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UNIVERSITY OF KANSAS

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UNIVERSITY

UNIVERSITY OF KANSAS PUBLICATIONS
University of Kansas Science Bulletin - Vol. XXXIV -Part I

October 1, 1951

Lawrence, Kansas I I

ANNOUNCEMENT

The University of Kansas Science Bulletin (continuation of the
Kansas University Quarterly) is issued in parts at irregular inter-
vals. Each volume contains from 300 to 600 pages of reading mat-
ter, with necessary illustrations. Exchanges with other institutions
and learned societies everywhere are solicited. All exchanges should
be addressed to:

The University of Kansas Science Bulletin,

Library of the University of Kansas,

Lawrence, Kan.

PUBLICATION DATES

The actual date of publication ( i. e., mailing date ) of many of the
volumes of the University of Kansas Science Bulletin differs so
markedly from the dates bourne on the covers of the publication or
on the covers of the separata that it seems wise to offer a corrected
list showing the mailing date. The editor has been unable to verify
mailing dates earlier than 1932. Separata were issued at the same
time as the whole volume.

Vol. XX— October 1, 1932. Vol. XXIX, Pt. I— July 15, 1943;
Vol. XXI— November 27, 1934. Pt. II— Oct. 15, 1943.

Vol. XXII— November 15, 1935. Vol. XXX, Pt. I— June 12, 1944.
Vol. XXIII— August 15, 1936. Pt. II— June 15, 1945.

Vol. XXIV— February 16, 1938. Vol. XXXI, Pt. I— May 1, 1946.
Vol. XXV— July 10, 1939. Pt. II— Nov. 1, 1947.

Vol. XXVI— November 27, 1940. Vol. XXXII— Nov. 25, 1948.

Vol. XXVII, Pt. I— Dec. 30, 1941. Vol. XXXIII, Pt. I— April 20, 1949.
Vol. XXVIII, Pt. I— May 15, 1942; Pt. II— Mar. 20, 1950.

Pt. II— Nov. 12, 1942.

Editor Edward H. Taylor

Editorial Board Charles D. Michener, Chairman

Ernest Griswold
Frank E. Hoecker
Russell C. Mills
Paul G. Roofe
Rufus H. Thompson

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

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THE PUBLICATION OF THE RESULTS OF

RESEARCH BY MEMBERS OF THE

UNIVERSITY OF KANSAS

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OCT 16 iL

HARVARD
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Volume XXXIV, Part I

University of Kansas Publications

Lawrence, October 1, 1951

PRINTED BY

FERD VOILAND. JR.. STATE PRINTER

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1951

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S-A -L.

Contents of Volume XXXIV, Part I

No. PAGE

1 A Brief Review of the Snakes of Costa Rica. Plates I to

XXIII. Text figs. 1-7 Edward H. Taylor, 3

2. A New Veracrucian Salamander. Text figs. 1 to 3.

Edward H. Taylor, 189

3. A New Species of Leiolopisma (Reptilia: Sauria) from

Mexico Hobart M. Smith, 195

4. Notes on a Small Herpetological Collection from Guerrero.

Plate XXIV Charles W. Hall, 201

5. A New Subspecies of the Mexican Moccasin, Agkistrodon

bilineatus. Plate XXV.

W. Leslie Burger and William B. Robertson, 213

6. An Anatomical Study of Certain Salamanders of the Genus

Pseudoeurycea. Plates XXVI to XXVIII. .Irwin L. Baird, 221

7. The Estrous Cycle in the Woodrat, Neotoma floridana.

Plates XXIX to XXXV Arthur 0. Chapman, 267

8. A Revision of the Genus Anisops (Notonectidae, Hemiptera)

Plates XXXVI to LVII George T. Brooks, 301

9. Concerning Oligocene Amphisbaenid Reptiles. Plates LVIII

to LXVII Edward H. Taylor, 521

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXIV, Pt. I] October 1, 1951 [No. 1

A Brief Review of the Snakes of Costa Rica

Edward H. Taylor

Abstract: The snakes of Costa Rica are reviewed, largely on the basis a
collection made by the author in Costa Rica in 1947. Synopses of all known
species are given and keys are added. The fauna as listed consists of 56 genera,
and 132 species and subspecies are recognized. The following forms are de-
scribed as new; Nothopsis torrcsi, Dipsas costaricensis, Pliocercus annellatus,
Micrurus pachecoi, and Micrurus alleni richardi.

INTRODUCTION

Early in June, 1947, at the instigation of Professor Reuben Torres
Rojas, of Cartago, Costa Rica, I received an invitation from the
Rector of the National University of Costa Rica to spend the summer
of 1947 studying the herpetological faunas in that country. The
welcome invitation was promptly accepted and after hasty prep-
arations, I was ready to leave for Costa Rica on June 18. I was
joined at this time by Richard C. Taylor, likewise bent on making
a herpetological collection in Costa Rica. After an uneventful air
journey we arrived on June 21. Three days were involved in getting
the equipment through customs and in obtaining the required per-
mits. The first day of collecting was June 24.

From this date until September 7, the greater part of the time
was spent in the field. Faunas of four principal areas were sampled.
These were: Turrialba and its immediate surroundings, in the east-
ern part of the plateau region; Isla Bonita, on the eastern slope of
Volcan Poas; the summit and western slopes of Cerro de la Muerte;
and Los Diamantes, in the eastern lowlands. A few other localities
were visited for shorter periods and of course less representative
collections were obtained than were taken in the aformentioned
places.

(3)

4 The University Science Bulletin

Considerable rain fell during these months, hampering collecting
and causing the loss of many man-hours to the collecting work.
The journey to the Pacific lowlands on the west slope of Cerro de la
Muerte was a pleasant exception. However, due to the fact that
this region was dry, collecting was poorer than in other localities
visited.

The night collecting everywhere was especially profitable, and
a part of practically every night provided its increment to the
collection. Collecting at high elevation on Mt. Irazu and Cerro
de la Muerte was hampered considerably by fog and rain coupled
with the low temperatures of these high peaks. Aside from driv-
ing the animals under cover, it likewise did much to render the
collector inactive.

While the greatest number of novelties is present in the amphibian
collection, certain forms among the snakes seem to merit new
designations, and they are given herein.

The collection of snakes from the four principal areas are as fol-
lows. The number of species, in a measure, represents the length
of time spent in collecting in each place.

Turrialba and surroundings

Constrictor constrictor imperator Spilotes p. pullatus

Nothopsis torresi sp. nov. Erythrolamprus bizonus

Geophis hoffmanni Leptodeira a. annulata

Geophis dolichocephalu Imantodes cenchoa semifasciatus

Xenodon bertholdi Tantilla shistosa

Dipsas costaricensis sp. nov. Tantilla annidata

Pliocercus annellatus sp. nov. Micrurus nigrocinctus rnosquitensis

Rhadinaea d. decorata Micrurus mipartitus midtifasciatus

Drymarchon corais melanurus Bothrops n. nummifer

Drymobius m. margaritiferus Bothrops schlegelii (nigroadspersus)

Dnjmobius rhombifer Bothrops lansbergii

Pseustes shropshirei

Pseustes poecilonotus chrysobronchus

Isla Bonita, Volcan Poas

Ninia oxynota Imantodes cenchoa semifascitatus

Dipsas annidata Imantodes inornatus

Chironius carinatus Clelia c. clelia

Chironius grandisquamis Bothrops picadoi

Lampropeltis doliata micropholis Bothrops lateralis
Leimadophis tacnitirus juvenilis

Taylor: Review of Snakes of Costa Rica 5

San Isidro El General

Geophis hoffmanni Dryadophis melanolomus alternatus

LampropelMs doliata micropholis Clelia petolarius

Coniophanes fissidens punctigulgaris Kenodon colubrinus

(Boquete camp)

Bothrops n. nigrovirdis
Micrurus n. nigrocinctus
Ninia metadata

Los Diamantes

Ninia tessellata Coniophanes fissidetis punctigularis

Trimetopon pliolepis Micrurus alleni richardi sp. nov.

Enulius sclateri Micrurus nigrocinctus mosquitensis

Leptodeira a. annulata Bothrops atrox
Oxybelis brevirostris

Miscellaneous Localities
Peralta, 300 m.

Thalerophis depresstrostris

Cervantes
Ninia maculata

Cartago
Thamnophis sirtalis chalceus

Santa Cruz on Volcan Turrialba, 6500 ft.

Bothrops lateralis
Geophis brachycephala
Geophis hoffmanni

PLAN OF WORK

One of the anticipated results of my acceptance of the invitation
to study in Costa Rica was the preparation of a work that might
present my findings, and at the same time serve as a primary text
for study and identification of the Costa Rican herpetological fauna.
No such work, purporting to fill such a purpose, has been prepared
for that country. The works of Dr. C. Picado (1931) and Senor
Carlos Viquez (1935. 1941) have dealt with only a limited part
of the snake fauna. The older work of Cope in 1875 listed the
known fauna but many species appear merely as names.

In consequence I have included for each species and subspecies,
regarded as having a valid place in the Costa Rican faunal list, a
description of an individual specimen or a brief description drawn
from the scattered literature. Keys are added to assist in identifica-
tion. No attempt is made to include an exhaustive treatment of

6 The University Science Bulletin

synonymies. Usually the earliest reference is recorded and one or
more others giving a report of the species in Costa Rica.

I contemplate the preparation of two other papers, one dealing
with the remaining reptiles ( crocodilians, turtles and lizards) and
the other treating of the amphibians. Already certain short papers
have appeared describing new frogs, salamanders, and lizards.

The half-tone illustrations presented are from photographs made
from preserved specimens submerged in water. The drawings are
the work of Miss Thespa Stapoulis.

This study is based primarily on two lots of Costa Rican snakes
now in the Museum of Natural History of the University of Kansas.
The first is that obtained by the Hall, Camp and Westman Costa
Rican Expedition of 1947. The second is that obtained by me
either by gift, purchase or collecting the same year. I have also
had at hand the collection made by Richard C. Taylor who has
allowed me the privilege of studying it. I have had access to most
of the snakes from Costa Rica now in the Museum of Comparative
Zoology, Harvard College, and those in the United States National
Museum of Washington, at which institutions I spent some time in
the summers of 1948 and 1949. I have likewise found occasion to
examine certain specimens in the Museum of Zoology University of
Michigan, Chicago Natural History Museum, The Academy of
Natural Sciences of Philadelphia and The American Museum of
Natural History. I desire to express my thanks especially to Mr.
A. Loveridge of Harvard, and Dr. Doris Cochran of the National
Museum for their many courtesies while I was studying in these
institutions, and to the directors and curators of other collections
for their kindness in making available certain specimens.

Specimens from the various collections are referred to in this
paper under the following initials:

A. M. N. H American Museum of Natural History

A. N. S. P Academy of Natural Sciences of Philadelphia

C. N. H. M Chicago Natural History Museum

EHT-HMS Edward H. Taylor-Hobart M. Smith collection

M. C. Z Museum of Comparative Zoology of Harvard College

M. Z. U. M Museum of Zoology, University of Michigan

M. N. H Museum of Natural History, University of Kansas

R. C. T Richard C. Taylor collection

U. S. N. M United States National Museum.

HISTORY OF COLLECTIONS AND LITERATURE

The first considerable collection of Costa Rican snakes made by
Dr. Carl Hoffmann, in Costa Rica prior to 1859 and forwarded to

Taylor: Review of Snakes of Costa Rica 7

the Royal Zoological Museum in Berlin, formed the basis of a report
by Prof. W. Peters* in 1859. The paper contained a listing of the
following 21 snakes. Eighteen of the snakes were of species pre-
viously described from neighboring countries. Three were de-
scribed as new. ( The original nomenclature is used ) :

Colobognathus Hoffmannii nov. gen. Liophis cobella Linne

et sp. Homalocranium melahocephalum
Hydromorphus concolor nov. gen. Dumeril, Bibron and Dumeril

et sp. Oxybelis Catisbyi Schlegel

Bothriechis nigroviridis nov. gen. Oxybelis aeneus Wagler

et sp. Erythrolamprus venustissimus Wied

Streptophorns Sebae Dumeril, Dipsas annulata Linne

Bibron, and Dumeril Himantodes cenchoa Linne

Herpetrodryas Boddaertii Schlegel Hydrophis bicolor Daudin

[Sentzen] Elaps semipartitus Dumeril, Bibron and
Herpetodryas Rappii Giinther Dumeril

Spilotes melanurus Schlegel Elaps circinalis Dumeril, Bibron and
Spilotes variabilis Merrem Dumeril

Leptophis ahaettdla Linne Bothrops bilineatus Wied
Leptophis margaritifer Schlegel

The specimens do not have the exact localities reported with the
exception of Oxybelis aeneus ( "Candalariogebirge und Punta de
Arenas"), Hydrophis bicolor from "Gulfo dulce," Bothrops bilineatus
and Bothriechis nigroviridis (Volcan Barba).

In 1861 Petersf described Typhlops (Hehninthophis) frontalis
from a specimen collected by Dr. Carl Hoffmann and forwarded to
Germany after his death, by Dr. von Frantzius.

Some 12 years after Peters' first report, a herpetological collection
made chiefly in the region about San Jose, Costa Rica by Dr. Van
Patten, reached Edward D. Cope $ at Philadelphia. Thirty-three
species of snakes were represented in the collection. The following
new species were described: Colobognathus dolichocephalus, Colo-
bognathus brachyccphalus and Rhadinaea serperastra.

Besides these three the following were included:

Candisona durissa Linnaeus Pelamis bicolor Daudin

Bothrops atrox Linnaeus Dryiophis brevirostris Cope

Bothriechis nigroviridis Peters Dryiophis acuminatus Wied

Bothriechis affinis Bocourt Thrasops ? mexicanus Dumeril,
Elaps nigrocinctus Girard Bibron and Dumeril

Elaps ornatissimus Jan (var. ) Leptognathus nebulatus Linnaeus

Elaps multifasciatus Jan Dipsas gemmistratus Cope

* Monatsb. Konig. Akad. Wiss. Berlin, 1859, pp. 275-278.
t Monatsb. Akad. Wiss. Berlin. Oct. 1800 (18(»1), pp. 517-521.

t Ninth Contribution to the Heroetology of Tropical America, Proc. Acad. Nat. Sci. Phila-
delphia, 1871, pp. 200-224.

8

The University Science Bulletin

Leptodeira annulata var.
Leptodeira annulata var.
Masticophis margaritiferus Schlegel
Masticophis boddaertii Schlegel
Herpetodryas carinatus Linnaeus
Spilotes melanurus Dumeril, Bibron

and Dumeril
Liophis epinephelus Cope
Coniophanes fissidens Giinther
Conophis lineatus Dumeril, Bibron

and Dumeril

Erythrolamprus venustissimus Linnaeus
Tantilla melanocephala Linnaeus
T ant ilia melanocephala Linnaeus (var.)
Stenorhina ventralis Dumeril, Bibron

and Dumeril
Stenorhina degenhardtii Berthold
Ninia maculata Peters
Ninia atrata Hallowell
Colobognathus hoffmannii Peters
Epicrates cenchria Linnaeus

Eight or nine, perhaps more, of the Van Patten species duplicate
specimens collected by Hoffmann. One of the species listed in this
paper as Tantilla melanocephala variety, is later described as Tan-
tilla armillata sp. nov*

William Gabb of Philadelphia, under the auspices of the Govern-
ment of Costa Rica, made a second important collection in south-
eastern and southern Costa Rica east of the Cordillera de Tala-
manca and on Pico Blanco ( 3565 m. ) , one of the highest peaks of
the range. This collection was much more extensive than that
made by Dr. Van Patten. It contained five turtles, 19 lizards, 35
serpents and 30 amphibians with a total of 37 forms presumably
new to science. In the paper * in which Cope reported the collec-
tion, sixty species and/or subspecies of serpents are listed; of these
35 were represented in the Gabb collection. The remaining 25
forms were either in the Van Patten collection or from a small col-
lection sent from Costa Rica by Mr. Riotte; or they were specimens
listed or reported by other authors. Thirteen species are described
as new. The list follows:

Helminthophis frontalis Peters
Xiphosoma annulatum sp. nov.
Boa imperator Daudin
Leptognathus annulata Giinther
Leptognathus argus sp. nov.
Leptognathus pictiventris sp. nov.
Leptognathus nehulata Linnaeus
Dipsas cenchoa Linnaeus
Sibon annulatum Linnaeus
Oxyrrhopus plumbeus Wied
Oxyrrhopus petolarius Linnaeus
Dryiophis brevirostris Cope
Dryiophis acuminatus Wied
Leptophis aeruginosus sp. nov.
Leptophis saturatus sp. nov.

Leptophis praestans Cope
Dendrophidium melanotropis sp. nov.
Drymobius margaritiferus Schlegel
Drymobius boddaertii Sentzen
Herpetodryas carinatus Linnaeus
Herpetodryas grand isquamis Peters
Spilotes pullatus Linnaeus
Spilotes corais melanurus Dumeril

Bibron and Dumeril
Spilotes chrysobronchus sp. nov.
Coniophanes fissidens Giinther
Pliocercus dimidiatus Cope
Rhadinaca decorata Giinther
Rhadinaea serperastra Cope
Erythrolamprus venustissimus Wied

Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1870 (1875), pp. 128-152.

Taylor: Review of Snakes of Costa Rica 9

Xenodon angustirostris Peters Colobognathus hoffmannii Peters
Stenorhina ventralis Dumeril, Bibron Elaps circinalis Dumeril, Bibron and

and Dumeril Dumeril

Tantilla armillata sp. nov. (from Teleurapsis schlegelii Berthold

Van Patten Coll.) Bothriechis nigroviridis Peters

Microdromus virgatus Giinther Bothriechis lateralis Peters

Ninia sebae tessellatus Cope Bothriechis affinis Boucourt

Contia pachyura sp. nov. Bothriopsis proboscideus sp. nov.

Contia calligaster sp. nov. Bothrops atrox Linnaeus

Catostoma psephotum sp. nov. Lachesis stenophrys sp. nov.

A collection sent to Edward D. Cope from Costa Rica by Sr. Don
Jose Zeledon was reported upon by Cope in 1879.* Three snakes
not previously listed from Costa Rica appear: Scolecophis zonatus
Hallowell, Coluber triaspis Cope, and Porthidiiim nasutum Bocourt.

Cope, in 1893f received and reported on a collection of reptiles

and amphibians from Mr. George K. Cherrie of San Jose, Costa

Rica, among which were three new species. In the following list

the first six species had not been previously listed for Costa Rica.

Rhadinaea ignita Cope Sibon septentrionale rubricatum
Drijmobius rhombifer Peters subsp. nov.

Drijmobius caeruleus Fischer Colobognathus hoffmannii Peters

Drijmobius percarinatus sp. nov. Oxybelis acuminata Wied

Synchalinus corrallioides gen. Elaps nigrocinctus Girard
et sp. nov.

In 1894 Cope \ reports on a series of specimens from the Museo
Nacional de Costa Rica. The following are listed, four of which
are new.

Trimetopon pliolepis sp. nov. Pogonaspis ruficeps sp. nov.

Drymobius paucicarinatus sp. nov. Enulius torquatus Giinther

Leptophis ultramarinus sp. nov.

In Otto Wettstein's "Ergebnisse der osterreichischen Costa Rica-
Expedition 1930; Die Amphibien und Reptilien,"j[ the following
snakes are listed as follows:

Leptotyphlops ( = Glauconia ) albifrons Wagler

Boa ( = Corallus) annulata Cope

Sibynophis (= Polyodontophis) venustissimus Giinther

Drymobius (Eudryas) boddaertii Sentzen

Drijmobius (Drymobius) margaritiferus Schlegel

Spilotes pullatus pullatus Linnaeus

Drymarchon corais melanurus Dumeril and Bibron

Chironius ( = Herpetodryas) fuscus grandisquamis Peters

Leptophis mexicanus Dumeril and Bibron

* Amer. Philos. Soc, vol. 18, no. 104, Aug. 11, 1879.

t Cope, Amer. Philos. Soc, 1893, p. 333.

t Proc. Acad. Nat. Sci. Philadelphia, 1894, pp. 194-206.

1 Sitz, Akad. Wiss. Wien, Math.-Natunv. Kl. Abt. 1. Bd. 143, Heft 1-2, 1934, pp. 1-39.

10 The University Science Bulletin

Leptophis occidentalis occidentalis Gunther

Leptophis bilineatus Gunther

Liophis ( = Rhadinaea ) cobella Linnaeus

Liophis ( = Rhadinaea ) pulveriventris Boulenger

Ophis ( = Xenodon) colubrinus Gunther

Urotheca elapoides Cope

Catostoma ( = Geophis = Dirosema ) brachycephala Cope

Imantodes cenchoa Linnaeus

Leptodeira ocellata Gunther ( = L. annulata personata Cope)

Clelia ( = Oxyrhopus ) cloelia Daudin

Erijthrolamprus aescidapii Linnaeus

Conioplumes (= Erijthrolamprus) imperialis imperialis Baird and Girard

Tantilla ( = Homalocranium ) virgata Gunther

Stenorhina degenhardtii Berthold

Micrurus (= Elaps) mipartitus midtifasciatus Jan

Micrurus ( = Elaps) nigrocinctus nigrocinctus Girard

Bothrops nasuta Bocourt (Amaral)

Bothrops nigroviridis nigroviridus Peters

Bothrops ophryomegas Bocourt (Amaral)

Bothrops schlegelii Berthold

Crotalus terrificus durissus Cope

In 1887 Cope in his "Catalogue of the Batrachians and Reptiles of
Central America and Mexico" listed the herpetological fauna of
Central America known at that time. The names of some 67
species and subspecies of snakes are given as occurring in Costa
Rica. Certain of these names are now regarded as synonyms of
others in the list, as for example, Phylothamnus aeruginosas Cope,
Leptophis bilineatus Gunther and Hapsidophrys saturatus (Cope)
are placed now in the synonymy of Thalerophis depressirostris
(Cope) by Dr. James A. Oliver, the latest reviewer of this group of
colubrine snakes.

Between April 1893, and October 1895, the section of the "Bio-
logia Centrali-Americana" dealing with Serpentes was published
by Gunther. Approximately 76 forms are reported for Costa Rica.
This is one of the most significant works dealing with Central
American Faunas, and it contains numerous excellent illustrations.
The collections in the British Museum form the basis for this work.

Another work of even more importance for the study of Central
American snakes is the great "Etudes sur les Reptiles;" Mission
Scientifique au Mexique et dans l'Amerique Centrale. The livrai-
sons nos. 8-17 dealing with the serpent fauna appeared between
1882 and 1909. The first 8 livraisons (nos. 8-15) were under the
authorship of Fermin Bocourt, the last two (nos. 16-17) were writ-
ten by Dr. F. Mocquard. This work contains hundreds of excellent

Taylor: Review of Snakes of Costa Rica 11

figures. Many Costa Rican specimens are listed. The collections
treated in this work were primarily those of the Paris Museum, espe-
cially those obtained by the Mission Scientifique. However, speci-
mens from other museums are included.

From 1893-1896 the Catalogues of the Snakes in the British Mu-
seum vols. I-III appeared and here again Costa Rican forms in the
British Museum are listed.

From 1896 to 1920, nearly a quarter of a century, little was pub-
lished that dealt directly with Costa Rican herpetological faunas.
In the latter year E. R. Dunn, then collecting for Harvard Univer-
sity was sent to Costa Rica. This marked a beginning of renewed
interest in this area. Later, other journeys were made by him to
Central America, and one or more other visits were made to Costa
Rica. No detailed reports of these collections have been made, but
a few forms from Costa Rica have been described from these or
other collections, among which may be mentioned are:

Thalerophis nebulosus Oliver Trimetopon simile Dunn

Lampropeltis triangulum gaigae Rliadinaea persimilis Dunn

Dunn Dipsas ruthveni Barbour and Dunn

Leimadophis epinephelus juvenilis Rhinobothryum bovaUii Andersson

Dunn Conophis nevermanni Dunn

Trimetopon viquezi Dunn Trimeresurus picadoi Dunn

Three works by Costa Rican authors dealing in part with the
herpetological faunas are: "Serpientes venenosas de Costa Rica
sus veninos seroterapia anti-ofidica" ( 1931 ) by C. Picado T. This
treats of the various species of the poisonous genera Lachesis, Botli-
rops, Crotahts, Micrurus and Pelamis. Numerous good illustrations
of these species are given. A few harmless species are figured that
seem to mimic the poisonous forms.

The other works are "Animales Venenosos de Costa Rica" ( 1935 )
by Lie. Carlos Viquez S. and "Nuestros Animales Venenosos"
( 1941 ) by the same author. The second book is in the nature of a
second edition of the first; but it is rewritten and revised with many
new figures added and it may be regarded essentially as a new
work. One of the features is a listing of the known snake species
from Costa Rica, compiled from literature. The list contains 113
names. This number must be reduced by approximately a dozen
names that are synonyms or repetitions. Maps showing distribu-
tion of a few forms are given.

12 The University Science Bulletin

GEOGRAPHICAL AND CLIMATIC CONSIDERATIONS

The country of Costa Rica, lying as it does athwart the tenuous
land mass that is Central America, has a fauna reflecting in great
measure the spread of forms from the continental part of North
America (Mexico) and from South America. It has served as a
portion of the roadway along which have settled many of the migrant
forms, more coming perhaps from the north than from the south.
Geological evidence points to the fact that at times the roadway has
not been a continuous route; there have been breaks in the land
mass, leaving one or possibly even a chain of islands in its stead.

Animals arriving in Central America, encountering new environ-
mental stimuli, often changed until no longer may the taxonomist
recognize in them the characters of their immediate ancestors living
or dead, but he must consider them as new local creations whose
ancestry may only be conjectured. Thus many of the individual
species now recognized in the fauna have developed in situ, de-
scendants of older residents or migrants.

Moreover the spread of forms from the north or the south has
taken place at various times and the degree of variation in local
populations may offer a chronometer for measuring the relative
time of arrival.

That the migration or spreading of species still continues is sug-
gested by the fact that only a single representative of the large
northern genus Thamnophis has reached Costa Rica and to our
knowledge, has as yet not reached Panama. Moreover, many
species have entered Panama from the south that have not been
encountered across the border in Costa Rica. In this series such
genera as Lygophis, Trypanurgos, Peropodum, Trachyboa and
Atractus may be cited. Certain of these, of course, may actually
occur and will eventually be reported for Costa Rica.

The fauna contains a few elements that did not arrive by land.
These are species that have braved ocean voyages, or not impossibly,
were brought unintentionally by man. Two forms appear to have
arrived via a water route. These are Tretanorhinus nigroluteus
nigroluteus and Pelamis platyurus. Of these, the first may be a
relatively recent coastal arrival and while as yet no specimen has
been found along the coastal or inland waters of Costa Rica, since
it occurs both north and south of the country, its presence there is
to be regarded as unquestionable. The second is a marine species
from South Asia and the Malay Archipelago and the fact that it
shows so few fixed differences from Asiatic members of the species,

Taylor: Review of Snakes of Costa Rica 13

may suggest an arrival in historic times, from, say, the Philippines
as a stow-away on Spanish galleons. We are aware that some-
thing of this sort has happened in the case of a land snake Typhlops
bra minus, and certain lizards Peropus mutilatus and Hemidactylus
frenatus in Mexico. The question whether the species could spread
from a point of landing, south along the Central and South American
coasts and/or north along the Central American and Mexican coasts
during post-Columbian times, I believe, can be answered in the
affirmative. I see nothing inconsistent in such a happening. More-
over, while certain species of sea serpents have definite migrations,
I cannot believe that a bottom-feeding snake could survive a slow
journey under its own power across the Pacific Ocean.

The country of Costa Rica is not large; it has a north-south length
of approximately two hundred and fifty miles, and a total estimated
area of 19,238 square miles.

Prior to the present geological uplift, which has connected North
America and South America by the Panama land bridge, and pre-
sumably Costa Rica to Nicaragua by a similar connection, it is highly
probable that Costa Rica and the adjoining high part of Panama
constituted an island separated by relatively narrow straits from the
high land of Nicaragua and likewise from the high land of eastern
Panama. At the point where Costa Rica bounds Nicaragua, the
land is less than 100 meters above sea level. However, beginning
near the northwestern border, and extending diagonally in a south-
easterly direction, is a mountain chain that for a hundred miles
maintains an elevation above 1000 meters, save for a single short
break. It is known variously as the Cordillera Volcanica, and the
northern part at least is called Cordillera de Guanacaste. In this
range is a series of mountain peaks, mostly volcanoes. The north-
ernmost is Volcan Orosi (1499 m. ). Then follow Volcan Rincon
de la Vieja ( 1502 m. ), Volcan Miravalles ( 1750 m. ), Volcan Tenorio
(1432 m.), and Volcan Canaste (Pelon) (1900 m.).

Near the southern end of this Cordillera, there is a break in the
continuity with the southern mountains (Cordillera Talamanca),
but it is contiguous on the east with a shorter, higher range having
only a slightly diagonal trend, the main axis being nearly east-west
in direction. This is called the Cordillera Central and maintains
throughout its length an elevation above 1500 m. with a series of
volcanic peaks from west to east as follows: Volcan Poas ( 2760 m. ) ;
Volcan Barba (2929 m.); Irazu (3452 m.); Turrialba (3421 m.), the
two latter being connected by a high isthmus above 2150 m. in ele-
vation.

14 The University Science Bulletin

Immediately to the south of these ranges is the central plateau
region. Here are to be found two rivers, the Rio Grande, draining
to the Pacific via the Gulf of Nicoya and Rio Reventazon, flowing
eastward into the Caribbean Sea. The minimum elevation of the
divide between these two rivers is 1566 m. and represents the eleva-
tion of the isthmus connecting the Irazu volcano with the great
southern Cordillera de Talamanca. Here too, on either side of the
central plateau are the chief cities, San Jose, Cartago, Alajuela,
Heredia, and Turrialba.

The directional trend of the Cordillera de Talamanca is to the
southeast, practically the same as that of the mountain chain in
northern Costa Rica. It maintains an elevation of 2150 m. for 110
miles save for a single break where a pass having an elevation
somewhat more than 1538 m. occurs. There is a series of peaks
in this chain as follows ( north to south ) : Cerro las Vueltas, 3087 m.;
Cerro Guerici, 3540 m.; Cerro de la Muerte, 3670 m.; Cerro Chirripo
Grande, 3S37 m., Cerro Durika, 3296 m.; Cerro Cruz del Obispo,
3099 m.; Pico Blanco, 3565 m. and Cerro Pando, 3162 m.

Northward a few peaks are isolated from the main range. Cerros
de Escaso and Cerros de Bustamente are highest, with elevations
2425 m. and 2419 m. respectively. This mountain mass appears
on certain maps as the Aguacate Mountains.

The peninsula of Nicoya jutting from the northwestern mainland
has highlands connecting with the mainland by land, none of which
reaches an elevation of 100 meters elevation and most of which is
much lower; while in the mountainous portion elevations up to
1017 meters are recorded. At the southern extremity of the country
on the Pacific side, the Peninsula de Osa as well as the mountain
mass known as Pico de Burica present similar relations to the main-
land. Elevations of 688 m. and 708 m. respectively are known. All
three of these areas must have experienced island or archipelagic
conditions in the past.

A study of Mexican faunae have shown marked differences in
the components of the faunae of various high mountains. Mount
Chiriqui, lying in northwestern Panama, isolated from the main
cordillera, seemingly has a fauna showing considerable differenti-
ation from that of neighboring mountains. Such differences prob-
ably likewise obtain in certain of the isolated peaks of the Costa
Rican mountains.

A series of outline maps have been made following the elevation
contours at sea level, at 1000 feet, 3000 feet, 5000 feet, 7000 feet

Taylor: Review of Snakes of Costa Rica

15

and 9000 feet. These, better than any description, show the degree
and extent of isolation of the mountain masses at the elevations
p'ven.

3000 FT CONTOUR

5000 FT CONTOUR

•-" \.S'

"S

.Ji.

0<J,

^P

7000 FT CONTOUR

O

. ■> ■

9000 FT CONTOUR

^

r\>-

V

l °

•

- \

~---*J

) j

Outline maps of Costa Rica showing outline of the entire country; and the
1000, 3000, 5000, 7000, and 9000 foot contours (indicated by heavier lines).

16 The University Science Bulletin

The northern and southern areas are connected at elevation of
1566 m. (5000 ft.) but above this there is a break and animals
adapted to elevations of 2000 m. and above are more or less
effectively isolated. At still higher elevations there are several
mountain "islands" separated from each other by varying distances.
Whether it will eventually be shown that each of these higher peaks
has endemic faunal elements is now only a matter of conjecture.
Certain of them certainly have. The age of the various mountains
(or volcanoes) will be a factor. One may presume the oldest ones,
other things being equal, will display modifications in the greatest
number of species. Unfortunately, fewer snakes become adapted
to the higher elevations than either amphibians or lizards.

The extent to which one may divide the country into faunal zones
and faunal areas purely on the study of the serpent fauna alone
cannot be determined with any degree of certainty at the moment,
since the distributional data are too meager and the collecting too
haphazard. Many specimens in collections lack data as to locality,
and if this is known, exact elevation data is often lacking.

The eastern, Caribbean, drainage area, and the Pacific drainage
area together with the dry portions of the central plateau, constitute
two areas differing considerably in amount of rainfall, and the con-
commitant differences in vegetation. The lowlands of the eastern
area have an annual rainfall in excess of 200 inches a year. This is
spread throughout the year with heavier fall in midwinter, Decem-
ber and January, and midsummer, June and July. As one ap-
proaches the summits of the cordilleras, the rainfall becomes less
but even here the vegetation is lush. On the summits of the higher
peaks the forest is absent but at least in the southern Cordillera
the summits have much short bamboo that forms often practically
impenetrable thickets. The ground itself is covered with moss and
lichens, offering shelter for a very considerable salamander popu-
lation.

The western Pacific slope is deprived of much of the moisture
because of the excessive precipitation on the eastern slope caused
by the high central mountains. Here the rainfall is confined very
largely to the summer months, May to August, while the remainder
of the year is dry, and months may pass with practically no rainfall.
The dry season has much wind which results in further moisture loss.

Costa Rica is divided into a series of provinces in such a manner
that a listing of the provincial faunas would contribute little to an
understanding of distributional problems. Hence little effort has
been made to associate place names with provinces.

Taylor: Review of Snakes of Costa Rica 17

FAUNA

The serpent fauna, for the size of the country and the present
state of our knowledge, is large. This is due, in a measure, to the
rugged and divergent characteristics of the topography, which allow
for the interplay of a great variety of environmental stimuli on this
plastic and definitely expanding group of reptiles. At the same
time it has available necessary isolating barriers to select and fix
the mutations that result. In this paper 132 species and subspecies
are considered. These are arranged in 56 genera, under seven
families.

The subfamilies of the Colubridae here treated are four; however
the group Colubrinae is further divisible into other subfamily groups
of perhaps equal rank with certain of those here treated. Certain
of the species of snakes that have been listed heretofore for Costa
Rica appear to be synonyms, and are so treated here; others so listed
may be legitimate members of the fauna, but since I am unable to
find verification of the record are not included. In this latter class
are certain forms listed by Lie. Carlos Viquez S°as follows: Anom-
alepis mexicana; Epicrates cenchria cenchria; Leimadophis taeniurus
epinephalus; Atractus quadrivirgatus [= AdeJphicos qiiadrivirgatus];
Urotheca lateristrigata [= Rhadinaea lateristrigata]; Urotheca el-
apoides elapoides [— Pliocercus elapoides elapoides]; Psendoboa
newiedii [= Clelia newiedii]; Micriiras elegans.

The report of Pseudoboa newiedii Dumeril, Bibron and Dumeril
[= Clelia newiedii] appears also in Amaral, Mem. Instit. Butantan,
tome 4, p. 207 ( 81 ) and may be based on a legitimate report. How-
ever, I have not found record of any specimen taken in the country.

Synchalinus corralliodes Cope is regarded by Dunn as being a
synonym of Phrynonax poecilonotus. I have not examined the type.

Ungaliophis continentalis F. Miiller is reported by Amaral ( Mem.
Inst. Butantan, 1929, IV, p. 145 (19) "Guatemala occidental ate
Panama). I have been unable to ascertain that the species has been
taken south of Nicaragua, f

* Nuestros Animates Venenosos, 1941, pp. (15-57.

tin the latest revision of the Boidae, Stull (Proc. Boston Soc. Nat. Hist., vol. 40, no. 8,
pp. 387-408) revives the generic name Peropodum F. Miiller for Ungaliophis dating the name
from Miiller, 1878, Verh. Nat. Ges. Basel, vol. 6, p. 652, pi. 1. It is true that on this page
Miiller did describe the genus Peropodum, and then proceeded to describe a snake without giv-
ing it a name, but the name appears elsewhere. In the listing of species on page 573, it ap-
pears "Boaedarum n. gen. et. sp. ?." The name Boaedarum is not intended as a generic name
but is the genitive plural form of the family name. However, on page 591, one finds the fol-
lowing :

"Nov. gen. Boid. Affin. Ungal (Anm. 13)
"Spec, guatemalensis.

"a. Retaluleu, costa grande v Guatemala, gesch. v. Dr. G. Bernoulli."
The (Anm. 13) refers to No. 13 of the Anmerkungen zum Katalog, which reads as follows:

2—3286

18 The University Science Bulletin

ACKNOWLEDGEMENTS

I wish to offer my gratitude to the Kansas University Endowment
Association, who generously provided a considerable portion of the
funds for defraying my personal expenses in travel to and from
Costa Rica, and travel and other expenses within the country.

I am deeply obligated to the President of the National Univer-
sity, Dr. Fernando Baudrit, for his generous invitation to become
a guest of that Institution and spend the summer in Costa Rica in
the study of the herpetological faunas.

My heartfelt thanks are offered to the genial, gifted, Prof.
Ruben Torres Rojas, my official host in the country who joined me
on certain memorable field expeditions; presented me with speci-
mens, and "smoothed the way" for me on numerous occasions while
I was entering and leaving the country.

I am under deep obligation to Dr. Ralph H. Allee. Director of the
Inter-American Institute of Agricultural Sciences at Turrialba, who
allowed me the privilege of making the Institute my headquarters,
in furnishing transportation and for other very numerous courtesies.
I am also under obligation to members of his staff: Dr. Frederick
Wellman and Mrs. Wellman, Dr. Guillermo Bonilla, Dr. Caceras,
Dr. Rhoades, Mr. James Forman, and others who were untiring in
their courtesy. A number of specimens collected by the Director's
son, Mr. David Allee, were presented to the collection.

A number of other persons likewise merit my sincere thanks. A
visit to the American Cinchona Plantation on Volcan Poas was ar-
ranged by Captain Hope, Production Manager of the plantation,
who also provided transportation. Mr. Deal Thornton, the Station
Director offered the facilities of the plantation and provided trans-
portation.

Mr. Harshberger and Mr. Virgil Cave, officials of the Pan-Ameri-
can Highway, offered the facilities of the camps at Millville, near
the summit of Cerro de la Muerte, and that at San Isidro El General.
Mr. Cave presented the collection with several important speci-
mens and provided much transportation.

To Mr. Wallace E. Manis, my genial and discerning host at Los
Diamantes (U. S. Department of Agriculture Rubber Station), I
offer my very cordial thanks.

"(13) Boidae von Guatemala Peropodum, n. sp. et genus ? (Hiezu Tafel I). Costa Grande."
Then follows a generic description and a species description of guatemalensis. Thus there can
be no question that both genus and species are correctly established. The name has been at-
tributed by Boulenger to Bocourt, Miss. Sc. Mex., Kept., p. 522, 1882. Accepting the syn-
onymy of Ungaliophis continentalis F. Miiller and Peropodum guaU maletisis F. Miiller, the lat-
ter name, not the former, must be used.

Taylor: Review of Snakes of Costa Rica 19

I am under obligation to several other persons, for specimens and
courtesies, among whom may be mentioned are Prof. Marco Tullio
Pacheco who collected with me and presented to me certain valu-
able specimens. Mr. Good and his sister, Miss Good, hosts at Finca
Dominica at Turrialba, graciously permitted me to obtain a fine
series of Basiliscus plumifrons from the shrubbery about the home.
Serior Antonio Machado of the American Cinchona Plantation pre-
sented the collection with several fine specimens.

To Mr. Richard Taylor, I am grateful for his companionship and
constant aid in the field, for the transportation by air of some forty
pounds of my equipment, and finally for permission to study the
collection which he made.

CHECKLIST OF THE SERPENTS OF COSTA RICA

The following list of genera, species, and subspecies are tenta-
tively recognized in this paper. The genera number 56, the species
and subspecies 132. ( Figures in parentheses refer to page numbers
in this volume. )

Family Anomalepidae ( 24 )

Genus Anomalepis Jan ( 24 )

dentatus Taylor ( 24 )
Genus Liotyphlops Peters ( 25 )

albirostris ( Peters ) ( 25 )
Genus Helminthophis Peters ( 26 )
frontalis ( Peters ) ( 26 )
Family Leptotyphlopidae ( 27 )

Genus Leptotyphlops Fitzinger (27)
albifrons (Wagler) (27)
Family Boidae ( 28 )

Genus Boa Linnaeus ( 28 )
annulate (Cope) (28)
Genus Constrictor Laurenti ( 29 )

constrictor imperator (Daudin) (29)
Genus Epierates Wagler ( 30 )
cenchriu mounts Gray (30)
Family Colubridae (30)

Subfamily Acrochordinae ( 30 )
Genus Nothopsis Cope ( 30 )
torresi sp. nov. (31)
Subfamily Sibynopbinae ( 35 )

Genus Seaphiodontophis Taylor and Smith ( 35 )
venustissimus ( Gunther ) ( 35 )
Subfamily Natricinae (36)

Genus Thamnopbis Fitzinger ( 36 )
sirtalis chalceus ( Cope ) ( 36 )
Subfamily Colubrinae (composite) (37)

20 The University Science Bulletin

Genus Geophis Wagler ( 38 )

[rhodogaster (Cope)] (39)

godmani Boulenger ( 40 )

hoffmanni (Peters) (40)

dolichocephala (Cope) (43)

moesta Gunther ( 44 )

brachycephala (Cope) (46)
Genus Ninia Baird and Girard (49)

sebae sebae Dumeril, Bibron and Dumeril (50)

atrata (Hallowell) (50)

macidata (Peters) (51)

tessellata Cope (53)

psephota (Cope) (55)

oxynota (Werner) (55)
Genus Dipsas Laurenti (58)

anthracops (Cope) (59)

pictiventris (Cope) (59)

argus (Cope) (60)

arHculata (Cope) (61)

annulata (Gunther) (61)

ruthveni ( Barbour and Dunn ) ( 62 )

costaricensis sp. nov. (62)
Genus Sibon Fitzinger (67)

nebtdatus (Linnaeus) (67)
Genus Neoparias Gunther (66)

bicolor Gunther (66)
Genus Tretanorhinus Dumeril, Bibron and Dumeril (66)

nigroluteus nigroluteus Cope ( 67 )
Genus Xenodon Boie (68)

bertholdi Jan (69)

colubrinus Gunther ( 69 )
Genus Enulius Cope ( 70 )

flavitorques (Cope) (70)

sclateri (Boulenger) (71)
Genus Helieops Wagler (74)

wettsteini Amaral (74)
Genus Hydromorphus Peters ( 74 )

concolor Peters ( 74 )
Genus Trimetopon Cope (75)

pliolepis Cope (76)
viquezi Dunn (78)

simile Dunn (79)
gracile (Gunther) (79)
Genus Hypsiglena Cope ( 80 )

torquata torquata (Gunther) (80)
Genus Amastridium ( 80 )
veliferum Cope (81)
Genus Spilotes Wagler ( 82 )

pullatus pullatus ( Linnaeus ) ( 82 )
pullatus mexicamis ( Laurenti ) ( 83 )

Taylor: Review of Snakes of Costa Rica 21

Genus Thalerophis Oliver ( 84 )

mexicanus mexicanus Dumeril, Bibron and Dumeril (84)

depressirostris ( Cope ) ( 85 )

nebulosus ( Oliver ) ( 85 )

richardi occidentalis (Giinther) (86)
Genus Dryadophis Stuart (87)

melanolomus altematus (Boeourt) (87)
Genus Drymobius Fitzinger (88)

margaritiferus margaritiferus (Seblegel) (89)

rhombifer (Giinther) (89)

chloroticus (Cope) (90)

melanotropis (Cope) (91)
Genus Dendrophidion Fitzinger (91)

paucicarinatus (Cope) (91)

dendrophis (Schlegel) (92)
Genus Pseustes Fitzinger (92)

poecilonotus chrysobronchus ( Cope ) ( 92 )

shropshirei Barbour and Amaral
Genus Chironius Fitzinger ( 95 )

carinatus ( Linnaeus ) ( 95 )

grandisquamis (Peters) (96)

melas (Cope) (97)
Genus Elaphe Fitzinger (97)

triaspis ( Cope ) ( 98 )

flavirufa flavirufa (Cope) (98)
Genus Drymaehron Fitzinger ( 99 )

corais melanurus Dumeril, Bibron and Dumeril (99)
Genus Mastieophis Baird and Girard ( 100)

mcntovarius Dumeril, Bibron and Dumeril (100)
Genus Leptodrymus Amaral ( 101 )

pulcherrimus ( Cope ) ( 101 )
Genus Leimadophis Fitzinger ( 102 )

taeniurus juvenilis Dunn (102)
Genus Lampropeltis ( 103 )

doliata micropholis Cope ( 104 )

doliata polyzona Cope ( 103 )

doliata gaigae Dune ( 104 )
Genus Pliocereus Cope (106)

dimidiatus Cope (106)

annellatus sp. nov. ( 107 )

Genus Liophis Wagler (110)

cobella Linnaeus (110)
Genus Bhadinaea Cope (111)

serperastra Cope (112)

calligaster (Cope) (113)

decorata decorata Giinther (113)

pulveriventris Boulenger (116)

verm iculaticeps ( Cope ) (116)

persimilis Dunn (117)

pachyura pachytira (Cope) (117)

paclujura decipiens (Giinther) (118)

22 The University Science Bulletin

Grooved back-fang series [Boiginae] (118)
Genus Trimorphodon Cope (120)

biscutatus biscutatus Dumeril, Bibron and Dumeril ( 120)
Genus Leptodeira Fitzinger ( 121 )

nigrofasciata Gunther ( 121 )

rubricata (Cope) 122)

annulata annulata (Linnaeus) (122)

ocellata Gunther (125)

rhombifera Gunther (126)

maculata (Hallowell) (126)
Genus Oxybelis Wagler (127)

fulgidus (Daudin) (127)

brevirostris (Cope) (128)

aeneus (Wagkr) (128)
Genus Imantodes Dumeril and Bibron (130)

inornatus Boulenger (130)

cenchoa semifasciatus Cope (132)

gemmistratus Cope (135)
Genus Clelia Fitzinger (135)

clelia clelia Daudin ( 135 )

petolarius ( Linnaeus ) ( 138 )
Genus Erythrolamprus Boie ( 140 )

bizomis Jan ( 140)
Genus Coniophanes Hallowell ( 141)

decipiens (Gunther) (141)

piceivittis Cope ( 142 )

fissidens punctigularis Cope (142)
Genus Bhinobothryum Wagler ( 144)

bovalii Andersson ( 144 )
Genus Conophis Peters ( 145 )

lineatus dunni Smith ( 145 )

nevermanni Dunn ( 145 )
Genus Stenorrhina Dumeril (146)

degenhardtii degenhardtii (Berthold) (146)

degenhardtii apiata Cope (147)

freminvillii freminvillii Dumeril, Bibron and Dumeril (117)
Genus Scoleeophis Fitzinger ( 148 )

atrocinctus ( Sehlegel ) ( 148 )
Genus Tantilla Baird and Girard ( 149)

annulata Boettger (149)

armillata Cope (152)

reticulata Cope (153)

virgata (Gunther) (154)

shistosa ( Boeourt ) ( 155 )

ruficeps ( Cope ) ( 156 )
Family Hydrophiidae (156)

Genus Pelamis Daudin (156)

platurus (Linnaeus) (156)

Taylor: Review of Snakes of Costa Rica 23

Family Elapidae ( 157 )

Genus Micrurus Wagler ( 157 )

mipartitus maltifasciata (Jan) (158)

nigrocinctus nigrocinctus (Girard) (160)

nigrocinctus mosquitensis Schmidt (164)

pachecoi sp. nov. (165)

chrki Schmidt (168)

alleni richardi subsp. nov. (169)
Family Viperidae ( 172 )

Genus Bothrops Wagler (composite) (172)

atrox atrox (Linaeus) (179)

nummifer ( Riippell ) ( 181 )

picadoi Dunn (180)

nasuta Bocourt (177)

ophryomegas Bocourt ( 178 )

lansbergii ( Schlegel ) ( 178 )

nigroviridis nigroviridis (Peters) (176)

lateralis (Peters) (175)

godmani (Giinther) (183)

schlegelii (Berthold) (173)
Genus Crotalus Linnaeus ( 183 )

terrificus durissus Linnaeus (183)
Genus Lachesis Daudin (184)

muta stenophrtjs Cope (184)

Key to Families of Costa Rican Snakes

1. Scales on midventral line of belly not, or scarcely, wider than adjoining scales. ... 2
Scales on belly several times wider than adjoining scales 5

2. Tail short strongly compressed vertically, forming an oar; marine snakes; eye

well developed Hydrophiidae

Tail very short, round; eye covered with head shields, concealed or faintly visible, 3

3. Teeth in lower jaw only: ocular scale reaching lip; scales in 14 rows around the

body Leptotyphlopidae

Teeth in upper jaw or in both jaws 4

4. Teeth in upper jaw only; ocular separated from lip by labial scale; hyobranchium

Y-shaped Typhlopidae*

Teeth in one or both jaws; ocular separated from lips by two scales; hyobranchium
W-shaped Anomalepidae

5. A deep pit between nostril and eye Viperidae

No pit between nostril and eye "

fi. Anterior part of maxillary provided with a stationary grooved fang Elapidae

Anterior part of maxillary without a grooved fang; however the posterior tooth in
maxilla may be grooved '

7. Vestiges of hind limbs; ventral scales relatively narrow, distinctly less than width

of body Boidae

No vestiges of hind limbs; ventral scales as wide as or wider than body, nor-
mally Colubridae

* Typhlopidae is included in the key, although no specimen of the family has been reported
in Costa Rica. It occurs to the north, in Guatemala and south in Colombia.

24 The University Science Bulletin

Family ANOMALEPIDAE

The recent findings of Tihen published under the name Dunn
and Tihen,* and of Tihen f strongly suggest the wisdom of recog-
nizing these small snakes under the family name proposed by
Taylor. |

Three genera, Anomalepis, Helminthophis and Liotyphlops, are
regarded as belonging in this family. § The three genera may be
differentiated by the following key.

Key to Costa Rican Anomalepidae

1. Rostral less than half width of head ; preoculars two 2

Rostral at least half width of head ; no preocular scales Liotyphlops

One subocular Helminthophis

Two suboculars Anomalepis

Genus Anomalepis Jan

Anomalepis JanH in Jan and Sordelli, L'iconographie Generate des Ophidiens, livr. 1 ( ?Dec.
1860), p. 1, pi. 5. fig. 1; Arch, per la Zool., vol. 1, p. 185, 1861.

Genotype: Anomalepis mexicanus Jan.

Anomalepis dentatus Taylor

Anomalepis dentatus Taylor, Proc. New England Zool. Club, vol. 17, 1939, pp. 90-91 (type
locality, Barro Colorado Island, Canal Zone**).

A small species of this genus reported from Costa Rica is referred
to dentatus. The species may be recognized by the following
characters: Very small snakes, known maximum size, 158 mm.;
teeth in both jaws; rostral visible from above for a distance equal
to its distance from frontal, its greatest width a little less than half

* Dunn and Tihen, Journ. Morph., vol. 74, no. 2, 1944. pp. 287-294, pi. 1.

f Tihen, Copeia, 1945, no. 4, Dec. 31. pp. 204-210.

t Taylor, Proc. New England Zool. Club, vol. 17, 1939, pp. 87-96, pi. 5.

§ See Smith and Warner, Herpetologica, vol. 4, pt. 6. Dec. 8. 1948, pp. 189, 193.

U Not "Jan and Sordelli" as Dunn insists, Bull. Mus. Comp. Zool. Harvard Coll., vol. 87,
No. 7, 1941, p. 511. According to Jan (Elenco sistematico degli Ofidi descritti e disegnati per
L'iconografie generale, 1863, p. 9) the name Anomalepis dates from the cited publication; while
the specific name is attributed to the Iconographie Generale des Ophidiens. livr. 1, dated De-
cember 1860 but presumably issued at a somewhat later date than that stated.

** In the original description, credit for the discovery of this form was attributed to James
Zetek, since the label in the container and the invoice sent with the specimen contained Mr.
Zetek's name. Dr. Emmet Reid Dunn has made a claim that he is the collector of the speci-
men that served as the type. Thinking Dr. Dunn's claim preposterous, I wrote to Mr. Love-
ridge of Harvard University from whom the specimen was borrowed. Mr. Loveridge admits an
error in cataloguing the specimen. Dr. Dunn is the collector.

In a paper by Dunn (Bull. Mus. Comp. Zool. Harvard College, 87, 1941) attempting Vo cor-
rect some typographical errors in my original paper, and to excuse certain ol his own previous'y
published errors, he is guilty of further errors and questionable taste.

1. Dunn states that the authority fo.' the name Anomalepis mexicanus is Jan and Sordelli.
I have attributed the name to Jan, and this is correct despite Dunn's statement. Jan Was
solely responsible for the taxonomy. Sordelli was the illustrator. Jan makes this clear by : s-
sociating his name alone with the species.

2. Dunn likewise makes this mistake on pages 511, 512, and likewise erroneously attributes
the name Leptognathus vagus to the two authors.

3. On page 516 Dunn refers to a species of Anomalepis as maxicanus. No such species has
been des<ribed.

Oth»r errors in Dunn's paper will be taken up when occasion demands.

Taylor: Review of Snakes of Costa Rica 25

the width of head; prefrontals large, longer than wide, about as
long as frontal; frontal broader than long, four times as large as the
supraocular scales. Five scales border frontal, two laterally, the
anterior of which is the supraocular; behind frontal is the post-
frontal; following the postfrontal are two rather large median scales
followed by two somewhat smaller median scales. The one touching
the postfrontal is largest and bordered laterally by a quadrangular
scale separated from the ocular by a somewhat smaller postocular.
Nasal divided, the lower anterior portion small; the nostril lying
diagonally between the two parts; anterior supralabial large, widely
separating nasals from mouth, its labial border narrower than labial
borders of other supralabials; second supralabial small but larger
than fourth and smaller than third which nearly equals first; loreal
bounded by three labials in front and below, by the larger nasal
and two preoculars above; two subequal suboculars and two post-
oculars, the upper much the larger; eye dimly visible; three infra-
labials; mental grooved but apparently not divided; when mouth
is closed, six scales appear to border upper labials. A pair of pre-
anal scales in contact anteriorly. Scale formula 28-24-22-22-24 (23
or 22). Dorsal scales from rostral to terminal caudal plate, 272.
Scales under tail, 7-9. Total length (maximum known), 158 mm.;
tail, 3 mm.

Genus Liotyphlops Peters

Liotyphlops Peters, Sitzb. Ges. naturf. Fr., 1881, p. 69.

Genotype: Liotyphlops albirostris Peters.

Whether more than a single species of Liotyphlops occurs in
Costa Rica is difficult to say. If there is but one, there may be some
doubt as to the proper name to apply to it. Typhlops (Idiotyphlops)
emunctus Garman, Rhinotyphlops albirostris Peters and Helmin-
thophis canellei Mocquard have been described from Panama.
Helminthophis bondensis Griffin, described from Bonda, Colombia,
has been reported in Panama also. The postulation that these
species are based on anomalous specimens and are synonyms of
albirostris needs confirmation.

Knowledge of the presence of the genus in Costa Rica is based
on the report of a Costa Rican specimen of Liotyphlops albirostris
in "Frankfort 7005a" (presumably Germany). There is a specimen
of Helminthophis eumunctus in the U. S. National Museum said to
be from Costa Rica, collected by J. A. McNeil. McNeil collected
the type (M. C. Z. No. 3971 ) in Panama. Perhaps the "Costa Rica"
locality is an error.

26 The University Science Bulletin

Liotyphlops albirostris (Peters)

Rhinotyphlops albirostris Peters, Monatsb. Akad. Wiss. Berlin, 1857, p. 402 (type locality,

Veragua).
Liotyphlops albirostris Peters, Sitz. Gesell. Xatuif. Freunde, 1881, p. 69; Boeourt, Etudes

sur les Reptiles; Mission Scientifique au Mexique et dans 1'Amerique Centrale, livr. 8, 1882,

pp. 501-502, pi. 30, fig. 1, la-lc (figure of type); Dunn, Proc. Biol. Soc. Washington, vol.

45, Oct. 1932, pp. 174-175 (Costa Rica).

Head narrowed and flattened somewhat; rostral well developed,
narrowed at level of nostrils, half ( or slightly more than half ) width
of head extending back beyond level of eyes, in contact with the
frontal, which is at least twice as broad as long; two large pre-
frontals extending to posterior level of rostral but not in contact
behind rostral; no preocular; two suboculars; eye distinguishable
under the suture between the prefrontal and the small ocular; four
upper labials, first largest touching rostral, second in contact nar-
rowly with the prefrontal; the second and third supralabials separ-
ated from ocular by a subocular; scales in a row from mouth to anus
approximately 392-414; 11-13 under tail; 22 scale rows around body.

Blackish or dark brown, each scale with a reddish brown border.

I regard Liotyphlops eumunctus (Garman) as a species distinct
from L. albirostris. The scales of the type are approximately 425
mouth to anus, and 12 subcaudals. There are no enlarged anals.
The scales following the frontal are three, the median transversely
elongate.

Genus Helminthophis Peters

Helminthophis Peters, Monatsb. Akad. Wiss. Berlin, 1800. p. 517; Tihen, Copeia, 1945, no. 4,
Dec. 31, pp. 204-210; Smith and Warner, Herpetologica, vol. 4, pt. 0, Dec. 8, 1948, pp.
189-193.

Genotype: Helminthophis frontalis.

Helminthophis frontalis (Peters)

Typhlops (.Helminthophis) frontalis Peters, Monatsb. Akad. Wiss. Berlin, 1860, p. 517, pi.,
fig. 1, a-c (type locality, Costa Rica).

Helminthophis frontalis Boeourt, Etudes sur les reptiles; Mission Scientifique au Mexique et
dans l'Amerique Centrale, livr. 8, 1882, pp. 502-503, pi. 311, figs. 2, 2a-b; Boulenger, Cata-
logue of the Snakes of the British Mus., 2nd ed., vol. 1, pp. 5-6, 1893; Amaral, Proc. New
England Zool. Club, vol. 9. 192 4, p. 25; Mem. Inst. Butantan, Tomo 4. 1929, p. 136;
Dunn, Proc. Biol. .Soc. Washington, vol. 45, 1932, pp. 173-175 (San Jose, Costa Rica).

This diminutive, cylindrical, wormlike snake, without widened
ventral scales, may be recognized by the following characteristics:
dark brown above, a little lighter below; the head, neck and anal
region white; rostral large and broad, somewhat less than half
width of head, not extending back to the level of eyes, separated
from frontal by the prefrontals, which form a common suture; fron-

Taylor: Review of Snakes of Costa Rica 27

tal broad; two preoculars and one subocular; eye distinguishable
under the small ocular; four upper labials; ocular in contact with
third supralabial or narrowly separated by the subocular; ocular
scale separated from frontal by supraocular; upper preocular
touches the second supralabial, lower preocular touches the second
and third supralabials. Diameter of body about 58 times in total
length; longitudinal scale rows 22; 23 transverse scale rows on tail.
Total length, 158 mm.

I have examined a single Costa Rican specimen, M. C. Z. 34879.
This specimen has the head and anterior part of the neck whitish
cream without trace of pigment. The scale formula is 20-22-22.
The scales from mouth to anus are approximately 476. Two large
preoculars, the lower largely covering eye; ocular does not touch
labial; upper preocular also separated from labials; lower preocu-
lar touches two labials.

Family LEPTOTYPHLOPIDAE

A single genus of this family of diminutive snakes is known to
occur in Costa Rica.

Genus Leptotyphlops Fitzinger

Leptotyphlops Fitzinger, Systema Reptilium, 1843, p. 24.

Genotype: Leptotyphlops nigricans.

One species, Leptotyphlops albifrons, occurs in Costa Rica. It is
a widespread form that, as now understood, extends throughout
much of Central America and over a considerable area in northern
South America.

Leptotyphlops albifrons (Wagler)

Stenostoma albifrons Wagler, in Spix, Seip. Brasil sp. now, 1824, p. 68, pi. xxv, fig. 3; Du-
nieril and Bibron, Erpetologie Generate 1844, vol. 6. p. 327.

Leptotyphlops (=: Glaucoma) albifrons Wettstein, Sitz. Akad. Wiss. Wien Math.-Naturw.
Klasse, Abt. 1, Bd. 143, Heft 1-2, 1934. p. 31 (Guanacaste: Bebedero).

This tiny, wormlike snake may be readily recognized by the brown
coloration, somewhat lighter on venter than on back, each scale
with darker edges, the color tending to form indistinct lines; a
cream-white spot on top of head; lips and end of tail white.

Snout rounded; supraocular present, large, in contact or not
with first labial; rostral scarcely reaching back to level of eyes,
which are distinguishable under the ocular scale; nasal completely
divided; ocular bordering lips between two labials; six infralabials;
14 scale rows around the body, lacking distinguishable ventrals;

28 The University Science Bulletin

body diameter 45 to 55 times in head-body length; tail in total
length 15-21 times. Total length, 275 mm.

In Costa Rica the species is known from Bebedero, in the low
area in Guanacaste. I have not examined any Costa Rican speci-
mens.

Family BOIDAE

Three genera of this family are known to occur, each represented
by a single known species. These may be identified by the following
synopsis.

Key to Costa Rican Species of Boidae

1. Small snakes, probably less than one meter in length; ventrals 260; subcaudals
82 ; scale rows about middle of body, 54 ; nasals separated by a pair of small inter -
nasals; usually two or three enlarged loreals with the same number of smaller
scales below them; 12 scales between eyes. Ash colored with darker oval figures

on either side Boa annuiata

Large snakes from two to four meters in length 2

2. 61-79 scale rows around body; ventrals 225-252; subcaudals 47-65; 15-18 scales
between eyes. Pale brown with numerous dark brown crossbars or blotches each
usually with a pair of cream-colored dashlike marks; a dark line in middle of head

and neck Constrictor constrictor imperator

Scales 45-51 about body; ventrals 233-238; subcaudals 55-58; 2 enlarged supra-
oculars separated by 4-6 scales; usually an enlarged loreal, with one or two smaller
below it; a pair of normal internasals and a pair of normal prefrontals. Nearly
uniform brown in color, the spots obsolescent or absent .... Epicrates cenchria maurus

Genus Boa Linnaeu

s

Boa Linnaeus (part), Systema Naturae, vol. 1, 1766, p. 373; Wagler, Natiirliches System dor
Amphibien, 1830, p. 169.

Genotype: Boa canina Linnaeus.
One species occurs in Costa Rica.

Boa annuiata (Cope)

Xiphisoma annulatum Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1875, p. 129,
pi. 28, fig. 6 (type locality, Costa Rica); Bocourt. Etudes sur les reptiles; Mission scien-
tifique au Mexique et dans l'Amerique Centrale, livr. 8, 1882, pp. 526-527, and livr. 9,
1883, pi. 31, fig. 4, 4a.

Corallus annulatus Boulenger, Catalogue of the Snakes in the British Museum, vol. 1, 1893,
p. 102.

This small boa may be recognized by the following characters:
Body strongly compressed, nearly three times as high as wide; scales
in 42 rows on anterior part of neck, 54 near middle of body; top of
snout covered with small scales, of which two, a little larger than
the rest, separate the nasals and border the rostral. Top of head
covered with small smooth scales, twelve to fourteen rows between
eyes; no enlarged supraoculars; supralabials 15-15, the eighth to
twelfth strongly pitted on the sutures; 18-19 infralabials, the ninth

Taylor: Review of Snakes of Costa Rica 29

to seventeenth pitted; six scales in loreal region; four supraoculars;
four postoculars; two suboculars; one or two preoculars; one series
of scales separates the orbit from the labials and bounds the labial
pits. Ventrals 260; anal single; subcaudals, 82.

A Harvard M. C. Z. specimen (No. 37862 from Costa Rica [no
specific locality] ) has the head, in front of eyes, covered by some
40 small scales; eye bordered by 12 scales, the preocular largest;
nasal divided; six scales occupy the loreal region; rostral higher
than wide; chin shields small, scarcely distinguished from adjoining
chin scales, the first pair touching only two labials. The second
row of scales, back from eye border, is somewhat enlarged. An-
teriorly the pattern consists of some H-shaped median dark spots
between which are elongate oval fawn lines; the sides of H's tend
to separate forming paired lateral spots having light centers and
dark edges which may be opposite each other or alternate, or again
may be joined mesially. The pattern tends to darken and become
indefinite in old specimens.

The species has been recorded in Costa Rica from Ontario Farm
and La Castilla on the lower Reventazon River, and Limon. The
type specimen has been examined. It is from the Caribbean slope
in the southeastern part of the country.

Genus Constrictor Laurenti

Constrictor Laurenti, Specimen medicum exhibens Synopsin Reptilium emendatum, 1768,
p. 107.

Genotype: Constrictor formosissimus.

A single representative of this genus of boas occurs in Costa Rica.
It is the largest snake known in the fauna.

Constrictor constrictor imperator (Daudin)

Boa imperator Daudin, Histoue Naturelle . . . reptiles, vol. 5, 180.3, pp. 150-152 (type local-
ity, Mexico).
Constrictor constrictor imperator Ihering, Rev. Mus. Paulista, vol. 8, 1910, p. 321.

One large specimen, M. N. H. No. 25754, was taken on the
I. A. I. A. farm about three kilometers southwest of Turrialba and
presented to me by the several workmen who captured the specimen.
It measures a little less than 6 feet in length. The scale formula is
60-70-65-37; ventrals 243, subcaudals 67 ( several near vent divided ) ;
anal single but greatly narrowed as are the three ventrals preceding
anals; spots on body, 21/2; on tail 4/2 with extreme tip black; supra-
labials 23-22; infralabials 24-25; 17 scale rows between eyes.

This snake is common in the lowland areas in Costa Rica, and
reaches elevations up to 3000 feet and perhaps somewhat higher.

30 The University Science Bulletin

Genus Epicrates Wagler

Epicrates Wagler, Naturliches System der Amphibien, 1830, p. 108.

Genotype: Epicrates cenchris Wagler.

Epicrates cenchria mauriLS Gray

Epicrates maurus Gray, Catalogue of the Specimens of Snakes in the collection of the Bulish

Museum, 1849, p. 90 (type locality. Venezuela).
Epicrates cenchria maurus Stull, Proc. Boston Soc. Nat. Hist., vol. 40, no. 8, 1935, p. 390.
Epicrates cupreus var. concolor Bocourt. Etude sur les reptiles; Mission Scientifique au Mex-

ique. . . . Livr. 8, 1882, pp. 525-520, pi. 31, fig. 3, 3a (part). (Costa Rica.)

Generally similar to Epicrates cenchria cenchria. Snout covered
with regular scales but frontal and parietal areas covered with
small irregular scales; the elongate loreal touching first labial but
separated or not separated from second and third by small scales;
supraoculars usually present but may be broken into more than a
single scale; two preoculars, four or five postoculars; shallow pits
present in upper and lower labials. Scales in 45-48 rows around
body; ventrals 233-238 subcaudals 55-58, single; length (of a Costa
Rican specimen ) 1300; tail 157.

Uniform brownish, the dorsal spots almost, or entirely, obsolete.

Family COLUBRIDAE

Four subfamilies are being recognized. These are the Natricinae,
Sibynophiinae, Acrochordinae, and Colubrinae. It is probable
that the Colubrinae as here considered is divisible into several sub-
family groups. However, owing to the inadequate knowledge of
the anatomy of several genera, I am not recognizing certain of such
groups that have been proposed ( i. e., Xenodontinae, Dipsadinae,
Boiginae, Dromicinae, Leptognathinae, Erythrolamprinae, Scyta-
linae, etc. ) .

The Natricinae, Sibynophiinae and Acrochordinae are repre-
sented each by a single genus, while no less than 28 genera are in-
cluded in the Colubrinae as here considered.

Subfamily Acrochordinae

This small subfamily has one or two Asiatic genera and a single
genus, Nothopsis, in the Western Hemisphere.

Genus Nothopsis Cope

Nothopis Cope, Proc. Acad. Nat. Sci. Philadelphia, Oct. 24. 1871. p. 201.

Genotype: Nothopsis rngosus Cope.

Two species have been described in the genus. These are
Not1w})sis rngosus Cope from Darien and N. affinis Boulenger from

Taylor: Review of Snakes of Costa Rica 31

Salidero, N. W. Ecuador. A third form is here described as torresi
from Turrialba, Costa Rica. The range of the genus is from Costa
Rica to Ecuador.

Nothopsis torresi sp. nov.

Plate I

Type— University of Kansas Museum of Natural History No.
28719; taken at the Morehead Finca. 5 miles southwest of Turrialba,
Costa Rica, July 16, 1947, by Edward H. Taylor.

Diagnosis.— A small snake, heavily keeled, the scales in 26 or 25
rows about middle of body; outer edges of smooth ventrals turned
up and finely striated, as are the lateral scales; body strongly com-
pressed; nasal very large, single, with an entrant suture; two inter-
nasals; prefrontal area covered with 20-25 small granular scales;
supraoculars large or divided, separated from single frontal by a
row of small scales; large parietals present with a lateral entrant
suture tending to separate the posterior parts from the main scale;
eight rows of temporals; five postoculars, labials separated from
the eye by two rows of small scales; four or five preoculars, one en-
larged; ventrals 149; subcaudals 71; eighteen maxillary teeth.

Description of type.— Rostral nearly triangular, somewhat con-
cave, directed upward and forward, barely visible from above; a
pair of enlarged internasals, narrowed anteriorly, touching rostral,
a little longer than wide, in contact for three fourths of their length;
a small pair of scales following internasals somewhat larger than
granular scales that separate them; remainder of area, usually oc-
cupied by prefrontals in most snakes, broken up into numerous
(approximately 20) keeled scales, two of which are rounded and
symmetrical, separated from each other and frontal; supraocular
small, posterior part wider than anterior, separated from frontal
and parietals by a single row of scales; frontal about as wide as
long, its edges irregular, with a short suture entering from the front;
parietals each nearly as large as frontal, their edges irregular; a pair
of "postparietal" scales following, partly fused to parietals.

Nasal large, smooth with a depressed area, in which nostril is
pierced; a suture from lower part of scale partly divides it; a flaplike
valve? present in nostril; area between nasal and eye occupied
by a number of smooth scales, one somewhat enlarged touches two
labials and nasal, and another enlarged one enters border of eye as
a preocular; four other granular preoculars and five postoculars
present; one or two rows of suboculars separate the labials from the

32

The University Science Bulletin

^

Plate I. Nothopsis torresi sp. 7iov. Type. MNH No. 28719, More-
he;id Finca, 5 miles southwest of Turrialba, Costa Rica; total length,
401 mm.

Taylor: Review of Snakes of Costa Rica 33

orbit; temporal region with eight rows of small keeled scales be-
tween labial and parietals; 9-10 supralabials, smooth save for the
upper edges of the posterior scales which are striate. Mental tri-
angular, narrower than rostral; first infralabials in contact behind
mental; 12-13 infralabials, but posterior scales, partly fused together
(two on right side, four on left); a single pair of chinshields; about
ten small scales separate chinshields from first ventral.

Scales heavily keeled on body, less so on head; scale rows around
back of head, 39; narrow part of neck, 26 (25); 26 (25) about
middle of body; 22 in front of anus; five or six median scale rows
larger than lateral scales; ventrals 149; anal single; subcaudals 71,
divided, except first; terminal spine of tail directed somewhat down
at tip and partly scaled dorsally.

Eye small, about 2 1/5 times in its distance from nostril.

Color— Head black; labials fawn; chin cream; dorsal ground color
lavender-brown, growing lighter low on sides; a series of about
thirty-two saddlelike, rather irregular, dark blotches which widen
laterally and are in contact with each other on anterior half of
body; posteriorly they may be together or they may fail to touch
dorsally; a series of somewhat more discrete dark spots are immedi-
ately below the saddles; numerous scattered dots on fawn-colored
ventrals; pattern continued on tail, but much reduced; scales bor-
dering ventrals on sides of neck nearly white; beginning in the
occipital region are two very short cream lines, which are separated
by a median black area five or six scales in width.

Measurements in mm.— Head length, 13; greatest width of head,
7; width at eyes, 5; total length, 401; tail, 117; body, 284.

Teeth— Eighteen (17) maxillary teeth all bent back strongly,
increasing a little in size from fore to aft, then followed after a
short diastema by two larger, heavier teeth.

Relationships— -Two species of the genus are known— Nothopsis
rugosas (type) and Nothopsis affinis. The present form may be
distinguished by the presence of an even or odd number of scale
rows; the reduction of the scales about the middle of body and the
increase in the number of maxillary teeth, approximately 18 instead
of 12.

When Boulenger described his Notliopsis affinis,0 he had before
him a specimen of Notliopsis from Carriblanca, Costa Rica, which
he identified as Nothopsis rugosa Cope, a species whose type locality
is "Isthmus of Darien." IV. affinis is reported as having 27 scale rows

* Ann. Mag. Nat. Hist., (ser. 7) vol. 15, 1905, p. 453.

3—3286

34 The University Science Bulletin

and the type of rugosa has 29. The species here described has a
pair of median scale rows which at intervals are fused on the me-
dian line in series of two to five scales. Consequently, the count is
26 scales where the median scales are not fused, 25 where they
are fused.

Whether Boulenger's Carriblancan specimen is actually a mem-
ber of the species here described cannot at this time be determined.

Remarks— Part of the scales on the head are smooth and shining
(rostral, nasals, internasals, scales in loreal region for the most
part, and labials). It is possible, but I think not probable, that this
is due to wear. The other scales have heavy keels and the scale
surface is very finely roughened by what appear to be extremely
numerous fine keels or striations.

I collected the specimen at night quite by accident, when I
grabbed at a small anole in a pile of leaves on the ground, near the
edge of a swampy area in the forest. The specimen made no at-
tempt to bite.

The following table presents in tabular form the differences in the
three species.

Comparative Data on Species of Nothopsis

Sub- Scale Supra- Maxillary Total

Ventrals eaudals row labials teeth length Tail

rugosa 158-161 97-100 28-30 12 12 381 ?

affinis 162 98 27 10 ? 320 100

torresi 149-153 71-92 (25-26) 9-10 18 401 117

The species is dedicated to Sr. Prof. Ruben Torres Rojas, noted
educator and scientist of Costa Rica, and my official host in the
country.

I have examined a female specimen from the Reventazon Valley
(M. C. Z. Harvard No. 15269) which I have referred with a question
to Nothopsis torresi. It differs from the type in that there is no
tendency of the two median dorsal scale rows to fuse, the formula
being 26-26-22. Some of the characters of this specimen are: 25
small scales on the top of rostrum, those following internasals some-
what larger and regular; the supraoculars are broken into two or
three scales, separated from the frontal by a row of small scales;
the frontal has a groove causing it to appear paired; parietals large,
separated mesially by small granular scales; supralabials 10-10; in-
fralabials 11-13; ventrals 153; anal single; subcaudals 92.

I have compared the new form with a specimen of IV. rugosa Cope
from Panama. This specimen has the scale formula 30-28 (29) -26;
ventrals 161; subcaudals 97.

Taylor: Review of Snakes of Costa Rica 35

Subfamily Sibynophiinae

A single generic representative of this subfamily, Scaphiodonto-
phis, is known in the Western Hemisphere.

Genus Scaphiodontophis Taylor and Smith

Scaphiodontophis Taylor and Smith, Univ. Kansas Sci. Bull., vol. 29, pt. 2, 1043, pp. 302-304,
fig. 1.

Genotype: Enicognathus annulatus Dumeril, Bibron and Dume-
ril.

Only a single species of this genus has been reported from Costa
Rica. The range of the genus extends from northern Veracruz in
Mexico to Colombia.

Scaphiodontophis venustissimus (Giinther)

Hi micognathus venustissimus (Giinther, Biologia Centrali-Americana, Reptilia and Batrachia,
Oct. 1804, p. 144, pi. 51, fig. C (type locality, Matagalpa Hda. Rosa de Jerico [3250 ft.],
Nicaragua).

Sibynophis venustissimus Dunn. Occ. Papers Boston Soc. Nat. Hist., vol. 5, 1930, p. 330 (in-
correctly attributes the spelling of this name by Giinther as venistussimus ; perhaps a lapsus
mentum) (Zent, Costa Rica).

Scaphiodontophis venustissimus Taylor and Smith. Univ. Kansas Sci. Bull., vol. 29, pt. 2, 1043,
pp. 300-311. fig. 5.

The species may be distinguished by the following diagnosis:
frontal as long as its distance to rostral or a little longer; loreal rhom-
bic; one preocular; two postoeulars, both touching parietal; tem-
porals 2 + 2 or 1 -f- 2; supralabials nine, the fourth, fifth and sixth
entering orbit; chinshields of both pairs about same length, the
anterior the larger; scales in 17-17-17 rows, smooth; ventrals 137-
152; anal divided; subcaudals 96+; maxillary teeth small, 43-56 in
a continuous series.

Coral (or reddish olive) above, with 8-11 white-edged, trans-
verse, black bands, which terminate at the ventrals, these often
broken and alternating; top of head generally whitish from rostral
to anterior part of parietals with an indistinct smoky spot on snout;
this followed by a yellow transverse band to near posterior end of
parietals, and this in turn followed by a black band, and a white
band across neck five scale rows back of parietals. Upper labials,
chin and venter whitish but with some pigmentation forming in-
distinct spots on the edges of ventrals; each subcaudal with a large
black spot. Red scales with black tips.

I have examined the following Costa Rican specimens: M. C. Z.
No. 15308, Bonilla, Costa Rica; M. C. Z. No. 15272, Zent, Costa Rica;
and U. S. N. M. No. 67350 from Columbaria, Costa Rica.

36 The University Science Bulletin

In the latter specimen the snout is blackish or blackish brown but
the interorbital region is somewhat lighter. This is followed by a
large black spot occupying the posterior three fourths of the parie-
tals and the first six or seven scale rows on neck. The body has
red bands separated by eight white-black-white triads (four of
which are broken mesially, the halves alternating). Ventrals 152;
the tail is injured, the tip missing.

Subfamily Natricinae

Onlv one genus, Thamnophis, reaches Costa Rica.

Genus Thamnophis Fitzinger

Thamnophis Fitzinger, Systema Reptilium, 1843, p. 26.

Genotype: Coluber sauritus Linnaeus.

Thamnophis sirtalis* chalceus (Cope)

Prymnomiodon chalceus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, 1860 (1861), p. 558

(type locality "Siam" in error).
Tliamnopliis sauritus chalceus Dunn, Herpetologica, vol. 1, 1940, pp. 192-193.

The genus Thamnophis has numerous species and subspecies in
the United States, and the number of forms becomes even larger
in Mexico. The Isthmus of Tehuantepec limits the greater number
of these, and the range of only one is known to reach southward as
far as Costa Rica.

The single specimen at hand, M. N. H. No. 25679, is one collected
by Sr. Ruben Torres Rojas and Sr. Marco Tullio Pacheco about two
miles south of Cartago. This specimen, which the collectors pre-
sented to me, has the following characteristics:

Rostral once and one-third times as wide as high; internasals as
long as the prefrontals; latter nearly one third wider than long;
frontal more than one fourth wider than long, the length more than
one fifth greater than its distance from tip of snout; length of parietal
one sixth longer than the frontal, equal to its distance from the inter-
nasals; nasal divided, touching two supralabials; loreal about as
long as high or a little longer; preoculars, 1-1, two to three times as
high as wide, rather narrowly separated from the frontal; post-
oculars 2-3 ( on one side the two lower scales are almost completely
fused); temporals 1 + 2 + 1 (1 + 2 + 2); supralabials 8-8, the
fourth and fifth bordering eye; infralabials 10-10, five bordering
first chinshields; second pair of chinshields longer than first pair,
separated by small scales.

* Klauber has suggested that sirtalis as a species name applies to this species. Copeia, No.
1, 1948, pp. 8-10.

Taylor: Review of Snakes of Costa Rica 37

Scales finely corrugated or striate, each bearing a strong keel, and
a small terminal notch. The scale formula is 19-17-17; ventrals 156,
subcaudals 88; anal single. In life the specimen was olive, with a
dim median line, and a very dim line on the third and fourth scale
rows.

The specimen was taken near a shallow pond on which water
hyacinths were floating.

I have also examined U. S. N. M. No. 38149 from San Jose, Costa
Rica. The following scale characters were recorded: scale formula
19-19-19-17, all rows keeled; ventrals 151; subcaudals 84; one pre-
ocular touching frontal on one side, not on other; two-three post-
oculars; loreal present; nasal divided; a small entrant suture in the
frontal; temporals 1 + 2 + 2; 8-8 supralabials, the fourth and fifth
entering orbit; 10-10 infralabials, five touching the first pair of chin-
shields, which are shorter than second pair and separated by a small
scale.

A median dark-edged light line present on back begins between
parietals; two small spots present on parietals.

Subfamily Colubrinae
Twenty-nine genera are treated under this subfamily.

Key to the Genera of the Colubrinae

1. Nasal scales in contact behind rostral (rarely very narrowly separated); nostril

directed upward 2

Nasal scales not in contact behind rostral, the nostril directed outward laterally. . . 3

2 Internasals fused into a single scale behind nasals Helicops

Internasals not fused into a single scale Tretanorhinus

3. Prefrontals fused into a single scale 4

Prefrontals not fused into a single scale 5

4. Nostrils in a single nasal scale directed upward; large snakes (900 mm.); coloration

uniform Hydromorphus

Nostrils in at least semidivided nasals, directed laterally; small snakes, less than
300 mm. ; (part of genus only) Trimetopon

5. Lacking a median mental groove, the chinshields transversely arranged ; a loreal

and preocular both entering orbit ; four postoculars Neoparias

A median mental groove present ; the chinshields longitudinally placed t>

6. The maxillary teeth nearly equal without posterior enlarged fangs, or the size re-
ducing posteriorly 7

Maxillary teeth increasing in size, the last one or two somewhat fanglike not

grooved and may or may not be separated from the series by an interval 14

Anal plate single 8

Anal plate double Eluphe

No preocular; loreal and prefrontal bordering orbit of eye y

A preocular, the loreal and prefrontal separated from orbit 12

No anterior temporal Geophis

Anterior temporal present 10

10. Scales 17-21 rows, strongly keeled Ninia

Scales 15-13 rows, smooth 11

11. Teeth diminishing in size posteriorly Sibon

Teeth not diminishing in size posteriorly Dtpsas

38 The University Science Bulletin

12. Scales smooth ; scale formula 19, 19(21 )-17 Drymarchon

Scales keeled 13

13. Four median scale rows keeled; scale rows uneven; scale formula 21-25-15. . .Pseustes
Dorsal scales, save outer row, keeled; scale rows in even numbers 14(16)-16-

10 Spilotes

14. Posterior fanglike teeth separated from other maxillary teeth by a toothless

interval 15

Posterior enlarged teeth of maxillary series not separated by a toothless interval
from the iemainder of the maxillary teetli 17

15. Body slender, the tail 30-39% of total length; small snakes; four or five maxillary

teeth followed by fang Enulius

Tail less than 25% of total length 1»'>

10. Scales arranged in oblique transverse rows; large thick-bodied snakes, length one

meter or more) Xenodon

Scales not arranged in oblique rows; smaller snakes, 800 mm Leunadoplus

17. No loreal present, tail one third total iength ; scales 17 rows, faintly keeled pos-
teriorly imastridium*

Loreal present IS

18. Scales about body in even numbers Chironius

Scales about body in odd numbers 19

19. No apical pits present 20

Apical pits present 21

20. Body banded black and red or black, yellow and red (the red and yellow fading

in preservative to cream or white Pliocercus

Body not banded; pattern, if present, of dark and light stripes or lines. . . .Rhadinaea

21. Eye with ellpitic pupil Hypsiglena

Eye with round pupil 22

22. Diminutive snakes probably not exceeding 250 mm. in length; scales in 15 rows

(part of genus only) Trimetopon

Medium to large snakes, all reaching 700 mm. or more in length 23

23. Body black crossed by whitish lines; belly yellow or coral red with transverse

black spots (cobella) Liopliis

Not so marked 2 4

24. Body banded in red, black and yellow or these bands tending to become obscured

(darker) Lampropeltis

Not so marked 25

25. Four black longitudinal stripes; rostral projecting, part visible above about hall

distance to frontal Salvadora

Not so marked 20

2G. Scales smooth 27

Scales keeled 28

27. Scales in 17-17-15 rows; 9 supralabials Dryadophis

Scales in 17-15-13 rows; 7 or 8 supralabials Masticophis

28. Scales in 17 rows around body 29

Scales in 15 rows reducing to 11 in front of anus Thalerophis

29. Maxillary teeth generally more than 33 Dendrophidion

Maxillary teeth generally less than 33 Drymobius

Genus Geophis Wagler

Geophis Wagler, Natiirliches System der Amphibien, 1838, p. 194.

( renotype: Catostoma chalybeum Wagler.

Six species are recognized as occurring in Costa Rica.

Key to Costa Rican Species of Geophis

1. Scales smooth in 17 rows, no supraocular; ventrals 135-141: subcaudals 3<i-43

rhodogastet
Scales in 15 rows, smooth or partly keeled; supraocular present or absent 2

* (Mirnetopon sapperi Werner, sometimes placed in the genus Amastridium is reputed to
have a minute groove visible at a magnification of 80 diameters.

Taylor: Review of Snakes of Costa Rica 39

2. Scales smooth; no supraocular; ventrals 144-145; subcaudals 27-28. Blackish
brown above, yellowish or whitish below; a few transverse blotches under tail

occasionally present on posterior part of belly godmani

Scales at least partially keeled ; a small supraocular present 3

3. Scales keeled at base of tail; supralabials five (rarely six): ventrals 129-133,
subcaudals 31-36; black above, below brownish yellow or whitish; a light nuchal

band in young hoffmanni

Scales keeled and striate on at least one third or more of body and tail ; color
variable ; supralabials six 4

4. Large red spots alternating or opposite on sides of body; ventrals 131-138; sub-
caudals 39-46; scales striated and strongly keeled on posterior half of body; head

narrow elongate dolichocephah

No large red spots on sides of body; scales striate and keeled on greater part

of body 5

5. Infralabials 7; ventrals 131-1411; subcaudals 32-39; blackish with a dim nuchal
collar; venter whitish with some pigment extending onto ventrals (rarely nearly

across) tnoesta

Infralabials 6 (7): ventrals 143-144; subcaudals 34-35. Black with a narrow lat-
eral orange stripe; venter smoky black, each scale with a lighter edge, .hrachycephala

[Geophis rhodogaster (Cope)]

Calophrys rhodogaster Cope, Proc. Acad. Nat. Sci. Philadelphia, Mar. 1868, p. 130, fig.

("Neighborhood of the city of Guatemala").
Geophis rhodogaster Bocourt, Etudes sur les reptiles; Mission Scientifique au Mexique et dans

l'Amerique Centrale, livr. 9, 1883, p. 531, pi. 31, fig. 12, a-c; Boulenger, Catalogue of the

Snakes in the British Museum, vol. 2, 1894, p. 317.

Eye small, measuring two thirds to three fourth its distance from
mouth; snout rounded, feebly projecting; rostral moderate, slightly
broader than deep, the part visible from above about one third its
distance from the frontal; internasals nearly twice as broad as long,
one half to two fifths of the length of the prefrontal; prefrontals
broader than long; frontal broader than long, as long or a little
longer than its distance from the end of snout; a little shorter than
parietals; no supraocular; frontal bordering eye; loreal nearly twice
as long as deep; one postocular; six supralabials, third and fourth
bordering eye; three infralabials border first chinshields, which
equal or are longer than the posterior, separated from mental;
scales in 17 rows, perfectly smooth; ventrals 135-144; anal single;
subcaudals 30-43.

Uniform dark brown above; upper lip, outer row of scales and
lower parts uniform orange.

The species is reported from Costa Rica by Boulenger, Joe. cit.
Giinther (Biologia Centrali-Americana; Reptilia and Batrachia,
1893, pp. 87-88) mentions a specimen in Smithsonian Institution
as a basis for a Costa Rican record. Boulenger, op cit., also gives
Costa Rica as within the range of rhodogaster, perhaps on the basis
of Giinther's statement. I regard the record as somewhat doubtful.

40 The University Science Bulletin

Geophis godmani Boulenger

Geophis godmani Boulenger, Catalogue of the Snakes in the British Museum, vol. 2, 1894,
p. 322, pi. 16, fig. 4 (type locality, Irazu, Costa Rica); Dunn, Notulae Naturae, no. 108,
1942, p. 4.

Eye small, a little shorter than its distance from the mouth; snout
obtusely pointed, very prominent; rostral rather large, a little higher
than broad, the portion visible above measuring two thirds to three
fourth its distance from frontal; internasals a little broader than
long, half as long as the prefrontals, which are a little longer than
broad; frontal as long as broad, as long as or a little shorter than
its distance from end of snout, shorter than parietals; no supraocu-
lar; prefrontal forming a narrow suture with the parietal between
frontal and eye; loreal once and a half times as long as broad, enter-
ing orbit; a minute postocular; six supralabials, the third and
fourth entering orbit; three infralabials touching chinshields which
are not longer than posterior and are separated from mental; scales
in 15 rows, all perfectly smooth; ventrals 144-145; anal single; sub-
caudals 27-28. Uniform blackish brown above, yellowish beneath;
a few transverse brown blotches under tail, sometimes also on bell v.
Total length, 400 mm.; tail, 55 mm.

Known in Costa Rica from the type locality. Volcan Irazu; from
Escazii, and Tierra Blanca.

Geophis hoffmanni (Peters)

Plate II, figs. 1-2

Colobognathus Hoffmanni Peters, Monatsb. Akad. Wiss. Berlin, 1859, p. 276, pi., fig. 2-2c

(type locality, Costa Rica).
Geophis hoffmanni Bocourt, Etudes sur les reptiles; Mission Scientifique au Mexique et dans

l'Amerique Centrale, livr. 9, 1883, pi. 31, figs. 8, 8a-c; Dunn, Notulae Naturae, no. 108,

1942, p. 4 (mentions 40 Costa Rican specimens, from 11 localities; also specimens from

Boquete, Panama).

Four specimens of Geophis hoffmanii were collected: Two on
the Pacific drainage near San Isidro El General, and two in the Cen-
tral Plateau: one at Turrialba and one at Cartago.

Description from No. 1032.— Portion of rostral visible above, a

PLATE II

Fig. 1. Geophis hoffmanni (Peters), RCT No. 346. Turrialba, Costa Rica,
July 13, 1945; total length. 201 mm. Ventral view.

Fig. 2. Geophis hoffmanni (Peters), MNH No. 1032. San Isidro El General,
August 22, 1947; total length, 192 mm. Ventral view.

Fig. 3. Ninia tessellata (Peters). RCT No. 1445, Los Diamantes, Costa Rica,
September 1-8, 1947; total length, 233 mm. Ventral view.

Fig. 4. Ninia tessellata (Peters), RCT No. 1364, Los Diamantes, Costa Rica,
September 1-8, 1947; total length, 279 mm.

PLATE II

42 The University Science Bulletin

little larger than an internasal; nostril between two subequal nasal
scales; internasals small, about one fifth as long as the prefrontals,
and about one sixth their area; frontal six-sided, its width minutely
greater than its length, its length about one-seventh longer than its
distance from tip of snout; length of parietal equal to its distance
from the rostral; loreal elongate nearly twice as long as high enter-
ing orbit; supraocular small, scarcely larger than postocular; no
preocular, one postocular; no anterior temporals, the fifth (and
last) labial broadly bordering the parietal; five supralabials, the
third and fourth border eye; six infralabials, the first three border
the first chinshields; these equally as long as second pair; second
chinshields partly separated by a large scale.

Scale formula, 15-15-15; all smooth except at the base of the tail,
at which point the scales are small ( 15 scale rows ) and keeled ( in
both sexes); ventrals, 133 (2 small scales between ventrals and
chinshields); subcaudals, 31; anal single.

Total length, 192 mm.; tail, 29 mm.; width of head, 3.5 mm.;
length of head to end of parietals, 6 mm.

The color in life is generally bluish black, more or less iridescent
on dorsal and lateral surface of body and head; no neck band; ven-
trals flesh white, the white extending onto the outer scale row, for
the most part covering less than half the scale row; a peppering of
smoky black pigment on the ventrals usually extending across the
outer third on each side; darker under the tail but some light color
on each subcaudal.

The other specimens differ but little in color from the one de-
scribed. One specimen has the ventrals without pigment for nearly
two thirds the length of the body. In another, smoky pigment en-
croaches on the white part of the first scale row. The squamation
of the specimens is very uniform. All have the scales smooth save
at the base of the tail where there are 15 keeled scale rows reduced
in size. The scale formula is the same in all. The relation of the two
pairs of chinshields is the same, and a median scale tends to sepa-
rate the posterior part of the second pair, both pairs being tubercular
in males. Five is the constant number of supralabials, the last very
large, broadly bordering the parietal. There are no primary tem-
porals lying between the parietals and labials. One secondary
temporal borders the parietal behind the fifth labial in all.

It would appear that these specimens, one from the Caribbean
drainage (Turrialba No. 346) and two from the Pacific drainage
( San Isidro El General No. 880, No. 1032 ) and one from the central
plateau region (Cartago No. 1020), represent a species that shows

Taylor: Review of Snakes of Costa Rica 43

very little variation despite the diverse geographical areas it occu-
pies. I have examined several Harvard specimens as follows:

A Harvard M. C. Z. specimen, No. 15304, from Navarro, Costa
Rica has four supralabials, the third and fourth being fused; infra-
labials five; ventrals 134, subcaudals 29; lateral scales keeled at the
base of the tail.

No. 15267 £ , has five supralabials and six infralabials; a small
supraocular present, not in contact with the small postocular; ven-
trals 126; tail broken; keels present at base of tail.

No. 15271 $ , from La Palma, Costa Rica has the ventrals with
brownish pigment tending to cross the ventrals; ventrals 133; sub-
caudals 28; keels near tail base scarcely indicated.

Nos. 19327 s and 19328 5 from Suretka, Costa Rica, have re-
spectively 124 ventrals, 37 subcaudals; 134 ventrals. 30 subcaudals.
In the former specimen the loreal narrowly enters orbit; in the latter,
it is excluded from orbit. Both specimens have five supralabials
and six infralabials.

Two specimens in the U. S. National Museum are from Cartago
and San Jose, Costa Rica.

Geophis dolichocephala <Cope)

Plate III. fin. 2
Colobognathus dolichocephahis Cope, Proc. Acad. Nat. Sci. Philadelphia, 1871 (Oct. 24). p. 211

(type locality, San Jose, Costa Rica).
Colostoma dolichocephalw Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1875,

p. 147.
Elapoidis dolichocephahis Cope, U. S. Nat. Mus. Bull. no. 32, 1887, p. 85 (San Jose).
Geophis dolichocephalus Boulenger, Catalogue of the Snakes in the British Museum, vol. 2,

1894, pp. 320-321 (Cartago, Costa Rica).
Geophis brachycephalus (part.) Dunn. Notulae Naturae, no. 108, 1942, p. 5.

A specimen of this species (No. 25732) was taken in a pile of
chips at the edge of a tiny rivulet on the Morehead finca, five miles
southwest of Turrialba, Costa Rica.

Color brownish black above with a series of 21 brick-red blotches
on each side. Anteriorly blotches scarcely distinguishable and
separate; on the latter two thirds of the body they are confluent
mesially or separated by a very narrow dark line. On sides the
spots, which are rarely more than three scales long, reach to first
scale row. A small red spot present near angle of jaw. Belly flesh
white with some olive reflections; head blue-black with metallic re-
flections. The lower labials and chinshields black, and tail black
without red spots. Caudals black with indefinite lighter areas on
each scale. The dark color of sides extends onto ventrals a greater
or lesser distance. Most scales of outer row have a narrow diagonal
cream spot, which in the aggregate suggests a zigzag line.

44 The University Science Bulletin

The scale characters of this species follow: rostral visible above
broader than high; internasals small, subtriangular, about one fourth
the total length of the prefrontals, about one third of the median
suture between prefrontals; frontal very broad, only minutely longer
than broad, almost quadrangular; supraocular diminutive, as wide
as long or minutely longer; parietals elongate, (4.8 x 3.3 mm. ), shorter
than their distance from snout tip; nasal divided, the nostril pierced
chiefly in anterior part; an elongate loreal entering eye and replacing
the missing preocular; a small postocular; no anterior temporal;
one posterior temporal; 6 supralabials, the third and fourth entering
the orbit; 7-7 lower labials, four touch first pair of chinshields;
second pair of chinshields smaller than first; eyes small, contained
in distance from snout tip about three and one-third times. Scales
smooth or very finely striate on neck; striate and keeled over re-
mainder of dorsal surface except outer enlarged scale row; scale
formula, 15-15-15; ventrals 142; anal single, subcaudals 48. Total
length, 359 mm.; total ventrals and subcaudals 190; snout to vent,
283 mm.; tail, 76 mm.; head length, 11.6 mm; head width, 6 mm.

The type of this form is described as having only 13 scale rows
but Boulenger (loc. cit. footnote) states that Cope has checked the
type and finds 15 scale rows present. The type has 131 ventrals,
39 subcaudals; total ventrals and subcaudals 170, which is 20 less
than this specimen. Boulenger lists two specimens having ventrals
135, 138 and subcaudals ? and 46.

Geophis moesta Giinther

Geophis moestus Giinther, Ann. Mag. Nat. Hist., ser. 4, vol. 9, 1872, p. 15 (type locality,
Cartago, Costa Rica); and Biologia Centrali-Aniericana, Reptilia and Batrachia, May 1893,
pp. 90-91, pi. 33. fig. C

Geophis hofimanni Boulenger, Catalogue of the Snakes in the British Museum, vol. 2, 1894,
pp. 319-320 (all or part).

Geophis brachycephalus part. Dunn, Notulae Naturae, no. 108, 1942, p. 4.

Dunn, op. cit., has recently placed this form in the synonymy of
Geophis brachycephalus. This I regard as an error since the species
appear to be distinct.

The following are the characteristics of M. N. H. No. 25738 taken
J2 mi. NE of Santa Cruz, south slope of Volcan Turrialba: scales
15-15-15; supralabials 6; infralabials 7-7; scales keeled throughout

PLATE III

Fig. 1. Dipsas annulata (Giinther), M.N.H. No. 25702, Isla Bonita, Costa
Rica, August 5, 1947; total length, 501 mm.

Fig. 2. Gcophis dolichocephala (Cope). M.N.H. No. 25732. Morehead
Finea, 5 miles SW Turrialba, Costa Rica, July 14, 1947; total length, 359 mm.

PLATE III

46 The University Science Bulletin

latter three fourths of body; anterior chinshields larger and longer
than posterior (or nearly of same size), which are partially or en-
tirely separated by a median scale in one specimen. Ventrals, 131;
subcaudals, 39; anal single. A black spot on anterior chinshields;
supraocular present, twice as large as the single postocular; a light
band across back part of head and anterior part of neck (tending
to disappear in old adults); posterior nasal much larger than an-
terior; fifth labial touches parietal; one secondary temporal; three or
four infralabials touch first chinshields.

A second female specimen (R.C.T. No. 839), from about one-half
mile east of Santa Cruz, has 140 ventrals and 32 subcaudals. In
this the infralabials are 7-6; the posterior chinshields are not sepa-
rated and are only a little smaller than the first pair.

Both specimens are blackish with a dim nuchal collar, the venter
being whitish with some dark pigment on outer edges of ventrals.
Posteriorly in one specimen (No. 25738) the dark color extends
almost across the ventrals; the subcaudals likewise blackish, each
scale with a narrow transverse light line across the middle. There
is a brownish area on the top of head. When submerged in water,
No. 839 shows the outer borders of body scales slightly darker than
their median parts. The infralabials are white in both specimens
and some lighter color may be evident on the anterior scales of the
outer row.

Geophis brochycephala (Cope)

Plate IV. figs. 2-3. text fig. 1

Colobognathus brachycephalum Cope, Proc. Acad. Nat. Sci. Philadelphia, 1871, p. 211 (type

locality, Costa Rica).
Catostoma brachycephalum Cope, Jonrn. Acad. Nat. Sci. Philadelphia, 2d ser., vol. 8, 1875

and 1876, p. 117.
Dirosema brachycephalum Boulenger, Catalogue of the Snakes in the British Museum, vol. 2,

1894, p. 299.
Geophis brachycephalus* (part.) Dunn. Notulae Naturae, no. 1(18, 1942, p. 4.

Two specimens of this rare snake were found under small logs on
the side of Volcan Turrialba at an elevation of near 6000 feet, about
three kilometers above the village of Santa Cruz. The general
coloration is blue black with an orange-red lateral stripe (not of
solid color) two or three scales wide anteriorly but posteriorly about
one scale wide; venter dusky, each ventral scale lighter edged.

These specimens were coiled tightly, almost into a ball under the
same logs as were found a number of leeches equally as large, or
larger, black in color with red markings low on sides and in ventral

* Dunn includes dolichocephalum, moesta, quadrangularis and championi as synonyms of
this species.

Taylor: Review of Snakes of Costa Rica

47

regions. These were likewise coiled into a ball-like mass and occupy-
ing small excavations or depressions below the logs as were the
snakes. In each case where a snake was found, one or more leeches
was present. On superficial examination they are very similar to
the snakes. Under some logs only the leeches were found, invariable
rolled into a knotted ball-like mass. I found no evidence that the
leeches preyed on the snakes. Since they are larger or equally
as large as the snakes, I think it unlikely that the snakes use the
leeches for food. However when placed in a sack, a leech fastened
on one of the snakes. I regard this as a matter of chance.

A B

Fig. 1. Geophis brachycephala M.X.H. No. 25735. Collected 3
miles NE Santa Cruz, on the southern slope of Volcan Turrialba.
A, Head, dorsal view; B, Head, lateral view; C, Head, ventral view.
X4.

The scale characters of the two specimens (R. C. T. No. 840 and
M. N. H. No. 25735 respectively) are given: Scale formulae, both,
15-15-15; ventrals, 144, 143; subcaudals, 34, 35; anal, 1; no pre-
oculars; postocular, 1-1; supraocular, 1-1; supralabials, 6-6; supra-
labials enter eye, 3, 4; infralabials, 6-6; touch chinshields, 3-3; total
length and tail length, 245-35; 230-24.

Area of frontal visible above smaller than or nearly as large as an
internasal; latter scales less than one third the length of the pre-
frontals; latter very large, about five times area of internasals;
frontal hexagonal, about 1/6 wider than long, its length greater
than its distance from tip of snout; parietals as long as their distance
from the tip of the snout; nasal divided; no anterior temporal, the
fifth labial bordering the parietal; one posterior temporal; loreal

«_ '**

>- .3"

■ •

.-.-:'

Plate IV. Fig. 1. Coniophanes fissidens punctigularis Cope, Los Diamantes,
1 mile south Guapiles, Costa Rica, September 2. 1947; about natural size.
RCT No. 1369.

Fig. 2. Geophis brachycephala (Cope), RCT No. 840, Volean Turrialba,
Costa Rica; elevation, approximately 6000 feet; total length, 245 mm. Ventral
view.

Fig. 3. Geophis brachycephala. Same as fig. 2 dorsal view (retouched).

Taylor: Review of Snakes of Costa Rica 49

two, to two and one-half times, as long as high; first chinshields
about as long as but larger than second pair (in one specimen dis-
tinctly longer); scales smooth anteriorly, keeled and finely striated
on posterior half of body and tail.

Black above, with a salmon to orange-red lateral stripe made up
of flecks, extending from neck to tail, but growing more faint pos-
teriorly. The stripe covers much of the second and third rows but
posteriorly it is on the third and fourth. Occasionally scales of the
scale row above the stripe are colored likewise, suggesting a series
of spots connected by the stripe. Venter smoky black with a
whitish line across each ventral not strongly contrasted; each sub-
caudal smoky black with a lighter area; a blackish spot on anterior
chinshields.

On No. 840 the lateral stripe is less distinct than in No. 25735.
The type is said to have a yellowish collar, a character that may be
confined to younger specimens.

Two specimens of the species examined at Harvard ( M. C. Z.
Nos. 15321 and 28070) have respectively 142 ventrals, 43 sub-
caudals; 141 ventrals and 34 subcaudals; supralabials 6, 6; infra-
labials 6, 7; the scales keeled and striate on body; there is an irregu-
lar orange stripe on sides of body in both.

Genus Ninia Baird and Girard

Ninia Baird and Girard, Catalogue of North American Reptiles in the Museum of Smithsonian
Institution, Pt. 1, Serpents, 1853, p. 49.

The small ground snakes of the genus Ninia are numerous in
Costa Rica, six forms being recognized. All the members of the
genus are small and slender, rarely reaching half a meter in length.

Synopsis of Costa Rican Forms of Ninia

1. Scales in 17 rows 2

Scales in 19-21 rows 3

2. Dorsal coloration uniform black ; venter black, part of the ventrals with red
marks; 6 supralabials; a sharp median keel, the body triangular in cross section;

ventrals 162 ; subcaudals 73 psephota

Dorsal coloration varied, consisting of numerous blackish or brownish bands nar-
rowing on sides, separted by narrow whitish, or brownish-white zigzag lines
usually considerably less than width of one scale-length ; a more or less distinct
series of dark spots low on sides alternating with the larger dark bands; nuchal
band approximately one scale wide, incomplete mesially; top of head dark with
some small lighter flecks or dashlike marks ; belly checkered with black and greenish
white; a sharp keel along middorsal line; 6 supralabials (rarely 5); ventrals 158;
subcaudals 68 + 1 ■ oxynota

3. Belly checkered with black and yellow (white or greenish white) 5

Belly never checkered with black and yellow 4

4. Above uniform black with a yellow collar; belly immaculate or with some scat-
tered pigment ; ventrals 129-151 ; caudals 34-67 ; supralabials 7-8 atrata

4—3286

50 The University Science Bulletin

Not uniform black above ; reddish or reddish brown above with black cross bars
(rarely lacking); a yellow collar followed by a black one; ventrals 131-147; sub-

caudals 36-70 ; supralabials and infralabials 7 sebae sebae

5. Brown above on body with a series of about 25 (or less) zigzag transverse black
bands, sometimes light -edged, or posteriorly broken medially and alternating, be-
coming smaller toward, and on tail ; belly checkered, the dark marks tending to
form a single median, or two outer bands, or the marks may be scattered. . . .maculata
Brownish with a broken series of fifty or more narrow, dark zigzag transverse
marks usually broken anteriorly, usually fused on most of body ; lower scale rows
may have much black pigment appearing as black flecks; upper labials with round-
ing white spots; neck colored like body; belly checkered irregularly with black and
greenish white ; sometimes a small subocular present tessellata

Ninia sebae sebae (Dumeril, Bibron and Dumeril)

Streptophorus sebae Dumeril, Bibron and Dumeril, Erpetologie Generale, vol. 7, pt. 1, 1854,

lip. 515-517 (type locality, Mexico).
Ninia sebae Dunn, Proc. Nat. Acad. Sci., vol. 21, 1935, pp. 10-11 (Costa Rica and numerous

other localities).
Ninia sebae sebae Smith and Taylor, Bull. U. S. Nat. Mus. no. 187, 1945, p. 100.

I have seen no specimens of this species from Costa Bica. It is
reported by Dunn, loc. cit., from "upper Costa Bica."

The color is red or reddish above usually with black cross-bands,
which tend to break mesially, and may alternate on the posterior
part of the body. Towards the tail the bands may be reduced to
small dots. There are usually 25 (or less) present or indicated on
the body. There is a strongly-defined black collar preceded by a
yellow band. The venter is immaculate or with some scattered pig-
ment not, or only rarely, forming small spots, never checkered like
maculatas.

See synopsis of the genus for diagnostic scale characters.

Ninia at rata (Hallowell)

Coluber atratus Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1854, p. 245 (Colombia [Vene-
zuela] less than 200 mi. from Caracas).

Ninia atrata Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1S75. p. 145 (San Jose,
Costa Rica); Dunn, Proc. Nat. Acad. Sri., vol. 21, 1935, p. 11; Dunn and Bailey, Bull.
Mus. Comp. Zool., vol. 86, no. 1, 1939, p. 7.

? Streptophorus atrata Boulenger, Catalogue of the Snakes in the British Museum (Natural
History), vol. 1. 1893. pp. 293-295 (Cartago, Costa Rica).

See synopsis of the genus for a diagnosis of this form.

The Cartago specimens reported by Boulenger have 142 ventrals
and 62 subcaudals in the male; 141, 155, 140 ventrals, and sub-
caudals 50, 53, 52 in three females. Dunn lists two specimens from
Cariblanco and Cope reports the species from San Jose. I have
seen two specimens in the U. S. National Museum from San Jose,
Costa Bica (Nos. 38147-48). Both are typical in color.

Two specimens in Harvard M. C. Z. (Nos. 15292-93 from Cari-
blanco) have the ventral-subcaudal counts respectively, 130-34
and 126-44. The light color is five scales wide and includes the

Taylor: Review of Snakes of Costa Rica 51

posterior parts of the parietals, some pigment being present; supra-
labials 8, with small black spots at the back of each scale. The
belly is completely immaculate except where the dorsal coloring
edges onto the ventrals; mental and first infralabials with some
black markings.

Ninia maculata I Peters I

Plate V, figs. 1-3

Slreptophorus maculata Peteis, Monatsb. Akad. Wiss. Berlin, 1861. p. 924 (type locality,
Costa Rica): Bocourt, Etudes sur les reptiles; Mission Seientifique an Mexique at dans
l'Amerique Centrale, livr. 9, 1883, p. 584, pi. 33, fig. 3.

A series of specimens (M. N. H. Nos. 25690-98) was obtained at
Cervantes, a village situated between Turrialba and Cartago at the
site of an old lava flow from Volcan Irazu. The small snakes were
ensconsed under lava rocks.

The series shows a certain uniformity of color and markings, the
color being brighter and the pattern more distinct in younger speci-
mens. The ground color is reddish brown in younger specimens
with small narrow cross bands usually some of which are broken
and may tend to alternate on the two sides; towards the posterior
part of the body the marks become less distinct, often represented
by only small blackish lateral dots. Older specimens have more
dark pigment scattered in the brown so that the dark markings are
less distinct. The ventral surface is light with a single median
series of quadrangular (rarely triangular) black spots, or two
series, one on each side; or the spots may be irregularly scattered.
See plate no. V. On the first scale row, very small black spots are
present at irregular intervals, each smaller than a single scale. The
underside of the tail is black.

There is a well-defined black nuchal band usually about as long
as three scale-lengths; a band colored like the body is in front of
this, three scale-lengths long but on the sides of the head it runs
forward to the eye covering the outer edges of the parietals. The
dorsal surface of the head is blackish or blackish brown, with occa-
sionally a distinct black spot on the posterior tips of the parietals.

The following scale characters obtain: the part of rostral visible
above forming a broad triangle, its length less than that of inter-
nasals; latter less than one third area of prefrontals; frontal shield-
shaped usually distinctly hexagonal, as broad as long, its length
equal to its distance from the tip of the snout; parietal large, its
length equal to its distance from tip of snout; nasal not completely
divided; loreal elongate, nearly twice as long as high; no preocular;

I --.-..

/^wr^

-A n^o

^

Plate V. Fig. 1. Ninia maculata (Peters), RCT No. 297, Cervan-
tes, Costa Rica, July 9, 1947; total length. 307 mm.

Fig. 2. Ninia maculata (Peters), RCT No. 1452, Cervantes, Costa
Rica, August 26. 1947; total length, 229 mm.

Fig. 3. Ninia maculata (Peters). RCT No. 303. Cervantes, Costa
Rica, July 9, 1947; total length. 268 mm.

Taylor: Review of Snakes of Costa Rica 53

postoculars two; temporals 1+2+3; supralabials seven, third and
fourth entering eye; infralabials seven, the first four bordering the
first chinshields, of which the first pair is nearly double the length
and area of second pair; no scales intervene between ventrals and
second chinshields. Scales striated strongly and all rows bearing a
strong medial keel. Scale formula 19-19-17(19). Ventrals in the
males range from 132-138; subcaudals 52-58; ventrals in females,
138-144; subcaudals 51-58. Total counts are 189-191 for males,
192-198 for females. Males have strong series of tubercles on men-
tal, first pair of infralabials, the first chinshields and on some of
the supralabials.

One specimen, also referred to this species, was collected on the
Pacific drainage slope near the western base of Cerro de la Muerte
at San Isidro El General. It differs in having the neck band nearly
double the width of that of the Cervantes specimens and connecting
bv an isthmus to the black color on the head. The ventrals of this
specimen are 139; the subcaudals 53.

It is probable that Ninia maculata is more closely related to Ninia
sebae than to other Costa Rican species. The heavy black quad-
rangular markings on the venter of N. maculata will readily separate
the two forms.

A series of Costa Rican specimens are in the U. S. National Mu-
seum. The variation in ventral count is 137-144, the average being
about 140 (six specimens counted); subcaudals 48 to 55. Three
exact localities are represented: Arriba, San Jose and Navarro.

Three specimens from Navarro, Costa Rica in the Harvard col-
lection (M. C. Z. Nos. 15315-15317) have the following ventral and
subcaudal counts respectively: 134-57, 141-48, 140-52.

Ninia tessellata Cope

PI. 2, figs. 3 and 4

Ninia sebae tessellata Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875),
p. 145 (type locality, ? Sipurio, Costa Rica by inference).

tStreptophorus subt essellat us Werner, Mitt. Naturh, Mus. Jahrg. 20. 1909 (1910), pp. 215-
216 (type locality, "Carriblanco" Costa Rica); ? Amaral, Mem. Inst. Butantan, vol. 4,
1929, p. 10.

Ninia maculata Dunn, part, Proc. Nat. Acad. Sci., vol. 21, 1935, p. 11 (Canal Zone, Nic-
aragua, Costa Rica).

Five specimens of this species were obtained at Los Diamantes
(R.C.T. Nos. 1364, 1445, 1447, 1448, and M.N.H. No. 25739) Sept.
1-8, 1947. All were found under logs.

Rostral about one third wider than high, barely visible above;
internasals small, their length in that of prefrontals two and one

54 The University Science Bulletin

fourth times; prefrontals distinctly longer than wide entering the
orbit; frontal about one fifth longer than wide, its length a little
greater than its distance from snout tip, at least four times as wide
as supraoculars; parietals elongate, equal to their distance from
tip of snout; scales bordering parietals behind are not, or scarcely,
enlarged; nasal divided; loreal large, quadrangular, entering orbit;
preoculars normally absent ( in some specimens where present they
are segmented from the prefrontal if above loreal, from the third
labial if below the loreal); two postoculars, the upper the larger;
head scales finely striated with distinct tubercles on loreals; in
males two anterior supralabials, mental, first four infralabials and
both pairs of chinshields with tubercles. Scales of body strongly
striate, the striation occurring also on outer edges of the ventrals.
The scale formula 21-19-19-19, all rows strongly keeled; first chin-
shields double the area and the length of second pair. Two scale
rows bordering the medial row are slightly larger than the medial
scales; all three median rows tend to form a continuous keel, that
of the median row not, or but slightly more elevated than the other
two rows.

The ventrals and subcaudals total 175 to 184, the lowest numbers
being those of the two females ( 175-178 ) ; the males vary between
182-184. The largest specimen measures: total length 279 mm.,
tail 59 mm. (R.C.T. No. 1364).

Color dark brown with a large series of narrow dark marks cross-
ing back usually continuous but sometimes broken on median line.
Occasionally dark marks on opposite sides alternate; black bands
rarely wider than a scale length, but they follow the scales and the
line tends to zigzag from scale to scale. Venter greenish with
quadrangular markings; occasionally (as in No. 1364) median series
of quadrangular marks distorted and each ventral with its posterior
edge dark. Subcaudals nearly black but with some traces of lighter
markings; top of head dark; labials with lighter cream areas more
or less pigmented.

Data on Xixia tessellata Cope

Pre-

Post-

Sub-

Supra-

Infra-

No.

Sex

oculavs

oculars

Temporals

Ventrals

caudals

labials

labials

1364

9

1-1

2-2

1 + 2

130

45

7-7

7-7

1445

9

0-0

2-2

1 + 2

134

44

7-7*

7-7

25739

6

0-0

2-2

1 + 2

125

57

7-7

7-7

1447

S

1-1

2-2

1 + 2

127

53

7-7

7-7

1448

S

2-1

2-2

1 + 3

129

55

7-7

7-7

•The Gth labial is unequally divided transversely; also a segmented part from internasal
present on one side.

Taylor: Review of Snakes of Costa Rica 55

I have examined the following specimens: Harvard M.C.Z. Nos.
19348-49, Salamanca, Costa Rica. They are typical in having 50 or
more dark spots or bands; supralabials and infralabials 7-7. The
three dorsal scale rows have keels somewhat more prominent than

the other rows.

Four specimens from Guapiles (Nos. 15276, 77, 80, 81) agree in
detail with the described specimen in most characters. The ventral-
subcaudal counts are respectively: 132-47, 129-51, 131-53, and

130-53.

Two specimens in the U. S. National Museum from Arriba, Costa
Rica ( Nos. 93835-36 ) have the ventral-subcaudal counts respectively
126-54 and 127-55. The supralabials are 7-7, the infralabials 6-6
in both.

The types (Nos. 32568-32569, U.S.N.M.) have the supralabials
and infralabials both 7-7. The ventral-subcaudal counts are respec-
tively 128-60 and 130-60.

Ninia psephota (Cope)

Colostoma psephotum Cope, Journ. Acad. Nat. Sei. Philadelphia, ser. 2, vol. 8, (1876) 1875,
p. 146 ("From higher points on the Pico Blanco, chiefly in the rainy zone at from 5000 to
7000 ft.").

Ninia psephota {part.) Dunn, Proc. Nat. Acad. Sci., vol. 21, 1935, p. 12.

This species was originally described in a genus, Catostoma, in
which Cope included two species of small snakes from Costa Rica
now regarded as belonging in the genus Geophis. However, it
seems without question to belong to the genus Ninia.

The type was collected by the William Gabb Expedition on the
higher parts of Pico Blanco, a high mountain in southern Costa
Rica, at an elevation of from 5000-7000 ft.

The characters of the species are: scales in 17 rows, all striate, and
all keeled except outer, those of the median row having stronger
keels than the others; tail triangular in cross section; forehead some-
what convex; internasals four-sided; frontal with convex anterior
border; prefrontal and large loreal entering orbit; nasal undivided;
postoculars two, no preocular; temporals 1 + 2 + 2; supralabials
six, third and fourth entering orbit, the fourth directly below eye,
the sixth longest; infralabials six, first pair in contact behind mental;
anterior scales of chin tuberculate in males; anterior chinshields
much longer than posterior; ventrals 162, anal single; subcaudals
73. Above uniform black; below black with half or less of an oc-
casional scutum red, forming a tessellated pattern; only a few spots
on the subcaudals. Total length 480 mm.; tail 128 mm.; to rictus oris
10 mm.; width of head (posterior) 8 mm.

56 The University Science Bulletin

Ninia oxynota (Werner)

Plate VI, figs. 2-3

Streptophortis oxynotus Werner, Mitt. Naturh. Mus. Hamburg, Jahrg. 26, 1909 (1910), (type
locality "Carriblanco" Costa Rica).

This species, described from a pair of specimens, has been
synonymized with Ninia psephota (Cope), but I regard this as a
very doubtful arrangement. The species psephota has a uniform
black coloration on back and sides, while the belly is black bearing
numerous red marks. In oxynotus, the body is banded with grayish
black or grayish blue, these bands separated by narrow brownish-
white transverse lines less than a scale wide; the bands narrow
somewhat low on the sides and a series of black spots alternate with
them; the belly is tessellated in checkerboard style in black and
greenish white.

That psephota and oxynota are related is suggested by the dor-
sal keel and the reduced number of labials.

Three specimens of this species are at hand, all taken at a point
about 5 kilometers southwest of Isla Bonita at an elevation of from
6000 to 6500 ft. M. N. H. No. 25741: rostral one third wider than
high, very narrowly visible above; internasals small, their length a
little less than half length of the prefrontals, and about one third
of their area; prefrontals about one fifth to one fourth longer than
wide, entering the orbit; frontal as wide as long, equal in length to
its distance from rostral, two and two-thirds times as wide as supra-
ocular; parietals as long as their distance from the rostral; scales
bordering parietals behind, enlarged; nasal divided; loreal large,
nearly rectangular in shape, broadly entering eye; no preoculars
(in M. N. H. No. 25740 a preocular is present on one side); two
postoculars, the upper very large, the lower small, about one sixth
the area of the upper; temporals 1 + 1+2 ( No. 25740, temporals
1+2); supralabials 6-6 (in No. 25740, 7-7), the third and fourth
enter the eye in both; infralabials six, the first four touch the first
chinshields, which are nearly double size of second pair; diameter
of eye less than distance from eye to nostril. The scales are dimly

PLATE VI

Fig. 1. Trimetopon pliolepis Cope, RCT No. 655, 5500 feet elevation; Isla
Bonita, Costa Rica ; total length, 242 mm.

Fig. 2. Ninia oxynota, RCT No. 641, approximately 6 miles west Isla Bonita,
August 3, 1947; total length, 395 mm. Ventral view.

Fig. 3. Same dorsal view.

PLATE VI

r^flP

I

58 The University Science Bulletin

striate but distinctly keeled. Scale formula 17-17-17; ventrals 161;
anal single; subcaudals, 70.

The color characters: General dorsal color brownish black with
narrow transverse lighter bands of brownish varying from 54-64 in
number. These indicated largely by lighter scale edges on one, or
one and a half rows. Ventral surface irregularly banded with
greenish cream and black. Sometimes a whole ventral scale light,
or black, but more often scales parti-colored, the black being in
angular spots; supra- and infralabials spotted black and greenish
cream as are the chinshields. Dorsal head scales with eight brown-
ish or cream markings.

This species has not been found outside the confines of Costa
Rica. There it seems to occur only in mountains.

There are several Costa Rican specimens in Harvard M. C. Z.
No. 15282 j , Navarro, has 157 ventrals and 76 subcaudals; No.
15283 $ , Navarro, 152 ventrals, 73 subcaudals; No. 15313 5 , "Costa
Rica," 164 ventrals, 75 subcaudals; No. 28073 $ , Estrella, 163 ven-
trals, 70 subcaudals. All of the specimens have six supralabials
and six infralabials.

The following table records the chief variation in the three speci-
mens collected:

Table of Data on Ninia oxynota

Pre-

Post-

Sub-

Bands

No.

Sex

oculars

oculars

Temporals

Ventrals

caudals

on body

641

9

0-0

2-2

1+2

161

64+ 1

56 (53)

25740

S

0-1

2-2

1+2

160

69+ 1

55

25741

$

0-0

2-2

1 + 1 + 2

161

69+ 1

64

Genus Dipsas Laurenti

Dipsas Laurenti, Specimen medicum exhibens synopsin reptilium . . . , 1708, p. 89.

Genotype: Dipsas indica Laurenti.

Six species occur in Costa Rica. One form described as Dipsas
ruthveni (Barbour and Dunn) from Aguacate Mts., Costa Rica, is
seemingly closely allied to Dipsas anthracops of Nicaragua and
they may possibly be the same species. However they are here
recognized as separate forms.

Key to Costa Rican Species of Dipsas

1. Scales in 13 rows on middle of body anthracops

Scales in 15 rows on middle of body 2

2. Ventral -subcaudal count 300 or more 3

Ventral -subcaudal count 290 or less 5

3. Four pairs of chinshields, the first separated from mental by first labials; 9 supra-
labials, fourth and fifth enter orbit; ventrals 215; subcaudals 135; yellow with

Taylor: Review of Snakes of Costa Rica 59

broad brown annuli, wider on anterior part of body articulata

Three pairs of chinshields ; less than 9 supralabials 4

4. Head longer; eight upper labials, fifth and sixth enter orbit; ventrals (?), sub-
caudals 121. Dorsal color (?), belly yellow with brown cross-bands on anterior
part of body, the posterior part with alternating brown spots; throat and lips

spotted brown pictiventris

Head shorter; seven labials, the fourth, fifth and sixth border orbit; ventrals
202; subcaudals 121; first chinshields touch mental; brown cross-bands ante-
riorly ; posteriorly with alternating spots argus

5. Scales 15-13; part of dorsal scales at least slightly enlarged; ventrals 1G5, sub-
caudals 79. Banded brown and whitish, the anterior bands three times width

of interspaces (25 body bands, 13 caudal) ruthveni

Scales 15-15; median dorsal scale row enlarged or not; chinshields touch mental. . . 6

6. Four pairs of chinshields, first small. Pale brown with dark brown rings; head
brown spotted, two larger spots on parietals; ventrals 164-169, subcaudals 113-

114 annulata

Three pairs of chinshields, the first largest. Greenish with a median series of
light spots ; ventrals 161, subcaudals 94 costariu nsis

Dipsas anthracops (Cope)

Leptognathus anthracops Cope, Proc. Acad. Nat. Sci. Philadelphia, 1868, pp. 108, 136 (type

locality, "Central America").
Tropidodipsas anthracops Boulenger, Catalogue of the Snakes in the British Museum, vol. 3,

1896, p. 297 (no specimen, Nicaragua).
Dipsas anthracops Dunn, Notulae Naturae no. 108, 1942, p. 7 ("Pacific side Costa Rica").

Twenty three yellow rings on body, twelve on tail separated by
9/x2 rows of scales anteriorly, and four rows posteriorly; yellow rings
wider behind; these may alternate on belly, which is otherwise un-
spotted; no white marks on head; temporal region and nape yellow.
Ventrals 177, subcaudals 76 ( may be partly undivided ) ; chinshields
normal with groove between; first two pairs wider than long, third
pair wider than long; temporals 1 -f- 2 -\- 3; no preocular; postocu-
lars two; seven supralabials, eight infralabials. Scales in 13 rows,
smooth; dorsals not enlarged, first labials in contact behind mentals.

This species has been reported on the Pacific side of Costa Rica
by Dunn, loc. cit.

Dipsas pictiventris (Cope)

Leptognathus pictiventris Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875),
p. 130, pi. 28, fig. 8 (type locality, E. Costa Rica); Giinther, Biologia Centrali-Americana ;
Reptilia and Batrachia. Oct. 1894, p. 142; Boulenger, Catalogue of the Snakes in the Brit-
ish Museum, vol. 3, 1896, p. 459 (thought to be a synonym of argus); Cope, Bull. U, S.
Nat. Mus. No. 32, 1887, p. 66.

S[ibynomorphus] pictiventris Barbour and Dunn, Proc. Biol. Soc. Washington, vol. 34, 1921,
p. 158 (Costa Rica).

Head longer and narrower than argus; scales in 15 rows, smooth,
the lateral scales a little smaller than dorsals, but median series not
enlarged; the elongate anterior chinshields touch mental, the first
infralabials not being in contact; internasals somewhat triangular,
frontal wide, nasal single; a presubocular below loreal; postoculars

60 The University Science Bulletin

two; supralabials eight, fifth and sixth enter orbit; temporals 1 + 2;
seven infralabials; three pairs of chinshields, first elongate, one half
longer than second pair; second and third pairs short and wide.

Belly yellow with brown cross bands on the anterior part of body;
the posterior part with large alternating brown spots. Back oc-
casionally crossed by spots. Throat and lips brown spotted. Ven-
trals ?, subcaudals 121.

The type specimen described by Cope is defective, and the all-
important number of ventrals cannot be determined; at least 161
are present but it is certain that this does not represent the total
number.

Boulenger has regarded argus and pictiventris as synonyms. An
examination of the types leaves me with the opinion that they are
distinct. The very bad condition of the type of pictiventris makes
it impossible to ascertain all pertinent characters.

Dipsas argus (Cope)

Leptognathus argus Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875),
p. 130, pi. 27, fig. 1, pi. 28, fig. 7 (type locality. Sipurio, Costa Rica); Boulenger, Cata-
logue of the Snakes in the British Museum (Natural History), vol. 3, 1896, pp. 458-459.

S[ibynomorphus\ argus Barbour and Dunn, Proc. Biol. Soc. Washington, vol. 34, Dec. 21,
1921, p. 158 (Costa Rica).

This is an extremely slender snake, the body compressed; head
wide, the snout short; rostral triangular, as high as wide, very small,
scarcely visible from above; internasals small, the prefrontals large
bordering orbit; frontal with parallel sides, equal to length of an-
terior border; supralabials seven, the fourth, fifth and sixth border
the orbit; 2 postoculars; temporals 1-2; infralabials seven, the first
pair not in contact behind the mental, four touching first chinshields;
three pairs of chinshields, the two anterior pairs elongate, longer
than wide, the third pair quadrate, smaller; scales in 15 rows,
smooth, larger above than on sides, the median row not abruptly
larger than those adjoining.

Ventrals 202,* anal single, subcaudals 121.

Color above, from the third row of scales, greenish ash, with two
series of alternating light ocelli with black borders; lower on side
the third scale row yellow with a series of black-edged ocelli like
those of the back; on venter, blackish; speckled on the posterior half
of the length. A large black-bordered ocellus on the nape; head
vermiculated with black; lips yellow with black specks.

* I have counted only 202 ventrals in the type; 212 as given in the type description.

Taylor: Review of Snakes of Costa Rica 61

Dipsas articulata Cope

Vipsas brevis (not of Dumeril, Bibron and Dumeril) Cope, Proc. Acad. Nat. Sci. Philadelphia,
1860, p. 266 (Cbcuyas de Veraguas, New Granada).

Leptognathus articulata Cope, Proc. Acad. Nat. Sci. Philadelphia, 1868, pp. 107, 135 (Ver-
aguas, Costa Rica [this may be in error]).

S[ibynomorphus] articulata Barbour and Dunn, Proc. Biol. Soc. Washington, vol. 34, Dec. 21,
1921, p. 158 (Costa Rica).

Body slender, strongly compressed, vertebral scales moderately
enlarged. Loreal and prefrontal border eye; no preocnlar; two
postoculars; temporals 2 + 3; nine snpralabials, the fourth and fifth
enter orbit; first pair of infralabials in contact behind mental; four
pairs of chinshields. Scales in 15 rows; ventrals 215; anal single;
subcaudals 135. Yellow with broad brown annuli surrounding body,
wider on anterior part of body than on posterior, and wider than
the interspaces of yellow; top and sides of head, supralabials in
front of eye, and all infralabials brown; some short, white or yellow
lines on head. Total length about 675 mm.; tail 225 mm.

I have not examined a Costa Rican specimen. R is retained in
the list because of the report of Barbour and Dunn, op. cit.

Dipsas annulata (Giinther)

Leptognathus annulatus Giinther, Ann. Mag. Nat. Hist., ser. 4, vol. 9, 1872, p. 30 (type lo-
cality, Cartago, Costa Rica); Biologia Centrali-Airiericana ; Reptiles and Batrachians, 1895,
p. 141, pi. 49, fig. C.

Leptognathus annulata Boulenger. Catalogue of the Snakes in the British Museum, vol. 3,
1890, p. 4 ."> 7 .

SUbynomorphus] annulata, Barbour and Dunn, Proc. Biol. Soc. Washington, vol. 34, Dec. 21,
1921, p. 158 (Costa Rica).

A female specimen (No. 25702) was captured at Isla Bonita
(American Cinchona Plantation) by Sr. Antonio Machado, Aug. 5,
1947, and presented to the collection.

Body distinctly compressed with broad thick head and large, pro-
tuberant eyes. Rostral not or scarcely visible from above, as high
as wide; internasals subtriangular, about one third the length of
prefrontals; latter very large, much wider than long, bending down
on side and forming a part of orbit boundary; frontal much longer
than wide, pentagonal, and about one sixth longer than its distance
from tip of snout; parietals large (3.5x5 mm.), shorter than their
distance from snout tip; supraoculars rather narrow, more or less
angular behind, bending down somewhat behind eye; nasal single,
pierced by the nostril, from which a small suture emerges termi-
nating on the upper border; loreal, which replaces the preocular,
longer than high; 2 postoculars; temporals 1+2+2; supralabials
7-7, increasing in size to sixth, the fourth and fifth entering orbit;
infralabials 7-9; a small pair of scales inserted between the mental

62 The University Science Bulletin

and first very large pair of chinshields, each in contact with two
labials; chinshields touch 4-5 infralabials, the last being very large;
second pair of chinshields in contact, little more than half size of
first pair; a third pair of chinshields in contact, somewhat larger
than second pair, precedes first ventral; ventrals 169; anal single;
subcaudals 113. Scale formula, 15-15-15, the scales all smooth ap-
parently without trace of pits. The median scale row much en-
larged.

Total length, 501 mm.; snout to vent, 330 mm.; tail, 171 mm.;
width of head, 8.6 mm.; head length, 12.6 mm.

Banded red brown and raspberry pink, the brown bands becom-
ing darker low on sides, and black below, the pink bands becoming
white below; 27 red-brown bands on body, 18 on tail; these are two
or three scales wide on median line, being a little broader on tail;
on venter bands narrow and usually the width of two ventrals,
while the white may occupy parts of three or four ventrals. Head
brownish with an indefinite lighter pattern; the temporal areas
being, on the whole, lighter than remainder. A distinct cream spot
on fourth supralabial and a cream spot covering parts of fifth and
sixth; infralabials and chin cream, with numerous brownish blotches.

Compared with the type description, the differences are indeed
small. The type has a small subocular below the loreal that is
wanting in this specimen. The type has 164 ventrals, and 113 sub-
caudals while this specimen has 169 ventrals and the same number
of subcaudals. The specimen is somewhat larger, 501 mm. total
length, compared to 440 mm., the length of the type.

Dipsas ruthveni (Barbour and Dunn)

Sibynomorphus ruthveni Barbour and Dunn, Proc. Biol. Soc, Washington, vol. 34, 1921, p. 158
(type locality, Aguacate Mts., Costa Rica); Werner, Zool. Jahrb., 1929, p. 189 (no speci-
mens).

This form is close to Dipsas anthracops and it may be necessary
to reduce it to the synonymy of that species.

Body strongly compressed anteriorly, less compressed along mid-
body region; tail diminishing suddenly in diameter behind vent;
strongly compressed posteriorly; rostral pentagonal scarcely visible
above; prefrontal large, broadly entering orbit; frontal longer than
wide, about equal to its distance from tip of snout; nasal partially
divided; loreal twice as long as high, entering orbit; no preocular;
two postoculars; temporals 14-2; seven or eight supralabials, fourth
and fifth enter orbit; first lower labials in contact behind mental; a
pair of elongate chinshields; scales in 15 or 13 rows without apical

Taylor: Review of Snakes of Costa Rica

63

pits; the scale rows bordering the median row may be slightly en-
larged.* Ventrals 165, subcaudals 79.

Banded brown and whitish; the brown bands anteriorly thrice as
wide as the light bands, posteriorly nearly equal and of solid color;
white bands clouded by groups of small darker streaks and flecks;
bands diagonal, tending to alternate on belly; 25 bands on head and
body; 13 on tail. Total length 425 mm.; tail 110 mm.

Dipsas costaricensis sp. nov.

Plate VII, text fig. 2

Type.-Uus. Nat. Hist. Univ. Kansas No. 25703, collected five
miles southwest of Turrialba, on the Morehead finca, July 16, 1947;
E. H. Taylor, collector.

Diagnosis. — Body moderately compressed, not noticeably slender;
first labials not in contact behind mental; no preocular, median
scale row not sensibly enlarged; ventrals 161, subcaudals 94.
Lichen-green with a median series of light spots and with lateral
and ventral ocellilike spots.

Fig. 2. Dipsas costaricensis sp. nov. Type. MNH No.25703. More-
head Finca, Wi miles SW Turrialba, Costa Rica. A. Head dorsal view;
B. Head lateral view; C. Head ventral view.

Description of the type.— Body more or less compressed, triangu-
lar in cross section, the head not much enlarged; eyes small; rostral
small, not or scarcely visible from above, slightly broader than high;
internasals small, a little less than one third length of the prefron-
tals, and about one fifth of their area; latter very large, a little

* The original description states "scales in "15 rows without apical pits, median row tint
sensibly enlarged or 13 rows with rows on each side of the median row sliehtly enlarged."
This is not clear but since there was but a single specimen, both conditions must obtain in
the same specimen.

64

The University Science Bulletin

If "%:

■ ■ .

w*

Plate VII. Dipsas costaricensis sp. nov., MXH No. 25703, More-
head Finca, 5 miles SW Turrialba, Costa Rica. Total length, 47 mm.

Taylor: Review of Snakes of Costa Rica 65

broader than long, entering orbit for a distance equal to that of the
loreal; frontal about one sixth longer than its distance from tip of
snout, about one third longer than wide; parietals large (3.24 x 5
mm. ) shorter than their distance from tip of snout; supraocular
curving slightly, widest but not angular, posteriorly; nasal partially
divided by a suture from nostril to upper edge of scale; loreal
nearly twice as long as high, entering orbit; no preocular; 2-1 post-
oculars, the upper very large (the two fused on left side); tem-
porals, 1+2+2 (2+2+2); supralabials, 7-6, the fourth, fifth and
sixth (fourth and fifth) entering orbit; infralabials 6-6, four touch-
ing first pair of chinshields which are in contact with mental; sec-
ond pair of chinshields in contact with each other, smaller than first
pair; third pair in contact, irregularly shaped.

Scales smooth, without apical pits, the median dorsal row not en-
larged; scale formula: 15-15-15; ventrals 161; subcaudals 94; anal
single.

Color in life.— Lichen-green, the color interrupted on the scale
edges by a variegated brown or black; a dorsal series of 33 brownish
yellow blotches, variegated with dark brown and dusky markings
on body; on tail there are 15-16 but these are rather indefinite; low on
the sides and encroaching on the ventrals is a series of cream-white,
dark edged, ocellilike markings; chin whitish with a faint yellow
wash; neck and anterior fourth of body nearly pure white with
small greenish or black flecks forming an irregular, indistinct,
median line; there is more yellow color on latter three fourths of
body; there are similar dark markings becoming more dense under
tail with some greenish and yellow flecks. On dorsal surface the
head is nearly uniform black-brown with some indistinct reddish
brown marks; a divided white blotch below eye; chin with a distinct
pair of black spots on the fourth labials and a few other less distinct
flecks of greenish black.

Measurements.— Total length, 477 mm.; tail, 147 mm.; width of
head, 9 mm.; length of head, 13.5 mm.; diameter of body, 10 mm.

Remarl s— This form differs from argus in having a much smaller
ventral-subcaudal count (255 compared to 323 in argus); from pic-
tiventris in a much lower number of subcaudals (94 compared to
121 in pictiventris) and a thicker body; from articulata in having
mental touching chinshields and much lower ventral-subcaudal
count (255 compared to 350 in articulata); from annulata in having
the mental separated from the first chinshields, smaller ventral-

5—3286

66 The University Science Bulletin

subcaudal count (255 compared to 277), and the median dorsal
scale row not enlarged; from ruthveni by having the scale rows con-
stantly 15, not reducing to 13, and a different color pattern.

Genus Neoparias Giinther

Neoparias Giinther, Biologia Centrali-Aroericana ; Reptilia and Batraehia. July. 1895, p. 178.

Genotype: Neoparias bicolor Giinther.

A single Costa Rican species is referred to this genus.

This genus has quite incorrectly been associated with Leptogna-
thus (Sibynomorphus or Dipsas); the striking character of the
chinshields in Neoparias readily separate the two genera.

Neoparias bicolor Giinther

Neoparias bicolor Giinther, Biologia Centrali-Americana : Reptilia and Batraehia, July 1895,
pp. 178-179, pi. 56, fig. C (type locality, Chontales Mines, Nicaragua).

Leptoynathus bicolor Boulenger, Catalogue of the Snakes of the British Museum, vol. 3, 1896,
p. 460.

S[ibynomorphus] bicolor Barbour and Dunn. Proc. Biol. Soc. "Washington, vol. 34, 1921, p. 158
(Costa Rica).

Body slender, strongly compressed; tail slender; scales of the
vertebral series not enlarged; scales on chin transverse, lacking a
median groove; rostral small, scarcely visible above; internasals less
than half size of prefrontals; frontal short, wide, about equal to its
distance to rostral, only a little shorter than parietals; nasal appar-
ently completely divided; a large square loreal entering eye, and a
small preocular above it not reaching frontal; three or four post-
oculars; temporals 1 + 2 + 3 + 3; supralabials 10-11, the fourth to
seventh, or the third to sixth enter orbit; infralabials 11, the first pair
fused together behind the small mental; three median azygos scales
on chin, the first touching five scales; scales smooth, in 15 rows;
ventrals 195; subcaudals 129.

Deep black with white rings completely encircling the body, the
first around the neck, 14 around the trunk, 12 around the tail; the
white rings half as long as the black interspaces; length about 516
mm.; tail about 192 mm. (from type description).

The species is arboreal, and perhaps because of this habit is very
rare in collections.

Genus Tretanorhinus Dumeril, Bibron and Dumeril

Tretanorhinus Dumeril, Bibron and Dumeril, Erpetologie Generate ou Histoire Naturelle Com-
plete des Reptiles, vol. 7. pt. 1, 1854, pp. 348-349.

Genotype: Tretanorhinus variabilis Dumeril, Bibron and Dumeril.
Only a single species of this genus is believed to occur in Costa
Rica.

Taylor: Review of Snakes of Costa Rica 67

Tretanorhinus nigroluteus nigroluU us Cope

Tretanorhinus nigroluteus Cope, Proc. Acad. Nat. SO. Philadelphia, 1SC1. p. 2<is (type locality,

Aspinwall, Panama [fide Dunnl ).
Tretanorhinus nigroluteus nigroluteus Dunn, Copeia, 1939, on. 4. p. 216.

The presence of this species in Nicaragua and Panama strongly
suggests its presence likewise in Costa Rica, although I know of no
Costa Rican specimens that have been reported.

The species has the following characteristics: blackish or olive
brown on dorsum, usually with small alternating dots on each side,
the dots variable in size, usually covering about one scale; two lower
scale rows lighter in color tending to form a stripe anteriorly; venter
light yellowish or whitish, sometimes with dots or a suggestion of a
midventral darker line; a dark streak on sides of head.

Nasals in contact behind rostral or narrowly separated; internasals
distinct, small, longer than wide; frontal as long as its distance from
end of snout, shorter than parietals; eight supralabials, the fourth
entering the orbit; normally two preoculars, two postoculars and
usually two loreals; ventrals of southern specimens 133-149; the
subcaudals 56-80; scales in 21 rows.

This form is aquatic and is said to frequent salt water, mangrove
swamps.

Genus Sibon Fitzinger

Sibon Fitzinger, Neue Classification der Reptilien, 1826, p. 31,

Genotype: Coluber nebulatus Linnaeus.
A single species is known.

Sibon nebulatus Linnaeus

Coluber nebulatus Linnaeus, Systerna naturae, ed 10, vol. 1, 1758, p. 222 (type locality, "Af-
rica [presumably in error] ).

Coluber sibo7i Linnaeus, Systerna naturae, ed. 10, vol. 1, 1758, p. 222.

Sibon nebulatus Taylor, Univ. Kansas Sci. Bull., vol. 26, 1939 (1940). pp. 473-474.

TSibon annulata Dunn and Bailey, Bull. Mus. Ccmp. Zool., Harvard Coll., vol. 86, 1939, p. 9
(Costa Rica).

Leptognathus nebulata Cope, Journ. Acad. Xat. Sci. Philadelphia, 1876 (1875) p. 131 (south-
eastern Costa Rica).

This very widespread species is not common in collections and is
relatively little known. The following characters will permit a
diagnosis: maxillary short, with 15-16 short teeth decreasing in size
posteriorly; rostral visible above as a narrow line, not reaching level
of dorsal surface of head; internasals and prefrontals normal, the
latter entering orbit; frontal pentagonal, as long as (or a little longer
than) its distance from tip of snout; parietals a little longer than
wide, their length less than, or approximately equal to their dis-

68 The University Science Bulletin

tance from prefrontals; nasal completely divided; loreal large, rec-
tangular, entering orbit; no preocular; postoculars two or three;
temporals 1+2; supralabials seven, the fourth and fifth or fourth,
fifth and sixth entering orbit; five or six infralabials touch anterior
chinshields, which are longer than second pair; a third pair of chin-
shields may be present, the scales of which are wider than long.

Scale rows 15-15-15, or the preanal count may reduce to 13; scales
smooth without pits. Ventrals 170-197; subcaudals 73-102; anal
single; probably not exceeding a meter in length; known length,
790 mm.; head large and distinct from neck; eye large, the pupil
vertically elliptic, the body somewhat compressed.

Gray or whitish above marked with numerous irregular transverse
black or brown markings which may extend across body or may be
broken into two or three parts, the edges of which are very irregular;
some scales bordering spots may be pure white; a nuchal dark spot
bordered by white dots; head marked with dark spots; a dark line
from eye to jaw; upper labials whitish; chin and lower surfaces white
with a series of more or less alternating quadrangular dark spots on
the sides of the venter with small flecks scattered between.

Two specimens in the U. S. National Museum numbered 30621,
30622 are a part of the early collection made by William Gabb.
They agree in most characters with the given description. No. 3C621
has 185 ventrals and 99 subcaudals, the anal single. There are ap-
proximately 40 bands. The head is brown with numerous white
flecks, which tend to form a symmetrical pattern. No. 30622 has
171 ventrals and 85 subcaudals. There are 41 bands. The maxillary
teeth are 14, slender, curved, none grooved. The posterior part of
the maxillary forms an elevated ridge.

It is presumed that the two listed specimens were taken in south-
eastern Costa Rica, where much of the William Gabb Collection
was made.

The species has been reported also from Guapiles in the eastern
lowlands.

Genus Xenodon Boie

Xenodon Boie, Isis, 1827, p. 540.

Genotype: Xenodon inornatus Boie (fide Fitzinger).
Two forms are known to occur in Costa Rica.

Key to Costa Rican Species of Xenodon

Body marked with large paired lateral spots, discrete anteriorly, often touching

posteriorly bertholdi

Body marked with large irregular saddlelike blotches colubrinus

Taylor: Review of Snakes of Costa Rica 69

Xenodon bertholdi Jan

Xcnodon Bertholdi Jan, Arch, per la Zool., 1863, pp. 318-319 (same in separate, pp. 108,
1(19) (type locality said to be "Mexico." Very doubtful).

The type description of this species gives the following charac-
ters: one preocnlar, three postocnlars; temporals 1+2; supralabials
eight; ten infralabials, six touching the chinshields; 19 scale rows
and 13 rows behind anus; ventrals 153, anal single; snbcandals 42.
On each side of the trunk is a series of large oval spots separated
from each other mesially.

The color pattern of a snake captured at Turrialba (K. U. No.
25167) closely approaches the color characters of the type of X.
bertholdi. In this specimen the spots are outlined in black, and
bordered by light ash-gray; six of the fourteen spots are separated
on the middorsal line, the others being narrowly in contact.

In this specimen a pair of gray lines, spotted and flecked with
black, passes back from the eye to a point on the neck across jaw
angle. The area between these lines is rather brownish fawn. A
character that may be of significance is that the supraoculars are
not truncate on their posterior border, but extend a noticeable dis-
tance behind the posterior end of the frontal. A pair of black spots
are present on the prefrontals. The labials are brownish fawn and
the chin and neck as far back as fifth ventral are white. The anal
scale is white. The venter is covered with flecks of pigment, scat-
tered irregularly. The subcaudal area has a peppering of black at
the base, but is nearly cream-white near the tip.

A specimen (M. C. Z. No. 19238) from Limon, Costa Rica has a
paired series of large spots on the anterior half of body, while be-
tween them are small dark medial spots. Paired spots on posterior
part of body fused mesially. Ventrals 145; subcaudals 44; scales,
19-19-17 with single apical pits on dorsal scales; supralabials 8-8;
infralabials 10-11, five (or six) touching first chinshields which are
twice size of second pair.

A specimen in the U. S. National Museum, No. 37482 (or 3) from
Esparta, Costa Rica, has 150J2 ventrals and 46+ subcaudals.

If the type of this species actually is Mexican, it bespeaks a wide
range for the form.

Xenodon colubrinus Giinther

Xenodon colubrinus Giinther, Catalogue of Colubrine Snakes in the Collection of the British
Museum, 1858, pp. 55-,r)6 (type locality, Para, Brazil).

The type of Xenodon colubrinus has the following scale charac-
ters: scales smooth in 19 rows; one preocular, two postoculars; two
nasals, the nostril very large; anal single; 151 ventrals, 45 subcaudals;

70 The University Science Bulletin

14 lozenge-shaped, pale-edged blotches on the trunk, four on tail.
Head elongate, snout protruding, angular in front. Rostral just vis-
ible above; frontal about as long as wide; parietals short; loreal
large, nearly square; eight supralabials, the fourth and fifth touch
eye. Head dusky; a brown marbled streak from the back edge of
eye to mouth angle; belly yellowish brown marbled.

The following additional characters obtain: rostral one and one-
third times as wide as high; frontal as long as wide, the length reach-
ing to middle of internasals; parietals a little wider than long, their
length equal to that of frontal; supraoculars truncate behind, not
extending back beyond level of frontal; first chinshields one and
one-half times size of second pair which are nearly separated bv the
scales following. The scales on sides form diagonal series.

The two Costa Rican specimens acquired by gift from Mr. Virgil
Cave agree in general with the characteristics of the type. It is dif-
ficult to state the variational limits of this species since no recent
revisional work has been done on the genus, and the name as now
understood is probably composite.

Table of Data for Xenodon colubrinus Ginther

No. Sex

Ventrals

Sub- i
caudals

Supra -
labials

Infra -

labials

Laliials
enter
nl -bit

Labials

contact

chinshields

851 9
1001 $

151

144

45

9-9

8-8

11-11
11-11

/4th 5th

\5,h6,h
4,h5,h

5-5
5-5

Table of D

ATA FOR

Xenodon

colubrinus Ginther ( concluded )

Scale
No. formula

Anal

Preocular

Postocular

Temporals

Bands
on body

Teeth

851 17-19-17
1001 17-19-17

1
1

1-1
1-1

2-
3-

1
-1

1 + 2

1 + 2

13
13

14 + 2

13 + 2

Genus Enulius Cope

Enulius Cope, Proc. Amer. Philos. Soc, vol. 11, 1871, p. 559.

Genotype: Enulius murinus Cope = Enulius flauitorques Cope.
One species is known in Costa Rica.

Enulius flavitorques Cope

Liophis flavitorques Cope, Proc. Acad. Nat. Sci. Philadelphia, 1868 (1869), p. 307 (type lo-
cality, Magdalina River, Colombia).

Drepanodou ? flavitorques Boulenger, Catalogue of the Snakes in the British .Museum, vol. 3,
1896, p. 639 (no specimens).

Enulius flavitorques Dunn. Proc. Acad. Nat. Sci. Philadelphia, vol. 89, 1937 (1938), pp. 415-
417 (Costa Rican records Marivalles, Barranca and Cartago).

I have seen no specimen of this form from Costa Rica. The sali-
ent characters are as follows:

Taylor: Review of Snakes of Costa Rica 71

Rostral projecting and well visible above; frontal very broad, with
a long-produced posterior angle; internasals about as long as wide;
prefrontals considerably wider than long; nasals higher than long,
narrower than loreal; preocular higher than long; loreal higher than
long, the suture straight above, angulate below; postoculars two;
temporals 1 + 2; snpralabials seven, all higher than long except last;
third and fourth enter orbit; infralabials seven, fourth elongate, first
pair in contact behind the mental; posterior chinshields half as long
as anterior, separated by a scale; scales in 17 rows. Ventrals 188;
anal divided; subcaudals 105. Total length about 420 mm.; tail
about 132 mm. Uniform dark brown above with a broad yellow
half-collar crossing the posterior part of the parietal shields and two
transverse rows of scales; venter dirty yellow.

There is some doubt that the form Emilias minimis Bocourt
[ — Enulius longicaudata (Cope)] is a synonym of this species.
It is not impossible that the latter is the proper name for the Costa
Rican species, but the matter cannot be decided now.

This species is reported as having single apical pits on the scales.
It may readily be separated from Enulius sclateri, which has four
(two pairs) apical pits on the dorsal scales.

I have examined casually a series of specimens of this species from
Honduras and Panama. There is considerable difference in the
head marking and nuchal band. Its presence in Costa Rica is at-
tested by Dunn's records, loc. cit., from Marivalles, Barranca and
Cartago.

Enulius sclateri (Boulenger)

Plate VIII, text fig. 3

Leptocalamus sclateri Boulenger, Catalogue of the Snakes in the British Museum, ed. 2, vol. 2,
1891, p. 251, pi. xii. fig. 1 (type locality, "South America"); vol. 3, 1890, p. 641 (Guasima,
Costa Rica).

Enulius slateri (sic) Dunn, Proc. Acad. Nat. Sci. Philadelphia, vol. 89, 1937 (1938), p. 417;
Smith and Taylor, U. S. Nat. Mus. Bull. no. 187, 1945, p. 61.

This species is known in literature from the type description, and
from some brief notes given by Dunn, loc. cit. The specimen here
described was captured at Los Diamantes (near Guapiles, Costa
Rica) by Richard C. Taylor. It was discovered by chopping to
pieces a rotting log in which the snake was concealed. The follow-
ing characters obtain:

Rostral at least twice as broad as high, its border along internasals
almost straight rather than angular, the length of part visible above
a little more than half length of internasals; suture between inter-
nasals equal or larger than that between prefrontals, but the scales
much shorter than prefrontals which enter eye; frontal much longer

72

The University Science Bulletin

than broad, four-sided, its length one and one-half times greater
than its distance from tip of snout; parietals elongate, a little longer
than the frontal; supraocular reduced, but extending above more
than half of the eye; nasal divided, anterior part the larger, con-
taining most of nostril; loreal elongate forming part of border of
eye; two postoculars, the upper somewhat the larger; temporals,
1 + 2, the anterior rectangular, touching both postoculars; seven
supralabials in following order of size: 1, 2, 5, 4, 3, 7, 6, ( 1, 2, 4, 5,
3, 7, 6 ) , third and fourth entering orbit; six lower labials in following
order of size: 2, 1, 5, 4, 3, 6, first labials in contact behind mental;
first chinshields in contact anteriorly, separated posteriorly by a
median diamond-shaped scale; second pair of chinshields widely
separated; two scale rows separate first ventrals from labials.

A B C '

Fig. 3. Enulius sclateri. R.C.T. No. 1368. Collected, Los Diamantes,
near Guapiles, Costa Rica. A, Head, dorsal view; B. Head, lateral view;
C, Head, ventral view. X 4.

Scale formula, 19-15-15-15; scales smooth, the anterior 20 trans-
verse scale rows each with four scale pits, two anterior, two pos-
terior, the latter appearing slightly the smaller; no pits apparent on
other scales; ventrals 145; anal divided; caudals 96 + 1.

Above blue-black, the scales on sides showing slightly lighter
edges- chin, throat and venter as far as anal opening, white, the lat-
eral color not extending onto the edges of ventrals; usually a whitish
area on the scales of the outer row; caudals with a median dark line
beginning behind vent; the color of outer scale rows encroaches
somewhat on edges of subcaudals so that they appear grayish with
some dim lighter areas. The head is pure white with a dark ring

Taylor: Review of Snakes of Costa Rica

73

surrounding the eye and a black spot covering first labial, rostral,
internasals, nasals, and a small triangular area on the prefrontals.

The measurements (in mm.) are as follows: width of head, 6;
length of head, 10.2; total length, 345; tail, 135.

The specimen is a male and it agrees with the type specimen in
most characters. The known ventral range is 132-151; subcau-
dals 97-98.

Plate VIII. Enulius sclateri (Boulenger), RCT No. 1368. Los Diamantes,
Costa Rica, September 6, 1949; total length, 345 mm.

74 The University Science Bulletin

I have examined two Harvard specimens, M. C. Z. Nos. 28079-80,
from Peralta, Costa Rica. These agree with the described speci-
mens." Specimens are known from Guasimo and Finca Hamburg.

Genus Helicops Wagler

Helicops Wagler, Naturliches System der Amphibien, 1830, p. 170 (part.)

Genotype: Helicops angulatus Linnaeus.

A single species has been reported from Costa Rica.

Helicops wettsteini Amaral

Helicops wettsteini Amaral, Bull. Antiv. Inst. Anier., vol. 3, no. 2, p. 40 (San Juan de Vinas
[1000 m. alt.] base of Volcan Turrialba, central Costa Rica).

This species may be readily recognized by the contact of the nasal
scales behind rostral.

Eye small; rostral wider than high, visible from above; nasals in
contact behind rostral; internasal single; prefrontals separate; frontal
once and two-thirds times as wide as long, as long as its distance
from the end of snout, shorter than the parietals; loreal higher than
long; one preocular, two postoculars; temporals 1 + 2; supralabials
eight (8-9 in paratype), the fourth and fifth border orbit; anterior
chinshields longer than posterior, touching five or six lower labials;
scales in 19 rows, smooth; ventrals 128; anal divided, subcaudals
37-38. Total length, 375 mm., tail 50 mm.

The color is uniform dark plumbeous above; light underneath
with a median interrupted black streak and a blackish line along
outside border of ventrals.

The species is known from the two types, collected at Juan Vinas
(San Juan de Vinas, 1000 m. elev. ).

Genus Hydromorphus Peters

Hydromorphus Peters, Monatsb. Akad. Wiss. Berlin, 185!». p. 270.

Genotype: Hydromorphus concolor Peters.
A single species is known from Costa Rica.

Hydromorphus concolor Peters

Hydromorphus concolor Peters, Monatbs. Akad. Wiss. Berlin, 1859, pp. 27ii-277 (type local-
ity, Costa Rica).

This medium-large snake still remains rare in collections. The
uniform color, smooth scales, small eye, single prefrontal, and the
nostril directed upward piercing a single scale, readily identify the

* A specimen, Harvard M. C Z. No. 34382 (Canal Zone, Atlantic Side), differs in having the
rostral, prefrontals, internasal, anterior part of frontal, and area about eye, black except that
the nostrils are whitish; second, third and fourth labials white except about eye; two small
black areas on lower lips; the neck ring includes the back of head and four transverse scale

Taylor: Review oe Snakes of Costa Rica 75

species. The following characters are those of the type.

Head depressed, with broadly rounded snout; nasals separated
by a pair of very small internasals; prefrontals much broader than
long; frontal slightly longer than broad, as long as its distance from
the rostral, much shorter than parietals; loreal entering orbit, twice
as long as high; one small preocular; two postoeulars; temporals
1 + 2; six supralabials, third entering orbit; four labials touch first
pair of chinshields; posterior chinshields reduced; scales in 17 rows,
smooth; ventrals 175; anal divided; subcaudals 31.

Dark grayish brown on dorsal surfaces, the sides and venter paler,
the scales being partly yellowish. Total length 850 mm.; tail 86 mm.

A specimen in the U. S. National Museum (No. 13537) from
San Jose, Costa Rica has the rostral much broader than high, one
half or more of its surface visible above; internasals fused but with
a small entrant suture; prefrontals fused into one scale, 2/2 times
wider than long; frontal small, as wide as long, shorter than its
distance from tip of snout; parietals equal to their distance from
tip of snout; nasal single, pierced in the anterior half of scale; the
loreal and preocular both enter orbit; two postoeulars; supraoculars
small; temporals 1 -4- 2 + 2 ( a small scale severed from the front
temporal on one side); supralabials 6-6, the third only entering
orbit; infralabials 9-8, four touching chinshields; two pairs of chin-
shields, the first largest, the second pair separated by two scales;
scale formula 19-17-15; ventrals 174/2; subcaudals 40; anal divided.

The specimen is uniform dark brown, the scale centers lighter
on sides of body.

Genus Trimetopon Cope

Trimetopon Cope, Proc. Amer. Phil. Soc, vol. 22, 1884, p. 177.

Genotype: Ablabes gracilis Giinther.

This genus as here considered may be composite. Four forms are
known to occur in Costa Rica. The forms of Trimetopon (sensu
strictu), having 15 scale rows and a fused prefrontal are kept with
forms having the prefrontals separated and 17 scale rows.

Key to Costa Rican Species of Trimetopon

1. Prefrontals fused ; scales in 15 rows 3

Prefrontals fused or separate ; scales in 17 rows 2

2. Prefrontals separate; seven supralabials; throat white, venter red; a black streak

from eye To suture of fourth and fifth labials; ventrals 163; subcaudals 33 viquezi

Prefrontals fused; eight supralabials; ventrals 154 ^ ; subcaudals 09; throat and
venter yellow; no apical pits; first chinshields twice size of second pair pliolepis

3. Ventrals 122, subcaudals 09; one preocular, two postoeulars; seven supralabials;
black above, and white below on venter; head white as far forward as middle of

frontal including third labial ; a black spot on seventh labial simile

Ventrals 141-149; subcaudals 60-69 (05); a tendency to form lateral dark stripes;

a neck band; single apical pits; chinshields subequal in size; yellowish below. ..gracile

76 The University Science Bulletin

Trimetopon pliolepis Cope

Trimetopon pliolepis Cope, Proc. Acad. Nat. Sci. Philadelphia, 1894, p. 201; Boulenger, Cat-
alogue of the Snakes in the British Museum, vol. 3, p. 636.

Trimetopon gracile (part.) Dunn. Copeia, 1937, no. 4, pp. 214-215 (The part referable is the
Trimetopon plioplcpsis [sic] stated by Dunn to have appeared in Cope, Proc. Amer. Philos.
Soc. 22, 1885:17". This is incorrect since the species was not described until 1894, and in
another publication. See above).

Rostral not visible above, small internasals much wider than long;
anterior border of frontal slightly convex forward; lateral border of
frontal shorter than the parietal border, "which forms less than a
right angle with that of the opposite side"; loreal longer than high;
one preocular well separated from frontal; one postocular; temporals
1 + 1; supralabials 8, fourth and fifth entering orbit, all longer than
high except eighth; infralabials 8; posterior chinshields half size of
anterior pair; scales 17, without pits ( fossae ) .

Total length 287 mm.; tail 76 mm.; ventrals 154; anal divided;
subcaudals 69.

Dark brown above, the scales with a paler, minutely speckled
center except those of third row; this, together with the more re-
stricted pale centers of the scales of the first and second rows gives
the appearance of a dark lateral band which tips the ventrals.
Below uniform yellow; a narrow yellow collar borders but does not
cross the extremities of the parietals; each upper labial with a large
yellow spot next the border; that at the top of the sixth and the
front of the seventh has the effect of a postocular band. ( From
type description. )

Cope does not specifically state that the prefrontals are fused but
one infers this since he does not mention any difference in this re-
spect from gracile when he compares the forms.

Two specimens were collected in Costa Rica. R.T.C. No. 655
was taken at Isla Bonita at approximately 5500 ft. elevation and
R.C.T. No. 1458 was taken at Los Diamantes at about 900 ft. eleva-
tion by Richard C. Taylor. The difference in elevation is nearly
4600 feet. Both are referred to this species despite certain discrep-
ancies in characters.

No. 1458 2 • The following scale characters obtain: part of ros-
tral visible above, a broad triangle, much smaller than an internasal;
internasals small, the common suture one-fourth times as long as
the fused prefrontals; latter scale not twice as wide as long, minutely
entering eye, its posterior edge nearly a straight transverse line; pen-
tagonal, its width four-fifths of the length, its length one-fourth
greater than its distance from the snout tip; parietals very large,

Taylor: Review of Snakes of Costa Rica 77

the length one-fifth to one-fourth greater than its distance from snout
tip; nasal at least partially separated; loreal present, longer than
high; a small preocular; supraocular about one third of the width
of the frontal; one postocular; temporals 1 + 1 + 2; supralabials
eight, the fourth and fifth border eye; infralabials 7-8, there being
four (or five) touching the anterior chinshields; latter scales twice
as long as posterior and more than twice their area; scale formula,
17-17-17, all smooth without pits; ventrals 141; subcaudals 59; anal
divided; total length, 228 mm.; tail 60 mm.

Above brown, the edges of the scales having deeper brown or
blackish edges, the centers being light brown or fawn resulting in
a pattern of indistinct dark and light lines. These dark lines vary in
thickness; one on lower and one on upper edge of first scale row;
one on edges of third and fourth rows; a narrower line on edges of
sixth and seventh; the median scale row is almost entirely dark.

The light lines on first and second rows are most distinct; two
lower dark lines merge into a single dark line on base of tail; head
brown flecked and mottled with fawn; a pair of narrow light spots
converge on the back point of frontal; first supralabials with irregu-
lar cream spots; a larger more prominent spot on sixth labial but
extending onto adjoining scales; dark spots on infralabials; a distinct
cream bar across neck; chin, venter, and under side of tail cream;
the extreme outer edges of ventrals dark brown.

No. 655, differs in a few characters. The anterior part of frontal
forms a curved line. On right side third and fourth labials are fused,
thus only third labial enters eye; on left side the third labial trans-
versely severed thus making an "extra" loreal. The part of rostral
visible above smaller than in preceding specimen; labial markings
nearly the same, and light markings on frontal borders only dimly
evident; temporals 1 + 1 + 1; scales 17-17-17; ventrals 143; sub-
caudals 73. Total length, 242 mm.; tail 73 mm. The frontal region
is somewhat broader, and the dark stripes are only dimly evident.
The length of the labials in relation to height varies somewhat from
the type.

I examined two specimens at the Harvard Museum of Compara-
tive Zoology from La Palma, Costa Rica collected and identified by
Dunn as Trimetopon gracile.

These are Harvard M.C.Z. No. 28049 (with 148)2 ventrals, tail in-
complete; 8-8 supralabials, 8-8 infralabials; prefrontals fused; scales
17-17-17 rows; second chinshields small, about one-third size of first
pair; keels present above anal region ) and No. 2S050 ( ventrals 148;
subcaudals 69; a white line from eye to lip).

78 The University Science Bulletin

A specimen in the U. S. National Museum, No. 75036 from San
Jose, Costa Rica has the following characters: rOstral narrowly visi-
ble above; internasals small, their length in prefrontal length ap-
proximately 2-2 5 times; prefrontals fused, twice (or more) size of
frontal; frontal equals its distance to snout tip; parietals elongate,
one-sixth longer than snout; loreal longer than high; one preocular,
one postocular, the latter distinctly the larger; temporals 1-1; 8-8
supralabials; 8-8 infralabials, five touch first chinshields which are
thrice area of second pair; first labials form a suture; 153 ventrals
(the first separated from the posterior chinshields by four scales);
anal divided; subcaudals 70 -f- 1.

A nuchal band, divided; a pair of symmetrical white spots on back
of frontal and a similar pair on edges of parietals; an irregular white
line crosses snout on rostral and internasals, goes onto labials termi-
nating at lip; a light spot below eye to seventh labial; head other-
wise brown flecked; chin immaculate except mental and the two
labials on each side; body scales brown with elongate dashlike lines
of varying width thus appearing as light and dark stripes of varying
width and distinctness; body color encroaches on ventrals but venter
chiefly immaculate cream white.

Trimetopon viquezi Dunn

Trimetopon viquezi Dunn, Copeia, 1937, no. 4, p. 215 (type locality, Siquirres, Costa Rica).

The following characters appear in the type description: Nasal
semidivided; two prefrontals; one preocular, two postoculars; tem-
porals 1-1; supralabials seven, the third and fourth border orbit;
infralabials eight, four touching the anterior chinshields which are
separated from the mental and are longer than the posterior; dorsal
scales in 17-17 smooth rows, save a few keeled scales on sides above
anus; ventrals 161 + 2; caudals 33 +. The total length is 209 +
(part of tail missing).

Black above, each scale with a white streak at base except those
of the vertebral and the fourth row, resulting in a dark vertebral
line and a dark lateral line; throat white; belly and under side of tail
red; a black streak from eye to suture of labials four and five; a white
streak from eye onto the sixth and seventh labials; a white streak
from side of neck crosses the seventh labial; anterior labials spotted
black and white.

I have not seen the type, which is from Siquirres, Costa Rica.

Taylor: Review of Snakes of Costa Rica 79

Trimetopon simile Dunn

Trimetopon simile Dunn, Occ. Papers Boston Sue Nat. Hist., vol. 5, 1930, p. S3] (type local-
its-. Reventazon, Costa Rica [= La Junta], later stated to be Siquirres).

Prefrontals fused into a single large scale. Scales in 15 rows; one
preocular; two postoculars; temporals 1 + 1; supralabials seven, the
third and fourth border eye; infralabials seven, four touching the an-
terior chinshields, which are longer than posterior pair.

Ventrals 122; anal divided; subcaudals 69. Black above, white on
ventral surface; head white as far forward as middle of frontal and
including third labial; a black spot on seventh labial. Total length,
155 mm.; tail, 50 mm.

I have seen only the type.

Trimetopon gracile (Gunther)

AMabes gracilis Gunther, Ann. Mag. Nat. Hist., ser. 4, vol. 9, Jan. 1872, p. 18, pi. 3, fig. D

("Elevated country near Cartago").
Trimetopon gracile Boulenger, Catalogue of the Snakes in the British Museum, vol. 2, pp. 184-

185 (Boulenger gives the count of ventrals as 141-149 for the two cotypes, and 60-65 for

the subcaudal counts; that given in the type description is ventrals 149, subcaudals 69);

Dunn, Copeia, 1937, no. 4, Dec. 31, pp. 214-215.

The characters of the type are: body and tail slender, subcylin-
drical; head narrow, not distinct from neck; rounded rostral very
broad and low; very small narrow internasals; prefrontals fused
into a single large scale; frontal broad and long, five-sided with the
posterior angle produced and pointed; parietal length equal to
frontal and prefrontal length together; nostril between two small
nasals; loreal longer than high; one preocular not reaching upper
surface of snout; one postocular; seven supralabials, the third and
fourth border eye; temporals 1 + 1+2; first infralabials in contact;
chinshields, two pairs subequal in size; scales 15 rows, smooth, with
single apical pits; ventrals 149, subcaudals 69. Length approxi-
mately 300 mm.

Upper parts nearly uniform blackish brown, the anterior and
lateral scales somewhat lighter in the center; an indistinct narrow
brownish collar; lower parts yellowish.

Boulenger, loc. cit., adds certain data, presumably taking into
consideration a second specimen: "seven or eight upper labials,
third and fourth or fourth and fifth entering the eye; four or five
lower labials in contact with the anterior chin-shields, which are
longer than the posterior."

As for the color, Boulenger states, "Dark brown above, with five
blackish longitudinal lines; a more or less distinct yellowish collar;

80 The University Science Bulletin

lateral scales lighter in the centre; labials yellowish, spotted with
black; lower parts uniform whitish."

Giinther's figure shows the anterior and posterior chinshields as
of approximately the same size, and the parietal longer than its
distance from the tip of the snout.

Genus Hypsiglena Cope

Hypsiglena Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, 1800, p. 240.

Genotype: Hypsiglena ochrorhynchus Cope.
A single species occurs in Costa Rica.

Hypsiglena torquata torquata (GiAnther)

Leptodeira torquata Gunther, Ann. Mag. Nat. Hist., ser. 3, vol. 5, Feb. 1800, p. 170, pi. 10,

fig. A.
Leptodeira torquata torquata, (part.) Dunn, Proc. Nat. Acad. Sci., vol. 22, 1930, p. 094.
Comastes quincunciatus Jan, Iconographie Generale des Ophidiens, livr. 38, 1871, pi. 1, fig. 1

(Costa Rican specimen listed).

Knowledge of the species as a member of the Costa Rican fauna
is based on two reports: One is that of Jan, loc. cit.; the other is
that of a Costa Rican specimen in the Museo Nacional de Costa
Rica.

The following characters distinguish the species: snout rounded,
obtuse, rostral much broader than deep, the part visible above one
fourth to one third its distance from frontal; frontal once and a
third to once and a half as long as broad, as long as or a little longer
than its distance from end of snout; loreal longer than deep; one
preocular; a presubocular; two postoculars; temporals 1+2; eight
(exceptional^ seven) upper labials, fourth and fifth (third and
fourth) entering eye; five infralabials touch first chinshields, which
are shorter than posterior pair; scales in 21 rows; ventrals 169-174;
anal divided; subcaudals 41-55.

Grayish or pale brown above, with a dorsal row, or two alternat-
ing series of dark brown spots separated by yellowish interspaces;
sides with two or three alternating series of small dark brown spots;
a large dark brown blotch on the nape preceded by a yellow collar;
a dark brown streak on each side of the head passing through the
eye; head speckled with dark brown; lower parts white. Total
length 4C0 mm.; tail 60 mm.

Genus Amastridium Cope

A mast rid turn Cope, Proc. Acad. Nat. Sci Philadelphia, 1800, p. 370.

Genotype: Amastridium veliferum Cope.

Only one species has been reported from Costa Rica.

Taylor: Review of Snakes of Costa Rica 81

Amastridium veliferum Cope

Amastridium veliferum Cope, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 370 (type locality,

"Cocuyas de Veraguas, N. Grenada" = Coyucas, Panama); Proc. Amer. Philos. Soc, vol.

23, 1886, p. 495; Amer. Nat., vol. 26, 1892, p. 481, and vol. 28, 1894, p. 840; Rept.

U. S. Nat. Mus., 1898, pi. 24, fig. 13; Boulenger, Catalogue of the Snakes in the British

Museum, ed. 2, vol. 2, 1894, p. 352; Dunn, Proc. U. S. Nat. Mus., vol. 65, 1924, p. 1-3

(Cariblanco, Costa Rica, etc.).
? Fleischmannia obscura Boettger, Katalog der Reptilien-Sammlung in Museum der Sencken-

bergischen Naturforschenden Gesellschaft in Frankfurt Am. Main. Teil II, (Schlangen)

1898, p. 69 (type locality, San Jose, Costa Rica).
tPhrydops melas Boulenger, Ann. Mag. Nat. Hist., ser. 7, vol. 15, May 1905, pp. 453-454

(Carriblanco, Costa Rica).
Phydrops melas (sic) Dunn, Copeia, 1391, no. 4, p. 163.

The characters by which this snake may be diagnosed are as fol-
lows: 12 maxillary teeth, the last one or two largest, not grooved;
pupil round; no loreal; scales in 17 rows, smooth anteriorly, lightly
keeled posteriorly, lacking pits, subcaudals divided.

Rostral scarcely visible above; internasals two-thirds times as
long as the prefrontals; frontal longer than its distance from snout
tip; one preocular; temporals, 1+2; seven supralabials, third and
fourth enter eye; nine infralabials; ventrals 123-127; subcaudals
79-85; anal divided.

Above and below reddish brown, paler on center of ventrals;
every fourth scale of the fifth row on each side pale, the adjacent
scales on the fourth and sixth rows generally darker; top of head
much lighter, varied anteriorly, palest behind eye and above
labials; latter dark with a few light spots. Total length 355 mm.;
tail, 120 mm.

I have examined a Harvard specimen (M. C. Z. No. 15319).
The following characters are in evidence: scale formula 17-17-17;
no loreal present; nasal single, the posterior end slightly higher
than anterior; eye larger than its distance from rostral; one large
squarish preocular, two postoculars; 7-7 supralabials, the third and
fourth enter orbit; infralabials 9-9, 4 touch chinshields; fifth supra-
labials touches the parietal narrowly, the first temporal thus being
separated from the postoculars; frontal narrow, twice as long as
wide; temporals 1+2 (the two upper fused on one side); ventrals
136; anal divided; 48+ subcaudals.

Color of head and neck yellow-brown, the dark body color run-
ning forward to a point on the posterior edge of the parietals; fourth
and seventh scale rows with darker lines made up of short dashlike
marks; venter dark. A row of cream colored dots on the fifth scale
row scarcely discernible.

6—3286

82 The University Science Bulletin

Records of the snake for Costa Rica are: a specimen in the Mu-
seum of Comparative Zoology, Harvard College, from Cariblanco,
Costa Rica; that of the type specimen of Fleischmannia obscura
from San Jose, Costa Rica and the type of Boulenger's Phrydops
mclas from Cariblanco, Costa Rica.

Genus Spilotes Wagler

Spilotes Wagler. Natiirliches System der Amphibien. 1830, p. 170.

Genotype: Coluber pullatus Linnaeus.

Two forms of Spilotes occur in Costa Rica. These are usually
considered as subspecies of pullatus. There is, however, doubt that
mexicanus and pullatus intergrade where they meet. This group
definitely is in need of a revision.

Key to Forms of Costa Rican Spilotes

Yellow markings confined largely to the anterior part of body; latter part nearly

uniform black ; supralabials 7 (usually) pullatus pullatus

Yellow markings of body extending to end of tail; supralabials 8 pullatus mexiacanus

Spilotes pullatus pullatus (Linnaeus)

Coluber pullatus Linnaeus, Museum Adolphi Friderici Regis, 1754, p. 35, pi. 20, fig. 3 (Asia,

in error).
Spilotes pullatus Wagler, Natiirliches System der Amphibien, 1830, p. 179.
Spilotes pullatus pullatus Dunn and Bailey, Bull. Mus. Comp. Zool. Harvard Coll., vol. 86,

1939, p. 18.

Two specimens of this large tree snake were captured, one
(M. N. H. No. 25757) June 26, 1947, along the Reventazon River
about 3 km. southwest of Turrialba, and one (M. N. H. No. 25756)
July 19, near the same locality.

The head in each is banded with yellow but the yellow markings
on body are greatly reduced, only traces being evident in a few
small dashlike marks on a few scales. On the anterior part of the
body, the ventrals are about half yellow; farther back the yellow is
reduced so that only half of a ventral may be affected, and at ventral
number 106 and 122 respectively the color ceases to have any
yellow and from there to end of tail the color is uniform black. The
ventral counts of the two specimens are 226, 221; subcaudals, 113 +
and 74 -f- respectively.

The following characters also obtain: rostral scarcely visible above;
internasals about one-fourth shorter than prefrontals; length of
frontal slightly less than its distance to end of snout, about as wide
as long, one-fourth shorter than parietals; length of latter scales
less than their distance to internasals; loreal very small, about as
long as high; preocular very large, minutely separated from frontal

Taylor: Review of Snakes of Costa Rica 83

scale; two postoculars; temporals. 1 + 1 + 1 (1 + 1); supralabials,
7-7, fourth and fifth entering eye; infralabials, 8-8, four touch chin-
shields, which are much larger than second pair. Scale rows keeled
except outer, the formula 14-16-10; paired or single apical pits are
present.

The tail of No. 25757 appears to be perfectly normal, the terminal
scute being well developed. However its thickness at the tip sug-
gests a well-repaired break. Moreover the number of subcaudals
is 39 less than in the other specimen and that one has the extreme
tip missing. No. 25756 has a more or less symmetrical pair of scales
segmented from the inner parts of the parietals, that on the right
being partially fused to the parent scale.

A specimen in the U. S. National Museum (No. 32650, Costa Rica)
has the head largely black, with a neck band of yellow. A yellow
stripe crossing head just behind the eyes is interrupted across the
top of the head. There are yellow spots on rostral, nasals, pre-
frontals, internasals, supraoculars and frontal. The ventrals are
black and yellow, the yellow predominating anteriorly, the latter
half of body and tail uniform black. Ventrals 218, subcaudals 118.

Spilotes pullatus mexicanus (Laurenti)

Cerastes rtiejncanus Laurenti, Specimen Medicum exhibens Synopsin Reptilinm emendatum,

1768, p. 83.
Spilotes pullatus 7nexicanus Amaral, Mem. Inst. Butantan, vol. 4. 1929, pp. 282-28-1 (Mexico

to Nicaragua).
Spilotes pullatus auribundus Cope. Proc. Acad. Nat. Sci. Philadelphia, vol. 13, 18eil, p. 300.

Amaral in his review of the Genus Spilotes characterizes this pre-
sumed subspecies as having a loreal, eight supralabials, the fourth
and fifth entering eye orbit; temporals 1 + 1 or 1 + 2; ventrals
204-222; subcaudals 115-138; scale rows 18 (19) around middle of
body.

The presence in Costa Rica of specimens of this subspecies, if
indeed it is related subspecifically to pullatus, is attested by certain
specimens in the U. S. National Museum: Nos. 37724, 37725, and
37726, Esparta, Costa Rica.

No. 37725 has the supralabials 8-8; scale rows 16-16-16-12; ventrals
220; subcaudals 134. The yellow banding of the head and body
continues on to the tail, the black interspaces are of about 7 scale
lengths, the yellow 3 or 4 scale lengths.

No. 37726 is a young specimen with the body coloring forming
strongly-defined body bands, and in appearance is very different
from that of the adult.

84 The University Science Bulletin

Genus Thalerophis Oliver

Thalerophis Oliver, Copeia, 1947, p. 64.

Genotype: Coluber richardi Bory St. Vincent.

The group of tree snakes long known under the genus Leptophis
are here treated under the genus Thalerophis since the name Lep-
tophis is not available. The species belonging in the genus have
recently been reviewed by Dr. James Oliver * and several nominal
species formerly believed to occur in Costa Rica are reduced to
synonymy. Thus aeruginosus Cope, bilineata Gunther and satura-
tus Cope are regarded as synonyms of T. depressirostris Cope; and
ultramariniis is regarded as a synonym of T. richardi occidentalis
Gunther.

The following four species are recognized as occuring in Costa
Rica: depressirostris, mexicanus mexicanus, nebulosus, richardi oc-
cidentalis, and these may be differentiated on the basis of the fol-
lowing synopsis.

Synopsis of Costa Rican Species of Thalerophis

1. Loreal normally present 2

Loreal normally absent 3

2. A strong black stripe from snout through eye and onto side of body; body scale
rows keeled except outer; ventrals: in males 148-169, average about 157; in fe-
males 154-174, average about 161; maxillary teeth 20-25 (average 21), last two

enlarged mexicanus mexicanus

No black stripe beginning on snout ; a short stripe beginning behind eye terminates
back of the jaw angle; two paravertebral scale rows keeled, others smooth; ven-
trals 144-158; nine supralabials ; maxillary teeth 33-36, the last three or four
enlarged depressirostris

3. A dark stripe on lores continued back of eye to jaw angle; a greenish blue, or
blue dorsolateral stripe on third and fourth scale rows (or third, fourth and fifth) ;
ventrals: males 150-160, female 158; maxillary teeth 28-29, the last three en-
larged nebulosus

No dark mark in front of eye; top of head dark blue or greenish blue, which
color extends over the body; ventrals: males 158-177, females 161-181; maxillary
teeth 18-25 (average about 21) ; supralabials 8 or 9 richardi occidentalis

Thalerophis mexicanus mexicanus (Dumeril, Bibron and Dumeril)

Leptophis mexicanus Dumeril, Bibron and Dumeril, Erpetologie Generate . . . , vol. 7, pt. 1,

p. 536 (type locality, Mexico.)
Thalerophis mexicanus mexicanus Oliver, Bull. Amer. Mus. Nat. Hist., 1948, pp. 211-217.

Rostral broader than high barely visible from above; internasals
usually a little shorter than the prefrontals; frontal once and one
third to once and two thirds as long as broad, as long as its distance
from the end of the snout, shorter than parietals; nasal elongate, at
least partially divided; loreal present, at least twice as long as high;
one preocular, very large but not reaching frontal; two postoculars;

* The Relationships and Zoogeography of the Genus Thalerophis. Oliver, Bull. Amer. Mus.
Nat. Hist., vol. 92, 1948, pp. 101--2so.

Taylor: Review of Snakes of Costa Rica 85

temporals 1+2; eight supralabials, fourth and fifth entering eye;
usually 10 infralabials. Scales in 15-15-11 rows, keeled save on
outer rows.

Green or dark greenish blue above; a narrow black stripe along
the side of head passing through eye, continues along the side of
the body and tail, varying somewhat in width and intensity. When
outer epidermis is present the color on sides and dorsum is brown-
ish to bronze-brown; when it is removed the general color is green-
ish blue to greenish or olive brown; belly white.

This species has been reported from San Jose; other specimens
are reported, labeled only "Costa Rica" without definite locality.

Thaler ophis depressirostris (Cope)

Philothamnus depressirostris Cope, Proc. Acad. Nat. Sei. Philadelphia, vol. 12, 1860, p. 557

(Cocuyas de Veraguas, Colombia).
Diplotropis bilineata Giinther, Ann. Mag. Nat. Hist., ser. 4. vol. 9, 1872, p. 24, pi. 6, fig. B

(type locality, Costa Rica).
Leptophis aeruginosus Cope, Journ. Acad. Nat. Sci. Philadelphia, vol. 8, 1876, p. 132 (type

locality, Costa Rica).
Leptophis saturatus Cope, Journ. Acad. Nat. Sci. Philadelphia, vol. 8, 1876 (1875) p. 133 (type

locality, Sipurio, Costa Rica).
Tlialeruphis depressirostris Oliver, Bull. Amer. Mus. Nat. Hist., vol. 92, 1948, pp. 203-207,

figs. 1G, 4, 8, and plate 16 (figure on right).

This species is represented by a single specimen, M. N. H. No.
25704, collected by Prof. Marco Tullio Pacheco, and presented to
me. It was found near Peralta at an elevation of 300 meters. The
following scale characteristics are present:

Rostral visible above as a small triangle; internasals as long as
prefrontals; frontal more than one fourth longer than wide, its
length about equal to its distance from end of snout; length of
parietal about equal to its distance from intemasal; nasal nearly
equally divided; loreal twice as long as wide; preocular large, well
separated from frontal; two postoculars; temporals 1+2; supra-
labials 8-9, the fifth and sixth (fourth and fifth) bordering eye; in-
fralabials 10-10, five bordering the chinshields; scale formula, 17-
15-11; the scales smooth except the two rows bordering the single
median row; indistinct apical pits on some scales; ventrals 147; tip
of tail lost; anal divided.

Thalerophis nebulosus (Oliver)

Leptophis nebulosus Oliver, Occ. Papers Mus. Zool., Univ. Michigan, no. 462, 1942, p. 12,
fig. 4 (second figure p. 192) (type locality. Cariblanco, Costa Rica).

Loreal absent; a single preocular, touching or narrowly missing
contact with frontal; frontal large, length approximately equal to
that of interparietal suture; temporals 1 + 2; eye equal to its distance

86 The University Science Bulletin

from anterior border of nostril; eight supralabials normally; infra-
labials ten (usually); scale formula, 15-15-11, keeled save on outer
row. Ventrals 150-160; subcaudals 145-151; anal plate divided.

Top of head bluish green; a narrow black stripe beginning on
upper edge of first or second labial passing back through eye and
for a short distance on the side of the body. Body color cream-
white on outer scale rows, the next three rows bright blue; scales on
head, below the black stripe, white or cream; venter and chin
cream.

Only the type is known from Costa Rica.

Thalerophis richardi occidentalis (Giinther)

Ahaetulla occidentalis Giinther, Proc. Zool. Soc. London. 1859, p. 412 (type locality, Guayaquil

and western Equador).
Thalerophis richardi occidentalis Oliver, Bull. Amer. Mus. Xat. Hist., vol. 92, 1948, pp. 241-

245.
Leptophis ultramarinus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 46, 1894, p. 204 (type

locality, Pozo Azul, Costa Rica).

Loreal shield normally absent; normally one preocular in contact
with the frontal; two postoculars; supralabials eight or nine, frontal
large; temporals 1+2; infralabials usually ten; scale rows 15-15-11,
all keeled save outer one or two rows, and the keels may be lacking
from the median row; ventrals: males 152-177, females 161-182;
subcaudals: males 153-189, females 161-175.

Color of head dark blue to greenish blue, this color extending
onto body and tail, extending even to outer edges of ventrals; infra-
labials, chin and venter pale greenish blue to yellowish white.

Reported from Barranca, Buenos Ayres, Colorado Bar, Limon,
Pozo Azul, and Siquirres.

I have examined an uncatalogued specimen of this species in the
U. S. National Museum collected 2 mi. SE of Turrialba, Costa Rica.
It has the following characters: loreal absent; supralabials 9-9, the
fifth and sixth entering orbit; one preocular, two postoculars; 10-11
infralabials, five touching the first chinshields which are shorter
than second pair; three labials touch prefrontals; eye large, equal to
its distance to front edge of nostril; frontal equal to its distance to
snout tip; ventrals 172; subcaudals 173+ ; scales 15-15-11, the scale
rows bordering the median row strongly keeled as is the adjoining
row; outer scales smooth but each with a median black line simu-
lating a keel.

Taylor: Review of Snakes of Costa Rica 87

Genus Dryadophis Stuart

Dryadophis Stuart, Copeia, No. 1, 1939, p. 55.

Genotype: Coluber boddaerti Sentzen.

Only a single species is known to occur in Costa Rica.

Dryadophis melanolomus alternatus (Bocourt)

Coryphodon alternatus Bocourt, Bull. Soc. Philom., ser. 7, vol. 8, 1884. pp. 133-142, and
E'tudes sur les Reptiles; Mission Scientifique au Mexique et dans l'Amerique Centrale,
livr. 11, 1888, pi. 45, fig. 3a-e (not mentioned in text).

Drymobius alternatus Cope, Bull. U. S. Nat. Mus., no. 32, 1887, p. 70.

Dryadophis melanolomus alternatus Stuart, Misc. Publ. Mus. Zool., Univ. Michigan, no. 49,
Mar. 17, 1941, pp. 81-86.

The specimen described by Bocourt, or at least the specimen
figured by Bocourt, for Dryadophis alternatus is strikingly different
from the species considered under this name by Stuart in his revision
of the genus, loc. cit. Bocourt's figure is of a snake with well-defined
quadrangular dorsal blotches alternating with a lateral series of
somewhat smaller quadrangular blotches. Under this name Stuart
treats of a snake that is uniformly colored. R is presumed that the
young have a pattern differing from adults, and the specimen de-
picted by Bocourt was a juvenile.

Of the five specimens at hand, R. C. T. Nos. 417 and 656 are from
Turrialba, elev. about 1900 ft.; R. C. T. No. 968 from San Isidro
El General, elev. about 2000 ft.; and R. C. T. Nos. 1437 and 1438
from Los Diamantes, about 2 kilometers south of Guapiles, at an
elevation of about 800 feet.

Specimens from Los Diamantes were olive in life; the ventral
color being immaculate pinkish. The sides of the ventrals are
subangular; however there is no decided keel present. The outer
edges of the ventrals were gray-olive in life. With a loss of the outer
epidermis the specimens now are uniform gray-blue. When placed
under water, the scales appear slightly darker on their edges. The
pink color is still present on the ventrals. Neither of the two speci-
mens displays any transverse banding or longitudinal lineation.
One of these has an undivided anal scale.

The two specimens, Nos. 417, 656 differ somewhat. In life they
were olive above without markings, the venter with a yellow wash
and the throat and chin bluish gray with small yellow spots. All
trace of olive and yellow has now disappeared, leaving the dorsum
dull grayish blue, the venter dirty white with the edges of the ven-
trals clouded with gray. The chin spots are cream-white. An
indistinct median zigzag line is present under the tail.

88 The University Science Bulletin

The small specimen from San Isidro El General when first exam-
ined was olive in color but with some slight indication of darker
crossbars on the anterior part of the body, and a faint indication of a
light lateral stripe. In the preserved specimen with outer epidermis
removed, no trace of the crossbands remains; but submerged in
water there are visible certain darker and lighter stripes; a dim
light stripe is present on the fourth and slightly on the fifth scale
row. The third scale row is bordered by darker color, tending to
form a dark longitudinal line. The first and second scale rows are
a little lighter than surrounding scales so that a dim lighter stripe
is evident here. The lower edges of the first row and outer edges
of the ventrals are darker than the remainder of the scale and another
dim, dark line is evident. Ventrals angular, the outer, turned-up
edges blue-gray, the ventral portion being finely mottled with light
gray. On the angular keel a light line is more or less evident.

Data on Dryadophis melanolomfs alternates (Bocourt)

Labials
No. Sex Ventrals Subcaudals Supralabials enter eye Infralabials

1438 $ 182 97 9-9 4,5,6 10-10

1437 S 180 96 9-9 4,5,6 10-10
417 S 175 91 9-9 4,5,6 11-11
656 S 179 101 9-9 4.5,6 10-11
968 S 176 108 8-9 f 4,5,6 9-10

\(3,4,5)

Data on Dryadophis melanolomfs alternates (Bocourt) (concluded)

Labials
touch Ventrals Ventrals

No. chinshields Temporals Scale rows Anal keeled color

1438 5 2 + 2 17-17-15 2 slightly reddish
1437 5 2 + 2 17-17-15 1 slightly reddish

417 5 2 + 2 17-17-15 2 not gray

656 5 2 + 2 17-17-15 2 not grav

968 5 2 + 2 17-17-15 2 not gray

Genus Drymobitjs Fitzinger

Drymobius Fitzinger, Systema Reptilium, 1843, p. 26.

Genotype: Herpetodryas margaritiferus Schlegel. Four species
are recognized in Costa Rica.

Key to Species of Drymobius in Costa Rica

1. Body with a series of brown saddlelike blotches, alternating with smaller lateral

spots ; ventrals 146-152 ; subcaudals 88-91 rhombifer

Body not marked with blotches, but generally black or green 2

2. Body black, each scale with a dashlike green or yellow-green mark; ventrals 146-

149 ; subcaudals 109-117 margaritiferus margaritiferus

Body green ; belly more or less yellowish in life 3

3. Body green, the color encroaching on ventrals; keels of three median rows black

?nelanotropis
Body green, the keels of the median rows not black chloroticus

Taylor: Review of Snakes of Costa Rica 89

Drymobius margaritiferus margaritiferus (Schlegel)

Herpetodryas margaritiferus Schlegel, Essai sur la physionomie des serpens, vol. 2, 1837, p. 184.
Drymobius margaritiferus [margaritiferus] Bocourt, Etudes sur les reptiles; Mission Scientifique
au Mexique et dans l'Amerique Centrale, livr. 12, 1890, pp. 71G-718, pi. 49, fig. 2.

This common, widespread species is represented by three speci-
mens. M.N.H. No. 25701, an adult male, was captured as it at-
tempted to cross the road in front of our car, five kilometers south-
west of Turrialba. A young specimen ( R.C.T. No. 205 ) was taken
three kilometers west of Turrialba. A third specimen (M.N.H. No.
25165) in the University collection was obtained five kilometers
southwest of Turrialba in 1947 by Mr. Howard Westman.

The characteristics of the young (No. 205) follow: a greenish
white longitudinal mark on the frontal; a pair of greenish white lines
curving behind last labial and extending to parietal; following this
a large series of vertical greenish white and black lines running up
on the sides to near the mid-line, a line occasionally meeting its fel-
low, but more commonly alternating with it. These lines gradually
more indistinct on latter half of body where light and dark colors
tend to form indefinite longitudinal lines, the most distinct being
the greenish white line on the two outer scale rows that terminate
at anus; ends of ventrals blackish; the free edge of the scales may be
blackish for a greater or lesser distance, rarely crossing ventral;
supralabials with black sutures; infralabials and chin greenish
cream.

In the two adults the color is blackish, each scale with a small
dashlike greenish or greenish-white spot near the center, the green-
ish spots becoming regularly larger on lower (outer) rows; black
on ventrals often across the posterior part of scale; spots on No.
26165 slightly larger and more distinct than those on No. 25701.

I have examined four Costa Rican specimens in the U. S. National
Museum; three of these have a ventral scale count of 148; one has
147.

Data on Drymobius margaritiferus margaritiferus

No.

Sex or
age

Supralabials

Infralabials

Ventrals

Subcaudals

Scale
formula

205
25701
25165

yg-

s

9- 9
9- 9
9-10

11-10

11-11

10-11

148
146
149

109
117

17-17-15
17-17-15
17-17-15

Drymobius rhombifer (Gunther)

Coryphodon rhombifer Gunther, Proc. Zool. Soc. London, 1860, p. 236 (type locality, Esmer-

aldas, Ecuador).
Drymobius rhombifer Peters, Monatsb. Akad. Wiss. Berlin, 1879, p. 777.

Two specimens of this species (M.N.H. Nos. 25734, 25744) were
obtained at Turrialba. The following characters are present in the

90 The University Science Bulletin

species; rostral about one fifth wider than high, the area visible
above, small, its length about one third the length of internasals;
length of latter three fourths of prefrontals, which are one fifth wider
than long; frontal one fourth longer than wide, its length equal to
its distance from tip of snout or slightly less; parietals short, their
width two thirds of their length; nasal divided; posterior part higher
and shorter than anterior; loreal a little longer than high; one pre-
ocular not reaching frontal; two postoculars; temporals 2 + 2; su-
pralabials 9-9, the fourth, fifth and sixth entering eye; infralabials
9-9, five touching first chinshields, which are a little shorter than
second pair; scale formula: 17-17-15, keeled lightly; ventrals 152;
subcaudals 88; anal divided.

A series of dark-brown, saddlelike blotches present reaching to
ventrals where they border a black ventral spot; between these
latter are small ( often ) triangular spots touching two or three ven-
tral black spots, the apex reaching as high as fifth lateral scale row.

Belly with an irregular series of black blotches along the outer
part of ventrals rarely more than one scale wide; between these on
middle of venter small scattered black spots. On the brown blotches,
each scale edged with black. Chin and throat immaculate, lower
part of supralabials nearly white.

The smaller specimen is quite similar. The ventrals are about
146 ( injured ), the subcaudals 91. The dorsal head scales have deep
black spots or punctations. In both specimens there is a somewhat
lighter area on each parietal.

This is a South American species that has entered Costa Rica.
The greater part of the range is in Colombia, Ecuador and Peru.

Drymobius chloroticus (Cope)

Dendrophidium chloroiicum Cope, Proc. Amer. Philos. Soc, vol. 23, 1886, p. 278 (Guatemala).
Drymobius chloroticus Boulenger, Catalogue of the Snakes in the British Museum, vol. 2, 1894,
pp. 10-17.

Two specimens were collected at Isla Bonita, one, R.C.T. No. 702,
an adult female, and R.C.T. No. 654, a young male.

Data taken from the larger, female specimen follows: part of ros-
tral visible above less than half length of internasals; latter scales
only a little shorter than the prefrontals; frontal one fourth longer
than wide, its length equal to its distance to end of snout or slightly
more; length of parietal equal to its distance from internasal; nasal
divided, the anterior part the larger; loreal nearly twice as wide as
high; one preocular, very large, widely separated from frontal; two
postoculars; temporals 2 + 2; supralabials, 9-9, the fourth, fifth and

Taylor: Review of Snakes of Costa Rica 91

sixth border eye; infralabials, 10-11, the first five touching first chin-
shields, which are about three fifths the length of second pair; ven-
trals, 186; anal divided; subcaudals, 121. Scales keeled, the formula,
17-17-15 with paired pits. Maxillary teeth about 31, increasing a
little in length and size posteriorly.

The second specimen, No. 654, has the infralabials 9-9; the ven-
trals 186; the subcaudals 100, the tail seemingly complete.

In preservative the specimens are uniform bluish gray above, the
belly greenish yellow, each ventral and subcaudal bordered by
blackish. In life, olive on dorsal and lateral regions, greenish yellow
on venter; upper labials ( except upper edges ) and chin, cream yel-
low.

Drymobius melanotropis (Cope)

Dendrophidium melanotropis Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876

(1875), p. 134, pi. 26, fig. 1 (type locality, Costa Rica).
Drymobius melanotropis Gaige, Hartweg and Stuart, Occ. Papers Mus. Zool. Univ. Michigan,

no. 357, 1937, p. 13 (Rio Siquia Nicaragua).

Scales in 17 rows, with apical pits present, all keeled except the
two outer rows, the keels on the paravertebral rows strongest; ros-
tral broader than deep; frontal bell-shaped, wide in front, contracted
behind, as long as the parietals; loreal much longer than high; one
preocular, one subocular; two postoculars; temporals 2 + 2; nine
supralabials, the fourth, fifth and sixth bordering orbit; ventrals 152-
163, not angulate; subcaudals 91-96; anal divided.

Green above and on sides, the color occupying the outer fourth
of the ventral scales. Yellow on venter; keels of three median rowj
black.

Genus Dendrophidion Fitzinger

Dendrophidion Fitzinger, Systema Reptilium, 1843, p. 26.

Genotype: Herpetodryas dendrophis Schegel.
Two species are known from Costa Rica.

Key to the Costa Rican Sfecies of Dendrophidion

Ventrals 150-175; subcaudals 113-155; olive brown, uniform or with blackish
cross-bands enclosing rounded whitish spots, or with whitish black-edged cross-
bands; upper lip whitish or yellow dendrophis

Ventrals 183; subcaudals 127; color brown, without markings; below yellow;

upper lip partly yellow paucicarinatus

Dendrophidion paucicarinatus (Cope)

Drymobius paucicarinatus Cope, Proc. Acad. Nat Sci. Philadelphia, 1894, pp. 202-203 (type
locality, La Candelaria, Costa Rica).

Scales in 17 rows, five median rows faintly keeled, others smooth;
one preocular, not reaching frontal; two postoculars; loreal sub-

92 The University Science Bulletin

quadrate, longer than high; temporals 1 + 2; four and one-half
scales bordering each parietal; supralabials nine, fourth, fifth and
sixth border orbit, the eighth and ninth longer than high; eye
large, its diameter equaling length of muzzle from its border to the
nostril, and equaling a little more than half interocular distance;
frontal wide in front, contracting rapidly posteriorly, the lateral
borders very little concave; ten infralabials; anterior chinshields
shorter than posterior; ventrals 183; subcaudals 127; anal divided.
Color above brown without markings, below yellow; ends of ven-
trals and a narrow transverse line near base of ventrals lead colored;
much of upper lip yellow.

Dendrophidion dendrophis (Schlegel)

Herpetodryas dendrophis Schlegel, Essai sur la physionomie des Serpens, vol. 2, 1837, p. 196.
(Also in another work having the same title and date, this name and a synopsis appears on
p. 153 with the following data, "Herpetodryas dendrophis. 15 Rangees d'ecailles carenees
et lanceolees. Dessous de la queue aplati ; ventre convexe. 140 + 196? Dessus brun-
olivatre, marque de nomhreuses bandes etroites transverales foneees, que renferment des
taches claires. De Cayenne.")

Dendrophidium dendrophis Cope, Proc. Amer. Philos. Soc., vol. 23, 1886, p. 278.

Scales in 17 rows with apical pits. Maxillary teeth in a continuous
series 33-50 subequal. Normally supralabials nine, the fourth, fifth
and sixth bordering orbit; one preocular, two postoculars; temporals
normally 2 + 2, but variable; ventrals 150-175; subcaudals 113-155.

"Olive-brown above, uniform or with blackish cross-bands en-
closing rounded whitish spots, or with whitish black-edged cross-
bands; upper lip whitish; ventrals and subcaudals olive on the sides,
whitish, uniform or dotted or edged with olive in the middle."

Genus Pseustes Fitzinger

Pseustes Fitzinger, Systema Reptilium, 1843, p. 27.

Genotype: Dipsas dieperinkii Schlegel [ = Pseustes sulphureus
sulphureus (Wagler) ] .

One species is known in Costa Rica.

Pseustes poecilonotus chrysobronchus (Cope)

Spilotes chrysobronchus Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875),
li. 136, pi. 28, figs. 11a, lib (head only); U. S. Nat. Mus. Bull. no. 32, 1887, p. 71;
Bocourt, Etudes sur les reptiles ; Mission Scientifique au Mexique, et dans l'Amerique Cen-
trale, livr. 11, 1888, pp. 695-696, pi. 48, figs. 3a, 3b (after Cope) (data from Cope);
Giinther, Biologia Centrali-Americana ; Reptilia and Batrachia, Feb. 1894, p. 119.

Phrynonax chrysobronchus Boulenger, Catalogue of the Snakes in the British Museum (Natural
History), vol. 2, p. 22 (data from Cope).

Plirynonax poecilonotus chrysobronchus Amaral, Mem. Inst. Butantan, T. 4, 1929, p. 156
(Costa Rica).

Cope has compared this form with Spilotes fasciatus Peters but
decides the two are distinct. Boulenger likewise regards the two

Taylor: Review of Snakes of Costa Rica 93

as distinct. Giinther however suggests that the two forms may be
identical.

The single specimen in the collection was obtained by Mr. Charles
R. Camp, 5 km. SE of Turrialba, elev. 1950 ft., Province Cartago,
Costa Rica.

Head flattened, distinct from neck; rostral a little broader than
deep; internasals about two thirds the area of prefrontals; latter
scales border loreal but are separated from the supraocular; frontal
as long as its distance from the tip of snout, about one fifth longer
than wide; parietals short, about as wide as long, equally as long
as frontal.

Nasal single, pierced by nostril, but a groove from nostril, to
labial suggests a partial division; loreal small, a little longer than
wide; one large preocular reaching frontal narrowly, and bordering
two labials below; supraocular large, longer than wide; two post-
oculars; temporals -=— f- 2; eight upper labials, the fourth, fifth, and
sixth bordering eye, eighth greatly elongated; 12 lower labials,
seven touching first pair of chinshields; these about one fifth shorter
than second pair.

Scale formula, 21-25-17-15; scales with paired apical pits, and
tending to form diagonal rows on the sides; four median dorsal rows
with indistinct keels. Ventrals 211; subcaudals 132; anal single;
snout to vent 1115 mm.; tail, 436 mm.; total length 1551 mm.; width
of head 43 mm.; snout to rictis oris, 40 mm.

Scales pinkish fawn, each bordered with brown or black. Some-
times entire scale is bordered, sometimes only a part, the border
varying in width, and thus developing a more or less distinct pattern
of indistinct, dark, diagonal blotches joining medially; supralabials,
infralabials, chin, throat and anterior one fourth of venter yellowish
white, there being a few black dots on outer edge of ventrals, which
more posteriorly encroach on the middle of ventrals until they form
continuous transverse black borders on each ventral, and still
farther back tend to cover the entire scale. Top and side of head
dark.

Variation: The total ventral-subcaudal count for one of the co-
types is 337 (ventrals, 220, subcaudals, 117). In the described spec-
imen the total is 343 (ventrals 211, subcaudals 132). This specimen
is a female; I presume that the cotype specimen is also a female.
The tail in the cotype is proportionally shorter and the body longer
( 1670 mm., to 422 mm. ) than in this specimen ( 1551 mm., to 436
mm.).

94 The University Science Bulletin

There are certain scale differences between the cotype and the
Turrialba specimen. Thus in the former there are 8 supralabials
(seven with one partially divided on one side, the third, fourth,
and fifth labials bordering orbit ) , while in the latter there are eight
on both sides, the fourth, fifth and sixth bordering orbit. The cotype
has 1-2 preoculars not reaching the frontal; in this they are 1-1 touch-
ing frontal on both sides.

Pseustes shropshirei (Barbour and Amaral )

Phrynonax shropshirei Barbour and Amaral, Occ. Papers Boston Soc. Nat. Hist., vol. 5, Sept.

12. 1924, p. 131 (vicinity of Gatun, Canal Zone of Panama).
Phrynonax poecilonotus shropshirei Amaral, Mem. Inst. Butantan, Tomo 4, 1929, pp. 317-318,

fig. 4.

This form was originally characterized as follows:

Rostral slightly wider than deep (8:7), just visible above; inter-
nasals two thirds as long as the prefrontals (4:6); frontal as long as
wide, as long as its distance from end of snout, a little shorter than
parietals (10:12); loreal longer than deep; one preocular touching
frontal; 2 postoculars; temporals 2+; seven or eight upper labials,
fourth and fifth or fourth, fifth and sixth entering orbit, the eighth
much the longest; seven to eight lower labials border first chin-
shields which are much shorter than the posterior ( 10 : 17 ) ; scales
in 25 rows (21-24-25-24-21-19-17-15), the median rows feebly
keeled; ventrals 211, obtusely angulate laterally; anal single; sub-
caudals 116-f-, divided.

Blackish brown above, irregularly barred with yellow; dorsal
scales either entirely black or black-edged; head dark brown above,
a wide border on lip yellow, the labials blackish on their upper
edges; lower surfaces yellowish gradually changing to pure black
posteriorly, including tail; anterior ventrals with dark outer edges.
Total length 1630 mm.; tail 430 mm. ( Data from type description. )

A specimen, R. C. T. 682 § , Turrialba, is referred to this species.
It agrees in all essential detail. The head and neck, for 6-8 inches,
are coal black save outer two scale rows which have some white;
beyond this the pattern of transverse marks consisting of black-
edged yellowish scales continues to tail; ventral surface of posterior
part of body and tail nearly uniform black; ventrals 214; subcaudals
116+ (tip missing). Scale rows 21-23-25-13(15); supralabials
8-8; infralabials 11-11, seven touching the first chinshield. Total
length 1714 mm.; tail 469 + mm. (tip missing).

Since the specimen of poecilonotus chrysobronchus was obtained
at this locality (within three miles), I would suspect that the two
are worthy of specific rank.

Taylor: Review of Snakes of Costa Rica 95

Genus Chironius Fitzinger

Chironius Fitzinger, Neue Classification der Reptilien ... 1826, p. 31.

Genotype: Coluber carinatus Linnaeus.

Three species of this genus occur in Costa Rica. They may be
distinguished by the following key.

Key to the Costa Rican Species of Chironius

1. Dorsal scales smooth ; three labials enter orbit ; scales 10-8 melas

Dorsal scales keeled ; two or three labials enter orbit 2

2. Scale formula, 12-8; two labials enter orbit ; (rarely not keeled) carinatus

Scale formula, 10-8, three labials enter orbit grandisquamis

Chironius carinatus (Linnaeus)

Coluber carinatus Linnaeus. Museum Adolphi Friderici Regis, I, 1754, p. 31; and Systems

Naturae, vol. 1, 1766, p. 384.
Chironius (arinatus Fitzinger. Neue Classification der Reptilien .... 1826, p. 60; Dunn and

Bailey, Bull. Mus. Comp. Zool. Harvard Coll., vol. 86, 1939, p. 18 (Cana, Panama).

Specimens of Chironius carinatus may be identified as to genus
by the large scales in even number of rows, varying from 12 rows
anteriorly to 8 on the posterior half of the body. A second easily
observed character is the high dorsal ridge on the body, flat on top,
covered by two scale rows, each scale bearing a sharp keel. An-
teriorly the scales tend to form diagonal rows, but posteriorly this
is not the case.

Our collections contain two specimens caught along the Reven-
tazon River on the farm of the Inter-American Institute of Agricul-
ture, two kilometers southwest of Turrialba, and one at Isla Ronita
on Volcan Poas.

The following scale characters obtain in M. N. H. No. 25745,
Turrialba: rostral visible above; frontal longer than its distance from
the snout, one-fourth longer than wide; parietals short, about one-
fourth times longer than wide, their length equal to that of the
frontal or slightly longer; nasal divided, the anterior part larger;
loreal low, much longer than high; one very large supraocular not
reaching the frontal; two postoculars; temporals 1+2; supralabials
9-9, the fifth and sixth entering orbit; infralabials 11-11, five touch-
ing anterior chinshields, which are about same length as posterior
pair; scale rows 12-12-8; ventrals 147, subcaudals 133.

A specimen in the U. S. National Museum collected by George K.
Cherrie in "Costa Rica" has 153 ventrals and 136+ subcaudals; scale
rows 12-12-8, the keels not strongly evident (the outer epidermis
being absent); supralabials 9-9; the fourth, fifth and sixth border
orbit (the fourth barely touches); infralabials 10-11, the six anterior
touching the first chinshields. Above uniform olive green, the belly

96 The University Science Bulletin

lighter olive; chin, labials and throat, for a short distance, yellowish.

Another specimen (U. S. N. M. No. 14063) has the median scale
rows very strongly keeled; scale formula, 12-12-8; ventrals 145, sub-
caudals 123; anal divided.

A third Costa Rican specimen is uniform olive-green and has the
median rows keeled; ventrals 146, subcaudals 138. I failed to
record the museum number of this specimen.

Chironius grandisquamis (Peters)

Spilotes grandisquamis Peters, Monatsb. Akad. Wiss. Berlin, 1868, p. 451 (type locality, Costa

Rica).
Herpetodryas grandisquamis Cope, Journ. Acad. Nat. Sci. Philadelphia, vol. 8, 1876 (1875),

p. 135; Bocourt, Etudes sur les Reptiles; Mission Scientifique au Mexique et dans l'Amer-

ique Centrale, livr. 12, 1890, p. 732, pi. 43, fig. 5.

One very large specimen (No. 703) was obtained at Isla Bonita.
In preservative the specimen is coal black above, especially on the
latter two thirds of the body and tail ( probably olive in life ) . The
lower part of the supralabials, the chin and throat are whitish ( prob-
ably yellowish in life ) . The light color of the ventrals begins to be
clouded at the 54th ventral, and at the 80th the ventrals have be-
come entirely dark.

This snake may be differentiated readily from Chironius carinatus
by the scale formula 10-8 (instead of 12-8), and by the fact that
three labials, the fourth, fifth and sixth (instead of the fifth and
sixth only) enter the orbit and the second pair of chinshields are
nearly a half longer than the first pair (instead of being equal or
only slightly larger). The two median scale rows are elevated and
strongly keeled, and the other scale rows are rather lightly keeled,
except the outer which is smooth.

Some of the scale characters of this specimen are: rostral visible
above as a narrow line; internasals very large, their length only a
little less than that of the prefrontals; frontal distinctly shorter than
its distance to the tip of the snout, and at least one-third shorter
than the short parietals; length of latter less than their distance from
the internasals; nasal not completely divided; loreal not noticeably
longer than high; one very large preocular not reaching frontal; two
postoculars; temporals, 2-2; supralabials, 9-9, fourth to sixth border-
ing orbit; infralabials 10-10, five touching the anterior chinshields;
ventrals 161; subcaudals 127 +; anal divided. Single pits are
present on a few scales on anterior part of body.

Length 2164 + mm., of which the tail is 750 + mm. (probably
not more than one or two centimeters missing). The stomach con-
tained a large specimen of Eleutherodactyhis partially digested.

Taylor: Review of Snakes of Costa Rica 97

A specimen in the U. S. National Museum from "Costa Rica" ( No.
14061 ) has 162 ventrals, 128 subcaudals. The yellow color of venter
extends to the 75th ventral.

Chironius melas (Cope)

Herpetodryas melas Cope, Proc. Amer. Philos. Soc, vol. 23, 1886, p. 278 (Nicaragua).
Chironius fuscus (part.) Amaral, Mem. Inst. Butantan, Tome 4, 1929, p. 161 (35).

The characters of this species are as follows:

Scales in ten longitudinal series, all smooth, those of the median
row larger than those of the lateral rows and rather smaller than the
parietal scales; parietals rather short and wide, openly emarginate
behind; nine snpralabials, all longer than high, the fourth, fifth and
sixth entering orbit; nasals well developed; loreal square; one pre-
ocular, two postoculars; temporals 1 -f- 1 -j- 1; muzzle rather short;
eye large, its diameter equal to its distance from the nostril; infra-
labials ten, five touching first chinshield; second pair of chmshields
longer than first; ventrals 158; anal divided; subcaudals 139. Total
length 1210 mm.; tail 470 mm.

Shining black except on the snpralabials and anterior or half of
venter which are cream; the edges of light ventrals are black; here
and there black scales have white edges, especially back of jaw
angle, (data from type specimen).

I have examined a Costa Rican specimen classified as this species
in the U. S. National Museum collected by C. N. Riotte. It agrees
with the type in the characters of squamation save that the anal is
single. Ventrals 163; subcaudals 141. The sides of the frontal
slightly concave; frontal 1/4 to 1/3 longer than its distance from
snout; parietals short, 1/6 longer than the frontal; head dark, the
labials cream; snout somewhat lighter than back of head; anteriorly
a series of transverse lines formed of brown dots; there are 20 of
these in a distance of 5 inches; farther back the dots are larger, the
interspaces narrower. The specimen is discolored to the extent that
one cannot be certain of the posterior coloration. It may be that this
specimen is incorrectly referred here. The anal plate may, however,
represent an anomaly.

Genus Elaphe Fitzinger

Elaphe Fitzinger in Wagler, Descriptiones et icones amphibiorum, pt. 3, 1833, text to, and
pi. 27.

Genotype: Elaphe parreysii = quatuorlineata.
Two species occur in Costa Rica. These may be distinguished
by the following key:

7—3286

98 The University Science Bulletin

Key to the Costa Rican Species of Elaphe

Scales 31-35-21 ; young pale brown with one or two dorsal series of about 50 dark
brown black-edged spots, alternating with smaller lateral spots. Adults uniform
brown, below yellowish triaspis

Scales 27-33-21; a series of large red or chestnut brown spots on each side which
may alternate or join to foim a zigzag band; below, yellowish, usually with
brown spots ftavirufa flavirufa

Elaphe triaspis (Cope I

Coluber triaspis Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 18. 1866, p. 128 (type locality,

Belize = British Honduras).
Elaphe triaspis Amaral, Mem. Inst. Butantan, Tome 4, 1929, p. 158 (33); Smith, Copeia,

No. 3, 1941, p. 136.

Rostral about as wide as high, visible from above; internasals
much shorter than prefrontals; frontal one and one half times as
long as wide, its length equal to its distance from end of snout; loreal
present; preocular one, large, reaching frontal or not; two postocu-
lars; temporals 2 -\- 3, 3 4- 3, or 3 + 4; supralabials eight or nine,
normally eight, the fourth and fifth bordering eye; infralabials 13,
four or five bordering first chinshields, which equal the posterior
pair in length; scale formula, 31-35-21, the posterior rows faintly
keeled. Ventrals 243-282; anal divided; subcaudals 73-119. Eye
relatively small.

Young pale brown above with one or two dorsal series of about
51 large dark brown black-edged spots and an alternating lateral
series of smaller spots; two or three dark brown stripes on occiput
and neck; two curved dark stripes across snout. Adult uniform
brown, lower parts uniform yellowish. Total length more than one
meter.

According to Smith, ( he. cit. ) a specimen from Costa Rica
( U.S.N. M. No. 9777), is one of the paratypes of Coluber mutabilis
Cope. The specimen has 259 ventrals; subcaudals 114; scale for-
mula 29-31-25-21, with low keels on the scales of the posterior part
of the body; temporals 4 -f- 5; supralabials 8-8. The specimen
measures 1080 mm. in total length; the tail, 247 mm.

Elaphe flavirufa flavirufa (Cope)

Coluber flavirufus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 18, 1866 (1867), p. 319 (type

locality, Yucatan).
Elaphe flavirufa Viquez, Nuestros Animales venenosas, 1941, p. 65 (Costa Rica).
Elaphe flavirufa flavirufa Smith, Copeia, 1941, no. 3, p. 132, fig. 2.

Rostral broader than deep, just visible above; internasals shorter
than prefrontals, frontal one and one half to one and one third
times as wide as long, nearly equal to its distance from snout tip,
shorter than parietals; loreal present, longer than high; preocular

Taylor: Review of Snakes of Costa Rica 99

one, large ( may be divided ) ; postoculars two; temporals 2 -|- 3,
3 + 3, or 3 + 4; eight or nine supralabials, fourth and fifth, or some-
times also sixth, enter orbit; infralabials, 13-13, four or five touching
the anterior chinshields; scale formula 27-33-21; the dorsals faintly
keeled; ventrals 252-266; subcaudals 102-119; anal divided.

Yellowish or pale brown above, with a dorsal series of about 51
reddish or chestnut-brown black-edged transverse spots which may
alternate or be confluent into a zigzag band; a lateral series of spots
alternating with the larger dorsal series; two curved bands across
snout, and some symmetrical marks on the crown and occiput; two
longitudinal bands on the occiput and nape, sometimes confluent
in front and behind; lower surface yellowish with or without small
brown spots.

Total length, 1220 mm.; tail 260 mm.

I have not examined any Costa Rican specimens of this species.

Genus Drymarchon Fitzinger

Drymarchon, Fitzinger, Systema Reptilium, 1943, p. 26.

Genotype: Coluber cordis Daudin.

Only a single species of this genus of very large snakes is known
in Costa Rica.

Drymarchon corais melanurus (Dumeril, Bibron and Dumeril)

Spilotes melamtnis Dumeril, Bibron and Dumeril, Erpetologie Generale, vol. 7. pt. 1, 1854,

pp. 224-225 (type locality, Mexico).
Drymar.-hon corais ?nelanurus Ruthven. Misc. Publ. Mus. Zool., Univ. Michigan, no. 8, 1922,

p. 65.

In Costa Rica this species is second in size to Constrictor constric-
tor imperator. The specimen at hand, M. N. H. No. 25755, from
2/2 km. west of Turrialba, measures 2634 mm. (8 ft. 8 in.); tail
410 mm.

Rostral visible above; internasals large, their common suture two
thirds that of prefrontals; frontal small, as broad as long, about
equal to its distance to middle of the internasals or a little longer;
supraoculars two thirds width of frontal and about one sixth longer;
parietals one third longer than wide, their length equal to their dis-
tance from the middle of the internasals; nostril large between two
nasals, the posterior much the larger; loreal a little longer than
high; preocular large, much higher than wide, widely separated

1

from the frontal; postoculars two; temporals 2 -| ; supralabials

2

8-8, the fourth and fifth border eye, the fifth and seventh touch

100 The University Science Bulletin

above the smaller triangular sixth; infralabials 8-8, four touch the
first chinshields, which are larger and longer than second pair.

Color above drab brown, many scales having cream edges; some
dim grayish markings on outer part of many of the scales but not
bordering the edges. Beginning about midway of the body, the
median rows develop small gray-black flecks; these grow more
numerous posteriorly so that latter fifth of body and tail shining
black; on sides one can discern an indefinite black coloration on
skin between scales, occasionally also visible on edges of some
scales. Head above uniformly colored like anterior part of body,
but the fourth to seventh supralabials with a black posterior border;
there is a pair of diagonal stripes on neck, joining a black ventral
scale; ventral surface flesh color, the outer posterior edges of ven-
trals bearing a narrow black stripe or spot, and on outer edges
( about 1/7 of width ) the body color encroaches on each ventral;
this is sharply delineated from ventral coloration. After middle of
body is passed ventrals become gradually darker until the latter
fifth of body is shining coal black.

I have observed a Costa Rican specimen in the U. S. National
Museum (No. 61947), without specific locality.

Genus Masticophis Baird and Girard

Masticophis Baird and Girard, Catalogue of North American Reptiles . . . , 1853, p. 98.

Genotype: Masticophis ornatus Baird and Girard.
One species is known from Costa Rica.

Masticophis mentovarius (Dumeril, Bibron and Dumeril)

Coryphodon mentovarius Dumeril, Bibron and Dumeril, Erpetologie Generate .... 1854, vol.

7, pt. 1, p. 187 (type locality, between Coban and Clusec, Guatemala).
Masticophis mentovarius Ortenburger, Occ. Papers Mus. Zool., Univ. Michigan, no. 139, p. 2;

Dunn, Copeia, 1933, p. 214 (La Palma, Costa Rica); Smith, Copeia, 1942, no. 2, pp. 85-88.
Coluber vientovarius Brongersma, Studies on the Fauna of Curasao, Arubu, Bonaire and the

Venezuelan Islands, vol. 2, 1940, pp. 6-7.

A specimen purporting to be this species has been reported from
La Palma, Costa Rica. Smith believes it is confined to the drier
western part of Central America.

Smith (loc. cit.), has pointed out that the form occurring in
Venezuela and Colombia is to be regarded as subspecifically dis-
tinct from the northern one, on the basis of juvenile coloration, and
revives Peter's name suborbitalis. Until juvenile specimens are col-
lected in Costa Rica, one cannot be certain whether the northern
mentovarius mentovarius, or mentovarius suborbitalis is the form
that occurs.

Taylor: Review of Snakes of Costa Rica 101

The species may be distinguished by the following synopsis:
scale formula 17-15-13; ventrals 189-208; subcaudals 111-119; seven
supralabials ( the normal fourth and fifth labials of the other species
are fused below eye to bring about the reduction to seven); infra-
labials ten, rarely eleven; two preoculars, lower very small; two
postoculars, temporals 2 + 2 + 2 usually; anterior chinshields wider
and shorter than posterior; loreal a little longer than high.

Dark brown to brown-gray; sides of neck may show some trace
of longitudinal stripes on rows one and two. Posteriorly the body
becomes lighter, and in some specimens almost every scale has one
or more elongate small gray marks; head uniform dark brown, with
dark markings on the side of the head; infralabials immaculate or
covered with large blotches; anterior belly marks vary from small
gray dots to large black blotches; posteriorly the belly is usually im-
maculate cream.

Genus Leptodrymus Amaral

heptodrymus Amaral, Bull. Antiv. Inst. Amer., vol. 1, 1!>27. p. 29.

Genotype: Salvadora grahamiae Baird and Girard.
A single species is known from Costa Rica. The genus ranges
from southern United States to Costa Rica.

Leptodrymus pulcherrimus (Cope)

Mast ico phis pulcherrimus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 26, 1874, p. 65

(western side of Central America).
Leptodrymus pulcherrimus Bogert, Amer. Mils. Novitates, no. 1352, 1947, pp. 1-14.

Rostral broadly visible above; internasals about as long as wide,
as long as prefrontals; frontal longer than its distance from the
snout tip, shorter than the parietals; nasal divided, the nostril chiefly
in the posterior part; loreal twice as long as high; one large pre-
ocular, not reaching frontal; three postoculars, subequal; temporals
2 + 1 + 1 + 2 (2 + 1 + 2 + 2); supralabials 9-10, the fifth and
sixth entering the orbit; infralabials 10-11, five (or six) bordering
first chinshields, which are a little shorter than second pair.

Scale formula: 17-17-15, with paired pits; scales of median dorsal
row smallest; ventrals 203, subcaudals 126+ (tip missing).

Top of head olive; ground color gray on sides; two black stripes
begin on snout, pass through eye and on body to tail; chin and
labials cream; venter cream.

The ventrals throughout the known range of the species vary from
199 to 210; subcaudals vary from 148 to 152.

102 The University Science Bulletin

I have examined a Costa Rican specimen in the U. S. National
Museum. Bogert, loc. cit., lists a specimen from Pacaca, Costa Rica.
These records are the southernmost known for the species.

Genus Leimadophis Fitzinger

Leimadophis Fitzinger, Systems Reptilium, 1843, p. 26.

Genotype: Natrix almadensis Wagler.
One species occurs in Costa Rica.

Leimadophis taeniurus juvenalis Dunn

Leimadophis taeniurus juvenalis Dunn, Copeia, 1937, no. 4, p. 213 (type locality, San Jose,

Costa Rica).
? Liophis (Rhadinaea) cobella Wettstein, Sitzber. Akad. Wiss. Wien, vol. 143, 1934, p. 341.

This is a common snake in Costa Rica. Six specimens were taken,
the series including both young and adult. The snake when annoyed
flattens its body greatly, thus spreading the scales and exposing
the bright red color on the skin and scale edges between the black
bands. The red color is largely concealed when the body is not
distended. Sometimes the snakes rear the head in cobralike fashion
and attack menacingly.

The general coloration of the snake is black dorsally. Thirty-
five bands present on the body, separated by lighter bands, in which
the skin between the scales, as well as the concealed edges of most
of the scales, is red; head black, labials yellow; below, chin and
anterior part of neck creamy white, the cream washed with orange
anteriorly, but posteriorly venter deep coral to orange red; some-
what triangular black spots on outer sides of ventrals (sometimes
the black spots alternate). These are absent on anterior part of
neck. First row of scales deep olive green anteriorly, grayish post-
eriorly on latter three fourths of body.

There are 23 maxillary teeth, followed after an interspace by two
enlarged fangs. One of the specimens, No. 699, contained in its
stomach a partly digested Eleuthcrodactylus of an undetermined
species.

Dunn, loc. cit., has identified Wettstein's specimens of Liophis
(Rhadinaea) cobella from Bebedero and Irazu as probably be-
longing in this species.

Table of Data on Leimadophis taeniurus juvenalis

Sub-

Supra -

Infra -

Scale

Museum

No.

Sex

Ventrals

caudals

labials

labials

formula

R.C.T.

651

(vs.)

140%

54

8-8

10-10

17-17-15

R.C.T.

699

9

145

62

8-8

10-10

17-17-15

R.C.T.

729

(yg.)

146

55

8-8

10-10

17-17-15

M.N.H.

25750

6

142

52

8-8

10-10

17-17-15

M.N.H.

25751

9

146%

54

8-9

10-10

17-17-15

M.N.H.

25752

6

145

53

8-8

10-10

17-17-15

Taylor: Review of Snakes of Costa Rica 103

Genus Lampropeltis Fitzinger

Lampropeltis Fitzinger, Systema Reptilium, 1843, p. 25.

Genotype: Herpetodryas getulus Schlegel.

There is a possibility that three separable forms occur in Costa
Rica. These are Lampropeltis doliato polyzona, L. d. micropholis
and L. d. gaigae. The first is said to intergrade with the second, and
if true, both forms may likewise occur in typical state. The status
of the third may be questioned since the young seemingly cannot
be told certainly from the young of the other subspecies. The adults,
which often reach a length of six feet, may become black, losing
most, if not all, of the color markings.

Key to the Subspecies of Lampropeltis doliata

*

1 . Adults black ; dorsal scale rows usually 19, rarely 21 gaigae

Adults ringed, not black; dorsal scales rare'y 19, almost always 21 or 23 2

2. Dorsal scale rows 21 or 23 in both sexes; caudals average 48 or more; usually two
anterior temporals; light areas between black rings narrower; snout black with

a light cross-bar polyzona

Dorsal scales rarely 19, usually 21 in males, never 23 in males, 21 to 23 in fe-
males; caudals average 46 or less, females average under 44; usually one anterior
tempoial ; light areas broader ; snout spotted micropholis

Lampropeltis doliata polyzona Cope

Lampropeltis polyzona Cope, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 258 (Quatupe, near
Jalapa, Mexico); Blanchard, U. S. Nat. Mus. Bull., no. 114. 1921, pp. 139-149.

Lampropeltis triangulum polyzona Dunn, Occ. Papers Mus. Zool.. Univ. Michigan, no. 353,
Apr. 28, 1937, pp. 1-11.

This form is a black, yellow ( cream ) and red snake belonging to
the harmless "coralillos." The bands are arranged in triads, black-
yellow - black separated by interspaces of red ( usually black
dotted); the number of these triads varies tremendously in num-
ber, from 17 to 37 as understood by Blanchard, Joe. cit. ( average of
25 fide Stuart, Occ. Papers Mus. Zool., Univ. Michigan, no. 309,
March, 1935, pp. 1-6). The head is black to near the end of the
parietals and there is usually a yellow band across the snout. The
anterior temporals are normally 2-2. A curving yellow band is
continuous across back of head covering back part of parietals and
then passing under chin. The ventrals of southern (Nicaraguan)
specimens vary between 223-231 $ and 228-234 $ . (Data largely
from Dunn, 1937. )

* Data from Dunn, Occ. Papers Mus. Zool., Univ. Michigan, No. 353, Apr. 2S, 1937, pp.
1-11.

104 The University Science Bulletin

[Lampropeltis doliata gaigae Dunn]

Lampropeltis triangulum yaigae Dunn, Occ. Papers Mus. Zool., Univ. Michigan, no. 353. Apr.
28, 1IK-S7, pp. 9-11 (Boquete, Chiriqui, Panama: also reported from Volcan Barha. Monte
Redondo, Volcan Irazii and Paloma. The subspecies name is probably intended for gaigeae
and presumably intended to honor Mrs. Helen Thompson Gaige, since the type is said to
have been collected by "the Gaiges").

This color form has been named from certain Panamanian and
Costa Rican specimens, all coming from considerable elevation
(3900-6500 ft.). The young are brilliantly colored like lowland
specimens, but the adults are black. I suspect it would be impos-
sible to separate the forms save on fully adult specimens. Pre-
sumably the number 19 for scale rows is more common than 21 or
23. From the data offered, the scale characters have the same
variations as the presumed intergrades of mieropholis and poly-
zona. It is conceivable that one or both forms listed as belonging
to mieropholis may belong here.

Lampropeltis doliata mieropholis Cope

Plate IX

Lampropeltis mieropholis Cope, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 257 (Panama).
Coronella doliata formosa (part.) Jan. Iconographie Generale des Ophidiens, hvr. 14, Dec.

1861, pi. 4, "fig. B*" (composite, but fig. B chosen as type. See following reference).
Lampropeltis mieropholis Blanchard, U. S. Nat. Mus. Bull. No. 114, 1921, pp. 149-152.
Lampropeltis triangulum mieropholis Dunn, Occ. Papers Mus. Zool., Univ. Michigan, no. 353,

Apr. 28. 1937, pp. 1-11.

Dunn, loc. cit., has suggested that the form Lampropeltis doliata
mieropholis intergrades with L. d. polyzona to the north. Of
the two specimens of Lampropeltis available in the collection, one
R.C.T. No. 1026 was taken from the Pacific slope of Cerro de la
Muerte at an elevation of approximately 5500 ft., and the other
from Isla Bonita (M.N.H. No. 25164) on the eastern slope of
Volcan Poas, at an elevation of approximately 5500 ft.

The general characters of these specimens are as follows (where
data vary, the first is No. 1026, the second, No. 25164 ) : rostral large,
the length of the part visible above greater than length of intemasals;
common suture of latter scales as long as that of prefrontals; frontal
very broad, equal or a fifth larger than the distance to tip of snout;
the parietals equal to their distance from middle of intemasals;
nasal divided; loreal trapezoidal, longer than high; a large pre-
ocular separated from frontal; two postoculars; temporals 1+2
(1+3), but with a tiny intercalated scale bordering the parietal on
left side; supralabials 7-7, third and fourth bordering orbit; infra-
labials 9-9, four touching anterior chinshields, which are nearly

Taylor: Review of Snakes of Costa Rica

105

Plate IX. Lampropeltis doliata micropholis Cope, RCT No. 1026, Pa-
cific slope of the Cerro de la Mueite, Costa Rica; approximate elevation
5500 feet. About natural size.

106 The University Science Bulletin

double area of second pair. Scale rows 21-19-19-15(14), (21-19-
19-17), all smooth with widely separated pits; ventrals 231, sub-
caudals 48 (230, 51).

Color pattern of No. 1026 is shown in Plate No. 9. The triads
(black-yellow-black) are 19, separated by red areas of greater
length than the triad except for the last five; head black with a nar-
row light band across snout (on prefrontals and internasals) and a
narrow curved light band on back of head, bordered by a similar
black band and involving back ends of parietals; the light band
curving forward to near eye and across throat and chin where it
widens; one postocular with a white area, as is true of the preocular;
first four supralabials black or with a black area; five anterior infra-
labials same; seven white rings on tail, only first two or three forming
triads; red absent on posterior part of tail; red scales with dark spots
indicated; venter with red and white areas immaculate.

In No. 25164 the curving band is indicated by a cream spot on
back ends of parietals and adjoining body scales, but black of head
joined, by a broad bridge, with the curving nuchal black band; the
whitish area on the fifth and sixth supralabials, and those across
chin, have some darker markings. There are 17 triads, the inter-
vening red areas always greater than length of combined parts of
triads; ventral red and yellow areas with much black; dorsal scales
of red bands with discrete black spots on anterior tip of scales; on
white (yellow) bands, spots are larger and irregular, almost ob-
scuring the band.

I have referred these forms to micropholis on the basis of the
reduced number of triads, the single anterior temporal and the wide
bands of red.

Genus Pliocercus Cope

Pliocercus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, 1860, p. 253.

Genotype: Pliocercus elapoicles Cope.
One species is recorded from Costa Rica.

Key to Species of Pliocercus ix Costa Rica

Rings on body fewer, 14, on tail S: ventrals 127, subcaudals 120, 3 preoeulars

dimidiatus
Rings on body more numerous, 22'/2, on tail approximately 11; ventrals 137,
subcaudals ( ?) ; 2 preoeulars annellatus

Pliocercus dimidiatus Cope

Pliocercus dimidiatus Cope, Proc. Acad. Nat. Sci. Philadelphia, 1865, p. 190 (type locality,
Arriba, Costa Rica) and Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol 8, 1876 (1875),
p. 138; U. S. Nat. Mus. Bull. no. 32, 1887, p. 79.

Elapochrus dimidiatus Giinther, Biologia Centrali-Americana, Oct. 1893, p. 107 (Costa Rica).

Taylor: Review of Snakes of Costa Rica 107

Urotheca elapoides (part.) Boulenger. Catalogue of the Snakes of the British Museum, 2nd ed.,

1894, pp. 182-183; Werner, Zool. Jahrb., Bd. 57, 1929, p. 122.
Urotheca dimidiata Gaige, Hartweg, and Stuart, Occ. Papers Mus. Zool., Univ. Michigan, no.

357, 1937, p. 16.
I'iiocercus euryzona dimidiatus Dunn and Bailey, Bull. Mus. Comp. Zool. Harvard Coll., vol.

86, no. 1, 1939, pp. 11-12.

The type of this black and red banded snake has the following
characters: Top of rostral round, curved back on upper surface;
internasals small, about one half the length of the prefrontals; lateral
borders on the frontal nearly parallel, the scale pentagonal; parietals
large; temporals 1 + 1; loreal nearly a rhomb; three preoculars, the
lower cut from a labial; two postoculars; supralabials nine, the fifth
and sixth enter orbit; nine infralabials; chinshields equal; scales in
17-17 rows; ventrals 127, subcaudals 120.

Red above crossed by 14 black rings on body and eight and part
of another on tail; these separated by nearly equal spaces below,
and rather narrower spaces above ( 3/2 scales ) ; a black nuchal band
8 scales wide involving posterior part of the parietals; a red band
across back of head; anterior part of head black, the first two labials
whitish.

I have examined the type specimen.

[Urotheca lateristriga (Berthold), now placed in the genus Rhad-
inaea has been reported from Costa Rica but the reports were pos-
sibly based on species other than lateristriga.]

Pliocercus annellatus sp. nov.

Plate X. text fig. 4

Typc—U. K. M. N. H. No. 25370; Morehead Finca, five miles south-
west of Turrialba, Costa Rica. E. H. Taylor, collector; 1947.

Diagnosis — Related to Pliocercus dimidiatus but having 22M body
bands instead of 14, two instead of three preoculars; eight supra-
labials, the fourth and fifth entering orbit, instead of nine with the
fifth and sixth entering orbit.

Description of type specimen— Rostral low, visible above as little
more than a line; internasals small, wider than long, their common
suture half that between prefrontals; latter scales longer than wide,
with a distinct rounded canthus, the snout rather narrowed; frontal
about one-fourth longer than wide, its length equal to its distance
from the rostral, the anterior border straight, the sides almost paral-
lel; parietals longer than frontal, their length a little less than dis-
tance to middle of internasals; nasal divided; loreal somewhat
trapezoidal; two preoculars, the upper large, but not touching

108

The University Science Bulletin

frontal, the lower small, wedged in between the third and fourth
supralabials; two subequal postoculars; supraoculars slender; tem-
porals 1 + 1 + 2(1 + 2 + 2); supralabials 8-8, the fourth and fifth
border orbit; 9-9 infralabials, five touching the first pair of chin-
shields, which are as long or slightly longer than second pair; scales
smooth, lacking apical pits, in 17-17-17 rows; ventrals 137, the first
separated from chinshields by one pair of scales; subcaudals 73 +
with a portion of the tail missing.

Fig. 4. Pliocercus annellatus sp. nov. Type. MXH Xo. 25370. Morehead
Finca, 5 miles SW Turrialba, Costa Rica. A. Head, dorsal view; B, Head,
lateral view; C, Head, ventral view.

Dorsallv banded red and black; the black bands 22/2 in number
(including nuchal), five or six scale-lengths long, the red bands two
or two and a half scales. Color white or cream-white on the neck
and chin, gradually becoming tomato-red in the middle and posterior
parts of body; the black bands occupy three or four ventrals, the
red two or three; tail partly missing but 6/2 bands present (three
or four more bands probably present on the part missing); caudal
black bands a little longer than those on body; head deep black to
level of middle of parietals except for a white spot on first two supra-
labials; white band crosses back of head not reaching the posterior
tip of parietals; chin and lower lip cream save for first four infra-
labials, anterior tips of first chinshields, and parts of the mental,
which are black; the nuchal black band interrupted mesially.

Measurements in mm.: Total length, part of tail missing, 699;
snout to vent 464; length of head 23; width of head, 14.

Taylor: Review of Snakes of Costa Rica

109

Remarks: The other form of this genus that might be expected
to occur in costa Rica is Pliocercus euryzonus euryzonus. That form
may be readily separated from annellatus by the pattern of the black
bands which on the dorsal surface leaves bands of red only one, or
more frequently, only one-half scale length wide; the band across
occiput usually is of the same width. The type description of this
form states that the snout is broadened at the muzzle; the temporals

+m*

Plate X. Pliocercus annellatus sp. nov., MNH No. 25370. Morehead Finca,
5 miles SW Turrialba, Costa Rica; total length. 699 mm.

110 The University Science Bulletin

are said to be "one large and four small," the supralabials are nine,
the fifth and sixth entering orbit; 10 infralabials present; the num-
ber of bands on body is 19; on tail, 11.

It would appear that Pliocercus annellatus represents a species
intermediate between dimidiatus and euryzonus. It is conceivable
that it does bear a subspecific relationship to one or the other of the
two, but this is surmise.

Genus Liophis Wagler

Liophis Wagler, Natiirliches System der Amphibien, 1830, p. 187.

Genotype: Coluber cobella Linnaeus.

A single species of this genus has been reported in Costa Rica.
The genus is South American and there may be some doubt that
the records here presented are authentic.

Liophis cobella (Linnaeus)

Coluber cobella Linnaeus, Museum Adolphi Friderici regis, 1754, p. 24. and Systema Naturae,

vol. 1, 1766, p. 378 (type locality, Asia [in errorl ).
Rhadinaea cobella Boulenger, Catalogue of the Snakes in the British Museum, vol. 2, 1894,

pp. 166-167. (South American localities.)
Liophis cobella Peters, Monatsb. Akad. Wiss. Berlin, 1859, p. 276 (Costa Rica).
t Liophis (= Rhadinaea) cdbella Wettstein, Sitz. Akad. Wiss. Wien, Math.-naturw. Klasse,

Abt. 1, Band 143, Heft V2, 1931, p. 34 (Costa Rica); Dunn, Copeia, 1937, no. 4, p. 213.

The reference by Wettstein ( loc. cit. ) of two specimens, one
adult from Volcan Irazu, 2400 m., and a half grown one from Bebe-
dero, to the species Liophis (Rhadinaea) cobella, is I believe the
second record of the species for Costa Rica. That of Peters, loc. cit.,
is the first. Wettstein states that he has compared the specimens
with numerous specimens of cobella in the Vienna Museum.

The specimens are bright gray brown, with bands of black. The
outer scale edges bear light streaks especially on the anterior part
of the body. The venter is bright orange (perhaps red in life); the
ventrals are 149-147; the subcaudals 58.

Boulenger, op. cit., gives the following characters: rostral broader
than deep, visible from above; intern asal as long as broad or a little
broader, approximately as long as the prefronals; frontal once and
a half to once and three fourths as long as broad, about as long as
its distance from end of snout, a little shorter than parietals; loreal
as long as high or a little longer; preocular one, postoculars two;
temporals 1+2; supralabials eight, fourth and fifth border eye; five
infralabials touch first chinshields which are as long as or a little
longer than second pair. Scales in 17 rows. Ventrals 143-163; anal
divided; subcaudals 45-57. Tail 4% to 6 times in total length.

Taylor: Review of Snakes of Costa Rica 111

Brown or blackish above with whitish lines and crossbands; belly
yellow ( coral red in life ) with transverse black spots or crossbands,
always more or less marked in young, which have a blackish nuchal
collar edged with whitish, and a pair of whitish dots on parietal
shields; upper labials yellowish.

Genus Rhadinaea Cope

Rhadinaea Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 15, 18G3, p. 100.

Genotype: Taeniophis vermiculaticeps Cope.

This genus of small terrestrial snakes is represented by a rather
extraordinary number of species, no less than nine having been re-
ported. A species, Rhadinaea guntheri Dunn," a new name for
Tachymenis decipiens Giinther,f is treated in the genus Conio-
phanes, a genus similar to Rhadinaea in many ways, but having
grooved posterior teeth. I suspect that the name should be sup-
pressed since Tachymenis decipiens and Rhadinaea decipiens are
probably not congeneric. If not a member of the genus Coniophanes,
it might well be treated under a generic name of its own. Refer-
ences of Rhadinaea lateristrigata for Costa Rica are seemingly
based on other species.

The genus Rhadinaea has recently undergone a partial revision
by Dr. J. Bailey (Occ. Papers Mus. Zool., Univ. Michigan, No. 412,
1940).

Synopsis op Costa Rican Species of Rhadinaea

1. Scales in 19 rows; eight supialabials, the 4th and 5th entering the eye; one pre-
ocular; ventrals 164-169; subcaudals 60-78; dark brown with six longitudinal
yellow or cream lines; belly yellowish with black dots on ends of ventrals. .serperastra
Scales in 17 rows 2

2. No narrow discrete continuous light lines on sides of body (discrete line may be

present on neck then replaced merely by indefinite dim lighter lines) 3

One or two narrow discrete light lines on side of body 6

3. Six or seven supralabials, 3rd and 4th enter eye; nasal single; tail not more than
one fourth of total length; one preocular, two postoculars ; the two pairs of chin-
shields of about equal length; ventrals 152, subcaudals 46; dark brown above,
with a narrow black vertebral stripe; two lateral paler stripes; a black stripe along
the ends of the ventrals; labials black bordered; belly yellow, with black crescents

on front of each ventral suture; a black line under tail calliyaster

Eight supralabials, 4th and 5th enter eye 4

4. No continuous line passing through eye and to snout tip; length of tail more
than one third of total length ; two preoculars (rarely one); ventrals 117-137;
subcaudals 90-120; anterior chinshields shorter than posterior; a slightly darker
median stripe, black edged, about three scales wide; a lateral black-edged light
yellow stripe, cream anteriorly, on 4th or 4th and 5th rows; lower sides darker
forming a broad dark stripe; ventrals more or less black-spotted on ends; a black-
edged stripe on parietal and an ocellus on side of neck decorata decorata

Dark lines on head and body continuous, passing thiough eye 5

* Copeia, 1938, no. 4, p. 198.

t Biologia Centrali-Americana, Mar. 1895, p. 163, pi. 53, fig. A.

112 The University Science Bulletin

5. Lower surface of body pinkish, the ventrals finely sprinkled all over with black-
ish ; one preocular, two postoculars ; anterior chinshields shorter than posterior ;
ventrals 132, subcaudals 66; reddish brown above, a black lateral line between
4th and 5th scale rows, edged with yellow on anterior part of body; this line
widens on neck and head passing through eye, joining its fellow on end of snout ;

upper lip white; length, 430 mm. ; tail, 140 mm pulveriventris

Lower surface of body yellowish white, the ends of the ventrals bearing deep brown
dots ; one preocular, two postoculars ; anterior chinshields shorter than posterior ;
ventrals 117, subcaudals 79; yellow-brown above with two deep brown dorsal
streaks, separated by width of one scale, diverging on neck and extending to the
outer posterior angle of supraocular shield ; another dark streak on each side of
head and body passing through eye; head brown vermiculated with yellowish;
total length 330, tail 115 vermiculaticeps

6. Two discrete lateral light lines 7

One discrete light lateral line (may be dim before shedding) ; subcaudals less than
70; one preocular, two postoculars; temporals 1 + 1; eight supralabials, fourth
and fifth entering eye; ventrals 133, subcaudals 50; total length 335 mm.; tail

in total length 3% times; scales of outer row with gray tips forming an indistinct
lateral line. Black above, the lower lateral scale rows a shade of rufous ; head
black-brown; labials yellow bordered above by blackish; below yellow, each ven-
tral with black ends pachyura pachyura

7. Two narrow light lateral lines on sides, the lower occupying the middle of first scale
row, the upper, the middle of the fifth ; tail three fourths of body length ; head
narrow, elongate; ventrals 131-151, subcaudals 113; supralabials eight, fourth and
fifth enter orbit; one preocular (rarely a "presubocular") ; cne or two postoculars;
temporals 1 + 1 + 2 : above brownish black, lower parts yellow with the ends of
the ventrals blackish; markings in females fainter; an occipital light band

pachyura decipiens
Blackish dorsally, on belly immaculate red ; a black line on edge of ventrals and
outer edge of first scale row from throat to tail ; immediately above this, a white
line from throat on middle of row one to end of tail ; a white line on fifth scale
row; light ocellae on outer corner of parietals, a larger one behind temporals join-
ing lateral line; total length 440 mm., tail 188; seven or eight supralabials; ven-
trals 137-143. subcaudals 96-112; temporals 1 + 1; one preocular, two post-
oculars (one "presubocular" usually present), 3rd and 4th or 4th and 5th labials
enter orbit ; no occipital light ring persimilis

Rhadinaea serperastra Cope

Rhadinaea serperastra Cope, Proc. Acad. Nat. Sci. Philadelphia, 1871, p. 212 (type locality,
Costa Rica); Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875), p. 140.

Scales in 19 rows, smooth, lacking pits; temporals 1+2+3; nasal
divided, the posterior part larger; supralabials eight, the fourth and
fifth border eye; loreal square; preocular one; postoculars two;
supralabials eight; chinshields subequal in length; ventrals 164;
subcaudals 78; anal divided. Maxillary teeth equal.

Dark brown with six longitudinal yellow or white lines of which
the first and second are brightest; a white line on each side pro-
duced by a series of dark spots on ends of ventrals; labials black
with yellow spots; head dark brown above with a pale shade across
frontal, and two [spots?] just behind the parietals; chin and belly
yellowish (after Cope).

A specimen in Harvard M.C.Z. (No. 28069) is from San Jose,
Costa Rica. There are present 170 ventrals and 65 subcaudals.

Taylor: Review of Snakes of Costa Rica 113

Other characters are very similar to those of the type. There is a
whitish line from eye to near jaw angle. The arrangement of the
lines is as follows: a median dark line; three scale rows on each
side of middle, each with a dim darker line; a discrete white
line on the fifth and sixth rows; a broader lateral black stripe on
third, fourth and fifth rows with a fine white line through the middle;
light lines on first, second and third rows, that on first scale row
bordered above and below by darker lines; each labial has a white
spot.

Rhadinaea calligaster (Cope)

Contia calligaster Cope, Journ. Acad. Nat. Sci. Philadelphia, vol. 8. 1876 (1875), p. 146 (type
locality. Pico Blanco, southern Costa Rica).

Scales acuminate in 17 rows, smooth, lacking apical pits; supra-
labials seven, third and fourth border the orbit; nasal single; loreal
subquadrate; preocular one; postoculars two; temporals 1 + 1 -(- 2
( 1 + 2 -f 2 ) ; infralabials eight, fourth and fifth largest, the first
pair barely in contact behind mental; ventrals 152, anal divided, sub-
caudals 46; chinshields equal. Maxillary teeth gradually increasing
in length to the posterior.

Color above dark brown with a narrow vertebral black band; two
lateral paler bands, the lower on the first and second scale rows, the
upper on the fourth and fifth, the space between black. A black
band along the edge of ventrals; belly yellow, each ventral with a
black crescent or with a black base; labials black-bordered.

Known from Pico Blanco, Costa Rica; the exact elevation on the
mountain unknown (5000-7000 ft).

The types (U.S.N.M. Nos. 30606-30607) at the present time are
in a poor state. Another specimen in the U. S. National Museum
(No. 38335) has 147 ventrals, 60 subcaudals. The general color
is violet-brown.

Rhadinaea decorata decorata (Giinther)

Plate XI

C crone1 la decorata Giinther, Catalogue of the Colubrine Snakes in the Collection of the British

Museum, 1858, pp. 35-36 (type locality, "Mexico").
Rhadinaea decorata Cope, Proe. Amer. Philos. Soc, vol. 22, 1885, p. 381.
Rhadinaea decorata decorata Bailey, Oce. Papers Mus. Zool. Univ. Michigan No. 412, May 6,

1940, pp. 7-8, plate 2, fig. 5 (Costa Rican specimen).

A specimen of this species was captured in the mountains one
mile west of La Suiza at about 2100 ft. elev. It was found coiled

8—3286

114

The University Science Bulletin

Plate XI. Rhadinaea decorata decorata, MNH No. 25733, La Suiza,
Costa Rica, approximate elevation 2400 feet. Total length, 421 mm.

Taylor: Review of Snakes of Costa Rica 115

under a small flat rock in a gully. The scale characters of this speci-
men are as follows:

Eye moderately large; the head distinct from neck; rostral dis-
tinctly broader than high, the scale scarcely visible from above;
length of internasals in that of prefrontals about one and one third
times; width of frontal at least three fifth of its length; length of
frontal a third longer than its distance from the end of snout; pari-
etals elongate, equal to their distance from the tip of snout; nasal
divided, the posterior part much the largest; loreal nearly square;
one preocular ( partially divided on both sides ) and a small "presub-
ocular" below preocular; two postoculars, only one touching the
temporal; formula for temporals, 1 + 2 + 3; supralabials 8-8; the
fourth and fifth entering orbit; infralabials 10-10, five touching
chinshields; first pair of chinshields about one sixth shorter than the
second pair; latter in contact for nearly half their length; scales
without apical pits, finely striated but not keeled. Ventrals 126;
subcaudals 94 -f-; anal divided; total length, 421 mm.; tail, 160 mm.

Top and upper part of the sides of head brownish cream with
minute blackish flecks, the bones semitransparent so that the brain
can be seen dimly. Lips and chin white. A black line borders
the upper labials to near the jaw angle; an elongate black-edged
light stripe on outer border of parietal, and another rounded ocellus-
like spot on beginning of neck. Middle scale row and two adjoining
rows blackish gray beginning on neck; a light cream stripe covering
sixth row and parts of adjoining rows ( more posteriorly, all or nearly
all of these rows). Below this there is a broad, dark gray line ex-
tending onto the outer edge of ventrals; this color on ventrals termi-
nates in a black dot on anterior ventrals, which may be dim or
almost indistinguishable posteriorly. Anteriorly the first scale row
may bear a narrow light line. Below yellowish about neck, chang-
ing to orange then to tomato-red on the remainder of body.

In the collection of the U. S. National Museum, I examined six
specimens from "Costa Rica", one from Suretka, C. R. The ventral-
subcaudal counts on No. 9788 are 120, 88 +; Nos. 30609-30612 have
respectively 116, 72 -f; 117, ?; 123, 96; 114, ?. The Suretka speci-
men has 119 ventrals, and 110 subcaudals.

These agree in all essential characters with the specimen de-
scribed except that sometimes there is a small light spot in front of
the upper eye level; sometimes this is absent.

Specimens in Harvard M.C.Z. represent the following localities:
Guapiles, Suretka, St. Cecilia and Limon.

116 The University Science Bulletin

The species has a wide distribution throughout Central America
and southern Mexico.

Rhadinaea pulveriventris Boulenger

Rhadinaea pulveriventris Boulenger, Catalogue of the Snakes in the British Museum, vol. 3
(addenda and corrigenda), 1896, pp. 635-636 (type locality, Costa Rica).

Liophis (— Rhadinaea) pulveriventris Wettstein, Sitz. Akad. Wiss. Wien, Math.-naturw.
Klasse, Abt. 1, Bd. 143, Heft 1-2, 1934, pp. 34-35 (Volcan Irazii, 1500 m.).

Eye rather small; rostral just visible from above; internasals
broader than long, much shorter than the prefrontals; frontal longer
than its distance to end of snout, shorter than parietals; loreal longer
than deep; one preocular; two postoculars; temporals 1 + 2; supra-
labials 8-8, fourth and fifth entering orbit; five infralabials touch an-
terior chinshields, which are shorter than posterior; scales in 17
rows; ventrals 132; anal divided; subcaudals 66.

Reddish brown above; a black lateral line between the fourth and
fifth scale row, edged with yellowish on the anterior part of body;
these lines widen on neck and head and passing through eye join
each other on the end of the snout; upper lip white; belly pinkish
yellow, finely speckled all over with blackish.

Wettstein, loc. cit., reports a specimen from Costa Rica. It agrees
with the type in essential characters. The following data are from
Wettstein's report: Maxillary teeth, 18 + 2; mandibular, 20-21;
scale rows, 17; ventrals 149; subcaudals 55; supralabials 8, the 4th
and 5th entering eye; first chinshields shorter than second pair, and
in contact with four infralabials (5 in type); one preocular; two
postoculars; temporals 1 + 2; total length, 444 mm.; tail, 106 mm.

A specimen in Harvard, M.C.Z. No. 15261, from Navarro, Costa
Rica, has 129 ventrals and 67 subcaudals; the scale-row formula is
17-17-15. In this specimen the venter is almost immaculate. The
pigment flecks are reduced, there being very few along the edges
of ventrals. Total length, 500 mm.; tail, 145. In other essential
characters the specimen agrees with the type.

Rhadinaea verrriiculaticeps I Cope)

Taeniophis vermiculaticeps Cope, Pioc. Acad. Nat. Sci. Philadelphia, 1860, p. 249 (type local-
ity, Veragua, Panama).
Rhadinaea vermiculaticeps Cope, Proc. Acad. Nat. Sci. Philadelphia, 1863, p. 101 (Costa Rica).

Scales in 17 rows; one preocular, two postoculars; eight supra-
labials, the fourth and fifth entering eye; frontal elongate; loreal as
long as deep; ventrals 117; anal divided; subcaudals 79.

Yellowish brown on dorsal surfaces with two deep brown dorsal
streaks, separated by the width of one scale, diverging on the neck

Taylor: Review of Snakes of Costa Rica 117

and extending to the outer posterior angle of the supraocular shield;
another dark streak on side of head and body passing through eye;
head generally brown above with vermiculations of yellow; labials
whitish narrowly edged with brown; chin and belly yellowish white,
each ventral with dots on outer borders. Total length 330 mm.;
tail 115 mm.

I find one record ( Cope, loc. cit. ) of the species from Costa Rica
without definite locality.

Harvard M.C.Z. No. 42764 from Piedras Pecora Ridge, Panama,
has a slight peppering of black on the outer edges of ventrals; a
whitish line passing through eye, black bordered. Upper and lower
labials with vermiculate blackish flecks, and black sutures.

Rhadinaea persimilis Dunn

Rhadinaea persimilis Dunn, Copeia, 1938, no. 4, Dec. 10, pp. 197-198 (type locality. La Loma,
1500 ft. altitude. Prov. Bocas del Toro, Panama) (also reported from Cartago and La Suiza,
Costa Rica and Eden Mine. Nicaragua).

Scales in 17 rows; temporals 1 + 1; preocular one, postoculars
two; a small subocular usually present; supralabials seven or eight,
third and fourth, or fourth and fifth entering orbit; infralabials seven
or eight, four, in contact with chinshields. Males 137-143 ventrals;
subcaudals 96-112.

The color above indefinite gray-brown the scales more or less
edged or flecked with black; belly red, immaculate; a black line
along edge of ventrals, and adjoining scale row, bordered above
by a white line in the middle of first scale row; another white line
on middle of the fifth scale row; a light spot on outer anterior corner
of parietals, one on second temporal, and one between temporal and
the lateral line. A median light spot on fifth vertebral scale behind
parietals. Supralabials with black upper borders and faint flecks
on rostral and anterior supralabials.

Tail more than one third of total length. Greatest known total
length, 440 mm.; tail 188 mm.

The species has been reported from La Suiza and Cartago in
Costa Rica.

I have examined the type.

Rhadinaea pachyura pachyura (Cope)

Contia pachyura Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875), p. 145

(type locality, Sipurio, Costa Rica).
Rhadinaea pachyura Dunn, Copeia. No. 4, 1938, p. 198; Notulae Naturae, 1942, No. 108, p. 5.

Scales in 17 rows, smooth and usually lacking pits (sometimes
with one pit ) ; head elongate, the rostral not prominent; intemasals

118 The University Science Bulletin

wider than long; prefrontals as long as wide; frontal elongate but
shorter than parietals; loreal large, higher than long; one preocular;
two postoculars, subequal; temporals 1 + 1; supralabials eight, the
fourth and fifth entering eye; chinshields equal; tail long, three and
three-fifths times in total length, and thickened to near end; ventrals
133; subcaudals 50; total length 335 mm.

Color black, the lower scale row with a rufous shade. Scales
of the first row with gray tips; head and a nuchal area brown, with
a black line from eye above labials; latter yellowish, unspotted or
with spots; belly yellowish, each scale with rather large black spots
on sides.

Of this species, I have examined a specimen from Panama, but
have seen none from Costa Rica. The type is from Sipurio, Costa
Rica. Dunn, op. cit., states that Rhadinaea dedpiens intergrades
with R. pachyura.

Rhadinaea pachyura decipiens (Gunther)

Ahlabes decipiens Gunther, Biologia Centrali- Americana ; Reptilia and Batrachia, 1893, p. 105,

pi. 37, fig. A (type locality. Irazii, Costa Rica).
iUrotheca lateristnaa Boulenger, Catalogue of the Snakes in the British Museum, ed. 2, vol. 2,

1894, pp. 181-18-2 (part.).
Rhadinaea pachyura decipiens Dunn, Copeia, No. 4, 1938. p. 198; Notulae Naturae, No. 108.

p. 4-5 (El Cedral de Navarro, San Jose, and La Palma, Costa Rica).

Scales in 17 rows without pits; temporals, 1 + 1 + 2; preoculars
one or two; postoculars two; usually a small presubocular; supra-
labials eight, fourth and fifth entering eye; four infralabials in con-
tact with the anterior chinshields; loreal present as long as high;
frontal elongate, longer than its distance to end of snout; ventrals
131-151; anal divided; subcaudals 113.

Blackish brown above with two narrow yellow lateral lines, the
lower on outer scale row most distinct; upper line on fifth scale row;
a broad yellow band crosses the temporal region and posterior half
of parietals; lower parts yellow, with the corners of the ventral
scales blackish.

( Subfamily Boiginae )

The following group of snakes, consisting of the genera Trimor-
phodon, Leptodeira, Rhinobotliryum, Imantodes, Oxybelis, Cono-
phis, Clelia, Coniophanes, Erijthrolamprus, Stenorrhina, Scole-
cophis and Tantilla, are characterized by having the posterior
maxillary teeth enlarged and grooved. They are often placed in
the Subfamily Boiginae (Family Boigidae, [auctores]). It may be
regarded as doubtful that a close relationship is indicated by the

Taylor: Review of Snakes of Costa Rica 119

dental character of grooved teeth since the groove varies consider-
ably in its depth as well as its position on the tooth. Characters
other than dentition show differences of such magnitude as to sug-
gest that the grooved posterior fangs have developed several times
in the evolution of the group. Hence the subfamily is only a term
of convenience.

Some of these snakes are known to be mildly poisonous; none
are known to be dangerous to man. The genera of the group like-
wise vary in size, Tantilla and Scolecophis being small snakes, while
Clelia and Trimorphodon reach a length of considerably more than
a meter. Species of Oxijbclis have long sharp-pointed heads while
those of Imantodes have short blunt heads. Both are aboreal in
habit with slender compressed bodies. Trimorphodon and Lepto-
deira are at least partially arboreal. Coniophanes, C onoph i s,
Stenorrhina, Tantilla, Scolecophis, Clelia, Rhinobothryum and
Erythrolamprus are, so far as I know, terrestrial in habit. Species
of Rhinobothryum and Erythrolamprus in general coloration tend to
resemble the poisonous snakes of the genus Micrurus and may some-
times be mistaken for species of this group. The name "Corales"
is applied to all snakes banded black and red, or black, yellow, and
red in Costa Rica.

Key to the "Opisthoolyph" Colubridae of Costa Rica

1. Anal scale entire Clelia

Anal scale divided -

2. Internasal and anterior nasal fused Stenorrhina

Internasal and anterior nasal not fused 3

3. Loreal usually absent I

Loreal present 5

4. Small, short, cylindrical snakes, terrestrial ; the head short, scarcely or not distinct

from body Tantilla

Elongate arboreal snakes with very slender bodies; head distinct from body,
much -elongated (loreal present in one species) Oxybelis

5. Two loreals present (rarely more); 10-11 maxillary teeth, the anterior distinctly
longer than posterior, followed after an interspace by two moderately enlarged

grooved fangs Trimorphodon

A single loreal present ; maxillary teeth variable, the interval present or not be-
tween maxillary teeth and grooved fangs 6

C. Pupil iound 7

Pupil vertically elliptic 8

7. Maxillary teeth 20-25 forming a continuous series the last three to five largest,
grooved; scales 15-17 rows smooth or feebly keeled with apical pits, arranged in

oblique transverse rows ; usually unicolor Oiyh, lis

Maxillary teeth 10, slightly increasing in size posteriorly, followed after an inter-
space by two very large grooved fangs; scales 17-19, smooth, lacking apical pits;
pattern of stripes Conophis

8. Pattern of red, black, and light (white or yellow) bands 9

Pattern banded or spotted (large or small spots) lacking red color in

9. Scales in 15 rows, smooth, without apical pixs; ventrals less than 210; subcaudals
less than 7o ; maxillary teeth subequal, followed after an interspace by feebly
grooved fangs Erythrolamprus

120 The University Science Bulletin

Scales in 21 rows; smooth anteriorly, partly keeled posteriorly; ventrals more
than 200; subcaudals more than 70; maxillary teeth equal, followed after an
interval by two grooved fangs Rhinobothryum

10. Completely banded with rings of black and yellow; scales in 15 rows, smooth,

lacking apical pits Scolecophis

Not banded with rings of black and yellow 11

11. Pattern unicolor or with some darker longitudinal stripes: or a pattern of black,

and cream, stripes; scales in 17-25 rows, smooth, lacking pits Coniophanes

Lacking such patterns 12

12. Body moderately slender; scales in 19-25 rows, smooth or faintly keeled with
apical pits; ventrals less than 215: spots or wide bands of darker color. . . .Leptodeira
Body especially slender and elongated ; vertebral row of scales often enlarged ;
ventrals more than 215: pattern of spots forming blotches or saddles; or forming
numerous small punctations Imantodes

Genus Trimorphodon Cope

Trimorphodon Cope, Proc. Acad. Nat. Sci. Philadelphia, 1861, p. 297.

Genotype: Trimorphodon hjrophanes (Cope).
One species is known from Costa Rica.

Trimorphodon biscutatus biscutatns ( Dumeril, Bibron and Dumeril )

Dipsas biscutata Dumeril, Bibron and Dumeril, Erpetologie Generate, vol. 7, pt. 2, 1854,

pp. 1153-1154 (type locality, Mexico).
Trimorphodon biscutatus Cope, Proc. Acad. Nat. Sci. Philadelphia, 1861, p. 297.

The presence of this species in Costa Rica is attested by two
specimens in the Museum of Comparative Zoology, Harvard Col-
lege, Nos. 16186, Puntarenas, Costa Rica, and 15306, Esparta, Costa
Rica.

The larger of the two specimens has the following characters:
rostral visible above as a broad triangle; internasal suture a little
more than a half the length of the prefronal suture; frontal a fourth
longer than wide, a little shorter than its distance from tip of snout;
three preoculars, the lower small and more or less below eye (sub-
ocular ) ; three postoculars; 3 + 4 temporals, three loreals, the two
upper largest, more or less quadrangular; supralabials 9-9, the fourth
and fifth enter orbit; infralabials 12-13, four or five touching first
chinshields, which are double the size of second pair; scale formula
25-27-19-18, the six median scale rows on latter half of the body
keeled; ventrals 262; subcaudals 85.

Pale grayish or brownish gray, with dark black-edged crossbars
or black-edged markings with a light center. Head with chevron-
like dark bands separated by similar light areas; a series of light
black-edged spots on alternating (or every third) ventrals; these
may connect with the transverse spots of dorsum; anteriorly a
median white stripe 40 mm. long. The specimen is a very large
one, having a total length of 1510 mm., the tail 238 mm.

Taylor: Review of Snakes of Costa Rica 121

Genus Leptodeira Fitzinger

Leptodeira Fitzinger, Systema Reptilium, 1843, p. 27.

Genotype: Coluber annulatus Linnaeus.

Six species of the genus have been reported from Costa Rica.

Key to Species of Leptodeira in Costa Rica

1. Body with 16 or less black bands; scales in 19 rows; body with the black bands
three to four times broader than interspaces of whitish ground color; head black

with a white collar; tail with eight black bands nigrofasciata

Body with 20 or more black bands or spots -

2. Spots on body more than 40 3

Spots on body less than 40 5

3. Body red, banded with black; below salmon color, dusted with brown; a few un-
divided subcaudals follow anus; scales in 23 rows; 61 dorsal black spots. . .rubricata
No red on body, but body usually fawn or light brown; 40-55 black spots or
bands across body. Arboreal forms with somewhat compressed bodies 4

4. Scales of vertebral row (or rows) enlarged; 19-21 scale rows; about 52 small

spots along back; lateral alternating spots dim annulata annulata

Scales of vertebral row not enlarged; scales in 21-23 rows; spots larger, approxi-
mately 38 in number, somewhat quadrangular, usually edged with lighter color
suggesting ocelli ; secondary lateral spots present ocellata

5. Spots less than thirty, terrestrial forms; scale rows 23-25; crescent light spot on
head interrupted medially by a black stripe connecting dark of head with neck

band. One preocular (usually) ; abdomen yellowish white rhombifera

Spots 20, scales in 21 rows; one preocular; one postocular; white below*. . . .maculata

Leptodeira nigrofasciato Giinther

Leptodeira nigrofasciata Giinther, Ann. Mag. Nat. Hist., ser. 4, vol. 1, 1868, p. 425 (type
locality, Nicaragua).

The type description gives the following data on the species:
scales in 19 rows; internasals nearly square, two thirds the size of
the prefrontals; frontal equal to its distance from tip of snout, and
a little shorter than the occipitals; loreal as high as long; one pre-
ocular, not quite reaching the frontal; a minute preocular on one
side; one postocular; eight supralabials, the third, fourth and fifth
border eye; none touching the parietal; temporals 1+2+3; ventrals
174, subcaudals 74.

Body with 16 elongate crossbands, which are from three to four
times longer than the interspaces of whitish ground color, and con-
fluent on posterior part of trunk. Head black above, separated by
a white collar from the first black band; tail with about eight black
bands. Lower parts whitish.

Two specimens have been reported from Costa Rica, one from a
specific locality (Turrialba).

* Characters of the type.

122 The University Science Bulletin

Leptodeira rubricata (Cope)

Sibon septentrionale rubricatum Cope, Proc. Amer. Philos. Soc, vol. 31, pp. 333-347 (type
locality, Boca Mala, Costa Rica).

Snout short, body robust; head with temporal region swollen,
not especially distinct from neck; supralabials eight, the fourth and
fifth enter orbit; two preoculars, two postoculars; scales in 23 rows;
five undivided subcaudals follow vent; ventrals 178.

Above bright red; on venter light salmon, dusted with brown; 61
transversely oval black spots on the back which cover 12 rows of
scales transversely and two and one half to three and one half
longitudinally; small black spots alternate with them on third row
of scales, and a less definite row of smaller spots alternate with
these on second and third rows; a black band extends from eye to
last labial plate and behind and above it a parallel black band ex-
tends from the parietal plate. The extremities of these bands are
fused with the first dorsal spot, the interspaces being red; supra-
labials red with a black spot in the center of each plate; infralabials
black spotted; top of head dark brown bordered posteriorly on out-
line of parietal plates by a red crescent; this bordered posteriorlv bv
a black crescent, and cut by a median black stripe, which connects
the dark brown of the head with the anterior four black dorsal
spots. Tail blackish red, spotted above; total length, 660 mm.; tail
133 mm.

This seemingly is a species of the Pacific lowlands. It would ap-
pear to be easily recognized by the red coloration; however, only
the type is known.

Leptodeira annulata annulata (Linnaeus)

Piute XII

Coluber anmilatus Linnaeus, Amoen. Acad., Tom 1, no. 5. p. 120; Systema Naturae, 10th eil.,
1758, vol. 1, p. 224; Andersson, Bihanj; till K.S.U. vet.-Akad. Handl.. Bd. 21. afd. 4,
1899, p. 21.

Leptodeira annulata annulata Dunn, Proc. Nat. Acad. Sci., vol. 22, 1930, pp. 692-093 (both
coasts; up to 2000 meters elevation at Tierra Blanca, Costa Rica).

The type locality of this form is not known. The six cotypes have
the following scale counts: 5 with 19 rows, one with 21. Ventrals
185-193; subcaudals 80-91.

This subspecies occurs in Costa Rica and differs from the more
northern form, polysticta, in having the scales of the vertebral series
enlarged and in lacking the longitudinal dark mark on the neck.

Three specimens of Leptodeira are referred to this species. They
are M.N.H. No. 25742, taken about five miles southwest of Turrialba;
M.N.H. No. 25743 and R.C.T. No. 1462, taken at Los Diamantes,

Taylor: Review of Snakes of Costa Rica 123

one mile south of Guapiles, Costa Rica. No. 25742 is young and the
markings are more distinct than in the older specimens, in which
some details of the pattern are obscured. Data on markings are
taken from that specimen.

Top of head brown; a pure-white horseshoe-shaped mark begins
behind eye and passes around behind the parietals (encroaching
slightly on the sides of parietals); the width of the mark on the
mid-line equal to from four to five scale lengths; behind this a brown
spot sending forward narrow arms that reach the eye; supralabials
with a light peppering of brown pigment; chin whitish with some
little pigment on anterior part; about 52 spots along middle of back;
on the sides, alternating with the median spots, can be discerned
faint brownish narrow vertical spots; the dorsal ground color is lav-
ender brown, somewhat mottled. Ventrals immaculate save for an
occasional brownish speck on outer edge of scales.

The older specimens have the nuchal white band completely ob-
scured, being pigmented as much as other dorsal areas between
spots on the body.

Characters of head scales not listed in the following table are:
rostral visible above as a narrow line; internasals at least two thirds
the length of prefrontals; latter scales about as wide as long, sepa-
rated from supraoculars by upper preocular; frontal, in contact with
preocular, three fourths as wide as long, its length a little greater
than its distance from end of snout; parietal nearly as wide as long,
equal to its length from internasal-prefrontal suture, or a little
longer; nasal at least partially divided; loreal longer than high; two
preoculars, two postoculars, the upper much the larger in each case.

Three Costa Rican specimens in the U. S. National Museum, col-
lected by William Gabb were examined (Nos. 32565-67). No.
32566 $ has 203 ventrals and 79 + subcaudals; No. 32565 has 197
ventrals and 93 subcaudals; the scale rows are 21 in the middle of
body in both. No. 32567 is in a bad state, and counts were not
attempted.

A Harvard M.C.Z. specimen from Limon, Costa Rica has 197
ventrals, 82 subcaudals; scale formula, 21-23-19-16. While this is a
young specimen, it lacks the large white crescentic area described
in the Turrialba specimen.

Scale Data ox Leptodeira an.nil.ata annulata

No.

Sex

Ventrals

Subcp.udal>

Preocular

Postocular

Supra -
labials

25742

25743

1462

(yg.)

9

201
199
199

92

59+

94

2
2
2

2
2

2

8-8
9-8
8-9

]24

The University Science Bulletin

Plate XII. Leptodeira annulata annulata, RCT No. 1462, Los Diamantes,
Costa Rica, September 7, 1949. approximately natural size.

Taylor: Review of Snakes of Costa Rica 125

Scale Data on Leptodeira annulata annulata (concluded)

Labials

Labials

Infra -

Scale

Body

contact

enter

No.

labials

formula

bands

Temporals

chinshields

orbit

25742

11-11

21-21-15

52

(1 + 2 + 3
\l + 2 + 2

6-6

4-5

25743

11-11

19-23-15

49

(1 + 3 + 4
11+3 + 3

6-6

5-6

1462

10-11

21-21-15

50

1+2 + 3

5-5

4-5

Leptodeira ocellata Gunther

Leptodeira ocellata Gunther, Biologia Centrali-Amerieana ; Reptilia and Bafrachia, May 1895,

pp. 172-173, pi. 55, fig. B (type locality, Chontales Mines, Nicaragua; Costa Rica).
Leptodeira rhombifera (part.) Dunn, Proc. Nat. Acad. Sci., vol. 22, 1936, pp. (593-694.

The following data are taken from the type description: scales
in 23 rows; internasals about one third of the prefrontals; frontal
a little shorter than parietal; loreal large, square; preoculars two,
the lower very small, the upper reaching the frontal, or nearly so;
rarely a portion of a labial forms a small presubocular; two post-
oculars; eight supralabials, of which the fourth and fifth border eye;
temporals 1 -f- 2 -\- 3; the two pairs of chinshields of equal length;
ventrals 165-170.

Ground color reddish, or brownish olive; a series of dark brown
transverse or irregularly rounded spots along the back, which do
not descend to the sides and are about 38 in number on the trunk;
sides with a series of round brown spots, the spots being opposite
to the interspaces of the ground color and accompanied by smaller
irregular spots; neck without a distinct collar and longitudinally di-
vided in the middle by a narrow line proceeding from the first dorsal
spot. Upper side of head mottled brown; a brown temporal band.
Lower parts yellowish.

A specimen from Miramar, Costa Rica (U. S. National Museum
No. 75037) has the following characters: a median groove in an-
terior part of frontal; two preoculars, the upper preocular barely
separated from frontal; frontal pentagonal, the sides parallel; supra-
labials 8-8; infralabials 10-10, five touching first chinshields; scale
formula 21-23-17; ventrals 171, subcaudals 90. There are 32 bands
or spots on dorsum of body; approximately 28 on tail, those near tip
indistinct. A series of small lateral spots alternate with these; ven-
ter immaculate; a bilaterally symmetrical mark on the frontal and
a tridentlike mark on neck; some irregular spotting on parietals.
The dorsal spots usually touch the fourth scale row.

]2H The University Science Bulletin

Leptodeira rhombifera Gunther

Leptodeira rhombifera Gunther, Ann. Mag. Nat. Hist., scr. 4, vol. 9, 1872. p. 32 (type locality,
Rio Chisoy near Cubulco. Guatemala).

This species has been reported from Costa Rica but the reports
may be based on other forms. I append a brief description from
Gunther.

Scales in 25 rows; internasals very small; frontal as long as the
snout; loreal longer than high; one preocular barely touching
frontal; two postoculars; eight upper labials, of which the fourth
and fifth enter the orbit; temporals 1 + 2 + 3; chinshields rather
elongate, the pairs equal in length; body slender; head broad and
depressed; ventrals 170.

Brownish above, the body with about 26 large subrhombic dark
brown spots edged with black; yellowish cross bands brightest on
median line separate the rhombic spots; upper part of head brown
powdered with black; a black stripe with a yellowish margin on
each side connects the crown of the head with the first rhombic
spot. Abdomen yellowish, subcaudals powdered with brown.

Leptodeira maculata (Hallowell)

Megalops maculatus Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1861, p. 488 (type locality,

"Tahiti," [in error] ; possibly Costa Rica).
Anoplophallus maculatus Cope, Amer. Naturalist, 1893, p. 480.

fLeptodeira personata Cope, Proc. Acad. Nat. Sci. Philadelphia, 1868 (1869), p. 310.
Leptodeira septentrionalis maculatus (part.) Dunn, Proc. Nat. Acad. Sci., vol. 22, 1936, p. 397.
Leptodeira maculata Taylor, Univ. Kansas Sci. Bull., vol 25, 1938 (1939) pp. 337-342.

Six specimens in the Museo Nacional de Costa Rica, [presum-
ably] from Costa Rica have been identified as belonging to this
species.* I have not seen the specimens.

It seems well to regard the record of this species in Costa Rica as
doubtful until the form is rediscovered there, f The species is a
northern one, occurring widely in Mexican lowlands.

The following characters of the type will assist in identifying
maculata. Body moderately slender, and relatively short; prefron-
tals double the length of the internasals (the prefrontals are fused
for more than half their length [abnormal?] ); loreal elongate, much
longer than high; one preocular well separated from frontal, with a
curved rather than an angular front edge; one postocular (seem-

* Reported by Dunn in Proc. Nat. Acad. Sci., vol. 22, 1936, p. 697. This paper, purport-
ing to be a revision of the Genus Leptodeira has attempted to unite the genus Hypsiglena with
Leptodeira. and has in numerous instances wrongly synonymized forms. Maculata has been
made a subspecies of septentrionalis. These two species occur together over much of the area
of their distribution, and are not closely related within the genus.

t I am not wholly convinced that the foim described by Hallowell is identical with Lepto-
deira personata Cope. I had hoped to find the Costa Rican species but no specimens were taken.

Taylor: Review of Snakes of Costa Rica 127

ingly); fourth and fifth labials enter eye; first light band six scales
long; no elongate stripe on nape; 20 dark spots across body, 9 + on
tail; ventrals 171; 61 -f- subcaudals, the tip of tail missing. Scales
in 21 rows; anal divided.

Some of the spots are broken and tend to alternate or are irregu-
lar. The contrast between the spots and the light areas between
them, very pronounced. The head markings are indiscernible save
that the frontal appears a little darker than other dorsal head scales
and the edges are somewhat lighter.

Genus Oxybelis Wagler

Oxybelis Wagler, Natiirliches system der Amphibien, 1830, p. 183.

Genotype: Oxybelis acuminatus = aeneus (Wagler).
Three species occur in Costa Rica.

Key to Species of Oxybelis in Costa Rica

1. .Scales in 15 rows (15-15-13); ventrals 177, subcaudals 162; supralabials 6, brevirostris
Scales in 17 rows 2

2. Ventrals 198-217; subcaudals 139-165; supralabials 9-10 fulgidus

Ventrals 174-203; subcaudals 150-188; supralabials 8 aeneus

Oxybelis fulgidus (Daudin)

Coluber fulgidus Daudin, Histoire naturelle des Reptiles, vol. 6, 1803, p. 352, pi. 80 (Port au

Prince, San Domingo ex errore).
Oxybelis fulgidus (part. )Dumeril, Bibron and Dumeril, Erpetologie Generale .... vol. 7,

pt. 2, 1854, pp. 817-819; Boulenger, Catalogue of the Snakes in the British Museum, vol. 3,

1896, pp. 191-192.

Snout very elongate, pointed, its length three times diameter of
eye, flat at end and prominent; rostral scarcely visible above; inter-
nasals elongate, a little shorter than elongate prefrontals; frontal
narrower than supraoculars, nearly twice as long as broad, sub-
equal in length to prefrontals and parietals; loreal absent; nasal
single, very small; prefrontal contacts two or three labials, not enter-
ing eye; preocular one, not reaching frontal; postoculars one or two;
temporals large, 1 + 2; supralabials nine or ten, usually the fourth,
fifth and sixth (fifth, sixth and seventh) enter the eye; four infra-
labials border anterior chinshields, which are but little more than
half size of posterior pair. Scales in 17 rows, the dorsals feebly
keeled. Ventrals 198-217; anal divided; subcaudals 139-165.

Bright green above, yellowish green below and on upper lip;
a yellowish white line along each side of venter and in subcaudal
area. Reaches a length in excess of one and a half meters.

Ranges from Yucatan, Mexico, into South America.

]28 The University Science Bulletin

Oxybelis brevirostris (Cope)

Plate XIII

Dryophis brevirostris Cope, Proc. Acad. Nat. Sci. Philadelphia, I860, p. 555 (type locality,
Veragua, Panama); and Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875),
p. 132, pi. 26, fig. 2 (southeastern Costa Rica).

Oxybelis caerulescens Jan, Elenco Systematieo degli Ofidi, 1863, p. 88.

Cxybelis brevirostris Boulenger, Catalogue of the Snakes in the British Museum, 2nd ed., vol.
3, 1896, p. 190; Andersson, Goteborg Kungl. Veten. Vitt. Samh. Hand., vol. 17, 1914
(1916); Medd. Gdteborgs Mus. Zool. Afdel. 9, p. 36 (Siquirres, Costa Rica).

A specimen of this very slender tree snake was captured at Los
Diamantes by Richard C. Taylor (No. 366). It agrees with speci-
mens reported by Cope in having the scales almost, if not entirely,
smooth.

Body slender, compressed, rostral visible above; the suture be-
tween rostral and internasals a transverse line; internasals one sixth
shorter than prefrontals; length of frontal equals parietal length and
also equals its distance from rostral, its width in length at least
twice; parietals followed by two greatly enlarged upper secondary
temporals, which are separated mesially by a smaller scale; nostril
in single elongate nasal; no loreal; a large preocular touches frontal;
supraoculars as long as, but much wider than, frontal; one postocu-
lar; temporals 1 + 2, extremely large; supralabials 6-6 in following
order of size: 1, 2, 3, 5, 4, 6; infralabials 7-7; two pairs of chin-
shields, the posterior narrower, but a little longer than the first pair;
scale formula 15-15-13, smooth, the vertebral row somewhat en-
larged; ventrals 177, subcaudals 162; anal single; total length 872
mm.; tail 348 mm.

Color uniform greenish tending to become purplish or lavender
in preservative. I have examined also two specimens from Costa
Rica collected by William Gabb ( U. S. National Museum Nos.
30559-30560. The ventrals are 178 and 176 respectively; the sub-
caudals 75 +, 159.

Oxybelis aeneus (Wagler)

Dryiniis aeneus Wagler, in Spix. Serpentum Brasiliensium, 1824, p. 12. pi. Ill (type locality

near Ega, Brazil).
Oxybelis aeneus auratus (part.) Bogart and Oliver, Bull. Amer. Mus. Nat. Hist., vol. 83, 1945,

pp. 381-392.
Oxybelis acununatus Boulenger, Catalogue of the Snakes in the British Museum, vol. 3, 1896,

pp. 192-193 (Irazu, Costa Rica).

The status of this name for this form is somewhat uncertain.
Presumably following Schmidt's suggestion (Field Mus. Nat. Hist.
Zool. Ser., vol. 24) Bogert and Oliver, loc. cit., combine several rec-
ognized species into a single "species" extending from Arizona to

Plate XIII. Oxybelis brevirostris
Costa Rica, September 4.

(Cope), RCT No. 1366. Los Diamantes,
1947. Approximately natural size.

9—3286

130 The University Science Bulletin

Brazil. They state, "Whereas this material is representative it is,
of course quite inadequate for any satisfactory study of a variable
form ranging over large portions of two continents. The conclu-
sions we are able to draw therefore must be considered tentative
. . ." They have however shown that the name aeneus of Wag-
ler, antedates acuminatus of Wied, the name under which this spe-
cies has long been known.

The following characters will identify this form in Costa Bica:
snout narrow, elongate, sharply pointed, the rostral usually not vis-
ible from above; snout three to three and one-half times eye di-
ameter; internasals slightly shorter than the prefrontals; frontal
long, narrow, the width slightly more than half of length; parietals
slightly longer than frontal; nasal slender, small, single; no loreal;
prefrontal touching three supralabials; one large preocular; post-
oculars two; temporals large, 1 + 2; supralabials eight, five and six
enter eye; nine infralabials, four in contact with first chinshields.

Scales in 17-17-13 rows, smooth or faintly keeled; ventrals 174-
203; subcaudals 150-188; anal divided. Variable in color; green,
bronzy, grayish or reddish above, uniform or flecked with brown;
a black line on the side of the head; yellowish or brownish on
venter, speckled or streaked with brown, often with scattered black
dots.

Beported from "Irazu" Costa Bica by Boulenger.

Genus Imantodes Dumeril and Bibron

Imantodes Dumeril and Bibron, Mem. Acad. Inst. France, vol. 23, 1853, p. 507.

Genotype: Coluber cenchoa Linnaeus.

Three species of the genus occur in Costa Bica.

Key to Species of Imantodes in Costa Rica

1. Scales of the median vertebral row less than twice the width of adjoining scales;

ventrals 202, subcaudals HE inomatux

Scales of median vertebral row twice (or more) as wide as the adjoining scales. ... 2

2. Scales of median row three to four times width of adjoining scales: ventrals 233-

252 ; subcaudals 157-171 cenchoa semifasciatus

Scales of median row twice width of adjoining scales: ventrals 220-237; sub-
caudals 114-140 gemmistratus

Imantodes inornatus Boulenger

Plate XIV

Himantodes inornatus Boulenger, Catalogue of the Snakes in the British Museum (Natural
History), vol. 3, 1890, p. 88, pi. 5, fig. 1 (type locality, Hda. Rosa de Jerico, 3250 ft.,
Nicaragua ).

A specimen of this species (M.N.H. No. 25158) was acquired at
TsUi Bonita (American Cinchona Plantation) on the east slope of
Vol can Poas.

Taylor: Review of Snakes of Costa Rica

131

ift% '■

ss

;

Plate XIV. Imantodes inornatus, MNH No. 25158. Isla Bonita, Costa Rica,

about natural size.

132 The University Science Bulletin

Head broad with a very narrow neck; rostral about twice as wide
as high, the part visible above forming a very narrow line; inter-
nasals large, their length about two thirds that of the prefrontals
and about one half of their area; prefrontals minutely wider than
long; frontal slightly less than twice as long as wide, narrower than
the supraoculars, equal to its distance from snout tip; supraoculars
very large, of greater area than frontal, about one-third longer than
wide, about one-seventh shorter than parietals; latter scales rela-
tively short, longer than wide, their length equal to their distance
to a point little beyond middle of prefrontals; nostril in a single
nasal; loreal as long as high; a single preocular, very narrow and
high; two postoculars; temporals 1 -f- 2 -f- 3.

Supralabials 8-8, in the following ascending order of size: 1, 2,
4, 3, 8, 5, 6, 7; third, fourth, and fifth supralabials enter orbit; eye
very large, equal or minutely less than its distance from the end of
snout, the pupil vertical; anterior chinshields a little shorter than
posterior; infralabials 9-9, the first six border first pair of chinshields.
Scales smooth, the median row slightly, but not abruptly enlarged;
scale formula, 17-17-17; ventrals 202; subcaudals, 115; anal divided.
Total length, 821 mm.; snout to vent, 586 mm.; tail, 235 mm.

Above fawn with scattered black flecks or dots on head, a pair on
the posterior part of the parietals largest; a dim elongate mark on
neck behind parietals. Body with indistinct, extremely narrow,
darker, transverse markings across the back where thev tend to
contact the upper points of a zigzag line indicated only by minute
black flecks; the markings on tail similar but less distinct. Ventrals
with a peppering of black; pigment concentrated so as to form a
somewhat darker median line; chin lacking pigment. This speci-
men agrees in rather remarkable detail with the type description.

A second specimen examined (U. S. N. M. No. 15318) has 204
ventrals and 96 subcaudals. Otherwise it agrees with the type in
essential characters.

Imantodes cenchoa semifasciatus Cope

Plate XV

Himantod.es semifasciatus Cope. American Nat., 1894, p. 613.

Intantodes cenclwa semifasciata Smith, Pioc. U. S. Nat. Mus., vol. 92, 1942, pp. 384-385.

This extremely slender, greatly elongated snake was found usually
in trees either under bark or coiled in growing bromelias. Its large
eyes suggest that it is of nocturnal habits. The specimens were
extremely gentle, moving with great deliberation and allowing
themselves to be handled without showing fear.

Of the six specimens of this species that I have at hand, the three

>**■■/

;.

I

€*

f.

\

Plate XV. I7nant0d.es cenchoa semifasciata RCT No. 1370, Los Dia-
mantes, Costa Rica. September 1-8. 1947; about natural size.

134 The University Science Bulletin

from rather high elevation (5-6000 ft. elev. ) do not offer constant
differences from those taken in lowlands ( 800 to 900 ft. ) .

R. C. T. Nos. 644, 829, 830 are from the region about Turrialba.
R. C. T. No. 1370, M. N. H. Nos. 25700 and 25680 are from Los
Diamantes, 1 mi. south of Giiapiles.

There is no variation in the number of the supralabials, the
fourth and fifth enter the orbit in all. Five is the normal
number of infralabials touching the first chinshields, six occuring
two times in twelve. The temporals however are irregular; formulae

such as 3 4- 3 + 4, 2 + 2 + 3, 2 + 2 + 4, 2 + 3 + 3, i±l 3 + 3,

4 + 4, and 1 + 2 + 3 occur.

Males are characterized by tubercles on the chinshields and occa-
sionally on adjoining infralabial scales. Sometimes the second pair of
chinshields is separated by one or more small scales; sometimes
they are together.

The spots over the anterior part of the body tend to reach the
ventrals, but all do not. Practically all spots are brown narrowly
bordered by blackish brown and this color is in turn bordered by
scales that are cream, and lacking pigment for the most part. There
may be a few irregular black spots on sides near the edges of the
ventrals, opposite the main spots or possibly alternating with them.

This species may readily be diagnosed by the following characters
in addition to those given: head two to three times as wide as the
narrowest point of neck; eye very large, nearly equal to its distance
from the tip of snout; a small loreal present; scales smooth without
apparent apical pits, the scales of the median row somewhat quad-
rangular, three to four times the width of adjoining row; the scale
formula is 17-17-17.

I have examined four specimens in the U. S. National Museum
from "Costa Rica". The number of spots varies from 42-47 on body.

There are two specimens, Nos. 15309, Zent, Costa Rica and 19338
from Suretka, C. R., in the Harvard M.C.Z.

The species is known from numerous localities in Costa Rica
where it is called Bejuquilla. The name, however, is doubtless also
applied at times to other slender snakes.

Data on Imantodes cenchoa semifasciatus

Supra-

Infra-

Pie-

Post-

Sub-

Body

No.

Sex

labials

labials

ocular

ocular

Ventrals

caudals

Anal

spots

644

9

8-8

9-10

1-1

2-2

246

1

43

829

s

8-8

10-10

1-1

2-2

252

166

2

48

830

S

8-8

11-12

1-1

2-3

252

91+

2

49

1370

$

8-8

10-10

1-1

2-2

249

168

2

48

25700

S

8-8

10-10

1-1

3-3

248

171

2

25680

9

8-8

9-10

1-1

2-2

233

157

2

. .

Taylor: Review of Snakes of Costa Rica 135

Imantodes gemmistratus Cope

Himantodes cenchoa Cope (wee Linnaeus), Proc. A.cad. Nat. Sci. Philadelphia, vol. 12, 1860,

p. 264 (near Isalco, San Salvador).
Himantodes gemmistratus Cope, Proc. Acad. Nat. Sri. Philadelphia, vol. 13, 1861, pp. 296-

297 (type locality, near Izalco, Salvador).
Imantodes gemmistratus Smith, Proc. U. S. Nat. Mus., 1942, vol. 92, pp. 38i-386 (Chiapas,

Nicaragua, Costa Rica, Panama).

This form, fide Smith, is characterized by having most of the me-
dian dorsal scales twice the width of the adjoining row. Usually
three labials border the orbit; one preocular, two postoculars; ven-
trals 220-237; subcaudals 114-146.

The bands across body are about 42 in number, usually complete
throughout the length of the body.

Cope's type specimen is described as: head short, thick, the tem-
poral regions swollen; scales of the median dorsal series diamond-
shaped, longer than broad; one temporal in contact with the post-
oculars; sixth infralabial largest; the spots on body 42, tending to be
connected by a median dorsal vitta; head pale brown, with a few
irregular spots; a single apical pit in each scale.

Genus Clelia Fitzinger

Clelia Fitzinger, Neue Classification der Reptilien, 182(3, p. 56.

Genotype: Clelia daudinii Fitzinger, substitute name for Coluber
clelia Daudin.

Two species are known from Costa Rica.

Key to the Species of Clelia in Costa Rica

Scale rows 17-17-17(15); supralabials 7 (8); juvenile coloration: head and neck
black with a nuchal light band several scales wide; body reddish, each scale with
a black dot. Adults blackish clelia clelia

Scale rows 19-19-17; labials 8, 9; juvenile coloration: black with a series of red
transverse bands, complete or broken, three or foui scales wide petalarius

Clelia clelia clelia (Daudin)

Plate XVI

Coluber clelia Daudin, Histoire Naturelle Generale et Particuliere des Reptiles, vol. 6, Year 11

(1803), pp. 330-331, pi. 78 (type locality, Surinam).
Clelia clelia clelia Smith. Proc. U. S. Nat. Mus., vol. 92. 1942, p. 394; vol 93, 1943, pp. 402-

403.

Three specimens of this widespread species were taken, as fol-
lows: R.C.T. No. 636 (yg. ), 4 mi. E of Isla Bonita at an elevation of
about 3600 feet, and M.N.H. No. 25746 (yg.) about 5 miles west-
southwest of Isla Bonita at an elevation of about 6300 feet, Aug. 3,
1947. A third adult specimen, M.N.H. No. 25747, was taken at Isla

136

The University Science Bulletin

Plate XVI. Clelia clelia clelia (Daudin), MNH No. 25746, 5 miles
west Isla Bonita. Costa Rica ; approximately natural size.

Taylor: Review of Snakes of Costa Rica 137

Bonita on the highway, elevation about 5400 feet. The specimens
differ considerably in the character of the marking on the body.

No. 25746: In this specimen the characteristics are as follows: top
and sides of head, including parietals, postoculars, and four anterior
supralabials brownish black; a cream band on anterior nuchal
scales and temporals, with clouding on some of the temporals and
on scales touching posterior edge of parietals; width of this band
equal to width of four scale-rows; a dorsal blackish spot eight or
nine scales wide on neck, which, on its anterior part, reaches to ven-
trals but posteriorly only to fourth scale row; general color above
pinkish; on ventral surfaces cream white; body scales with a black-
ish area on posterior part of each scale, except outer row less regu-
larly spotted; tail blackish toward tip, ventrals immaculate save for
an occasional fleck on outer edges of the scales. Careful scrutiny
of body shows a suggestion of bands in a few groups of black flecks
on dorsum and sides.

No. 636: The general coloration follows the preceding with the
following exceptions. The color above pinkish, the edges of the
scales usually darker than centers, there being more pigment on
dorsal areas, less on sides; no distinct black spot on each scale; on
outer scale row and outer edge of ventrals dim grayish marking
suggesting a discontinuous irregular grayish line.

No. 25747, an adult, is blue black, the color extending onto ven-
trals on each side for one fourth of their width throughout greater
part of length. Middle half of ventrals immaculate cream-white
with a rosy tinge; chin and infralabials light; supralabials a little
lighter than head.

The following scale characters are the same in all the specimens:
Supralabials 7-7, the third and fourth entering eye; infralabials 8-8,
the first five bordering first chinshields. The temporals, however,

are variable. The formulae follow: No. 636, 2 + —,2 + 2 + 3-

2 + 2

No. 25746, 2 + ~-^ 5 No. 25747, 2 + 3.

The following table gives further scale variation:
Data on Clblia clelia clelia Daudix

No.

Sex

Ventral?

Caudals

Scale formula

636

$

209

82

17-17-15

25746

9

222

71

17-17-17

25747

$

228

61

17-17-17

138 The University Science Bulletin

Clelia petolarius (Linnaeus)

Plate XVII

Coluber petolarius Linnaeus, Museum Adolphus Friderici Regis, 1754, p. 35, pi. ix, fig. 2;

and Systems Naturae, 1766, 1, p. 387.
Oxyrhopus petolarius Boulenger, Catalogue of the Snakes in the British Museum, vol. 3, 1896,

pp. 101-103; Cope. Journ. Acad. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875), p. 132

(Sipurio, Costa Rica).

A single Costa Rican specimen is at hand, collected on the Pa-
cific slope at San Isidro El General (R.C.T. No. 969).

Rostral narrowly visible, its width a little greater than its height;
internasals small, about one third area of prefrontals which are
very nearly as broad as long; frontal subtriangular, its length about
one-sixth longer than wide, and equal to its distance from the
middle of the prefrontals, much more than twice as wide as the
supraoculars; length of parietal about equal to its distance from
the internasal; nostril between two nasals; loreal elongate, nearly
twice as long as high; one preocular touching frontal; two postocu-
lars; temporals 2 + 3; supralabials 8-8, the fourth and fifth enter
orbit; infralabials 10-10, five touching first pair of chinshields,
which are a little larger but about same length as posterior pair;
scale formula: 24 (about head) -19-19-17, with paired apical pits.
Ventrals 201; subcaudals 79 + 1.

Color above blackish; an occipital reddish band covering ex-
treme tips of parietals and the following four scale rows; this in
turn followed by a broad black collar, eleven scales wide; behind
this 29 orange-red bands, most of which are continuous across back,
but nine or ten separated mesially tending to alternate on opposite
sides; ten transverse bands on tail most of which are continuous;
red bands are two to three scale rows wide, the intervening black
bands six to eight scale rows wide posteriorly (anteriorly, four to
five). The belly and chin white; subcaudal region grayish to black,
the red bands encircling the tail. The reddish orange bands have
the scale edges darkened, occasionally spotted.

The specimen was caught under a small log, about a half mile
west of San Isidro El General on the Pacific side of Costa Rica, by
Richard C. Taylor.

A specimen in the U. S. National Museum listed by Cope, op. cit.,
has been examined. There are 202 ventrals and 92 subcaudals in
this specimen; scale rows 21-19-19-17; one preocular; two postocu-
lars; temporals 2 + 3; supralabials 8-8; infralabials 10-10; loreal 2/2
times as long as wide; anterior chinshields double area of second
pair. There are 25 light bands on body.

Taylor: Review of Snakes of Costa Rica

139

Plate XVII. Clelia petolarius (Linnaeus), RCT No. 969, San Isidro El General,
Costa Rica. August 27, 1947; about natural size.

140 The University Science Bulletin

Genus Erythrolamprus Boie

Erythrolamprus Boie, Isis, vol. 19, 1826, p. 981.

Erythrolamprus Wagler, Natiirliches System der Amphibien, 1830, p. 187 (part.).

Genotype: Erythrolamprus agilis Wagler (fide Fitzinger).
A single form is known to occur in Costa Rica.

Erythrolamprus bizonus Jan

Erythrolamprus aesculapii bizona Jan, Arch. Zool. Anat. Fis., vol. 2, fasc. 2, 1863, pp. 314-
315 (Bahia, Messico, Popayan, Cayenne, Brasile, Montevideo, Colombia).

Erythrolamprus venustissimus Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876
(1875, published as separate), p. 141 (Sipurio, Costa Rica).

Erythrolamprus aesculapii (part.) Boulenger, Catalogue of the Snakes in the British Museum,
vol. 3, 1896, pp. 200-201 (lists a specimen from Irazu, Costa Rica, ventrals 197. sub-
caudals 50); Picado, Serpientes Venenosas de Costa Rica, 1931, pp. 32-33, fig. 8 (right).

The use of the name bizonus for the Costa Rican form of Erythro-
lamprus aesculapii Linnaeus (£. venustissimus Wied) is that of
Bailey, mentioned in Dunn and Bailey, Bull. Mus. Comp. Zool.,
Harvard College, vol. 86, no. 1, 1939, p. 12.

Jan has applied this name to a form presumably ranging from
Mexico south through Colombia, Cayenne, Brazil and Uruguay.
Dunn and Bailey state that, as the original name was somewhat
composite, they restrict the name to Colombian specimens with the
bands double even on the neck.

I have examined several Costa Rican specimens in the U. S. Na-
tional Museum and a summary of the scale counts taken is given
in the accompanying table. The scale characters are as follows:

The rostral visible above as a narrow triangle; length of common
internasal suture about 2/2 times in that between prefrontals; frontal
about one-fifth longer than its distance to snout tip, nearly as long
as parietals; nasal partially divided, the suture above nostril; loreal
higher than wide; a large preocular; two postoculars; temporals
1+2; seven supralabials, the third and fourth entering orbit; eight
or (usually) nine infralabials, four touching the first chinshields,
which are longer than second pair; scale formula 15-15-15, scales
smooth; ventrals 181-201, subcaudals 49-59.

Color above red with a series of 13-15/2 paired black bands en-
circling body separated by a narrow red band. A black spot on the
head involves supraoculars, frontal, and two thirds of the area of
parietals. The labials and scales on tip of snout and some scales
on back of head with blackish marks.

Total length (of U. S. N. M. No. 13536) 598 mm.; tail 87 mm.
(San Jose, Costa Rica).

Taylor: Review of Snakes of Costa Rica 141

Table of Scale Counts for Costa Rican Ertthrolampeus bizona

Body Tail Yen- Sub- Supra- Infra-

No. Locality triads triads trals caudals labials labials

9784 Costa Rica 13% 4% 189 53 7-7 9-7

9779 Costa Rica 15% 4 198 49 7-7 9-9

13536 San Jose, C.R. 151/.. 4 198 56 7-7 8-9

14013 Costa Rica 15% 4 198 59 7-7 9-9

30677 Sipurio.CR. 15% 4 194 58 7-7 9-9

8312* Costa Rica 15 4 201 57 7-7 9-9

Genus Coniophanes Hallowell

Coniophanes, Hallowell, in Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, 1860, p. 248.

Genotype: Coronella fissidens Giinther.

Three Costa Rican species are referred to this genus.

Key to Costa Rican Species of Coniophanes

1. Scales in 17 rows; upper parts grayish brown; two narrow yellowish lines on each
side, one on the first scale row, the other originating from a spot on neck, and

continuing on body as a series of almost contiguous dashlike spots elect /inn.-;

Scales in 21-25 rows 2

2. Scales in 25 rows; a light stripe beginning on head passing eye and continuing with
the dorsolateral light lines of body (rarely interrupted on neck); a black dorsal
stripe present, five to seven scales wide; separated from a lateral black stripe three
scales and two half scale rows wide, by a cream stripe somewhat less than three

scales wide piceivitlis

Scales in 21 rows; above brownish, the median dorsal scales each with a small
blackish posterior spot; all scales of body slightly darker on edges; a dim lateral
line on 4th and 5th scale row, and a light line on sixth row ; venter peppered with
fine pigment fissidens punctigularis

Coniophanes decipiens (Giinther I

Tachymenes decipiens Giinther, Biologia Centrali-Americana ; Reptilia and Batrachia, 1895,

p. 163, pi. 53. fig. A (type locality, Irazii, Costa Rica).
Rhadinaea guntheri Dunn, Copeia, 1938, no. 4, p. 198 (substitute name for decipiens).

Dunn's reference of this snake with grooved back fangs to the
genus Rhadinaea, I consider untenable and the substitute name
which he proposed after this action, I believe, should be suppressed.

Body slender, the tail about one half length of body (tail mu-
tilated in the type); snout short; internasals short and small; frontal
shorter than parietals, but as long as or a little longer than snout;
one preocular, one or two postoculars; temporals 1+2, widely
separated from the postoculars; the sixth, or fifth and sixth labials
forming sutures with the parietal; seven supralabials, third and
fourth bordering orbit; scales in 17 rows; ventrals 165-173; grooved
posterior fangs present.

Upper parts dark grayish-brown; two narrow yellowish lines on
each side, one of which runs on the first row of scales; the other
originates from a spot on each side of the neck and is continued as
a series of elongate, dashlike spots which are almost contiguous; a

* No. 8312 is in M. C Z.. Harvard. Others in I . S. National Museum.

142 The University Science Bulletin

white spot on the antero-lateral part of the parietal; supralabials
blackish brown on upper parts, yellowish on lower parts. Venter
uniform yellowish.

If the species is not congeneric with Coniophanes, it may be
necessary to consider this form under a new generic designation.

Coniophanes piceivittis Cope

Coniophanes piceivittis Cope, Proc. American Philos. Soe., vol. 11, Ju!y, 1869 (type locality
Chihuitan, Oaxaca): Bailey. Michigan Acad. Sci.. Arts. Lett., vol. 24, pt. 2, 1938 (1939),
p. 31 (Bebedero, Costa Rica).

This species, so far as I am aware has its place in the list of Costa
Rican snakes on the basis of a specimen in Vienna * purporting to
come from Bebedero, Costa Rica. Unless further specimens are
found it might be well to re-examine this specimen to confirm the
identification or to question the origin of the specimen. The point
nearest to Bebedero, at which the species is definitely known to
occur as recorded by Bailey, is nearly 300 mi. north of Costa Rica
in the northern part of Honduras!

A diagnosis of the species is given: scale rows 25; a small "subpre-
ocular"; a broad median stripe and a broad lateral dark stripe four
or five scales wide, the dark stripes having sharply defined edges;
ventrals, males 15S-168, females 160-173; subcaudals, males 82-91,
females 78-84; scale formula: 21-23-25-17. Anal ridges present in
mature males. Eight supralabials, fourth and fifth entering eye;
infralabials 10, six touching chinshields; temporals 1 + 2.

Head with a light stripe through top of eye, continuous with the
dorsolateral light lines or interrupted on neck. Labials and chin
spotted with dark brown or black; body with a black dorsal stripe
five to seven scales wide; a black lateral stripe on either side three
and two half scales to five and two half scales wide, separated from
the dorsal stripe by a narrow dorsolateral light stripe one and two
half scales wide. The lower edge of the lateral dark stripe sharply
defined contrasting with the light lower rows; lowest two rows
faintly flecked with black; belly nearly immaculate.

Coniophanes fissidens punctigularis Cope

Coniophanes punctigularis Cope. Proc. Acad. Nat. Sci. Philadelphia, vol. 12, 18t!0, p. 248
(type locality, Honduras); Smith, Proc. U. S. Nat. Mus., vol. 91, 1941, pp. 107-109,
map, fig. 33.

Coniophanes fissidens fissidens (parr.). Bailey, Michigan Acad. Sci., Arts Letters, vol. 24, pt. 2.
1938 (1939), pp. 14-23, pi. 1. fig. 4.

Two specimens are in the collection from Costa Rica. The fol-
lowing characters obtain: R.C.T. Nos. S/2, 1369; sex, § , (yg. );

•Bailey (Pap. Michigan Acad. Sci. Arts Lett., vol. 24, pt. 2. 1938 [1939], p. 31) quotes
from notes of E. R. Dunn.

Taylor: Review of Snakes of Costa Rica 143

loreal quadrangular; preoculars, 2-1, 1-1; postoculars, 2-2, 2-2; tem-
porals, 1 + 2, 1 + 2, 1 + 3 + 3, 1 + -^-; supralabials, 8-8, 8-8; labials
enter eye, 4-5, 4-5; 4-5, 4-5; infralabials, 10-9, 10-9; infralabials touch
chinshields, 5-4, 5-5; scale formulae, 21-21-17, 22-21-17. Ventrals
120, 123; subcaudals 46 +, 69; anals, 2, 2.

Color markings very distinct in the young No. 1369 from Los
Diamantes, Costa Rica. Above brownish, the median scale row
with a black spot in posterior part of each scale so as to simulate a
median discontinuous dark line; all scales with slightly darker edges
so that a fine, but distinct reticulated pattern is evident; fourth and
fifth rows with dark flecks so that a dim lateral or dorsolateral line
is evident, the color growing lighter toward ventrals.

Head light brown with some indefinite blackish dots on parietals;
labials cream, bordered by a blackish line above; posterior seventh
and eighth labials are brown above, the cream covering only their
lower fourth (or less); cream area flecked with black; two minute,
blackish-bordered, cream ocelli on temporals and a larger one
somewhat behind these at same level. At about fifth ventral a
black-edged cream line begins on the sixth scale row and continues
back. The black border is soon lost and the line can be traced with
little difficulty to end of body; chin and entire underside of body
light cream peppered with black dots, eight or ten on each ventral;
those on outer edge of ventrals largest.

The larger specimen, No. 972, is darker and venter more salmon
pink; spot on outer edge of ventrals not larger than the others and
may be indistinct or absent; lateral stripe not clearly discernible
since all scales are more pigmented; ocelli and light lateral stripe
evident, the stripe being only on anterior part of body.

The larger specimen was killed by Richard C. Taylor near San
Isidro El General with the same shot that also killed a specimen of
Dryadophis melanolomus alternatus. Only a single snake had been
seen, and consequently there is no data on the behavior of the two
species or known reason for the two specimens being so closely ap-
proximated.

A specimen in the U. S. National Museum has 115 ventrals and 70
subcaudals.

Known variation in ventrals and subcaudal counts are: g 114-
119, subcaudals 70-80; 5 121-127, 63-75.

Railey, loc. cit., has reported the species from the following locali-
ties in Costa Rica: Siquirres; El General, Surubres, Suretka, and
Palmilla.

144 The University Science Bulletin

Genus Rhinobothryum Wagler

Rliinobotltryum Wagler, Natiirliehes System der Amphibien, 1830, p. 180.

Genotype: Rhinobothryum macrorhinum Wagler.
Only one species is known from Costa Rica.

Rhinobothryum bovallii Andersson

Rhinobothrium bovallii Andersson, Medd. Goteborgs Mus. Zool. Afdel., 9 (Goteborgs Kungl.

Vetensk. Vitt. Semh.) Band 17. 1914 (1916), pp. 32-33 (type locality, Siquirrres, Costa

Rica).
Rhinobothryum bovallii Dunn and Bailey, Bull. Mus. Comp. Zool., Harvard Coll., vol. 86,

1939, pp. 16-17 (Limon, Costa Rica, and localities in Panama).

This rare back-fanged snake with grooved teeth may be dis-
tinguished by the following characters: rostral large, wedged in
between internasals, the portion visible above shorter than its dis-
tance from frontal; latter broader than long, much shorter than
parietals, much shorter than its distance from tip of snout; loreal
longer than high; one preocular, not reaching to upper surface of
head; two postoculars; temporals, 2-2; eight supralabials, the fourth
and fifth bordering eye; fifth to seventh much higher than others;
ten infralabials, four or five touching the anterior chinshields; pos-
terior chinshields about half area of first and separated by two scales,
nearly equal to anterior in length; scales in 21 rows, smooth an-
teriorly, keeled posteriorly on back and on tail; on sides the five
outer scale rows are smooth; ventrals 240-246, notched and dis-
tinctly angular laterally; anal single; subcaudals (divided) 115.
Two posterior teeth of maxilla enlarged, grooved.

Top of head black, the scales outlined with yellow margins, the
sides white (red) each scale with a dark spot; a narrow light band
on back of head followed by a dark band 12 scales long, not cross-
ing throat; body banded with black or blackish brown and red
bands, separated from each other by narrow white bands; in middle
of body the red bands cover six or seven up to nine or ten scale-
lengths, the dark bands six and the white only two scale-lengths.
Red scales all tipped with black; on belly, red becomes white and
the dark and white alternate; posterior ventrals have dark spots in
areas corresponding to the red color. Five black bands on tail
as broad as the red intervening bands. Length 1190 mm.; tail
290 mm.

The species is known from Limon and Siquirres, Costa Rica. I
have examined a specimen (Harvard M.C.Z. No. 42790 from Es-
peranza Ridge, Panama.

Taylor: Review of Snakes of Costa Rica 145

Genus Conophis Peters

Conophis Peters, Monatsb. Akad. Wiss. Berlin, I860, p. 519.

Genotype: Conophis vittatus Peters.

Two species have been reported from Costa Rica. They may be
differentiated by the following key.

Key to Costa Rican Species of Conophis

Upper labials immaculate; head light with three broad brown stripes; body light.

with brown stripes lineatus dunni

Upper labials with black borders ; head black with two white lines ; body blackish

with two white lines on each side. . . nevermanni

Conophis lineatus dunni Smith

Conophis lineatus Cope (nee Dumeril and Bibron), U. S. Nat. Mus. Bull., 32, 1887, p. 77 (San

Jose, Costa Rica); Dunn, Copeia, 1937, no. 1, p. 214 (Barranca, Tivives, Esparta, San

Jose and Cartago).
Conophis lineatus similis Smith (nee Boeourt), Journ. Washington Acad. Sci., vol. 31, no. 3,

Mar. 15, 1941, pp. 123-124.
Conophis lineatus dunni Smith, Proc. U. S. Nat. Mus., vol. 92, 1942, p. 395. (Managua,

Nicaragua.)

I have examined a paratype of this form from Esparta, Costa Rica.

Snout projecting, part of rostral visible above, one third to one
half its distance from rostral; frontal approximately as long as wide,
a little longer than its distance from tip of snout; approximately
as long as the parietals; one preocular, not touching frontal; two
postoculars; loreal present; temporals 2 + 2; supralabials 8(7), the
fourth and fifth enter eye; infralabials 8-8 to 10-11; temporals
2 + 2 + 3; scale rows 19-19-17; ventrals 169-174; subcaudals 64 <? ,
67-69 2 .

Chin and labial borders heavily pigmented; a dark stripe along
first scale row; lateral dark stripe, passing through eye, restricted
to the third and fourth scale rows throughout the length of the body
and of solid color ( without a broad light median area between the
two dark edges); a dark stripe on seventh (or sixth posteriorly)
scale rows; no secondary dark stripes on either paravertebral rows
or those adjacent to them laterally; second scale row white anteriorly;
posteriorly with a dotted secondary line; ends of ventrals pigmented
in some parts of body.

Conophis nevermanni Dunn

Conophis nevermanni Dunn, Copeia, 1937, no. 4, pp. 214-215 (type locality, Rio Poas de
Aserri, Costa Rica).

The type description gives the following data: Scutation similar
to Conophis lineatus. Scale rows 19-17, smooth, lacking apical pits;
nasal divided; preocular one, postoculars two; temporals 2 + 2;
10—3286

146 The University Science Bulletin

supralabials eight, the fourth and fifth border orbit; infralabials 9
or 10, five touching anterior chinshields; ventrals 179-184 ( 182 + 2);
subcaudals 70.

Black on back and sides, light on venter; a light line from snout
continues on sixth scale row; a similar line on second scale row;
upper labials edged with black.

Maximum known size, 820 mm.

Reported from type locality and possibly from Bebedero, Costa
Rica (see Dunn, loc. cit.). The type locality is said to be "a few
miles south of San Jose."

Genus Stenorrhina Dumeril

Stenorrhina Dumeril, Mem. Acad. Inst. France, vol. 23, 1853, p. 490.

Genotype: Stenorrhina ventralis Dumeril.

The snakes of this genus may be readily distinguished from other
snakes by the fusion of the anterior nasal with the internasal scale.

The material that I have studied leaves some doubt as to the num-
ber of recognizable forms of this genus that occurs in Costa Rica.
This genus is in need of revision and I believe that Dr. J. Bailey has
such a revision under way.

At least three forms appear to be present, which may be distin-
guished by the following key.

Key to Forms of Stenorrhina in Costa Rica

1. Belly strongly pigmented, each ventral scale with a darker transverse band

d. degenhardtii
Belly immaculate cream-white without or with slight trace of pigment; a black
temporal stripe present or indicated 2

2. Five lines on body more or less distinct /. freminvillii

No lines on body, scales dark or blackish on bases d. apiata

Stenorrhina degenhardtii degenhardtii (Berthold)

Calamaria degenhardtii Berthold, Abhand. Gesell. Wiss. Gottingen, vol. 3, 1846. p. 8, pi. 1,

fig. 2, 4 (type locality, Mexico and Cential America).
Stenorhina degenhardtii Jan, Arch. Zool. Anat. Fis., vol. 2, 1862, p. 63; Boulenger, Catalogue

of the Snakes in the British Museum, vol. 3, 1896, pp. 229-230.
Stenorhina ventralis Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875)

p. 141 (Old Harbor, Costa Rica).

Maxillary teeth 15-17, anterior ones small, equal, separated from
two enlarged posterior teeth with a groove on outer side. Head not
distinct from neck; nostrils between two nasals, the anterior of
which is fused with internasals; snout obtusely pointed; rostral
broader than deep, the part visible above measuring one half to
two thirds of its distance from frontal (rarely equal); internasals
about as long as prefrontals or a little shorter; frontal one and one-
third to one and one-half times as long as wide, as long as or a little

Taylor: Review of Snakes of Costa Rica 147

longer than its distance from snout tip; as long as or a little shorter
than parietals; loreal present or absent, if present small; when ab-
sent fused to an adjoining scale; prefrontals often touching labials;
one preoculars, two postoculars; temporals 1+2, 2+3, or 2+4;
seven snpralabials, third and fourth enter eye; seven or eight infra-
labials, three or rarely four labials touching first pair of chinshields,
which are longer than posterior pair. Scales in 17-17-17 smooth
rows without apical pits; ventrals 158-171; subcaudals 33-36; anal
divided; length 750 mm.; tail 100 mm.

A Costa Rican specimen has upper parts brown with more or
less distinct darker spots and irregular crossbars; belly more or less
black spotted. In one specimen there are no well-defined markings.

Two Costa Rican specimens (Nos. 32582-83) collected by Wil-
liam Gabb are in the U. S. National Museum collection. Both are
now very soft but the dark ventral coloration is distinct in both.
No. 32582 has 159 ventrals, 33 subcaudals.

Stenorrhina degenhardtii apiata Cope

Stenorhina d[egenhardtii] apiata Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2. vol. 8,
1876 (1875), p. 142 (El Barrio, Oaxaca).

This subspecies is included in the Fauna of Costa Rica on the
basis of a specimen in the U. S. National Museum (No. 9774) col-
lected by Zeledon in "Costa Rica," no specific locality known.

The general characters of this form are as follows: rostral large,
more than half its surface visible from above, equal to about three
fourths its distance from frontal; internasal fused with the nasal;
posterior part of the combined scale elongated; frontal six-sided,
elongate; nearly a third longer than its distance from snout tip; a
large preocular, VA times as high as long; two postoculars; temporals
1-2; 7-7 supralabials, the third and fourth enter orbit; first chin-
shields larger than second, and tending to separate the second pair
for some distance; scale rows 17-17-17, the count around posterior
part of head, 24; ventrals 174, subcaudals 36.

The specimen is nearly uniformly colored, each scale with a
darker basal spot or area; belly immaculate cream; labials cream
except for a black line that crosses side of head.

Stenorrhina freminvillii freminvillii Dumeril, Bibron and Dumeril

Stenorhina freminvillii Dumeril. Bihron and Dumeril, Erpetologie Generate, vol. 7, pt. 2, 1854,

pp. 868-869, pi. 70. fig. 1, 2 (Mexico).
Stenorrhina freminvillii freminvillii Smith, Proc. U. S. Nat. Mus., vol. 93, 1943, p. 472.

A single specimen, Harvard M. C. Z. No. 15294, from Costa Rica
is referred to this subspecies.

148 The University Science Bulletin

The part of rostral visible above large; internasal fused to nasal;
a small loreal present; one preocular, two postoculars; temporals
1-2; supralabials 7-7; infralabials 7-7, three labials touching the
anterior chinshields; second chinshields small, less than half of first;
mental very narrow; ventrals 182, subcaudals 38; anal divided;
scale rows 17-17-17 without apical pits.

Head brown with some black spotting; labials cream save on
their upper edges, which have a dark longitudinal mark. Body
brownish with 5 indistinct dotted lines on back; venter immaculate
cream.

Genus Scolecophis Fitzinger

Stolecophis Fitzinger, Systems Reptilium, 1843, p. 25.

Genotype: Calamaria atrocincta Schlegel.
A single species is known from Costa Rica.

Scolecophis atrocinctus (Schlegel)

Calamaria atrocinctus Schlegel, Essai sur la Physionomie des Serpents, vol. 2, 1837. p. 47.

Scolecophis atrocinctus Cope, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 259.

fPseudoboa petola Picado, Serpientes Yenenosas de Costa Rica, 1931, pp. 33-34, fig. 9 (right).

The species has a range extending from Honduras to Costa Rica.
I have examined a single male specimen from Costa Rica now in
the U. S. National Museum, collected by Zeledon.

Rostral narrowly visible above; internasals small, their length
in prefrontal length about 2/2 times; frontal distinctly longer than its
distance to the end of snout; parietals large, one fourth longer
than their distance to snout tip; nasal divided; a small loreal, nearly
square; one preocular; two postoculars; temporals 1 + 1; supra-
labials 7-7; anterior chinshields twice size of second pair; eye small,
its diameter twice in distance to anterior edge of rostral; temporals
1+1+2; ventrals 191; anal divided; subcaudals 53; scales 15-15-15,
smooth with a single, very small, apical pit.

Body ringed in black and yellow (white in preservation). On
the dorsum the black transverse rings cover 3/2 to 4 scale rows, the
yellow, 2 to 2/2 rows; on venter the white occupies two ventrals, the
black three ventrals; head blackish with a band across snout chiefly
on the prefrontals, first two labials and the anterior edge of frontals;
a triangular white spot from the labials reaching above eye level
involving all of fifth and parts of adjoining labials; tip of chin black
to the middle of the first chinshields; a small black round spot at
the beginning of the throat; first white neck band two scales wide
followed by a black band five scales wide; a total of 36 black bands
on bodv, two on head and seven on tail.

Taylor: Review of Snakes of Costa Rica 149

The maxillary teeth 14, with two posterior small grooved fangs;
eight anterior maxillary teeth grooved laterally (perhaps all maxil-
lary teeth are so grooved since the six following teeth are missing).

( lenus Tantilla Baird and Girard

Tartiilla Baird and Girard, Catalogue of North American Reptiles. 1853, p. 132.

Genotype: Tantilla coronata Baird and Girard.
Six species are regarded as occurring in Costa Rica. They ma}'
be differentiated by the following key.

Key to Costa Rican Species of Tantilla

1. Body reddish brown, with narrow black-bordered cream lines from middorsal line
to ventrals, usually alternating; first labials touch or are separated; ventrals 148-

155 : subcaudals 60-65 annulata

Holy not so marked with transverse bars i!

2. Body brownish or reddish; head and part of neck brown followed by a narrow
white collar severed medially by a longitudinal median black line; white on venter;
nasal separated from preocular or not; ventrals 150-171; subcaudals 44-52 . .armillata
Body not so marked 3

3. Three longitudinal black-edged strips on body; frontal pentagonal; first pair of
lower labials separated (rarely touch) bringing mental and first chinshields to-
gether; chestnut brown, the scales darker on edges; a pale collar across back of

head; ventrals 148; subcaudals 67 reticulata

Not so marked 4

4. Nasal undivided (rarely partly divided); ventrals 180; subcaudals 71; grayish
above with a white collar; two black-edged dorsal brown stripes and one lateral
run length of body; sometimes a dark line on each side of venter; below white, virgata

5. Only a single pair of chinshields; first infralabials in contact: ventrals 146;
subcaudals?; light brown above, yellow below; a median brownish stripe and a
narrow yellow stripe on adjacent borders of the third and fourth scale rows,
bounded above by a single dark brown row, and below by several rows with

dark centers ; black spot below eye ruficeps

Two pairs of chinshields; first lower labials separated: ventrals 121-135; sub-
caudals 38; above grayish brown, with an indistinct lighter collar across parietals ;

no white spots on head shistosa

Tantilla annulata Boettger

Plate XVI 11

Tantilla annulata Boettger. Zool. Anz., 1892, p. 419 (type locality. Nicaragua): Dunn, Proc.

Acad. Nat. Sci. Philadelphia, vol. 92, 1940, p. 119; Dunn and Bailey, Bull. Mus. Coin]..

Zool. Harvard Coll., vol. 86, Oct. 1939, p. 19 (between Peralta and Turrialba, Costa Rica).
Tantilla semicinctum Barbour and Amaral, Bull. Antiv. Inst. Amer., vol. 1, no. 4, 1928, p. 100

(fide Dunn) (Not of Dumeril, Bibron and Dumeril).
Homalocranium annulatum Giinther, Biologia Centrali-Americana, 1895, p. 150 (description

of type specimen); Boulenger, Catalogue of the Snakes in the British Museum, ed. i. 1896,

p. 217; Werner, Mitt. Naturh. Mus. Hamburg, Jahrg. 26, 1909. pp. 238-239, fig. 11.

A single specimen (M.N.H. No. 25705) of this rare form was
obtained at Turrialba, July 17, 1947. Its characters are given in
detail since it differs from the type in certain points.

Rostral distinctly wider than high, part visible above as wide as
an internasal, angular posteriorly; internasals half as long as pre-
frontals; frontal hexagonal, about one-sixth longer than wide, the

150

The University Science Bulletin

lateral angles extremely obtuse, its length about one fourth greater
than its distance from tip of snout, its width less than twice that of

Plate XVIII. Tantilla annulata Boettger. MNH No. 25705, Turrialba, Costa

Rica, July 17, 1947; 502 mm.

posterior ends of supraoculars; parietals large, their length very
slightly less than their distance from tip of snout; nostril between
two nasals, which are narrowed at point of mutual contact, the
posterior smaller, in contact with the preocular; no loreal; one

Taylor: Review of Snakes of Costa Rica 151

preocular; two postoculars, the lower smaller; temporals 1 + 1 + 1
(on left side the two posterior are partially fused); supralabials,
7-7, in the following order of size: 1, 3, 2, 5, 4, 6, 7; infralabials
6-6, in the following order of size: 2, 6, 1, 3, 5, 4, the first four touch
chinshields; first pair of labials separated, leaving the very large
mental in contact with first chinshields, which are double the size
of the second pair; latter narrowly in contact (in the type the first
infralabials are in contact, and the mental is separated from the
first chinshields).

Scale formula: 15-15-15, the scales smooth with a single mi-
nute pit on most of the dorsal scales, situated some distance in
from the posterior median border," visible under magnification,
ventrals, 148, separated from the posterior chinshields by one scale;
subcaudals 59+1; anal divided.

Snout yellowish white (the rostral somewhat grayish), the color
extending to mouth, and posteriorly to middle of prefrontals; this
followed by a broad black band, which extends back to posterior
tips of parietals, and laterally across upper and lower labials to
chinshields; a white spot covering fifth upper and fifth lower labials
as well as edges of adjoining scales; remainder of chin white; a
narrow yellowish band, one scale wide, crosses extreme posterior
parts of parietals and adjoining scale row, interrupted mesially by
a single black scale; this band followed by a broad black-brown
nuchal band at least six transverse scale-rows wide and bordered
behind by another narrow yellowish band; this in turn bordered
by a blackish band two scales wide. General body color following
this is reddish brown, to brown lower on sides, the median part of
each scale slightly darker than the outer parts, thus leaving an
impression of very faint longitudinal lines on each scale row ( a
little more evident posteriorly); a series of yellow vertical blotches
the width of a scale length and bordered laterally by black of
nearly same width, reach to middorsal line, alternating (usually)
with a similar series on opposite side; these yellow and black spots
separated usually by eight to ten scales. Dorsal coloration en-
croaches on ventrals so that outer fourth of ventrals colored like
sides; a few scattered dark flecks on median part of ventrals; me-
dian part of venter and subcaudal region salmon to coral red;
yellow bands tend to widen somewhat low on sides; there are 12
or 13 of these on body, five or six on tail. Maxillary teeth, 20, sub-

* The presence of these minute "pits" lias been noted by me in certain other species of
Tantilla. I am uncertain whether or not these are homologous with the one, two or four
"apical" pits that may occur on other snakes.

152 The University Science Bulletin

equal, followed, after a slight diastema, by two smaller grooved
fangs; 25-26 mandibular teeth.

Total length, 502 mm.; tail, 108 mm.; snout to vent, 394 mm.;
width of head, 10.2.; length of head, 15 mm.; width of body,
11.5 mm.

The above description differs somewhat from the brief descrip-
tion of the type. In that specimen, the markings are confined to
the anterior third of the body, and the ventral color is yellow
(faded?) instead of red.

From a young specimen described by Dunn ( loc. cit. ) , the Costa
Rican specimen, while agreeing in the general pattern, differs much
in color and in having the first labials separated. In Dunn's speci-
men the ventrals are 155, subcaudals, 65. There are 13 marks on
each side of body; six on tail.

A specimen reported by Werner, op. cit., agrees in much better
detail with my specimen, both as regards squamation and color
pattern. This specimen is 590 mm. in length; ventrals 149; sub-
caudals 59 + 1. No locality data are given.

Dunn and Bailey ( loc. cit. ) , report a specimen from between
Peralta and Turrialba, Costa Rica. This has 151 ventrals; sub-
caudal count not given ( perhaps a defective tail ) .

Tantilla armillata Cope

Tantilla armillatum Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (,1875),

p. 143 (type locality, middle Costa Rica).
Tantilla melanocephalum var. Cope, Proc. Acad. Nat. Sci. Philadelphia. 1871, p. 205.
Hvmalocraiuum armillatum Giinther, Biologia Centrali- Americana ; Reptilia and Batrachia,

1895, p. 149, pi. 52, tig. C (Cartago, Costa Rica).

Slender, small snake with scales in 15 rows; head flat, depressed;
eye less than half the length of the snout; length of prefrontals
three times the length of internasals; frontal wide, much shorter
than the parietals; one preocular, two postoculars: nasals notched
slightly below, the posterior touching the preocular (or narrowly
separated from it by the prefrontal ) ; temporals 1 + 1+2; seven
supralabials, the seventh much the largest; six infralabials, the first
pair form a suture behind the mental, first four touch first chin-
shield, the fourth also touching second pair of chinshields. Ven-
trals 159-171; anal divided; subcaudals 44-52.

Above chocolate-brown, head and nape for five scales black,
with a yellow spot on end of muzzle, one on posterior part of each
parietal plate, two on lip behind eye, and one below each nostril;
the black is bordered posteriorly by a yellow collar of two scales
length, which is also bordered by black behind except where it
sends off on the third and fourth rows of scales on each side a

Taylor: Review of Snakes of Costa Rica 153

narrow light band which extends to tail. Below this, and also on
median scale row, is a narrow line of brown. Below immaculate
white; total length as given by Gunther, eight and three-fourths
inches.

I have examined two Costa Rican specimens of this species as
follows: Harvard M.C.Z. No. 15285 from Surubres (Surubreo),
Costa Rica has a ventral count of 174, the subcaudals defective.
The head and neck, for six scale rows back of parietals, generally
black with some cream spots followed by a cream band two scales
wide; a discontinuous band forming four spots, one on last labials,
one covering parts of back end of parietal and temporal; two small
yellow spots on extreme anterior part of parietals; a spot on back
edge of rostral and internasals; a spot covering parts of first and
second labials, and a spot on fifth labial; venter immaculate, light
lateral band present.

A specimen in the U.S. National Museum (No. 9787) from
"Costa Rica" has 169 ventrals, 47 subcaudals. The head spotting
differs somewhat in detail from that described above; a light stripe
follows the third and fourth scale rows to tail; a row of dark dots
follows the first scale row.

It may eventually be necessary to regard this form as a sub-
species of Tantilla melanocephala, but I lack necessary material
to establish this.

Tantilla reticulata Cope

T[antilla] reticulata Cope, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 77 (type locality,
Coyuoas de Yeiagua, New Grenada).

Rostral broad, visible from above; nasal divided, the nostril
chiefly in prenasal; postnasal touching first two supralabials, pre-
ocular, internasals, and prefrontals; one preocular, two postoculars;
temporals 1 + 1 + 1; frontal plate broad, angular in front, pro-
jecting posteriorly for half its length between parietals; supra-
labials seven, last largest, third and fourth border the orbit; an-
terior chinshields touch mental separating first pair infralabials;
scales in 15 rows; ventrals 148, anal divided; subcaudals 67.

Color chestnut brown above, much darker posteriorly than an-
teriorly, the color extending upon the edges of the ventrals; anteri-
orly the scales have somewhat darkened edges, presenting a reticu-
lated appearance; median dorsal row of scales lighter, forming a
pale line which disappears on tail; third and fourth rows on each
side lighter, forming indistinct lines; a pale yellow-brown collar
crosses posterior parts of parietals; head scales clouded and edged

154 The University Science Bulletin

with darker; a deep brown mark extending from parietals to the
month across the yellowish labials; on under side pale yellow,
deepening posteriorly.

I have examined a specimen from Reventazon (river?), Costa
Rica ( Harvard M. C. Z. No. 15266). The detail of the color pattern
follows: A brown line on first scale row and ventral edge; a cream
line covering upper edge of first row and part of second; a brown
line covering upper half of second and one half of third; a cream
line on upper half of third, fourth and lower half of fifth rows; a
brown line on upper half of fifth; a cream line on sixth row; lower
part of seventh row dark; a median light stripe on middorsal row
and upper part of seventh scale row. The nuchal cream band is
interrupted mesially; head dark with a white spot on snout, and one
on first labials; a black spot below eye; brown flecks on several of
the infralabials; venter immaculate yellow or whitish; ventrals 176;
subcaudals 66.

Tantilla virgata (Giinther)

Microdromus virgatus Giinther, Ann. Mag. Nat. Hist., ser. 4, vol. 9, 1873, p. 17, pi. 4, fig. B

(type locality, Cartago, Costa Rica, four cotypes).
Homalocranium virgatum Bocourt, Etudes sur les reptiles; Mission Scientifique au Mexique
, livr. 9, 1883, p. 584, pi. 30, fig. 4 ; Giinther, Biologia Centrali-Americana, Reptilia

and Batrachia, 1895, p. 154, pi. 52, fig. A; Boulenger, Catalogue of the Snakes in the

British Museum, vol 3, 1896, p. 223.
Homalocranium sexfasciatum Fischer, Abhandl. d. naturw. Verein, Bremen, Bd. 7, 1882, pp.

225-220, pi. 14, figs. 8-10 (Costa Rica).
Tantilla sexfasciata Cope, Bull. U. S. Nat. Mus. no. 32, 1887, p. 83.
Tantilla virgata Cope, Bull. U. S. Nat. Mus., no. 32, 1887, p. 84.

Nasal usually undivided; rostral just reaching upper surface of
snout; internasals a little less than half of the prefrontals; frontal
five-sided, longer than broad; parietals as long as their distance to
rostral; nasal not or but slightly in contact with preocular; one
preocular, two postoculars; seven supralabials, third and fourth
border orbit; temporals 14-1; first pair of infralabials separated
or narrowly in contact; anterior chinshields large; posterior small,
scalelike; ventrals 154-175, subcaudals 57-71; scales in 15-15-15 rows;
total length 315 mm.; tail 75 mm.

Upper parts grayish or yellowish brown; a white collar present;
three or four stripes on each side of body; a pair of brown bands
edged with black two scales broad, run from collar to middle of tail;
a similar band along each side of body, and sometimes a narrow
blackish line along each side of abdomen. Lower parts uniform
yellowish white. Upper labials white, with a black spot below eye
and on rostral shield.

Known from Cartago, Costa Rica.

Taylor: Review of Snakes of Costa Rica 155

Tantilla schistosa (Bocourt)

Homalocranion schistosum Bocourl , Etudes sur les reptiles: Mission scientifique au Mexique et

dans l'Amerique Centrale, livr. 9, 1883, p. ;">84, pi. 30>, figs. 10, lOa-lOe (type locality.

Haute Verapaz, Guatemala).
Tantilla schistosa Cope, U. S. Nat. Mus. Bull., no. 32, 1887, p. 83; Dunn and Bailey, Bull

Mus. Comp. Zool., Harvard College, 1939, pp. 5, 19 (Altos de Cangrejal, near San Jose.

C. R.).

A specimen of this Tantilla (M. N. H. No. 25731) was captured
along the Reventazon River at Turrialba, under a log where also
was obtained a Micrurns nigrocinctus mosquitensis. The Tantilla
was placed in the same container with the Micrurus, but on return-
ing to the laboratory, it was missing and was later discovered in the
stomach of the Micrurus. Digestive fluids have injured the specimen
so that accurate identification is impossible. The following miscel-
laneous characters can be discerned, however, and the species ap-
pears to be very close, if not identical, to Bocourt's schistosa.

Part of rostral visible above smaller than internasals; latter scales
about one half length of prefrontals, which are broader than long;
frontal hexagonal, about one-third longer than its distance from end
of snout, one-fourth longer than wide, the front border forming a
very obtuse angle; parietals about one-fifth longer than their dis-
tance from end of snout; scale rows 15-15-15.

The character of the scales on the sides of the head and the lips
cannot be determined with certainty; the first pair of labials are
separated, the large anterior chinshields touching the mental. A
light yellowish band crosses the neck and encroaches on the pos-
terior part of the parietals. Olive above (becoming brown in
preservatives), each scale with a lighter center and a darker outer
border, thus giving a geometrically reticulated pattern (which is
distinct if the specimen is submerged in water). Chin and throat
bluish white; anterior one fourth of body dull creamy white gradu-
ally becoming, first dimly, then strongly, salmon pink; edges of ven-
trals lighter; under side of tail uniform salmon pink.

A specimen in Harvard M. C. Z. (No. 15302) is from Altos del
Cangrejal, Costa Rica. The following scale characters obtain: ven-
trals 140; subcaudals 29; one preocular; two postoculars; temporals
1+1; seven upper labials, the third and fourth border orbit; five
infralabials, three touching first chinshields. Generally brown
above, the scale edges a little darker.

156 The University Science Bulletin

T ant ilia ruficeps (Cope)

Poyanaspis ruficeps Cope, Proc. Acad. Nat. Sci. Philadelphia, 1894, p. 204 (type locality,

Costa Rica).
Tantilla ruficeps Amaral, Mem. Inst. Butantan, vol. 4, 1929, p. 221 (Costa Rica).

Rostral visible from above; frontal rather long, hexagonal; nostril
between two nasals, the posterior separated from preocular by pre-
frontal; one preocular, two postoculars; temporals 1-1; seven supra-
labials, the third and fourth bordering orbit; first infralabials in
contact behind mental; a single pair of chinshields touching four
lower labials. Ventrals 146; anal divided.

Light brown above; median dorsal row of scales deep brown; a
narrow yellow line on adjacent borders of third and fourth rows
of scales which is bounded above by a dark brown line; upper sur-
face of head reddish brown, with a pale spot at end of each parietal
shield; upper lip yellow, and a black spot below the eye; lower
parts yellow. Length 223 mm.

The species presumably is known from the type specimen only.

Family HYDROPHIIDAE

A single genus representing this marine serpent family has
reached the Western Hemisphere. It is distributed along the west-
ern coast from California to Chile.

Genus Pelamis Daudin

Pelamis Daudin, Histoire naturelle . . . des Reptiles, vol. 7, year XI, (?1803), pp. 357-362.

Genotype: Anguis plat lira Linnaeus.

One species is known from the Pacific coastal waters.

Pelamis platiirus (Linnaeus)

Hydrus bicolor Schneider, Historiae Amphibiorum naturalis et literariae, Fasc. Primus, 1799,

p. 242 (type locality, Pine Island, Pacific Ocean).
Pelamis bicolor Ficndo, Serpientes venenosas de Costa Rica, 1931, pp. 23-27, fig. 6 (Costa

Rica).
Pelamis platurus Gray, Ann. Philos., 1825, p. 15.

This marine serpent, the only species known on the American
coasts, has been reported for the Pacific coast of Costa Rica, where
it is reputed to be abundant. The high, compressed type of tail is
found on no other American snake. This serpent may be recog-
nized by the following characters: upper half of body blackish,
lower half dirty white or yellowish, the two separated by a narrow
cream line; posterior part of body somewhat spotted; tail yellow
with a number of black spots.

The tail strongly compressed into an "oar" about 10 times as high
as wide. Scales on median ventral surface of body and tail about

Taylor: Review of Snakes of Costa Rica 157

size of dorsal scales. Frontal as long as its distance from tip of
snout, nearly equal to a parietal; nostril in nasal directed dorsally;
one preocular, two or three postoculars; temporals numerous; seven
to nine supralabials, the fourth or fourth and fifth entering orbit.
Scales 45-57 about middle of body, smooth in young and females,
with one to three tubercles in males. The species reaches a length
of three feet.

I have examined the following Costa Rican specimen: U. S. N. M.
No. 37490. Some of the characteristics of this specimen are: scale
formula, 42-47-49-35; temporals 2+2+3+4; supralabials 9-9; ven-
trals 295, preceded by 13 scales between chinshields and first dis-
tinguishable ventral; 49 subcaudals; 16 dorsal scale rows black;
next four cream, remainder dirty white or light tan; spots tend to
form dorsal and ventral rows on tail which tend to dovetail with
some yellow color between.

U. S. N. M. Nos. 96038, 96040, 96041, Port "Parlear," Costa Rica:

No. 96038. Scale formula, 41-46-47-33; scales with two or three
tubercles, sometimes absent on dorsal scales; rostral partly divided
by a suture; 8-9 supralabials; ventrals 340; subcaudals 43.

No. 96040. Scale formula, 42-46-47-35; 9-9 supralabials; tern-
portals 2+2+3+4; an elongate subocular; ventrals 320; sub-
caudals 48.

No. 96041. Scale formula, 42-46-49-37; ventrals 320, subcaudals
41; 7-7 upper labials; 11-11 infralabials; temporals 2+2+3+4; one
subocular on one side.

Family ELAPIDAE
Only a single genus, Micrurus, is known to occur in Costa Rica.

Genus Micrurus Wagler

Micrurus Wagler, in Spix, Serpentium Brasiliensium . . . , 1825, p. 48.

Genotype: Micrurus spixi Wagler.

Four species and six forms are known to occur in Costa Rica.
They may be recognized by the following key.

Key to Forms of Micrurus in Costa Rica

1. Two-color species, black and red (pink or whitish); long, moderately slender
snakes ; ventrals 269 ; subcaudals 26 ; 62 dark black bands on body, four on

tail mipartitlts mitltifasciatus

Three-color species, black, yellow (cream), red '-

2. Subcaudals more than 50 (53-58); ventrals 192-199; 13-16 black rings on body;

7-9 on tail : no keeled supra -anals clarki

Subcaudals less than 50 3

3. Seventeen or more black rings on body -*

Sixteen or less rings on body •>

158 The University Science Bulletin

4. Body slender; 18 body rings, 4 on tail; ventrals 2 241; subcaudals 32; keeled

supra-anal scales alleni richardi

Body thicker; 22 body rings, 5 or 6 on tail; ventrals <|> 215; subcaudals <j>

40 pachecoi

5. Black rings 10-12 on body ; 3-5 on tail, usually totaling 15; ventrals g 189-
193; $ 208-210; subcaudals ^ 47-49; 9 35; length 698 mm.

nigrocinctus mosquitensis
Black rings 16 or. body; 5 on tail; ^ ventrals 194-204, subcaudals 47-48; J
ventrals 214-218; subcaudals 34-36; length 596 mm nigrocinctus itigrocinctus

Micrurus mipartitus multifasciatus (Jan)

Plate XIX

Elaps multifasciatus Jan, Rev. Mag. Zool., 1858, p. 521; Gunther, Ann. Mag. Nat. Hist., ser.

4, vol. 9, 1872, p. 36; and Biologia Centrali-Americana, Reptilia and Batrachia, 1895,

p. 184.
E'aps mipartitus Picado, Serpientes venenosas de Costa Rica, 1931, p. 32 fig. 9 (non Dumeril,

Bibron and Dumeril).

Only a single specimen (M.N.H. No. 25187) of this large red and
black banded coral snake is in the material at hand. It was found
at Turrialba on the I. A. LA. farm in trash, Aug. 1, 1947. The speci-
men furnishes the following data: head rather flat, not, or scarcely,
distinct from neck; eye diminutive, its greatest diameter ( two mm. )
about four times in distance from eye to end of snout, and about
two and two-tenths times in its distance from edge of upper lip.
Rostral broader than deep, part visible from above shorter than
internasals; latter scales less than half length of prefrontals; frontal
scale narrow but at least one-third wider than supraocular, its length
equal to its distance from middle of internasals; parietals large, their
length equal to their distance from rostral; nasal divided; loreal
longer than wide, bordering eye and forming a broad suture with
supraocular; no preocular present; two postoculars; temporals 1 + 1
(1 + 2), the anterior not especially narrow; seven supralabials,
third and fourth very high entering orbit, in following ascending
order of size: 1, 2, 4, 5, 7, 3, 6; infralabials seven, four normally
touching first chinshields, which are a little broader but about same
length as second pair (scales on right side of chin region abnormal
due to injury).

Scale rows 15-15-15; ventrals, 269; anal divided; subcaudals, 26,
the fourth and fifth undivided; 62 dark bands on body, four on tail
with extreme tip black, and deflected downward.

Picado (loc. tit.,) reports a total length of 1130 mm. for the spe-
cies.

Taylor: Review of Snakes of Costa Rica

159

W

Plate XIX. Micrurus mipartitus multifasciatus (Jan), MNH No. 25187,
Turrialba, Costa Rica, August 1, 1947.

160 The University Science Bulletin

Micrurus nigrocinctus nigrocinctus (Girard)

Plate XX

Elaps nigrocinctus Girard, Proc. Acad. Nat. Sci. Philadelphia, 1854, p. 226, and U. S. Xaval
and Astronomical Expedition, vol. 2, Zoology, 1855, p. 210, pi. 35 (type locality, Taboga
Island, Bay of Panama).

Micrurus nigrocinctus yugrocinctus Schmidt, Smithsonian Misc. Coll., vol. 89. no. 1, Mar. 16.
1933, p. 18 (Panama localities); Zool. Ser. Field Mus. Nat. Hist., vol. 20, Dec. 11, 1933,
p. 33 (.Panama Canal Zone, southwestern Panama, Pacific slope of Costa Rica, and Nic-
aragua); Dunn, Notulae Naturae, no. 108, 1942, p. 8 (Boruca, Buenos Ayres, Costa Rica.
One of these, Amer. Mus. Nat. Hist. No. 17365, served as a paratype for it. mosquitensis
Schmidt).

Schmidt gives the following range of scales in a series from
Panama: Ventrals and subcaudals respectively, males 194, 196, 204;
47, 47, 48, with totals 241, 243, 245. For females, 214, 215, 215, 218;
34, 36, 36, 35, with totals 248, 251, 251, 253.

A specimen M. N. H. No. 25190 taken on the Pacific slope of
Cerro de la Muerte at Boquete Camp is referred to this species. The
ventrals are 207, the subcaudals 55, totaling 262, of which the third
to tenth are undivided. There are 16 black bands on the body, and
five on the tail.

Scale characters of this specimen are as follows: part of rostral
visible above nearly equal in length to median internasal suture,
much wider than high; length of internasals equal to two thirds
length of prefrontals; latter scales one fifth wider than long, forming
an acute angle laterally; frontal nearly double width of supraoculars
which are narrowed posteriorly, the widest part being near the an-
terior end; eye very small, 3/2 times in its distance from end of snout;
frontal at least one fourth longer than its distance from the end of
snout; one fifth shorter than parietals; length of latter equal to their
distance from rostral; nasal divided; loreal as high as long, forming
equal contacts with eye and supraocular; two postoculars; tem-
porals 1 + 1 + 2, the upper tertiary largest; seven supralabials;
seven infralabials, three in contact with first chinshields which are
about one fourth shorter than second pair.

Scales smooth save that the lateral scales in front of anus and four
lateral caudal rows are distinctly keeled, as is the outer edge of the
first subcaudals. Scale formula 15-15-15; ventrals 207; anal divided;
55 subcaudals, of which the third to tenth are undivided.

Tip of snout black, the color covering first three labials and upper
part of fourth, the supraoculars, frontal, and a narrow margin of
parietals that border frontals. There are 16 black bands on body,
their width equal to four scale lengths; anterior edges of black bands
straight, posterior edge dentate; on each side a yellow band two

Taylor: Review of Sxakes of Costa Rica

161

mm

Plate XX. Micrurus nigrocinctiis nigrodnctus (Girard), M.N.H. No. 25190.
Boquete Camp on Pan-American Highway. Pacific slope, Cerro de la Muerte;
total length, 596 mm.

11—3286

162 The University Science Bulletin

scales wide borders the black; red bands between yellow bands, all
scales of which have a black spot; on ventral surface body bands
occupy two or three ventral scales, the red and yellow, eight to
eleven; there are five black bands on tail, each the width of five or
six subcaudals.

Usually on each of the ventral red areas there is a single small
black fleck. The yellowish band on the back of the head terminates
one scale row back of the parietals; it is continuous under chin,
covering all of it save the anterior portions of the first three infra-
labials. Total length, 596 mm.; snout to vent, 502 mm.; tail, 94 mm.

This specimen differs from other Costa Rican and Panamanian
specimens that I have examined in having several of the subcaudals
single.

The type specimens of Elaps nigrocinctus Girard * were collected
on Taboga Island, Bay of Panama, Panama, by Lieut. J. M. Gilliss.
The two specimens are U. S. N. M. No. 7347 (2 specimens), the
larger $ (figured) and a smaller § cotype. The figured specimen
has 16 body bands (including the nuchal) and four on tail, the
terminal band being very small. The yellow borders of the black
bands are scarcely indicated in the larger cotvpe, but the smaller
specimen has one or one and one-half scale rows of yellow border-
ing the black bands. The smaller specimen has 14 bands on neck
and body, four on tail. The ventral counts of the two specimens
are respectively 213 ventrals, 36 4- 1 subcaudals; 214 ventrals, 35
subcaudals. Girard reports 217 -f- 1 ventrals ( probably counting
the small scales on chin posterior to chinshields); subcaudals 18.
The latter figure is an error probably for 38 which would be cor-
rect if the small scale lateral to the anal is counted. Total length,
760 mm.; tail, 72 mm.

The mainland specimens of nigrocinctus agree fairly well with
the small Taboga Island form although there is usually a somewhat
larger number of body bands ( 15 to 17 ) and the ventral count is
higher, the subcaudal count lower; however, the total ventral-sub-
caudal counts are 244-256, similar to the total in the types. This
variation may represent merely a sex difference.

0 Proc. Acad. Nat. Sci. Philadelphia, vol. 7, 1854, p. 226; and U. S. N. and Astro Exped.,
vol. 2, Zool. 18.55, pi. 35, figs. 1-6.

Taylor: Review of Snakes of Costa Rica

163

Plate XXI. Micrurus nigrocinctus mosquitensis Schmidt, RCT No. 471, La
Suiza, July 15, 1947; about natural size.

164 The University Science Bulletin

Micrurus nigrocinctus mosquitensis Schmidt

Plate XXI. text fig. 5

Micrurus nigrocinctus mosquitensis Schmidt. Zool. Ser. Field litis. Nat. Hist., vol. 20, 1933,

p. 33 (type locality. Limon, Costa Rica).
Elaps fulvius Picado, Serpientes Venenosas de Ccsta Rica, 1931, pp. 28-29, fig. 8 (left) (non

Elans fulvius [Linnaeus]).

Five specimens of this species were collected in the Caribbean
drainage of Costa Rica. Two are from the lowlands (elevation less
than 900 ft. ) at Los Diamantes, one mile south of Guapiles ( RCT
no. 1442, MNH no. 25191). Two were taken at Turrialba (RCT
no. 15, MNH no. 25166), and one at La Suiza, five to six miles south-
west of Turrialba (RCT no. 471). This series of specimens agrees
in general in the following color characters:

Fig. 5. Micrurus nigrocinctus mosquitensis Schmidt,
RCT No. 471. La Suiza. Costa Rica. Head dorsal
view X 3.

Anterior part of head black, this color reaching back and includ-
ing most of supraoculars, entire frontal, most of fourth supralabial,
and a narrow black stripe behind eye on postoculars; a light yellow-
ish band crossing back of head reaches one scale-length back of
parietals, passes under chin to near median ventral line, where it
is interrupted by a black or blackish longitudinal stripe connecting
blackish area on front of chin with first black neck band. Paired
chinshields are black with some light flecks.

Body with 10-12 bands varying in length from six to nine scale-
lengths. Anterior edge of each band straight, posterior edge den-
tate; yellow bands bordering black, washed with brownish fawn,
their length equal to three scale-lengths. Intervening spaces dull
red above, orange or tomato-red below; each scale with a very
distinct black spot often covering more than half its area; red

Taylor: Review of Snakes of Costa Rica

165

ventrals have small black flecks or blotches. On venter, black bands
have a length of six ventrals on anterior part, and five ventrals on
posterior part of body; each black band bordered by immaculate
yellow for a width of two ventrals. Tail with from three to five
bands, having a length of nine or ten scales, and covering nine or
ten pairs of subcaudals ventrally except where the last black spot
is terminal in which case band may be small.

The two Turrialba specimens, from elevations of 1750 and 1950
ft. have practically no dark blotching or spotting on the ventrals
and all caudal red bands lack many of the black spots on the dorsal
and lateral scales. The yellow head band is dim but in those from
near Turrialba the area is suffused with smoky fawn.

The figure given ( pi. XXI ) shows the general characteristics of the
form, particularly the wide yellow bands bordering the black.

I have examined the type and certain paratypes* of mosquitensis.

Scale

Data on

MlCBURUS NIGROCINCTUS MOSQUITENSIS

Sub-

Black bands

Supra -

Infra -

No. Sex

Ventrals

caudal

s

Total

body and tail

labials

labials

15 $

189

47

236

11 + 4

7-7

6-6

471 9

210

35

245

12 + 3

7-7

6-6

1442 9

208

35

243

12 + 3

7-7

6-6

25166 $

(192)*

48

240

10 + 5

7-7

6-6

25191 3

193

49

broken :

Hid

242
part missing.

11 +5

7-7

6-6

* Estimate, since

the body is

Table of

M

EASUREMENTS OF

M

ICRURUS

NIGROCINCTUS

MOSQUITENSIS

Total

Snout

No.

Sex

length

Tail

to vent

15

6

494

78

316

471

9

568

58

510

1442

9

698

75

623

25166

<5

750*

114

636*

25191

S

480

73

407

* Estimate.

Micrurus pachecoi subsp. nov.

Plate XXII. text fig. 6

Type. — Univ. Kansas Mus. Nat. Hist. No. 25188; Guanacaste,
Costa Rica; collector and date unknown.

Diagnosis.— A large species with 22 bands on body and 5 on tail,
6 counting a black tip. Ventrals 215*2; subcaudals 40; very few or
no black spots on red areas; dark pigment dispersed, slightly

* One of the numbers listed as a paratype, U. S. N. M. No. 9784, is a specimen of Erythro-
lamprus aesculapii; it is possible No. 9785 is intended since this is a coral snake with 19
bands on body and six on the tail. I do not believe this to be a mosquitensis, however; the
diagnosis of the form mentions body bands "10-19 on the body," and this may be the source
of the "19" since all specimens that I regard as authentic mosquitensis have a much lower
count of black bands.

166

The University Science Bulletin

Plate XXII. Micrurus pachecoi sp. nor., MNH No. 25188. Guanacaste, Costa
Rioa, August 1947; total length, 984 mm.

Taylor: Review of Snakes of Costa Rica

167

heavier near posterior tip of scales; yellow bands narrow, pig-
mented, about one scale-length wide. Temporals 1 + 2. ( Since
specimen is female it is impossible to say whether the male may
have the keeled scales on sides above anus. )

Description of the type.— Rostral injured, slightly visible from
above; mternasals a little more than half the length of the prefron-
tals which are about one fifth wider than long, forming an acute
angle between loreal and posterior nasal; supraoculars equal to or
only slightly less than width of frontal; latter scale almost twice as
long as wide, about one fourth or one fifth longer than its distance
from tip of snout; length of parietals equal or slightly greater than
their distance from snout tip; nasal divided, loreal longer than wide,

Fig. 6. Micrurus pacht ?coi sp. nov. Type. MXH No.
25188, Guanacaste, Costa Rica.

entering orbit and forming a rather large suture with the supraocu-
lar; no preocular; two postoculars; temporals 1 + 2 + 2, upper pos-
terior largest; seven upper labials in the following ascending order
of size: 1, 2, 4, 3, 5, 7, 6, third and fourth entering orbit; infralabials
seven, four bordering the first chinshields, which are about one fifth
shorter than second pair; scales smooth, 15-15-15; ventrals 215)2;
subcaudals 40; 22 bands on body, 5 on tail, also a black tip.

Measurements in mm.— Total body length, 984; tail 102; snout to
vent 882.

168 The University Science Bulletin

Color. — The black bands vary in width between four and five
scale-lengths, while on the ventral surface they occupy two or three
ventrals. The caudal bands are wider and may cover five to seven
subcaudals; the width of the intervening red blotches is from six to
eight scale-lengths, occupying six to eight ventrals on the under
surface. The usual yellow borders on the black bands are obsolete
or nearly so, save that the outer scale may be cream or yellow, very
rarely the ventral scale preceding or following the black bands may
be partially white. Some of the red scales have black dots but
these are not very conspicuous since the whole scale has a brown-
ish wash. The ventrals have a few black flecks on the posterior
part of the body. The red bands on the tail are very narrow, one
or two scale-lengths in width dorsally; on the subcaudal region the
band is as wide as three subcaudals and a black spot is present in
the center. Head and snout black, including frontal supraoculars,
postoculars, first, second and the upper part of the third and fourth
labials. The four anterior infralabials are dull black but the broad
yellowish head band is continuous across chin. Dorsally the band
is clouded with smoky black, lighter on the supralabials.

Remarks.— The specimen was presented to me alive by Prof.
Marco Tullio Pacheco, and it is for him that the species is named.

The peninsula which juts out from the northern part of the west
coast of Costa Rica includes much of the province of Guanacaste.
It is connected with the main land mass by relatively low land, sug-
gesting that it was formerly an island. I suspect that this species of
coral snake was developed when island conditions obtained in the
region. The reduction of discrete black spots on the red scales,
the larger size of the supraocular scales with reference to the fron-
tal, the increased number of black bands on body and tail, and the
temporal formula 1+2, suggest the wisdom of regarding the form
as having full specific rank rather than as a subspecies of the re-
lated M. nigrocinctus, until such time as intergradation is demon-
strated. It may, however, be related to that species through nigro-
cinctus coibensis whose range is Goiba Island off the Pacific coast
of Panama.

Micrurus clarki Schmidt

Micrurus clarki Schmidt, Zool, Series Field Mus. Nat, Hist., vol. 20, Oct. 31, 1936, pp. 211-
212 (type locality, Yavisa, Darien, Panama).

This coral snake lacks the supra-anal tubercles in the male. It
may be recognized by the following description: body form and
general appearance that of Micrurus corallinus; supralabials 7-7;

Taylor: Review of Snakes of Costa Rica 169

infralabials 7-7; preoculars 1-1; postoculars 2-2; temporals 1+1,
1 + 1; ventrals 199, subcaudals 58; parietals angulate at sutures
of temporals and post-temporals, and produced at the postero-
lateral angles into distinct points; 13 black rings on body, usu-
ally the width of three ventrals, and seven on tail; the nuchal
black ring has a length of six scales, narrowed on belly to three
ventrals; yellow, bordering black, one scale-length wide; scales of
yellow and red zones uniformly spotted with black at tips; black
of head extending to tips of parietals; yellow nuchal scales heavily
black-margined; black caudal rings separated by yellow rings which
are immaculate beneath but heavily black-mottled above; red ven-
trals immaculate. (From type description).

I have examined a Costa Rican specimen listed as a paratype.
This has 189 ( 192 ) ventrals and 53 subcaudals. The black rings are
16 + 8 ( 9 ) on body and tail. This specimen, collected by Sr. Jose
Zeledon, is without specific locality data.

Micrurus alleni richardi subsp. nov.

Plate XXIII, text fig. 7

Type. — Univ. Kansas Mus. Nat. Hist. No. 25189, collected by
Edward H. Taylor and Richard C. Taylor, Sept. 8, 1947, at Los
Diamantes about 2 km. southeast of Guapiles, Costa Rica.

Diagnosis. — A slender elongate form brilliantly colored with yel-
low-edged black bands on a red body. Black bands, 18 on body,
four on tail; ventrals, 241; subcaudals, 32; space between black
bands 2/2 to 2% the length of a black band; scales of red areas with
a black spot on each scale; narrow yellow areas with lighter spot-
ting; ventral parts of the red and yellow bands immaculate. Black
of snout extending back onto partietals.

Description of the .species.— Rostral slightly visible above; inter-
nasals about half the length of prefrontals; latter scales very slightly
wider than long laterally forming an angle between nasal and
loreal; frontal narrow, one and two-thirds as long as wide, sharply
angled posteriorly, considerably (K) longer than its distance from
tip of snout; parietals narrow, elongate, about one-sixth longer than
frontal, about equal to their distance from internasals; nasal divided;
loreal large, forming anterior border of eye and a broad suture with
the supraocular; latter scale nearly as broad anteriorly as posteriorly;
eye small, about 3.2 times in its distance from snout tip and about
once and a half in its distance from the lip; no preocular; two post-
oculars, upper not noticeably larger than lower; temporals 1 + 1;

170

The University Science Bulletin

a large scale borders the posterior part of parietal; supralabials 7-7,
in the following ascending order of size: 1, 2, 4, 3, 5, 7, 6; infra-
labials 7-7, four bordering anterior chinshields; posterior chin-
shields narrower and about one-fourth longer than anterior. Scales

Plate XXIII. Micrurus alleni richardi subsp. nov., MNH No. 25189, Los
Diamantes, 2 km. SE Guapiles, Co.sta Rica, September 8. 1947; total length,
888 mm.

Taylor: Review of Snakes of Costa Rica

171

smooth except on side just anterior and posterior to vent, which
are keeled. Scale formula, 15-15-15; ventrals, 241; subcaudals, 32;
anal divided.

Measurements in mm. — Total length, 888; snout to vent, 815; tail,
73.

Color. — There are 18 black bands on body, and one crossing an-
terior part of head but incomplete below; four black bands on tail.
Body black bands usually as wide as five or six scale-lengths, their
anterior borders usually straight, the posterior borders dentate;
black bands bordered by one or two rows of yellowish or yellowish
white scales with usually a dark spot; intervening bands broad,

Fig. 7. Micrurus alleni richardi subsp. nov. Type.
MNH Xo. 25189. Los Diamantes, Costa Rica.

red, each scale of which bears a black spot; these bands are more
than double the width of the black bands, being equal to 10-12
scale-lengths; on ventral surface the black covers three or four
ventrals on body, five to six subcaudals; the red and white (yel-
low ) together cover nine or ten ventrals except the first two, which
are 15 and 12 respectively; under tail the red occupies four or five
pairs of subcaudals; snout and anterior part of infralabials black.
This includes first three supralabials, most of the supraoculars, all
of the frontal, and some of anterior mesial part of the parietal;
posterior edge of the black is bordered by a clouded fawn border
that extends to back edge of parietals. Thus the yellow head band
is partially interrupted on the median dorsal line but is continuous
on sides of head and chin.

Remarks.— The colors of this specimen were extremely brilliant
in life. This specimen was apprehended after it had entered a crev-

172 The University Science Bulletin

ice in a stony bank of a small river, and only a small part of the
body was in evidence. In the same area within a few hundred yards,
specimens of typical Micrurus nigrocinctus mosquitensis Schmidt
were found.

The fact that this form occurs together with Micrurus nigro-
cinctus mosquitensis leads me to believe that both forms cannot
belong to nigrocinctus. The elongation of the body, and the great
increase in the ventral-subcaudal count causes me to consider alleni
as a distinct species, although it was described as a subspecies of
nigrocinctus originally.

The differences between this presumed subspecies and a. alleni
consists in narrower yellow borders on the black bands, the black
spotting on the yellow bands, more elongate red areas, and the
immaculate condition of the yellow and red areas on the venter.
(Compare the figure given herewith of alleni richardi with the
figure of alleni alleni (nigrocinctus alleni Schmidt) given by
Schmidt (Zool. Series Field Mus. Nat. Hist., vol. 20, 1936, p. 210).

The species is named for Richard C. Taylor who assisted in the
capture of the type.

Family VIPERIDAE

Three genera, Lachesis, Bothrops (composite) and Crotalus, are
recognized as occurring in Costa Rica.

Key to Costa Rican Genera of Viperidae

1. A series of rattles at termination of tail Crotalus

No series of rattles at tip of tail 2

2. Subcaudals partly double, partly quadruple Lachesis

Subcaudals never quadruple, but occasionally partly or wholly double or com-
pletely single (composite) Bothrops

Genus Bothrops Wagler

Bothrops Wagler, in Spix, Serpentum Barsiliensium . . . , 1824, p. 50.

Genotype: Coluber lanceolatus Lacepede.

The genus as here used is a complex of several genera or sub-
genera, most, if not all, of which have already been named. In-
sufficient comparative material prevents me from attempting to
resegregate the species into their proper genera.

There are ten species known in Costa Rica.

Key to the Species of Bothrops in Costa Rica

1. Supralabials seven, much enlarged; subcaudals divided; large terrestrial snakes

reaching two meters in length atrox atrox

Supralabials nine or more; subcaudals all or part (picadoi and nummifer), single. . 2

2. Two or three more or less erect pointed scales between supraocular scale and

eye ; tail prehensile, arboreal schlegelii

No erect spinelike scales between supraoculai scale and eye 3

Taylor: Review of Snakes of Costa Rica 173

3. Second labial forms part of border of loreal pit: green with a cream line on firsi

scale row (young with a blotched pattern) lateralis

Second labial not forming part of border of loreal pit; no cream line on first scale
row i 4

4. Rostral scale at least one and a half to one and two-fifths times as high as wide,
rising considerably above level of head to form a proboscislike appendage, di-
rected upward ; one canthal between internasal and preocular 5

Rostral scale reaching no higher than level of head, or at most to height of
canthal ridge "

5. Ventrals 130-145 ; color pattern of more or less distinct blotches nasuta

Ventrals 147-159; pattern of more or less regular small black marks lansbergii

6. Combined supraocular width one fourth (or less) width of head between eyes ;

huge snakes with body scales strongly keeled (and beaded in adults) 7

Combined supraocular width usually equal to one half width of head (very rarely
one third) ; scales keeled ; but never strongly beaded 8

7. Body very thick, short (700-800 mm.); rostral separated or at least partially sep-
arated from nasal by a row of intercalated scales nummifer nummifer

Body longer, thicker (1000 mm. or more); rostra! not even partially separated
from nasal by intercalated scales picadoi

8. Color dark green, most scales with black markings; tail prehensile nigroviridis

Color not green; most scales lacking black markings: tail not prehensile 9

9. Dorsal scales on head more or less irregularly enlarged, smooth ; black line from

eye god man i

Dorsal scales on middle of head nearly uniform, those on snout somewhat larger,

all keeled; a raised canthal ridge; no distinct black line behind eye; pattern of
quadrangular blotches ophryomegas

Bothrops schlegelii (Bertholdi

Trigonocephalus schlegelii Bert hold, Abh. Ges. Wiss. GSttingen. vol. 3, 1846, p. 13, pi. 1.

figs. 5-6 (type locality, Colombia).
B[othrops] schlegelii Jan, Elenco systematico degli ofidi, 1863, p. 127.
Bothrops (Teleuraspis) nigroadspersus Steindachuer, Sitz. Akad. Wien, Bd. 42, Abt. 1, 1870.

p. 348, pi. 8.
Bothrops ichlegeli Pieado, Serpientes Venenosas de Costa Rica .... 1931, pp. 76-80, figs.

32-35 (Santa Clara, Sarapiqui, Miravalles, Volcan Poas).

Two of the so-called color phases of this species were acquired
at Turrialba, Costa Rica. The specimen (M.N.H. No. 25162 J1 ) is
grayish to brownish black with a series of reddish-brown lighter
spots, about 22 on body and 6 or 8 on the tail ( indistinct posteriorly
on tail). The neck is whitish becoming gradually yellowish on an-
terior half and reddish brown on the posterior part, strongly de-
fined under tail. On the sides of the ventrals and usually includ-
ing one lateral scale is an irregular series of white spots, often
more or less quadrangular; on the anterior part of the body these
can be seen outlined faintly by pigment on the medial part of
ventrals.

The following scale characters are present: Scale formula
30-21-21-17, the outer row lacking keels; ventrals 160, subcaudals
53; dorsal head scales keeled very strongly, as well as those on
temporal region; canthal scales elevated forming low spines. Supra-
labials 8-8, second almost entirely fused with scale forming anterior
border of pit; three small loreals; fourth labial separated from

174 The University Science Bulletin

elongate subocular by two scale rows; infralabials 10-11, first two
touching first chinshields, others separated by very small scales;
second pair of chinshields separated from each other by a small pair
of scales; two elongate, more or less erect pointed scales emerge
between supraocular and eye, bordered at their base by a few
minute scales; scales bordering canthus, especially two anterior,
sharp-edged and partly erect; supraoculars large, longer than wide;
posterior part of nasal free, somewhat erect; three preoculars; nasal
very large, at most only partially divided.

M.N.H. no. 25159. Aside from being a brilliant yellow (faded
with small black spots in evidence, in the preservative) the scale
formula is 33-23-25-19-15, which compares poorly with that of the
male specimen (30-21-21-19-17). The ventral-subcaudal counts do
not differ greatly, the ventrals ( $ ) being 154/2, the subcaudals 54;
160-53 in the male specimen, with totals of 208*2 for the female and
213 for the male. Probably in the majority of snake genera, when
the scale rows on body normally remain the same in both sexes, the
female has a longer snout-to-vent measurement proportionally and
the number of ventrals is greater, the subcaudals fewer than in
corresponding males of a given population. Where a difference
normally exists in the scale rows about body, between the sexes,
and the larger number occurs in the females, the number of ven-
trals may be expected to be equal to those in males, and likewise
the totals of the ventrals and subcaudals should be nearly equal.
It would appear that the added scale rows represent a compensa-
tion so that adequate space is provided for the young in the female,
rather than by the more usual procedure of lengthening the body
cavity by moving the anal opening a distance of several ventrals
to the rear.

For a very long time Schlegel's B. nigroadaspersus has been con-'
sidered a synonym of B. schlegelii. If two forms are not involved it
may be that the major differences indicated by the two forms are
sexual, since the dark specimen taken at Turrialba is a male, the
brilliant yellow one a female.

It is not so easy to account for an increase in head size. Presum-
ably larger head size would allow engulfing larger prey and it is
not difficult to see the survival value of this, in allowing a wider
choice of food with the same effort. However, I strongly suspect
that in this case something more than a sex difference is involved.

The measurements of the two specimens are placed here for
direct comparison.

Taylor: Review of Snakes of Costa Rica 175

Measurements of Bothrops schl.egki.ii

Total

Snout

Head

Head

No.

Sex

length

to vent

Tail

length

width

25159

9

448

366

•82

27

21

25162

$

445

365

80

21

17

It is to be noted that a somewhat wider gape, made possible by
the larger size, would allow a somewhat larger labial surface and
hence the added infralabials. The greater roughness of the head
scales of males may represent a secondary sex character such as
keeling of scales on sides of anus, rugosities on chins, etc., that are
encountered in males of other genera. The more elongate tail is a
factor in its prehensility.

In Costa Rica the name Oropel is applied to the yellow form,
while the names Bocaraca or Toboba are applied to the darker-col-
ored forms.

Bothrops lateralis (Peters)

Bothriechis lateralis Peters, Monatsb. AUad. Wiss. Berlin, 1852, p. 074 (type locality, Costa

Rica).
Bothrops lateralis Picado, Serpientes venenosas de Costa Rica .... 1931, pp. 8.3-87, figs.

38. 39 (Altuias de Orosi ; Santa Maria de Dota ; Montanas de Navarro).

Five specimens of this well-known, easily-recognized species are
at hand. The two young specimens (M.N. H. Nos. 25160, 25161)
were collected at an elevation between 5000-6000 feet on Volcan
Poas at Isla Bonita, and two adults ( M. N. H. Nos. 25687. 256S8 )
were collected from the same locality; M. N. H. No. 25163 was taken
2 miles above Santa Cruz on Volcan Turrialba at an elevation of
between 6500-7000 feet.

The following characters are evident in the adults: rostral nearly
triangular, a little broader than high; nasal single with a slight
suture evident; second labial enters pit and forms anterior boun-
dary; loreals, three (rarely four), the upper largest between nasal
and upper preocular; supralabials separated from subocular by one
scale row.

The scale formula for both sexes is (33-30) 21-21-17; supralabials
10 (rarely 9 or 11); infralabials 12 (more infrequently 11, or 10),
three of these touching the anterior chinshields; four pairs of chin-
shields, all save first pair separated by small scales; usually 3 (rarely
4) postoculars; subocular enlarged.

Body scales rather pointed behind, keeled save on outer row. No.
25163 is uniform green above with a few small yellow flecks; green-
ish yellow on ventral surface, median area more yellow than outer
parts; a whitish line beginning on neck borders the outer ventral

176 The University Science Bulletin

edge and lower edge of first scale row; labials yellowish, chin
largely cream; tail bluish green to blue.

Nos. 25687 and 25688 agree closely with the preceding save that
the dorsal yellow or light flecks tend to form a transverse series of
spots on each side at rather regular intervals, their edges occasion-
ally black, and tending to alternate on the two sides; approximately
22-24 on body, 8 on tail. Young specimens ( 25160-61 ) are light lav-
ender (in preservative) with a series of black bars, lighter-edged
behind; these may be continuous across back or alternate (there are
23 on body, 6-f- on tail); between the bars there are other indefi-
nite black markings. These color differences are those of age only.

The species is known in Costa Rica under the name Culebra Lora.
It has been reported from numerous localities.

I have examined specimen A. M. N. H. No. 17372 and specimen
U. S. N. M. No. 37730, both from "Costa Rica."

Table of Data on Bothrops lateralis (Peters)

No.

Ssx

Ventrals

Subcaudals

Total length

Tail length

25688

$

166

66

632

105

25687

9

155

58

760

110

25160

9

164

61

255

44

25161

9

169

61

234

43

25163

9

161

54

715

109

Bothrops nigwviridis nigroviridis (Peters)

Bothriechis nigroviridis Peters, Monatsb. Akad. Wiss. Berlin, 1859. p. 278, pi. — , fig. 4.
Bothrops nigriviridis F. Miiller, Verh. Nat. Ges. Basel, Bd. 6, 1878, p. 401; Pieado, Serpientes

Venenosas de Costa Rica . . . , 1931, pp. 80-85, figs. 3G-37 (Yolcan de Barba ; vertientes

del Rio Sarapiqui; faldas de Poas, altos de La Palma).
Lachesis nigroviridis Boulenger, Catalogue of the Snakes in the British Museum, vol. 3, 1896,

p. 568.

A single specimen (M. N. H. No. 25699 of this species was ob-
tained at Isla Bonita at about 5500 ft. elevation. This form has a
rather short head, more or less rounded, the rostral very slightly
broader than high; nasals large, single, but sometimes partially
divided; five or six loreals; three preoculars, upper large, separ-
ated from nasal by upper enlarged loreal; scales on snout imbri-
cate, more or less symmetrical, smooth, about four or five rows
between the enlarged supraoculars; latter scales large, the outer
edges slightly elevated; one scale below supraoculars bordering
eye; one or two postoculars; a large subocular in contact with or
separated by one scale from lower preocular; supralabials 9-9, the
second separated from pit by a large scale, the third separated from
it by a small scale; infralabials 9-9, the first three touch the first
pair of chinshields which are larger than two succeeding pairs; all

Taylor: Review of Snakes of Costa Rica 177

pairs of chinshields without intercalated scales; scale formula, 25-
21-19-19-17, all keeled except outer row; ventrals, 143; subcaudals
(single) 50; anal single.

The specimen here described is somewhat discolored by preserva-
tive. In life, however, the species is green, strongly speckled with
black; a black streak on head from canthus to angle of mouth pass-
ing above the eye; top of head may be streaked with black; yellowish
beneath, some or all of the shields black edged.

The shed epidermis of this snake is strongly pigmented but in
each scale there is a small unpigmented area. Dr. Picado ( loc. cit. ) ,
gives excellent photographs of several living specimens, and notes
that it occurs at elevations of more than 2000 m. The Costa Rican
name is Vibora de Arbol.

The following specimens have been examined also: A. M. N. H.
No. 17283, Volcan Barba; U. S. N. M. Nos. 32580-32581, Pico Blanco.

Bothrops nasuta Bocourt

Bothrops nasutus Bocourt, Ann. Sci. Nat., ser. 5, vol. 10, 18(38, p. 202 (Pansas, bank of Polo-
chic, Guatemala); Amaral, Bull. Antiv. Inst. Amer., vol. 3, 1929, pp. 25-27, pi. 3, a-d,
text fig. 7 (Zent, Limon and Sipurio, C. R.) ; ? Picado, (part.) Serpientes Venenosas de
Costa Rica . . . , 1931, pp. 69-73, figs. 30, 31.

Snout pointed, turned up at end with a sharp canthus rostralis;
rostral usually VA times as high as wide; canthus formed from one
elongate intemasal, one canthal and upper part of preocular, their
edges rough. Nasal not completely divided; a pair of elongate in-
ternasals, elevated anteriorly; upper head scales imbricate, strongly
keeled; large supraoculars separated by from five to seven scale
series; two or three scale rows between labials and eye; subocular
sometimes broken; temporals all keeled; supralabials nine to eleven,
none bordering the sensory pit; scales of body 23-27 (23-25); ven-
trals 130-145; anal entire; subcaudals single, 24-35; body slender,
tail short, not prehensile.

Yellowish brown, pale brown or grayish above, with a dorsal
series of 13-20 large, dark brown, black-edged rhomboidal or squar-
ish spots, usually divided on median line by a narrow white or
orange line; occasionally the spots on the two sides tend to alter-
nate; sides of head blackish; belly powdered with brown with or
without whitish spots.

The species is known to occur on the Carribean drainage area.
In Costa Rica it is known under the name Toboba Chinga.

I have examined the following specimens: A. M. N. H. Nos.
12—3286

178 The University Science Bulletin

17287, 17308, Sarapiqui; No. 17332, Sipurio; U. M. M. Z. No. 17306,
San Jose.

It has also been reported from Zent and Limon on the east coast.

Bothrops ophryomegas Boconrt

Bothrops ophryomegas Bocourt, Ann, Sci. Nat., ser. 5, vol. 10, 1868, p. 201 (type locality,
Occidental slope of Esciuntla range, Guatemala); Amaral, Bull. Antiv. Inst. Amer., vol. 3,
no. 1, 1929, pp. 23-24.

Data for this is taken from Amaral's redescription (loc. cit.).
Body long and slender, tail short, not prehensile; head wide, snout
not turned up; canthus rostralis raised and sharp, formed by one
internasal, two canthals and upper part of preocular; rostral slightly
higher than wide; internasals short and straight; canthals double
and small; supraoculars large with ridgelike border; head scales
keeled; supralabials 9-10; body scales in 25-27 rows; ventrals 166-
173; subcaudals entire, 32-39.

Brown above with 26 to 40 small black markings placed in pairs
along the vertebral line alternating with or opposite to those of
other side; flanks with darks spots in pairs; corresponding to ver-
tebral markings; belly light, speckled or transversely marked with
brown; sides blotched with dark; head with a dark line behind
eye and dark markings on the top; lips with light spots.

I have examined specimens of this species as follows: U. M. M. Z.
No. 83183 Esparta; No. 83184 Puntarenas; U. S. N. M. No. 37728
Esparta and No. 37729 Jimenez.

This form is regarded by Amaral as belonging to the arid, west
coast areas of Central America.

Bothrops lansbergii (Schlegel)

Trigonocephaly lansbergii Schlegel, Mag. de Zool., 1841, 1-3, Rept. pi. 1 (type locality, Tur-

baco, Colombia).
Bothrops lansbergii Jan, Elenco sistematico degli ofidi, 1863, p. 127 (Costa Rica, etc.);

Amaral, Bull. Antiv. Inst. Amer.. vol. 1, no. 1, 1927, p. 22, and ibid, vol. 3, no. 1, pp.

19-27, fig 1; Dunn, ibid, vol. 2, no. 2, 1928, pp. 29-30; Loveridge, ibid, vol. 2, no. 3,

pp. 64-65.

This species is represented in my material by a single female
specimen (M. N. H. No. 25689) obtained at Turrialba, July 11,
1947. The characters of this specimen are: body rather thick and
short; tail short; rostral vertical, the width in height one and one
half times; three ridged canthals, the anterior elongate, bordering
its fellow on median line behind rostral; a small median scale fol-
lows their common suture; third scale on canthus is the very large
upper preocular; supraoculars border orbit, one and a third times
as long as wide; six scales between the supraoculars; head scales

Taylor: Review of Snakes of Costa Rica 179

dimly keeled, except perhaps the first row on snout; nasal large,
with a small entrant suture from top, the posterior part of scale
concave; seven small and two large loreal scales; second preocular
bordering upper part of pit broken into three scales; lower pre-
ocular small; two rows of small scales between anterior labials and
pit; four rows of scales between labials and eye (or 3 rows and
subocular, latter may be broken); supralabials 10-9, fourth dis-
tinctly largest; infralabials 11-10, three touching first pair of chin-
shields which are more than twice as long and at least three times
area of second pair; anterior chin scales, labials, nasal and rostral
more or less covered with small tubercles; scale formula; 39-25-23-
23-19; ventrals, 139; subcaudals, 28, single; anal single.

Dorsal color bluish to ash gray with small discrete black spots,
the ventral coloration being lavender; a few white flecks present
low on sides, extending occasionally onto ventrals; dark spots ar-
ranged in groups of four as if marking the four corners of a quad-
rangle, and opposite or (occasionally) alternate with a similar
grouping on opposite side; 15 present on each side. On underside
of jaw some light lavender marks, the one beginning back of jaw
angle forming an irregular line for a short distance.

The name by which this species is known in Costa Rica is La
Tamaga; it is said to reach half a meter in length only rarely. This
specimen has a total length of 294 mm.; tail, 31 mm.

Males and females differ somewhat in the character of the can-
thus. In females this rough, serrated appearance is lacking almost
completely. The known ventral range is 130-145; subcaudal range,
24-35, according to Amaral.

Bothrops atrox atrox (Linnaeus)

Coluber atrox Linnaeus, Museum Adolphus Fricleriei regis, vol. 1, 1754, p. 33, pi. 22, fig. 2,

and Systema Naturae, vol. 1, 176G. p. 383 (type locality ?).
Bothrops atrox {part.) Dumeril, Bibron and Dumeril, Erpetologie Generale, vol. 7, 1854,

pp. 1507-1509.

This great pit viper, known commonly under the name La Tercio-
pelo, is, according to Sr. C. Picado-T., the most feared snake in the
country of Costa Rica, because of its deadly bite and great size. It
is reported as reaching a length a little greater than two meters.

A specimen, M. N. H. No. 25677, was killed close to the central
station at Los Diamantes (rubber plantation) near Guapiles in
early September 1947, and presented to us by Mr. Wallace Manis,
the Director of the plantation and our genial host.

Scale data on this specimen follow: rostral as wide as high, vis-
ible above as a tiny triangle bent flat on tip of snout; two canthal

180 The University Science Bulletin

scales form a bordering keel together with upper edge of the upper
preocular; supraoculars very large, twice as long as wide, separated
from each other by six or seven scale rows; upper surface of snout
a little more elevated than interocular area; nasal completely di-
vided; two loreals, upper largest; second labial forming anterior
border of pit, the scale showing a partial division; supralabials
7-7, all very large save first two; subocular irregular, elongate,
lower preocular divided vertically; two scales between subocular
and second labial; one scale row between fourth labial and subocu-
lar; dorsal head scales keeled; lateral scales of head smooth save
the upper temporals; ten lower labials, three touching first chin-
shields, which are a little longer and considerably larger than sec-
ond pair; scales strongly keeled; the scale formula, 33-27-27-19;
anal single; ventrals 192; subcaudals divided, 67; total length, 1773
mm.; the tail, 223 mm.

It has been reported from numerous localities in the lowlands.

Bothrops pieadoi (Dunn)

Trimeresurus nummifer pieadoi Dunn, Proc. Biol. Soc. Washington, vol 52. 1939, pp. 165-168,

(La Palma, C. R., elev. 4500 ft.).
Bothrops pieadoi Smith and Taylor, Bull. U. S. Nat. Mus., No. 187, Oct. 5, 1945, pp. 182-183.

This large species, which in size stands between nummifer and
atrox, seems to occupy a region on the central plateau of Costa Rica
and the surrounding mountains. The specimens in the collection
M. N. H. (Nos. 25672-75) were obtained at Isla Bonita (A. C.P.)
on the eastern slope of Volcan Poas at an elevation between 5000
and 6000 ft. In this area it is confused with the memo de piedra
(nummifer), under which name it is known.

Rostral more than once and a half as wide as high, reaching
level of snout, visible above as a rather narrow line; nasals com-
pletely divided, the anterior larger part bordering rostral without
any intercalated scales; supraocular broken up and a small narrow
elongate part of it, differentiated from other head scales, borders
eye and is separated from its fellow by eleven rows of lightly keeled
head scales; canthal scales not forming a shelf; a large loreal be-
tween very large preocular and posterior nasal; one to three small
loreals below this scale; subocular elongate, narrow; three post-
oculars; a large triangular upper preocular, and small lower which
borders pit; supralabials 9-10, the first, second and third separated
from scales surrounding pit by one scale row, fourth and fifth sepa-
rated from labials by three or four scales; infralabials 9-11, two touch
first pair of chinshields which are more than twice length of second

Taylor: Review of Snakes of Costa Rica

181

or third pairs, and three times area of second pair; the first pair of
infralabials are separated in four out of six cases, and in these ex-
ceptions they are very narrowly in contact. All scales are keeled
or beaded, the keel widening and forming a low rounded mound
posteriorly on the scale. This is especially prominent on the fe-
males, less so in males.

The type of this species which I have examined is a young speci-
men with the markings differing slightly from specimens here re-
corded. One other specimen, R. C. T. No. 695, is included on the
data sheet.

Table of Data for Bothrops picadoi (Dunn)

Scale

Sub-

Supra -

Infra-

No.

Sex

formula

Ventrals

caudals

labials

labials

695

6

28-25-21

146

38

9^10

11-11

25675

9

30-25-21

148

32

10-10

11-11

25676

9

30-25-21

152

23+

10-10

11-11

25673

9

33-25-21

146

33

10-10

11-12

25672

9

33-25-21

149

33

9- 9

11-11

25674

6

31-25-21

146

40

9- 9

10-10

Table of D.

\ta for Both

ROPS PICADOI

(Dunn) (concluded)

First

Infralabials

infralabials

touch

Total

Divided

No.

separated

chinshields

length

Tail

subcaudals

695

no

2

1202

120

11

(5, 14-24)

25675

yes

2

955

89

0

25676

yes

3

930+

68+

(1

25673

yes

2

257

27

1)

25672

no

2

269

30

L3

(12-25)

25674

yes

2

249

29

2

(last two)

Bothrops nummifer nummifer (Riippell)

Atropos nummifer Riippell, Verz. Mus. Senck., Amph., 1845, p. 21 (type locality, Mexico).
Bothrops nummifer Jan, Elenco Systematico degli Ofidi, 1863, p. 126.

This extremely thick, short snake is known in Costa Rica under
the name mono de piedra. It ordinarily reaches but little more than
half a meter in length, but the body is as thick as that of picadoi
of nearly double its length. The most distinctive characteristic of
the snake is the strong ridge on the back, especially prominent on
the anterior half of the body, bearing three scale rows, a median
and two adjoining lateral rows all very heavily keeled or tubercled.
One specimen, M. N. H. No. 25678, was obtained at the Inter-Amer-
ican Institute of Agriculture, three km. west of Turrialba.

In this the following scale characters obtain: rostral triangular,
not visible above, distinctly wider than high, separated from nasal
on right side by three intercalated scales, on left by two scales only,
thus allowing rostral and nasal to touch at one point; nasal divided

182 The University Science Bulletin

completely, the anterior part the larger and somewhat convex; dor-
sal head scales heavily keeled, the anterior ones tending to bend
down to touch rostral in front, and in canthal and supraorbital
regions, to form slightly projecting shelves; an elongate loreal bor-
ders canthal edge touching first supraocular and posterior nasal;
eight or nine small loreals; supraocular broken into several small
scales, not or scarcely larger than adjoining scales; subocular nar-
row, elongate; two or three postoculars; only one large preocular;
the two scales below it are pushed forward more than their length
in front of the eye; supralabials 9-10; third separated from facial
pit by two or three scale rows, fourth and fifth separated from
subocular by three rows; infralabials 12-13, three touch first chin-
shields which are double size of two succeeding pairs.

Scales keeled, the scale formula being 30-25-19; ventrals 122; sub-
caudals 35, first two pairs divided, others single. The total length
is 790 mm.; tail, 91 mm.; head width, 45 mm.; head length, 57 mm.;
height of body, 55 mm.

The basal pattern gray with 16 saddlelike darker brown rhombs
on back, each with a somewhat lighter center and each with a nar-
row lateral extension onto the ventrals; low on sides a series of
small black spots alternating with the rhombs; frequently rhombs
are contiguous middorsally and bordered by lighter gray color, es-
pecially along the upper parts; base of tail with two discernible
rhombs but the remainder of tail black above and below; a diag-
onal black stripe from eye to behind jaw angle; chin, throat and
anterior part of body white on ventral surface; but posterior part
of body clouded with pigment and numerous more or less quad-
rangular black spots.

The specimen when in captivity was quiet, save that on being
disturbed by the approach of anyone, the head was moved in a
series of slight rapid jerks. When dissected it was found to contain
27 embryos.

This is a wide-ranging species occurring as far north as San Luis
Potosi, Mexico. Known variation in ventrals, 121-135; subcaudals
26-37, the higher numbers being from the northern part of the
range.

The species has been reported in the literature from El General,
Monte Redondo, Chitaria, Cariblanco, Peralta, Guapiles, and Si-
quirres, in Costa Rica.

Taylor: Review of Snakes of Costa Rica 183

Bothrops godmani (Giinther)

Bothriechis godmanni Giinther, Ann. Mag. Nat. Hist., vol. 12, 18C3, p. H < ; 4 . pi. 6, fig. G, G'
(near Duefias, and on other parts of the tableland of Guatemala). (The speeies was named
for Godman. The original "Godmanni" constitutes an error of transliteration.)

Bothrops godmanni Amaral, Mem. Inst. Butantan, tome 4, 1929, p. 235 (or 109).

This species, recently collected in Panama, occurs to the north of
Costa Rica and perchance will eventually be found in Costa Rica. I
know of no records to date. It may be characterized as follows:
rostral broader than high, extending to upper level of head, equally
as high as the anterior nasals, but broader than high; supraoculars
large, often with other somewhat enlarged scales (frontal or pa-
rietals) appearing; number of scales between supraoculars from
three to seven; scales of top of head keeled; nasal divided, the
anterior part largest and not separated from rostral by intercalated
scales; a large preocular separated by the large loreal from the
posterior nasal; supralabials 9-9, the second not joined to pit scale,
or bordering pit; a single series of scales between labials and elon-
gate narrow subocular; 10-11 infralabials, four touching chinshields;
second chinshields small, separated by a pair of scales; scale rows
23-21-21-19(17), keeled except outer row; ventrals 146; anal sin-
gle; subcaudals 36, none divided.

Above olive gray with a black stripe beginning behind eye and
reaching onto neck, bordered above and below by whitish, touch-
ing two or three supralabials; side of head, labials, and chin gray-
ish white; body gray-olive with a series of median blotches of
brownish black, darker edged, often contiguous or confluent; a
series of lateral spots usually bordered with whitish gray. On outer
edge of ventrals a series of spots that may be partly edged with
whitish, becoming obsolete or merged with dark ventral coloration.
Throat grayish white; body becoming rapidly darker until on the
latter half body nearly uniform black. (U. S. N. M. No. 24782.)

The measurements are: total length 550 mm.; tail 65 mm.

The known variation in ventrals is from 142 to 148- subcaudals
28 to 36.

Genus Crotalus Linnaeus

Crotalus Linnaeus, Systema Naturae, 10th ed., vol. 1, 1758, p. 214.

Genotype: Crotalus horridus Linnaeus.

A single species is known to occur in Costa Rica.

Crotalus terrijicus durissus Linnaeus

Crotalus durissus Linne, Systema Naturae, vol. 1, p. 372, 1766 (type localitv unknown).
Crotalus tcrrificus Picado, Serpientes Venenosas de Costa Rica, 1931 pp 43-53 has 1 and
14-19. ' s"

This large rattlesnake formerly had a range well onto the plateau,

184 The University Science Bulletin

specimens having been obtained some distance east of Cartago. It
has become rare or has disappeared largely from plateau country
according to Dr. Picado.

The species may be readily recognized by the presence of a
series of rattles on the tip of the tail. In the very young specimens
only a small button is present.

A specimen, U. S. N. M. No. 13534, from San Jose, Costa Rica,
has been examined. The scale formula is 26-25-25-19; ventrals 172,
subcaudals 25; the first, and last three subcaudals are divided,
others single; 14 infralabials; four rows of scales between labials
and eye; internasals and prefrontals present; supraoculars large,
separated anteriorly by two scales, posteriorly by five scales; 25
scales across head between last supralabials.

Genus Lachesis Daudin

Lachesis Daudin, Histoire Naturelle des Reptiles, vol. 5, 1803, p. 349.

Genotype: Lachesis tnuta (Linnaeus.)

A single species is known to occur in Costa Rica.

Lachesis muta stenophrys Cope

Crotalus mutus Linnaeus, Systema Naturae, vol. 1, 1766, p. 373 (? Surinam).

Lachesis stenophrys Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875),

p. 152 (Sipurio, Costa Rica).
Lachesis muta Picado, Serpientes Venenosas de Costa Rica, San Jose, Costa Rica, 1931, pp. 37-

43, figs. 10-12 (Rio Banana).

This large Bushmaster or Cascabela muda seems to be a rare
snake in Costa Rica. It reaches a greater size than any other of the
poisonous snakes of the country. Picado shows a photograph of a
specimen 2150 mm. in length.

This species may be recognized by the following characters: ros-
tral triangular, broader than deep, the muzzle short and depressed;
nasal divided; scales on top of head flat, hexagonal and faintly
keeled; 12 series of scales between the supraoculars which are of
moderate size, but narrow; nine supralabials, of which the third is
largest; second low, separated from the p?!t; pit bordered by three
scales, the superior of which borders the two preoculars; the in-
ferior, wider, stands on the third labial and the anterior, which is
subcrescentic, and rests on the second supralabial; four rows of
scales separate orbit from labials; infralabials 13, first and second
touch chinshields; only one pair of chinshields, squarely truncate
anteriorly, narrowly rounded behind. Ventrals 200; subcaudals 49
(32 double, 17 quadruple); caudal spine well developed.

Taylor: Review of Snakes of Costa Rica 185

Color in preservative, fawn-brown with 23 reddish-brown me-
dium rhombs on the dorsal region. The lateral corners of these are
dark spots, sometimes isolated and do not extend below fifth row
of scales; on middle of body rhombs with pale centers; posteriorly
they are darker and become confluent into a zigzag band. Tail dark
brown, with narrow, light cross-bands. Lower surfaces greenish yel-
low except throat and chin which are white. A black band extends
from eye above labials, and is broken upon neck into a series of
black spots. Top of head uniform brown.

Boulenger has placed Cope's L. stenophrys in the synonymy of
L. mutus. However the very low ventral count (Boulenger gives
200-230 throughout the range with counts of 202 and 206 for
Panama [200 for type of stenophrys] and higher counts 223-226 for
contintental South America) suggests that it should be considered
a separable form. Also there are some differences in color and pat-
tern.

BIBLIOGRAPHY
Amaral, Afranio do

1927. Studies of Neotropical Ophidia. V. Notes on Bothrops lansbergii and
B. brachystoma. Bull. Antiv. Inst. Amer., vol. 1, no. 1, 1927, p. 22.

1929. Studies of Neotropical Ophidia. XII. On the Bothrops lansbergii
group. Bull. Antiv. Inst. Amer., vol. 3, 1929, pp. 19-27, pis. 1-3
and text fig. 7.

1929. Studies of Neotropical Ophidia. XIII. A new colubrine snake in the
collection of the Vienna Museum. Bull. Antiv. Inst. Amer., vol. 3,
no. 2, 1929, p. 40. (Helicops wettsteini described).

1929. Estudos sobre ophidios neotropicos XVIII. Lista remissiva dos
ophidios da regiao neotropica. Mem. Inst. Butantan, Tomo IV,
1929, pp. 129-271. (This purports to be a complete list of Neo-
tropical snakes. )

Boulenger, George Alhert

1893, 1894, 1896. Catalogue of the Snakes in the British Museum (Nat-
ural Historv), vols. I-III. Vol. I, 1893, pp. I-XII1, 1-448, pis. 1-28,
text figs. 1-26; Vol. II, 1894, pp. I-XI, 1-382, pis. 1-20, text figs.
1-24; Vol. Ill, 1896, pp. I-XIV, 1-727, pis. 1-25, text figs. 1-37.

Cope, Edward Drinker

1871. Ninth Contribution to the Herpetology of Tropical America. Proc.
Acad. Nat. Sci. Philadelphia, 1871, pp. 200-224, pi. 17, figs. 1-7
( Bhadinaea serperastra, Colobognathus bracliycephalus, and Col-
obognathus dolichocephahis are the new snakes described. A total
of 34 snakes are listed for the country. ) .

1876. On the Batrachia and Reptilia of Costa Rica. Journ. Acad. Nat.
Sci. Philadelphia, ser. 2, vol. 8, 1876 ( 1875), pp. 93-154, pis. 23-28.
(A most important work. Sixty species of snakes are listed. Sev-
eral forms are described as new. The larger number of specimens
were from a collection made by William Gabb in southern and
southeastern Costa Rica. )

1879. Eleventh Contribution to the Herpetology of Tropical America.
Proc. Amer. Philos. Soc, vol. 18, no. 104, Aug. 11, 1879, pp. 261-
277. (Lists three snakes: Scolecophis zonatus Hallowell; Coluber
triaspis Cope and Porthidium nasutum Bocourt. )

186 The University Science Bulletin

1887. Catalogue of Batrachians and Reptiles of Central America and
Mexico. U. S. Nat. Mus. Bull. 32, 1887, pp. 1-96.

1893. Second Addition to a Knowledge of the Batrachia and Reptilia of
Costa Rica. Proc. American Philos. Soc, vol. 31, Dec. 3, 1893
( 1894), pp. 333-347. (This paper treats of a collection made or at
least sent by Mr. George K. Cherrie of the National Museum of
San Jose. Three forms are described as new; several are new
to Costa Rica; and three have been previously reported from
Costa Rica. )

1894. Third Addition to a Knowledge of the Batrachia and Reptilia of
Costa Rica. Proc. Acad. Nat. Sci. Philadelphia, 1894, pp. 194-206.
(This deals with collection from the Museo Nacional de Costa
Rica. ( Four forms are described as new, and one form reported for
first time for the country. Trimetopon pliolepis, Drymobius pauci-
carinatus, Leptophis ultramarinus, and Poganaspis ruficeps are de-
scribed. Enulius torquatus is reported.

Dumeril, A., Bocourt, Fermin, and Mocquard, Dr. F.

1870. Etudes sur les reptiles; Mission Scientifique au Mexique et dans
lAmerique Centrale. 1870-1909, livraisons 1-17, pp. 1-1012, pis. 99.

Dunn, E. R.

1924. Amastridium a Neglected Genus of Snakes. Proc. U. S. Nat. Mus.,
1924, pp. 1-3.

1928. Notes on Bothrops lansbergii and Bothrops ophryomegas. Bull.
Antiv. Inst. Amer., vol. 2, no. 2, 1928, pp. 29-30.

1930. New Snakes from Costa Rica and Panama, Occ. Papers Boston Soc.
Nat. Hist., vol. 5, Sept. 15, 1930, pp. 329-332. ( Describes Trime-
topon simile from Costa Rica. )

1931. Some Central American Snake Genera, Copeia, no. 4, 1931, p. 163.

1932. Notes on the Blind Snakes from Lower Central America. Proc. Biol.
Soc. Washington, vol. 45, Oct. 11, 1932, pp. 173-176.

1932. The Status of the Snake Genus Rhadinaea Cope. Occ. Papers Mus.
Zool., Univ. Michigan, no. 251, Oct. 20, 1932, pp. 1-2.

1935. The Snakes of the Genus Ninia. Proc. Nat. Acad. Sci., vol. 21,
no. 1, Jan., 1935, pp. 9-12.

1936. Notes on North American Leptodeira. Proc. Nat. Acad. Sci., vol.
22, 1936, pp. 689-698.

1937. The Snake Genus Enulius Cope. Proc. Acad. Nat. Sci. Philadelphia,
vol. 89, 1937 (Jan. 14, 1938), pp. 415-418.

1937. Notes on Tropical Lampropeltis. Occ. Papers Mus. Zool. Univ.
Michigan, no. 353, Apr. 28, 1937, pp. 1-11.

1937. The Amphibian and Reptilian Fauna of Bromeliads in Costa Rica
and Panama. Copeia, no. 3, Nov. 19, 1937, pp. 163-167.

1937. New or Unnamed Snakes from Costa Rica. Copeia, no. 4, Dec. 31,

1937, pp. 213-215. {Lemwdoplvs taeniurus juvenalis, Conophis

nevermanni, Trimetopon viquezi.)

1938. A New Rhadinaea from Central America. Copeia, no. 4, Dec. 10,

1938, pp. 197-198. (Describes Rhadinaea persimilis and proposes
Rhadinaea giintheri for Tachymenis decipiens [Giinther].)

1939. A new Pit Viper from Costa Rica. Proc. Biol. Soc. Washington,
vol. 52, Oct. 11, 1939, pp. 165-166. (Describes Trimeresurus
nummifer picadoi [= Bothrops picadoi].)

1940. Notes on some American Lizards and Snakes in the Museum of
Goteborg. (sic. ) Herpetologica, vol. 1, 1940, pp. 189-194.

1940. New and Noteworthy Herpetological Material from Panama. Proc.
Acad. Nat. Sci. Philadelphia, vol. 92, 1940, pp. 105-122, pi. 2.

Taylor: Review of Snakes of Costa Rica 187

1942. New or Noteworthy Snakes from Panama. Notulae Naturae, Acad.
Nat. Sci. Philadelphia, no. 108, 1942, pp. 1-8.

Dunn, E. R., and Bailey, Joseph R.

1939. Snakes from the Uplands of the Canal Zone and of Darien. Bull.
Mus. Comp. Zool., Harvard Coll., vol. 86, no. 1, Oct. 1939, pp. 1-22.

Dunn, E. R., and Tihen, J. A.

1944. The Skeletal Anatomy of Liotyphlops alhirostris. Journ. Morph.,
vol. 74, Mar. 1944, pp. 287-294, pi. 1.

Fowler, H. W.

1916. Cold-blooded Vertebrates from Costa Rica and the Canal Zone.
Proc. Acad. Nat. Sci. Philadelphia, vol. 68, 1916, pp. 389-414.

Gunther, Albert C. L. G.

1872. Seventh account of new Species of Snakes in the Collection of the
British Museum. Ann. and Mag. Nat. Hist., ser. 4, vol. 9, 1872,
pp. 13-37, pis. 3-6. ( New forms are Ablabes gracilis, Geophis
moestus, Genus Microdromus, Microdromus virgatus, Leptognathus
annulatus, Genus Diplotropis, DipJotropis bilineata.)
1885-1902. Biologia Centrali-Americana; Reptilia and Batrachia, 1885-
1902, the part dealing with snakes, pp. 85-195, published between
Apr. 1893 and Aug. 1895, pis. 33-59. (The Costa Rican collections
of the British Museum were made largely by Mr. H. Rogers and
and Mr. C. F. Underwood. Species reported by other workers are
given; a total of approximately 76 forms are listed. )

Oliver, J. A.

1942. A Check List of the Snakes of the Genus Leptophis, with Descrip-
tions of New Forms. Occ. Papers Mus. Zool. Univ. Michigan, no.
462, 1942, pp. 1-19.

1948. The Relationships and Zoogeography of the Genus Thalerophis
Oliver. Bull. Amer. Mus. Nat. Hist., vol. 92, art. 4, Nov. 22, 1948,
pp. 157-280, text figs. 1-13, pis. 16-19, tables 1-13.
Peters, W.

1859. t)ber die von Hrn. Dr. Hoffmann in Costa Rica gesammelten und
aus das konigl. zoologische Museum gesandten Schlangen. Monatsb.
konig. preuss. Akad. Wiss. Berlin, 1859 March, pp. 275-278, 1 pi.
(Three new genera and three new species are described: Colobo-
gnathns Hoffmanni, Hydromorplius concolor and Bothriechis nigro-
viridis. Fourteen other species are listed. )

1861. Drei neue Schlangen des k. zoologischen Museums aus America vor
und fiigte hieran Bemerkungen iiber die generelle Unterscheidung
von anderen bereits bekannten Arten. Monats. konig. preuss. Akad.
Wiss. Berlin, Oct. 1860 (1861), pp. 517-521, 1 pi. (Typhlops
[Helminthopsis] frontalis sp. nov. is described from Costa Rica.)

PlCADO T, C.

1931. Epidermal microornaments of the Crotalinae. Bull. Amer. Anti\ .

Inst, vol. 4, no. 4, 1931, p. 104-105, figs. 1-10.
1931. Serpientes Venenosas de Costa Rica, sus Veninos Seroterapia Ofi-

dica. 1931. San Jose de Costa Rica. pp. 1-219.
Viquez S, lic. Carlos

1935. Animales Venenosos de Costa Rica. San Jose, 1935, pp. I-IV,

1-313, figs. 1-83 (pages 1-84 deal with serpents).

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ous text figs., unnumbered.

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Inst. Def. Cafe Costa Rica, vol. 11, 1942, pp. 608-611 (not ex-
amined ) .

188 The University Science Bulletin

Werner, Franz

1908. fiber neue oder seltene Reptilien des Naturhistorischen Museums
in Hamburg. Hamburg Jahrb. Wiss. Anst., vol. 26 (1908), Beih.
2, 1909 (1910), pp. 205-247. (Two new Costa Rican species de-
scribed: Streptophorus stibtessellatus and Streptophorus oxynotus.)

Wettstein, Otto

1934. Ergebnisse der osterreicbischen Costa Rica— Expedition 1930. Die
Amphibien und Reptilien. Sitz. Akad. Wiss. Wien. Math.-Naturw.
Abt. 1, Bd. 143, Heft 1-2, 1934, pp. 1-39. (30 species of Costa
Rican snakes are listed. )

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXIV, Pt. I] October 1, 1951 [No. 2

A New Veracrucian Salamander

BY

Edward H. Taylor,
Department of Zoology, University of Kansas

Recent collections made in Mexico by Mr. Walter W. Dalquest*
for the Museum of Natural History of Kansas University contain
a series of salamanders belonging to the genus Bolitoghssa, from
the lowlands of southern Veracruz. The species is regarded as new
and is herewith described.

The specimens were collected at night on the lower edge of a
great limestone cliff, the exposed section of the outcrop being
"nearly one hundred feet in thickness." Individuals occupy the
small nooks and crannies in the limestone in the daytime and at
night they move about the face of the cliff presumably in search
of food. They are not or only slightly disturbed by the light and
are captured with comparative ease.

No other species of salamanders, except Bolitoghssa phtydactyla
( Gray ) , has been taken in the immediate vicinity of the type local-
ity; however, Bolitoglossa rufescens probably occurs also. The
little-known Bolitoglossa yucatana (Peters) may occur in this area,
as it occupies terrain in limestone regions of the Yucatan peninsula.
The differences from this form are discussed later, since the two
forms seemingly belong in the same section of the genus.

Bolitoglossa veracracis sp. nov.

Type.— U.K.M.N.H. No. 26941; collected at a point 35 km. SE
of Jesus Carranza (Santa Lucretia), elev. approx. 350 feet, Vera-
cruz, Mexico, April 6, 1949. Walter W. Dalquest, collector.

* The Museum wishes to acknowledge financial assistance from the University of Kansas
Endowment Association for Mr. Dalquest 's collecting in Mexico.

(189)

190

The University Science Bulletin

Paratypes.-V.KM.N.H. Nos. 26933-26940; 26942-26953; same
date, locality and collector.

Diagnosis. — A medium salamander, maximum known length
(snout to end of vent), 55 mm.; in adults, width of head in head-
body length 5.5-6.25 times; length of head in head-body length
5 to 5.7 times; costal grooves 13, counting one in axilla and one in
groin; tail with a strong basal constriction; tail less than head-body
length; hand and foot palmate, spread widely, somewhat emargi-

Bolitoglossa veracrucis sp. nov.

Fig. 1, UKMNH No. 26953 paratype. Fig. 2, No. 26938 paratype. Fig. 3,
No. 26941 type. All from 35 km SE of Jesus Carranza (Santa Lucretia), Vera-
cruz. ( elev. approx. 350 ft. ) ( See text for actual size ) .

nate between digits; vomerine teeth usually in a single continuous
or in a somewhat irregular row, 12-19 in number, curving back
slightly and nearly meeting mesially; four premaxillary teeth pierce
lip in males.

Description of the type. — Head broad, depressed, the snout
truncate protruding beyond mouth more than 1 mm.; in dorsal
profile the end of snout slightly concave between the subnarial

Taylor: A New Veracrucian Salamander 191

swellings; seen in lateral profile there is an emargination of the line
of upper lip immediately behind snbnarial swellings; latter some-
what bulbous, slightly pendant; canthus rostralis not present, the
snout rounded, the lores sloping gradually to lip; eye rather large,
slightly longer than median length of snout; width of an eyelid
contained in interorbital distance approximately li times; no
orbitolabial groove; posterior parts of eyelids concealed under a
short diagonal fold; line of the mouth forming an angle under
posterior part of eye; a postorbital groove faintly indicated; a ver-
ticle groove indicated back of jaw angle which may be traced on
side of head and on throat; an ample gular fold, very slightly
curved across neck, scarcely traceable on the sides of neck; eleva-
tion made by the posterior part of the epibranchial cartilage,
passes above arm to second costal groove; nostrils small; skin on
head porous; musculature of the posterior part of head and begin-
ning of neck causing distinct and regular elevations of the skin
dorsally and laterally; skin on lateral costal folds with some longi-
tudinal wrinkling: the upper parts of the costal grooves form zig-
zag lines reaching to near the median dorsal line; about 25 caudal
grooves discernible; tail quadrangular in cross section; an indistinct
groove on under surface of tail.

Arm reaching middle of eye; hand palmate, tip of third finger
webbed but extending forward as a slight triangle; tips of other
fingers scarcely free; foot palmate, the edges with slight emargi-
nations between tips of toes; lateral cloacal walls covered with pa-
pilla in male.

Tongue bolitoglossid; no trace of a sublingual fold; four premaxil-
lary teeth penetrating edge of upper lip (one missing in type);
vomerine teeth 14-15, in two transverse series, bending back a little
and forming an angle mesially, the two series separated mesially
by a distance equal to that between two teeth; paravomerine teeth
tending to form a single patch narrowed anteriorly, expanded pos-
teriorly with a slight median posterior notch; maxillary teeth small,
30-32; there is a hiatus anteriorly between maxillary and premaxil-
lary series.

Measurements in mm.— Snout to end of vent 50; tail 46; width of
head 8.0; length of head to behind jaw angle 10; length of head to
gular fold 13.5; arm 12.3; leg 13; axilla to groin 26.5.

Color and markings. — Lavender-brown with irregular series of
clay or fawn-colored spots scattered from snout to tail tip, the spots
varying in size; spots small low on side; venter and under side of
limbs and tail lighter lavender-brown, the individual chromato-

192

The University Science Bulletin

phores separate, with numerous small cream or yellow spots (not
longitudinal); a small lighter glandular area behind insertion of
thigh; a white line along lower edge of lower eyelid; limbs spotted
like body; cloacal walls of males without pigment (female with
cloacal walls pigmented).

Table of Measurements of Adult Bolitoglossa veracrucis

Snout to

Axilla

Snout to

end of

Tail

Head

Head

to

gular

No.

Sex

vent

length

width

length

Arm

Leg

groin

fold

26940

$

55

9

10.3

12.5

14

32

14

26941

$

50

46

8

10

12.3

13

26.5

13.5

26942

s

50

42

9

10

12.4

12

28

13

26938

9

50

42

8.4

9.8

12.5

13.2

28

13

26947

$

47.5

, ,

8

9.5

11.5

11.5

25.5

12.2

26939

$

46

7.7

8

11

11

25

11.9

Table

of Data on Bolitoglos

3A VERACRUCIS

Costal

Vomerine

Maxillary

Pre-

No.

Sex

grooves

teeth

teeth "

maxillary

26940

9

13-13

17-19

31-29

6

26941

$

13-13

14-15

30-33

3

(+D

26942

$

13-13

14-13

35-29

4

26938

$

13-13

19-17

37-39

6

26947

$

13-13

12-14

27-26

5

(+D

26939

$

13-13

17-15

24-26

4

Variation.— The series of specimens range in size from 22-55 mm.,
snout-to-end-of-vent measurement. They vary considerably in color
but this variation does not seem to have any relation to sex or age.
Some of the younger specimens are largely fawn on the dorsal sur-
faces, with a few black flecks. Some of the young are colored much
as are the adults. The costal grooves are 13 in all if one counts a
groove in axilla and groin; occasionally a partial division is sug-
gested in the last costal fold.

Dentition is lacking in the young but specimens above 45 mm.
snout-to- vent length seem to have the adult dentition. The num-
ber of teeth in females is a little greater than in the males, there
being normally six premaxillaries penetrating the gum in females
and four normally in the males penetrating the upper lip. There
seemingly is a higher count of vomerine teeth in females than in
males as suggested by the table of data. The subnarial swellings
are less bulbous and less pendant in females, and the cloacal wall
has numerous pigmented folds instead of papilla. It is presumed
that the considerable series taken in the same locality and at the
same time contains specimens of fully grown animals. That some of
the animals attain larger size than those given here is to be ex-
pected; I doubt, however, that this difference will be very consider-
able.

Taylor: A New Veracrucian Salamander 193

Relationship— The salamanders here described belong in the
genus Bolitoglossa and probably in the section of the genus that in-
cludes Bolitoglossa odonelli (Stuart), Bolitoglossa mulleri (Broc-
chi), and not impossibly also Bolitoglossa yucatana (Peters). The
two former species are mountain forms that differ from the one here
described in having fewer teeth, shorter limbs in proportion to the
axilla to groin measurements, a different basal coloration and dif-
ferent dorsal markings. Both are presumably larger species. The
total length is 123 mm. for odonelli; for mulleri, as given by Stuart
(1943), 145. The tail length of odonelli is practically that of head-
body length (61-62 mm. in the type) and varying but little in the
paratypes. Stuart has omitted several important characters from
his description such as the presence of absence of the sublingual
fold, the paravomerine teeth, the maxillary teeth, etc.

From yucatana this form differs in being of smaller size, vomerine
teeth not an "irregular patch of teeth," no ventral linear markings,
markings not concentrated in two dorsolateral lines, long limbs in
proportion to axilla-to-groin length (one to one and a half costal
folds between adpressed limbs instead of five folds), snout de-
pressed instead of "quite high," the tail shorter than body.

Stuart ( 1943) * calls attention to my error in using punctatus when

mulleri is intended, a curious error resulting from the dropping of a

complete line of text. Punctatus definitely does not belong in this

group.

* While it is not pertinent I wish to call attention to a confusing error of
Stuart, p. 16, where he states that Taylor has described Bolitoglossa nigrofilaves-
cens from Chiapas. This name has not been used by Taylor.

LITERATURE CITED
Stuart, L. C.

Taxonomic and Geographic Comments on Guatemalan Salamanders of
the Genus Oedipus. Misc. Publ. Mus. Zool. Univ. Michigan No. 56, 1943,
pp. 1-33, pis. 1-2.

13—3286

THE UNIVERSITY OP KANSAS

SGIENGE BULLETIN

Vol. XXXIV,, Pt. I] October 1, 1951 [No. 3

A New Species of Leiolopisma (Reptilia: Sauria)

from Mexico*

Hobakt M. Smith

Abstract. Leiolopisma caudaeqiiinae is described from Horsetail Falls,
near Monterrey, Nuevo Leon, Mexico, the type in the Museum of Natural His-
tory, University of Illinois. The status of the other four North American
members of its group is discussed and L. forbesorum and L. gemmingeri are
regarded as subspecies. A key to the five fonns is appended.

A Leiolopisma recently received as a gift to the University of
Illinois Museum of Natural History from Mr. J. P. Craig, and col-
lected at Horsetail Falls in Nuevo Leon, Mexico, differs markedly,
but chiefly in leg proportions, from its closest geographic relative,
L. forbesorum of high elevations in the state of Hidalgo. A query
to the authority on Mexican Leiolopisma, Dr. Edward H. Taylor,
revealed that for a number of years he has possessed specimens of
the long-legged northern species from San Luis Potosi. Upon his
suggestion the form is named herewith. I am indebted to him for
the loan of comparative material and assistance in preparing this
account, and to J. P. Craig for the gift of the type.

Leiolopisma caudaequinae f sp. nov.

Holotype. Univ. 111. Mus. Nat. Hist. No. 10131, an adult male
from Horsetail Falls, 25 miles south of Monterrey, Nuevo, Leon, col-
lected by J. P. Craig, April 19, 1946. Paratypes.- Two, E. H. Taylor
Coll. Nos. 23886, 23892, from a locality 10 miles west of Naranjo,
San Luis Potosi, on the Antiguo Morelos-San Luis Potosi highway,
along a small creek. Hypoparatypes. Four, E. H. Taylor Coll.
Nos. 23888-23891 (all juveniles), from the same locality as the para-
types.

* Contribution from the Museum of Natural History and the Department of Zoology, Uni-
versity of Illinois, Urbana, Illinois.

t In reference to Horsetail Falls, the type locality.

(195)

196 The University Science Bulletin

Diagnosis. — A member of the genus Leiolopisma similar and re-
lated to L. silvicolum, having a divided frontoparietal, 61-70 dorsals,
a brown lateral stripe involving ear, 28-32 scale rows, and large,
strong limbs overlapping in adults when adpressed and the fore-
limb reaching eye; differing from silvicolum in having usually 1 pair
of nuchals, 17-19 lamellae under the fourth toe, and a continuous,
unbroken dorsolateral light stripe.

Description of holotype.—A mature male; frontal elongate, width
4/5 of length, its distance from tip of snout % its length, which is
about 5/6 length of frontoparietals and interparietal combined;
internasal very large, almost as long as broad, in contact with
frontal by a suture almost as long as that with rostral; prefrontals
a fourth larger than nasals; frontoparietal divided, the median
suture 2/3 length of interparietal; parietals in contact behind fron-
tal with a suture 1/5 length of interparietal. Four supraoculars,
anterior narrowly in contact with prefrontal, 2nd narrowly in con-
tact with frontoparietal, 3rd smallest; 7 superciliaries, anterior larg-
est. Nasal entire; postnasal 2/3 size of nasal ( divided into 2 super-
imposed scales on 1 side); a nearly square loreal % larger than an-
terior superciliary; one large (lower) and 3 (4 on one side) small
preoculars; 2 presuboculars above 4th supralabial, a little smaller
than large lower preocular; 2 tiny suboculars above 5th supralabial;
two postsuboculars in contact with 6th supralabial; 3 large and
several small postoculars; nine upper and 12 lower palpebrals.
Seven supralabials, 6th slightly larger than 7th, 4th smallest; mental
with a longer labial border than rostral; 6 infralabials, 5th longest,
1st smallest; postmental nearly twice as large as mental, in con-
tact laterally with 2 infralabials; 3 chinshields on each side, the
scales of the anterior pair in contact medially, those of 2nd sepa-
rated by 1 scale, the 3rd by 3; a small postgenial, longer than broad.

Ear large, 1.6 x 1.1 mm., tympanum deeply sunken; anterior tem-
poral very slightly larger than lower secondary; upper secondary
temporal very large, 2/3 size of parietal; two relatively large post-
labials followed by 3 vertical rows of small scales in front of ear; 1
large pair of nuchals, in contact with secondary temporal; on one
side a smaller nuchal behind the other; two tertiary temporals,
lower largest, between nuchal and postlabials.

Dorsal and lateral scales with several minute keels or pits, es-
pecially prominent on lateral scales; 63 scales from interparietal to
level of posterior edge of hind leg; 30 scale rows at middle of body;
17 lamellae on 4th toe.

Smith: New Species of Leiolopisma 197

Limbs large, overlapping 3 scale-lengths, forelimb reaching rear
corner of eye opening; median subcaudals a little wider than adja-
cent scales, and a little longer, every other scale in contact laterally
with two scales (a large and a small), the alternating scales in con-
tact with one on each side; regenerated portion of tail with broad
median subcaudals each in contact laterally with one scale.

Ground color a dark tan; dorsal surface not marked; a fine dor-
solateral light line, lacking melanophores, with a well-defined lateral
border (formed by a very dark brown lateral band), and a dimly
evident median border, following median edge of the 4th scale
row, originating on snout and terminating near base of tail; a lateral,
very dark brown, continuous band ( not interrupted by light streaks )
below this, occupying 2 and 2 half scale rows, its lower edge rather
irregular and tending to blend gradually with the adjacent slate
color; sides below band sparsely pigmented; belly not pigmented;
a few melanophores on sides of ventral surfaces of neck and head;
a poorly defined light line from snout below eye and through ear to
arm insertion.

Variation. — The paratypes are very similar to the type. Varia-
tions in important features are indicated in the accompanying table.
The four hypoparatypes likewise are very similar, and agree in all
diagnostic characters, even limb proportions. Since a differential
growth rate occurs in these lizards, the most reliable proportions are,
however, those of the adult, to which generalizations here are re-
stricted. The dorsal scales number 62, 64 and 67 in hypoparatypes
which can be counted, and the scale rows around middle of body
28, 28, 30, 32 (all carefully checked several times). The nuchals
are 1 pair in two, 1-% pairs in one, 2 pairs in one; the members of
the anterior pair are separated from upper secondary temporal in 3,
from each other in 1. The lamellae under the 4th toe are 18 (two)
and 19 (two).

Comparisons.— From forbesorum the present form differs chiefly
in length of foreleg (reaching orbit) and extent of overlap of ad-
pressed limbs ( touching or overlapping as much as 7 scale lengths ) .
In forbesorum the foreleg does not extend forward beyond a point
halfway between eye and ear, and the adpressed limbs are sepa-
rated by from 1 to 8 scales. The most conspicuous difference is in
limb proportions, those of caudaequinae being conspicuously larger
and longer than those of g. forbesorum. In addition the new species
possesses a different curve of variation in scale row count around
middle of body (28, four; 30, two; 32, one), although it overlaps

198 The University Science Bulletin

that of g. forbesorum (26, two; 27, one; 28, seven) so greatly that
it has no diagnostic value, and even its significance is questionable.

L. gemmingeri, like forbesorum, has short limbs. L. silvicolum,
the only other species in Mexico of this group, has long limbs as
in caudaequinae, but the nuchals are usually in 2 rows, the lamel-
lae under the 4th toe 20, the dorsolateral light stripe is broken, and
the scale rows more regularly 30 or 32.

Relationships. — The "Oligosoma" group of Taylor ( Copeia, 1937,
p. 11) in North America consists of laterale, gemmingeri, forbes-
orum, caudaequinae and silvicolum. It is characterized by hav-
ing a divided frontoparietal, and a peculiar variation in size of the
paraventral subcaudals, so that many scales of the median row
are in contact laterally with 2 scales instead of the normal single
scale. The only other North American members of the genus
belong to the "Mocoa" group of Taylor ( loc. cit. ) and are charac-
terized by a single frontoparietal, and normal median subcaudals
each of which is in contact laterally with a single scale.

Of the "Oligosoma" group, laterale is easily distinguished from
others by having the upper tertiary temporal (occasionally split)
in contact with the parietal and usually 3 pairs of nuchals. All
others of the group have the upper tertiary temporal separated
from the parietal by contact of the upper secondary temporal and
nuchal (sometime split), and usually 2 or fewer pairs of nuchals.
The other four species fall naturally into two groups, one charac-
terized by long limbs that overlap when adpressed, the foreleg reach-
ing the eye, and scale rows usually 28 or 30, sometimes 32 (silvi-
colum, caudaequinae) the other having short limbs failing to
overlap or reach the eye, and scale rows usually 26 or 28, sometimes
25 or 30 ( gemmingeri, forbesorum ) . Each group is represented by a
northern species (caudaequinae, forbesorum), having usually one
pair of nuchals, separated by a distance of 150 ( in the short-legged
group) to 300 miles from a southern species having usually two
pairs of nuchals. For the most part all four species appear to oc-
cupy their ranges to the exclusion of all others of the group, and
probably even the other group does not usually (if ever) occur
sympatrically. Allopatry in species of this genus in North America
seems to be virtually invariable.

One apparent exception to this rule deserves comment. A single
short-legged specimen of L. forbesorum bears the same data as the
series of caudaequinae from San Luis Potosi (EHT No. 23887). It
agrees well with others of the species (now known also from the
type locality at La Placita, Hidalgo, and possibly Rio Verde, San

Smith: New Species of Leiolopisma 199

Luis Potosi * ) except in scale row count, which is 30 at the middle
of the body. Counts and measurements are given in the accom-
panying table. Possibly some error occurred in recording the lo-
cality data for the specimen, but no indication of such error exists
in the catalogue, on the tag or in the condition of the specimen.

L. forbesorum has been recorded no nearer the known range of
gemmingeri than La Placita, Hidalgo; the nearest locality known for
gemmingeri is Tequeyutepec, Veracruz, about 150 miles distant.
They occupy much the same habitat, both occurring at relatively
high elevations. Localities recorded for gemmingeri are: Oaxaca:
"Tehuantepec,f" 17 mi. N. Niltepec, Ixcuintepec, Tres Cruces, San
Jose Manteca, Cafetal Concordia; Veracruz: Jalapa, Orizaba, La
Perla, Tequeyutepec; Hidalgo: Zacualtipan.}:

All of the differences between these are relatively minor and the
curves of variation either are known to overlap or assuredly can
be expected to do so. The differences are: number of nuchals
(gemmingeri usually 2 pairs, forbesorum usually 1 pair, the number
somewhat variable in both f ) ; lateral light line along head and neck
(poorly defined in gemmingeri, fairly well defined in forbesorum);
axilla-groin/snout- vent percentage (59% in gemmingeri, 56% in
forbesorum); maximum snout-vent length (65 mm. in gemmingeri,
54 mm. in forbesorum) .

In view of the fact that these two forms occur in the same habitat,
in areas known to be closely approximated if not actually adjacent,
and differ incompletely, it is not unreasonable to regard them as
subspecies (Leiolopisma gemmingeri gemmingeri (Cope) and Leio-
lopisma gemmingeri forbesorum Taylor).

The long-legged species, silvicolum and caudaeq ulnae, differ from
each other to a somewhat greater degree than the preceding races
do from each other ( see Comparisons ) . Unfortunately only the type
and paratype of the arboreal silvicolum have been described in
detail; but little information is available for the specimens from
Oaxaca. The known degree of difference undoubtedly will be al-
tered by larger series. L. silvicolum, known only from San Jose de
Gracia, Veracruz, and Cuicatlan, Oaxaca, does not so clearly occupy

* This specimen (U. S. Nat. Mus. No. 32372), labeled simply "Rio Verde" (presumably
San Luis Potosi, but not certainly) cannot now be found.

t Undoubtedly in reference to the area, or Isthmus; the species does not occur at or near
the city of this name.

t Neither Doctor Taylor nor I have seen this specimen (a Cope record) ; if correctly assigned
it brings the ranges of the two species much closer, into the same range. Even if referable to
forbesorum, the result is the same so far as range approximation is concerned.

K Nuchals on the two sides 0-1 to 3-3 in 21 gemmingeri, 89% 1-2 or more (0-1. one; 1-1,
one; 1-2, three; 2-2, twelve; 2-3, two; 3-3, two); nuchals 0-0 to 2-3 in 12 forbesorum,
88% 1-1 or less (0-0, one; 0-1, two: 1-1, seven; 2-2, one; 2-3, one).

200 The University Science Bulletin

the same habitat as caudaequinae of Nuevo Leon and San Luis
Potosi, although all these localities are on the Atlantic drainage and
at relatively low elevations. That these forms, like those of the
short-legged group, may actually be subspecies is quite possible,
but information currently available does not at the present time war-
rant assumption of subspecific status.

Key to Mexican Members of the "Oligosoma" Group of Leiolopisma

1. Upper tertiary temporal (occasionally split) in contact with parietal; usually 3

or more pairs of nuchals laterale

Upper tertiary temporal separated from parietal by contact of upper secondary
temporal and nuchal ; usually less than 3 pairs of nuchals 2

2. Limbs, when adpressed, touching or overlapping in adults, forelimb reaching rear

corner of eye ; scale rows at middle of body often 30 or 32 3

Limbs, when adpressed, separated from each other by 1 or more scale lengths;
scale rows at middle of body rarely 30 and apparently never 32 4

3. Usually one pair of nuchals or less ; lamellae under 4th toe 19 or fewer ; dorso-
lateral light stripe continuous caudaequinae

Usually two pairs of nuchals; lamellae under 4th toe 20 (constant?); dorsolateral
light stripe dark spotted, interrupted silvicolum

4. Usually nuchals on the two sides 1-2 or more (89%); lateral light line along
head and neck poorly defined; axilla -groin/snout-vent percentage 59; Go mm.

maximum snout-vent measurement gemmingeri gemmingeri

Usually nuchals on the two sides 1-1 or less (88%); lateral light line along head
and neck fairly well defined; axilla-groin/snout-vent percentage 56; 54 mm.
maximum snout-vent measurement gemmingeri forbesorum

Table of Variation in Leiolopisma*

Number 10131 23892 23886 23887

Sex $ $ 5 $

Snout to vent 49.0 47.0 49.0 52.0

Snout to eye 3.5 3.6 3.4 3.3

Snout to ear 9.0 9.0 8.3 8.5

Snout to foreleg 18.0 17.6 17.0 17.5

Eye to foreleg 11.8 11.3 11.3 11.0

Axilla to groin 25.8 24.0 26.5 30.3

Tail 71.0 78.0

Head width 7.1 6.7 6.8 6.2

Head length to nuchal 8.7 8.7 8.3 8.2

Arm 11.3 11.5 11.0 10.5

Leg 17.8 17.5 16.5 16.2

Longest toe 7.0 6.6 6.2 6.1

Scales behind ear 35 36 38 38

Scales around neck 30 30 32 32

Scales 2 mm. behind axilla 35 35 36 35

Scales about middle of body 28 28 30 30

Scales in row from parietal to rear of thighs. .. . 63 61 62 70

Lamellae under 4th toe 17 17 18 18

Legs overlapping by scales 3 7 0 -8

Nuchals 1-2 1-2 1-1 1-1

Anterior nuchal contacting upper 3-2 temporal, 0-+ 0 - -f- 0 - + 0 - +

Anterior nuchals separated from each other.... 0 0+0

* No. 10131, holotype, Nos. 23886, 23892, paratypes, of L. caudaequinae; No. 23887, L. g.
forbesorum from the same locality as the paratypes of the preceding. Measurements in mm.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIV, Pt. I] October 1, 1951 [No. 4

Notes on a Small Herpetological Collection
from Guerrero

BY

Charles W. Hall

Arstract. A collection of snakes and lizards made in the vicinity of Chil-
pancingo, Guerrero by W. Wilmot Brown is reported. A single new subspecies,
Coniophan.es piceivittis taylori is described.

This paper is based on a small collection of Mexican reptiles ac-
quired by the Kansas University Museum of Natural History from
Mr. W. Wilmot Brown. The collecting was done within a 15 km.
radius of Chilpancingo, Guerrero. Unfortunately neither exact
localities nor approximate elevations are appended but undoubt-
edly certain specimens came from considerably higher elevations
than the city itself. Chilpancingo is located on the Mexico-Aca-
pulco Highway at an elevation of approximately 1380 meters. The
city lies on the southern drainage of the Sierra Madre del Sur. The
famous collecting locality, Omilteme, (elevation 2460 meters) is
located about ten kilometers west of Chilpancingo.

To the west of the city, the mountains of the Sierra Madre del Sur
range rise rapidly to reach a maximum elevation of 3700 m. and
extend westward for about 275 km. where the range terminates at
the Rio Balsas. To the eastward the mountains are somewhat lower
than at Chilpancingo. This low pass divides the Sierra Madre del
Sur from its extension into Oaxaca. The range is separated from
the Mexican Plateau on the north by the Balsas Basin and is isolated
on the south by the Pacific Ocean.

It is in this area that a great number of endemic species are found.
Taylor * has reported 64 endemic species of reptiles and Amphibia in
this region ( 1940 ) . The most surprising character of the fauna has
to do with the absence of all salamanders save for what is regarded

* Taylor, E. H., "Island" Faunas on the Mexican Plateau; Proc. Eighth Amer. Sci. Cong.,
vol. Ill, Biol. Sci., pp. 503-504.

(201)

202 The University Science Bulletin

as the most primitive Mexican plethodontid, Pseudoeurycea belli.
One can reasonably account for the absence of other forms of pletho-
dontids only by postulating salt water isolation for the area for a
considerable period of time, as it hardly seems possible that the
Balsas Basin could act as such a barrier. To what extent geological
evidence will bear this out, I do not know.

The following species are present in this collection:

SAURIA

Barisia gadovii gadovii (Boulenger)

SERPENTES

Leptotyphlops maximus Loveridge

Drymarchon corais rubidus Smith

Elaphe chlorosoma (Giinther)

Lampropeltis doliata blanchardi Stuart

Masticophis flagellum Hneatus (Bocourt)

Salvadora mexicana (Dumeril, Bibron, and Dumeril)

Storeria storerioides ( Cope )

Thamnophis chrysocephalus (Cope)

Thamnophis eques eques (Reuss)

Trimorphodon latifascia Peters

Leptodeira maculata (Hallowell)

Coniophanes piceiviltis tuylori subsp. nov.

Sonora michoacanensis michoacanensis (Duges)

Oxybelis auratus (Bell)

SYSTEMATIC REPORT

Barisia gadovii gadovii (Boulenger)

Gerrhonotus gadovii Boulenger, 1913, Ann. Mag. Nat. Hist., ser. 8, vol. 12, p. 564.
Barisia gadovii gadovii Till en, Univ. Kansas Sci. Bull., vol. 33, 1949, pp. 230-231.

Three specimens of Barisia gadovii gadovii (KUMNH Nos. 23792,
23793, and 23794) are said to have been collected in "the vicinity of
Chilpancingo," Guerrero. However one supposes that they came
from the mountain west of, and at a higher elevation than the city
itself. The type locality, Omilteme, Guerrero, is at an elevation of
2460 meters and about 10 kilometers west from Chilpancingo.

Examination of these specimens revealed the following data:
postrostral absent in specimens No. 23793 and No. 23794, but
present in specimen No. 23792 separating the two supranasals; a
small pair of supranasals separating internasals from nasals; a pair
of canthals lying on either side and broadly in contact with fronto-
nasal; three lateral supraoculars; five medial supraoculars; supra-
ciliaries 5-4 in No. 23792, 5-5 in No. 23793, 4-4 in No. 23794; one pre-

Hall: Snakes from Guerrero 203

ocular; two suboculars; two postoculars; four primary and four
secondary temporals; supralabials 9-10 in each specimen; infra-
labials 8-8 in Nos. 23792 and 23794, 7-8 in No. 23793; postmental
paired, followed by three large pairs of chinshields.
Further data is given in the table.

Scale Counts and Measurements (in mm.) of Barisia gadovii gadovii

(Boulenger)

No 23792 23793 23794

Snout to vent 66 73 83

Tail length 115 138 119

Right forelimb 15 16 17

Left forelimb 17 17 17

Right hindlimb 20 21 22

Left hindlimb 21 22 23

Snout to ear 14 17 16

Axila to groin 37 39 48

Snout to forelimb 23 25 26

Parietal to above anus 44 48 51

Number of caudals 79 90 65

Dorsal scales at mid-body 16 16 18

Ventral scales at mid-body 12 12 12

Leptotyphlops maximus Loveridge

Leptotyphlops maximus Loveridge, Proe. Biol. Soc. Wash., vol. 45, 1932, pp. 151-152; Taylor,
Copeia, 1939, No. 1, pp. 4-5, figs. 3-4; Klauber, Trans. San Diego Soc. Nat. Hist., vol.
9, 1940, pp. 120-12-2, map 1.

There is one specimen of this large blind snake in the collection,
which gives the following data: supralabials 2; infralabials 3; 14
scale rows around body; ventrals 219; caudals 15; anal single; total
length 210 mm; tail length 9 mm; tail length contained in body
length 23 1/3 times.

In alcohol the specimen is brown above with the underside pink.
A casual glance gives one the impression that the scales contain a
pattern fringing their edges. However on closer inspection, the
scale edge is seen not to coincide with the pattern. Upon removal
of the outer epidermis, there is seen the uncornified part of the
scale lying beneath. It appears as if the outer epidermis has grown
caudad from the deeper portion of the underneath scale, its posterior
edge becoming free as occurs in the growth of a fingernail.

Drymarchon corais rubidm Smith

Drymarchon corais rubidus Smith, Journ. Washington Acad. Sci., vol. 31, 1941, pp. 474-476,
map fig. 2 (part).

Two specimens in this collection are referred to this subspecies
(K.U.M.N.H. Nos. 23799 and 26728). Head scales normal; suprala-
bials eight, the antepenultimate labial in contact with the temporal

204 The University Science Bulletin

but not with the postocular; one preocular; two postoculars; tem-
porals 2-2; anal entire.

The color is in accord with that given by Smith ( loc. cit. ) except
the female, which I judge to be half-grown, has retained a pattern
on the dorsal surface. The pattern, a series of pink chevronlike
bars, one scale row in width, commences on the nape and continues
throughout the body but disappears on the tail. The bands are not
discrete, but are infiltrated with the black color. The smaller,
male specimen is uniformly black on the dorsal surface with only
a faint suggestion of a pattern.

Data on Dkymarchon corais rubidus Smith

No.

Sex

Scale
formula

Ventrals

Caudals

Infra-
labials

Total
length

Tail
length

13799
!6728

9
$

17-17-15
19-17-15

198
197

69
59

8-9

8-8

1500
1130

260
185

Elaphe chlorosoma Giinther

Coluber chlorosoma Giinther, Biologia Centiali-Americana, Rept., 1894, pp. 115-116, pi. 41.
Eiaphe chlorosoma Stejneger and Barbour, A Checklist of North American Amphibians and
Reptiles, ed. 1, 1917, p. 82; Smith, Copeia, 1941, No. 3, pp. 134-135.

There are three specimens of Elaphe chlorosoma in this collec-
tion, two adults and fortunately one juvenile, KUMNH Nos. 23781-
23783. The juvenile has a somewhat larger count of spots on the
body than those mentioned by Smith {loc. cit.). He reported 57
spots, this one having approximately 66. The fact that this juvenile
and two adults were collected from the same general area, seem
to confirm in some measure the opinion of Smith (loc. cit.) that
Elaphe matabilis (Cope) represents the juvenile pattern of chlor-
osoma. However, as shown in the table, the juvenile has a lower
ventral count and higher caudal count.

The three specimens are identical in the following characteris-
tics: infralabials 10; preoculars 1; postoculars 2; loreal 1; temporals
3, 4; chinshields 2; anal divided. The wide range in the ventral
count found in these three specimens (247-278, a difference of 31
scales) is especially noteworthy and may suggest that two forms
are involved.

Data on Elaphe chlorosoma Giinther

No.

Sex

Scale

formula

Ventrals

Caudals

Total
length

Tail
length

Supra-
labials

23781

9

33-35-25

267

88

1260

210

9-10

23782

9

31-35-23

278

93

1170

210

8

23783

S

31-35-23

247

108

380

80

8

Hall: Snakes from Guerrero

205

Lampropeltis doliata blanchardi Stuart

Lampropeltis polyzona blanchardi Stuart, Occ. Pap. Mus. Zool. Univ. Michigan, No. 309, 1935,
pp. 1-6. Taylor, Univ. of Kansas Sci. Bull., vol. 26, 1939 (1940), p. 467, pi. 49.

Lampropeltis triangulum blanchardi Dunn, Occ. Pap. Mus. Zool. Univ. Michigan, No. 353,
1937, p. 8.

Lampropeltis doliata blanchardi Klauber, Copeia, No. 1, pp. 10-11, 1948.

Nine specimens of Lampropeltis doliata blanchardi Stuart are in
this collection. They are identical in the following characteristics:
supralabials 7, infralabials 8, preoculars 1, postoculars 2, tem-
porals 2 + 3, chinshields 2, loreal 1, anal single.

The snout is totally black with no evidence of a white snout band.
These specimens show a slightly higher average ventral count than
that given by Smith,* 212 as compared to 208.

Lampropeltis doliata blanchardi Stuart

Scale

Total

Tail

No.

Sex

formula

Yentrals

Caudals

length

length

23772

$

21-21-19

211

51

650

93

23773

9

21-19-19

216

46

740

102

23774

9

21-21-19

209

47

370

52

23775

9

21-19-19

218

50

900

129

23776

S

21-21-19

216

28

780

75

23777

S

21-19-19

210

54

900

132

23778

$

21-21-19

211

53

870

130

23779

9

21-21-19

212

49

810

109

23780

9

21-21-19

211

49

550

84

Masticophis jlagellum lineatus (Bocourt)

Bascanion lineatus Bocourt, Mission scientifique au Mexique et dans 1'Amerique centrale;

Etude sur les reptiles, livr. 12, 1890, pp. 700-701, pi. 48, fig. 1.
Masticophis lineatus Ortenburger, Occ. Pap. Mus. Zool. Univ. Michigan, No. 139, 1923, p. 2;

Mem. Univ. Michigan Mus. Zool., vol. 1, 1928, pp. 134-138, p. 125.
Coluber striolatus Mertens, Zoologica (Stuttgart), vol. 32, 1934, p. 190 (substitute name for

Coluber lineatus (Bocourt), a secondary homonym of Coluber lineatus Linnaeus (Lygophis

lineatus).

A series of six specimens was obtained in the Chilpancingo region
(KUMNH Nos. 24104-24109). The dorsal color, at a glance, ap-
pears to be bluish gray, but upon closer inspection of the specimens,
the dorsal scales are seen to be tipped with black and small cream-
colored lines are faintly discernible. The venter is cream with the
outer margins of each ventral scale taking on the gray dorsal color.
The lips and chin are heavily mottled. The supralabials of each
specimen are 8-8, with the fourth and fifth entering the orbit; pre-
oculars 2; postoculars 2; loreal 1; anal divided; scale formula 19-
17-13.

* Smith, H. M., Proc. of the Rochester Acad. Sci., Sept. 10, 1942, vol. 8, pp. 199-260.

206 The University Science Bulletin

Data on Masticophis flagellitm lineatus (Bocourt)

Infra -

Total

Tail

No.

Sex

Ventrals

Caudals

labials

Temporal

length

length

24104

$

192

129

10

2-3

1570

460

24105

$

194

77

10

2-3

1460

300

24106

s

194

108

10-11

2-2
2-3

1440

380

24107

$

192

129

10

2-2
2-2

1450

440

24108

9

193

97

10

2-3

1340

340

24109

9

195

116

10-11

2-2
2-3

1235

350

Salvadora mexicana (Dumeril, Bibron, and Dumeril)

Zamenis mexicanus Dumeril, Bibron, and Dumeril, Erpetologie generate, vol. 7, pt. 1, 1854,
pp. 695-696. — Bocourt, Mission scientifique au Mexique et dans l'Amerique Centrale;
Etude sur les reptiles, livr. 11, 1888, p. 664, pi. 46, fie. 5.

Salvadora mexicana Gunther, Ann. Mag. Nat. Hist., ser. 3, vol. 12, 1863, p. 349. — Bogert,
Copria, 1939, No. 3, pp. 144-145.

There is only one specimen of Salvadora mexicana in this collec-
tion (KUMNH No. 23784). The specimen is an adult male, measuring
1213 mm. total length, with a tail length of 460 mm. In alcohol the
ground color is cream with four longitudinal brown or black stripes
running almost the full length of the snake, fading out only near the
tip of the tail and interrupted in the neck regions by eight or ten cross
bars. There are some black spots on the outer margins of the ven-
trals in the anterior third of the body. The following scale char-
acters are in evidence: scale formula 17-17-13, ventrals 192, cau-
dals 140; supralabials 9, infralabials 10, preoculars 1, postoculars 2,
temporals 2 + 2, chinshields 2, loreal 1, anal divided.

Storeria storerioides (Cope)

Tropidoclonium storerioides Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 17, 1865, pp. 190-

191.
Storeria storcriodes Garman, Mem. Mus. Comp. Zool., vol. 8, No. 3, 1883, p. 29; Taylor and

Smith, Univ. Kansas Sci. Bull., vol. 25, 1938 (1939), pp. 249-251, fig. 3 (head only):

Smith, Proc. U. S. Nat. Mus., vol. 93, 1943, p. 473.
Hemingenius variabilis Duges, Proc. Amer. Philos. Soc. vol. 25, 1888, pp. 182-183, fig. 2

(type locality, Guanajuato; type in Mus. Alfredo Duges, Univ. Guanajuato, Mexico).

There is one specimen ( KUMNH No. 23788 ) of Storeria storeri-
oides in this collection, having the following characteristics:
scale formula 15-15-15; ventrals 128; caudals 50; anal divided; su-
pralabials 7-6; infralabials 7; loreal 1; preoculars 2; postoculars 3;
temporals 1 + 2; chinshields 2 pairs; total length 270 mm; tail
length 60 mm; coloration reddish brown with dark brown chevron-
like bands faintly discernible on the upper 1/3 of the body.

Hall: Snakes from Guerrero 207

Thamnophis chrysocephalus (Cope)

Eutaenia chrysocephalus Cope, Proc. Amer. Philos. Soc, vol. 22, 1S84 (1885) pp. 173-174.
Thamnophis chrysocephalus Smith, Zoologica, vol. 27, 1942, p. 104; Proc. U. S. Nat. Mus.,

vol. 93, 1943, pp. 478-479.
Thamnophis eburatus Taylor, Herpetologica, vol. 1, 1940, pp. 187-189, pi. 19, text fig. 2

(type locality, Cerro San Felipe, Oaxaca; type, E. H. Taylor-H. M. Smith coll. No. 5556).

This collection contains one specimen of Thamnophis chryso-
cephalus ( KUMNH No. 23785 ) . It is a female measuring 580 mm.
total length and 150 mm. tail length. On the upper surface the
color, in alcohol, is a midnight blue. The venter is blue, spotted
with pink. The underside of the chin and infralabials is an im-
maculate pink. The supralabials are also spotted with pink.

Thamnophis eques eques (Reuss)

Coluber eques Reuss, Zool. Misc., 1834, pp. 152-155, pi. 8, fig. 2.

Thamnophis eques eques Gloyd and Smith, Bull. Chicago Acad. Sci., vol. G, 1942, p. 234;
Smith, Zoologica, vol. 27, 1942, pp. 106-108.

Thamnophis cyrtopsis cyclides Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 13, 1861, pp. 229-
230 (type locality unknown, "Cape St. Lucas"; type, USNM No. 5023).

Entaenia pulchrilatus Cope, Proc. Amer. Philos. Soc. vol. 22, 1884 (1885), p. 174 (type local-
ity, Guanajuato; type USNM No. 9899).

Two specimens collected in the Chilpancingo region are referred
to this subspecies (KUMNH Nos. 23797 and 23798).

Head scales normal; supralabials eight, the fifth and sixth in
contact with lower postocular; infralabials eleven, five in contact
with anterior chinshields; one preocular; two postoculars; anal
single; scale formula 19-19-17. A very distinct median light stripe,
involving only the vertebral row of scales, runs the entire length of
the body; dark spots present on scales below the lateral light line,
at least anteriorly, visible without spreading the scales.

Data on Thamnophis eques eques

Total

Tail

No.

Sex

Ventrals

Caudals

length

length

23797

9

151

85

410

110

23798

S

150

83

810

210

Trimorphodon latifascia (Peters)

Trimorphodon biscutata latifascia Peters, Monatsb. Akad. Wiss. Berlin. 1869, p. 877.
Trimorphodon latifascia Taylor, Univ. Kan. Sci. Bull., vol. 25. 1938 (1939), pp. 364-365,

pi. 36, fig. 2; and vol. 26, 1939 (1940), p. 479, pi. 52; Smith, Proc. U. S. Nat, Mus.,

vol. 91, 1941, p. 160, map, fig. 38.

There are four Guerrero specimens of this rare Trimorphodon
in the collection, two adult females, one adult male, and one juvenile.
Heretofore the known range was Morelos and southeastern Puebla.

The only discernible difference between these specimens and the
group described by Taylor* is in the number of spots. The num-

* Taylor, E. H., Univ. Kan. Sci. Bull., vol. 26, 1939 (1940), p. 479, pi. 52.

208 The University Science Bulletin

bers recorded by Taylor are 13-15 on the body and 5-7 on the tail,
with the highest total ( body -f- tail ) being 22, and the lowest total
being 19. The four specimens in this collection have a range of
15-17 spots on body and I'A-WA on the tail, with a high total of 27K
and a low total of 23. The ground color of the adults is tan, ( on the
juvenile, pink), the spots themselves ranging from grey in their
middle to black at their periphery.

Data on Trimorphodon latifascta (Peters)

Number 24100 24101 24102 24103

Sex 9 9 $ 9

Ventrals 218 215 215 215

Caudals 66 74 78 59

Scale formula 23-23-16 23-23-17 23-23-16 23-23-16

Supralabials 8-9 8-8 8-10 8-S

Infralabials 13-13 12-12 13-13 12-13

Loreals 3 3 3 3

Preoculars 3 3 3 3

Postoculars 3 3 3 3

Spots on body 17 15 17 16

Spots on tail 8 8 101/. IV-i

Leptodeira metadata (Hallowell)

Megalops maculatus Hallowell, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, 1860 (1861), p. 468.
Leptodeira septentrionalis metadata Dunn, Proc. Nat. Acad. Sci., vol. 22, 1936. p. 697.
Leptodeira maculata Taylor, Univ. Kansas Sci. Bull., vol. 25, 1938 (1939), pp. 337-342, figs.

6-7, pi. 31, fig. 1, pi. 32, pi. 33, figs. 1-3; Smith, Proc. U. S. Nat. Mus., vol. 93. 1943,

pp. 440-441.
Leptodeira personata Cope, Proc. Acad. Nat. Sci. Philadelphia, vol.- 20, 1868 (1869). p. 310

(type locality, Mazatlan, Sinaloa).

There is a single male specimen of Leptodeira maculata in the
collection (KUMNH No. 23786). The specimen has the following
characteristics: scale formula 21-23-17; ventrals 186; caudals 79;
anal divided; supralabials 8; infralabials 9-10; preoculars 2; postoc-
ulars 2; temporals 1 + 2; chinshields 2; total length 580 mm; tail
length 130 mm. There are 32 black blotches on the body and 17 on
the tail. The dorsal blotches on the body reach the second or third
scale row, the majority reaching only the third. In alcohol the
ground color is tan to gray with black bands.

Coniophanes piceivittis taylori, subsp. no v.

Type. -EHT-HMS No. 23523, Agua del Obispo, Gro., 1940, E. H.
Taylor collector.

Explanation of Plate XXIV

Fig. 1. Coniophanes piceivittis taylori subsp. nov. Type specimen. Dorsal
view of head. Actual head width 8 mm.

Fig. 2. Coniophanes piceivittis taylori. Dorsal color pattern (enlarged, dia-
grammatic).

Fig. 3. Coniophanes piceivittis piceivittis Cope. Dorsal color pattern (en-
larged, diagrammatic).

PLATE XXIV

14—3286

210 The University Science Bulletin

Paraty pes. -EHT-HMS Nos. 23524, 8 m. east of Chilpancingo,
Gro., E. H. Taylor collector; 5197, Km 357 on Mexico-Acapulco
Highway, Gro., E. H. Taylor collector; KUMNH No. 23789, Chil-
pancingo, Gro., W. W. Brown collector.

Diagnosis.— Similar to Coniophanes piceivittis piceivittis in gen-
eral characters but differing in having an elongate frontal, its length
equal to VA times distance of frontal from tip of snout (in p. picei-
vittis shorter than its distance to tip of snout) and with somewhat
shorter prefrontals; color pattern in general similar to p. piceivittis
but differing in having head brown above instead of black; in having
a small cream-colored, black-bordered ocellus (See Fig. 1) on
parietals (no such spot present in p. piceivittis). The inferior
lateral stripe commencing on middle of fourth scale row and oc-
cupying two and two half scale rows as compared with p. piceivittis
which has the lateral stripe commencing on middle of third scale
row and occupying three and two half scale rows. Separating
lateral stripe from median dorsal stripe is an intervening space of
cream ground color, which occupies two and one half scale rows
as compared to VA scale rows in p. piceivittis; median dorsal black
stripe occupies five and two half scale rows, while that of p. picei-
vittis with six and two half-rows, or seven complete scale rows ( See
Fig. 2).

Description of type.— Rostral VA times as wide as high, plainlv
visible from above; internasals two-thirds length of prefrontals;
prefrontals as wide as long; frontal VA times as long as distance from
frontal to tip of snout; parietals VA times as long as the distance
from snout to frontal. Nasal divided, posterior part larger; loreal
squarish, a little longer than high; two preoculars, upper large, lower
small; two postoculars, the upper being the larger; temporals
1 + 2 + 3, the anterior temporal bordering both postoculars. Supra-
labials eight, ascending order of size: 2, 3, 1, 4, 5, 6, 8, 7; fourth
and fifth bordering the eye; infralabials nine, first four bordering
the first pair of chinshields; scale formula 21-25-19; all scales smooth,
except (in male specimens) keeled scales present on the first two
scale rows on sides above anal region; scales without apical pits.
Ventrals 169, caudals 90; anal divided.

Color.— Pattern consisting of three longitudinal black stripes on
a ground color of cream, the stripes commencing on snout and con-
tinuing to within the second or third scale from the tip of the tail.
Lateral black stripe commencing in the middle of the fourth scale
row and occupying two and two half scale rows; separating lateral
from medial dorsal stripe is an intervening space of two and one-

Hall: Snakes from Guerrero 211

half scale rows of cream ground color; median dorsal black stripe
occupies five and two half scale rows. Moving anteriorly, the
median dorsal black stripe becomes brown on entering the parietal
scales and continues as a brown stripe to the rostrum; a small cream
colored ocellus with a black periphery lying in the center of the
parietal region; from eye forward, intervening light stripe peppered
with black spots. Venter immaculate; dark brown or black spots
on chin and infralabials.

Measurements in mm. — Head length ( snout to jaw angle ) 14;
head width 8; total length 340; tail length 98.

Variation.— The paratypes agree in most characters except in scale
counts. A table is appended to show the variations.

Remarks.— The geography of this area does not differ greatly
from that in which the typical subspecies is known to occur in
Oaxaca, Chiapas and in northern Central America. The specimen
referred to by Bailey,* reported to have been collected in Costa
Rica, must be verified.

The specimens of p. taylori which I have before me are some-
what smaller than the specimens of p. piceivittis which I have ex-
amined. Whether or not this is due to the age of the specimens,
I do not know. All were collected at an altitude of from 800-1400
meters elevation.

Data on Coniophanes piceivittis taylori

Museum EHT-HMS EHT-HMS EHJ-HMS KUMNH

No 23523 23524 5197 23789

Sex (or age) £ 9 Juv. $

Scale formula 21-25-19 23-25-19 23-21-25-A 23-25-19

Ventrals 169 171 165 170

Subcaudals 90 87 85 78-

Supralabials 8-8 8-8 8-8 8-8

Infralabials 9-9 9-10 10-10 9-10

Total length (mm.).... 340 330 188 240

Tail length (mm.) 98 85 48 58

Sonora michoacanensis michoacanensis (Duges)

Contia 7>iichaocanensis Duges, in Cope, Proe. Amer. Philos. Soc, vol. 22, 1884 (1885), pp. 178-

179 ("Michoaean").
Sonora michoacanensis michoacanensis Stickel, Proe. Biol. Soe. Washington, vol. 56, 1943,

pp. 113-110.
Sonora erythura Taylor, Herpetologiea, vol. 1, 1937, pp. 69-71, pi. 6, fig. 1, (type locality,

10 mi. south of Taxco, Guerrero).

There are two specimens of Sonora michoacanensis michoacan-
ensis in this collection (KUMNH Nos. 23790, 23791). Scale rows
15-15-15; supralabials 7; infralabials 7; preoculars 1; postoculars 2;

* Bailey, Joseph R., Mich. Acad, of Sci., Arts and Letters, vol. 24, part II; 1938, pub. 1939,
pp. 29, 30, 31.

i

212 The University Science Bulletin

chinshields 2; loreal present; anal divided. Anteriorly on the body
the color is arranged in triads composed of two black bands sepa-
rated by a white band of about the same width. An orange band
two or three times as wide as either the black or the white bands
separates these from the next black and white triad. Near the
middle of body, the orange gives way to black, presenting black
and white diads on the posterior part of the body. However oc-
casionally some orange may be visible in the black annuli on the
ventral surface. The tail is solid orange.

Taylor (loc. cit.) has described this Guerrero species under the
name of Sonoro erythrura, having compared his specimen with the
figures given by Giinther in (1895) Biologia Centrali- Americana.
Reptilia and Batrachia, pi. 36, fig. C, named Homalocranium michoa-
canense and found it different. He was unaware that these speci-
mens of Giinther actually represented an undescribed form, which
has since been named Sonora michoacanensis mutabilis by Stickel
(loc. cit.) (pp. 116-117).

Data on Sonora michoacanensis michoacanensis

No.

Sex

Yentrals

Caudals

Temporals

Total
length

Tail
length

Annuli

23790
23791

9
9

176
177

41
42

1 2

1 1

290
340

50
60

16
15

Oxybelis auratus (Belli

Dryinus auratus Bell, Journ. Zool., vol. 2, 1825, pp. 324-320, pi. 12; "Mexico." (It appears
that auratus antedates acuminatum Wied. The specific and subspecific relationship of Mex-
ican forms is still open to question.)

Oxybelis aeneus auratus Bogert and Oliver, (part), Bull. Amer. Mus. Nat. Hist., vol. 83, 1945,
pp. 381-392, figs. 10-11.

Coluber acuminatus Wied, Abbildungen zur Naturgeshichte Brasilien, Lief. 14, 1822 (1830?),
pi. 1.

Oxybelis acuminatus Steindachner, R^ise der osterreichisehen Fregatte Novara .... Reptilien.
1867, p. 72. Boulenger, Catalogue of the Snakes in the British Museum, vol. 3, 1896,
pp. 192-193. Bocourt, Mission Scientifique an Mexique et dans l'Amerique centrale; Etude
sur les reptiles, livr. 15, ls!»7, pp. 83S-840, pi. 65, fig. 4. Taylor, Univ. Kansas Sci. Bull.,
vol. 27, 1H41, p. 138. jil. 6, figs. 7. 8, 9.

Oxybelis fulgidus Crimmins (nee Daudin), Copeia, 1937, No. 4, p. 233.

Two specimens obtained from the Chilpancingo region referred
to this species, give the following data: preoculars 1, postoculars 2,
supralabials 8-8, scale formula 17-17-13. The coloration is normal
for the species.

Data on Oxybelis atratus (Bell)

No.

Sex

Vent ra Is

Caudals

Infra -
labials

Body

length

Tail
length

23795
23798

9

$

195
192

177
184

9- 9
9-10

1235
1220

500
510

THE UNIVERSITY OP KANSAS

SGIENCE BULLETIN

Vol. XXXIV, Pt. I] October 1, 1951 [No. 5

A New Subspecies of the Mexican Moccasin,
Agkistrodon bilineatus *

W. Leslie Burcer and William B. Bobertson

Two specimens of Agkistrodon bilineatus Giinther from northeast-
ern Mexico represent a distinct subspecies for which no name has
yet been proposed. The localities where these specimens were col-
lected are in a semi-arid area covered with desert shrub vegeta-
tion, as contrasted to the humid, lowland jungle where A. bilineatus
is usually found. The nearest locality records for typical A. bilin-
eatus are several hundred miles distant from those known for the
new subspecies.

Our attention was first called to this interesting situation by Dr.
Edward H. Taylor, who collected the specimen which we have
designated holotype of the new subspecies, and to whom we are
indebted for permission to study it and other specimens of the
species in his collection. For advice and assistance, including the
loan of specimens, we wish in addition to thank Dr. Howard K.
Gloyd, Dr. Hobart M. Smith, and Mr. Clifford H. Pope.

We have used Gloyd and Conant's ( 1943 ) scheme of description
of races of Agkistrodon eontortrix as a model for the following de-
scription.

Agkistrodon bilineatus taylori new subspeciesf

Agkistrodon bilineatus Taylor. Univ. Kansas Sci. Bull., vol. 2(i, no. 14, 1940, p. 486. — Schmidt
and Owens, Zool. Ser. Field Mus. Nat. Hist., vol. 29, no. 6, 1944, p. 113.

Holotype— E. H. Taylor Collection No. 5514, a young male from
km. 833, 21 kilometers north of Villagran, Tamaulipas, collected
alive by Edward H. Taylor on June 9, 1938, on the highway pave-
ment about dark. Paratype. — Chicago Natural History Museum No.

* Contribution from the Museum of Natural History and the Department of Zoology, Uni-
veisity of Illinois, Urbana, Illinois.

t Named for Dr. Edward H. Taylor in recognition of his many contributions to our knowl-
edge of the Mexican herpetofauna.

(213)

214 The University Science Bulletin

28794, adult male from south of Linares in the region of the bound-
ary between the state of Tamaulipas and Nuevo Leon, Mexico. | The
exact locality is unknown but undoubtedly is no more than a few
miles from the type locality.

Diagnosis.— This subspecies differs from Agkistrodon bilineatns
bilineatns in the following respects : ( 1 ) lower light line on side of
head not bordered below by a dark line posterior to the second su-
pralabial; (2) fewer subcaudals, 51-54 in males; and (3) a pattern
of dorsal crossbands which remains distinct in adults.

Description of holotype.—Head shields arranged as in Agkistrodon
bilineatns bilineatns: paired and bilaterally symmetrical internasals,
prefrontals, supraoculars, and parietals, and a single rostral and
frontal; posterior border of parietals irregular and broken by several
sutures which extend a short distance anteriorly. Nasals 2, the an-
terior large and crescent-shaped, the posterior smaller and approxi-
mately square with much rounded corners. Loreal about as long
as high, smaller than upper preocular and about the same size as
posterior nasal. Three preoculars, the uppermost larger, the middle
one forming the upper and posterior borders of pit, and the lower-
most very small. Separated from orbit by the last is an elongate
scale which forms the lower border of pit. One large subocular and
two postoculars. Supralabials 8, the first smaller, irregularly quad-
rangular, and lying entirely below anterior nasal; the second larger
and elongated upward to form anterior border of pit; the third,
fourth, fifth, sixth, and seventh largest and about equal in size; the
posterior part of third and most of fourth beneath eye; the eighth
shorter than others. Infralabials 11 on each side, the anterior pair
meeting immediately posterior to mental. Temporals irregular, 4 to
6 in each vertical row. A single pair of enlarged chinshields fol-
lowed by four pairs of scales, the last which is separated from the
labials by 4 longitudinal rows of scales on each side.

Dorsal scales keeled on the back, the keels reduced on the sides
and absent from the lower two rows on each side; scale rows odd
in number, progressively reducing bilaterally toward the tail as fol-
lows : f

$ Due to an error in the museum catalogue, locality data for the latter specimen was er-
roneously recorded by Gloyd and Conant (1943) ; but the record was subsequently corrected by
Schmidt and Owens (1944).

H In the following scale count formula the Clark and Inger recount system (1942) is used
with several modifications. Changes of an anomalous nature are enclosed in brackets and the
number of rows covering the loci where the usual three counts are made are printed in bold-
faced type. In a number of specimens the anterior and mid-body counts are the same (this
is the case in all instances where only two numbers are in bold-faced type). An abbreviated
formula omitting the number of the scale rows involved in each change is used for all except
the holotype and paratype.

Burger and Robertson: A New Subspecies

215

li 10 15

(7+8) (5 + 6) (5 + 6)

20— —27— —25— —23

(6 + 7) (5 + 6) (5 + 6)

7 10 15

22—

(1 + 5) (5)
28 30

-23-

(4 + 5)
32

-22

(5)
40

23

66 101 133
(4 + 5) (5)

il— —19

(4 + 5) (4 + 5)
64 99 133

The ventrals are 133; subcaudals 54, the last 19 divided. The
anal is entire but has the posterior left corner almost cut off by
a suture extending from the left anterior corner toward the middle
of the posterior edge. Terminal spine short and blunt, scarcely
protruding beyond the last pair of subcaudals. The snout-vent
length is 383 mm. and the tail length, 82 mm.

"Head grayish-black above, more grayish on the sides of head
and chin; a yellowish-white line from tip of snout along the canthus
rostralis to angle of the jaw, where it joins a line originating on
the anterior nasal which runs across the labials and across angle of
the jaw; below the white line on labials, the lip is edged with
amber-orange; a vertical stripe on rostral which is white with am-
ber-orange center; this connects, when mouth is closed, with a
stripe extending back from mental to the posterior chinshields and
then bifurcates, the lines continuing back to first widened ventral.
The lines are here joined by another white line; two amber-orange
lines extend from sixth lower labial diagonally to the bifurcating
lines.

"Body generally lavender gray, traversed by lighter gray, irregular,
saddle-like blotches, with interrupted amber-orange borders, which
join and form a large amber-orange spot low on side; intervening
areas may be divided by a very dim medial band, which joins a
black, light-edged spot on the ventrals. Belly generally dark with
amber-orange spots and reticulations. Tail yellow-green." §

Dark dorsal crossbands 12 in number on the body, the first con-
tinuous with the darker portions of the head, the last extending
onto the base of the tail, all wide and slightly constricted medially,
averaging 10 scales in width on the sides and 8 at the midline. Each
anterior and posterior border of the bands fuses with a lateral belly
spot. Each band has a slightly lighter median band across the
center which expands on the sides to enclose a third lateral ventral
spot for each band. This middle lateral spot is usually more exten-
sive than the others, reaching to or beyond the middle of the ven-
trals and often joining the corresponding spot of the opposite side.
Posteriorly the dark ventral coloring becomes extensive and all of
the lateral ventral spots tend to be connected. The dark brown ver-

§ Color of the holotype in life, quoted from Taylor (1940, p. 486).

216 The University Science Bulletin

tical band on each side of the light rostral stripe extends posteriorly
along the upper lip only to the middle of the second supralabial.

Variation. — The two specimens of taylori," the holotype and para-
type, agree closely in coloration and scutellation. The lower light
head stripe of the paratype has a dark border only on the first and
adjacent third of the second supralabial on the left and on the first
supralabial on the right. The dorsal crossbands are distinct, as in
the holotype, but slightly narrower averaging 9 scales wide on the
sides and 7 scales at the midline. Ventrals are 133 in number,
caudals 51, snout-vent length 553 mm., tail length 125 mm., supra-
labials 8, infralabials 10. The dorsal scale rows are as follows:

11 14 85 106 133
(7 + 8) (6 + 7) (4 + 5) (5)

27— —25 23 — 21— —19

(7 + 8) (6 + 7) (4 + 5) (5)

11 11 77 106 133

Discussion.— The two specimens of this subspecies were com-
pared with ten specimens of the typical subspecies from the follow-
ing localities: j | MEXICO: no definite locality (CNHM 42147,
CNHM 56376); Colima: no definite locality (CAS 5253), Tecoman
(CAS 11333); Michoacdn: Apatzingan (CNHM 39093), El Sabino,
Uruapan (EHT-HMS 5357); Oaxaca: Rincon Bamba (EHT-HMS
28019); Yucatan: Libre Union (CNHM 36253), Merida (CNHM
19425). BRITISH HONDURAS: Belize (CNHM 4196). In addi-
tion the summary of the data for 46 specimens of bilineatus given by
Gloyd and Conant ( 1943, p. 168 ) proved very useful.

The dark lower border of the lower head stripe in all of the speci-
mens of bilineatus which we have examined extends to at least
the sixth supralabial and usually to the seventh or eighth supra-
labial. This is in marked contrast to the condition in taylori in
which the dark border does not extend beyond the middle of the
second supralabial (see pi. 25).

With one exception the numbers of ventrals and caudals in these
specimens of bilineatus fall within the ranges given by Gloyd and
Conant ( 1943, p. 168 ) . The numbers of subcaudals of the two males
of taylori, 54 and 51, fall well below the range in number of sub-
caudals of male bilineatus ( 59-68 ) . The exception mentioned above,
a male bilineatus iXNHM 39093) with 69 subcaudals, extends this

° Hereafter the single names taylori or bilineatus refer to the subspecies unless otherwise in-
dicated.

|| The following abbreviations are used for the collections from which specimens were ex-
amined: Chicago Academy of Sciences (CAS). Chicago Natural History Museum (CNHM),
Edward H. Tayloi -Hobart M. Smith Collection (EHT-HMS).

Burger and Robertson: A New Subspecies 217

range slightly. The number of ventrals in both specimens of taylori,
133, is well within the range of variation of male bilineatus.

In adult bilineatus the dorsal crossbands exhibited by juveniles
are obscured by the general darkening of the ground color (see
Gloyd and Conant, 1943, fig. 11). The dark adult coloration is ac-
quired in this subspecies at a rather small size, probably during the
second or third year. A male of bilineatus (CNHxVI 1133), 587 mm.
in snout-vent length, shows the typical melanistic coloration. In
contrast taylori retains distinct cross bands for a longer time, pos-
sibly throughout life ( see pi. 25 ) . The crossbands of the paratype,
with a snout-vent length of 553 mm., are distinct with no indication
of becoming obscured. For this reason the color pattern of A. b.
taylori bears a general resemblance to that of A. contortrix picti-
gaster and of juvenile A. piscivorus leucostoma, the geographic-
ranges of both of which it closely approaches ( see Gloyd and Con-
ant, 1943, figs. 10, 13). These three forms also show convergence
in number of subcaudals. The exact phylogenetic significance of
these trends is not certain. However, it seems very unlikely that the
species bilineatus through its subspecies taylori will be found to in-
tergrade with either the copperhead or cottonmouth.

The specimens available to us do not indicate any subspecific
difference in proportionate tail length. Measurements of larger
series of both subspecies would be required, as ontogenetic changes
in proportionate tail length are obviously involved. Similarly no
subspecific differences could be found in the number of dorsal scale
rows. To illustrate the variation several abbreviated scale row
formulae of bilineatus are given below:

13 91 120 138
25— —23— —21— —19

14 89 114 138

14 84 96 132
25— —23 21— —19

15 85 97 132

23 105 132

25 23 21 —20

CNHM

36258

2

CNHM

39093

rt

EHT-HMS

28019

tf

9

97

9

11

10

12

27-

15 17 104 133 132

If the variation in the location of each scale row reduction is
noted, no definite difference is apparent between bilineatus and tay-
lori (see Table 1). The average point of each scale row reduction
may be found to be more posterior in bilineatus when larger series
of each subspecies are examined.

218

The University Science Bulletin

Table I. — Ordinal Number of Ventral Where Each Dorsal Scale Row

Reduction Occurs *

Scale Row Reduction

Subspecies

27#->-25

*-" VW r *>&

23#->-21

21#->-19

-4. 6. taylori

10-11

11-15

64-85

99-106

9-15

13-23

84-105

96-133f

* Based on only 2 specimens of taylori (both males) and three of bilineatus (2 males and 1
female).

t In one specimen no last reduction occurred anterior to the anus on one side.

In the three characteristics mentioned in the diagnosis, taylori is
apparently completely distinct. Its affinity with A. bilineatus is,
however, unmistakable. Future collecting will probably reveal in-
tergading populations somewhere in the Atlantic coastal plain be-
tween central Tamaulipas and Yucatan. Even in the event that such
intergrades fail to appear, we feel that the relatively minor nature
of the differences between taylori and bilineatus indicate existence
of a subspecific relationship between them. We are in accord
with Mayr's (1947, p. 166-167) opinion that, in the case of forms not
known to occur together, the degree of morphological difference as-
sociated with reproductive isolation in forms of the same group
known to be species should be applied as a yardstick to determine
whether the forms in question are best to be considered as species
or subspecies. Only in the event that actual overlap without
intergradation was found to occur would one be justified in con-
sidering taylori a distinct species.

LITERATURE CITED

Clark, Philip J., and Robert F. Inger

1942. Scale reductions in snakes. Copeia, 1942, no. 3, p. 163-170.

Gloyd, Howard K., and Roger Conant

1943. A synopsis of the American forms of Agkistrodon (copperheads and
moccasins). Bull. Chicago Acad. Sci., vol. 7, no. 2, p. 147-170,
fig. 1-16, map 1-2.

Mayr, Ernst

1942. Systematics and the origin of species. New York: Columbia Uni-
versity Press. XIV + 334 p., 29 figs.

Schmidt, Karl P., and David W. Owens

1944. Amphibians and reptiles of northern Coahuila, Mexico. Zool. Ser.
Field Mus. Nat. Hist., vol. 29, no. 6, p. 97-115.

Taylor, Edward H.

1940. Some Mexican serpents. Univ. Kansas Sci. Bull., vol., 26, no. 14,
p. 445-487, pi. 49-52, fig. 1-9.

Plate XXV. Fig. 1. Dorsal view of the holotvpe of Agkistrodon bilineatus
taylori subsp. now Total 678 mm. E.H.T.-H.M.S. No. 5514. Fig. 2. Lateral
view of head of holotvpe of A. b. taylori (enlarged). Fig. 3. Lateral view of
head of Agkistrodon bilineatus bilineatus, E.H.T.-H.M.S. from YA Sabino,
Uruapan, Michoacan.

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXIV, Pt. I] October 1, 1951 [No. 6

An Anatomical Study of Certain Salamanders of
the Genus Pseudoeurycea

BY

Irwin L. Baird
TABLE OF CONTENTS

PAGE

Introduction 221

Anatomy of Pseudoeurycea Bellii 223

External Anatomy— Female 223

External Anatomy— Male 226

Buccal Cavity 226

Viscera 228

Skull 229

Hyobranchial Apparatus 233

Vertebrae 234

Skeleton of the Girdles and Limbs 236

Hyobranchial Musculature 237

Musculature of the Mandible 241

Musculature of the Body Wall 243

Musculature of the Pectoral Girdle and Forelimb 245

Musculature of the Pelvis and Thigh 248

Comparison of Other Species with Pseudoeurycea Bellii 252

Summary and Conclusions 258

Literature Cited 260

INTRODUCTION

The aims of this investigation have been three in number: first,
to describe in some detail, certain features of the anatomy of sala-
manders of the genus Pseudoeurycea Taylor; second, to determine
which anatomical features might best be used in determining the
taxonomic relationships within the genus; and last, to inquire into
the phylogenetic relationships of the several species studied. So far
as I have been able to ascertain, the general anatomy of this group
has never been recorded, nor have features other than those of the

(221)

222 The University Science Bulletin

external and skeletal anatomy been used in estimating the relation-
ships within the group.

Pseudoeurycea, as proposed by Taylor ( 1944 ) , is composed of
eighteen species and subspecies, and these, on the basis of distribu-
tion and characteristics of the skeleton and external anatomy, are
arranged in five species groups. These are confined to the Mexican
Plateau, south from Zacatecas and Nuevo Leon, except that two
species, P. rex and P. goebeli, occur in Guatemala.

For reasons of continuity and clarity, this paper has been divided
into two parts. Part I is purely descriptive and deals with the anat-
omy of P. bellii, the most widely distributed of the species in the
genus. The description of the external anatomy and buccal cavity is
based upon the examination of fifteen specimens taken near Omil-
teme, Guerrero, Mexico; descriptions of the myology, body organs
and cartilaginous parts are from dissections made of a female ( Cata-
log Number 24274 ) * and a male ( No. 16339 ) from the same locality.
A cleaned and disarticulated skeleton ( No. 24275 ) and a cleared and
stained specimen (No. 16346) constituted the material from which
skeletal descriptions were made.

Part II consists of brief statements of anatomical differences found
in certain other species of the genus. For this phase of the work,
a male and female of one species from each of the five species groups
suggested by Taylor (op. cit.), were dissected and their anatomy
compared to the structures considered in P. bellii; cleaned skeletons
and cleared specimens were used for the study of the osteology.
Additional material available for dissection was checked for cer-
tain differences observed in the first specimens of the species group
examined; cleared specimens and skeletons were studied and exter-
nal characteristics recorded where possible. The specimens used
are listed below.

Gadovii group: P. gadovii from Veracruz (state), Mexico; ten
preserved specimens (two dissected), two skeletons, one cleared
specimen; P. imguidentis from Oaxaca (state), Mexico, two pre-
served specimens (one dissected), one skeleton. Smithi group:
P. smithi from Oaxaca (state), Mexico, ten preserved specimens
(two dissected), one skeleton, two cleared specimens. Cephalica
group: P. cephalica cephalica from Puebla (state), Mexico, ten pre-
served specimens (two dissected), one skeleton, two cleared speci-
mens. Leprosa group: P. leprosa from Puebla (state), Mexico, ten
preserved specimens (two dissected), five skeletons, five cleared

♦All catalog numbers refer to the EHT-HM3 collection of Dr. E. H. Taylor.

Baird: The Genus Pseudoeurycea 223

specimens; P. cochranae from Oaxaca (state), Mexico, one preserved
specimen (dissected), one cleared specimen; P. robertsi from Mexico
(state), Mexico, one preserved specimen (dissected); P. rex and
P. goebeli from near Tecpan, Guatemala, one preserved specimen
each ( partially dissected ) .

Unless otherwise indicated, the teminology of the muscles is that
accepted by Francis ( 1934 ) .

I am indebted to Dr. Edward H. Taylor, Professor of Zoology,
University of Kansas, for guidance, criticism and suggestions con-
cerning this work, and for the use of literature and specimens with-
out which it could not have been undertaken. Professors E. Ray-
mond Hall and A. Byron Leonard, University of Kansas, and Dr.
Dorthea S. Franzen, Assistant Professor of Zoology, Washburn Uni-
versity, have given valuable assistance by criticism and suggestions
during the preparation of the manuscript and the illustrations.

ANATOMY OF PSEUDOEURYCEA BELLII
External Anatomy — Female

The ground color in the female is bluish black, marked dorsally
and laterally with distinct maculations which are red-orange in life,
but creamy yellow in preserved specimens. The head bears one
pair of maculations, usually in the parietal region immediately pos-
terior and medial to the eye. Posterior to the nuchal groove the
dorsum is marked with a series of paired spots, each pair divided
by the dorsal midline and separated from the adjacent pair by an
intervening costal groove. Cranially, in the region of the pectoral
girdle, dorsal spots closely approximate the dorsal midline and
frequently fuse across it; posterior to this area, maculations are
separate and distinct.

In some specimens, indistinct and irregular spotting may occur
laterally in the region of the sulcus lateralis. Rarely, for macula-
tions are normally confined to the trunk, the first five or six seg-
ments of the tail bear marks similar to those on the trunk.

The head, broad and depressed, is only slightly longer than wide
and the snout is exceptionally blunt. Maximum width of the head
and maximum gape are attained at the posterior border of the eye.

The nostrils are as far apart as are the anterior extremities of the
orbits, and from the ventro lateral margin of each nostril, a naso-
labial groove extends ventrad to terminate at the mid-point of the
subnarial swelling directly beneath. The latter is round and promi-
nent; its center lies the width of the nostrils above the margin of the
upper lip.

224 The University Science Bulletin

The greatest diameter of the eye is contained at least three times
in the snout to gular fold measurement. The eyes themselves are
prominent, directed forward and slightly laterad, and are situated
approximately two thirds their own length posterior to the tip of
the snout. A postocular groove extends caudad from the posterior
angle of the eye and disappears at the juncture of the gular fold
with the nuchal groove. The postocular groove is crossed by a
transverse groove which lies approximately midway between the
angle of the mouth and the angle of the jaw. This groove begins
lateral to the dorsal midline of the head, and roughly paralleling
the posterior margin of the skull, passes ventrad onto the ventral
surface of the head to approach the groove of the opposite side.
The postocular groove is usually less evident posterior to this point.

The gular fold is quite evident ventrally, but at the level of the
angle of the jaw the fold of skin disappears. From this point the
nuchal groove extends dorsad and curves slightly craniad on the
dorsolateral aspect of the cervical region and terminates just short
of the dorsal midline at the level of the juncture of the atlas with
the first trunk vertebra.

Dorsally the skin of the head is densely pitted, and pierced by
pores of small subcutaneous glands situated beneath the pits. On
the upper lip and on the under side of the head, pits are reduced
or absent, but the pores, slightly reduced in size, are extremely
dense. Thinness of the skin on the ventral surface allows the dis-
cernment of rounded subcutaneous glands in this region. In most
preserved specimens, large tan and smaller cream-colored glands
can be differentiated.

Anteriorly the trunk is approximately the same width as the head,
but at the level of the first two costal folds, the width is consider-
ably reduced. Posterior to this point the trunk again broadens; the
degree of this depending upon the age, sex and state of reproduc-
tive activity of the animal. From a point anterior to the pelvic re-
gion, the trunk tapers gently toward the constriction marking the
junction of the trunk with the tail. This constriction is normally
situated immediately posterior to the anus at the level of the pos-
terior margin of the second caudal vertebra. The anus itself appears
as a longitudinal slit in the venter immediately posterior to the pel-
vis. The cloacal lips are marked with a series of small folds of skin.
The twelve or thirteen well-marked costal grooves are on the lateral
surfaces of the body and except for a narrow strip along the mid-
ventral line, are quite visible on the ventral surface. The skin of
the dorsal and dorsolateral surfaces of the trunk is densely pitted

Baird: The Genus Pseudoeurycea 225

and subcutaneous glands may be discerned through the skin. Large
reddish tan glands may be differentiated from smaller, creamy yel-
low glands. On the venter, pitting is much reduced and glands are
fewer; reduction of the number of the smaller glands is particularly
noticeable. About the anus, however, subcutaneous glands are
present in considerable numbers, and their pores are large.

Immediately posterior and dorsal to the base of the thigh there
is an elongate oval area with pigment much reduced and with a
large concentration of small light-colored subcutaneous glands. This
concentration probably serves some specialized function for it is
fixed in position, general structure and appearance in a series of
fifteen specimens checked, and at no other point of the body is an
equally dense concentration of glands visible. Though this area is
mentioned in the literature, no name has been found designating
it; it is suggested therefore, that it be termed the post-iliac gland.

Extending caudad and dorsad from the dorsal extremity of the
gular fold, a ridge marking the position of the m. subarcualis rectus
I ( and the epibranchial cartilage which it surrounds ) , passes dorsal
to the insertion of the forelimb and terminates at the anterior part
of the third costal fold.

Both forelimbs and hindlimbs are stout and long enough to be
well adapted to walking; one to four costal folds separate the digits
of the adpressed limbs. The hand is broad and depressed, the pal-
mar aspect presenting a roughly rounded outline. The four digits,
in order of length 3-2-4-1, show no webbing of the two distal pha-
langes in the second, third and fourth digits, and the distal phalanx
of the reduced first digit is also free. Each of the digits carries a
prominent, ventr ally-projecting lobe or pad of tissue which covers
the ventral surface of the distal phalanx. Five digits are present
on the foot, in order of length 3-4-2-5-1. These tend to be more
widely spread than the digits of the hand but the distal phalanges
carry pads similar to those mentioned above. The reduced first
and fifth digits have but one phalanx free, whereas the other three
have two or two and one-half unwebbed phalanges.

The tail is long and powerful, constricted at the base, and roughly
round in cross-section. In large specimens the anterior part of the
tail is greater in diameter than the terminal part of the trunk, but in
smaller specimens, the greatest diameter may occur near the third
or fourth caudal fold. The tail is deeply marked laterally by caudal
grooves, which become slightly less evident toward its posterior
extremity. From its greatest diameter near its anterior limit, the
tail is tapered smoothly to its tip.
15—3286

226 The University Science Bulletin

External Anatomy— Male

In males, coloration and markings are similar to those in the fe-
male, but there is a more marked lightening of the ground color on
the lateral and ventral surface in most specimens. The nostrils are
slightly more elongate transversely, tapering toward their lateral
margins, and the nasolabial groove divides into several faint fur-
rows at the ventral or antero-ventral limit of the mildly pendent sub-
narial swelling. The latter attains its maximum prominence im-
mediately ventral to the margin of the upper lip.

A submental gland almost round in outline, and smooth and flat-
tened on its ventral surface is situated behind the anterior extremity
of the lower jaw. The antero-posterior diameter of the gland is ap-
proximately one sixth the distance from the snout to the gular fold.
Removal of the gland reveals that it is dorsally rounded and fits,
posterior to the mandibular symphysis, in the curve of the lower
jaw. Since neither ducts nor openings of such structures could be
found associated with the submental gland it has been tentatively
concluded that the glandular product must be exuded through the
minute pores distributed densely in the skin covering the gland.

Except that the cloacal lips are papillate and the dorsal macula-
tions near the posterior limit of the trunk are the site of heavy con-
centrations of subcutaneous glands, no other differences in the ex-
ternal appearance of the two sexes were noted.

Buccal Cavity

The mucous membrane of the floor of the mouth is marked by
rugose longitudinal folds which are continuous anterior to the
tongue. Posterior to the mandibular symphysis and immediately
anterior to a depressed area where the membrane is smooth, these
folds are accentuated to form a distinct curved sublingual fold.
Posterior to this, the stalk of the boletoid tongue projects into the
buccal cavity through the center of the depressed area, and is affixed
to the body of the tongue anterior to its midpoint, in the curve of a
ventrally-projecting U-shaped ridge of tissue. The latter fits well
into the depression in the floor of the mouth when the tongue is fully
retracted. The dorsal surface of the tongue is densely covered with
large papillae arranged roughly in rows radiating from the center
of the surface.

The mandible carries from 34 to 44 ( average 40) short teeth fused
to the medial side of the dentary. The tip of each tooth curves
sharply inward, and each tooth carries a subterminal point on its

Baird: The Genus Pseudoeurycea 227

anterior surface, just proximal to the terminal tip. The extremities
of both the terminal and subterminal points are marked with red-
dish brown. Anterior teeth are approximately vertical or slant but
slightly inward but immediately anterior to the articulation of the
mandible, the teeth are slanted almost horizontally into the buccal
cavity.

The palatine surface of the mouth is covered with heavy mucous
membrane and is bounded anteriorly and laterally by the upper lip
and the teeth of the premaxilla and maxillae. The premaxillary
dentition shows marked sexual dimorphism; in adult males there
are but four teeth on the premaxilla, while in females, thirteen teeth
are usually present. The premaxillary teeth in males are greatly
elongated, penetrate the upper lip and no subterminal point is vis-
ible on them. Premaxillary dentition of this sort is evidently not
present on young individuals, for in two immature males, premaxil-
lary teeth were similar in number and structure to those of females.
The premaxillary teeth in females closely resemble those of the man-
dible, and form a row aligned with the rows of maxillary teeth but
separated from the latter by a short diastema. Maxillary teeth
varied in number from thirty-four to fifty in the specimens examined,
the larger numbers present in older specimens. The females showed
the widest range of variation in the number of teeth present on the
maxilla and limited data tend to indicate that teeth may be added
on this element with increase in size and age. From thirty-five to
forty maxillary teeth were present in the males examined. The
internal nares are situated approximately twice their own antero-
posterior diameter postero-lateral to the premaxilla, and approxi-
mately half that distance anterior to the rows of prevomerine teeth.
The openings of the internal nares are oval, but each opening is
extended by a cleft at its lateral margin. This cleft, which I term the
latichoanal fissure, follows the outline of the slit in the prevomer, ex-
tends to the lateral extremity of the prevomerine tooth row, then is
continued caudad by a groove in the mucous membrane. This
groove is medial to the base of the fold of mucous membrane, bor-
dering the maxillary teeth. At the posterior limit of the maxilla, the
groove turns caudo-mediad and disappears almost immediately.

The lateral extremities of the rows of prevomerine teeth are sepa-
rated from the midpoints of the maxillae by approximately the di-
ameter of the internal naris, and from each other medially, by
approximately half that distance. Laterally the prevomerine teeth
are in a transverse row, but medial to the internal nares, the rows

228 The University Science Bulletin

curve caudally as they approach the midline. From eighteen to
twenty-one teeth were present in prevomerine rows in the speci-
mens examined.

Two patches of paravomerine * teeth lie approximately twice the
diameter of the internal naris posterior to the medial extremities of
the prevomerine rows. The patches are separated medially by a
narrow grove, are wider posteriorly than anteriorly, and extend to
the level of the angle of the jaw. The teeth of these patches are
arranged in oblique rows resembling caudally-directed chevrons,
but no constancy could be found in either the number of rows or
the number of teeth. Paravomerine and prevomerine teeth are simi-
lar to the teeth of the mandible.

Viscera

Though the viscera probably merit detailed study, seasonal vari-
ations in the reproductive organs, and daily variation in the alimen-
tary canal, make comparisons impractical unless large series of
specimens are available. I am, however, calling attention to a few
of the structures noted in the specimens examined.

The stomach is large and extends caudad for fully three fourths
the total length of the coelom. The duodenum extends craniad along
the right wall of the stomach and comes to lie dorsally and to attach
to the liver near its posterior margin. The ileum and colon are of
approximately equal length and each is slightly longer than the
duodenum. The ventral mesentery is but slightly reduced and is
present along the full length of the coelom. Extending along the
midventral line, the mesentery attaches anteriorly to the ventral-
surface of the liver and clearly shows continuity with the gastro-
hepatic and hepatoduodenal ligaments at the posterior margin of
that organ. The dorsal edge of the mesentery is not attached for a
distance equal to the width of one costal fold posterior to the liver,
but at that point it envelops the urinary bladder and attaches to the
ventral surface of the colon throughout its length.

The structures of the urogenital systems of male and female are
delicate and difficult to dissect; consequently a histological study of
serial transverse sections probably will have to be made to deter-
mine all the relationships. In both sexes the mesonephroi appear
to be loosely organized structures composed of tiny, coiled tubules.
In females, the mesonephroi are broad, lateral to the cloaca, and
taper slowly toward their anterior extremities just craniad to the

* This name was first proposed by Dr. Hobart M. Smith and was published by Dr. Edward
H. Taylor (1944:206).

Baird: The Genus PsErDOEURYCEA 229

anterior limits of the ovaries. In males, the mesonephroi are well
defined only at the posterior limit of the coelom; anteriorly, slender
extensions of the organs are attached to the medial surface of the
male duct.

In females, there is a slender straight oviduct extending retro-
peritoneally to the cloaca, from a well-defined ostium attached to
the transverse septum and ventral mesentery. Lateral to the an-
terior part of the mesonephros, a tiny duct (probably the mesone-
phric duct), is separable from the medial surface of the oviduct,
but in the region of the broader posterior part of the mesonephros,
that duct cannot be discerned. In the latter region, well-defined
urinary tubules leave the mesonephros and appear to enter directly
into the oviduct; however, the mesonephric duct may be present,
fused to the oviduct. In males, somewhat caudal to the position of
the ostium in the female, a tiny opening with slightly flared lips
communicates with a delicate, straight, retroperitoneal duct. Im-
mediately cranial to the anterior limit of the testis, this duct is con-
tinuous with a large convoluted duct suspended by a distinct mesen-
tery. With the anterior extension of the mesonephros attached to
its medial wall, this duct extends caudad, receives the efferent duc-
tules from the anterior lobe of the testis, and from the mesonephros,
receives several urinary tubules before entering the cloaca.

Anteriorly there is marked similarity in the internal organization
of the cloacae of the male and female. In both sexes the lining is
thrown into deep longitudinal folds, the urogenital ducts open into
the dorsolateral walls, and the urinary bladder opens into the ven-
tral wall of the cloaca. Near the cloacal lips however, the lining of
the cloaca becomes papillate in the male, while in the female, the
folds are reduced but otherwise unchanged.

Three pairs of tubular glands empty into the cloacae of males.
The ventral cloacal glands are gray in color and are divided into
two parts by the mm. ischio-caudalis. The abdominal glands are
rust-red. They are composed of longer tubules than are the cloacal
glands, and are situated antero-dorsal to the cloaca. The pelvic
glands, short and creamy white, are situated posterior and dorsal
to the cloaca. All open directly through the cloacal wall.

Skull

The skull is broad and depressed, is approximately as wide as it
is long, and its greatest depth (measured from the ventral surface
of the parasphenoid to the dorsal surface of the parietal), is con-
tained approximately three and one-half times in its maximum

230 The University Science Bulletin

length. With the bones of the lower jaw in their normal position,
the skull appears roughly ovoid from above. The fused premaxillae,
the maxillae and the mandibles form the margins curving to the
point of greatest width at the posterior limits of the quadrates. The
otic-occipital region extends cauded and narrows slightly to form
the broader posterior curvature of the skull. Viewed laterally the
skull is bluntly wedge-shaped, its depth increasing from its anterior
to its posterior limits.

The skull is composed of the parasphenoid, the fused premaxillae,
and paired nasal, maxillary, prevomer, prefrontal, frontal, parietal,
otic-occipital, squamosal, quadrate, and orbito-sphenoid bones, and
the pterygoid cartilages. These elements are considered in the fol-
lowing paragraphs.

The premaxillae are fused into a single bone and a pair of frontal
processes arise from a medially thickened elevation on its dorsal
surface. The frontal processes are thin, parallel and moderately
spatulate toward their posterior extremities. They enclose between
them a well-defined fronto-premaxillary fontanelle which extends
cauded to the level of the anterior margin of the orbit. Laterally
the premaxilla is united to the maxilla by cartilaginous connective
tissue, while the maxilla articulates with the prefrontal and nasal
by means of a roughly rectangular frontal process, notched anteriorly
at its base. The external nares are situated at the level of this notch
and are approximately midway between the frontal processes of
the premaxilla and maxilla; no septo-maxillary element is present.
Posteriorly, the frontal process of the maxilla forms the ventral half
of the anterior margin of the orbit, and dorsally articulates with the
poorly-ossified prefrontal and nasal. The former is affixed to the
posterior fourth of the frontal process and forms the dorsal half of
the anterior margin of the orbit, while the latter is affixed to the
anterior half of the frontal process, roofs the nasal capsule and forms
the dorsal margin of the external nares. Both the prefrontal and
nasal articulate with the frontal and frontal process of the premaxilla.
The frontal, roughly leaf-shaped anteriorly, narrows to form the
anterior half of the dorsal margin of the orbit, and overlaps the
parietal posteriorly. The fronto-parietal articulation is accomplished
by means of a dentate process on the posterior margin of the frontal.
The two frontals are strongly articulated by irregular dovetailing
processes carried on the medial margin of each.

The parietal is bluntly pointed at its lateral margin anteriorly.
Tt shows medially, even more distinctly than the frontal, irregular
articulating processes, and is depressed postero-medially and is

Baird: The Genus Pseudoeurycea 231

flared postero-laterally for articulation with the dorsal surface of the
otic-occipital. Approximately at its midpoint, the parietal carries a
ventral projection on its lateral margin. This projection lies medial
to (and serves to strengthen), the dorsal cartilaginous articulation
of the orbito-sphenoid with the otic-occipital.

The otic-occipital is covered antero-medially by the parietal, and
the postero-lateral part of its dorsal surface is depressed, and slightly
ridged by the vertical semicircular canals. The postero-lateral part
of the dorsal surface of the otic-occipital is rounded and furnishes
articulation for the proximal half of the squamosal. Dorso-medially
each otic-occipital presents a well-ossified process; these two pro-
cesses are united at the midline by a cartilaginous remnant of the
synotic tectum.

Antero-laterally, beneath the midpoint of the lateral margin of the
parietal, the otic-occipital carries an oval process, which articulates
with cartilage extending from the dorsal posterior process of the
orbitosphenoid. The bone, rectangular and well-ossified, forms the
lateral wall of the cranial cavity and articulates anteriorly with the
nasal capsule by dorsal and ventral processes. The ventral posterior
articulation of the orbito-sphenoid is accomplished by means of a
short cartilage which joins with a process extending anteriorly from
the base of the otic-occipital. Both of the posterior articulating pro-
cesses of the orbito-sphenoid are short and have only a narrow
groove between them. The base of this groove is formed by narrow
processes extending from each articulation and fusing posterior to
the large optic foramen.

Immediately lateral to the posterior ventral articulation of the
orbito-sphenoid, the same projection of the otic-occipital articulates
with the postero-medial surface of the pterygoid cartilage. This
rectangular element is thick and solid at the point of its articulation
with the otic-occipital, but laterally it thins, is more flexible, and is
attached to the antero-medial surface of the quadrate. Anteriorly,
the pterygoid cartilage is markedly thinner. It forms the postero-
ventral margin of the orbit and attaches to the postero-medial aspect
of the maxilla.

The quadrate is short, heavy and well ossified. Viewed laterally
it appears to be rectangular, but examination of its internal surface
reveals that the major part of the bone is in the form of an irregular
cylinder constricted at its midpoint. The rectangular appearance
is imparted by a thin bony flange on its antero-lateral margin. The
quadrate articulates proximally with a lateral projection of the
otic-occipital, and distally with the lower jaw.

232 The University Science Bulletin

The squamosal is thin and shaped to conform with irregularities
of the elements to which it is attached. The proximal half of the
squamosal is narrow and closely applied to the dorsolateral surface
of the otic-occipital, posterior to the ridge of the semicircular canal.
The distal half of the squamosal widens and is attached to the proxi-
mal half of the lateral surface of the quadrate.

In its postero-lateral wall, just posterior to its articulation with
the quadrate, the otic-occipital has a large round opening, the fe-
nestra vestibulae. This opening is covered by a tough membrane to
which is fused the bony operculum. No columella auris is carried
on the operculum.

The crista muscularis is immediately postero-dorsal to the fenestra
vestibulae and this roughened ridge marks the position of the hori-
zontal semicircular canal. Ventral to each crista muscularis and
ventro-lateral to the margin of the foramen magnum, there is a short
sessile occipital condyle. Antero-medial to these, the otic-occipi-
tals are united in the median plane by processes ossified in the region
of the embryonic hypochordal commissure.

The broad, rounded, posterior part of the parasphenoid is attached
to the ventral surface of the otic-occipitals, anterior to the occipital
condyles and medial to the process articulating with the orbito-
sphenoid. Anterior to this attachment, the parasphenoid tapers to-
ward its spatulate attachment to the ventral surface of the nasal
capsules. The parasphenoid is attached to the ventral margins of
the orbito-sphenoids throughout its length, and loosely attached to
its ventral surface are the two paravomers, discontinuous processes
of the pre vomers.

The postero-medial extremity of the prevomer is firmly fused to
the ventral surface of the anterior part of the parasphenoid. From
this area the dental process of the prevomer curves laterally, forming
the posterior margin of the opening for the internal naris and the
latichoanal fissure. The prevomer extends craniad from its articu-
lation with the parasphenoid and forms the medial margin of the
narial opening, then curves ventrad and widens abruptly to form
an antero-lateral wing. The latter attaches loosely to the premaxilla
and maxilla, and forms the anterior margin of the opening for the
internal naris and the latichoanal fissure. Medially the prevomer
articulates with that of the opposite side by means of a thin carti-
lage.

The dentary is approximately round in cross section in its anterior
third, but posterior to that part, its medial surface is deeply grooved
to receive Meckel's cartilage. Except at its anterior and posterior

Baird: The Genus Psetjdoetjrycea 233

extremities, this cartilage is completely enclosed between the dentary
and angulare. At its posterior extremity the cartilage articulates
with the quadrate.

The angulare is pointed anteriorly to fit the groove in the medial
surface of the dentary, and is solidly attached to the dorsal and
ventral margins of that groove. The angulare forms a coronoid pro-
cess on the dorso-medial margin of the posterior fourth of the mandi-
ble, and its ventral margin extends laterally to invest the ventral
surface of the posterior part of Meckel's cartilage.

Hyobranchial Apparatus

In a general way the hyobranchial apparatus corresponds with the
description given by Smith (1920:536), for that of Eiirycea bis-
lineata. The cartilaginous elements differ somewhat in their con-
figuration however, and neither a lingual cartilage nor an os
thyroideum is present.

The thin, free, dorsally-curved, anterior extremities of the cerato-
hyals extend into the sublingual fold and lend weak support to that
structure. Posteriorly the flattened parts of the cartilages form the
lateral margins of the opening for the stalk of the tongue, then
broaden posterior to that opening to form its postero-lateral mar-
gins and offer broad surfaces for the attachment of the m. supra-
peduncularis. Caudal to the attachments of that muscle, the cera-
tohyals become rounded and extend caudo-laterad deep to the
two superficial layers of hyobranchial muscles. Just posterior to
the level of the angles of the jaws, the ceratohyals curve sharply
dorsad and craniad; their distal extremities come to lie immediately
posterior to the fused quadrates and squamosals, and are attached
to the mid-points of the posterior surfaces of those bones by short
stout hyoquadrate ligaments.

The broad anterior extremity of the basibranchial bears small
antero-lateral horns and is situated and firmly attached within the
body of the tongue. Posteriorly the basibranchial is rodlike and
extends caudo-ventrad for half its own length, within the stalk of
the tongue. At the base of that structure the basibranchial receives
the proximal articulations of the first ceratobranchials on its ven-
tral surface, then extends caudad to articulate with the second
ceratobranchials at its posterior extremity.

The first ceratobranchials are S-shaped and approximately as long
as the basibranchial. They are flattened at their proximal articula-
tions, but posteriorly they twist mediad about their own axes and
become medially concave. The distal third of each is closely ap-

234 The University Science Bulletin

plied and attached to the lateral surface of the distal half of the
second ceratobranchial. The latter is heavy, shorter than the basi-
branchial, and each of the pair extends caudo-laterad to attach, in
company with the first ceratobranchial, to the proximal extremity
of an epibranchial.

The eipbranchials are flexible, slender and approximately three
times as long as the basibranchial. They lie entirely within the
sheaths formed by the mm. subarcualis rectus I, and are tapered
toward their distal extremities.

Vertebrae

The vertebral column consists of one cervical vertebra, fourteen
trunk vertebrae, one sacral vertebra and a variable number of ver-
tebrae posterior to the sacrum. These elements are joined by a
modification of the amphicoelous type of intervertebral articula-
tion; the centra are biconcave but show calcification of the cartilage
contained within their cavities. The amount of cartilage calcified
is variable, and according to Taylor (1944:204), is correlated with
the age of the specimen. In none of the vertebrae examined was
the calcification complete to the lip of the centrum, but in every
case the concavity of the centrum was partly filled with soft partially
calcified cartilage.

The cervical vertebra differs markedly from others in the series,
for the centrum is present only posteriorly. The ventral surface of
the centrum bears a pair of longitudinal bony ridges, which extend
to the posterior limit of the paired lateral projections furnishing
articulation with the occipital condyles. The anterior faces of these
projections are slightly concave, and from between them, a broad
odontoid process extends craniad to fit into the foramen magnum
and articulate with the walls of that opening. The facets of the
odontoid process are convex and situated on its ventro-lateral sur-
faces. A heavy neural arch is present dorsally and that structure
bears a pair of well-developed postzygapophyses on its posterior
margin and a low, rounded, cartilaginous neural spine dorsally.
The trunk vertebrae are not notably modified one from another;
the eighth vertebra is here described as typical. The centrum is
long, slender and biconcave, and its dorsal neural arch is low and
broad. The neural spine is long and low and is formed by a thin
plate of bone affixed vertically to the mid-dorsal line of the neural
arch. The zygapophyses are broad and flat; the articulating sur-
faces of the prezygapophyses are directed dorso-medially, and those
of the postzygapophyses are directed ventro-laterallv. Medial to

Baird: The Genus Pseudoeurycea 235

the postzygapophyses the neural arch carries two small triangular
projections on its posterior margin. The neural canal is rounded
dorsally, flattened ventrally and is completely roofed by the neural
arch. Paired diapophyses and parapophyses extend caudad and
slightly laterad from each trunk vertebra except the last. These
processes are fused except at their bases and at their distal extremi-
ties where they diverge slightly to articulate with the ribs. The last
trunk vertebra bears only parapophyses; the diapophyses are con-
siderably reduced.

The ribs are bicipital, and those of the first two or three trunk
vertebrae may be slightly forked at their distal extremities. Pos-
terior to that, ribs are not modified distally. The last pair of ribs
is reduced to simple cartilaginous stubs carried on the parapophyses
of the last trunk vertebra.

The sacrum is similar to the trunk vertebrae in most respects, but
the parapophyses and diapophyses are markedly heavier, are not
as closely approximated as those of the trunk vertebrae and are con-
siderably expanded at their lateral extremities. The sacral rib is
heavy, deeply forked proximally, and expanded distally for articu-
lation with the ilium.

Posterior to the sacrum, the vertebrae differ markedly from those
discussed above. The first two vertebrae of this series are transi-
tional, and show the abrupt loss of characteristics typical of the
trunk vertebrae, and the concurrent addition of structures peculiar
to typical caudal vertebrae. Since these two vertebrae are unlike
each other, differ from other vertebrae of the body, and are in a
region considered to be a part of the trunk, it is suggested that they
be designated precaudal vertebrae.

The first precaudal vertebra has the parapophyses and diapo-
physes considerably reduced and fused. The neural arch is low
and no haemal arch or spine is present. The second precaudal
vertebra has no parapophyses or diapophyses which are recognizable
as such, but bears a winglike lateral projection near the base of each
prezygapophysis. An incomplete haemal arch is present on the
ventral surface of the centrum, and from this a strong, deeply bi-
furcate haemal spine projects caudad and ventrad to lie ventral to
the anterior part of the centrum of the first caudal vertebra. The
haemal spine of the second precaudal vertebra is notably longer,
heavier and directed more caudally than the haemal spines of the
caudal vertebrae. The pair of projections medial to the postzyga-
pophyses are broad and project dorsally.

In general shape and structural relationships the caudal vertebrae

236 The University Science Bulletin

are quite similar one to another, but because of progressive reduction
toward the tip of the tail, the second caudal vertebra is here de-
scribed. The neural arch, centrum and zygapophyses bear the same
general relations as described for the trunk vertebrae, but the neural
spine is low and the pair of projections medial to the postzygapophy-
ses are enlarged and extend dorso-caudally. From the ventro-lateral
surface of the prezygapophyses, a dentoid spine projects laterally,
its distal half extending past the lateral limit of the prezygapophysis.
Attached to the ventral surface of the centrum in its posterior two
thirds is a heavy haemal arch from which a bifurcate haemal spine
projects caudo-ventrally past the level of the posterior limit of the
centrum. Attached to the ventral surface of the haemal arch and to
the antero-ventral surface of the haemal spine, is a thin ventrally-
projecting lamella of bone. This lamella is rather large and pro-
jects anteriorly past the anterior margin of the haemal arch.

The Skeleton of the Girdles and Limbs

In general structure and configuration, the skeletal elements of the
pectoral and pelvic girdles and their appendages have typical sala-
mandroid form. For that reason, they are only partially described
here.

The sternal cartilage is delicate, has approximately the shape of
an equilateral triangle and though attached to the coracoid car-
tilages by fasciae, is not otherwise articulated with them. The
scapulo-coracoids are well-ossified but appended cartilages are
broad and thin. The glenoid fossa is deep and bony.

The elements of the forelimbs are stout and articulated with
heavy cartilages. The phalangeal count is 1.2.3.2; the proximal
phalanges are hourglass-shaped, and the distal phalanges are ter-
minally broadened to assume the shape of a broad-based "T".

The roughly rectangular puboischiac plate is heavy and formed
mostly of two well-ossified ischia joined in the median plane by a
cartilaginous symphysis. The cranial margins of the ischia are
fused with narrow but heavy pubic cartilages, which form the
anterior fourth of the plate. The ilium, short, heavy and largely
ossified, articulates with the puboischiac plate by way of heavy
cartilage. Dorsally the ilium bears a heavy, round, platelike iliac
cartilage, and to the internal surface of this element, the sacral rib
is attached by a cartilaginous enlargement of its distal extremity.
The acetabulum is deep; its medial and medio-dorsal wall is formed
by the ventral part of the ilium, its postero-ventral wall by the
lateral cartilaginous part of the ischium, and its antero-ventral wall

Baird: The Genus Pseudoeurycea 237

by the pubic cartilage. The anterior and posterior walls of the
acetabulum are formed by thin cartilages extending from the ilium
to the pubic cartilage and to the ischium.

The elements of the hindlimb are stout and firmly articulated.
Structure of the bones is typically salamandroid, the phlangeal count
is 1.2.3.3.2, and the phlanges have the same shape as described in the

hand.

Hyobranchial Musculature
superficial layer

M. Intermandibularis Anterior: This muscle, considered to be
a modification of the larval muscle of the same name, is the most
anterior of the superficial muscles in this region. It is situated an-
terior to the cranial extremity of the median raphe, and attaching
laterally to the fasciae of the anterior parts of the two intermandi-
bularis posterior muscles, it extends across the ventral midline with-
out interruption of its fibers. It is thin, narrow and short and al-
though it is easily distinguished from the m. intermandibularis pos-
terior, the m. intermandibularis anterior is difficult to separate along
its posterior margin because of its close attachment to the posterior
muscle.

M. Intermandibularis Posterior: This muscle is composed of
two distinct parts. The more anterior of these originates from the
dorso-medial aspect of the mandible, just posterior and lateral to the
origin of the m. geniohyoideus lateralis and from the fascia of that
muscle. This part extends from its origin to insert on the antero-
lateral margin of the median raphe. The posterior part of the m.
intermandibularis posterior, a sheetlike muscle (as is the anterior
part), originates along a line on the dorso-medial surface of the
mandible. This line of origin begins at a point one half the width
of the posterior part of the muscle caudal to the anterior part, and
extends to a point just anterior to the caudal limit of the mandible.
From its origin the posterior part of the m. intermandibularis pos-
terior extends mediad, fanning slightly before it inserts on the
lateral margin of the median raphe.

M. Quadrato-pectoralis — Driiner ( 1901 ) : The m. quadrato-
pectoralis is the most caudad of the superficial muscles in this re-
gion. It originates, as a narrow heavy band, ventral to the attach-
ment of the hyoquadrate ligament, from the postero-medial surface
of the ventral half of the quadrate, then fans out as it extends craniad
and mediad. The anterior half of the muscle passes dorsal to the
posterior part of the m. intermandibularis posterior and inserts

238 The University Science Bulletin

by a common aponeurosis with the posterior fourth of that muscle.
The posterior half of the m. quadrato-pectoralis inserts chiefly along
the lateral margin of the median raphe, but a few fibers near the
posterior border of the muscle turn caudad to insert on the pectoral
fascia, and at least one tiny slip attaches to the gular fold in com-
mon with the medial margin of the m. gularis.

M. Interhyoideus : This muscle, deep to the m. intermandibularis
posterior and the m. quadrato-pectoralis, originates from the an-
terior surface of the lateral extremity of the ceratohyal. From its
origin the m. interhyoideus extends craniad and mediad, fanning
and thinning. It inserts dorsally on the aponeurosis of the posterior
part of the m. intermandibularis posterior; the insertion extends one
fourth the width of the aponeurosis anterior and posterior to the
midpoint of that structure. The m. interhyoideus twists 180 degrees
in passing from its origin to its insertion; the fibers originating most
cranially insert caudally on the aponeurosis.

M. Gularis — Smith (1920): The m. gularis originates from the
fascia cephalo-dorsalis on the dorsolateral aspect of the head. Its
line of origin coincides with the line traced by the postocular groove,
and extends from the angle of the mouth, to the ventral limit of the
nuchal groove. From its origin the m. gularis curves obliquely
caudad to insert on the skin of the gular fold from just lateral to the
mid-ventral line to a point near the dorsal extremity of the fold.

second layer

M. Geniohyoideus Medialis — Smith (1920): This muscle orig-
inates from the ventral surface of the dentary just lateral to the
mandibular symphysis. The muscle extends caudad from that point
showing little change in its form throughout its first half, but pos-
terior to that, it spreads and becomes thinner (particularly toward
the midline ) . The insertion is made by means of fleshy slips, chiefly
on the myocommata at the superficial anterior boundary of the m.
rectus cervicis. The thin medial part of the m. geniohyoideus
medialis slightly overlaps the midline and the same part of the m.
geniohyoideus medialis of the opposite side; these medial parts of
the two muscles insert by a thin common aponeurosis. The lateral
fourth of the muscle inserts by fleshy slips on the fascia of the m.
subarcualis rectus I.

M. Geniohyoideus Lateralis — Smith (1920): This muscle orig-
inates from the ventral aspect of the dentary lateral to the mandibu-
lar symphysis and posterior to the origin of the m. geniohyoideus
medialis. The origins of the muscle last named and of the m.

Baird: The Genus Pseudoeurycea 239

geniohyoideus lateralis are so close together that, upon first impres-
sion, the two muscles seem to have a common origin. Laterally the
origin of the m. geniohyoideus lateralis extends along the postero-
ventral margin of the dentary to a point slightly anterior and
medial to the origin of the anterior part of the m. intermandibularis
posterior.

The m. geniohyoideus lateralis extends caudad from its origin,
fanning slightly laterally. The medial part of the muscle lies dorsal
to the m. geniohyoideus medialis and passes dorsal to the ceratohyal
to insert on the fascia of the floor of the mouth; the lateral part of
the m. geniohyoideus lateralis inserts by fleshy slips on the antero-
ventral margin of the ceratohyal, lateral to its midpoint.

M. Rectus Cervicis: This muscle is a direct continuation of the
m. rectus abdominis; its origin has been described above. From its
origin, the m. rectus cervicis extends craniad as a broad sheet cover-
ing the ventral surface of the throat. Immediately posterior to the
first spiral of the m. abdominohyoideus, the m. rectus cervicis sends
a rounded slip dorso-cranially to insert on the mucous membrane of
the floor of the mouth, dorsal to the basibranchial. The major part
of the m. rectus cervicis crosses the proximal tip of the epibranchial
and the ceratobranchial, then turns dorsad to insert by fleshy slips
on the postero-ventral surface of the ceratohyal from just caudal to
its anterior tip to a point at the level of the posterior limit of the
basibranchial. In this course, the m. rectus cervicis crosses five
myocommata, the anterior two of which are well marked.

A part of the m. rectus cervicis, probably comparable to the m.
rectus cervicis superficialis described by Francis (1934:68), orig-
inates from the dorsal surface of the sternal cartilage near its antero-
lateral margin. From that point the slip passes obliquelv laterad
and craniad, ventral to the main mass of the m. rectus cervicis, until
at the third myocomma crossing the muscle, the fibers of the two
become inseparably intermingled.

deeper muscles

M. Subarcualis Rectus I: This muscle originates by a series of
discrete slips from the postero-ventral margin of the ceratohyal, just
medial to its lateral extremity. The most medial of these slips is
immediately lateral to the lateral border of the m. rectus cervicis.
Fibers of the muscle pass caudad from the origin forming a distinct
band on the dorsal surface of the main mass of the muscle and
extending for approximately two thirds its total length. From the
base of the epibranchial cartilage however, fibers depart from the

240 The University Science Bulletin

longitudinal band and "with new fibers formed at metamorphosis"
(Smith, 1920:541), form a complicated spiral around the epibran-
ehial. This arrangement results in the formation of a muscular sac
which extends just posterior to the posterior extremity of the epi-
branchial. Contraction of the muscle forces the cartilage from the
sheath, much as water is forced from a rubber bulb, for there is no
insertion of muscle fibers on the cartilage.

M. Abdominohyoideus — Smith ( 1920 ) : This muscle originates
from the entire ventral surface of the ischium, deep to the m. pubo-
ischio-femoralis externus. From its spatulate origin, the m. abdom-
inohyoideus passes craniad leaving the pubo-ischiac plate between
the medial and lateral origins of the m. rectus abdominis, and enters
a sheathlike passage between the mm. obliquus externus (deep por-
tion ) and trans versus abdominis. Within this sheath the m. abdom-
inohyoideus continues craniad just lateral to the m. rectus abdom-
inis, and at the level of the second costal fold, splits into two easily
separable slips. These continue in close association as the m. ab-
dominohyoideus passes deep to the pectoral girdle, but at the level
of the anterior myocomma of the m. rectus cervicis, the smaller, flat,
dorsal slip curves, comes to lie ventral to the main mass of the
muscle, and the two parts diverge. The small flattened slip passes
deep to the m. rectus cervicis and inserts on the ceratohyal in com-
mon with that muscle. The main bundle of the m. abdominohyoi-
deus, round in cross-section, turns mediad, and immediately pos-
terior to the basibranchial is thrown into a spiral. The muscle makes
one and three-fourths turns in the spiral, crosses the proximal half
of the second ceratobranchial ventrally, curves dorso-medially across
the proximal third of the first ceratobranchial dorsally, then comes
to lie dorsolateral and parallel to the basibranchial. In this relation
to the cartilage, this part of the m. abdominohyoideus extends within
the stalk of the tongue to insert on the anterior tip of the basibran-
chial and on connective tissue within the tongue itself.

M. Hyoglossus: This muscle arises from the anterior tip of the
basibranchial cartilage. It is minute and poorly defined, and (so
far as could be demonstrated by the dissection methods employed)
radiates from its origin and inserts on connective tissue in the
tongue.

M. Suprapeduncularis — Smith ( 1920) : This muscle is a thin flat
band of fibers extending between the cranial portions of the two
ceratohyals, immediately posterior to the stalk of the tongue. The
muscle is firmly affixed to the dorsal surface of each ceratohyal near
its antero-lateral margin, and the anterior margin of the muscle

Baird: The Genus Pseudoeurycea 241

serves to bound the posterior lip of the opening through which the
stalk of the tongue protrudes.

Musculature of the Mandible

M. Depressor Mandibulae: This muscle corresponds in a gen-
eral way with the same muscle in other salamanders, but in this
group it is specialized to the extent of having three distinct heads.
Two of these have their origins separated widely enough to bring
about a considerable difference in the angle through which the
muscles act upon the mandible, and the two muscles can be separ-
ated with ease from their origins to their insertions. Furthermore,
each muscle is innervated by a separate twig from the seventh
cranial nerve. Consequently, the m. depressor mandibulae here is
divided into two parts as follows:

M. Depressor Mandibulae Posterior ( new name ) : This tri-
angular, sheetlike muscle lies deep to the m. gularis, and originates
from the fascia cephalo-dorsalis along a line immediately dorsal to
the ventral margin of the m. longissimus capitis of Nishi ( Francis,
1934:93). The fibers are firmly attached to the skin at the nuchal
groove, which is the posterior limit of the origin. The anterior limit
of the origin is immediately posterior to the level of the angle of
the jaw.

The fibers of the m. depressor mandibulae posterior converge as
they extend cranio-ventrad and insert on the angulare, at the pos-
tero-ventral angle of the mandible, by means of a short tendon. The
anterior margin of the muscle is proximate to the posterior margin
of the m. depressor mandibulae anterior throughout the greater
part of its length, but the two are not fused and are quite separable.

M. Depressor Mandibulae Anterior (new name) : This muscle
is composed of superficial and deep heads that are clearly separable
throughout, although supplied by the same twig of the seventh
cranial nerve. The superficial head, large, heavy and flattened, lies
just posterior to the m. levator mandibulae externus and just ventral
to the m. longissimus capitis. The origin of this superficial head is
from the common aponeurosis of the mm. longissimus capitis and
levator mandibulae externus, from the fascia of the latter muscle,
and from the otic-occipital along the ridge of the anterior vertical
semicircular canal. A few fibers on the internal surface of the
muscle may originate from the lateral surface of the squamosal.

Extending ventrad and slightly caudad, the fibers of the super-
ficial head of the m. depressor mandibulae anterior diverge slightly

16—3286

242 The University Science Bulletin

toward the midpoint of the muscle, then converge markedly in pass-
ing to insert by a fleshy slip on the postero-lateral margin of the
dentary, and on the exposed posterior surface of Meckel's cartilage
immediately dorsal to the insertion of the m. depressor mandibulae
posterior.

The deep head of the m. depressor mandibulae anterior originates
from the lateral and postero-lateral surfaces of the quadrate. Con-
verging from a thin sheet to form a rounded bundle, the fibers of the
deep head curve ventro-medially around the postero-lateral sur-
face of the quadrate and insert on the posterior surface of Meckel's
cartilage and the posterior margin of the angulare. The insertion is
fleshy, medial to the insertion of the superficial head and is dorsal
to the insertion of the m. depressor mandibulae posterior.

M. Levator Mandibulae Externus: The dorsal third of this
heavy, roughly rectangular sheet of muscle originates from the fascia
of the superficial portion of the m. levator mandibulae anterior,
along a line crossing transverse to the axis of that muscle at its curva-
ture. The middle third of the m. levator mandibulae externus arises
from a common aponeurosis with the anterior margin of the m.
longissimus capitis, and the ventral third of the muscle arises from
the antero-lateral surface of the squamosal. The m. levator mandi-
bulae externus curves ventrad, its anterior margin bordering the
postero-ventral margin of the eye, and inserts on the mandible. The
anterior fibers of the muscles are closely attached to the skin at the
angle of the mouth, but insert on the lateral margin of the dentary
by means of a broad tendon. Posteriorly the insertion of the m.
levator mandibulae externus is fleshy and is attached to the latero-
dorsal margin of the dentary. Some of the deeper fibers insert in
common with or proximate to the mm. levator mandibulae anterior
(superficial portion) and levator mandibulae posterior.

M. Levator Mandibulae Posterior: This muscle is short,
heavy, broad and somewhat triangular in outline. It lies deep to
the m. levator mandibulae externus and originates from the anterior
and antero-medial surfaces of the quadrate. With its antero-dorsal
margin overlapping the posterior margin of the m. levator mandi-
bulae anterior (superficial portion), the in. levator mandibulae
posterior extends cranio-ventrad to insert, the more anterior fibers
in common with the m. levator mandibulae anterior (superficial
portion), and the posterior fibers on Meckel's cartilage anterior to
the articulation with the quadrate and on the dorsal margin of the
coronoid process.

Baird: The Genus Pseudoeurycea 243

M. Levator Mandibulae Anterior (superficial portion):
This muscle, a heavy, cordlike bundle of fibers, originates from a
short heavy tendon attached to the neural spine of the atlas. From
its origin the muscle extends craniad, becoming heavier as it passes
dorsal to the medial border of the otic region, then curves ventrad
to pass caudad to the orbit. The muscle passes almost directly
ventrad from this curvature, and inserts by means of a short
aponeurosis on the dorsal margin of the angulare anterior to its
posterior limit.

M. Levator Mandibulae Anterior (deep portion): This
muscle is short and heavy. It has a broad origin from the dorso-
lateral surfaces of the frontal and parietal at the postero-dorsal
margin of the orbit, but the fibers converge abruptly as they pass
ventrad. The fibers terminate on a long triangular tendon, which
passes ventrad, deep to the m. levator mandibulae anterior (super-
ficial portion ) , and the articulation of the mandible with the quad-
rate, and inserts on the medio-ventral surface of the angulare di-
rectly ventral to the articulation.

Musculature of the Body Wall

In Pseudoeurycea as in all salamanders, the muscles of the body
showing little specialization preserve the myotomic arrangement
shown in the embryo. The muscles described below are either
composed of a series of myotomes fused to act as a united whole,
or still show their myotomic arrangement by the presence of re-
duced myosepta, here called myocommata.

M. Obliquus Externus (superficial portion): This muscle,
the most superficial layer of the muscles of the body wall, orig-
inates by tendinous slips from the fascia cephalo-dorsalis and the
skin. The muscle extends from the level of the base of the forelimb
to the tenth costal fold. Its fibers pass almost transversely ventrad
(except at the posterior extremity of the muscle), and insert by
means of a thin aponeurosis extending from just ventral to the
lateral margin of the m. rectus abdominis, to the linea alba. As
this aponeurosis passes superficial to the m. rectus abdominis, and
at the linea alba, it is closely attached to the skin.

The most posterior of the series of myotomes forming the m.
obliquus externus passes caudad and ventrad and makes two dis-
tinct insertions. The more dorsal fibers insert upon the fascia of
the groin immediately anterior to tl e base of the hindlimb, whereas
the ventral group of fibers passes dseper to insert in common with
the posterior part of the m. obliquus externus (deep portion).

244 The University Science Bulletin

M. Rectus Abdominis: The m. rectus abdominis is broad, ex-
tending from approximately midway between the sulcus lateralis
and the ventral midline, to the lateral margin of the linea alba; the
muscle originates in three parts. The first part, a thin superficial
layer, arises from a delicate sheet of connective tissue covering the
m. pubo-ischio-femoralis externus, and is approximately three
fourths the total width of the muscle. A second part originates
from the anterior margin of the pubic cartilage just lateral to its
midline. From this narrow origin, fibers fan slightly to mingle with
those of the other two parts, but chiefly serve to form the deeper
part of the muscle lateral to the linea alba. The third part of the
m. rectus abdominis originates from a thin tendon, affixed to the
latero- ventral aspect of the pubic cartilage and also serving for the
insertion of the two posterior myotomes of the m. obliquus externus
(deep portion), and the posterior myotome of the m. obliquus
externus (superficial portion). From this tendon fibers of the m.
rectus abdominis fan slightly and mingle with those of the super-
ficial part of the m. rectus abdominis, but form chiefly the lateral
third of the muscle.

The insertion of the m. rectus abdominis is difficult to ascertain
for there is a more or less distinct break in the continuity of its
fibers at each myocomma. Posterior and ventral to the base of the
forelimb, however, fibers of the superficial part of the muscle inter-
mingle with those of the m. pectoralis. The deeper part of the m.
rectus abdominis immediately lateral to the linea alba, terminates
in part on the dorsal surface of the sternal cartilage and contributes
to the superfical part of the m. rectus cervicis. All of the remain-
ing fibers of the in. rectus abdominis pass dorsal and lateral to the
sternal cartilage and arbitrarily may be said to terminate at the
myocomma near the anterior extremity of the sternal cartilage.
Here these fibers may be said to give rise to the major part of the
m. rectus cervicis.

M. Obliquus Externus (deep portion): This muscle originates
from the lateral processes of the trunk vertebrae, from the ribs and
from the fascia of the sulcus lateralis. The anterior margin of the
muscle is at the level of the anterior margin of the m. obliquus ex-
ternus (superficial portion), and the posterior margin is a distance
equal to the width of two costal folds anterior to the posterior mar-
gin of that muscle. Throughout its length the m. obliquus externus
(deep portion) is composed of myotomes in which the fibers are
directed almost longitudinally; the only notable deviation from this

Baird: The Genus Pseudoexjrycea 245

occurs toward the ventral margin of the muscle where some fibers
are directed slightly ventrad.

The m. obliquus externus (deep portion) has no single distinct
insertion. There is a distinct break in the continuity of its fibers at
each myocomma; from each myotome a few fibers at the ventral
margin pass deep to the lateral margin of the m. rectus abdominis
and mingle with the fibers on the internal surface of that muscle;
and the two posterior myotomes of the muscle form a distinct part,
the fibers of which insert by means of a thin tendon on the latero-
ventral margin of the pubic cartilage,

M. Transversus Ardominis: The anterior margin of this muscle is
at the same level as the anterior margin of the oblique muscle. The
nine myotomes composing the anterior part of the m. transversus
abdominis originate from the ventro-lateral surfaces of the lateral
processes and centra of the trunk vertebrae. The last three or four
myotomes forming the posterior part of the muscle, originate from
the fascia of the sulcus lateralis. The fibers of the muscle extend
ventrad and slightly craniad, terminating in a thin aponeurosis,
which lies ventral to the lateral margin of the m. rectus abdominis.
This aponeurosis, closely attached to the internal surface of the m.
rectus abdominis and to the parietal peritoneum, extends to attach
on the linea alba.

Musculature of the Pectoral Girdle and Forelimb

dorsal muscles

(The muscles of this group are all deep to the m. subarcualis
rectus I, even though specific mention of the presence of this muscle
may not be made in individual descriptions. )

M. Dorso-humeralis : This large, thin, triangular muscle origi-
nates from the fascia cephalo-dorsalis and the skin, from immediately
dorsal to the dorsal margin of the suprascapular cartilage to the
third costal fold. The fibers of the muscle converge markedly as
they extend toward the head of the humerus, the more anterior fibers
passing almost ventrad as they cross the suprascapular cartilage,
while the more posterior fibers are directed cranio-ventrad. Dorsal
to the proximal limit of the upper arm, the fibers of the m. dorso-
humeralis terminate on a narrow flat tendon, which passes deep to
the m. anconaeus humeralis lateralis, just dorsal to its origin, and
inserts on the antero-dorsal aspect of the humerus just distal to its
head.

M. Dorsalis Scapulae: This muscle, its posterior margin deep to
the m. dorso-humeralis and its antero-dorsal margin deep to the m.

246 The University Science Bulletin

cucullaris, originates from the lateral surface of the suprascapular
cartilage. The fibers of the muscle converge as they pass ventrad
from the fan-shaped origin and immediately anterior to the insertion
of the m. dorso-humeralis, they insert on the humerus.

M. Cucullaris: This muscle arises in two parts. The more an-
terior of these parts, designated as the cucullaris major, arises from
the posterior surface of the quadrate immediately dorsal to the at-
tachment of the ceratohyal, from the crista muscularis and from the
operculum. This stout head, round in cross-section, passes caudo-
ventrad to cross the posterior part of the m. cucullaris and inserts
on the anterior surface of the base of the scapula.

The posterior part of the m. cucullaris, designated as the cucullaris
minor, arises from a slip fused with the ventral surface of the m.
longissimus capitis immediately posterior to the skull (this slip is
almost worthy of consideration as a separate head ) , from the fascia
cephalo-dorsalis, and from the antero-dorsal part of the lateral sur-
face of the suprascapular cartilage. From this exceptionally broad
origin, fibers of the cucullaris minor converge on a short flat tendon
deep to the insertion of the cucullaris major, and by means of that
tendon, insert on the lateral margin of the procoracoid cartilage
near its base.

M. Operculars: This muscle originates from the postero-medial
lip of the fenestra vestibulae as a slender round bundle of fibers. It
passes obliquely caudad and laterad from its origin, becoming more
superficial toward its insertion. The muscle passes immediately
ventral to the ventral margin of the dorsal muscle mass, and an-
teriorly, it is deep to the mm. cucullaris ( both parts ) and depressor
mandibulae posterior. Caudal to the posterior border of the muscle
last named, the m. opercularis is covered by only the m. cucullaris
minor. The m. opercularis broadens and thins toward its spatulate
insertion on the antero-dorsal part of the lateral surface of the
suprascapular cartilage.

ventral muscles

M. Pectoralis: This is the most posterior of the ventral pectoral
muscles. It is roughly triangular, sheetlike and is thinner anteriorly
than posteriorly. The posterior third of the muscle may arbitrarily
be said to originate from the myocomma of the third costal groove,
for at this level superficial fibers from the m. rectus abdominis curve
laterad and contribute to the m. pectoralis. The middle third of the
muscle originates from the linea alba, and the anterior third from
the ventral surface of the sternal cartilage near its midline. All fibers

Baird: The Genus Pseudoeurycea 247

from this fan-shaped origin converge on a short strong tendon
which inserts on the posterior surface of the crista ventralis humeri.

M. Supracoracoideus: This fan-shaped muscle originates from
the ventral surface of the coracoid cartilage along its medial border,
and the posterior part of the muscle is overlapped by the m. pec-
toralis. The fibers of the m. supracoracoideus converge toward the
proximal extremity of the humerus and insert on the crista ventralis
humeri in common with the m. pectoralis.

M. Coraco-radialis Proprius: This muscle situated just deep to
the m. supracoracoideus, originates on the ventral surface of the
coracoid cartilage. The head of the muscle is thin and fan-shaped,
and its line of origin is continuous with that of the m. coracobrachia-
lis brevis. The fibers of the m. coraco-radialis proprius converge on
a flat tendon which divides just medial to the shoulder joint. The
lateral slip of this tendon inserts on the crista ventralis humeri, but
the medial part of the tendon is long and enters the upper arm, pass-
ing between the mm. coracobrachialis longus and humero-antibra-
chialis to insert on the medial aspect of the radius just beyond its
proximal limit.

M. Procoracohumeralis : This muscle originates in two parts.
The superficial head, thin and spatulate, arises from the ventral sur-
face of the procoracoid cartilage just caudal to its anterior limit and
passes caudad to insert on the anterior margin of the crista ventralis
humeri. The deep part of the m. procoraco-humeralis is quite dis-
tinct from the part above. It originates from the ventral surface of
the procoracoid cartilage near its midpoint, and passing deep to the
superficial head, extends caudad to insert on the capsule of the
shoulder joint and the proximal margin of the crista ventralis humeri.

M. Coracobrachialis Longus et Brevis: The long head (caput
longum) of this muscle arises from the posterior part of the dorsal
surface of the coracoid cartilage. The heavy bundle of fibers ex-
tends to insert along the middle third of the posterior surface of
the humerus.

The short head (caput brevis) arises from the posterior third of
the ventral surface of the coracoid cartilage and extends laterad
over the shoulder joint to insert along the base of the posterior sur-
face of the crista ventralis humeri.

The two heads are closely fused throughout, and are separable
only with difficulty.

M. Subscapulars: This muscle originates on the internal sur-
faces of the scapulo-coracoid and two of its cartilages. The origin
is fan-shaped and the thin sheet of fibers is attached to the ventral

248 The University Science Bulletin

part of the suprascapular cartilage, the scapula, the procoracoid
cartilage and the coracoid. One slip of the muscle also originates
from the posterior margin of the scapula. From the origin, fibers of
the muscle converge to curve laterally at the posterior margin of the
scapula and insert on the crista dorsalis humeri.

Musculature of the Pelvis and Thigh
ventral and lateral muscles

M. Pubo-ischio-tibialis: This muscle is distinctly divided into
proximal and distal parts, continuous by way of a tendinous aponeu-
rosis. The proximal part, the most superficial of the muscles on the
ventral surface of the pelvic girdle, originates as a thin sheet from
the pubo-ischiac plate immediately lateral to its midventral crest.
The anterior margin of this part lies caudal to the anterior margin
of the m. pubo-ischio-femoralis externus and the caudal margin is
inseparably fused with the anterior margin of the m. ischio-flexorius.
The proximal part of the muscle narrows slightly as it passes laterally
over the pubo-ischiac plate to terminate in an aponeurosis at the
level of the acetabulum. The anterior part of this aponeurosis at-
taches to the origin of the m. pubo-tibialis, forming a postero-medial
support for that muscle, while the posterior part of the aponeurosis
extends but a short distance before becoming continuous with the
distal part of the m. pubo-ischio-tibialis. The distal part of the
muscle, tapering slightly, extends distad in a midventral position in
the thigh and inserts on the proximal half of the postero-ventral sur-
face of the tibia.

M. Ischio-flexorius: This muscle too is composed of proximal
and distal parts. The proximal part, almost inseparably fused with
the posterior margin of the proximal part of the m. pubo-ischio-
tibialis, lies superficially upon the posterior part of the pubo-ischiac
plate and originates from the postero-lateral margin of that element.
The fibers of this part converge as they pass laterad and terminate
in a tendon just posterior and lateral to the aponeurosis dividing the
m. pubo-ischio-tibialis, and immediately lateral to the aponeurosis
of the m. caudali-pubo-ischio-tibialis as it crosses to insert upon the
m. pubo-ischio-tibialis. The slender, spindle-shaped distal part of
the m. ischio-flexorius extends from the distal extremity of this
tendon and passes along the femur posterior to the m. pubo-ischio-
tibialis, to insert by a delicate aponeurosis on the fascia of the flexor
surface of the m. flexor primordialis communis.

M. Pubo-tibialis: This muscle originates by a tendinous aponeu-
rosis from the antero-lateral margin of the pubis immediately cranial

Baird: The Genlts Pseudoetjrycea 249

to the anterior margin of the m. pnbo-ischio-femoralis-externns. The
postero-medial aspect of the origin is strengthened by the attachment
of a portion of the aponeurosis of the proximal part of the m. pubo-
ischio-tibialis. From its origin the m. pubo-tibialis passes distad
along the femur, just anterior to the distal part of the m. pubo-ischio-
tibialis, and inserts on the proximal fourth of the antero-ventral sur-
face of the tibia.

M. Ischio-caudalis: In the female this short, heavy muscle lies
immediately lateral to the cloaca. It originates from the haemal
spine of the second precaudal vertebra and from the myoseptum at
the junction of the trunk and tail, and passes straight forward to
insert on the posterior margin of the ischium just lateral to the ven-
tral midline. Some of the fibers in contact with the lateral wall of
the cloaca are firmly attached to it, but there is no break to indicate
a distinct insertion thereto.

In the male the ischio-caudalis originates wholly from the ventral
margin of the myoseptum mentioned above. The muscle is thin
and ribbonlike, and from its origin, following the curve of the cloacal
lip, it passes between the dorsal and ventral parts of the cloacal
gland. The insertion is like that in the female.

M. Caudali-pubo-ischio-tibialis: In the female this muscle lies
immediately lateral to the m. ischio-caudalis. Its origin is from the
same parts as the ischio-caudalis, but slightly dorsal to the origin of
that muscle. The caudali-pubo-ischio-tibialis is stout and straplike
and passes antero-laterally to insert by means of a delicate aponeu-
rosis upon the postero-lateral quarter of the fused proximal parts of
the m. pubo-ischio-tibialis and m. ischio-flexorius.

In the male the muscle is thin and ribbonlike, and only a small
slip from the dorsal margin originates from the haemal arch; the
remainder of the muscle originates from the myoseptum. The body
of the muscle is more superficial and further separated from the m.
ischio-caudalis than it is in the female, due to its passage over the
lateral surface of the dorsal part of the cloacal gland in this sex.

M. Caudali-femoralis: This muscle, situated just dorsal to the
m. caudali-pubo-ischio-tibialis, originates in common with, and im-
mediately dorsal to, the two preceding. In the female it extends
antero-laterally, increasing slightly in diameter near its mid-point,
then tapers toward its insertion. The latter is made by a narrow ten-
don which passes dorsal to the posterior margin of the distal part of
the m. ischio-flexorius, and inserts on the external trochanter of the
femur.

250 The University Science Bulletin

In the male the origin of the m. caudali-femoralis is similar to that
in the female, but the muscle is not as stout as in the other sex, and
curves about the lateral aspect of the dorsal part of the cloacal gland
in passing to its insertion.

M. Ilio-caudalis : This muscle is situated between the dorsal
margin of the m. caudali-femoralis and the ventral margin of the m.
dorsalis trunci. It originates chiefly from the myoseptum at the
junction of the trunk and tail, though some of the deeper fibers arise
from the lateral processes of the first and second caudal vertebrae.
As a short thick bundle of fibers, the ilio-caudalis passes craniad to
insert on the internal surface of the dorsal extremity of the ilium
near its posterior border. The fibers of the muscle are interrupted
by one myocomma in each sex, and the muscle is proportionally
smaller in the male than in the female.

M. Pubo-ischio-femoralis Externus: This muscle, except for its
anterior fourth, is covered by the fused proximal portions of the m.
pubo-ischio-tibialis and m. ischio-nexorius. It originates from the
pubo-ischiac plate along a line extending the full length of that
element, immediately lateral to its mid-ventral crest. The m. pubo-
ischio-femoralis externus, a broad thick sheet at its origin, narrows
considerably as it passes laterad, crossing ventral to the origin of the
m. abdominohyoideus. Deep to the juncture of the proximal and
distal part of the m. pubo-ischio-tibialis, the m. pubo-ischio-fem-
oralis externus turns abruptly dorsad to insert directly upon the
trochanter.

M. Pubi-femoralis: This muscle is short and heavy, and origi-
nates in two parts. The origin of the ventral head is from the dorsal
aspect of the aponeurosis of the m. pubo-ischio-tibialis where it is
attached to the origin of the m. pubo-tibialis. The dorsal head
originates on the pubic cartilage from the process immediately an-
terior to the acetabulum. Fibers from the two heads unite and pass
caudo-distad, crossing the posterior extremity of the insertion of the
m. pubo-ischio-femoralis internus, and insert on the middle third of
the antero-ventral surface of the femur.

M. Ischio-femoralis : This short stout muscle originates from the
lateral margin and postero-dorsal surface of the ischium posterior to
the acetabulum. It passes cranio-laterad to insert on the posterior
surface of the head of the femur.

dorsal muscles

M. Iliofibularis : This long slender muscle originates from a
short heavy tendon attached to the postero-lateral surface of the

Baird: The Genus Pseudoeurycea 251

ilium immediately ventral to the union of that bone with the iliac
cartilage. From its origin the m. iliofibularis extends distad in the
thigh, crossing the posterior border of the femur, and passes ventro-
lateral to the knee capsule. The muscle inserts on the postero-
ventral margin of the fibula, just distal to the knee.

M. Extensor Iliotibialis : This muscle is composed of two parts,
ovoid in cross-section and separate throughout the greater part of
their extent. Both parts are slender. The posterior head originates
in common with the m. iliofibularis; the anterior head originates at
the same level from the antero-lateral surface of the ilium. The
two parts immediately approximate along the middorsal surface of
the femur, but show no sign of fusion except just proximal to the
tendon of insertion. This tendon, broad at its attachment to the
muscle, passes distad on the dorsal surface of the knee capsule (to
which it shows no attachment) and enters the shank between the
mm. extensor digitorum communis and extensor cruris tibialis.
Much narrower at its distal extremity, the tendon inserts on the
anterior spine of the tibia.

M. Pubo-ischio-femoralis Internus: This muscle originates
as a broad sheet from just lateral to the middorsal line of the pubic
cartilage for its full length, and from the antero-medial part of the
dorsal surface of the ischium. The fibers of the muscle converge
as they pass laterad from their origin. At the angle formed by the
union of the ilium with the pubic cartilage, the m. pubo-ischio-
femoralis internus twists; the posterior part of the muscle curves
about the base of the ilium immediately cranial to the in. extensor
iliotibialis, and the posterior part of the muscle curves ventrad over
the antero-lateral margin of the pubic cartilage. The m. pubo-
ischio-femoralis internus covers the anterior surface of the head
of the femur and inserts on an elongate area bounding that structure
ventrally and distally. The longest fibers of the muscle insert on
the dorsal surface of the femur and extend no farther distad than
the mid-point of that bone.

A few fibers from the antero-dorsal surface of the m. pubo-
ischio-femoralis internus insert on a long delicate tendon which
extends distad to its insertion on the antero-dorsal surface of the
knee capsule. This part of the muscle probably represents the
m. pubo-extensorius of Low (1926) and Francis (1934).

M. Iliofemoralis: This muscle originates as a thin fan-shaped
sheet from the postero-medial surface of the ilium and the antero-
lateral part of the dorsal surface of the ischium. As the fibers ex-
tend postero-dorsally from their origin, they converge into a round

252 The University Science Bulletin

bundle at the posterior angle formed by the junction of the ilium
with the ischium. At this point the m. iliofemoralis turns abruptly
ventrad over the margin of the ischium anterior to the m. ischio-
femoralis and posterior to the origin of the m. iliofibularis, and then
extends ventrad and distad on the posterior surface of the femur to
insert on the external trochanter in common with the m. caudali-
femoralis.

COMPARISON OF OTHER SPECIES WITH PSEUDOEURYCEA

BELLII

In making the following comparisons with P. bellii, I have ex-
cluded differences that I consider the probable result of age, sea-
sonal modification or individual variation. Secondary sexual modi-
fications have been noted and allowance made for them.

External Anatomy: The general coloration of gigantea* closely
resembles that of bellii, but no maculations are present in the
parietal region. In cochranae, robertsi and gobeli there are ground
colors of gray or tannish gray and some degree of broad orange or
reddish striping. Irregular, light, dorsal and dorso-lateral flecking
on a slate or slate-black ground color is characteristic of leprosa
and cephalica; rex is an almost solid grayish tan, and smithi and
gadovii are grayish tan dorsally but markedly lighter ventral to
the sulcus lateralis.

The relative width of the head, as shown by the relation of the
width of the eyelid to the interorbital width, is reduced from the
condition in bellii in all species examined. In bellii the width of
the eyelid is contained approximately 1.6 times in the interorbital
width; in rex, 1.5; in leprosa, 1.4; in robertsi, 1.2; in cochranae,
cephalica and goebeli, 1.0; in smithi, .93; in gadovii, .86; and un-
guidentis, .84.

The postocular groove is straight in cephalica, touches the gular
fold ventral to its juncture with the postocular groove in gadovii,
and in smithi, its anterior extremity touches the angle of the eye.
The first gular grooves can be traced completely across the ventral
surface of the throat in all species except smithi, in which they are
indistinct and cannot be traced beyond the angle of the jaw. These
grooves are not discernible above the postocular groove in smithi
or in gadovii. The gular fold is deep in cephalica, reduced in
gigantea, and is barely marked by folding of the skin in leprosa.

The costal grooves are well-marked and meet ventrally in
gadovii, smithi and unguidentis; they extend almost to the mid-

* Color names are general and do not refer to a specific standard.

Baird: The Genus Pseudoeurycea 253

ventral line in cephalica, but are faint and are not visible on the
venter in rex, goebeli or leprosa. In gadovii, the costal grooves ex-
tend and fork slightly above the sulcus lateralis, and in smithi and
unguidentis, they are continued above the sulcus by Z-shaped
grooves, which extend to the middorsal line.

A distance equal to the width of three to four costal folds sep-
arates the digits of the adpressed limbs in smithi and rex; in
leprosa, robertsi and cochranae the distance is equal to the width
of two to three costal folds; and in gadovii and unguidentis, the
distance varies from equal to the width of one to two costal folds
to overlapping of the digits. The distance separating the digits
of the adpressed limbs in cephalica is constant and equal to the
width of one to two costal folds. The webbing of the digits in-
volves the greater part of the proximal phalanges, and may be ex-
tended to the basal part of the second phalanges in cephalica.
In unguidentis, gadovii and smithi, the webbing at the base of the
proximal phalanges is slightly less evident than in bellii.

The tail is as long as, or slightly shorter than, the distance from
the snout to the posterior margin of the anus in smithi, gadovii
and unguidentis, but in the other species examined, the tail is only
slightly longer than the length of the trunk alone. The constriction
at the base of the tail is not well marked in leprosa, goebeli or rex.

The ridge of the m. subarcualis rectus I extends to touch the third
costal groove in smithi, gadovii and unguidentis; in leprosa,
robertsi, cochranae, rex and goebeli, the ridge extends to midway
between the second and third costal grooves, and in cephalica its
posterior extremity touches or falls just short of the first costal
groove.

The postiliac gland is faint in smithi, and in smithi, gadovii and
unguidentis, there is marked enlargement of the parietal region of
the head, more evident in males than in females.

Buccal Cavity: The palatine surface of the mouth differs mark-
edly in smithi, for midway between the openings of the internal
nares, there is a rounded opening approximately as large as the
narial opening. This marks the termination of the duct of a well-
developed internasal gland located in the fronto-premaxillary fon-
tanelle. In gadovii and unguidentis, the opening is present but
minute, and is surrounded by a narrow area where the mucous is
unusually porous. In other species examined, only the porosity in
the mucous membrane can be discerned between the nares; the
internasal gland is more highly developed in all of these than in
bellii, but has no distinct duct.

254 The University Science Bulletin

The internal nares are round in smithi and cephalica; in the latter
the narial openings are unusually large and are situated anterior
to the lateral extremities of the prevomerine tooth-rows. The
latichoanal fissure is deep in cephalica and extends past the poster-
ior limit of the maxilla to terminate near the posterior limit of the
pharynx. In nnguidentis, gadovii and smithi, the fissure originates
from the postero-lateral margin of the narial opening.

Only six to eight premaxillary teeth are present in females of
leprosa, robertsi, cochranae, rex, goebeli, gadovii, smithi and un-
guidentis; in males of the three species last named, usually only three
enlarged premaxillary teeth penetrate the lip. There is probably
some variation in the number of enlarged teeth in males of the other
species, for in the specimens examined, from two to five were noted,
although the usual number was four. The premaxillary teeth in
males of gadovii show a marked enlargement of the subterminal
point and from the lateral aspect, resemble an inverted "Y" in shape.
In males of smithi and unguidentis, these teeth show even more
modification, for the subterminal point curves caudo-ventrally and
comes to lie antero-ventral to the terminal point.

The maxillary and mandibular teeth are noticeably larger in
gadovii, smithi and unguidentis and the number of these teeth is
considerably reduced. Only 22 to 24 mandibular teeth are present
in each, and the maxillary teeth number from 18 to 24. Excepting
the teeth of the premaxilla, the teeth have no discernible subterminal
points in these species.

In leprosa, goebeli and rex, 11 to 13 teeth are present in the
slightly curved prevomerine series; in cochranae and robertsi the
prevomerine rows are nearly straight and carry 9 to 11 teeth. The
prevomerine rows are broadly curved in gadovii, unguidentis and
smithi; 10 or 11 prevomerine teeth are present in the two species
first named, and 12 to 14 are present in smithi. In cephalica, curva-
ture of the prevomerine rows is barely discernable, and the 13 to
15 teeth are arranged in short discontinuous series. In cephalica, all
teeth are tipped with black and the tongue is attached at its anterior
extremity.

The rows of teeth on the paravomers are arranged to form distinct
cranially-directed chevrons in gadovii, smithi and unguidentis.

Skull: The proportion of the over-all length of the skull to its
greatest width ( measured at the points of articulation of the quad-
rates), is increased over the condition in bellii in all species ex-
amined. From 1.5 in bellii, the proportion rises to 1.8 in leprosa and
ceplialica, 2.0 in gadovii, 2.1 in smithi and 2.25 in unguidentis.

Baird: The Genus Pseudoevrycea 255

In leprosa the frontal processes of the premaxilla are round and
slender anteriorly, flattened at their posterior extremities and are
parallel to each other. These processes articulate only loosely with
the nasals, posterior parts of the prefrontals and anterior tips of the
frontals. In cephalica these processes are slightly heavier, diverge
mildly toward their posterior extremities, and are more firmly at-
tached to the frontal. The frontal processes of the premaxilla in
unguidentis are similar in shape to those of cephalica, but posteri-
orly, one half their length overlaps and articulates with the frontal.
In gadovii these processes are slightly compressed laterally at their
posterior extremities, which articulate with the frontal and form a
slight lateral wall for the posterior part of the fronto-premaxillary
fontanelle. In smithi the frontal processes of the premaxilla are
heavy and laterally compressed throughout their length. Posteriorly
thev articulate with ventrally-curved processes of the frontals, and
together with those parts, form lateral walls for the fronto-premaxil-
lary fontanelle. In unguidentis, gadovii and smithi, the premaxilla
is firmly articulated with the maxilla.

There is a progressive increase in the degree of ossification of the
prefrontals and nasals, and a concurrent increase in the relative
proportion of the size of the former to the size of the latter in
cephalica, unguidentis, gadovii and smithi. The ridge of the pos-
terior vertical semicircular canal is prominent in ungindentis, and
the medial extremity of the anterior ridge bears a small but distinct
crest. The posterior vertical semicircular canal is crested through-
out its length in gadovii, and in smithi the crest is high, extends
the full length of both the anterior and posterior ridges, and is
medially continuous between the two. In gadovii and smithi the
posterior margin of the parasphenoid is angular rather than curved,
and the articulation with the ventral surfaces of the otic-occipitals
is markedly stronger.

A small oval opening is present between the medial margins of
the prevomers in leprosa and ceplialica. This opening is progres-
sively increased in size, and the cartilage forming the articulation
between the prevomers is progressively reduced in unguidentis.
gadovii and smithi. This intervomerine foramen lies posterior to
the fronto-premaxillary fontanelle in leprosa and ceplwlica, but is
ventral to the posterior part of the fontanelle in unguidentis and
gadovii. In smithi the intervomerine foramen lies directly beneath
the fronto-premaxillary fontanelle.

The coronoid process of the angulare is markedly longer and
higher in smithi, gadovii and unguidentis, and in these species the

256 The University Science Bulletin

articulating part of Meckel's cartilage shows slight calcification, but
the articulare does not appear as a separate element.

Other Skeletal Parts : The length of the hyobranchial apparatus
in relation to the length of the head and trunk is slightly reduced
from the condition in bellii in all species examined. Less reduction
is shown in this respect in smithi, gadovii and unguidentis than in
leprosa and other members of its species group. In cephalica, there
is not only reduction in the relative length of the hyobranchial
apparatus, but the relationships between the lengths of its com-
ponent parts are changed as well. The ceratobranchials are mark-
edly shorter than the basibranchial and the epibranchials are less
than twice as long as the basibranchial in that species.

In smithi, gadovii and unguidentis the neural spine of the cervical
vertebra is markedly more ossified, and in these three forms the
neural spines of the trunk vertebrae are markedly lower than in
bellii. Neural spines are present only on the first three trunk verte-
brae in cephalica, and in leprosa, neural spines are completely ab-
sent. Only in cephalica are there cartilages similar to those replac-
ing the last pair of ribs in bellii; in all other species examined, the
last pair of ribs is completely absent and the lateral processes have
no evidence of articulating surfaces. The first two or three pairs of
ribs are broadened distally but not forked, in leprosa, cephalica,
gadovii, smithi and unguidentis.

In gadovii and smithi the ischia are relatively larger, and are
notched posteriorly, leaving a heavy posteriorly-projecting process
behind the acetabulum. In gadovii the synchondrosis between the
ischia is narrow, and in smithi the ischia are nearly in contact in
the median plane.

Hyorranchial Musculature: The m. intermandibularis anterior
is fused to the m. intermandibularis posterior only toward its lat-
eral extremities in leprosa, cochranae and robertsi; in gadovii the
muscle has distinct attachment to the dentary rather than to the
fascia; and in smithi and unguidentis, the muscle is reduced or
absent. The m. geniohyoideus lateralis is attached dorsally to the
floor of the mouth but has no insertion in that area in smithi, gadovii
or unguidentis. The m. abdominohyoideus differs markedly in
cephalica; in that species, the muscle is reduced and no spiral coil-
ing is present at the posterior limit of the basibranchial. The m.
suprapeduncularis is also reduced in cephalica.

Mandirular Musculature: The m. depressor mandibulae poste-
rior is markedly broader at its origin in leprosa, robertsi and coch-
ranae; in these forms the posterior part of the muscle lies dorsal to the

Baird: The Genus Pseudoeurycea 257

anterior margin of the suprascapular cartilage. In smithi and
unguidentis the muscle is slightly enlarged and originates closer
to the m. depressor mandibulae anterior. The latter is also en-
larged in smithi and originates partly from the lateral extremity
of the crista muscularis. The mm. levator mandibulae externus is
thin in cephalica but slightly wider than in bellii. In unguidentis,
all of the mandibular levators show some enlargement. The m.
levator mandibulae externus is heavier, and the levator mandibulae
anterior (deep portion) originates slightly anterior to the point
of origin noted in bellii. The m. levator mandibulae externus,
levator mandibulae posterior, and levator mandibulae anterior
(superficial portion) fuse slightly near their insertions. In gadovii
the enlargement of these muscles is still more marked, the m. levator
mandibulae anterior (deep portion) originates from the anterior
half of the dorsal margin of the orbit, the insertions of the muscles
are made along the entire posterior half of the mandible, the fusion
of the three muscles extends as far dorsally as one third the length
of the m. levator mandibulae externus. In smithi this group of
muscles reaches its greatest development. All muscles of the
group are greatly enlarged, originate and insert as in gadovii, and
are fused inseparably beneath the m. levator mandibulae externus.
In gadovii and smithi, the development of mandibular levators
is noticeably greater in males than in females.

Musculature of the Girdles and Limbs: The m. opercularis is
slightly reduced in leprosa, robertsi, cochranae and cephalica; in
unguidentis, gadovii and smithi the muscle is reduced, partly fused
to the m. dorsalis trunci, and originates more medially on the otic-
occipital. The m. cucullaris is considerably enlarged in gadovii
and smitJii.

In the pelvic girdle and limb, the m. ischio-caudalis is enlarged
and originates wholly from the myoseptum between the trunk and
tail in cephalica and gadovii. The m. iliocaudalis is partly fused
with the m. dorsalis trunci in unguidentis, and in gadovii and
smithi, the two muscles are inseparably fused. The two heads
of the m. extensor iliotibialis are fused slightly in the distal two
thirds of their length in unguidentis and gadovii, and in smithi
they fuse immediately distal to their origins. The m. pubo-ischio-
femoralis internus is narrower at its origin in both smithi and
gadovii; in the former the origin is from the anterior half of the
puboischiac plate, while in the latter, the origin covers the anterior
two thirds of the plate.

17—3286

258 The University Science Bulletin

SUMMARY AND CONCLUSIONS

The general anatomy of Pseudoeurycea bellii is recorded for the
first time. A concentration of subcutaneous glands postero-dorsal
to the insertion of the hindlimb is designated the postiliac gland;
a cleft, which curves caudo-laterad from the lateral margin of the
opening of the internal naris, is termed the latichoanal fissure; and
it is suggested that the name precaudal vertebrae be used to desig-
nate the transitional vertebrae immediately posterior to the sacral
vertebra. The m. depressor mandibulae is found to be divided
into two muscles, each supplied by a separate twig from the seventh
cranial nerve; these two muscles are designated as the m. depressor
mandibulae posterior and m. depressor mandibulae anterior.

Anatomical comparisons of certain other species (gigantea, rex,
goebeli, cochranac, robertsi, eephalica, gadovii, unguidentis and
smithi) with P. bellii, show unquestionably that all are properly
referred to the same genus. The structural relationships of elements
of the skull are identical; the only differences found in the cranial
structure involve the degree of ossification of the bony parts, the
positional relationships of certain structures and the development
of crests on the otic-occipital. Variations in the vertebrae, in bony
elements of the girdles and limbs, and in the anatomy of soft parts
certainly do not fall outside generic limits as they are now under-
stood.

Within the genus, four lines of development are indicated. It is
inferred that all four lines stem from a common ancestral stock
probably referrable to the genus Pseudoeurycea. The most primi-
tive line, represented by bellii and gigantea, shows no well-marked
sexual dimorphism, has cartilaginous remnants of the last pair of
ribs, and shows little development of the internasal gland and its
foramen between the vomers. Pseudoeurycea leprosa, rex, goebeli,
cochranae and robertsi, fall into a second group which is but slightly
more advanced. No well-marked sexual dimorphism is evidenced
in this group, but the last pair of ribs is completely lost, the elements
of the skull are well ossified, and there is better development of
the internasal gland and its foramen than in bellii or gigantea.

Pseudoeurycea eephalica and its subspecies represent a third line
of development and show more specialization than either of the
two groups mentioned above. A primitive characteristic is the
retention of the cartilaginous remnants of the last pair of ribs;
nevertheless the hyobrancial apparatus is modified, the digits are
more fully webbed than in any other species of the group, and there

Baird: The Genus Pseudoeurycea 259

is more external sexual dimorphism evidenced in this species than
by any other. These three characteristics are considered to be
modern as opposed to primitive. The characteristic arrangement
of the prevomerine teeth into discontinuous short series (re-
sembling patches in some) is further evidence of specialization,
probably the result of some food habit requiring strengthening
of the holding abilities of these teeth, although the actual food
habits of this species are not known.

In Pseudoeurycea unguidentis, gadovii and smithi there is a
progressive increase in the size of the mandibular levators and
secondary sexual modification of these muscles as well. Three
series of progressive structural modifications are clearly correlated
with the enlargement of the mandibular levators. The modifica-
tions are: ( 1 ) increase in the size of the crests on the otic-occipital;
(2) increase in the size of the coronoid process of the angulare;
and (3) increase in the degree of the calcification of the posterior
part of Meckel's cartilage. Also there is a progressive increase
in the degree of development of the subterminal point on pre-
maxillary teeth in males, and the internasal gland is progressively
better developed and reaches its peak in smithi. Fusion of the m.
iliocaudalis to the m. dorsalis trunci in each of these three species
is additional indication of their close relationship.

In final analysis, the genus Pseudoeurycea Taylor comprises a
series of species that form a natural group. Anatomical findings
indicate that there are four species groups within the genus, the
species of each group having undergone a series of modifications
which clearly differentiate them from species of the other three
groups. The group represented by P. cephalica is perhaps the
most specialized externally. The group represented by P. gadovii,
unguidentis and smithi have undoubtedly undergone development
along the line of specialization of the mandibular levators, and the
progressive nature of this development and similarity of sexual
dimorphism in the premaxillary teeth warrants their inclusion in
the same species group. The inclusion of these three species in the
same species group differs from the arrangement proposed by
Taylor (1944:209) in that he placed smithi, separately, in a fifth
species group.

260 The University Science Bulletin

LITERATURE CITED
Druner, L.

1901. Studien zur Anatomic des Zungenhein-, Kiemenbogen- und Kehl-
kopfmuskels der Urodelen. Zool. Jahrb. Abt. f. Anat, vol. 15(1),
pp. 435-622, 11 pis.

Dunn, E. R.

1926. The salamanders of the family Plethodontidae. Smith College
Fiftieth Anniversary Publ., viii - 441 pp., 3 pis., 86 maps.

Francis, E. T. R.

1934. The anatomy of the salamander. Clarendon Press, Oxford, xxxi -
376 pp., 24 pis., 84 figs.
Low, J. W.

1926. Contributions to the development of the pelvic girdle. Proc. Zool.
Soc, London, 1926, pp. 913-29, 3 figs.

Smith, L.

1920. The hyobranchial apparatus of Spelerpes bislineatus. Jour. Morph.,
vol. 33, pp. 527-83, 29 figs.

Taylor, E. H.

1944. The genera of plcthodont salamanders in Mexico, Pt. 1. Univ.
Kansas Sci. Bull., vol. 33, pt. 1, 1944, pp. 189-232, 4 pis., 2 figs.

PLATE XXVI

Dissection of the hyobranchial musculature of Pseudoeurycea bellii. (X 7).

a. mandible

b. m. rectus cervicis (cut)

c. ceratobranchial I

d. basibranchial

e. ceratobranchial II

f. ceratohyal

g. m. abdominohyoideus (major part)
h. m. abdominohyoideus (slip)

i. m. subarcualis rectus I

j. epibranchi:il

Baird: The Gents Psevdoeurycea

261

PLATE XXVI

262 The University Science Bulletin

PLATE XXVII

Figs. 2-14. Disarticulated bones of the skull and the cervical vertebra of

Pseudoeurycea bellii. (X^)-

Fig. 2. Premaxilla — lateral.

Fig. 3. Premaxilla— dorsal.

Fig. 4. Maxilla— lateral.

Fig. 5. Frontal — dorsal.

Fig. 6. Prevomer — ventral.

Fig. 7. Orbito-sphenoid — medial.

Fig. 8. Squamosal — lateral.

Fig. 9. Parietal — dorsal.

Fig. 10. Quadrate — medial.

Fig. 11. Parasphenoid — ventral.

Fig. 12. Atlas— ventral.

Fig. 13. Atlas— dorsal.

Fig. 14. Otic-occipital — dorsal

PLATE XXVII

2.

4.

7.

10.

12,

13

14.

264 The University Science Bulletin

PLATE XXVIII

Figs. 15-19. Lateral views of representative vertebrae of Pseudoeurycea bellii.

(X6).

Fig.. ,15. Tenth trunk vertebra.

Fig. 16. Sacral vertebra.

Fig. 17. First precaudal vertebra.

Fig. 18. Second precaudal vertebra.

Fig. 19. Fourth caudal vertebra.

Baird: The Genus Pseudoei'Rycea

265

PLATE XXVIII

15.

16.

17.

18.

19.

THE UNIVEKSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIV, Pt. I] October 1, 1951 [No. 7

The Estrous Cycle in the Woodrat, Neotoma

floridana

Arthur O. Chapman
Department of Anatomy0

Abstract: This species of woodrat has year round, spontaneous estrous
cycles that are usually four to six days long. Leucocytes are almost always
present in the vaginal smears and epithelium, especially at the caudal end of
the vagina. Cornification does not occur, except at the external orifice which
may contribute cornified cells to the smears at any time of the cycle. There is
usually extensive mucification of the epithelium in the middle and cephalic
regions of the vagina at the time of ovulation, followed by a predominance of
vacuolated cells in the smears. Later stages of the cycle are similar to those
in other rodents.

INTRODUCTION

The vaginal smear has been applied to many mammals since
Stockard and Papanicolaou (17) first used it to indicate the stages
of the estrous cycle of the guinea pig. Outstanding contributions
with this method were also made by Long and Evans ('22) on the
rat and Allen ( '22 ) on the mouse.

The stages in the cycle are designated in most rodents by the
relative numbers of cornified and "pavement" epithelial cells and
by the presence or absence of leucocytes in the smears. The time
of heat, or estrus, is recognized by the presence of only cornified
cells in the smears. Of the wild rodents, a similar vaginal smear
picture was found by Clark ('36a) in the deer mouse; by Clark
('36b) in the cotton rat; by Foster ('34) and Coco ('42) in the
ground squirrel; by Deanesly and Parkes ('33) in the grey squirrel;
and by Parkes ('31) in the Chinese hamster.

Deanesly ('38), Kent and Smith ('45), Sheehan and Bruner
('45), and Ward ('46) have found some differences in the degree
of cornification and time of heat in the golden hamster.

* Now at the Department of Anatomy, University of Nebraska College of Medicine,
Omaha, Nebraska.

( 267 )

268 The University Science Bulletin

Donat ('33) and Wood ('35) made vaginal smears on two sub-
species of the woodrat, Neotoma fuscipes. Leucocytes and epithelial
cells were found to dominate the smears most of the time and
cornified cells appearing only in small numbers. The woodrats
were considered to remain in the diestrous interval, and they con-
cluded that the female does not have a regular cycle in captivity,
or where there is no stimulus from a male.

A more thorough investigation seemed desirable, in order to com-
pare the woodrat with other rodents and learn more about the
changes during the estrous cycle.

MATERIALS AND METHODS

The woodrats used in this study were caught with live traps in
the vicinity of Lawrence, Douglas County, Kansas. They were kept
separated in individual metal cages in a room that was dark most
of the time. Some of them were placed under a small roof outside
during warm weather. One female was kept for five months in a
large outdoor cage that had been constructed especially to simulate
natural conditions. Two woodrat houses of sticks and stones had
been transported intact and placed on the ground of the cage for
her to live in.

Purina Fox Chow and water were ^available to the captive wood-
rats at all times; and supplemented by lettuce, cabbage, carrots,
grass or some other green feed once or twice a week. The diet
seemed to be adequate for good health and growth.

Vaginal smears were made from thirteen adult female woodrats,
some being made during every month of the year except September
and October. The smears were made once daily, except during one
series from woodrats No. 3, No. 6, and No. 8, when they were made
twice a day for a month. Nine adult females were sacrificed and
the reproductive organs removed for sectioning. Eight of them
were killed at known stages of the estrous cycle, after a series of
daily smears had been made. One that was in early pregnancy was
sacrificed the same day she was caught.

The genital tracts were stretched gently and laid on a flat cork,
where they remained adhered and straight while being fixed in
Bouin's solution. Serial sections were made of the ovaries and
oviducts at 10 micra and stained with Harris' hematoxylin and eosin.
Longitudinal sections were made through the lumen of the vagina
for its entire length, including the cervix and a short portion of the
two uterine horns. The contents of the lumen remained in position
and could be seen in the sections. That made it possible to tell

Chapman: The Woodrat Neotoma Floridana 269

where cells were desquamating and mucus was being formed. Sev-
eral sections were made and stained with hematoxylin and eosin, a
rapid trichrome stain (Pollak, '44), and Mayer's muci-carmine as a
test for mucin. Half of the vagina from one woodrat was fixed in
absolute alcohol and stained with Best's carmine for glycogen, and
the other half was fixed with formalin, cut with the freezing micro-
tome and stained with Sudan IV for fat.

A cone-shaped woodrat holder was made of half inch hardware
cloth into which they could be maneuvered head first and held
snugly while taking the vaginal contents. After wetting a medicine
dropper with physiological salt solution, it was inserted about one-
half inch into the vagina and suction applied while withdrawing
slowly. When no sample was obtained on the first try, a drop of
the saline was expressed into the lumen and withdrawn by the
dropper to wash out a sample of the cells. This method has the ad-
vantage of taking most of the contents of the lumen, thus obtaining
a sample from all parts of the vagina and not having much left there
to be included with the next smear.

The sample was then spread on a microscope slide and immersed
for five minutes or more in a solution of half and half 95 per cent
ethyl alcohol and ether to fix the smear. Most of the smears were
stained with the rapid trichrome, as a counter-stain to Harris' hema-
toxylin, and some with hematoxylin and eosin. A few duplicate
smears were fixed with osmic acid fumes and Flemming's solution or
fixed in formalin and stained with Sudan IV to test for fat in the
vacuolated cells.

The types of epithelial cells in the smears are described, based
on Hartman's ( '44 ) classification. Four of the types vary in number
during the cycle and are used to designate the stages. Unless other-
wise indicated, descriptions of epithelial cells in smears and sections
of the vagina are based on the trichrome stain.

Four stages in the estrous cycle are based on vaginal smears and
a description made of the vaginal epithelium of the woodrats killed
during each stage. A summary of each of these animals is given in
Table I. The relative thickness of the vaginal epithelium is indi-
cated by -j- to -j — | — | — \-, and the cell types that make up the
superficial layers of the epithelium are listed for each part of the
vagina. Follicles over 0.8 mm. in diameter seem to be significant
to the cycle in progress and are tabulated. The youngest sets of
corpora lutea are briefly described and the eggs in the oviducts
listed.

270 The University Science Bulletin

The corpora lutea are divided into the following age groups:
(a) those with blue lutein cells and cavities that become progres-
sively smaller; ( b ) those with light blue lutein cells and connective
tissue in the centers; (c) large ones with pale lutein cells and a
few fibrocytes; (d) those with pale lutein cells and many fibro-
cytes; (e) small ones with few lutein cells and mostlv fibrous tissue.
The corpora lutea that are known to be forming from follicles that
have not ovulated appear to develop later in the cycle and are
apparently supplementary to the ones formed from ovulated
follicles.

The writer wishes to .acknowledge the assistance of Drs. Homer
B. Latimer and Paul G. Roofe in reading the manuscript and mak-
ing many suggestions, and Paul N. Wilkinson for making the photo-
micrographs.

CELL TYPES OF THE VAGINAL EPITHELIUM

Hartman ('44) describes the cell types found in vaginal smears
from the rat, using a similar trichrome stain. Most of the types can
be recognized in the woodrat and his system will be used in order
to recognize the similarities and have a point of departure for the
difference between the two rodents. The following description of
the cell types will be based on their appearance in the woodrat,
using the trichrome stain.

Cell types I, II and III are considered to be representative of the
"cornified series," and the rest we shall group in the "oval series."
The cells of the "cornified series" are larger and flatter than the
others. Those of the "oval series" correspond to those of the
"atrophic series" in monkeys and women (de Allende, Shorr and
Hartman '43, Plate I, type I), and the small cells found in the dog
during anestrum (Evans and Cole '31, Fig. 2).

Cell Type I. These are Hartman's "scales" and the cornified cells
of other authors. They are large, red or orange, having no visible
or barely visible nuclei, and often showing wrinkles, folds or lines
through the cytoplasm.

Cornified cells often make up about five or ten percent of the
cells in the smear (Fig. 2a), but seldom more than twenty percent.
No consistant variation in number is recognized in the woodrat and
they are not used as an indicator of any stage in the vaginal cycle.
In the sections they are seen only at the external orifice of the
vagina, where thev are continually being dehisced in layers like
the skin.

Cell Type II. These are Hartman's "shorr cells," which he de-

Chapman: The Woodrat Neotoma Floridana 271

scribes in the rat as being large with "blue or blue-green to gray
cytoplasm, and large vesicular nuclei." A few large cells may be
seen in smears from woodrats at any time of the cycle that fit his
description, but the size of the nuclei grades down to the small dark
nuclei of the more prevalent precornified cells. Both kinds probably
arise from the transition zone between the cornified area at the
external orifice and the vagina proper (Fig. 9).

Cell Type III. These are pre-cornified cells with large baso-
philic granules in the cytoplasm. They apparently come from the
stratum granulosum seen under the cornified layer at the external
orifice and the transition zone between that cornified area and the
vagina proper. These, like Types I and II, may be seen in the
smear at any stage of the cycle.

Cell Type IV. Hartman calls them "mucous cells," but his de-
scription fits the vacuolated cells seen in the smears and lumen of
the woodrat, which are negative for mucin. This type may be sub-
divided into two classes:

Class I. These are mucous cells, which are seen forming a
superficial layer of tall columnar cells along the middle and cephalic-
regions of the vagina during Stages 1 and 2 (Figs. 3 and 7).
Muci-carmine stains them red, as well as the strands of mucus
secreted by them into the lumen. Only fragments of these cells have
been recognized in the lumen and smears, and they appear to be
used up in the process of secretion.

Class 2. These are vacuolated cells, which stain green or blue.
When they first appear during the cycle there are a few small
vacuoles scattered around in the cytoplasm (Fig. 5). By the time
they are plentiful enough to indicate Stage 2, the vacuoles are
large and appear to replace the cytoplasm, compressing the nuclei
to one side or to the center (Fig. 6). They resemble the "foam
cells" Evans and Cole ('31) found in the dog during anestrum.

The vacuoles can not be seen until the cells are sloughed off into
the lumen, when Bouin's fixative and trichrome or hematoxylin and
eosin stains are used on sections of the vagina. For that reason
the cells are not identified in the epithelium, except in the part of
the vagina of Woodrat No. 26 that was fixed in alcohol. Sudan IV
stain, on frozen sections of formalin-fixed tissue from the middle
part of the vagina from that same woodrat, shows many cells deep
to the mucous layer which contain small fat droplets. Muci-
carmine colors the strands of mucus among the cells in the lumen,
but the vacuoles are negative.

272 The University Science Bulletin

In the smears, the vacuoles are prominent when alcohol-ether has
been used as a fixative, and they look like the spaces left when fat
droplets have been dissolved away. For some unknown reason,
however, the vacuoles are negative for fat with osmic acid fixation,
and with Sudan IV on formalin fixed smears.

Cell Type V. These are Hartman's "phagocytized cells," con-
taining polymorphonuclear leucocytes. One or more leucocytes
may occur in vacuoles in any of the cells of the oval series, (Figs.
4b and 12a), but they are usually in the vacuolated cells or the
"dense" cells ( Type VI, Class 1 ) . Sometimes a cell is seen to have
many leucocytes in a large vacuole.

Cell Type VI. These are called "other epithelial cells" by Hart-
man. He divides them into six classes, some of which are important
in the woodrat.

Epithelial Cells, Class 1. Hartman describes them as "dense
cells, often red-purple in color, with full nuclei." The woodrat has
many cells fitting that description at certain stages of the cycle
( Fig. 2d ) . Some are seen to take the red stain in a small part of
the cytoplasm around the nuclei, where it often looks like a dense
flocculate (Fig. 6a); others have more of the cytoplasm stained
red, tending to obscure the nuclei; and others are completely red
and the nuclei not visible. The nuclei can be seen when hema-
toxylin and eosin stain is used, even though the cytoplasm takes a
dense red.

There are other cells that resemble the red staining dense cells,
except that they stain green. They occur in the smears at the same
time of the cycle, and the color difference does not seem to be suffi-
cient reason for placing the green staining ones in a separate class.

Epithelial cells, Class 2. Hartman describes them as degenerating
"deep cells" with misshapen pyknotic nuclei. In the woodrat there
are two kinds that fit in this class: One staining light green (pink
with H and E ) and the other red-orange ( dark red with H & E ) .
In both kinds the nuclei are small, dark, often irregularly shaped
or fragmented, and the cells are small.

At no time during the cycle are they seen to form a definite
layer in the sections of the vagina, but are mixed at random with
other types near the surface of the lumen during Stages 2 and 3
(Figs. 7a and 10a).

One or more of either kinds of the "deep" cells may be seen in
lacunae in the epithelium. Sometimes leucocytes are seen with
them in larger spaces. In the lumen and in the smears many of

Chapman: The Woodrat Neotoma Floridana 273

them appear to be inside other cells, usually surrounded by a
vacuolar space. Some cells are seen to contain two or three of them
(Fig. 8a).

Epithelial cells, Class 3. These are Hartman's "red epithelial
cells". Some smears from woodrats contain many red cells, but
they do not appear consistently during the cycle and are probably
due to variations in stain differentiation.

Epithelial cells, Class 4. These are Hartman's "proestrous cells."
The cells of the woodrat which appear during Stage 1 ( Figs. 2c
and 4a) are among the largest of the "oval series," but are small
compared with the cells of the "cornified series." They stain green
or bluish-green, have large vesicular nuclei, and are not flattened
when seen on the surface of the epithelium during Stage 1 ( Fig. 1 ) .
Later in the cycle they are seen in decreasing numbers in a more
flattened form, and are probably degenerating. The nuclei become
smaller and darker and the cytoplasm more dense. Some of them
appear to be intermediate forms of "dense'cells that stain green,
blue or gray, instead of red and are hard to differentiate.

Epithelial cells, Classes 5 and 6 are not recognized in the woodrat.

STAGES IN THE ESTROUS CYCLE

It would be desirable to use the names proestrum, estrus, met-
estrum and diestrum in designating the stages of the cycle, but it
is hard to make accurate indications of the beginning and end of
each of these standardized stages in the woodrat. They are not
tame enough for the mating response, which can be elicited by
hand in some laboratory rodents (Ball, '37; Blandau, Boling and
Young, '41; Young, '37), nor would they mate while under observa-
tion. Thus, there was no convenient method of telling when physio-
logical estrus occurred.

Four stages are based on the relative numbers of certain cell
types in the vaginal smears. In woodrats sacrificed during each
stage the ovarian condition is summarized and related to the vaginal
sections and smears ( Table I ) . One killed in early pregnancy is
included in the descriptions. These stages can usually be followed
in succession by taking smears once daily, but occasionally one of
the shorter stages may be missed. A continuous series of smears,
taken twice daily at 9 a. m. and 5 p. m. from July 10 to August 12,
1946, on woodrats No. 3, No. 6, and No. 8, are used as a basis for
interpreting subsequent smears and for the following stages:

18—3286

274 The University Science Bulletin

Stage 1

The vaginal smears (Figs. 2, 4, and 5) have "proestrous cells"
(Type VI, Class 4) appearing in large numbers compared with
the scanty smear of Stage 4. They usually constitute 30 to 60 per
cent of those present. Various degenerating cells of Stage 4 are
also found during the first part of the stage and vacuolated cells
(Type IV, Class 2) and "dense" cells (Type VI, Class 1) appear
later. Mucus is usually abundant.

This is the shortest stage, usually lasting 12 to 30 hours. When
smears are taken once a day, the woodrats are seldom found to be
in this stage for two days in a row, and it is sometimes missed.

The vaginal epithelium is thick during Stage 1, especially at the
caudal part ( Fig. 1 ) . That region is lined with green staining "pro-
estrous cells," where they can be seen desquamating into the lumen
in large numbers. Leucocytes, many of which appear distorted, are
distributed among the epithelial cells of that part of the vagina and
can be seen infiltrating into the lumen.

The middle and cephalic parts of the vagina and cervix have
columnar-shaped mucous cells forming the superficial layer of the
epithelium (Fig. 3). The nuclei are dark, pyknotic and basilar in
position and the cytoplasm stains red with muci-carmine.

There is usually much mucus in the lumen of the vagina for its
entire length. With muci-carmine stain, red strands appear to be
extending from the mucous cells toward the external orifice.

At the external orifice, cornified cells (Type I) can be seen des-
quamating in layers. Under the stratum corneum is a distinct
stratum granulosum and a deeper stratum germinativum. These
layers blend into the epithelium of the vagina proper. Near the
point of junction can be seen flat, green cells on the surface, super-
ficial to the cornified layer. This region at and near the orifice is
apparently the source of the cells of the cornified series (Types I,
II and III).

Ovulation occurs during stage 1. Of the three woodrats killed
during this stage (Table I), woodrat No. 7 has three large follicles
which appear to be almost ready to ovulate and woodrats No. 10
and No. 20 had both recently ovulated.

Stage 2

Vaginal smear samples usually contain a large amount of cellular
material. The vacuolated cells (Type IV, Class 2) are predominant
in the smear during this time, making up 30 to 60 percent of those

Chapman: The Woodrat Neotoma Floridana 275

present (Figs. 6 and 8). "Proestrons" cells (Type VI, Class 4) and
"dense" cells (Type VI, Class 1) are also abundant early in the
stage and "deep" cells ( Type VI. Class 2 ) appear later.

The vagina has a medium thick epithelium. The caudal end has
a uniform epithelium of green cells that show a little degeneration
and flattening at the surface. At the middle and cephalic regions it
is lined mostly with "dense" cells (Type VI, Class 1) with many of
the "deep" cells (Type VI, Class 2) distributed among them. The
epithelium of the cervix and that between the folds in some parts of
the cephalic region have tall mucous cells persisting. "Deep" cells
are also seen distributed among and deep to the mucous cells
(Fig. 7).

The cytoplasm of the mucous cells appears to be stringing off with
the mucus into the lumen, as if they are being used up by a process
of holocrine secretion. Muci-carmine stains the cytoplasm and
strands of mucus red. Vacuolated cells in the lumen are negative
for mucin.

Leucocytes appear in relatively large numbers in the lumen, even
though the epithelial cells are abundant. In addition to being in-
filtrated among the epithelial cells, they appear in lacunae along the
surface, except at the cervix. Both green and red staining "deep"
cells are also found in the lacunae and are seen to be located inside
epithelial cells in the lumen, similar to the ones containing leuco-
cytes.

Desquamation is going on in all parts except at the cervix, and is
especially rapid in the middle and parts of the cephalic region.

Two woodrats ( No. 2 and No. 21 ) were sacrificed during Stage 2
(Table I ). Both have new corpora lutea and a corresponding num-
ber of eggs in the terminal portion of the oviducts, indicating that
ovulation had occurred recently.

Stage 3

"Deep" cells that stain both red and green make up 40 to 70 per-
cent of those present in the smears for one or two days of the cycle
( Fig. 12 ) . There are also "dense" cells and vacuolated cells present
in large numbers, but the latter become fewer as the stage pro-
gresses. It is considered to be over when the vaginal sample be-
comes scarce or when the relative number of cells is greatly reduced.

The vaginal epithelium is thin in all areas ( Figs. 10 and 11 ) except
the caudal part, which is of medium thickness (Fig. 9). The latter
region has a layer of flattened cells on the surface. Many leucocytes
are infiltrated through the epithelium and into the lumen.

276 The University Science Bulletin

The middle part of the vagina appears to have lost most of the de-
generating cells from the surface, with just a thin layer of dense
flattened cells along the lumen (Fig. 10). The cephalic region and
cervix are a little thicker and the "deep" cells are still present among
the remaining flattened degenerating cells along the surface, but
not forming a definite layer. A few areas of the cervix (Fig. 11)
have some mucous cells left, but they appear to be mostly gone.
These cells take a light pink color with muci-carmine, but the con-
tents of the lumen are negative for mucin. The cephalic region and
cervix appear to be the only areas where desquamation is abundant

Some leucocytes can be seen among the degenerated cells on the
surface and in lacunae there, but seldom deeper in the epithelium of
the cephalic region and cervix.

Cells containing leucocytes (Type V) are seen among the super-
ficial cells, but some of the larger lacunae open on the surface and
release the leucocytes into the lumen. The "deep" cells also are in
lacunae, alone or with leucocytes. The degenerating epithelial cells
are mostly sloughed off by the time Stage 3 is ended.

Woodrat No. 11 was sacrificed during this stage (Table I). There
are young corpora lutea with cavities and light blue luteal cells that
probably represent the last ovulation, but no eggs can be found in
the oviducts. The corpora lutea that are forming from follicles that
have not ovulated appear to start developing later in the cycle.
Three follicles in one of the ovaries seem to be earmarked for the
next ovulation.

Stage 4

This is the interval of relative quiescence, or diestrum. There is
usually a scanty smear obtained, containing pale, degenerated
cells that are difficult to identify. When mucus is aspirated into
the dropper the cells are usually trapped and obscured by the thick
strands. The stage usually lasts one or two days, but may be much
longer, as occurred in woodrats No. 6 and No. 22 before they were
sacrificed ( Table I ) . The smear does not vary much during a pro-
longed stage, but those two animals may not have a vaginal picture
that is typical of stage 4.

In both of these woodrats there is a thin epithelium lined with
flattened degenerated cells, most of which are hard to classify.
Some are recognized as "dense," "deep" and mucous cells. The
caudal region has a thicker epithelium which is lined with un-
flattened, undegenerated cells. The cervix has a thin epithelium
similar to that of the cephalic and middle regions. Many leucocytes
are infiltrated among the epithelial cells of the caudal part and a

Chapman: The Woodrat Neotoma Floridana 277

few are seen in the layer of degenerating cells along the lumen of
the rest of the vagina. There are not many cells in the lumen.

The few mucous cells remaining in and near the cervix are stained
faintly with muci-carmine, but there is no mucus in the lumen.

Woodrat No. 6 had remained in Stage 4 for five days. The ab-
sence of any large follicles that might soon ovulate would indicate
that it may have continued for a while longer before a new cycle
would begin. Three sets of corpora lutea would seem to indicate
that previous cycles had taken place normally.

Woodrat No. 22 had been in this stage for 21 days before death.
The complete absence of corpora lutea and large follicles are fur-
ther evidence that the cycle was arrested and abnormal. The wood
rat had been losing weight and appeared to be sick.

Early Pregnancy

Woodrat No. 17 was captured March 19, 1947, and sacrificed the
same day, immediately after taking a vaginal smear. There are
three embryos less than 1 mm. long in the left horn and one in the
right horn of the uterus.

The vaginal smear has a mixture of the usual cells seen during
Stage 3 of the estrous cycle; and some green oval cells with pyknotic
nuclei are also abundant. These cells are different from any of the
cell types of the normal cycle, being larger than the "deep" cells
and having small, dark, round nuclei.

The vaginal epithelium is of medium thickness in general, with
considerable variation at different areas. The anterior half, includ-
ing the cervix, is lined with columnar-shaped mucous cells and some
red and green staining "deep" cells mixed in among them. The
mucous cells stain red with muci-carmine. The rest of the epithe-
lium has a more or less flattened layer of "dense" cells, the cells with
round pyknotic nuclei like the ones in the smear, and "deep" cells
distributed among them. There appears to be more desquamation
and leucocytic infiltration in the posterior part. Some mucus and a
few cells are seen in the lumen.

The ovaries are of about the same size as those of non pregnant
woodrats. There are many growing and atretic follicles under 0.8
mm. in diameter and no eggs in the oviducts. One ovary has one
large corpus luteum with pale lutein cells that have fibrocytes be-
tween them. The other ovary has three large corpora lutea similar
to the one in the first.

278 The University Science Bulletin

REPRODUCTIVE FUNCTION UNDER DIFFERENT

CONDITIONS

Since the woodrat does not have a true cornified cell stage, it is
not surprising that Donat ( '33 ) and Wood ( '35 ) failed to recognize
cyclic changes with a few smears. There is a possibility of species
differences, however.

They suggest that the cycle might be different while the animals
are kept in captivity. We have some evidence to indicate that the
woodrats have cycles under natural conditions similar to those in
captivity, especially during the spring and summer. Woodrat No.
13, which was kept in the large outdoor cage that was constructed
to simulate natural conditions, was found to show a smear cycle of
four or five days during March and April, and a cellular sequence
like the woodrats kept in small cages. Woodrat No. 21 was caught
in one of the live traps in the morning, March 30, 1947, and daily
smears were begun that afternoon. She was in Stage 2 that day,
continued through the rest of the cycle in about the usual way, and
reached Stage 1 of the next cycle on the fifth day after being cap-
tured. The next two cycles were both 6 days long, indication that
the one in progress when the woodrat was captured corresponded
very closely to those that followed.

DISCUSSION

Considering proestrum to be the period of growth of the repro-
ductive tissues in preparation for heat, or estrus, cetestrum to be
the degenerating and sloughing off period, and diestrum to be the
period of quiescense; we can correlate the stages of the woodrat to
this scheme to a certain extent.

Physiological estrus probably occurs at the time of ovulation, as
in other rodents, and that includes at least the latter part of Stage 1.
Proestrum is apparently the first part of Stage 1 and is very short,
unless growth starts in the vaginal epithelium late in Stage 4 as in
Woodrat No. 22 ( Table I ) . Physiological estrus may extend into
Stage 2, which corresponds to the cornified cells stage of some
rodents, except that vacuolated cells appear instead. Stages 2 and 3
correspond to metestrum of other rodents, in that the thickness of
the vagina becomes much reduced (Table I). Stage 4 is a period
of quiescence and fits well into the description of diestrum.

The vacuolated cells are not recognized in the epithelium without
special stains and it is difficult to tell where they come from. The
fact that that they are negative for mucin indicates that they, like
the other cell types seen in the smears and lumen, are sloughed off

Chapman: The Woodrat Neotoma Floridana 279

where there is no superficial layer of mucous cells. They have be-
gun to appear in the smears at the time of ovulation, during Stage
1, and the desquamation probably takes place later than the forma-
tion of a particular cell type. That would place the vacuolated cell
formation approximately during ovulation and heat, and corres-
ponds to cornification in most other rodents. Drying of smears
causes the epithelial cells to lose their true appearance (Papanico-
laou, '33 and Hartman, '44), and causes vacuoles to be masked.
Some workers may not have recognized them because of inferior
methods of fixation.

Selle ('22) describes vacuolated cells during his Stage 1 of the
guinea pig cycle, both in the smears and forming a superficial layer
for the whole length of the vagina. He apparently did not use a
specific test for mucin, and it is hard to tell if he had mucous cells or
vacuolated cells, or both. The cornified layer was seen to form deep
to the superficial layer of "vacuolated cells", and the latter would be
removed when the cornified cells sloughed off in layers. He de-
scribes a comparatively sudden end to his Stage 1, as the cornified
cells started sloughing off abruptly. Without a cornified layer form-
ing beneath the superficial mucous layer in the woodrat, the latter
layer persists in the cephalic part of the vagina, and desquamation
does not take place over the entire vagina at once until late in Stage
3. That and the lack of layer formation of desquamating cells,
accounts for the mixture of more than one cell type in smears.

Cole (-30) found cornification in the vestibule, but not in the
vagina proper of the cow. He also describes a superficial layer of
tall, columnar, mucus-secreting cells during proestrum. As the
cycle progressed the layer became reduced in height and the whole
epithelium became thinner in some unknown way, but not by des-
quamation. The superficial layer of mucous cells is not sloughed off
in the woodrat, but appears to be used up in the process of secretion.
Cole (30, Fig. 13) also describes a "vacuolar degeneration" deeper
in the vagina, which resembles the vacuoles seen in the part of the
vagina of Woodrat No. 26 which was fixed in alcohol.

Wilson ('26) also describes a superficial layer in the vagina of
the sow with "vacuolar degeneration", and no cornification. Stains
for fat or mucin may be helpful in correlating the cellular changes
in some of these animals.

Hartman ('44) found the "mucous cells" to occur in the vaginal
smears from the rat during the third day of the cycle, which is two
days after the "scales" dominate the smear. He also found phago-

280 The University Science Bulletin

sytized cells and some deep cells in the smear at that time. They
seem to occur later in the cycle than the time when vacuolated cells
are first found in smears from woodrats. The time relationship
of the woodrat fits that of the castrated rats he injected daily with
0.005 mg. of estradiol dipropionate. Two days after injections were
started there were small green "mucous cells" present in the smear
in large numbers, and on the next day there were only scales, corni-
fied cells and "shorr" cells present. He considers these small green
vacuolated cells to be different from the "mucous cells" that occur in
the natural cycle on the third day. Most of the vacuolated cells in
the smears from woodrats are green, especially the ones appearing
early in the cycle. Mucification and vacuolation occur at about the
same time in the woodrat, and the hormones mediating one process
may also effect the other.

Even though the vacuolated cells in the smears are negative for
fat with both osmic acid and Sudan IV, it seems possible that the
vacuoles contain fat that is not colored by the dyes after being
sloughed off into the lumen. The frozen sections of the vagina from
woodrat No. 26 show only relatively small fat droplets in the epi-
thelium that are colored with Sudan IV. The part fixed in alcohol
shows large vacuoles, which are of about the same size as most of
the ones in the vacuolated cells of smears.

The color difference between the two kinds of "deep" cells is
striking and probably significant, but we do not know why some
are pale green (pink with H & E) and the others are red-orange
( dark red with H & E ) . Both kinds appear in the vaginal epi-
thelium and smears during late Stage 2, become abundant during
Stage 3 and persist in small numbers in the first part of Stage 4.
Since they are distributed among the other cell types, without form-
ing a layer, their development appears to be related to the time of
the cycle more than their position in the epithelium. There seems
to be a "selective degeneration" in the epithelium. We also wonder
how they get inside other epithelial cells, where they are seen fre-
quently in the lumen and smears. Wilson ('26) describes vacuoles
in the vaginal epithelium of the sow which contain "remnants of
degenerating cells" and occasional leucocytes during the first week
of estrus.

The caudal part of the vagina proper, except the external orifice,
has epithelial changes that are different from those of other regions.
The "proestrous" cells are desquamated from that region during
Stage 1 and it is probably the source of vacuolated cells during that
stage. Mucous cells or "deep" cells are not seen there at any

Chapman: The Woodrat Neotoma Floridana 281

time, and the amount of desquamation is much reduced in that area
by Stage 2. Leucocytes may be seen infiltrating through the epi-
thelium at any time of the cycle, and it appears to be the only source
of them during Stages I and II. Late in Stage II and during the rest
of the cycle the leucocytes are also seen infiltrating and in lacunae
in the epithelium of the rest of the vagina proper.

Shorr ('40) considers the trichrome stain to be a very sensitive
indicator of the degree of cornification, by staining the cells pro-
gressively greener, then suddenly red-orange as the estrogen level
is raised. That does not apply to cells of the "oval series" in the
woodrat, and most of the cells of the vagina belong in that group.
The red staining of some of the major tvpes of cells is a common
feature of the degeneration process, yet it is doubtful if it is a
cornification. There are no signs of basophilic cells as precursors
of these cells, as has been described deep to the stratum corneum by
Long and Evans ('22) and Selle ('22), and as seen at the external
orifice of the vagina in the woodrat.

These and other questions concerning changes in the vaginal
epithelium may be answered in the future with specific stains and
histochemical procedures. This animal might also lend itself to
endocrine research, in an effort to learn if hormones cause such
processes as vacuolation and mucification.

SUMMARY AND CONCLUSIONS

Vaginal smears have been made from thirteen adult female wood-
rats. Nine were sacrificed and the microscopic structure of the
vaginas and ovaries described. Eight of them were killed at known
stages of the cycle and one was in early pregnancy.

Cornification does not occur in the vaginal epithelium at any
time during the cycle, except at the external orifice. Longitudinal
sections of the vagina and cervix show regional variations in the
epithelium and contents of the lumen. Leucocytes are always pres-
ent in the smears, and variations in their number are not found
to occur consistently during the cycle.

The cell types in the vaginal smears are described and some of
them are used as indicators of four stages in the estrous cycle, and
correlated with the vaginal epithelium and ovaries of woodrats
sacrificed:

Stage 1 corresponds to the "proestrous" cells stage of other
rodents, but the vaginal epithelium has a superficial layer of colum-
nar mucous cells and ovulation takes place during that period.

Stage 2 has vacuolated cells predominant in the thick smear,

282 The University Science Bulletin

desquamation at the caudal and middle regions of the vagina and
corpora lutea forming from recently ovulated follicles in the ovary.

Stage 3 smear is still thick and contains many cells, mostly small
round ones from deeper in the epithelium and desquamation is
taking place along the whole length of the vagina.

Stage 4 has a scanty smear, containing a few degenerated epi-
thelial cells and varying amounts of mucus. The corpora lutea con-
tinue to grow and the lutein cells appear lighter.

While in captivity the woodrat has year round spontaneous
estrous cycles of three to eight days. Most of them last four to six
days. Vaginal smears during March and April from one female kept
in a large outdoor cage also show regular cycles during those two
months.

The woodrat sacrificed during early pregnancy has a vaginal epi-
thelium similar to Stage three of the estrous cycle.

REFERENCES CITED

Allen, E. 1922. The oestrous cycle in the mouse. Am. J. Anat., 30: 297-371.

Ball, J. 1937. A test for measuring sexual excitability in the female rat.
Comp. Psychol. Monog., 14: 1-37.

Blandau, R. J., J. L. Boling, and W. C. Young. 1941. The length of heat
in the albino rat as determined by the copulatory response. Anat. Rec,
79: 453-463.

Clark, F. H. 1936a. The estrous cycle of the deer-mouse, Peromyscus
maniculatus. Contr. Lab. Vert. Genetics, Univ. Mich., no. 1: 1-9.

Clark, F. H. 1936b. The estrous cycle of the cotton rat. Contr. Lab. Vert.
Genetics, Univ. Mich., no. 2: 1-2.

Coco, R. M. 1942. Notes on vaginal smears in the ground squirrel, Citellus
tridecemlineatus. Proc. Louisiana Acad. Sci., 6: 83-85.

Cole, H. H. 1930. A study of the mucosa of the genital tract of the cow,
with special reference to the cyclic changes. Am. J. Anat., 46: 261-301.

de Allende, I. L. C, E. Shorr, and C. G. Hartman— 1943. A comparative
study of the vaginal smear cycle of the rhesus monkey and the human.
Carnegie Inst. Wash. Contr. Embryol., 31: 1-26.

Deanesly, R., and A. S. Parkes. 1933. The reproductive processes of cer-
tain mammals. IV. The oestrous cycle of the Grey Squirrel, Sciurus
carohnensis. Trans. Roy Soc. London, 222: 47-75.

Deanesly, R. 1938. The reproductive cycle of the golden hamster. Proc.
Zool. Soc. Lond., Series A, 108:31-39.

Donat, F. 1933. Notes on the life history and behavior of Neotoma fuscipes.
J. Mamm., 14: 19-26.

Evans, H. M., and H. H. Cole. 1931. An introduction to the study of
oestrous cycle in the dog. Mem. Univ. Calif., 9: 65-119.

Foster, M. A. 1934. The reproductive cycle in the female ground squirrel.
Citellus tridecemlineatus. (Mitchell). Am. J. Anat., 54: 487-511.

Hartman, C. G. 1944. Some new observations on the vaginal smear of the
rat. Yale J. Biol. & Med., 17: 99-112.

Chapman: The Woodrat Neotoma Floridana 283

Kent, G. C, Jr., and R. A. Smith. 1945. A study of the estrous cycle in the
golden hamster. Anat. Rec., 92: 263-271.

Long, J. A., and H. M. Evans. 1922. The oestrous cycle in the rat and its
associated phenomena. Mem. Univ. Calif., 6: 1-148.

Papanicolaou, G. N. 1933. The sexual cvcle in the human female. Am.
J. Anat, 52: 519-637.

Parkes, A. S. 1931. The reproductivity processes of certain mammals. I. The
estrous cycle of the Chinese hamster. Proc. Roy. Soc, Series R., 108:
138-147.

Pollak, O. J. 1944. A rapid trichrome stain. Arch. Path., 37:294.

Selle, R. M. 1922. Changes in the vaginal epithelium of the guinea-
pig during the oestrous cycle. Am. J. Anat., 30: 429-449.

Sheehan, J. F., and J. A. Bruner. 1945. The care, breeding habits and
vaginal smear cycle of the laboratory hamster, Cricctus auratus. Turtox
News, Chicago, 23: 65-68.

Shorr, E. 1940. A new technic for staining vaginal smears. Science, 91:
321-322.

Stockard, C. R., and G. N. Papanicolaou. 1917. The existence of a typical
oestrous cycle in the guinea-pig with a study of its histological and
physiological changes. Am. J. Anat., 22: 225-283.

Ward, M. C. 1946. A study of the estrous cycle and the breeding of the
golden hamster, Cricetus auratus. Anat. Rec, 94: 139-159.

Wilson, K. M. 1926. Histological changes in the vaginal mucosa of the sow
in relation to the oestrous cycle. Am. J. Anat., 37: 417-431.

Wood, F. D. 1935. Notes on the breeding behavior and fertility of Neotoma
fuscipes macrotis in captivity. J. Mamm., 16: 105-109.

Young, W. C. 1937. The vaginal smear picture, sexual receptivity, and time
of ovulation in the guinea-pig. Anat. Rec, 67: 305-325.

I'M

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PLATE XXIX

Fig. 1. Vaginal epithelium during Stage 1, Woodrat #7; a longitudinal sec-
tion at the posterior end. X 300. The epithelium is thick and appears to have
very little degeneration. Leucocytes, some of which are distorted, are infiltrated
among the epithelium cells.

Fig. 2. Vaginal smear of Stage 1, Woodrat #7, the last one before death.
X 300. Showing a mixture of cells of different types. Arrows indicate some of
the cell types: (a) cornified cell and (b) "precornified" cell, which come from
the external orifice of the vagina; (c) "proestrous" cells and (d) dense cells.

Chapman: The Woodrat Neotoma Floridana

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PLATE XXIX

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PLATE XXX

Fig. 3. Vaginal epithelium of Stage 1, Woodrat #10. X 300. A longitudinal
section showing the epithelium on both sides of the lumen in an area of de-
hiscence between the caudal and middle regions of the vagina. Tall columnar
mucous cells line the lumen on the right and some low mucous cells persist on
the left, where "proestrous" cells and leucocytes are sloughing off.

Fig. 4. Vaginal smear of Stage 1, Woodrat #7, the last one before death.
X 300. "Proestrous" cells are numerous, but pale staining (a). Many "deep"
cells have persisted from the previous stage and appear small anil dark, (b)
A "phagocytized" cell.

Chapman: The Woodrat Neotoma Floridaxa

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PLATE XXXI

Fig. 5. Vaginal smear of Stage 1, Woodrat #20. taken just before death.
X 300. Most of the "proestrous" cells are pale and difficult to see. Some vac-
uolated cells are present, a few in the center of the field have several small
vacuoles in the cytoplasm.

Fig. 6. Vaginal smear of Stage 2, Woodrat #2, taken just before death.
X 300. Vacuolated cells are numerous, with some "deep" cells and a few "pro-
estrous"' cells persisting from the previous stage. The larger dark cells are the
"dense'' cells, (a) one which has the appearance of a flocculate in the cytoplasm.

Chapman: The Woodrat Neotoma Floridana 291

PLATE XXXI

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292 The University Science Bulletin

PLATE XXXII

Fig. 7. Vaginal epithelium of Stage 2, Woodrat #21. X 300. Longitudinal
section with a transverse fold, at the central region of the vagina. Many epi-
thelial cells and leucocytes are seen in the lumen. Some mucous cells are per-
sisting in the fold. Lacunae in the epithelium contain leucocytes and "deep"
cells, (a) A "deep" cell in one of the smaller lacunae; (b) a "deep" cell inside
another epithelial cell in the lumen.

Fig. 8. Vaginal smear of late Stage 2, Woodrat #21, the last one before
death. X 300. "Deep" cells and vacuolated cells are most abundant, (a) An
epithelial cell containing two "deep*' cells.

Chapman: The Woodrat Neotoma Floridana 293

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PLATE XXXIII

Fig. 9. Stage 3, Woodrat #11. Longitudinal section of the vaginal epithe-
lium near the external orifice. X 300. The epithelium is still quite thick in
that area, and large numbers of leucocytes are infiltrated through it and into
the lumen. The transitional zone between the vaginal epithelium and the
cornified layer of the external orifice is seen at the left.

Fig. 10. Stage 3, Woodrat #11. Longitudinal section of the epithelium in
the central region of the vagina. X 300. The epithelium is very thin, lined
with "dense" cells and a few "deep" cells among them, (a) A "deep" cell in
a lacuna.

Chapman: The Woodrat Neotoma Floridana 295

PLATE XXXIII

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296 The University Science Bulletin

PLATE XXXIV

Fig. 11. Stage 3, Woodrat #11. Longitudinal section of the epithelium at
the cervix. X 300. The epithelium is thin. Some mucous cells can still be
seen, especially in the folds, but the surface of the epithelium is mostly "dense"
cells and some "deep" cells distributed among them.

Fig. 12. Stage 3, Woodrat #11. Vaginal smear. X 300. "Deep" cells make
up most of the epithelial elements in the field, (a) A "phagocytized" cell,
which contains one leucocyte.

Chapman: The Woodrat Neotoma Floridana

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PLATE XXXV

Fig. 13. Stage 4, Woodrat #22. Longitudinal section of the vagina in the
central region. X 300. The epithelium is thin and shows little evidence of
either growth or desquamation.

Fig. 14. Stage 4. Woodrat #22. Vaginal smear taken just before death.
X 300. Leucocytes and degenerated epithelial cells are trapped in the strands
of mucous.

Chapman: The Woodrat Neotoma Floridana

299

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THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIV, Ft. I] October 1, 1951 [No. 8

A Revision of the Genus Anisops * (Notonectidae,

Hemiptera)

BY

George T. Bfooks f
TABLE OF CONTENTS

PAGE

Introduction 304

Review of the Literature 305

The Family Notonectidae 306

The Genus Anisops 307

Systematics of the Genus Anisops 308

Phylogeny 309

Biology and Development 310

Distribution 311

Techniques and Terminology 312

Taxonomy of Anisops (with Key to Species) . . 313

Anisops agalia Hutchinson 318

Anisops stali Kirkaldy 319

Anisops campbelli n. sp. 322

Anisops amaryllis Hutchinson 324

Anisops elstoni n. sp 326

Anisops eros Hutchinson 327

Anisops kampalensis Hutchinson 328

Anisops paracrinita n. sp 329

Anisops hacked n. sp 331

Anisops hyperion Kirkaldy 332

Anisops tuherculata Poisson 334

Anisops nodulata n. sp 336

Anisops apicalis Stal 337

Anisops canadensis perplexa Poisson 339

Anisops edepol Kirkaldy 341

* Submitted to the Department of Entomology and the Faculty of the Graduate School of
the University of Kansas in partial fulfillment of the requirements for the degree of Doctor
of Philosophy.

t Associate Professor of Entomology, Department of Biology, Southern University, Baton
Rouge, Louisiana.

(301)

302 The University Science Bulletin

PAGE

Anisops timorensis n. sp 343

Anisops occipitalis Breddin 344

Anisops coutierei n. sp 346

Anisops pellucens Gerstaecker 347

Anisops malkini n. sp 349

Anisops evansi n. sp 350

Anisops gratus Hale 352

Anisops gobana Poisson 353

Anisops varia Fieber 355

Anisops robusta Hutchinson 358

Anisops sikoroensis n. sp 360

Anisops fijiensis n. sp. 361

Anisops adonis Hutchinson 362

Anisops jaczewski Hutchinson 364

Anisops semita n. sp. 366

Anisops canaliculata n. sp 367

Anisops psyche Hutchinson 369

Anisops gen]i Hutchinson 371

Anisops nivea (Fabricius) 373

Anisops tahitiensis Lundblad 376

Anisops tasmaniaensis n. sp 378

Anisops lipovskyi n. sp 379

Anisops deanei n. sp 381

Anisops philippinensis n. sp. 383

Anisops cleopatra Distant 384

Anisops praetexta Hutchinson 386

Anisops barbata n. sp 387

Anisops grandis Poisson 389

Anisops tanalensis n. sp. 391

Anisops windi n. sp. 392

Anisops leucothea Esaki 394

Anisops doris Kirkaldy 395

Anisops wakefieldi White 397

Anisops hancocki Hutchinson 399

Anisops decipiens Hutchinson 401

Anisops barrenensis n. sp 402

Anisops hungerfordi Poisson 404

Anisops ares Hutchinson 405

Anisops paranigrolineata n. sp 407

Anisops nigrolineata Lundblad 409

Anisops assimilis White 411

Anisops thienemanni Lundblad 413

Anisops nasuta Fieber 416

Anisops cavifrons n. sp 417

Anisops batillifrons Lundblad 420

Anisops sardea Herrich-Shaffer 423

Anisoj)s madagascarensis Poisson 428

Anisops bouvieri Kirkaldy 430

Brooks: The Genus Anisops 303

PAGE

Anisops extendofrons n. sp 432

Anisops graciloides n. sp. 434

Anisops gracilis Hutchinson 436

Anisops poweri Hutchinson 437

Anisops leesoniana Hutchinson 438

Anisops breddeni Kirkaldy 439

Anisops kempi n. sp 441

Anisops crinita n. sp 443

Anisops majiensis n. sp. 444

Anisops debilis Gerstaeker 446

Anisops exigera Horvath 447

Anisops vitrea Signoret 449

Anisops hypatia Hutchinson 452

Anisops balcis Hutchinson 453

Anisops biroi n. sp 454

Anisops rigoensis n. sp 456

Anisops waltairensis n. sp 457

Anisops ali Distant 459

Anisops alluaudi Poisson 459

Anisops aphrodite Kirkaldy 460

Anisops calcaratus Hale 461

Anisops canadensis canadensis Noualhier 462

Anisops ciliata Stal 462

Anisops endymion Kirkaldy 463

Anisops hyalina Fieber 464

Anisops lanceolata Poisson 465

Anisops letitia Hutchinson 466

Anisops mauricensis Poisson 467

Anisops meulenaerei Poisson 468

Anisops milloti Poisson 469

Anisops ocularis Hale 469

Anisops pugnax Poisson 470

Anisops worthingtoni Jaczewski 471

Bibliography 472

Illustrations — Plates I — XXII

304 The University Science Bulletin

Abstract: This paper constitutes a revision of the genus Anisops of the
family Notonectidae, Hemiptera. Though it is primarily taxonomic, material
on the biology of genus is included.

Prior to this work sixty-four species and two subspecies were known. Thirty-
one new species are described in this paper. Two color forms are raised to
specific level and two species are taken out of synonomy and placed again on
specific level. Four species are placed into synonomy.

The subgeneric designations as established by Hutchinson in 1929 are
omitted as no evidence was found to support their validity.

The new species are: A. campbelli (India), A. elstoni (Australia), A.
paracrinita (Australia), A. hackeri (Australia), A. nodulata (Philippine Isls.),
A. timorensis (Timor), A. coutierei (Djibouti, Africa), A. malkini (Australia),
A. evansi (Tasmania), A. sikoroensis (Madagascar), A. fijiensis (Fiji Islands),
A. semita (Australia), A. canaliculata (Australia), A. tasmaniaensis (Tas-
mania), A. lipovskyi (India). A. deanei (Australia), A. philippinensis (Philip-
pine Isls.), A. barbata (India, Java), A. tanalensw (Madagascar), A. ivindi
(Australia), A. barrenensis (Australia), A. paranigroUneata (India), A. cavi-
jrons (India), A. extcndofrons (India), A. graciloides (South Africa), A. kempt
(India, Burma, Siam), A. crinita (Corfu, India, New Caledonia), A. majiensis
(Maji-Chumvi, Africa), A. birai (New Guinea), A. rigoensis (New Guinea),
and A. waltairensis (Australia).

Up to and including this paper ( exclusive of the synonyms ) there have been
described ninety-four species and two subspecies.

ACKNOWLEDGMENTS

I wish to express my gratitude to all who have assisted me in
the preparation of this paper. I am deeply indebted to Dr. H. B.
Hungerford, who suggested this study and without whose careful
guidance its completion would not have been possible.. To Dr. R. H.
Beamer, Dr. K. C. Doering, and Dr. C. D. Michener I extend my
sincere thanks for the advice and constructive criticism they so
willingly gave. I highly appreciate the help given me by the de-
partmental artists in the preparation of my plates.

, INTRODUCTION

This paper is a taxonomic study of the genus Anisops ( Notonec-
tidae, Hemiptera), designed to alleviate many of the problems
which have originated through poor descriptions and inaccurate
determinations. It covers ninety-six species, thirty of which are
new. In addition to having material from the Francis Huntington
Snow Collection, a vast amount of material from other museums
was placed at my disposal. The Museums, which made their ma-
terial available to me, are as follows: British Museum, Paris Mu-
seum, Royal Museum of Natural History of Belgium, Basel Natural
History Museum of Switzerland, Indian Museum at Calcutta, United

Brooks: The Genus Anisops 305

States National Museum, Harvard Museum of Comparative Zoology,
and the Chicago Natural History Museum. In addition, I have
studied material from the collections of Lutz, Usinger, California
Academy of Science, and the Department of Agriculture, Sydney,
New South Wales, Australia. To each of these I wish to express my
gratitude for without their cooperation, this study would have been
impossible.

REVIEW OF THE LITERATURE

In 1840, Spinola (38) established the genus Anisops. Franz X.
Fieber (11) in 1851, redescribed the genus and all known species
belonging thereto. During the next half-century many species were
described; some were valid, others not, for unfortunately it was
the practice of the taxonomists of that time to describe and deter-
mine species largely on the basis of color. This resulted in much
confusion not only in regard to the accuracy of determination but
also in ascertaining the possible geographical distribution of a
species.

In 1904, Kirkaldy (30) published his "t)ber Notonectiden." In
this paper three important changes were made. First, he exam-
ined types when possible and indicated synonymous species. Sec-
ond, he introduced the use of structural characteristics in species
determination. Third, he divided the genus into two genera. The
generic name Buenoa was proposed for those species inhabiting
the Western Hemisphere; the name Anisops was retained for the
species of the Eastern Hemisphere. However, this separation was
not made on geographical distribution alone, but on the fact that
the males of the species of the Western Hemisphere all have two-
segmented front tarsi (the third is present, but so small as not to
be readily discernible) while the males of the species of the East-
ern Hemisphere have one-segmented front tarsi.

In 1923, H. M. Hale (14) published "Some Aquatic Hemiptera
from Western Australia" which included descriptions of the species
of that area along with a key. In 1929, G. E. Hutchinson (23) pub-
lished "A Revision of the Notonectidae and Corixidae of South
Africa." In this publication keys and descriptions for species of
that area were provided. Lundblad (32) in 1933 published "Zur
Kenntnis der Aquatilen und Semi-Aquatilen Hemipteren von Su-
matra, Java, and Bali," in which many of the species of that area
were described adequately for the first time. Though each of these
above mentioned papers is extremely valuable, together they ac-
counted for only about half of the species known in 1933.

20—3286

306 The University Science Bulletin

Through the years two problems have consistently plagued the
workers in this genus. First, most of the older descriptions were
based on color and it is almost impossible to determine species on
this basis. Second, the finding of suitable characters to separate
the species has been particularly difficult. As has been pointed
out by previous workers, notably Lundblad (32) and Hutchinson
(23), only the males can be relied on for accurate determination.
Unfortunately, the male genital capsule is not of specific value in
this genus and other characters for separation of the species must
be found.

Jaczewski (28) noted that the shape of the eyes together with
that of the vertex appeared to be of specific importance as well as
the shape and size of the rostral prong. Poisson in many of his
descriptions has made extensive use of the chaetotaxy of the male
front legs and Hutchinson (23) has indicated the specific value of
the facial tubercle. Lundblad ( 32 ) made use of all these characters
in his specific descriptions.

After examining many species, I find the above characters to-
gether with the shape of the labrum and front femur of the males
afford a ready means of species determination.

This paper represents the first generic revision since Kirkaldy's
paper of 1904. I have tried to account for every named species, but
in some cases the descriptions were based on females and it is im-
possible to assign a species to such a description, as females of
different species are often almost identical; in other instances the
material available yielded no specimens which matched the de-
scriptions.

I was extremely fortunate in having at my disposal many speci-
mens, which were compared with types by D. H. B. Hungerford
in 1928. This has been extremely valuable, especially in cases of
unavailable types and poor descriptions.

THE FAMILY NOTONECTIDAE

The family Notonectidae includes those truly aquatic forms which
differ from all others, except Pleidae and Helotrephidae, in the
constant habit of swimming on their backs. The members of this
family are deep-bodied, flat ventrally, and convex dorsally. The
eyes are large reniform, and twice sinuate, occupying most of the
dorsal surface of the head. Ocelli are absent. The antennae arise
on the latero-ventral surface of the head, immediately posterior to
the eyes; each antenna consisting of three or four segments. Beak
four-segmented. Front and middle pairs of legs adapted for grasp-

Brooks: The Genus Anisops 307

ing, hind legs for swimming. Tarsi (except the one-segmented
fore tarsi of the male Anisops) three-segmented counting a small
basal segment which is inconspicuous. Tarsi possessing two apical
claws which may be greatly reduced on the hind pair of legs. Ven-
tral abdominal segments with median longitudinal keel, which
has hairs, at least on its lateral margin. Sides of the venter with
hairs directed medianly.

This family consists of eight genera which may be separated by
the following key as advanced by Hungerford (22).*

Key to Genera of the Notonectidae

A. Hemelytral commissure without definite hair-lined pit at anterior end

(Subfamily Notonectinae)
B. Intermediate femur with anteapical pointed protuberance and antennae

4-segmented (Tribe Notonectini)

C. Anterolateral margins of prothorax not foveate Notonecta

CC. Anterolateral margins of the prothorax foveate Enithares

BB. Intermediate femur without anteapical pointed protuberance. Antennae

3- or 4-segmented (Tribe Nychini)

C. Intermediate tarsus with two well -developed segments and a very
small basal one. Sides of prothorax not foveate. Infra-coxal

plates bare, but margined with hair Neonychia

CC. Intermediate tarsus with one well -developed segment. Infra-coxal
plates covered with hair.

D. Antennae three segmented Nychia

DD. Antennae four segmented Martarega

AA. Hemelytral commissure with definite hair-lined pit at anterior end

(Subfamily Anisopinae)
B. Ventral abdominal keel not extending onto last abdominal segment.
Male genital capsule cleft behind. Males without stridular protuberance

on front tibia. Females with short gonapophyses Paranisops

BB. Ventral abdominal segment extending onto last abdominal segment.

C. Male with anterior tarsus 2 -segmented Buenoa

CC Male with anterior tarsus 1 -segmented Anisops

THE GENUS ANISOPS

The genus Anisops is readily distinguishable from all other genera
in the family by the following three combination of characters:
the presence of a pit at the anterior end of the hemelytral commis-
sure, the extension of the ventral abdominal keel onto the last
abdominal segment, and the one-segmented front tarsi of the males.

The members of this genus are slender insects and usually small,
though some species attain a length of about 12 mm. The eyes are
large and not holoptic (except A. breddeni Kirkaldy and A. kempi
n. sp. ) . The anterior half of the dorsal interocular space has a me-
dian longitudinal depression between two slightly swollen areas.
The back is convex longitudinally with the thorax being deeper
than the other parts of the body. The hemelytra are not coriaceous

* Later modified to include the genus Paronychia. Still later changed to Neonychia
(see Journ. Kansas Ent. Soc, vol. XXIII. p. 73, 1950).

308 The University Science Bulletin

as in Notonecta and Enithares and there is little difference in the
texture of the membrane and the remainder of the wing though
the two areas are definitely delimited. The underside of the abdo-
men has a median longitudinal keel whose lateral margins bear
long hairs. On either side of the abdomen next to the connexivum
is a longitudinal trough. The inner margins of the connexivum
bear long hairs which together with those of the keel serve to trap
air in the trough. All the tarsi are two-segmented with the excep-
tion of the male front tarsi which are one-segmented.

Only the males possess a stridulatory apparatus, which consists of
a tibial comb and a rostral prong (to be described in detail later).
The claspers of the genital capsule are asymetrical and of uniform
shape for the genus, the right one being broad, while the left is
deeply excavate on the posterior margin and hook-shaped at the
apex. ( Plate XXXVI figs. 3, 4. )

An interesting feature of the internal anatomy of both males and
females is the presence of two ventral rows of cells containing hemo-
globin (35). Each row is divided into distinct groups correspond-
ing in number to the abdominal spiracles and attached to the latter
by the tracheal trunk. These cells are tracheal cells, around and in
which the tracheoles ramify. A similar group of cells have been
observed by Bare (2) in the genus Buenoa.

SYSTEMATICS OF THE GENUS ANISOPS

Hutchinson in 1929 divided the genus into three subgenera,
namely Microanisops, Anisopoides, and Anisops. The first two are
monotypic.

The subgenus Microanisops was erected on the following char-
acters: brachypterous wings and elytra, a short posterior tarsus
with the tibia being one and three-fourths the length of the latter,
and the presence of a subapical black spot on the hemelytra. The
subgeneric type is Anisops (M.) apicalis Stal. In a study of a male
and female of this species determined by myself, I do not find the
hemelytra to be brachypterous, but extending the full length of the
body; however the wings are reduced and appear non-functional.
Moreover, the posterior tibia is not one and three-fourths the length
of the posterior tarsi but only one and two-sevenths, a relationship
also found by Jaczewski (23). Anisops (M.) apicalis is not unique
in having the subapical black spot as I have observed similar spots
on specimens of A. (A.) ares Hutchinson. The single good character
for such a group would be the brachypterous wings. However,
Anisops (Anisops) hungerfordi Poisson which occurs in central

Brooks: The Genus Anisops 309

Africa along with Anisops (M.) apicalis, is so closely related to this
species that they may be forms of the same species. They differ
primarily in the character of the scutellum which is greatly reduced
in size in Anisops (M.) apicalis, a condition which is to be expected
in brachypterous forms. Also, the wings of the Anisops (A.) hunger-
fordi are not brachypterous. A closer investigation of the biology
of the two species would be necessary before determining whether
or not they are forms of the same species.

The type of the subgenus Anisopoides is Anisops (Anisopoides)
agalia Hutchinson and the subgenus is based on the lack of rostral
prongs. The presence of the stridulatory comb on the tibia sug-
gests that this species arose from forms having rostral prongs and
which have been lost through specialization.

In my study of the genus, I have found no forms to add to either
of the subgenera of Hutchinson. Both subgenera appear to be only
individual specializations and not worthy of subgeneric designation.
Therefore, I have chosen not to use the subgeneric groups as es-
tablished by Hutchinson as I do not believe such to be valid.

The term "form" has been used by previous workers to describe
specimens having a different color pattern from that of the type
of the species. Only in two cases, A. sardea f. madagascarensis Pois-
son and A. pellucens f. grandis Poisson are such differences sup-
ported by morphological differences. In both of these cases I have
raised the so-called "forms" to species level. Otherwise, I have dis-
regarded the "form" as a subspecific designation.

PHYLOGENY

The species of Anisops exhibit many specializations. Most of
them do not indicate phyletic tendencies as they occur, oftentimes,
in completely unrelated forms.

The only specialization that appears to indicate a phyletic line is
an increase in the size of the eyes. On the basis of this character,
the species of the genus fall in one of two groups. In one the
enlargement of the eyes is accompanied by a subsequent narrowing
of the interocular space from the synthlipsis forward. The peak of
this series is reached in A. breddeni Kirkaldy and in A. kempt n. sp.,
both of which have holoptic eyes. This group embraces the
majority of the species. Within this group there is a small discrete
unit of eight species, so closely allied to one another and repre-
senting such a unique line of specialization as to be worthy of
note. This small group is characterized by males possessing an
excavate frons and peculiarly curved middle tarsal claws in which

310 The University Science Bulletin

the claws are turned strongly inward at the base with the posterior
claw thicker than the anterior one.

The second major group on the basis of the enlargement of the
eyes is somewhat smaller than the foregoing and is one in which the
enlargement of the eyes proceeds laterally. This is accompanied
by the narrowing of the anterior width of the vertex until the dorsal
interocular space becomes almost parallel-sided with the synthlipsis
remaining broad. The peak of this series is reached in A. evansi
n. sp. and A. tasmaniaensis n. sp. in which the head is at least as
wide as the pronotal humeral width and the synthlipsis is three-
fourths the anterior width of the vertex.

Some of the specializations, which appear in completely unrelated
forms are: the dorso-ventral enlargement of the male front femur;
the projecting of the interocular space into an anterior cephalic
horn; the facial tubercle becoming depressed, grooved, excavate
or laterally compressed, and the elongation of the pronotum.

BIOLOGY AND DEVELOPMENT

Hale (13, 14) and Poisson (35) have both studied the biology
and development of Anisops, using Anisops thienemanni Lundblad
(= Anisops hyperion Hale, nee. Kirkaldy) and Anisops sardea Her-
rich-Schaffer respectively.

The members of this genus of backswimmers live in freshwater
pools, lakes, and ponds; however at least one species, A. sardea is
adapted to marine life as well. As with the other members of the
family they swim on their backs and are predaceous, feeding on
small crustaceans and mosquito larvae, which they hold securely be-
neath the bristles arming the margins of the fore and middle legs.

In order to replenish an air supply the insects ascends to the
surface of the water where the venter is opened by the spreading
apart of the guard hairs of the connexivum and keel. By the sudden
closing of the guard hairs air is gathered in the longitudinal air
troughs, after which the insect quickly darts down a few inches into
the body of the water where it maintains almost perfect balance.
Tendencies to rise or sink are offset by strokes of the swimming legs.
Poisson points out that when fatigued the insect will sink to the
bottom of the water and there remain on its back for a short period.

Preceding copulation there is an elaborate courtship during which,
as recorded by Hale ". . . the male stridulating the while, poises
below and a little behind the female, and in this position accom-
panies her every movement; finally with the extended posterior

Brooks: The Genus Anisops 311

legs quivering with excitement, he attempts to clasp her from be-
low."

This stridulatory apparatus consists of a stridulatory comb borne
on an expanded basal portion of the inner surface of each anterior
tibia and a pair of chitinous prongs, one on each side of the third
rostral segment. The stridulation is accomplished by rubbing the
comb over the rostral prong.

The eggs are laid singly in stems of aquatic plants and to effect
oviposition the female gouges a hole in the stem of the selected
plant and inserts the egg leaving a small portion of the anterior
surface exposed at the mouth of the cavity. The egg is oval and
elongate and Poisson notes that the eggs of A. sardea bore two small
appendages at the anterior end which he regarded as rudimentary
pneumatic appendages.

The eggs hatch in about three weeks. There are five nymphal
instars, the first four occupying each a little more than a week, with
the fifth a little longer. Adulthood is reached in about two months.
Hale points out that there are at least two generations a year.

The newly hatched nymphs are helpless until the air troughs are
filled with air. During this period movement is awkward. A period
of three days may elapse before the surface film is attained and
the air troughs filled. Possibly during this period respiration is
effected through the skin.

Anisops has almost perfect balance in the water and rarely clings
to the plant stems as in the case of Notonecta and Enithares. This
ability is likewise encountered in Buenoa which is quite similar to
this genus in many respects.

DISTRIBUTION

The geographical range of Anisops extends throughout Africa,
Madagascar and neighboring islands, the Mediterranean area, ex-
tending eastward to the islands of the South Pacific, thence north-
ward into China and Japan, with A. sardea carrying its range along
the entire eastern coast of Asia and, while this record is somewhat
questionable, Fieber (11, 12) records it from Unalaska in the West-
ern Hemisphere.

Although many species have been recorded from Asia as well as
Africa, I have been able to find only two species whose distribution
is so widespread, Anisops sardea which occurs throughout most of
the generic range and Anisops crinita n. sp. which occurs in India
as well as on the Mediterranean island of Corfu.

312 The University Science Bulletin

TECHNIQUES AND TERMINOLOGY

In order to interpret correctly the species descriptions, and to use
them successfully, the reader should be acquainted with the tech-
nique and terminology employed herein.

The measurements of the head, pronotum and their respective
parts were taken with the insect in a perfectly horizontal position.
To view and measure the rostral prong, the specimen should be
placed in a lateral position and the length of the prong measured
from the base of the third rostral segment to the apex of the prong,
along the posterior margin of the latter. To secure accurate meas-
urement of the prong, the bug should be tilted until the prong is
horizontal. The length of the third rostral segment is measured
along its posterior margin.

To observe accurately and closely the chaetotaxy of the male
front leg, the leg should be removed. This is best accomplished by
placing the tip of a pin at the base of the coxa on its inner surface
and applying a little outward pressure. The break is cleaner and
accomplished with much less pressure if the specimen is dry and has
not been relaxed. The leg should then be placed in a caustic potash
solution until clear enough to observe. This clearing can be ac-
complished in a few minutes if the solution is hot, though not boil-
ing. If placed in a cold solution, one day will be sufficient for
clearing. It will be noted that my plates do not show the thick cov-
ering of hairs along the anterior half of the inner surface of the leg.
These were purposely omitted so that the characteristic setae would
not be obscured. The relative lengths of the parts of the legs are
measured along the longest part of each segment.

Different workers in describing members of this genus have used
different terms for the same structure. In the preparation of this
manuscript, the most descriptive terms have been employed. Below
is a short list of words and phrases with which the reader should
be acquainted to fully understand the descriptions.

1. Anterior width of the vertex — the width of the interocular space lying be-
tween the anterior margins of the eyes as seen from above.

2. Facial tubercle — that portion of the frons immediately above the labrum.

3. Frons — the lower portion of the interocular space from the labrum to the
anterior margin of the head.

4. Interocular space — that portion of the head between the eyes.

5. Rostral prong — the pair of projections borne on the third rostral segment,
one on each side.

6; Synthlipsis — the narrowest dorsal portion of the interocular space immedi-
ately anterior to the pronotum.

7. Stridulatonj ridge — the expanded portion of the inner basal surface of the
anterior tibiae of the males bearing the stridulatory comb.

Brooks: The Genus Anisops 313

TAXONOMY OF THE GENUS ANISOPS

Genus Anisops Spinola

( Type A. sardea Herrich-Shaffer )

1840. Anisops Spinola, Essais sur les Insectes Hemipteres, Rhyngotes ou Heteropteren,
p. 58.

1851. Anisops, Fieber, Abhandl. Kongl. Bohmischen Ges. der Wiss., vol. V, pt. 7, pp. 481-
482.

1865. Anisops, Stal, Hemiptera Africana, vol. Ill, p. 191.

1904. Anisops, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, p. 111.

1926. Anisops, Jaczewski, Ann. Zool. Mus. Polonici Hist. Nat., vol. II, p. 81.

1929. Anisops, Hutchinson, Ann. South African Mus., vol. XXV, pt. 3, pp. 376-382.

Of the ninety-four species herein described, sixteen species,
namely Anisops ali Distant, Anisops alluaudi Poisson, Anisops aphro-
dite Kirkaldy, Anisops calcaratus Hale, Anisops canadensis Noual-
hier, Anisops ciliata Stal, Anisops endymion Kirkaldy, Anisops hya-
line Fieber, Anisops lanceolata Poisson, Anisops letitia Hutchinson,
Anisops mauricensis Poisson, Anisops meulenaerei Poisson, Anisops
milloti Poisson, Anisops ocularis Hale, Anisops pugnax Poisson, and
Anisops worthingtoni Jaczewski, were not present in the material
studied. In such cases I have presented the original descriptions
and, when possible, I have included copies of the drawings on my
plates. However, they were omitted from the key. The following
is a key to the males of Anisops.

Key to Males of Anisops

1. Syntlilipsis wide, one third or more the anterior width of the vertex 2

Synthlipsis narrow, less than one third the anterior width of the vertex 69

2. (1 ) Third rostral segment with prongs 3

Third rostral segment without prongs A. agaJia Hutchinson

3. (2) Along the median longitudinal axis, the head is one half or more as long as

the pronotum 4

Along the median longitudinal axis, the head is less than one half of the
length of the pronotum 57

4. (3) Greatest width of the head less than seven times the anterior width of the

vertex 5

Greatest width of the head more than seven times the anterior width of
the vertex 42

5. (4) Facial tubercle either longitudinally grooved, or excavate, or laterally com-

pressed forming a median carina 32

Facial tubercle neither longitudinally grooved, nor excavate, nor laterally
compressed forming a median carina 6

6. (5) Interocular space projected anteriorly into a short cephalic horn 7

Interocular space not projected anteriorly into a short cephalic horn 8

7. (6) Base of middle tibia with an inward projection which bears on its inner sur-

face an oval patch of short, stout, closely set setae .4. stali Kirkaldy

Base of middle tibia not so formed A. campbelli n. sp.

8. (6) Over 6.5 mm. in length 24

Less than 6.5 mm. in length 9

314 The University Science Bulletin

9. (8) Synthlipsis one half or more the anterior width of the vertex 20

Synthlipsis less than one half the anterior width of the vertex 10

10. (9) Apex of front femur rounded or truncate 17

Apex of front femur accuminate 11

11. (10) Inner surface of front tarsus without median small setae. .A. amaryllis Hutchinson

Inner surface of front tarsus with one or more median small setae 12

12. (11) Inner surface of front tarsus with one basal median setae .4. elstoni n. sp.

Inner surface of front tarsus with a median row of small setae 13

13. (12) Inner surface of front tarsus with a median row of three small setae in

basal third 1 4

Inner surface of front tarsus with a median row of more than three small
setae extending beyond the basal third 15

14. (13) Apex of rostral prong more or less rounded; stridulatory comb with longest

teeth subapical A. eros Hutchinson

Apex of rostral prong accuminate; stridulatory comb with longest teeth

at apex -4. kampalensis Hutchinson

1.3. (13) Facial tubercle with a large patch of long erect hairs .4. paracrinita n. sp.

Facial tubercle without a patch of hairs 10

10. (15) Labium with basal width twice its median length -4. hacked n. sp.

Labrum with basal width less than twice its median length, .4. Hyperion Kirkaldy

17. (10) Anterior margin of pronotum bearing a large prominent tubercle behind the

synthlipsis -4. tuberculata Poisson

Anterior margin not bearing a large tubercle 18

15. (17) Third rostral segment bearing two nodules on its anterior median line

.4. nodulata n. sp.

Third rostral segment not bearing nodules on anterior line 19

19. (18) Posterior margin of pronotum straight, not medianly emarginate. ...4. apicalis Stal

Posterior margin of pronotum medianly emarginate

.4. canariensis perpleia Poisson

20. (9) Apex of front femur truncate or rounded 22

Apex of front femur accuminate 21

21. (20) Facial tubercle bearing a short erect spur .4. edepol Kirkaldy

Facial tubercle not bearing a short erect spur 4. elstoni n. sp.

22. (20) Second rostral segment with a flap-like extension along basal half of posterior

surface A. timorensis n. sp.

Second rostral segment not so formed 23

23. (22) Viewed laterally the anterior margin of the third rostra! segment overlaps

at its apex Ihe base of the fourth .4. occipitalis Breddin

Viewed laterally the third rostral segment does not overlap the base of

the fourth -4. coutierei n. sp.

2 1. (8) Large, robust species over 9.0 mm. in length A. pellucens Gerstaeker

Less than 9.0 mm. in length 25

25. (24) Apex of front femur accuminate 26

Apex of front femur rounded or truncate 30

20. (25) Anterior margin of fore tibia with a row of eleven prominent setae extendiai;

from basal third to apex -4. malkini n. sp.

Anterior margin of fore tibia without suih a row of setae 27

27. (26) Greatest width of head equal to the pronotal humeral width .4. evansi n. sp.

Greatest width less than the pronotal width 28

28. (27) Inner surface of fore tarsi with only a median large, stout setae at base

A. gratus Hale

Inner surface of fore tarsi with a median longitudinal row of small setae. ... 29

Brooks: The Genus Anisops 315

29. (28) Greatest width of head five times the anterior width of the vertex

.4 hyperion Kirkaldy

Greatest width of head less than five times the anterior width of the

vertex -4. gobana Poisson

3D. (25) Dorsal margin of anterior femur with a slight concavity at apical third 31

Dorsal margin of anterior femur without a slight concavity A. varia Fieber

31. (30) Greatest width of head equal to the pronotal humeral width; pronotal lateral

margins convex and converging .-1 robusta Hutchinson

Greatest width of head less than the pronotal humeral width ; pronotal
lateral margins almost straight and diverging A. occipitalis Breddin

32. (5) Large species, 9.0 mm. or more in length; frons with a median longitudinal

row of erect hairs A- sikorensis n. sp.

Less than 9.0 mm. in length ; frons without a median row of hairs 33

33. (32) Facial tubercle laterally compressed forming a median carina 34

Facial tubercle not laterally compressed forming a median carina 30

34. (33) Rostral prong longer than third rostral segment -4 fijiensis n. sp.

Rostral prong shorter than third rostral segment 35

35. (34) Fore tarsus two thirds the length of the fore tibia .4. adonis Hutchinson

Fore tarsus less than two thirds the length of the fore tibia

.4. jaczewski Hutchinson

36. (33) Inner surface of anterior tibia with two pairs of short stout setae at its

anterior margin midway between the base and apex 37

Inner surface of anterior tibia without such setae 38

37. (36) Base of middle tibia with an inward projection which bears on its inner sur-

face an oval patch of short, stout, closely-set setae .4. stali Kirkaldy

Base of middle tibia not so formed A. campbelli n. sp.

35. (36) Facial tubercle with a median longitudinal groove 39

Facial tubercle not so 40

39. (38) Ridges bordering longitudinal groove provided with long erect hairs

A. seniita n. sp.

Ridges bordering longitudinal groove not provided with long erect hairs

.4. canaliculata n. sp.

40. (38) Middle tarsal claws strongly curved inward at base; posterior claw thicker

than anterior one ■* l

Middle tarsal claws not strongly curved inward at base ; posterior and ante-
rior claws the same width A psyche Hutchinson

41. (40) Frontal excavation bordered by two ridges on each side .4 genji Hutchinson

Frontal excavation bordered by only one ridge .4 nivea (Fabricius)

42. (4) Greatest width of head equal to or more than pronotal humeral width 54

Greatest width of head less than pronotal humeral width 43

43. (42) Facial tubercle laterally compressed forming a median carina 44

Facial tubercle not laterally compressed 46

44. (43) Apex of front femur truncate or rounded 45

Apex of front femur accuminate -4 fijiensis n. sp.

45. (44) Vertex not extending beyond the anterior margins of the eyes

A. tahitiensis Lundblad

Vertex extending slightly beyond the anterior margins of the eyes

A. lipovskyi n. sp.

46. (43) Synthlipsis less than one half the anterior width of the vertex 47

Synthlipsis one half or more the anterior width of the vertex 50

47. (46) Inner surface of fore tarsi provided with a median row of five small setae

extending from base to apex A. deanei n. sp.

Inner surface of fore tarsi not provided with a median row of setae ex-
tending from base to apex 48

316 The University Science Bulletin

48. (47) Rostral prong shorter than third rostral segment A philippinensis n. sp.

Rostral prong longer then third rostral segment 49

49. (48) Inner surface of fore tarsi without a row of small setae A. cleopatra Distant

Inner surface of basal fourth of fore tarsi with a short row of four setae

A. praetexta Hutchinson

50. (46) Large species ; over 8.0 in length 51

Less than 8.0 in length 52

51. (50) Facial tubercle with two median tufts of hairs .4. barbata n. sp.

Facial tubercle without two tufts of hairs A. grandis Poisson

52. (50) Apex of front femur rounded or truncate A. tanaknsis n. sp.

Apex of front femur not accuminate 53

53. (52) Rostral prong slightly shorter than third rostral segment. .A. philippinensis n. sp.

Rostral prong at least as long as the third rostral segment A windi n. sp.

51. (42) Apex of front femur rounded A. leucothea Esaki

Apex of front femur accuminate 55

55. (54) Pronotal lateral margins parallel A. doris Kirkaldy

Pronotal lateral margins diverging 56

56. (55) Greatest width of head seven to eight times the anterior width of the

vertex ; over 7.0 mm. in length .4. tatmaniaensis n. sp.

Greatest width of head ten times the anterior width of the vertex ; less
than 6.0 mm. in length A, phiiippinensis n. sp.

57. (3) Facial tubercle swollen into a triangular eminence; apex accuminate

A. ivakefieldi White

Facial tubercle not so formed 58

58. (57) Facial tubercle excavate or longitudinally grooved 59

Facial tubercle not as above 60

59. (58) Facial tubercle deeply excavate .4. hancocki Hutchinson

Facial tubercle longitudinally grooved 4. canaliculata n. sp.

60. (58) Pronotal lateral margins almost parallel 68

Pronotal lateral margins diverging 61

61. (60) Median length of pronotum three fourths the humeral width 67

Median length of pronotum less than three fourths the median length ...... 62

62. (61 ) Front femur with apex curved 64

Front femur with apex accuminate 63

63. (62) Pronotum with a deep median longitudinal depression; greatest width of

head eight tenths pronotal humeral width .4. gratus Hale

Pronotum without median longitudinal depression; greatest width of head
nine tenths pronotal humeral width .4. decipiens Hutchinson

64. (62) Less than 6.0 mm. in length 65

6.0 mm. or more in length 66

65. (64) Front femur greatly enlarged dorso-ventrally ; dorsal margin strongly con-

vex .4. barrenensis n. sp.

Front femur nol greatly enlarged; dorsal margin not strongly convex

.4. hungerfordi Poisson

60. (64) Dorsal and ventral margins almost parallel for the basal three fourths of

(heir length .4. ares Hutchinson

Dorsal and ventral margins converging from the base .4. varia Fieber

67. (61) Viewed laterally, the interocular space not swollen beyond the margins of

the eyes .4. paranigrolineata n. sp.

Viewed laterally, the interocular space is swollen beyond the margins of
the eyes A. nigrolineata Lundblad

Brooks: The Genus Anisops 317

68. (60) Greatest width of head equal to pronotal humeral width A. assimilis White

Greatest width of head less than pronotal humeral width

A. thienemanni Lundblad

69. (1) Interocular space produced anteriorly into a cephalic projection 70

Interocular space not produced anteriorly into a cephalic projection 76

70. (69) Frons not excavate for its entire length .4. nasuta Fieber

Frons excavate • 1

71. (70) Cephalic projection rounded at apex 72

Cephalic projection more or less accuminate 73

72. (71) Basal third of fore tibia with a large procumbent spine which lies on the

dorsal surface .4. cavifrons n. sp.

Basal third of fore tibia without such a spine .4. batillifrons Lundblad

73. (71) Inner surface of fore tibia with five prominent setae extending from basal

fourth to apex 74

Inner surface of fore tibia without such setae 75

74. (73) Basal two tibial setae of anterior leg spatulate; tip ot the cephalic projection

not black .4. sardea Herrick -Shaffer

Basal two tibial setae of anterior leg not spatulate ; tip of the cephalic
projection black A. madagascarensis Poisson

75. (73) Viewed on inner surface, subapical fifth of fore tibia angulate, A bouvieri Kirkaldy

Viewed on inner surface, apex of tibia rounded .4. extendojrons n. sp.

76. (69) Facial tubercle excavate, with lateral margins carinate .4. nivea Fieber

Facial tubercle not excavate, at most only slightly depressed 77

77. (76) Stridulatory ridge transversely striated 78

Stridulatory ridge not transversely striated 79

78. (76) Facial tubercle with faint median depression; rostral prong rounded at

apex A. gruciloides n. sp.

Facial tubercle not faintly depressed ; rostral prong accuminate at apex

.4. gracilis Hutchinson

79. (76) Longer claw of front leg more than one half as long as front tarsi 80

Longer claw of front leg not more than one half as long as fore tarsi 81

80. (79) Facial tubercle greatly swollen. . . .4. poweri Hutchinson

Facial tubercle not greatly swollen .4. leesoniana Hutchinson

81. (79) Eyes holoptic in basal half of dorsal surface 82

Eyes not holoptic 83

82. (81) Pronotal lateral margins parallel .4. breddeni Kirkaldy

Pronotal lateral margins diverging A. kempi n. sp.

83. (81) Greatest width of head seven times or more the anterior width of the

vertex ^1

Greatest width of head less than seven times the anterior width of vertex. ... 84

84. (83) Facial tubercle covered with hairs 85

Facial tubercle not covered with hairs 86

85. (84) Hairs of facial tubercle long and erect -4. paracrinita n. sp.

Hairs of facial tubeicle short and procumbent .4. crinita n. sp.

86. (84) Inner surface of anterior tibia provided with stout setae along the apical

two thirds of its anterior margin 87

Inner surface of fore tibia not so armed 88

87. (86) Synthlipsis extremely narrow, approximately one tenth the anterior width

of the vertex; apex of front femur rounded A. majiensis n. sp.

Synthlipsis one sixth the anterior width of the vertex ; apex of fore femur
accuminate .4. debilis Gerstacker

318 The University Science Bulletin

8S. (86) Over 6.5 mm. in length 4. gratus Hale

Less than 6.5 mm. in length 89

89. (88) Rostral prong shorter than third rostral segment; apex rounded

A. exigera Horvath

Rostral prong longer than third rostral segment ; apex accumulate 90

90. (89) Facial tubercle flat -4. vitrea Signoret

Facial tubercle slightly raised, not flat -4. hypatia Hutchinson

91. (83) Facial tubercle slightly raised and flattened on ventral surface 92

Facial tubercle not flattened on ventral surface 93

92. (91) Small species, less than 5.5 mm. in length A. vitrea Signoret

Over 6.0 mm. in length -4. balcis Hutchinson

93. (91) Anterior tarsus without a median row of small setae on inner surface 94

Anterior tarsus with a median row of small setae on inner surface 95

94. (93) Dorsal margin of third rostral prong appears to arise near the apex of

third rostral segment A. biroi n. sp.

Dorsal margin of third rostral prong appears to arise in basal half of third
rostral segment A. rigoensis n. sp.

Ofi. (93) Anterior tarsus with a median row of five small setas on inner surface

A. deanei n. sp.

Anterior tarsus with a median row of three small setae on inner surface

A. waltairensis n. sp.

Anisops agalia Hutchinson

(PI. XXXVI, fig. 7)

1929. Anisops agalia Hutchinson, Ann. South African Mus., vol. XXV, pt. 3, pp. 404-406,
PI. XXIX, fig. 4, PI. XXXII, fig. 7, 7a.

1933. Anisops agalia, Hutchinson, Internat. Rev. ges. Hydrob. und Hydrog., vol. 28, pt.
5/6, pp. 446, 459, 460 (ecological notes).

Size. — Males, length 5.9 mm., greatest body width 1.5 mm.;
females, length 6.6 mm., greatest body width 2.1 mm.

Shape. — Fusiform species; greatest body width about two fifths
the body length.

Color. — General facies stramineous. Eyes brown. Legs stramine-
ous. Abdominal venter dark brown.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the greatest width of the head slightly
less than the humeral width of the pronotum and almost four times
the anterior width of the vertex; synthlipsis wide, one third the an-
terior width of the vertex; along the median longitudinal axis the
head is two thirds the length of the pronotum. Pronotum with its
humeral width almost twice its median length; lateral margins di-
verging and almost one half the median length; posterior margin
convex, medianly emarginate. Facial tubercle slightly raised. La-
brum short with its basal width one and one half the median length;
apex truncate, almost one-third the basal width. Third rostral seg-
ment lacking rostral prongs. Stridulatory comb (PI. XXXVI, fig. 7b)

Brooks: The Genus Anisops 319

of approximately six, even-length teeth. Anterior tibia with its
inner surface covered with procumbent spatulate setae. Chaetotaxy
of the front leg as shown on Plate XXXVI. The relative lengths of
the parts of the legs are as follows : *

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 128 86

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with the greatest width of the head ap-
proximately eight tenths the pronotal humeral width and four times
the anterior width of the vertex; synthlipsis wide, more than one
third the anterior width of the vertex; along the median longitudinal
axis the head is almost one-half the length of the pronotum. Pro-
notum with its humeral width twice its median length; lateral mar-
gine diverging and slightly more than one half the median length;
posterior margin convex, medianly emarginate. The relative lengths
of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 119 56 31

Middle leg 100 78 36 22

Hind leg 100 79 43 46

Location of types. — Type, allotype, and paratypes from Cape,
Transvaal in the South African Museum.

Comparative notes. — By its lack of rostral prongs, A. agalia
Hutchinson is different from all other known species of Anisops.
However, it appears similar to A. hypatia from which it can be
readily separated by the fact that the males of A. hypatia lack the
spatulate front tibial hairs as found on the males of A. agalia as well
as by the presence of rostral prongs.

Data on distribution:

South Africa

Vlei Roxana, 1-21-1927, C. F. Mobray, one male, one female, gift to H. B. H.
from G. E. Hutchinson. (F. H. Snow Coll.)

Anisops stall Kirkaldy

(PI. XXXVII, fig. 9; PI. LVII, fig. 107)

1855. Notonecta australis Stal, Ofversigt af Kongl. Vet. Akad. For., vol. VII, p. 89.
(nee Oliver).

1859. Notonecta australis, Stal, Eugenies Resa, p. 267.

1904. Anisops stall, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 113, 132.

1923. Anisops staii, Hale, Rec. South Australian Mus., vol. II, no. 3, pp. 414-415, fig. 36S.

1925. Anisops staii, Hale, Archiv for Zoologi utgivit av. K. Svenska Vetenskapsakademien,
vol. 17, no. 20, p. 17 (ecological note).

* The single male specimen at my disposal was minus the two hind pairs of legs.

320 The University Science Bulletin

1933. Anisops stali, Lunciblad, Arch, fiir Hydrob., Suppl., vol. XII, p. 146 (list of Indo-
australian species of this genus).

1934. Anisops stali, Hungerford, Bull. Brooklyn Ent. Soc, vol. XXIX, p. 69 (ecological
note).

Size. — Males, length 9.0 mm. -10.6 mm., greatest body width
2.5 mm. -2.8 mm.; females, length 8.5 mm. -10.2 mm., greatest body
width, 2.4 mm.-2.9 mm.

Shape. — Large fusiform species; greatest body width about four
tenths body length.

Color. — Brown form: General facies stramineous. Eyes brown.
Pronotum may be orange or tinged with orange. Hemelytra may be
hyaline and appear darker due to underlying dark dorsal body sur-
face. Gray form : General facies gray; eyes brown or gray. Hemel-
ytra hyaline and appear gray due to the black dorsal body surface.
Legs of both forms stramineous. Abdominal venter dark brown or
black with keel and segmental margins of the connexivum stramine-
ous.

Male structural characteristics. — Viewed from above, the out-
line of the head is rounded with the vertex extending beyond the
anterior margins of the eyes; greatest width of the head eight to
nine tenths the pronotal humeral width and three to four times the
anterior width of the vertex; synthlipsis wide, two thirds the anterior
width of the vertex; along the median longitudinal axis the head
is one half as long as the pronotum. Pronotum with its humeral
width twice its median length; lateral margins diverging and slightly
more than one half the median length; posterior margin convex,
medianly emarginate. Frons concavely depressed with apex ex-
tending slightly beyond the anterior margins of the eyes to form a
short cephalic horn; facial tubercle with a horseshoe-shaped pro-
truding area immediately anterior to the labrum, surrounded by a
deep depression. Labrum long, with its median length one and
three fourths its basal width; apex more or less accuminate. Ros-
tral prong (PI. LVII fig. 107) shorter than the third rostral segment;
apex more or less accuminate. Anterior margin of the third rostral
segment with long procumbent hairs on each side, extending to the
apex. Stridulatory comb ( PI. XXXVII fig. 9b ) with a highly vari-
able number of teeth, varying from fifteen to twenty-five. Chaeto-
taxy of male front leg as shown on Plate XXXVII. Base of the
middle tibia with a short inward projection which bears on its inner
surface a circular group of thickly-set, stout, even-length setae;

Brooks: The Genus Anisops 321

basal third of middle tibia with a strong concavity on anterior mar-
gin. The relative lengths of the parts of the legs are as follows:

lsl 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 65

Middle leg 100 84 34 23

Hind leg 100 83 33 30

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head eight to nine
tenths the pronotal humeral width and three to four times the an-
terior width of the vertex; synthlipsis wide, slightly more than one
half the anterior width of the vertex; along the median longitudinal
axis the head is almost one half the length of the pronotum. Pro-
notum with its humeral width almost twice its median length; lateral
margins diverging and slightly more than one half the median
length; posterior margin convex, medianly emarginate. The relative
lengths of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 125 48 29

Middle leg 100 83 36 22

Hind leg 100 77 35 30

Location of types. — Type material from Australia, in the Stock-
holm and Paris Museums.

Comparative notes. — This species along with Anisops pellucens
Gerstacker are two of the largest species within this genus. How-
ever, they are not readily confused with one another as the males of
A. pellucens lack the short cephalic horn, and the circular group of
basal setae on the middle tibiae as found on the males of A stali.

Data on distribution:
Australia

New South Wales, one male ( F. H. Snow Coll. )

Broken Hill, X-4-44, C. E. Chadwick, two males (Dept. of Agric, N. S. W.,
Austral. )

New South Wales, Coolebah, 1904, one male, one female (Dept. of Agrie.,
N. S. W., Austral.)

New South Wales, Hay, Lea, one male (Dept. of Agric., N. S. W., Austral.)

New South Wales, Brewarrina, June, 1944, W. W. Froggatt, one female
(Dept. of Agric., N. S. W., Austral.)

West Australia, Wiluna, Oct. 19, 1931, Harv. Austral. Expecl., P. J. Darling-
ton, seven males, four females ( Harv. Mus. Comp. Zool. )

Queensland, Brisbane, Dec. 1922, H. Hacker, one female ( F. H. Snow Coll. )

Queensland, Duaringa, two males, two females ( F. H. Snow Coll. )

21—3286

322 The University Science Bulletin

Cordilla Downs, one male, one female ( F. H. Snow Coll. )
Cordilla Downs, VII-16-'88, F. Archer, one female ( F. H. Snow Coll. )
Townesville, Jan. 1945, B. Malkin, one female (U. S. Nat. Mus. )
Australia, II-'47, Verreaux, one female ( F. H. Snow Coll. )

Timor

Soe, 1935, Buhler, five males, five females ( F. H. Snow Coll. ) ; twelve
males, fourteen females ( Basel Nat. Hist. Mus., Switzerland ) .

Soe, June, 1935, C. Buhler and Meyer, three males, one female (Basel Nat.
Hist. Mus., Switzerland).

Celebes

Lahendong, six females (Basel Nat. Hist. Mus., Switzerland).

Okinawa

Okinawa, Sept. 30, 1945, W. D. Fields, four males, one female ( U. S. Nat.
Mus. ) .

Philippine Islands
Mindanao, Baker, one female ( U. S. Nat. Mus.).

Anisops campbelli n. sp.

(PI. XXXVII, fig. S)

Size. — Males, length 7.3 mm. -8.1 mm., greatest body width 2.1
mm-2.2 mm.; females, length 8.1 mm. -8.4 mm., greatest body width
2.2 mm. -2. 5 mm.

Shape. — Fusiform, with greatest body width between one third
and one half the body length.

Color. — Broivn form: General facies stramineous. Eyes brown.
Vertex may be tinged with dark brown and hemelytra may be hya-
line at apex, appearing dark brown as it overlies the dark brown
dorsal surface of the abdomen. Legs stramineous. Abdominal ven-
ter black with keel and segmental margins of the connexivum stra-
mineous. Gray form: General facies gray; legs testaceous; abdomi-
nal venter dark brown with keel and segmental margins of the con-
nexivum gray.

Male structural characteristics. — As viewed from above, the out-
line of the head is rounded with the vertex extending slightly beyond
the anterior margin of the eyes; greatest width of head nine tenths
the pronotal humeral width and three to four times the anterior
width of the vertex; synthlipsis wide, slightly less than two thirds
the anterior width of the vertex; along the median longitudinal axis
the head is one half the pronotal length. Pronotum with a faint
median ridge on anterior half; humeral width at least twice its me-
dian length; lateral margins diverging and one half of the median
length; posterior margin convex, medianly concave. Viewed ven-

Brooks: The Genus Anisops 323

trally the frons is slightly concave with the facial tubercle flat; apex
delinited by a transverse ridge which projects slightly beyond the
margin of the head. Rostral prong (PI. XXXVII fig. 8b) shorter
than third rostral segment; apex accuminate. Third rostral segment
with a procumbent tuft of hairs along each side of the dorsal sur-
face. Labrum long, with median length one and one-fourth times
its basal width; apex truncate. Stridulatory comb (PI. XXXVII fig.
8c) of approximately eighteen teeth which increases in height from
base to apex. Anterior margin of innner surface of front tibia with
a short row of two pairs of stout setae. Chaetotaxy of the male front
leg as shown on Plate XXXVII. The relative lengths of the parts of
the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 133 60

Middle leg 100 81 54 21

Hind leg 100 83 35 30

Female structural characteristics. — Viewed from above, the an-
terior outline of the head is rounded with the anterior margin of the
vertex extending slightly beyond that of the eyes; greatest width of
the head slightly more than eight tenths the pronotal humeral width
and three to four times the anterior width of the vertex; synthlipsis
wide, two thirds the anterior width of the vertex; along the median
longitudinal axis the head is slightly more than one half the pronotal
length. Pronotum with a faint median ridge on the anterior half;
humeral width slightly less than twice the median length; lateral
margins diverging almost three fourths the median length; posterior
margin convex, medianly emarginate. The relative lengths of the
parts of the legs are as follows:

1st 2nd

Femur Tiliia Tar. Seg. Tar. Seg.

Fore leg 100 125 44 24

Middle leg 100 82 35 23

Hind leg 100 87 34 31

Location of types. — Male holotype, female allotype, three males,
one female paratype, Chikkaballapura, S. India, T. V. Campbell in
the Snow Entomological Collection. Other paratypes are as fol-
lows. In the Snow Entomological Collection: one male, three fe-
males, Lucknow, India, 1-22-08, Mus. Collr. R. H. In the Indian
Museum at Calcutta: Kandaghat, Simla Hills, 3500'-4600', Sta. 2,
Pond, VIII-'25, B. Chapra; one male, two females, Punjab, Salt
Range, Sta. 9, S. L. Hora; four males, three females, Darjiling Dist.
E. Himalayas, Nam Ting Pokri above Sitong nr. Mangpur 4800',
VII-2-18, S. Kemp; one male, Lucknow, II-4-08, Mus. Collr. R. H.;

324 The University Science Bulletin

five females, Lucknow, II-5-08, Mus. Collr. R. H.; one male, Luck-
now, 1-16-08, Mus. Collr. R. H.; two males, one female, Lucknow,
1-17-08, Mus. Collr. R. H.

Comparative notes. — Though somewhat smaller, this species ap-
pears closely related to Anisops stall Kirkaldy, as all the distinctive
features as borne by the latter species are also present on A. camp-
belli though much reduced in prominence. The two species may be
readily separated since the middle tibiae of the males of the latter
species lack the inward basal projection bearing the circular group
of thickly set, stout, even-length setae as found on A. stall.

Data on distribution:

Burma

S. Shan States, Yawaghee, Heho, 3800, III — 29-17, Gravely, one male, one
female (Bueno Coll. of F. H. Snow Coll.).

India

Darjeeling, Le Moult, one male, one female (Bueno Coll. of F. H. Snow
Coll.).

Kalka, base of W. Himalayas, V — 16-11, N. Annandale, two males, one
female ( F. H. Snow Coll.).

S. India, Bangalore, 3000, X — 16-10, Annandale, one female (F. H. Snow
Coll.).

Patiala State, Dhurampor, base of Simla Hills, VII — 21-11, one female
(F. H. Snow Coll.).

Anisops amaryllis Hutchinson

(PI. XXXVI, ;ig. G)

1928. Anisops amaryllis Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. I, text fig. 5.

1930. Anisops amaryllis, Hutchinson, Ann. Mag. Nat. Hist., vol. VI, Ser. 10, pp. 58-59
(ecological note).

1933. Anisops amaryllis, Jaczewski, Linn. Sci. Jour., Zool., vol. XXXVII, p. 345 (eco-
logical note).

Size — Males, length 5.4 mm. -6 mm., greatest body width 1.5 mm.-
1.8 mm.; females, length, 5.3 mm. -6.3 mm., greatest body width
1.5 mm. -1.9 mm.

Shape — Small, subfusiform species; lateral margins in anterior
half of body almost parallel, converging in posterior half.

Color — Light form: General facies testaceous. Eyes brown. Hem-
elytra may appear dark gray as it is hyaline and overlies the dark
brown or black dorsal body surface. Legs stramineous. Abdominal
venter dark brown or black with keel and segmental margins of
the connexivum stramineous. Dark form: General facies black.
Eyes dark brown. Vertex and pronotum testaceous. Scutellum
dark brown or black; lateral margins may be crimson. Hemelytra
hyaline and appear dark gray as they overlie the black dorsal body

Brooks: The Genus Anisops 325

surface; margins bordering the scutellum may be tinged with crim-
son. Leg stramineous. Abdominal venter with keel and segmental
margins of the connexivum stramineous.

Male structural characteristics — Viewed from above, the outline
of the head is rounded with the greatest width of the head nine
tenths the pronotal humeral width and almost six times the anterior
width of the vertex; synthlipsis wide, one third the anterior width
of the vertex; along the median longitudinal axis the head is slightly
shorter than the pronotum. Pronotum with its humeral width
slightly more than twice its median length; lateral margins diverg-
ing and one-half the median length; posterior margin convex, me-
dianly emarginate. Rostral prongs ( PI. XXXVI, fig. 6b ) as long as
the third rostral segment; apex more or less rounded. Stridulatory
comb (PI. XXXVI, fig. 6c) of approximately twelve even length
teeth. Chaetotaxy of the male front leg as shown on Plate XXXVI.
The relative lengths of the parts of the legs are as follows:0

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 117 73

Middle leg 100 77 38 25

Female structural characteristics — Viewed from above, the out-
line of the head is rounded; greatest width of the head eight-tenths
the pronotal humeral width and five times the anterior width of
the vertex; synthlipsis wide, slightly less than one-half the anterior
width of the vertex; along the median longitudinal axis the head is
slightly more than one-half the length of the pronotum. The rela-
tive lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 125 58 28

Middle leg 100 87 43 26

Hind leg 100 84 33 33

Location of types. — Male type from British East Africa (now
Kenya Colony) Nairowa Pass, C. S. Betton in the British Museum.

Comparative notes. — This species appears closely allied to
Anisops eros Hutchinson from which it can be separated by the
shape of the stridulatory comb and the chaetotaxies of the front
legs of the males. In the latter species the distal teeth of the
stridualtory comb are elongate, more than twice as long as the basal
elements, whereas in A. amaryllis all the teeth of the comb are
approximately the same length. The row of three small spines as

* None of the males at my disposal possessed a complete hind leg.

326 The University Science Bulletin

found on the basal inner surface of the male front tarsi of A. eros
is lacking on the tarsi of the male of A. amaryllis.

Data on distribution:
Africa

Central Africa, one male, (Basel Nat. Hist. Mus., Switzerland).

Uganda, Lake Naivasha, E. B. Worthington, exchange from the British
Museum, one male, one female (F. H. Snow Coll.).

South Rhodesia, Mazoe Valley, VI-10-28, A. Cuthberson, two males, one
female ( F. H. Snow Coll. ) .

Anisops elstoni n. sp.

(PI. XXXVI, fig. 5)

Size. — Males, length 4.3 mm. -4.6 mm., greatest body width 1.2
mm. -1.3 mm.; females, length 5.1 mm.-5.5mm., greatest body width
1.4 mm. -1.6 mm.

Shape. — Small, subfusiform species; anterior half of body with
lateral margin only slightly converging, subparellel, those of pos-
terior half converging.

Color. — General body color stramineous. Eyes brown. Scutellum
may be orange or tinged with orange with anterior margin black.
Hemelytra hyaline and may have margins adjacent to scutellum
orange. Legs stramineous. Abdominal venter dark brown or black
with keel and segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width nine tenths pronotal
humeral width, six times the anterior width of vertex; synthlipsis
wide, one half the anterior width of the vertex; along the median
longitudinal axis the head is at least three fourths the pronotal
length. Pronotum with a large median depression; humeral width
slightly more than twice the median length; lateral margins di-
verging and at least one half the median length; posterior margin
convex, medianly concave. Facial tubercle flat. Labrum with a
few short hairs; basal width equal to median length; apex rounded.
Rostral prong longer than third rostral segment. Stridulatory comb
(Pi. XXXVI, fig. 5c) composed of about thirteen teeth; apical two
greatly reduced in size. Chaetotaxy of the male front leg as shown
on Plate XXXVI, fig. 5a. The relative lengths of the parts of the
legs are as follows :

1st 2nd

Femur Tibia Tar. See. Tar. Ses.

Fore leg 100 115 77

Middle leg 100 79 48 24

Hind leg 100 79 31 33

Brooks: The Genus Anisops 327

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width nine tenths the pronotal
humeral width, five to six times the anterior width of the vertex;
synthlipsis wide, slightly more than one third the anterior width of
the vertex; along median longitudinal axis the head is slightly more
than three fourths the pronotal length. Pronotum with a faint
median depression; humeral width slightly more than twice its
median length; lateral margins diverging and one half the median
length. The relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 118 57 36

Middle leg 100 80 37 26

Hind leg 100 79 33 33

Location of types. — Male holotype, female allotype, three male
and one female paratypes, S. Australia, Myponga, A. H. Elston, in
the Snow Entomological Collection. Other paratypes are as fol-
lows. In the Snow Entomological Collection: one male, six females
Australia, Brisbane, Queensland, Dec. 1933, H. Hacker; one male,
Suifu Szecheun, China, D. C. Graham. In the Harvard Museum of
Comparative Zoology: three males, three females, New South Wales,
The Dorrigo, 3000', Feb. 1932, Darlington, Harv. Austrl. Exped.

Comparative notes. — A very small species of the same general
appearance as Anisops exigera Horvath. However the males of
this species have a much narrower synthlipsis, one fifth to one sixth
the anterior width of the vertex, whereas in A. elstoni the synthlipsis
is wide, at least one third the anterior width of the vertex. The
rostral prong of A. exigera is shorter than the third rostral segment
and rounded at the apex while that of A. elstoni is longer than the
third rostral segment and accuminate at the apex.

Data on distribution. — Known only from type series.

Anisops eros Hutchinson*

(PI. XXXVII, fig. 10)

1928. Anisops eros Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. I, pp. 160-161, text
fig. 4.

1933. Anisops eros. Jaczewski, Linn. Soc. Jour. Zool., Vol. XXXVIII, p. 345 (ecological
note).

1933. Anisops eros, Hutchinson, Internat. ges. Hydrob. nnd Hydrog., vol. 28, pt. 5/6,
p. 462 (ecological note).

1937. Anisops eros, Poisson, Ann. Soc. Ent. France, vol. CVI, p. 117, text fig. 2.

1948. Anisops eros. Poisson, Rev. Francaise Ent., vol. XV, p. 168 (ecological note).

Size. — Male, length 4.8 mm., greatest body width 1.2 mm.
Shape. — Small fusiform species; greatest width midway the body
length.

* Only a single male specimen of this species was in my material.

328 The University Science Bulletin

Color. — General facies testaceous. Eyes gray brown. Legs
stramineous. Abdominal venter dark brown with keel and segmen-
tal margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the anterior margin almost straight;
greatest width of the head nine tenths the pronotal humeral width
and slightly more than six times the anterior width of the vertex;
synthlipsis wide, slightly more than one half the anterior width of
the vertex; along the medial longitudinal axis the head is almost as
long as the pronotum. Pronotum with its humeral width slightly
more than twice its median length; lateral margins diverging and
almost one half the median length; posterior margin convex, me-
dianly emarginate. Facial tubercle only slightly raised. Labrum
with its median length slightly less than its basal width; apex
rounded. Rostral prong (Pi. XXXVII, fig. 10b) longer than the
third rostral segment; apex more or less rounded. Stridulatory
comb (PI. XXXVII, fig. 10c) of approximately fourteen teeth which
increase in length toward the middle. Chaetotaxy of the front leg
as shown on Plate XXXVII. The relative lengths of the parts of the
legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 127 91

Middle leg 100 80 40 25

Hind leg 100 80 33 33

Location of types. — Type material from Uganda, Kampala, III-
20-1927, G. L. R. Hancock, in the British Museum.

Comparative notes. — This species is very similar to Anisops
amaryUis Hutchinson from which it can be successfully separated
by the fact that the males of A. eros have a short row of three small
setae on the inner surface of the basal half of the fore tarsi, which is
lacking on the fore tarsus of A. amaryUis.

Data on distribution:
Africa

Tanganyika Terr., Amani, Feb. 19, 1940, F. X. Williams, one male ( Harv.
Mus. Comp. Zool. ) .

Anisops kampalensis Hutchinson*

(PI. XXXVIII. fig. 15)

1928. A?iisops kampalensis Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. I, pp. 162-
163, text fig. 6.

1933. Anisops kampalensis, Hutchinson, Internationale der gesamten Hydrobiologie und
Hydrographie, vol. 28, prt. 5/6, p. 462 (ecological note).

Size. — Males, length 4.9 mm., greatest body width 1.3 mm.

* Only two male specimens were available for study.

Brooks: The Genus Anisops 329

Shape. — Small, robust, subfusiform species; greatest body width
midway the body length.

Color. — General facies black. Eyes brown. Vertex, pronotum
testaceous, the latter with irregular hyaline areas on posterior mar-
gin, such areas appear black due to the color of the underlying
scutellum. Scutellum testaceous with anterior half black. Hemely-
tra hyaline and appear black due to the black color of the dorsal
body surface. Legs stramineous. Abdominal venter black with
keel and segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head nine tenths the
pronotal humeral width and five times the anterior width of the
vertex; synthlipsis wide, one third the anterior width of the vertex;
along the median longitudinal axis the head is one half the length
of the pronotum. Pronotum with its humeral width almost twice the
median length; lateral margins diverging and one half the median
length; posterior margin convex, medianly emarginate. Facial
tubercle slightly raised. Labrum broad, basal width equal to the
median length; apex rounded. Rostral prong (PL XXXVIII, fig.
15b ) as long as the third rostral segment; apex accumulate. Stridu-
latory comb (Pi. XXXVIII, fig. 15c) of approximately eleven teeth,
the apical four longer than the basal eight. The relative lengths of
the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. S>eg.

Fore leg 100 117 88

Middle leg 100 82 37 27

Hind leg 100 82 30 30

Location of types. — Type material from Kampala, Uganda in the
British Museum.

Comparative notes. — This species is very similar to A. adonis
Hutchinson, however, the males lack the carinate frons of the latter
species.

Data on distribution:
Africa

Huat Owanhui, Bessou (Mission) Amont de Fort de Passel, Ouacha, 1804,
Dr. J. DeCorse, one male (F. H. Snow Coll.).

Mission Tilho, Ouacha, 1910, Dr. R. Gaillard, one male (Paris Museum).

Anisops paracrine ta n. sp.

(PI. XXXVIII, fig. 12)

Size. — Males, length 4.8 mm., greatest body width 1.3 mm.; fe-
males, length 4.8 mm., greatest body width 1.3 mm.

330 The University Science Bulletin

Shape. — Small, fusiform species; greatest body width about mid-
way its body length.

Color. — General facies brown. Eyes brown. Vertex, pronotum
and scutellum testaceous, the latter dark brown on apical half.
Hemelytra hyaline and appearing brown due to the underlying
brown dorsal body surface. Legs stramineous. Abdominal venter
dark brown with keel and segmental margins of the connexivum
stramineous.

Male structural characteristics. — Viewed from above, the out-
line of the head is rounded with the greatest width eight tenths the
pronotal humeral width and slightly more than six times the anterior
width of the vertex; along the median longitudinal axis the head is
almost as long as the pronotum. Pronotum with its humeral width
slightly more than two and one half times its median length; lateral
margins diverging and three fifths the median length; posterior mar-
gin convex, medianly emarginate. Facial tubercle with a triangular
patch for long hairs whose outer ones are erect. Labrum covered
with long hairs; basal width slightly more than its median length;
apex rounded. Rostral prong (PI. XXXVIII, fig. 12b) shorter than
the third rostral segment; apex accuminate. Stridulatory comb ( Pi.
XXXVIII, fig. 12c) of eleven even-length teeth. Chaetotaxy of the
front leg as shown on Plate XXXVIII. The relative lengths of the
parts of the legs are as follows.

1st 2nd

Femur Tibia Tar. Seg. Tar. .Seg.

Fore leg 100 120 90

Middle leg 100 76 40 26

Hind leg 100 71 33 33

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with its greatest width almost nine-
tenths the pronotal humeral width and slightly more than four times
the anterior width of the vertex; synthlipsis wide almost one half the
anterior width of the vertex; along the median longitudinal axis the
head is more than one half the median length. Pronotum with its
humeral width two and one fourth times the median length; lateral
margins diverging and slightly more than one half the median
length; posterior margin convex, medianly emarginate. The relative
lengths of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 126 60 40

Middle leg 100 81 40 25

Hind leg 100 81 33 33

Brooks: The Genus Anisops 331

Location of types. — Male holotype Queensland, Australia, Coen.
C. York, May '32, Harv. Austrl. Exped. Darlington, in the Harvard
Museum of Comparative Zoology. Female allotype, one male and
one female paratype, Moore Id. Australia, Feb. 20, 1945, B. Milkin
in the United States National Museum.

Comparative notes. — Very similar to Anisops crinita n. sp. but
the two species may be readily separated on the basis of the males.
The male of A. paracrinita has long hairs on the facial tubercle and
for the most part these are erect, whereas the facial tubercle of A.
crinita has short procumbent hairs. Also the synthlipsis of the latter
is much narrower being only at most one fifth the anterior width of
the vertex. For comparison of the front legs of the males see PL
XXXVIII, fig. 12a and PL I, fig. 68a.

Data on distribution. — Known only from type series.

Anisops hacker i n. sp.

(PI. XXXVIII. fig. 13)

Size — Males, length, 5.5 mm. -5.7 mm., greatest body width 1.4
mm. -1.6 mm.; females, length 5.7 mm.-6.0 mm., greatest body width
1.5 mm. -1.7 mm.

Shape — Short, robust, subfusiform species; lateral margins of
anterior half of body almost parallel.

Color — Eyes brown; vertex and pronotum testaceous, the former
may be tinged with orange, the latter may have its posterior margin
hyaline and appear the black color of the underlying surface of the
scutellum. Scutellum orange, black along its apical margin. Hem-
elytra hyaline and appearing gray as it overlies the black dorsal
body surface. Legs stramineous. Abdominal venter black with
keel and segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of head nine tenths the
pronotal humeral width; five times the anterior width of the vertex;
synthlipsis wide, slightly less than one half the anterior width of
the vertex; along the median longitudinal axis the head is three
fifths the pronotal length. Pronotum with its humeral width slightly
more than twice its median length; lateral margins diverging and
three fifths the median length; posterior margin convex, slight
median emargination. Facial tubercle simple only slightly raised.
Labrum with its basal width slightly more than its median length;
apex rounded. Rostral prong (PL XXXVIII, fig. 13b) as long as
the third rostral segment; apex accuminate. Stridulatory comb

332 The University Science Bulletin

(PL XXXVIII, fig. 13c) of approximately twenty-four teeth. Chaeto-
taxy of the male front leg as shown on Plate XXXVIII. The rela-
tive lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 105 79

Middle leg 100 86 38 25

--&

Unfortunately neither of the two male specimens at my disposal
possess complete hind legs.

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest head width nine tenths the
pronotal humeral width; five times the anterior width of the vertex;
synthlipsis wide, one-half the anterior width of the vertex; along the
median longitudinal axis the head is three fifths as long as the
pronotum. Pronotum with its humeral width slightly more than
twice its median length, lateral margins diverging and slightly
more than one half the median length; posterior margin convex,
medianly emarginate. The relative lengths of the parts of the legs
are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 123 61 31

Middle leg 100 82 41 27

Hind leg 100 84 34 33

Location of types. — Male holotype, female allotype, one male,
two female paratypes, Brisbane, Australia, Dec. 1932, H. Hacker,
in the Snow Entomological Museum.

Comparative notes. — This species appears closely related to Ani-
sops barrenensis n. sp., only slightly larger and about the same
shape. However, the males of A. hacked lack the large swollen
front femur as found on A. barrenensis, and the second rostral seg-
ment does not extend almost the length of the third rostral segment
as in the latter species.

Data on distribution. — Known only from type series.

Anisops hyperion Kirkaldy

(PI. XXXVIII, fig. 1C>)

1898. Anisops hyperion Kirkaldy, Wiener Ent. Zeit., vol. XVII, p. 141.
1904. Anisops hyperion, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. Ill, 132.
1914. Anisops hyperion. Distant, Nova Caledonia, Zoologie, vol. I, p. 386 (ecological note).
1923. Anisops hyperion, Hale, South Australian Naturalist, vol. IV, no. 3, pp. 124-128,
text fig. 1 (determined in error; actually Anisops thienemanni Lundblad).

1923. Anisops hyperion, Hale, Rec. South Australian Mus., vol. II, no. 3, pp. 403-412,
text fig. 365 (determined in error, actually A?iisops thienemanni Lundblad).

1924. Anisops hyperion. Hale, South Australian Nat., vol. V, no. 4, p. 135 (determined in
error; actually Anisops thienemanni Lundblad).

Brooks: The Genus Anisops 333

1926. Anisops hyperion, Esaki, Ann. Mus. Nat. Hungarici, vol. XXIV, p. 188 (eco-
logical data; determination doubtful).

1930. Anisops hyperion, Hutchinson, Proc. Zool. Soc. London, vol. XXIX, p. 145, text
fig. 2 (determined in error; actually Antisops thiene?nanni Lund.).

1933. Anisops hyperion, Lundblad, Archiv. fiir Hydrobiologie Suppl., vol. XII, p. 145 (a
list of the Indo-australian and Pacific forms of this genus).

Size. — Males, length 6.0 mm.-6.5 mm., greatest width of body
1.5 mm.-1.6 mm.; females, length 6.0 mm. -6.6 mm., greatest body
width 1.5 mm.-1.8 mm.

Shape. — Fusiform species; greatest body width midway the length
of the body.

Color. — General facies stramineous. Eyes brown. Scutellum
may be red orange or only have its margins tinged with orange.
Legs stramineous. Abonminal venter dark brown with keel and
segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the out-
line of the head is rounded with the greatest width of the head al-
most nine tenths the pronotal humeral width and five to six times
the anterior width of the vertex; synthlipsis wide, one third the an-
terior width of the vertex; along the median longitudinal axis the
head is nine tenth the length of the pronotum. Pronotum with its
humeral width slightly more than twice its median length; lateral
margins diverging and a slightly more than one half the median
length; posterior margin convex, medianly emarginate. Facial
tubercle slightly raised. Labrum with its basal width slightly more
than the median length; apex rounded; surface of the labrum pro-
vided with a few scattered hairs. Rostral prong (Pi. XXXVIII, fig.
16b) almost as long as the third rostral segment; apex accuminate.
Chaetotaxy of the male front leg as shown on Plate XXXVIII. The
relative lengths of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 108 76

Middle leg 100 86 41 27

Hind leg 100 85 35 35

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head almost nine
tenths the pronotal humeral width and five and one half times the
anterior width of the vertex; synthlipsis wide, almost one half the
anterior width of the vertex; along the median longitudinal axis the
head is two thirds as long as the pronotum with its humeral width
at least twice its median length; lateral margins diverging and at
least one half the median length; posterior margin convex, medianly

Tibia

1st
Tar. Seg.

2nd
Tar. Seg.

112

83

51
39

36
26

334 The University Science Bulletin

emarginate. The relative lengths of the parts of the legs are as
follows : *

Femur

Fore leg 100

Middle leg 100

Location of types. — Kirkaldy's type material consisting of two
females from Rockhampton, Queensland, Australia in the Hamburg
Museum. In the Kirkaldy Collection of the Snow Entomological
Collection is a male specimen bearing in Kirkaldy's handwriting the
label "hyperion." I am designating this specimen the male allotype.
It is from Victoria, Alexandria in Australia.

Comparative notes. — Kirkaldy mentioned in 1904 that possibly
he had two species mixed. Such was true, for collected along with
A. hyperion was a new species which I have named A. cleanei.
These two species are superficially very similar; can best be separ-
ated on the character of the male rostral prong, which in A. deanei
has it dorsal margin extending almost to the apex of the third rostral
prong whereas that of A. hyperion extends only slightly beyond
the basal half of the third rostral segment. Also the apex of the
third rostral segment of A. deanei is considerably wider than the
base of the fourth, whereas in the case of A. hyperion the two are
approximately the same width.

Data on distribution:
Australia

Victoria, Alexandria, F. L. Billinghurst, six males, three females, Kirkaldy
Collection ( F. H. Snow Coll. )

Anisops tuberculata Poisson

(PI. XXXVIII, fig. 11; PI. LVII, fig. 104)

1928. Ar}isops tuberculata Poisson, Bull. Soc. Ent. France, vol. 33, p. 74, text fig. 17.

1929. Anisops tuberculata, Poisson, Faune des Colonies Francaises, vol. Ill, pp. 150-152,
text figs. 17, 18, 19.

1948. Anisops tuberculata, Poisson, Rev. Francaise Ent., vol. XV, pp. 167-177, (ecological
note).

Size. — Males, length 5.5 mm. 6.0 mm., greatest body width 1.6
mm. -1.7 mm.; females, length 5.4 mm. -6.6 mm., greatest body width
1.5 mm. -1.7 mm.

Shape. — Subfusiform species; greatest body width about midway
the body length.

Color. — General facies red brown. Hemelytra with hyaline areas
which appear black due to the underlying color of the dorsal body

* Unfortunately none of the female specimens at my disposal possessed a complete hind leg.

Brooks: The Genus Anisops

335

surface. Legs red brown. Abdominal venter dark brown with keel
and segmental margins of the connexivum red brown.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of the head almost equal to
the pronotal humeral width and five to six times the anterior width
of the vertex; synthlipsis wide, one third the anterior width of the
vertex; along the median longitudinal axis the head is almost two
thirds the length of the pronotum. Pronotum with its humeral
width twice the median length; lateral margins diverging and two
thirds the median length; posterior margin convex, medianly emar-
ginate; anterior margin between the margins of the eyes bearing a
large tubercle. Facial tubercle slightly raised. Labrum with its
basal width one and one half the median length; apex rounded.
Rostral prong (PI. LVII, fig. 104) longer than the third rostral seg-
ment; apex rounded. Stridulatory comb (PL XXXVIII, fig. lib)
highly irregular, apical two teeth large and spatulate. Chaetotaxy
of the male front leg as shown on Plate XXXVIII. The relative
lengths of the parts of the legs are as follows:

1st 2nd

Femur Til >ia Tar. Seg. Tar. Seg.

Fore leg 100 105 64

Middle leg 100 85 40 37

Hind leg 100 86 33 33

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head nine tenths the
pronotal humeral width and between five and six times the anterior
width of the vertex; synthlipsis wide, one third the anterior width of
the vertex; along the median longitudinal axis the head is slightly
more than one half the length of the pronotum. Pronotum with its
humeral width twice its median length; lateral margins diverging
and slightly more than one half the median length; posterior margin
convex, medianly emarginate. The relative lengths of the parts of
the legs are as follows:

1st. 2nd

Femur Tibia Tar. Seg, Tar. Seg.

Foreleg 100 115 51 31

Middle leg 100 85 40 25

Hind leg 100 86 34 34

Location of types. — Type material from Cameron in the Paris
Museum.

Comparative notes. — This species, by its large tubercle on the
anterior margin of the male pronotum is not readily confused with
any other of the genus. However, in general body shape it is similar
to A. debilis, though it is slightly larger.

336

The University Science Bulletin

Data on distribution:
Africa

French Congo, Lake Onango, F. C. Good, Holland Collection, exchange
from Carneige Museum, nine males, six females ( F. H. Snow Coll.).

Cameron, W. Africa, Samgmelina, IV-16th to 19th-32 five males, one female
(F. H. Snow Coll.).

Congo Francaise, N'Gomo, Bas Ogoque, 1906, E. Haug, one male, one
female ( Paris Museum ) .

Congo, Ogaque, Sam Kita, 1910, R. Ellenberger, one female (Paris Museum).

Anisops nodulata n. sp.

(PI. XXXVIII, iig. 14)

Size. — Males, length, 4.6 mm.-4.8 mm., greatest body width 1.2
mm. -1.3 mm.; females, length, 4.8 mm. -5. 4 mm., greatest body width
1.3 mm. -1.6 mm.

Shape. — Small fusiform species, with greatest width almost mid-
way the body length.

Color. — General facies stramineous. Eyes brown. Hemelytra
may be hyaline and such areas appear black as they overlie the black
dorsal body surface. Legs stramineous. Abdominal venter black
with keel and segmental margins of the connexivum stramineous.

Male structural characteristics. — View from above, the outline of
the head is rounded with its greatest width eight to nine tenths the
pronotal humeral width, five times the anterior width of the vertex;
synthlipsis wide, slightly more than one third the anterior width of
the vertex; along the median longitudinal axis the head is three
fourths the pronotal length. Pronotum with its humeral width
slightly more than twice its median length; lateral margins diverging
and one half the median length; posterior margin convex, medianly
emarginate. Facial tubercle flattened and slightly concave. Lab-
rum short; basal width one and two thirds its median length; apex
rounded. Rostral prong longer than third rostral segment which is
medianly carinate on anterior surface, carina with two nodules (Pi.
XXXVIII, fig. 14b ) . Anterior femur with dorsal and ventral margins
almost parallel; apex rounded (Pi. XXXVIII, fig. 14a). Stridulatory
comb (PI. XXXVIII, fig. 14c) of approximately fifteen teeth, longest
ones midway between base and apex. Chaetotaxy of male front leg
as shown on Plate XXXVIII. The relative lengths of the parts of
the legs are as follows:

Femur

Fore leg 100

Middle leg 100

Hind leg 100

Tibia

108
87
91

1st

2nd

Tar. Seg.

Tar. Se;

80

, ,

40

25

32

28

Brooks: The Genus Anisofs 337

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width nine tenths the pronotal
humeral width, four to four and one half times the anterior width of
the vertex; synthlipsis wide, almost one half the anterior width of
the vertex; along the median longitudinal axis the head is slightly
more than one half the pronotal length. Pronotum with humeral
width slightly more than twice the median length; lateral margins'
diverging and slightly more than one half the median length; pos-
terior margin convex, medianly emarginate. The relative lengths
of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 50 33

Middle leg 100 84 36 29

Hind leg 100 87 30 30

Location of types. — Male holotype, female allotype, four male
paratypes, Philippine Islands, Pangasinan, rice fields, 11-19-34, Z.
Abalos in the Snow Entomological Collection. Other paratypes are
as follows. In the Snow Entomological Collection: two females,
Philippine Isl. Dagupan Pangasinan, Sept. 2, 1936, Roman Abalos;
two females, Philippine Isl., Bagnio, N. T. Prov. Apr. 26, 1936,
Roman Abalos; two males, four females, New Guinea, Rigo, 1889,
L. Loria.

Comparative notes. — This species is about the same size as A. ex-
igera Horvath, from which it differs in having a much wider synth-
lipsis, front femur of the males rounded and a nodulate carina on
the anterior surface of the third rostral segment.

Data on distribution:
Australia

New Queensland, Palm Is. G. F. Hill, one male (Bueno Coll. of F. H. Snow
Coll).

Anisops apicalis Stal

(PI. XXXIX, fig. 17)

1855. Anisops apicallis Stal, Ofversigt at Kongl. Vetenskaps-Akademines Fordhandlinger.
vol. XII, p. 89.

1865. Anisops apicalis, Stal, Hemiptera Africana, vol. Ill, p. 192.

1904. Anisops apicalis, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 117-118, 132.

1926. Anisops apicalis, Jaczewski, Annalibus Zoologicis Musei Polonici Historiae Naturalis,
vol. V, No. 2, p. 91, text figs. 50, 51, 52, 53.

1929. Anisops apicalis, Hutchinson, Ann. South African Mus., vol. XXV, pp. 406-408,
PI. XXIX, fig. 5; Plate XXX, fig. 21.

1930. Anisops apicalis, Hutchinson, Proc. Zool. Soc. London, vol. XXIX, pt. 2, p. 450
(ecological note).

1930. Anisops apicalis, Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. VI, p. 59 (eco-
logical note).

22—3286

338 The University Science Bulletin

1933. A7iisops apicalis, Jaczewski, Linn. Sci. Jour. Zool., vol. XXXVII, p. 345 (ecological
note).

1933. A7iisops apicalis, Hutchinson, Internat. Rev. ges. Hydiob. Hydrog., vol. 28. pt.
5/6, p. 440.

1948. Anisops apicalis, Poisson, Mem. Inst. Scient. Madagascar, Ser. X, vol. I, p. 108
(ecological note).

Size. — Males, length 4.8 mm., greatest body width 1.4 mm.; fe-
males, length 4.5 mm., greatest body width 1.6 mm.

Shape. — Small, robust, fusiform species with greatest body width
slightly before the middle of the body.

Color. — General facies stramineous. Eyes dark brown. Hemely-
tra with a subapical transverse brown band. Legs stramineous.
Abdominal venter dark brown with keel and segmental margins of
the connexivum stramineous.

Male structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of the head equal to the
pronotal humeral width and five and one half times the anterior
width of the vertex; synthlipsis wide, slightly more than one third
the anterior width of the vertex; along the median longitudinal axis
the head is slightly more than two thirds the length of the pronotum.
Pronotum with its humeral width almost twice its median length;
lateral margins almost parallel only slightly diverging, three fifths
the median length; posterior margin almost straight. Scutellum
greatly reduced; basal width one and three fourths the median
length. Wings greatly reduced and appear nonfunctional. Facial
tubercle flat with lateral margins posterior to the eyes converging.
Labrum narrow and long; median length one and one third its basal
width; apex rounded. Rostral prong (PI. XXXIX, fig. 17b) equal
to the length of the third rostral segment; apex more or less rounded.
Anterior leg (Pi. XXXIX, fig. 17a) with dorsal and ventral femoral
margins almost parallel, apex truncate; stridulatory comb (PI.
XXXIX, fig. 17c ) of approximately nineteen long slender teeth which
decrease in length at the apex. Chaetotaxy of the male front leg
as shown on Plate XXXIX. The relative lengths of the parts of the
legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 119 79

Middle leg 100 81 33 28

Hind leg 100 95 36 35

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head slightly less than
the humeral width of the pronotum and almost five and one half
times the anterior width of the vertex; synthlipsis wide, slightly
more than one third the anterior width of the vertex; along the

Brooks: The Genus Anisops 339

median longitudinal axis the head is as long as the pronotum. Pro-
notum short with its humeral width slightly more than three times
its median length; lateral margins only slightly diverging and two
thirds the median length; posterior margin straight. The relative
lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Foreleg 100 112 55 36

Middle leg 100 87 39 28

Hind leg 100 97 25 25

Location of types. — Type material from Caffraria in the Stockholm
Museum.

Comparative notes. — This species appears very similar to Anisops
hungerfordi Poisson from which it can be separated on the basis of
the scutellum which is greatly reduced in A. apicalis, basal width
one and three fourths its median length; whereas in A. hungerfordi
the basal width is only one and two sevenths the median length.
Also the wings of the latter species are not brachypterous.

Data on distribution:
Africa

Ugando, Entebbe, VIII-4-1929, G. H. E. Hopkins, gift to H. B. Hungerford
from G. E. Hutchinson, one male, one female (F. H. Snow Coll.).

Afrique Orient. Angl., Zanzibar, Riviera Mivera, 1904, Ch. Alluaud, one
male (Paris Museum).

Anisops caneriensis perplexa Poisson

(PI. XXXIX, fig. 20)

1929. Anisops perplexa Poisson, Bull. Soc. Hist. Nat. Afrique du Nord, vol. XX, pp. 87-
89, figs. 1, 2, 3, 113.

1933. Anisops perplexa, Hutchinson, Internal, ges. Hydrob. und Hydrog., vol. 28, pt. 5/6,
p 452 (ecological note).

1934. Anisops perplexa, Poisson, Bull. Soc. Hist. Nat. Afrique du Nord, vol. XXV, pp.
135-137.

1948. Anisops canariensis perplexa, Poisson, Inst. Rech. Sahar. Univ. D'Alger, p. 5. text
figs. 14, a, c. (Places A. perplexa as a subspecies of A. canariensis Noualhier.)

Size. — Males, 6.0 mm., greatest body width 1.5 mm.; females,
length 5.9 mm. -6.3 mm., greatest body width 1.5 mm.-1.6 mm.

Shape. — Slightly fusiform species; greatest body width about
midway the length of the body.

Color. — General facies stramineous. Abdominal venter dark
brown with keel and segmental margins of the connexivum stram-
ineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the anterior margin almost straight;
greatest width of the head almost equal to the pronotal humeral
width and almost six times the anterior width of the vertex; synth-

340

The University Science Bulletin

lipsis wide, slightly more than one third the anterior width of the
vertex; along the median longitudinal axis the head is eight tenths
the pronotal length. Pronotum with its humeral width almost twice
the median length; lateral margins diverging and slightly more than
one half the median length; posterior margin convex, medianly
concave. Facial tubercle slightly raised. Labrum short, basal width
one and one half the median length. Rostral prong (PL XXXIX,
fig. 20b) longer than the third rostral segment; apex accumulate.
Stridulatory comb (PL XXXIX, fig. 20c) of approximately twenty-
three teeth which decrease slightly in length from base to apex.
Chaetotaxy of the male front leg as shown in Plate XXXIX. The
relative lengths of the parts of the legs are as follows : *

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 115 78

Middle leg 100 86 38 25

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of the head nine tenths
the pronotal humeral width and slightly more than five times the
anterior width of the vertex; synthlipsis wide, one half the anterior
width of vertex; along the median longitudinal axis the head is three
fifths the length of the pronotum. Pronotum with its humeral width
slightly more than twice the median length; lateral margins of the
pronotum diverging and three fifths the median length; posterior
margin convex, medianly emarginate. The relative lengths of the
parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 121 54 31

Middle leg 100 78 37 22

Hind leg 100 78 28 28

Location of types. — Type material from Algeria is in Poisson's
Collection.

Comparative notes. — This subspecies differs from A. canariensis
canadensis by the fact that the anterior tarsi of the males have a
median row of four small setae in the basal half of the inner sur-
face. It is remarkably similar to A. debilis Gerst. and differs from
it primarily by the width of the synthlipsis which is one third the
anterior width of the vertex instead of one sixth the anterior width
of the vertex as in A. debilis.

Hind legs on two male specimens in my material missing.

Brooks: The Genus Anisops 341

Data on distribution:
Africa

Algerie, S. Culomb-Baehae, 1912; P. Germain, one male, one female in the
F. H. Snow Collection, fourteen males in the Paris Museum.

Tripolitaine, Marais de Tadjouran, Pres de Tripoli, 1901, Ch. Alluaud, one
male (Paris Museum).

Anisops edepol Kirkaldy

(PI. XXXIX, fig. 19; PI. LIV, fig. 94)

1899. Anisops edepol Kirkaldy, Ann. Soc. Ent. France, vol. LXVIII, p. 107.

1904. Anisops edepol, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 119, 132. Referring
also to this species :

1899. Anisops erebus, Kirkaldy, Ann. Soc. Ent. France, vol. LXVIII, pp. 107-108.

1904. Anisops erebus, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 119-132.

1948. Anisops erebus, Poisson, Mem. Inst. Sci. Madagascar, Ser. A, vol. I, text fig. 18,
p. 109. (Poisson mentions that the male of the species has not been described.)

1906. Anisops Kirkaldyanus, Bergroth, Wiener Ent. Zeit., vol. XXV, pp. 112 (Bergroth
proposes the name Kirkaldyanus for edepol).

Size. — Males, length 4.5 mm.-4.8 mm., greatest body width 1.6
mm. -1.8 mm.; females, length 5 mm.-5.5 mm., greatest body width
1.8 mm.-2.1 mm.

Shape. — Small, robust fusiform species, greatest body width
about two fifths the body length.

Color. — Brown form: General facies stramineous. Eyes brown.
Legs stramineous. Abdominal venter dark brown with keel and
segmental margins stramineous. Dark form: General facies black.
Eyes brown. Vertex, pronotum flavus. Hemelytra hyaline and ap-
pear black due to the underlying black body surface. Legs
stramineous. Abdominal venter black with keel and segmental
margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the out-
line of the head is rounded with its greatest width nine tenths the
pronotal humeral width and six times the anterior width of the
vertex; synthlipsis wide, slightly more than one half the anterior
width of the vertex; along the median longitudinal axis the head
is only slightly shorter than the length of the pronotum. Pronotum
with its humeral width over twice its median length; lateral mar-
gins only slightly diverging and over one half the median length;
posterior margin straight. Facial tubercle swollen and bearing a
median, short, ventrally directed spur. Labrum very short; basal
width one and one half the median length; apex rounded. Rostral
prong (PI. LIV, fig. 94) short and narrow; shorter than third
rostral segment; apex rounded. Stridulatory comb (Pi. XXXIX,

342

The University Science Bulletin

fig. 19b) of approximately seven teeth which increase in length
from base to apex. Chaetotaxy of the male front leg as shown on
Plate XXXIX. The relative lengths of the parts of the legs are as
follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 135 85

Middle leg 100 85 35 30

Hind leg 100 90 25 25

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head eight tenths the
pronotal humeral width and five times the anterior width of the
vertex; synthlipsis wide, slightly more than one half the anterior
width of the vertex; along the median longitudinal axis the head is
slightly more than half the length of the pronotum. Pronotum with
its humeral width slightly more than twice the median length; lat-
eral margins only slightly diverging and slightly more than one half
the median length; posterior margin straight. The relative lengths
of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 131 55 42

Middle leg 100 85 40 30

Hind leg 100 93 30 30

Location of types. — Part of the type material from Madagascar
in the Paris Museum. One male and one female cotype (exchange
from Paris Museum) in the F. H. Snow Collection. The female
type of A. erebus Kirkaldy in the F. H. Snow Collection.

Comparative notes. — The robustness and small size of this species
makes it appear superficially similar to A. apicalis Stal. However
the males of A. eclepol lack the apically enlarged front femur with
its truncate apex as found on the males of A. apicalis. Also the
male facial tubercle of the latter species is not swollen and bearing
the short projection as on A. edepol. The wings of A. edepol are
not brachypterous.

Remarks: I was fortunate in having at my disposal the female
type of A. erebus and a male and female cotype of A. edepol. Ex-
amination of the two groups showed immediately that they were
identical. Therefore I am placing A. erebus as a synonym of
A. edepol as the latter has page priority.

Data on distribution:

Madagascar

Madagascar, P. Camboue, one male, one female (eo-types) (F. H. Snow
Coll.).

Tananarive, 1921, R. Decary, one male, one female (Paris Museum).

Brooks: The Genus Anisops

343

Region du Sud-est, Vallee du Fanjakira Isaka, 1901, Ch. Alluaud, one male,
one female (Paris Museum).

Maroantsetra, purchased fr. Dr. O. Standinger, two females (F. H. Snow
Coll.).

Diego-Suarez, 1893, Ch. Alluaud, one female (type of A. erebus) ( F. H.
Snow Coll. ) .

Anisops timorensis n. sp.*

(PI. XXXIX, fig. 18)

Size. — Male, length 5.4 mm., greatest body width 1.5 mm.

Shape. — Small, subfusiform species, lateral margins in posterior
half of body converging, those of anterior half almost parallel.

Color. — General facies gray. Eyes gray brown. Vertex, pronotum,
and scutellum testaceous; pronotum mostly hyaline and appearing
black as it overlies the black anterior margin of the scutellum. Hem-
elvtra, with claval margins tinged with crimson; remainder hyaline
and appearing dark gray as it overlies the black dorsal body sur-
face. Legs stramineous. Abdominal venter black with keel and
segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with anterior margin almost straight; great-
est width of head nine tenths the pronotal humeral width, slightly
more than five times the anterior width of the vertex; synthlipsis
wide, one half the anterior width of the vertex; along the median
longitudinal axis the head is three fifths the pronotal length. Pro-
notum with its humeral width slightly more than twice its median
length; lateral margins diverging and three fifths the median length;
posterior margin convex, medianly emarginate. Facial tubercle only
slightly raised. Labrum moderately long, median length equal to
its basal width; apex rounded. Rostral prong ( PI. XXXIX, fig. 18b )
with apex accuminate. Basal half of third rostral segment covered
with an extension of the second rostral segment along its posterior
margin. Front femur (PI. XXXIX, fig. 18a) with apex rounded.
Stridulatory comb (PL XXXIX, fig. 18c) of approximately twenty-
four teeth. Chaetotaxy of the male front leg as shown on Plate
XXXIX. The relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tii>ia Tar. Seg. Tar. Seg.

Fore leg 100 111 83

Middle leg 100 77 45 23

Hind leg 100 83 35 35

Location of types. — Male holotype, Baguia, Timor, in the Basel
Naturhistorisches Museum in Switzerland.

Comparative notes. — Though much smaller, it is verv similar in

° Only a single male specimen available for study.

344 The University Science Bulletin

general appearance to A. occipitalis Breddin. However the males
of the latter lack the extension of the second rostral segment along
the third rostral segment as found on the male of A. timorensis.
For comparison of the chaetotaxies of the front legs of the males of
the two see Plate XXXIX, fig. 18a and Plate XXXIX, fig. 22a.
Data on distribution. — Known only from type locality.

Anisops occipitalis Breddin

(PI. XXXIX, fig. 22)

1905. Anisops occipitalis Breddin, Mitteil. Naturh. Mus. Hamburg, vol. XX, p. 152.
1933. Anisops occipitalis, Lundblad, Arch, fiir Hydrob., Suppl. vol. XII, pp. 145, 158-
160, text fig. 154.

Size. — Males, length 6.6 mm. -7.2 mm., greatest body width 1.9.-2.1
mm.; females, length 6.6 mm.-7.2 mm., greatest width of body 1.6
mm.-2.1 mm.

Shape. — Bobust, slightly fusiform species; lateral margins in an-
terior half of body only slightly converging.

Color. — General facies stramineous or testaceous. Scutellum may
be testaceous, black, or black only on apical half, posterior half
testaceous or stramineous. Legs stramineous. Abdominal venter
black with keel and segmental margins of the connexivum stra-
mineous.

Male structural characteristics. — Viewed from above, the head is
broad, with its outline rounded; greatest width of head eight to
nine tenths the pronotal width and six and one half times the an-
terior width of the vertex; synthlipsis wide, one half the anterior
width of the vertev; along the median longitudinal axis the head is
slightly more than two thirds the length of the pronotum. Pronotum
with its humeral width two and one half times its median length;
lateral margins diverging and about two thirds the median length;
posterior margin convex, medianly emarginate. Facial tubercle
only slightly raised. Labrum with its basal width slightly more
than its median length; apex rounded; covered with short procum-
bent hairs. Bostral prong (PI. XXXIX, fig. 22b) equal to or slightly
longer than the third rostral segment; apex accumulate. Stridula-
tory comb (Pi. XXXIX, fig. 22c) of about twenty teeth which de-
crease in width from base to apex. Chaetotaxy of the front leg as
shown on Plate XXXIX. The relative lengths of the parts of the legs
are as follows:

Femur

Fore leg; 100

Middle leg 100

Hind leg 100

1st

2nd

Tibia

Tar. Seg.

Tar. Seg.

111

73

76

40

24

82

29

27

Brooks: The Genus Anisops 345

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with its greatest width eight to nine
tenths the pronotal humeral width and five to six times the anterior
width of the vertex; synthlipsis wide, slightly more than one half
the anterior width of the vertex; along the median longitudinal axis
the head is two thirds the length of the pronotum. Pronotum with
its humeral width slightly more than twice its median length; lateral
margins diverging and two thirds the median length; posterior
margin convex, medianly emarginate. The relative lengths of the
parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. .Seg.

Fore leg 100 113 50 29

Middle "leg 100 77 38 23

Hind leg 100 81 29 29

Location of types. — Breddin's type female from Buitenzorg, Java,
in the Hamburg Museum.

Comparative notes. — Very similar in size and general appearance
to Anisops leucothea Esaki from which it can best be distinguished
on the basis of the males. The male of A. leucothea have the
greatest width of the head equal to the pronotal humeral width
and the lateral margins of the pronotum are almost parallel whereas
in this species the greatest width of the head is about nine tenths the
pronotal humeral width and the lateral margins of the pronotum are
diverging.

Data on distribution:

Java

Buitenzorg, Apr. to Dec. 1896, D. G. Fairchild, fifteen males, fourteen
females, Kirkaldy Collection (F. H. Snow Coll.).

Buitenzorg, 1875, G. B. Ferrari, one female, Kirkaldy Collection ( F. H.
Snow Coll. ).

New Guinea

Port Moresby, Guigno, 1889, L. Loria, two males, three females, Kirkaldy
Collection ( F. H. Snow Coll.).

Rigo, Luglio, 1889, L. Loria, seventeen males, eleven females, Kirkaldy
Collection (F. H. Snow Coll.).

New Caledonia

Noumea, A. Fauvel, one male, Kirkaldy Collection ( F. H. Snow Coll. ) .

Australia

Sir Graham Moore Id. 11-20-1945, B. Malkin, two males, one female ( U. S.
Nat. Mus.).

Barren River nr. Barren Waters, IX-2-38, R. G. Wind, one male ( F. II.
Snow Coll.).

346 The University Science Bulletin

Anisops coutierei n. sp.

(PI. XXXIX, fig. 21)

Size. — Males, length 5.4 mm. -5. 8 mm., greatest body width 1.4
mm.-1.6 mm.; females, length 5.8 mm. -6. 8 mm., greatest body width
1.5 mm. -1.8 mm.

Shape. — Subfusiform species; greatest body width about midway
the body length.

Color. — General facies testaceous. Legs stramineous. Abdominal
venter dark brown with keel and segmental margins of the connex-
ivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of head nine tenths the pro-
notal humeral width and five to six times the anterior width of the
vertex; synthlipsis wide, one half the anterior width of the vertex;
along the median longitudinal axis the head is one half the pronotal
length. Pronotum with its humeral width twice its median length;
lateral margins diverging, almost three fifths the median length;
posterior margin convex, medianly emarginate. Facial tubercle
only slightly raised. Labrum short and broad; basal width one and
one half the median length; apex rounded. Rostral prong (PI.
XXXIX, fig. 21b) longer than the third rostral segment; apex ac-
cumulate. Stridulatory comb, (PI. XXXIX, fig. 21a) of approximately
twenty teeth, decreasing slightly in length at apex. Chaetotaxy of
the male front leg as shown on Plate XXXIX. The relative lengths
of the parts of the legs are as follows :

1st 2nd

Femur Tiliia Tar. Seg. Tar. Seg.

Fore leg 100 108 72

Middle leg 100 85 40 25

Hind leg 100 79 32 32

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of the head nine tenths
the pronotal humeral width and five to six times the anterior width
of the vertex; synthlipsis wide, one half the anterior width of the
vertex; along the median longitudinal axis the head is three fifths
the length of the pronotum. Pronotum with its humeral width
slightly more than twice its median length; lateral margins diverging
and three fifths the median length; posterior margin convex, me-
dianly emarginate. The relative lengths of the parts of the legs are
as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 114 51 36

Middle leg 100 83 38 25

Hind leg 100 77 30 30

Brooks: The Genus Anisops 347

Location of types. — Male holotype, female allotype, one male and
one female paratypes Africa, Djibouti, 1897, H. Coutiere, in the
Paris Museum. Two male and two female paratypes ( same collec-
tion data as above) in the F. H. Snow Collection.

Comparative notes. — Very closely related to Anisops debiUis
Gerst. from which it can be distinguished by its wider synthlipsis
which is one half the anterior width of the vertex instead of one fifth
the anterior width of the vertex. Also the male front femur is
slightly rounded at the apex whereas in A. debilis it is accumulate.

Data on distribution. — Known only from type series.

Anisops pellucens Gerstaecker

(PI. XL, fig. 26)

1873. Anisops pellucens Gerstaecker, Decken's Reise in Ost Africa III, pt. 2, pp. 424-425.

1929. Anisops pellucens, Hutchinson. Ann. South African Mus., vol. XXV, pt. 2, pp. 384-
385, PI. XXIX, fig. 2; PI. XXX, fig. 10; PI. XXXI, fig. 2.

1930. Anisops pellucens f. pellucens, Hutchinson. Proc. Zool. Soc. London, vol. XXIX,

pt. 2, p. 445.

1930. Anisops pellucens f. splendida, Hutchinson, Proc. Zool. Soc. London, vol. XXIX,

pt. 2, p. 445.

1930. Anisops pellucens f. pellucens. Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. VI,
pp. 57-58 (ecological note).

1933. Ayiisops pellucens, Jaczewski, Linn. Soc. Jour., Zool., vol. XXXVIII, p. 345.

1937. Anisops pellucens, Poisson, Ann. Soc. Ent. France, vol. CVI, p. 118, fig. 5 (discusses
the color forms of Hutchinson).

1937. Anisops pellucens f. rubroscutellata, Poisson. (loc. cit.), p. 119.

1937. Anisops pellucens f. grandis, Poisson. (loc. cit.), p. 120. (I have raised this to
specific level.)

1940. Anisops pellucens f. pellucens. Poisson, Bull. Mus. Royal Hist. Nat. Belgique, vol.
XVII, no. 29, p. 19, (ecological note).

1941. Anisops pellucens, Poisson, Rev. Franc. Ent.. vol. VIII, p. 97, (ecological note).
1948. Anisops pellucens f. pellucens, Poisson, Mem. Inst. Sci. Madagascar, Ser. A, vol. I,

pt. 2, p. 108 (ecological note).

Referring to this species also :

1899. Anisops nivea, Kirkaldy, Ann. Soc. Ent. France, vol. LXVIII, p. 105 (determined
in error).

1901. Anisops nivea, Kirkaldy, The Entomologist, vol. XXXIV, p. 5, (determined in error).

1904. Anisops nivea, Kirkaldy, Wiener Ent. Zeit.. vol. XXIII, p. 119, 132 (determined in
error).

1908. Anisops nivea, Kirkaldy, Sjostedf : YVissenschaften Ergebn. der Schwed. Zool.
Exped., Hemiptera, vol. XII, p. 24 (determined in error).

1928. Anisops nivea, Hutchinson. Ann. Mag. Nat. Hist., Ser. 10, vol. I, p. 164 (deter-
mined in error).

1929. Anisops nivea, Poisson, Faune des Colonies Francaise, vol. Ill, pp. 154-155, text
fig. 22.

1898. Anisops ciliata, Kirkaldy, Ann. Mus. Civ. Stor. Nat. Genova, vol. XXIX (deter-
mined in error).

1892. Anisops scutellaris, De Carlini, Ann. Mus. Civ. Stor. Nat. Genova Ser. 2, vol. XII
(XXXII), p. 11 (determined in error: no description given, only size; probably this species).

1895. Anisops scutellaris, De Carlini, {loc cit.), vol. XXXV, p. 19 (determined in error;
no description given, only size; probably this species).

Size. — Males, length 9.0 mm. -9.6 mm., greatest body width 2.4
mm.-2.8 mm.; females, length 9.6 mm. -9.9 mm., greatest body
width 2.2 mm. -2.7 mm.

348

The University Science Bulletin

Shape. — Large fusiform species, widest about midway the length
of the body.

Color. — General facies stramineous or gray. Eyes brown. Scu-
tellum may be red with anterior margin black, entirely red or testa-
ceous, or entirely black. Hemelytra gray or stramineous and hyaline
and appear darker due to the underlying darker dorsal body surface.
Legs stramineous. Abdominal venter brown or dark brown with
keel and segmental margins of the connexivum testaceous or stra-
mineous. The latter may be tinged with red.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of the head eight to nine
tenths the pronotal humeral width and six times the anterior width
of the vertex; synthlipsis wide, one half or more the anterior width of
the vertex; along the median longitudinal axis the head is slightly
more than one half the length of the pronotum. Pronotum with its
humeral width almost twice its median length; lateral margins di-
verging and one half the median length; posterior margin convex,
medianly emarginate. Facial tubercle slightly raised with a few,
small scattered hairs. Labium short and broad; basal width one
and eight tenths the median length. Rostral prong ( Pi. XL, fig. 26b )
slightly shorter than the third rostral segment; apex more or less
blunt. Stridulatory comb ( PL XL, fig. 26c ) of approximately twenty-
two teeth which increase slightly in length from the base to the apex.
Chaetotaxy of the front leg as shown on Plate XL. The relative
lengths of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 80

Middle leg 100 87 38 25

Hind leg 100 80 30 28

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of the head eight tenths
the pronotal humeral width and five to six times the anterior width
of the vertex; synthlipsis wide, almost two thirds the anterior width
of the vertex; along the median longitudinal axis the head is ap-
proximately one half the length of the pronotum. Pronotum with
its humeral width slightly more than twice the median length;
lateral margins diverging and one half the median length; posterior
margin convex, medianly emarginate. The relative lengths of the
parts of the legs are as follows:

lsl 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 117 50 25

Middle leg 100 90 37 25

Hind leg 100 77 27 27

Brooks: The Genus Anisops 349

Location of types. — Gerstaecker's type material in the Berlin
Museum.

Comparative notes. — This species along with Anisops letitia
Hutchinson is the largest known species from the African mainland.
Though only slightly larger than A. letitia it can be readily dis-
tinguished from it by the fact that the latter species has the posterior
margin of the pronotum convex and not medianly emarginate, as in
A. pellucens.

Data on distribution:
Africa

Angola, St. Paul de Loando, two females (U. S. Nat. Mus. ).

St. Paul, two females (F. H. Snow Coll.).

? — Paru, one male (F. H. Snow Coll.).

East Africa, IV-28-1948, F. X. Williams (Harv. Mus. Comp. Zool.).

Tanganyika Terr. Amani, Feb. 1948, F. X. Williams, two males, two females
(F. H. Snow Coll.), four males, twelve females (Harv. Mus. Comp. Zool.).

Reg. de Zinder, Dungass, Mission Tikho, 1910, Dr. R. Gaillard, one female
(Paris Museum).

Abyssinia, Plaine Danakil, Maro, 1903, Dr. J. Rogers, one female (Paris
Museum ) .

Souden-Egyptian, Roseires, Haut Nil Bleu, 1907, Ch. Alluaud, one male, one
female (Paris Museum).

Senegal, one male (Paris Museum).

Africa Orient. Anglo, Kisoumou, Victoria-Kyanza, 1904, Ch. Alluaud, two
males, five females ( Paris Museum ) .

Belgium Congo: Eala, XI-1935, J. Ghesquiere, three females (Belgium
Museum).

South Africa: Belagoa, Coll. Juond., one male (Basel. Nat. Hist. Mus.
Switzerland ) .

Anisops malkini n. sp.*

(PI. XL, fig. 23)

Size. — Male, length 6.9 mm.; greatest body width 1.6 mm.

Shape. — Fusiform species with greatest width about midway the
body length.

Color. — General facies stramineous. Eyes brown. Legs stramin-
eous. Abdominal venter black with keel and segmental margins
of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of the head nine tenths the
pronotal humeral width, five to six times the anterior width of the
vertex; synthlipsis wide, slightly more than one half the anterior
width of the vertex; along the median longitudinal axis the head is
three fourths the pronotal length. Pronotum with its humeral width

* Only one male specimen was available for study.

350 The University Science Bulletin

almost twice its median length; lateral margins diverging and one
half the median length; posterior margin convex, medianly emar-
ginate. Frons depressed and flat, terminated by a transverse ridge
at its apex. Labrum short, with the basal width one and one fifth
its median length; apex rounded. Rostral prong (PL XL, fig. 23b)
slightly shorter than the third rostral segment; apex accuminate.
Stridulatory comb ( PL XL. fig. 23c ) of about fifteen teeth with the
longest midway between the base and apex. Chaetotaxy of the
front leg as shown on Plate V. The relative lengths of the parts of
the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 68

Middle leg 100 84 40 22

Hind leg 100 82

Unfortunately, the first and second tarsal segments of the hind
legs were lacking.

Location of types. — Male holotype. Darwin NT, Australia, III-
25-45, B. Malkin, in the United States National Museum.

Comparative notes. — Though much larger than Anisops crinita
n. sp., this species has the flattened frons as possesed by A. crinita.
There is no danger of confusing the two for in addition to the size
difference, A. malkini does not have the hairy frons of A. crinita
and also the chaetotaxies of the front legs of the males is decidedly
different.

Data on distribution: Known only from type locality.

Anisops evansi n. sp.

(PI. XLI, fig. 27)

Size. — Males, length 6.9 mm., greatest body width 1.6 mm.; fe-
males, length 6.7 mm., greatest body width 1.4 mm.

Shape. — Fusiform, broadheaded species; greatest body width
midway the body length.

Color. — General facies gray. Eyes brown. Legs stramineous.
Abdominal venter dark brown with keel and segmental margins of
the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the vertex extending slightly beyond
the anterior margin of the eyes; greatest width of the head slightly
more than the pronotal humeral width, slightly more than six times
the anterior width of the vertex; synthlipsis wide, three fourtiis the
anterior width of the vertex; along the median longitudinal axis
the head is slightly shorter than the pronotum. Pronotum with its

Brooks: The Genus Anisops

351

humeral width approximately one and five sevenths its median
length; lateral margins almost parallel only slightly diverging and
slightly more than one half the median length; posterior margin
convex, medianly emarginate. Facial tubercle flat, only slightly
raised. Labrum short with long hairs arising at base and extending
almost the length of the labrum; basal width one and one third the
median length; apex rounded, almost truncate. Rostral prong (PL
XLI, fig. 27b) shorter than third rostral segment; apex accuminate.
Stridulatory comb (PL XLI, fig. 27c) of approximately twenty teeth,
basal seven longer than the apical thirteen. Chaetotaxy as shown
on Plate XLI. The relative lengths of the parts of the legs are as

f°ll0WS: 1st 2nd

Femur Tibia Tar. t?eg. Tar. Seg.

Fore leg 100 133 86

Middle leg 100 81 34 22

Hind leg 100 78 32 32

Female structural characteristics. — Viewed from above, the out-
line of the head is more or less conical with the vertex extending
slightly beyond the anterior margins of the eyes; greatest width of
head nine tenths the pronotal humeral width, four times the anterior
width of the vertex; synthlipsis wide, two thirds the anterior width
of the vertex; along the median longitudinal axis the head is slightly
more than two thirds the pronotal length. Pronotum with its hu-
meral width twice its median length; lateral margins diverging and
slightly more than one half the median length; posterior margin
convex, medianly emarginate. The relative lengths of the parts of
the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 131 56 31

Middle leg 100 88 38 23

Hind leg 100 83 30 30

Location of types. — Male holotype, female allotype. Tasmania,
Lake Dakerton, 11-1940, J. W. Evans, in the Snow Entomological
Collection.

Comparative notes. — Though slightly smaller than Anisops tas-
maniaensis n. sp., A. evansi appears closely related to it. The males
of the two species serve as a ready means of separation for that of
the latter has the head wider than the pronotum whereas in the
former the head is at most only equal to the pronotal width. Also
the males of Anisops tasmaniaensis lack the long hairs as found on
the labrum of A. evansi. For comparison of the chaetotaxies of the
front legs of the males see Plate XLI, fig. 27a, and Plate XL VII,
fig. 58a.

Data on distribution. — Known only from type series.

352 The University Science Bulletin

Anisops gratus Hale

(PI. XLII, fig. 31)

1923. Anisops gratus Hale, Rec. South Australian Mus. vol. II, no. 3. pp. 413-414, text
fig. 369.

1933. Anisops gratus, Lundblad, Archiv fur Hydrob., Suppl. vol. XII, p. 145 (a list of
Indo-australian and Pacific forms of this genus).

Size. — Males, length 7.2 mm. -7. 8 mm., greatest body width 1.9
mm. -2.1 mm.; females, length 7.2 mm.-7.8 mm., greatest body width
1.9 mm. -2.1 mm.

Shape. — Fusiform species; greatest body width about midway the
body length.

Color. — General facies stramineous or testaceous. Eyes brown.
The testaceous areas are often tinged with orange on the scutellum.
Legs stramineous. Abdominal venter testaceous or stramineous
with irregular brown areas appearing on each sternite.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded, with the anterior margin of the head almost
straight; greatest width of head nine tenths the pronotal humeral
width and four to five times the anterior width of the vertex; synth-
lipsis wide, approximately one third the anterior width of the vertex;
along the median longitudinal axis the head varies from almost one
half to one half the length of the pronotum. Pronotum with its
humeral width almost twice its median length; lateral margins
diverging and one half the median length; posterior margin convex,
medianly emarginate; dorsal surface with a median longitudinal de-
pression. Facial tubercle only slightly raised. Labrum with its
basal width one and one fifth its median length; apex rounded. Ros-
tral prong ( PI. XLII, fig. 31b ) longer than the third rostral segment;
apex accuminate. Stridulatory comb (PI. XLII, fig. 31c) of approxi-
mately sixteen teeth. Chaetotaxy of the front leg as shown on Plate
XLII. The relative lengths of the parts of the legs are as follows:

1st -llul

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 111 65

Middle leg 100 73 35 20

Hind leg 100 85 37 37

Feiyiale structural characteristics. — Viewed from above, the out-
line of the head is rounded with the vertex extending slightly beyond
the anterior margins of the eyes; greatest width of the head eight
tenths the pronotal humeral width and three to four times the an-
terior width of the vertex; synthlipsis wide, slightly more than one
third the anterior width of the vertex; along the median longitudinal

Brooks: The Genus Anisops 353

axis the head is almost one half the pronotal length. Pronotum with
its humeral width slightly more than twice its median length; lateral
margins diverging and eight tenths the median length; posterior
margin convex, medianly emarginate, dorsal surface with the same
median depression as found on the males. The relative lengths of
the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 118 50 27

Middle leg 100 89 41 22

Hind leg 100 45 38 38

Location of types. — Type material, Broken Hill, New South Wales,
Australia, F. W. Shepherd in the South Australian Museum. Two
male, and four female cotypes, Broken Hill, New South Wales,
Australia, Fred W. Shepherd in the Snow Entomological Collection.

Comparative notes. — This species is of about the same size as A.
doris Kirk, but the males lack the broad rounded head and the nar-
row anterior width of the vertex as found on the males of A. doris.
For comparison of the chaetotaxies of the front legs of the males
see PL XLII, fig. 31a and PI. XLVI, fig. 53a.

Data on distribution:
Australia

New South Wales, Broken Hill, Fred W. Shepherd, three females (F. H.
Snow Coll. ) .

New South Wales, Broken Hill, in nine mile dam, IV-23-44, C. E. Chadwick,
one male, five females (Dept. of Agrie., N. S. W., Austrl. ).

New South Wales, Broken Hill, IV-10-44, C. E. Chadwick, nine females
(Dept. of Agric. N. S. W., Austrl.).

New South Wales, Broken Hill, 1944, C. E. Chadwick, two females (Dept.
of Agric. N. S. W., Austrl.).

New South Wales, Imperial Dam, XII-31-1925, F. W. Shepherd, thirty-two
males, eleven females (F. H. Snow Coll.).

Bordertown, one female (F. H. Snow Coll.).

Sir Graham Moore Id. 11-20-45, B. Malkin, two females (F. H. Snow Coll.).

Anisops gobana Poisson

(PI. XLI, fig. 28)

1940. Anisops gobana Poisson, Bull. Mus. Roy. Hist. Nat. Belgique, vol. XVI, no. 27,
pp. 10-12, text figs. 13-16.

Size. — Males, length 6.9 mm. greatest body width 2.1 mm.; fe-
males, length 7.2 mm., greatest body width 2.1 mm.

Shape. — Subfusiform species; lateral margins almost parallel in
anterior half, converging in posterior half.

Color. — General facies black. Eyes brown. Vertex and pro-
notum stramineous, the latter with a faint dark brown median longi-
23—3286

354 The University Science Bulletin

tudinal stripe. Scutellum black, with lateral margins testaceous.
Hemelytra hyaline and appears black due to the black dorsal body
surface, margins bordering scutellum and hemelytral commissure
red. Legs stramineous. Abdominal venter black with keel and
segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of head nine tenths the pro-
notal humeral width and slightly more than four times the anterior
width of the vertex; synthlipsis wide, slightly more than one third
the anterior width of the vertex; along the median longitudinal axis
the head is three fifths the length of the pronotum. Pronotum with
humeral width slightly more than twice the median length; lateral
margins diverging and one half the median length; posterior margin
convex, medianly emarginate. Facial tubercle slightly raised. La-
brum long, median length equal to basal width; apex more or less
accuminate. Rostral prong (PI. XLI, fig. 28b) shorter than the
third rostral segment; apex accuminate. Stridulatory comb (PI.
XLI, fig. 28c) of approximately eight teeth. Chaetotaxy of the male
front leg as shown on Plate XLI. The relative lengths of the parts
of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 125 90

Middle leg 100 81 39 23

Hind leg 100 80 42 (lacking)

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of the head eight tenths
the pronotal humeral width and slightly more than four times the
anterior width of the vertex; along the median longitudinal axis the
head is slightly more than one third the length of the pronotum.
Pronotum with its humeral width slightly more than twice the
median length; lateral margins diverging and one half the median
length; posterior margin convex, medianly emarginate. The relative
lengths of the parts of the legs are as follows:

1st 2nd

Femur Tilna Tar. Seg. Tar. Seg.

Fore lea; 100 125 86 37

Middle leg 100 80 40 25

Hind leg 100 82 37 37

Location of types. — Type material, Ethiopia, Cuba, 1934-1935, R.
de Meulenaere, in the Royal Museum of Natural History in Brussels,
Belgium. One male and one female paratypes (same collection
data as above) in the F. H. Snow Entomological Collection.

Comparative notes. — This species is very similar in general ap-

Brooks: The Genus Anisops 355

pearance to Anisops varia Fieber. However, the synthlipsis is nar-
rower, being about one third the anterior width of the vertex instead
of one half as in A. varia. Also the head is slightly narrower in this
species, only eight tenths the pronotal humeral width instead of nine
tenths as found on A. varia.

Data on distribution. — Known only from type series.

Anisops varia Fieber

(PI. XLII, fig. 32)

1851. Anisops rarius Fieber, Abhand. Kongl. Boh. Gesell. Wiss., vol. V, pt. 7, pp. 483-484.

1851. Anisops various f. sugillata, Fieber, (loc. cit.), p. 483.

1851. Anisops varius i. scutellata, Fieber, (loc. cit.), p. 483.

1853. Atiisops varius, Heirich- Shaffer, Die Wanzenartigen Insecten, vol. IX, p. 10 (list of
the species of Anisops).

1888. Anisops varius, Horvath, Rev. Ent., vol. VII, p. 189 (= Notonecta nanula Walker).

1899. Anisops varius, Kirkaldy, Ann. Soc. Ent. France, vol. LXVIII, p. 100 (= Anisops
perpulcher Stal).

1899-1900. Anisops varia, Puton, Rev. Ent., vol. LXVIII, p. SO.

1904. Anisops varia, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 119, 132.

1905. Anisops varius, Distant, Trans. Linn. Soc. London, vol. XIII, pt. I, p. 38 (ecological
note).

1910. Anisops varius. Distant, Fauna of British India, vol. V, p. 332 (ecological note).

1922. Anisops varia, Lindberg, Notulae Entomologicae, vol. II, p. 47, text fig. 3.

1920. Anisops varius, Esaki, Ann. Mus. Nat. Hungarici, vol. XXIV, p. 188 (ecological
note).

1926. Anisops varia, Jaczewski, Ann. Mus. Polonici Hist. Nat., vol. XXIV, p. 188 (eco-
logical note).

1928. Anisops varia, Dover, Treubia, vol. 10, p. 71 (ecological note).

1929. Anisops varia, Hutchinson, Ann. South African Mus., vol. XXV, pp. 393-400, PI.
XXX, fig. 17; PI. XXXII, fig. 2 (says the .4. perpulcher kalahariensis Schumacher and A. per-
pulcher piumbeus Schumacher are general forms).

1929. Anisops varia varia. Hutchinson, Ann. South African Mus., vol. XXV, p. 399 (= A.
varia sugillata Fieber).

1929. Anisops varia scutellata, Hutchinson, Ann. South African Mus., vol. XXV, p. 399
(= A. perpulcher Stal).

1930. Anisops varia varia, Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. VI, p. 58 (eco-
logical note).

1932. Anisops varia varia, Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. IV, p. 324
(ecological note).

1933. Anisops varia, Hutchinson, Internat. Revue der ges. Hydrob. Hydrog., vol. 28,
pt. 5/6, p. 438 (ecological note).

1933. Anisops varia, Lundblad, Arch, fur Hydrob., Suppl. vol. XII, p. 146 (list of Indo-

australian and Pacific members of this genus).

1935. Anisops varia varia, Poisson, Mus. Nat. Hist. Nat., vol. Ill, p. 208 (ecological note).

1936. Anisops varia, Jaczewski, Ann. Zool. Musei Polonici. vol. XI, p. 201 (ecological note).
1939. Anisops varia varia, Poisson, Bull. Soc. Ent. France, vol. 44. p. 42 (ecological note).
1948. Anisops varia varia, Poisson, Inst. Rech. Sahar. Univ. Alger, p. 7 (ecological note).
194S. Anisops varia scutellata, Poisson, (loc. cit.), p. 7 (ecological note).

Referring to this species also:

1858. Anisops perpulcher, Stal, Ofver. Kongl. Vetensk. Akad. Fordhandl., vol. XX, p. 89.

1869. Anisops perpulcher, Stal, Hemiptera Africana III, p. 192.

1892. Anisops perpulcher, De Carlini, Ann. Mus. Stor. Nat. Genova, Series 2a, vol. XII,
(XXXII), p. 11, (ecological note).

1913. Anisops perpulcher. Schumacher, Denksch. der Medic. Ges. Jena, vol. 17. p. 83.

1913. Anisops perpulcher piumbeus, Schumacher, (loc. cit.), p. 83 (determined in error).

1913. Anisops perpulcher kalahariensis, Schumacher. {Inc. cit.) p. 83 (determined in error).

356 The University Science Bulletin

1925. Anisops perpulcher. Hesse, Ann. South African Mas., vol. XXII, p. 137 (determined
in error).

1929. Anisops hoggarica, Poisson, Bull. Soc. Hist. Nat. Afrique du Xord, vol. XX, pp. 89-
91, text figs. 4, 5, 6 (determined in error).

1933. Anisops hoggarica, Hutchinson, Internat. ges. Hydrob. Hydrog., vol. 28, pt. 5/6,
p. 453 (ecological note).

1934. Anisops hoggarica, Poisson, Mem. Soc. Hist. Nat. Afrique du Nord, vol. 4, pp. 137-
139, text figs. 4, 5, 6 (determined in error).

1939. Anisops hoggarica, Poisson, Bull. Soc. Ent. France, vol. 44, p. 43 (ecological note)

1941. Anisops hoggarica, Poisson, Rev. Francaise Ent., vol. VIII, p. 77 (ecological note).

1948. Anisops varia hoggarica, Poisson, Inst. Recher. Sahar. Univ. d'Alger, p. 6.

1850. Anisops scutallaris, Herrich -Shaffer, Die Wanzenartigen Insecten, vol. IX, p. 41,
fig. 90G.

189G. Anisops Scutellaria, Matsumara, Trans. Sapporo Nat. Hist. Soc, vol. I, p. 28 (eco-
logical note).

1915. Anisops scutellaris, Matsumura, Ent. Mag., Kyoto, vol. I, p. 110 (ecological note).

1915. Anisops scutellaris, Esaki, Ent. Mag. Kyoto, vol. I, pp. 31, 81 (ecological note).

1870. Notonecta nanula. Walker, The Zoologist (II), vol. V, p. 2381.

Size. — Males, length 6.0 mm.-6.2 mm., greatest body width 1.8
mm. -2.9 mm.; females, length 6.0 mm. -6.6 mm., greatest body width
1.9 mm. -2.1 mm.

Shape. — Snbfusiform species; greatest body width about midway
the body length.

Color. — General facies dark gray or black. Eyes dark brown.
Vertex and pronotum testaceous, the latter may have irregular
brown spots. Scutellum dark brown or black, lateral margins may
be tinged with red or scutellum may be entirely red. Hemelytra
hyaline and appear dark gray or black due to the color of the
dorsal body surface; margins bordering scutellum and hemelytral
commissure may be tinged with red. Legs stramineous. Abdominal
venter black with keel and segmental margins of the connexivum
stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the anterior margin almost straight;
anterior margins of eyes extending slightly beyond that of the
vertex; greatest width of head nine tenths the pronotal humeral
width and six times the anterior width of the vertex; synthlipsis
wide, one half the anterior width of the vertex; along the median
longitudinal axis the head is almost one half the length of the
pronotum with its humeral width slightly less than twice the
median length; lateral margins diverging and one half the median
length; posterior margin convex, medianly emarginate. Facial tu-
bercle slightly raised. Labrum long; median length equal to its
basal width; apex rounded. Rostral prong (PI. XLII, fig. 32b)
shorter than third rostral segment; apex more or less accuminate.
Anterior femur with apex more or less curved. Stridulatory comb
(Pi. XLII, fig. 32c) of approximately twenty-one teeth. Chaetotaxy

Brooks: The Genus Anisops 357

of the male front leg as shown on Plate XLII. The relative lengths
of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 119 75

Middle leg 100 82 36 25

Hind leg 100 82 29 31

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head almost nine
tenths the pronotal humeral width and five times the anterior width
of the vertex; synthlipsis wide, slightly more than one half the
anterior margin of the vertex; along the median longitudinal axis
the head is almost one half the length of the pronotum. Pronotum
with its humeral width slightly more than twice the median length;
lateral margins diverging and slightly more than one half the me-
dian length of the pronotum; posterior margin convex, medianly
emarginate. The relative lengths of the parts of the legs are as
follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 53 33

Middle leg 100 80 36 24

Hind leg 100 80 28 28

Location of types. — Fieber's type material in the Berlin Museum.

Comparative notes. — This species is very similar to Anisops ares
Hutch, but does not have the long pronotum of this species. In
A. varia the pronotum is twice the length of the head whereas in
A. ares the pronotum is three times the length of the head.

Remarks. — This species has a rather wide recorded distribution
occurring in Asia as well as in Africa. However, in the vast Pacific
and Asiatic material available to me, not one specimen of this species
was found. I am of the opinion that this species is composite, with
the Asiatic form being another species entirely. There is need of a
close examination of the type and the Asiatic forms placed with
this species.

Data on distribution:
Africa

South Africa, Pine Mission, III-3-1926, one female, gift to H. B. H. fr. G. E.
Hutchinson (F. H. Snow Coll.).

South Africa, Blaaukrantz Valley, pools in dried stream bed, IX-3-1926,
gift to H. B. H. fr. G. E. Hutchinson, one female ( F. H. Snow Coll.).

Egypt, Alexandria, E. de Bergevin, one male, one female (U. S. Nat. Mus. ),
two females, five males ( F. H. Snow Coll. ) .

Abyssinia, Colonia Erytree, A. Thery, one male (F. H. Snow Coll.).

358 The University Science Bulletin

Mocambique, Vallee du Pungoue, Guengere, 1906, G. Vasse, four females
( Paris Museum ) .

Afrique, Orient. Angl., Naivasha, 1904, Ch. Alluaud, one female (Paris
Museum).

Basin Inf. du Zambeze, Vallee du Nuza, 1905, G. Vasse, one female (Paris
Museum ) .

Sengal, Heudelot, one female (Paris Museum).

Cape-Town, E. Simon, 1898, Noualhier, two females (Paris Museum) one
female (F. H. Snow Coll.).

Reg. de Zinder, Maradi (Mission, Tilmo) 1910, Dr. R. Gaillard, two females
( Paris Museum ) .

Tchad, N'Guigmi, 1919, Dr. Noel, one male (Paris Museum).

Soudan, Nioro, 1909, F. de Zeltner, one male (F. H. Snow Coll.).

Sahel Soudanais, Goumbou 1909, F. de Zeltner, one male (Paris Museum).

Belgian Congo, Musoca, IX-1939, H. J. Bredo, one male, one female (Bel-
guium Museum).

Erythraea, Ghenda, two females (Bueno Coll. of F. H. Snow Coll.).

Syria

Syrien, Kaifa, Reuter, one male, one female (Kirkaldy Coll. of the F. H.
Snow Coll. ).

Anisops robusta Hutchinson

(PI. XLI, fig. 30)

1930. Anisops robusta Hutchinson, Proc. Zool. Soc. London, vol. XXIX, pp. 446-447, text
figs. 3, a, b, c.

1933. Anisops robusta, Hutchinson, Internat. ges. Hydrob. Hydrog., vol. 28, pt. 5/6, p. 462
(ecological note).

Size. — Males, length 7.3 mm., greatest body width 2.2 mm.;
females, length 7.3 mm., greatest body width 2.4 mm.

Shape. — Robust, subfusiform species; greatest body width mid-
way the body length.

Color. — General facies gray. Eyes gray brown. Hemelytra
hyaline and appears dark gray due to the dorsal dark body surface.
Legs stramineous. Abdominal venter dark brown with keel and
segmental margins stramineous.

Male structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of the head equal to the
pronotal humeral width and five times the anterior width of the
vertex; synthlipsis wide, almost one half the anterior width of the
vertex; along the median longitudinal axis the head is slightly more
than one half the length of the pronotum. Pronotum with its
humeral width one and two thirds the median length; lateral mar-
gins convexly curved and converging, one half the length of the
median length; posterior margin convex, medianly emarginate.
Facial tubercle slightly raised. Labrum long, median length equal
to the basal width; apex accumulate. Rostral prong (PI. XLI, fig.

Brooks: The Genus Anisops 359

30b) slightly shorter than third rostral segment; apex accuminate.
Fore leg (PL XLI, fig. 30a) with dorsal and ventral femoral mar-
gins almost parallel, apex rounded; stridulatory comb (PL XLI,
fig. 30c) of approximately eleven teeth which increase in length
from base to apex. Whether or not two apical teeth are missing
from my specimen is not clear for Hutchinson (24) figures such a
condition in his drawings. Chaetotaxy of the front leg as shown on
Plate XLI. The relative lengths of the parts of the legs are as fol-
lows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 105 69

Middle leg 100 87 41 27

Hind leg 100 87 32 32

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head nine tenths the
pronotal humeral width and four times the anterior width of the
vertex; synthlipsis wide, almost one half the anterior width of the
vertex; along the median longitudinal axis the head is over one half
the length of the pronotum. Pronotum with its humeral width
slightly more than twice the median length; lateral margins di-
verging and almost three fourths the median length, posterior
margin convex, medianly emarginate. The relative lengths of the
parts of the legs are as follows:

1st 2nd

Femur Tihia Tar. Seg. Tar. Seg.

Fore leg 100 105 48 26

Middle leg 100 84 41 23

Hind leg 100 80 35 35

Location of types. — Type material from Wouranboulchi, Abys-
sinia, in the British Museum.

Comparative notes. — The peculiar shape of the front femur and
the convexly curved, converging lateral margins of the pronotum
of the males of this species make it quite unique. Anisops ares
Hutchinson is perhaps the most similar species, however, the fore
femur of the males is not parallel sided for its entire length and the
lateral margins of the pronotum are at most subparallel.

Data on distribution:
Africa

Abyssinia, Serpent Lake, Wouranboulchi, X-4-1926, J. Omer-Cooper one
male, one female, exchange from the British Museum (F. H. Snow Coll.).

Abyssina, Wouranboulchi nr. Djem-Djem. X-5-1926, J. Omer-Cooper, one
male (F. H. Snow Coll.).

Ahyssinia, Wouranboulchi, marsh, X-5-1926, J. Omer-Cooper, one female
(F. H. Snow Coll.).

360 The University Science Bulletin

Anisops sikoroensis n. sp.

(PI. XLII. fig. 34; PI. LVI. fig. 103)

Size.— Males, length, 10.5 mm., greatest body width 3 mm.; fe-
males, length 10 mm., greatest body width 3 mm.

Shape. — Large fusiform species, with greatest body width about
one third the body length.

Color. — General facies red brown. Abdominal venter darker with
keel and segmental margins of the connexivum red brown.

Male structural characteristics. — Viewed from above, the out-
line of the head is rounded with the greatest width of the head
eight tenths the pronotal humeral width and five to six times the
anterior width of the vertex; synthlipsis wide, slightly more than
one half the anterior width of the vertex; along the median longi-
tudinal axis the head is one half as long as the pronotum. Pronotum
with its humeral width twice the median length; lateral margins
diverging and almost three fourths the median length; posterior
margin convex, medianly emarginate. Facial tubercle with a slight
median groove which extends to about midway the length of the
frons, apical two thirds of groove bearing a median longitudinal
row of long erect hairs. Labrum with basal width one and one
fourth the median length; apex rounded. Rostral prong (PI. LVI,
fig. 103) shorter than third rostral segment; apex more or less
accuminate. Stridulatory comb (PI. XLII, fig. 34b) of approxi-
mately sixteen even-length teeth. Chaetotaxy of male front leg as
shown on Plate XLII. The relative lengths of the parts of the legs
are as follows: lst 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 125 87

Middle leg 100 84 37 20

Hind leg 100 87 36 30

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head eight tenths the
pronotal humeral width and six times the anterior width of the
vertex; synthlipsis wide, two thirds the anterior width of the vertex;
along the median longitudinal axis the head is slightly more than
one half the length of the pronotum. Pronotum with its humeral
width slightly more than twice the median length; lateral margins
diverging, two thirds the median length; posterior margin convex,
medianly emarginate. The relative lengths of the parts of the legs
are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 125 50 33

Middle leg 100 81 31 22

Hind leg 100 81 32 26

Brooks: The Genus Anisops 361

Location of types.— Male holotype, female allotype, Madagascar,
Annanarivo, Sikoro, 1898, Noualhier, in the Paris Museum. One
male paratype (same collection data as above) in the F. H. Snow
Collection.

Comparative notes. — Very similar in general appearance to
A. grandis Poisson, however, the latter lacks the groove on the frons
and the median frontal row of erect hairs as found on the males of
this species.

Data on distribution.— Known only from type series.

Anisops fijiensis n. sp.

(PI. XL, fig. 24)

Size.— Males, length 4.6 mm.4.9 mm., greatest body width 1.2
mm.-1.4 mm.; females, length 4.9 mm.-5.4 mm., greatest body width
1.3 mm. -1.5 mm.

Shape.— Small fusiform species, greatest body width about mid-
way the body length.

Color.— General facies dark gray. Eyes brown. Vertex and pro-
notum testaceous or stramineous, the latter may be hyaline on its
posterior margin and appear the color of the underlying scutellum.
Scutellum black, orange, or black with orange or testaceous apex.
Hemelytra hyaline and appear dark gray as it overlies the black
dorsal body surface. Legs stramineous. Abdominal venter black
with keel and segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest head width nine tenths the pronotal
humeral width, six to seven times the anterior width of the vertex;
synthlipsis wide, approximately one half the anterior width of the
vertex; along the median longitudinal axis the head is three fourths
the pronotal length. Pronotum with its humeral width twice its
median length; lateral margins diverging and almost one half the
median length; posterior margin convex, medianly emarginate.
Facial tubercle laterally compressed forming a slight carina which
runs anteriorly to apex of frons. Labium short; basal width one and
three fourths its median length; apex rounded. Rostral prong
( PL XL, fig. 24b ) as long as third rostral segment; apex accumulate.
Stridulatory comb (PI. XL, fig. 24c) of approximately twenty teeth
which gradually decrease in length from base to apex; apical tooth
almost one half the length of the basal tooth. Chaetotaxy of the
male front leg as shown on Plate XL. The relative lengths of the
parts of the legs are as follows :

362 The University Science Bulletin

1st. 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 84

Middle leg 100 80 37 26

Hind leg 100 82 30 31

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest head width nine tenths the pro-
notal humeral width, six times the anterior width of the vertex,
synthlipsis wide, one half the anterior width of the vertex; along the
median longitudinal axis the head is slightly more than one half the
pronotal length. Pronotum with humeral width slightly more than
twice the median length; lateral margins diverging and almost
one half the median length; posterior margin convex, medianly
emarginate. The relative lengths of the parts of the legs are as
follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 128 55 36

Middle leg 100 80 36 27

Hind leg 100 82 31 31

Location of types. — Male holotype, female allotype, five male and
six female paratypes, Fiji Islands, Taveuni, in shallow fresh water
swamp above high tide, VIII-1934, R. W. Paine in the Snow Ento-
mological Collection. Other paratypes are as follows. In the Snow
Entomological Collection: four males, three females, Fiji Islands,
Taveuni, from shallow muddy pool in coconut plantation, VIII-1934,
R. W. Paine; three males, two females, Fiji Islands, IV-19-39, R. A.
Lever.

Comparative notes. — Though somewhat smaller, this species by
its carinate frons and coloration resembles Anisops tahitiensis Lund-
blad. Anisops fijiensis, however, lacks the enlarged truncate apex
of the male front femur as found on A. tahitiensis Lundblad.

Data on distribution. — Known only from type series.

Anisops adonis Hutchinson

(PI. XLI, fig. 29)

1928. Anisops adonis Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. I, pp. 157-159,
text fig. 2.

1933. A7>isops adonis, Hutchinson, Internat. Rev. ges. Hydrob. Hydrog., vol. 28, pt. 5/6,
p. 462 (ecological note).

Size. — Males, length 5.2 mm.-5.5 mm., greatest body width 1.5
mm. -1.6 mm.; females, length 5.1 mm.-5.4 mm., greatest body width
1.5 mm. -1.8 mm.

Shape. — Small fusiform species; greatest body width about mid-
way the body length.

Brooks: The Genus x\nisops

363

Color. — Brown form: General facies stramineous. Eyes brown.
Legs stramineous. Abdominal venter dark brown with keel and
segmental margins of the connexivum stramineous. Dark form:
General facies dark brown or black. Eyes dark brown. Vertex and
pronotum stramineous. Scutellum dark brown or black. Hemelytra
hyaline, appear black due to the underlying black dorsal body
surface. Legs stramineous. Abdominal venter black with keel and
segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of the head eight tenths the
pronotal humeral width and six times the anterior width of the
vertex; synthlipsis wide, slightly more than one third the anterior
width of the vertex; along the median longitudinal axis the head is
three fourths the length of the pronotum. Pronotum with is humeral
width two and one half times the median length; lateral margins
diverging and slightly more than one half the median length; pos-
terior margin convex, medianly emarginate. Facial tubercle later-
ally compressed with a faint median carina that extends anteriorly
on the frons. Rostral prong (PI. XLI, fig. 29b) shorter than the
third rostral segment; apex more or less rounded. Labrum long
with its median length equal to its basal width; apex rounded; pro-
vided with a few long scattered hairs. Stridulatory comb (PI. XLI,
fig. 29c) of approximately seven, even-length, short, cylindrical
teeth. Chaetotaxy of the front leg as shown on Plate XLI. The
relative lengths of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 128 83

Middle leg 100 88 44 29

Hind leg 100 82 37 34

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with its greatest width eight tenths the
pronotal humeral width and four and one half times the anterior
width of the vertex; synthlipsis wide, one half the anterior width
of the vertex; along the median longitudinal axis the head is
two thirds the length of the pronotum. Pronotum with its humeral
width almost two and one half times the median length; lateral
margins diverging and three fourths the median length; posterior
margin convex, medianly emarginate. The relative lengths of the
parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 125 67 50

Middle leg 100 85 40 27

Hind leg 100 82 36 33

364 The University Science Bulletin

Location of types. — Male type and female allotype, N. Nigeria,
Zungeru, in the British Museum.

Comparative notes. — This species appears closely allied to A.
jaczewski Hutchinson, both have a laterally compressed frons and
similar chaetotaxies on the male front legs. However, the males of
the latter have a short front tarsus, only about one half the length of
the fore tibiae and a narrower synthlipsis, about one third the an-
terior width of the vertex, whereas in the case of A. adonis the
males have the front tarsi two thirds the length of the tibiae and the
synthlipsis is two thirds the anterior width of the vertex.

Data on distribution:
Africa

Soudan, Rock pool, R. Yei Equatoria, Dec. 11, 1937, J. G. Myers, four
males, four females (F. H. Snow Coll.); eleven males, twenty-seven females
(British Nat. Hist. Museum).

Nigeria, Olokemiji, J. C. Birdwell, two males, six females (U. S. Nat. Mus. ).

Nigeria, Olokemiji, Iboda, one male (U. S. Nat. Mus.).

Anisops jaczewski Hutchinson

(PI. XL, fig. 25)

1928. Anisops jaczewski Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. I, p. 304.

1929. A?iisops jaczewski, Hutchinson, Ann. South African Mus., vol. XXV, pt. 3, pp. 403-
404, PI. XXX, fig. 18; PI. XXXII, figs. 5, 8.

1939. Anisops jaczewski, Poisson, Bull. Soc. Ent. France, vol. 44, p. 43 (ecological note).

Referring to this species also :

1926. .4 n is ops vitrea, Horvath, Arch, for Zool., vol. 18A, no. 31, p. 2 (determined in error,
actually A. jaczewski Hutchinson).

1926. Anisops vitrea. Jaczewski, Analibus Zoologicis Musei Polonici Historiae Naturalis,

vol. V, p. 86, text figs. 42, 43, 44 (determined in error; actually A. jaczewski Hutchinson).

Size. — Males, length 4.5 mm. -4.7 mm., greatest body width 1.2
mm.; females, length 4.8 mm., greatest body width 1.4 mm.

Shape. — Small subfusiform species; lateral margins in anterior
half of body almost parallel, lateral margins of posterior half strongly
converging.

Color. — General facies gray. Eyes brown. Vertex, pronotum,
and scutellum testaceous. Hemelytra hyaline and appears dark gray
due to color of the dorsal body surface. Legs stramineous. Abdomi-
nal venter dark brown with keel and segmental margins of the con-
nexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of head eight tenths the
pronotal humeral width and six times the anterior width of the
vertex; synthlipsis wide, one third the anterior width of the vertex;
along the median longitudinal axis the head is three fourth the
pronotal length. Pronotum with its humeral width twice its median

Brooks: The Genus Anisops 365

length; lateral margins diverging and slightly more than one half
the median length; posterior margin convex, medianly emarginate.
Facial tubercle laterally compressed forming a median carina which
runs forward on the frons. Labrum short, basal width one and one
third the median length; apex rounded. Rostral prong ( PI. XL, fig.
25b) shorter than the third rostral segment; apex more or less
accuminate. Stridulatory comb (Pi. XL, fig. 25c) of approximately
ten teeth which increase slightly in height from base to apex.
Chaetotaxy of the male front leg as shown on Plate XL. The relative
lengths of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 110 60

Middle leg 100 86 36 28

Hind leg 100 76 32 30

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of the head eight tenths
the pronotal humeral width and four and one half times the anterior
width of the vertex; synthlipsis wide, slightly more than one third
the anterior width of the vertex; along the median longitudinal axis
the head is slightly more than one half the length of the pronotum.
Pronotum with its humeral width twice the median length; lateral
margins diverging and slightly more than one half the median
length; posterior margin convex, medianly emarginate. The rela-
tive lengths of the parts of the legs are as follows:

ls1 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 46 30

Middle leg 100 78 33 28

Hind leg 100 82 31 29

Location of types. — Type material from Transvaal, Africa in the
British Museum.

Comparative notes. — This species is closely related to A. adonis
Hutch. However the male front tarsus is shorter than in the case
of the latter, being slightly more than one half the length of the
fore tibia whereas the fore tarsus of A. adonis is fully two thirds the
length of the fore tibia. Also the teeth of the stridulatory comb of
A. adonis do not overlap one another as in the case of A. jaczewski
but are distinct and separate from one another.

Data on distribution:
Africa

Transvaal, Main Drift, Limpopo R. Messina, V-29-1927, one male, gift to
H. B. H. fr. G. E. Hutchinson (F. H. Snow Coll.).

Khami River below Ninr, VI-27-1921, one female, gift to H. B. H. fr. G. E.
Hutchinson ( F. H. Snow Coll. ) .

366 The University Science Bulletin

Sudan Africa, Rock pool, R. Yei Equatoria, Dec. 11, 1937, J. G. Myers,
four males, four females (F. H. Snow Coll.) eight males, fourteen females
(Brit. Mus.).

Basin du Chari, Rievere Gribanjui, Mission Chari-Tchad, Jan. 1904, DeCorse,
two males, one female ( Paris Museum ) .

Cameron, 1920, Dr. Noel, one female, one male (Paris Museum).

Moyen Chari, Fort Archambault, Mission Chari-Tchad, 1-1904, Dr. J. De-
Corse, one male, one female (Paris Museum).

Rahr-al-Chazal, Souch-Nassili, 1912, Dr. R. Gaillard, one male, one female
(Paris Museum).

Portuguese East Africa, C. W. Howard, two males (U. S. Nat. Mus.).

Anisops semita n. sp.

(PL XLII, fig. 33)

Size. — Males, length 4.2 mm.-4.5 mm., greatest body width 1.1
mm. -1.2 mm.; females, length 4.1 mm. -4.5 mm., greatest body width
1.1 mm. -1-2 mm.

Shape. — Small, fusiform species; greatest body width about mid-
way its body length.

Color. — General facies stramineous. Eyes brown. Legs stramin-
eous. Abdominal venter black with keel and segmental margins
of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of head nine tenths the
pronotal width, slightly more than five times the anterior width of
the vertex; synthlipsis wide, almost one half the anterior width of
the vertex; along the median longitudinal axis the head is slightly
more than two thirds the length of the pronotum. Pronotum with
its humeral width slightly more than twice its median length;
lateral margins diverging and approximately one half the median
length of the pronotum; posterior margin convex, medianly emargi-
nate. Facial tubercle with a longitudinal depression forming two
lateral ridges which bear long erect hairs. These two rows of hairs
continue down the lateral margins of the labrum to the apex of the
rostrum. Labrum broad; basal width twice its median length; apex
rounded. Rostral prong (PI. XLII, fig. 33b) as long as the third
rostral segment; apex accuminate. Stridulatory comb (PI. XLII,
fig. 33c ) of approximately nine teeth, the basal four twice the length
of the apical five. Chaetotaxy of the male front leg as shown on
Plate XLII. The relative lengths of the parts of the legs are as
follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 76

Middle leg 100 95 37 25

Hind leg 100 78 30 30

Brooks: The Gents Anisops 367

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest head width nine tenths the
pronotal humeral width, four to five times the anterior width of the
vertex; synthlipsis wide, one half the anterior width of the vertex;
along the median longitudinal axis the head is slightly more than
one half the length of the pronotum. Pronotum with its humeral
width twice its median length; lateral margins diverging and slightly
more than one half the median length; posterior margin convex,
medianly emarginate. The relative lengths of the parts of the legs
are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 121 52 28

Middle leg 100 78 45 26

Hind leg 100 77 32 31

Location of types. — Male holotype, Australia, N. Queensland,
Barren River nr. Barren Waters, IX-28-38. R. G. Wind, in the
Snow Entomological Collection. Female allotype, three male and
ten female paratypes, Queensland, Coen C. York, May, '32, Harv.
Austral. Exped., Darlington, in the Harvard Museum of Compara-
tive Zoology. Other paratypes are as follows: One male, two fe-
males, Queensland, Coen C. York, May, '32, Harv. Austral. Exped.,
Darlington, in the Snow Entomological Collection.

Comparative notes. — This species is closely related to Anisops
canaliculata n. sp. and can only be distinguished from it on the
basis of the males which have the two rows of erect hairs which
extend from the ridges of the facial tubercle to the fourth rostral
segment.

Data on distribution. — Known only from type series.

Anisops canaliculata n. sp.

(PI. XLIII, fig. 35)

Size. — Males, length 4.5 mm.-4.6 mm., greatest body width 1.3
mm.-1.4 mm.; females, length 4.9 mm. -5.4 mm., greatest body width
1.3 mm. -1.4 mm.

Shape. — Small species, only slightly fusiform; lateral margins in
anterior half of body subparallel, only slightly converging, converg-
ing in posterior half of body.

Color. — General facies testaceous. Eyes brown. Pronotum and
scutellum may be irregularly tinged with orange. Scutellum may
have two triangular black patches, one on each side of the anterior
half, projecting back from the posterior pronotal margin. Legs
testaceous. Abdominal venter black with keel and segmental mar-
gins of the connexivum testaceous.

368 The University Science Bulletin

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with its greatest width nine tenths the pro-
notal humeral width and at least four times the anterior width of the
vertex; synthlipsis wide, at least one third the anterior width of the
vertex; along the median longitudinal axis the head is at least one
half as long as the pronotum. Pronotum with its humeral width
at least twice its median length; lateral margins diverging and one
half the median length; posterior margin convex, medianly emar-
ginate. Facial tubercle with a median longitudinal groove. Labrum
short; basal width one and one half times its median length, apex
rounded; faint median depression extending from base to apex.
Rostral prong (PI. XLIII, fig. 35b) slightly shorter than third rostral
segment and with apex accumulate. Stridulatory comb of approxi-
mately ten teeth with basal five twice as long as apical five. Chaeto-
taxy of the male front leg as shown on Plate XLIII, fig. 35a. The
relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. -seg.

Fore lest 100 127 61

Middle leg 100 80 36 24

Hind leg 100 82 33 35

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with its greatest width nine tenths the
pronotal humeral width and at least four times the anterior width
of the vertex; synthlipsis wide, slightly less than one half the ante-
rior width of the vertex; along the median longitudinal axis the head
is three fifths the length of the pronotum. Pronotum with its hu-
meral width twice its median length; lateral margins diverging and
one half the median length; posterior margin convex, medianly
emarginate. The relative lengths of the parts of the legs are as
follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 127 52 35

Middle leg 100 75 36 26

Hind leg 100 75 30 32

Location of types. — Male holotype, female allotype, three male
and seven female paratypes, N. Queensland, Barren River nr. Barren
Waters, 9-28-38, R. G. Wind in the Snow Entomological Collection.
Other paratypes are as follows: In the Harvard Museum of Com-
parative Zoology: one male, six females, Queensland, Australia,
Townesville, Mar. '32, Harv. Austr. Exped., Darlington.

Comparative notes. — Its small size, wide synthlipsis, and de-
pressed facial tubercle are strongly suggestive of A. nivea (Fabri-

Brooks: The Genus Anisops 369

cius). However, the males of the two species serve to distinguish
them quite readily. The males of A. canaliculata lack the three tufts
of labral hairs and the peculiarly curved tarsal claws of the middle
leg as found on A. nivea. For comparison of the chaetotaxies of the
male front legs see PI. XLIII, fig. 35a and PI. XLIII, fig. 39a.

Data on distribution:

Australia
New Queensland, Townesville, 1920, G. F. Hill, five males, six females
(Bueno Coll. of F. H. Snow Coll.).

Anisops psyche Hutchinson

(PI. XLIII: fig. 38)

1928. Anisops psyche Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. 1, pp. 159-160, text
fig. 3.

1929. Anisops psyche, Hutchinson, Ann. South African Mus., vol. XXVr, pp. 400-402,
PI. XXX, fig. 20; PI. XXXII, fig. 3.

1930. Anisops psyche, Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. VI, p. 58 (eco-
logical note).

1930. Anisops psyche, Hutchinson, Proc. Zool. Soc. London, vol. XXIX, pt. 2, pp. 449-
450 (ecological note).

1932. Anisops psyche, Hutchinson, Proc. Zool. Soc. London, vol. XXXI, pt. 1, p. 125
(ecological note).

1934. Anisops psyche*. Poisson, Mus. Nat. Hist. Natur., vol. HI, p. 211 (ecological note).

Size. — Males, length 5.7 mm., greatest body width 1.5 mm.; fe-
males, length 5.7 mm., greatest body width 1.5 mm.

Shape. — Small, fusiform species; greatest body width midway the
body length.

Color. — General facies brown. Eyes brown. Vertex and pro-
notum testaceous, the latter may be hyaline and appear the color
of the scutellum. Scutellum orange or testaceous. Hemelytra
hyaline and appear the brown color of the underlying scutellum.
Legs stramineous. Abdominal venter dark brown with keel and
segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the anterior margin almost straight;
greatest width of the head eight tenths the pronotal humeral width
and almost five times the anterior width of the vertex; synthlipsis
wide, one half the anterior width of the vertex; along the median
longitudinal axis the head is almost one half the length of the pro-
notum. Pronotum with its humeral width twice the median length;
lateral margins diverging and three fifths the median length; pos-
terior margin convex, medianly emarginate. Facial tubercle exca-
vate with lateral margins carinate. Labrum short, basal width twice
the median length; apex rounded. Rostral prong (PI. XLIII, fig.

24—3286

370 The University Science Bulletin

38b) shorter than the third rostral segment; apex accuminate.
Stridulatory comb (PL XLIII, fig. 38c) of approximately eight even-
length teeth. Keel of second abdominal segment laterally expanded
and triangularly excavate; lateral margins carinate and bearing long
erect hairs; excavation lined with short, procumbent hairs. Con-
nexivum of the third abdominal segment with a row of nodules on
its ventral surface which becomes progressively smaller toward the
apex, nodules continued onto the connexivum of the fourth ab-
dominal segment. Chaetotaxy of the front leg as shown on Plate
XLIII. The relative lengths of the parts of the legs are as follows: *

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120

Middle leg 100 82 36 28

Hind leg 100 83 36 36

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head nine tenths the
pronotal humeral width and five times the anterior width of the
vertex; synthlipsis wide, slightly more than one half the anterior
width of the vertex; along the median longitudinal axis the head is
at least one half the length of the pronotum. Pronotum with its
humeral width almost three times the median length; lateral margins
diverging and slightly more than one third the median length;
posterior margin convex, medianly emarginate. The relative lengths
of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 122 55 35

Middle leg 100 82 36 28

Hind leg 100 83 34 34

Location of types. — Type material from Kampala, Uganda in the
British Museum.

Comparative notes. — Very closely allied to Anisops hancocki
Hutchinson from which it differs by the males not having the fore
femur greatly enlarged dorso-ventrally just before the base. For
comparison of the chaetotaxies of the front legs of the males see
Plate XLIII, fig. 38a and Plate XLVI, fig. 52a.

Data on distribution:
Africa

Kueni River, VI-27-1927, one male, gift to H. B. H. fr. G. E. Hutchinson
(F. H. Snow Coll.).

Salisbury, Makahiri River, VII-3-1927, one female, gift to H. B. H. fr. G. E.
Hutchinson (F. H. Snow Coll.).

* Tarsi of single male specimen lacking.

Brooks: The Genus Anisops 371

Anisops gen]i Hutchinson

(PI. XLIII. fig. 36)

192V. Anisops genji. Hutchinson, Ann. Mag. Nat. Hist., Ser. 9, vol. XIX, pp. 377-378,
text fig. 2.

1929. Anisops genji, Hutchinson, Ann. South African Mus., vol. XXV, p. 383, PI. XXX,
fig. 5.

1933. Anisops genji, Lundblad, Arch, fur Hydrob., Suppl. vol. XII, p. 14") (a list of Indo-
australian and Pacific forms of this genus).

1933. Anisops genji, Lundblad, Sartryck ur Entomogolisk Tidskrift Haft. 3-4, pp. 265-
267, text figures 8, A, B. C, D, E, F, G, & PI. 15 lower right fig.

1941. Anisops genji, Hoffmann, Lingnan Sci. Jour., vol. XX, p. 59 (catalogue).

Referring to this species also:

1896. Anisops Scutellaria, {nee Herrich-Shaffer) filler, Proc. U. S. Nat. Mus., Vol. VII,
p 275 (believed by Hutchinson to be Anisops genji).

1928. Anisops fieberi, Jaczewski, Ann. Mus. Zool. Polonici, vol. VII, p. 113, PI. XVI, figs.
18, 19, 20, 21 (his drawings are of Anisops genji Hutch, and not of Anisops nasuta Fieb. (.4.
fieberi Kirk.).

Size. — Males, length 5.8 mm. -6.6 mm., greatest body width. 1.6
mm.-1.8 mm.; females, length 6.1 mm.-6.6 mm., greatest body width
1.6 mm.-1.8 mm.

Shape. — Fusiform species; greatest body width about one third
its length.

Color. — Eyes gray or brown. Vertex and pronotum testaceous,
the latter may be hyaline on its posterior margin and appear the
color of the underlying scutellum. Scutellum testaceous, black, or
black with its apex testaceous or orange. Hemelytra hyaline and
appear the underlying color of the dorsal body surface which may
be black, or testaceous with the apical abdominal segments alter-
nating brown and black. Legs stramineous. Abdominal venter
black with keel and segmental margins of the connexivum stramin-
eous or brown.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the vertex extending slightly beyond the
anterior margins of the eyes; greatest width of the head nine tenths
the pronotal humeral width and four to five times the anterior width
of the vertex; synthlipsis wide, one third the anterior width of the
vertex; along the median longitudinal axis the head is slightly more
than one half as long as the pronotum. Pronotum with its humeral
width almost twice its median length; lateral margins diverging and
almost one half the median length; posterior margin convex, me-
dianly emarginate. Frons triangularly excavate, bordered on each
side by two carinae, the outer of which meet apically to form an
accuminate apex. Anterior interocular space with a median carina
which meets the accuminated apex of the frons. Labrum short;
basal width one and two thirds its median length; apex accumulate;

372

The University Science Bulletin

each basal angle of the labium bears a long tuft of hairs which curve
anteriorly along the outer carinae of the frons. Rostral prong (PI.
XLIII, fig. 36b ) slightly shorter than the third rostral segment; apex
accumulate. Stridulatory comb (PI. XLIII, fig. 36c) of approxi-
mately thirteen even-length teeth. Chaetotaxy of the male front leg
as shown on Plate XLIII. The relative lengths of the parts of the
legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 103 64

Middle leg 100 81 42 22

Hind leg 100 85 36 36

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with the anterior margin of the vertex
extending slightly beyond that of the eyes; greatest width of the
head nine tenths the pronotal humeral width and five to six times the
anterior width of the vertex; synthlipsis wide, slightly more than
one third the anterior width of the vertex; along the median longi-
tudinal axis the head is slightly more than one half the length of the
pronotum. Pronotum with its humeral width slightly more than
twice its median length; lateral margins diverging and over one half
the median length; posterior margin convex, medianly emarginate.
The relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 114 57 33

Middle leg 100 83 42 25

Hind leg 100 81 35 35

Location of types. — Male type, two male and one female para-
types, "Japan" in the collection of G. E. Hancock.

Comparative notes. — The excavate facial tubercle, labral hairs,
and the curved middle tarsal claws of the males strongly relate this
species to A. nivea (Fabricius). However, the excavation of A.
genji extends the length of the frons whereas on A. nivea it is re-
stricted to the facial tubercle. Also the excavation of the latter
species is bordered on each side by only one carina whereas in
A. genji it is bordered on each side by two carinae. In addition, the
median tuft of labral hairs as found on A. nivea is lacking in this
species.

Data on distribution:
Japan

Shimbara Peninsula, Unzen, 2200', Sept. 4, 1937, E. Suenson, eighteen
males, fifty-six females (F. H. Snow Coll.).

Kobe, Baker, two males, nine females ( U. S. Nat. Mus. ).

Brooks: The Genus Anisops 373

China

Soochow, fr. N. Gist. Gee, five males, ten females (F. H. Snow Coll.).

Shanghai, E. Suenson, one female (F. H. Snow Coll.).

Peking, X-13-25, P. W. Claassen, Tsing Huo Col, twenty-one males,
twenty-two females (F. H. Snow Coll.).

Peking, X-13-24, P. W. Claassen, Tsing Huo Col., eight males, six females
(F. H. Snow Coll.).

Peking, X-15-24, P. W. Claassen, Tsing Huo Col., three males, three fe-
males ( F. H. Snow Coll. ) .

Peking, XI-10-24, P. W. Claassen, Tsing Huo Col., three males, ten females
(F. H. Snow Coll.).

Manchuria, Yablonya St., VII-20-1940, via Weyman, one male, one female
(F. H. Snow Coll.).

Manchuria, Dairen, X-8-40, Michael Weyman, two males (F. H. Snow
Coll.).

Shanghai, Aug. 20, 1939, via Weyman, one male, one female (F. H. Snow
Coll.).

Manchuria, X-15-38, M. J. Nikitin, one male ( F. H. Snow Coll.).

Soochow, Chenfu F. Wu, one male (F. H. Snow Coll.).

Peking, Chenfu F. Wu, two females (F. H. Snow Coll.).

Foochow, Chenfu F. Wu, one female ( F. H. Snow Coll. ) .

Hangchow, Chenfu F. Wu, one female ( F. H. Snow Coll.).

Szechwan, Chenfu F. Wu, one male (F. H. Snow Coll.).

Suifu Szechwan, D. C. Graham, twelve males, thirteen females (F. H. Snow
Coll.).

Anisops nivea (Fabricius)

(PI. XLIII, fig. 39; PI. LIV, fig. 93)

1775. Notonecta nivea Fabricius, Systema Entomologiae. Flenaburgi et hipsiae, p. 690.

1794. Notonecta nivea, Fabricius, Entomologia Systema, vol. IV. p. 58.

1803. Notonecta nivea, Fabricius, Systema Rhyngatorum, p. 103 (list of species of Noto-
necta).

1810. Anisops nivea, Spinola, Essais sur les Insectes Hemipteres Rhyngotes ou Heterop-
teres, p. 58 (determined in error, actually A. sardea Herrich-Shaffer).

1840. Anisops nivea, Rambur, Faune entomologique de l'Andalouise, vol. II, p. 191 (de-
termined in error, actually A. sardea Herrich-Shaffer).

1847. Anisops nivea, Costa, Atti del Reale Instituta L'Incoraggiamento Alle Scienze Nat-
urali di Napoli, vol. VII, p. 148 (determined in error, actually A. sardea Herrich-Shaffer).

1848. Anisops niveus, Amyot, Entomologie Franciase, Rhynchotas, p. 338 (determined in
error, actually A. sardea Herrich-Shaffer).

1851. Anisops niveus, Fieber, Abhandl. Konigl. bohm. Ges. Wiss., vol. 7, p. 484 (deter-
mined as A. niveus though in parenthesis he named it A. nasuta).

1899. Anisops niveus, Kirkaldy, Ann. Soc. Ent. France, vol. LXVIII, p. 105 (determined
in error, probably A. pellucens Gerst).

1901. Anisops niveus, Kirkaldy, The Entomologist, vol. XXXIV. p. 5 (determined in er-
ror).

1904. Anisops niveus, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 118-119, 132 (deter-
mined in error, probably A. pellucens Gerst.).

1905. Anisops nivea, Matsumura, Trans. Sapporo Nat. Hist. Soc, vol. I, p. 28 (deter-
mined in error, actually A. nasuta Fieber).

1906. Anisops niveus, Distant. Fauna of British India, Rhynchota, vol. Ill, p. 46 (deter-
mined in error).

1908. Anisops nivea, Kirkaldy, Y. sjostedt : Wissenschaften Ergebn. der schwed. zool.
Exped. Hemiptera. XII, p. 24 (determined in erroi . probably A. pellucens Gerst).

374 The University Science Bulletin

1915. Anisops niveus, Esaki, Ent. Mag., Kyoto, vol. I, p. 31 (determined in error, actually
A. nasuta Fieber).

1915. Anisops niveus, Matsuniura, Ent. Mag., Kyoto, vol. I, p. 110.

1918. Anisops niveus, Paiva, Rec. Ind. Mus., vol. XIV, p. 27 (ecological note).

1919. Anisops niveus, Paiva, Rec. Ind. Mus., vol. XV, p. 13 (ecological note).

1926. Anisops niveus, Esaki, Ann. Mus. Nat. Hungarici, vol. XXIV, p. 187 (determined
in error).

1927. Anisops niveus, Dover, Jour. Bombay Nat. Hist. Soc, vol. XXXII, p. 615.

1927. Anisops niveus, Torre Bueno, Bull. Brooklyn Ent. Soc, vol. XXII, p. 30 (determina-
tion questionable).

1928. Anisops nivea, Esaki, Insects of Samoa II, p. 76 (mentioned in footnote the correct
location of type).

1929. Anisops niveus, Hutchinson, Ann. South African Mus., vol. XXV, p. 385 (mentions
the need of a closer study of the type, as the species seems to be composite).

1929. Anisops niveus, Poisson, Faune des Colonies Francaises, vol. Ill, p. 154 (determined
in error, actually A. pellucens Gerst).

1933. Anisops niveus, Hoffmann, Lingnan Sci. Jour., vol. XII, pp. 255-256 (catalogue; of
little value for at least two species are mixed).

1933. Anisops niveus, Wu, Lingnan Sci. Jour., vol. XII, pp. 213-214 (catalogue; of little
value for at least two species are mixed).

1933. Anisops niveus, Lundblad, Arch, fiir Hydrob., Suppl. vol. XII, pp. 145, 163-166,
text fig. 56.

1935. Anisops niveus, Wu, Catalogus Insecturum Sinensium, vol. II, p. 576.

1941. Anisops niveus, Hoffmann, Lingnan Sci. Jour., vol. XX, pp. 60-61 (catalogue; this
species omitted as it is composite ; therefore distribution questionable).

Size. — Males, length 4.8 mm. -5.1 mm., greatest body width 1.2
mm.-1.4 mm.; females, length 4.8 mm.-5.4 mm., greatest body width
1.3 mm.-1.8 mm.

Shape. — Fusiform species, greatest body width about one half
the length of the body.

Color. — General facies pearlaceous. Eyes brown. Legs stramin-
eous. Abdominal venter dark brown with keel and segmental mar-
gins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the greatest width of the head nine
tenths the pronotal humeral width and four to five times the anterior
width of the vertex; synthlipsis wide, approximately one third the
anterior width of the vertex; along the median longitudinal axis the
head is seven eighths the length of the pronotum. Pronotum with
its humeral width slightly more than twice the median length; lateral
margins diverging and slightly more than one half the median
length; posterior margin convex, medianly emarginate. Facial
tubercle with a triangular excavation bordered on each side by a
raised carina. Labium broad and short; basal width almost twice
its median length; apex more or less accumulate; base with a trans-
verse row of long hairs with the outer one on each side grouped
together to form a tuft which curves anteriorly on the excavate
facial tubercle, the inner group erect. Rostral prong (PL LIX, fig.

Brooks: The Genus Anisops 375

93 ) longer than the third rostral segment; apex accumulate. Stridu-
latory comb (PL XLIII, fig. 39b) of approximately fifteen even
length teeth. Middle tarsal claws strongly curved inward at base;
posterior claw thicker than anterior one. Chaetotaxy of the front
leg as shown on Plate XLIII. The relative lengths of the parts of the
legs are as follows:

lsl 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 102 79

Middle leg 100 85 40 25

Hind leg 100 78 33 33

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with the greatest width of the head
eight to nine tenths the pronotal humeral width and four to five
times the anterior width of the vertex; synthlipsis wide, almost one
half the anterior width of the vertex; along the median longitudinal
axis the head is three fifths the length of the pronotum. Pronotum
with its humeral width slightly more than its median length; lateral
margins diverging and six tenths the median length; posterior mar-
gin convex, medianly emarginate. The relative lengths of the parts
of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 116 60 33

Middle leg 100 79 38 22

Hind leg 100 83 33 33

Location of types. — Fabricius' type material from India located
in the Keil Museum.

Comparative notes. — This species is closely related to A. genji
Hutch., but may be readily separated from it on the basis of the
males. The males of A. nivea have the excavation restricted to the
facial tubercle whereas in A. genji the excavation extends almost
the full length of the frons. The excavation is bordered by one
carina on each side in the former species whereas in the latter the
excavation is bordered by two carinae on each side.

Bibliographical notes. — Kirkaldy in 1904 (30) as a result of a
study of the type of Anisops ciliata Stal and what he erroneously
believed to be the type of A. niveus ( Fabricius ) declared the two
to be the same and placed several other species into synonomy there-
with. It has since been shown by Lundblad (32) that what
Kirkaldy thought was the type of A. nivea was not the type at all.
Many of the references that Kirkaldy placed with A. nivea I have
been able to place with their correct species, others defy determina-
tion due to poor or no accompanying description.

376 The University Science Bulletin

Notonecta ciliata Fabricius which both Stal and Kirkaldy felt
was an Anisops, is actually an Enithares. However, Anisops ciliata
Stal from the He de France is a valid Anisops species. Anisops
hyalinus Fieber was first placed as a possible synonym of A. ciliata
by Stal and later by Kirkaldy as a possible variety of A. ciliata only
ever, it is much too large for A. nivea, and resembles A. ciliata only
in size. I have found no species in my vast Asiatic material which
fits the description of A. hyalinus, so I shall present a translation of
the original description. Though highly inadequate, it is the only
description of the species.

Data on distribution:

Sumatra
Sumatra, Thienemanni, one male (gift from Lundblad) (F. H. Snow Coll.)

India

Ganjam Dist., Barkuda, Chilka Lake, Madras Pres., VIII-18, F. H. Gravely,
fifteen males, fifteen females (F. H. Snow Coll.); seventeen males, nine females
( Indian Museum ) .

Ganjam Dist., Barkuda, Chilka Lake, Madras Pres., XI-23-19, N. A., eight
males, twenty females (Indian Museum).

Ganjam Dist., Barkuda, Chilka Lake, Madras, N. A., one female (Indian
Museum ) .

Ganjam Dist., Bambha, Madras, IX-20-13, Annandale, one male, seventeen
females (Indian Museum), four males, three females (F. H. Snow Coll.).

Madras, Barkuda Id., Chilka Lake, VIII-4-7, one male, one female (Bueno
Coll. of F. H. Snow Coll.).

Br. India, Coimbatore, L. V. Newton S. J. one male, one female (Bueno
Coll. of F. H. Snow Coll.).

Anisops tahitiensis Lundblad

(PI. XLIII, fig. 40)

1934. Anisops tahitiensis Lundblad, Bull. Bernice Pauki Bishop Mus. Poly. Ethn. Nat.
Hist,, vol. 113, pp. 121-123, text figs. 1-5.

Size. — Males, length 5.1 mm. -5. 5 mm., greatest body width 1.3
mm. -1.5 mm.; females, length 5.1 mm. -5.5 mm., greatest body width
1.3 mm. -1.5 mm.

Shape. — Slightly fusiform species; greatest body width almost
midway the body length.

Color. — Gray form: General facies peralaceous. Legs stramine-
ous. Abdominal venter dark brown with keel and segmental mar-
gins of the connexivum stramineous. Brown form: Eyes brown.
Vertex and pronotum testaceous, the latter may be hyaline on its
posterior margin and appear dark brown or black due to the under-
lying color of the scutellum. Scutellum testaceous or dark brown

Brooks: The Genus Anisops 377

with its lateral margins orange. Hemelytra hyaline and appear
stramineous or dark brown depending on the color of the dorsal
body surface. Legs stramineous. Abdominal venter dark brown
or black with keel and segmental margins of the connexivum
stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the anterior margin almost straight;
greatest width of the head nine tenths of the pronotal humeral width
and seven times the anterior width of the vertex; synthlipsis wide,
slightly more than one third the anterior width of the vertex; along
the median longitudinal axis the head is slightly more than three
fourths the length of the pronotum. Pronotum with its humeral
width approximately twice its median length; lateral margins di-
verging and from one half to three fifths the median length;
posterior margin convex, medianly emarginate. Facial tubercle
laterally compressed to form a median carina which proceeds
anteriorly for the basal two thirds of the frons. Labrum with basal
width one and one half times its median length; apex rounded.
Rostral prong (Pi. XLIII, fig. 40b) longer than the third rostral
segment; apex accumulate. Labrum with its basal width one and
one half its median length. Stridulatory comb (Pi. XLIII, fig. 40c)
of approximately twenty-one teeth. Chaetotaxy of the male front
leg as shown on Plate XLIII. The relative lengths of the parts of
the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 111 77

Middle leg 100 84 77 27

Hind leg 100 81 30 31

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with the greatest width of the head
nine tenths the pronotal humeral width and five and one half times
the anterior width of the vertex; synthlipsis wide, one half the
anterior width of the vertex; along the median longitudinal axis the
head is three fifths as long as the pronotum; lateral margins diverg-
ing and one half as long as the pronotum; posterior margin convex,
medianly emarginate. The relative lengths of the legs are as fol-
lows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 115 50 34

Middle leg 100 77 36 25

Hind leg 100 87 31 32

378 The University Science Bulletin

Location of types. — Lundblad's type material, Tahiti, July 20,
1925, L. E. Cheesman, is in the British Museum.

Comparative notes. — This species is about the same size and
general appearance as Anisops cleopatra Distant from which it
can be separated on the basis of the males. The males of A.
cleopatra lack the carinate frons and the rounded apex of the
anterior femur. Also the head of the males of A. tahitiensis is
much larger, the eyes being more voluminous, and almost straight
on its anterior margin whereas the head of A. cleopatra has a
curved anterior margin and the eyes are not voluminous.
Data on distribution:

Guadalcanal
Jan., 1945, P. H. Eschmeier, four males, eight females (F. H. Snow Coll.).
1944, L. J. Lipovsky, twenty-three males, twenty-seven females ( F. H.
Snow Coll.).

New Guinea
Rigo, 1889, L. Loria, two males, one female (F. H. Snow Coll.).

Andaman Islands
nr. W. Goast IsL, III-1914, C. J. Rogers, five males, eight females (Indian
Museum).

New Hebrides
March 15, 1943, Oman, two males, four females (U. S. Nat. Mus. ).

Philippine Islands
Mindanao, Davao, Raker, two males, one female (U. S. Nat. Mus.).

Okinawa
Sept. 18, 1945, W. D. Fields, six males, eight females (U. S. Nat. Mus.).

Anisops tasmaniaensis n. sp.

(PI. XLVII, fig. 5S)

Size. — Males, length 7.5 mm. -7.9 mm., greatest body width 2.4
mm.-2.5 mm.; females, length 7.8 mm.-7.9 mm., greatest body width
2.1 mm.-2.3 mm.

Shape. — Moderately large species; fusiform, with greatest body
width midway the body length.

Color. — General facies stramineous. Eyes brown. Legs stramin-
eous. Abdominal venter black with keel and segmental margins
of the connexivum brown.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of head equal to pronotal
humeral width, slightly more than seven times the anterior width
of the vertex; synthlipsis wide, three fourths the anterior width of
the vertex; along the median longitudinal axis the head is almost
three fourths the pronotal length. Pronotum with its humeral width
twice its median length; lateral margins diverging and slightly more

Brooks: The Genus Anisops 379

than one half the median length; posterior margin convex, medianly
emarginate. Facial tubercle slightly raised. Labrum long, me-
dianly length equal to its basal width; apex rounded. Rostral prong
(PL XLVII, fig. 58b) slightly shorter than third rostral segment;
apex accuminate. Stridulatory comb (PL XLVII, fig. 58c) of ap-
proximately twenty-two teeth; basal five slightly shorter than the
others. Chaetotaxy of front leg as shown on Plate XLVII. The
relative lengths of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 127 83

Middle leg 100 83 41 24

Hind leg 100 SO 32 32

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded, with the vertex extending slightly be-
yond the anterior margins of the eyes; greatest width of the head
nine tenths the pronotal humeral width, five times the anterior width
of the vertex; synthlipsis wide, two thirds the anterior width of the
vertex; along the median longitudinal axis the head is almost three
fourths the pronotal length. Pronotum with its humeral width twice
the median length; lateral margins diverging and slightly more than
one half the median length; posterior margin convex, medianly
emarginate. The relative lengths of the parts of the legs are as

follows: 1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 126 58 32

Middle leg 100 76 42 21

Hind leg 100 81 34 32

Location of types. — Male holotype, female allotype, two male and
one female paratypes, Tasmania, Lake Leake, XI-1937, J. W. Evans,
in the Snow Entomological Collection.

Comparative notes. — Though not quite as broad as Anisops bar-
hata n. sp., it appears very similar to it. However, the males of
this species lack the tufts of hairs as found on the male facial tu-
bercle of A. barbata. For comparison of the male front legs of the
two, see Plate XLVII, fig. 58a and Plate XLVI, fig. 54a.

Data on distribution. — Known only from type series.

Anisops lipovskyi n. sp.

(PI. XLIII, fig. 37)

Size. — Males, length 5.4 mm.-6.0 mm., greatest body width 1.5
mm. -1.7 mm.; females, length 6. mm., greatest bodv width 1.6 mm.

Shape. — Small, fusiform species with its greatest width midway
its body length.

380

The University Science Bulletin

Color. — General facies pearlaceous. Eyes brown. Hemelytra
partly hyaline and such areas appear darker as they overlie the
dark brown dorsal body surface. Legs stramineous. Abdominal
ventor dark brown with keel and lateral margins stramineous.

Male structural characteristics. — As viewed from above, the out-
line of the head is rounded, with the anterior margin of the vertex
extending slightly beyond that of the eyes; greatest width of head
nine tenths the pronotal humeral width and seven times the anterior
width of the vertex; synthlipsis wide, approximately one third the
anterior width of the vertex; along the median longitudinal axis
the head is at least one half the pronotal length. Pronotum with
its humeral width about twice its median length; lateral margins
diverging and one half the median length; posterior margin convex,
medianly emarginate. Facial tubercle laterally compressed form-
ing a median carina which extends anteriorly on the frons to the
apex. Rostral prong (PI. XLIII, fig. 37b) longer than third rostral
segment; apex accuminate. Labrum short; basal width one and
one third its median length, apex broad and rounded. Anterior
femur (PI. XLIII, fig. 37a) truncate at apex. Stridulatory comb
(PL XLIII, fig. 37c) of about twenty-eight teeth; greatest width at
basal third. The relative lengths of the parts of the legs are as
follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 110 71

Middle leg 100 81 38 28

Hind leg 100 82 39 39

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width nine tenths the pronotal
humeral width and at least five times the anterior width of the
vertex; synthlipsis wide, slightly less than one half the anterior
width of the vertex; along the median longitudinal axis the head is
approximately three fifths the pronotal length. Pronotum with
humeral width at least twice its median length; posterior margin
convex, medianly emarginate. The relative lengths of the parts of
the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 114 51 33

Middle leg 100 78 39 26

Hind leg 100 83 31 31

Location of types. — Male holotype, female allotype Jor Pokri,
Sitong, Darjiling Dist. E. Himalayas, India, X-22-17, N. A. and
F. G. Paratypes are as follows. In the Indian Museum: One male,

Brooks: The Genus Anisops 381

Jor Pokri, 4800', Darjiling, Dist. E. Himalayas, India, VII-6-18,
S. Kemp; one male, one female, Jor Porki 4800' nr. Sitong, Dar-
jiling Dist. E. Himalayas, India, VII-6-18, S. Kemp. In the Snow
Entomological Collection; one male, Calcutta, India, in pool, XI-
15-11; two males, Jor Pokri, 4800' Darjiling Dist. E. Himalayas,
India, VI-6-18.

Comparative notes. — This species appears closely related to Ani-
sops tahitiensis Lundblad, the males of both species have a carinate
frons, an apically truncate femur and similar chaetotaxies on the
front legs. The head serves best to distinguish the two, that of
Anisops tahitiensis being broad and almost straight along its an-
terior margin whereas in Anisops lipovskyi the head is more or less
conical shape with the vertex extending slightly beyond the eyes
at the anterior margin.

Anisops deanei n. sp.

(PI. XLIV, fig. 45)

Size. — Males, length 5.4 mm. -6.0 mm., greatest body width 1.3
mm. -1.5 mm.; females, length 5.7 mm. -6.3 mm., greatest body width,
1.5 mm. -1.7 mm.

Shape. — Fusiform, greatest body width midway the body length.

Color. — General facies stramineous. Eyes brown or gray. Scu-
tellum may be irregularly tinged with crimson. Hemelytra hyaline
and appear stramineous, red, or dark brown, depending on the
dorsal body surface color. Legs stramineous. Abdominal venter
dark brown with keel and segmental margins of the connexivum
stramineous. Some forms are completely gray with the abdominal
venter as above.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of the head nine tenths the
pronotal humeral width, approximately seven times the anterior
width of the vertex; synthlipsis variable, from slightly less than
one third to almost one half the anterior width of the vertex; along
median longitudinal axis head three fourths the pronotal length.
Pronotal humeral width at least twice its median length; lateral
margins diverging and slightly more than one half the median
length; posterior margin convex, medianly emarginate. Facial
tubercle only slightly raised; bearing scattered hairs. Labrum with
basal width only slightly more than median length, apex rounded.
Rostral prong ( PL XLIV, fig. 45b ) slightly longer than third rostral
segment. Anterior femur slightly curved at apex. Stridulatory
comb (PI. XLIV, fig. 45c) of twenty-one to twenty-four even-length

382 The University Science Bulletin

teeth, apical one shorter than remainder. Chaetotaxy of front leg
as shown on Plate XLIV. The relative lengths of the parts of the
legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 119 74

Middle leg 100 81 38 26

Hind leg'. 100 82 34 33

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width nine tenths the pronotal
humeral width, six times the anterior width of vertex; width of
synthlipsis variable from approximately one third to almost one half
the anterior width of the vertex; along the median longitudinal axis
the head is slightly more than three fourths the pronotal length.
Pronotum with humeral width at least twice its median length;
lateral margins diverging and more than one half the median length;
posterior margin convex, medianly concave. The relative lengths
of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 123 62 35

Middle leg 100 78 39 27

Hind leg 100 87 36 39

Location of Types. — Male holotype, female allotype, sixteen males
and ten female paratypes, New South Wales, Began River, F. Arm-
strong, via Mr. Deane in the Snow Entomological Collection. Other
paratypes are as follows. In the Snow Entomological Collection:
two males, three females, Queensland, St. George, Dist. 5, 1923, G.
H. Williams; sixteen males, eleven females, Victoria Alexandria, F.
L. Billinghurst (Kirkaldy Collection) in the Snow Entomological
Collection.

Comparative notes. — This species is very similar to Anisops Hype-
rion Kirkaldy.* The males of the two species offer a means of ready
separation. The apex of the male front femur of A. deanei is slightly
curved, whereas that of A. Hyperion is accumulate. Also the apex
of the third rostral segment of the male of A. deanei is decidely
wider than the base of the fourth, whereas in A. Hyperion the two
areas are of about the same width. For comparison of the chaeto-
taxies of the front legs of the males see Plate XLIV, fig. 45a and
Plate XXXVIII, fig. 16a.

Data on distribution:

Australia
Victoria, Bacchus Marsh, 1904, six males, two females ( Bueno Coll. of F. H.
Snow Coll.).

* nee. Anisops hyperion H;ile.

Brooks: The Genus Anisops

383

Anisops philippinensis n. sp.

(PI. XLIV, fig. 44)

Size. — Males, length 5.4 mm.-5.8 mm., greatest body width 1.2
mm.-1.3 mm.; females, length 5.8 mm.-6.0 mm., greatest body width
1.5 mm. -1.6 mm.

Shape. — Small, broad-headed species; body fusiform with greatest
width about midway the body length.

Color. — General facies dark gray. Vertex, pronotum, and scu-
tellum testaceous, the latter may be black with only the anterior
margin testaceous. Hemelytra hyaline and appearing dark gray
as it overlies the black dorsal body surface. Legs stramineous.
Abdominal venter black with keel and segmental margins of the
connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the anterior margin almost straight;
greatest width of head nine tenths to almost equal the pronotal
humeral width, ten times the anterior width of the vertex; synthlipsis
wide, one third the anterior width of the vertex; along the median
longitudinal axis the head is nine tenths, to equal the pronotal length.
Pronotum with its humeral width slightly more than twice its median
length; lateral margins diverging and at least one half the median
length; posterior margin convex, medianly emarginate. Facial
tubercle only slightly raised. Labrum with its basal width only
slightly more than its median length; apex more or less accuminate.
Rostral prong ( Pi. XLIV, fig. 44b ) slightly shorter than third rostral
segment; apex accuminate. Stridulatory comb (Pi. XLIV, fig. 44c)
of approximately seventeen teeth whose lengths increase from base
to apex. Chaetotaxy of the male front leg as shown on Plate XLIV.
The relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 83

Middle leg 100 85 40 28

Hind leg 100 87 30 33

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with its greatest width nine tenths the
pronotal humeral width and seven times the anterior width of the
vertex; synthlipsis wide, one half the anterior width of the vertex;
along the median longitudinal axis the head is almost three fifths as
long as the pronotum. Pronotum with its humeral width slightly
more than twice its median length; lateral margins almost three

384 The University Science Bulletin

fifths the median length; posterior margin convex, medianly emargi-
nate. The relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 123 51 35

Middle leg 100 78 37 28

Hind leg ". 100 84 31 33

Location of types. — Male holotype, female allotype, forty-eight
male and twenty female paratypes, Philippine Isl., Mindanao, Lake
Linau, N. slope, Mt. Apo, Davao Prov. 7800', 46, H. Hoagstraal and
F. G. Werner, Zoological Expedition, in the Chicago Natural History
Museum. Other paratypes are: ten males, ten females, same col-
lection data as above, in the Snow Entomological Collection.

Comparative notes. — Of about the same size and general appear-
ance as Anisops tahitiensis Lundblad from which it can be readily
separated by the fact that the males do not posses the carinate frons
and the apically rounded front femur as found on the males of A.
tahitiensis.

Data on distribution. — Known only from type series.

Anisops cleopatra Distant

(PI. XLIV, fig. 43)

1914. A?risops cleopatra Distant, Nova Caledonia, Zoologie, vol. I, p. 386, PI. XI, fig. 8.

19-23. Anisops cleopatra, Hale, Rec. South Austr. Mus., vol. II, no. 3, p. 413 (only a men-
tion in a footnote).

1928. Anisops cleopatra, Esaki, Insects of Samoa, Part II, pp. 77-78, text fig. 5 (ecological
note).

1933. Anisops cleopatra, Lundblad, Arch, fiir Hydrob., Suppl. vol. XII, pp. 145, 171-173,

text figs. 59-62.

Size. — Males, length 5.4 mm. -5.8 mm., greatest body width 1.4
mm. -1.7 mm.; females, length 5.8 mm.-6.3 mm., greatest body width
1.5 mm. -1.8 mm.

Shape. — Fusiform species, with greatest body width about mid-
way the body length.

Color. — General facies dark gray or black. Eyes dark brown or
gray. Vertex and pronotum testaceous, the latter hyaline on
posterior half and appearing the black color of the underlying
anterior margin of the scutellum. Scutellum black with apex and
lateral margins testaceous or red orange. Hemelytra hyaline and
appearing dark gray or black due to the underlying color of the
dorsal body surface. Legs stramineous. Abdominal venter black
with keel and segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head nine tenths

Brooks: The Genus Anisops 385

the pronotal humeral width and seven times the anterior width of
the vertex; synthlipsis wide, one half the anterior width of the
vertex; along the median longitudinal axis the head is two thirds
as long as the pronotum. Pronotum with humeral width slightly
more than twice its median length; lateral margins diverging and
three fifths the median length; posterior margin convex, medianly
emarginate. Facial tubercle only slightly raised. Labrum with its
basal width slightly more than its median length; apex truncate
and almost one third the basal width. Rostral prong (Pi. XLIV,
fig. 43b ) longer than the third rostral segment; apex truncate, though
very narrow in width. Stridulatory comb Pi. XLIV, fig. 43c) of ap-
proximately twenty-four teeth. Chaetotaxy of the male front leg
as shown on Plate XLIV. The relative lengths of the parts of the
legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 104 69

Middle leg 100 85 40 30

Hind leg 100 87 31 32

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of the head nine tenths
the pronotal humeral width and six times the anterior width of the
vertex; synthlipsis wide, slightly more than one half the anterior
width of the vertex; along the median longitudinal axis the head
is almost one half the length of the pronotum. Pronotum with its
humeral width two and one half times its median length; lateral
margins diverging and three fifths its median length; posterior mar-
gin convex, medianly emarginate. The relative lengths of the parts
of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 123 58 32

Middle leg 100 85 38 28

Hind leg 100 88 32 38

Location of types. — The type material from New Caledonia
should be in the British Museum. However, neither Esaki (8) nor
Lundblad (32) were able to find it. The latter is of the opinion
that it may have been destroyed.

Comparative notes. — This species is about the same size and
general appearance as A. tahitiensis Lundblad from which it can
be readily separated by the fact that the males lack the laterally
compressed facial tubercle and the carinate frons as found on A.
tahitiensis.

25—3286

386 The University Science Bulletin

Data on distribution:

New Caledonia

Nr. Noumea, July, 1940, F. X. Williams, eight males, ten females (F. H.
Snow Coll.).

Noumea, A. Fauvel, one female, Kirkaldy Collection (F. H. Snow Coll. K
New Caledonia, Marie Schastien, Kirkaldy Collection ( F. H. Snow Coll. ) .

Anisops praetexta Hutchinson

(PI. XLIV, fig. 41)

1929. Anisops praetexta Hutchinson, Ann. South African Mus., vol. XXV, pt. 3, pp. 402-
403, PI. XXX, fig. 19; PI. XXXII, fig. 4, 4a,

1934. Anisops praetexta, Poisson, Bull., Soc. Zool. France, vol. XXIX, p. 100 (ecological
note).

Size. — Males, length 5.4 mm., greatest body width 1.4 mm.; fe-
males, length 5.1 mm.-5.4 mm., greatest body width 1.4 mm. -1.6 mm.

Shape. — Small fusiform species, greatest body width about mid-
way the body length.

Color. — General facies brown. Vertex, pronotum, and scutellum
testaceous. Hemelytra hyaline and appears dark brown due to the
color of the dorsal body surface. Legs stramineous. Abdominal
venter dark brown with keel and segmental margins of the connexi-
vum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of the head almost nine
tenths the pronotal humeral width and slightly more than seven
times the anterior width of the vertex; synthlipsis wide, one third the
anterior width of the vertex; along the median longitudinal axis the
head is almost as long as the pronotum. Pronotum with its humeral
width two and one third times its median length; lateral margins
diverging and slightly more than one half the median length; pos-
terior margin convex medianly emarginate. Facial tubercle slightly
raised. Labrum with its basal width equal to its median length;
apex rounded. Rostral prong ( PL XLIV, fig. 41b ) longer than third
rostral segment. Stridulatory comb (PL XLIV, fig. 41c) of approxi-
mately twelve teeth. Chaetotaxy of male front leg as shown on
Plate XLIV. The relative lengths of the parts of the legs are as
follows :

1st _ 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 87

Middle leg 100 80 40 25

Hind leg 100 84 33 34

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head nine tenths the

Brooks: The Genus Anisops

387

pronotal humeral width and five to six times the anterior width of
the vertex; synthlipsis wide, slightly more than one third the anterior
width of the vertex; along the median longitudinal axis the head is
two third the length of the pronotum. Pronotum with its humeral
width two and one half times its median length; lateral margins
diverging and one half the median length; posterior margin convex,
medianly emarginate. The relative lengths of the parts of the legs
are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 117 52 37

Middle leg 100 83 40 25

Hind leg 100 85 30 33

Location of types. — Type material from Southern Rhodesia in the
South African Museum.

Comparative notes. — Very similar to Anisops amaryllis Hutch,
and superficially the two species are almost identical. However,
A. amaryllis does not have the short row of small setae on the basal
inner surface of the male fore tarsi, as found on A. praetexta.

Data on distribution:

Africa

Central Africa, one male, one female (Basel Nat. Hist., Switzerland) one
female (F. H. Snow Coll.).

Anisops barbata n. sp.

(PI. XLVI, fig. 54)

Size. — Males, length 8.6 mm. -9.3 mm., greatest body width 2.4
mm. -3 mm.; females, length 8 mm. -9.1 mm., greatest body width
2.8 mm.-3 mm.

Shape. — Robust, slightly fusiform species with the greatest body
width about midway the body.

Color. — Gray form: General facies gray. Eyes brown. Hemely-
tra may be partly hyaline and such areas appear darker as they
overlie the dark brown dorsal body surface. Legs testaceous. Ab-
dominal venter black with keel and segmental margins of the con-
nexivum testaceous. Stramineous form: General facies stramineous.
Eyes brown. Legs stramineous; abdominal venter dark brown with
keel and segmental margins of the connexivum stramineous.

Male structural characteristics. — As viewed from above, the out-
line of the head is rounded with the anterior margin of the eyes
slightly in advance of that of the vertex; greatest width of the head
nine tenths the pronotal humeral width and slightly more than
eight times the anterior width of the vertex; synthlipsis wide, at

Ms.s The University Science Bulletin

least two thirds the anterior width of the vertex; along the median
longitudinal axis the head is approximately one half the length of
the pronotum. Pronotum with its humeral width twice its median
length; lateral margins diverging and approximately one half the
median length; posterior margin convex, medianly emarginate;
dorsal surface with a faint median depression and a concavelv
curved ridge on either side, halfway between the median line and
the lateral margin. Facial tubercle with two tufts of hairs, one on
either side of the median line, which extend posteriorly to the base
of the labrum. Labrum with its basal width slightly more than its
median length, and with its apex more or less accuminate. Rostral
prongs ( PI. XLVI, fig. 54b ) with their length slightly less than that
of the third rostral segment; apex accuminate. Chaetotaxy of the
male front leg as shown on Plate XLVI. The relative length of the
parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 119 86

Middle leg 100 84 33 22

Hind leg 100 77 30 25

■-&

Female structural characteristics. — As viewed from above, the
outline of the head is rounded with its greatest width approxi-
mately nine tenths the pronotal humeral width and slightly more
than six times the anterior width of the vertex; synthlipsis wide,
two thirds the anterior width of the vertex; along the median longi-
tudinal axis the head is less than one half the pronotal lengths.
Pronotum with humeral width twice its median length; lateral mar-
gins diverging and one half the median length; posterior margin
convex, medianly emarginate; dorsal surface with same markings
as male only fainter. The relative lengths of the parts of the legs
are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 126 56 38

Middle leg 100 83 36 21

Hind leg 100 79 33 25

Location of types. — Male holotype, female allotype, nine male,
six female paratypes, Rambha, Ganjam Dist. Madras, India, IX-20-
13, Annandale, in the Indian Museum at Calcutta. Other paratypes
are as follows: In the Snow Entomological Collection: Seven males,
seven females (same collection data as above); one male, one fe-
male, Barkuda, Chilka Lake, Madras Pres., India, VIII-19, F. H.
Gravely; one male, Java, VIII-1939. In the Indian Museum at Cal-
cutta: One male, Waltair, India, 1-27-21; eight females, Barkuda,

Brooks: The Gents Anisops 389

Chilka Lake, G an jam Dist., Madras, India, IX-23-19, N. A.; two
males, Rambha, Ganjam, fr. an ornamental fountain, XII-30-18,
N. A.; three males, one female, Bangalore, S. India, ca 3000 ft.,
X-13-10, Annandale; two females, Markiuppon, S. India, ca 2500
ft.; one male, Barkul, Puri Dist. Orissa, India, XI-9 to 13-12, Gravely;
one female, Lucknow, 1-16-13, Mus. Collr. R. H.; one female Sata-
pare, Lake Dhilha, Orissa, IV-16-13, Annandale; two females, Ber-
hampur Murshidabad, Bengal, Sept. '12, Southwell. In the Usinger
Collection; two males, one female, Java, VIII-1939.

Comparative notes. — This species is of about the same size and
superficial appearance as Anisops stali Kirkaldy. It has, however,
two tufts of hairs on the facial tubercle that are lacking on the
facial tubercle of the latter species. Also A. stali has a slightly
depressed frons and a faint anterior cephalic projection, both are
lacking on A. barbata. For comparison of the chaetotaxies of the
male front legs see Plate XLVI, fig. 54a and Plate XXXVII, fig. 9a.

Data on distribution:

Burma

S. Shan States, Yawaghee, Heho, 3800', III-7-17, Gravely (F. H. Snow Coll.).
India

Kalka, base of Simla Hills, 2400', VII-16-11, one male, two females (F. H.
Snow Coll.).

S. India, Bangalore, 3000', X-16-10, Annandale, three males, one female
(F. H. Snow Coll.).

Bengal, Madhupur, IV-28-to V-7-11, C. Paiva, nine males, four females ( F.
H. Snow Coll.).

India, Kalka, base of W. Himalayas, V-16-I1, Annandale, three males, one
female (F. H. Snow Coll.).

Madras, Ganjam Dist., Rambha, IX-20-13, Annandale, nine males, fifteen
females (F. H. Snow Coll.).

Patiola State Dhurampur Kooa, base of Simla Hills, VII-2-11, one female
(F. H. Snow Coll.).

Brindia, Pulney Hills, L. V. Newton, S. J., two males, three females ( Bueno
Coll. of F. H. Snow Coll.).

Java
Java, Le Moult, one female (Bueno Coll. of F. H. Snow Coll.).

Formosa
Formosa one male (F. H. Snow Coll.).

Anisops grand is Poisson

(PI. XLV. fig. 50)
1937. Anisops pellucens grandis Poisson. Ann. Soc. Ent. France, vol. XVI, p. 130.

Size. — Males, length 9.0 mm.-9.3 mm., greatest body width 2.7
mm.; females, length 9.3 mm. -10.0 mm. greatest body width 2.7
mm-3.0 mm.

390 The University Science Bulletin

Shape. — Large, fusiform species; greatest body width midway
the body length.

Color. — General body color testaceous. Eyes brown. Legs stra-
mineous. Abdominal venter dark brown with keel and segmental
margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of the head eight tenths the
pronotal humeral width and seven to eight times the anterior width
of the vertex; synthlipsis wide, slightly more than one half the an-
terior width of the vertex; along the median longitudinal axis the
head is almost one half the length of the pronotum. Pronotum with
its humeral width almost twice the median length; lateral margins
diverging and one half the median length; posterior margin convex,
medianly emarginate. Facial tubercle slightly raised. Labrum
with its basal width one and three tenths the median length; apex
rounded. Rostral prong (Pi. XLV, fig. 50b) slightly shorter than
the third rostral segment; apex more or less accuminate. Stridu-
latory comb (Pi. XLV, fig. 50c) of approximately twenty teeth.
Chaetotaxy of the male front leg as shown on Plate XLV. The rela-
tive lengths of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 127 91

Middle leg 100 89 33 22

Hind leg 100 82 32 29

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head nine tenths the
pronotal humeral width and slightly more than five times the an-
terior width of the vertex; synthlipsis wide, two thirds the anterior
width of the vertex; along the median longitudinal axis the head
is one half the length of the pronotum. Pronotum with its humeral
width slightly more than twice the median length; lateral margins
diverging and almost one half the median length; posterior margin
convex, medianly emarginate. The relative lengths of the parts of
the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 128 55 36

Middle leg 100 89 33 25

Hind leg 100 78 28 23

Location of types. — The type material for Aiiisops pellucens
grandis Poisson from Tananarive Madagascar is in the Paris Mu-
seum.

Comparative notes. — This species is very closely related to Ani-

Brooks: The Genus Anisops 391

sops pellucens Gerst. and can be successfully separated only on the
basis of the males. The males of A. grandis have a narrower an-
terior width of the vertex, one seventh to one eighth the greatest
width of the head, instead of one sixth as in A. pellucens. Also the
males lack the apical spine on the inner surface of the anterior mar-
gin of the fore tibia as found on A. pellucens. The rostral prongs
are shaped somewhat differently, that of Anisops pellucens being
much wider in its apical half than that of A. grandis.

Remarks. — I am raising Anisops pellucens grandis Poisson to spe-
cific level on the basis of the above mentioned differences from A.
pellucens.

Data on distribution:

Madagascar

Tananarive, 1921, R. Decary, one male, one female (F. H. Snow Coll.);
one male, one female (Paris Museum).

Madagascar, 1899, Grandidier, four females (Paris Museum).

Marais du Fimerna, 1905, F. Geay, five females (Paris Museum).

Region de L'Androy Ambouvombe, 1-15-1901, Dr. F. DeCorse, one female
(Paris Museum).

Madagascar, two males, two females ( U. S. Nat. Mus. ).

Madagascar, Diego Suarez, 1893, Ch. Alluaud, one male (U. S. Nat. Mus.).

Betaimisaratha, exchange fr. Horvath, two females (F. H. Snow Coll.).

Madagascar, one male, six females (Bueno Coll. of F. H. Snow Coll.).

Anisops tanalensis n. sp.*

(PI. XL1V, fig. 46)

Size. — Males 5.8 mm.-6.3 mm., greatest body width 1.6 mm.-
1.9 mm.

Shape. — Subfusiform species; greatest body width midway the
length of the body.

Color. — General facies dark gray. Eyes dark brown. Vertex
and pronotoum testaceous, the latter may have a brown median
spot. Scutellum black with lateral margins testaceous or testaceous
with only apical half black. Hemelytra hyaline and appear the
black color of the dorsal body surface. Legs stramineous. Abdomi-
nal venter dark brown or black with keel and segmental margins of
the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the anterior margin almost straight;
greatest width of head almost as wide as the pronotal width and
slightly more than seven times the anterior width of the vertrex;
synthlipsis wide, one half the anterior width of the vertex; along the

* No females were present in the material at my disposal.

392 The University Science Bulletin

median longitudinal axis the head is two thirds the length of the
pronotum with its humeral width almost twice the median length;
lateral margins diverging and one half the median length; posterior
margin convex, medianly emarginate. Facial tubercle slightly
raised. Labrum long; median length almost equal to the basal
width; apex rounded. Rostral prong (PI. XLIV, fig. 46b) longer
than the third rostral segment; apex accuminate. Anterior femur
rounded at the apex. Stridulatory comb (PI. XLIV, fig. 46c) of ap-
proximately thirteen teeth; apical three shorter than basal ten.
Chaetotaxy of the male front leg as shown on Plate XLIV. The
relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 115 92

Middle leg 100 85 40 28

Hind leg 100 86 31 34

Location of types. — Male holotype, Madagascar, Foret Tanala,
Reo de Ranonafana, Anjorojoro, 1901, Ch. Alluaud, in the Paris Mu-
seum. Paratypes are as follows: one male, Madagascar, Diego-
Suarez, 1901, Ch. Alluaud, in the Paris Museum; one male (same
collection data as type) in the Francis H. Snow Collection.

Comparative notes. — This species appears closely related to
Anisops alluaudi Poisson and differs from it by the fact that the
males have the apex of the front femur rounded, whereas that of
A. alluaudi is accuminate.

Data on distribution. — Known only from type series.

Anisops windi n. sp.

(PI. XLV, fig. 48)

Size. — Males, length 5.2 mm. -5.4 mm., greatest body width 1.4
mm.; females, length 4.8 mm., greatest body width 1.3 mm.

Shape. — Small, slightly fusiform species, greatest body width
almost midway the body length.

Color. — Eyes brown. Vertex and pronotum testaceous. Scutel-
lum testaceous, orange, or black with lateral margins crimson.
Hemelytra hyaline and may appear black or stramineous, depending
on the color of the underlying color of the dorsal body surface.
Hemelytra may be tinged with crimson along scutellar margins.
Legs stramineous. Abdominal venter black with keel and seg-
mental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the anterior margins of the eyes ex-

Brooks: The Genus Anisops 393

tending slightly beyond that of the vertex; greatest width of the
head nine tenths the pronotal humeral width, approximately nine
times the anterior width of the vertex; synthlipsis wide, nine tenths
the anterior width of the vertex; along the median longitudinal axis
the head is four fifths the length of the pronotum. Pronotum with
its humeral width slightly more than twice its median length; lateral
margins diverging and slightly more than twice its median length;
posterior margin convex, medianly emarginate. Facial tubercle
only slightly raised; bearing long procumbent hairs. Labrum
hairy; basal width equal to median length; apex rounded. Ros-
tral prong (PL XLV, fig. 48b) longer than third rostral segment;
apex accumulate. Stridulatory comb (Pi. XLV, fig. 48c) of approxi-
mately seventeen, almost even-length, teeth. Chaetotaxy of the
males front leg as shown on Plate XLV. The relative lengths of the
parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 127 82

Middle leg 100 79 38 27

Hind leg 100 86 30 36

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with its greatest width nine tenths the
pronotal humeral width and five times the anterior width of the
vertex; synthlipsis wide, slightly more than one half the anterior
width of the vertex; along the median longitudinal axis the head is
three fifths the pronotal length. Pronotum with its humeral width
almost twice the median length; lateral margins diverging and one
half the median length; postertior margin convex, medianly emargi-
nate. The relative lengths of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 109 * *

Middle leg 100 88 44 26

Hind leg 100 81 31 31

Location of types. — Male holotype, female allotype, Australia,
N. Queensland, Barren River near Redlyach, IX-21-28, R. G. Wind,
in the Snow Entomological Collection. One male paratype. Rocky
Scrub, Mcllwraith Rge. C. York, Queensland, Australia, Harvard
Exped., Darlington, in the Harvard Museum of Comparative Zool-
ogy.

Comparative notes. — This species appears somewhat similar to
A. barrenesis n. sp. from which it can be distinguished on the basis

* Lacking on the female at my disposal. Specimen mutilated.

394 The University Science Bulletin

of the males which lack the greatly swollen anterior femur as found
on the latter.

Data on distribution. — Known only from type series.

Anisops leucothea Esaki

(PI. XLV, fig. 47)

1926. Anisops leucothea Esaki, Insects of Samoa, Part II, pp. 76-80, text fig. 6.
1933. Anisops leucothea, Lundblad, Arch, fur Hydrob., Suppl. vol. XII, p. 145 (a list of
Indo-australian and Pacific forms of this genus).

Size. — Males, length 7.2 mm. -7.3 mm., greatest width of body
2.1 mm. -2.2 mm.; females, length 6.9 mm. -7.6 mm., greatest body
width 1.5 mm. -2.3 mm.

Shape. — Robust, slightly fusiform species; lateral margins in an-
terior half of body almost parallel, converging in posterior half of
body.

Color.— ^General facies stramineous. Eyes brown. Legs stra-
mineous. Abdominal venter dark brown with keel and segmental
margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the head is
broad with its outline rounded; greatest width of head equal to the
pronotal humeral width and seven to eight times the anterior width
of the vertex; synthlipsis wide, over one half the anterior width of
the vertex; along the median longitudinal axis the head slightly
more than half the length of the pronotum. Pronotum with its
humeral width one and two thirds its median length; lateral margins
more or less parallel with their length two thirds the median length;
posterior margin convex, medianly emarginate. Facial tubercle only
slightly raised. Labrum long, median length slightly more than its
basal width; apex rounded; base of the labrum set with long hairs
that extend the full length of the labrum. Rostral prong ( PL XLV,
fig. 47c ) slightly shorter than the third rostral segment; apex accumi-
nate. Stridulatory comb ( PI. XLV, fig. 47b ) of approximately nine-
teen teeth. Chaetotaxy of the male front leg as shown on Plate XLV.
The relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 103 72

Middle leg 100 81 48 27

Hind leg 100 80 25 28

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head nine tenths the
pronotal humeral width and six times the anterior width of the
vertex; synthlipsis wide, slightly more than one half the anterior

Brooks: The Genus Anisops 395

width of the vertex; along the median longitudinal axis the head is
three fifths as long as the pronotum. Pronotum with humeral width
slightly more than twice the median length; lateral margins diverg-
ing and one half the median length; posterior margin convex,
medianly emarginate. The relative lengths of the parts of the legs
are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 114 57 36

Middle leg 100 85 42 37

Hind leg 100 81 28 28

Location of types. — Male and female types, six male and eight
female paratypes, Upolu: Mulifanua, XI-9-1925, in the British Mu-
seum.

Comparative notes. — Very similar to Anisops occipitalis Breddin
and only the males can be relied on for accurate separation. On
the males of A. leucothea Esaki the width of the head is equal to the
pronotal humeral width and the lateral margins of the pronotum
are almost parallel whereas on the males of A. occipitalis the head is
only nine tenths the pronotal humeral width and the lateral pro-
notal margins are diverging. Also the head of the former is seven
to eight times the anterior width of the vertex while in the latter
the head is approximately six times the anterior width of the vertex.

Data on distribution:
Wallis Island

Uvea, Lake Kikila, 1903, Dr. Joly, one male ( compared with type by Dr.
Hungerford ) , two females ( F. H. Snow Coll. ) ; one male five females ( Paris
Museum).

Anisops dor is Kirkaldy

(PI. XLVI, fig. 53)

1904. Anisops doris Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 112, 132.
1923. Anisops doris, Hale, Rec. South Australian Mus., vol. II, no. 3, p. 402-403, text fig.
364, PI. XI, fig. 11 (evidently had a mixture of two or more species).

*1924. Anisops doris, Hale, South Australian Nat., vol. V, no. 4, p. 135 (note on migra-
tion).

1933. Anisops doris. Lundblad, Arch, fiir Hydrob., Suppl. vol. XII, p. 145 (a list of Indo-
australian and Pacific forms of this genus).

1934. Anisops doris, Hungerford, Bull. Brooklyn Ent. Soc, vol. XXIX, p. 68 (ecological
note; inaccurate as composite species, neither of which A. doris Kirkaldy).

*1935. Anisops doris, Hale, Trans. Royal Soc. South Australia, vol. LIX, p. 249 (ecological
ncte).

Size. — Males, length 7.5 mm. -7.8 mm., greatest body width 1.8
mm.; females, length 7.2 mm.-7.4 mm., greatest body width 1.8 mm.-
1.9 mm.

* Questionable, due to previous inaccuracy in determination.

396 The University Science Bulletin

Shape. — Broad-headed, fusiform species; greatest body width
about midway the body length.

Color. — General facies stramineous. Eyes brown. Legs stra-
mineous. Abdominal venter dark brown with keel and segmental
margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the greatest width of the head slightly
more than the pronotal humeral width and about eight and one
half times the anterior width of the vertex; synthlipsis wide, at least
one third the anterior width of the vertex; along the median longi-
tudinal axis the head is nine tenths the length of the pronotum.
Pronotum with its humeral width slightly less than twice its median
length; lateral margins slightly convex, almost parallel; two thirds
the median length of the pronotum. Frons narrow, about the width
of the synthlipsis; facial tubercle only slightly raised. Labrum with
its basal width slightly more than its median length; apex rounded.
Rostral prong (PL XL VI, fig. 53b) shorter than the third rostral
segment; apex accuminate. Stridulatory comb (PL XLVI, fig. 53c)
of approximately fifty-four closely appressed teeth. Chaetotaxy
of the front leg as shown on Plate XLVI. The relative lengths of the
parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 131 78

Middle leg 100 80 33 27

Hind leg 100 81 30 35

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of the head nine tenths
the pronotal humeral width and six times the anterior width of the
vertex; synthlipsis wide, one half the anterior width of the vertex;
along the median longitudinal axis the head is two thirds the
median length; lateral margins diverging and three fourths the me-
dian length; posterior margin convex, medianly emarginate. The
relative lengths of the parts of the legs are as follows:*

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 123 56 35

Middle leg 100 89 38 30

Location of types. — Male type, two male and two female cotypes,
Australia, Victoria, Alexandria, F. L. Billinghurst in the Kirkaldy col-
lection of the Snow Entomological Collection.

* Neither of the two females available for study possessed hind legs.

Brooks: The Genus Anisops 397

Comparative notes. — This species is about the same size as Ani-
sops grains Hale. However, the two may be readily separated on
the basis of the males which in A. doris Kirkaldy have the head
greater in width than the pronotum humeral width and the anterior
width of the vertex is greatly reduced, slightly less than one eighth
the width of the head, whereas in A. gratns the greatest width of
the head is only nine tenths the pronotal humeral width and the
anterior width of the vertex is at least one fifth the greatest width of
the head.

Data on distribution:

Australia

Victoria, Bacchus Marsh, 1-9-14, one male, two females ( Bueno Coll. of F.
H. Snow Coll. ) .

Anisops wakefieldi White

(PI. XLVII, fig. 55; PI. LVII, fig. 106)

187S. Anisops wakefieldi White, The Entomologist Month. Mag., vol. XV, p. 161.

1897. Anisops wakefieldi, Hutton, Trans. New Zealand Inst., vol. XXX, pp. 179-180.

1904. Anisops wakefieldi, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. Ill, 132.

1908. Anisops wakefieldi, Kirkaldy, Trans. New Zealand Inst., vol. XLI, p. 27 (ecological
note).

1914. Anisops wakefieldi, Myers, Trans. New Zealand Inst., vol. 56, pp. 468-469.

1933. Anisops wakefieldi, Lundblad, Arch, fur Hydrob., Suppl. vol. XII, p. 146 (a list of
Indo-australian and Pacific forms of this genus).

Size. — Males, length 7.5 mm., greatest body width 1.9 mm.; fe-
males, length 8.1 mm.-8.4 mm., greatest body width 2.1 mm. -2,2 mm.

Shape. — Subfusiform species; lateral margins of the anterior third
of the body almost parallel, only slightly converging; lateral margins
strongly converging in the posterior two thirds of the body.

Color. — Brown form: General facies stramineous. Eyes brown.
Abdominal venter dark brown with keel and segmental margins of
the connexivum and stramineous. Dark form: General facies gray.
Eyes brown. Vertex and pronotum testaceous, the latter with the
posterior half hyaline and appears black as it overlies the black
anterior portion of the scutellum. Scutellum black with its lateral
margin stramineous. Hemelytra hyaline and appear dark gray as
they overlie the black dorsal body surface. Legs stramineous. Ab-
dominal venter black with keel and segmental margins of the con-
nexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with its greatest width nine tenths the pro-
notal humeral width and slightly more than four times the anterior
width of the vertex; synthlipsis wide, three fifths the anterior width
of the vertex; along the median longitudinal axis the head is almost

398 The University Science Bulletin

one half the length of the pronotum. Pronotum with its humeral
width one and one half its median length; lateral margins diverging
and one-half the median length; posterior margin convex, medianly
emarginate. Facial tubercle swollen into a ventrally directed tri-
angular eminence, with its apex accumulate. Labrum with its
basal width only slightly more than its median length; apex rounded.
Rostral prong (Pi. LVII, fig. 106) greatly reduced, only one fourth
the length of the third rostral segment; apex truncate. Stridulatory
comb (PI. XLVII, fig. 55b) of approximately eighteen teeth. Chae-
totaxy of the male front leg as shown on Place XLVII. The relative
lengths of the parts of the legs are as follows:*

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 110 70

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded, with its greatest width eight to nine
tenths the pronotal humeral width and four times the anterior width
of the vertex; synthlipsis wide, two thirds the anterior width of the
vertex; along the median longitudinal axis the head is slightly more
than one third the length of the pronotum. Pronotum with its
humeral width one and three fourths its median length; lateral mar-
gins diverging and less than one half the median length; posterior
margin convex, medianly emarginate. Facial tubercle swollen to
form a triangular shaped eminence as in the males, but not as large
or as accuminate. The relative lengths of the parts of the legs are
as follows:

1st 2nd

Feraui Tibia Tar. Seg. Tar. Seg.

Fore leg 100 102 46 38

Middle leg 100 83 35 24

Hind leg 100 80 31 31

Location of types. — White's type material from Canterbury and
Otago, New Zealand in the Perth Museum.

Com))arativc notes. — This species closely resembles A. assimilis
White. However, the pronotum of the male of the latter species has
its lateral margins almost parallel whereas in A. wakefieldi the pro-
notal lateral margins are diverging. Also the males of the latter
lack the greatly swollen facial tubercle as found on the former.
Data on distribution:
New Zealand

Queenstown, 1875, Hutton, one male, ( F. H. Snow Coll.).
Nelson, 1876, Filhal, two females (F. H. Snow Coll.).
New Zealand, one female (F. H. Snow Coll.).

* Only the front leg was intact on the single male specimen at my disposal.

Brooks: The Genus Anisops 399

An isops hancocki Hutchinson

(PI. XLVI, fig. 52 ; PI. LV, fig. 96)

1928. Anisops hancocki Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. I, pp. 163-104,
text fi;:. 7.

1933. Anisops hancocki, Hutchinson, Internat. ges. Hydiob. unci Hydrog., vol. 28, pt. 5/0,
p. 462 (ecological note).

Referring to this species also :

1939. Anisops murati, Poisson, Bull. Soc. Ent. France, vol. 44, pp. 43-44, text figs. 2, 3.

Size. — Males, length 6.3 mm. -6.9 mm., greatest body width 1.9
mm. -2.3 mm.; females, length 6.4 mm. -6.9 mm., greatest body width
1.9 mm.-2.2 mm.

Shape. — Robust, fusiform species; greatest body width about
midway the body length.

Color. — General facies brown. Eyes brown. Vertex, pronotum
and scutellum testaceous, the latter may be brown with only apex
testaceous and may have its lateral margins red. Hemelytra hyaline
and appear dark brown due to the brown color of the dorsal body
surface. Legs stramineous. Abdominal venter dark brown with
keel and segmental margins of the connexivum stramineous. One
specimen is flavus, probably a teneral form.

Male structural characteristics. — Viewed from above, the anterior
margin of the head is almost straight; laterally curved; greatest
width of head nine tenths the pronotal humeral width and four and
one half times the anterior width of the vertex; synthlipsis wide,
slightly more than one half the anterior width of the vertex; along
the median longitudinal axis the head is short, less than one half
the length of the pronotum. Pronotum with its humeral width
twice its median length; lateral margins diverging and one half the
median length. Posterior margin convex, medianly emarginate.
Facial tubercle excavate; lateral margins carinate. Labrum broad;
basal width one and two thirds the median length; apex rounded;
a long tuft of hairs at each basal angle. Rostral prong ( PL LV, fig.
96) shorter than third rostral segment; apex more or less rounded.
Stridulatory comb (PI. XLVI, fig. 52b) of approximately ten teeth
which decrease in length at the apex. Keel of the second abdominal
segment enlarged laterally and excavate medianly; lateral margins
carinate and bearing long erect hairs; excavation lined with short
procumbent hairs. Connexivum of the third abdominal segment
with a row of nodules on its inner margin extending from base to
apex; basal two nodules greatly enlarged. Chaetotaxy of the male
front leg as shown on Plate XLVI. The relative lengths of the parts
of the legs are as follows:

400 The University Science Bulletin

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 108 87

Middle leg 100 83 42 29

Hind leg 100 80 31 31

Female structural characteristics. — Viewed from above the out-
line of the head is rounded; greatest width of head eight tenths
the pronotal humeral width and almost four times the anterior
width of the vertex; synthlipsis wide, one half the anterior width of
the vertex; along the median longitudinal axis the head is slightly
more than one half the length of the pronotum. Pronotum with
its humeral width two and one third the median length; lateral
margins diverging and slightly more than one half the median
length; posterior margin convex, medianly emarginate. The rela-
tive lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 118 57 28

Middle leg 100 81 50 23

Hind leg 100 81 33 33

Location of types. — Male type and one male paratype, Uganda,
Kampala, 111-20-1927, G. L. Hancock, in the British Museum.

Comparative notes. — This species is very similar to A. psyche
Hutchinson and differs from it by having the fore femur of the male
greatly enlarged dorso-ventrally just before the base, a condition
lacking in A. psyche. For comparison of the chaetotaxies of the
front legs of the males see Plate XLVI, fig. 52a and Plate XLIII, fig.
38a.

Remarks. — A. murati Poisson, according to the drawings and de-
scription of Poisson appears to be identical in every respect to
Anisops hancocki. So I am placing this species as a synonym of the
later.

Data on distribution:
Africa

Uganda, Kampala, XI-20-1929, G. E. Hopkins, one male (F. H. Snow Coll.).

Orient. Angl. Sambourou, Wa-Nyika, 1904, Ch. Allnaud, one male (F. H.
Snow Coll.).

East Africa, IV-21-1949, F. X. Williams, two males (Harv. Mus. Comp.
Zool.).

Soudan, Nioro, 1909, F. de Zeltner, one male (Paris Museum).

Sudan, Rock pool, R. Yei Equatoria, Dec. 11, 1937, J. G. Myers, one male,
two females (Rrit. Mus.).

Ogaden, Ouabi-Chebeli, Imi Mission, 1903, Du Rourg du Rozas, one male
( Paris Museum ) .

Brooks: The Genus Anisops 401

Anisops decipiens Hutchinson

(PI. XLVIII, tig. 62)

1930. Anisops decipiens Hutchinson, Proc. Zool. Soc. London, vol. XXIX, pt. "2, pp. 448-
449, text figs. 4, a, b, c.

1932. Anisops decipiens, Hutchinson, Proc. Zool. Soc. London, vol. XXXI, pt. 1, pp. 125-
12(5 (ecological note).

Size. — Males, length 6 mm., greatest body width 1.8 mm.; females,
length 7.4 mm., greatest body width 2.1 mm.

Shape. — Robust, slightly fusiform species; greatest width at
humeral margins of pronotum.

Color. — General facies dark brown. Eyes dark brown. Vertex
and pronotum testaceous, the latter irregularly hyaline and with
a median brown spot. Scutellum dark brown, lateral margins crim-
son. Hemelytra hyaline and appearing the dark brown color of the
underlying dorsal body surface. Legs stramineous. Abdominal
venter dark brown with keel and segmental margins of the con-
nexivum stramineous.

Male structural characteristics. — Viewed from above, the head is
short with the outline rounded, anterior margin almost straight;
greatest width of the head nine tenths the pronotal humeral width
and five times the anterior width of the vertex; synthlipsis wide, one
half the anterior width of the vertex; along the median longitudinal
axis the head is slightly more than one third the length of the pro-
notum. Pronotum with its humeral width twice the median length;
lateral margins diverging and less than one half the median length;
posterior margin convex, medianly emarginate. Facial tubercle
only slightly raised. Labrum with its basal width almost one and
one third the median length; apex rounded. Rostral prong (Pi.
XLVIII, fig. 62c) equal to the length of the third rostral segment;
apex accuminate. Stridulatory comb (PI. XLVIII, fig. 62b) of ap-
proximately seven stout teeth which increase in length at the middle.
Chaetotaxy of the front leg as shown on Plate XLVIII. The relative
lengths of the parts of the legs are as follows:*

Femur

Fore leg 100

Middle leg 100

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of the head eight tenths
the pronotal humeral width and slightly more than four times the

* Unfortunately, the single male specimen in my material was minus the fore tarsi and the
hind legs.

26—3286

1st

2nd

?ibia

Tar. Seg.

Tar. Seg

119

81

38

23

1st
Tar. Seg.

2nd
Tar. Seg.

56
36

33

21

402 The University Science Bulletin

anterior width of the vertex; synthlipsis wide, one half the anterior
width of the vertex; along the median longitudinal axis the head is
one half the length of the pronotum. Pronotum with its humeral
width slightly more than twice its median length; lateral margins
diverging and almost one half the median length; posterior margin
convex, medianly emarginate. The relative lengths of the parts of
the legs are as follows :f

Femur Tibia

Fore leg 100 122

Middle leg 100 82

Location of types. — Male type, female allotype, and one female
paratype, Asbysinnsi, Wouranboulchi, XI-4th to 7th-1926, J. Omer-
Cooper, in the British Museum.

Comparative notes. — Superficially similar to Anisops ares Hutch-
inson from which it distinguishes itself by the fact that the males
lack the dorso-ventrally expanded front femur with the curved apex
as found on the males of A. ares.

Data on distribution:

Africa

Abyssinia, Djem-Djem Forrest, circa. 8000', X-21-1926, J. Omer-Cooper,
one male, one female, gift to H. B. H. from G. E. Hutchinson (F. H. Snow
Coll.).

Anisops barrenensis n. sp.

(PI. XLIV, fig. 42)

Size. — Males, length, 4.8 mm. -5.1 mm., greatest body width
1.4 mm. -1.6 mm.; females, length 4.8 mm.-5.1 mm., greatest body
width 1.4 mm. -1.6 mm.

Shape. — Small, robust, fusiform species; greatest body width
midway the body length.

Color. — General facies gray. Eyes brown. Vertex may be tinged
with orange. Hind margin of pronotum may be hyaline and appear
black as it overlies the black anterior portion of the scutellum.
Legs stramineous. Abdominal venter black with keel and seg-
mental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the anterior margin almost straight;
greatest width of head nine tenths the pronotal humeral width;
four and one half times the anterior width of the vertex; synthlipsis

t The female specimen in my material was minus the hind iegs.

Brooks: The Genus Anisops

403

wide, slightly more than one half the anterior width of the vertex;
along the median longitudinal axis the head is slightly less than
one half the pronotal length. Pronotum with its humeral width
slightly less than twice the median length; lateral margins diverg-
ing and one half the median length; posterior margin convex,
medianly emarginate; dorsal surface with a faint median ridge and
a convexly curved ridge on each side between median line and
lateral margin. Facial tubercle slightly raised and bearing a few
hairs. Posterior margin of third rostral segment covered for al-
most its entire length by an extension of the second rostral seg-
ment (PI. XLIV, fig. 42b). Rostral prong accuminate. Labrum
covered with small fine hairs; basal width only slightly greater
than median length; apex truncate. Stridulatory comb (Pi. XLIV,
fig. 42c) of approximately sixteen teeth. Chaetotaxy of the male
front leg as shown on Plate XLIV. The relative lengths of the legs
are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 116 83

Middle leg 100 81 39 28

Hind leg 100 86 31 36

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head nine tenths
the pronotal humeral width, slightly more than four times the
anterior width of the vertex; synthlipsis wide, more than one half
the anterior width of the vertex; along the median longitudinal
axis the head is one half the pronotal length. Pronotum with its
humeral width slightly more than twice its median length; lateral
margins diverging and one half the median length; posterior mar-
gin convex, medianly emarginate, dorsal ridges as in the males.
The relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. .Seg. Tar. Seg.

Fore leg 100 116 50 28

Middle leg 100 80 37 26

Hind leg 100 86 31 36

Location of types. — Male holotype, female allotype, twenty-one
male and twenty-nine female paratypes. N. Queensland, Barren
River nr. Barren Waters, IX-9-28, R. Wind, in the Snow Entomo-
logical Collection. Other paratypes are as follows. In the Harvard
Museum of Comparative Zoology: one male, two females, Queens-
land, Australia, Coen C. York, May 32, Darlington, Harv. Austrl.
Exped.

404 The University Science Bulletin

Comparative notes. — Very similar to A. hackeri n. sp. from which
it differs by the fact that the males of the latter do not have the
greatly expanded front femur as found on A. barrenensis.

Data on distribution. — Known only from type series.

Anisops hungerfordi Poisson

(PI. XLVI, fig. 51)

1935. Anisops hungerfordi Poisson, Mus. Nat. Hist. Nat., vol. Ill, p. 213, text fig. 28,
A, B. C.

Size. — Males, length 5.1 mm., greatest body width 1.5 mm.; fe-
males, length 5.4 mm., greatest body width 1.5 mm.

Shape. — Small, robust, slightly fusiform species; greatest body
width about midway the body length.

Color. — General facies stramineous. Eyes brown. Legs stramine-
ous. Abdominal venter dark brown with keel and segmental mar-
gins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of the head nine tenths the
pronotal humeral width and six times the anterior width of the
vertex; synthlipsis wide, one third the anterior width of the vertex;
along the median longitudinal axis the head is almost one half the
length of the pronotum; lateral margins diverging and almost one
half the median length; posterior margin straight. Facial tubercle
flat. Labrum long with its median length almost one and one third
the median length; apex rounded. Rostral prong (Pi. XLVI, fig. 51b)
longer than third rostral segment; apex rounded. Stridulatory comb
(PI. XLVI, fig. 51c) of approximately fifteen teeth which decrease in
length in apical half. Chaetotaxy of the male front leg as shown on
Plate XLVI. The relative lengths of the parts of the legs are as
follows:*

Femur

Fore leg 100

Middle leg 100

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with the greatest width nine tenths the
pronotal humeral width and four to five times the anterior width
of the vertex; synthlipsis wide, slightly more than one third the
anterior width of the vertex; along the median longitudinal axis the
head is slightly more than one half the pronotal length. Pronotum
with its humeral width slightly more than twice the median length;
lateral margins diverging and almost one half the median length;

* Hind leg not complete on single male specimen in my material.

Tibia

1st
Tar. >Seg.

2nd
Tar. Seg.

121
76

81
37

27

Brooks: The Genus Anisops 405

posterior margin straight. The relative lengths of the parts of the
legs are as follows:

1st 2nd

Femur Tibia Tar. Scg. Tar. Seg.

Fore leg 100 117 50 33

Middle leg 100 76 37 28

Hind leg 100 91 32 32

Location of types. — Type material from Bourille, Ethiopia in the
Paris Museum.

Comparative notes. — This species is very similar to A. apicalis
Stal and differs from it primarily by the fact that the wings are not
brachypterous; also the scutellum is larger, its basal width only one
and one fourth the median length instead of one and three fourths
the median length as in A. apicalis.

Data on distribution:
Africa

Belgian Congo, Mucosa, XI-1939, H. S. Bredo, one male, two females (Roy.
Mus. Nat. Hist. Belg.).

Belgian Congo, Mucosa, VIII-3-1927, H. H. Bredo, two females (Roy Mus.
Nat. Hist. Belg.).

Belgian Congo, Elizabethville, 1-10-1938. H. S. Bredo (Roy. Mus. Nat.
Hist. Belg.).

Anisops ares Hutchinson

(PI. XLVII, fig. 57)

1926. Anisops ares Hutchinson, Ann. Mag. Nat. Hist.. Ser. 10, vol. I, pp. 104-106, text
fig. 8.

1933. Anisops ares, Hutchinson, Internat. ges. Hydrob. unci Hydrog., vol. 28, pt. 5/6,
p. 462.

Referring to this species also :

1928. Anisops coxalis, Poisson, Bull. Soc. Ent. France, no. 4, p. 74.

1929. Anisops coxalis, Poisson, Fauna des Colonies Francaises, vol. Ill, pp. 152-154,
figs. 20, 21.

1948. Anisops coxalis, Poisson, Rev. Francaise Ent., vol. XV. p. 168 (ecological note).

Size. — Males, length 6.5 mm. -7.1 mm., greatest body width 1.9
mm. -2.1 mm.; females, length 6.0 mm. -6.3 mm., greatest body width
1.2 mm. -1.4 mm.

Shape. — Robust, slightly fusiform species; greatest body width at
the humeral width of the pronotum.

Color. — Light form: General facies stramineous. Eyes brown.
Apical third of hemelytra dark brown. Legs stramineous. Ab-
dominal venter dark brown with keel and segmental margins of the
connexivum stramineous. Dark form: Eyes brown. Vertex brown.
Vertex and pronotum testaceous, the latter may be hyaline on its
posterior half and appear the black color of the underlying scu-
tellum. Scutellum dark brown or black. Hemelytra hyaline with

406 The University Science Bulletin

posterior third dark brown or black; hyaline areas appear dark
brown or black depending on the underlying color of the dorsal
body surface. Legs stramineous. Abdominal venter black with
keel and segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the lateral
margins of the head are curved and the anterior margin almost
straight; greatest width of the head only slightly narrower than the
pronotal humeral width and six times the anterior width of the
vertex; synthlipsis wide, at least one half the anterior width of the
vertex; along the median longitudinal axis head is short, only one
third the pronotal length. Pronotum long, humeral width one and
two thirds the median length; lateral margins slightly diverging and
one half the median length; posterior margin convex, medianly
emarginate. Facial tubercle slightly raised. Labrum short, basal
width slightly more than the median length; apex rounded. Rostral
prong (PI. XLVII, fig. 57b) longer than the third rostral segment;
apex accuminate. Fore leg (PI. XLVII, fig. 57a) with the dorsal and
ventral margins of the femur almost parallel for the basal three
fourths of their length; apex rounded; stridulatory comb (Pi. XLVII,
fig. 57c) of approximately eleven teeth, the apical five slightly
longer than the basal six. Chaetotaxy of the male front leg as shown
on Plate XLVII. The relative lengths of the parts of the legs are as
follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 105 73

Middle leg 100 82 33 24

Hind leg 100 82 33 30

Female structural characteristics. — Viewed from above, the out-
line of the head is curved laterally and almost straight on its anterior
margin; greatest width of head nine tenths the pronotal humeral
width and six times the anterior width of the vertex; along the
median longitudinal axis the head is one half the length of the pro-
notum. Pronotum with its humeral width slightly more than twice
its median length; lateral margins diverging and one half the median
length; posterior margin convex, medianly emarginate. The rela-
tive lengths of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 117 56 33

Middle leg 100 77 35 27

Hind leg 100 86 37 37

Location of types. — Male type, female allotype, three male and
two female paratypes Uganda, Kampala, 111-20-1927, in the British
Museum.

Brooks: The Genus Anisops 407

Comparative notes. — Very similar to Anisops varia Fieber but
can easily be separated on the basis of the males. The males of the
latter species have a much shorter pronotum which is twice the
length of the head instead of three times the length of the head as
in A. ares. Also, the male front femur of A. varia has its dorsal and
ventral margin converging from the base, instead of the almost
parallel condition as found on A. ares.

Remarks. — Anisops ares Hutchinson was described from a form
occurring in Uganda and A. coxalis was described from Cameron.
The material at my disposal reveals that A. ares is a species of wide
distribution occurring in South Africa, Tanganyika, Uganda west to
Cameron, and that A. coxalis is a synonym of A. ares.

Data on distribution:
Africa

South Africa, Natal, Mahira Forrest, IX- 1920, through R. A. Drummer, four
males, five females (U. S. Nat. Mus. ).

Uganda, Kampala, 1-20-30, G. E. Hopkins, gift to H. B. H. from G. E.
Hutchinson, one male, one female (F. H. Snow Coll.).

Tanganyika, Mpala, Oberthur, one male, three females (Paris Museum).

Tanganyika, Amani, Feb. 1948, F. X. Williams, one female ( Harv. Mus.
Com. Zool. ).

Cameroons, W. Africa, Sangmelina, 4-16-32, A. I. Good, fifteen males,
twenty females ( F. H. Snow Coll. ) .

Cameroons, W. Africa, Sangmelina, Oct. 17, 1934, A. I. Good, twenty-five
males, fourteen females ( F. H. Snow Coll.).

Nigeria, Abakaliki, 1-26-49, B. Malkin, one male, four females ( Calif. Acad,
of Science).

Anisops paranigrolineata n. sp.

(PI. XLVIII, fig. 59)

Size. — Males, length 6.0 mm., greatest body width 5.4 mm.; fe-
males, length 6.0 mm. -6.3 mm., greatest body width 1.8 mm. -1.9 mm.

Shape. — Fusiform species, with eyes much wider than the an-
terior width of the pronotum; greatest body width about midway
the body length.

Color. — Brown form: General facies stramineous. Eyes brown.
Legs stramineous. Abdominal venter black with keel and seg-
mental margins of the connexivum stramineous. Dark form: Eyes
dark brown. Vertex stramineous. Pronotum testaceous with
posterior margin appearing black as it is hyaline and overlies the
black anterior margin of the scutellum. Hemelytra hyaline and
appearing black as it overlies the black dorsal body surface. Legs
stramineous. Abdominal venter black with keel and segmental mar-
gins of the connexivum stramineous.

408 The University Science Bulletin

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of head nine tenths the
pronotal width, five to six times the anterior width of the vertex;
synthlipsis wide, almost nine tenths the anterior width of the vertex;
along the median longitudinal axis the head is less than one third
the length of the pronotum. Pronotum with its humeral width one
and one fourth its median length; lateral margins diverging and al-
most one half the median length; posterior margin convex; dorsal
surface with a faint median ridge and a longitudinal convexly
curved ridge on each side, near the lateral margin. Facial tubercle
slightly raised and spread out laterally. Labrum greatly reduced
in length; basal width almost four times the median length; rounded
along anterior margin. Rostral prong (Pi. XL VIII, fig. 59b) shorter
than third rostral segment, apex rounded. Anterior femur greatly
enlarged dorso-ventrally at base; apex accumulate. Stridulatory
comb (PI. XLVIII, fig. 59c) of approximately eighteen teeth.
Chaetotaxy of the male front leg as shown on Plate XLVIII. The
relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 132 86

Middle leg 100 73 43 25

Hind leg 100 83 33 34

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width nine tenths the pronotal
humeral width, though much wider than the anterior width of the
pronotum; greatest width of head slightly more than five times
the anterior width of the vertex; synthlipsis wide, almost eight
tenths the anterior width of the vertex; along the median longi-
tudinal axis the head is almost one third the length of the pronotum.
Pronotum with its humeral width one and one half its median
length; lateral margins concave and diverging, almost one half the
median length; posterior margin convex; dorsal ridges present as
in male though somewhat fainter. The relative lengths of the parts
of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 128 57 36

Middle leg 100 74 37 27

Hind leg 100 84 33 33

Location of types. — Male holotype, female allotype, Mahabale-
shwar, Satara Dist. India, 4200', V-13 to 16-12, F. H. Gravely, in
the Indian Museum at Calcutta. Other paratypes are as follows.
In the Indian Museum at Calcutta: one male, one female, Satara

Brooks: The Genus Anisops 409

Dist, Nachal, W. Chats, 2000', IV-30-12, F. H. Gravely. In the
Snow Entomological Collection: two males, two females, Mahabale-
shwar, Satara Dist. India, 4200', V-13 to 16-12, F. H. Gravely.

Comparative notes. — This species is closely related to Anisops
nigrolineata Lundblad and differs from it in not having its inter-
ocular space swollen and extending beyond the margin of the eyes.

Data on distribution:

India

Kumaun, W. Himalayas, Nauhuchia Tal, 4200, V-5-11, S. W. Kemp, one
female ( F. H. Snow Coll. ) .

Coimbatore, one male, one female ( F. H. Snow Coll. ) .

Anisops nigrolineata Lundblad

PI. XLVII, fig. 50; PI. LVII, fig. 105)

1933. Anisops nigrolineata Lundblad, Arch, fur Hydrob., Suppl. vol. XII, pp. 145, 160-
163, fig. 55.

Size. — Males, length 5.4 mm. -6.3 mm., greatest body width 1.5
mm. -18 mm.; females, length 5.4 mm. -6. 6 mm., greatest body width
1.5 mm. -1.8 mm.

Shape. — Fusiform species with greatest body width at apical third
of body.

Color. — Dark form: General facies black. Eyes brown. Vertex
pronotum testaceous with the latter irregularly hyaline and such
areas appear black as they overlie the black scutellum. Scutellum
black, may be testaceous on the apical half. Hemelytra hyaline and
appear black as it overlies the black dorsal body surface. Legs
dark brown. Abdominal venter black with keel and segmental
margins of the connexivum brown. Pale form: General facies
pearlaceous. Eyes brown. Legs stramineous. Abdominal venter
black or dark brown with keel and segmental margins of the con-
nexivum pearlaceous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the vertex extending beyond the anterior
margins of the eyes; greatest width of the head nine tenths the pro-
notal humeral width and four to five times the anterior width of the
vertex; synthlipsis wide, one half the anterior width of the vertex;
along the median longitudinal axis the head is slightly more than
one third the length of the pronotum. Pronotum long; humeral width
one and two fifths the median length; lateral margins diverging and
slightly more than one third the median length; posterior margin
convex, medianly emarginate; dorsal surface with a median longitu-

410 The University Science Bulletin

dinal ridge and a convexly curved longitudinal ridge on each side,
midway between the median line and the lateral margin. Facial
tubercle flat and spread out laterally. Viewed laterally the inter-
ocular space protrudes beyond the margins of the eyes from midway
the dorsal length to the base of the labrum; flattened on its outer
surface. Labrum greatly reduced in length; basal width slightly
more than twice its median length; apex rounded. Rostral prong
(PI. LVII, fig. 105) slightly shorter than the third rostral segment;
apex truncate. Stridulatory comb (PL XLVII, fig. 56b) of approxi-
mately twenty-two teeth, the apical fourteen slender and long, more
than twice the length of the basal eight. Chaetotaxy of the front
leg as shown on Plate XLVII. The relative length of the parts of
the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 125 78

Middle leg 100 58 29 24

Hind leg 100 87 30 30

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with the vertex extending slightly beyond
the anterior margin of the eyes; greatest width of head eight to nine
tenths the pronotal humeral width and four to five times the anterior
width of the vertex; synthlipsis wide, almost one half the anterior
width of the vertex; along the median longitudinal axis the head is
one third as long as the pronotum. Pronotum long, humeral width
only one and one half the median length; lateral margins diverging
and slightly more than one third the median length; posterior margin
almost straight, only slightly concave; dorsal surface with the same
ridges as found on the males. The relative lengths of the parts of
the legs are as follows:

1st 2nd

Femur Tiliia Tar. Seg. Tar. Seg.

Fore leg 100 126 54 35

Middle leg 100 74 37 30

Hind leg 100 85 33 34

Location of types. — Type material from Java in the Stockholm
Museum.

Comparative notes. — The long pronotum of this species is also a
characteristic of A. thienemanni Lundblad, another species from
Java. However, the pronotum of the males of the latter species has
its lateral margins almost parallel whereas in A. nigrolineata the pro-
notal lateral margins are diverging. Also the males of A. thiene-
manni lack the protruding interocular space as found on the males of

Brooks: The Genus Anisops 411

this species. For comparison of the male front legs of the two
species see Plate XLVII, fig. 56a, and Plate XLVIII, fig. 63a.

Data on distribution:
Burma

Shingbwiyang, III-7-1944, Lt. L. C. Kuitert, sixteen males, twenty-one fe-
males (F. H. Snow Coll.).

Shingbwiyang, XI-15-1944, Capt. L. C. Kuitert, nine males, eight females
(F. H. Snow Coll.).

Myitkyina, XI-18-44, Capt. L. C. Kuitert, two males (F. H. Snow Coll.).

Myitkyina, X-3-45, Capt. L. C. Kuitert, one male, ( F. H. Snow Coll.).

Myitkyina, V-29-45, Capt. L. C. Kuitert, one female (F. H. Snow Coll.).

Myitkyina, IX-30-45, B. McDennott, two males (F. H. Snow Coll.). Burma,
1944, L. C. Kuitert, one male, two females (F. H. Snow Coll.).

India

Baluchistan, Quetta, 5700', pond in residence garden, Sta. 3, XI-10, B. S.
six males, eleven females (Indian Museum).

Central Province, Puchapur, III-4-19, C. P. and F. H. G., one male, one
female (Indian Museum).

Batnagiri Dist. 400', pophil, Vashishti, V-lst to 2nd-12, F. H. Gravely, four
males (Indian Museum).

Batnagiri Dist., 0-300', Chiplum Vashishti, V-3rd to 5th-12, F. H. Gravely,
two males ( Indian Museum ) .

Satara Dist., Nechal, W. Ghats, IV-30-12, F. H. Gravely, one male, two
females (Indian Museum).

Satara Dist., Medha Yanna Valley, 2200', IV-17th to 23rd-12, F. H. Gravely,
three males, four females (Indian Museum).

Satara Dist., Belvak, Koyna Valley, 2000', IV-28th to 30th-12, F. H. Gravely,
one female ( Indian Museum ) .

Bengal, Bangamati, Chittagong Hill Tracts, VH-llth to 16th-15, B. Hodgard,
three males, three females (Indian Museum).

S. India, Coimbatore, 11-14-13, four males ( Bueno Coll. of F. H. Snow Coll.).

Kumaun, W. Himalayas, Sat. Tal., 4500', V-9 to 10-11, S. W. Kemp, one
male (F. H. Snow Coll.).

Sibuyan Island
Sibuyan Isl. Baker, one male, one female ( U. S. Nat. Mus. ).

Anisops assimilis White

(PI. XLVIII, fig. 61)

1878. Anisops assimilis White, Entomologists Month. Mag., vol. XV, p. 161.

1897. Anisops assimilis, Hutton, Trans. New Zealand Inst., vol. XXX, p. 180.

1904. Ajiisops assimilis, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 112, 132.

1908. Anisops assimilis Kirkaldy, Trans. New Zealand Inst., vol. XLI, p. 27 (a check list
of Hemiptera of Maorian subregion).

1926. Anisops assimilis, Myers, Trans. New Zealand Inst., vol. 56, pp. 468-469.

1928. Anisops assimilis, Esaki, Insects of Samoa, vol. II, p. 79.

1933. Anisops assi7niiis, Lundblad, Arch, fiir Hydrob., Suppl. vol. XII, pp. 14.">, 173-174,
text fig. 63 (compared with Anisops thienemanni Lundblad).

Size. — Males, length 6.6 mm., greatest body width 1.8 mm., fe-

412 The University Science Bulletin

males, length 6.3 mm. -7. 8 mm., greatest width of body 1.7 mm.-
2.1 mm.

Shape. — Robust, only slightly fusiform species with lateral mar-
gins in the anterior half of the body almost parallel-sided and con-
verging in posterior half.

Color. — General facies stramineous. Eyes brown. Posterior half
of hemelytra may be hyaline and appear darker as it overlies the
dark brown dorsal surface of the abdomen. Legs stramineous.
Abdominal venter black or dark brown with keel and segmental
margins of the connexivum stramineous.

Male structural characteristics. — As viewed from above, the out-
line of the head is rounded with its greatest width equal to the
pronotal humeral width and six times the anterior width of the
vertex; synthlipsis wide, approximately four fifths the anterior width
of the vertex; along the median longitudinal axis the head is less
than half the length of the pronotum. Pronotum with its median
length three fourths the humeral width; pronotal lateral margins
diverging; posterior margin convex, medianly emarginate. Facial
tubercle slightly swollen and raised. Labrum arched along median
longitudinal line; basal width equal to median length; apex ac-
cumulate. Rostral prong (Pi. XLVIII, fig. 61b) with its apex
accumulate and its length equal to the length of the posterior mar-
gin of the third rostral segment. Anterior leg with dorsal and
ventral femoral margins sub-parallel only slightly converging for
the basal three fourths of their lengths, apex rounded. Stridulatory
comb (PI. XLVIII, fig. 61c) of approximately twenty-eight teeth,
the basal fourteen teeth twice as long as the apical fourteen, with
an abrupt decrease in length between the two groups. Chaetotaxy
of the male front leg as shown on Plate XLVIII, fig. 61a. The rela-
tive lengths of the parts of the legs are as follows :

Femur

Tibia

1st
Tar. Seg.

2nd
Tar. Seg.

100

116

Fore leg

Unfortunately, the single male specimen that I had for study was
rather mutilated, possessing only the femur and tibia of the right
fore leg.

Female structural characteristics. — As viewed from above, the
outline of the head is rounded with its greatest width nine tenths
the pronotal humeral width and four times the anterior width of
the vertex; synthlipsis wide, four fifths the anterior width of the
vertex; along the median longitudinal axis the head is slightly less
than one half the length of the pronotum. Pronotum with its

Brooks: The Genus Anisops 413

humeral width slightly less than twice its median length; lateral
margins diverging, posterior margin convex, medianly emarginate.
The relative length of the parts of the legs are as follows :

Femur

Tibia

1st

Tar. Seg.

2nd
Tar. Seg

Fore leg

Middle leg

100

100

100

80

62
40

31
25

Unfortunately I had no specimens which had a complete hind
leg.

Location of types. — Type material studied by White is in the
Perth Museum, Scotland.

Comparative notes. — By its shape, size and structural character-
istics this species appears quite closely related to Anisops thiene-
manni Lundblad. A. assimilis, however, possesses a more globose
head which is equal to the pronotum in width, whereas in A. thiene-
manni, the width of the head is less than that of the pronotum. For
comparison of the chaetatoxy of the male front legs see Plate XLVIII,
fig. 61a and Plate XLVIII, fig. 63a.

Data on distribution:

New Zealand

Queenstown, 1875, Hutton, one male, three females (F. H. Snow Coll.).
Silver Stream, 1-14-22, J. C. Myers, four females ( F. H. Snow Coll.).

Anisops thienemanni Lundblad

(PI. XLVIII, fig. 63)

1933. Anisops thienemanni Lundblad, Arch, fiir Hydrob., Suppl. vol. XII, pp. 167-168.
text fig. 57.

Referring to this species, also:

1923. Anisops hyperion\, Hale, South Australian Nat., vol. IV, no. 3, pp. 124-128, text
fig. 1, (determined in error).

1923. Anisops hyperion, Hale, Rec. South Australian Mus., vol. II, no. 3, pp. 403-412,
text fig. 365 (determined in error).

1924. Anisops hyperion, Hale, The South Australian Nat., vol. V, no. 4, p. 135 (deter-
mined in error).

1930. Anisops hyperion, Hutchinson, Proc. Zool. Soc. London, vol. XXIX, p. 145, text
fig. 2 (determined in error).

1935. Anisops hyperion, Hale, Trans. Royal Soc. South Australia, vol. XIX, p. 249 (de-
termined in error).

Size. — Males, length 6.0 mm. -7.2 mm., greatest body width, 1.6
mm.-2.1 mm.; females, length, 6.0 mm. -6.9 mm., greatest body width
1.8 mm. -1.9 mm.

Shape. — Bobust species, anterior half with lateral margins almost
parallel, lateral margins converging in posterior half.

Color. — General facies testaceous. Eyes brown. Anterior margin
of pronotum may be tinged with orange; may be hyaline on pos-

414 The University Science Btjlletin

terior and appear the black color of the underlying anterior margin
of the scutellum. Scutellum may be tinged with orange. Hemelytra
hyaline and appear black or testaceous depending on the color of
the underlying dorsal body surface. Legs stramineous. Abdominal
venter black with keel and segmental margins of the connexivum
stramineous.

Male structural characteristics. — Viewed from above, the head is
short and broad; outline rounded; greatest width of head almost
equal to the pronotal humeral width and four and one half times
the anterior width of the vertex; synthlipsis wide, slightly more
than one half the anterior width of the vertex; along the median
longitudinal axis the head is one third or less the length of the
pronotum. Pronotum long; its humeral width only one and two
fifths the median length; lateral margins almost parallel, only
slightly diverging and about one half the median length; posterior
margin almost straight, medianly emarginate. Facial tubercle only
slightly raised. Labrum long, with its median length slightly more
than its basal width; apex more or less accuminate. Rostral prong
(PI. XLVIII, fig. 63b) shorter than the third rostral segment; apex
accuminate. Anterior femur with dorsal and ventral margins almost
parallel for the basal three fourths of their length; dorsal margin
curved in apical fourth; apical fifth of inner surface with a curved
ridge extending from dorsal margin to almost midway the width.
Stridulatory comb (PI. XLVIII, fig. 63c) of approximately twenty-
four to thirty-four teeth. Chaetotaxy of male front leg as shown on
Plate XLVIII. The relative lengths of the parts of the legs are as
follows:

1st 2nd

Femur Tibia Tar. 8eg. Tar. Seg.

Fore leg 100 113 74

Middle leg 100 75 32 18

Hind leg 100 78 31 31

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with the vertex extending slightly be-
yond the anterior margins of the eyes; greatest width almost nine
tenths the pronotal humeral width and four times the anterior
width of the vertex; synthlipsis wide, slightly more than one half
the anterior width of the vertex; along the median longitudinal axis
the head is almost one half the length of the pronotum. Pronotum
with its humeral width less than twice its median length; lateral
margins diverging and one half the median length; posterior margin
convex, medianly emarginate. The relative lengths of the parts of
the legs are as follows:

Brooks: The Genus Anisops

415

1st 2nd

Femur Tibia Tar. Seg. Tar. Scg.

Fore leg 100 128 63 31

Middle leg 100 77 38 23

Hind leg 100 84 32 32

Location of types. — Type material from Diengplateau, Java in
the Stockholm Museum.

Comparative notes. — A. thienemanni is very closely related to
Anisops assimilis White, from which it can be separated only on
the basis of the males. The head of the male of A. assimilis is wider
than or at least equal to the pronotal humeral width, whereas in
A. thienemanni the head of the male is not as wide as pronotal
humeral width and the eyes are not as voluminous as in the case
of the male A. assimilis.

Data on distribution:
Australia

New South Wales, Began River, F. Armstrong, via Mr. Deane, eight males,
lour females ( F. H. Snow Coll. ) .

New South Wales, Imperial Dam, nr. Broken Hill, Fred W. Shepherd, one
male, one female (F. H. Snow Coll.).

New South Wales, Imperial Dam, XII-30-1925, F. W. Shepherd, three
males, five females (F. H. Snow Coll.).

New South Wales, Concobolin, X-17-'00, three males, two females ( Dept.
of Agrie., N. S. W., Austral.).

Broken Hill, XI-3-43, Chadwick, one male (Dept. of Agric., N. S. W.,
Austral. ) .

Broken Hill, in nine mile dam, 111-12-44, C. E. Chadwick, two females,
one male (Dept. of Agric., N. S. W., Austral.).

Broken Hill, in nine mile dam, IV-23-44, C. E. Chadwick, one male, two
females (Dept. of Agric, N. S. W., Austral.).

Broken Hill, 111-30-44, C. E. Chadwick (Dept. of Agric, N. S. W.,
Austral.).

New South Wales, Effing, Sydney, XII-27, Harv. Austral. Exped. Darling-
ton, one female (Harv. Mus. Comp. Zool.).

West Australia, Wiluna, Lake Violet, four males, nine females ( Harv. Mus.
Comp. Zool.).

West Australia, Rottnest Isl., Oct. '24, Harv. Austral. Exped. P. J. Darling-
ton, two males, nine females (Harv. Mus. Comp. Zool.).

West Australia, Wiluna, Oct. 1, 1931, Harv. Austral. Exped., P. J. Darling-
ton, one male, four females ( Harv. Mus. Comp. Zool. ) .

West Australia, Mullewa, Sept. 13, 1931, Harv. Austral. Exped., P. J. Darl-
ington, two females (Harv. Mus. Comp. Zool.).

West Australia, Geralton, Oct. 11, Harv. Austral. Exped., P. J. Darlington,
three females (Harv. Mus. Comp. Zool.).

Adelaide, H. M. Hale, ten males, twelve females (F. H. Snow Coll.).

Swamp near Camberra, III-'32, J. Evans, one male (F. H. Snow Coll.).

South Australia, Port Willunga, H. M. Hale, four males, four females
(F. H. Snow Coll.).

416 The University Science Bulletin

South Australia, Adelaide, Nov. '30, Harv. Austral. Exped., P. J. Darling-
ton, one male, one female ( Harv. Mus. Comp. Zool. ) .

South Australia, one female ( Harv. Mus. Comp. Zool. ) .

East Australia, one female (Harv. Mus. Comp. Zool.).
Tasmania

Tasmania, Hobart, IV-1939, two males, nine females, J. W. Evans ( F. H.
Snow Coll.).

Anisops nasuta Fieber

(PI. XLVIII, fig. 60; PI. LV, fig. 98)
1851. Anisops nasuta Fieber, Abhandl. Konigl. bohm. Ges. Wiss., vol. V, pt. 7, pp. 484-
485 (Fieber believed this species to be .4. nivea Fabr. ; but he simultaneously named it A.
nasuta in parenthesis).

1905. Anisops nasutus, Breddin, Societas Entomologica, vol. XVII, pp. 153-154.

1933. Anisops nasuta, Lundblad, Arch, fiir Hydrob., Suppl. Vol XII, pp. 145, 168-171,
text fig. 58.

1941. Anisops nasuta, Hoffmann, Lingnan Sci. Jour., vol. XX, pp. 59-60 (catalogue).

Referring to this species, also :

1901. Anisops fieberi, Kirkaldy, The Entomologist, vol. XXXIV, p. 5.

1901. Anisops fieberi, Breddin, Abhandl. Nat. Ges. Halle, vol. XXIV, p. 103, (ecological
note).

1904. Anisops fieberi, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 116, 132.

1906. Anisops fieberi, Distant, Fauna of British India, Rhynchota, III, p. 46.

1923. Anisops fieberi, Hale, Rec. South Australian Mus., vol. 11, no. 3, pp. 400-401, text
fig. 363.

1925. Anisops fieberi, Hale, Arch. Zool. utgivet av K. Svenska Vetensk., vol. XVII, no. 26,
p. 17 (ecological note).

1926. Anisops fieberi, Esaki, Ann. Mus. Nat. Hungarici, vol. XXIV, p. 188 (.4. niueus
Matsumura (nee. Fabricius) and Anisops Kuroiwae Matsumura synonomized here). (Ecological
note also.)

1928. Anisops fieberi, Esaki, Insects of Samoa, pt. II, pp. 76-77.

1931. Anisops fieberi, China, Jour. Federated Malay states, vol. VIII, p. 261.

1933. Aiusops fieberi, Wu, Lingnan Sci. Jour., vol. XII, p. 213 (catalogue).

1933. Anisops fieberi, Hoffmann, Lingnan Sci. Jour., vol. XII, p. 256 (catalogue).

1935. Anisops fieberi, Wu, Catalogue Insectorum Sinensium, vol. II, pp. 575-576.

1905. Anisops niveus, (nee. Fabr.) Matsumura, Trans. Sappora Nat. Hist. Soc, vol I, p. 28.
1915. Anisops niveus, (nee. Fabr.) Esaki, Ent. Mag., Kyoto, vol. I, p. 31.

1915. Anisops Kuroiwae, Matsumara, Ent. Mag., Kyoto, vol. I, p. 109.

1928. Anisops nasuta, Dover, Treubia, vol. X, p. 71 (determined in error).

1928. Anisops fieberi, Jaczewski, Ann. Mus. Zool. Polonici, vol. VII, p. 113, Plate XVI,
figs. 18, 19, 20, 21 (determined in error, actually A. genji Hutchinson).

1915. A7iisops niveus var. ogasawarensis, Matsumura, Ent. Mag., Kyoto, vol. I, pt. 3,
p. 109.

1930. Anisops ogaswarensis, Esaki, Bull. Biogeog. Soc. Japan, vol. I, no. 3, p. 214.

1933. Anisops ogasawarensis, Lundblad, Arch fiir Hydrob., Suppl. vol. XII, p. 145 (list
of Indo-australian and Pacific forms of this genus).

Size. — Males, length 6.0 mm. -7.8 mm., greatest body width 1.5
mm. -1.8 mm.; females, length, 6.0- mm. -6.9 mm., greatest body width
1.4 mm. -1.9 mm.

Shape. — Slightly fusiform species; greatest width of the body
about midway the body length.

Color. — General facies stramineous or pearlaceous. Eyes brown.
Scutellum may be tinged with orange. Legs stramineous. Ab-

Brooks: The Genus Anisops 417

dominal venter black with keel and segmental margins of the con-
nexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the vertex extending beyond the an-
terior margins of the eyes; greatest width of the head nine tenths
the pronotal humeral width and slightly more than four times the
anterior width of the vertex; synthlipsis narrow, less than one third
the anterior width of the vertex; along the median longitudinal axis
the head is seven tenths the length of the pronotum. Pronotum
with its humeral width twice its median length; lateral margins
slightly diverging and three fifths the median length; posterior mar-
gin convex, medianly emarginate. Frons produced anteriorly into a
short cephalic horn; apex of projection with a median oval depres-
sion, bordered laterally by a carina on each side; facial tubercle with
a median depression. Labrum short and broad; basal width one and
one half times the median length; apex broad and rounded; base
provided with a few long hairs which extend almost the full length
of the labrum. Rostral prong (PL LV, fig. 98) shorter than the
third rostral segment; apex accuminate. Stridulatory comb (PL
XLVIII, fig. 60b ) of approximately fourteen teeth which increase in
length toward the middle. Chaetotaxy of the male front legs as
shown on Plate XLVIII. The relative lengths of the parts of the
legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 102 58

Middle leg 100 81 34 24

Hind leg 100 81 29 29

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with the greatest width of the head nine
tenths the pronotal humeral width and five times the anterior width
of the vertex; synthlipsis wide, slightly more than one third the an-
terior width of the vertex; along the median longitudinal axis the
head is one half as long as the pronotum. Pronotum with humeral
width slightly more than twice its median length; lateral margins
diverging and one half or more the median length; posterior margin
convex, medianly emarginate. The relative lengths of the parts of
the legs are as follows:

_ 1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 118 42 31

Middle leg 100 82 35 25

Hind leg 100 87 31 31

27—3286

418 The University Science Bulletin

Location of types. — Fieber's type material was placed in the Ber-
lin Museum. Lundblad (32) makes no mention of the location of
the type and Dr. H. B. Hungerford, who, while in Europe in 1928,
recorded the location of many types of Anisops species, does not
note it as being in the Berlin Museum.

Comparative notes. — This species is superficially similar to A.
batillifrons Lundblad. However, the cephalic projection of the
males is not excavate for its entire length as in A. batillifrons and
the apex of the projection is not bordered by a raised carina as in
the case of the latter. Also the males lack the peculiarly curved
middle tarsal claws as found in A. batillifrons.

Data on distribution:
New Guinea

Port Moresby, Guigno, 1889, L. Loria, eight males, seventeen females,
Kirkaldy Collection (F. H. Snow Coll.).

Erima Astrolobe B. '97, Biro, one female, Kirkaldy Collection (F. H. Snow
Coll.).

Bigo, Luglio, 1889, L. Loria, seven males, ten females, Kirkaldy Collection
(F. H. Snow Coll.).
Celebes

Lahendong, two males, one female (Basal Nat. Mus., Switzerland).
Friendly Islands

Niva Pou Island, 1930, Lt. H. C. Kellers, M. C, seven males, one female
(U. S. Nat. Mus.).
Australia

N. Queensland, Barren Biver nr. Barren Waters, IX-28-38, B. G. Wind,
nine males, ten females (F. H. Snow Coll.).

Darwin, one male, one female (F. H. Snow Coll.).

Moore Id., Feb. 20, 1945, B. Malkin, eight males, nineteen females (U. S.
Nat. Mus.).

Townesville, Jan. 9, 1945, B. Malkin, two males, two females (U. S. Nat.
Mus. ) .

Guam

Guam, J. B. Thompson, one male (F. H. Snow Coll.).

Anisops cavifrons n. sp.

(PI. L, fisr. 73)

Size. — Males, length 5.1 mm. -5.7 mm., greatest body width 1.3
mm. -1.6 mm.; females length 5.4 mm. -6.0 mm., greatest body width
1.5 mm. -1.8 mm.

Shape. — Fusiform species, greatest body width about midway
the body length.

Color. — General facies pearlaceous, hemelytra hyaline and ap-

Brooks: The Genus Anisops

419

pearing gray as it overlies the dark dorsal body surface. Legs
testaceous. Abdominal venter dark brown with keel and segmental
margins of the connexivum testaceous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with vertex projecting beyond the anterior
margins of the eyes; greatest width of the head eight to nine tenths
the pronotal humeral width and slightly more than four times the
anterior width of the vertex; synthlipsis narrow, one fourth to one
fifth the anterior width of the vertex; along median longitudinal
axis head approximately three fifths as long as pronotum. Pronotum
with humeral width slightly less than twice its median length; lat-
eral margins diverging and more than one half the median length;
posterior margin convex, medianly emarginate. Frons triangularly
excavate; apex rounded; bordered on each side by two carinae, the
inner meet in the apical sixth of the frons to form a median com-
missure. Labrum short; basal width one and three fourths the
median length; apex broad and rounded. Rostral prong (PI. L,
fig. 73b) as long as the third rostral segment; apex accumulate.
Stridulatory comb (Pi. L, fig. 73c) of approximately fourteen teeth
which decrease in height from base to apex. Basal third of pos-
terior margin of fore tibia with a large procumbent spine which
curves medianly on dorsal suface. Tarsal claws of male middle
leg curved strongly inward at base; posterior claw thicker than
anterior one. Chaetotaxy of the front leg as shown on Plate L.
The relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 107 71

Middle leg 100 82 36 32

Hind leg 100 74 30 30

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width nine tenths the pronotal
humeral width, four to five times the anterior width of vertex;
synthlipsis wide, slightly more than one third the anterior width of
the vertex; along median longitudinal axis head more than one half
the pronotal length. Pronotum with humeral width twice the
median length; lateral margins diverging and slightly more than
one half the median length; posterior margin convex, medianly
emarginate. The relative lengths of the parts of the legs are as
follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 125 51 31

Middle leg 100 82 41 27

Hind leg 100 80 33 30

420 The University Science Bulletin

Location of types. — Male holotype, female allotype, twenty- three
male and eighty-six female paratypes, Lahori, India, Temple tank,
1-8-19, in the Indian Museum at Calcutta. Other paratypes are as
follows: In the Indian Museum: two males, ten females, Salt Range,
Punjab, India, Sta. 13, VII-29, S. L. Hora; one male, Puenjukara Isl.
Cochin backwater nr. Ernakulane, F. H. Gravely; three males, four
females, Kandaghat, Simla Hills, VIII-25, B. Chopra; six males, seven
females, Marihan, Village Tank, XII-30-12, Resee; three males, nine
females, Bannu; seven males, six females, Madhupur, Bengal,
X-16-09, C. Paiva; one male, two females, Ratnagiri Dist. Chiplum,
Vashishti V. O"-300", V-3 to 5-12, F. H. Gravely. In the Snow
Entomological Collection twenty-five males, twenty-five females,
Lahori, Temple tank, 1-8-19. In Lutz collection: two males and two
females, Cantonment Dehra Dun, U. P. India, V-10-1946, Jai L.
Uniyal.

Comparative notes. — This species is very closely related to Anisops
batillifrons Lundblad and appears identical to it in almost every
superficial detail. However, the basal third of the posterior margin
of the male fore tibia of A. batillifrons lacks the large procumbent
spine as found on A. cavifrons. For comparison of the chaetotaxies
of the front legs of the males see Plate L, fig. 73a and Plate L, fig. 71a.

Data on distribution:
India

Simla Hills, Ghuma (Ponds), 3700', IX-6 to 8-1925, B. Chopra, four males,
sixteen females ( F. H. Snow Coll. ) .

Kandaghat, Simla Hills, ponds, 3500' to 4600', Sta. 2, VIII-25, B. Chopra,
one male, seven females ( F. H. Snow Coll.).

Br. India, Coimbatore, L. V. Newton, S. J., nine males, nineteen females
(Bueno Coll. of F. H. Snow Coll.).

Br. India, Pulney, L. V. Newton, S. }., one male (Bueno Coll. of F. H.
Snow Coll.).

Inde, Darjeeling, le Moult, one male (Bueno Coll. of F. H. Snow Coll.).

Anisops batillifrons Lundblad

(PL L. fig. 71: PI. LVI, fig. 101)

1933. Anisops batillifrons Lundblad, Ann. Mag. Nat. Hist., vol. XII, ser. 10, pp. 463-
464, text fig. 8.

1935. Anisops balillifrons, Wu, Catalogus Insectorum Sinensium, vol. 2, p. 576.
1941. Anisops batillifrons, Hoffmann, Lingnan Sci. Jour., vol. 20, p. 59 (catalogue).

Size. — Males, length 5.6 mm. -6.4 mm., greatest body width 1.5
mm. -1.6 mm.; females, length 5.4 mm. -6.3 mm., greatest body width
1.3 mm. -1.8 mm.

Brooks: The Genus Anisops 421

Shape. — Fusiform species; greatest body width about two fifths'
the body length.

Color. — General facies pale, stramineous or gray. Eyes brown.
Hemelytra hyaline and appearing stramineous or gray depending
on the color of the underlying body surface. Legs stramineous.
Abdominal venter dark brown or black with keel and segmental
margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the lateral
outline of the head is rounded with the vertex extending beyond the
anterior margins of the eyes to form a short cephalic horn; greatest
width of the head nine tenths the pronotal humeral width and
slightly less than four times the anterior width of the vertex; synth-
lipsis narrow, less than one fourth the anterior width of the vertex;
along the median longitudinal axis the head is approximately three
fifths as long as the pronotum. Pronotum with its humeral width
slightly less than twice its median length; lateral margins diverg-
ing and more than one half the median length; posterior margin
convex, medianly emarginate. Frons triangularly excavate; apex
rounded; bordered on each side by two carinae, the inner two
meet in the apical sixth of the frons to form a median commissure.
Labrum short; basal width one and three fourths the median length;
apex broad and rounded. Rostral prong (Pi. LVI, fig. 101) slightly
longer than the third rostral segment; apex accuminate. Stridula-
tory comb (Pi. L, fig. 71b) of approximately thirteen teeth, the
basal six much longer than the apical seven. Middle tarsal claws
strongly curved inward at base, the posterior claw much thicker
than the anterior one. Chaetotaxy of the male front leg as shown
on Plate L. The relative lengths of the parts of the legs are as fol-
lows:

l*t 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 114 68

Middle leg 100 81 39 20

Hind leg 100 82 35 36

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with the greatest width of the head
nine tenths the pronotal humeral width and four to five times the
anterior width of the vertex; synthlipsis wide, slightly less than one
third the anterior width of the vertex; along the median longitudinal
axis the head is slightly more than one half the pronotal length.
Pronotum with its humeral width slightly more than twice its
median length; lateral margins diverging and slightly more than

422 The University Science Bulletin

one half the median length; posterior margin convex, medianly
emarginate. The relative lengths of the parts of the legs are as
follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 123 54 34

Middle leg 100 90 45 30

Hind leg 100 76 34 32

Location of types. — Type material from Foochow, China in the
Stockholm Museum.

Comparative notes. — This species is very closely related to
Anisops bouvieri Kirkaldy and the females of the two are indis-
tinguishable. However, a study of the males will afford ready
separation. The cephalic horn of Anisops bouvieri is accuminate
at the apex and extends one fifth its ventral length beyond the
anterior margins of the eyes, whereas in this species the apex of
the cephalic horn is rounded at the apex afid extends only one
twelfth its ventral length beyond the anterior margins of the eyes.
Also the males of A. batillifrons lack the three tufts of labral hairs
as found on A. bouvieri.

Data on distribution:

Formosa

Rokki, VI-17-34, L. Gressitt, one male, one female (F. H. Snow Coll.).
Rokki, V-20-34, L. Gressitt, ten males, ten females (F. H. Snow Coll.).
Taihoku, IX-9-22, R. Takahashi, nine females ( Bueno Coll. of F. H. Snow
Coll.).

Formosa, three males, ten females ( Bueno Coll. of F. H. Snow Coll. ) .
Heito, V-ll-34, L. Gressitt, one male, two females (F. H. Snow Coll.).
Yantempo, V-26-07, two females (F. H. Snow Coll.).

Iriomote Island
Iriomote Isd., VIII-25-34, L. Gressitt, one female (F. H. Snow Coll.).

Hainan Island
Fan Ta, VI-4-35, L. Gressitt (F. H. Snow Coll.).

Burma

Myitkyina, 111-31-45, L. C. Kuitert, two males, eight females (F. H. Snow
Coll.).

Myitkyina, XII-18-44, L. C. Kuitert, one female ( F. H. Snow Coll.).

Myitkyina, IX-30-45, B. McDermott, one male (F. H. Snow Coll.).

Shingbwiyang, VIII-14-44, L. C. Kuitert, ten males, twenty-three females
(F. H. Snow Coll.).

Shingbwiyang, VIII-17-44, L. C. Kuitert, one male (F. H. Snow Coll.).

Rangoon, Churchill Rd. Miss Northop, one male ( U. S. Nat. Mus. ).

Assam
Ledo, VII-3-44, L. C. Kuitert, six males, four females (F. H. Snow Coll.).

Brooks: The Genus Anisops 423

Tinsukia, IV-9-44, D. E. Hardy, ten males, twelve females ( F. H. Snow
Coll.).

Margaldal, Mozbat, 1-4-11, S. W. Kemp, four males, nine females (F. H.
Snow Coll.).

India

Darjeeling Dist. Sitong Ridge, 4700', E. Himalayas, three males, six females,
N. A. and F. G. (Indian Museum).

Darjeeling Dist. Mongpoo pond, 4000', E. Himalayas, two males, three fe-
males, N. A. and F. G. (Indian Museum).

Philippine Islands
Baeoor, P. L. Stang, one female ( F. H. Snow Coll. ) .

Mindanao, Dapitan, Baker, three males, nine females ( U. S. Nat. Mus. ).
Laguna de Bay, VIII-19-1936, two males (Usinger Coll.).

Okinawa Island
Okinawa Id., Aug. 18, 1945, W. D. Fields (U. S. Nat. Mus.).

Anisops sardea Herrich-Shaffer

(PI. XLIX, fig. 64; PI. LVI, fig. 102)

1850. Anisops sardeus Herrich-Shaffer, Die Wanzenartigen Insecten, vol. IX, p. 41, fig. 294.

*1870. Anisops sardeus, Paluinbo, Mina, Bibl. Natur. Sieil. Ent., vol. VII, p. 16.

1879. Anisops sardeus. Berg, Hemiptera Argentina, p. 119 (A sardeus appears identical
with .4. productus Fieb. to him).

1904. Anisops sardea, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 114-116, 132.

1906. Anisops sardeus. Distant, Fauna of British India, Rhynchota, vol. Ill; pp. 40-41,
text fig. 27 (ecological note).

1918. Anisops sardeus, Paiva, Rec. Indian Mus., vol. XIV, p. 28 (ecological note).

1925. Anisops sardea, Hesse, Ann. South African Mus., vol. XXIII, p. 137 (catalogue).

1926. Anisops sardea, Jaczewski, Ann. Zool. Mus. Polonici Hist. Nat., vol. V, p. 83, 83-86,
text figs. 34, 35, 36, 37.

1927. Anisops sardeus, Dover, Jour. Bombay Nat. Hist. Soc, vol. XXXII, p. 615 (eco-
logical note).

1928. Anisops sardea, Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. I, p. 164 (ecological

note).

1929. Anisops sardeus, Poisson, Bull. Soc. Hist. Nat. Afrique du Nord, vol. XX, p. 87
(ecological note).

1929. Anisops sardea, Hutchinson, Ann. South African Mus., vol. XXV, pt. 3, pp. 381-
384, PI. XXIX, fig. 13; PI. XXX, figs. 7, 8, 9.

1930. Anisops sardea, Hutchinson, Proc. Zool. Soc. London, vol. XXIX, pt. 2, p. 445 (eco-
logical note).

1930. Anisops sardea, Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. VI, p. 57 (ecological
note).

1933. Anisops sardea, Hutchinson, Internat. ges. Hydrob. und Hydrog., vol. 28, pt. 5/6,
pp. 452, 460 (ecological notes).

1933. Anisops sardea, Jaczewski, Linnean Soc. Jour., Zool., vol. XXXVIII, p. 343 (eco-
logical note).

1934. Anisops sardea, Jaczewski, Ann. Mus. Zool. Polonici. vol. X, no. 14, p. 280 (eco-
logical note).

1934. Anisops sardea, Poisson, Memoires Soc. Hist. Nat. Afrique du Nord, vol. XXV,
p. 134 (ecological note).

1935. Anisops sardea sardea, Poisson, Mus. Nat. Hist. Nat., vol. III. pp. 208-209, text
figs. 22B, 23A, 24B.

* Reference not seen by me.

424 The University Science Bulletin

1935. Anisops sardea madagascarensis, Poisson, 'loc. cit.) pp. 2091210, text figs. 22A,
23B, 24B. (I have raised this to specific level.)

1936. Anisops sardea, Jaczewski, Ann. Mus. Zool. Polonici, vol. XI, pp. 186-187 (eco-
logical note).

1939. Anisops sardea sardea, Poisson, Bull. Soc. Ent. France, vol. 44, p. 43 (ecological
note).

1941. Anisops sardea sardea, Poisson, Rev. Francaise Ent., vol. VIII, p. 77 (ecological
note).

Referring to this species also:f

1840. Anisops nivea, Spinola, Essais sur Hemipteres, p. 60 (determined in error).

1840. Anisops nivea, Rambur, Faune Entcmologique de l'Andalouise, vol. 2, no. 5, pp.
190-191 (determined in error).

1843. Anisops nivea, Amyot and Serville, Hemipteres, p. 454, PL VIII, fig. 8 (determined
in error).

1848. Anisops nivea, Costa, Atti Reale Inst. Incorag. Alle Sci. Nat. Napoli, vol. Ill, p. 148
(determined in error).

1848. Anisops nivea, Amyot, Entomologie Francaise, Rhynchota, p. 338 (determined in
error).

1855. Anisops nata!ensis, Stal, Ofversift Kongl. Vetensk. Akad. Forhand., vol. XII, p. 89
(determined in error).

1851. Anisops productus, Fieber, Abhandl. Bohm. Ges. Wiss., vol. V, pt. 7, p. 484.

1860. Anisops productus, Fieber, Die Europaische Herniptera, p. 100.

1862. Anisops productus,, Schawn, Peter's Reise nach Mossambique, V. p. 51.

1865. Anisops productus, Stal, Herniptera Africana III, p. 191.

*1872. Anisops productus, Marshall, Ent. Month. Mag., vol. VIII. p. 191.

1873. Anisops producta, Gerstaecker, Decken Reise's in Ost Africa III, no. 2, p. 425.

1878. Anisops producta, Horvath, Entomologische Nachrichten, vol. IV, p. 174.

*1880. Anisops producta, Puton, Synopsis des Heteropteres de France, p. 217.

1886. Anisops producta, Puton, Exploration Scientifique de la Tunisie, p. 8 (ecological
note).

1892. Anisops producta, De Carlini, Ann. Mus. Civ. Storin Nat. Genova, vol. XII
(XXXII), p. 11 (ecological note).

1893. Anisops producta, Noualhier, Ann. Soc. Ent. France, vol. LXII, p. 17 (ecological
note).

1899. Anisops producta, Kirkaldy, Ann. Soc. Ent. France, vol. LXVI1I, p. 106 (ecological
note).

1883. Anisops producta, Lethierry, Ann. Mus. Civ. Storia Nat. Genova, vol. XVIII, p. 2
(ecological note).

1892. Anisops producta, Liethierry, Bull. Soc. Ent. France, vol. XVII, p. 208 (ecological
note).

*1894. Anisops producta, Giglia-Tos, Boll. Mus. Zool. Anat. Comp., vol. IX, p. 11.

1905. Anisops producta, Breddin, Mitt. Naturh. Mus.. vol. XXII, pp. 153-154.

1919. Anisops producta, Hungerford, Kansas Sci. Bull., vol. XXI, p. 176.

1922. Anisops producta, Lindberg, Notulae Entomologicae, vol. II, pp. 47-48, text figs.
5, 6.

1926. Anisops producta, Poisson, Acad. Sci. Paris, vol. 181, p. 684.

1926. Anisops producta, Poisson, Arch. Zool. Exper. Gen., vol. 65, pt. 4, pp. 181-208,
text figs. I-XVII (bionomics).

1926. Anisops producta, Poisson, Bull. Soc. Hist. Nat. Afrique du Nord, vol. XXII,
p. 269 (ecological note).

1936. Anisops producta, Lindberg, Soc. Scient. Fennica Comment. Biol., vol. VI, no. 7,
pp. 41-42 (ecological note).

° Reference not seen by me.

f The following reference as noted by Kirkaldy in 1904 (30) was unavailable: ? IV. alba
Forskal, Deer. Anim. Orient, p. XXIII. The name A. compressa as noted by Herrich-Shaffer
and Kirkaldy ( 30 ) evidently is a manuscript name, for there is no reference to it in the litera-
ture.

Brooks: The Genus Anisops 425

Size. — Males, length 7.5 mm. -8.4 mm., greatest body width 1.8
mm. -1.9 mm.; females, length 7.2 mm.-7.5 mm., greatest body width
1.9 mm. -2.0 mm.

Shape. — Subfusiform species; greatest body width about midway
the length of the body.

Color. — General facies stramineous or testaceous. Eyes brown.
Hemelytra may be hyaline and appear darker due to dark brown
body surface. Legs stramineous. Abdominal venter dark brown
or black with keel and segmental margins of the connexivum stra-
mineous or testaceous.

Male structural characteristics. — Viewed from above, the inter-
ocular space is produced anteriorly into a prominent cephalic horn
with its apex accumulate; lateral margins of the head rounded;
greatest width of the head nine tenths the pronotal humeral width
and three times the anterior width of the vertex; synthlipsis narrow,
approximately one sixth the anterior width of the vertex; along the
median longitudinal axis the head is slightly more than three fourths
the length of the pronotum. Pronotum with its humeral width al-
most twice the median length; lateral margins diverging and almost
one half the median length; posterior margin convex, medianly
emarginate. Frons, which forms the ventral margin of the cephalic
horn, excavate for its entire length and bordered laterally by two
carinae; apex more or less accumulate. Labium short; basal width
slightly more than twice the median length; apex accumulate; each
basal angle bearing a tuft of erect hairs whose tips are curved
medianly and meet one another, forming a loop. Rostral prong (Pi.
LVI, fig. 102) slightly shorter than the third rostral segment; apex
accumulate. Stridulatory comb (Pi. XLIX, fig. 64b) of approxi-
mately eighteen teeth which decrease slightly in length from base to
apex. Middle tarsal claws strongly curved inward at base; posterior
claw thicker than anterior claw. The chaetotaxy of the front leg as
shown on Plate XLIX. The relative lengths of the parts of the legs
are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 118 72

Middle leg 100 83 42 20

Hind leg 100 84 30 30

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded, with the vertex extending slightly be-
yond the anterior margins of the eyes; greatest width of head nine

426 The University Science Bulletin

tenths the pronotal humeral width and approximately four times
the anterior width of the vertex; synthlipsis wide, one third the an-
terior width of the vertex; along the median longitudinal axis the
head almost as long as the pronotum. Pronotum with its humeral
width twice the median length; posterior margin convex, medianly
emarginate. Facial tubercle with a slight triangular depression.
The relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 118 55 33

Middle leg 100 86 42 23

Hind leg 100 80 31 30

Location of types. — The location of the type material of A. sardea
appears to be unknown. As it was part of Herrich-Shaffer's own
collection it may be in the Berlin Museum, though Dr. H. B. Hun-
gerford does not record it in the notonectid material he observed
there in 1928.

Comparative notes. — Though somewhat larger, this species, by
the presence and shape of the male cephalic projection is very
similar in appearance to A. bouvieri Kirk. Besides being larger it is
different in many other respects from Kirkaldy's species. It lacks
the laterally expanded keel of the third abdominal sternite as pos-
sessed by A. bouvieri. A. bouvieri does not have the marginal row
of prominent setae on the inner surface of the fore tibia of the male
of A. sardea.

Data on distribution:
India

Baluchistan, Yakmach Cagai, XI-13-18, B. S., one male (F. H. Snow Coll.).

Punjab, Salt Pangi, Sta. 9. S. L. Hora, two males, two females (F. H.
Snow Coll.), seven males, three females (Indian Museum).

Moriban, Village tank, XII-20-15, Resee, four females (Indian Museum).

S. India. Chikkaballpura, T. V. Campbell, two females (F. H. Snow Coll.).

Kuman Lakes, one female (Indian Museum).

Bannu, one female (Indian Museum).

Lucknow, 1-22-08, Mus. Collr. R. H., one male, four females (Indian
Museum).

S. India, Coromandel, 2500' X-23-10, Mus. Collr., one male (Indian Mu-
seum ) .

Turkey

Erdemli, VIII-27-47, two males, two females (F. H. Snow Coll.).
Syria

Syria, Kaifa, Reuter, exchange fr. Budapest Museum, two females ( F. H.
Snow Coll. ).

Brooks: The Genus Anisops 427

Corfu

Corfu, J. Sahlberg, gift fr. Horvath, Kirkaldy remnant, one male, one female
(F. H. Snow Coll.).
Albania

Mustajebg Velipoja, one male ( F. H. Snow Coll. ) .
Africa

Egypt, Dr. Tuton, one male (U. S. Nat. Mus. ).

Egypt, Mariout, 1881, Letournex, two males, two females (Paris Museum).

Egypt, R. Amle, 1881, Letournex, three females (Paris Museum).

Basse Egypte, Le Caire et Env. 1907, Ch. Alluaud, one male, three females
(Paris Museum).

Soudan Egyptian, Poseires, Haut Nil Bleu, Dec. 1907, Ch. Alluaud, one
male, one female (Paris Museum).

Soudan, Nioro, Oct. 1909, F. de Zeltner, one male (Paris Museum).

Algerie, Bone, Hgemn, one female ( F. H. Snow Coll. ) .

Algerie, Philippville, A. Thery, three females, one male (Bueno Coll. of
F. H. Snow Coll.).

Abessynien, one female (Bueno Coll. of F. H. Snow Coll.).

Abessynien, Dire Doana, two females ( Bueno Coll. of F. H. Snow Coll. ) .

Cairo, four females ( Bueno Coll. of F. H. Snow Coll. ) .

Algerie, Constantine, E. de. Bergevin, one female (Paris Museum).

Morco, Si Rahal, 1913, R. Pallary, one male, one female (Paris Museum).

Moroc, Oued, Mharhar pris Bow Berein, 1903, C. Buchet, three males, three
females (Paris Museum).

Moroc, Andiere, Boegel Merare, 1903, C. Buchet, one female (Paris Mu-
seum ) .

Tchad, N'Guigmi, 1919, Dr. Noel, one female, one male (Paris Museum).

Tchad, Mission Tihlo, 1910, Dr. Gaillard, two females (Paris Museum).

Bas Chari, Fort Lamt, Mission Chari-Tchad, Aug. 1904, Dr. J. DeCorse,
three males, six females ( Paris Museum ) .

Reg. de Zinder, Dungass, Mission Tilho, Oct. 1910, Dr. R. Guillard, five
males, seven females (Paris Museum), one male, one female, Jan. 1910 (Paris
Museum ) .

Sahel Soudanais, Goumbou, 1909, F. de Zeltner, one male, three females
( Paris Museum ) .

Env. de Zinder, Guidimouni, Mission Tilho, Nov. 1910, Ch. Alluaud, one
male, one female (Paris Museum).

Mocambique, Vallee du Pungoue, Guengere, 1906, G. Vassee, one male,
three females ( Paris Museum ) .

Kanem, Sud de Nyouri, 1904, A. Chevalier, two males (Paris Museum).

Congo Francaise, Liberville, 1898, C. Chalot, one male (Paris Museum).

Haut Oubangui, Bessou (Mission), 1904, Dr. J. DeCorse, one female (Paris
Museum).

Massahori, 66 K. est du Tchad, 1904, A. Chevalier, one female (Paris Mu-
seum ) .

Afrique Orient. Angl. Maji-Chumi, (Wa-Nyika) 1904, Ch. Alluaud, one
female ( Paris Museum ) .

428 The University Science Bulletin

Tombouctore, 1900, A. Chevalier, one male, two females (Paris Museum).

Abyssinia, Hora Abjata, small pond in marsh, XI-18-1926, J. Omer-Cooper,
one female ( F. H. Snow Coll.).

E. Africa, P. H. Uhler Collection (U. S. Nat. Mus.).

Tanga, Sjostedt, gift from Sweeny (Kirkaldy remnant) (F. H. Snow Coll.).

Transvaal, Klobers Dam, Setagali Dist, Mafering, Jan. 1927.

B. Lamb, gift to H. B. Hungerford fr. G. E. Hutchinson, one male, one
female ( F. H. Snow Coll.).

Kilimandjoro, Sjostedt, Natron-sjoarne, gift fr. Sweeny, one female ( F. H.
Snow Coll. ).

French Sudan, Bamako, purchased fr. Dr. O. Standlinger, one male, five
females ( F. H. Snow Coll.).

Oran, E. Morrison, one male, three females ( F. H. Snow Coll.).

Canary Islands
He de Canaries, 1897, C. Buchet, two males, two females (Paris Museum).
Islas Canarias, Tenerife, A. Cabrera, one male, six females (Bueno Coll. of
F. H. Snow Coll.).

Tenerife, two females (Bueno Coll. of F. H. Snow Coll.).

Anisops madagascarensis Poisson

(PI. XLIX, fig. 67)

1935. Anisops sardea madagascarensis Poisson, Bull. Mus. Nat. Hist. Natur., vol. HI,
pp. 208-211, text figs. 22A, 23A, 24B.

Size.— Males, length 8.1 mm. -9.0 mm., greatest body width 1.9
mm.-2.1 mm.; females, length 7.3 mm.-7.8 mm., greatest body width
1.9 mm. -2.1 mm.

Shape. — Subfusiform species; greatest width of the body slightly
before midway the body length.

Color. — General facies testaceous. Eyes brown. Hemelytra may
be hyaline and appear darker due to dark brown body surface. Legs
stramineous. Abdominal venter dark brown, with keel and seg-
mental margins of the connexivum stramineous. Tip of cephalic
horn black.

Male structural characteristics. — Viewed from above, the inter-
ocular space is produced anteriorly into a prominent cephalic horn
with its apex accuminate; lateral margins of the head rounded;
greatest width of the head eight tenths the pronotal humeral width
and three times the anterior width of the vertex; synthlipsis narrow,
one sixth the anterior width of the vertex; along the median longi-
tudinal axis the head is three fourths the length of the pronotum.
Pronotum with its humeral width one and two thirds the median
length; lateral margins diverging and three fifths the median length;
posterior margin convex, medianly emarginate. Frons, which forms

Brooks: The Genus Anisops 429

the ventral margin of the cephalic horn, excavate for its entire
length and bordered laterally by two carinae; apex more or less
accumulate. Labrum short; basal width slightly more than twice
the median length; apex accuminate; each basal angle bearing a tuft
of erect hairs which curve their tips medianly and meet one another
forming a loop. Rostral prong as in A. sardea (PI. LVI, fig. 102)
longer than third rostral segment; apex accuminate. Stridulatory
comb ( PL XLIX, fig. 67b ) of approximately seventeen teeth, which
decrease slightly in length from base to apex. Middle tarsal claws
strongly curved inward at the base, posterior claw thicker than an-
terior one. Chaetotaxy of male front leg as shown on Plate XLIX.
The relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 119 64

Middle leg 100 88 42 19

Hind leg 100 83 33 33

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded, with the vertex extending slightly be-
yond the anterior margins of the eyes; greatest width of the head
nine tenths of the pronotal humeral width and almost four times the
anterior width of the vertex; synthlipsis narrow, slightly more than
one fourth the anterior width of the vertex; along the median longi-
tudinal axis the head is slightly more than one half the length of the
pronotum. Pronotum with its humeral width twice its median
length; lateral margins diverging and almost one half the median
length; posterior margin convex, medianly emarginate. Facial
tubercle with a faint triangular depression. The relative lengths of
the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 118 54 29

Middle leg 100 81 41 21

Hind leg 100 82 34 32

Location of types. — Type material from Madagascar in the Paris
Museum.

Comparative notes. — This species is almost identical with A.
sardea H.-S. from which it can be best separated on the basis of
the chaetotaxy of the male front leg. The setae of the fore tibia in
this species are never spatulate whereas the second and third basal
setae of Anisops sardea are spatulate. The tip of the cephalic horn
in all the specimens that I have observed is black, a condition
pointed out by Poisson (loc. cit.). This peculiarity is not en-
countered in A. sardea.

430 The University Science Bulletin

Remarks. — This species was first described as a sub-species by
Poisson ( loc. cit. ) . However, I am placing it on specific level on the
basis of the above differences between it and A. sardea H.-S.

Data on distribution:
Madagascar

Tananarive, 1921, R. Decary, five males, nineteen females (Paris Museum);
two males, two females (F. H. Snow Museum).

Marais du Finerena, 1905, F. Geay, two males, eight females (Paris
Museum ) .

Madagascar, 1897, E. Dorr, two females (Paris Museum).

Madagascar, 1897, F. de Zeltner, one female (Paris Museum).

Plaines du Sakondry, 1906, F. Geay, one female (Paris Museum).

Madagascar, P. Camboue, two females (Paris Museum).

Tananarive, purchased fr. Prof. Lamberton Nov. 1931, one male, one female
(F. H. Snow Coll.).

Maroantsetra, purchased fr. Dr. O. Standinger, one female (F. H. Snow
Coll.).

Mocquereya, Chutes de Samilia, Riv. N. Gamie, G. Severin, one male ( U. S.
Nat. Mus. ).

Madagascar, one male, two females (Rueno Coll. of F. H. Snow Coll.).

Anisops bouvieri Kirkaldy

(PI. XLIX, fig. 65; PI. LV, fig. 100)

1904. Anisops bouvieri Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 116, 132.
1933. Anisops bouvieri, Hoffmann. Lingnan Sci. Jour., vol. 12, p. 256 (catalogue).
1933. Anisops bouvieri, Lundblad, Arch, fur Hydrob., Suppl. vol. XII, p. 145 (a list of
Indo-australian and Pacific members of this genus).

1935. Anisops bouvieri, Wu, Catalogus Insectorum Sinensium, vol. II, p. 575.
1941. Anisops bouvieri, Hoffmann, Lingnan Sci. Jour., vol. 20, p. 59 (catalogue).

Size. — Males, length 6.0 mm.-6.3 mm., greatest body width 1.5
mm. -1.8 mm.; females, length 5.7 mm.-6.0 mm., greatest body width
1.7 mm. -1.8 mm.

Shape. — Fusiform species; greatest body width about two fifths
the body length.

Color. — General facies pearlaceous. Eyes gray. Abdominal ven-
ter dark brown with keel and segmental margins of the connexivum
pearlaceous.

Male structural characteristics. — Viewed from above, the lateral
outline of the head is rounded; vertex extended anteriorly into a
short cephalic horn; greatest width of the head nine tenths the pro-
notal humeral width and slightly more than three times the anterior
width of the vertex; synthlipsis narrow, almost one fourth the
anterior width of the vertex; along the median longitudinal axis the
head minus the cephalic horn is two thirds the length of the pro-
notum. Pronotum with its humeral width one and four fifth times

Brooks: The Genus Anisops

431

the median length; lateral margins diverging and two thirds the
median length; posterior margin convex, medianly emarginate.
Viewed ventrally the frons which forms the ventral margin of the
cephalic horn is triangularly excavate; bordered laterally by two
carinae; apex more or less accuminate. Labrum short and broad;
basal width slightly more than one and one half its median length;
apex accuminate; bears three tufts of hairs, one at each basal angle
and one at the apex. Rostral prong (PL LV, fig. 100), as long as
the third rostral segment; apex accuminate. Stridulatory comb
(PL XLIX, fig. 65b) of approximately twelve teeth which decrease
in length from base to apex. Middle tarsal claws strongly curved
inward at base, posterior claw thicker than anterior one. The keel
of the third abdominal segment is flattened and bordered on its
lateral and apical margins by stout erect hairs. Chaetotaxy of the
male front leg as shown on Plate XLIX. The relative lengths of
the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 102 75

Middle leg 100 74 38 33

Hind leg 100 78 32 28

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head nine tenths the
pronotal humeral width and slightly more than three times the
anterior width of the vertex; synthlipsis wide, almost one half the
anterior width of the vertex; along the median longitudinal axis the
head is two thirds as long as the pronotum; lateral margins diverg-
ing and almost three fifths the median length; posterior margin
convex, medianly emarginate. The relative lengths of the parts of
the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 114 57 29

Middle leg 100 90 45 22

Hind leg 100 84 32 28

Location of types.— Male type from Cochin China, in the Paris
Museum.

Comparative notes. — This species is closely related to Anisops
batillifrons Lundblad, however the cephalic horn is more or less
accuminate and extends one fifth its ventral length beyond the
anterior margin of the eyes, whereas in A. batillifrons the cephalic
horn is rounded at the apex and extends only one twelfth its ventral
length beyond the anterior margin of the eyes. In addition.

432 The University Science Bulletin

A. batillifrons lacks the tufts of labral hairs that are found on this
species.

Data on distribution:

New Guinea

Simbang Huon Gulf, 1898, Biro, one male, twenty-four females (F. H.
Snow Coll.).

Siam
Brangkok, Sept. 17, 1926, H. M. Smith (U. S. Nat. Mus.).

Burma
Mohnyin, X-27-44, L. C. Kiuitert, two males, three females (F. H. Snow
Coll.).

Assam
Tinsukia, IV-9-44, D. E. Hardy, one male (F. H. Snow Coll.).

India

Barkuda, Chilka Lake, Ganjam Dist. Madras Pres. VIII-19, F. H. Gravely,
twenty-five males, twenty-five females ( F. H. Snow Coll. ) ; thirty males, thirty-
six females (Indian Museum).

Barkuda, Chilka Lake, Ganjam Dist. Madras Pres. VII-23-19, N. A. one
male, twenty-two females (Indian Museum).

Waltair, 1-27-21, fourteen males, nine females (Indian Museum).

Calcutta, Eden Gardens at light, X-26-11, F. H. Gravely, one male, two
females (Indian Museum).

Calcutta, Eden Gardens at light, X-ll, F. H. Gravely, one female (Indian
Museum ) .

Calcutta, VIII-9-'06, two males (Indian Museum).

Bengal, Berhampore, Court, IV-13-10 S. W. K., one male (Indian Museum).

Bengal, Madhupur, X-16-09, C. Paiva, two males, seven females (Indian
Museum ) .

Bengal, Berhampur Murshidabad, Sept. '12, Southwell, one male, five fe-
males ( Indian Museum ) .

East Bengal, Damukdia Ghat, VII-22-17, Mus. Collr. four males, two fe-
males (Indian Museum).

Central Province, Purhapura, III-4-19, C. P. and F. H. G., twelve males,
seven females (Indian Museum).

Puri Dist. Barkul, Orissa, XI-9th to 13th-12, Gravely, six males, twenty-one
females (Indian Museum).

Madras, Ganjam Dist, Bambha, IX-20-13,, Annandale, seventeen males, five
females (Bueno Coll. of F. H. Snow Coll.).

Andamans, fresh water tank, III-31-ll, C. Paiva, three males, one female
(Bueno Coll. of F. II. Snow Coll.).

Orissa, Lake Chilka, Satpara, IX-10-13, Annandale, seventeen males, two
females ( Bueno Coll. of F. H. Snow Coll. ) .

Anisops extendofrons n. sp.

(PI. XLIX, fig. 66)

Size. — Males, length 6.4 mm.-6.8 mm., greatest body width 1.5

mm. -1.7 mm.; females, length 6 mm., greatest body width 1.8 mm.

Shape. — Subfusiform species; lateral margins in anterior half of

Brooks: The Genus Anisops 433

body, with exception of the head, almost parallel; posterior lateral
margins converging.

Color. — General facies pearlaceous. Eyes brown. Legs testa-
ceous. Abdominal venter dark brown or black, with keel and seg-
mental margins of the connexivum testaceous.

Male structural characteristics. — Viewed from above, the lateral
outline of the head is rounded with the anterior interocular space
extended forward in a cephalic horn; greatest width of the head nine
tenths the pronotal humeral width, slightly more than three times
the anterior width of the vertex; synthlipsis narrow, less than one
third the anterior width of the vertex; along the median longitudinal
axis the head is approximately three fourths the pronotal length.
Pronotum with its humeral width almost twice its median length;
lateral margins diverging and slightly more than two thirds the
median length; posterior margin convex, medianly concave. Viewed
ventrally the frons, which forms the ventral surface of the cephalic
horn, is triangularly excavate, bordered laterally by two carinae;
apex more or less accuminate. Viewed laterally the cephalic horn
extends one third its ventral length beyond the anterior margin of
the eyes. Labrum short; basal width almost twice its median
length; apex accuminate; bears three tufts of anteriorly curved hairs,
one at each basal angle and one at the apex. Rostral prong slightly
shorter than third rostral segment; apex accuminate. Fore tibia
(PI. XLIX, fig. 66b) with anterior margin curved at apex; stridula-
tory comb of approximately fourteen teeth which decrease in height
from base to apex. Middle tarsal claws turned strongly inward at
base; posterior claw thicker than anterior claw. Chaetotaxy of the
male front leg as shown on Plate XLIX. The relative lengths of the
parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 123 96

Middle leg 100 92 46 23

Hind leg 100 78 30 29

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest head width nine tenths the
pronotal humeral width, slightly less than four times the anterior
width of the vertex; synthlipsis wide, almost one half the anterior
width of the vertex; along the median longitudinal axis the head is
one half the pronotal length. Pronotum with humeral width twice
its median length; lateral margins diverging and three fifths the

28—3286

434 The University Science Bulletin

median length; posterior margin convex, medianly emarginate. The
relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 114 60 29

Middle leg 100 80 40 22

Hind leg 100 78 35 33

Location of types. — Male holotype, female allotype, five male and
three female paratypes, Waltair, India, 1-27-21 in the Indian Mu-
seum at Calcutta. Other paratypes are as follows: In the Indian
Museum at Calcutta: one male, Puenjikara Isl. Cochin Backwater
nr. Ernakulane, India, X-14, F. H. Gravely; one male, one female,
Satara Dist. 2000', Nechal, W. Chata, F. H. Gravely. In the Snow
Entomological Collection: five males, three females, Waltair, India,
1-27-21.

Comparative notes. — Extremely closely related to Anisops bou-
vieri Kirk, and superficially the two species appear the same. How-
ever, careful examination of the males will show distinct differences.
The anterior margin of the fore tibiae lack the subapical angle as
found on A. bouvieri and also the long spine borne at the angle.
The keel of the third abdominal segment of A. extendofrons is not
flattened and bordered with hairs as in A. bouvieri, but covered with
hairs.

Data on distribution:

India
Br. India, Coimbatore, L. V. Newton S. J., one male (Bueno Coll. of F. H.
Snow Coll. ).

Anisops graciloides n. sp.

(PI. LI, fig. 79)

Size. — Males, length 7.0 mm. -8.2 mm., greatest body width 1.5
mm.-2.1 mm.; females, length 7.5 mm., greatest body width 1.9 mm.

Shape. — Slender, fusiform species; greatest width about mid-
way the body length.

Color. — General facies testaceous. Eyes brown or gray. Legs
stramineous. Abdominal venter irregularly colored dark brown
and gray.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of head nine tenths the
pronotal humeral width and six and one-half times the anterior
width of the vertex; synthlipsis narrow; one fifth the anterior width
of the vertex; along the median longitudinal axis the head is slightly
more than two thirds the length of the pronotum. Pronotum with

Brooks: The Genus Anisops

435

its humeral width almost twice its median length; lateral margins
only slightly diverging and slightly more than one half the median
length; posterior margin convex, medianly emarginate. Facial
tubercle slightly compressed laterally with a slight median depres-
sion. Labrum long, basal width slightly less than its median length;
apex rounded. Rostral prong (PI. LI, fig. 79b) shorter than third
rostral segment; apex rounded. Stridulatory ridge ( PI. LI, fig. 79a )
transversely striated. Stridulatory comb (Pi. LI, fig. 79c) of ap-
proximately sixteen teeth which increase slightly in length from
base to apex. Chaetotaxy of front leg as shown on Plate LI.
The relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. .Seg.

Fore leg 100 112 56

Middle leg 100 95 41 28

Hind leg 100 92 34 34

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head nine tenths the
humeral width and almost four times the anterior width of the
vertex; synthlipsis wide, one third the anterior width of the vertex;
along the median longitudinal axis the head is almost two thirds
the length of the pronotum. Pronotum with its humeral width
twice the median length; lateral margins diverging and one half
the median length; posterior margin convex, medianly emarginate.
The relative lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 113 50 38

Middle leg 100 89 42 27

Hind leg 100 83 34 34

Location of types. — Male holotype and female allotype, Salisbury,
S. Rhodesia, X-16-15, in the Snow Entomological Collection. Two
male paratypes Africa nr. Kabale, Uganda, L. Bungymi, Feb. 1948,
R. X. Williams in the Harvard Museum of Comparative Zoology.

Comparative notes. — This species is closely related to Anisops
gracilis Hutchinson from which it can be separated by the fact that
the males have the facial tubercle slightly compressed laterally
and bearing a faint median depression. This condition is lacking
on the males of A. gracilis. Also the rostral prong has a rounded
apex instead of the accuminate apex of A. gracillis.

Data on distribution. — Known only from type series.

436 The University Science Bulletin

Anisops gracilis Hutchinson*

(PI. LI, fig. 74)

1929. A7iisops gracilis Hutchinson, Ann. South African Mus., vol. XXV, pt. 3, pp. 386-
388, PI. XXX, fig. 12; PI. XXXI, fig. 4; PI. XXXII, fig. 12.

Size. — Males, length 6.9 mm., greatest body width 1.8 mm.

Shape. — Slender, fusiform species; greatest width about halfway
the body length.

Color. — General facies stramineous. Eyes brown. Abdominal
venter dark brown with keel and segmental margins of the con-
nexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of head nine tenths the pro-
notal humeral width and five and one-half times the anterior width
of the vertex; synthlipsis narrow, one fourth the anterior width of
the vertex; along the median longitudinal axis the head is three
fourths the length of the pronotum. Pronotum with its humeral
width slightly less than twice the median length; lateral margins
slightly diverging and one half the median length; posterior margin
convex, medianly emarginate. Facial tubercle slightly raised. Lab-
rum long, median length equal to the basal width; apex rounded.
Rostral prong (Pi. LI. fig. 74b) longer than third rostral segment;
apex accuminate. Stridulatory ridge (Pi. LI. fig. 74c) with trans-
verse striations; stridulatory comb of approximately fourteen even-
length teeth. Chaetotaxy of the male front leg as shown on Plate
LI. The relative lengths of the parts of the legs are as follows :f

1st

2nd

Femur

Tibia

Tar. Seg.

Tar. Seg.

Fore leg;

100

113

62

. .

Hind les

100

S3

31

33

Location of types. — Type material from South Africa in the South
African Museum.

Comparative notes. — The body shape and coloration are similar
to that of A. balcis Hutchinson from which it can be separated by
the fact that the males of A. gracilis lack the flattened facial tubercle
so characteristic of A. balcis. The latter species does not have a
striated stridulatory ridge as found on the former species.

Data on distribution:
Africa

South Africa, Messina, Limpopo gorge, V-27-1921, gift to H. B. H. fr. G. W.
Hutchinson, one male (F. H. Snow Coll.).

* Only a single male specimen of this species was in my material,
t Minus the middle pair of legs.

Brooks: The Genus Anisops 437

*

Anisops poweri Hutchinson

(PI. L, fig. 72; PI. LIV, fig. 95)

1929. Ayusops poweri Hutchinson, Ann. South African Mus., vol. XXV, pt. 3, pp. 398-
390, PI. XXX, fig. 13; PI. XXXI, fig. 6, 6a.

1933. Anisops poweri, Hutchinson, Internat. ges. Hydrob. und Hydrog., vol. 28, pt. 5/6,
p. 460 (ecological note).

Size. — Males, length 5.7 mm., greatest body width 1.5 mm.

Shape. — Small, fusiform species; greatest body width about mid-
way the body length.

Color. — General body color stramineous. Eyes brown. Legs
stramineous. Abdominal venter dark brown with keel and seg-
mental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of the head is eight tenths
the pronotal humeral width and slightly more than three times the
anterior width of the vertex; synthlipsis narrow, slightly more than
one fourth the anterior width of the vertex; length of the median
longitudinal axis the head is slightly more than one half the length of
the pronotum. Pronotum with its humeral width almost twice the
median length; lateral margins diverging and two thirds the median
length; posterior margin convex, medianly emarginate. Facial
tubercle greatly swollen. Labrum with its basal width one and one
half the median length; apex rounded. Rostral prong (Pi. LIV,
fig. 95) slightly shorter than the third rostral segment. Stridulatory
comb broken off of the legs of the specimen in my material. Chae-
totxy of the male front leg as shown on plate L. The relative length
of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 105 56

Middle leg 100 87 41 26

Hind leg 100 81 32 36

Location of types. — Type material from Kimberly Cape, South
Africa in the South African Museum.

Comparative notes. — The long claws of the male front leg are
found on only one other species, A. leesoniana Hutch, from which it
may be readily distinguished on the basis of the male facial tubercle
and the fore tarsus. The facial tubercle of A. poweri is greatly
swollen and the basal inner surface of the fore tarsus bears a median
row of three long spines. Neither of these conditions is found on
A. leesoniana.

* Only a male specimen was present in my material.

438 The University Science Bulletin

Data on distribution:

Africa

Orange River, Diep Klof, east of Aliwil N., IX-5-28, gift to H. B. H. fr. G. E.
Hutchinson, one male ( F. H. Snow Coll. ) .

Anisops leesoniana Hutchinson*

(PI. L, fig. 70)

1929. Anisops leesoniana Hutchinson, Ann. South African Mus., vol. XXV, pt. 3, pp. 390-
3S1, PI. XXX, fig. 14; PI. XXXI, fig. 7.

Size. — Males, length 4.8 mm.; greatest body width 1.3 mm.

Shape. — Small, fusiform species; greatest body width midway the
body length.

Color. — General facies stramineous. Abdominal venter dark
brown with keel and segmental margins of the connexivum stra-
mineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the greatest width of the head almost
equal to the pronotal humeral width and seven times the anterior
width of the vertex; synthlipsis narrow, one sixth the anterior width
of the vertex; along the median longitudinal axis the head is three
fourths the length of the pronotum. Pronotum with its humeral
width almost twice its median length; lateral margins almost parallel
and one half the median length; posterior margin convex, medianly
emarginate. Facial tubercle slightly raised. Labrum with lateral
margins strongly converging; basal width equal to the median
length; apex more or less accumulate. Rostral prong (PL XV, fig.
70b) slightly shorter than third rostral segment; apex accuminate.
Stridulatory comb (PI. L, fig. 70c) of approximately twenty-three
teeth, with the apical one slightly shorter than the remainder.
Chaetotaxy of the male front leg as shown on Plate L. The relative
lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 105 70

Middle leg 100 86 41 25

Hind leg 100 87 31 31

Location of types. — Male holotype from South Rhodesia in the
South African Museum.

Comparative notes. — This species and A. poweri Hutchinson are
unique in having the extremely long front tarsal claws, more than
one half the length of the fore tarsi. Anisops leesoniana differs in
many respects from A. poweri however; the facial tubercle is not

* Only a single male specimen in my material and it is badly mutilated.

Brooks: The Genus Anisops 439

greatly swollen as in A. poweri, and the fore tarsal claws are not
curved as in the case of the latter.
Data on distribution:

Africa
South Africa, Rikatle, Lourenco Marques. Coll. Junod., one male (Basel
Nat. Hist. Mus., Switzerland).

Anisops breddeni Kirkaldy

(PI. LI, fig. 78; PI. LV, fig. 99)

1901. Anisops breddeni Kirkaldy, The Entomologist, vol. 34, pp. 5-6.

1901. Anisops breddeni, Breddin, Abhand. Naturf. Gesell. Halle, vol. XXIV, p. 103 (eco-
logical note).

1904. Anisops breddeni, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 117, 132.

1910. Anisops breddeni, Distant, Fauna of British India, Rhynchota, vol. V, p. 333, fig.
194 (ecological note; expresses doubt as to its belonging in this genus).

1928. A7iisops breddeni, Dover, Treubia, vol. X, p. 171 (ecological note, disagrees with
Distant and believes it should remain in the genus).

1933. Anisops breddeni, Lundblad, Archiv. fiir Hydrob. Suppl. vol. XII, p. 145 (a list of
Indo-australian and Pacific members of this genus).

Size. — Males, length 5.7 mm. -6.6 mm., greatest body width 1.2
mm. -1.6 mm.; females, length 5.4 mm.-5.7 mm., greatest body width
1.2 mm. -1.5 mm.

Shape. — Slender, slightly fusiform species; greatest width about
one half the body length.

Color. — General facies pearlaceous. Eyes brown. Legs stra-
mineous. Abdominal venter dark brown with keel and segmental
margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is more or less conical with the anterior margin of the
vertex extending slightly beyond that of the eyes; greatest width of
the head almost equal to the pronotal humeral width and five to
six times the anterior width of the vertex; eyes holoptic for the
basal three eighths of the median length of the head; along the
median longitudinal axis the head is slightly more than one half
the length of the pronotum. Pronotum with its humeral width
almost twice its median length; lateral margins parallel and two
thirds the median length; posterior margin convex, medianly emar-
ginate. Scutellum greatly reduced, basal width only slightly more
than median length. Facial tubercle swollen. Labrum strongly
convex transversely; basal width equal to its median length; apex
rounded. Rostral prong (PI. LV, fig. 99) accuminate. Rostrum
with the second rostral segment produced into a flaplike process
that extends behind the third and fourth segments, not adjacent to
the fourth and the apical half of the third segments. Fore leg (PI.
LI, fig. 78a) with femur greatly expanded dorso-ventrally, with
dorsal margin convex; stridulatory comb (Pi. LI, fig. 78b) highly

440 The University Science Bulletin

irregular, apical fourth and fifth teeth flattened and capitate. Chae-
totaxy of the male front leg as shown on Plate LI. The relative
lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. .Seg.

Fore leg 100 114 71

Middle leg 100 85 40 33

Hind leg 100 77 31 28

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded with the vertex extending slightly be-
yond the anterior margin of the eyes, greatest width of head nine
tenths the pronotal humeral width and slightly more than five
times the anterior width of the vertex; eyes holoptic for the basal
third of the median length of the head; along the median longi-
tudinal axis the head is three fourths the length of the pronotum.
Pronotum with humeral width slightly more than twice the median
length; lateral margins slightly diverging and almost as long as
the median length of the pronotum; posterior margin convex,
medianly emarginate. Second rostral segment produced into a
short spur on its apical posterior margin. The relative lengths of
the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 57 36

Middle leg 100 86 40 27

Hind leg 100 81 34 34

Location of types. — The type material from Celebes in the Bred-
din Collection of the Hamburg Museum. One female co-type,
Celebes, Lura See bei Lura, VIII-95, Dr. Sarasin, in the Kirkaldy
Collection of the Snow Entomological Collection.

Comparative notes. — Very closely related to Anisops kempi n. sp.
from which it can best be separated on the basis of the males.
The males of A. breddeni Kirkaldy have the lateral margins of
the pronotum almost parallel while in the former species they are
distinctly diverging. Also the scutellum is much smaller in the
case of A. breddeni, with its basal width only slightly more than
its median length whereas in A. kempi the basal width of the pro-
notum is one and one third the median length. The outline of the
head of A. breddeni is more or less conical whereas that of A. kempi
is rounded.

Data on distribution:

India

Puenjikara Island, Cochin Backwater, nr. Ernakulane, X-14, F. H. Gravely,
one male (F. H. Snow Coll.); one male (Indian Museum).

Brooks: The Genus ANisors 441

Barkuda, Chilka Lake, Ganjam Dist. Madras Pres. VIII-19, F. H. Gravely
( F. H. Snow Coll. )

Calcutta, Lakes, Sta. 4, three females (Indian Museum).

Burma
Bangoon, VIII-21-15, Hodgard, three males (Indian Museum); one female
(F. H. Snow Coll.)

Ceylon
Mt. Lavinia, Mar. 9, '96, Madarasz, one male, one female ( F. H. Snow Coll. )

Anisops kempi n. sp.

(PI. LI, fig. 75)

Size. — Males, length, 5.7 mm. -6.6 mm., greatest body width 1.3
mm.-1.6 mm.; females, length 5.4 mm.-6.0 mm., greatest body width
1.4 mm. -1.6 mm.

Shape. — Slender, slightly fusiform species, with greatest width
about midway its length.

Color. — General facies stramineous. Eyes brown. Legs stra-
mineous. Abdominal venter dark brown with keel and segmental
margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest head width nine tenths the pro-
notal humeral width and five to seven times the anterior width of
the vertex; eyes holoptic in basal third of head; along median longi-
tudinal axis head approximately two thirds the pronotal length.
Pronotum with its humeral width about one and one half its median
length; lateral margins diverging and two thirds median length;
posterior margin convex, medianly emarginate. Scutellum with
basal width one and one third its median length. Facial tubercle
slightly raised. Labrum short; basal width only slightly more than
median length; apex rounded. Rostral prong (Pi. LI, fig. 75b) ac-
cumulate at apex. Second rostral segment extended into flaplike
process behind third and fourth rostral segments, not adjacent to
the fourth or apical half of the third rostral segments. Fore leg
(PI. LI, fig. 75a) with femur greatly expanded dorsoventrally;
stridulatory comb highly irregular, with four prominent teeth, the
middle two flattened and capitate (PI. LI, fig. 75c). Chaetotaxy
as shown on Plate LI. The relative lengths of the parts of the
legs are as follows: . . „ ,

° 1st 2nd

Femur Tibia Tar. Seg. Tar. >Seg.

Fore leg 100 125 69

Middle leg 100 82 41 23

Hind leg 100 82 32 26

442 The University Science Bulletin

Female structural characteristics. — Viewed from above the out-
line of the head is rounded; greatest head width nine tenths pro-
notal length, five times the anterior width of the vertex; eyes hol-
optic in basal third of the head; along the median longitudinal axis
the head is approximately two thirds the pronotal length. Pronotum
with humeral width slightly more than twice its median length;
lateral margins diverging and almost one half the median length;
posterior margin convex, medianly emarginate. The relative
lengths of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 133 50 33

Middle leg 100 86 38 25

Hind leg 100 86 34 30

Location of types. — Male holotype, female allotype, Assam, Tin-
sukia IV-9-44, D. E. Hardy in the Snow Entomological Collection.
The paratypes are as follows: In the Snow Entomological Col-
lection: four males, two females, Drappa nr. Calcutta, India,
1-22-11, Kemp. In the Indian Museum at Calcutta: One male, Cal-
cutta, at light, VI-7-11, F. H. G.; one male, four females, Calcutta,
XII-2-07; two males, one female, Calcutta, Eden gardens, at light,
X-14-11, F. H. Gravely; one female, Garia nr. Calcutta, XII-13-10;
one female, Lower Burma, Amherst Dist, Kawkareih to third Camp,
XI-21-11 to XII-1-11, F. H. Gravely. In the United States National
Museum: one male, Siam, Bangkok, Sept. 17, 1926, H. N. Smith.

Comparative notes. — This species appears very similar to Anisops
breddeni Kirkaldy. However, the pronotal lateral margins of A.
kempi diverge whereas in A. breddeni they are almost parallel. Also
the head of the latter is more or less conical with the vertex extend-
ing beyond the anterior margins of the eyes, while in A. kempi the
outline of the head is rounded. Also the latter species has a much
larger scutellum which has a basal width one and one third its me-
dian length, whereas in A. breddeni the scutellum is small with its
basal width only slightly more than its median length.

Data on distribution:
India

Orissa, Satpara, IX-10-13, N. A., one male (F. H. Snow Coll.) United Prov.
Cannapore Dist. X-l to 8-1911, J. W. C, two males, four females (F. H. Snow
Coll.).

Brooks: The Genus Anisops 443

Anisops crinita n. sp.

(PI. L. fig. 68)

Size. — Males, length 4.9 mm.-5.4 mm., greatest body width 1.3
mm. -1.5 mm; females, length 4.8 mm.-5.7 mm., greatest body width
1.3 mm. -1.6 mm.

Shape. — Small, fusiform species; greatest body width about mid-
way the body length.

Color. — General facies stramineous or testaceous. Eyes brown
or gray. Hemelytra may have hyaline areas which appear darker
due to the underlying dark brown dorsal body surface. Legs stra-
mineous. Abdominal venter dark brown or black with keel and
segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width nine tenths the pronotal
humeral width, five times the anterior width of the vertex; synthlipsis
narrow, between one sixth and one seventh the anterior width of the
vertex; along the median longitudinal axis the head is slightly more
than three fifths the pronotal length. Pronotum with its humeral
width approximately twice the median length; lateral margins di-
verging and at least one half the median length; posterior margin
almost straight, with only slight median emargination. Frons
slightly depressed, with facial tubercle flat and covered with small
procumbent hairs. Labrum short, covered with long hairs; basal
width one and one half its median length; apex broad and rounded,
almost truncate. Rostral prong (Pi. L, fig. 68c) shorter than third
rostral segment; apex truncate. Stridulatory comb ( PI. L, fig. 68b )
of approximately six to nine even-length teeth. Chaetotaxy of the
male front leg as shown on Plate L. The relative lengths of the
parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. i>eg.

Fore leg 100 125 75

Middle leg 100 SO 40 23

Hind leg 100 79 30 30

Female structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width nine tenths the pronotal
humeral width; four to six times the anterior width of the vertex;
synthlipsis variable, from one fifth to one sixth the anterior width
of the vertex; along the median longitudinal axis the head is three
fifths the pronotal length. Pronotum with humeral width slightly
more than twice its median length; lateral margins diverging and
one half the median length; posterior margin convex, medianly

444 The University Science Bulletin

emarginate. The relative lengths of the parts of the legs are as
follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 125 47 29

Middle leg 100 81 38 20

Hind leg 100 80 28 30

Location of types. — Male holotype, female allotype, one male
and one female paratypes, Corfu, J. Sahib, in the Kirkaldy Col-
lection of the Snow Entomological Collection. Other paratypes
are as follows: In the Snow Entomological Collection: twelve
males, five females, Daulatabad, Seistan Village, pond, XII-20-18,
Sta. 29, B. S.; one male, New Caledonia, Noumea, A. Faval. In
the Indian Museum at Calcutta: thirteen males, five females, Dau-
latabad, Seistan Village, pond, XII-20-18, Sta. 29, B. S.; three males,
five females, Baluchistan, Zangi Nawar, 20 mi. w. of Nushki, XII-25
to 29-18, Sta. 35, B. S., N. A. & S. K.; two males, three females,
Ganjam Dist. Barkuda, Chilka Lake, Madras Pres. VIII-18, F. H.
Gravely; one male, Baluchistan, Quetta, 5700", pond in res. garden,
Sta. 3, XI-10, BS; one male, one female, Berhampur, Murshidabad,
Bengal, Sept. '12, Southwell.

Comparative notes. — About the same size and general appearance
as A. nivea ( Fabricius ) . However, the males of A. crinita lack the
excavate facial tubercle and peculiarly twisted tarsal claws as found
on the males of A. nivea. Also the labral hairs of A. crinita are not
organized into definite tufts as those of A. nivea.

Data on distribution. — Known only from type series.

Anisops majiensis n. six

(P. LII, fig. 83)

Size. — Males, length 6 mm., greatest body width 1.5 mm.; females,
length 6.3 mm., greatest body width 1.6 mm.

Shape. — Fusiform species; greatest width about midway the body
length.

Color. — General facies stramineous. Abdominal venter dark
brown with keel and segmental margins of the connexivum stra-
mineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width slightly less than the pro-
notal humeral width and five times the anterior width of the ver-
tex; synthlipsis extremely narrow, one tenth the anterior width of
the vertex; along the median longitudinal axis the head is eight
tenths the pronotal length. Pronotum with its humeral width

Brooks: The Genus Anisops 445

twice the median length; lateral margins diverging and slightly
more than three fifths the median length; posterior margin convex,
medianly emarginate. Facial tubercle slightly raised. Labrum
short and broad; basal width twice the median length; apex more
or less accumulate. Rostral prong (PI. LII, fig. 83b) longer than
the third rostral segment; apex accumulate. Fore leg (PL LII,
fig. 83a) with anterior femur enlarged at apex and rounded; stridu-
latory comb (PL LII, fig. 83c) of approximately twenty-four teeth,
with basal fifteen almost twice the length of the apical nine. Chae-
totaxy of the male front leg as shown on Plate LII. The relative
lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 110 55

Middle leg 100 82 36 23

Hind leg 100 82 32 32

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head only slightly
less than the pronotal humeral width and five to six times the an-
terior width of the vertex; synthlipsis wide, one third the anterior
width of the vertex; along the median longitudinal axis the head
is almost as long as the pronotum. Pronotum with its humeral
width one and one half the median length; lateral margins di-
verging and slightly more than one half the median length; posterior
margin convex, medianly emarginate. The relative lengths of the
parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 121 71 54

Middle leg 100 91 46 27

Hind leg 100 81 33 33

Location of types. — Male holotype, female allotype, Afrique,
Orient, Angl. Maji-Chumvi, (Wa-Nyika) 1904, Ch. Alluaud, in the
Paris Museum.

Comparative notes. — The chaetotaxy of the male front legs is
strongly suggestive of a relationship with A. debilis Gerst. How-
ever, the latter species does not have the extremely narrow (one
tenth the anterior width of the vertex) synthlipsis, nor the apically
enlarged and rounded front femur as found on the males of this
species.

Data on distribution. — Known only from type series.

446 The University Science Bulletin

Anisops debilis Gerstaeker

(PI. Lll, fig. 80)

J 873. Anisops debilis Gerstaeker, Decken's Reise in Ost Africa III, pt. 2, p. 425.
1901. Anisops debilis, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 119, 132.
1926. Anisops debilis, Jaczewski, Annal. Zool. Musei Polonici Hist. Nat. vol. V., no. 2,
pp. 86-88, text fig. 38, 39, 40, 41.

1929. Anisops debilis, Hutchinson, Ann. South African Mus., vol. XXV, pt. 3, pp. 391-
392, PI. XXX, fig. 15.

1930. Anisops debilis, Hutchinson, Ann. Mag. Nat. Hist., Ser. 10, vol. VI, p. 58 (ecological
note).

1933. Anisops debilis, Hutchinson, Inter, ges. Hydrob. Hydrog., vol. 28, p. 45, pt. 5/6,
p. 461.

1935. Anisops debilis, Poisson, Mus. Nat. Hist. Nat., vol. Ill, pp. 211-213, text figs.
26, 27.

1937. Anisops debilis, Poisson, Ann. Soc. Ent. France, vol. CVI, pp. 117-118, text fig. 3.

1939. A?iisops debilis, Poisson, Bull. Soc. Ent. France, vol. 44, p. 43 (ecological note).

Size. — Males, length 5.8 mm. -6.3 mm., greatest body width 1.5
mm.-1.7 mm.; females, length 6.3 mm., greatest body width 1.5 mm.

Shape. — Slender, fusiform species; greatest body width about
midway the body length.

Color. — General facies gray. Eyes brown. Eyes and pronotum
testaceous. Scutellum testaceous with two triangular hyaline areas
on each side of the anterior half. Hemelytra hyaline and appear
dark gray due to the underlying black dorsal body surface. Legs
stramineous. Abdominal venter dark brown or black with keel and
segmental margins of the connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of head nine tenths the pro-
notal humeral width and six times the anterior width of the vertex;
synthlipsis narrow, between one fourth and one fifth the anterior
width of the vertex; along median longitudinal axis the head is more
than one half the length of the pronotum. Pronotum with its
humeral width twice the median length; lateral margins diverging
and three fifths the median length; posterior margin convex, medi-
anly emarginate. Facial tubercle slightly raised. Labrum with its
basal width one and one third the median length; apex rounded.
Rostral prong (PL LII, fig. 80b) longer than the third rostral seg-
ment; apex accuminate. Stridulatory comb (PI. LII, fig. 80c) of
approximately twenty-one teeth which decrease slightly in length
from base to apex. The chaetotaxy of the male front leg as shown
on Plate LII. The relative lengths of the parts of the legs are as
follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 114 71

Middle leg 100 86 43 35

Hind leg 100 80 31 31

Brooks: The Genus Anisops 447

Female structural characteristics. — Viewed from above the out-
line of the head is rounded; greatest width of the head nine tenths
the pronotal humeral width and five times the anterior width of the
vertex; synthlipsis narrow, less than one third the anterior width of
the vertex; along the median longitudinal axis the head is slightly
more than one half the length of the pronotum. Pronotum with its
humeral width twice the median length; lateral margins diverging
and almost one half the median length; posterior margin convex,
medianly emarginate. The relative lengths of the parts of the legs
are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 115 54 31

Middle leg 100 82 41 32

Hind leg 100 80 30 30

Location of types. — Type material from East Africa in the Berlin
Museum.

Comparative notes. — The narrow synthlipsis and shape of the
rostral prong make this species similar to A. vitrea Signoret. How-
ever, this species is a larger one than A. vitrea and the males lack the
flattened facial tubercle as found on the A. vitrea.

Data on distribution:
Africa

French Sudan, Bamako, March 1929, A. Bang Baas, purchased fr. Dr. O.
Standinger, two males, five females (F. H. Snow Coll.).

Uganda, Katona, IX-1913, Muyenju, exchange fr. Horvath, one male, one
female (F. H. Snow Coll.).

Messina, Limpopo, V-25-'92, gift to H. B. H. fr. G. E. Hutchinson, two fe-
males (F. H. Snow Coll.).

Ogaden, Quabi-Chebeli, Imi Mission, 1902, Du Bourg de Bozas, two males,
one female ( F. H. Snow Coll. ) ; six females, one male ( Paris Museum ) Soudan,
Mioro, Oct. 1909, E. De Zeltner, five males, one female (Paris Museum).

Env. de Zinder, Guidimauni (mission Tilho) 1910, Dr. R. Gaillard three
males ( Paris Museum ) .

Central Afr. Uganda, Entebbe, H. Rolle, one male, three females (Bueno
Coll. of F. H. Snow Coll.).

Anisops exigera Horvath

(PI. LI, fig. 77)

1919. Anisops exigera Horvath, Abhand. Senckenberg. naturf. Gesell., vol. XXXV, p. 314.
1933. Anisops exigera, Lundblad, Arch, fur Hydrob. Suppl., vol. XII, p. 145 (a list of
Indo-australian and Pacific forms of this genus).

Size. — Males, length 4.3 mm. -4.5 mm., greatest body width 1.2

mm.; females, length 4.3 mm. -4.5 mm., greatest body width 1.2 mm.

Shape. — Small, subfusiform species with lateral margins in an-

448 The University Science Bulletin

terior half of body almost parallel, lateral margins of posterior
half converging.

Color. — General facies pearlaceous. Legs stramineous. Abdom-
inal venter dark brown with keel and segmental margins of the
connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the greatest width of the head eight
tenths the pronotal humeral width and six times the anterior width
of the vertex; synthlipsis narrow, from one fifth to one sixth the
anterior width of the vertex; along the median longitudinal axis
the head is almost as long as the pronotum. Pronotum with its
humeral width slightly more than twice its median length; lateral
margins diverging and one half as long as the median length; pos-
terior margin convex, medianly emarginate. Facial tubercle slightly
raised, Labrum with its basal width one and one fourth times its
median length; apex truncate, with its length almost one half the
basal width. Rostral prong (PI. LI, fig. 77b) shorter than the
third rostral segment; apex rounded. Stridulatory comb (PI. LI,
fig. 77c) of approximately ten distinct teeth which increase in
length toward the middle. Chaetotaxy of the front leg as shown
on Plate LI. The relative lengths of the parts of the legs are as
follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 114 77

Middle leg 100 78 33 22

Hind leg 100 78 30 30

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of the head nine tenths
the pronotal humeral width and six times the anterior width of
the vertex; synthlipsis narrow, one fifth the anterior width of the
vertex; along the median longitudinal axis the head is two thirds
the length of the pronotum. Pronotum with its humeral width
slightly more than twice its median length; lateral margins di-
verging and slightly more than one-half the median length; posterior
margin convex, slightly convex medianly. The relative lengths of
the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 124 44 32

Middle leg 100 77 35 22

Hind leg 100 77 31 31

Location of types. — Type material from New Guinea, Simbang
Huon Gulf, 1898, Biro, in the Budapest Museum in Hungary.

Brooks: The Genus Anisops 449

Comparative notes. — This species is about the same size and gen-
eral facies as Anisops n'wea Fabr. from which it can be readily sepa-
rated at once on the basis of the males which lack the excavate
frons as found on the males of A. n'wea. Both sexes of A. exigera
possess a much narrower synthlipsis, only one fifth to one sixth the
anterior width of the vertex, where as the synthlipsis of A. nivea
is much wider, being approximately one third the anterior width
of the vertex.

Data on distribution:

New Guinea

Simbang Huon Golf, 1898, Biro (determined by Horvath), one male (F. H.
Snow Coll. ).

India

Puenyikara Isl., Cochin Backwater nr. Ernakulane, X-14, F. H. Gravely, nine
males, fourteen females (F. H. Snow Coll.).

Central Province, Varcam Tank nr. Pachmachi, Satpura Hills, 3000', Sta.
16, C. P. and F. H. G., thirteen males, nine females (Indian Museum).

Morihan, XI-20-15, Rescee, five females (Indian Museum).

Anisops vitrea Signoret

(PI. LI, fig. 76)

1860. Anisops vitrea Signoret, Ann. Soc. Ent. France, vol. VIII, no. 3, p. 792.

1895. Anisops vitrea, De Carlini, Ann. Mus. Civ. Stor. Nat. Genova, Series 2a, vol. XV
(XXXV), p. 19 (ecological note).

1899. Anisops vitrea, Kirkaldy, Ann. Soc. Ent. France, vol. LXVIII, pp. 160-107 (=per-
sephone Kirkaldy).

1904. Anisops vitrea, Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 119. 132.

1926. Anisops vitrea, Horvath, Arch, fur Zool., vol. ISA, no. 31, p. 2 (determined in error;
actually ^4. jaczewski Hutchinson).

1926. Anisops vitrea, Jaczewski, Ann. Zool. Musei Polonici Hist. Nat., vol. V, p. 86 text
figs. 42, 43, 44 (determined in error; actually A. jaczewski Hutchinson).

1928. A?iisops vitrea, Hutchinson, Ann. Mag. Nat. Hist., Ser. 10. vol. I, pp. 303-304,
text figs. 1, a, b (cites the error of Horvath and Jaczewski and names the species observed by
them A. jaczewski).

Referring to this species also

1898. Anisops persephone Kirkaldy, Wiener Ent. Zeit., vol. XVIII, p. 142.
1913. Anisops aldabrana, Distant, Trans. Linn. Soc. London, vol. 16, p. 1S9, PI. XIII.
fig. 30.

1927. Anisops aldabrana, Hutchinson, Ann. Mag. Nat. Hist., Ser. 9, vol. 19, p. 376, text
figs. 1, a b.

Size. — Males, length 4.5 mm. -5.1 mm., greatest body width 1.2
mm. -1.3 mm.; females, length 4.5 mm. -5.1 mm., greatest body width
1.2 mm. -1.3 mm.

Shape. — Small, subfusiform species; greatest body width about
one third the length of the body.

Color. — General facies dark gray. Eyes brown. Vertex and pro-
notum stramineous, the latter may have a median longitudinal band

29—3286

450 The University Science Bulletin

of orange. Scutellum testaceous or orange with anterior margin
dark brown. Hemelytra hyaline and appear dark gray due to the
dark color of the underlying body surface. Legs stramineous. Ab-
dominal venter dark brown with keel and segmental margins of the
connexivum stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the greatest width of the head eight
tenths the pronotol humeral width and six to seven times the anterior
width of the vertex; synthlipsis narrow, from one sixth to one eighth
the anterior width of the vertex; along the median longitudinal axis
the head is almost as long as the pronotum. Pronotum with its
humeral width slightly more than twice the median length; lateral
margins diverging and one half the median length; posterior margin
convex, medianly emarginate. Facial tubercle slightly raised and
flattened on ventral surface; flattened area surrounded by more or
less triangular, faint ridge. Labrum short and with a few procum-
bent hairs; basal width almost twice the median length. Rostral
prong ( PI. LI, fig. 76b ) longer than the third rostral segment; apex
accuminate. Stridulatory comb (PL LI, fig. 76c) of approximately
twenty teeth, teeth slightly shorter at apex than at base. Chaetotaxy
of the male front leg as shown on Plate LI. The relative lengths of
the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 80

Middle leg 100 85 41 27

Hind leg 100 75 33 33

Female structural characteristics. — Viewed from above the out-
line of the head is rounded; greatest width of the head almost nine
tenths the pronotal humeral width and seven times the anterior
width of the vertex; synthlipsis narrow, about one fourth the anterior
width of the vertex; along the median longitudinal axis the head is
two thirds the length of the pronotum. Pronotum with its humeral
width slightly more than twice the median length; lateral margins
diverging and one half the median length; posterior margin convex,
medianly emarginate. The relative lengths of the parts of the legs
are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 120 54 32

Middle leg 100 85 41 27

Hind leg 100 76 33 33

Location of types. — Type material in Stockholm Museum.

Brooks: The Genus Anisops 451

Comparative notes. — Though not quite as robust as Anisops
edepol Kirk., this species is of approximately the same length. How-
ever, the two are by no means similar. They can be readily dis-
tinguished by the fact that the latter species has the posterior margin
of the pronotum almost straight instead of convex and medianly
emarginate as in the case of A. vitrea. Also the males of A. vitrea
lack the short spur on the facial tubercle as found on the males of
A. edepol.

Remarks. — I was fortunate in having at my disposal some of the
specimens of Anisops aldabrana Distant collected by J. F. Fryer at
the same time as the type. Though a trifle smaller than A. vitrea,
this species is identical in every other respect and therefore is being
placed as a synonym of A. vitrea.

Data on distribution:
Madagascar

Tananarive, purchased from Prof. C. Lamberton, thirty-six males, thirty-
four females ( F. H. Snow Coll. ) .

Moroantsetra, March, 1929, A. Bang Baas, purchased from Dr. O. Stand-
linger, nineteen males, twenty-nine females (F. H. Snow Coll.).

Madagascar, 1897, F. de Zeltner, one female (Paris Museum).

Reg. du Sud-est, Vallee du Fanjahira Isaka, 1901, Ch. Alluaud, one male
(Paris Museum).

Boeni, Maevatanano, Oct.-Nov. 1899, Dr. J. DeCorse, two females (Paris
Museum ) .

Madagascar, two males, ( Bueno Coll. of F. H. Snow Coll.).

Madagascar, 1897, E. Dorr, four females (Paris Museum).

Prov. D'Ankavandra, 1-98, S. Hure, one male, one female (Paris Museum).

Marais du Finerena, 1905, G. Geay, three males, three females (Paris
Museum).

Antanimora, 1901, Ch. Alluaud, two males, four females (Paris Museum).

Annanariva, Sikora, 1896, Noualhier, three males, one female (Paris Mu-
seum ) .

Diego-Suarez, five males (U. S. Nat. Mus. ).

Diego-Suarez, 1893, Ch. Alluaud, five males, eleven females (U. S. Nat.
Mus. ) ; one female ( Paris Museum ) .

Aldabra Island
Aldabra, IX-'08, J. C. F. Fryer, one male, three females ( F. H. Snow Coll. ) .

Mauritius
Roduit, 111-25-1948, R. Manet, three males, one female (F. H. Snow Coll.).

La Reunion

Plides Palmistes, 1897, Ch. Alluaud, one male, (Bueno Coll. of F. H. Snow
Coll.).

452

The University Science Bulletin

Anisops hypatia Hutchinson

(PI. LII, fig. 82)

1929. Anisops hypatia Hutchinson, Ann. South African Mus. vol. XXV, pt. 3, pp. 399-
400, PI. XXX, fig. 16; PI. XXXII, figs. 2, 2a.

Size. — Males, length 5.4 mm., greatest body width 1.5 mm.; fe-
males, length 5.7 mm., greatest body width 1.7 mm.

Shape. — Small, fusiform species; greatest body width midway the
body length.

Color. — General facies stramineous. Eyes brown. Abdominal
venter dark brown with keel and segmental margins stramineous.

Male structural characteristics. — Viewed from above the outline
of the head is rounded with the anterior margin almost straight;
greatest width of the head equal to the pronotal humeral width
and five times the anterior width of the vertex; synthlipsis narrow,
slightly more than one fourth the anterior width of the vertex; along
the median longitudinal axis the head is slightly shorter than the
length of the pronotum. Pronotum with its humeral width twice
its median length; lateral margins diverging and one half the median
length; posterior margin convex, medianly emarginate. Facial
tubercle raised. Labrum with basal width almost one and one-half
the median length; apex rounded. Rostral prong (PI. LII, fig. 82b)
as long as the third rostral segment; apex accumulate. Stridulatory
comb (PI. LII, fig. 82c) of approximately fourteen teeth which in-
crease in length from base to apex. Chaetotaxy of the male front
leg as shown on Plate LII. The relative lengths of the parts of the
legs are as follows:

1st 2nd

Femur Tilna Tar. Seg. Tar. Seg.

Fore leg 100 124 92

Middle leg 100 80 39 24

Hind leg 100 81 36 38

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head slightly less
than the pronotal humeral width and five times the anterior width
of the vertex; synthlipsis wide, one third the anterior width of the
vertex; along the median longitudinal axis the head is almost equal
to the length of the pronotum. Pronotum with its humeral width
twice its median length; lateral margins diverging and one half the
median length; posterior margin convex, medianly emarginate. The
relative lengths of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 118 55 35

Middle leg 100 94 42 32

Hind leg 100 85 33 34

Brooks: The Gents Anisops 453

Location of types. — Type material from Cape Province, Africa
in the South African Museum.

Comparative notes. — This species is of the same general appear-
ance as A. amaryllis Hutchinson, however the head of the males is
much wider, being equal to the pronotal humeral width instead of
only nine tenths the pronotal humeral width as in A. amaryllis. Also
the latter species lacks the short basal row of three small spines as
found on the inner surface of the fore tarsi of A. hypatia.

Data on distribution:
Africa

Concordia, pond nr. Knysna, XII-23-1926, Gift to H. B. H. fr. G. E.
Hutchinson, one male, one female (F. H. Snow Coll. ).

Anisops balcis Hutchinson

(PL LII, fig. 84; PI. LV. fig. 97)

1930. Anisops balcis Hutchinson, Proc. Zool. Soe. London, vol. XXIX, pit. 2, pp. 447-
448, text fig. 4, a, b, c.

1930. Anisops balcis, Hutchinson, Ann. Mag. Nat. Hist. vol. VI, Series 10, p. 59 (eco-
logical note).

1932. Anisops balcis. Hutchinson, Ann. Mag. Nat. Hist. vol. IX. Ser. 10, p. 324 (eco-
logical note).

1933. Anisops balcis, Hutchinson, Internal, ges. Hydrob. und Hydros., vol. 28, prt 5/6,
p. 462 (ecological note).

1933. Anisops balcis. Jaczwski, Linn. Society Jour. Zool., vol. XXXVIII, p. 345 (ecological
note).

Size. — Males, length 6.8 mm., greatest body width 1.8 mm.; fe-
males, length 6.6 mm., greatest body width 1.8 mm.

Shape. — Slender and slightly fusiform species; greatest body
width slightly before midway the body length.

Color. — General facies testaceous. Abdominal venter dark brown
with keel and segmental margins of the connexivum testaceous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded; greatest width of the head eight tenths the
pronotal humeral width and seven times the anterior width of the
vertex; synthlipsis narrow, one fourth the anterior width of the ver-
tex; along the median longitudinal axis the head is three fourths the
length of the pronotum. Pronotum with its humeral width almost
twice the median length; lateral margins almost parallel, only
slightly diverging and two thirds the median length; posterior mar-
gin convex, medianly emarginate. Facial tubercle slightly raised
and triangularly flattened. Labrum short and broad; basal width
one and one half its median length; apex rounded. Rostral prong
( PI. LV, fig. 97 ) longer than the third rostral segment; apex accumi-
nate. Anterior femur enlarged dorso-ventrally in apical half; apex
truncate. Stridulatory comb (Pi. LII, fig. 84b) of about twenty-two

454 The University Science Bulletin

teeth, increasing in length at the middle. Chaetotaxy of the front
leg as shown on Plate LI I. The relative lengths of the parts of the
legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 112 75

Middle leg 100 85 39 26

Hind leg 100 78 32 32

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head eight tenths
the pronotal humeral width and five and one half times the anterior
width of the vertex; synthlipsis narrow, slightly more than one
fourth the anterior width of the vertex; along the median longi-
tudinal axis the head is almost two thirds the length of the pronotum.
Pronotum with its humeral width slightly more than twice the me-
dian length; lateral margins diverging and three fifths the median
length; posterior margin convex, medianly emarginate. The rela-
tive lengths of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 114 57 33

Middle leg 100 84 32 27

Hind leg 100 78 32 32

Location of types. — Type material from Lake Zwai, c. 5500',
XI-4-1926, J. Omer-Cooper, in the British Museum.

Comparative notes. — This species is similar in body shape to A.
gracilis Hutchinson from which it can be distinguished by the fact
that the males lack the transversely striated ridge as found on the
latter species. The males of A. gracilis do not possess the flattened
facial tubercle found on the males of this species.

Data on distribution:
Africa

Uganda, Hippo Pt. Lake Victoria, Feb. 29, 1948, F. X. Williams, two males
(Harv. Mus. Comp. Zool. ).

Uganda, Lake Baringo, 1931, E. W. Worthington, one male ( F. H. Snow
Coll.).

Abyssinia, Hora Abjata, small pond in Marsh, XI-18-1926, J. Omer-Cooper,
one male, one female, exchange from British Museum (F. H. Snow Coll.).

Anisops biroi n. sp.

(PI. XLV, fig. 49)

Size. — Males, length 4.2 mm.-4.5 mm., greatest body width 1.2
mm. -1.3 mm.; females, length 4.3 mm., greatest body width 1.2 mm.

Shape. — Small, fusiform species, with greatest width about two
fifths the body length.

Brooks: The Genus Anisops 455

Color. — General fades stramineous. Eyes brown. Pronotum
may be hyaline and appear the brown color of the underlying scutel-
lum. Legs stramineous. Abdominal venter dark brown with keel
and segmental margins of the connexivum stramineous.

Male structural characteristics. — As viewed from above, the out-
line of the head is rounded with its greatest width nine tenths the
pronotal humeral width and approximately nine times the anterior
width of the vertex; synthlipsis narrow, approximately one fifth
the anterior width of the vertex; along the median longitudinal axis
the length of the head is equal to or slightly less than that of the
pronotum. Pronotum with its humeral width at least twice its me-
dian length; lateral margins diverging and slightly less than one half
the median length; posterior margin convex, medianly emarginate.
Facial tubercle simple, neither raised nor excavate Rostral prong
(PI. XLV, fig. 49b) at least as long as third rostral segment; apex
accumulate. Labrum short and spinose; basal width slightly more
than median length; apex accuminate. Stridulatory comb (Pi.
XLV, fig. 49c) of approximately twenty-two teeth. Chaetotaxy of
the male front legs as shown on Plate XLV. The relative lengths
of the parts of the legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 125 81

Middle leg 100 83 36 26

Hind leg 100 83 30 33

Female structural characteristics. — As viewed from above, the out-
line of the head is rounded; greatest width of the head nine tenths
the pronotal humeral width and eight times the anterior width of
the vertex; synthlipsis wide, one third the anterior width of the
vertex; along the median longitudinal axis the head is seven tenths
the length of the pronotum. Pronotum with its humeral width
two and one half times the median length; lateral margins diverging
and one half the median length; posterior margin convex, medianly
emarginate. The relative lengths of the parts of the legs are as fol-
lows:

1st 2nd

Femur Tibia Tar. Seg, Tar. Seg.

Fore leg 100 120 54 34

Middle leg 100 78 34 28

Hind leg 100 83 31 33

Location of types. — Male holotype, female allotype, two male
and one female paratypes, New Guinea, Seleo Berlinhafen, '96, Biro
in the Kirkaldy Collection of the Snow Entomological Collection.

Comparative notes. — Very similar to A. rigoensis n. sp. from which

456 The University Science Bulletin

it differs in having its rostral prongs at least as long as the third
rostral segment whereas in A. rigoensis the rostral prongs are shorter
than the third rostral segment. Also A. biroi lacks the short row of
three small setae on the inner surface of the male anterior tarsi as
found on the males of A. rigoensis.

Data on distribution. — Known only from type series.

Anisops rigoensis n. sp.

(PI. LII, fig. 81)

Size. — Males, length 4.8 mm. -5.1 mm., greatest body width
1.3 mm.; females, length, 5.4 mm. -6.0 mm., greatest body width
1.3 mm. -1.5 mm.

Shape. — Small, fusiform species, with greatest body width about
four tenths its body length.

Color. — General facies gray. Eyes gray or dark brown. Vertex,
pronotum, and scutellum testaceous. The latter may be black and
may have its hemelytral margins tinged with crimson. Hemelytra
hyaline and appear dark gray, gray, or testaceous depending on the
color of the underlying body surface. Legs stramineous. Abdom-
inal venter black with keel and segmental margins of the connexivum
stramineous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded, with its greatest width almost nine tenths
the pronotal humeral width and eight to nine times the anterior
width of the vertex; synthlipsis variable, from one fifth to one third
the anterior width of the vertex; along the median longitudinal
axis the head is equal to or longer than twice its median length;
lateral margins diverging and slightly more than twice its median
length; posterior margin convex, medianly emarginate. Facial
tubercle only slightly raised. Labrum with its basal width slightly
more than its median length. Rostral prong (PI. LII, fig. 81b)
slightly longer than the third rostral segment; apex accumulate.
Stridulatory comb ( PI. LII, fig. 81c ) of approximately twenty teeth;
basal eight much shorter than others. Chaetotaxy of the front leg
as shown on Plate LII. The relative lengths of the parts of the legs
are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 126 88

Middle leg 100 70 32 22

Hind leg 100 85 31 31

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head nine tenths the

Brooks: The Genus Anisops 457

pronotal humeral width and approximately seven times the anterior
width of the vertex; synthlipsis narrow, between one fifth and one
sixth the anterior width of the vertex; along the median longitudinal
axis the head is three fourths the pronotal length. Pronotum with
its humeral width slightly more than twice its median length; lateral
margins diverging and slightly more than one half the median
length; posterior margin concave, medianly emarginate. The rela-
tive lengths of the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 130 64 42

Middle leg 100 85 40 25

Hind leg 100 83 34 36

Location of types. — Male holotype, female allotype, three male
and six female paratypes, New Guinea, Rigo, Luglio, 1889, L. Loria
in the Snow Entomological Collection. Other paratypes are: one
male, three females, New Guinea, Lemian Berlinhafen, '96, Biro
and one female New Guinea, Friedrich-Wilh.-hafen, '96, Biro in
the Kirkaldy collection of the Snow Entomological Collection.

Comparative notes. — Very similar to Anisops birio n. sp. and
is separated best from this species on the character of the rostral
prong, which in A. biroi extends its dorsal margin almost to the
apex of the third rostral segment whereas in A. rigoensis the dorsal
margin of the prong extends scarcely beyond the base. For com-
parison of the chaetotaxies of the front legs of the males see PI. LII,
fig. 81a and PL XLV, fig. 49a.

Data on distribution. — Known only from type series.

Anisops waltairensis n. sp.

(PI. L, fig. 69)

Size. — Males, length 4.5 mm. -4.7 mm., greatest body width 1.2
mm. -1.4 mm.; females, length 4.8 mm. -5.1 mm., greatest body width
1.5 mm. -1.6 mm.

Shape. — Small, fusiform species; greatest body width about mid-
way the body length.

Color. — General facies pale, may be stramineous or pearlaceous.
Legs testaceous. Abdominal venter with keel and segmental mar-
gins of the connexivum stramineous or pearlaceous.

Male structural characteristics. — Viewed from above, the outline
of the head is rounded with the greatest width almost nine tenths
the pronotal humeral width and approximately eight times the an-
terior width of the vertex; synthlipsis narrow, about one eighth

458 The University Science Bulletin

the anterior width of the vertex; along the median longitudinal axis
the head is nine tenths the pronotal length. Pronotum with humeral
width two and one half times the median length; lateral margins
diverging and slightly more than one half the median length; pos-
terior margin convex, slightly concave medianly. Facial tubercle
only slightly raised. Labrum long; median length equal to its
basal width. Rostral prong (PI. L, fig. 69b) slightly shorter than
the third rostral segment; apex more or less accumulate. Stridula-
tory comb (Pi. L, fig. 69c) of approximately eleven teeth which in-
crease in length toward the middle. Chaetotaxy of the male front
leg as shown on Plate L. The relative lengths of the parts of the
legs are as follows:

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 118 85

Middle leg 100 82 42 24

Hind leg 100 75 33 33

Female structural characteristics. — Viewed from above, the out-
line of the head is rounded; greatest width of head eight to nine
tenths the pronotal humeral width and almost eight times the an-
terior width of the vertex; synthlipsis narrow, one seventh the an-
terior width of the vertex; along the median longitudinal axis the
head is three fourths the pronotal length. Pronotum with humeral
width two and one half times the median length; lateral margins
diverging and slightly more than one half the median length; pos-
terior margins convex, medianly concave. The relative lengths of
the parts of the legs are as follows :

1st 2nd

Femur Tibia Tar. Seg. Tar. Seg.

Fore leg 100 116 57 32

Middle leg 100 80 40 20

Hind leg 100 85 37 30

Location of types. — Male holotype, female allotype, Waltair,
India, 1-27-21 in the Indian Museum. Paratypes are as follows.
In the Indian Museum: two males, Kalka, Base of Simla Hills, 2400",
VII-16-11, Mus. Collr.; one female, Kalka, base of Simla Hills, 2400",
in small pond of rain water, Annandale. In the Snow Entomological
Collection: one male, one female, Waltair, India, 1-27-21; one male
Kalka, Base of Simla Hills, 2400", VII-16-11, Mus. Collr.; one female
Kalka, Base of Simla Hills, 2400", in small pond of rain water,
Annandale.

Comparative notes. — About the same size and general appearance
as A. exigera Horvath, from which it differs in having a much nar-
rower synthlipsis, one eighth the anterior width of the vertex and

Brooks: The Genus Anisops 459

in having its rostral prong accumulate at the apex whereas in
A. exigera the synthlipsis, though narrow is about one sixth the an-
terior width of the vertex and the rostral prong is truncate at the
apex.

Data on distribution. — Known only from type series.

Anisops ali Distant

1911. Anisops ali Distant, The Entomologist, vol. 44, p. 107.

1933. Anisops ali, Lundblad, Arch, fiir Hydrob. Suppl., vol. XII, p. 146 (a list of Indo-
australian and Pacific members of this genus).

As no specimens of this species were available for study I have
chosen to present a copy of the original description.

"Ochraceous, the hemelytra more or less shaded with fuliginous;
eyes black; interocular space narrow, margins parallel, not or very
slightly narrowing at base, above with a distinct central longitudinal
impression, the margins of which are ridged, beneath narrow, paral-
lel, of the same breadth throughout; pronotum with a distinct waved
carinate line on the anterior area, commencing on the lateral margins
behind eyes and then roundly directed upward and united between
the inner posterior angles of the eyes; abdomen beneath fuscous.
This species is allied in general coloration to A. sardea, Herr.-Schaff.,
and A. fiebri, Kirk. From both of these, however, it is distinctly
separated by the narrow parallel interocular space, and by the dis-
tinct carinate waved line to the pronotum. Long. 5 to 6M millim."

Hab. Ceylon; Diyatalawa (E. E. Green).

Location of types. — Female type located in the British Museum.

Anisops alluaudi Poisson

(PI. LIU, fig. 85)
1945. Anisops alluaudi Poisson, Bull. Soe. Ent. France, vol. 50, pp. 92-93, text fig. 6.

This species was not present in the material at my disposal.
Therefore I shall present a translation of the original description and
copies of the drawings.

Anisops Alluaudi n. sp.

Vertex two times wider than the synthlipsis; head (eyes included)
two times longer than its posterior width. Pronotum 1.8 times wider
than long, its posterior margin concave. Scutellum 1.17 times longer
than pronotum.

Eyes grayish brown; pronotum and scutellum brownish black,
abdominal tergites, anterior and posterior black, the intermediate
yellowish, bordered with black; hemelytra hyaline, not trimmed with
red.

460 The University Science Bulletin

Male: Third rostral segment presenting the long lateral spurs.

Anterior leg: F-100, T-120, t-91, c-29.

The tibial comb composed of 10-12 spatulate teeth, regularly
distributed; the tibia carries on its internal and lower side a short
spine, there are three others, linearly distributed on the internal and
basal side of the tarsi.

Length 7 mm.

Distribution: two males (1), La Reunion; Plaine des palmistes
(Ch. Alluaud).

Location of types. — Type material destroyed during the war.

Anisops aphrodite Kirkaldy

1900. Anisops aphrodite Kirkaldy, Ann. Soc. Ent. Belgique, vol. XLIV, p. 435.
1905. Anisops aphrodite, Bergroth, and Shouteden. Ann. Soc. Ent. Belgique, vol. XLIX,
pp. 388-389 (cites Kirkaldy's omission of this species from his "Uber Notonectiden" in 1904).

Kirkaldy (30) omitted this species from his Uber Notonectiden.
Whether or not he considered it as a synonym of a previously de-
scribed species or whether it was just an oversight on his part
has been a question confronting all subsequent workers. The type
material from Kinchassa is in the Perth Museum. As yet the spe-
cies has not been redescribed and as the type material was unavail-
able to me I can present only a translation of the original descrip-
tion.

Anisops aphrodite. — Vertex approximately four times wider than
the synthlipsis. Scutellum % longer than the pronotum.

Male. Anterior tibia one-fifth longer than the femur and one
fourth longer than the tarsus, the latter approximately three times
as long as the claws which are shaped as fingers. Middle femur
three eighths longer than the tibia, the latter three-eighths longer
than the tarsus, first tarsal segment two-fifths longer than the sec-
ond, the latter two times as long as the falciform claws.

Female. Anterior tibia one-sixth longer than the tarsus, first tar-
sal segment one-third longer than the second, the latter one-third
longer than the claws. Middle tibia one-fourth longer than the tar-
sus, first tarsal segment one-third longer than the second, the latter
two times as long as the falciform claws.

Male, female length. 5!? mm., width 1% mm.

Kinchassa ( Waelbroeck, Type), Lemba (Gilmont) and Boma
( Tschoffen ) .

Hemelytra hyaline, transparent. Abdomen black above and be-
neath, genital segments flavus.

Brooks: The Genus Anisops 461

Anisops calcaratus Hale

1923. A/iisops calcaratus Hale, Rec. South Asutralian Mus. vol. II, no. 3, pp. 416-417,
text fig. 369.

1933. Anisops calcaratus, Lundblad, Arch, fiir Hydrob. Suppl., vol. XII, p. 145 (a list of
Indo-australian and Pacific members of the genus).

As no male specimens of this species were available for study I
have chosen to present a copy of the original description.

Male. Head, including eyes, narrower than the pronotum; noto-
cephalon with a median groove, which does not extend to hinder
margin of the head and with a swelling on each side; synthlipsis
less than 1.5 in vertex, 3.5 times in the width of an eye; notocephalic
swellings united at the front of the head and continued on to face
as a strong, median carina; eyes large, prominent, projecting slightly
in front of vertex. Pronotum sordid testaceous, about 1.5 times
wider than long, with a coarse, median carina reaching to posterior
margin; lateral margins divergent; hinder edge evenly convex.
Scutellum testaceous, wider than long, a little longer than the prono-
tum. Metanotum testaceous, with a black spot on each side; upper
side of abdomen testaceous, posterior segments black. Anterior
femora stout, the superior edge with a knife-like ridge, which grad-
ually rises until it attains a point beyond the middle of the length
of the thigh, where it abruptly terminates; summit of this ridge
set with very small prostrate spines; tibiae strong, the anterior end
of the inner face with a distinct spur, in the apex of which is set
a short, stout spine; tarsi damaged. Length, nearly 9 mm.; width
2.4 mm.

Female. Eyes not so prominent, and notocephalon wider than in
male. Synthlipsis 1.5 in vertex, 3 times in the width of an eye.
Pronotum about twice as long, with a coarse, median carina, hinder
margin convex. Scutellum pale yellow to orange; sometimes with
a black spot on each side anteriorly; 1.5 times as long as the prono-
tum. Anterior femora not ridged on superior edge; anterior tibia
one-fourth longer than the tarsi, the first tarsal segment nearly
twice as long as the second, which is less than twice the length of
the longest claw. Length, 8 mm. to 9.5 mm.; width 2.5 mm. to 3 mm.

Hob. South Australia; Bordertown (J. G. O. Tepper, type lo-
cality); Queensland: Cunnamulla (H. Hardcastle).

The type which is somewhat damaged, is the only male of this
distinct species as yet received. The convex posterior margin of the
pronotum distinguishes it from all other Australian forms, excepting
possibly A. endymion Kirk.; in the description of the last named
species the character of the hinder edge of the pronotum is not

462 The University Science Bulletin

stated but the synthlipsis is described as about half of the width
of an eye and but slightly narrower than the vertex, while the
pronotum is relatively longer and the scutellum shorter than in the
female of A. calcaratus.

Location of types. — Male type located in the South Australian
Museum.

Anisops canariensis, ccmariensis Noualhier

1893. Anisops canariensis Noualhier, Ann. Soc. Ent. France, p. 18.
1899. Anisops canariensis, Puton, Rev. Ent., vol. XVIII, p. 80.
1904. Anisops canariensis, Kirkaldy, Wiener Ent. Zeit., vol. XXIII. pp. 117, 132.
1922. Anisops canariensis, Lindberg, Notulae Entomologicae, vol. II, pp. 47, 48, text fig. 4
(brief description and sketch of head).

1932. Anisops canariensis, Poisson, Bull. Soc. Ent. France, vol. XXVII, p. 42.

194S. Anisops canariensis canariensis, Poisson, Inst. Rech. Sahariennes Universite D'Alger,
p. 5, text figs. 14, b. d (places A perplexa Poisson as a subspecies of A. canariensis Noualhier).

This species was not present in the material at my disposal.
Therefore, I shall present a translation of the original.

Anisops canariensis n. sp. — Canary: pools of Tamaraceite, moun-
tain stream between Tafira and San Lorenzo; Tenerife; Laguna.

Testaceous, very pale dorsally, with the elytra translucent. Ven-
trally, varies from black to testaceous. Head of male and female
similar, the former having only the clypeus a little strumose. Eyes
very large; their sides almost parallel, seen from above; as advanced
as the frons (while, with A. producta, the frons is always visible in
regarding the insect in profile, here it is the anterior border of the
eye which forms, in this position, the anterior border of the head).
Pronotum a good third shorter than the scutellum. The remainder,
as with A. producta, from which in addition it distinguishes itself
by the smaller size and the less pronounced dorsal body longitudinal
curvature. Length 6/2-7 mm.

Anterior tibiae enlarged, with a red tooth at the internal base.

Location of types. — Type material from the Canary Islands in the
Paris Museum.

Anisops ciliata Stal

1868. Anisops ciliatus Stal, Ofver. Kongl. Yeten. Akad. Fordhandl., vol. VII, p. 137. (nee
Nolonecta ciliata Fabricius).

1898. Anisops ciliata, Kirkaldy, Annal. Mus. Civ. Stor. Nat. Genova, vol. XXXIX, p. 145
(determined in error; probably A. pellucens, Gerstaecker).

1933. Anisops ciliata. Lundblad, Arch, fur Hydrob., Suppl. vol. XII, p. 164 (points out
that Notonecta ciliata Fabricius is actually an Enithares).

This species was not present in the material at my disposal.
Therefore, I shall present a translation of the original description.
Dirty, yellowish-white; with the penultimate joint of the rostrum

Brooks: The Genus Anisops 463

above, with all the last joint, the venter, all these dark, with a slant-
ing white band before the coxae, connexivum spotted; ventral carina
yellowish-white.

The head seen from above or from the side not projecting very
far before the eyes, with the vertex and frons narrow, with the base,
on this side below the middle rather narrow and almost equally
broad, with that base very lightly furrowed. Thorax light, a
little narrowed in front. Scutellum light. Hemelytra seen against
the light, very lightly punctuated. The claws of the front tarsi
somewhat short, the claw on the outside scarcely longer than the
inner claw.

Very much akin to Anisops austrolis, with the back of the abdo-
men pale, with the head lightly furrowed, somewhat narrower ver-
tex and frons, with thoracic carina broad, obtuse, missing in front,
with anterior margin broadly angulate, apex projecting in a broad
angle between the eyes ( not truncate in the same place ) , with the
exterior claw of the front tarsi scarcely longer than the inner claw.

Location of types. — Lundblad (loc. cit.) states that Stal's speci-
mens of Anisops ciliatus which were in the Copenhagen Museum
have been destroyed.

Remarks: — This species along with many others was inaccurately
placed as a synonym of Anisops nivea ( Fabricius ) by Kirkaldy ( loc.
cit. ) . In an effort to clear up the confusion that has resulted it has
been necessary to take these species out of synonomy and place them
again as valid species. This I have done in regard to Anisops
ciliata.

Anisops endymion Kirkaldy

1904. Anisops endymion Kirkaldy, Wiener Ent. Zeit., vol. XXIII, pp. 112, 132.

1923. Anisops endymion, Hale, Rec. South Australian Mus., vol. II, p. 417 (translation
of the original description).

1924. Anisops endymion. Hale, Proc. Linn. Soc. New South Wales, vol. XLIX, pt. 4,
p. 465 (believes that this species may be a Paranisops due to pronotol distortion between the
eyes and pigmented hemelytra).

1933. Anisops endymion, Lundblad, Archiv. fur Hydrob., Suppl. vol. XII, p. 145 (a list
of Indo-australian and Pacific forms of this genus).

As no specimens of this species were available for study, I have
chosen to present a translation of the original description.

Elytra ash-coloured, transparent; posterior half of exocorium and
the clavus smoke-coloured, anterior half of clavus, basal margin of
corium and the basal half of exocorium, black. Veins of wings pale.
Metanotum brownish-black, lateral margins pale. Legs pale. Ab-
domen above dull, pale, in the centre black. Below black. Vertex
longitudinally grooved, hardly broader on the anterior margin than

464 The University Science Bulletin

the synthlipsis, the breadth of the latter barely half the width of an
eye.

Anterior margin of the pronotum between the eyes much more
distorted than in other species (the distorted portion rounded an-
teriorly); pronotum three-fourths broader than its length, one-half
times longer than the scutellum. Anterior and middle tibiae flat
and laterally expanded, broader at apex than at base; one-fifth times
longer than tarsi, first tarsal segment two-fifths longer than the other,
which is two and a half times as long as the claw. Length, 9 mm.;
breadth, 3 mm.

Hab. Australia: Swan River (Perth Museum, Scotland).

Only a single female of this distinct species is before me.

Anisops hyalina Fieber

1851. Anisops hyalinus Fieber, Abhandl. Konigl. Bohm. Gesells. Wiss., vol. V, pt. 7,
p. 482.

This species was not present in any of the material at my disposal,
so I have chosen to present a translation of the original description.

1. A. hyalinus m.

Back yellowish, basal and two end segments blackish. Upperside
of the anterior and middle femora black brown.

From east India ( Heifer, Mus. Berol. )

Length 4-/3-5 Lin. Dirty whitish yellow. Head almost truncate
in front. Vertex with a longitudinal furrow. Eyes approach one
another at the synthlipsis. Vertex very narrow, triangularly elon-
gate at the end. Third rostral segment as above, fourth entirely
brown black. Scutellum triangular, sides of equal length, sides
curved. The entire upperside finely punctate, fine and long curly
pubescence. Mesa- and meta-sternites black brown as the coxa.
Tibia and outer margin of the femora heavily haired. Abdomen
black, anal segment yellowish white, the penultimate segment with
yellowish spot. Median keel, a white spot in the lateral furrows
upon each segment. Connexivum with brown spots.

Bibliographical notes. — Stal (39) in 1868 indicated that this spe-
cies might be a synonym of A. ciliata Stal, and likewise Kirkaldy
(30) placed this species as a possible synonym of A. nivea (Fab.).
Both workers were incorrect, and due to the confusion that existed
in regard to A. nivea no taxonomist has redescribed the species,
though Esaki (7) pointed out that it was not a synonym of A. nivea
and that Kirkaldy had been mistaken. Whether or not the type is
still in the museum as mentioned by Fieber is questionable. It may
no longer be in existence.

Brooks: The Genus Anisops 465

Comparative notes. — The extremely large size (about 10 mm. a
line being one twelfth of an inch) places it within a very small
group. However, none of the species of that size have the trian-
gularly elongate vertex as noted by Fieber. Anisops sardea U.S.
which is the largest of the species possessing a cephalic horn, is
described by Fieber in the same paper as Anisops productus. It,
however, does not have the narrow vertex as posessed by A. hyalinns.
The author of this species does not mention whether or not he was
describing a female, but nevertheless this species was not present
in the vast Indian material I had for study.

Anisops lanceolata Poisson

(PI. LIII, fig. 87)

1949. Anisops lanceolata Poisson, Rev. Francaise Ent., vol. XV, p. 168, text figs. 7A, B,
C, D, E.

This species was not present in the material at my disposal.
Therefore, I shall present a translation of the original description
and copies of the figures.

Anisops lanceolata n. sp.

Body elongate, fusiform. General pigmentation of a flavus yellow
with the hemelytra of uniform color, hyaline, transparent and shin-
ing. Synthlipsis very narrow in the two sexes; narrower on the
males than on the females ( about ten to eleven times narrower than
the anterior width of the vertex ) . Scutellum a little longer medianly
than the pronotum.

Male: anterior leg, femur — 62, tibia — 63, tarsus — 40, claws — 13.

Tibial comb of disassociated elements, irregularly arranged and
unequally developed; one or two elements of the comb bifurcated.

Third rostral segment with developed spurs (rostral prong) and
profundly striated on the external border.

Length 6.8 mm.; 6.9 mm.

Distr. one male, one female; Crater of Monoun.

Location of types. — Types located in Poisson's collection.

Comparative notes. — This species with its very narrow synthlipsis,
shape of the anterior male femur, peculiar stridulatory comb and
the male pronotum with its lateral margins almost parallel is
strongly suggestive of a close relationship with Anisops breddeni
Kirkaldy from which it differs in not having the eyes of the males
holoptic.

30—3286

466 The University Science Bulletin

Anisops letitia Hutchinson

(PI. LIII, fig. 88)

1929. Anisops letitia Hutchinson, Ann. South African Mus., vol. XXV, pt. 3, pp. 385-
386, PI. XXIX, fig. 11; PI. XXXI, fig. 3.

This species was not present in the material at my disposal.
Therefore, I shall present a copy of the original description and
copies of the figures.

Anisops letitia n. sp.

Yellowish-white, with apical part of dorsum abdominis and venter
abdominis dark.

Fusiform, less than four times as long as wide, broadest before
the middle, at about the first fourth of the elytra; pronotum narrower
than the greatest width of the body, but slightly wider than that of
the head and eyes.

Head and eyes large; notocephalon with a longitudinal depres-
sion between two slightly swollen areas along its anterior half, not
reaching vertex and spreading out posteriorly before another in-
definite raised area. Facial tubercle well marked and often im-
pressed at the sides. Vertex about one fourth as wide as head and
eyes and two and one half to three and one half times as wide as the
synthlipsis. Pronotum smooth with a few pale hairs, less than twice
as wide as long, longer than the scutellum in some species (type)
shorter in others, carinated posteriorly, the carina not reaching the
posterior margin and variably produced anteriorly; posterior margin
convex, or at most slightly flattened, not sinuate or emarginate.

Scutellum fairly rugosely punctured.

Male: Third joint of rostrum with short prongs.

Anterior tibia just over one-half as long again as the tarsus, which
is about two and one-half times as long as the longer claw. Tibia
rather flattened, anterior margin with three spinous hairs basally,
posterior margin with two of its basal hairs elongate.

Stridulatory comb very small, about one-twentieth of the length
of the tibia, subparallel sided, of about sixteen lamellae. Tarsus
with a spinous hair and a few very small peg-like hairs basally.

Intermediate tibia just under two and one-third times as long as
the first (1 and 2) tarsal joint, which is about one and one-fourth
to one and one-half times as long as the third; the latter about
twice as long as the longer claw.

Posterior tibia one and one-third times as long as the tarsus.

Female: Eyes slightly less prominent than in the male.

Anterior tibia slightly shorter than in males, nearly two and one-

Brooks: The Genus Anisops 467

half times as long as the first ( 1 and 2 ) tarsal joint, which is about
one-half as long again as the second longer claw, rather over one-
half as long as the latter.

Intermediate tibia just over twice as long as the first (1 and 2)
tarsal joint, which is just under one and one-fourth times as long as
the third, the latter being about two and three-fifths as long as the
longer claw.

Posterior tibia about one and one-third as long as the tarsus.

Length: 8-9 mm.

Cape; Hovieson's Poort, near Grahanstown.

This very distinct species may be recognized at once by the con-
vex or straight posterior border to the elytra and the small stridula-
tory comb. It is evidently very local.

Location of types. — Type material located in the South African
Museum.

Comparative notes. — This species together with A. pcllucens
Gerst. are the two largest species known from South Africa. How-
ever, they are easily separated from one another on the basis of the
pronotum which in A. letitia is not medianly emarginate as in A. pel-
lucens. The synthlipsis of A. letitia is less than one half the anterior
width of the vertex whereas in A. pelhicens the synthlipsis is more
than one half the anterior width of the vertex.

Anisops mauricensis Poisson

(PI. LIII, fig. 86)
1945. Anisops mauricensis Poisson, Bull. Soc. Ent. France, vol. 50, p. 93, text fig. 7.

This species was not present in the material at my disposal.
Therefore, I shall present a translation of the original description
and copies of the figures.

Anisops mauricensis n. sp.

Vertex four times wider than the synthlipsis on the males and 2.5
times wider on the females. Head (eyes included) 1.3 times wider
than it is long up to the synthlipsis. Pronotum 1.3 times wider than
long. Scutellum 1.7 times longer than the pronotum.

General color yellowish brown. Abdominal stemites black in
the same way as the metanotum. Lateral spur of the third rostral
segment almost two times longer than the body of the segment.
Anterior tibia 1.6 times longer than the tarsi; the tibial comb carries
23-24 elongate teeth and the chaetotaxy of the tibia is composed in
particular of a short spine on the internal apical surface, followed
by three others on the tarsi.

Length 6.5 mm.

468 The University Science Bulletin

Distribution: one male, two females, He Mauritius, Curepipe
(Ch. Alluaud, 1897).

Observation: The species appears similar to A. cananariensis
Noualhier, but is differentiated, in particular, by the extreme nar-
rowness of the synthlipsis.

Location of types. — Type material located in Poisson's collection.

Anisops meulenaerei Poisson

(PI. LIII, fig. 89)

1940. Anisops meulenaerei Poisson, Bull. Mus. Royal Hist. Nat. Belgique, vol. XVI,
no. 27, pp. 8-10, text figs. 9, 10 ,11, 12.

This species was not present in the material at my disposal.
Therefore, I shall present a translation of the original description
and copies of the figures.

Anisops meulenaerei n. sp.

General pigmentation flavus.

Back of the abdomen flavus, with the margins of the tergites black.
Hemelytra hyaline and transparent. Eyes globose and slate colored.
Synthlipsis two times narrower than the anterior width of the vertex,
Pronotum 1.5 to 1.6 times wider than long; 2.85 times longer than
the scutellum. Third rostral segment provided with short spurs.

Anterior leg: claws 3.6 times shorter than the tarsi, which is 1.6
times shorter than the tibia and 1.1 times shorter than the femur;
the latter segment being itself 1.4 times shorter than the tibia.

Stridulatory apparatus with 23 long teeth and associated with a
strong, spiny tubercle situated at the beginning of a row of seven
long bristles. A strong spine at the extremity (tarsal) of the an-
terior femur. The anterior tarsus carries five spines distributed
regularly.

Length 9.5 mm., width 2.5-2.75 mm.

Distribution: Ethopia, Goba: one male, 1934-35 (R. de Meulen-
aerei), Mus. Roy. Hist. Belgian. 10,738 (type).

Comparative notes. — This species appears to be similar to Anisops
ares Hutchinson, from which it differs primarily in the chaetotaxy
of the male front leg as the latter species lacks the spur adjacent to
the stridulatory comb and also in the shape of the anterior male
femur which is parallel for its entire length with the apex rounded
while in A. ares Hutch, only the basal three-fourths is parallel.

Brooks: The Genus Anisops 469

Anisops milloti Poisson

(PI. LIV, fig. 90)

1948. Anisops milloti Poisson, Mem. Inst. Seient. Madagascar, Serie A, vol. I, p. 109,
text figs. 20, A. B.

This species was not present in the material at my disposal.
Therefore, I shall present a translation of the original description
and copies of the figures.

Anisops (A) milloti n. sp.

General color flavus (after standing in alcohol at 70 degrees),
eyes dark maroon. Abdominal sternites dark brown. Body very
fusiform. Head including the eyes, slightly wider than the pro-
notum and, viewed dorsally, a little shorter than the pronotum
(1.13 times shorter); pronotum 1.10 times shorter than the scutellum
and nearly two times wider at the base than its median length.
Synthlipsis four times narrower than the anterior width of the vertex.
Facial tubercle very prominent, overhanging the base of the rostrum.
Appendages of the third rostral segment well developed and
rounded at their extremity. Anterior tibia and middle tibia rather
dilated and flattened.

Male, anterior leg: femur — 22, tibia — 37.5, tarsus — 23, claws — 13.

Stridulatory comb rather well developed, represented by a groove
embelished with long thin bristles and with 7-8 spatulate setae.

Distribution: Three males Angavokely, Madagascar.

Comparative notes: The ventral projection of the facial tubercle
is suggestive of the similar condition found on the males of A. wakc-
fieldi White, however, in the latter case the projection is strongly
accuminate and not truncate as in this species. Also the rostral
prongs of the third rostral segment of the males of A. wakcfieldi are
very short, only about one-fourth the length of the third rostral seg-
ment whereas those of A. milloti appear to be at least as long as the
third rostral segment.

Location of types. — Type material located in Poisson's collection.

Anisops ocularis Hale

1923. Ariisops ocularis Hale, Rec. South Australian Mus. vol. II, no. 3, pp. 412-413, text
fig. 366.

1933. Anisops ocularis, Lundblad Arch, fur Hydrob. Suppl. vol. XII, p. 145 (a list of
Indo-australian and Pacific forms of this genus).

This species was not present in the material at my disposal.
Therefore, I shall present a translation of the original description.

Head, including the large and prominent eyes, very slightly nar-
rower than the pronotum; notocephalon with a swelling on each

470 The University Science Bulletin

side, not reaching to base of head, converging at vertex and con-
tinued as median carina on to the very narrow face; synthlipsis
about 1.5 in vertex, 5 times in the width of an eye. Pronotum pale
testaceous; as long as the head, twice as wide as long, with a feeble
median carina, which disappears posteriorly; hinder edge slightly
concavely incised. Scutellum testaceous, with a large, pale triangu-
lar patch near each anterior angle; wider than long, and about 1.25
times as long as the pronotum. Metanotum testaceous; upperside
of abdomen pale, with the posterior segments in parts black. Un-
derside of abdomen black, ventral carina and lateral edges ochra-
ceous. Anterior tibiae much expanded, the greatest width (near
the base) being .3 of the length; anterior tarsi 1.6 times in tibiae, 3
times longer than the longer claw. Length, 8 mm.; width, 2.5 mm.

Hob. Northern Territory: Darwin (W. K. Hunt).

The type is the only representative of this species before me. As
in A. doris, the eyes are large, the notocephalon is narrow, and the
pronotum is short; it differs, however, in the wider synthlipsis, the
markedly more robust form, the much more expanded anterior
tibiae of the male, etc. It resembles Distant's description and figure
of A. cleopatra from New Caledonia, but in that species the synth-
lipsis is "not more than half" as wide as the vertex and the size is
smaller.

Location of types. — Male type located in the South Australian
Museum.

Anisops pugnax Poisson

(PI. LIV, fig. 92)

1945. Anisops pugnax Poisson, Bull. Soc. Ent. France, vol. 50, p. 94, text fig. 8.

This species was not present in the material at my disposal.
Therefore, I shall present a translation of the original description
and copies of the figures.

Anisops pugnax n. sp.

Female: vertex 1.7 times wider than the synthlipsis. Head 2.5
times wider than long. Pronotum 2.3 times wider than long; its
posterior margin slightly concave. Scutellum 1.2 times shorter than
its basal width and 1.5 times longer than the pronotum.

Vertex, pronotum as well as the legs, yellowish. Scutellum black.
Scutellar border of the hemelytra as well as the internal margin of
the hemelytral pit, trimmed with carmin. Metanotum and the last
abdominal tergites black, the others yellowish; abdominal sternites
black.

Brooks: The Genus Anisops 471

Male: anterior tibia 1.2 times longer than the tarsus. The tibial
comb is formed of 12 irregular teeth.

Length: 7.5 mm. — male; 8 mm. — female.

Distribution: one male, one female, Mt. Kenia (C. & N. O.);
lower zone, prairies, 2000m. (C. Alluaud. 1908).

Location of types. — Types destroyed during the war.

Comparative notes. — The cylindrical distinct teeth of the stridula-
tory comb are similar to those found in A. adonis though in the
latter case there are approximately seven instead of the twelve of A.
pugnax. This species is much larger than A. adonis and the anterior
male tibia is one and two tenths the length of the anterior tarsus
whereas the anterior male tibia of A. adonis is one and one half the
length of the fore tarsus.

Anisops worthingtoni Jaczewski

(PI. LIV, fig. 91)

1933. Anisops worthingtoni Jaczewski, Linn. Soc. Jour., vol. XXXVIII. pp. 343-345, text
figs. 1, 2, 3, 4, 5.

1935. Ajiisops worthingtoni Poisson, Mus. Nat. Hist. Nat. vol. III. p. 211, fig. 25, a, b
(ecological Lote).

This species was not present in the material at my disposal.
Therefore, I shall present a copy of the original description and
copies of the figures.

Anisops (Anisops) worthingtoni sp. n.

Lake Rudolf, stn. 280, 1 male, 1 female.

Length 6.5 mm. Yellowish white, fairly slender.

Head of the male produced in front into a distinct prominence,
resembling that of the Australasian A. fieberi Kirk. (Hale, 1923),
much smaller, however, than the cephalic prominence in A. sardea
H. S. At its apex some elongated dark hairs are inserted. Face of
the male flattened. Head of the female simple, vertex forming a
very slight, but easily discernible convex prominence between the
eyes, when seen from above. Vertex in both sexes a little over three
times as wide as the synthlipsis. Rostral prongs of the males moder-
ately divergent.

Trochanter of the front legs of the male armed with two groups
of short, slightly curved pegs, one at the basal prominence, the
other at the lower margin. Tibial prong with about fifteen lamellae.
Outer margin of the tibia with two thickened spines and a short peg
inserted more distally. Inner margin with three long basal bristles
and a number of shorter ones. Tarsal joint with a continuous row
of short spines along its outer margin. Tibia one and one-fourth

472 The University Science Bulletin

times as long as the tarsus, which is twice as long as the longer claw.

Sinistral spine of the seventh abdominal segment of the male
thickened in its basal portion, curved and tapering towards the apex.
Parameres as shown in the text figure.

This species differs at once from all other African representatives
of the genus by the structure of the head and of the front legs in the
males, the females may be recognized by their slightly prominent
vertex.

I have pleasure in dedicating this species to Dr. E. B. Worthing-
ton, its discoverer.

Location of types. — Type material in Cambridge Museum.

BIBLIOGRAPHY ON ANISOPS

(Other references cited under species)

1. Amyot, C. J., and A. Serville. 1843. Hemipteres, p. 453.

2. Bare, C. O. 1928. Haemoglobin Cells and Other Studies of the Genus
Buenoa ( Hemiptera, Notoneetidae ) in: The University of Kansas Science
Bulletin, vol. XVIII.

3. Berg, C. 1879. Hemiptera Argentina, p. 119.

4. Bergroth, E. 1906. Systematische und Synonymische Bermerkunger
iiber Hemipteres in: Wiener Entomologische Zeitung, vol. XXV, pp. 1-12.

5. Breddin, P. 1901. Die Hemipteren von Celebes in: Abhandlungen der
Naturf'orschenden Gesellschaft zu Halle, vol. XXIV, p. 103.

6. Distant, W. L. 1906. Rhynchota from New Caledonia and the Sur-
rounding Islands in: New Caledonia, Zoologie, vol. I, pp. 371-391, Pit. I.

7. Esaki, T. 1926. Verzeichniss der Hemiptera-Heteroptera der Insel For-
mosa in: Annales Musei Nationalis Hungarici, vol. XXIV, pp. 136-139.

8. 1928. Aquatic and Semi-aquatic Heteroptera in: Insects of

Samoa, prt. 2, text figs. 1-6.

9. Fabricius, J. C. 1775. Systema Entomologiae, p. 690.

10. 1794. Entomologia Systema, vol. IV, p. 58.

11. Fieber, F. X. 1851. Rhynchotographieen in: Abhandlungen Kongl.
Bohmischen Gesellschaft Wissenschaften, vol. V, (7), pp. 469-486.

12. 1860. Die Europaichen Hemiptera, p. 100.

13. Hale, H. M. 1923. The Breedings Habits of a Backswimmer (A.
hyperion) in: The South Australian Naturalist, vol. IV, No. 3, pp. 124-128,
text fig. 1.

14. 1923. Studies in Australian Aquatic Hemiptera in: Records of

the South Australian Museum, vol. II, no. 3, pp. 397-424, Pits. X, XI; text
figs. 361-367.

15. 1924. Notes on Eggs, Habits, and Migration of some Australian

Aquatic Bugs in: The South Australian Naturalist, vol. V, no. 4, pp. 133-
143.

16. 1925. The Aquatic and Semi-aquatic Hemiptera in; Archiv Zool-

ogi utgivet av K. Svenska Vetenskapaakademien, vol. 17, no. 20, pp. 1-19,
text figs. 1-8.

Brooks: The Genus Anisops 473

17. Herrich-Shaffer, G. A. W. 1850. Die Wanzenartigen Insecten, vol. 9,
pp. 40-41, text fig. 904.

18. Hesse, A. J. 1925. Contribution to a Knowledge of the Fauna of S. West
Africa in: The Annals of the South African Museum, vol. 23, pp. 1-179,
Pits. I-VIII.

19. Hoffmann, W. E. 1933. A Preliminary List of the Aquatic and Semi-
aquatic Hemiptera of China, Chosen ( Korea ) , and Indo-China in : Ling-
nan Science Journal, vol. 12, pp. 243-258.

20. 1941. Catalogue of Aquatic Hemiptera of China, Indo-China,

Formosa, and Korea in: Lingnan Science Journal, vol. 20, pp. 1-78.

21. Horvath, G. 1888. Materiaux pour servir a l'etude des Hemipteres de
la faune palearctique in: Revue d'Entomologique, vol. VII, 75.

22. Hungerford, H. B. 1933. The Genus Notonecta of the World in: The
Universitv of Kansas Science Bulletin, vol. XXI, no. 9, pp. 1-195, Pits.
I-XVII.

23. Hutchinson, G. E. 1933. A Revision of the Notonectidae and Corixidae
of South Africa in: Annals of the South African Museum, vol. XXV, pp.
360-474, Pits. XXVII-XLL.

24. 1930. Report on Notonectidae, Pleidae, and Corixidae (Hemip-
tera, Mr. Omer-Cooper's Investigation of the Abyssinian Fresh Waters.
Dr. Hugh Scott's Expedition) in: Proceedings of the Zoological Society of
London, vol. XXIX, prt. 2, pp. 437-466.

25. 1932. Supplementary Report on Notonectidae, Pleidae, and Corixi-
dae (Hemiptera: Mr. Omer-Cooper Investigation of the Abyssinian Fresh
Waters (Dr. Hugh Scott's Expedition) in: Proceedings of the Zoological
Society of London, vol. XXXI, prt. 1.

26. 1933. The Zoo-geography of the African Aquatic Hemiptera in

Relation to Past Climatic Changes in: Sonderadruck aus Internationale
Revue der gesamten Hydrobiologie und Hydrographie, vol. 28, prt. 5/6,
pp. 436-468.

27. Hutton, F. W. 1897. On the Hemiptera of New Zealand in: Transac-
tions of the New Zealand Institute, vol. XXX, p. 179-180.

28. Jaczewski, T. 1826. Notes on Some West African Heteroptera in: An-
nalibus Zoologicis Musei Polonici Historae Naturalis, vol. V, pp. 62-106,
text figs. 1-72, Pits. I, II.

29. Kirkaldy, G. W. 1901. Miscellanea Rhvnchotalia in: The Entomologist,
vol. 34, pp. 93-135.

30. 1904. Uber Notonectiden in: Weiner Entomologische Zeitung,

vol. XXIII, pp. 93-135.

31. 1906. List of the Genera of Pagiopodus, Hemiptera-Heteroptera

in : Transactions of the American Entomological Society, vol. XXXII, no. 2.

32. Lundrlad, O. 1933. Sur Kenntnis du Aquatilen und Semiaquatilen
Hemipteren von Sumatra, Java, und Bali in: Sonder-Abdruck aus dem
Archiv fur Hydrobiologie, Suppl. vol. XII, pp. 1-489, text figs. 1-142, Pits.
I-XXI.

33. Myers, J. G. 1926. Biological Notes on New Zealand Heteroptera in:
Transactions and Proceedings of the New Zealand Institut, vol. LVI, pp.
468-469.

34. Poisson, R. 1925. Quelques Observations sur YAnisops producta Fieber
in: Comtes Rendus Hebdomadaires des seances de 1' Academic des Sciences,
Paris, vol. 92, pp. 4-7, text fig. 1.

474 The University Science Bulletin

35. UAnisops producta Fieber ( Hemiptera, Notonectidae ) Observa-
tions sur son Anatomie et sa Biologie in: Archives de Zoologie Experi-
mental et Generale, vol. 65, pp. 181-208, text figs. I-XVII.

36. Puton, A. 1880. Synopsis des Heteropteres de France, p. 217.

37. Rambur, J. D. 1840. Faune Entomoligique de l'Andalouise, vol. 2, no. 5,
p. 190.

38. Spinola, M. M. 1840. Essai sur les Insects Hemipteres Rhyngotes ou
Heteropteres, p. 58.

39. Stal, C. 1865. Hemiptera Africana, vol. Ill, p. 191.

40. White, F. B. List of the Hemiptera of New Zealand in: The Entomolo-
gist Monthly Magazine, vol. XV.

41. Wu, C. F. List of Kwangtung Hemiptera in: Lingnan Science Journal,
vol. XII, pp. 213-214.

42. Catalogue Insectorum Sinensium, vol. II, pp. 575-576.

INDEX

PAGE

adonis Hutchinson 362

agalia Hutchinson 318

ali Distant 459

alluaudi Poisson 459

amaryllis Hutchinson 324

aphrodite Kirkaldy 460

apicalis Stal 337

ares Hutchinson . . 405

assimilis White 411

balds Hutchinson 453

harbata n. sp 387

barrenensis n. sp 402

batillifrons Lundblad 420

biroi n. sp 454

bouvieri Kirkaldy 430

breddeni Kirkaldy 439

calcaratus Hale 461

campbelli n. sp 322

canaliculata n. sp 367

canadensis canadensis Noualhier, 462

canadensis perplexa Poisson 339

cavifrons n. sp 417

ciliata Stal 462

cleopatra Distant 384

coutierei n. sp 346

crinita n. sp 443

deanei n. sp 381

debilis Gerstaecker 446

decipiens Hutchinson 401

doris Kirkaldy 395

edepol Kirkaldy 341

elstoni n. sp 326

endijmion Kirkaldy 463

eros Hutchinson . 327

evansi n. sp 350

exigera Horvath 447

extendofrons n. sp 432

fijiensis n. sp .... 361

genji Hutchinson 371

gobana Poisson 353

gracilis Hutchinson 436

graciloides n. sp 434

grandis Poisson 389

gratus Hale 352

hacked n. sp 331

hancocki Hutchinson 399

hungerfordi Poisson 404

hyalina Fieber 464

PAGE

hypatia Hutchinson 452

Hyperion Kirkaldy 332

jaczewski Hutchinson 364

kampalensis Hutchinson 328

kempi n. sp 441

lanceolata Poisson 465

leesoniana Hutchinson 438

letitia Hutchinson 466

leucothea Esaki 394

lipovskyi n. sp 379

madagascarensis n. sp. 428

majiensis n. sp 444

malkini n. sp 349

rnauricensis Poisson 467

metdenaerei Poisson 468

milloti Poisson . . 469

nasuta Fieber 416

nigrolineata Lundblad 409

nivea ( Fabricius ) 373

nodulata n. sp 336

occipitalis Breddin . . 344

ocularis Hale 469

paracrinita n. sp 329

paranigrolineata n. sp 407

pellucens Gerstaecker 347

philippinensis n. sp . . 383

poweri Hutchinson 437

praetexta Hutchinson 386

psyche Hutchinson 369

pugnax Poisson 470

rigoensis n. sp 456

robusta Hutchinson 358

sardea Herrich-Shaffer 423

semita n. sp 366

sikoroensis n. sp 360

stall Kirkaldy 319

tahitiensis Lundblad 376

tanalensis n. sp 391

tasmaniaensis n. sp . . 378

thienemanni Lundblad 413

timorensis n. sp 343

tuberculata Poisson 334

varia Fieber 355

vitrea Signoret 449

icaltairensis n. sp 457

ivakefieldi White 397

windi n. sp 392

worthingtoni Jaczewski . 471

(475)

■4^6 The University Science Bulletin

PLATE XXXVI

Figure

1. Male Anisops — dorsal view.

2. Male Anisops — ventral view.

3. Left genital elasper.

4. Right genital elasper.

5. Anisops elstoni n. sp.

5a. Inner surface view of male left fore leg.

5b. Left lateral view of third and fourth male rostral segments.

5c. Enlarged view of left stridulatory comb.

6. Anisops amaryllis Hutchinson.

6a. Inner surface view of male left fore leg.

6b. Left lateral view of third and fourth male rostral segments.

6c. Enlarged view of left stridulatory comb.

7. Anisops agalia Hutchinson.

7a. Inner surface view of male left fore leg.
7b. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

477

PLATE XXXVI

vertex

synfhlips/s

pronof um
scufelli/m

hemelylrol pit

opaque zone

of membrane

3 LEFT GENITAL CLASPER

I.MALE ANISOPS (dorsum)

4. RIGHT GENITAL CLASPER

facial tubercle
Jabrum
rostral prong

keel

5 a

5. A.ELSTONI

2 MALE ANISOPS (venter)

7A AGALfA

478 The University Science Bulletin

PLATE XXXVII

Figure

8. Ayusops campbelli n. sp.

8a. Inner surface view of male left fore leg.

8b. Left view of third and fourth male rostral segments.

8c. Enlarged view of left stridulatory comb.

9. Anisops stall Kirkaldy.

9a. Inner surface view of male left fore leg.
9b. Enlarged view of left stridulatory comb.
10. Anisops eros Hutchinson.

10a. Inner surface view of male left fore leg.

10b. Left view of third and fourth male rostral segments.

10c. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

479

PLATE XXXVII

10. A EROS

480 The University Science Bulletin

PLATE XXXVIII

Figure

11. Anisops tuberculata Poisson.

11a. Inner surface view of male left fore leg.
lib. Enlarged view of left stridulatory comb.

12. Anisops paracrinita n. sp.

12a. Inner surface view of male left fore leg.

12b. Left view of third and fourth male rostral segments.

12c. Enlarged view of left stridulatory comb.

13. Anisops hackeri n. sp.

13a. Inner surface view of male left fore leg.

13b. Left view of third and fourth male rostral segments.

13c. Enlarged view of left stridulatory comb.

14. Anisops nodulata n. sp.

14a. Inner surface view of male left fore leg.

14b. Left view of third and fourth male rostral segments.

14c. Enlarged view of left stridulatory comb.

15. Anisops kampalensis Hutchinson.

15a. Inner surface view of male left fore leg.

15b. Left view of third and fourth male rostral segments.

15c. Enlarged view of stridulatory comb.

16. Anisops hyperion Kirkaldy.

16a. Inner surface view of male left fore leg.

16b. Left view of third and fourth male rostral segments.

16c. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

481

PLATE XXXVIII

I5.A. KAMPALENSIS

31—3286

482 The University Science Bulletin

PLATE XXXIX

Figure

17. Anisops apicalis Stal.

17a. Inner surface view of male left fore leg.

17b. Left view of third and fourth male rostral segments

17c. Enlarged view of left stridulatory comb.

18. Anisops timorcnsis n. sp.

18a. Inner surface view of male left fore leg.

18b. Left view of male rostrum.

18c. Enlarged view of left stridulatory comb.

19. Anisops edepol Kirkaldy.

19a. Inner surface view of male left fore leg.
19b. Enlarged view of left stridulatory comb.

20. Anisops canariensis perplexa Poisson.

20a. Inner surface view of male left fore leg.

20b. Left view of third and fourth male rostral segments.

20c. Enlarged view of left stridulatory comb.

21. Anisops coutierei n. sp.

21a. Inner surface view of male left fore leg.

21b. Left view of third and fourth male rostral segments.

21c. Enlarged view of left stridulatory comb.

22. Anisops occipitalis Breddin.

22a. Inner surface view of male left fore leg.

22b. Left view of third and fourth male rostral segments.

22c. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

483

PLATE XXXIX

2I<J 2I.A COUTIEREI

2 2a

22. A OCCIPITALIS

484 The University Science Bulletin

PLATE XL

Figure

23. Anisops malkini n. sp.

23a. Inner surface view of male left fore leg.

23b. Left view of third and fourth male rostral segments.

23c. Enlarged view of left stridulatory comb.

24. Anisops fijiensis n. sp.

24a. Inner surface view of male left fore leg.

24b. Left view of third and fourth male rostral segments.

24c. Enlarged view of left stridulatory comb.

25. Anisops jaczewski Hutchinson.

25a. Inner surface view of male left fore leg.

25b. Left view of third and fourth male rostral segments.

25c. Enlarged view of left stridulatory comb.

26. Anisops pellucens Gerstaecker.

26a. Inner surface view of male left fore leg.

26b. Left view of third and fourth male rostral segments.

26c. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

485

PLATE XL

25. A. JACZEWSKI

26. A. PELLUCENS

486 The University Science Bulletin

PLATE XLI

Figure

27. Anisops evansi n. sp.

27a. Inner surface view of male left fore leg.

27b. Left view of third and fourth male rostral segments.

27c. Enlarged view of the left stridulatory comb.

28. Anisops gobana Poisson.

28a. Inner surface view of male left fore leg.

28b. Left view of third and fourth male rostral segments.

28c. Enlarged view of the left stridulatory comb.

29. Anisops adonis Hutchinson.

29a. Inner surface view of male fore leg.

29b. Left view of third and fourth male rostral segments.

29c. Enlarged view of the left stridulatory comb.

30. Anisops robusta Hutchinson.

30a. Inner surface view of male fore leg.

30b. Left view of third and fourth rostral segments.

30c. Enlarged view of the left stridulatory comb.

Brooks: The Genus Anisops

487

PLATE XLI

2 7a

28o.

28. A. GOBANA

27. A. EVANSI

29. A ADONIS

488 The University Science Bulletin

PLATE XLII

Figure

31. Anisops gratus Hale.

31a. Inner surface view of male left fore leg.

31b. Left view of third and fourth male rostral segments.

31c. Enlarged view of left stridulatory comb.

32. Anisops varia Fieber.

32a. Inner surface view of male left fore leg.

32b. Left view of third and fourth male rostral segments.

32c. Enlarged view of left stridulatory comb.

33. Anisops semita n. sp.

33a. Inner surface view of male left fore leg.

33b. Left view of third and fourth male rostral segments.

33c. Enlarged view of left stridulatory comb.

34. Anisops sikorensis n. sp.

34a. Inner surface view of male left fore leg.
34b. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

489

PLATE XLII

32. A. VARIA

3 la

31. A. GRATUS

3 3o
33. A. SEMITA

34. A. SIKOROENSIS

34 a

490 The University Science Bulletin

PLATE XLIII

Figure

35. Anisops canaliculata n. sp.

35a. Inner surface view of male left fore leg.

35b. Left view of third and fourth male rostral segments.

35c. Enlarged view of left stridulatory comb.

36. Anisops genji Hutchinson.

36a. Inner surface view of male left fore leg.

36b. Left view of third and fourth male rostral segments.

36c. Enlarged view of left stridulatory comb.

37. Anisops lipovskyi n. sp.

37a. Inner surface view of male left fore leg.

37b. Left view of third and fourth male rostral segments.

37c. Enlarged view of left stridulatory comb.

38. Anisops psyche Hutchinson.

38a. Inner surface view of male right fore leg minus the tarsus.
38b. Left view of third and fourth male rostral segments.
38c. Enlarged view of right stridulatory comb.

39. Anisops nivea (Fabricius).

39a. Inner surface view of male left fore leg.
39b. Enlarged view of left stridulatory comb.

40. Anisops tahiliensis Lundblad.

40a. Inner surface view of male left fore leg.

40b. Left view of third and fourth male rostral segments.

40c. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

491

PLATE XLIII

39. A. NIVEA

40.A.TAHITIENSIS

492 The University Science Bulletin

PLATE XLIV

Figure

41. Anisops praetexta Hutchinson.

41a. Inner surface view of male left fore leg.

41b. Left view of third and fourth male rostral segments.

41c. Enlarged view of left stridulatory comb.

42. Anisops barrenensis n. sp.

42a. Inner surface view of male left fore leg.

42b. Left view of third and fourth male rostral segments.

42c. Enlarged view of left stridulatory comb.

43. Anisops cleopatra Distant.

43a. Inner surface view of male left fore leg.

43b. Left view of rostrum.

43c. Enlarged view of left stridulatory comb.

44. Anisops philippinensis n. sp.

44a. Inner surface view of male left fore leg.

44b. Left view of third and fourth male rostral segments.

44c. Enlarged view of left stridulatory comb.

45. Anisops deanei n. sp.

45a. Inner surface view of male left fore leg.

45b. Left view of third and fourth male rostral segments.

45c. Enlarged view of left stridulatory comb.

46. Anisops lanalensis n. sp.

46a. Inner surface view of male left fore leg.

46b. Left view of third and fourth male rostral segments.

46c. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

493

PLATE XLIV

494 The University Science Bulletin

PLATE XLV

Figure

47. Anisops leucothea Esaki.

47a. Inner surface view of left male fore leg.

47b. Enlarged view of left stridulatory comb.

47c. Left view of third and fourth male rostral segments.

48. Anisops uindi n. sp.

48a. Inner surface view of left male fore leg.

48b. Left view of third and fourth male rostral segments.

48c. Enlarged view of left stridulatory comb.

49. Anisops biroi n. sp.

49a. Inner surface of left male fore leg.

49b. Left view of third and fourth male rostral segments.

49c. Enlarged view of left stridulatory comb.

50. Anisops grandis Poisson.

50a. Inner surface view of left male fore leg.

50b. Left view of third and fourth male rostral segments.

50c. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

495

PLATE XLV

48.A.WINDI

4 7. A. LEUCOTHEA,

49 A. Bl

496 The University Science Bulletin

PLATE XLVI

Figure

51. Anisops hungerjordi Poisson.

51a. Inner surface view of male left fore leg.

51b. Left view of third and fourth male rostral segments.

51c. Enlarged view of left stridulatory comb.

52. Anisops hancocki Hutchinson.

52a. Inner surface view of male left fore leg.
52b. Enlarged view of left stridulatory comb.

53. Anisops doris Kirkaldy.

53a. Inner surface view of male left fore leg.

53b. Left view of third and fourth male rostral segments.

53c. Enlarged view of left stridulatory comb.

54. Anisops barbata n. sp.

54a. Inner surface view of male left fore leg.

54b. Left view of third and fourth male rostral segments.

54c. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

497

PLATE XLVI

51. A. HUNGERFORDI

52. A HANCOCKI

53o 53. A. DORIS

32— 32SG

54. A. BARBATA

498 The University Science Bulletin

PLATE XLYll

Figure

55. Anisops wakefieldi White.

55a. Inner surface view of male left fore leg.
55b. Enlarged view of left stridulatory comb.

56. Anisops nigrolineata Lundblad.

56a. Inner surface view of male left fore leg.
56b. Enlarged view of left stridulatory comb.

57. Anisops ares Hutchinson.

57a. Inner surface view of male left fore leg.

57b. Left view of third and fourth male rostral segments.

57c. Enlarged view of left stridulatory comb.

58. Anisops tasmaniaensis n. sp.

58a. Inner surface view of male left fore leg.

58b. Left view of third and fourth male rostral segments.

58c. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

499

PLATE XLVII

57 A. ARES

58.A TASMANIAENSIS

500 The University Science Bulletin

PLATE XLVIII

Figure

59. Anisops paranig rolineata n. sp.

59a. Inner surface view of male left fore leg.

59b. Left view of third and fourth male rostral segments.

59c. Enlarged view of left stridulatory comb.

60. Anisops nasuta Fieber.

60a. Inner surface view of male left fore leg.
60b. Enlarged view of left stridulatory comb.

61. Anisops assimilis White.

61a. Inner surface view of male right fore leg minus the tarsus.
61b. Left view of third and fourth male rostral segments.
61c. Enlarged view of right stridulatory comb.

62. Anisops decvpiens Hutchinson.

62a. Inner surface view of male right fore leg minus the tarsus.

62b. Enlarged view of right stridulatory comb.

62c. Left view of third and fourth male rostral segments.

63. Anisops thienemanni Lundblad.

63a. Inner surface view of male left fore leg.

63b. Left view of third and fourth male rostral segments.

63c. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

501

PLATE XLVIII

62 o.
62. A. OECIPIEVS

63. A. THIENEMANNI

502 The University Science Bulletin

PLATE XLIX

Figoue

64. Anisops sardea Herrich-Shaffer.

64a. Inner surface view of male left fore leg.
64b. Enlarged view of left stridulatory comb.

65. Anisops bouvieri Kirkaldy.

65a. Inner surface view of male left fore leg.
65b. Enlarged view of left stridulatory comb.

66. Anisops extendojrons n. sp.

66:\. Inner surface view of male left fore leg.

66b. Left view of third and fourth male rostral segments.

66c. Enlarged view of left stridulatory comb.

67. Anisops ynadagascarensis Poisson.

67a. Inner surface view of male left fore leg.
67b. Enlarged view of left stridulatorv comb.

Brooks: The Genus Anisops

503

PLATE XLIX

65. A. 80UVIERI

6 4. A. S AROEA

6ba.
66.A. EXTENDOFRONS

67.A. MADAGASCARENSIS

504 The University Science Bulletin

PLATE L

Figure

68. Anisops crinita n. sp.

68a. Inner surface view of male left fore leg.

68b. Left view of third and fourth male rostral segments.

68c. Enlarged view of left stridulatory comb.

69. Anisops waltairensis n. sp.

69a. Inner surface view of male left fore leg.

69b. Left view of third and fourth male rostral segments.

69c. Enlarged view of left stridulatory comb.

70. Anisops leesoniana Hutchinson

70a. Inner surface view of male left fore leg.

70b. Left view of third and fourth male rostral segments.

70c. Enlarged view of left stridulatory comb.

71. Anisops batillifrons Lundblad.

71a. Inner surface view of male left fore leg.
71b. Enlarged view of left stridulatory comb.

72. Anisops poweii Hutchinson — Inner surface view of male left fore leg.

73. Anisops cavifrons n. sp.

73a. Inner surface view of male left fore leg.

73b. Left view of third and fourth male rostral segments.

73c. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisofs

505

PLATE L

72. A. POWERI

RONS

506 The University Science Bulletin

PLATE LI

Figurei

74. Anisops gracilis Hutchinson.

74a. Inner surface view of male left fore leg.

74b. Left view of third and fourth male rostral segments.

74c. Enlarged view of left stridulatory comb.

75. Anisops kempi n. sp.

75a. Inner surface view of male left fore leg.

75b. Left view of male rostrum.

75c. Enlarged view of left stridulatory comb.

76. Anisops vitrea Signoret.

76a. Inner surface view of male left fore leg.

76b. Left view of third and fourth male rostral segments.

76c. Enlarged view of left stridulatory comb.

77. Anisops exigera Horvath.

77a. Inner surface view of male left fore leg.

77b. Left view of third and fourth male rostral segments.

77c. Enlarged view of left stridulatory comb.

78. Anisops breddeni Kirkaldy.

78a. Inner surface view of male left fore leg.
78b. Enlarged view of left stridulatory comb.

79. Anisops graciloidcs n. sp.

79a. Inner surface view of male left fore leg.

79b. Left view of third and fourth male rostral segments.

79c. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

507

PLATE LI

79 A. GRACILO/DES

508 The University Science Bulletin

PLATE LII

Figure

80. Anisops clebilis Gerstaecker.

80a. Inner surface view of male left fore leg.

80b. Left view of third and fourth male rostral segments.

80c. Enlarged view of left stridulatory comb.

81. Anisops rigoensis n. sp.

81a. Inner surface view of male left fore leg.

Sib. Left view of third and fourth male rostral segments.

81c. Enlarged view of left stridulatory comb.

82. Anisops hypatia Hutchinson.

82a. Inner surface view of male left fore leg.

82b. Left view of third and fourth male rostral segments.

82c. Enlarged view of left stridulatory comb.

83. Anisops majiensis n. sp.

83a. Inner surface view of male left fore leg.

83b. Left view of third and fourth male rostral segments.

83c. Enlarged view of left stridulatory comb.

84. Anisops balcis Hutchinson.

84a. Inner surface view of male left fore leg.
84b. Enlarged view of left stridulatory comb.

Brooks: The Genus Anisops

509

PLATE LII

83a. 83AMAJIENSIS

510 The University Science Bulletin

PLATE LIII

Figure

85. Anisops alluaudi Poisson.

85a. Inner surface view of male right fore leg.

85b. Enlarged view of right stridulatory comb.

85c. Right view of third and fourth male rostral segments.

86. Anisops mauricensis Poisson.

86a. Inner surface view of male right fore tibia and tarsus.
86b. Enlarged view of right stridulatory comb.

87. Anisops lanceolata Poisson.

87a. Inner surface view of male right fore leg.

87b. Diagramatic frontal view of third male rostral segment.

87c. Enlarged view of right stridulatory comb.

88. A?iisops letitia Hutchinson — inner surface view of right fore tibia.

89. Anisops meulenaerei Poisson.

89a. Inner surface view of right male fore femur.

89b. Inner surface view of right male fore tibia.

89c. Inner surface view of right male fore tarsus.

89d. Enlarged view of right stridulatory comb.

89e. Right view of third and fourth male rostral segments.

Brooks: The Genus Anisops

511

PLATE LIII

85. A. ALLUAUDI

86 a.
86.A MAURICENSIS

87 A.LANCEOLATA

88. A.LETITIA

89b.

89.A.MEULENAEREI

512 The University Science Bulletin

PLATE LIV

Figure

90. Anisops milloti Poisson.

90a. Inner surface view of male right fore leg.
90b. Enlarged view of right stridulatory comb.
90c. Right view of male head.

91. Anisops worthingtoni Jaczewski.

91a. Inner surface view of male left fore tibia and tarsus.
91b. Left view of male head.

92. Anisops pugnax Poisson.

92a. Enlarged view of left stridulatory comb.

92b. Inner surface view of left male fore tibia and tarsus.

93. Anisops nivea (Fabricius) — left view of male head and pronotum.

94. Anisops edepol Kirkaldy — left view of male head and pronotum.

95. Anisops poweri Hutchinson — left view of male head and pronotum.

Brooks: The Genus Anisops

513

PLATE LIV

94. A. EDEPOL

33-3286

514 The University Science Bulletin

PLATE LV

Figure

96. Anisops hancocki Hutchinson — left view of male head and pronotum.

97. Anisops balcis Hutchinson — left view of male head and pronotum.

98. Anisops nasuta Fieber — left view of male head and pronotum.

99. Anisops breddeni Kirkald}' — left view of male head and pronotum.
100. Anisops bouvieri Kirkaldy — left view of male head and pronotum.

Brooks: The Genus Anisops

515

PLATE LV

99. A BPEDDENI

516 The University Science Bulletin

PLATE LVI

Figure

101. Anisops batillifrons Lundblad — left view of male head and pronotum.

102. Anisops sardea Herrich-Shaffer — left view of male head and pronotum.

103. Anisops sikoroensis n. sp. — left view of male head and pronotum.

Brooks: The Genus Anisops

517

PLATE LVI

518 The University Science Bulletin

PLATE LVII

Figure

104. Anisops tuberculata Poisson — left view of male head and pronotum.

105. Anisops nigrolineata Lundblad — left view of male head and pronotum.

106. Anisops wakefieldi White — left view of male head and pronotum.

107. Anisops stali Kirkaldy — left view of male head and pronotum.

Brooks: The Genus Anisops

519

PLATE LVII

THE UNIVEKSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIV, Pt. I] October 1, 1951 [No. 9

Concerning Oligocene Amphisbaenid Reptiles

BY

Edward H. Taylor

Abstract: The paper deals with the amphisbaenid fauna of the Orellan,
Middle Oligocene of Logan County, Colorado, and with a single species from
the Orellan of Converse County, Wyoming. The following species are de-
scribed: Hijporhina galbreathi sp. nov., Rhineura anblyceps sp. nov., Rhineura
hibbardi sp. nov., Rhineura wilsoni sp. nov., Gilmoreia attenuatus gen. et sp. nov.
The family Hyporhinidae is recognized for the genus Hijporhina, and the fam-
ily Crythiosauridae is proposed for a Mongolian genus Crythiosaurus.

INTRODUCTION

The amphisbaenid reptiles have been variously treated by pre-
vious authors. They have been regarded as a Family, as a Super-
family, and as an Infraorder under the Lacertilia (Sauria); as a
Suborder of equal rank with the Sauria, and even as a reptilian
Order. Some authors have placed all the known species in a single
family, the Amphisbaenidae; while others have found it convenient
to recognize several families. More than 30 genera have been
named.

This aberrant group of vertebrates appears to stand, in relation
to the other living Reptilia, much as do the Gymnophiona to other
living Amphibia; and it is doubtful that the differences from
Caudata or other Amphibia displayed by the Gymnophiona are
more significant than those that obtain between the Amphisbaenia
and the other groups of the Reptilia. I am strongly inclined toward
a recognition of these considerable differences by placing them in
an Order of their own. A discussion of the matter is, however, not
the primary concern of this paper. Romer (1932) considers them
of Subordinal rank, as do Smith and Taylor ( 1950 ) . These authors
are followed here.

There is considerable difference of opinion as to the validity of
many of the genera proposed for the living species, and the number
recognized varies much with different authors. Loveridge (1940)

(521)

522 The University Science Bulletin

has recently reviewed the African genera, recognizing Trogonophis,
P achy calamus, Agamodon, Baikia, Blanus, Amphishaena, Placogas-
ter, Geocalamus, Monopeltis and Dalophia* Laurent has recently
revived two genera Chirindia and Cynisca placed by Loveridge in
the genus Amphishaena. He describes a new genus Tomuropeltis,
thus bringing the total African genera to 13.

The American genera Bipes, Mexico, Bhineura, Florida and
Oligocene of Colorado, etc., Lepisternon Brazil, Bronia Brazil,
Aulura, Brazil, Anopsibaena, Brazil, Mesobaena, Venezuela, Di-
phalus, Virgin Islands, Cadea, Cuba, are probably all recognizable
genera. However, the generic names Chirotes, Hemichirotes, Eu-
chirotes and Bimanus of Mexico should doubtless be treated as
synonyms of Bipes. I am uncertain as to the status of Aporarchus,
Sarea, Monotrophis Cephalopeltis and Ancylocranium.

Four genera known only from fossils are recognized. These are
Hyporhina, Ototriton, Crythiosaurus and Gilmoreia (the latter
herein described).

The question of the number of living families in the amphis-
baenids is likewise unsettled. I regard the families Amphisbaenidae,
Bipedidae and Trogonophidae (for the acrodont group) as valid.
It is not impossible that certain other groups of species merit fam-
ily status. All of the previously known American fossil forms except
Hyporhina are presumably referable to the Amphisbaenidae. I
consider Hyporhina worthy of family rank under the name Hypor-
hinidae based primarily on the presence of the postorbital bone.
While the anatomy of the Mongolian genus Crythiosaurus is not
wholly known, the characters given seem to warrant the erection
of the family Crythiosauridae, the characters being those of the
genus.

It is possible that Ototriton belongs to still another family. Camp,
Taylor and Welles ( 1942 ) placed Ototriton in the Dolichosauridae
and Lestophis (the latter known only from vertebrae) in the
Sauria.

The Amphisbaenia represent a group of vertebrates that now
have become highly specialized for subterranean life by total or
partial loss of limbs, and the aquisition of a serpentine form. Be-
cause of this they have survived long after the extinction of their
more generalized terrestrial tetrapod ancestors. One may conceive
of a populous tetrapod suborder (or order) existing in the past,
and practically worldwide in distribution, that had diversified into
several families. Competition for food, probably among terrestrial

* The listing of the genus Dalophis (Loveridge 1941, p. 357) for Dalophis jallae and Dal-
ophis pistillum is an error. These are properly Dalophia, and are so treated elsewhere.

Taylor: Oligocene Amphisbaenid Reptiles 523

members of the same order, forced certain forms in several fam-
ilies to seek underground food. It is presumed that later the ter-
restrial generalized forms succumbed to competition with mem-
bers of other orders.

Counterparts of these happenings are suggested in the Order
Gymnophiona of the Class Amphibia. Here only the limbless
burrowing forms have survived. In the Order Caudata, members
of certain suborders have survived probably because of their sub-
terranean habitat (Suborder Meantes). In other suborders certain
of the less specialized forms still exist. In the Mutabilia a few
species have become subterranean, while many of the families retain
the generalized terrestrial tetrapod form.

Similar surviving groups occur in the Sauria. The families Anely-
tropidae, Feylinidae, Dibamidae, Anniellidae, and Pygopodidae all
are regarded as burrowing survivors of former terrestrial groups
in which only relict forms are now known. In the following families
only a part of the members have taken on a serpentine form and
become subterranean: Scincidae (Brachymeles and several other
genera), Zonuridae (Chamaesaura) , Anguidae (Ophisaurus) , and
Tejiidae (Scolecosaarus) . In certain other genera this process of
specialization is already begun.

Certain characters in which Amphisbaenia differ completely (or
partly) from Sauria are:

1. Mode of progression (progression is made largely in a straight
line by slight vertical undulations ) .

2. Loss or reduction of the right lung, rather than the left.

3. Absence of evidence in the skull of a pineal* or parietal eye.

4. Absence of skull arches.

5. Ambulation of body.

6. Absence of most of the dermal bones of the orbital ring.

7. Overlapping (telescoping) of bones.

8. Phalangeal formula when present 3-3-3-3-3 instead of the more
typical 2-3-4-5-3.

9. A greater development of chondral elements in the skull.

The known fossil history of the Amphisbaenia on the North Ameri-
can continent begins with the discovery of Ototriton solidus Loomis
in the early Eocene ( Wasatchian ) . Even in the Eocene it would
appear that the group had already undergone a long evolution.
Ototriton solidus is one of the largest species of the group judging
by the skull that measures 31.5 mm. in length and 23.5 mm. in

* Reported as present by Gilmore and Jepsen (1945) ; said to be absent by Zangerl (1944).
I suspect that the opening on the median line between the supraoccipitals and parietal of
Hyporhina should not be interpreted as that of a pineal eye.

524 The University Science Bulletin

width, a size that is reached by only a few species living today.
It already displays most of the expected characteristics of amphis-
baenids, as well as its own peculiar specializations. Another species
Ototriton minor is about half as large. Two other forms from the
Middle Eocene, based on vertebrae alone, have been referred to the
genus Lestophis, a genus of doubtful status.

Passing over the Upper Eocene where specimens are as yet un-
known, we find several species and genera present in the Oligocene.
Above the Oligocene another great gap in our knowledge occurs,
and no specimen is known from the Miocene, the Pliocene, or the
Pleistocene. In the Recent however five living forms, belonging to
two genera, Rhineura and Bipes, are known on the North American
continent and one other genus, Ampliisbaena, with four or more
species, occurs in the West Indies. Thus the probability is strong
that Amphisbaenids were present and will eventually be found as
fossils in the three periods that have as yet no known species.

Some time ago three Amphisbaenid specimens in the paleonto-
logical section of the University of Kansas Museum of Natural
History were placed in my hands for study by Dr. R. W. Wilson,
Associate Curator of Paleontology, University of Kansas, and Mr.
Edwin C. Galbreath of that Institution. These specimens were
found in the summer of 1946 by a party consisting of Richard Rinker,
Joao Moojen, Edwin C. Galbreath, Theodore Downs, Russell R.
Camp, and Henry Hildebrand, under the direction of Dr. Claude
W. Hibbard, while investigating Oligocene faunas of Logan County,
Colorado. The three specimens represent the firstf reported finds
of amphisbaenids in the Oreodon Zone of Logan County, Colorado.
A number of other remains taken at the same time pertain to reptiles
of other groups and these are to be reported at a later time.

Through the courtesy of the authorities of the University of Colo-
rado Museum, I have obtained a loan of an amphisbaenid skull
collected by Dr. R. W. Wilson in 1940 at the same locality as those
collected by the University of Kansas party in 1946.

Through the courtesy of Dr. Lewis B. Kellum and Dr. Claude
Hibbard of the University of Michigan Museum of Paleontology, I
have had available for study three Amphisbaenid skulls from Logan
County, collected by Dr. Hibbard and party in 1948. A single speci-
men of an Amphisbaenid skull from the Oligocene of Wyoming was
loaned by Dr. C. Lewis Gazin of the U. S. National Museum.

t One specimen consisting of a vertebra taken in 1873 from the Horsetail Creek Beds
(Titanotherium Beds, Horizon A) of Logan County, was described by E. D. Cope as Platyrachis
coloradoensis. Cope did not recognize the relationship of the specimen to the amphisbaenids
and it remained for Gilmore (1928) to transfer the form to the genus Rhineura.

Taylor: Oligocene Amphisbaenid Reptiles

525

Mr. Galbreath in 1948 acquired four other skulls in the Logan
County area which he presented to the University of Kansas. These
also have been available for study.

I am of the opinion that certain of the specimens represent un-
described species. One of these is being placed in the genus
Hyporhina Baur and certain ones are being placed in the genus
Rhineura. One is regarded as representing a new genus.

Of the genus Rhineura, aside from a living species in peninsular
Florida, four fossils forms are recognized: these are Rhineura
hatched Baur, Rhineura minuta Gilmore, Rhineura sternbergii
Walker, all based on adequate material. A fifth species Rhineura
coloradoensis (Cope) (based on Platyrachis coloradoensis Cope)
may be regarded as being of uncertain status; and until its vertebrae
are found associated with a skull, its relationship to the species here
described cannot be ascertained.

The following outline presents data on the geological and geo-
graphical distribution of North American amphisbaenids.

Known Geological Distribution of the Amphisbaenia in
Continental North America

'Rhineura floridaria (Baird) ,
Bipes biporus (Cope), Baja Calit,

Mex.
Bipes canaliculatus (Bonneterre),

Guerrero, Mex.
Bipes tridactylus (Duges),

Guerrero, Mex.
fBipes sp. Arizona

Recent (Living forms)*

Pleistocene (Unknown)
Pliocene (Unknown)
Miocene (Unknown)

r.

Oligocene

Late. Whitneyan, Upper Brule) Rhineura hatcheri Baur
(Leptauchenia Zone) ] Hyporhina antiquaTS&ur

Middle.

Orellan, Lower Brule<
(Oreodon Zone)

.Early. Chadronian (Titano-
therium Zone)

Rhineura sternbergii Walker
Rhineura minuta Gilmore
Rhineura hibbardi sp. nov.
Rhineura wilsoni sp. nov.
Rhineura amblyceps sp. nov.
Rhineura hatcheri Baur
Hyporhina galbreathi sp. nov.
Gilmoreia attenuatus sp. nov.

Rhineura coloradoensis (Cope)

* These species were originally described as Lepidosternon floridana, Bipes canaliculatus,
Euchirotes biporus and Hemichirotes tridactylum. Another species probably occurs, (see
James in Long's Expedition to the Rocky Mountains, vol. 1, 1823, p. 484; Taylor, Copeia,
1938, no. 4, Dec. 10, p. 202.)

526

The University Science Bulletin

Eocene

-

Late. (Unknown)

Middle. Bridgerian, Bridger,
Black's Fork
member

Wasatchian, Wind
River, Lost Cabin
member

Early.

Wasatchian, Wind
River, Lysite
member

'Lestophis anceps (Marsh) Uinta

Co., Wyo.
Lestophis crassus (Marsh) Uinta
. Co., Wyo.

Ototriton minor Gilmore and
Jepsen, Natrona Co., Wyo.

Ototriton solidus Loomis,
Fremont Co., Wyo.

Geographical Distribution of Oligocene Amphisbaenidae

Whitneyan

(Leptau- Battle Draw Springs,

chenia Washington Co., S. D.

Zone)

Sioux Co., Neb.

Washington Co., S. D.
Orellan

(Oredon J Douglas, Converse Co., Wyo.
Zone)

Logan Co., Colo.

i

Chadronian
(Titano-
therium
Zone)

Logan Co., Colo.

Rhineura hatcheri Baur
Hyporhina antiqua Baur

Rhineura hatcheri Baur

Rhineura hatcheri Baur

Rhineura sternbergii Walker
Rhineura minuta Gilmore
Gilmoreia attenuatus sp. nov.

'Hyporhina galbreathi sp. nov.

Rhineura hibbardi sp. nov.
■s Rhineura amblyceps sp. nov.

Rhineura wilsoni sp. nov.
.Rhineura hatcheri Baur

Rhineura coloradoensis Cope

In the preparation of this paper I am under deep obligation to
Dr. Robert Wilson and Mr. Edwin Galbreath for assistance with the
stratigraphic nomenclature, and for reading and criticizing the
manuscript.

Dr. Byron Leonard has kindly prepared the photographs of the
skulls reproduced here. For this I am indeed grateful.

SYSTEMATIC TREATMENT

A total of fourteen skulls have been available in this study. They
vary considerably in completeness and state of preservation. All
but one of these are from the Orellan (Oreodon Zone) of Logan
County, Colorado. This single species is from the Orellan of Con-
verse County, Wyoming, likewise the type locality of Rhineura

Taylor: Oligocene Amphisbaenid Reptiles

527

minuta and Rhineura sternbergii, and is here described as a new
genus and species.

Key to Symbols Used in the Text Figures

BO

Basioccipital

OS

Orbitosphenoid

BS

Basiphenoid

OT

Otic

C

Coronoid

P

Parietal

CB

Corpora bigemina

PA(L)

Palatine

CH

Cerebral hemisphere

PF

Prefrontal

D

Dentary

PM

Premaxilla

EC

Extracolumella

PO

Paroccipital (in Rhineura)

EO

Exoccipiial

PO

Postcrbital (in Hyporhina)

EP

Epipterygoid

PT

Pterygoid

F

Frontal

PV(V)

Prevomer

FM

Foramen magnum

Q

Quadrate

FO

Fenestra ovale

S

Stapes

M

Maxillary

SA

Surangular (Supraangular)

N

Nasal

sec

Semicircular canal

0

Orbit.

so

Supraoccipital

oco

Occipital condyle

SQ

Squamosal

OLL

Olfactory lobe

Gilmoreia genus novum
A genus of fossil Amphisbaenid reptile characterized by a shallow
or absent orbital vacuity, not overhung by the frontal bone; elevated
portion of skull much narrowed, its surface smooth, lacking sculp-
ture. Section of skull posterior to its highest point much shortened;
no postfrontal; dental characters unknown. Type species Gilmoreia
attenuatus,

Gilmoreia attenuatus sp. nov.

Plate LVIII, figs. 3, 4, 5; text fig. 1, a, b, c.

Type.— United States National Museum, No. 16308 (field No. 40),
consisting of a skull, the anterior part, from about the middle of
the frontals, missing. The quadrate, extracolumella, pterygoid,
palatine, prevomer, and entire lower jaw missing.

Type locality. — Eight miles east of Douglas, Converse County,
Wyoming. G. F. Sternberg, collector, 1931.

Horizon. — Oligocene, Brule.

Diagnosis. — The deflected portion of the skull is much longer
than the remainder. The length from the anterior edge of the fora-
men magnum to the highest elevation of the skull is less than the
median length of deflected portion of the parietal. The skull is at-
tenuated anteriorly, its narrowest width being considerably anterior,
not posterior, to the point of highest elevation. The diamenter of
the brain cavity at the anterior level of the basioccipito-sphenoid

528

The University Science Bulletin

bone, is proportionally much larger than in Hyporhina or Rhinenra.
The outline of the squamosal is completely obliterated, the point
of articulation for the quadrate, however, strongly projecting. The
bones of the facial portion of the skull lacking sculpturing.

oco

Fig. 1. Gilmoreia attenu-atus gen. ct sp. nov. Type, U. S. Nat. Mus. No.
16308 (original No. 40). Oligocene, Orellan (Brule), 8 m, N Douglas, Converse
Co., Wyoming. G. F. Sternberg, collector, 1931. A. Lateral view; B. Dorsal
view; C. Ventral view. Actual length of fossil 7.5 mm.

Description of type. — The greatest width of the moderately in-
flated posterior part of the otic region of the skull (4.5 mm.) is
slightly greater than the length of the skull from the posterior edge
of the condyles to the level of the highest point (4.25 mm.); the
foramen magnum opens dorsally or, perhaps better, postero-dorsally.

The frontal bones are largely missing; their posterior ends, par-
tially separated by a tongue from the parietal, are narrow, their

Taylor: Oligocene Amphisbaenid Reptiles 529

length nearly twice their width. The width of the frontal at the
farthest (lateral) extension of the parietal is about 1.1 mm. It
is presumed that this point is posterior to the middle of the total
length of the frontal and that there is a very slight widening on the
anterior terminal portions of the skull as is typical of certain living
genera (Amphisbaena, Bipes).

The parietal is large, somewhat saddle-shaped and covers the
greater part of the posterior dorsal surface of the skull, overlapping
very considerably the deeper bones of the posterior part of the
brain case, leaving only a very narrow rim of the supraoccipital
and the posterior parts of the otic region bordering the foramen
magnum. The foramen magnum tends to notch the parietal, the
posterior parts appearing as two broadly rounded lobes. On the
median line a short sagittal ridge arises from the main bone, projects
back to dovetail on the median line with a very narrow median
crest of the supraoccipital. The parietal also covers the greater
portion of the lateral surface of the brain case terminating anteriorly
in a point. This forward extension is visible along the elevated dor-
sal surface as a very narrow line, bordering the outer edge of the
frontal ( broken away on left side ) .

The supraoccipital, exoccipital, basioccipitals, and the otic capsule
are fused into a single element, the former sutures now being com-
pletely obliterated. In the otic regions the two dorsal semicircular
canals of the ear are clearly visible as two slightly elevated whitish
ridges which meet nearly at a right angle at a point lateral to the
foramen magnum where it is widest. The third canal is partly in-
dicated on the posterior face of the skull.

The fenestra ovale is irregularly oval, and the stapes is present on
the left side, though sunken more deeply than is normal. The colu-
mellar part is largely missing, only the basal portion remaining.

Overhanging the anterior edge of the fenestra ovale is the facet
for the articulation of the quadrate. Presumably the squamosal is
completely fused with the outer part of the otic region. In most
cases the fusions in the skull are of chondral bone to chondral bone;
here it would appear that the fusion is of dermal bone to chondral
bone if the squamosal has even been present.

A short distance in front of the fenestra ovale is the longitudinally
oval opening along the side of the brain case characteristic of the
amphisbaenids, having an area exceeding that of the fenestra ovale;
its upper posterior border is formed by the fused prootic. Beginning
near the middle of the upper border is the orbitosphenoid that runs

34-3286

530 The University Science Bulletin

forward forming a suture with the basal part of the brain case, the
fused basisphenoid ( parasphenoid ) . The anterior extension of the
orbitosphenoid is broken away. A small foramen is evident in the
bone and two canals can be discerned radiating from it. It would
appear that the orbital vacuity is absent or if present is very shallow
and has been moved far forward in comparison with that of Rhi-
neura.

The basioccipital-exoccipital region bears a transverse dumbbell-
like condyle, the two outer knobs being much wider than the median
portion. It is presumed that both exoccipitals and basioccipital
bones contribute to the condyle. Seen directly from behind the
condyle occupies a position about mid-distant between top and
bottom and its transverse length equals one third of the greatest
width of the brain case. Directly lateral to the condyle knobs and
closely approximating them are large nerve foramina.

Seen from the ventral surface, anterior to the condyles are two
roughened elevated areas, the outer edges of which lie just below
the fenestra ovale. Presumably they mark the original positions of
the paroccipital* bones that are now missing. Somewhat farther
forward, on the outer edge of the bone are notches marking point of
contact with the posterior end of the missing pterygoid. Just an-
terior to the level of the notches the bone has been broken, but
anterior to the break it continues forward, having the appearance of
a spearhead. Toward its anterior end a saggittal ridge develops.
The anterior part of the bone is absent.

Measurements in mm. — Actual length of fossil, 7.5; estimated
skull length, 10; greatest width of brain case (otic region), 4.5;
narrowest width at level of posterior point of frontal, 1.5. From
highest elevation to foramen magnum, 2.7; from highest point to
end of condyle, 4.3; from highest point to the anterior end of parietal,
4.15; to posterior end of parietal 4.1.

Remarks. — The aberrant character of the species Rhineura minuta
described by Gilmore (Proc. U. S. Nat. Mus., vol. 86, 1938, pp. 12-
14, fig. 1) strongly suggests that the relationships might better be
shown by regarding it in a genus apart from Rhineura. The char-
acters which appear to separate the species from Rhineura are those
in which Rhineura differs from Gilmoreia. These are the absence
of a wide flaring deflected facial portion, the lesser angle of de-
flection, the narrow, sharp-edged elevation on the dorsal surface
lacking an overhanging edge, the smooth character of the anterior

* In the figure, the legend "PO" is not connected with the paroccipital.

Taylor: Oligocene Amphisbaenid Reptiles 531

dorsal part of the skull. In consequence there is a probability that
minuta may eventually prove to be a member of the genus Gil-
moreia.

The type locality of G. attenuatus is also the type locality of two
other species of amphisbaenids, Rhineura minuta Gilmore and
Rhincura stcrnbergii Walker. Since my skull is incomplete and the
other two practically complete, many points of comparison are lack-
ing. Based on the skull characters given by Gilmore and the ac-
companying figures, Gilmoreia attenuatus differs from Rhineura
minuta in being large, estimated length, 10 mm. (7.8 mm.). The
greatest width of the brain case 4.5 mm. ( 3.3 mm. ) ; width at orbits
1.5 mm. (?) (2.9 mm.); least width 1.5 mm. (1.6 mm.). (The
measurements of minuta in parenthesis. ) In G. attenuatus the skull
seen in lateral profile has the posterior part of the brain case nearly
horizontal then it suddenly rises to the highest point then slopes
gently (not curves) downward. In profile the dorsal outline of
minuta seems to curve from the foramen magnum to snout tip, not
reaching or forming an angular elevation. The transverse width of
the condyle is equal to one third of the total width of the brain case.
In R. minuta neither the size of the condyle nor its position are given
in the description. However, the figure shows the transverse width
of the condyle to be one half of the width of the skull and situated
lower than in attenuatus.

Characters on which the two species agree are, the smooth surface
of dorsal elevated part of skull; the narrowness of the elevated skull
part; and absence of the overhanging wings of the parietal and
frontal.

Compared with Rhineura sternhergii, the angle of defllection is
less sharp and the depth of the skull at the highest point is seemingly
less. The length of the posterior part of skull (highest point to end
of condyle) is 4.3 mm.; from highest point to anterior extension of
the parietal, 4. 15 mm.; while in stcrnbergii the distances from high-
est point to condyle end is practically double the extension of the
parietal from the highest point forward.

Genus Hyporhina Baur

Hyporhina Baur, American Naturalist, vol. 27, 1893, pp. 998-999.

Type species. — Hyporhina antiqua Baur.

This genus has been established on the following characters:

A postorbital arch formed by a postorbital bone; prefrontal small

excluded from the boundary of orbit by the frontal and maxillary;

nostril ventral to snout; four acrodont teeth in each maxillary; pre-

532 The University Science Bulletin

maxillary, nasals and frontals nearly meeting at a point near tip of
snout.

A specimen of a small amphisbaenid from the Middle Oligocene
of Logan County, Colorado, belongs in this genus. It bears some
points of resemblance to, and numerous differences from, Hypor-
hina antiqua from the Middle Oligocene, Washington County, South
Dakota. There is a postorbital arch closing the orbit of the eye;
and the frontals, nasals, and prefrontals almost meet at a point near
the tip of the snout. The contour of the skulls are in general sim-
ilar; however, the angle of deflection in the Colorado skull is some-
what less than in Hyporhina antiqua. The two most significant
differences are that the prefrontal bone is much larger than in
Hyporhina antiqua forming at least one third of the border of the
orbit, and there are six pleurodont maxillary teeth on each side,
instead of four "acrodont" teeth. The original description does not
mention this dental character but Charles W. Gilmore who studied
the type says: "The teeth appear to be acrodont."

If the teeth of Hyporhina are indeed acrodont one may doubt
the wisdom of considering this form with pleurodont teeth as be-
longing to the same genus. Such a dental character is considered
very significant, serving in Lacertilia to distinguish the two lizard
families Agamidae and Iguanidae from each other. Until the char-
acter of the teeth of Hyporhina antiqua can be established as acro-
dont beyond question, I shall consider the present species as con-
generic with Hyporhina antiqua.

Hyporhina galbreathi sp. nov.

Plate LVIII, figs. 6, 7, 8; Plate LIX, figs. 4, 5; text figs. 2, 3

Type. — University of Kansas Museum of Natural History, Paleo.
No. 8221, consisting of a skull, complete save for the absence of the
quadrate and a part of the squamosal. The lower jaw is absent.
Collected under the direction of Dr. Claude W. Hibbard, University
of Kansas Museum of Natural History Expedition of 1946.

Type locality. — Clyde Ward Ranch, Logan County, Colorado,
July 31, 1946.

Horizon. — Middle Oligocene (Orellan), White River Formation.

Diagnosis. — A species, related to Hyporhina antiqua Baur, but
differing from this form in having two spinelike lateral processes
on the anterior part of the parietal (not protruding); in having a
much larger prefrontal forming more than one third of the border
of the orbit, rather than a small one completely separated from the

Taylor: Oligocene Amphisbaenid Reptiles

533

orbit; six maxillary teeth present that are definitely pleurodont,
rather than four that are regarded as acrodont.

Description of the type. — The skull is diminutive (8.5 mm. long),
the frontal portion strongly deflected and wider than the posterior
part of the braincase. The premaxilla seen from above has a curved
surface, subtriangular in shape, about a third greater in its trans-
verse width than in its length. Posteriorly it is in contact with the

oco

_ pM

-N
-M

■F

-PF
-0

PO

-SQ
FM

_. OCO

Fig. 2. Hyporhina galbreathi sp. nov. Type, Kansas U. Mus. Nat. Hist.,
Paleontology No. 8221. Oligocene, Orellan, Clyde Ward Ranch, Logan Co.,
Colorado. Dr. Claude Hibbard and party, collectors, July 31, 1946. A. Lateral
view; B. Dorsal view; C. Ventral view. Actual length of fossil 8.5 mm.

nasals. When seen from below, it presents a flattened surface
external to the mouth, with two rather shallow diagonal grooves.
When seen laterally the snout is rather wedge-shaped. Latero-
posteriorly the element is bordered by the external nares, and its
edge is notched on each side by the tip of the maxillary process,

534 The University Science Bulletin

which forms the inner rim of the naris. Medially, nearly in a line
between the nares but at a different level, is a single tooth arising
from the premaxilla within the edge of the buccal cavity. Behind
this the premaxilla sends out two divergent processes that pass
along the sides of the anterior part of the maxillary tooth rows on
a level with the bases of the teeth, and are separated by the anterior
part of the pre vomers.

On the dorsal surface of the skull the nasals are narrowly in
contact mesially, from which point they diverge and pass forward
narrowing greatly, then widening somewhat as they turn outward
and under the edge of the snout to form the anterior border of the
nares. Small foramina are present on the naso-frontal border as
well as on the naso-premaxillary border. The paired frontals are
large, about twice as long as wide, separated narrowly from the
premaxillary, and in contact with each other along a straight median
suture for more than half their length. Posteriorly the bones have
a lateral process that extends downward to and forms a part of the
posterodorsal rim of the orbit. Seemingly they are in contact with
the maxillary by a very narrow pointed process posterior to the
orbit. The posterior parts of the frontals are heavily sculptured
with pits, foramina, and shallow grooves, while the anterior part
is shallowly and indistinctly sculptured. The maxillary bone is
longer than wide, forming the lower part of the orbit. It folds
under the lateral edge of the snout to the alveolar edge and forms
a considerable area on the underside of the skull. The teeth are
pleurodont in character and are set well back under the bone.
There are six maxillary teeth, the largest being the second from the
front.

The prefrontal is a small element lying somewhat above and
in front of the orbit, and forming more than one third of the
border of the orbit. The postfrontal is a slender bone inclosing
the orbit and fitting into a notch in the parietal, dorsally, and into
a similar notch in the maxillary, ventrally.

The single parietal is an extensive saddle-shaped bone, the an-
terior triangular part being elevated, and deflected downward
slightly. It pushes in between the posterior parts of the frontals.
A spinous process is present on each side at a level with the
posterior edge of frontal, touching the postorbital bone and border-
ing the posterior edge of the frontal. At the highest part of the
skull there are two symetrical, roughened depressions on the parietal
narrowly separated medially, which presumably serve for muscle

Taylor: Oligocene Amphisbaenid Reptiles 535

attachments. Posteriorly the parietal sends back two broad pro-
cesses which diverge and reach rather close to the posterior level
of the condyle. Laterally the parietal is in contact with the otic
region, the squamosal and the orbitosphenoid.*

The supraoccipital seemingly is fused with the exoccipitals the
basioccipital, and the bones of the otic capsule, so that none of
the individual elements are discrete. The combined element forms
the border of the foramen magnum, becoming thicker and more
massive on the sides. The posterior rim of the foramen bears the
occipital condyle, on each side of which is an articular knob. The
exoccipital area of the combined bone is thick and massive, with
a forward directed part that is closely joined to the squamosal and
lies above the ear opening. The squamosal appears as an "inlay"
bone on top of the otic region and is wedged in between the posterior
part of the parietal and the orbitosphenoid. A portion of the com-
bined occipital bone (prootic?) appears directly in front of the
squamosal.

The orbitosphenoid is a large bone, reaching forward from the
squamosal to the anterior edge of the orbital vacuity and forming
the cranial wall on the side of the skull. On the floor of the orbital
vacuity it is in contact with a section of the frontal.

The stapes covering the fenestra ovalis is oval in shape with a
cylindrical columella extending outward and forward. There is
no trace apparent of an extracolumellar element, although one may
have been present originally.

No trace of the division between the exoccipitals and basioc-
cipital is evident. The latter bone slants forward and downward.
The suture between this element and the basisphenoid (para-
sphenoid of Zangerl) is faintly evident at a point 1.8 mm. anterior
to the median depression between the condyles. The basisphenoid
is notched laterally at the point where its widest point narrows
suddenly to half its width and proceeds forward, the sides converg-
ing anteriorly. Here it contacts (passes above) the vomers which
extend forward to the median point of the premaxillaries. Two
roughened areas on either side of the posterior part of the basi-
sphenoid may represent the position of the missing paroccipitals
( "element X" of Zangerl ) .

The prevomers form a median suture anteriorly for about 1.15
mm. then diverge, the two branches bordering the anterior part of

* The nomenclature of this element may be questioned. Gilmore calls it the alisphenoid
but it appears to be more properly the orbitosphenoid, or a combined element. However, there
is no evidence that an optic foramen is present.

536 The University Science Bulletin

the parasphenoid ( presphenoid? ) , which is slightly deflected down-
ward and has a median ridge. The prevomers also contact the an-
terior part of the pterygoids and the palatines laterally. The ptery-
goids are narrow anteriorly, and extend back, passing to some
distance behind the palatines, where each sends np a small lateral
process that reaches almost to the stapes. External to the palatines
the pterygoids contact the ectopterygoid. The palatines occupy the
depressed area between the maxilla and the ectopterygoid on the
outside, and the vomerine process and the pterygoid on the inside.

Measurements in mm. — Greatest length, 8.5; greatest width of
braincase, 4; greatest width of deflected frontal area, 4.4; narrowest
skull width, 2.8; greatest elevation (without lower jaw), 3.8.

Remarks. — As already stated, this specimen differs from the
generic description in having more than four teeth on the maxillary,
and in having the prefrontal forming a large part of the boundary
of the orbit (more than one third). In appearance there is such
a striking general similarity in form that I am inclined to place
the species in Hyporhina at least for the present.

It is generally accepted that the Oreodon (Orellan) beds of the
White River formation in northeastern Colorado are older than the
Leptauchenia (Whitneyan) beds of South Dakota. It would then
appear that this species is older than Hyporhina antiqua (if the
latter is correctly allocated in time) but it is not at all impossible
that the two may be found to occur together since the time separa-
tion of the two zones is seemingly not great.

The presumed differences in the character of the teeth of this
species and Hyporhina antiqua would suggest that a much wider
separation between the forms exists than that between species. The
living amphisbaenid genera Trogonophis, Pachycalamus, and Agam-
odon have acrodont teeth. All are African in distribution. Other
living forms presumably have pleurodont teeth.

Referred Material

Among the fossils collected bv Mr. Edwin C. Galbreath in the
Oligocene of Logan County, Colorado, in 1948, are two other speci-
mens that are here regarded as referable to this species: K. U. M.
N. H. P. No. 8222 and No. 8219.

No. 8222: This specimen bears the following data: SWJ-t of Sec.
12, T. UN. R54W., Logan County by E. C. Galbreath (field No.
5850). It consists of a skull that retains only a small part of the
deflected facial portion, and the greater part of the right side of

Taylor: Oligocene Amphisbaenid Reptiles 537

the braincase. The front and outer part has been fragmented re-
cently as the broken edges show but little wear. Of the parts pres-
ent the following characteristics are discernible ( Plate LIX, fig. 4 ) :

Frontals. — Only the posterior parts of these elements are present
but these are considerable larger than the deflected facial part of
the parietal, partially wedged between them.

The surface sculpturing is similar to the type but differs some-
what in details. On the sides there are two or three grooves
terminating in deep pits. No fragments of the prefrontals are in
evidence but a portion of the orbital vacuity is overhung by the
frontal. This edge has a small rounded notch for contact with the
prefrontal. An examination of this area discloses no foramen that
might serve as a passageway for an optic nerve.

Parietal. — The parts of the parietal that remain, and particularly
the posterior extensions on each side of the foramen magnum, re-
semble the condition in the type. However, on the right side a
small thin squamosal element, lying above the otic-occipital region,
is larger than a similar element in the type. Abutting against the
lower posterior extension of this element is a quadrate in normal,
vertical, functioning position. Immediately posterior to the quad-
rate is the "stapes" with a broad irregularly oval base covering th'i
fenestra ovalis, and a slender columella rising from it to nearly two
thirds of the greatest length of the base. The outer termination of
the columella is somewhat expanded and may have supported an
extra-columella such as obtains in certain living, as well as in cer-
tain extinct, forms. On the median line of the parietal at or near
the point at which the bone contacts the supraoccipital there is a
rather large perforation,* a character evident or suggested in some
other fossil skulls, as well as in skulls of living forms.

The basal portion of the skull, from the basioccipital to a point
where the anterior extension of the cranial cavity is exposed in a
fracture, is a single element composed of (presumably) the basi-
occipital, basisphenoid and perhaps also a presphenoid. There
is no evidence of a break in its continuity. The superficial mem-
brane bones of the palatine region normally present ( the prevomers,
pterygoids, ectopterygoids, and palatines) are absent, leaving a
broad smooth chondral bone terminating anteriorly in three points,
of which the median is on a different plane than the other two.
It forms a suture with the frontals. Laterally for a considerable
portion of its length, the bone forms a suture ( not an overlapping )

*0f this opening Rainer Zangerl (1914) says, "space left by removal of cartilaginous part
of the anterior supraoccipital prong."

538

The University Science Bulletin

with the orbitosphenoid. Posterior to the frontal a fragment of the
upper part of the postorbital is discernible.

The association of this incomplete skull with the species H. gal-
breathi can scarcely be questioned. Measurements of equivalent
parts differ only by fractions of a millimeter. The one exception,
already mentioned, is the greater distinctness and apparentlv
larger size of the squamosal. This may be due to a partial fusion in
the type of a part of this bone to the underlying chondral elements.

---PT

B

Fig. 3. Hyporhina galbreathi sp. nov. (referred material showing part of a
cast of the braincase) Kansas U. Mus. Nat. Hist., Paleontology No. 8219. Ohg-
ocene, Orellan, Clyde Ward Ranch (SE*4 SW}4 Sec. 12, f. UN, R. 54W),
Logan Co., Colorado. Edwin C. Galbreath, collector. A. Dorsal view; B. Ven-
tral view. Actual length 8.4 mm.

As a result of this the terminal portion of the parietal on either side
of the dorsal foramen magnum differs somewhat from the type in
contour.

No. 8219: This fragmentary skull taken from the same locality
as the previous specimen, is associated with Hyporhina galbreathi
on the basis of structural similarity of the few elements of the brain-
case that still remain.

The vomero-palatine region is similar to that of the type save that
the posterior palatal prong of the premaxillary is a little longer and
slightly more slender. It bears the same general relationship to the
premaxillary and maxillary fragment. On the dorsal surface, the

Taylor: Oligocene Amphisbaenid Reptiles 539

close approximation of the frontals, nasals, and premaxillary to a
common medial point obtains, however the premaxillary actually is
in contact with the frontals, thus separating the nasals narrowly.
A fragment of the postorbital definitely indicates its generic as well
as its specific relationship. A bone chip from the dorsal surface,
comprising the anterior part of the parietal and the posterior part
of the frontals, shows marked similarity of this portion of the skull
to this region in the type specimen.

Probably the most significant feature of this specimen is that, due
to the removal of much of the roof of the skull, a considerable por-
tion of a cast of the brain is exposed.

In the mesencephalic region a pair of rounded oval corpora
bigemina may be distinguished, flanked on either side by the orbital
vacuities. This is at the narrowest point of the skull. Immediately
following this is a massive area the anterior part of which is trilo-
bulate, and separated from the mesencephalon by sulci. The two
outer lobules are relatively narrow; the median broad. Posteriorly,
across the middle of this region, there is no trace of lobulation or
sulci. Immediately in front of the point of entrance of the foramen
magnum there is a broad entrant superficial groove, which lies be-
low a dorsal perforation of the brain case situated on the median line
between the parietal and the supraoccipital bones. From this de-
pression a slight epiphysis seems to be present but actually this may
be an unnatural accretion of some sort. There is no apparent divi-
sion into metencephalic and myelencephalic parts.

Anterior to the mesencephalon the cerebral lobes push posteriorly
and laterally, terminating immediately in front of the orbital vacuity.
This widening of the brain in this region suggests that the telen-
cephalon has somewhat the shape of an arrowhead with the dien-
cephalon and part of the mesencephalon pushing in between the
lobes. (Plate LIX, fig. 5).

Genus Rhineura Cope

Rhineura Cope, Proc. Acad. Nat. Sci. Phila. 1861, p. 75.

Type species. — Rhineura floridana (Baird).

One living species and six fossil species are recognized.

Rhineura hibbardi sp. nov.

Plate LX, figs. 1, '1, 3; text rig. 4 A. B. C.

Type. — University of Michigan, Museum of Paleontology, no.
25431. Skull with lower jaws; bones on anterior facial portion of
skull fragmentary; condyle lacking. Collected by Dr. Claude W.
Hibbard and party, August 5, 1947.

540

The University Science Bulletin

Type locality. — Smith Ranch, Logan County, Colorado. Center
of Sec. 7, T.11N— R.53W.

Horizon. — Middle Oligocene, White River Formation, Oreodon
Zone.

PM

_0

_M

- EP
-PA

_BS

-PT
.SA

-_BO

Fig. 4. Rhmeura hibbardi sp. nov. Type. Univ. Michigan Mus. Paleo. No.
25431. Oligocene, Orel Ian, Smith Ranch (center sec. 7, T. UN, R. 53W),
Logan Co., Colorado. Dr. Claude Hibbard and party, collectors, Aug. 5, 1947!
A. Lateral view; B. Dorsal view; C. Ventral view. Actual length of' fossil
1G.6 mm.

Taylor: Oligocene Amphisbaenid Reptiles 541

Diagnosis. — A large member of the genus Rhineura, the skull
length reaching approximately 20 mm.; the width of the brain case
in the otic region 8 mm.; seven teeth in maxillary, the second largest
and somewhat caninelike; seven teeth in dentary, the fourth and
first being largest.

Description of the type skull. — The skull at present measures
16.6 mm. from the nasal region to the back of the otic capsule area.
The anterior part of the nasals has been largely chipped away, as
is true also of most of the premaxilla and the upper parts of the
maxillaries; the mold of the brain case appears partially exposed
in several places. The olfactory lobes running forward are par-
tially separated by the deeper fragments of the posterior part of
the premaxilla, whose suture with the nasals is clearly indicated;
the internasal suture can be discerned beginning from the pre-
maxillary, but it can be followed posteriorly for only a short distance.
Most of the dorsal surface of the nasals and frontals is missing; how-
ever, a few fragments of the frontals display the rugosities and
sculpturing typical of the genus Rhineura. The outer posterior
edge of the frontal overhangs the orbital vacuity for a maximum
width of slightly more than two millimeters. The parietal is a
saddle-shaped bone, bearing a median sagittal ridge that begins
just in front of the foramen magnum, and runs forward and upward
for 7 mm. Here the parietal surface becomes suddenly elevated
forming a roughened triangular area. Forward of this area the
facial part of the skull is deflected abruptly downward. The area
of the parietal on this deflected part of the skull consists of a pair of
lateral diverging prongs overhanging the orbital vacuities and a
forward median extension whose extent cannot be clearly deter-
mined. Anterior to the otic capsules the whole brain case is con-
stricted, the parietal extending far down on the sides where it is
bordered, anteriorly by the orbitosphenoid and posteriorly by a
portion of the occipito-otic complex, that probably corresponds to
the prootic. Posteriorly the parietal flairs out but the boundaries
with the supraoccipital behind are indistinct and partially broken.
For a considerable distance (perhaps half of its entire length)
it overlays the bones of the occipital-otic complex. This complex
is composed of the supraoccipital, basioccipital, exoccipitals and
otic capsules fused seemingly into a single entity, the sutures be-
tween the elements being no longer distinguishable. The squa-
mosal is fused solidly to the outer border of the capsule forming
a thick overhanging shelf with which the quadrate articulates. The

542 The University Science Bulletin

quadrate is present but has slipped downward exposing its own
articular surface. The anterior face of the quadrate rises consider-
ably higher than the articular face, and the whole upper part
is normally exposed as a somewhat rounded knob. Covering part
of the lower portion of the quadrate is a biscuit-shaped bone, the
extracolumella. This bone, seemingly, is not attached; that is,
it forms no sutures but is normally held by a ligament from the
columella. The fenestra ovalis is large, measuring, on one side of
the skull, 2.4 mm. in length, and 2.1 mm. high. The stapes with
the columella is missing entirely on the right side, while a portion of
the element is present on the left side.

The lower outer face of the maxillaries are more or less complete
while the upper surface is chipped away. Five teeth are actually
present on the right side with a space between the second and
third, and presumably a space following the last, making a total
of seven. On the left side there are seven teeth, the first being
broken at the base. The second of both series is much enlarged
and caninelike. Posteriorly the maxillary joins the ectopterygoid
but the actual extent of the suture cannot be traced since the bone
itself disappears behind the coronoid. The preoculars lie in front of
the orbit and make an undulating suture with the maxillary. They
border the whole anterior rim of the orbital vacuities.

Both dentaries are present, each bearing seven teeth of which
the fourth is largest. The first tooth of the series is larger than
the remaining five. The dentary joins a very large coronoid, which
has its beginning anteriorly at the level of the sixth tooth of the
dentary series. The greatest length of the coronoid element is
more than double its greatest height above the dentary. The den-
tary and coronoid join the surangular posteriorly, but the details
of the sutures are not wholly clear. The surangular bears a notch
near its upper center into which the quadrate articulates, while the
posterior part of the element bends sharply downward and tends
to cover partially the fenestra ovalis.

The basal part of the brain case consists of the basioccipital fused
laterally with the exoccipitals, and paroccipitals, and anteriorly
with the basisphenoid. The condylar protuberance, save for a small
fragment on the left side, is missing. Seen from the ventral surface
there is a smooth, relatively deep depression between the points of
fusion of the paroccipitals from which point the forward extension
of the basal part of the combined element bends downward. A
median ridge is evident on the median line; bordering the sides

Taylor: Oligocene Amphisbaenid Reptiles 543

of the palate (or anterior part of the ventral surface of the brain
case) are the pterygoids, which are strongly elevated rather than
flat as is the expected position of these bones. Anteriorly they join
with the ectopterygoid, and posteriorly they abut against a notch in
the side of the basioccipital. Palatines are present as are also the
prevomers, but this area of the skull is covered with a hard matrix
so most of the significant details cannot be given here.

Measurements in mm.— Estimated length 20. Actual length 16.6
(condyles and tip of snout missing); greatest width of brain case
8.1; least width 4; notch of foramen magnum to point of greatest
elevation 7.7; greatest depth of skull and jaws 7.8.

Remarks. — The lateral view of the photograph of the type skull
(plate LX, fig. 1) shows the fenestra ovalis, the orbital vacuity,
and a point between them intensely black. This is deep shadow
only, the skull being intact at these points.

Rhineura amblyceps sp. now

Plate LIX, fig. 1; Plate LXI. figs. 1, 2, 3. 4, 5; Plate LXII, figs. 1, 2, 3, 4, 5;
text figs. 5 A. B. C, 6 A. B. C.

T ?//><? .—University of Kansas Museum of Natural History ( Paleo. )
No. 7649, consisting of the basal portion of a skull, the anterior
facial portion of which, from about the middle of the frontals be-
ing entirely absent.

Type locality. — Clyde Ward Ranch, Logan County, Colorado,
July 29, 1946. *

Horizon. — Middle Oligocene, Orellan, Oreodon Zone.

Diagnosis. — A moderately large species of Rhineura, with an
estimated total skull length of approximately 18 mm., and the
greatest width of the brain case of about 7.7 mm. The orbitosphe-
noid is separated from the squamosal by a distance more than one
half of its length.

Description of the type. — The deflected facial portion of the skull
consisting of the posterior parts of the frontals and the anterior
part of the parietal, stands at an angle with the dorsal axis of the
skull. The sculpturing of the surface is extremely roughened, there
being deep pits, grooves, rounded knobs and irregular vermiform
elevations. The frontals, incomplete anteriorly, become narrowed
somewhat posteriorly, terminating in blunt points. They are sepa-
rated posteriorly by a triangular forward prolongation of the parie-
tal; the suture between the frontals and parietal being very irregu-
lar. The two lateral wings of the anterior part of the parietal form
a sinuous suture while the medial part is somewhat dovetailed.

544

The University Science Bulletin

Only after submersion in a clearing fluid can the complete suture
be discerned. The portion of the common median frontal suture
remaining, measures 3 mm. in length while the greatest length of
the remaining parts of the frontal is 4.3 mm. There is no part of
the orbit indicated, but the inner part of the orbital vacuity is
overhung by the remaining edge of the frontals and the anterior
tip of the parietal.

oco

— oco

Fig. 5. Rhineura amblyceps sp. nov. Type. Kansas Univ. Mus. Nat. Hist.
(Paleo.) No. 7469. Oligocene, Orellan, Clyde Ward Ranch, Logan Co., Col-
orado. Dr. Claude Hibbard and party, collectors, 1946. A. Lateral view; B.
Ventral view; C. Dorsal view. Actual length of specimen 12.5 mm.

Taylor: Oligocene Amphisbaenid Reptiles 545

The parietal is saddle-shaped, flaring broadly posteriorly, bending
low on the sides of skull, and reaching forward to the orbital vacuity.
The dorsal anterior part is elevated forming an area, triangular in
shape with three forward projecting prongs, the two outer are
longest and the median widest but shorter than the outer.

There is a median longitudinal ridge beginning near the foramen
magnum and running forward to the deflected facial portion of the
skull varying in its elevation. In the lower posterior part of the
triangular elevated area there are two symmetrical, oval, slightly de-
pressed areas with somewhat roughened surfaces (presumably for
attachment of muscles to hold the head at an angle ) . The posterior
extension of the parietal is missing, having been chipped away but
there is a depression following the posterior rim of the supraoccipital
that seemingly marks its original contours. These form two rounded
extensions, separated posteriorly by a median projection from the
supraoccipital which dove-tails with the parietal about two milli-
meters anterior to the forward edge of the foramen magnum. Else-
where the parietal overlies the combined otic-occipitals, the extent
of the overlap at places equaling half the length of the parietal
(perhaps more).

The posterior chondrocranial bones of the brain case have fused
into a continuous bony complex, composed of the supraoccipital,
exoccipitals, basioccipital together with the elements of the otic
capsule. On the ventral surface of the brain case the basisphenoid
is likewise fused to the combined bone and possibly also a pre-
sphenoid (it is presumed that these elements are all distinct in the
embryo or young animals). The limits of these various elements
is only to be conjectured (save for a remnant of a suture between
basioccipital and basisphenoid ) . ( The references to area and region
are based on probable positions of the various elements. ) The otic
areas are very large and dorsally are surmounted by a thin scalelike
bone partially fused, that I regard as the squamosal. The outer
edge of this bone is thickened and forms the point of contact for
the quadrate (this element is lost in this specimen).

The occipital condyle* is a curved, continuous "roll," with articu-
lar lateral knobs or facets. Presumably it is composed of the basioc-
cipital and parts of the exoccipitals. The condyle is elevated
strongly leaving the foramen opening almost wholly dorsal, sug-
gesting that the head must be held at an angle for the proper en-

* Since the description was drawn up the skull has suffered certain fractures and the loss of
the condyle.

35-3286

546 The University Science Bulletin

trance of the chord. Anterior to the condyle on the posterior ven-
tral surface, the basioccipital flares out suddenly then narrows
gradually to a point in front of the anterior level of the fenestra
ovale, where it is notched to receive the posterior ends of the ptery-
goids; but the sides remain nearly parallel for some distance be-
fore the bone terminates in (probably) three points (the tip of
this element is missing but the impression is more or less distinct ) .
Anteriorly the bone forms a suture with the orbitosphenoid and
along its middle parts with the occipital complex (prootic). Low
on the side anterior to the fenestra ovale is a large opening on the
side of the skull, bounded above by the prootic and below by the
sphenoidal element.

The anterior extension of the cranial cavity is exposed anteriorly.
It is surrounded by very heavy bone of the frontal and orbitosphe-
noid (see text fig. 6, Bf ).

Measurements in mm. of type. — Total length (estimated) 18; ac-
tual length of fossil 12.5; greatest width of braincase 7.7; narrowest
width of braincase 3.4.

Referred material. — Univ. Kansas Mus. Nat. Hist. (Paleo.) No.
7650 is referred with doubt to Rhineura ambhjceps. The braincase
is somewhat narrower, and while there are points of resemblance,
some differences are discernible. (Plate LXII).

Kansas Univ. Mus. Nat. Hist. (Paleo.) No. 9041. Fragmentary
skull. This seems to differ from the other described specimens in
having the posterior part of the pterygoids enlarged and much
thickened, with a thin, laterally compressed process reaching for-
ward to contact palatine and possibly also the ectopterygoid. A
fragment of the lower jaw (surangular) is pushed against the
fenestra ovale on the right side. A quadrate is in place, more hori-
zontal than vertical in position ( obviously not the normal position ) .
This bone shows some wear. There is no knob-like outer process
on its upper forward edge. ( Plate LXI ) .

Michigan Mus. Paleo. No. 25430, Oligocene, Orellan. Red zone
and sandstone. Smith pasture, center sec. 7, T.11N, R.53W, Logan
Co., Colorado. Actual length 14 mm. (Plate X). Skull, the brain-
case and facial portion largely complete. This is referred to Rhi-
neura ambhjceps with some hesitancy. Six teeth are present in the
maxillary, the second largest, the last three subequal in size with a
suggestion of a diastema between the third and fourth. The lower
jaw is fragmentary and two posterior teeth are in place. The quad-

t In this figure the element marked PO (paroccipital) is actually missing. A scar shows
its previous position.

Taylor: Oligocene Amphisbaenid Reptiles

547

rate is nearly vertical and there is a heavy knob-like structure on its
upper anterior edge. Running from the lateral floor of the orbital
vacuity to the coronoid is an ossified structure suggesting a cylin-
drical tendon. ( Plate LXVII ) .

oco

B

Fig. 6. Rhineura amblyeeps sp. nov. Referred specimen. Univ. Kansas
Mus. Nat. Hist. (Paleo.) No. 7650. Oligocene, Orellan, Clyde Ward Ranch,
Logan Co., Colorado. Dr. Claude Hibbard and party, collectors, July 31, 1946.
A. Lateral view; B. Ventral view; C. Dorsal view (relief of sculpturing on
frontals not adequately shown). Actual length 11.6 mm.

548 The University Science Bulletin

Rhineura wilsoni sp. nov.

Plate LVHI, figs. 1, 2; Plate LIX, figs. 2, 3; Plate LXIII, figs 1, 2, 3 ; text fig. 7, A, B, C

Type. — University of Kansas Museum of Natural History, Paleo.
No. 7651, consisting of a skull, nearly complete, but lacking squa-
mosal, quadrate, columella, extracolumella, and the entire lower
jaws. Collected under the direction of Dr. Claude W. Hibbard,
University of Kansas Museum of Natural History Expedition of
1946.

Type locality. — Clyde Ward Ranch, Logan County, Colorado,
July 1946.

Horizon. — Middle Oligocene, White River Formation, Oreodon
Zone.

Diagnosis. — A species characterized by the small depth of the
braincase, elongate prefrontals, and general slenderness of the
skull; maxillary teeth 7; nostrils ventral; area lateral to prevomers
somewhat inflated. Skull length 14.1. Deflected facial portion of
skull practically as wide as greatest posterior width.

Description of the type. — The skull is strongly humped near the
middle. The anterior part of the parietal, together with the remain-
der of the facial portion of the skull being strongly deflected down-
ward. Near the central part of the parietal there is a low sagittal
crest developed. This runs forward and terminates at a point where
two flattened oval surfaces diverge anteriorly. These two surfaces
are presumably points of muscle attachment. These surfaces are
slightly depressed but anterior to them the parietal surface becomes
very rugose, with pits and grooves, that tend to form a slightly radi-
ating pattern on the frontals. At the posterior end of the sagittal
crest of the parietal, the median point is wedged between two
points of the forward projecting part of the supraoccipital, which
bears a slight median crest. This is flanked laterally by the two
broad posterior extensions of the parietals that reach to within
one-third millimeter of the superior notch of the foramen magnum;
the bones are truncate posteriorly and a part is chipped away but
their posterior limit is marked by a low transverse ridge on the
supraoccipital. The parietals extend low on the sides of the brain-
case and form sutures with the prootic and apparently for a very
short distance with the orbitosphenoid, and broadly overlies the
forward extension of the combined occipital and otic elements
which are fused together. The anterior parietal suture with the
frontals is intricately dove-tailed with an outer diverging spine that
reaches forward perhaps as far as the prefrontal (the edges of the

Taylor: Oligocene Amphisbaenid Reptiles

549

supraorbital region are chipped and the extent of this element can-
not be determined ) ; a prefrontal is present on each side, the anterior
part wedged into the maxillary. It is somewhat triangular in shape,
however it is likely that portions of the edges of this element are
chipped away.

oco

--PM
-N

M

PV

BS

PO

BO

oco

PM

N

a h

PF

F

SO

I oco

Fig. 7. Rhineura wilsoni sp. nov. Type. Kansas Univ. Mus. Nat. Hist.
(Paleo.) No. 7649. Oligocene, Orellan, Clyde Ward Ranch, Logan Co., Col-
orado. Dr. Claude Hibbard and party, collectors, 1946. A. Lateral view;
B. Ventral view; C. Dorsal view. Actual length of skull 14.1 mm.

550 The University Science Bulletin

The frontals are once and one half as long as wide, the anterior
portions being wedged in between the maxillary and nasals. Nasals
are about three times as long as wide, extending to the end of
the snout and bending under to form the anterior rim of the nostril.
The premaxillary is about 2^ mm. wide on the superior aspect of
the snout. On the inferior aspect of snout the premaxillary narrows
between the two large nares, but I am uncertain whether they form
a part of the rim. Behind nostrils the element widens. The alveolar
surface which is more or less directed backward bears a single me-
dian tooth. Behind this and on a different level the bone again
narrows terminating in two blunt tips in contact with the prevomers.

The maxillary, laterally, rises considerably above the level of the
prefrontal and has contact with the prefrontal, frontal and nasal
on the superior aspect. Anteriorly a flattened extension of it forms
the outer and posterior border of the nostril, and a small curved
spinelike projection from it forms most of the inner nostril border.
In ventral aspect the maxillary borders the premaxillary, the pre-
vomers anteriorly, and the palatine, ectopterygoid, and apparently
also the orbitosphenoid, posteriorly. Three small pits (foramina)
are evident on the lateral aspect. A groove or depression passing
from nostril is ventral anteriorly, but becomes lateral low on the
side. Seen from the ventral aspect the alveolar surface faces slightly
inward. Seven teeth are present but only the bases remain, the
third and fourth being very much smaller than those preceding or
following. A deep pit exists behind the nostril which connects with
the groove along the alveolar surface. The broad inner shelf of
the maxillary overhangs the outer inflated wings of the prevomers.

The ventral part of the braincase consists of the basioccipital
fused laterally with the exoccipitals, paraccipitals, basisphenoid and
perhaps a presphenoid. Two lateral notches somewhat in front of
the level with the anterior edge of the fenestra ovalis, are for the
reception of the posterior extension of the pterygoids. These ele-
ments are missing from the skull. It is probable that anteriorly a
portion of the palatines remain.

Measurements in nun. — Total length 14.1; greatest width of brain-
case (through otic region) 6; narrowest width 3.1; greatest width
of facial portion of skull 5.8; greatest elevation of braincase 3.7.

Referred material. — A fragmentary specimen, consisting chiefly
of a braincase, is referred to this species with doubt. It is Univ.
of Michigan Museum of Paleontology No. 25429, Oligocene, Orellan,
from Smith Ranch, center sec. 7, T.11N, R.53W, Logan Co., Colo-

Taylor: Oligocene Amphisbaenid Reptiles 551

rado. It appears to approach Rhineura wilsoni closer than it does
the other species, and in spite of certain differences, I believe this
association the wisest disposition until more material of the forms
are available. ( Plate LVIII, figs. 1-2; Plate LIX, figs. 2, 3 ) .

Rhineura liatcheri Baur

Plate LXIV, figs. 1, 2, 3; Plate LXV, figs. 1, 2, 3; Plate LXVI ; figs 1, 2, 3; text fig. 8

Rhineura katcheri Baur, Amer. Naturalist, vol. 27, 1S93, p. 998, (Type, Princeton University-
No. 11389, Battle Draw Spring, Washington Co., South Dakota, Leptauchenia Zone) ;
Cope, Rep. U. S. Nat. Mus. 1898 (1900), p. 084; Hay, Bull. U. S. Geol. Surv. No. 179,
1902, p. 476; Eigenmann, Proc. Washington Acad. Sci., vol. 4, Sept. 30, 1902, pp. 533-
548, pis. 32-34; Douglass, Annal. Carnegie Mus., vol. 4, nos. 3 & 4, Apr. 1, 1908, pp.
283-284, text figs. 3-5; Nopcsa, Beit. Pal. Geol. Oester.-Ung., vol. 21, 1908, p. 46;
Eigenmann, Carnegie Inst. Washington Pub. 104, 1909, p. 4S ; O'Harra, South Dakota
School of Mines, Bull No. 9, p. 121, fig. 20; ibid. No. 13, 1920, p. 160; Zittel, and
Broili-Schlosser, Griindziige der Paleontologie, 4 ed., vol. 2. 1923, p. 257, fig. 360; Gil-
more, Mem. Nat. Acad. Sci., vol. 22, 3rd memoir, 1928, pp. 35-42, pi. 1, text figs. 17-19;
Proc. U. S. Nat. Mus., vol. S6, p. 15, fig. 3; Gilmore and Jepsen, Journ. Paleontology,
vol. 19, No. 1, Jan. 1945, p. 31.

This fossil species has had detailed treatment by Gilmore, 1928.
I have not had opportunity to examine the referred material, but
accept Gilmore's reference of two specimens to the Oreodon beds,
Badland Creek, Sioux Co., Nebraska ( Carnegie Mus. Nos. 423B and
423C ) ; and one in the Lower Oreodon beds, White River, Oligocene,
Cedar Draw, Cheyenne R., Washington Co., S. Dak. (Amer. Mus.
Nat. Hist. No. 12226). Whether the faunas of these two places are of
identical age as the Orellan of Logan Co., Colorado, remains to
be proved. However, it is generally believed that there is no very
considerable time interval involved between the three faunas.

In the material at hand I find certain skulls that appear to be
referable to the species Rhineura liatcheri judging by the published
descriptions and figures. These are discussed below.

Univ. Colorado Mus. No. 19852. Cedar Creek phase, White
River, Orellan (Middle of west half, sec. 7, T.11N, R.53W, VA miles
NE of Ward Ranch, 35 miles NW of Sterling, Logan Co., Colo.).
This skull is one of the most complete skulls I have examined.
Moreover, certain of the chondral bones of the braincase still have
sutures evident. Both lower jaws are in place, attached by their
respective quadrates, that on the right side has the posterior part
of the surangular absent. On the left side the protruding portion
of the stapes is missing and the extracolumella is lost from both
sides. A portion of the premaxilla has been chipped away.

The nasals are 3.2 mm. long and 1.2 mm. wide anteriorly. A short
median posterior projection of the premaxilla tends to separate
them anteriorly. The bones show little or no sculpturing or pitting.

552

The University Science Bulletin

The frontals, however, are heavily sculptured on their outer parts
with numerous pits ( and foramina ) while more mesially they show
some transverse grooves connecting with pits. They form dovetailed
sutures with the anterior part of the parietal along their posterior

oco

M N PM

-^ PM

OCO

Fig. 8. Rhineura hatcherii Baur. Univ. of Colorado Mus. No. 19852. Olig-
ocene, Orellan (Cedar Creek phase), middle of west half Sec. 7, T. UN, R.
53W, lVomi. NE of Ward's Ranch, 35 mi. NW Sterling, Logan Co., Colorado.
Dr. Robert Wilson, collector. A. Lateral view; B. Dorsal view; C. Ventral
view. Actual length of skull 12.2 mm.

Taylor: Oligocene Amphisbaenid Reptiles 553

borders. A median anterior parietal projection separates the pos-
terior ends of the frontals. Laterally the frontal overhangs the
orbital vacuity above its middle and forms a narrow lateral suture
with the prefrontal, and an elongate one with the maxillary; an-
teriorly an elongate spine projects forward from each frontal sepa-
rating the nasals, for half their length, from the maxillary.

The parietal is very large, saddle-shaped, with a median sagittal
crest (somewhat chipped away) from the anterior border of the
dorsal notch of the foramen magnum, and terminating where the
elevated portion begins to widen into a roughened triangular area
(presumably for muscle attachment). From the anterior edge of
this area the facial portion of the skull is deflected downward at
an angle. The anterior part of the parietal has a median projection
and two lateral arms that extend along the outer edge of the dorsal
deflected part of the skull, reaching forward to near the edge of
the orbital vacuity. Laterally the anterior part of the parietal
reaches to the orbital vacuity. Low on the sides the parietal con-
tacts two bones, the orbitosphenoid anteriorly and the prootic pos-
teriorly. The posterior extension of the parietal is chipped away
on both sides but a depression over the otic region suggests that the
original limits of the element approached near to the border of the
foramen magnum.

The supraoccipital dovetails mesially with the parietal, and on
each side the two wings extend forward and laterally, bordering the
foramen posteriorly. Normally it is largely covered by the parietal.
Laterally the supraoccipital forms sutures with the prootic and the
exoccipital.

The exoccipital is clearly defined above the suture, being evident
on the left side of the skull ( chipped away on the right side ) . It
borders the foramen magnum and extends back to form the outer
knob of the condyle, while a projection from the lower part forms
the posterior rim of the fenestra ovalis. Its relationship to the region
of the fenestra ovalis is not clearly defined. A deep foramen is evi-
dent at the lateral base of the condyle. Two of the semicircular
canals can be dimly discerned through the transparent bone as
whitish stripes. Lying on the anterior part of the exoccipital is a
superficial flake of bone, the squamosal, forming a lateral projec-
tion for the articulation of the quadrate. The basioccipital is clearly
defined on the ventral surface, its sides narrowing and forming
the median portion of the occipital condyle. Anteriorly on the ven-
tral surface it forms a suture with the basisphenoid and laterally

554 The University Science Bulletin

with the exoccipital. At the juncture of the three bones is a small,
somewhat elongate paroccipital forming the posterior ventral rim
of the fenestra ovalis.

The basisphenoid extends forward along the mesial ventral part
of the skull, forming sutures with the posterior part of the pterygoids
and overlaid by the latter bones and the palatines save for a narrow
median pointed part which disappears under the prevomers. I
cannot be certain that a separate presphenoid is present. Anteriorly
on the palate are the prevomers, the wings of which are not notice-
ably inflated.

The premaxillary is chipped away but on the ventral portion a
single tooth is evident, the base only remaining. The nostrils are
ventral, the dorsal rims being formed by the premaxillary, while a
spine from the maxillary forms most of the lower edge of the nostril.

The maxillary on the facial surface has a maximum width of 2.1
mm. and a total length of approximately 4 mm. An irregular row
of four pits appears near its lower edge, and two or three are present
near the middle of its lateral surface. Six teeth are present on the
alevolar surface, the second being largest. There is no appreciable
diastema in the series. At the posterior end of the maxillary the
ectopterygoid is present connecting pterygoid and maxillary.

The dentary is approximately 6 mm. in length, its greatest eleva-
tion being about 1.2 mm. It bears seven teeth, the first and fourth
being largest. From its dorsal edge arises a coronoid, its base
having a length of 1.5 mm. and an elevation of 2 mm. It forms a
suture with the inner surface of the surangular. The surangu-
lar is elongate extending for nearly half its length behind the articu-
lating fossae. I do not discern a splenial or an angulare since the
inner surface of the jaws cannot be seen, as the fossil is now pre-
pared. The quadrate is approaching a vertical position, its width
at the fossae being 1 mm., the lower edge rounded; and its greatest
length 2 mm., the upper part being divided into a broad surface
articulating with the squamosal and a small prong which lies against
the prootic. The surface directed toward the fenestra ovale is
strongly excavated. The stapes, present on the right side, is an
irregular oval with a median columellar prong 1 mm. long, directed
forward.

Measurements in mm. — Total length of skull (approximately)
12.6; total length of fossil 12.1; greatest width of brain case 6; great-
est width of facial portion 5; greatest depth of skull 4.1. (See
plate LXIV, figs. 1, 2, 3.)

Taylor: Oligocene Amphisbaenid Reptiles 555

Remarks.— The description of this young specimen (regarded
as young because of the lack of co-ossification of the chondral bones
of the brain case ) differs in no significant feature from the descrip-
tions of the type and the figures given by Gilmore (1928). The
smaller size of this specimen would seem to indicate an age differ-
ence.

I am also referring to the species two other skulls from the same
general locality. Univ. of Kansas Nat. Hist. Mus. Paleo. No. 8220
(Plate LXV). The skull has been broken across the middle and the
posterior parts are crushed somewhat. The parietal is more com-
plete than in the preceding specimen and extends farther back
covering the sutures between the supra- and exoccipital bones.
The posterior section of the surangular is present on each side
making contact with the quadrate. An extra-columella is present
on both sides, closely applied to the posterior excavation (notch)
of the quadrate. The sutures between the basioccipital and basi-
sphenoid are present and the tripartite character of the condyle can
be discerned. The maxillary teeth are 6-6, the second largest, the
fifth a little larger than the adjoining teeth. The measurements
approximate very closely those given for the preceding skull.

Univ of Kansas Nat. Hist. Mus. Paleo. No. 8960. Skull nearly
complete but with the posterior part of the braincase flattened.
The jaws are present, the posterior parts ( surangular ) being pushed
in, covering largely the fenestra ovalis. Sutures between the chon-
dral bones of the braincase can be discerned on the ventral side of
the skull, those separating paroccipitals, exoccipitals, basioccipital
and parisphenoid being distinct even on the condyle.

The maxillary bears seven teeth and there are seven teeth on the
dentary. However, the maxillary teeth seem anomalous. The sec-
ond upper tooth is largest and is followed by a pair of minute teeth
occupying space that would suffice for a tooth of normal size. The
last three teeth are subequal, the sixth being a trifle the larger. In
the lower jaw there are only six teeth actually present, but a tooth
is lost. On the right side the first and third (fourth) are enlarged
but a tooth is missing (presumably) behind the first; posterior to
the second large tooth there are three subequal teeth, equally
spaced. On the left jaw the first and fourth are largest, the two
intervening teeth being present. Behind the fourth there is a
narrow diastema scarcely large enough to permit a tooth as large
as the two posterior of the series. ( Plate LXVI. )

The skulls referred to Rhineura hatched all agree in having the
condyle placed close to the lower level of the skull ( See text figure

556 The University Science Bulletin

8 this paper; and Gilmore 1928, pi. 1, figs. 2 and 3). This allows
the foramen magnum opportunity to face posteriorly more than in
certain other species where the condyle is elevated and the foramen
magnum is largely dorsal. It would appear that the head of R.
hatched was held less elevated for direct entrance of the spinal
chord, than in these species ( amblyceps et al. ) .

BIBLIOGRAPHY

Baur, George.

1893. The discovery of Miocene amphisbaenians. American Nat., vol.
27, 1893, pp. 998-999. Hyporhina is described from the "Mio-
cene," but this is now regarded as Oligocene.

Bedriaga, J. VON.

1884. Amphisbaena cinerea Vand. and A. strauchi v. Bedr. Arch, fur
Naturg., vol. 1, 1884, pp. 24-35, pi. IV.

BOULENGER, GEORGE A.

1885. Catalogue of the lizards in the British Museum (Natural History),
2 ed., vol. 2, 1885, pp. 430-492, pis. 23-24.

Butler, Gerard W.

1895. On the complete or partial suppression of the right lung in the
Amphisbaenidae and of the left lung in snakes and snake-like
lizards and Amphibians. Proc. Zool. Soc. London, 1895, pp.
691-712, pi. 40.

Camp, C. L.

1923. Classification of the Lizards. Bull. Amer. Mus. Nat. Hist., vol.
48, 1923, pp. 289-481.

Camp, C. L., D. N. Taylor and S. P. Welles.

1942. Bibliography of fossil vertebrates, 1934-1938. Geol. Soc. Amer.
Special Paper No. 42, Nov. 1942, pp. 1-663.

Cope, Edward D.

1861. [Remarks on Rhineura floridana.] Proc. Acad. Nat. Sci. Philadel-
phia, 1861, p. 75.

1873. Synopsis of new Vertebrata from the Tertiary of Colorado.
Obtained during the summer of 1873. Washington Govt. Print.
Office, 1873, pp. 16-19.

1884. The Vertebrata of the Tertiary formations of the west. Rep. U. S.
Geol. Surv. Territ, vol. 3, 1884, pp. 770-781, pi. l-75a.

1900. The crocodilians lizards and snakes of North America. Ann. Rep.
U. S. Nat. Mus., 1898 (1900), pp. 151-1294, pis. 1-36, numerous
text figures.

Douglass, Earl.

1908. Some Oligocene lizards. Ann. Carnegie Mus., vol. 4, Nos. 3 and
4, 1908, pp. 278-285, text figs. 1-7.

ElGENMANN, C. H.

1902. The eyes of Rhineura floridana. Proc. Washington Acad. Sci., vol.
4, Sept. 30, 1902, pp. 533-548, pis. 32-34. (Mentions Hyporhina
antigua (sic) on p. 534.)

1909. Cave vertebrates of America; a study in degenerative evolution.
Carnegie Inst. Washington publ. 104, 1909, p. 48.

Fischer, E.

1900. Beitriige zur Kenntnis der Nasenhble und des Tranennasenganges
der Amphisbaeniden. Arch, micro. Anat., vol. 55, 1900, pp. 441-
478, pis. 21-24.

Taylor: Oligocene Amphisbaenid Reptiles 557

Gervais, Paul.

1853. Recherches sur l'osteologie de plusieurs especes d'amphisbenes et
remarques sur la classification de ces reptiles. Ann. Sci. Nat.
Paris, ser. 3, vol. 20, 1853, pp. 293-312, pis. 14-15.

(Note. — The plates are incorrectly numbered. The plate ex-
planation for plate 14, applies to plate 15, and vice versa.)

Gilmore, Charles W.

1915. Fossil turtles from the Uinta formation. Mem. Carnegie Mus., vol.

7, No. 2, pp. 101-161, pis. XVIII-XVII, 22 text figs.
1928. Fossil lizards of North America. Mem. Nat. Acad. Sci., vol. 22, 3d
mem., 1928, pp. i-ix; 1-201, pis. 1-27, text figs. 1-106.

(Part dealing with Amphisbaenids pp. 33-52, text figs. 16-28,
pi. 1.)
1938. Descriptions of new and little known fossil lizards from North
America. Proc. U. S. Nat. Mus., vol. 86, No. 3042, Dec. 16, 1938,
pp. 11-26, pi. 1, text figs. 1-9.

Rhineura minutus described (fig. 1) and Rhineura floridana and
Rhineura sternbergii figured.

1943. Fossil lizards of Mongolia. Bull. Amer. Mus. Nat. Hist., vol. 81,
art. 4, July 30, 1943, pp. 361-384, pi. 52, text figs. 1-22.
Crythiosaurus mongoliensis described (fig. 16).

Gilmore, Charles W., and G. L. Jepsen

1945. A new Eocene lizard from Wyoming. Tour. Paleont. ( Menasha,
Wis.) vol. 19, No. 1, 1945, pp. 30-34, 1 text fig.
Ototriton minor described.

Goodrich, E. S.

1930. Studies on the structure and development of vertebrates. London,
1930, pp. 1-837.

Hay, O. P.

Bibliography and catalogue of the fossil Vertebrata of North
America. Bull. U. S. Geol. Survey, No. 179, pp. 473-478.

HOFFSTETTER, R.

1942. Sur la presence d'Amphisbaenidae dans les gisements francais.
C. R. Seances Soc. Geol. France, 1942, 3-4, pp. 24-25.

James, Edwin.

1823. In Long's Expedition to the Rocky Mountains, vol. 1, 1823, p. 484.

Laurent, R.

1947. Note sur les Amphisbaenidae d'Afrique. Rev. Zool. Bot. Africa,
vol. 40, No. 1, 1947, pp. 52-63.

Loomis, F. B.

1919. A new amphibian from the Eocene. Amer. Jour. Sci., vol. 47, 1919,
pp. 217-219, text fig. 1.

Ototriton solidus described.

Loveridge, Arthur.

1941. Revision of the African lizards of the family Amphisbaenidae. Bull.
Mus. Comp. Zool. Harvard Coll., vol. 87, 1941, pp. 353-451.
(Includes excellent bibliography.)

Marsh, O. C.

1871. Notice of some new fossil reptiles from the Cretaceous and Tertiary-
formations; with note on a new and gigantic pterodactyl. Amer.
Jour. Sci., 3d ser., vol. 1, art. 66, 1871, pp. 447-459.
1871. [Communication on some new reptiles and fishes from the Cre-
taceous and Tertiary formations.] Proc. Acad. Nat. Sci. Phila-
delphia, 1871, pp. 103-105.

G[hjtosanrus] anceps = Lestophis anceps.

558 The University Science Bulletin

Nopcsa, VON F.

1903. Zur Kenntniss fossilen Eidechsen. Beitr. Pal. Geol. Osterreich-
Ungams, vol. 21, 1903, pp. 33-62, pi. 3, 5 text figs.

O'Harra, C. C.

1910. The Bad Lands formation of the Black Hills region. Bull. South

Dakota School of Mines, No. 9, 1910, p. 121, fig. 20.
1920. White Biver Bad Lands. Bull. South Dakota School of Mines,

No. 13, 1920, p. 160.

Peters, Wilhelm.

1882. liber eine neue Art und Gattung der Amphisbaenoiden, Agamodon
anguliceps. mit eingewachsenen Zahnen, aus Barava ( Ostafrica )
und neber die zu den Trogonophides gehorigen gattungen. Sitz-
ungb. konig. preuss. Akad. Wiss. Berlin, Bd. 26, 1882, pp. 579-
584, pi. 10.

Romer, Alfred S.

1932. Vertebrate paleontology. Univ. Chicago Press, 1932, pp. 1-687.

Schmidt, Karl P.

1927. New reptilian generic names. Copeia, 1927, No. 163, pp. 58-59.

Smalian, Carl.

1885. Beitriige zur Anatomie der Amphisbaeniden. Zeitsch. wiss. Zool.,
Bd. 42, 1885, pp. 126-202, pis. 5-6.

Smith, Hobart M., and Edward H. Taylor.

1948. An annotated checklist and kev to the amphibia of Mexico. U. S.
Nat. Mus. Bull. 194, 1948, pp. 1-118.

Taylor, Edward H.

1938. Does the amphisbaenid genus Bipes occur in the United States.
Copeia, 1938, No. 4, Dec, p. 209.

Walker, M. V.

1932. A new burrowing lizard from the Oligocene of central Wyoming.
Trans. Kansas Acad. Sci., vol. 35, 1932 (1933), pp. 224-231.
Rhineura sternbergii described.

Williston, Samuel W.

1925. The osteology of the reptiles. Harvard Press, Cambridge, Mass.,
1925, pp. 1-300, figs. 1-191.

Zangerl, Rainer.

1944. Contributions to the osteologv of the skull of the Amphisbaenidae.
Amer.. Midi. Nat., vol. 31, No. 2, Mar. 1944, pp. 417-454. text figs.
1-18.

1945. Contributions to the osteology of the post-cranial skeleton of the
Amphisbaenidae. Amer. Midi. Nat., vol. 33, No. 3, May 1945, pp.
764-780, figs. 1-9.

Wiegmann, August.

1836. Ueber die fusslosen Amphisbanen mit Brustschildern ( Lepidosternon
Wagl.) Arch, fur Naturg., vol. 2, pt. 1, 1836, pp. 152-158, Tab. 4,
figs. 3-6.

Zittel, K., and Broili-Schlosser.

1923. Grundzuge der Paleontologie, vol. 2, 1923, p. 257-360.

560 The University Science Bulletin

PLATE LVIII

Fig. 1. fRhineura wilsoni sp. nov. Referred materials Univ. Michigan Mus.
Paleontology No. 25429. Oligocene, Orellan, Smith Ranch, center sec. 7, T.
UN, R. 53W, Logan Co., Colorado. Dr. Claude Hibbard and party, Aug. 5,
1947. Ventral view. Actual length of fossil, 13 mm.

Fig. 2. Same. Dorsal view.

Fig. 3. Gilmoreia attenuatus sp. nov. Type. U. S. Nat. Mus. No. 16308.
Oligocene, Orellan (Brule), 8 mi. N Douglas, Converse Co., Wyoming. G. P.
Sternberg, collector. Dorsal view. Actual length of fossil 7.5 mm.

Fig. 4. Same. Lateral view.

Fig. 5. Same. Ventral view.

Fig. 6. Hyporhina galbreathi sp. nov. Tj-pe. Kansas U. Mus. Nat. Hist.
(Paleo.) No. 8221. Oligocene, Orellan, Clyde Ward Ranch, Logan Co., Col-
orado. Dr. Claude Hibbard and party, collectors. Ventral view. Actual length
of fossil S.5 mm.

Fig. 7. Same. Lateral view.

Fig. 8. Same. Dorsal view.

PLATE LVIII

36-3286

562 The University Science Bulletin

PLATE LIX

Fig. 1. fRhineura amblyceps sp. nov. (Referred material) Kansas U. Mus.
Nat. Hist, No. 9041. Oligocene, Orellan, Clyde Ward Ranch (Wy,, sec. 7, T.
UN, R. 53W), Logan Co., Colorado. Edwin C. Galbreath, collector, 1948
Lateral view. Actual length of fossil 10.6 mm.

Fig. 2. fRhineura ivilsoni sp. nov. (Referred material). Univ. Michigan
Mus. Paleo. No. 25429. Oligocene, Orellan, Smith Ranch (center sec. 7, T.
UN, R. 53W), Logan Co.. Colorado, Dr. Claude Hibbard and party, collectors,
Aug. 5, 1947. Lateral view. Actual length of fossil 12 mm.

Fig. 3. Same. Ventral view.

Fig. 4. Hyporhina galbreathi sp. nov. (Referred material). Kansas U. Mus.
Nat. Hist. (Paleo.) No. 8222. Oligocene, Orellan, Clyde Ward Ranch, SE%,
SWy4, S. 12, T. UN, R. 54W, Logan Co., Colorado. Edwin C. Galbreath. Col-
lector. Dorsal view. Actual length of fossil 6 mm.

Fig. 5. Hyporhina galbreathi sp. nov. (Referred material). Kansas U. Mus.
Nat. Hist, (Paleo.) No. 8219. Oligocene, Orellan, Clyde Ward Ranch, SE%,
SW%, S. 12, T. UN, R, 54W, Logan Co., Colorado. Edwin C. Galbreath, col-
lector, 1948. Dorsal view. Actual length of fossil 8.4 mm.

PLATE LIX

564 The University Science Bulletin

PLATE LX

Fig. 1. Rhineura hibbardi sp. nov. Type. Univ. Michigan Mus. Paleo.
No. 25431. Oligocene, Orellan, Smith Ranch (center sec. 7, T. 11N, R. 53W),
Logan Co., Colorado. Dr. Claude Hibbard and party, collectors, 1947. Lateral
view. Actual length of fossil 16.6 mm.

Fig. 2. Same. Dorsal view.

Fig. 3. Same. Ventral view.

Taylor: Oligocene Amphisbaenid Reptiles

565

PLATE LX

566 The University Science Bulletin

PLATE LXI

Fig. 1. Rhineura amblyceps sp. nov. Type. Kansas Univ. Mus. Nat. Hist.
(Paleo.) No. 7649. Oligocene, Orellan, Clyde Ward Ranch. Logan Co.. Col-
orado. Dr. Claude Hibbard and party, collectors, 1946. Lateral view. Actual
length of fossil 11 mm.

Fig. 2. Same. Ventral view.

Fig. 3. Same. Dorsal view (note figure in text drawn before loss of condyle).

Fig. 4. ?Rhineura amblyceps sp. nov. (Referred material) Kansas IT. Mus.
Nat. Hist. No. 9041. Oligocene, Orellan. Clyde Ward Ranch (W% sec. 7,
T. UN, R. 53W), Logan Co., Colorado. Edwin C. Calbreath, collector, 1948.
Ventral view. Actual length of fossil 10.6 mm.

Fig. 5. Same. Dorsal view.

PLATE LXI

568 The University Science Bulletin

PLATE LXII

Fig. 1. Rhineura amblyceps sp. nov. (Referred material) Kansas Univ.
Mus. Nat. Hist. (Paleo.) No. 7650. Oligocene, White River (Orellan), Logan
Co., Colorado. Dr. Claude Hibbard and party, collectors, July 31, 1946. Lat-
eral view of skull, actual length 12.1 mm.

Fig. 2. Same. Ventral view of skull.

Fig. 3. Same. Dorsal view.

PLATE LXII

570 The University Science Bulletin

PLATE LXIII

Fig. 1. Rhineura wihoni sp. nov. Type. Kansas Univ. Mus. Nat. Hist.
(Paleo.) No. 7649. Oligocene, Orellan, Clyde Ward Ranch, Logan Co.. Col-
orado. Dr. Claude Hibbard and party, collectors, 1946. Lateral view. Actual
length of skull 14.1 mm.

Fig. 2. Same. Ventral view.

Fig. 3. Same. Dorsal view.

572 The University Science Bulletin

PLATE LXIV

Fig. 1. Rhineura hatcheru Baur. (Referred material) Univ. of Colorado
Museum (Paleo.) No. 19852. Oligocene, White River (Orellan), Cedar Creek
Phase; middle of west half sec. 7, T. UN. R. 53W, iy2 miles NE of Clyde Ward
Ranch, 35 miles NW of Sterling, Logan Co., Colorado. Dr. Robert Wilson,
collector, 1940. Lateral view of skull. Actual length of fossil 12.1 mm.

Fig. 2. Same. Ventral view of skull.

Fig. 3. Same. Dorsal view of skull.

Taylor: Oligocene Amphisbaenid Reptiles

573

PLATE LXIV

574 The University Science Bulletin

PLATE LXV

Fig. 1. Rhineura hatcherii Baur. (Referred material) Kansas Univ. Mus.
Nat. Hist. (Paleo.) No. 8220. Oligocene, Orellan. SE%, S. 21, T. UN, R. 53W.
Logan Co., Colorado. Edwin C. Galbreath, collector, 1948. Lateral view of
skull. Actual length of fossil 12.2 mm.

Fig. 2. Same. Ventral view of skull.

Fig. 3. Same. Dorsal view of skull.

Taylor: Oligocene Amphisbaenid Reptiles

575

PLATE LXV

576 The University Science Bulletin

PLATE LXVI

Fig. 1. Rhineura hatcherii Baur. (Referred material) Kansas Univ. Mus.
Nat. Hist. (Paleo.) No. 8960. Oligocene, Orellan. SEYi, S. 3, T. UN, R. 54W,
Logan Co., Colorado. Edwin C. Galbreath, collector, 1949. Lateral view of
skull. Actual length of fossil 12.2 mm.

Fig. 2. Same. Ventral view of skull.

Fig. 3. Same. Dorsal view of skull.

Taylor: Oligocene Amphisbaenid Reptiles

571

PLATE LXVI

37-3286

578 The University Science Bulletin

PLATE LXVII

Fig. 1. fRhim urn amblyceps. (Referred material) Univ. Michigan Mns.
Paleontology No. 25430. Oligocene, Orellan (Brule), Red zone and Sandstone.
Smith pasture, center sec 7. T. UN, R. 53W, Logan Co., Colorado. Dr. Claude
Hibbard and party, collectors, Aug. 6, 1947. Lateral view of skull. Actual
length of fossil 14 mm.

Fig. 2. Same. Ventral view of skull.

Fig. 3. Same. Dorsal view of skull.

PLATE LXVII

□

23-3280

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