A Revision of Snakes of the Genus Conophis (family Colubridae, From Middle America)

Part 1

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Typographical corrections are listed at the end of this version. The list of publications has been compiled after the article's text.

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UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY

Volume 15, No. 6, pp. 251-295, 9 figs.

October 4, 1963

A Revision of Snakes of the Genus Conophis (Family Colubridae, from Middle America)

BY JOHN WELLMAN

UNIVERSITY OF KANSAS LAWRENCE 1963

UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY

Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr.

Volume 15, No. 6, pp. 251-295, 9 figs. Published October 4, 1963

UNIVERSITY OF KANSAS Lawrence, Kansas

PRINTED BY JEAN M. NEIBARGER. STATE PRINTER TOPEKA. KANSAS 1963

29-5936

A Revision of Snakes of the Genus Conophis (Family Colubridae, from Middle America)

JOHN WELLMAN

CONTENTS

PAGE

INTRODUCTION 253

ACKNOWLEDGMENTS 254

MATERIALS AND METHODS 254

GENUS Conophis Peters 255 Key to the Species and Subspecies 257 Analysis of Characters 257 Scutellation 258 Size and Proportions 258 Color Pattern 260 Sexual Dimorphism 260 C. lineatus 262 C. lineatus dunni 262 C. lineatus lineatus 267 C. lineatus concolor 270 C. nevermanni 272 C. pulcher 274 C. vittatus 277 Skull 282 Dentition 288 Vertebrae 288 Hemipenes 289 Food and Feeding 289 Effect of Poison 290

TAXONOMIC RELATIONSHIPS AND EVOLUTION 291

SUMMARY 292

LITERATURE CITED 293

INTRODUCTION

Part 2

Need for a comprehensive systematic review of the snakes of the genus Conophis was pointed out by Stuart (1954a, b). Since these snakes appeared to be of zoogeographic importance in the Central American region, I undertook the review as set forth on the following pages.

ACKNOWLEDGMENTS

For permission to examine specimens, and for information concerning specimens in their care, I am grateful to Mr. L. C. Battersby and Miss Alice G. C. Grandison, British Museum (Natural History); Mr. Charles M. Bogert and Dr. Richard G. Zweifel, American Museum of Natural History; Dr. Doris M. Cochran, United States National Museum; Prof. William B. Davis, Agricultural and Mechanical College of Texas; Dr. Josef Eiselt, Naturhistorisches Museums, Vienna; Prof. Norman Hartweg and Prof. Laurence C. Stuart, Museum of Zoology, University of Michigan; Dr. Robert F. Inger, Chicago Natural History Museum; Dr. Alan E. Leviton, California Academy of Sciences; Mr. Edmond V. Malnate, Academy of Natural Sciences, Philadelphia; Prof. George S. Myers, Stanford University Natural History Museum; Mr. Wilfred T. Neill, Ross Allen's Reptile Institute; Mr. Neil D. Richmond, Carnegie Museum; Dr. William J. Riemer, University of Florida Collections; Prof. Robert C. Stebbins, Museum of Vertebrate Zoology, University of California; Prof. Hobart M. Smith, University of Illinois Natural History Museum; and Dr. Ernest E. Williams, Museum of Comparative Zoology, Harvard.

Prof. William E. Duellman supplied invaluable information and guidance in my study. I am grateful to Prof. E. Raymond Hall for use of facilities of the Museum of Natural History and editorial assistance. I thank Prof. Laurence C. Stuart and Prof. Edward H. Taylor for information and suggestions. My own field experience in Middle America came as a result of assisting Professor Duellman in his own researches supported by a grant from the National Science Foundation (NSF-G 9827). For these things I am deeply grateful. Specimens that I have seen alive were collected by field companions Dale L. Hoyt and Jerome B. Tulecke. Finally, I am grateful to my wife, Margaret L. Wellman, for much help including typing much of the manuscript.

MATERIALS AND METHODS

Of the 325 specimens of the genus Conophis available to me, representing most of those in museum collections, scale counts were made in the usual manner on 309. Ventrals were counted following the system proposed by Dowling (1951:97-99); the anal plate was not included. The anteroposterior position of the place where reduction occurs in the number of the dorsal rows of scales is designated by citing the number of the ventral scale directly beneath that place.

Measurements were taken to the nearest millimeter by means of a millimeter stick. Body length is the distance from the tip of the snout to the posterior edge of the anal plate; tail length, from the latter point to the tip of the tail; and total length, the sum of the body plus tail.

Descriptions of color are based on preserved specimens. Where descriptions of the color of living individuals are given, the data were taken from Kodachrome slides made available to me by William E. Duellman. Due to the transient nature of the longitudinal dark stripes in these snakes, no standard terminology has been devised, except that the posterior continuations of the stripes which on the head pass through the eye are termed lateral stripes; the posterior continuations of the median stripe of the head are termed dorsolateral stripes. A paravertebral stripe is one that is present on the scale-row on either side of, but not including, the mid-dorsal (vertebral) scale-row.

In order to reduce confusion in the discussion of variation, the numbers designating the rows of dorsal scales are written as 1st, 2nd, whereas the numbers designating the stripes are written as first, second.

Except in three dried skeletons, teeth were counted on dentigerous bones in situ. Since teeth are often missing, the sockets were counted in order to obtain an accurate count.

In accounts of the species and subspecies, the observed range of variation is followed by the mean in parentheses; in some instances the mean is followed by the standard deviation, also in parentheses. An example is 65-79 (70.6 +- 3.93).

Each synonymy includes all generic and specific combinations known to me that have been used for the genus, and, in addition, references to catalogues, checklists, and reports of collections.

Localities of occurrence that are not plotted on the distribution maps are recorded in italic type under Specimens Examined. In the list of Specimens Examined the localities and specimens are listed in the following order: countries in alphabetical order; states or departments in alphabetical order in each country; localities in alphabetical order in each state or department; museum numbers in numerical order after the abbreviations of names of museums. When more than one specimen bears a single catalogue number, the number of specimens is given in parentheses following the museum catalogue number. Specimens for which data are given only as to country or to state or department are listed first after the name of that political unit under "no specific locality."

The abbreviations for the museum collections are:

AMNH American Museum of Natural History ANSP Academy of Natural Sciences of Philadelphia BMNH British Museum (Natural History) CAS California Academy of Sciences CNHM Chicago Natural History Museum ERA-WTN E. Ross Allen-Wilfred T. Neill, Ross Allen's Reptile Institute KU University of Kansas Museum of Natural History MCZ Museum of Comparative Zoology, Harvard MVZ Museum of Vertebrate Zoology, University of California NMW Naturhistorisches Museums Wien, Vienna SU Stanford University Natural History Museum TCWC Texas Cooperative Wildlife Collection, Agricultural and Mechanical College of Texas UF University of Florida Collections UIMNH University of Illinois Museum of Natural History UMMZ University of Michigan Museum of Zoology USNM United States National Museum

Family COLUBRIDAE

Subfamily Xenodontinae

Genus =Conophis= Peters

Tomodon (part) Dumeril, Bibron and Dumeril, Erpetologie Generale, 7(pt.2):936, February 7(pt.2):936, February 25, 1854 (lineatus and vittatus); Salvin, Proc. Zool. Soc. London, 28:455, 1860 (pulcher).

Psammophis (part), Guenther, Catalogue of Colubrine Snakes in the Collection of the British Museum, London, 1858:135 (lineatus).

Conophis Peters, Monatsb. Akad. Wiss. Berlin, 1860:519-520, pl., fig. 3 (vittatus); Cope, Proc. Acad. Nat. Sci. Philadelphia, 13:300, December 28, 1861 (lineatus concolor); Proc. Acad. Nat. Sci. Philadelphia, 18:318-319, February 20, 1867 (lineatus concolor); Proc. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876 (pulcher); Bocourt in Dumeril, Bocourt and Mocquard, Mission Scientifique au Mexique et dans l'Amerique Centrale, 2:643-644, pl. 38, fig. 5, 1886 (lineatus lineatus); Cope, Proc. Amer. Philos. Soc., 23:489, October 28, 1886; Hoffmann, Klassen und Ordnungen des Thier-Reichs. Reptilien. Bd. 6, 3:1707, 1890; Cope, Trans. Amer. Philos. Soc., 18:207, April 15, 1895; Dunn, Bull. Antivenin Inst. Amer., 2(1):21, 24, April, 1928; Copeia, no. 4:214, December 31, 1937 (nevermanni).

Tachymenis (in part), Garman, Bull. Essex Inst., 16:33, January 9, 1884 (vittatus and lineatus).

Erythrolamprus (in part), Ditmars, Bull. Antivenin Inst. Amer., 2(2):27-29, June.

Coniophanes (in part), Wettstein, Sitz. Akad. Wiss. Wien, mathem-naturw. kl. 143:37-38, 1934 (nevermanni).

Historical summary.--In 1854 Dumeril, Bibron and Dumeril described and figured Tomodon lineatum from America. In 1860 Peters described and figured as a new genus and species, Conophis vittatus, based on a specimen that he had obtained from a dealer in Hamburg. The provenance of this specimen is not known, for it was discovered aboard a ship near the mouth of the Mississippi River. It was not until 1871 that Cope included lineatus in the genus Conophis. Cope (1861) proposed the name Conophis vittatus (nec Peters, 1860). Later (1900) he changed its name to Conophis lineaticeps. Early uncertainty of the relationships of the species lineatus caused Guenther (1858) to place it in the genus Psammophis. With the exception of Garman (1884a and 1884b) who placed lineatus in the genus Tachymenis, and Wettstein (1934) who reported five specimens of Conophis nevermanni as Coniophanes i. imperialis, all specimens reported after 1876 were placed in the genus Conophis.

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