny granules, intervals 3 and 5 with a low granular elevation not far
from the base and another behind the middle, the posterior callus on interval 5
sharply conical and bearing a low tuft of fulvous setae, with a deep impression
behind the callus. Legs red-brown, with macular groups of recumbent yellow
setae and sparse stiffer subrecumbent dark setae ; front tibiae broadly and
roundly dilated near the apex, the inner apical angle rounded off.
Length 5-5 mm., breadth 2-4 mm.
N.E. Burma : Kambaiti, 7000 ft., 2$?, v-vi. 1934.
Ochyromera brevicornis sp. n.
(J$. Derm black, the elytra becoming red-brown towards the base ; pro-
thorax unevenly clothed with recumbent yellow setae, with a bare median stripe ;
elytra with much denser similar setae and a broad common dark band before the
middle on which the setae are much sparser.
Head subglobular, not constricted, with coarse subreticulate punctures and
sparse setae ; frons five-sevenths of the width of the rostrum, with a rather large
shallow median fovea ; eyes moderately convex, but not exceeding the line of the
temples. Rostrum rather stout, slightly shorter than the head and pronotum in
$ and slightly longer in $, very gradually widening from base to apex in $, in $
parallel-sided to beyond the antennae and then slightly widening to the apex ;
with coarse confluent punctures in £, a median carina that becomes narrower
and sinuous at the base ; and with two narrow sinuous carinae on each side ; the
punctures less confluent in 9, the lateral carinae less distinct and the median one
not nearly reaching the base. Antennae short, red-brown with the club fuscous ;
funicle a little shorter than the scape, with joint 1 somewhat longer than 2, 3-7
BURMA AND BRITISH INDIA — COLEOPTERA \ CURCULIONIDAE 425
transverse, especially 6 and 7, which are twice as broad as long and disk-like.
Prothorax transverse (n : 13), rounded laterally, widest behind the middle, with
a broad constricted collar ; dorsum gently convex longitudinally, with coarse
reticulate punctures, the interspaces being very narrow. Scutellum triangular,
with close shallow punctures and sparse setae. Elytra parallel from the broad
roundly-rectangular shoulders to beyond the middle, the apices separately
rounded ; dorsum sloping steeply forwards at the base, there being a deep
impression within the shoulder on stria 5 and interval 1 being still more deeply
depressed on the basal fourth, so that there is a low obtuse elevation between
striae 1 and 5, the posterior calli forming large rounded tubercles covered with
dense setae, and the broad rounded shoulders projecting very slightly laterally ;
the narrow indistinct striae containing small deep separated punctures, the
intervals with a row of shining granules, those on 3, 5, 7 smaller and more
numerous, those on 2, 4, 6 larger and widely separated, each granule bearing a
puncture containing a short stiff suberect black seta. Legs red-brown, the knees
and front tibiae blackish, the tarsi honey-brown, the yellow setae long and
rather sparse ; front tibiae rugose, curved, widening from the base to three-fourths,
where there is on the lower edge a long, sharp spine, the inner apical angle
rounded off in <$ but forming a sharp right-angle in $.
Length 5-3-6-0 mm, breadth 2-3-2-5 mm.
N.E. Burma : Kambaiti, 7000 ft., 1^2$, vi. 1934.
Ochyromera cognata sp. n.
$. Very similar superficially to 0. brevicornis, sp.n., but the elytra consider-
ably longer in proportion ; derm red-brown, the clothing similar, except that the
dark band on the elytra is less distinct.
Structure similar except as follows : — Head with frons four-sevenths of the
width of the rostrum ; eyes somewhat less convex. Rostrum with the lateral
carinae less distinct. Antennae with the two basal joints of the funicle equal,
3 and 4 longer than broad, 5 and 6 as long as broad, 7 transverse. Prothorax with
the spaces between the punctures appreciably broader. Elytra sloping much less
steeply at the base and the three basal depressions much shallower ; the granules
on the intervals much flatter and with smaller setigerous punctures, those on
intervals 3, 5, 7 larger and much fewer. Legs entirely red-brown ; front tibiae
with the dorsal edge forming a flat regular curve, the lower edge broadly dilated at
a little beyond the middle into a large sharp rectangular tooth, the inner apical
angle rounded off.
Length 60 mm., breadth 2-5 mm.
N.E. Burma : Kambaiti, 7000 ft., ij, v. 1934.
Ochyromera coronata sp. n.
$. Derm red-brown, with rather thin uniform pale brown recumbent
pubescence on the elytra, the middle of the prothorax almost bare, the sides and
pleurae with dense longer pubescence.
Head elongate, subcorneal, with coarse reticulate punctures ; frons five-
sevenths of the width of the rostrum, with a shallow median depression ; eyes
almost flat, about as long as their distance from the prothorax. Rostrum straight,
elongate, shorter than the head and prothorax (8:9), parallel-sided from the
base to beyond the antennae, then slightly widening to the apex, with coarse
longitudinal punctures, without any carinae. Antennae honey-brown ; funicle
much shorter than the scape (3 : 4), joint 1 longer than 2, 3 longer than broad, 4
and 5 as long as broad, 6 and 7 transverse, 7 broader than the others. Prothorax
426 SIR GUY A. K. MARSHALL
very nearly as long as broad, strongly but unevenly rounded laterally, widest at
one-fourth from the base, with a broad apical collar and a basal constriction ;
dorsum very rugose, the disk bearing a coronet of six conical tubercles arranged
in a circle, and in front of this a short longitudinal carina on each side of the
apical collar. Sciitellum with a deep sulcus on each side, subtriangular, rugulose,
sparsely setose, with a shallow impression down the middle. Elytra with the
shoulders obtusely projecting laterally, parallel thence to beyond the middle,
with the apices separately rounded ; the dorsum rather steeply declivous at the
base, the striae with small close punctures ; interval 3 with a densely pubescent
low curved costa at the base, close behind it on 5 a small obtuse tubercle, a large
tubercle on 4 at about the middle, and the posterior callus large and conical, all
the intervals with unevenly distributed small granules, those on 8 larger and
those on 6 larger still, the granules bearing short subrecumbent pale setae. Legs
red-brown, with fairly close recumbent yellow setae ; front tibiae gently curved on
the upper edge, the lower edge with a broad sharp acute-angled tooth at three-
fifths from the base, the inner apical angle sharply produced.
Length 6-5 mm., breadth 2-8 mm.
N.E. Burma : Kambaiti, 7000 ft., 1$, vi. 1934.
Ochyromera sp. — S. Shan States, Taunggyi, 1500 m., 1$, viii-ix.
Key to Burmese Species of Ochyromera.
1 (6). Head subglobular, temples shorter than an eye ; rostrum curved ;
scutellum without a sulcus on each side ; prothorax without
dorsal tubercles.
2 (3). Front tibiae roundly dilated near apex, without any tooth on the
lower edge ; elytra with a small conical tubercle at the middle of
interval 3, the posterior calli sharply conical . . sericea sp. n.
3 (2). Front tibiae with a sharp tooth on the lower edge ; elytra without
a tubercle on interval 3, the posterior calli obtusely rounded.
4 (5). Front tibiae with a narrow spine-like tooth on the lower edge at
three- fourths from base ; funicle with joint 1 longer than 2, the
distal joints very short and disk-like . . . brevicornis sp. n.
5 (4). Front tibiae with a broad rectangular tooth at two-thirds from
base ; funicle with the two basal joints equal, the distal ones
bead-like ....... cognata sp. n.
6 (1). Head subconical, temples as long as an eye ; rostrum straight ;
scutellum with a deep sulcus on each side ; prothorax with a
ring of six conical tubercles on the disk . . coronata sp. n.
Endaeus signatus sp. n.
cJ. Derm dark red-brown, with the following black markings on the elytra :
a broad dentate oblique band starting at one-third from the base on intervals
5-7 and ending at the top of the declivity on intervals 2-3 ; a sub-basal marking on
intervals 3-5, the part on 3 much longer than the others, but that on 5 some-
times uniting with the band behind ; and a spot at the apical junction of intervals
4-6 ; metastemum and venter mainly blackish.
Head not constricted behind the eyes, with fine close punctures and rather
sparse pubescence ; frons two-thirds the width of the rostrum, with a small
median fovea ; eyes moderately convex. Rostrum shorter than the pronotum
(6 : 7), very stout, curved, parallel-sided in the basal half and somewhat broader
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 427
beyond the antennae, with tine longitudinally confluent punctation and a low
narrow median carina. Antennae red-brown ; funicle not widening distally, joint
1 as long as but much thicker than 2, 3-7 transverse, moniliform and subequal.
Prothorax transverse (7 : 10), strongly rounded laterally, widest at the middle,
not or very shallowly constricted at the apex, which is much narrower than the
subtruncate base ; dorsum with dense fine punctation and remote larger punctures,
and with fine transverse recumbent pubescence and sparse short suberect setae,
there being a row of these projecting over the front margin. Elytra broadly ovate,
very slightly widened beyond the middle, broadly rounded behind, with the
apices separately rounded, the posterior calli obsolescent ; the striae with small
separated punctures that hardly diminish behind, the intervals rather shiny,
feebly convex, with thin pubescence and a row of minute granules, each bearing
a short stiff suberect yellow seta. Legs red-brown, sparsely pubescent ; front
femora with a large triangular tooth with a row of stiff erect setae beyond it, the
tooth on the other pairs rather smaller ; front tibiae deeply sinuate on the basal
half, the wider apical part carinate on its lower edge. Venter with ventrites 2
and 3 somewhat angulate laterally.
Length 3-5-4-0 mm., breadth 1-6-1-7 mm.
N.E. Burma : Kambaiti, 7000 ft., 5^, iv-vi. 1934.
Differs from the previously described Oriental species in its colouring and,
especially, in having seven joints in the funicle instead of six.
Endaeus sp. — Kambaiti, 1$, vi.
Endaeus sp. — Kambaiti, i$, iv.
Subfamily Acicnemidinae
Acicnemis spp. — Kambaiti, two specimens of different species, iv-vi.
METRACHODES gen. n.
The species described below appeared to be congeneric with Trachodes
aculangulus Heller 10,08, from India, but as the genus was founded by Germar on
an Alaskan species, ptinoides Germ.,* the generic allocation seemed somewhat
dubious. A comparison of Germar's species with the five available Old World
species (Itispidus L., hystrix Gyll., oblongus Keitt., ovatus Reitt., and acittangulus
Hllr.) has shown that they are not even closely related. The following are some
of the salient differences.
In Trachodes the front coxae are contiguous and the femora are unarmed ;
the antennal scrobes are visible laterally right up to the base and the antennae
are inserted at one-third from the apex of the rostrum ; and the elytra have
stria 10 complete.
In the other species the front coxae are separated and the femora bear a
strong tooth ; the antennal scrobes pass rapidly beneath the rostrum and the
antennae are inserted behind the middle ; and stria 10 is much abbreviated.
The affinities of typical Trachodes are clearly with the small wingless
Hylobiinae, to which it should be transferred. On the other hand, the Old
World species belong to the subfamily Acicnemidinae ; indeed, I am unable to
find any character by which to distinguish oblongus from Acicnemis, in which it
should now be included, and this also applies to Trachodes elongatus Reitt.,
judging from the description. The other species mentioned have a different
fades, the elytra being much shorter and broader, without any true humeral
calli, the scutcllum is concealed, and they have no functional wings.
* This is an error; the genotype was hispidui I so thai W eb u Falls -i-* .1 synonym
of Trachodes. G.A.K.M.
428 SIR GUY A. K. MARSHALL
A new genus Metrachodes is therefore proposed for them, with the genotype
Trachodes hispidns L. Pascoe erected the genus Semelima for a wingless species
of Acicnemis from Borneo ; but this differs from Metrachodes in its elongate
facies, the hind tibiae extend beyond the apex of the elytra, and joint 2 of the
tarsi is longer than 3. Whereas in Metrachodes the hind tibiae do not exceed the
elytra, and joints 2 and 3 of the tarsi are equal.
This segregation of the species of "Trachodes" into two different subfamilies
necessitates the abolition of the subfamily Trachodinae. The only other genus
included in it in the Catalogus Coleopt. is Ancylocnemis Mshl., which should be
transferred to the Acicnemidinae.
Metrachodes albicollis sp. n.
$. Derm dull black ; prothorax densely covered above and below with
yellowish white scales, except for a large subquadrate black patch in the middle
of the base ; elytra with fuscous scaling, a pale humeral patch which extends
backwards for a short distance on interval 7, and an ill-defined pale subapical
band, and with tufts of erect black scales.
Head with dense whitish scaling ; frons much narrower than the base of the
rostrum, with a single erect scale by each eye and a deep median fovea. Rostrum
a little longer than the pronotum, parallel-sided from the base to the antennae,
there rather abruptly narrowed and then gradually widening again to the apex,
the basal part with dense whitish scaling, the apical part bare, shiny and with
sparse minute punctures. Antennae red-brown, with the club black, inserted
a little behind the middle of the rostrum ($) ; funicle with joint 1 as long as but
much thicker than 2, 3-5 transverse and subequal, 6 and 7 as long as broad ;
club rather broadly ovate. Prothorax about as long as broad, strongly rounded
laterally, widest at the middle, constricted at the base, which is only slightly
wider than the apex and truncate, the basal angles right angles, the apical margin
slightly arcuate, the postocular lobes feeble ; dorsum strongly convex longitudin-
ally, highest at the middle, lower at the base than at the apex, the large deep
punctures concealed by the dense scaling, except in the quadrate basal black
patch, with a transverse row of four tufts of erect scale-like setae at a little in
front of the middle and two similar tufts on the apical margin. Elytra broadly
ovate, widest before the middle, rapidly narrowing behind to the obtusely
acuminate apex, without any humeral or posterior calli ; the somewhat irregular
striae with very large punctures basally, which diminish greatly behind ; the
intervals rather sinuous and uneven, the alternate ones with an irregular row of
erect scale-like setae, usually two or three together, and a much larger tuft
before the middle on interval 3 containing 10-12 setae. Legs red-brown, with
the distal half of the femora blackish ; the strongly clavate femora with a small
sharp tooth on the posterior pairs and a longer one on the front pair ; tibiae
comparatively slender, shallowly bisinuate on the lower edge (without any
angular projection), with a very small apical mucro ; joint 3 of the tarsi not
truly bilobate, the apical margin being only shallowly sinuate.
Length 2-6 mm., breadth i-i mm.
N.E. Burma : Kambaiti, 7000 ft., 2$, iv-vi. 1934.
Subfamily Alcidodinae.
Alcidodes artivittis sp. n.
cJ9- Derm rather shiny black, sparsely clothed with short yellowish setae,
some of which are bifid or trifid ; prothorax with five narrow stripes of plumose
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 429
scales varying from whitish to fulvous, three dorsal, the others on the pleurae, the
median stripe not narrower than the others ; elytra with the following markings
formed of similar scales : a round spot on interval 3 at one-fourth from the base,
an oblique narrow stripe running from stria 9 at one-fourth from the base to
stria 5 at a little before the middle, an exactly parallel stripe behind this from
stria 9 to stria 1 at the top of the declivity (these stripes partly broken by the
striae), a longitudinal stripe on interval 3 starting just behind the previous one
and reaching the apex, where it joins a stripe of about the same length on interval
9 ; underside with sparse yellowish plumose scales which are denser at the sides
of the sternum and form four rows of very indefinite spots on the venter.
Head densely punctate, sometimes with a small median fovea. Rostrum
long and slender, slightly curved, subcylindrical, a little widened at the apex
and at the insertion of the antennae (at middle in $, beyond in $), with dense
small punctures on the basal half ; length in $ one and a half times the median
line of the pronotum, in $ nearly twice the pronotum. Prothorax somewhat
transverse, widest at the base, subcorneal, with the sides gently rounded, very
shallowly constricted near the apex, the dorsal apical margin truncate, the
postocular lobes moderate ; dorsum flat longitudinally, with small separated
granules of unequal sizes. Scutellum not enclosed, small, oval, shiny. Elytra
elongate, a little wider at the shoulders than the prothorax, parallel to near the
middle, then gradually narrowing behind, with a broad shallow transverse
depression near the base ; the shallow striae with rather large close subquadrate
punctures, which are as broad as the narrow rugulose intervals. Legs rather long
and slender, black, with yellowish setae ; femora with a small sharp denticulate
tooth ; tibiae straight, the front pair sharply angulated before the middle, the
hind pair with the corbel ascending the dorsal edge. Sternum with the front
intercoxal space about one-fourth the width of the median one.
Length 7-0-8-0 mm., breadth 2-5-3-0 mm.
Upper Burma: Nam Tamai, 4000 ft., 1$, viii. 1938 (R. Kaulback— type) ;
Kambaiti, 7000 ft., 1$, v. 1934 ; Sadon, 4000 ft., i<J, vi-vii. 1934 (Dr. R.-
Malaise). India : N. Manipur, 1$, 1889 (Doherty).
Type in the British Museum, cotypes in the Stockholm Museum.
Nearly allied to A. westermanni Boh., which has all the stripes much broader
and less regular ; the pronotum bears punctures bordered by lunate granules ;
the elytra are broader and much less narrowed behind, the punctures in the striae
smaller and narrower than the intervals ; and the femora have a fringe of suberect
setae on the lower edge.
Alcidodes basipennis sp. n.
$9. Derm black, opaque ; prothorax with five narrow stripes of fulvous
scales, three being dorsal and the others on the pleurae ; elytra with the following
pale yellowish markings : a spot on interval 3 at one-third from the base, a very
oblique narrow band from interval 9 at one-third from the base to about the
middle of 4, and a similar but much less oblique band near the top of the declivity,
which is often straight but usually somewhat bisinuate and runs from interval 9
to the suture ; underside with the following fulvous marks : a broad oblique
stripe on the metasternum, a spot at the base of the metepisternum, and lateral
spots on the ventrites ; all the scales plumose.
Head densely punctate, bare, usually with a smooth median spot in place of
a fovea. Rostrum of <J long and slender, one and a half times the length of the
novit. zool., 42, 3. 1948. 82
430 SIR GUY A. K. MARSHALL
pronotum, closely and strongly punctate from the base to the antennae (at one-
third from the apex) and more finely beyond, only slightly widened at the apex ;
rostrum of $ longer (nearly twice the pronotum), rather more slender, more
widened at the apex, with the antennae at a little beyond the middle. Prothorax
transverse (7 : 10), subconical, widest at the base, rapidly narrowing with a
slight curve to the very shallowly constricted apex, the dorsal apical margin
truncate, the postocular lobes slight ; dorsum fiat longitudinally, with small
separated granules of various sizes, the spaces between the fulvous stripes
almost bare. Scutellum not enclosed, very small, bare. Elytra wider than the
prothorax, widest at the shoulders and very gradually narrowing posteriorly,
the sutural area to interval 3 more or less flattened from the base to two-thirds,
the basal margin from the suture to interval 4 raised into a rather broad flat
strigose elevation ; the striae with rather large deep punctures, many of the
septa between them being somewhat raised and uniting laterally with those in
adjoining striae to form short transverse ridges, so that the whole surface appears
uneven ; the intervals irregular, convex and rugulose. Legs rather long, black,
with very short sparse pale setae ; femora rugosely punctate, with a sharp
crenulate tooth ; tibiae with the corbels ascending the dorsal edge, the front
tibiae rather deeply bisinuate on the lower edge, but the submedian projection
rounded, not angulate.
Length 7-2-9-0 mm., breadth 2-7-3-5 mm.
N.E. Burma : Kambaiti, 7000 ft., 23^ 17$, iv-vi. 1934.
The allied species, scenicus Fst. and artivittis sp.n., may both be distinguished
by the absence on the elytra of the elevation at the base and the transverse ridges
on the disk ; further they both have a white stripe on the apical part of interval 3.
Alcidodes trigonophorus Mshl. — Kambaiti, 4^ 6$, iv-v.
Alcidodes westermanni Boh. — Sadon, 1200 m., 1$, vi-vii.
Alcidodes decursus Pasc. — Sadon, 1$, vi-vii.
Alcidodes pectoralis Boh. — Sadon, 1$, vi-vii.
Alcidodes affaber Auriv. — Punkaung, between Sadon and Mytikyina, 1$, vii.
Alcidodes fremitus Fst. — Inle Lake, 900 m., i^, xi.
Alcidodes sp. — Kambaiti, i<^, v.
Subfamily Cryptorrhynchinae.
Dcsmidophorus hebes F. var. aterrimus Aur. — Moulmein, 3^, xi ; S. Shan States,
Inle Lake, S. end, 3000 ft., 1$ 3$, ix ; 40 km. E. of Taunggyi, 1^, ix-x.
It may be noted that D. morbus Pasc. (Siam) was erroneously sunk by
Hubenthal (Ent. Bl. 13, 1917 : 114) as a variety of hebes ; it is really a very
distinct species allied to schenklingi Hub. In the Catalogus Coleopt. the name is
wrongly spelt morphosus.
Desmidophoras confucii Boh. — Washaung, 700 ft., 20 km. E. of Myitkyina,
2q- v"-
Thisus burmanus sp. n.
$$. Derm dull black or piceous, with fairly dense grey or brownish scaling ;
the dorsal tufts formed of thick short blackish setae, except one at the top of the
elytral declivity which is yellowish.
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 43I
Head covered by the pronotum, with the frons transversely impressed.
Rostrum as long as the pronotum, moderately stout, strongly curved, not dilated
at the apex, but narrowing dorso- vent rally beyond the antennae, which are
inserted at one-third from the apex ; the basal half rugose and densely squamose,
the apical half shiny, bare and finely punctate. Antennae piceous to red-brown ;
funicle widening distally, the two basal joints of equal length, 3-7 transverse,
moniliform. Prothorax nearly as long as broad, widest at the base, narrowing
very slightly with a feeble curve to the broad shallow apical constriction ; dorsum
set with close deep subreticulate punctures, with the transverse row of four
tufts at about the middle and two smaller ones at the strongly arcuate apical
margin, which conceals the head. Scutellum small, convex, bare. Elytra much
longer than broad (3 : 2), almost parallel from the prominent shoulders to beyond
the middle, produced downwards at the apex in $ ; dorsum convex longitudinally,
highest at the middle, very steeply declivous behind ; the shallow striae containing
coarse punctures but more or less obscured by the small round convex scales ;
the intervals broader than the striae, interval 1 with a row of short distant peg-
like erect setae ; 3 with an elongate low elevation near the base, a shorter one at
the middle, and a conical one at the top of the declivity, all these bearing tufts of
erect peg-like setae ; 5 with a short basal elevation and three others which
alternate with those on 3 and similarly tufted. Legs black or piceous with
blackish scales and sparse short erect pale setae, the tarsi red ; femora with a
rather broad triangular tooth ; tibiae bisinuate beneath and shallowly sinuate
dorsally.
Length 3-5-4'0 mm., breadth I-8-2-0 mm.
N.E. Burma : Kambaiti, 7000 ft., 1^ i2, v-vi. 1934.
The genotype, T. biguttatus Pasc. is a larger insect with much broader
elytra (3 : 2-6) ; the prothorax is more transverse, much more convex longitudinally
and does not conceal the head, and the tufts in the transverse row (not mentioned
by Pascoe) are much lower and more diffuse ; the similar elevations on the elytra
are much lower and inconspicuous ; the tooth on the femora is much smaller,
Thisus sp. — Kambaiti, i2, vi.
Thisns sp. — Kambaiti, itf, v.
YPSILEPIDUS gen. n
Head concealed from above by the pronotum ; frons much wider than the
rostrum ; eyes coarsely faceted. Rostrum about as long as the pronotum in both
sexes, moderately curved, parallel-sided, not constricted at the base ; scrobes
passing rapidly beneath the rostrum, not uniting at the base, their apices shortly
visible from above ; mandibles decussate, strongly toothed. Antennae inserted at
the apical third in (J, a little further back in 2 '. scape slender in the basal half,
strongly and abruptly clavate ; funicle with joint 1 as long as but much thicker
than 2 ; club pubescent, the sutures indistinct. Prothorax as long as broad,
shallowly bisinuate at the base, the postocular lobes feeble. Scutellum concealed.
Elytra rather short and broad, with the sides strongly rounded, without true
shoulders, the posterior calli small and obtuse ; stria 10 abbreviated. Wings not
functional. Legs rather slender ; femora clavate, with a small tooth ; tibiae
not compressed, nearly straight, with a long uncus and no mucro ; tarsi with
joint 2 transverse, shorter than 3, which is bilobate, the claws rather slender.
Sternum with the pectoral furrow bare, not extending behind the front coxae,
which are widely separated ; mesosternal process declivous, a little wider than
432 SIR GUY A. K. MARSHALL
the prosternal ; metasternum as long as a median coxa, the metepisterna
indistinguishable. Venter with the intercoxal process broadly rounded ; ventrite
2 as long as I and twice as long as 3 and 4, its basal margin rounded.
Genotype : Ypsilepidus thisoides sp.n.
Belongs to the group of genera round Colobodes Schonh. and most nearly
resembles Thisus Pasc. ; but from all these it may be distinguished by the absence
of functional wings, of the scutellum, and of the metepisternal suture.
Ypsilepidus thisoides sp. n.
(J$. Derm red-brown, with uniform grey scales intermixed with particles of
a brown substance which is probably a natural exudation ; underside bare, shiny.
Rostrum of ^ moderately stout, slightly widened at the apex, densely
clothed from the base to the antennae with U-shaped or Y-shaped scales, which
conceal the sculpture, the apical part bare, shiny, with very fine shallow
punctures ; rostrum of ^ a little narrower, with the scaling restricted to the basal
third. Antennae red-brown ; funicle with joint 3 a little longer than broad, 4-7
moniliform, transverse, 7 the broadest. Prothorax as long as broad, parallel-
sided from the base to the middle, then narrowing to the unconstricted apex ;
dorsum with large deep close punctures, with an abbreviated smooth median
line, but the 'sculpture more or less hidden by V-shaped scales and brown
exudation ; at about the middle a transverse row of four tufts of stout erect
rod-like setae and two other tufts on the apical margin. Elvtra (5 : 4) strongly
rounded laterally, widest at about the middle ; the dorsal outline sloping steeply
at the base, flat in the middle, and steeply declivous behind ; the striae with
rather large deep punctures but normally covered by scaling ; interval 3 with a
tubercle near the base, a much smaller one not far behind it, a much larger one
at the top of the declivity, and a small one on the declivity ; interval 5 with a
small sub-basal tubercle (on a level with that on 3), another at the middle, and a
third on the declivity ; all these tubercles are covered by a tuft of stout erect
peg-like brown setae ; the scales are all U-shaped or Y-shaped. Legs black, with
narrow pale scales, some of which are suberect, tarsi red-brown. Underside with
very sparse punctures.
Length 3-6-4-0 mm., breadth 2-0-2-2 mm.
N.E. Burma : Kambaiti, 7000 ft., ij 1$, vi. 1934.
The form of the scales is very unusual, and it seems possible they have been
developed to keep in place the brown substance (presumably an exudation) that
occurs among them, which may serve a procryptic purpose.
Ocoblodes sp. — Kambaiti, 1$. vi.
Tadius erirrhinoides Pasc. — Rangoon, 1^ 1$, xii ; Siam, Medaw (Burma frontier),
1$, xi.
Mecistocerus guttatus sp. n.
^$. Derm rather dull black ; prothorax with a rather narrow irregular
sublateral stripe of fulvous scales and with variable paler scaling on each side of
the median carina ; elytra with blackish scaling and numerous variable fulvous
spots ; underside with sparse isolated small pale scales, and a small spot in the
hind angles of ventrites 2-4.
Head with fine punctation laterally, the frons almost impunctate in the
middle with a broad deep median sulcus that encroaches on the vertex. Rostrum
longer than the pronotum, deeply constricted at the base, with the usual coarse
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 433
basal punctures and short median carina, the punctate area being only slightly
more extensive in $, in which however, the base is squamose, being bare in 9-
Antennae inserted at the middle of the rostrum (9) or at three-eighths from the
apex (£), dark red-brown to blackish ; funicle rather densely pubescent, joints
1 and 2 equal, 3-5 longer than broad, 6 and 7 subglobular or transverse.
Prothorax nearly as long as broad, subparallcl-sided almost to the middle, then
narrowing in a curve to the comparatively narrow apical constriction ; dorsum
only slightly convex longitudinally, with rather large deep reticulate punctures
(somewhat larger on the pleurae) and a shiny carina that reaches from the base
almost to the apex, the narrow interspaces with fine sparse punctures, without
conspicuous setae. Scutellum trapezoidal, convex, bare, shiny. Elytra broad,
parallel from the roundly rectangular shoulders to the middle, the posterior calli
distinct, the apex not produced downwards in $ ; the shallow striae with rather
shallow subquadrate separated punctures that contain no scale and are greatly
diminished behind ; the intervals broader than the punctures, comparatively
smooth, interval 3 not costate near the base, the setae very short and appressed.
Legs, including the tarsi, black ; femora with black scaling and sparse short pale
recumbent scale-like setae, the posterior pairs with a pale subapical dorsal patch,
the tooth large and triangular ; tibiae black, with the apical half pale on the
posterior pairs, feebly bisinuate beneath, the external apical angle of the hind
pair rounded. Metasternum with large shallow reticulate punctures laterally,
shiny and almost impunctate in the middle with a complete shallow median
sulcus.
Length 8-5-12-0 mm., breadth 3-5-5-1 mm.
N.E. Burma : Kambaiti, 7000 ft., 2$ 8$, vi. 1934.
This unusually large species is characterised by the broad shiny impunctate
area in the middle of the metasternum.
Mecistocerus latisetis sp. n.
cJ9. Derm black, with almost uniform dense brown scaling ; underside with
sparse small pale scales, one in each puncture.
Head densely squamose, with a comparatively small deep frontal fovea.
Rostrum longer than the pronotum (5:4), rather slender, constricted at the base,
rugosely punctate and tricarinate in the basal half and rather densely squamose
near the base. Antennae inserted at about the middle of the rostrum (9) or a
little beyond it (<$), red-brown ; funicle sparsely pubescent, the two basal joints
equal, 3-5 longer than broad, 6 and 7 globular or slightly transverse. Prothorax
nearly as long as broad, parallel-sided in the basal half, then narrowing in a
curve to the broad shallow apical constriction ; dorsum gently convex
longitudinally, the small dense punctures almost entirely concealed by scaling,
without any smooth median line or carina, and unevenly set with very thick
short suberect setae. Scutellum small, somewhat transverse, convex, with a few
minute setae. Elytra broad, parallel from the roundly rectangular shoulders t<>
about one-third, the posterior calli distinct, the apex not produced downwards
in 9; the elongate separated punctures partly concealed by scaling, diminishing
behind and each containing a yellowish scale ; the intervals broader than the
punctures, flat, with a row of very broad triangular truncate squamiform setae,
which are either black or yellow, interval 3 not elevated at the base. Legs slender,
black, the tarsi red-brown, with dense uniform light-brown Milling ; femora with
a triangular tooth ; tibiae almost straight, the outer apical angle of the hind pair
434 SIR GUY A- K- MARSHALL
rectangular, the hind unci of J with a small tooth on the dorsal edge. Metasternum
laterally opaque and with rather small separated punctures, in the middle, shiny,
with much smaller punctures and a complete median sulcus.
Length 6-0-6-5 mm., breadth 2-6-3-0 mm.
N.E. Burma : Kambaiti, 7000 ft., ig 1$, v-vi. 1934.
Readily distinguished by the unusually broad fan-like setae on the elytra.
Mecistocerus nubilus sp. n.
<J$. Derm black ; elytra densely squamose, mottled with black, brown and
grey ; scaling much less dense on the prothorax owing to the narrow spaces
between the punctures ; underside with sparse small pale scales.
Head densely squamose, with a very large bare triangular frontal fovea,
which is broad in front and narrows to a point behind. Rostrum of $ longer than
the pronotum (5 14), constricted at the base, with four shallow punctate sulci
and a narrow median carina on the basal third ; that of <$ rugosely punctate to
beyond the middle and with three narrow sinuous carinae. Antennae inserted at
about the middle of the rostrum in $ and at one-third from the apex in $,
entirely red-brown ; funicle thinly pubescent, the two basal joints equal, 3 and
4 longer than broad, 5 as long as broad, 6 and 7 more or less transverse. Prothorax
as long as broad in <$, transverse (4 : 5) in $, parallel-sided in the basal third,
then narrowing in a curve to the shallow apical constriction ; dorsum only
slightly convex longitudinally, with large deep reticulate punctures (those on the
pleurae similar), the narrow interspaces of which are squamose, the scales on the
disk being mostly black, with an irregular narrow pale stripe on each side, the
very short setae recumbent and inconspicuous, except near the apex where they
are suberect. Scutellum trapezoidal, widest behind, with an impression in the
middle of the anterior margin, bare, shiny. Elytra rather broad, parallel from the
roundly rectangular shoulders to beyond the middle, the posterior calli distinct,
the apex not produced downwards in § ; the shallow striae with deep round
punctures that are partly overlapped by scaling, being separated by at least
their own length in $ but closer in <J, and diminishing greatly behind ; the
intervals broader than the punctures, even, except towards the base where they
tend to be subgranulate, interval 3 slightly raised near the base, the suberect
scale-like setae very short. Legs black, with the tarsi red-brown ; femora with a
rather large triangular tooth, the scaling pale basally, black towards the apex
with pale setae and a large subapical dorsal pale patch, but in smaller specimens
the scaling may be almost entirely pale ; tibiae with black scales on the basal
half and pale apically, the external apical angle of the hind pair rounded.
Metasternum opaque and with very large shallow punctures laterally, shiny and
with smaller punctures in the middle, and with a complete median sulcus.
Length 5-5-9-5 mm., breadth 2-0-4-0 mm.
N.E. Burma : Kambaiti, 7000 ft., 2<J 7$, v-vi. 1934.
The nearest ally is corticeus Fst., which is a broader insect with the shoulders
of the elytra more sloping and the sides slightly rounded, and interval 3 is not
elevated near the base ; the funicle of the antennae is densely pubescent and thus
appears much thicker ; the venter is much more densely squamose, the scales
being dark brown in the middle of ventrites 3 and 4 and pale elsewhere, while in
the middle of I and 2 the punctures are very large and close, whereas in nubilus
they are here very small and sparse.
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 435
Mecistocerus integrirostris sp. n.
9. Derm rather shiny black to piceous, with the apex of the rostrum and the
apical margin of the prothorax red-brown ; prothorax with a narrow median
stripe of yellowish scales on the basal two-thirds and sparse scales laterally which
sometimes form an indefinite stripe ; elytra with irregular broken bands of
yellowish scales ; underside with only a minute narrow scale in each puncture.
Head with dense small punctures, the frons without any sulcus or fovea.
Rostrum not constricted laterally at the base, which is punctured like the head,
the lateral areas opaque on the basal half, the disk being shiny and sparsely
punctate. Antennae inserted well beyond the middle($), the scape and the basal
funicular joint honey-brown, the remainder more or less fuscous ; funicle not
pubescent, joint 1 much longer than 2, the rest bead-like, 7 strongly transverse.
Prothorax nearly as long as broad, moderately rounded laterally, widest at a
little behind the middle, broadly but shallowly constricted at the apex ; dorsum
longitudinally convex on the basal half, with dense deep reticulate punctation
(similar to that on the pleurae) and a very short median carina, the punctures on
the disk containing a recumbent scale. Scutellum round, opaque, rugulose.
Elytra rather narrow, parallel to beyond the middle, produced downwards at the
apex in $ ; the striae with close quadrate punctures that almost disappear
apically ; the intervals mostly a little narrower than the striae, rugulose, minutely
granulate towards the base, with a row of suberect spatulate setae, interval 3 with
an elongate granular elevation near the base and much smaller ones adjoining it
on 4 and 5. Legs black to piceous, with uniform sparse pale scales, the tarsi
paler ; femora with rather large close shallow punctures and a small sharp tooth ;
tibiae almost straight, the front pair very shallowly sinuate on the dorsal edge,
the corbels of the hind pair with the outer angles slightly projecting. Metastemum
with comparatively small dense punctures and with only a narrow median
sulcus on the posterior half.
Length 3-0-4-0 mm., breadth 1-0-1-5 mxn-
N.E. Burma : Kambaiti, 7000 ft., 5$, v-vi. 1934.
This small species differs from all those hitherto recorded from Burma in
having the rostrum unconstricted at the base and joint I of the funicle distinctly
longer than 2.
Mecistocerus cariniceps sp. n.
1^$. Derm dull black ; prothorax with a transverse patch of fulvous scales
on each side of the front margin ; elytra with a broad irregular (sometimes
macular) band of fulvous scales across the top of the declivity, the anterior part
with variable spots of similar scales ; underside with an oblong scale in each
puncture.
Head rugosely punctate, with an irregular median carina, the frons with
only a small median puncture. Rostrum a little longer than the pronotum (7 : 6),
strongly constricted at the base ; dorsum of $ with four broad, coarsely punctate
sulci from the base to beyond the middle separated by three narrow carinae, the
sulci shorter and narrower in $ and the interspaces broader. Antennae inserted
at a little beyond the middle of the rostrum in $, at two-thirds from the base in
$, red-brown ; funicle thinly pubescent, the two basal joints equal, 3-6 longer
than broad, 7 as long as broad. Prothorax very nearly as long as broad, parallel-
sided from the base to the middle, then narrowing in a curve to the rather deep
apical constriction ; dorsum gently convex longitudinally, with large deep
honeycomb punctures (those on the pleurae smaller and shallower), the narrow
436 SIR GUY A. K. MARSHALL
interspaces tending to run together in sinuous oblique lines, and with a rather
strong sinuous median carina ; most of the punctures with an oblong suberect
scale. Scutellum nearly round, bare and shiny. Elytra elongate, parallel from the
roundly rectangular shoulders to two-thirds, the posterior calli low but distinct,
the apex not produced downwards in $ ; the shallow striae with deep punctures
that diminish behind and are separated by about their own length, each puncture
containing a recumbent scale and most of the discal ones with their anterior
margin subgranulate ; the intervals as broad as the punctures on the basal half
and much broader behind, finely rugulose and with a row of suberect scale-like
setae, intervals 3 and 5 with a low granulate elevation near the base. Legs black,
with the tarsi red-brown ; femora with a large triangular tooth, front pair with
dense fulvous scales except at the apex which is dark brown, posterior pairs with
mixed fulvous and brown scales and a broad fulvous band ; tibiae almost straight,
the basal half with brown, the apical half with pale scales, the corbels of the hind
pair obliquely rounded externally. Metasternum opaque and reticulately
punctured laterally, shiny in the middle, with small distant punctures and only
a narrow sulcus in the middle of the apical half.
Length 6-0-8-0 mm., breadth 2-3-3-0 mm.
N.E. Burma : Kambaiti, 7000 ft., i<?5?, v-vi. 1934.
Distinguished from the other Burmese species by the sculpture of the head
and the granules in front of the punctures on the elytra. Superficially it most
resembles M. subcylindricus Fst., but the latter differs also in having a longer
and more smooth prothorax, the shoulders of the elytra are narrower and more
sloping, the legs are shorter and much thicker, and the metasternum is equally
coarsely punctate throughout.
Mecistocerns spp. — Kambaiti, seven specimens of seven different species, v-vi.
Mecistocerus sp. — Sadon, 1$, vi-vii.
Rhadinomerus stricticollis sp. n.
9- Derm dull black ; prothorax with some fulvous scaling between the bare
granules ; elytra with irregular tufts and short transverse bands of short suberect
oblong fulvous scales.
Head bare on the vertex, with coarse shallow punctures ; frons opaque,
shallowly rugulose, with sparse scales and a deep median sulcus that extends on
to the base of the rostrum, and a deep furrow adjoining the eyes. Rostrum about
as long as the pronotum, deeply constricted at the base, with four punctate sulci
on the basal half, the median space narrowing to a point before reaching the base.
Antennae inserted at the middle of the rostrum, red-brown ; funicle pubescent,
joint 1 as long as but much thicker than 2, 3 and 4 subequal and longer than
broad, 5 as long as broad, 6 and 7 transverse. Prothorax as long as broad,
rounded laterally, widest at the middle, strongly constricted near the apex, the
constriction continued deeply across the disk, truncate at the base ; dorsum
strongly convex, highest behind the middle, coarsely punctate, with the intervals
partly subgranulate, there being a low median carina on the anterior half, with
a large granule on each side of it close behind the constriction ; scaling confined
to the punctures, with sparse short suberect clavate setae. Scutellum very small,
bare. Elytra ovate, with the shoulders reduced and obliquely rounded, the
posterior calli small ; the separated punctures rather large, subquadrate, partly
filled with scaling and diminishing behind, the rows being somewhat irregular in
parts, the interspaces in the rows as long as or longer than the punctures and
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 437
often bearing large flattened granules ; the intervals rather irregular, sinuous and
subgranulate in parts, interval 3 being slightly higher than the others and raised
at the base into a large elongate granular tubercle. Legs with fairly dense elongate
brown scales ; femora with only a small tooth that is partly obscured by scales ;
tibiae straight. Underside : metasternum depressed in the middle but without
any median sulcus, set with large contiguous punctures, each containing a short
scale ; ventrite 1 with large separated punctures, 2-4 impunctate.
Length 7-0 mm., breadth 3-0 mm.
N.E. Burma : Kambaiti, 7000 ft., 2$, v-vi. 1934.
Most nearly allied to the Sumatran, Berosiris pauper Pascoe, which is a
Rhadinomerus, and the sloping shoulders of the elytra in both species suggest
that they may be incapable of flight. R. pauper lacks the very deep subapical
constriction of the prothorax, which is simply punctate, without any trace of
granules or median carina ; the frontal sulcus does not extend on to the rostrum ;
and the metasternum has a deep median furrow.
The deep subapical constriction of the pronotum distinguishes this species
from all its known congeners.
Rhadinomerus impar sp. n.
(^$. Derm rather shiny black or piceous ; prothorax with the sides and
pleurae rather thinly clothed with yellowish scales, the disk dark with an
indefinite pale median stripe ; elytra variegated with yellowish scales which tend
to form irregular broken transverse bands.
Head closely punctate, without any frontal fovea or sulcus. Rostrum a little
longer than the pronotum, constricted at the base, confluently punctate to
beyond the middle in $ and with three narrow carinae ; in $ punctate only to
the middle and with the carinae less distinct. Antennae inserted at a little beyond
the middle in $ and at about one-third from the apex in $, with the scape and
joint 1 of the funicle flavous, the rest red-brown ; funicle not pubescent, joint 1
slightly longer and much thicker than 2, the rest bead-like and transverse.
Prothorax a little broader than long, moderately rounded laterally, widest behind
the middle, with a broad shallow apical constriction ; dorsum slightly convex
longitudinally in the basal half, with a broad shallow depression apically, and
with dense small reticulate punctures, without any smooth median line, the
scale-like setae (mostly black) obliquely raised and some projecting over the
apical margin. Scutellum round, bare, opaque. Elytra oblong-ovate, rather broad
(9 : 5'5). parallel from the prominent shoulders to well beyond the middle, the
subapical depressions distinct ; the striae containing close subquadrate punctures
(partly obscured by scaling) that diminish behind, the larger punctures with a
minute shiny granule on each side ; the intervals as broad as the striae on the
basal half and broader behind, bearing small granules and a row of suberect
scale-like setae (some black, some yellowish), interval 3 with an elongate granular
tubercle near the base, a much smaller one adjoining it on 4, and a trace of one
on 5. Legs piceous, with the tarsi flavous, thinly clothed with uniform grey
scaling ; the femoral tooth very small and sharp ; tibiae straight, the uncus quite
straight in £, somewhat curved in 9- Underside : metasternum densely punctate,
without any fovea or sulcus ; ventrites 1 and 2 with small separated punctures,
3 and 4 opaque, impunctate.
Length 3-0-5-0 mm., breadth 11-20 mm.
N.E. Burma : Kambaiti, 7000 ft., 3c?4$. v-vi. 1934.
438 SIR GUY A. K. MARSHALL
Rhadinomerus favicollis sp. n.
cJ$. Derm black ; prothorax with scattered suberect spatulate pale setae
(probably one in each puncture normally) ; elytra irregularly variegated with
black and fulvous to yellowish bands or patches of scales.
Head coarsely punctate, with a few erect scales near the eyes, frons with
a minutely punctate smooth median area including a broad deep sulcus.
Rostrum as long as the pronotum, constricted at the base, coarsely punctate
and tricarinate basally, the median carina distinct in$, almost obliterated in $.
Antennae inserted at slightly beyond the middle in $ and at about one-third from
the apex in (J, red-brown ; funicle not pubescent, joints 3 and 4 as long as broad
and subequal, 5-7 transverse and bead-like. Prothorax a little broader than long,
almost parallel-sided to beyond middle in g, slightly rounded in $, broadly and
shallowly constricted at apex ; dorsum longitudinally convex basally and
depressed in front, closely set with deep subreticulate punctures, like a shallow
honeycomb, with a very narrow abbreviated median carina. Scutellum round,
bare, shiny. Elytra elongate, parallel from the roundly rectangular shoulders to
the middle, the subapical calli well-marked in $, less so in $ ; the striae with small
close subquadrate punctures that diminish behind ; the intervals broader than
the striae, rugulose, with a row of suberect scale-like setae (black or yellowish),
interval 3 with a short elevation near the base and a shorter one on 5. Legs black
or piceous, the tarsi paler ; femora with a small stout tooth, thinly squamose and
with a pale subapical band ; tibiae with the basal half dark, the apical half with
pale scaling, the lower edge shallowly bisinuate, corbels of the hind pair with the
outer angle rectangular and slightly projecting. Underside : metasternum opaque
and densely punctate, except in the middle where it is shiny with separated
punctures, without any median fovea or sulcus ; ventrites 1 and 2 with small
sparse punctures, 3 and 4 almost impunctate.
Length 4-5-5-2 mm., breadth 1-2-2-0 mm.
N.E. Burma : Kambaiti, 7000 ft., i<$ 2$, v-vi. 1934.
Rhadinomerus furcatipes sp. n.
cJ§. Derm rather shiny black with scattered sparse groups of narrow
yellowish scales.
Head with strong close punctures, the frons with a large deep median fovea.
Rostrum slightly longer than the pronotum, constricted at the base, with coarse
confluent punctation on the basal half and a smooth median line. Antennae red-
brown ; funicle not pubescent, joints 3 and 4 subequal and slightly longer than
broad, 5-7 transverse. Prothorax slightly broader than long, rounded laterally,
widest behind the middle, broadly constricted at the apex ; dorsum longitudinally
convex basally and depressed in front, with rather large reticulate punctures and a
fine short median carina, with sparse narrow elongate clavate scales. Scutellum
rounded, rugulose and setose. Elytra oblong-ovate, parallel from the roundly
rectangular shoulders to the middle, more narrowed apically than in the two
preceding species, especially in $, the posterior calli feeble in $, a little more
prominent in $ ; the striae with small close punctures that diminish behind ; the
intervals broader than the striae, slightly convex, shagreened, with a row of
suberect scale-like setae, interval 3 with a short basal elevation and a much
smaller one on 5. Legs black, evenly covered with sparse pale scales, the tarsi
red-brown ; femoral tooth small on the hind pair, more or less rudimentary on
the anterior pairs ; tibiae straight, exterior angle of corbels rounded, the mucro
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 439
on the posterior pairs straight and bifurcate in $, simple and curved in $.
Underside : metasternum opaque and closely punctate laterally, shiny and with
more distant punctures in the middle, there being a fairly impunctate area in the
middle of the base, with a shallow median depression, but no fovea or sulcus ; all
the ventrites with separated strong punctures.
Length 4-7-5-0 mm., breadth 1-7-2-0 mm.
N.E. Burma : Kambaiti, 7000 ft., 2$ 19, vi. 1934.
Rhadinomerus spp. — Kambaiti, five specimens of five different species, v-vi.
Key to the Burmese Species of Rhadinonomerus.
1 (2). Apical area of prothorax separated off by a deep sulcus ; elytra
with obliquely sloping shoulders .... slricticollis sp.n.
2 (1). Apical area of pronotum simply depressed ; elytra with prominent,
roundly rectangular shoulders.
3 (4). Prothorax widest in front of the middle, without a median carina ;
elytra with dense yellowish scaling . . . contemptits Fst.
4 (3). Prothorax widest at or behind the middle ; pale scales on elytra
forming spots or irregular bands.
5 (6). Prothorax coarsely granulato-punctate, without a median carina,
widest at middle ; intervals on elytra narrower than the
punctures ; metasternum coarsely punctate throughout, with a
median fovea at base and apex ; ventrite 1 with a row of very
large punctures along the whole base . . . granulicollis Fst.
6 (5). Prothorax not granulate, with a short median carina (except
impar, sp. n.) ; intervals on elytra not narrower than the
punctures ; metasternum without median foveae ; ventrite 1
without a basal row of very large punctures.
7 (8). Frons without a fovea or sulcus ; pronotum without a median
carina ........ impar sp.n.
8 (7). Frons with a deep fovea or sulcus ; pronotum with a short median
carina.
9 (12). Setae on upperside scale-like and suberect ; intervals on elytra not
granulate ; interval 3 with a short elevation near base.
10 (11). Frons with a deep sulcus ; scutellum bare, shiny ; femora with a
pale subapical band ; hind tibiae with the outer angle of the
corbel rectangular and slightly projecting, uncus on posterior
pairs of $ normal ...... favicollis sp.n.
11 (10). Frons with a deep fovea ; scutellum rugulose, setose ; femora
uniformly squamose ; hind tibiae with the outer angle of the
corbel rounded, uncus on posterior pairs of ^bifurcate furcatipes sp.n.
12 (9). Setae on upperside hair-like and erect ; intervals on elytra finely
granulate, interval 3 without a basal elevation . . conciliates Fst.
The characters of R. contemptus and conciliates have been drawn from the
descriptions only.
Rhadinopus tecticollis sp. 11.
$$. Derm black, variegated with dense pale brown, black and whitish
scales without any definite pattern ; underside with a broad suberect whitish
scale in each puncture.
Head with pale brown and whitish scales and behind each eye a large patch
of short erect black peg-like scales ; frons in front about as wide as the rostrum
44° SIR GUY A. K. MARSHALL
near the base and rather rapidly narrowed behind, with a small fovea. Rostrum
with the antennae inserted at about one-third from the base in both sexes ; the
basal area with coarse confluent punctation and a smooth median carina that
does not ascend the frons ; the anterior part with very sparse fine punctures in
§, these being distinctly larger in ^J. Antennae red-brown, with joint 2 of the
funicle a little longer than I, 3 somewhat longer than broad and equal to 4, 5-7
subequal and bead-like ; club broadly ovate in one aspect, somewhat compressed
in the other. Prothorax nearly twice as broad as long, widest close to the base and
there gently rounded, then narrowing rapidly to the apex, which is shallowly
constricted ; dorsum raised in the middle of the disk into a tectiform ridge, with
a complete narrow median carina, the deep pentagonal punctures with their
interspaces rather broader than usual and opaque ; the paler scales broad and
recumbent, the black ones erect and peg-like. Scutellum nearly round, squamose,
its cavity as long as broad. Elytra comparatively short and very broadly ovate,
with only a small subapical impression and no posterior calli ; the deep striae
with large punctures, which are partly concealed by scaling, the intervals a little
broader than the striae, flat and not carinate even posteriorly, except interval 9,
interval 1 with a smooth line along the sutural edge ; scales dense, smaller than
those on the pronotum, all the intervals with a row of numerous suberect scale-
like setae, the punctures in the striae containing a long narrow scale. Legs with
fairly dense brown and whitish scales ; femora convex on the dorsal edge, with a
very small second tooth ; tibiae with the sharply carinate dorsal edge only
slightly curved and for the greater part parallel with the lower edge.
Length 5-0-5-5 mm., breadth 3-0-3-2 mm.
Burma : Tenasserim, Malvedaung, 1000 ft., 30 km. S. of Ye, 2$ 1$, xi. 1934.
Distinguished from all the previously described species by the tectiform
pronotum and the second femoral tooth.
Rhadinopus indutus sp. n.
cj$. Derm piceous, with dense pale fawn-coloured scaling above and a few
tufts of black peg-like setae ; underside with small isolated recumbent pale
scales.
Head rugosely punctate, with a small recumbent scale in each puncture ;
frons rapidly narrowing posteriorly, with a small fovea. Rostrum with the
antennae inserted at two-fifths from the apex in both sexes ; dorsum rugosely
punctate on the basal half in $, on the basal third in 9- Antennae red-brown, with
the two basal joints of the funicle subequal, 3 longer than 4, 5-7 subequal and a
little longer than broad ; club somewhat compressed. Prothorax nearly twice as
broad as long, widest at the base, rapidly narrowing to the apex with the sides
slightly curved, shallowly constricted at the apex, which is half as wide as the
base ; dorsum flat along the median line, with large dense punctures and a narrow
sinuous median carina, the punctures partly hidden by large flat scales ; across
the middle a row of four small tufts, each containing three or four erect stout
peglike black setae, and two longer and denser tufts on the apical margin.
Scutellum with a small median tubercle. Elytra triangular, widest at the
shoulders and rapidly narrowing to the apex with the sides slightly curved, with
a distinct subapical constriction and well-marked posterior calli ; the punctate
striae concealed beneath the broad overlapping scales ; the intervals flat, broader
than the striae, interval 3 slightly higher than the others with a row of three or
four tufts of erect peg-like black setae, 9 costate at the apex only, 1 with a smooth
line along the sutural edge, 5 and 7 with a row of distant erect scale-like pale
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 44I
setae. Legs with fairly dense, pale fawn scaling; femora nearly straight on the dorsal
edge, with only one small tooth beneath ; tibiae with the upper and lower edges
parallel and without any external angulation at the apex.
Length 3-8-4-0 mm., breadth 2-0-2-1 mm.
N.E. Burma : Kambaiti, 7000 ft., i<J 1$, v-vi. 1934.
From the previously described species of the genus indutus differs in its
small size, more triangular elytra with their flat intervals and concealed striae,
the tuberculate scutellum, entirely bare pectoral channel, etc.
Rhadinopus sp. — Kambaiti, 1$, vi.
Rhadinopus sp. — Myitkyina, 1$, vii.
Rhadinopus sp. — Tenasserim, Mekane, 90 km. E. of Moulmein, 200 m., i<3\ xi.
Zeugenia hislrionica Pasc. — Mekane, 1$, xi.
Zeugenia sp. — Kambaiti, 1$, vi.
Orochlesis spp. — Mekane, two specimens of different species, xi.
Cryptorrhnchyus sp. — S. Shan States, Inle Lake, 1$, ix.
Cryptorrhynchus sp. — 40 km. E. of Taunggyi, 1^, ix-x.
Genus ? (Cryptorrhynchinae) — four specimens of three different genera, Kambaiti,
v-vi, and Washaung, vii.
LEPIDARCUS gen. n.
Head convex, not hidden by the pronotum ; frons narrower than the rostrum
between the antennae ; eyes coarsely faceted. Rostrum roundly dilated near the
base and constricted at the actual base ; scrobes passing beneath the rostrum
basally and there dilated. Antennae inserted at a little beyond the middle of the
rostrum in both sexes ; scape about as long as the funicle, clavate ; funicle with
joints 1 and 2 equal ; club elongate, widest beyond the middle, segmented.
Prothorax transverse, bisinuate at the base, the apical margin arcuate, the
postocular lobes distinct. Scutellum exposed, squamose. Elytra much wider at
the shoulders than the prothorax, with ten complete striae, the subapical calli
prominent. Legs short, slender ; femora only slightly clavate, not sulcate
beneath, with a small tooth, the hind pair not exceeding the subapical calli on the
elytra ; tibiae not compressed, straight, with a strong uncus and no mucro ; tarsi
with joint 2 longer than broad, as long as 3, the claws slender. Sternum with the
pectoral furrow bare, the mesosternal cavity horseshoe-shaped, much higher than
the metasternum, overhanging at the apex, reaching the middle of the median
coxae ; metasternum longer than a median coxa. Venter with the intercoxal
process narrow, sharply acuminate; posterior margin of ventrite 1 quite straight ,
2 longer than 3 but shorter than 3 and 4.
Genotype : Lepidarcus malaisei sp. n.
This genus comes nearest to the Papuan Hyparinus Pasc, in which however
the mesosternal receptacle is on a level with the metasternum and reaches the
hind margin of the median coxae ; the intercoxal process of the venter is broad
and forms an obtuse angle ; the femora exceed the elytra and the tibiae are
strongly compressed ; the antennae are inserted far behind the middle of the
rostrum, and joint 2 of the funicle is longer than 1.
Lepidarcus malaisei sp. n.
cJ$. Derm black ; upperside with dense fawn scaling ; pronotum with two
indefinite darker discal stripes from the median tufts to the base ; elytra with a
more or less diamond-shaped large common blackish area, extending to stria 4 in
the middle and narrowing to stria 1 at the base and to stria 2 at the top of the
442 SIR GUY A. K. MARSHALL
declivity, being sometimes invaded by fawn scales ; metasternum and the basal
ventrite with isolated fawn scales in the punctures, the remainder being almost bare.
Head with the rugose punctures entirely concealed by the dense overlapping
concave scales. Rostrum as long as the pronotum, rather strongly curved, the
basal half with large subreticulate punctures and a smooth median line, the apical
half smooth with sparse small punctures ; rostrum of $ similar, except that the
apical half is a little narrower with smaller closer punctures. Antennae honey-
brown ; funicle with joints 3 and 4 subequal and slightly longer than broad, 5-7
transverse, moniliform, 7 being the widest. Prothorax transverse (5 : 6), widest
at the middle, narrowing slightly to the base, much more so in front,
shallowly constricted at the apex ; dorsum with a broad shallow median
longitudinal depression, a transverse row beyond the middle of four tufts of stout
erect scales (the median ones larger) and two more at the apex, the whole
punctation concealed by very large overlapping flat scales, much larger than those
on the elytra. Scutellum densely squamose. Elytra parallel from the roundly
rectangular shoulders to beyond the middle ; the dorsal outline rising shortly
from the base, then flat to the top of the declivity, which slopes steeply to the
apex ; the rows of large deep punctures (each containing a narrow scale)
completely hidden by large overlapping scales, except near the suture ; interval
2 with three erect tufts of dense black scales, near the base, at the middle and at
the top of the declivity, interval 4 with two tufts in between those on 2, other
intervals with a row of very short erect scale-like setae. Legs with the femora
black, the tibiae and tarsi red-brown ; femora with dense grey scaling, with two
dorsal black spots, a large median and a smaller subapical one ; tibiae with grey
scaling and the apical half black dorsally.
Length 5-0-5-5 mm., breadth 2-0-2-2 mm.
N.E Burma : Kambaiti, 7000 ft., 4^ 1$, v-vi. 1934.
Lepidarcus sp. — Kambaiti, i(J, v.
Subfamily Lobotrachelinae.
Genus LOBOTRACHELUS Schanh.
The small weevils of this genus appear to be numerous in the Old World
Tropics, and as they are difficult to discriminate, it has seemed advisable to
describe here all the Burmese species of which adequate material is available and
to supply a key which may facilitate their identification.
Faust (Ann. Mus. Stor. nat. Genova, 1895, 34 : 294) recorded nine species
from Burma, but the occurrence of one of these (angulatus Mot.) seems dubious ;
of the remainder four are known to me from the description only : ineptus Gyll.,
teucaspis Fst., ruficornis Mot., and ingratus Fst., and of these only leucaspis is
included in the key. Dr. Malaise found six new species, and five more are
described from the British Museum collection. Thus twenty-one species have now
been recorded, of which eighteen appear in the key, and there are eight additional
species of which I have seen single specimens only.
Key to the Burmese Species of Lobotrachelus.
1 (4). Clothing of elytra consisting entirely or mostly of scales.
2 (3). Dorsal scaling pale grey ; elytra each with three large round black
spots (the discal one sometimes indistinct) ; legs very long and
slender, hind femora exceeding the elytra by almost half their
length ; rostrum of 5 with a few scales at base only and a row of
fine erect hairs on each side ..... sexnotatus sp. n.
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 443
3 (2). Elytra with an indefinite pattern of pale brown and white scales ;
legs much shorter, femora broad, the hind pair only shortly
exceeding the elytra ; rostrum of $ squamose nearly to apex,
without erect hairs ...... lepidotus sp. n.
4 (1). Clothing of elytra consisting entirely of setae, or very rarely
(Jrenatus) with a few markings of narrow scales.
5 (8). Tooth on front femora forming a broad lamina.
6 (7). Elytra with markings of white setae but no white scales ;
pronotum without a white band at base and apex laminatipes sp. n.
7 (6). Elytra with markings of narrow white scales : a sutural stripe, a
long stripe on interval 5, a lateral patch before middle ;
pronotum with a white band at base and apex . . frenatus sp. n.
8 (5). Teeth on front femora in the form of a simple spine, or more or less
obsolete.
9 (14). Front femora with two teeth.
10 (13). Teeth on front femora placed one behind the other but not in
alignment.
11 (12). Elytra clothed with dark setae except for a basal band of white
setae on intervals 1-4 ; tarsi testaceous . . . bidens sp. n., (J$
12 (11). Elytra with rather dense uniform grey setae ; tarsi black
plumbeus Mot., $
13 (10). Teeth on front femora placed side by side . . . plumbeus Mot., $
14 (9). Front femora with a single tooth or none.
15 (22). Scutellum entirely concealed by the median basal lobe of the
pronotum ; many intervals on the elytra with only a single row
of setae.
16 (17). Ventrite 5 of $ with a deep round bare punctate depression
occupying nearly the whole surface, each side with an angulate
projection bearing a tuft of dense setae ; rostrum of $ very
long, as long as the elytra ; antennae with the scape and joint
1 of the funicle testaceous, the rest black . . morosus Fst.
17 (16). Ventrite 5 of $ without any depression or projections, closely
squamose ; rostrum of $ much shorter than the elytra ; antennae
entirely testaceous.
18 (21). Rostrum forming a distinct angle with the frons ; elytra without
bands of white setae ; tibiae wholly or partly red.
19 (20). Rostrum and all the tibiae red ; prothorax with a fairly large
patch of white scales in the anterior and posterior angles
rufitibiis sp. n.
20 (19). Rostrum black, with the extreme apex red ; tibiae black, the
anterior pairs red on the apical third, the hind pair only at the
apex ; prothorax with only a fringe of single white scales in the
hind angles and no white scales in the anterior ones parens sp. n.
21 (18). Dorsal outline of the rostrum forming a continuous line with the
frons ; elytra with a basal and postmedian band of sparse
whitish setae ; tibiae black ..... asperulus Fst.
22 (15). Scutellum visible behind the prothoracic lobe ; intervals on elytra
with two or more rows of setae or none.
444 S1R GUY A- K- MARSHALL
23 (28). Front femora with the tooth absent or rudimentary.
24 (27). Pronotum only slightly convex longitudinally in the middle line,
the discal punctures comparatively large, much wider than the
intervals between them.
25 (26). Upper surface mostly clothed with long coarse grey setae ; suture
of elytra with a complete stripe of intercrossed grey setae ;
rostrum of $ not deeper at the antennae than near the base
vacivus sp. n., $.
26 (25). Upper surface mostly clothed with short dark setae, with
scattered groups of sparse whitish setae ; suture of elytra
without such a stripe ; rostrum of $ much deeper at the
antennae than near the base . . . simplicipes sp. n., ^9-
27 (24). Pronotum strongly convex longitudinally, the discal punctures
smaller than the intervals . . . convexicollis sp. n., $.
28 (23) . Front femora with a single sharp tooth.
29 (36). Disk of pronotum dull, with large shallow ocellated punctures, each
containing a seta, the intervals narrower than the punctures.
30 (31). Basal lobe of pronotum clothed with sparse grey setae ; scutellum
with dense whitish scales ; suture of elytra with a complete
stripe of intercrossed grey setae .... vacivus sp. n.,§.
31 (30). Basal lobe of pronotum with a patch of dense silvery-white
scales ; scutellum bare or with dark setae ; suture of elytra
without such a stripe.
32 (33). Vertex of head opaque ; upper edge of the area of white scales on
the propleurae sinuate in the middle ; rostrum of <J not deeper
at the antennae than at the base ; median area of metasternum
and basal ventrite of $ with dense erect hairs . . subfasciatus Mot.
33 (32). Vertex of head shiny ; upper edge of the white area on the
propleurae quite straight ; rostrum of <J deeper at the antennae
than at the base ; median area of metasternum and basal
ventrite of <§ with flat scales.
34 (35)- Sides of prothorax curved ..... leucaspis Fst.
35 (34)- Sides of prothorax straight ..... urenae Mshl.
36 (29). Disk of pronotum shiny, with small simple punctures without a
seta, the intervals wider than the punctures.
37 (38). Elytra much broader at the shoulders than the prothorax, with
the intervals rugulose and a common bisinuate band of sparse
white setae behind the middle reaching stria 7 ; middle of
metasternum with plumose scales .... laporteae Mshl.
38 (37). Elytra only slightly broader at the shoulders than the prothorax,
with the intervals smooth, and having behind the middle at
most a few white setae on interval 3 ; middle of metasternum
with simple recumbent setae .... nudus sp. n.
Lobotrachelus sexnotatus sp. n.
$$. Derm black, with dense pale grey scaling above and below ; pronotum
with two round black spots in the middle of the disk (clothed with short brown
setae), but these almost obliterated in <$ ; elytra each with three large round
black spots (clothed with brown setae), one on the humeral callus, another before
the middle between striae 1 and 4 (sometimes partly obliterated) , and the third on
the declivity between striae 2 and 4.
BURMA AND BRITISH INDIA — COLEOPTERA I CURCULIONIDAE 445
Head with dense scaling, forming an erect crest along the frons, which is
comparatively broad. Rostrum of $ as long as the pronotum, gradually widening
in the apical half, porrect to the antennae (at two-sevenths from the apex) then
curving downwards, much deeper at the antennae than at the base, compressed
dorsally into a sharp high median carina and densely squamose throughout ;
rostrum of 9 slightly longer, with the antennae a little further back, not deeper
there than at the base, dorsum with scales only at the base and a row of short
erect hairs on each side, the carina much less prominent. Antennae testaceous.
Prothorax transverse (2 : 3), with the sides straight, the median basal lobe rather
short, densely squamose, not carinate and not concealing the scutellum ; dorsum
moderately convex longitudinally, highest at the middle, with dense shallow
ocellated punctures (concealed except in the discal spots) and no smooth median
line. Scutellum exposed, squamose. Elytra with the fine striae largely obscured
by the dense elongate scales, the broad intervals strongly rugulose. Legs unusually
long and slender, the hind femora exceeding the elytra by nearly half their length,
the front pair without a tooth or with a rudimentary one, the posterior pairs with
a small sharp tooth.
Length 3-0-3-5 mm., breadth 1-5-1-7 mm.
N.E. Burma : Kambaiti, 7000 ft., 1$ 3$, v. 1934.
Lobotrachelus lepidotus sp. n.
$. Derm black to piceous, with more or less dense scaling forming an
indefinite pattern ; prothorax with the median lobe pure white, two large round
dark spots in the middle of the base and two fainter ones in front of them, two
indefinite whitish patches on each side of these, which either merge into the
white on the pleurae or are separated from it by pale fawn-coloured scales ; elytra
with pale fawn scales, a subquadrate whitish patch at the base of intervals 1-3, an
indefinite common arcuate dark band crossing the suture before the middle and
extending up to the shoulder on each side, and behind this some variable short
whitish stripes usually on intervals 3, 5 and 7 ; underside with dense whitish
scales.
Head with dense fawn scales ; frons forming a wide angle with the rostrum, a
little broader than the apex of the scape, the scales suberect. Rostrum of $
elongate, as long as the pronotum (with the lobe), nearly straight, fairly closely
squamose almost to the apex. Antennae inserted only a little beyond the middle
of the rostrum, testaceous, elongate, all the joints longer than broad.
Prothorax somewhat transverse (5 : 6), with the sides gently rounded, the basal
lobe not carinate and partly concealing the scutellum ; dorsum slightly convex
longitudinally, with dense ocellated punctures (mostly concealed) and without
any smooth median line. Scutellum squamose. Elytra with narrow striae
containing a row of pale setae, the intervals rugosely punctate, the dark scales
very narrow, the white ones the broadest. Legs piceous, with dense whitish
scales, the tarsi paler ; femora short and broad, with a sharp tooth, the hind pair
only shortly exceeding the elytra.
Length 3-5 mm., breadth 1-5 mm.
Burma, 3$ (^4. K. Weld Downing).
Type in the British Museum .
novit. zool., 42, 3. 1948. 88
446 SIR GUY A. K. MARSHALL
Lobotrachelus laminatipes sp. n.
<J9. Derm black to red-brown ; prothorax dorsally with rather sparse whitish
setae, becoming dark on the disk, the basal lobe with dense narrow white scales,
the pleurae with a large irregular patch of broader white scales at base and apex
(sometimes uniting in the middle) ; elytra with a nearly complete sutural stripe of
interlocking white setae (but often much abbreviated), a broad common basal
band of white setae extending to interval 5 or 6, a similar band behind the middle,
extending in the typical form to interval 4 and narrowing outwards, but this may
be more or less completely replaced by dark setae, whereas in southern (Tavoy)
specimens it may extend broadly to interval 8 ; underside with dense white
scales.
Head with shallow setigerous punctures which are denser behind the eyes ;
Irons narrow, with short erect whitish setae. Rostrum of £ shorter than the
pronotum (5:6), not dilated at the apex, of equal depth throughout, almost
straight, with rather dense white scales, dorsum compressed into a sharp ridge on
the basal half ; rostrum of $ longer than the pronotum (7 : 6), a little narrower,
bare except for a few pale setae at the base, with very fine sparse punctures and
a less distinct median carina. Antennae testaceous, inserted at two-fifths from
the apex of the rostrum in £ and a little further back in $. Prothorax transverse
(2 13), with the sides straight, the basal lobe densely squamose, not carinate,
partly concealing the scutelJum ; dorsum gently convex longitudinally, with
dense shallow ocellated punctures and no smooth median line. Scutellum small,
squamose. Elytra with deep, shallowly punctate striae, the intervals flat, shiny,
with sparse minute granules and 2-3 rows of setae. Legs black, with stripes of
white setae ; front femora with the tooth expanded into a lamina about twice as
long as deep, the distal end of which is sharply pointed and the proximal end
rounded, the shape varying somewhat ; posterior femora with the tooth normal
and sharp, the hind pair shortly exceeding the elytra.
Length 2-0 mm., breadth i-o mm.
Burma: Ruby Mines, 3^ (Doherty-type) ; Tenasserim, Tavoy, 2^ 1$;
Washaung, 20 km. E. of Myitkyina, 700 ft., 1$ 1$, vii. 1934 (Dr. R. Malaise) ;
Momeit, 1$ (Doherty).
Type in the British Museum.
Lobotrachelus frenatus sp. n.
$. Derm black, with dark setae above and lines and bands of narrow white
scales ; pronotum with a narrow white band at the base and a slightly wider one
just behind the apical margin, these joining up laterally with the broad white
area on the pleurae ; elytra without any basal band, but with a complete white
sutural stripe and another on interval 5 that reaches neither base nor apex, a
lateral white patch (divided by the striae) before the middle between stria 9 and
the margin, and two or three lines of separate white scales along the apical margin ;
underside with dense white scales.
Head with sparse pale setae and shallow punctures ; frons with two rows of
narrow suberect white scales. Rostrum of $ about as long as the pronotum, bare
except for a row of fine recumbent setae on each side, with sparse fine punctures
and a narrow median carina. Antennae inserted at a little beyond the middle of
the rostrum, testaceous. Prothorax transverse (3:4), with the sides straight, the
basal lobe squamose but not covering the scutellum ; dorsum gently convex
longitudinally, with dense ocellated punctures without any smooth median line.
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 447
Scutellum squamose. Elytra with deep, shallowly punctate striae, the flat
intervals shallowly rugulose and with two rows of setae. Legs as in
L. laminatipes sp. n.
Length 2-0 mm., breadth i-o mm.
Burma : Ruby Mines, i$ (Doherty).
Type in the British Museum.
Lobotrachelus bidens sp. n.
cJ$. Derm black, clothed above mainly with short dark setae ; prothorax
with a narrow band across the base formed of narrow white scales, these being
denser on the basal lobe, and the pleurae with a broad stripe of broader white
scales, the upper edge of which is sinuate ; elytra with a broad common basal
band of white on intervals 1-4, usually projecting further backwards on interval
1 ; underside with dense white scales.
Head with fine shallow punctures and sparse dark setae ; frons of <J with the
usual crest of erect white scales, which is reduced to a few setae in $. Rostrum of
<J a little longer than the pronotum, slightly curved, a little deeper at the
antennae (one-third from the apex) than near the base, the dorsum compressed
into a sharp median ridge and densely clothed with white scales nearly to the
apex ; rostrum of 9 somewhat longer and more slender, of equal depth throughout
and entirely devoid of scales. Antennae testaceous. Prothorax transverse (2:3),
with the sides straight, the basal lobe only partly covering the scutellum ; dorsum
gently convex longitudinally, with dense shallow ocellated punctures and no
smooth median line. Scutellum squamose. Elytra with deep, shallowly punctate
striae, the intervals slightly convex, with numerous granules and 2-3 rows of
setae. Legs black, with stripes of long white setae, the tarsi testaceous ; front
femora with two small teeth, one behind the other but not in the same line, the
posterior tooth at or behind the middle of the femur.
Length 1-7-2-0 mm., breadth 0-7-1-0 mm.
Burma : Karen Mts., i,^ (Doherty-type) ; Ruby Mines, 5000-7000 ft., i$
(Doherty) ; Kambaiti, 7000 ft., 29, vi. 1934 (Dr. R. Malaise).
Type in the British Museum.
Lobotrachelus plumbeus Mot. — Kambaiti, 4^ 10$, v-vi.
Neither Motchulsky nor Faust mentions the curious sexual dimorphism in
the teeth of the front femora.
Lobotrachelus rufitibiis sp. n.
cJ9- Derm black ; prothorax with sparse yellowish setae, a small spot of
narrow white scales on the median basal lobe and longer spots at the apex and
base of the pleurae ; elytra with lines of pale yellowish setae and a complete
sutural stripe of intercrossing white setae ; underside with dense white scaling.
Head of <$ with numerous fine yellowish setae, that of $ bare, shiny and
impunctate dorsally ; frons narrow, with two lines of suberect yellowish setae.
Rostrum of £ shorter than the pronotum (4 : 5), red, with a fine median carina,
two median lines of narrow white scales on the basal three-fifths and pale setae on
each side ; rostrum of 9 longer than the pronotum (6 : 5), more slender, widening
at the apex, with only sparse setae and no scales. Antennae testaceous. Prothorax
transverse (5 : 7), with the sides straight, the basal lobe concealing the scutellum ;
dorsum convex longitudinally in g and highest behind the middle, almost flat in
9, with very shallow ocellated punctures and no smooth median line. Scutellum
44§ SIR GUY A. K. MARSHALL
concealed. Elytra with deep, shallowly punctate striae, the intervals flat, finely
and sparsely subgranulate, many of them bearing (wholly or partly) only a
single row of setae and double rows elsewhere, the setae being rather stouter in $.
Legs with lines of white setae, the femora black, the tibiae and tarsi red ; all the
femora with a small sharp tooth.
Length 1-9 mm., breadth i-o mm.
Burma : Tenasserim, Tavoy, 1^ 1$ (Doherty).
Type in the British Museum.
It seems probable that this species is the same as that tentatively attributed
to angulatus Mot., by Faust [Ann. Mus. Stor. nat. Genova, 1895, 34 : 295), but
Motchulsky's species was described from a high altitude in Ceylon, and its
occurrence in Burma seems very unlikely.
Lobotrachelus parcus sp. n.
$. Derm black, with sparse grey setae ; prothorax with only a fringe of
single white scales in the posterior angles and no scales on the median lobe or in
the anterior angles ; elytra with lines of grey setae (mostly forming single rows
on the intervals) and a complete sutural stripe of interlocking grey setae ; under-
side with dense whitish scaling.
Structurally very close to L. rufitibiis, sp. n., the description of which applies
to it, except as follows : Head of $ somewhat more punctate on the vertex.
Rostrum black, red only at the extreme apex. Prothorax more transverse (5:8),
more sloping in front. Legs with the tibiae black, the anterior pairs red on the
apical third, the hind pair at the apex only.
Length 2-0 mm., breadth i-o mm.
Burma : 2$ [A. K. Weld Downing).
Lobotrachelus simplicipes sp. n.
cJ$. Derm black, with numerous short black setae and a few sparse longer
white setae ; prothorax with a spot on each side of the front margin formed of
separated white setae, a similar pair behind the middle, and behind these a pair
of transverse spots on the hind margin, a few white setae on the median lobe, and
the pleurae in <$ with separated grey scales throughout with a denser band along
the basal margin, in $ with black setae and only a spot of whitish scales in the
basal angle ; elytra with the following markings of sparse whitish setae : a
quadrate spot behind the scutellum, a small spot at the base of interval 3, an
abbreviated stripe (may be much reduced or absent) on 5 and 7, and a few isolated
setae elsewhere ; underside with fairly dense whitish scales.
Head with dense whitish scales on the frons in <$ and blackish ones in $.
Rostrum of J longer than the pronotum (4:3), straight from the base to the
antennae (three-eighths from the apex) then strongly deflected, much deeper at
the antennae than at the base, the dorsum compressed into a sharp median
ridge, with sparse narrow whitish scales ; rostrum of $ longer (5:3), regularly
curved, not deeper at the antennae (inserted at the middle), the dorsum not
compressed, with a low median carina and bare of scales. Antennae testaceous.
Prothorax transverse (2:3), with the sides almost straight from the base to near
the apex and there feebly sinuate, the basal lobe leaving the scutellum fully
exposed ; dorsum nearly flat longitudinally, with dense shallow punctures and no
smooth median line. Scutellum densely squamose. Elytra with deep, shallowly
punctate striae, the broad intervals flat and finely rugulose, with three-four rows
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 449
of dark setae but only two white setae where these are present. Legs black, the
tarsi reddish ; femora of <$ with whitish setae, the front pair without a tooth, tie
others with the tooth very small or rudimentary ; femora of $ with black setae
and only a dorsal patch of whitish setae on the posterior pairs, the tooth on the
posterior pairs small, that on the front pair rudimentary.
Length 2-4-1-0 mm., breadth 2-6-1-1 mm.
N.E. Burma : Kambaiti, 7000 ft., 4c? 2$, vi. 1934.
Lobotrachelus vacivus sp. n.
cJ$. Derm black, with thin grey scaling above, the elytra with -a large
subquadrate patch of dark setae before the middle between striae 1 and 4 and a
similar one on the declivity ; prothorax with dense greyish white scales on the
pleurae, the upper edge of the patch being ill-defined ; underside with dense
whitish scales.
Head with dense whitish scales on the frons in $ and darker ones in $.
Rostrum of <$ a. little shorter than the pronotum, gently and regularly curved, not
deeper at the antennae than at the base, the dorsum not compressed, with fairly
dense whitish scales almost to the apex ; rostrum of $ slightly longer and more
slender, bare, shiny and impunctate. Antennae testaceous, those of $ inserted only
slightly beyond the middle of the rostrum, those of $ at one-third from the apex.
Prothorax transverse (7 : 10), with the sides very slightly rounded, the basal
lobe not covering the scutellum ; dorsum gently convex longitudinally, with
dense shallow ocellated punctures without any smooth median line. Scutellum
densely squamose, entirely exposed. Elytra with deep, shallowly punctate striae,
the broad intervals gently convex, with minute low granules and two-four rows
of setae. Legs black with lines of whitish setae, only the base of the tarsi reddish ;
front femora with the tooth rudimentary in $, fully developed in $, tooth on
posterior pairs distinct in both sexes.
Length 2-5-2-7 mm., breadth 1-5-1-6 mm.
N.E. Burma : Kambaiti, 7000 ft., 4c? 3$, v-vi. 1934.
Lobotrachelus convexicollis sp. n.
cJ. Derm red-brown, with short dark setae ; prothorax with a dense fringe
of long white scales in the posterior angles and sparse whitish setae along the
anterior margin of the pleurae, and narrow white scales on the basal lobe ; elytra
with a short stripe of whitish setae at the base of interval 3 and sometimes on 2
also, and a common band of similar setae behind the middle extending to interval
4, but no white setae along the apical margin ; underside with separated whitish
scales.
Head bare on the vertex, with rather sparse punctures, the frons very
narrow, with a low ridge of dark scales. Rostrum of $ as long as the pronotum,
stout, widening from near the base to the apex, not deeper at the antennae
(inserted beyond the middle) than near the base ; dorsum compressed on the
basal half to form a sharp median ridge, with longitudinally confluent punctures
and sparse pale setae. Antennae testaceous, unusually short and stout, joints
4-7 of the funicle being transverse. Prothorax transverse (2 : 3), with the sides
straight ; dorsum strongly convex longitudinally, with small simple punctures
which are narrower than the intervals between them and with a transverse
basal impression on each side, without any smooth median line. Scutellum
partly covered by the pronotal lobe, bare. Elytra with deep striae, the intervals
45° SIR GUY A. K. MARSHALL
flat and rugulose, with three-four rows of setae. Legs black, with the usual white
setae, the tarsi red-brown ; posterior pairs of femora with a small sharp tooth,
the front pair without a tooth.
Length 2-3 mm., breadth i-o mm.
Burma : Ruby Mines, 2^ (Doherty).
Type in the British Museum.
Lobotrachelus urenae Mshl. — Sadon, 4000 ft., i<$ 1$, vi-vii. It is possible that
this species may prove to be synonymous with leucaspis, Fst.
Lobotrachelus laporteae Mshl. — Sadon, 1$, vi-vii.
Lobotrachelus 5 spp. — Kambaiti, five specimens, iv-vi.
Lobotrachelus nudus sp. n.
<J$. Derm black, almost devoid of setae above and without definite white
markings ; underside with rather thin grey scaling laterally and setae in the
middle.
Head bare, with small separated punctures ; frons with a low carina without
scales. Rostrum of $ as long as the pronotum, gently curved, stout, not dilated
apically, not deeper at the antennae (two-sevenths from the apex) than near the
base ; dorsum strongly convex transversely with a complete narrow median
carina, bare, with punctures that are subconfluent longitudinally ; rostrum of $
of the same length, more slender, nearly straight and more finely punctate.
Antennae testaceous, rather short and stout, joints 4-7 of the funicle transverse.
Prothorax transverse (3 : 4), gently rounded laterally, the basal lobe unusually
short ; dorsum gently convex longitudinally, sloping anteriorly, very shiny,
with numerous small simple widely-spaced punctures and a smooth median line,
entirely bare, except for a few short white setae on the basal lobe and a short
fringe of single narrower white scales in the basal angles ; pleurae with the
punctures rather larger than those on the disk and bare except for a few white
scales on the anterior margin. Scutellum entirely exposed, bare. Elytra unusually
narrow at the shoulders, being there only a little wider than the prothorax, with
deep remotely-punctate striae, the broad intervals flat, bare and impunctate, except
for a few white setae at the base of intervals 2 and 3, a few on 2 behind the middle,
and numerous punctures and minute dark setae throughout interval 1. Legs
black, with sparse white setae, the tarsi red-brown ; the tooth on the anterior
pairs of femora comparatively long and sharp, that on the hind pair much
smaller.
Length 2-5-3-0 mm., breadth 1-0-1-5 mm-
Assam : Manipur, 2<J (Doherty-type) . N.E. Burma : Sadon, 4000 ft., r$,
vi-vii. 1934 (Dr. R. Malaise).
Type in the British Museum.
Subfamily Zygopinae.
Metialma suturella sp. n.
(J$. Derm black ; prothorax with a rhomboidal patch of sparse narrow pale
scales in the middle of the basal half, enclosing a median stripe of dense yellowish-
white broader scales, the whole surrounded by four coalescing black spots, the
lateral areas variegated with sparse narrow pale scales enclosing an ovate black
spot at about the middle and a smaller one in front of it, the posterior angles with
a spot of dense yellowish-white scales, the pleurae with dense uniform whitish
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 451
scales ; scutellurri with dense yellowish scales ; elytra variegated with white,
yellowish, brown and blackish, narrow scales, the suture with a stripe of dense
yellowish scales from the base to beyond the middle, followed by a black spot
and then an apical white one of about the same length ; underside with dense
whitish scaling, with a large quadrate black spot at the base of the metepisterna
and a smaller one adjoining it on the metasternum, the usual black median area
on the venter extending on to ventrite 2.
Rostrum of $ black on the basal half and red from the apex almost or quite
to the antennae, the basal part unevenly clothed with rather dense whitish
scales ; rostrum of $ with the red area shorter, from the apex about half-way to the
antennae, the scales on the basal half yellowish, narrower and much sparser ; the
basal part rugulose and five-carinate in both sexes. Antennae red, often rather
darker distally, inserted somewhat beyond the middle of the rostrum in both
sexes ; funicle with joint 1 about as long as 2 and 3. Prothorax somewhat
transverse, widest at the base and narrowing with a slight curve to the very
shallow apical collar, which is much deeper dorsally, the basal angles rounded,
without any smooth median line. Elytra oblong-ovate, gradually narrowing from
the shoulders to near the apex and then abruptly narrowed ; the transverse basal
and apical impressions deep, the suture depressed in the basal third, and interval
2 strongly tilted inwards basally. Legs with a small sinuation on the front
femora just beyond the tooth ; tibiae with the apical half red, the front pair with
the dorsal edge almost straight on the apical three-fourths. Pygidium with
rather sparse fine setae and with whitish or yellowish scales on the lateral margins
only, and a median carina in both sexes ; ventrite 5 with a tuft of setae on
each side.
Length 3-0-4-0 mm., breadth 1-5-2-0 mm.
N.E. Burma : Kambaiti, 31^ 16$, iv-vi. 1934.
Metialma mimica sp. n.
cJJ. Extremely close to M. suturella, sp. n., the description of which applies
to it except in the following particulars : — Rostrum rather stouter and much more
strongly curved, entirely black in both sexes, except for the red mandibles.
Antennae with joint 2 of the funicle about as long as 1. Prothorax slightly more
transverse, with the pale spot in the hind angles rather larger. Legs with the tooth
on the front femora smaller and much less sharply pointed. Pygidium with
dense yellowish scales and setae.
Length 4-0-4-5 mm., breadth 2-0-2-2 mm.
N.E. Burma : Kambaiti, 7000 ft., 11^ 15$, v-vi. 1934.
Metialma saeva Fst. — Kambaiti, 11^ 7$, iv-vi ; Sadon 1$, vi-vii.
Metialma ignorata Fst. — Punkaung, between Sadon and Myikyina, 1^, vii.
Metialma 7 spp. — Kambaiti, 7 exx., v— vii.
Metialma 2 spp. — Sadon, 2 exx, vi-vii.
Metialma sp. — Taunggyi, 1$, viii-ix.
Metialma cordata sp. n.
(J$. Derm black ; prothorax with the large spot in the middle of the basal
half broadly ovate, formed of narrow yellowish scales and enclosing behind an
obovate spot of much denser broader yellowish-white scales, the whole surrounded
by four subrotund black spots that are sometimes separate but more usually
fused together, the remaining surface with separated narrow light brown and
452 SIR GUY A. K. MARSHALL
whitish scales enclosing an oval lateral black spot behind the middle and a
smaller round one in front of it ; scutellum with dense whitish or pale yellow
scales ; elytra with mixed white, yellowish and brown scales and variable
indefinite dark spots, without any distinct sutural stripe, except for an elongate
white spot at the apex and a much shorter black spot in front of it ; underside with
dense whitish or yellowish scales, with a black spot at the base of the metepisterna,
and the median black spots on the venter extending on to ventrite 2 but often
bearing scattered pale scales, especially in <$.
Rostrum longer than the pronotum (10 : 7) in both sexes, red from the
antennae to the apex (<J) or only narrowly at the apex ($), rugulose and five-
carinate from the base to the antennae in both sexes, and here with fairly dense
whitish scaling in <J and with thinner grey or brownish scaling in $. Antennae
entirely red, inserted well beyond the middle of the rostrum in both sexes ;
funicle with joint 1 slightly longer than 2. Prothorax somewhat transverse,
broadest at the base, gradually narrowing to the middle with the sides straight,
then curving in to the apex where there is a shallow collar, which is rather deeper
dorsally, the basal angles rounded ; the dorsum gently convex longitudinally,
but in the centre of the disk the scales are convergently suberect, so that in lateral
view the dorsum may appear to be subconical in the middle. Elytra strongly
cordate, rapidly narrowing from the broad shoulders in an even curve to the apex,
the transverse apical impression being very shallow and short and the posterior
calli feeble, the transverse basal depression broad and distinct, the suture
moderately depressed in the basal third and interval 2 somewhat tilted inwards
basally. Legs without any sinuation beyond the tooth of the front femora ; front
tibiae strongly curved dorsally from the base nearly to the apex. Pygidium
setose in the middle and with broader yellowish scales at the sides ; a median
carina on the basal half in $, which is more or less obsolete in $ ; ventrite 5 with a
broad triangular projection on each side in $ only.
Length 3-o-3-5 mm., breadth i^-i-g mm.
India : Kumaon, Ranikhet, 6^ 6$ (H. G. Champion) ; Almora, Khaula,
4500 ft., 3(J 3$ (H. G. C.-type) ; Darjeeling, Gopaldhara, 3400-4700 ft., 12^ 5$,
iv-vi. 1914, vi. 1916, x. 1917, iv-xi. 1918 (H. Stevens) ; Tista Valley, Gielle
Khola, 600 ft., 1$, vi. 1919 (H. S.).
Burma : Kambaiti, 7000 ft., 3$ 1$, iii-v. 1934 (Dr. R. Malaise) ; Momeit,
i(J (Doherty) ; Tenasserim, Tavoy, 2$ (Doherty).
Metialma jejuna sp. n.
cJ$. Derm black, with fairly dense, irregularly mingled, white, yellow and
dark brown scales ; prothorax with a large circular spot of such scales in the
middle of the basal half, partly enclosing behind a spot of much denser
yellowish-white scales on the basal lobe ; this circular spot surrounded by an
almost bare black quadrifid area (formed by the fusion of four spots), the rest of
the surface with variegated scales enclosing a small oval dark spot in the middle
of the dorso-lateral margin, a small yellowish spot in the basal angles, the lower
half of the pleurae with denser broader yellowish-white scales, and the scutellum
with dense similar scales ; elytra variegated throughout, without any definite
sutural marking except an elongate whitish spot at the apex ; metepisterna with
a subquadrate dark spot on the basal third, the rest with dense whitish scaling ;
venter mostly with separated pale yellowish scales, the usual dark median area
on ventrites 3 and 4 broad and extending on to 2.
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 453
Rostrum similar in the two sexes, only slightly longer in §, black with the
apex narrowly red, rugosely punctate, five-carinate and sparsely squamose from
the base to the antennae. Antennae inserted beyond the middle of the rostrum in
both sexes, red with the club fuscous ; funicle with joint 1 slightly longer than 2.
Prothorax slightly transverse, widest at the base, narrowing with a slight curve to
the apex, with a shallow apical collar which is deeper dorsally, the basal angles
rounded, sometimes with a narrow irregular abbreviated smooth median line.
Elytra rather oblong-ovate, narrowing gradually from the oblique shoulders,
the transverse apical depression rather deep, the transverse basal depression
broad and distinct, interval 1 depressed in the basal third and 2 tilted inwards
there. Legs with a small sinuation on the front femora just beyond the tooth ;
front tibiae with the dorsal edge almost straight on the apical three-fourths.
Pygidium with broader whitish scales laterally and a narrow setose area in the
middle about the median carina, which is distinct in both sexes ; ventrite 5 with
a small tuft of setae on each side.
Length 4-5-50 mm., breadth 2-0-2-2 mm.
N.E. Burma : Kambaiti, 7000 ft., 3^ 2$, v. 1934.
Closely allied to M. saeva Fst., but apart from the characters mentioned in
the key, that species differs in having the spot in the middle of the pronotum
elongate, the dark area round it having its outer margins straight and not lobate,
and the suture is less depressed at the base.
Metialma levirostris sp. n.
cJ9- Derm black, with pale brown and grey setiform scales intermixed ;
prothorax with a transverse black band on each side of the base, two small round
black spots a little in front of the middle of the disk and a very indistinct one
just behind and between them, a yellow to whitish spot in the basal angles, and
an elongate spot of dense white scales in the middle of the base ; scutellum with
dense white scales ; elytra with a small white sutural spot close to the apex and
blackish scales before and behind it and a small elongate spot of dense white
scales on the margin opposite the hind coxae ; pygidium, metasternum and venter
(except the usual black median area on ventrites 3 and 4) with dense white scales,
and a quadrate black spot on the base of the metepisterna, the mesepimera with
sparse light brown scales and a few white or yellowish scales in the upper angle.
Rostrum of $ a little longer than the pronotum (8 : 7), black, opaque and
strongly punctate nearly to the apex, with three fine carinae and pale scales on
the basal half ; that of $a little longer (9:7). smooth and shiny, almost impunctate
and bare except close to the base, without any carinae. Antennae entirely red,
inserted at (<$) or behind the middle of the rostrum ($) ; joint 1 of the funicle as
long as 2 and 3. Prothorax somewhat transverse, parallel-sided in the basal half,
then rapidly narrowing in a curve to the apex, which is not constricted, the
basal angles subrectangular ; dorsum moderately convex longitudinally, highest
at a little beyond the middle, only shallowly depressed at the apex, without any
smooth median line. Elytra rather narrowly ovate or subelliptical, the shoulders
being much less prominent than usual, the broad transverse basal impression
shallow and sometimes obsolescent, the posterior calli distinct, interval I
depressed only quite close to the scutellum, 2 flat near the base. Legs with the
front femora sinuate on the lower edge just beyond the tooth ; front tibiae
strongly curved dorsally on the basal two-thirds. Pygidium densely clothed with
454 SIR GUY A- K- MARSHALL
broad white scales in both sexes ; ventrite 5 with a triangular projection on each
side of the pygidium in $ and a narrower one in $.
Length 3-0-3-5 mm., breadth 1-4-1-5 mm.
India : Manipur, 2^ (Doherty-type) .
Burma : Kambaiti, 7000 ft., ig 3$ (Dr. R. Malaise) ; Ruby Mines, ig 2$
(Doherty).
Type in the British Museum.
This is the only species of the genus known at present in which the rostrum
of the $ is smooth throughout.
Key to the Burmese Species of Metialma.
1 (14). Each elytron with a transverse depression at the apex and an
obtuse callus in front of it ; rostrum and tibiae never entirely
red.
2 (3). Metepisterna without any blackish basal spot ; elytra with a
whitish sutural stripe from base to near middle ; antennal club
black ........ saeva Fst.
3 (2). Metepisterna with a large subquadrate blackish spot at base.
4 (7). Elytra with a sutural stripe of dense yellowish scales from base to
beyond middle ; tibiae with the apical half red ; antennal club
red.
5 (6). Rostrum of $ red from apex nearly to antennae, that of 9 red for
only half that distance ; pygidium sparsely setose with only a
few whitish scales on the lateral margins . . suturella sp. n.
6 (5). Rostrum of both sexes stouter, more strongly curved, black with
only the mandibles red ; pygidium with dense yellowish scales
and setae ........ mimica sp. n.
7 (4). Elytra without any distinct sutural stripe ; tibiae black, rarely
with the extreme apex red.
8 (13). Antennae inserted beyond middle of rostrum in both sexes ;
rostrum of $ coarsely punctate and carinate from base to
antennae ; elytra without a spot of dense white scales on the
lateral margin ; apex of femora black.
9 (10) . Front tibiae straight dorsally on the apical three-fourths ; elytra
■ oblong-ovate ; antennal club fuscous ; ventrite 5 of <$ with the
external angles not projecting beyond the pygidium jejuna sp. n.
10 (9). Front tibiae strongly curved ; elytra more cordate ; antennal club
red ; ventrite 5 of <$ with the external angles projecting well
beyond the pygidium.
11 (12). Elytra more abruptly narrowed at the apex, the blackish area on
the suture in front of the apical white spot as long as or longer
than the latter ; rostrum of $ red at apex only ; ventrite 5 of $
produced on each side into a narrow tufted process ; pygidium
of 9 with a complete median carina ; tarsal claws simple
ignorata Fst.
12 (11). Elytra more regularly narrowed at apex in an even curve, the
blackish subapical spot on the suture much shorter than the
white apical spot ; rostrum of <$ red from antennae to apex ;
ventrite 5 of $ with a broad triangular process on each side ;
pygidium of $ with only a short carina at the base ; claws with
a minute basal tooth ...... cordata sp. n.
BURMA AND BRITISH INDIA — COLEOPTERA I CURCULIONIDAE 455
13 (8). Antennae inserted at (rf) or behind ($) middle of rostrum ;
rostrum of $ smooth, without carinae, and simply punctate
near base ; elytra with a spot of dense white scales on the
margin opposite the hind coxae ; apex of femora red levirostris sp. n.
14 (1). Elytra without any apical depression or subapical callus ; rostrum
and tibiae entirely red in both sexes ; metepisterna without any
basal black spot ; ventrite 5 of $ with a small conical projection
on each side (<J unknown) ..... ruj irostris Fst.
Subfamily Ceuthorrhynchinae.
Mecysmoderes armirufus sp. n.
^$. Derm dull black, the underside of the head, the sides and lower surface
of the prothorax, and the shoulders of the elytra (broadly in 9. more narrowly in
<J), red ; prothorax with the black discal area almost bare, the red lateral area
thinly clothed with yellowish setiform scales, and the pleurae with a broad stripe
of densely overlapping broader cream-coloured scales ; elytra with the black
parts bare, the red shoulders with sparse yellowish setiform scales, and the edge of
the suture with a single row of dense small broader whitish scales, which become more
numerous and sparser beside the prothoracic spine and along the basal margin ;
mesepisterna and mesepimera black with a narrow border of white scales ;
metepisterna black with a posterior border of white scales and a large black area
adjoining it on the metastemum, the rest of which is covered with dense white
scales ; venter with dense white scales, except for a large round black area at the
side of ventrites 1 and 2 ; pygidium black.
Head deeply depressed between the eyes, with large reticulate punctures and
sparse pale setae. Rostrum of £ as long as the elytra along the suture, strongly
curved, slender, bare, somewhat flattened and obsoletely punctate dorsally with
a complete narrow median carina, the lateral areas tricarinate behind the
antennae and with longitudinally confluent punctures in front ; rostrum of $> of
the same width but very nearly as long as the whole body, and the median carina
confined to the basal third. Antennae red, inserted at a little behind the middle
of the rostrum {£) or at about one-third from the base ($) ; scape produced at its
apex into a short transparent lamina, with two or three setae beside it ; funicle
with joints 2 and 3 equal and very long, longer than 1 or than 4-6 together.
Prothorax with the sides shallowly sinuate in the apical half (as seen from above) ,
the anterior part of the dorsum being strongly compressed laterally, the apical
margin arcuate dorsally, the basal spine two-sevenths the length of the suture of
the elytra ; dorsum somewhat convex longitudinally, with strong reticulate
punctures, a complete low median carina and a short carina on each side of it in the
apical depression. Elytra about as broad as long, widest at the broadly rounded
shoulders, jointly subtruncate at the apex, broadly depressed transversely at the
base (but the basal margin not elevated) and with a shallow transverse depression
behind the middle ; the narrow deep striae bare, with catenulate punctures ; the
intervals about twice as broad as the striae, each with a single row of small granules,
these being irregularly duplicated on interval 5, which is slightly higher than the
others, intervals 8 and 10 broadly depressed behind the shoulders ; the granules
bearing an inconspicuous recumbent dark seta. Legs black, with the front pair
entirely and the tarsi of the posterior pairs, red ; femora rugosely punctate, with
456 SIR GUY A. K. MARSHALL
sparse pale setae, the hind pair with a broad band of white scales ; the dorsal
fringe of the hind tibiae as long as the first tarsal joint. Sternum with the proster-
nal furrow twice as wide as the space between the front coxae ; mesosternum not
impressed.
Length 3-5-4-5 mm., breadth 2-5-3-0 mm.
N.E. Burma : Kambaiti, 7000 ft., 8^6$, iv-v. 1934.
Allied to M. stylicomis Mshl. 1934, which differs considerably in colour and
scaling ; further, the rostrum is much shorter, joint 3 of the funicle is much
longer than 2, the granules on the elytra are numerous and irregular, the prosternal
furrow is as wide as the space between the front coxae, etc.
Mecysmoderes levipes sp. n.
(J$. Derm black, the rostrum (except the base) and the apical margins of
the prothorax and elytra, red ; prothorax with a few pale setae on the disk and
an ill-defined curved lateral stripe formed of transverse pale setae, the prosternum
with dense whitish scales ; elytra rather thinly clothed with narrow dark scales,
interval I with a velvety stripe of dense black scales beside the prothoracic
spine and a much shorter whitish or yellowish spot immediately behind it, a
small whitish spot at the base of interval 2 and a longer one at the base of 6, and
sparse irregular narrow whitish scales scattered on the disk, interval 8 with a
small spot of broad white scales behind the shoulder and a longer one at its apex ;
side-pieces of mesosternum black with a narrow margin of pale scales, metepis-
terna black with an apical band of pale scales, metasternum and venter with
dense white scales, pygidium black.
Head with reticulate punctures and sparse pale setae ; frons deeply depressed
between the eyes. Rostrum a little longer than the head and pronotum (without
the spine) in both sexes, with the antennae at the middle, the basal half with
confluent punctures and three fine carinae, the apical half a little narrower with
fine sparse punctures in $, impunctate in $. Antennae red ; scape not produced
at the apex but with two short setae ; funicle with joints 1 and 3 equal, 2 a little
longer, 4 and 5 longer than broad, 6 globular. Prothorax feebly rounded laterally,
with a shallow apical collar (deeper dorsally), the apical margin angulate in the
middle, the basal spine one-fourth the length of the suture ; dorsum moderately
convex longitudinally, with dense reticulate punctures, the margins of which
tend to form irregular longitudinal carinulae, with a narrow median carina and
a short carina on each side of it anteriorly. Elytra a little longer than broad,
shallowly depressed at the base, the striae much narrower than the rugosely
granulate flat intervals, which are of equal height. Legs with the femora edentate,
black to piceous, the apices narrowly red, the basal two-thirds with sparse
whitish setiform scales ; tibiae and tarsi entirely red. Sternum with the
prosternal furrow wider than the space between the front coxae ; mesosternum
not impressed.
Length 2-5 mm., breadth 1-5 mm.
N.E. Burma : Kambaiti, 6$ 10$, v-vi. 1934.
Of the Indo-Burmese species with unarmed femora the nearest is M.
nigroru/us, Mot., which apart from its colouring differs in having the frons very
shallowly depressed ; the pronotum is much less compressed anteriorly, without
the lateral carinae ; the elytra are much narrower at the shoulders, which are
BURMA AND BRITISH INDIA — COLEOPTERA ! CURCULIONIDAE 457
obliquely rounded, and they are not depressed at the base, the margins of which
are much more oblique ; and the mesosternum is impressed to receive the apex
of the rostrum.
Mecysmoderes spp. — Kambaiti, 2$ 1$, v-vi.
Ceuthorrhynchus 2 spp. — Kambaiti, i$ i^, v.
Ceuthorrhynchus sp. — Sadon, i£, vi-vii.
Subfamily Baridinae.
Baris tumida sp. 11.
<$. Derm dull black, bare.
Head separated from the rostrum by a distinct impression, with numerous
shallow punctures but no median fovea. Rostrum as long as the pronotum, rather
stout, strongly curved, not widened at the apex ; the lateral areas very coarsely
and confluently punctate, with indications of two irregular carinae, the dorsum
with uneven fine punctures which become a little larger at the base, without any
median carina. Antennae inserted at about one-fourth from the apex of the
rostrum, black, with the basal half of the scape red ; funicle widening distally,
joint 1 slightly longer than 2 and 3, 4-7 transverse. Prothorax nearly as long as
broad, parallel-sided from the base to the middle, then gradually narrowing to
the deep abrupt subapical constriction, which is continued more shallowly across
the disk, the apex truncate, the gular margin rather deeply sinuate in the middle ;
dorsum rugosely sculptured with irregular strong curved ridges, with an
abbreviated median carina and a large shallow impression on each side at about
the middle. Scutellum small, almost round, smooth with two very shallow
impressions, and tilted forwards. Elytra very rugose and uneven, subovate,
with the sides sinuous (as seen from above), wider at the somewhat sloping
shoulders than the base of the prothorax, and rather broadly truncate at the
apex ; the deep striae partly sinuous and containing distant punctures ; the
broad intervals very uneven and coarsely punctate, interval 3 with an elongate
tubercle near the base and a similar one behind the middle, 4 with a smaller
tubercle adjoining the former, 5 with a small tubercle behind the middle and a
larger one on the declivity, a prominent subapical tubercle at the junction of
intervals 3 and 8, and some small ones on 7 and 9. Legs black, with the tarsi red-
brown ; femora coarsely punctate, with short recumbent pale setae ; tibiae not
gradually widening distally, shallowly bisinuate on the lower edge, the front pair
with a small mucro ; tarsi with joint 4 more dilated than usual.
Length 4-0 mm., breadth 17 mm.
N.E. Burma : Kambaiti, 7000 ft., 2$, v. 1934.
A very distinct species owing to the tuberculate elytra and carinulate
pronotum.
Baris aspersa sp. n.
(J. Derm black to piceous ; prothorax with a lateral stripe of sparse narrow
yellow scales on each side and a few scattered scales on the disk ; elytra with
numerous scattered groups of a few narrow yellow scales, the largest being
behind the middle on interval 3, at the base of which is a short stripe of dense
scales ; underside bare, or with a minute pale seta in each puncture.
Rostrum about as long as the pronotum, stout, bent at the base, strongly
curved dorsally, parallel-sided throughout, with coarse confluent punctures and
an abbreviated smooth median line. Antennae inserted at one-fourth from the
apex of the rostrum, piceous, except the narrow part of the scape which is red ;
458 SIR GUY A. K. MARSHALL
funicle with joint 2 as long as broad, 3-7 strongly transverse. Prothorax transverse
(8 : 11), widest at the base, narrowing from there to two-thirds, with the sides
straight and the apical collar rather abruptly constricted, the base shallowly
bisinuate ; dorsum with strong separated punctures, which become larger
laterally, without any smooth median line. Scutellum small, subtriangular, bare.
Elytra broadly ovate, much wider at the sloping shoulders than the base of the
prothorax ; the deep narrow striae with indistinct distant punctures, except in
the basal third of stria 1 and the basal fourth of 2 where the punctures are large
and close, stria 10 disappearing behind the middle ; the disk without impressions,
the posterior calli obtusely prominent, the broad flat intervals with a rather
irregular row of punctures, which are larger and closer on the basal half, becoming
small and widely separated behind, those on 3, 5, 7 more or less duplicated. Legs
red-brown, with small pale setae in the punctures ; femora rather coarsely
punctate ; tibiae not widening distally, shallowly sulcate, the front pair not
mucronate.
Length 3-5-4-0 mm., breadth 1-5— 1-7 mm.
N.E. Burma : Sadon, 4000 ft., 5<J, vi-vii. 1934.
A somewhat aberrant species owing to the undilated tibiae, the abbreviation
of stria 10 on the elytra, the unequal punctation in striae 1 and 2, and the well-
developed posterior calli.
Baris spp. — Kambaiti, six specimens of five species, iv-vi.
Baris spp. — Sadon, three specimens of three species, vi-vii.
Baris spp. — Taunggyi, two specimens of two species, viii-ix.
Baris sp. — Tenasserim, Sukli, one specimen, x.
Baris blennus sp. n.
^9- Derm dull black, with dark recumbent setae and a row of distant
narrow white scales on the alternate intervals of the elytra ; underside with a
small narrow white scale in each puncture.
Head smooth, with numerous small separated punctures. Rostrum a little
longer than the head and pronotum in both sexes, opaque and with dense
punctures throughout in $, more shiny and with rather smaller punctures in $ ;
lower surface with a small median tooth near the base. Antennae piceous with
the base of the funicle paler, inserted at one-sixth ($) or one-fifth ($) from the
apex of the rostrum ; funicle with joint 2 longer than broad, 3-7 transverse.
Prothorax a little broader than long, parallel-sided from the base to the middle,
then narrowing in a curve to the abrupt apical constriction, the base shallowly
bisinuate ; dorsum with dense confluent punctures, the intervals forming sinuous
longitudinal ridges, with sparse narrow pale scales. Scutellum small, triangular,
bare, with a median impression. Elytra broadly ovate, wider at the sloping
shoulders than the prothorax, with a faint transverse impression at one-fourth
from the base ; the deep striae with very shallow distant punctures, stria 10
complete ; the intervals broader than the striae, flat, with an irregular row of
shallow punctures, most of which are nearly as wide as the interval, and each
containing a recumbent brown seta, the single narrow white scales on the
alternate intervals widely separated. Legs red-brown, with sparse short white
setae ; femora coarsely punctate ; tibiae not widening distally. Pygidium of £
convex, with small separated punctures, that of $ depressed and rugosely
punctate on the apical two-thirds.
Length 3-0-3-5 mm., breadth 1-2-1-5 mm.
N.E. Burma : Kambaiti, 7000 ft., 11$ 5$, v-vi. 1934.
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 459
Athesapeuta gyrosicollis sp. n.
$. Derm black, the rostrum, antennae and legs red-brown ; prothorax
clothed with yellowish-white to pale yellow scales, with a large median area from
the base to two-thirds apparently bare, but really sparsely covered with dark
narrow inconspicuous scales except for an irregular median stripe of pale scales ;
elytra with a broad basal band of yellowish scales from interval 2 to 9 and an
equally broad one behind the middle (leaving interval 1 bare), these being
united in perfect specimens by rather sparser scales on intervals 5-9, and with
sparse scales about the apex ; underside with subcontiguous yellowish scales,
which are much denser and overlapping on the mesepimera and metepisterna.
The dorsal scales are easily abraded, so that the prothorax often appears to have
only a broad lateral stripe on each side and the elytra only two widely separated
bands.
Rostrum a little longer than the pronotum, rather strongly curved, sub-
cylindrical, gradually widening from the antennae (inserted at one-fourth from
apex) to the apex, with sparse small punctures, which form four dorsal lines and
two less regular lateral ones on each side. Antennae with joint 1 of the funicle as
long as 2-f-3-}-4, 5-7 transverse. Prothorax a little broader than long,
widest close to the base and narrowing with a strong curve to the apical collar ;
dorsum convex longitudinally, highest behind the middle, the small punctures
all longitudinally confluent, forming numerous curved ridges, which become
somewhat coarser on the pleurae ; the narrow oblong scales all lie transversely and
do not overlap. Scutellum transverse, suboblong, broadly depressed in the middle,
and bare or with a few short setae. Elytra a little wider at the sloping shoulders
than the base of the prothorax and gradually narrowing thence to the broadly
rounded apex ; the dorsum without impressions but with deep impunctate striae
that show through the scales, when these are not disarranged, the broad flat
intervals with irregular, transversely wrinkled punctures. Legs with non-con-
tiguous scales, which on the femora are short at the base and much longer at the
apex, except on the hind pair on which they are long and dense throughout.
Length 47-5-5 mm., breadth 2-2-2-6 mm.
S. Shan States : Taunggyi, 5000 ft., 3<J, viii-ix. 1934 (type) ; Pekkong,
3000 ft., ig, x. 1934.
This insect is distinguished from all previously described Athesapeuta by the
vermiculate sculpture of the prothorax.
PHRISSODERES gen. 11.
Head separated from the rostrum by a shallow depression. Rostrum strongly
bent downwards near the base, comparatively slender, wider beyond the antennae,
not deeper than broad at the base, nor narrowed dorso-ventrally at the apex,
without carinae ; scrobes passing below the rostrum before the middle and
uniting towards the base ; mandibles dentate, interlocking, with a small angle
externally. Antennae inserted at one-fourth from the apex of the rostrum in both
sexes ; scape slender, moderately clavate, with a few sparse setae ; funicle only
slightly widening distally, joint 1 much longer than 2, /distinct from the club. Protho-
rax transverse, shallowly bisinuate at the base, without postocular lobes. Elytra sub-
triangular, much wider at the shoulders than the prothorax, with ten complete deep
striae and obtuse posterior calli. Legs slender; femora only slightly clavate, not sulcate
or toothed beneath ; tibiae not dilated distally, sulcate and without a mucro ;
tarsi with joint 3 deeply bilobate, 2 longer than broad, the claws long, slender
460 SIR GUY A. K. MARSHALL
and divergent. Sternum with a very large deep subhexagonal excavation on the
presternum, which is truncate behind, the gular margin sinuate ; mesosternum
deeply depressed, the intercoxal process short, vertical and much broader than
a coxa. Abdomen with the pygidium entirely concealed, the two basal ventrites
fused throughout, ventrite 2 much longer than 3-7-4.
Genotype : Phrissoderes costalis, sp. n.
A very distinct genus characterised by the very large cavity on the prosternum
the wide vertical mesosternal process, and the anteriorly compressed pronotum,
which is entirely covered with costulae radiating in all directions from the middle
of the base.
Phrissoderes costalis sp. n.
(J$. Derm uniform shiny black, entirely bare.
Rostrum very similar in the two sexes, rather slender ; the subcylindrical
basal part in <J with two rather strong rows of punctures dorsally, the apical part
with stronger confluent punctures, the lateral areas with coarse irregular furrows
and costae ; rostrum of $ with the dorsal punctures on the basal area minute,
but those on the apical part as coarse as in $. Antennae red-brown, with the
club fuscous ; funicle with joints 3-7 transverse, the club large, as long as joints
2-7. Protkorax narrowing from the base to beyond the middle, with the sides
almost straight, then curving in rapidly to the apical collar ; dorsum much more
strongly compressed in front in $ than in $, with branching costulae radiating in
all directions from the middle of the base, and similar coarser sculpture on the
pleurae. Scutellum with a broad median depression in $, smaller and almost
flat in <$. Elytra subtruncate at the base, parallel or very shallowly sinuate from
the shoulders to one-third, then rapidly narrowing to the apex, where they are
separately rounded, the disk with a shallow transverse impression at one-fourth
from the base ; in $ the broad deep striae contain small distant punctures, each
puncture having a minute granule on each side of it, in $ the striae are much
broader and the punctures much larger ; the intervals in $ are broader than the
striae, flat, with a row of microscopic punctures (sometimes obsolescent), in ^ the
intervals are not broader than the striae and strongly costate. Legs black, with
sparse short appressed setae, the tarsi red ; femora with large close punctures.
Underside with round isolated punctures which become confluent towards the
sides of the metasternum and form longitudinal ridges ; venter with a deep
lateral fovea on the line of junction of ventrites 1 and 2.
Length 2-8-3-4 mm., breadth 1-4-1-7 mm.
N.E. Burma : Kambaiti, 7000 ft., i<$ 5$, v-vi. 1934.
It is very unusual among the weevils for the apex of the rostrum to be more
strongly punctured than the base, especially in the female.
Chelonebarus burmanus sp. n.
(J. Derm red-brown ; pronotum rather darker, with a broad lateral stripe
of dense oblong yellowish scales divided in the middle by a large transverse bare
patch, and with a few scales along the median line ; elytra with a loose patch of
similar scales at the base of intervals 2 and 3, and a few scales near the base of 10
behind the shoulder, and a few elongate narrow scales in the striae (these may
normally form continuous rows in the striae, as some seem to have been abraded).
Head mostly concealed within the prothorax, with close shallow punctures.
Rostrum 1-5 times as long as the pronotum, rather strongly curved, subcylindrical,
BURMA AND BRITISH INDIA — COLEOPTERA I CURCULIONIDAE 461
only slightly widened at the apex, closely and strongly punctate nearly to the apex,
with a complete fine low median carina and a regular row of close subquadrate
punctures on each side of it on the basal half, a fine irregular carina externally to
these, and the lateral areas with larger irregular subconfluent punctures.
Antennae inserted at one-third from the apex of the rostrum, red-brown ; funicle
with joint I a little longer than 2-|-3-|-4, 2 and 3 equal and slightly longer
than broad, 4-7 transverse. Prothorax transverse (2 : 3), widest at the base,
rapidly narrowing with a slight curve to the shallowly constricted apex, which is
about one-third the width of the deeply bisinuate base ; dorsum strongly convex
longitudinally, with coarse confluent punctation, the intervals forming sinuous
ridges, without any smooth median line. Scutellum oval, bare, with a median
impression. Elytra very broadly ovate, nearly" as broad as long (2-8 by 2-5 mm.),
much broader at the prominent shoulders than the base of the prothorax,
slightly widening behind the shoulders, then rapidly narrowing to the broadly
rounded apex ; dorsum comparatively flat transversely, highest near the base
and sloping uniformly from there to the apex, with a broad transverse depression
behind the shoulders and large obtuse conical posterior calli ; the striae narrow
and deep, containing shallow spaced punctures, except in the basal fourth of
striae 1 and 2 where the large close punctures quite obliterate the striae, stria 7
ceasing behind the humeral callus ; the broad flat intervals opaque and shagreened,
1-3 with large punctures on the basal third, elsewhere the intervals with a row
of minute shiny granules, the humeral calli also with small flat granules. Legs
red-brown, coarsely punctate, each puncture with a short stout white recumbent
seta, the femora with a row of short fine erect setae on the basal half of the
lower edge.
Length 3-0-3-9 mm., breadth 1-9-2-5 mm.
N.E. Burma : Kambaiti, 7000 ft., 2^, v-vi. 1934.
The discovery of this genus in Burma was unexpected, the only other
species (C. partimpunctatus Mshl. 1940) having been described from Kenya. It
differs in having a much shorter rostrum, the pronotum simply punctate, elytra
with much less prominent posterior calli and uneven punctation, etc.
PSILARTHRUS gen. n.
Head with fine sparse punctures, separated from the rostrum by a shallow-
impression. Rostrum varying in thickness and curvature, not compressed at the
base and not narrowed dorso-ventrally at the apex, without any sulci or carinae ;
scrobes passing completely below the rostrum near the base, not or very shortly
extending beyond the antennae ; mandibles dentate and interlocking, not
projecting when closed. Antennae inserted beyond the middle of the rostrum in
both sexes ; scape slender, abruptly clavate ; funicle not or but slightly widening
distally, joint 1 longer than 2, 7 distinct from the club. Prothorax with a collar-
like constriction at the apex, only shallowly bisinuate at the base. Elytra black,
shiny, with prominent shoulders and ten deep narrow striae containing catenulate
punctures, with posterior calli. Legs very slender ; femora moderately clavate,
not sulcate or toothed beneath ; tibiae narrow, subcylindrical, straight, not
widening distally, without any sulci, impunctate or with only obsolescent
punctures ; front or anterior pairs of tibiae sometimes mucronate in $, the hind
corbels open ; tarsi with joint 3 deeply bilobate, the claws divergent from the base.
Sternum without any prosternal sulcus, the front margin of the prosternum
truncate or sinuate, its hind margin without any median process ; mesosternum
novit. ZOOL., 42, 3. 1948. 84
462 SIR GUY A. K. MARSHALL
depressed, the side-pieces fused. Abdomen with the pygidium shortly exposed,
ventrite 2 as long as or longer than 3+4, more or less fused with 1 in the
middle.
Genotype : Psilarthrus dentipes sp. n.
Distinguished from the known Oriental genera of Baridini by the slender,
non-sulcate tibiae, a very unusual character in the tribe.
Psilarthrus dentipes sp. n.
(5*$. Derm shiny black, the antennae and tarsi, and the apex of the rostrum
in 9. reddish.
Head without a frontal fovea. Rostrum strongly curved, a little longer than
the head and pronotum in $ (10 : 9), still longer in £ (13 : 9), slightly widened in
the apical part ; dorsum with only a few sparse punctures near the base, those at
the sides much coarser and more or less confluent, the punctures smaller and
sparser in 9- Antennae red to red-brown, the club fuscous ; funicle with joint 1 as
long as 2+3, 5-7 transverse. Prothorax nearly as long as broad, widest at the
base, narrowing very slightly from the base to the middle and then curving in to
the apical collar, the gular margin truncate ; dorsum with small strong punctures
of varying density, some on the disk usually separated by more than their
diameters, and with a smooth median line ; the lateral areas with sparse narrow
whitish scales. Elytra narrowly ovate, very shallowly sinuate laterally behind
the shoulders and thence narrowing to the apex ; dorsum without discal
impressions but with the suture distinctly raised behind the middle, the
subapical calli well developed, and a slight elevation at the joint apex of intervals
3 and 8 ; the intervals broad, flat, shiny and impunctate, with widely spaced long
triangular whitish scales on 3, 5 and 7, which are apparently rather easily
abraded. Legs with the front coxae very narrowly separated, the space being no
wider than the narrowest part of the antennal scape ; femora with fine sparse
punctures containing a minute white seta, but more coarsely punctate at the
apex ; tibiae not widened at the apex, without a mucro in either sex, the front
pair in $ with a large sharp triangular tooth at a little beyond the middle of the
lower edge.
Length 3'3-4'0 mm., breadth 1-2— 1-5 mm.
N.E. Burma : Kambaiti, 7000 ft., 16J 13$, v-vi. 1934 (Dr. R. Malaise-type).
Assam : Manipur, i$ (Doherty).
Psilarthrus costicollis sp. n.
cJ$. Derm shiny black, the antennae and tarsi red.
Head with sparse minute punctures and no frontal fovea. Rostrum about
as long as the head and pronotum in both sexes, moderately stout, slightly less
so in $, parallel-sided from base to antennae (inserted at two-sevenths from
apex in $, three-tenths in §,), beyond which it is somewhat wider, the apical part
being parallel-sided in <§ and gradually widening distally in $ ; dorsum with very
fine sparse punctures, the sides coarsely punctate, being more confluent in $.
Antennae with joint 1 of the funicle slightly shorter than 2+3, 5-7 transverse.
Prothorax nearly as long as broad, very gradually narrowing from the base to
two-thirds or almost parallel-sided, the apical constriction rather abrupt, the
basal angles acute and very slightly projecting ; dorsum moderately convex
longitudinally, closely set with large strong punctures which are much wider than
the intervals between them, and with a low smooth median costa, which is usually
complete but sometimes abbreviated, and very sparse whitish setae. Elytra
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 463
ovate, the sides rather deeply sinuate behind the shoulders ; dorsum without
impressions, the subapical calli low and obtuse, the intervals broad, flat and
impunctate, with very sparse elongate white scales on 3, 5 and 7. Legs with
small separated punctures on the femora, but more coarsely punctate towards
the apex ; tibiae not widened at the apex, without a mucro or tooth in either sex.
Length 3-0-3-5 mm., breadth 1-4-1-5 mm.
Assam : Manipur, itf i$ (Doherty-type) . N.E. Burma : Kambaiti, 7000 ft.,
2(J 2$, v-vi. 1934 (Dr. R. Malaise).
Type in the British Museum ; cotypes in the Stockholm Museum.
Psilarthrus mucronatus sp. n.
J$. Entirely black and shiny, except for the red tarsal claws and tibial
unci ; upper surface entirely bare.
Head with fine sparse punctures and a shallow median fovea. Rostrum a
little longer than the head and pronotum (13 : 10) in both sexes, somewhat
widened beyond the antennae (at one quarter from the apex in £, a little further
back in 9), the apical part being parallel-sided in $ and gradually widening
distally in $, with small subconfluent punctures dorsally and rugosely punctate
laterally in $, the punctures smaller in $. Antennae with joint 1 of the funicle as
long as 2+3, 6 and 7 transverse. Prothorax very nearly as long as broad,
gently rounded laterally, widest behind the middle, the basal angles not projecting,
the apical constriction gradual and not abrupt ; dorsum moderately convex
longitudinally, closely punctate, without any smooth median line or costa, the
intervals narrower than the punctures, which are partly subconfluent on the
disk and much more strongly so laterally, so as to form curved ridges. Elytra
ovate, distinctly sinuate laterally behind the shoulders ; dorsum with a shallow
transverse impression on each side at one-fourth from the base and another
behind the middle, the subapical calli distinct, the intervals broad and flat,
devoid of scales and with a rather irregular row of very fine shallow punctures.
Legs with the space between the front coxae broader than the antennal club ;
femora with the punctures rather stronger and closer near the base, fine and
sparse distally ; front tibiae somewhat widened at the apex, with a strong mucro
in both sexes, the uncus and mucro being widely separated in ^ and closer in 9-
Length 4-0-4-5 mm., breadth 1-5-1-6 mm.
Burma : Ruby Mines, 1^2? (Dohert v-type) ; Kambaiti, 7000 ft., 1$ 2$,
v. 1934 (Dr. R. Malaise).
Type in the British Museum ; cotypes in the Stockholm Museum.
Psilarthrus coracinus sp. n.
$$. Derm entirely black, except for the red claws and tibial unci.
Head with fine separated punctures and a small frontal fovea. Rostrum of ^
half as long again as the head and pronotum, comparatively slender, bent at the
base and less curved distally than in the other species, somewhat widened and
parallel-sided beyond the antennae (inserted at one-third from the apex), the
dorsum more strongly and closely punctate than in the other species, more
coarsely punctate laterally ; rostrum of $ unusually long and slender, being
nearly twice as long as the head and pronotum, scarcely widened beyond the
antennae (inserted at only a little beyond the middle), the punctures much finer
and sparser. Antennae comparatively long and slender, with joint 1 of the funicle
as long as 24-3 (<J) or 24-34-4 ($), only 7 slightly transverse. Prothorax
nearly as long as broad, parallel-sided from the base to the middle, then narrowing
464 SIR GUY A. K. MARSHALL
with a curve to the shallow apical constriction ; dorsum almost flat longitudinally,
rather unevenly punctate, the intervals on some parts of disk being as wide as or
wider than the punctures, but without any smooth median line and entirely
without scales, the punctures on the pleurae much denser and coarser. Elytra
rather broadly ovate, almost parallel to the middle, with only a shallow
sinuation behind the shoulders and rather broadly rounded at the apex ; dorsum
with a shallow impression on intervals 3-5 at one-fourth from the base, the
subapical calli obsolescent, the broad intervals feebly convex, the minute
punctures forming small transverse wrinkles near the base but evanescent
behind, entirely devoid of scaling. Legs with the space between the front coxae
as broad as the antennal club ; femora with rather large shallow punctures ; front
tibiae somewhat widened at the apex, with a distinct mucro in $ and a minute
one in $, both widely separated from the uncus.
Length 3-5-4-5 mm., breadth I-5-I-6 mm.
N.E. Burma : Kambaiti, 7000 ft., 2$ 1$, v-vi. 1934.
Psilarthrus squamifer sp. n.
<£$. Derm shiny black, the tarsi red-brown to piceous, the tarsal claws,
tibial imci, and the base of the antennal scape, red.
Head with small separated punctures, without any frontal fovea. Rostrum
similar in the two sexes, as long as the prothorax only, comparatively short and
stout, moderately curved, somewhat widened beyond the antennae which are
much nearer to the apex than usual, the apical part being about as long as the
antennal club ; dorsum with fine separated punctures, those on the sides coarse
and confluent. Antennae comparatively short, with joint 1 of the funicle slightly
longer than 2+3, 3-7 transverse. Prothorax nearly as long as broad, widest
at the very slightly projecting basal angles, very gradually narrowing to the
middle, with the sides straight in $ and feebly curved in $, then curving in to
the rather abrupt apical constriction ; dorsum slightly convex longitudinally,
with strong close even punctures, which are much broader than the intervals,
without any smooth median line ; with sparse narrow whitish scales, especially
towards the sides. Elytra rather broadly ovate, feebly rounded laterally, the
post-humeral sinuation very shallow ; dorsum without discal impressions, the
subapical calli distinct, the broad intervals flat and impunctate, with a row of
sparse long whitish scales on 3, 5, 7 and 9. Legs with rather large shallow punctures
on the femora ; front tibiae of <J not widened at the apex with a minute tooth on
the lower edge at one-third from the apex which is surrounded by a few com-
paratively long erect setae, and at the apex a very small mucro which is united
at the base with the uncus ; front tibiae of $ without the tooth or mucro.
Length 2-6 mm., breadth 1-2 mm.
N.E. Burma : Kambaiti, 7000 ft., 1$ 1$, v. 1934.
Key to the Species of Psilarthrus.
1 (2). Gular margin of presternum truncate ; space between the front
coxae not wider than the narrowest part of the antennal scape ;
front tibiae without a mucro in both sexes, those of $ with a
large triangular tooth a little beyond the middle, tarsi red ;
elytra with sparse isolated large whitish scales . . dentipes sp. n.
2 (1). Gular margin of presternum sinuate ; space between the front
coxae wider than the scape ; front tibiae of (J without any large
triangular tooth on the lower edge.
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 465
3 (4). Prothorax with a low median costa ; scape of antennae and tarsi
red ; front tibiae without a mucro ; elytra with sparse narrow
whitish scales ....... costicollis sp. n.
4 (3). Prothorax without any median costa ; antennae and tarsi black
or mainly black.
5 (6) . Prothorax with the punctures confluent laterally, forming curved
ridges ; front tibiae with a strong mucro in both sexes ; elytra
without scales ...... mucronatus sp. n.
6 (5). Prothorax with the punctures not confluent and without lateral
ridges ; front tibiae of 9 at most with a minute rudimentary
mucro.
7 (8). Rostrum dissimilar in the sexes, the distance-of the antennae from
the apex greater than the length of the antennal club ; antennal
scape entirely black ; elytra without scales ; front tibiae of <J
without any tooth on the lower edge and the mucro widely
separated from the uncus, that of 9 with a minute mucro
coracinus sp. n.
8 (7). Rostrum similar in the sexes, shorter and stouter, the distance
of the antennae from the apex about equal to the antennal club ;
scape red at the base ; elytra with sparse large whitish scales ;
front tibiae of <$ with a minute tooth at one-third from the apex,
the small mucro united with the base of the uncus, 9 without
any mucro ........ squamifer sp. n.
DOLICHAULAX gen. n.
The species upon which this genus is based agrees entirely with Psilarthrus,
gen. n., except in regard to the structure of the scrobes on the rostrum, which do
not pass beneath the rostrum basally but are fully visible laterally right up to
the eyes, and moreover they extend deeply beyond the antennae almost to the
apex. This condition of the scrobes is quite exceptional, and I have been unable
to find a similar case in any other Old World Baridinae.
Genotype : Dolichaidax curvirostris sp. n.
Dolichaulax curvirostris sp. n.
9. Derm black beneath, the head and pronotum black with the apical
margin of the latter red-brown, the elytra chestnut-brown, the legs reddish with
the femora and tibiae sometimes piceous ; prothorax and elytra with sparse
whitish scales.
Head with fine separated punctures, without any frontal fovea. Rostrum
of 9 long and very strongly curved, i-6 times as long as the head and pronotum,
comparatively slender, cylindrical and only slightly widened close to the apex,
the antennae being inserted at a little beyond the middle ; dorsum shiny and
impunctate, the lateral areas with small close shallow punctures at the base and
a single line of punctures from there to the antennae. Antennae red-brown with
the club fuscous ; scape long, slender and abruptly clavate ; funicle not widened
distally, joint r as long as 2+3, 3 and 4 longer than broad. 5-7 as long as or
slightly longer than broad. Prothorax nearly as long as broad, parallel-sided from
the base to the middle, then narrowing with a curve to the shallow apical
constriction ; dorsum almost flat longitudinally, closely and strongly punctate,
without any smooth median line, and with separated oblong whitish scales
466 SIR GUY A. K. MARSHALL
laterally and a few on the disk. Elytra oblong-ovate, parallel from the prominent
shoulders to the middle and obtusely rounded at the apex ; dorsum without
discal impressions, the fine deep striae with very shallow small punctures, the
broad flat intervals impunctate, 3, 5, 7 and 9 with large separated triangular
whitish scales (often partly abraded), the alternate intervals with small narrow
scales, the subapical calli well-developed and obtuse. Legs slender ; femora with
fine sparse punctures, each containing a very short fine recumbent pale seta ;
tibiae impunctate, not widened at the apex, without any mucro.
Length 3-0 mm., breadth i-o mm.
N.E. Burma : Kambaiti, 7000 ft., 3$, v-vi. 1934.
CALYPTOPYGUS gen. n
Head not separated from the rostrum by an impression. Rostrum slender,
cylindrical, not compressed at the base, nor narrowed dorso-ventrally at the apex,
without carinae ; scrobes passing below the rostrum near the base ; mandibles
dentate and interlocking. Antennae inserted at a little beyond the middle of the
rostrum in both sexes ; scape slender, abruptly clavate ; funicle with joint 1
longer than 2, 7 distinct from the club. Prothorax as long as broad, shallowly
bisinuate at the base, truncate at the apex, without any postocular lobes, the
dorsum laterally compressed in front. Elytra only a little wider than the prothorax
with very sloping shoulders and ten deep complete striae, the posterior calli,
obsolescent. Legs slender ; femora moderately clavate, not sulcate or toothed
beneath ; tibiae not dilated distally, with fine shallow sulci dorsally, and without
a mucro ; tarsi with joint 3 deeply bilobate, the claws separated. Sternum
without any prosternal sulcus, the gular margin truncate, and the hind margin
not produced in the middle ; the mesosternum only slightly lower than the
metasternum. Abdomen with the pygidium entirely concealed in both sexes ;
ventrite 2 longer than 3 and 4 and fused in the middle with 1.
Genotype : Calyptopygus elliplicus sp. n.
Distinguished from Baris by the outline of the slender rostrum being
completely continuous with that of the head and by the entirely concealed
pygidium. These characters also distinguish it from the two preceding new
genera, which it somewhat resembles, though the facies is different owing to its
broader prothorax and narrow elytral shoulders.
Calyptopygus ellipticus sp. n.
(J$. Derm entirely black, shiny ; the upperside bare, except for sparse
minute narrow whitish scales at the sides of the pronotum ; underside with a
small whitish scale in each puncture.
Head with fine sparse punctures. Rostrum of <$ as long as the pronotum,
moderately curved and almost parallel -sided throughout, the antennae being
inserted at three-eighths from the apex ; dorsum shiny, with an irregular line of
punctures on each side from the base nearly to the apex, the lateral areas with
coarser confluent punctures and a short sulcus above the scrobe ; rostrum of 9 of
the same length but very slightly more slender and with the apex red, all the
punctures much reduced and the antennae slightly farther from the apex.
Antennae black ; funicle only slightly widened distally, joint 1 as long as 24-3,
4-7 transverse. Elytra only a little wider at the shoulders than the base of the
prothorax (10-5 : 8), parallel to beyond the middle, broadly rounded at the apex ;
dorsum rather flat, without discal impressions, the broad intervals with a row of
distinct small separated punctures. Legs rather slender, entirely black (except
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 467
the red claws), with sparse small punctures, each containing a short recumbent
white seta ; front femora with a fringe of erect pale setae on the lower edge,
which are rather longer in <J; front tibiae of (J with a small triangular tooth on the
lower edge a little beyond the middle.
Length 3-2-3-5 mm., breadth 11-12 mm.
N.E. Burma : Kambaiti, 7000 ft., 3 $ 5$, iii-vi. 1934.
Subfamily Cossoninae.
Eutornus lateralis sp. n.
(J9- Derm of head, prothorax and underside black, rostrum black basally
and red-brown in front, elytra red-brown, with the suture, lateral margins and
apical declivity usually blackish.
Head very shallowly constricted behind the eyes, with rather strong close
punctures and a short shallow frontal stria. Rostrum stout, moderately curved,
1-5 times as long as broad, fairly closely punctate and with a short interantennal
stria, the punctures being somewhat finer in $. Antennae inserted at a little
behind the middle of the rostrum in both sexes, red, with the funicle darker ;
scape strongly curved and gradually widening distally. Prothorax longer than
broad, much longer than the head and rostrum (3 : 2), widest near the base,
feebly rounded laterally, and deeply constricted near the apex ; dorsum com-
paratively strongly punctured, with an abbreviated impunctate median line, the
punctures being a little smaller than those in the striae on the elytra and much
larger than those on the intervals. Elytra with closely punctate striae, stria I
and the basal half of 5 being deeper than the others ; the intervals almost flat
on the disk, with a single row of very small distant punctures. Legs with the
front tibiae sinuate on the dorsal edge.
Length 3-5-4-7 mm., breadth 0-9-1-1 mm.
N.E. Burma : Kambaiti, 7000 ft., 15^ 21$, iv-vi. 1934.
Of the five other Burmese species the most nearly allied are antennalis Fst. and
congener Woll. The former differs in having the prothorax much more finely
punctate and the apical constriction shallower, the punctures in the striae on the
elytra are about twice as large as those on the pronotum, and in the male the
antennal scape is abruptly incrassate. E. congener has the prothorax red, with
much finer punctures and a shallower constriction, and joint 4 of the tarsi is
widened towards the base, which is not the case in lateralis.
Cossonus corax sp. n.
Derm shiny black, the tarsi, the stem of the scape and sometimes the
funicle, red.
Head with sparse minute punctures on the vertex, the frons with much
larger separated punctures and a deep median fovea ; eyes gently convex, their
curvature not continuous with that of the temples. Rostrum with the head as
long as or slightly longer than the pronotum, porrect in the basal half and sloping
steeply at the apex, gradually widening from the base to the antennae and there
abruptly dilated, the apical area being as long as broad; dorsum with numerous
small separated punctures which are much finer on the apical area. Prothorax
a little broader than long, gently rounded laterally, widest behind the middle,
rather deeply constricted apically, but the constriction not continuous across the
disk ; dorsum without any median or basal depressions, with rather large
punctures, which are smaller and dense laterally, becoming larger, irregular and
468 SIR GUY A. K. MARSHALL
sparser in the middle of the disk, leaving variable impunctate patches. Elytra
much wider at the shoulders than the prothorax, flat as far as interval 3, the
striae with strong close punctures which become much smaller on the declivity,
where the striae are much deeper ; intervals 1 to 4 or 5 flat, broader than the
striae, the lateral ones much narrower, all with an irregular row of minute
punctures that do not become larger behind. Legs with small sparse punctures
on the femora ; front tibiae not angulate beneath.
Length 5-0-7-0 mm., breadth 1-0-2-2 mm.
N.E. Burma : Kambaiti, 7000 ft., four specimens, v-vi. 1934.
Allied to C. carinensis Fst., which differs in its much shorter rostrum, flatter
eyes, and much more densely punctate pronotum, which normally has a shallow
median depression.
Cossonus stricticeps sp. n.
Derm shiny black, the tarsi, the stem of the scape and the base of the funicle
red.
Head constricted immediately behind the eyes, the vertex impunctate, the
frons with strong punctures and a large elongate fovea. Rostrum rather long,
together with the head longer than the pronotum (7:6), porrect in the basal
half and strongly curved downwards at the antennae, comparatively narrow
near the base, widening to the antennae and there abruptly dilated, the sub-
quadrate apical part being slightly broader than long ; dorsum with rather strong
punctures, which are sparser on the basal half and denser in front. Prothorax as
long as or very slightly longer than broad, almost parallel-sided from the base to
the middle then curving in to the rather shallow apical constriction, which is not
continuous across the disk ; dorsum without the usual longitudinal median
impression, but with a transverse depression in the middle of the base, the
middle of the disk with large irregular remote punctures which become much
denser laterally. Elytra much wider at the shoulders than the prothorax, flat as
far as interval 3, the striae containing very large round punctures which becomes
very much smaller behind, the striae being much deeper there ; the intervals
much narrower than the striae, except 2 (and sometimes 3) behind the middle,
and bearing an irregular row of minute punctures, which become much coarser
on the declivity. Legs with the punctures on the femora fine and sparse but
denser towards the apex and dorsally ; front tibiae not angulate beneath.
Length 6-5 mm., breadth 2-0 mm.
N.E. Burma : Kambaiti, 7000 ft., two specimens, v. 1934.
Distinguished from the other Burmese species by the constriction behind
the eyes, the parallel-sided prothorax, and the strongly punctate elytra.
Oxydema sp. — Taunggyi, 1^, viii-ix.
? Eremotes 2 spp. — Kambaiti, 2 ex., v-vi.
Tomicoproctus sp. — Kambaiti, 1 ex., vi.
Genus ?. — Kambaiti, 2 ex., iv-v.
Xenomimetes opacicauda sp. n.
Derm piceous brown, bare, the prothorax usually darker ; antennae and
tarsi red.
Head with small sparse punctures on the vertex, those on the frons larger
and much denser. Rostrum as long as broad, parallel-sided, with small shallow
punctures and a shallow median sulcus. Prothorax a little longer than broad,
widest not far from the base, almost parallel-sided in the apical half, the sides
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 469
with a small sinuation in the middle and gently rounded posteriorly, the apical
constriction well-marked ; dorsum with dense elongate punctures and an
abbreviated smooth median line, the punctures longitudinally confluent,
especially on the sides and pleurae. Elytra cylindrical, parallel from the shoulders
to the very deep subapical constriction below the subrectangular posterior calli
on interval 9, the apices being, as usual, subexplanate, broadly divergent and
with the margins feebly denticulate ; dorsum rather shiny, but the apical
declivity quite opaque, with the intervals broader, flatter, finely rugulose and
with short recumbent setae ; the striae containing deep close oblong punctures,
the intervals about as broad as the striae with a single row of small punctures,
except on the first three intervals, of which 1 is transversely wrinkled and
depressed at the base, and 2 and 3 are confusedly punctate on the basal half,
uniting at the base to form a rugulose callus.
Length 3-3-4-0 mm., breadth o-8-i-o mm.
N.E. Burma : Kambaiti, 7000 ft., five specimens, v-vi. 1934.
In the closely allied A*, himalayensis Stebb. and sikkimensis Mshl. (both
originally described in E remotes) the punctures on the pronotum are not elongate,
the elytra are not more opaque on the declivity than on the disk and the posterior
calli are not so angularly prominent.
Subfamily Stromboscerinae.
Xerodermia sp. — Kambaiti, 1$, iii.
Dryophthoroides sp. — Kambaiti, 1^, iii.
Subfamily Calandrinae.
Cyrtotrachelus buqueti Guerin. — S. Shan States, Pekkong, 3000 ft., i<^, x.
Cercidocerus carinensis Faust. — S. Shan States, Taunggyi, i^J, ix-x ; Pekkong,
i<Ji$. x.
Cercidocerus dohertyi Giinther. — Taunggyi, 2$, ix-x.
Olidognathus quadrimaculatus Buq. — Kambaiti, 1^ 19. v.
Otidognathus sp. — Kambaiti, 1^, v.
Otidognathus amoenulus sp. n.
<J$. Derm black ; elytra with a variable irregular reddish-yellow transverse
band near the base from stria 1 to the margin, then continuing to the base and
inwards to stria 6, so as to half-encircle the black humeral callus ; behind this a
similar broad band running obliquely from about the middle of the lateral margin
to behind the middle of stria 1 ; underside entirely black or with the mesepimera
and the anterior part of the prosternum red.
Head strongly and closely punctate ; frons almost parallel-sided, fiat, with
or without a deep median fovea. Rostrum a little shorter than the pronotum,
that of $ with distinct irregular punctures without any granules, its upper
surface forming a continuous line with the frons ; that of $ less dilated at the
base, with minute obsolescent punctures and with a fine median carina on the
apical half, the dorsal outline forming a wide angle with the frons. Antennae
black ; funicle with joint 2 longer than broad, 6 as long as broad ; club with the
pubescent part only narrowly projecting, the glabrous part as long as broad in <£,
much longer in $. Prothorax widest at its strongly arcuate base and narrowing
with a curve to the apical collar ; dorsum with rather strong uneven punctation,
which is denser in the middle of the anterior half, a deep impunctate transverse
sulcus close to the base and on the posterior half of the disk a broad deep oblique
47° SIR GUY A. K. MARSHALL
punctate depression on each side, these almost meeting on the middle line at
one-fifth from the base ; the lateral areas of the apical collar almost impunctate.
Scutellum unusually large, nearly forming an equilateral triangle, with a few
shallow punctures at the base. Elytra widest at the oblique shoulders and rapidly
narrowing behind ; all the striae impressed, 1-5 with the punctures very small
and shallow, those in 6-9 much stronger ; the intervals with an irregular row of
minute punctures. Legs, with minute sparse punctures, red, with the knees,
apex of tibiae and the tarsi black, or the tibiae and femora may be half black.
Pygidium closely punctate, with a low median carina.
Length 5-0 mm., breadth 2-0 mm.
N.E. Burma : Kambaiti, 7000 ft., 1^ 1$, v-vi. 1934.
Closely allied to the Assamese 0. safeties Gthr., which differs as follows : the
rostrum of the £ bears two rows of granules ; the pubescent part of the antennal
club is nearly or quite as long as the glabrous part ; the pronotum is more finely
and sparsely punctate ; the scutellum is much smaller and narrower ; the elytra
are much more narrowed behind and the shoulders more prominent.
Adapanetus malaisei sp. n.
cJ. Derm bare, rather opaque, very variable in colour ; head and rostrum
black ; prothorax black beneath, fulvous above, with the basal and apical
margins and a variable median stripe black, the latter normally narrow in front
and dilated behind, but varying in width, and rarely the whole dorsum black ;
elytra normally fulvous, with the suture and lateral margins black, a transverse
black spot near the base between striae 2 and 5 (often much reduced or entirely
absent), an oblong subapical spot between striae 3 or 4 and 6, and sometimes a
transverse black band at the middle ; rarely the elytra may be black with six
fulvous spots ; underside rarely entirely black, usually with a stripe at the sides
of the metasternum, another on the metepisterna, and the basal margin of the
ventrites, reddish.
Head with comparatively strong irregular punctures and a deep frontal
fovea. Rostrum shorter than the pronotum (7 : 10), the dilated basal part
parallel-sided, punctate like the head, and without any angular projection on the
lower edge near the base ; the narrow anterior part moderately curved, opaque
and minutely punctate. Antennae black, shiny ; scape only slightly longer than
the funicle ; club of $ hatchet-shaped, with its basal two-thirds chitinous and
shiny, the spongy terminal part with its apical margin broadly truncate. Prothorax
a little longer than broad, widest near the base, the basal angles rounded, the
sides straight in the middle or very shallowly sinuate, roundly narrowing in front
to the deep apical constriction which is continued across the disk ; dorsum with
irregular strong punctures which are densest near the apex and in the middle of
the base, and in front of the middle an indefinite broad transverse area which
is impunctate or very sparsely punctate. Scutellum elongate, triangular,
impunctate and very shallowly impressed in the middle. Elytra somewhat wider
at the rounded shoulders than the prothorax, widest at the shoulders, very
slightly narrowing behind, and separately rounded at the apex ; the narrow
deep striae with fairly strong punctures, striae 7 and 8 continued to the base, 8
very shallowly ; the convex intervals with a row of distinct separated punctures.
Legs fulvous, with the apex and base of the femora and tibiae black, the tibiae
rarely all black, the tarsi black ; structurally as in the genotype.
Length 8-0-9-0 mm., breadth 2-7-3-0 mm.
N.E. Burma : Kambaiti, 7000 ft., 9$, v-vi. 1934.
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 47I
The only other known species, A . sericoclava Giinther 1936, differs in having
the head, rostrum and pronotum much more finely punctate ; the rostrum is
separated from the head by a narrow transverse depression and its basal part is
strongly dilated from the base to the antennae ; the antennal club of the male is
silky throughout and its apical margin is broadly rounded ; the elytra are jointly
rounded at the apex, the intervals being flat and minutely punctate.
Adapanetus sericoclava Giinther. — Kambaiti, 3^ 1.9. iv-vi.
Aplotes histrio sp. n.
<J9- Derm red ; the head dorsally, the apex of the rostrum and the area
round the scrobe, blackish ; prothorax with the following black markings : an
incomplete median stripe, an elongate spot in the basal angles, a variable stripe
on the pleurae and another one passing over the coxa ; black markings on the
elytra : a little before the middle a transverse spot between striae 2 and 5 or 6,
a small one on interval 9 (rarely 8 and 9), and an oblique band above the declivity
between striae 3 and 9, and just in front of this a small spot on interval 3, which
is often absent and rarely (in 9) extending inwards to the suture ; underside with
very variable black markings, often absent.
Head coarsely punctate, with a shallow punctate sulcus immediately behind
the eyes ; frons one-third as wide as the base of the rostrum. Rostrum rather
strongly curved, slender, the dilated basal part irregularly punctate and with a
shallow median stria ; the slender anterior part impunctate in $ except near the
base, shorter and slightly stouter in $ with very shallow punctures on the sides.
Antennae black ; funicle with joint 1 longer than 2, 3 and 4 as long as broad, 5
and 6 transverse ; club with the pubescent part occupying fully half the length.
Prothorax of $ almost parallel-sided, only slightly widening towards the base,
with an abruptly constricted apical collar, that of 9 distinctly widening basally ;
dorsum coarsely reticulately punctate, without any median sulcus but with a
large shallow discal depression on the basal half and an abbreviated median
carina. Scutellum plane, without any median stria. Elytra only slightly wider at
the shoulders than the prothorax in <$, distinctly broader in 9. subparallel from
there to the middle, with a broad shallow transverse impression close to the base
extending from the scutellum to the shoulder ; the striae with strong separated
punctures on the basal half which diminish behind, interval 1 with a dense row of
small punctures, the others with large shallow irregular punctures on the basal
third or half which become small and sparse or even obsolescent behind. Legs
red, with the knees and tarsi black ; femora with rather large separated punctures
usually filled with tomentum ; tarsi with joint 4 normal. Pygidium of $ coarsely
punctate, without a median carina, obtusely rounded at the apex ; that of 9
truncate at the apex, with a tuft of setae on each angle.
Length 6-5-8-5 mm., breadth 2-5-3-0 mm.
N.E. Burma : Kambaiti, 7000 ft., 11$ 119. iv-vi. 1934.
Nearly allied to the Japanese A. roelofsi Chev., which, however, has the head,
apex of the rostrum and the tarsi red, whereas the basal margin of the prothorax
and the humeral callus of the elytra are black ; the head has no sulcus behind the
eyes ; the pubescent part of the antennal club is much shorter than the glabrous
part ; the pronotum bears a broad median sulcus, the punctures are smaller and
hardly reticulate, and the apical constriction is much less abrupt ; the pygidium
of the male with the apical margin broadly sinuate and with a tuft of setae in
each angle.
472 SIR GUY A. K. MARSHALL
Aplotes pollex sp. n.
(J9- Colour similar to that of A. histrio sp. n., except that the head and
rostrum are entirely red, and there is a small black spot on the shoulders of the
elytra.
Head coarsely punctate, with a sulcus behind the eyes ; front half as wide as
the base of the rostrum. Rostrum as in A. histrio except that the apex is not
black, and that of the <$ is distinctly punctate above and laterally on the basal
two-thirds. Antennae red, only the club black ; funicle with the two basal joints
subequal, the rest more or less transverse ; club with the pubescent part occupying
a little less than half the length. Prothorax differing from that of histrio in being
more bell-shaped, similar in the two sexes, the dorsum rather less coarsely
punctate, with a broad median sulcus that is dilated behind. Scutellum with a
distinct median stria. Elytra similar in the two sexes, as wide at the shoulders as
in histrio 9 but narrowing more rapidly behind, the basal depression shallower and
interrupted by interval 3; the punctures in the striae and on the intervals much
smaller. Legs red, the knees not black, the tarsi red to red-brown ; the fourth
tarsal joint produced into a sharp angle on the underside behind each claw.
Pygidium of <J with the apex broadly truncate and with sparse erect pale setae,
that of $ narrower at the apex and without erect setae.
Length 7-0-10-0 mm., breadth 2-4-3-8 mm.
N.E. Burma : Kambaiti, 7000 ft., 2431 20$, iv-vi. 1934.
This species closely resembles superficially the female of A. histrio sp. n.,
but it may be distinguished from all the known Burmese species by the unusual
structure of the claw-joint of the tarsi.
Aplotes sp. — Kambaiti, 1$, iv
Aplotes sp. — Sadon, 4000 ft.,
1 9. vi-vii.
Diocalandra caelata sp. n.
(J9- Derm dull black ; the extreme apex of the rostrum and the apical
margin of the prothorax reddish ; elytra with a fulvous stripe on intervals 6-8
from the base nearly to the middle, where it unites with the anterior margin of
a large quadrate fulvous spot on intervals 2-6, and a smaller oblong spot on
interval 3-5 extending from near the base to one-third, the basal half of which
is usually connected with the lateral stripe.
Head coarsely punctate, the frons as long as broad, with a deep median
sulcus and a row of erect setae on each side. Rostrum of $ with the head as long
as the pronotum, the dilated basal part almost parallel-sided and shorter than
the head ; dorsum with two rows of minute granules and a very fine median
carina ; rostrum of 9 longer, the narrow anterior part without granules or carina,
the former replaced by punctures. Antennae blackish with the scape red ; scape
strongly curved ; funicle with joint 2 longer than 1, the rest as long as or slightly
longer than broad ; club with the pubescent apical part as long as the shiny
basal part. Prothorax 1-5 times as long as broad, almost parallel-sided from
the base to two-thirds, then narrowing gradually to the deep subapical constric-
tion ; dorsum opaque, with dense reticulate punctation and a very shallow large
depression in the middle of the basal half, and on each side of this behind the
middle a small group of very short erect setae. Elytra scarcely wider at the
shoulders than the prothorax and narrowing very gradually from there to the
apex ; the1 striae containing large quadrate punctures, the alternate intervals
BURMA AND BRITISH INDIA — COLEOPTERA : CURCULIONIDAE 473
more raised and bearing a rather sparse row of erect clavate white setae. Legs
comparatively long and slender, black, the basal half and apical margin of the
femora red ; femora rugosely punctate, front tibiae finely serrate on the lower
edge. Pygidium coarsely punctate, with sparse erect clavate white setae.
Length 3-5-4"5 mm., breadth o-8-i-o mm.
N.E. Burma : Kambaiti, 7000 ft., 3(j4$, v-vi. 1934.
Most nearly allied to D. elongata Roel., which in the Catalogus Coleopt. is
placed in Calandra (=Sphenophorus). This Japanese species differs in having
the rostrum much shorter, of the same length in the two sexes, that of the male
having no granules ; the antennae are red and the pubescent part of the club is
much shorter than the bare part ; the setae on the elytra are much more numerous
and closely set ; and the legs are shorter and stouter.
Calandra oryzae L. — Myitkyina, 1$ 1$, hi ; Sadon, 2$, vi-vii.
Periphemns sp. — Tenasserim, Malvedaung, 100 ft., 1$ 19, xi.
474
MECEDANUM
475
A SYNOPSIS OF THE GENUS MECEDANUM ERICHSON
(COLEOPTERA, COLYDIIDAE)
By H. E. HINTON, Ph.D.,
Department of Entomology, British Museum (Natural History).
(With five text-figures.)
(~)NLY five species of Mecedanum have been described, three from Madagascar
and two from Africa. In this paper four additional species are described,
three from Africa and one from the Seychelles Islands. The presence
of a species of Mecedanum in the Seychelles Islands (Silhouette) was
reported long ago by Grouvelle (1918), and the new species here described from
those islands is based on the two damaged specimens examined by Grouvelle. A
list of the species, including the new additions, is as follows : —
Total number of specimens examined
5i
In his catalogue of the family, Hetschko (1930) includes a further species
M. erichsoni Broun, which he records from New Zealand. The citation following
this name refers to the original description of Lithostygnus minor Broun (1893).
In the Zoological Record for 1893, M. erichsoni Sharp is listed immediately
beneath Lithostygnus minor Broun, so that by a mistake in transcribing the
names, the compiler of the catalogue was left with a M. erichsoni Broun from
New Zealand and a M. erichsoni Sharp from Madagascar. M. erichsoni Sharp
was unnecessarily renamed by Hetschko, and the synonymy of the species is
now as follows :—
Mecedanum erichsoni Sharp.
1893. Mecedanum erichsoni Sharp, Knt. num. Mag. 29 : 257.
1929. Mecedanum sharpi Hetschko, Wien. en!. Ztg. 46 : 84.
1930. Mecedanum sharpi Hetschko. Coleopt. Cat. 107 : 10.
Almost nothing is known of the biology of these curious-looking Colydiidae.
Observations made by Mr. G. H. Thompson in the Gold Coast on M. antennatum
(Kraatz), M. filum sp. n., M. tomentosum sp. n., and .1/. giganteum (Kraatz)
suggest that there is a definite association between the species of Mecedanum and
various species of Platypodidae (Platypus, Crossotarsus) and possibly some
species of Scolytidae. All of the species mentioned above have been found by
Mr. Thompson in the galleries of species of Platypus and Crossotarsus, but whether
they feed on the fungi in the galleries or whether they prey on the makers of the
galleries is not clear.
In all species of Mecedanum there is an extremely fine plate projecting from
the sides of the first five abdominal segments. This semi-membranous plate is
finely striated, and it must rub against the striated inner ventral surface of the
476 H. E. HINTON
elytra when the abdomen and elytra are moved against each other. It is possible
that the semi-membranous abdominal plates and the striated inner surface of
the elytra are the two components of a stridulating organ.
Key to the Species of Mecedanum.
1. Front of head without median longitudinal carinae or gibbosities . . 2
— Front of head with one or more median longitudinal carinae or
gibbosities .......... 4
2. Elytra with fourth and fifth carinae joined together before apex ;
third, fifth, and seventh intervals distinctly carinate on middle of
elytra. Clypeus and anterior part of head with punctures as coarse
as facets of eyes, often elongate, and above eyes often longitudinally
confluent so that surface is partly strigose. Length, 9-0-10-0 mm.
Madagascar . . . Mecedanum punctatum Grouvelle (1906)
— Elytra with fourth carina ending freely before apex, not joined to
fifth ; third, fifth, and seventh intervals only feebly convex on middle
of elytra. Clypeus and anterior part of head with punctures round
or oval, never elongate and longitudinally confluent ... 3
3. Antenna with club 5-segmented. Head with a transverse ridge
opposite anterior margin of eyes ; middle of head opposite anterior
margins of eyes very nearly as high as margins above eyes.
Pronotum narrowest at base, not distinctly narrowed before base
but with sides more or less evenly converging from apex to base.
Abdomen with punctures of fourth sternite much finer than those
of third and fifth sternites except for a posterior row of coarse,
deep, often confluent punctures. Length, 8-0 mm. W. Africa,
Uganda ..... Mecedanum antennatum (Kraatz, 1895)
— Antenna (fig. 1) with club distinctly 3-segmented. Head without a
transverse ridge opposite anterior margins of eyes and here very
much lower than margins above eyes, which are very strongly
elevated. Pronotum broadly and deeply sinuate before base so that
it is distinctly narrower in front of base than at base. Abdomen
entirely coarsely punctate like third sternite or at most with an
oval area on each side which is only finely punctate. Length,
4-5-6-9 mm. Belgian Congo, Sierra Leone, Gold Coast
. Mecedanum filum sp. n.
4. Front of head with three longitudinal carinae. Length, 8-0-13-0 mm.
Madagascar . . . Mecedanum carinifrons Grouvelle (1906)
— Front of head with a single median longitudinal cariniform elevation
or with a single very broad longitudinal gibbosity ... 5
5. Elytra (fig. 4) with third and fourth carinae ending freely before apex,
neither joined to the fifth (ninth interval) ; second to fifth carinae
nearly as strongly raised on basal half as at apex. Pronotum with
sides of median impression narrowly and strongly elevated,
cariniform. Head, pronotum, and carinate intervals of elytra
densely and finely but conspicuously tomentose. Length, 9-5 mm.
Gold Coast ..... Mecedanum tomentosum sp. n.
MECEDANUM
477
EXPLANATION OF TEXT-FIGURES
Figs. 1-3 Mecedanum fllum, sp. n. (1) Antenna. (2) Dorsal view of male genitalia after the
median lobe has been pulled out. (3) Ventral view of median lobe Lines next to figures refer
to a length of 0.20 mm.
Fig. 4 Mecedanum tomentosum, sp. n. Lateral view of apex of right elytron.
Fiq. 5 Mecedanum centralis, sp. n. Lateral view of apex of right elytron.
NOVIT. ZOOL., 42, 3. I948. 85
47»
H. E. HINTON
— Elytra (fig. 5) with third or fourth carina joined to fifth (ninth
interval) ; second to fifth carinae noticeably decreasing in height
towards base so that on basal half they are usually but little more
convex than the even intervals. Pronotum with sides of median
impression rounded or elevated but never produced as narrow
carinae. Head, pronotum, and carinate intervals of elytra
glabrous or sparsely hairy, without trace of tomentum ... 6
6. Elytra (fig. 5) with fourth carina ending freely before apex, not joined
to third or fifth 7
— Elytra with fourth carina joined to fifth before apex .... 8
7. Head with anterior margin of clypeus moderately deeply, arcuately
emarginate ; coarsest punctures of clypeus and frons much finer than
facets of eyes and seldom separated by less than three diameters ;
median carina approximately half as broad as head between carina
and elevated margins above eyes ; top of median carina convex.
Eyes separated beneath head by much less than twice their horizontal
diameter (30 : 22). Pronotum with narrow channel in bottom of
median impression restricted to anterior seventh ; hypomera with
oval punctures usually separated by less than their diameters.
Metasternal sides without apparent hairs between the setiferous
punctures. Length, 5-0 mm. Seychelles Islands. Mecedanum scotti sp. n.
— Head with anterior margin of clypeus truncate or nearly so ; coarsest
punctures of clypeus and frons as coarse or coarser than facets of
eyes and rarely separated by more than two diameters ; median
carina two-thirds as broad as head between carina and elevated
margins above eyes ; top of median carina flat ; Eyes separated
beneath head by twice (90 : 40) or nearly twice (68 : 40) their
horizontal diameters. Pronotum with a nearly complete, narrow,
parallel-sided channel in bottom of median longitudinal impression ;
hypomera with round punctures usually separated by one or more
diameters. Metasternal sides very finely and densely pubescent,
nearly tomentose. Length, 10-0-12-5 mm. Uganda, Congo. .
Mecedanum centralis sp. n.
8. Antenna with club 5-segmented, the seventh segment being scarcely
narrower than the eighth and much broader than the sixth. Front
of head with median gibbosity very strongly raised and flat-topped.
Apex of elytra deeply and broadly, truncately emarginate. Length
8-5-13-5 mm. Togo, Gold Coast, Uganda
Mecedanum giganteum (Kraatz, 1895)
— Antenna with club 4-segmented, the seventh segment being much
narrower than the eighth and scarcely broader than the sixth. Front
of head with median gibbosity feebly elevated. Apex of elytra at
most feebly, triangularly emarginate. Length, 8-2-9-5 mm.
Madagascar .... Mecedanum erichsoni Sharp (1893)
Mecedanum antennatum (Kraatz).
1895. Colydium antennatum Kraatz, Dtsch. ent. Z . 1895 : 158.
This species was described from two specimens taken in Togo. Before me
are three specimens, which agree well with the original description, with
MECEDANUM 479
data as follows : I, Uganda : Bunnga, 20. v. 1926 (G. L. R. Hancock) and 2,
Gold Coast : Mpraeso District, 19-24. ix. 1945, in galleries of Platypodidae in
Albizzia zygia and Trichilia hendelotii (G. H. Thompson).
Mecedanum filum sp. n. (Figs. 1-3).
Male: Length, 4-5-6-9 mm. ; breadth, 0-57-0-96 mm. Cuticle moderately
shining and black or nearly black with antennae and legs dark rufo-piceous.
Head without a median longitudinal carina or a transverse ridge ; in some of the
larger specimens the middle of the head between the eyes is very feebly convex.
Lateral margins narrowly and very strongly elevated to a point opposite
posterior margins of eyes, the most strongly elevated part of each margin being
opposite anterior margin of eye. Clypeus with anterior margin truncate or nearly
so and anterior angles rounded and moderately strongly projecting forwards ;
anterior tentorial pit (or channel in this position) oblong-oval or with sides
subparallel, and usually nearly twice as broad as largest puncture of head.
Surface of frons and clypeus with deep or moderately deep punctures which
are half as coarse to as coarse as facets of eyes and are very irregularly distributed ;
basal part of head with punctures broadly oval, as coarse as facets of eyes, and
usually separated by about one diameter, the surface between the punctures
being more strongly shining than that of frons and clypeus. Eyes separated
above by slightly more than twice their horizontal diameter (o-6o : 0-26 mm.)
and beneath head by about one and a half diameters (0-38 : 0-26 mm.). Antenna
(fig. 1) with a distinct 3-segmented club. Pronotum at broadest point, which is at
about apical fifth, less than half as broad as long (0-90 : 2-02 mm.) and apex
broader than base (0-82 : o-68 mm.). Sides distinctly margined very nearly to
base, and base completely, distinctly margined ; sides broadly, moderately
deeply sinuate before base so that narrowest part of pronotum is a short distance
before base. Median impression broad and deep with a deep narrow channel
extending along its bottom from apex to near base ; secondary channel deepened
and slightly broadened at middle and near apex ; sides of median impression
rounded, neither elevated nor carinate. Surface laterad from rounded and nearly
impunctate dorsal margins of the median impression with oval to irregular
punctures which are two to three times as coarse as facets of eyes ; in large
specimens these punctures are seldom separated by as much as one diameter, but
in some small specimens they are usually separated by about one diameter.
Elytra twice as long as pronotum (4-0 : 2-0 mm.) and very nearly parallel from
broadest point behind humeri (0-96 mm.) to near apex. Apex of elytra deeply,
very narrowly but often irregularly emarginate ; inner apex of each elytron with
a large, ventrally directed tooth, the margin laterad from this tooth being finely
toothed or crenate for a short distance. On basal two-fifths the alternate intervals
are not more than feebly convex except at extreme base where the second and
third (third and fifth intervals) are broadly carinate. Apex with first, second,
and third carinae joining fifth, which here forms the dorsal apical margin, the
second carina being joined to the fifth more or less opposite the junction of the
epipleuron and fifth. Third carina much lower than fifth at the junction of the
two, so that in some specimens — which have third less elevated than usual — this
carina appears to be separated from fifth by a short distance. Seventh interval
(fourth carina of other species) ending some distance before apex, where it is as
far from third as from fifth ; at apex only feebly convex, nowhere distinctly
carinate. Surface of carinae with sparse, erect, testaceous setae and also with
numerous punctures. Strial punctures very deep, flat-bottomed, subquadrate
480 H. E. HINTON
usually broader than flat intervals, and usually separated longitudinally by one
or slightly less than one diameter. Hypomera with punctures round to broadly
oval, as coarse as facets of eyes to half again as coarse, and usually separated by
slightly less than one to slightly more than one diameter ; surface between
punctures not pubescent (mag. X 75). Metasternum with extreme sides non-
tomentose and so closely punctate that surface is rugose. Abdomen with first
four stemites punctate something like hypomera ; surface between punctures
with a very dense, reticulate, alutaceous microsculpture ; fourth sternite in many
but not all specimens with an oval area of varying size on each side, which is
free of coarse punctures but has numerous fine punctures and is more strongly
shining than adjacent areas. Legs with first segment of front tarsi distinctly
shorter than combined length of two following segments (o-n : 0-16 mm.).
Middle and hind tarsi with first segment only slightly shorter than combined
length of two following segments (0-16 : 0-19 mm.).
Female : Externally similar to male.
Type : A male in the British Museum (Nat. Hist.). Belgian Congo : 18
miles S.W. of Elizabethville, i. 1928 (H. S. Evans).
Paratypes : 14, with same data as type. 7, Sierra Leone : 1, Kent, ii. 1928
(E. Hargreaves) ; 4, Njala, 4. i. 1933, under bark of dead rubber tree (E.
Hargreaves) ; I, Njala, at light, 7. x. 1925 (E. Hargreaves) ; and I without
further data. 15, Gold Coast (G. H. Thompson) ; 6, from Sunyani District and 9
from Mpraeso District, i-iii, vi, viii, ix, xi. 1945-46, on Albizzia zygia, Bosquiea
angolensis, Cassia siamea, Ficus exasperata, Lonchocarpus sericeus, Terminalia
ivoriensis, and Trichilia heudelotii timber or trees scorched by fire. These
specimens were commonly found under loose bark, and two were found in the
tunnels of Platypodidae (Crossotarsus sp.).
Comparative notes : The distinct three-segmented antennal club (fig. 1) will
serve to distinguish it from any other known species of Mecedanum.
Mecedanum tomentosum sp. n. (Fig. 4).
Male : Length, 9-4 mm. ; breadth, 15 mm. Cuticle nearly entirely mat, not
shining ; black or nearly black, with tarsi dark rufo-piceous and elytra moderately
pale brownish red. Cuticle nearly everywhere densely, scarcely visibly (mag. X 75)
pubescent, but head, pronotum, and carinate elytral intervals with the hairs
longer, denser, greenish yellow, and very distinct ; cuticle also with sparse, long,
nearly erect setae which are exceptionally long and regularly spaced on the
carinate elytral intervals ; fourth abdominal sternite moderately shining and with-
out the fine, dense pubescence of other sternites. Head with median longitudinal
carina extending from a point near anterior margin of clypeus to a point opposite
anterior margins of eyes, where it is very nearly as high as elevated margins
above antennae ; surface of median carina strongly convex. Clypeus with
anterior margin very broadly, shallowly, arcuately emarginate, with the antero-
lateral angles rounded but prominent. Tomentum of head absent on a narrow
anterior belt on clypeus, on top of median carina, and on top of lateral elevated
margins ; surface of head and clypeus -with numerous round or oval punctures
which are coarser than facets of eyes. Eyes more finely faceted than in any
other species known to me ; separated above by three times their horizontal
diameters (0-93 : 0-30 mm.) and separated beneath by more than two diameters
(0-77: 0-30 mm.) . Antenna with club 5-segmented, but seventh segment only slightly
broader than sixth although distinctly so. Pronotum at broadest point, which is
at about apical fourth, about half as broad as long (1-26 : 2-51 mm.) and base only
MECEDANUM 481
slightly narrower than apex (i-oi : r-20 mm.). Median impression deep and
broad with its sides strongly and narrowly carinate like the alternate intervals
of the elytra ; lateral marginal line deep and complete nearly to base ; lateral
margins thickened and cariniform, so that with sides of median impression the
pronotum is quadricarinate ; sides on anterior three-fifths between submedian
carinae and lateral margins broadly and longitudinally gibbous. Surface through-
out with round or nearly round punctures which are half again to twice as coarse
as facets of eyes and are usually separated by a little less than one to about two
diameters ; on basal third and in median impression the coarse punctures are
sparser and only about as broad as facets of eyes. Elytra more than twice as long
as pronotum (6-0 : 2-5 mm.), at base much broader than base of pronotum
(1-48 : i-oi mm.), and feebly broadened behind humeri to apical two-thirds,
where they become slightly narrower ; from middle feebly and gradually
broadened to about apical sixth, where they are broadest. Apex of elytra deeply
and truncately emarginate. Alternate intervals strongly but not equally carinate
from base to apex ; at base the second carina (third interval) is the most strongly
elevated and the third (fifth interval) the second most strongly elevated. First
carina (sutural interval) most strongly elevated from basal third to apical two-
fifths, where it has a few broad dorsal tubercles which are feebly to strongly
elevated. Second carinae feebly diverging from base, strongly diverging for a
short distance on apical declivity, and thence nearly parallel to apex where each
is joined to the fifth carina. Third and fourth carinae ending freely before apex
(fig. 4). Strial punctures very deep, often flat-bottomed, and usually two to three
times as coarse as facets of eyes. Legs with first segment of front tarsi distinctly
(7 : 8-5) and first segment of middle and hind tarsi scarcely (8-5 : 9-0) shorter
than combined length of two following segments.
Female : Unknown.
Type : A male in the British Museum (Nat. Hist.). Africa : Gold Coast :
Mpraeso District, 5. ix. 1945, in tunnels of Platypus sp. in a mahogany (Khaya
grandifolia) log (G. H. Thompson).
Comparative notes : The very distinctive appearance of this species is due to
the dense olive-green tomentum of the head, pronotum, and carinate intervals of
the elytra, no other species of the genus possessing tomentum on its dorsal
surface. In no other species are the margins of the median pronotal impression
elevated to form narrow carinae, and in no other are the elytral carinae strongly
elevated from apex to base. A further interesting feature of M. tomentosum is the
sparsely punctate and strongly shining fourth abdominal sternite which contrasts
strongly with the densely punctate and mat sternites before and behind.
Mecedanum scotti sp. n.
1918. Mecedanum sp., Grouvelle, Trans, ent. Soc. Land. 1918 : 28.
Female: Length, 5-0 mm. ; breadth, o-8 mm. Cuticle strongly shining and
moderately dark rufo-piceous ; fine hairs everywhere sparse. Head with median
longitudinal carina extending from a point opposite anterior tentorial pits to a
point opposite middle of eyes, posteriorly slightly more strongly elevated than
margins above eyes, dorsal surface strongly convex, and with breadth equal to
about half of distance between carina and elevated margins above eyes. Clypeus
with anterior margin broadly, moderately deeply, and arcuately emarginate.
Surface of frons and clypeus strongly polished and with both sparse microscopic
punctures and punctures about one-half to two-thirds as coarse as facets of eyes ;
482 H. E. HINTON
larger punctures largely confined to the clypeus, where they are rarely separated by
as little as one diameter ; head behind highest point of median carina with very
dense, deep, round to broadly oval punctures which are as coarse as facets of eyes.
Eyes separated above by twice their horizontal diameter (0-46 : 0-22 mm.) and
beneath head by little more than one diameter (0-30 : 0-22 mm.). Antenna with
segments from fourth onwards becoming progressively broader towards apex ;
seventh segment not sufficiently narrower than eighth to make a distinct 4-
segmented club ; inner surface of third to sixth segments with numerous long,
pale-testaceous hairs. Pronotum at broadest point, which is at about apical sixth,
slightly more than half as broad as long (0-82 : 1-48 mm.) and much broader than
base (0-82 : 0-65 mm.). Sides with a distinct marginal line extending from apex
nearly to base, but with margins cut off by marginal line not conspicuous.
Median impression broad, moderately shallow, and with latero-dorsal margins
distinct but rounded, not carinate ; pit slightly behind middle in bottom of median
impression deep and distinct ; apical fifth or sixth of impression with a deep
secondary channel which may be entire or interrupted. Surface with deep, round
to broadly oval punctures which are usually half again as coarse as facets of eyes
and are seldom separated by more than two-thirds of one diameter, but on
declivious part of apex are much finer and distinctly sparser ; surface between
coarse punctures with a few microscopic punctures, Elytra twice as long as
pronotum (3-07 : 1-53 mm.). Apex of elytra not or only very feebly emarginate and
from a lateral view regularly and distinctly toothed on apical part of epipleuron,
as the latter is regularly notched. Alternate intervals feebly convex at base but
becoming more strongly convex towards apex, so that on apical declivity first
(sutural), third, fifth, and ninth are distinctly carinate and seventh interval (fourth
carina of other species) is very feebly carinate. First three carinae joined to
fifth which is continued to suture as dorsal apical margin ; fourth carina ending
freely considerably before apex where its apex is nearly as far from fourth as from
fifth carina. Strial punctures deep, subquadrate, as broad or nearly as broad as
intervals, and usually separated longitudinally by slightly less than one to one
diameter. Hypomera with oval punctures very nearly as coarse as those of
pronotum and seldom separated by distinctly more than one diameter. Meta-
sternum with punctures of extreme sides oval, about as coarse as facets of eyes, and
usually separated by half or less than half of one diameter ; surface between
coarse punctures densely, scarcely visibly alutaceous and also with numerous
fine punctures but nevertheless moderately strongly shining, without apparent
hairs and not mat as in other species of the genus. Legs with first segment of
front tarsus about as long as the combined length of the two following segments.
Middle and hind tarsi with first segment very slightly longer than combined
length of two following segments.
Type : In the British Museum (Nat. Hist.). Seychelles Is. : Silhouette,
forest at edge of Mare aux Cochons plateau, ca. 1000 ft., ix. 1908 (H. Scott). The
apex of the elytra and all of the abdomen are lost.
Paratype : A female with same data as type. Part of the head and one
elytron are missing.
Comparative notes : Its small size and strongly shining, moderately pale
rufo-piceous instead of black or nearly black cuticle will serve to distinguish it at
first sight from any other Mecedannm. In structural details it most nearly
resembles M. centralis sp. n., but it is not very closely related to the latter, as is
evident from the differences between the two listed in the key. The species is
named after its collector, Dr. Hugh Scott, F.R.S.
MECEDANUM 483
Mecedanum centralis sp. n. (Fig. 5).
Male : Length, 12-5 mm. ; breadth, 2-n mm. Cuticle moderately strongly
shining and black or nearly black with legs dark rufo-piceous. Head with median
longitudinal carina extending from a point opposite anterior part of tentorial pits
to slightly beyond anterior margins of eyes, posteriorly nearly as strongly
elevated as margins above eyes, two-thirds as broad as head between carina and
elevated margin above eye, and with dorsal surface flat. Clypeus with anterior
margin truncate or nearly so, with anterior angles rounded and moderately
strongly produced forwards ; anterior tentorial pits parallel-sided, curved, and
about five times as long as broad. Surface of frons and clypeus with moderately
deep punctures which are usually round, slightly finer than facets of eyes to half
again as coarse, and are irregularly distributed, being separated by less than one
to two or even more diameters ; head behind median carina with coarse punctures
seldom separated by more than one diameter ; surface between coarse punctures
shining, densely microscopically alutaceous, and with a few very fine punctures.
Eyes separated above by three times their horizontal diameter (1*23 : 0-4 mm.
and beneath head by slightly more than two diameters (0-90 : C40 mm.).
Antenna with a 5-segmented club, the seventh segment being considerably
broader than the sixth, which is not noticeably broader than the fifth. Pronotum
at broadest point, which is at about apical fifth, only slightly more than half as
broad as long (r86 : 3-42 mm.) and apex broader than base (1-56 : 1-42 mm.).
Sides deeply margined from apex very nearly to base ; base broadly, shallowly,
and completely margined. Median impression broad and moderately deep ;
dorsal sides distinct but rounded and not carinate ; bottom of impression with a
deep, narrow, parallel-sided channel which extends from apex nearly to base ;
and median pit round, very deep, and scarcely broader than secondary channel
in median impression. Surface of pronotum with round punctures which are
half again to twice as coarse as facets of eyes and are usually separated by one-
half to two-thirds of their diameters ; surface between coarse punctures with
only microscopic punctures ; surface only microscopically punctate on narrow
dorsal part of sides of median impression. Elytra more than twice as long as
pronotum (8-39 : 3-42 mm.) and base broader than base of pronotum (178 :
1-42 mm.). Apex of elytra not or only scarcely noticeably emarginate. First
carina (sutural interval) only feebly convex on basal third and elsewhere only
moderately strongly carinate ; before apical declivity with four or five broad
but low gibbosities. Second carina (third interval) strongly carinate at base,
from basal fifth to about apical third only moderately convex, and thence strongly
carinate to apical margin to which it is joined shortly beyond junction of
epipleuron and fifth carina. Third carina joined to fifth before apex (fig. 5).
Fourth carina (seventh interval) ending freely some distance before apex. Strial
punctures very deep, subquadrate, as broad as even intervals, and usually
separated longitudinally by their own diameters. Hypomera with deep, round
punctures slightly finer than those of pronotum and usually separated by one to
two diameters ; surface between punctures with dense, scarcely visible (mag. X 75)
hairs which make the surface mat. Sides of metasternum similarly sculptured
and pubescent and also mat. Abdomen punctate and finely pubescent like
hypomera ; disk of first three sterntites not mat like sides and with the very fine
pubescence sparse or absent ; each side of fourth sternite with a large triangular
area which is strongly shining and free of the very fine pubescence. Legs with
first segment of front tarsus as long as combined length of two following segments.
484 H. E. HINTON
Middle and hind tarsi with first segment distinctly longer than combined length
of the two following.
Female : Unknown.
Type : A male in the British Museum (Nat. Hist.). Uganda : Entebbe, iii.
1929 (G. D. Hale Carpenter).
Paratype : Male, Belgian Congo : 18 miles S.W. of Elizabethville, 1927
(H. S. Evans). The paratype is only 10-5 mm. instead of 12-5 mm. long, its
elytra are slightly less strongly sculptured and more strongly shining, the top of
the median carina of the head is shallowly but distinctly concave, and nearly
the whole of the fourth sternite is shining and free of the microscopic pubescence.
The genitalia of the type and paratype are identical in all essentials. The
proximal teeth of the ejaculatory duct are slightly longer in the type, but these
and other slight differences are almost certainly attributable to the differences
between the two in absolute size.
Comparative notes : Of the known species, this is nearest to M. scotti, from
which it may be distinguished as shown in the key.
Mecedanum giganteum (Kraatz).
1893. Colydium giganteum Kraatz, Dtsch. ent. Z. 1895 : 159.
The types of this species are from Togo. In the collection before me there
are four females with data as follows : 1, Uganda : Budongo Forest, ix. 1934
(T. H. E. Jackson) ; 1, Uganda : Jinga, iv. 1923 (V. G. L. van Someren) ; and 2,
Gold Coast : Mpraeso District, 17. viii. 1945 and 13. i. 1946, in dead trees
(Albizzia zygia and Cassia siamea) associated with Scolytids and Platypodids
(G. H. Thompson) . These range in length from 8-5 mm. to 13-5 mm. All have
the inner apices of the third to seventh, and, to a lesser extent, the eighth
antennal segments clothed with very long, golden-testaceous hairs.
485
NOTES ON ASTEROIDS IN THE BRITISH MUSEUM
(NATURAL HISTORY)— I
The Species of ASTROPECTEN.
By D. DILWYN JOHN, M.Sc,
Department of Zoology, British Museum (Natural History).
(With four plates.)
' I HE large and valuable collection of Asteroids in the British Museum (Natural
•*- History) is being revised. This paper, describing what has seemed worthy
of description concerning the collection of Astropecten, will, it is hoped, be the
first of a series. It is therefore numbered as such.
The following species of Astropecten are represented in the Museum collection ;
those of which the types are held are marked with an asterisk (*) :
*acanthifera Sladen
andersoni Sladen
antares Doderlein
antillensis Liitken
aranciacus Linn.
"annates Gray
articulates (Say)
bispinosus Otto
brasiliensis M. and Tr.
*brevispinus Sladen
duplicates Gray
*formosus Sladen
granulates M. and Tr.
granulates natalensis n. subsp.
griegi Koehler
hartmeyeri Doderlein
hemprichi M. and Tr.
* hermatophilus Sladen
mimbelUs Sladen
indicus Doderlein
irregularis (Pennant)
"irregularis pontoporaeus Sladen
javanicus Liitken
johnstoni (Delle Chiaje)
kagoshimensis de Loriol
mammilatus Koehler
marginatus Gray
"mauntianus Gray
"mesactus Sladen
michaelseni Koehler
mindanensis Doderlein
*monacanthus Sladen
"pectinatus Sladen
platyacanthus Philippi
polyacanthus M. and Tr.
preissii M. and Tr
pusillulus Fisher
regalis Gray
sanctae helenae Mortensen
scoparius M. and Tr.
* sphenoplax Bell
spinuhsus Philippi
triseriatus M. and Tr.
* triseriatus fijiensis n. subsp.
vappa M. and Tr.
variegates Mortensen
velitaris von Martens
verrilli cali/ornicus Fisher
"zebra Sladen
Something is written below of fifteen of these forty-nine species. Five
(granulatus, indicus, monacanthus and zebra and hartmeyeri) are discussed chiefly
because the Museum specimens show that, within each, the range of variation is
greater than had previously been described. In one of them (granulatus) variation
is correlated with locality and a new subspecies is created. In four other species
(brevispinus, mesactus, pectinatus and sphenoplax) re-examination has shown the
presence of many more actinal intermediate plates than was hitherto known ;
they extend much farther down the arm than is normal in the genus. The Museum
specimens of triseriatus have made possible a fuller description of that little-
known form; one of them is described as a new subspecies. A. michaeheni
Koehler is shown to include A . dahomensis Doderlein ; some notes on the Museum
specimens are added. Some Museum specimens of A . antillensis appear to show
that the species has a greater range than was previously known.
Before the war a search was made for Gray's Asteroid types. Of those of his
species of Astropecten which, according to Doderlein (1917), are still valid as species
486 D. DILWYN JOHN
or subspecies, the types of only three were found. They are those of A . brasiliensis
armatus, A. brasiliensis erinaceus and A. mauritianus. They are redescribed and
figured.
Astropecten granulatus Miiller and Troschel
(Plate I, figs, i and 2.)
Astropecten granulatus Miiller and Troschel. 1842, Syst. Ast. : 7S ; Sladen, 1889 : 215. pi. 35. figs. 3-4,
pi. 39, figs. 4-6 ; Koehler, 1910 : 266 ; Doderlein, 1896 : 305. pi. 18, figs. 30 and 30a ; Doderlein,
1917 : 148, pi. 5. fig. 7; pi. 14, figs. 1, 3 and 3a; pi. 17, figs. 2, 2a and b ; Fisher, 1919 : 73;
Clark, H. L., 1923 : 250 ; Livingstone, 1932 : 242, pi. 8, figs. 2 and 3 ; Clark, H. L., 1938 : 60,
pi. 1, fig. 1 (in colour).
The locality from which the types of this species came is not known, but all
subsequent records, with the exception of that of Clark (1923), have been from
Australian, E. Indian or Philippine seas. Clark recorded it from off the coast of
Natal and from off Cape Agulhas (one "young and dubious specimen"), in South
African waters. But in 1938 he changed his opinion, saying : " Comparison of
the South African specimens (those he described in 1923) with the material from
Broome indicates that it is quite different and further study suggests that it is
nearer to A. notograpttts Sladen than to any species hitherto described. Hence
granulatus should be deleted from the list of South African sea-stars."
There are in the British Museum (Natural History) fourteen South African
specimens which I am satisfied are of the same species as those Clark ascribed to
A . granulatus. They were presented in 1904 by the Cape of Good Hope Govern-
ment under the following reference numbers :
Ref. No. 49 — 1 specimen Ref. No. 10639 — ° specimens
101 — 1 ,, 10723 — 2
? 171 — 3 specimens 10894 — 1 specimen
A search of the correspondence between the Marine Biologist at the Cape
(Dr. Gilchrist) and Professor Jeffrey Bell concerning the Echinoderms sent to the
British Museum from the Cape between 1897 and 1904 shows that each specimen
had a reference number and that lists, giving localities against reference numbers,
accompanied Dr. Gilchrist's letters. Some of the lists have been found, but not
any including one of the reference numbers above. One of them, however, is to
be found in Bell's paper on the Asteroidea (1905, p. 244). It is 10639 and the
accompanying data read : "Cape Natal W. by N. 4} miles, large dredge, 47 fms.,
sand and shells." Professor Bell named these specimens Astropecten pontoporeus.
It must, unfortunately, remain unknown exactly where the other specimens
came from.
There are twelve specimens from Australian waters : the "Challenger"
specimen ; the Barrier Reef Expedition specimen figured by Livingstone ; two
specimens from Broome received through Dr. H. L. Clark from the Museum of
Comparative Zoology ; one from 36 fathoms on the north side of Holothuria
Bank and four from Roebuck Bay (Admiralty) ; and three specimens from
"West Australia " (Australian Museum, Sydney).
A comparison of the two sets of specimens, the Australian augmented by
ample description and illustration in published papers, does not convince me that
they are of two species : —
NOTES ON ASTEROIDS IN THE BRITISH MUSEUM
487
Note. — One of the S. African specimens (Ref. No. 10639) is too distorted to measure.
There are differences, some of which are brought out in the table. The number
of superomarginal plates is greater in Australian than South African specimens :
of two specimens of R = 35 mm. the South African has twenty-two, the Australian
tvventy-nine ; a South African specimen of R=37 mm. has nineteen marginal
plates, while an Australian specimen of R=38 mm. has twenty-nine. The covering
of the superomarginal plates of the South African specimens is a little less dense,
that of the inferomarginal plates a little more dense, than those of Australian
specimens, but they are of the same nature ; and the difference in the infero-
marginal plate covering is not greater than that between Aru Islands and Thursday
Island specimens shown in Doderlein's figs, za and 3 (1917, pi. 17). The second
of the two marginal spines of the inferomarginal plates is in most, but not in all,
South African specimens bigger in proportion to the first than it is in Australian
specimens, being often two-thirds and sometimes three-quarters as long.
The only difference that I see in the general facies is that of the relative
proportions of the inferomarginal spines ; that the superomarginal plates of
Australian specimens encroach more on the abactinal surface than do those of
South African specimens ; and that two of the South African specimens have
spines on some of the superomarginal plates. ( )ne of them is remarkable in having
large spines, more than 2 mm. long, on the first superomarginal plates in some
488
D. DILWYN JOHN
interradii but not in others. The spines on the more distal plates of both specimens
are smaller, about half the length of the plates on which they occur. They are
broad based and pointed — i.e., sharply conical.
Clark (1923, p. 251) wrote : "One of the smaller African specimens shows six
superomarginal spines," but he does not state where they occurred. Their very
irregular occurrence in the two British Museum specimens is shown in the following
table :—
Many characters are shared in common by the two sets of specimens. The
paxillae, with all their variations of age, are essentially the same. A characteristic
feature is the raised rim which the outer spinelets form around the shorter central
granules and the unequal size of the latter (Doderlein's fig. 2a, pi. 17, 1917,
shows this very well.) The rim is least apparent in the very large Australian
specimen (R=86 mm.) in which all the spines are of roughly equal length and have
flattened polygonal ends ; but it tends to be present even there. The number of
spines or granules present on the largest paxillae varies from one or two central
and seven to ten peripheral in the smallest specimens, to ten to seventeen central
and fifteen to seventeen peripheral in those of medium size. In the largest
specimen (R=93 mm.) there are twenty-five to thirty in the centre and about
twenty-five around the edge. The shape of the larger paxillae where they are
crowded tends to be polygonal.
The characteristic spines which occur among the squamules on the ventral
surfaces of the inferomarginal plates are present in nearly all the British Museum
specimens, Australian and South African alike. They are flattened and blade-like
with an irregular outline, but each tapers to a point. Their occurrence is very
irregular. In smaller, and sometimes in medium-sized, specimens they are present
on the first plate only and lie in a row down its centre. In larger specimens they
occur too on neighbouring plates, but have moved to the aboral edge ; and in
very large specimens they may extend to the end of the ray (Doderlein, 1917,
pi. 14, fig. 1).
The order of the adambulacral spines is the same in both sets of specimens,
but the aboral spine of the second row is much less enlarged in South African than
in Australian specimens ; it is only slightly bigger than the adoral.
More South African specimens are required before it can be said whether the
form occurring there is best called a separate species. In the meantime it seems
NOTES ON ASTEROIDS IN THE BRITISH MUSEUM
489
wise to regard it as a new sub-species — subsp. natalensis, distinguished by its
smaller number of superomarginal plates, which may carry spines, and by other
differences, detailed above (in the covering of the marginal plates, the relative
proportions of the two spines of the inferomarginal plates, etc.), which are of
degree and not of kind ; and, even so, are not constant.
The Australian specimen of R=46 mm. which has the unexpectedly large
minor radius of 13 mm. has unusual numbers of actinal intermediate plates. In
two interradii there are five ; in the other three there are six, seven and nine
respectively. The nine are arranged as a chevron of seven enclosing a pair in
its angle.
Astropecten indicus Doderlein.
Astropecten indicus Doderlein, 1888, Zool. Jb. Abt. 1 : 828, pi. 31, figs. ia-d ; Koehler, 1910 :
27, pi. 4, figs. 8-15 ; D6derlein, 1917 : 146, pi. 14, figs. 4-40.
Astropecten koehleri Loriol, 1899, Mim. Soc. Phys. Genive : 19. pi. 1, fig. 5 ; Doderlein, 1917 : 122,
pi. 11, figs. 1— la.
Astropecten pleiacanthus Bedford, 1900, Proc. Zool. Soc. Lond. : 292, pi. 24, figs. ga-c.
There are in the collection ten specimens which appear to me to show that
Koehler (1910) was right in uniting A. koehleri with A. indicus, a conclusion with
which Doderlein (1917) did not agree. He thought koehleri to be sharply marked
off from indicus by (a) the possession of well-developed spines on the inner margins
of the more proximal superomarginals, and (b) the presence of well-developed
rows of spines along the distal edges of the inferomarginals. How he can have held
the first, (a), after Koehler's careful description of a series I do not know. The
second, (b), I find a more puzzling point.
The ten specimens are as follows : —
They show, in one direction, an even wider variation in the spination of the
superomarginals than did Koehler's series. No. 1, which is very juvenile, has no
trace of spines. Nos. 2 and 3 each has a small spine on practically every supero-
marginal ; they are inner spines on the first three plates and pass to the outer edge
on the fourth to sixth plate. No. 4 is similar except that on most arms the inner
490 D. DILWYN JOHN
spine is confined to the first two plates, and the outer series begins on the second
plate, which therefore has two spines. No. 5 is a dry specimen which has lost
many spines. It seems to have been similar to Nos. 2 and 3. One of its arms is
regenerated beyond the fourth superomarginal. The spines borne on the regener-
ated plates are relatively much bigger than those of the adult arm.
No. 6, the paratype of Bedford's A. pleiacanthus , is described below.
No. 7 has large inner spines on the first three to four plates, inner and outer
spines on each of the mid-arm plates, single outer spines on the last six or seven
plates. No. 8 is rather different, for the spines are very small. There is one to each
plate except that on some arms there are one to four unarmed plates between the
first three, which have inner spines, and the outer plates, which have spines on the
external edge. Some plates at the extremity of the arm are also unarmed.
Nos. 9 and 10 show spination greater than any Koehler described. No. 9 has
large single spines on the first three superomarginals. In mid-arm each plate has
up to six unequal spines or enlarged granules. They form an irregular row near
the distal edge with odd spines or granules standing proximal to it. Four or five
plates at the tip of the arm usually have but one, outer, spine. One arm is
regenerated beyond the third superomarginal. The regenerated portion bears one
spine on each of the twelve plates. No. 10 is similar but even more spiny, for most
of the first superomarginals bear two strong spines. For the largest specimen it
has a curiously low number of marginal plates (20).
The other characters of these specimens (except for the juvenile No. 1) are
those of indicus as described by Doderlein (1917).
No. 6 is different and it was long before I decided to treat it as of the same
species. (It is the smaller of the two specimens upon which Bedford based his
description of A. pleiacanthus and it is the one photographed by him.) It has
spines on all the superomarginals and they are bigger than those of any of the
specimens described above. Two spines occur, very irregularly on different arms,
on some of the plates between the third and the eighth ; the inner spine of the
pair is much smaller than the outer. In this there is no difference in kind from the
series described above ; but there is such a difference in the armature of the infero-
marginals. The first six to nine possess strong spines arising among the scale-
like spinelets of the ventral surface. There are three to four on the first, two on
the second, and, usually, one thereafter. They lie in the mid-line on the first
plate, near the distal edge of the others.
Koehler described these spines as constituting one of the characters of
A. indicus and shows them in figs. 11 and 12 of his pi. 4. In the other specimens
of the Museum series (with the exception of No. 1) there are enlarged spinelets in
corresponding positions on the first, second and sometimes on the third to fifth
plates ; but they are so different in size from those of No. 6 that they are not
immediately apparent as its spines are. To regard No. 6 as a separate species
because it has these spines would be to suppose that intermediate stages between
it and the others of the series do not occur. On the other hand, I cannot assert
that they do.
No. 6, however, has another difference. The other specimens of the series
(again excepting No. 1) have a rudimentary spine standing proximal to and above
the large inferomarginal spine. It is variable in size ; in some of the larger
specimens it is double. It is not present at all in No. 6.
NOTES ON ASTEROIDS IN THE BRITISH MUSEUM 491
Astropecten monacanthus Sladen.
Aslropecten monacanthus Sladen, 1883, J . Linn. Soc. (Zool.), 17 : 263 ; 1889 : 216, pi. 33,
figs. 7 and 8 ; pi. 37, figs. 10-12 ; Koehler, 1910 : 37, pi. 3, figs. 9-1 1 ; pi. 5, fig. 11 ; Doderlein,
1917 : 150, pi. 14, figs. 5-56 ; pi. 17, fig. 9 ; Fisher, 1919 : 74 ; Macan, 1938 : 336 ; H. L. Clark,
1938 : 63.
There are in the British Museum twenty- three specimens, including the type,
of this species. They come from Arabia, the Maldives, India, the Andamans,
Burma, the East Indies and Philippines, and (one large specimen) from Pin Tang
in China. Included in this number are seven juvenile specimens (R=9~i5 mm.)
the adambulacral spines of which do not have the characteristic shape of those
of the adult. They are regarded as monacanthus because they were taken with
adults and because most of them have the distinctive disk markings of the species
(see below). There are, on the other hand, two juvenile specimens (R=n and
15 mm.) in which the outer adambulacral spines are widened and flattened as in
the adult. The largest specimen previously recorded was of R=53 mm. (Macan).
There are specimens of R=62 and 91 mm. in the Museum.
If nothing else were said of this series, this should be recorded : that ten
specimens of it have spines or enlarged granules on the superomarginals. Koehler
described their occurrence and figured them. But two more recent descriptions
of the species have seemed to deny their presence : Doderlein's diagnosis says
superomarginals "without spines," although on another page (p. 146) he notes
that there may be spines ; and Fisher writes of "unarmed granulate supero-
marginals."
The spines may be very small — no more than enlarged granules ; but they
may, on the other hand, be anything up to half the length of the plate from which
they arise. Their presence is not, so far as this series shows, correlated with size.
The largest specimen (R=9i mm.) possesses them on some plates near the end
of the arms, but they are very small. A specimen of R=52 mm. has none. The
largest are on a specimen of R=37 mm. They occur on four specimens of
R=9-i4 mm., on three of them on the first and second superomarginals only.
It may be significant that all the specimens on which they occur come from
Indian seas.
Their occurrence is very irregular. They may be only near the end of the
arm ; or from the fourth or fifth plate to the end ; or only on mid-arm plates ; or
irregularly anywhere ; or (in the juvenile specimens referred to above) on the
first and second plates only. They are sometimes double.
Fourteen of the specimens, including five of the juveniles, bear characteristic
markings on the dorsal side. (All are in spirit.) A dark line lies parallel and near
to the margin of the proximal part of each arm on each side, those of contiguous
arms being joined around the arm angle to form a chevron. In some of the larger
specimens there is also a dusky ring around the centre of the disk from which a
band runs down the centre of each ami. The largest specimen has this pattern
alone. There may also be dark bands across the arms two-thirds of the way down.
Twelve of the specimens, including some that are fairly large (up to
R=62 mm.), possess an epiproctal cone.
There is nothing to add to Fisher's account of the adambulacral armature
except that in the Pin Tang specimen the adoral spine of the outer series is more
broadly spatulate than in any figure, or in any specimen that I have seen.
The largest paxillae of the biggest specimen have up to forty central and
thirty peripheral granules. The number of superomarginals of this specimen,
492 D. DILWYN JOHN
being forty-seven, appears very high. Four specimens of between R=42 and
45 mm. have twenty-four to thirty superomarginals ; a specimen of R=62 mm.
has thirty-six.
Astropecten zebra Sladen and Astropecten hartmeyeri Mtiller and Troschel.
A stropecien zebra Sladen, 1883. /. Linn. Soc. (Zool), 17 : 261 ; 1889 : 212, pi. 36, figs. 3 and 4 ;
pi. 39, figs. 7 and 9 ; Doderlein, 1896 : 306, pi. 18, figs. 31 and 31a ; Loriol, 1899 : 9 ; Bell,
1904 : 149 ; Koehler, 1910 : 44 ; Brown, 1910 : 29 ; Doderlein, 1917 : 155, pi. 14, figs. 8 and
8a ; Clark, H. L., 1921 : 27 ; Livingstone, 1932 : 242 ; Clark, H. L., 1938 : 63.
Astropecten coppingeri Bell, 1884, "Alert" Report : 132.
Astropecten zebra var. rosea Sladen, 1883, /. Linn. Soc. (Zool.), 17 : 263 ; 1889 : 214.
Astropecten zebra var. sibogae D6derlein, 1917, Siboga Exped. Monog. 46a : 156, pi. 6, fig. 4 ; pi. 14,
figs. 7 and ya.
Astropecten hartmeyeri Doderlein, 1917, Siboga Exped. Monog. 46a; 156, pi. 5. fig. 8; pi. 14, figs.
6-6c ; Clark, H. L., 1938 : 62.
Astropecten hartmeyeri subsp. siamensis Ddderlein, 1926, K. svenska Vetensk.-Akad. Handl., (3) 2 :
5, pi. 1, fig. 2.
H. L. Clark (1938, p. 63), in writing of A. hartmeyeri, states "whether hart-
meyeri is really distinct from zebra Sladen seems to me rather doubtful, but the
only specimens of zebra available for comparison are too young to be of real
service. Larger collections from North-eastern Australia are necessary for a
final decision."
In the British Museum collection there is a series of twenty-four specimens,
a description of which may help to solve the problem. They are as follows : —
I. From North-eastern Australia.
A. Torres Strait, Challenger Station 186. Four specimens, the types of
A. zebra.
B. Torres Strait, Challenger Station 187. Three specimens, the types of
the var. rosea.
C. Thursday Island, Torres Strait, "Alert" Collection. One specimen, the
type of A. coppingeri.
D. Torres Strait, "Alert" Collection. Three specimens, labelled coppingeri.
Di. Prince of Wales Channel, Torres Strait, "Alert" Collection. One
specimen, labelled coppingeri.
E. Holothuria Bank, Admiralty Collection. Two specimens.
F. Holothuria Bank, 34 fms., Admiralty Collection. Three specimens.
G. North-west side of Holothuria Bank, 39 fms., Admiralty Collection.
Two specimens.
H. South of Cape Kimberley, 4 fms., Great Barrier Reef Expedition.
Two specimens.
II. From Western Australia.
J. Magnetic Shoal, off Cossack, Admiralty Collection. Two specimens.
K. Broome. One specimen of A. hartmeyeri, presented by Dr. H. L. Clark.
In the following discussion the specimens will sometimes be referred to as of
A, B, C, etc. The variation described is tabulated, so far as is convenient, in
the table on p. 495.
Doderlein distinguishes hartmeyeri from zebra by (1) the fact that its supero-
marginal plates beyond the first three to six nearly always bear small spines on
the outer edges, whereas those of zebra do not ; and (2) the fact that it has
NOTES ON ASTEROIDS IN THE BRITISH MUSEUM 493
pedicellariae on the first inferomarginal plates in place of the large marginal spine ;
small pedicellariae on the lower distal edges of some of the superomarginal
plates ; and pedicellariae among the paxillae of the dorsal surface, usually near
the marginal plates ; whereas zebra has them on the adambulacral and actinal
intermediate plates only.
The occurrence of the superomarginal spines and pedicellariae is described as
extraordinarily variable.
Discussion of Series.
Superomarginal spines. In the largest of the type specimens (A) the inner
spines of the arm angle extend to the sixth plate, becoming progressively smaller.
They reach the third plate in the types of the var. rosea (B). In one of the largest
specimens (R=23 mm.) of C and D they extend to the third plate on some rays ;
in the smallest (R=n mm.) there are very small spines on the first plates only.
In none of these specimens are there spines on the outer edges of the supero-
marginal plates. It is otherwise with most of the specimens of Di, E, F, G and H.
In Di, by far the largest specimen in the series (R=I07 mm.), there are outer
spines on the fourteenth and fifteenth plates on one side of one arm, none
on the other side nor on any other arm. In both specimens of E there are spines
on all the superomarginal plates except one or two at the ends of the arms ; they
move from the inner to the outer edge on the third and fourth plate. In the three
specimens of F they extend to respectively, in order of size, the twelfth, seven-
teenth and sixteenth plates. There are, here and there, gaps in their occurrence :
in the smallest specimen they are wanting from the third and fourth plates on
most arms. In one of the specimens of G there are inner spines on the first and
second, and none on any other plates ; in the second specimen there are small
outer spines from the fourth or fifth to the seventh, eighth or ninth plates. H are
the specimens listed by Livingstone (1932). One has only large spines on the inner
edges of the first and second plates ; the other has, in addition, small outer spines
occurring irregularly as far out as the eleventh plate ; some plates, including the
first, have two spines.
In the larger specimen of J (R=46 mm.) small outer spines extend to the
eighteenth superomarginal plate on some rays. They are not always continuous.
The change from inner to outer edge takes place on the fourth to seventh plate.
In the smaller specimen (R=i2 mm.) the only spines present are very small ones
on the inner edges of the first plates in some interradii. In the large specimen
(A. hartmeyeri, K) only one or two of the plates of some of the margins are armed
with very small outer spines. Examples on different margins are : the ninth and
tenth plates, the twelfth and fourteenth plates, the twelfth plate.
Pedicellariae. In the three larger type specimens of zebra (A) the first supero-
marginal plate bears in place of the large marginal spine a small pedicellaria com-
posed of three to six spines ; and in one of them there are eight small pedicellariae
on the dorsal surface among the paxillae near the marginal plates. These, un-
described by Sladen, are among the distinguishing features of hartmeyeri. In the
smallest specimen (R = n mm., Sladen's "young phase") the marginal spine of the
first inferomarginal plate is present and there is no pedicellaria. The types of
rosea (B) resemble those of zebra in the occurrence of pedicellariae except that none
possesses any in the paxillar field, and the largest has none on the adambulacral
plates. In C and D the large spine of the first inferomarginal plate is missing ; it
is replaced by a pedicellaria in the larger specimens, but not in the smaller. There
novit. zool., 42, 3. 1948. 86
494 D- DILWYN JOHN
are no pedicellariae among the paxillae. The smallest (R=u mm.) has pedicel-
lariae only on the actinal intermediate plates. In the large specimen Di there are
pedicellariae on the actinal intermediate, adambulacral and first inferomarginal
plates ; they are also present in small numbers in the paxillar areas near the
marginal plates and on the lower distal corners of some superomarginal plates.
In the group of specimens (E, F and G) from the Holothuria Bank the first
inferomarginal plate carries a spine and not a pedicellaria. Otherwise they vary.
The specimens of F have no pedicellariae at all, not even on the actinal inter-
mediate plates. Those of G have the spines of the actinal intermediate plates
grouped in a pedicellaria-like way and there are what appear to be incipient
pedicellariae on one or two adambulacral plates of one specimen. The specimens
of E, which are not very much bigger than those of G and are smaUer than some
of those of F, are rich in pedicellariae. They are present on the actinal inter-
mediate, and are numerous on the adambulacral, plates. They are also found on
the paxillar area : in the smaller there are about ten, all near the marginal plates ;
in the larger there are over forty grouped for the most part near the marginal
plates at the arm angles. The Barrier Reef Expedition specimens (H) have neither
large marginal spines nor pedicellariae on the first inferomarginal plates and no
pedicellariae in the paxillar area. Both have them on the actinal intermediate
plates, but in the bigger only are they present on a very few adambulacral plates.
Turning to the specimens from Western Australia : the large specimen from
Cossack (J) has pedicellariae on actinal intermediate and adambulacral plates, in
place of spines on the margins of the first inferomarginal plates, and on the dorsal
surface (about fifty in number) mostly near the marginal plates ; and in addition
there are small, low pedicellariae on the lowermost and distal-most corners of
some superomarginal plates, more especially those in the proximal part of the
arm. The smaller specimen (R=i2 mm.) has pedicellariae on the actinal inter-
mediate plates and in the arm angles.
The Broome specimen has pedicellariae on the actinal intermediate, adam-
bulacral and first inferomarginal plates as well as many small ones on the dorsal
surface near the marginal plates. It has none on the superomarginal plates.
Other Characters. The general facies and the remaining characters of all the
specimens are the same. Three sets show a peculiar colour : those Sladen named
var. rosea are pink ; three specimens (F) from the Holothuria Bank are a rich
straw yellow ; the Barrier Reef Expedition specimens are very dark grey. Yet
these, as well as the remainder which are the usual museum colour, show the dark
pattern characteristic of zebra. Doderlein created a variety of zebra, var. sibogae,
on one specimen (R=i8.5 mm.) with the aboral spines of the inferomarginal
plates missing. I find them to be wanting in all but the first two or three proximal
plates of specimens below R=20 mm. It is worth noting that Doderlein describes
his specimen of var. sibogae as having a pedicellaria in place of a spine on the outer
edge of the first inferomarginal plate ; that De Loriol saw no pedicellariae in his
specimens from Sumatra ; that Rudmose Brown mentions none in examples
from the Mergui Archipelago ; that Koehler found none on the adambulacral
plates of specimens from Madras, Burma and the Andamans. Doderlein does not
describe pedicellariae in his single specimen of the subspecies of hartmeyeri,
siamensis from the Gulf of Siam. There are no specimens from these regions in the
British Museum collection.
There are three small specimens with no more precise locality than "Australia"
which have not been described. Bell referred to them (1884, p. 133). R=io.5, 17
NOTES ON ASTEROIDS IN THE BRITISH MUSEUM
495
and 20 mm. None of them has outer spines on the superomarginal plates. The
smallest has an inner spine on the first plate only, the second on the first and second,
and the largest on the first, second and third. The smallest has no pedicellariae ;
but the first inferomarginals are without marginal spines so that it may be sup-
posed that pedicellariae would have developed there later. That with K = ij mm.
has pedicellariae on the actinal intermediate plates and they are incipient on
the adambulacral plates. There are pedicellariae on the actinal intermediate,
adambulacral and first inferomarginal plates as well as on the paxillar area of the
largest specimen.
Conclusions. Though it cannot be said with certainty that hartmeyeri and
zebra are one, it has been shown that the characters used to separate them do not
hold good, and are, apart from that, extremely variable. This account is prepared
as a contribution to the final solution of the problem.
TABLE OF VARIATIONS
X denotes M present."
496 D. DILWYN JOHN
Astropecten brevispinus Sladen
(Plate I, figs. 3 and 4.)
Astropecten brevispinus Sladen, 1883, /. Linn. Soc. (Zool.). 17 : 249 ; 1889 : 198, -pi. 33, figs. 1 and 2
pi. 37, figs. 1-3 ; Goto, 1914 : 221 ; Doderlein, 1917 : 60.
Sladen's description of A. brevispinus is excellent and complete except that
he described only the spinelets of the actinal intermediate plates and not the
plates themselves. They are of exceptional interest.
In the adult specimen (R=32 mm.) there are four chevrons in each inter-
radius. In the first there are nineteen plates on either side reaching to the twelfth
inferomarginal, which is about three-quarters of the way down the arm. In the
second and third chevrons there are on either side seven and four plates reaching,
respectively, the fourth and second inferomarginal. The fourth chevron is of six
plates, three on either side ; they are in contact with the first inferomarginal.
The larger actinal intermediate plates in the interradial area are strongly arched
in the middle.
In the "young phase" (R=i6.5 mm.) there are three chevrons having respec-
tively six, three and single plates on either side. The first row reaches the third
inferomarginal ; the second and third are in contact with the first inferomarginal.
The possession of these plates, running far down the arm in the adult, might
cause the species to fall nearer Persephonaster than Astropecten in Fisher's key
(1911, p. 39). But Goto (1914, p. 221) could not be certain of their presence, and
did not explicitly surmise it, when he placed the species in Persephonaster.1 He
did so because of the close relations be saw between it and his Persephonaster
triacantkus. He placed the latter in Persephonaster despite its "very strong external
resemblance to Astropecten" because, above all, it possessed an anus. "The only
single character of decisive value that distinguishes the genus (Persephonaster)
from Astropecten is the presence of the anus," he wrote (p. 221). On the other hand,
Fisher (1911, p. 38) says of the anus that "No more unstable character can be
conjured up to separate Astropecten and its near relatives from Plutonaster,
Dytaster, or Dipsacaster . . . there is good evidence that the character is variable
within a genus (Astropecten, etc., etc.)." I cannot see an anus in Astropecten
brevispinus.
There is no doubt that Sladen's Astropecten brevispinus and Goto's Perse-
phonaster triacanthus are very closely related. Their general facies is undoubtedly
more Astropecten than Persephonaster and so are the nature of the adambulacral
and mouth-plate armature, and of the paxillae. It therefore seems better to call
them so.
Astropecten mesactus Sladen
(Plate I, figs. 5 and 6.)
Astropecten mesactus Sladen, 1883, /. Linn. Soc. {Zool.), 18 : 267; 1889 : 219, pi. 34, figs. 5 and
6 ; pi. 38, figs. 7-9 ; Doderlein, 1917 : 59.
non Studer, 1884, Abh. preuss. A had. Wiss. : 46.
Sladen described this as a "very abnormal Astropecten" which he admitted
"not without hesitation" into the genus. "The character of the abactinal paxillae
(the pedicel being represented only by a broad tubercular eminence of the plate)
and the great development of the actinal interradial areas are altogether unlike
any Astropecten with which I am acquainted" (1889, p. 221). Sladen's description
1 Doderlein (1921, p. 26) gives a list of the species of Persephonaster known at that time. He
omits the three species described by Goto (1914).
NOTES ON ASTEROIDS IN THE BRITISH MUSEUM 497
of it is of his usual excellence except that he does not fully describe the actinal
intermediate plates. They extend two-thirds of the way down the arms.
There are four clearly marked chevrons. The first has twenty-two plates in
each limb, reaching the twelfth inferomarginal. The second, with nine plates
(including a single plate at the apex), and the third with four plates on each side,
reach respectively the fourth and second inferomarginals. The fourth chevron is
of four plates, two in each limb, the outer touching the first inferomarginals.
There appear to be no other references to this species than those given above.
Astropecten pectinatus Sladen
(Plate II, fig. 5.)
Astropecten pectinatus Sladen, 1883, /. Linn. Soc. (Zoo!.), 17 : 251 ; 1889 : 202, pi. 33, figs. 3
and 4 ; pi. 37, figs. 4-6.
Description. There are ten specimens in the Museum collection. Eight are
"Challenger" specimens as follows : Port Jackson, five ; St. 161, off entrance to
Port Philip, two ; St. 162, Bass Strait, one. The other two, dried specimens in
good condition, were collected by J. B. Wilson, Esq., at Port Philip Heads in or
before 1885.
I find Sladen to have been wrong in his description of the actinal intermediate
plates, which is as follows : "The actinal intermediate plates, which are small and
very few in number, are confined to the immediate interradial area. The spinelets
that cover them are small, more or less subspatulate in form, and radiate apart."
One of the smallest specimens (Port Jackson, R=28 mm., r=8 mm., with
eighteen superomarginals) has two rows of actinal intermediate plates on either
side. The first extends far down the arm to the seventh inferomarginal and con-
sists of thirteen plates. The most distal are very small and cannot be seen until
the spines of the adjacent inferomarginals and adambulacrals are removed. The
second row is of three plates (including the unpaired plate at the apex of the
chevron formed by the two rows in one interradius) . The actinal intermediate
plates carry tufts of spinelets. Those of the inner row are of smaller spines and
fewer in number (seven to eight) on the innermost plates than on those farther
out in the interradial area. The plates of the second row have a greatly enlarged
and flattened central spinelet ; it is as big as those on the aboral edge of the
first inferomarginal plates.
In the smaller specimen from Port Philip Heads (R=35 mm., r=io.5 mm.,
nineteen superomarginals) there are three rows of actinal intermediate plates on
either side. The first is of twenty-three plates and extends to the thirteenth
inferomarginal. The second is of four plates. The third "row," if it may be so
called, is represented by one minute plate. The plates of the second row and some
of those of the first row in the interradial area (not down the arms) have the
greatly enlarged spinelet described in the specimen above ; on some plates it is
double.
There are similarly three rows of plates in the specimen upon which Sladen
based his description (R=48 mm., r=i4 mm., twenty-one superomarginals). But
the first is of only sixteen plates, reaching the ninth inferomarginal. The second
and third rows are of four and single plates respectively.
The largest specimen in the collection is the second from Port Philip Heads
(R=5& mm., r=i6 mm., twenty-one superomarginals). It has four rows of
actinal intermediate plates on either side. There are no less than twenty-seven
498
D. DILWYN JOHN
plates in the inner row, reaching the fourteenth inferomarginal plate. The second
row is of eight plates and it too extends into the arm, as far as the fourth infero-
marginal. The third and fourth rows are of three and two plates respectively,
those of the latter being very small. The plates of the second and third rows of
this specimen, and those of the second row of the preceding specimen, have an
enlarged blade-like spine like that described for the smaller specimens above.
The figures in the above descriptions are shown in the following table : —
Discussion. Astropecten pectinatus has been shown to have a large number of
actinal intermediate plates ; and the number of plates and of rows increases with
size. It is clear that had Doderlein known this he would have assigned the single
specimen from Tasmania upon which he based his species schayeri to pectinatus ;
for (1917, p. 62) he wrote : "Ich wiirde beide Formen zu einer Art vereinigen,
wenn nicht Sladen ausdriicklich angegeben hatte, dass A. pectinatus nur sehr
wenige Ventrolateralplatten besitzt."
For all that, I do not find it easy to declare schayeri a synonym of pectinatus ;
for Doderlein describes the adambulacral armature of the former as being so
different from that of the latter (which is essentially as Sladen described it) ; and
Clark (1938, p. 66) emphasises the description. I think, after a careful comparison
of my largest specimen of A. pectinatus and Doderlein's fig. 1, pi. 7 of A. schayeri,
of nearly the same size, that Doderlein in describing the spines on the ventral
surface of the adambulacral plate as up to seventeen in number may have taken
the spines of the adjacent actinal intermediate plate into his count. They would
not be equal in size — nor do they appear so in his photograph. This is assuming
that there are more plates in the first row than he describes ; which I think is
possible, for the distal plates in A . pectinatus are small and hard to see unless all
neighbouring spines are removed.
I cannot agree with Clark (loc. cit.) when he says that "the difference in the
large spines of the inferomarginal plates (of pectinatus and schayeri) is very great."
I find them to be quite similar. The inferomarginal armature of A. syntomus
(H. L. Clark, 1928, p. 372), with its double fringe of spines, is clearly different, but
the species must be closely related to pectinatus. It is worth adding that its actinal
intermediate plates are similarly armed ; as are those of Doderlein's A. schayeri.
The occurrence of spines on the superomarginal plates is, on the whole,
regular. But on one arm of one specimen (R=32 mm.) the third to seventh plates
are unarmed ; in another (R=48 mm.) many plates are unarmed, especially in
mid-arm ; in others an odd plate here and there has no spine. In the largest
specimen (R=56 mm.) some of the plates in the arm angle have two spines.
NOTES ON ASTEROIDS IN THE BRITISH MUSEUM 499
Astropecten sphenoplax Bell
(Plate II, figs. 1-3.)
Astropecten sphenoplax Bell, 18920, Sci. Proc. R. Dublin Soc. 7 : 522, pi. 22 ; 1892* : 68, pi. 9 •
Koehler, 1909 : 42, pi. 17, fig. 8 ; Mortensen, 1927 : 59.
Bell records seven specimens. I find six in the collection, one of which is dry.
Bell gave measurements of only three and did not include the largest among them.
The following are my measurements of these : —
The paxillae have not been described. They are small and crowded near the
centre of the disk and at the ends of the arms. On the proximal parts of the arms
they are arranged in transverse rows on either side, but are irregularly disposed
in the centre. In the distal part of the arm the arrangement is irregular through-
out. The larger paxillae bear forty to fifty fine spinelets with no difference
between those which are peripheral and those in the centre. In one specimen only
(the dry specimen) one central spinelet of a small proportion of the paxillae on
the outer part of the disk and in the proximal half of the arm is enlarged to many
times the size of its fellows. When shorn of their spinelets the pedicels of the
paxillae are seen to be low and rounded, not hour-glass shaped.
The superomarginal plates do not meet near the tips of the arm as Bell
states, though they come very near to doing so. Koehler described his two
specimens as having a spine, or the scar of a spine, on each superomarginal. It
is not so in these specimens : spines are of irregular occurrence and vary from one
individual to another. They are most frequently absent from the arm angles and
if present there are very small. Where they occur they are largest, up to 2 mm.
long, in the region of the fourth to tenth superomarginals and thereafter become
smaller. They are most frequently missing on the outer parts of the arms. In
the largest specimen they are present only on the fourth and fifth plates of some
arms, and are represented by tubercles on a few of the succeeding plates.
Apart from their larger spines the marginals are densely covered with very
fine spinelets which stand out at right angles to the plates and are not flattened
or scale-like.
The actinal intermediate plates are more numerous and extend farther into
the arms than previous descriptions have shown. Those of three specimens have
been examined. In the smallest (R=ca. 41 mm.) there are three rows, the first
of ten plates extending to the fifth inferomarginal ; the second, third and fourth
of five, three and single plates respectively. The dry specimen (R=52 mm.) has
twelve plates in the first row reaching the seventh inferomarginal ; the second
row has five plates, and the third has four plates on one side, five on the other.
Five plates irregularly arranged in the angle of the chevron formed by the third
rows represent the fourth rows.
500 D. DILWYN JOHN
It is curious that in the largest specimen (R= ca. 72 mm.) there are only
three rows. The first is of fifteen plates reaching the seventh inferomarginal, the
second of five, the third of two plates.
Expressed in another way, the actinal intermediate plates reach, in the three
specimens examined, between a third and half-way down the arms.
Astropecten triseriatus Muller and Troschel
Astropecten triseriatus Miiller and Troschel, 1843, Arch. Naturgesch., 9 (1) : riS ; Doderlein, 1917 :
125, pi. 5, figs. 2-3 ; pi. 11, figs. 5, 6-6a ; Clark, H. L., 1938 : 66.
Astropecten arenarius Perrier, 1876, Arch. Zool. exp. gin., 5 : 286.
Astropecten triseriatus myobrachius Fisher, 1925. Bull. Bishop Mus., Honolulu. 27: 69, pi. 5. fig. B.
The only specimens of this species previously described are as follows : —
From south Western Australia : the type (Muller and Troschel and Doder-
lein) ; part of one arm (H. L. Clark).
From north Western Australia : one specimen (Doderlein).
From an unknown locality : one specimen (Perrier).
From the Hawaiian Islands : one specimen of the subspecies myobrachius
(Fisher).
It is therefore worth while to give some details of the two specimens in the
British Museum collection, as well as a full account of a third which is described
as a new subspecies. The measurements of all the known specimens, including
those of the new subspecies, are brought together in the table (p. 501).
Specimen from Bass Strait, R=54 mm., r=i2 mm., thirty-one supero-
marginals. Only one arm is of normal length ; each of the others was broken
and is in part regenerated. None of the paxillae have enlarged central spinelets.
There is, as a rule, only one spine on the first four superomarginal plates ; it is
slightly longer than one plate, being 2 mm. long. The succeeding plates have two,
three or — towards the end of the arm — one spine. Where there are two or three
the outermost is the larger, being as long as one plate. The marginal spine of the
inferomarginal plates is as long as three plates. There are two inside it as described
by Doderlein, but the first of them is more than half as long as the marginal spine.
Only the more proximal of the inferomarginal plates have one or two larger spines
along the aboral edge. There is only one actinal intermediate plate on either side.
The madreporite is 3 mm. long ; a few small scattered spines arise from its
surface.
Specimen from an unknown locality, R=84 mm., r=i7 mm., thirty-seven
superomarginals. In some arm angles the first pair of superomarginals carry only
one spine each ; in others there is a second and smaller outer spine. The remaining
plates carry three, sometimes four, spines, of which the lowermost is the longest ;
it is nearly as long as two plates in the middle part of the arm, but shorter towards
the end. The three large spines of the inferomarginal plates have some of the
spinelets at their bases enlarged to spines. The row of aboral spines, three, two
or one in number, extends to about the eighteenth plate. There are two actinal
intermediate plates on each side. The madreporite is 4 mm. long with a few small
scattered spines.
NOTES ON ASTEROIDS IN THE BRITISH MUSEUM
501
MEASUREMENTS
Astropecten triseriatus fijiensis subsp. nov.
(Plate II, fig. 4.)
This new subspecies is based on a single dry specimen in good condition from
the Fiji Islands which came to the Museum in 1862 or earlier. There are no details
of depth, etc.
R=66 mm., r= 17 mm. R=3.o, r. Number of superomarginal plates,
twenty-five.
This subspecies differs from A. triseriatus in the smaller number of supero-
marginal plates, the great development of the first pair, and the unarmed condition
of the second pair, of superomarginal plates.
The larger paxillae have one to six central spinelets which are often of very
unequal size, and nine to twelve peripheral spinelets. Many of those on the disk
and near the base of the arms have one spinelet, usually but not always the central
one, strongly enlarged, or one or two less markedly enlarged. There are about
fourteen paxillae across the base of the arm.
The most striking feature of the superomarginal plates is the great size of
the first pair : they rise strongly above the level of the remainder, swelling out
as they do so, so that their dorsal width is many times their width at the base.
And each carries a larger spine, 5 mm. long, than any other plate. With one
exception the second superomarginal plates are narrow and unarmed. The third
and fourth, or the third, fourth and fifth, plates each carries one large spine about
4 mm. long. The remaining plates have smaller spines. There are usually three,
but sometimes two or four. They are irregular in size and may be so in arrange-
ment— i.e., they are not always in a straight line. The largest is about as long as
one plate. The spinelets at the base of the spine are enlarged.
The inferomarginal plates are covered with long spinelets with narrow bases
which gradually widen to rounded flattened ends. They are small in the furrows
between the plates, greatly enlarged around the bases of the spines described
below. Each plate carries on the outer margin one strong pointed spine
about as long as two plates. A similar but somewhat smaller and more flattened
spine stands inside and slightly aboral to it ; it is the first of an aboral row of
three or four. The remaining two or three are much shorter and are flattened,
pointed and blade-like. In the proximal part of the arm there is frequently a
second spine standing on the adoral side of the outermost spine of the aboral row ;
it is about half as long as its neighbour.
The adambulacral armature in the proximal part of the arm consists of three
long and equal furrow spines followed by three similar pairs of spines, one behind
the other, each a little smaller than the last ; the aboral spine of the first pair is
as long as the furrow spines, the adoral is slightly smaller. Farther out on the arm
there are only two pairs of spines behind the furrow series.
502 D. DILWYN JOHN
The combined mouth plates bear a pair of large spines at the inner angle ;
each is followed by five smaller spines along the furrow margin. The strongly
raised central portion is lenticular in shape and carries nine or ten spines on
either side, the space between them being free of spines, except on one plate
where one occurs.
There are two actinal intermediate plates on either side, each with a group
of nine to twelve long spines similar to those on the inferomarginal plates.
The madreporite is 3 mm. long and bears a group of low spines.
Astropecten michaelseni Koebier.
Astropecten michaelseni Koehler, 1914, Beitr. Meeersfauna Westafr., 2: 144, pi. 4. figs. 3-6, 8-1 1 ;
pi. 5. figs. 1. 2, 13. 16-17.
Astropecten dahomensis Doderlein, 1917, Siboga Exped. Monog. 46a : 77, pi. 16, figs. 3-3^ ; text-fig. Q.
There can be no doubt that Koehler's and Doderlein's species are the same.
I can find no reference to either of them since the original descriptions. There are
twelve specimens in the Museum collection, eleven of which were collected in
recent years on the Gold Coast by Dr. F. R. Irvine and Miss V. J. Foote of
Achimoto College, Accra. The twelfth came in 1890 and is from Gambia and so
extends the previously known range, which was from 7 deg. S. to 9 deg. N.
There is little to add to Koehler's excellent description based on nearly fifty
specimens. Two of the specimens, the one from Gambia and the other from the
Gold Coast, are bigger than the remainder, having each R=63 mm. In neither
are the central granules on the paxillae enlarged as in all the other, smaller,
specimens. But they differ in the arming of the superomarginal plates. In the
Gambia specimens there are no spines. On the ninth and succeeding plates an
enlarged granule, circular in outline, occupies the position normal for the external
spine. It rises hardly at all above the level of its smaller fellows until it reaches
the end of the arm. The Gold Coast specimen of the same size has strong spines;
on many plates, including some in the arm angle, there are three.
A specimen of R=56 mm. shows nine to ten actinal intermediate plates,
reaching to the sixth inferomarginal, on either side.
A note attached to a large specimen from Labadi describes it as having been
salmon red in life, and the species as being common along the coast. Some, if not
all, of the specimens come from rock pools. A batch of five taken at Elmina are,
in spirit, a very dark brown with tube-feet deep purple.
Astropecten antillensis Liitken.
Astropecten antillensis Liitken, 1859, Vidensk. Medd. naturh. Foren. Kbh. 1 : 47 ; Doderlein. 1917 :
105, pi. 2, figs. 9-10 ; pi. 9. figs. 7 and 8 ; Clark, 1933 : 16 ; Boone, 1933 : 75, pis. 31-32.
Sladen in the Challenger Report (p. 198) records A. brasiliensis from "Off
Bahia. Depth 7 to 20 fms." The jar so labelled in the Museum collection con-
tained nine specimens, of which four were brasiliensis, but the remaining five are
undoubtedly A. antillensis Liitken. Another jar contains two specimens of
antillensis labelled "from ? Bahia" which were presented in 1903 by the Earl of
Crawford. The species does not previously appear to have been recorded from
anywhere but the West Indies.
Another specimen (R=35 mm.), which came to the Museum in 1848 from
"S. America," is a typical A. antillensis except that there is a band of paxillae,
encircling the area of small paxillae on the centre of the disk, in which each has
a single central spinelet which is considerably enlarged.
NOTES ON ASTEROIDS IN THE BRITISH MUSEUM 503
Astropecten brasiliensis armatus Gray
(Plate III, figs. 1 and 2.)
Astropecten armatus Gray, 1840, Ann. Mag. Nat. Hist., 6 : 181 ; Fisher. 191 1 : 56, pi. 5. figs. 1 and 2 ;
pi. 7, figs. 3 and 6 ; pi. 50, fig. 4 : pi. 51. fig. 3.
Astropecten brasiliensis armatus Doderlein, 1917. Siboga Exped. Monog. 46a : 84 ; with earlier
references and synonymy: 170.
Description of Type Specimen. R=65 mm., r= 14 mm., R : r=4.6. The larger
paxillae have seven to nine central and twelve to fifteen peripheral spinelets.
The central spinelets are globose and unequal in size, the larger being slightly
bigger than those of the periphery, which are equal. None of the central spinelets
are fused together.
There are twenty-three superomarginal plates which are large and high,
particularly in the arm angle. The first bears a spine over 4 mm. long on its inner
edge. With some exceptions the second and third plates each bear a much smaller
spine on or near the inner edge. A series of strong outer spines begins on the second
or third plate and extends to, or to within one or two plates of, the end of the ray.
On a number of the second and third plates there are therefore two spines ; the
outer is stronger than the inner. Double spines occur irregularly and infrequently
farther out on the ray. The general surface is covered with upstanding spinelets
which are largest, as big as those of the paxillae, on the abactinal surface ; they
become smaller near the lower margin and fine and capillary towards the fasciolar
grooves.
The inferomarginals project appreciably beyond the superomarginals. They
are thickly covered with short flattened spinelets with truncated ends. They are
finer and very numerous on the adambulacral margin and slender and capillary
on the edges of the fasciolar grooves. There are two large spines on the outer
margin, the larger above and slightly adoral to the smaller. The larger may be as
long as three plates, the smaller slightly over half as long. These spines are not,
or are only slightly, flattened. Smaller spines arise around their bases and a row
of two or three spines, about as long as one plate, arises from the aboral margin
of the plate. There is no corresponding adoral row, but there is frequently a
single spine near the adoral edge and the adambulacral plates.
There are three furrow spines, of which the middle is the strongest and longest.
Their ends are faintly scooped in that each has a short longitudinal groove on the
under side. In the second row there is one enormous blade-like spine nearly
3 mm. long and, adorally to it, a spine less than half its length which is slender.
The end of the large spine is, in the proximal part of the ray, enlarged as shown
in Fisher's pi. 7, fig. 3, but not so strongly as in his pi. 51, fig. 3. It is very faintly
scooped at the end. In the "third row" there are three to five slender spines about
equal to the adoral member of the second row. They may be in a group or a row.
The preservation of the mouth-plates does not permit of an accurate descrip-
tion cf the armature ; what can be seen of it appears to agree with Fisher's
description.
There are four actinal intermediate plates on either side. They carry groups
of slender spinelets, but do not have the larger spines described in some specimens
by Fisher.
The madreporite is about 4 mm. in diameter and is separated by one row of
paxillae from the superomarginals. Very small spines arise from the ridges of
its striations.
504 D. DILWYN JOHN
Discussion. Gray describes under armatus a variety pulcher. The locality he
gives may, both in the 1840 paper and in his Synopsis (1866), be read as applying
only to the variety, and Doderlein (1917, pp. 169 and 170) has so read it. On the
other hand, since it is his custom elsewhere throughout the 1840 paper and the
Synopsis to give localities or to indicate where they are not known that it is so,
the locality of Puerto Portrero may be read as applying both to armatus and its
variety. It appears to me that Fisher (1911, p. 60) has done this. I also do so ;
and I think I am right because of evidence afforded by the specimen described
above. It is accompanied by a very old label reading "Puerto Portrero, C.A.,
sandy bottom, 9 fms." ; it has inferomarginals which are produced beyond the
superomarginals ; it has a second label in the writing of E. A. Smith (who was in
charge of the collection from 1867 to 1878) which reads " Astropecten armatus
Gray (type)" and the initials "E. A. S." Because the variety pulcher had "the
under series of marginal tubercles not produced," and because Smith stood much
nearer to the event than I stand, I believe this to be the type of armatus ; and,
because it has a label saying so, that it came from Puerto Portrero. No trace
of the variety pulcher has been found.
A further discussion of this subspecies appears below after the description
of A . brasiliensis erinaceus.
Astropecten brasiliensis erinaceus Gray
(Plate IV, figs. 1 and 2.)
Astropecten erinaceus Gray, 1840, Ann. Mag. Nat. Hist. 6 : 182 ; Boone, 1928 : 3, pi. 2.
Astropecten brasiliensis erinaceus Doderlein, 1917, Siboga Exped. Monog. 46a : 84 ; with earlier
references and synonymy : 169, pi. 8, figs. 4-43.
Description of Type Specimen. R=65 mm., r=i4 mm., R : r=4.6
The paxillae are similar to those of armatus.
The superomarginal plates number twenty-three. They are similar to those
of armatus except that the inner row of spines, present in armatus on the first
to third plates only, extends as far down the ray as to the thirteenth or fifteenth
plate. It is often absent from the fifth plate, which carries a large spine of the
outer row. The latter starts on the fourth plate and continues to the end, or to
near the end, of the ray. Most plates therefore have two spines ; the outer is
conspicuously stronger than the inner. The single spines on the first plates are the
strongest of all, being nearly 4 mm. long. On a few plates the single inner spine
is replaced by two.
The general covering and armature of the inferomarginals is similar to that
of armatus. The outermost of the spines along the aboral edge is enlarged so as
to appear, with the two marginal spines with which it stands in line, as one of a
series of three.
The arrangement and proportions of the adambulacral spines are the same
as in armatus except that the big spine of the second row is not so big. In the
proximal part of the ray where it is 2.5 mm. long its end is only very faintly, if
at all, expanded. In the distal part of the ray it is not expanded at all ; the free
end is narrower than the base. The ends of this large spine and of the furrow spines
are strongly grooved on the under, the outer, side.
There are only two, exceptionally three, actinal intermediate plates on either
side. They are armed with groups of slender spinelets like those on the adam-
bulacral edges of the adjacent inferomarginals. On some of them one of the spines
is larger than the remainder and compressed.
NOTES ON ASTEROIDS IN THE BRITISH MUSEUM 505
The mouth-plates and madreporite are as in armatus.
Discussion. Earlier writers had written of erinaceus as identical with armatus
(see Fisher, 1911, p. 60), but Doderlein nevertheless regarded them as distinct
and treated each as a subspecies of brasiliensis . Two recent authors have not
followed Doderlein though neither has given reasons for not doing so : Boone
(1928, p. 3) writes of erinaceus as a separate species, and Clark (1940, p. 322) of
armatus as a species in Fisher's sense (including erinaceus).
There are not in the British Museum collection any other specimens of
armatus or erinaceus than the specimens described above ; so that my only
possible contribution to the problem of whether or not they are distinct is that
of describing those specimens. If they are distinct, these are the types.
Doderlein describes the differences between armatus and erinaceus as being
in the number and arrangement of the superomarginal spines, and in the nature
of the enlarged adambulacral spine. Superomarginal spines may be so very
variable in so many species that I think they may be discounted in this discussion.
The differences which Doderlein describes between the large adambulacral spine
are shown, for what they may be worth, by the types described above.
D6derlein's picture of the under surface of the inferomarginals of erinaceus
(pi. 8, fig. 4a) shows no isolated spine near the adoral and adambulacral edge.
Such a spine occurs in the types of both erinaceus and armatus. It is not present
in Fisher's picture of the inferomarginals of armatus (191 1, pi. 7, fig. 3).
Astropecten mauritianus Gray
(Plate IV, figs. 3 and 4.)
Astropecten mauritianus Gray, 1S40, Ann. Mag. Nat. Hist. 6 : 182 ; 1S66, 3 ; Perrier, 1876 : 346 and
359 ; Doderlein, 1917 : 127 ; non Koehler. 1910 : 32. ; ? non Brown. 1910 : 29.
Astropecten mauritianus var. mascarena Doderlein, 1917, Siboga Exped. Mo-nog. 46a : 143. pi. 6,
fig. 3 ; pi. 13, figs. 2-26.
Astropecten hemprichii, Loriol, 1885, Mem. Soc. Phys. Genive, 29 (4) : 74, pi. 21. figs. 7 and 8.
Astropecten sp. Loriol, 1885, Mim. Soc. Phys. Genive, 29 (4) : 77, pi. 22, fig. 1.
Description of Type Specimen. R==57 mm., r=i8.5 mm., R : r=3'i.
Locality : Mauritius.
The specimen is very brittle. Two of the rays are broken off and parts of the
paxillar fields of those which remain have fallen in.
Most of the larger paxillae have eight or nine central, and fifteen or sixteen
peripheral, spinelets, with the central slightly larger than the peripheral. The
central spinelets may be more numerous, up to sixteen, and no bigger than the
peripheral. The paxillae are very small in the centre of the disk and at the ends
of the rays. In the proximal part of the ray those in the centre are larger than
those, which are regularly arranged, on the sides.
There are twenty-one superomarginal plates. They encroach strongly on the
abactinal surface particularly in the proximal part of the ray, where they have
a more rounded contour than in the distal part. They are covered with low-
rounded granules which are of the same diameter as, but lower than, the largest
of the paxillar spinelets. On each plate there are about six irregular rows of
granules, those near the margin smaller than those in the centre. The fasciolar
grooves are fringed by fine capillary spinelets. Each plate, except for the first
two or three, bears a small conical spine on the outer edge. On the
proximal plates it is small, less than half as long as a plate ; it is larger, but never
as long as a plate, in the distal part of the ray.
506 D. DILWYN JOHN
The inferomarginals do not project as far as the superomarginals in the
arm-angles, but do so slightly beyond them in the distal part of the ray. They
are covered, though not thickly, with narrow flattened spinelets with rounded
ends. Large flattened spines arise from among them. The most conspicuous form
a row of three or four along the aboral edge ; there is a less conspicuous row of
shorter spines along the adoral edge, and one or two arise from the middle of the
plate. On-the outer margin there is one long spine nearly as long as three plates.
Its base is rounded, but it is compressed towards the end which is sharply pointed.
Three flattened spines stand in a row parallel to the axis of the ray below the
marginal spine. The aboral is the longest, being half as long as the marginal spine.
The spinelets of the fasciolar groove are capillary.
There are three furrow spines of which the middle is the longest. It is not
much stouter than the other two and is only slightly compressed. There are two,
or exceptionally three, spines in the second row. The aboral is nearly as long as
the furrow spine ; it is longer and stouter than the adoral. A row or a group of
three to five much smaller spines comprise the third row.
The mouthplates are prominent. Parallel to the suture there is on either side
a row of about ten roughly equal spines, ending in a very large spine at the inner
end. The free margin on either side appears to have about seven spines which
grow in size from the outer to the inner end. The innermost stands beside the
largest of the first series and is only slightly smaller than it.
There are three actinal intermediate plates on either side. They are armed
with groups of spines somewhat similar in size to those of the third row of the
adambulacral plates.
The madreporite is small. A number of low irregular mound-like granules
are borne on its surface.
Description of Second Specimen. There is a second and larger specimen in
the Museum collection : R=82 mm., r= 22 mm., R : r=37 ; twenty-seven
superomarginal plates. It is labelled "Isle of France" (Mauritius). There are no
other data with it, but it is probably old for it is broken and brittle. It differs
from the type in a number of small ways.
The central spinelets of the paxillae are globose and markedly larger than the
peripheral. They are often arranged in a circle around one which is central. The
larger paxillae tend to be polygonal. This heightens the contrast between the
irregularly arranged paxillae down the centre of the rays and the smaller ones
which are regularly disposed on either side of them.
On one of the arms the superomarginal spines begin on the fourth plate, on
the other four on the fifth. In each of the five arm-angles one of the first supero-
marginal plates, but not the other, has a single enlarged granule like an incipient
spine.
On the inferomarginals there is a higher proportion of spines to spinelets
than in the type ; there are ten to twelve to each plate apart from the three below
the large marginal spine.
On the adambulacral plates one of the spines in the groups which comprise
the third row is often enlarged. Similarly on the actinal intermediate plates one
or two spines are considerably bigger than the others. The madreporite is almost
completely hidden.
Discussion. Perrier records that he studied Gray's types of this species at
the British Museum. He describes them as having strong conical spines on the
NOTES ON ASTEROIDS IN THE BRITISH MUSEUM 507
inner margins of the first and neighbouring superomarginal plates. It was an
error, difficult to understand and unfortunate in its effects, for it has been copied
by Loriol, Koehlcr, Brown and Doderlein. The latter would not have created
his variety mascarena were it not for this error : there is no doubt that the
specimens described under that name are mauritianus.
Brown states that he named his specimen from the Mergui Archipelago after
careful comparison with the specimens in the British Museum. It cannot have
been with the only two specimens of mauritianus or he could not have failed to
detect Perrier's mistake. It must remain doubtful what species he was describing
until his specimens can be re-examined.
Astropecten mauritianus is known only from Mauritius.
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1938. Echinoderms from Australia. Mem. Harv. Mus. comp. Zool., 55 : 1-596,
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5 pis.
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508 D. DILWYN JOHN
Fisher, W. K. 1911. Asteroidea of the North Pacific and adjacent waters. 1. Phan-
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1919- Starfishes of the Philippine Seas and adjacent waters. Bull. U.S. nat.
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1925. Sea Stars of Tropical Central Pacific. Bull. Bishop Mus., Honolulu,
27 : 63-88, 4 pis., 9 text-figs.
Goto, S. 1914. A descriptive monograph of Japanese Asteroidea. J. Coll. Set.
Tokyo, 29 (1) : 1-108, 19 pis.
Gray, J. E. 1840. A synopsis of the genera and species of the Class Hypostoma
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1866. Synopsis of species of Starfish in the British Museum. London.
Koehler, R. 1909. Echinodermes provenant des campagnes du yacht Princesse-
Alice. Result. Camp. sci. Monaco., 34 : 1-317, 32 pis.
1910. Echinoderma of the Indian Museum, Pt. 6. The shallow water Asteroidea.
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fauna Westafr., 1 : 127-303, 12 pis.
Livingstone, A. A. 1932. Asteroidea. Sci. Rep. Gr. Barrier Reef Exped., 4 (8) :
241-265, 12 pis.
Loriol, P. 1885. Catalogue raisonne des Echinodermes recueillis par M. V. de
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LtiTKEN, C. 1859. Bidrag til kundskab om de ved kysterna af Mellem-og Syd-
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25-96.
Macan, T. T. 1938. Asteroidea. Sci. Rep. John Murray Exped., 4 (9) : 323-435,
6 pis.
Mortensen, Th. 1927. Handbook of the Echinoderms of the British Isles. Oxford.
M6ller, J., and Troschel, F. H. 1842. System der Asteriden. Braunschweig.
1843. Neue Beitrage zu Kenntniss der Asteriden. Arch. Naturgesch. 9 (1) :
113-120.
Perrier, E. 1875-1876. Revision de la collection de Stellerides du Museum
d'Histoire Naturelle de Paris. Arch. Zool. exp. gen., 4 (1875) : 265-450 ;
5 (1876) : 1-104, 209-304.
Sladen, W. P. 1883. The Asteroidea of H.M.S. "Challenger" Expedition. Pt. 2.
/. Linn. Soc. {Zool.), 17 : 214-269.
1889. Report on Asteroidea. Rep. voyage Challenger (Zool.) 30.
Studer, Th. 1884. Verzeichniss der wahrend der Reise S.M.S. Gazelle urn die Erde
1874-76 gesammelten Asteriden und Euryaliden. Abh. preuss. Akad. Wiss.
(Anh. No. 2) : 1-64, 5 pis.
PLATE I.
i . I ,!> ,./>, i u n ■:> anulatu ■ natalensi s, ref. no [Q, abactinal surface, natural size [**i I
granulatus natalensis, rel 0639, R 22 mm., actinal surface, enlarged Fi
pinu ;, type, abactinal surface, enlai ged Fig. 4, type, actinal su
enlarged. Fig ;, Istropecten nti actus, holotype, abactinal surfaci enlai ed 1 ; >pecifti
>.■ ■ holotype portion of actinal surface showing actinal intermediate plates, enlai
PLAT] I!
l ig i . I ;tropt i ft n spht nop I ax, dry specimen, aba* tinal surface natural size, l;ij
/>/<m , dry specimen, actinal surface, natural size Fig ■; I i dn
side view <>f arm, enlarged l ig \, > to /■■ . . natural size.
Fig. g tstropet ten pectinatus, K 50 mm., portion oi actinal urfaci h n actinal intermediate
plates, enlarged.
ri \ 1 1: in
i I tropecten brasiliensis or mat u type, .iK<.im.<i surface, natural size.
brasitiensis armatus, type actinal surface, natural size.
;')g. J. J-//
PI.. VI I l\
fig lyAslropedenbrasiliensiserinaci i type abactinal surfi I uct->l Fig 2, \stropectenbrasiliensis
erinaceus. type, actinal surface reduced Fig 3 I
red I Fig 1 \ stropeclen maurilianm type ictinal surface redu
5«9
INDEX TO VOLUME 42
New taxonomic mines are printed in clarendon type.
abditus (Hexacylloepus), 223, 333
abdominalis (Hexacylloepus), 333,336—337
absens (Diopsis), 156
acanthifera (Astropecten), 485
Achias, 147-148, 150
Acicnemis, 427, 428
acicula (Heterelmis), 218, 2 kj, 220. 224,
373. 389-390, 39i. 395
aciculus (Hexacylloepus), 333
acrosticlialis (Homoneura), 137, 143
acuminatus (Amphialodes), 409
acuminatus (Potamophilus), 226
acutangulus (Trachodes), 427
acutirostris (Crinia), 73, 83-84
Adapanetus, 470-471
Adelotus, 1, 3, 6, 8, 9, 12, 30-31
adustus (Echinocnemus), 310
aequalis (Elsianus), 264
aethiopica (Diopsis), 156
affaber (Alcidodes), 430
affinis (Crinia), 79, 80, 8r
affinis (Euryscaplius), 212
affinis (Pterophrynus), 80
africana (Cyrtodiopsis), 157,160-161, 163
albatrox (Euporismus), 187-188, 201
albicollis (Metrachodes), 428
alboguttatus (Cyclorana), 16, 19, 20
albopunctatus (Cionus). 423
albopunctatus (Cyclorana), 9
albopunctatus (Heleioporus), 4, 6, 32, 33.
38-39
Alcidodes, 428-430
alni (Niphades), 408
alternans (Pamborus), 204
imoenulus (Otidognathus), 469-470
Amphialodes, 408-410
Amphialus, 409
amplipennis (Lara), 226
Vnaeropsis, 150
Vm j loi nemis, (.28
Aiicvronyx, 224, 231
andersoni (Astropecten), 485
angulatus (Euryscaplius), 210, 211
angulatus (Lobotrachelus), 442, 448
angustus (Microcylloepus). 22^, 304, 306,
307-3"". ;' I
antares (Astropecten), 485
antennalis (Eutomus), 467
antennatum (Mo edanum), 475, 476, 478-479
Anthonomus, j 2 j
Novn . zool., 4-', 3. 1948.
anthracinum (Carenum), ji<.
antillensis ( Astropecten), 485, 502
anuda (Homoneura). [43
apicalis (Diopsis), 151
apii alis (Elmis), 319
apicalis (Hexacylloepus), 223, m ;)
337. 339
apicalis (Neoelniis), 22^,, $22. 32$, 327-330
apicalis (Neoempheria), 107
Apion, 411, 423
Apionodes, 422 423
Aplotes, 471-472
Apoderus, 413-416
aranciacus (Astropecten), 485
araneolus (Cylloepus), 342, 346
araneolus (Limnius), 342, 345
arborea (Hyla), 10
arcurostris (Gasteroclisus), 399
armatus (Astropecten), 485, 486. 503-504,
5°5
armirufus (Mecysmoderes). 455-456
articulatus (Astropecten), 485
artivittis (Alcidodes), 428-429, 130
asper (Dyscerus), 402-403, 406
aspersa (Baris), 457-458
aspera (Neoelmis). 223, 322, 325, 328, 350
asperula (Ixalma), 411
asperulus (Lobotrachelus), 44 ^
assimilis (Scaraphites), 208
Astropecten. 485
Asyntona, 148
Athesapeuta. 4 S' '
atratus (Euryscaplius), 211
atrifacies (Homoneura). 136, 139-140
atronitens (Apoderus), 415
attenuatum (Monostomum), 215
australiaca (Rana), 1
australiacus (Heleioporus), 33, 39-40
australiensis (Stenolysmus), 173
australis (Bombinator), >>|
australis (Cyclorana). 3, 6, 9, 10, 16, 18-19,
20
australis (Heleioporus), 9
australis (Phryniscus), 94, 104
australis (Pseudophryne), 9, 96, 98, 101-102,
104
Australysmus, im, 20]
Austrolimnius, 22^. 2::
aval > I Lara). 22"
azteca (Neoelmisj. 22^, ^22. u\. 324. 327
51°
babadjanedesi (Sphyracephala), 151
bacchus (Scaraphites), 208, 209
Bagous, 411
Balaninus, 420-421
barberi (Cylloepus), 346, 347, 356, 359, 361,
365
bardus (Peribleptus), 399
Bans, 457-458
barnardi (Philoscaphus), 212
basipennis (Alcidodes), 429-430
Batrachopsis, 1
bennetti (Notaden), 9, 61, 62—63
Berosiris, 437
Betelmis, 227
bibroni (Pseudophryne), 9, 94, 96, 97, 99,
102-105
bicolor (Deporaus), 418
bicolor (Elsianus), 261, 263
bicostatus (Philoscaphus), 213
bidens (Lobotrachelus), 443, 447
biflagellata (Neoempheria), 127, 128, 129
biguttatus (Thisus), 430-431
bigutticollis (Dyscerus), 403-404, 406
bilobata (Neoempheria), 113
binodosus (Kobuso), 407
bioculata (Homoneura), 140
bipectinata (Neoempheria), 109, 110-111,
129
bipunctipennis (Teleopsis), 150
bispinosus (Astropecten), 485
bispinulus (Dyscerus), 407
bistrispinosus (Hoplapoderus), 412
bisulcatus (Peribleptus), 399
bivittatus (Copidorrhinus), 421, 422
blairi (Cylloepus), 223, 346, 349~352. 355
blanchardi (Pseudophryne), 100
blandus (Apoderus), 416
blenuus (Baris), 458
Bombinator, 94
Borborocaetes, 1
borgmeieri (Neoempheria), 11 3-1 14, 129
brachmanus (Paratrachelophorus), 417
bradleyi (Neoempheria), 117, 118
brasiliensis (Astropecten), 485, 486, 502,
503-505
brevicauda (Neoempheria), 112, 129
breviceps (Hoplapoderus), 412
brevicornis (Diopsis), 147
brevicornis (Ochyromera), 424-425, J26
brevicornis (Sphyracephala;, 151
brevipes (Cyclorana), 16, 21
brevis (Adelotus), 9, 30-31
brevispinus (Astropecten), 485, 496
brisbanensis (Pamborus), 204
brooksi (Pseudophryne), 97
Bufo, 10
bufo (Elmis), 296
bufo (Xenelmis), 223, 298-300
Bufonella, 95
burmanensis (Trachelolabus), 412
burmanus (Chelonebarus), 460-461
burmanus (Euscelophilus), 411-412
burmanus (Thisus), 430-431
caelata (Diocalandra), 472-473
caesa (Elmis), 319 '
caesa (Neoelmis), 322, 323, 330
Calandra, 473
californicus (Astropecten), 485
caligatus (Dyscerus), 404-405, 407
caliginosus (Hoplapoderus), 412
Calobata, 148
Calyptopygus, 466-467
Camariolius, 89
caniventris (Sapromyza), 144
caraibus (Hexanchorus), 232, 233
carbonarius (Euryscaphus), 21 1-2 12
carbonicolor (Apoderus), 415
Carenum, 210, 212
carinatus (Microcylloepus), 301, 302, 303, 304
carinatus (Philoscaphus), 212
carinensis (Cercidocerus), 469
cariniceps (Mecistocerus), 435-436
carinifrons (Mecedanum), 475, 476
Catatropis, 215
centralis (Heleioporus), 33, 35-36
Centralis (Mecedanum), 475, 476, 477,
478, 482, 483-484
Centrocorynus, 416
Cephaloconus, 145
Cercaria, 215-216
Cercidocerus, 469
ceres (Steganopsis), 133
Cestrotus, 134
Ceuthorrhynchus, 457
championi (Euops), 412
chaudoiri (Mystropomus), 205, 206
chaudoiri (Pamborus), 204
Chelonebarus, 460-461
Chelydobatrachus, 1
chiloensis (Austrolimnius), 286
Chiroleptes, 1, 2, 15
Chiroleptina, 2
Chrysopogon, 154
chrysops (Osmylus), 195
Cionus, 423
circularis (Diopsis), 161
citrinus (Kempynus), 192-193. 201
clathratus (Dyscerus), 406
INDEX
5"
clavicornis (Phanocerus), 223, 231, 245,
246-248, 249, 250, 251-254
clypeatus (Elsianus), 263, 264
coarctus (Rhynchites), 420
Codula, 154
cognata (Ochyromera), 425, 426
cognatus (Dyscerus), 404, 406
Colobodes, 432
conciliatus (Rhadinomerus), 439
confucii (Desmidophorus), 430
confusa (Trypaneoides), 136, 145, 146
confusus (Scaraphites), 208, 209
congener (Entornus), 467
connexus (Euosmylus), 197, 201
consobrinus (Cylloepus), 343, 344, 345, 346,
365
contemptus (Rhadinomerus), 439
convexlcollis (Lobotrachelus), 444, 449-
45°
convexiusculus (Limnodynastes), 44, 54-55
Copidorrhinus, 421-422
coppingeri (Astropecten), 492
coraclnus (Psilarthrus), 463-464, 465
corax (Cossonus), 467-468
cordata (Metialma), 451-452, 454
coriacea (Pseudophryne), 9, 96, 99, 100-101,
104
cornuta (Monocera), 135
coronata (Ochyromera), 425-426
corporaali (Apoderus), 414
corticeus (Mecistocerus), 434
Cossonus, 467-468
costa-limai (Neoempherla), 126-127, 12&>
129
costalis (Philoscaphus), 213
costalis (Phrissoderes), 460
costicollis (Psilarthrus), 462-463, 465
cothurnata (Diopsis), 149
cothumata (Pseudodiopsis), 150
crassus (Kobuso), 407
crenaticollis (Scaraphites), 208
Crinia, "1-3, 5-6, 8-9, 64, 71-91, 95. 100
crockeri (Homoneura), 137, 143
Crossotarsus, 475, 480
crux (Balaninus), 421
Cryptorrhynchus, 441
Cryptotis, 1
cultrlpes (Cyclorana), 17, 22
curranl (Cyrtodiopsis), 157, 158, 159, 163
curtulus (Austrolimnius), 286, 290, 295
curvirostrls (Dolichaulax), 465-466
cuvieri (Physalaemus), 89
cyclops (Tomapoderus), 413
Cyclorana, I, 2, 3, 6, 8, 10, 12, 15-24
Cycnotrachelus, 416
Cylloepus, 221, 223, 224, 229, 230, 261, 303,
333. 334. 342-368
cyprinus (Cephaloconus), 145
Cyrtodiopsis, 147, 149, 150, 153, 154, 155,
156-161
Cyrtotrachelus, 469
dahlii (Cyclorana), 16, 17-18
dahomensis (Astropecten), 485
dalmanni (Cyrtodiopsis), 156, 157, 158, 159,
163
dalmanni (Diopsis), 149, 156, 157
darlingtoni (Crinia), 72, 77
darllngtoni (Pamborus), 205
dealbatus (Peribleptus), 399
debilis (Niphades), 408
decursus (Alcidodes), 430
Demimaea, 421, 422
dendyi (Pseudophryne), 96, 104, 105
dentipes (Psilarthrus), 462, 464
Deporaus, 418
Desmidophorus, 430
Diasemopsis, 148, 149, 150, 152, 153, 154,
155. 156, 157
diemenensis (Austrolimnius), 290
dilatatus (Euryscaphus), 211-212
Dimorphognathus, 6
Diocalandra, 472-473
Dioides, 134, 146
Diopsina, 149, 150, 153, 154, 155, 156, 160-161
Diopsis, 147-151, 153. 154. '55. '56. 157. 158
Dipsacaster, 496
direptus (Dyscerus), 401-402, 406
discalis (Apoderus), 414-415
Disersus, 226, 230, 231
distincta (Homoneura), 141
diversa (Homoneura), 138, 143
dohertyi (Cercidocerus), 469
Dolichaulax, 465-466
dorsalis (Limnodynastes), 9, 40, 44-48
Dryopomorphus, 225
dumerili (Limnodynastes), 46-47
duplicatus (Astropecten), 485
Dupophilus, 229, 231
Dyscerus, 399-407
Dytaster, 496
ebeninus (Euryscaphus), 210
echinatus (Hoplapoderus), 412
Echinocnemus, 410, 41 1
Eidoporismus, 165, 200
elegans (Pamborus), 204
ellipticus (Calyptopygus), 406-467
512
INDEX
Elmis, 227, 230, 286, 287, 295, 300, 303, 319,
332. 342
elongata (Diocalandra), 473
elongatipes (Phymatapoderus), 416
elongatus (Trachodes), 427
Elsianus, 221, 223, 227, 230, 258, 261-286,
317. 331. 342.
Empheria, 107
Endaeus, 426-427
enderleini (Neoempheria), 122, 123, 125
englishi (Crinia), 73, 89-90
ephemera (Cercaria), 215
Ephippium, 154
Eremotes, 468
erichsoni (Mecedanum), 475, 478
erinaceus (Astropecten), 486, 504-505
erirrhinoides (Tadius), 432
erosus (Paratrachelophorus), 417
Esolus, 230, 321
ethelia (Minettia), 144
Eugnamptus, 418-420
Euops, 412
Euosmylus, 166, 188, 196-197, 201
Euporismus, 166, 187-188, 201
Eurydiopsis, 150
Euryscaphus, 203, 209-212
Euscelophilus, 411-412
Euthycus, 410
Eutornus, 467
evanescens (Neoempheria), 120-121
evanescens (Pleonazoneura), 107, 122
excisus (Kempynus), 189
extraneus (Stenolysmus), 172-174, 200
eyrei (Heleioporus), 33, 40-42
falcatus (Kempynus), 195, 196, 201
fasciatus (Isostenosmylus), 184, 185, 186,
200
fasciolatus (Mixophyes) , 6, 9, 13-15
faunus (Xanthochelus), 399
favicollis (Rhadinomerus), 438, 439
feae (Paramecolabus), 412
ferruginea (Diopsina), 155, 160, 161
ferruginea (Diopsis), 156
ferruginea (Hexacylloepus), 333, 335
fijiensis (Astropecten), 485, 501-502
filum (Mecedanum), 475, 476, 477, 479-
480
flavicornis (Neoempheria), 123, 124, 129
flavicoxa (Neoempheria), 115, 129
flavipes (Hexacylloepus), 333
fiavonotatus (Cycnotrachelus), 416
fletcheri (Lechriodus), 9, 26-27
fletcheri (Limnodynastes), 44, 56-57, 106
formosensis (Sciophila), 107
formosus (Astropecten), 485
formosus (Austrolimnius), 223, 286, 289,
290, 291-293
frenatus (Alcidodes), 430
frenatus (Lobotrachelus), 443, 446-447
frenatus (Peribleptus), 399
frosti (Philoria), 60
furcata (Monocera), 135, 146
furcatipes (Rhadinomerus), 438-439
furvus (Eugnamptus), 420
fusciceps (Isostenosmylus), 181-182, 183,
185, 200
fuscoapicata (Dioides), t34, 146
Gasteroclisus, 399
gemmosus (Hoplapoderus), 412
georgiana (Crinia), 1, 2, 9, 10, 73, 78-81, 82,
89
gibbicollis (Euscelophilus), 412
giganteum (Mecedanum), 475, 478, 484
gigas (Neoelmis), 322
gigas (Scaraphites), 206, 207
glauerti (Crinia), 73, 81-82
Glauertia, 3, 6, 9, 64, 65-68, 69, 95
gouldii (Myobatrachus), 9, 91-93
gracilipes (Hexanchorus) , 223, 231, 232, 233-
238, 239, 240-244
gracilis (Neoelmis), 321
granata (Xenelmis), 295
grandis (Elsianus), 223, 264, 269-271
granicollis (Carcilia), 410
granicollis (Niphades), 408
graniger (Amphialodes), 409-410
graniger (Elsianus), 223, 264, 267-269, 276-
277, 278-281, 282, 283, 284, 285
granosus (Hexacylloepus), 333
granulatus (Astropecten), 485, 486-489
granulicollis (Rhadinomerus), 439
granulosus (Hexacylloepus), 333
grayi (Heliorana), 48
grayi (Limnodynastes), 48
griegi (Astropecten), 485
grouvellei (Hexacylloepus), 333
guentheri (Pseudophryne), 95, 96-97, 104
guerini (Pamborus), 204
guttatus (Mecistocerus), 432-433
gyrosicollis (Athesapeuta), 459
hartmeyeri (Astropecten), 485, 492-495
haswelli (Crinia), 72, 77-78, 81, 89
hawaiiensis (Homoneura), 137, 143
hearseiana (Sphyracephala), 151
INDEX
513
hebes (Desraidophoms), 430
Heleioporus, 1. 4, 6, 8, 9, 13, 16, 25, 31-42
Heliorana, 48
Hemisus, 8
hemprichi (Astropecten), 485
hermatophilus (Astropecten), 485
heros (Scaraphites), 208, 209
Heterelmis, 218, 219, 220, 224, 229, 230,
368-396
heterocerus (Eylloepus), 223, 347, 352-355
Hexacylloepus, 221, 223, 229, 331-342,
371
Hexanchorus, 223, 226, 230, 231-244, 245
Hexechopsis, 148
hirtipes (Scaraphites), 207, 208, 209
hispidus (Trachodes), 427, 428
histrio (Aplotes), 471, 472
histrionica (Zeugenia), 441
Homoneura, 134, 135-144, 146
Hoplapoderus, 412
horni (Hexacylloepus), 223, 333, 340, 341-
342
horvathi (Homoneura), 137, 142, 144
humeralis (Scaraphites), 206-209
Hydora, 225
Hydrethus, 225
Hyla, 10
Hyparlnus, 441
Hyperolia, 68
Hyperolius, 68
hystrix (Hoplapoderus), 412
hystrix (Trachodes), 427
ichneumonea (Achias), 148
ichneumonea (Diopsis), 147, 151
ignita (Crinia), 79, 80, 81
ignorata (Metialma), 451, 454
illustris (Neoempheria), 126
imbellis (Astropecten), 485
imbricata (Cercaria), 215-216
imbricatus (Notocotylus), 215
immaculiventris (Homoneura), 136, 140
impar (Rhadinomerus), 437, 439
inaequalis (Microcylloepus), 220, 223, 304,
3°5. 3°9. 3IO-313. 3M. 315. 316. 3i7-3'9
incisus (Kempynus), 189-192, 196, 201
incisus (Osmylus), 189
indicus (Astropecten), 485, 489-490
indistinctus (Hexacylloepus), 333
Indobatrachus, 10
indutus (Rhadinopus), 440-441
ineptus (Lobotrachelus), 442
inermis (Cyclorana), 16, 17
ingratus (Lobotrachelus), 442
insulanus (Scaraphites), 207
insularis (Limnodynastes), 47
insularis (Paracycnotrachelus), 416
integrirostris (Mecistocerus), 435
intermedius (Scaraphites), 207
irregularis (Astropecten), 485
Isostenosmylus, 165-167, 174, 180-186, 200
Ixalma, 411
javanicus (Astropecten), 485
jejuna (Metialma), 452-453, 454
johannseni (Neoempheria), 126
johannseni (Pleonazoneura), 107, 121, 122
johnstoni (Astropecten), 485
johnstoni (Petropedetes), 59
kagoshimensis (Astropecten), 485
kastnerl (Neoempheria), 120, 121, 122,
123, 125
Kempynus, 166, 188, 189-196, 197, 199, 201
Kobuso, 407
koehleri (Astropecten), 489
lacustrls (Australysmus), 198-200, 201
laevigatus (Austrolimnius), 286
laevis (Crinia), 5, 9, 71, 72, 75-77, 78
Laglaisia, 150
laminatipes (Lobotrachelus), 443, 446,
447
lanei (Neoempheria), 117, 118-119, 129
languidus (Lixus), 399
laporteae (LobotracheUis), 444, 450
Lara, 225, 226, 230
Latelmis, 230
lateralis (Eutornus), 467
laticollis (Scaraphites), 206
laticosta (Homoneura), 137, 141
latipennis (Oedosmylus), 178, 179, 200
latipennis (Scaraphites), 208, 209
latipes (Dyscerus), 405-406, 407
latisetis (Mecistocerus), 433-434
latiusculus (Kempynus), 192, 194, 195, 201
lativola (Diopsis), 158
leai (Crinia), 72, 73-74
Lechriodus, 1, 6, 8, 9, 12, 24-29
lenaeus (Scaraphites), 208, 209
Lepidarcus, 441-442
lepidotus (Lobotrachelus), 443, 445
leucaspis (Lobotrachelus), 442
leucosticta (Sapromyza), 144
leverl (Homoneura), 135, J38-140
levlpes (Mecysmodere8), 456
514
INDEX
levirostris (Metlalma), 453-454, 455
limbata (Homoneura), 139
Limnius, 228, 230, 254, 288, 289, 296, 297,
342
Limnodynastes, 1, 4, 6, 7, 8, 9, 10, 12, 13, 25,
42-59, 106
lindneri (Neoempheria), 125, 126, 129
Lithostygnus, 475
Lixus, 399
lobata (Neoelmis), 322
Lobotrachelus, 442—450
longiclavis (Dyscerus), 406
longior (Heterelmis), 369, 370, 371
longipennis (Kempynus), 194-195, 201
longipes (Apionodes), 423
longipes (Microcylloepus), 300
longula (Heterelmis), 219, 220, 224, 371,
372- 373. 376-3§o. 382, 383, 384, 385, 387,
389, 392-396
longula (Neoelmis), 220, 223, 320, 322, 323-
326, 327, 328, 329, 330
loveridgei (Philoria), 60-61
Loxocera, 148
lucidus (Scaraphites) , 206, 207, 209
liiderwaldti (Neoempheria), 119,121-122,
123, 129
luridus (Austrolimnius), 286, 289
lutzl (Neoempheria), 128-129
Lysmus, 172
macilentus (Tetragonothorax), 399
macleayi (Euryscaphus), 210
macleayi (Pamborus), 204
macleayi (Scaraphites), 207
Macrelmis, 231
Macronychoides, 254
maculata (Neoelmis), 319
maculipennis (Achias), 148
maculipennis (Neoempheria), 107, 117, 118,
119, 129
major (Pseudophryne), 96, 98-99
malaisei (Adapanetus), 470
malaisei (Dyscerus), 399-400, 406
malaisei (Lepidarcus), 441-442
malaisei (Niphades), 407-408
malaisei (Paraplapoderus), 413
malaita (Maquilingia), 144
mammilatus (Astropecten), 485
Maquilingia, 144
marginatus (Astropecten), 485
marmorata (Uperoleia), 9, 10, 69-70, 95
martini (Scaraphites), 208, 209
mascarena (Astropecten), 507
mastersi (Philoscaphus), 213
mastersi (Scaraphites), 208
Mataeopsephenus, 227
materna (Maquilingia), 144
mauritianus (Astropecten), 485, 486, 505-507
Mecedanum, 475-484
Mecistocerus, 432-436
Mecysmoderes, 455-457
Megalabops, 149, 150, 151, 153, 154, 157,
Megalathoraca, 148
melanogaster (Steganopsis), 131, 132
melanopyga (Lechriodus), 9, 25-26
mesactus (Astropecten), 485, 496-497
Metacrinia, 3, 6, 9, 65, 93-94
Metialma, 450-455
Metrachodes, 427-428
michaelseni (Astropecten), 485, 502
Microcylloepus, 220, 223, 228, 230, 300-319
Microdiopsis, 149
micros (Xenelmis), 295, 297
mimlca (Metlalma), 451, 454
mindanensis (Astropecten), 485
Minettia, 144
minima (Elmis), 319
minor (Lithostygnus), 475
minutus (Austrolimnius), 286
Mixophyes, 1, 6, 8, 9, 10, 12, 13-15
mjobergi (Glauertia), 65, 66
mjdbergi (Pseudophryne), 69, 95
monacanthus (Astropecten), 485, 491-492
Monocera, 134-135, 146
Monostomum, 215
monstruosa (Monocera), 135
montana (Crinia), 73, 90-91
montanus (Austrolimnius), 289
montanus (Oedosmylus), 179-180, 200
montanus (Paracycnotrachelus), 417
morbus (Desmidophorus), 430
morenoi (Isostenosmylus), 186
morosus (Lobotrachelus), 443
mucronatus (Psilarthrus), 463, 465
miilleri (Neoempheria), 116-118, 129
musgravei (Neoelmis), 322
Mycomyia, 108
Myiodactylus, 172
Myobatrachus, 1, 3, 6, 7, 9, 10, 65, 91-93
myobrachius (Astropecten), 500, 501
Myrmecia, 94
Mystropomus, 203, 205-206
Nanophyes, 411
natalensis (Astropecten), 485, 489
Negritomyia, 154
neivai (Neoempheria), 109-110, 129
Nemoxenus, 399
INDEX
515
Neobatrachus, 1
Neoelmis, 220, 223, 229, 231, 317, 319-331
Neoerapheria, 107-129
neosignata (Homoneura), 140
Neosolus, 286
Neurocompsa, 107, 108
nichollsi (Metaerinia), 9, 93—94
nlchollsl (Notaden), 9, 62, 63-64
nichollsi (Pseudophryne), 93, 95
nlgrlcollis (Euops), 412
nigrifrons (Homoneura), 139
nigrifrons (Isostenosmylus), 185-186, 200
nigrigenibus (Paracycnotrachelus), 416-
417
nigritulus (Nanophyes), 411
nigrorufus (Mecysmoderes), 456
Niphades, 407-408
Notaden, 1, 3, 5, 6, 7, 9, 10, 13, 61-64
notatlventris (Homoneura), 137, 142
notatus (Apoderus), 416
notatus (Dyscerus), 407
Notocotylus, 215
notograptus (Astropecten), 486
nubilus (Mecistocerus), 434
nudus (Lobotrachelus), 444, 450
obesa (Heterelmis), 219, 220, 224, 373, 380,
381, 382-386, 387, 389, 395, 396
obesus (Cionus), 423
obesus (Euryscaphus), 210
obesus (Microcylloepus), 223, 304, 305-
307. 315
obliteratus (Euosmylus), 197, 201
oblongus (Austrolimnius), 289
oblongus (Trachodes), 427
obscura (Heterelmis), 218, 219, 220, 224, 373,
387-389, 390, 391, 396
occldentalls (Pseudophryne), 95, 97-98,
102
Ochyromera, 424-426
Ocoblodes, 432
Oedosmylus, 165, 166, 167, 174-180, 187, 200
olivaceus (Limnodynastes), 9
opacicauda (Xenomimetes), 468-469
opacus (Apoderus), 413-414
opacus (Pamborus), 203, 204
optatus (Cylloepus), 346, 347, 356, 359, 365
ordlnatus (Dyscerus), 401, 406
orien talis (Glauertia), 9, 65, 67-68
orientalis (Hoplapoderus), 412
orientis (Homoneura), 140
ornatipennis (Neoempheria), 119-120, 121,
129
ornatipennis (Neurocompsa), 107-108
ornatus (Limnodynastes), 9, 44, 48-50, 51
Orochlesis, 441
oryzae (Calandra), 473
Osmylus, 172, 189, 194
Otidognathus, 469-470
ovatus (Trachodes), 427
Oxydema, 468
Oxyglossus, 10
pacca (Pagiophloeus), 399
pacificus (Scaraphites), 208
Pagiophloeus, 399
pallidipes (Stenelmis), 224
pallidus (Oedosmylus), 175-176, 177, 179
palpalis (Cylloepus), 346
Pamborus, 203-205
pannosus (Eugnamptus), 419-420
papuanus (Lechriodus), 25, 27-28
Paracycnotrachelus, 416-417
Paradoxus (Myobatrachus), 93
Paragorgopsis, 148
Paramecolabus, 412
Paratrachelophorus, 417
parcus (Lobotrachelus), 443, 448
pardalls (Eugnamptus), 419
Paroplapoderus, 413
partimpunctatus (Chelonebarus), 461
pauper (Berosiris), 437
pectinatus (Astropecten), 485, 497-498
pectoralis (Alcidodes), 430
peguensis (Euops), 412
Pelmatops, 148
pelobatoides (Heleioporus), 33, 36-38
pereirai (Neoempheria), 129
Peribleptus, 399
Periphemus, 473
peronii (Limnodynastes), 4, 6, 9, 10, 44,
57-59. 106
Persephonaster, 496
Petropedetes, 6, 59
Phanoceroides, 225, 229, 246, 248, 392
Phanocerus, 223, 226, 229, 231, 232, 245-254
Philocryphus, 32
Philoria, 6, 8, 13, 59-61
Philoscaphus, 203, 212-213
Phractops, 15
Phrissoderes, 459-460
Phryniscus, 94, 104
Phrynodon, 6
Phymatapoderus, 415, 416
Physalaemus, 89
pictipennis (Dioides), 134
pictus (Heleioporus), 9, 33, 34
Pilolabus, 415
5x6
INDEX
pilosa (Neoempheria), 124-125
plana (Heterelmis), 219, 220, 224, 373, 383,
384, 385-386, 395, 396
platyacanthus (Astropecten), 485
platycephalus (Cyclorana), 9, 17, 22-24
platyceps (Lechriodus), 9, 25, 28-29
Platypus, 475, 481
plaumani (Microcylloepus) , 304
plaumanni (Hexacylloepus), 333
plaumannl (Neoempheria), 114, 115, 116,
118, 129
pleiacanthus (Astropecten), 489, 490
Pleonazoneura, 107, 125
Pleurodema, 10
plumbeus (Lobotrachelus), 443, 447
politus (Elmis), 287
politus (Euryscaphus), 210, 211, 212
pollex (Aplotes), 472
polyacanthus (Astropecten), 485
pontoporaeus (Astropecten), 485
pontoporeus (Astropecten), 486
Porismus, 165, 187
posticus (Dyscerus), 400-401, 406
Potamodytes, 225, 226
Potamophilinus, 225
Potamophilops, 226, 227
Potamophilus, 225, 226, 230
pradieri (Pamborus), 204, 205
praecelleus (Apoderus), 415
prainae (Lixus), 399
preissii (Astropecten), 485
propinquus (Centrocorynus) , 416
proximus (Cylloepus), 224, 347, 363, 365,
366
prostemalis (Neoelmis), 322
prothoracica (Elmis), 286
Pseudodiopsis, 149, 150, 152, 153, 154, 155,
156
Pseudophryne, i, 3, 5, 6, 7, 9, 65, 69,
94-105
Psilarthrus, 461-465
ptinoides (Trachodes), 427
pulchellus (Eidoporismus), 165, 200
pulcher (Astropecten), 504
pulverulentus (Isostenosmylus), 182-184,
186, 200
pumilus (Balanlnus), 420-421
punctatum (Mecedanum), 475, 476
puncticollis (Cylloepus), 224, 334, 346, 347,
355-359. 361, 365
puncticoxa (Neoempheria), 114, 115-116,
117
pusilla (Elmis), 303
pusillulus (Astropecten), 485
quadriguttata (Diopsis), 149
quadrimaculatus (Otidognathus), 469
quinqueguttata (Cyrtodiopsis) , 157,
159, 163
quinqueguttata (Diopsis), 156, 158
158-
Rana, 1, 2, 3, 6, 10
regalis (Astropecten), 485
regularis (Myrmecia), 94
regularls (Mystropomus), 205-206
Rhadinomerus, 436-439
Rhadinopus, 439-441
rhinoceros (Monocera), 135
Rhinophrynus, 10
Rhynchaenus, 423
Rhynchites, 420
Richardia, 148
Riolus, 230
roelofsi (Aplotes), 471
roelofsi (Echinocnemus), 411
roratus (Dyscerus), 403, 406
rosea (Astropecten), 492, 493, 494
rosea (Crinia), 72, 74-75
rostrata (Neoempheria), 124
rotundipennis (Scaraphites), 207, 209
rubra (Hyla), 10
ruficomis (Lobotrachelus), 442
rufipennis (Eugnamptus), 418-419
rufirostris (Metialma), 455
rufitibiis (Lobotrachelus), 443, 447-448
rufulus (Centrocorynus), 416
rugosa (Uperoleia), 5, 69, 70-71
russelli (Glauertia), 9, 65, 66-67
russelli (Homoneura) , 136, 140, 146
saeva (Metialma), 451, 453, 454
salmini (Limnodynastes), 9, 44, 55-56, 106
sanctae helenae (Astropecten), 485
sandersoni (Elsianus), 223, 264, 265, 271,
275-276
Sapromyza, 134, 144
satelles (Otidognathus), 470
scabrosus (Hexacylloepus), 223, 333, 337-
339, 341. 342
scalptus (Peribleptus), 399
Scaraphites, 203, 206-209
scenicus (Alcidodes), 430
schayeri (Astropecten), 498
schenklingi (Desmidophorus), 430
schevilli (Mixophyes), 15
Sciophila, 107
scoparius (Astropecten), 485
scotti (Mecedanum), 475, 478, 481-482, 484
INDEX
517
sculptipennis (Cylloepus), 224, 344, 346-349
scutellaris (Centrocorynus), 416
scutellaris (Elsianus), 223, 264, 265-267, 269,
Semelima, 428
semimarmorata (Pseudophryne), 94, 96, 99-
100, 104
semiruber (Deporaus), 418
serlcea (Ochyromera), 424, 426
sericoclava (Adapanetus), 471
sexnotatus (Lobotrachelus), 442, 444-445
sexualis (Cylloepus), 224, 346, 347, 361, 362-
365
shannoni (Neoempheria), 125-126, 129
siamensis (Astropecten), 494
sibogae (Astropecten), 494
signata (Homoneura), 140
signatifrons (Homoneura), 136, 139
signatus (Endaeus), 426-427
signifera (Crinia), 5, 6, 9, 40, 73, 78, 79, 80,
81, 84-91, loo
sikaiana (Homoneura), 142
silenus (Scaraphites), 206, 208, 209
simoni (Elmis), 319
simoni (Neoelmis), 322
simplex (Neoempheria), 112-113, 129
simplicipes (Lobotrachelus), 444, 448-449
smarti (Neoempheria), 122, 123
Smicronyx, 411
smithi (Elmis), 332
smithi (Hexacylloepus), 332-333
solomensis (Steganopsis), 132, 133
Solomonia, 135, 138-139
spenceri (Limnodynastes), 44, 50-52
Sphenophorus, 473
sphenoplax (Astropecten), 485, 499-500
Sphinxis, 423
Sphyracephala, 148, 149, 150, 151, 152, 153,
154. 155. 156
spinipes (Cylloepus), 224, 346, 347, 359-361,
365
spinipes (Microcylloepus), 304
spinipes (Rana), 1
spinosa (Neoempheria), 111-113, 129
spinulosus (Astropecten), 485
squamifer (Psilarthrus), 464, 465
Steganopsis, 131-133, 146
stellae (Euosmylus), 196-197, 201
Stenelmis, 224, 231, 263, 319, 345, 366
Stenolysmus, 167, 172-174
stenopterus(Stenosmylus), 169-170, 171, 172,
200
Stenosmylus, 165, 166, 167-172, 200
Stetholus, 225
stolata (Crinia), 79, 80
striata (Neoempheria), 109
striata (Sciophila), 107
striatoides (Elsianus), 223, 264, 265, 271, 274,
275
striatus (Elsianus), 262, 263, 264, 271, 272-
274, 276, 278, 281-283, 284. 285
strictlceps (Cossonus), 468
strictlcollis (Rhadinomerus), 436-437, 439
strigicollis (Carcilia), 410
stylicornis (Mecysmoderes) , 456
subclavata (Neoempheria), 120-121
subcostatus (Mystropomus), 205, 206
subcylindricus (Mecistocerus), 436
subfasciatus (Lobotrachelus), 444
subnuda (Homoneura), 137, 141-142
subplumbeus (Rhynchites), 420
subsulcatus (Euryscaphus), 211
subsulcatus (Hexacylloepus), 333
sudelli (Heleioporus), 33, 42
suffusus (Austrolimnius), 289
sulcatus (Hexacylloepus), 333
sulcicollis (Austrolimnius), 223, 286, 288,
289, 290, 293-295
sulcicollis (Euryscaphus), 210
surda (Minettia), 144
suspensa (Homoneura), 136, 138, 141, 143
suturella (Metlalma), 450-451, 454
syntomus (Astropecten), 498
tarsalis (Austrolimnius), 287, 289, 290
tarsalis (Elsianus), 263, 264
tarsalis (Heterelmis), 224, 373, 374-376
tarsalis (Hexanchorus), 232, 233, 239
tarsalis (Xenelmis), 296, 297, 298, 300
tasmaniensis (Crinia), 9, 73, 82-83, 89
tasmaniensis (Limnodynastes), 9, 10, 44, 52—
54, 106
tasmaniensis (Oedosmylus), 176-179, 200
tecticollis (Rhadinopus), 439-440
Teleopsis, 148, 149, 150, 151, 152, 153, 154,
155. 156
tenuis (Stenosmylus), 167-169, 170, 171, 200
tepperi (Philoscaphus), 213
terraereginae (Limnodynastes), 46
Tetragonothorax, 399
Themara, 148
thermarius (Hexanchorus), 232
thermarum (Microcylloepus), 300, 309
thienemanni (Notocotylus), 215
thlsoides (Ypsilepidus), 432
Thisus, 430-431
thoracica (Neoelmis), 320, 321
tibialis (Hexanchorus), 232, 233, 239
Tolrlolus, 223, 227, 230, 254, -261
Tomapoderus, 413
5i8
INDEX
romentosum (Mecedanum), 475, 476, 477,
480-481
Tomicoproctus, 468
Trachelolabus, 412
Trachodes, 427
triacanthus (Persephonaster), 496
trigonophorus (Alcidodes), 430
tripunctifacles (Steganopsis), 131-133,
146
triseriatus ( Astropecten) , 485, 500-502
troilus (Microcylloepus), 223, 305, 314-
316
tropicus (Neosolus), 286
truncata (Teleopsis), 157
Trypaneoides, 136, 144-145, 146
tuberculatum (Carenum), 212
tuberculatus (Philoscaphus), 213
tuberosus (Potamodytes), 226
tumida (Barls), 457
turneri (Stenosmylus), 171-172, 200
ungulatus (Limnius), 254
ungulatus (Tolriolus), 223, 255-257, 258-261
unispinosa (Neoempheria), 123-124
Uperoleia, 1, 5, 6, 7, 9, 10, 64, 65, 67, 68-71,
95
Urenae (Lobotrachelus), 444, 450
vacivus (Lobotrachelus), 444, 449
vappa (Astropecten), 485
varia (Homoneura), 142
variegatus (Astropecten), 485
varimana (Steganopsis), 132, 133, 146
varipennis (Neoempheria), 107, 116-118
varius (Carmariolius), 89
velitaris (Astropecten), 485
ventralis (Cylloepus), 346
verrilli (Astropecten), 485
verrucosa (Catatropis) , 215
verrucosum (Monostomum), 215
vetula, (Lixus) 399
viatrix (Homoneura), 136, 139
victoriana (Crinia), 72, 76-77
victoriensis (Austrolimnius), 289
villosa (Diopsis), 156
viridis (Pamborus), 204
vitticeps (Paroplapoderus), 413
vogell (Neoempheria), 127-129
waterhousei (Euryscaphus), 211
westermanni (Alcidodes), 429, 430
whitei (Cyrtodiopsis), 157, 159, 163
whitei (Diopsis), 156
wilsmorel (Heleioporus), 33, 36
Xanthochelus, 399
Xenelmis, 223, 228, 249, 295—300
Xenelmoides, 297
Xenomimetes, 468-469
Ypsilepldus, 431-432
zebra (Astropecten), 485, 492-495
zebra (Nemoxenus), 399
Zeugenia, 441
Zygocephala, 148
THE ZOOLOGICAL RECORD
An annual record of the zoological literature of the world, con-
taining a reference to every book or paper of importance that has
appeared in the previous year. It is divided into 19 sections, which
can he obtained separately, at the following prices :
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The price of the complete Annual Volume is £4, or £3 5s. to
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CONTENTS
Entomological Results from the Swedish Expedition 1934
to Burma and British India— Coleoptera : Curculionidae.
By Sir G. A. K. Marshall, F.R.S
PAGE
397
A Synopsis of the Genus Mecedanum Erichson (Coleoptera,
Colydiidae). By H. E. Hinton, Ph.D
475
Notes on Asteroids in the British Museum (Natural
History) — I. The Species of Astropecten. By D. Dilwyn
John, M.Sc
485
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