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UNIVERSITY OF KANSAS

SCIENCE BULLETIN

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MUS. CGKP. ZOOL LIBRARY

OCT 16

HARVARD 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

DEVOTED TO

THE PUBLICATION OF THE RESULTS OF

RESEARCH BY MEMBERS OF THE

UNIVERSITY OF KANSAS

MUS. COMP. ZOOL LIBRARY

OCT 16 iL

HARVARD UNIVERSITY

Volume XXXIV, Part I

University of Kansas Publications

Lawrence, October 1, 1951

PRINTED BY

FERD VOILAND. JR.. STATE PRINTER

TOPEKA. KANSAS

1951

23-3286

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

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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)