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AMERICAN MUSEUM NovJitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET NEW YORK, N.Y. 10024 U.S.A. NUMBER 2625 JUNE 30, 1977 ROBERT WAYNE VAN DEVENDER AND CHARLES J. COLE Notes on a Colubrid Snake, Tantilla vermifarmis from Central America * t,a*; *e?;,t.'TM- AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET. NEW YORK, N.Y. 10024 Number 2625, pp. 1-12, figs. 1-8, tables 1-3 June 30, 1977 Notes on a Colubrid Snake, Tantilla vermiformis, from Central America ROBERT WAYNE VAN DEVENDER1 AND CHARLES J. COLE2 ABSTRACT Tan tilla vermiformis has been known from the species: distribution, ecology, reproduction, only seven specimens, all from Nicaragua, only external morphology, osteology, hemipenes, one of which has precise locality data. We report chromosomes and relationships. We designate here the discovery of 25 specimens in Costa Rica USNM 32338 as the lectotype. and present new information on the biology of INTRODUCTION The genus Tantilla contains about 47 species species has been known previously from a total distributed between the central United States of only seven specimens, all from Nicaragua, and and northern Argentina (Peters and Orejas- only one of these previous specimens had precise Miranda, 1970). There is no comprehensive re- locality data (Wilson and Villa, 1973). Because view of the genus, and relationships among little is known about this species and because species and species groups are poorly known. studies of variation in local populations provide Study of these snakes is difficult for several rea- baselines for comparative work in systematics, sons: (1) Many species are known from few indi- we report the Costa Rican series in detail here viduals, so variation in scutellation, coloration and comment on various aspects of its biology: and other classical taxonomic characters is distribution, ecology, reproduction, external poorly known; (2) they are small, secretive, fos- morphology, osteology, hemipenes, chromo- sorial animals with scutellation simplified by somes, and relationships. scale fusion; (3) most investigators have not used Hallowell ("1860" [1861?]) described Lio- such potentially important characters as are avail- ninia vermiformis and stated that there were able in hemipenis morphology, osteology and three specimens from "Nicaragua." Cope (1861) karyotypes. examined the syntypes and placed the species in Since 1972, 25 specimens of Tantilla vermi- the genus Tantilla. Later, Cope (1875) men- formis (Hallowell, "1860" [1861?]) have been tioned that there were four syntypes of T. vermi- found on the Pacific versant near Los Angeles de formis, when he described Tantilla canula on the Tilaran, Guanacaste Province, Costa Rica. This basis of an additional specimen that previously 'Research Associate, Division of Amphibians and Reptiles, Museum of Zoology, the University of Michigan. 2Associate Curator, Department of Herpetology, the American Museum of Natural History. Copyright O The American Museum of Natural History 1977 ISSN 0003-0082 / Price $1.05 2 AMERICAN MUSEUM NOVITATES NO. 2625 he (Cope, 1866) had referred to T vermiformis. Nanne. Appreciation of permission to examine The four syntypes of T. vermiformis are now specimens in their care is extended to Dr. Arnold USNM 32338-32341. They were recatalogued G. Kluge and Dr. Ronald A. Nussbaum (UMMZ), from USNM 5792 in the year 1903, with the Dr. Tom Collins and Dr. William E. Duellman notation "Ret. from Cope's estate." The con- (KU), Dr. Jay M. Savage (CRE), Dr. John W. fusion regarding the syntypes and our examina- Wright (LACM) and Dr. George R. Zug and Dr. tion of a large number of specimens, including W. Ronald Heyer (USNM). In addition, we are re-examination of the type-series by one of us grateful to Dr. Thomas Moore, who identified (R.W.V.) make it desirable to designate a lecto- the insect parts, Mr. P. Dayanandan and Mr. V. type here. Baldwin for assistance in preparing the scanning There is no way to recognize which of the electron micrographs, Mr. Juan Carlos Barberis, four specimens of USNM 32338-32341 was not who rendered the illustrations of a hemipenis, in the original type-series of three specimens Ms. Carol R. Townsend, who made the chromo- mentioned by Hallowell ("1860" [1861?]), or some preparations, and to Dr. Roy W. Mc- whether in fact that statement was an error and Diarmid, Dr. Ronald A. Nussbaum, Mr. Paul E. actually there always were four syntypes. The Feaver and Dr. Douglas Ruby, who read an early original catalogue at the National Museum of draft of the manuscript. Natural History (USNM) states that all four specimens were originally catalogued there to- METHODS gether in 1861 (Alan H. Savitzky, personal commun.). Three of the four syntypes are signifi- The 25 T. vermiformis were collected in cantly incomplete. The heads and tails of USNM Guanacaste Province, Costa Rica between 1972 32339 and 32341 are missing, and USNM and 1975. Several living ones were sent directly 32340 is separated into three parts with some to the American Museum of Natural History, skin missing from between them. USNM 32338 is New York, for study of chromosomes and hemi- the only specimen that is essentially intact. It penes. The remaining specimens were preserved agrees with immediately for analysis of diet and scutellation. Hallowell's ("1860" [1861?]) de- Trunk vertebrae from one specimen were pre- scription of coloration and head scutellation, al- pared water though it does not have 122 ventrals and 26 sub- by maceration. Twenty-five mid- caudals as mentioned in the original description; body vertebrae were measured as described by USNM 32338 has 119 ventrals and 21+ sub- Auffenberg (1963). Other osteological prepara- caudals (tail nearly, but not, complete). None of tions were made from five additional specimens. the other syntypes matches the original descrip- Variation in all characters was determined and compared with that of the seven Nicaraguan tion more closely than this specimen,' however, specimens. All values are expressed as: Range so we designate USNM 32338 as the lectotype. (mean ± standard error [SE], N = sample size). ACKNOWLEDGMENTS The SE is omitted if sample size is fewer than 10. Statistical comparisons were based on t-tests, We thank Dr. Norman J. Scott, Dr. Robert using the 95 percent confidence level. Wilkinson, Mrs. Amy Van Devender and Dr. We used the Dowling (1951) method in count- James DeWeese for assistance in collecting and ing ventral scales. Subcaudals were counted on transporting specimens. Permission to collect and one side, beginning with the first scale in contact export these animals was provided by the Minis- with its opposite number. The terminal spine was terio de Agricultura y Ganaderia, Sylvestre y not counted. Condition of the anal plate was Pescado, through the courtesy of Sefior Herbert noted on all specimens. Head scales counted were supralabials, infralabials, preoculars, postoculars, 'In the original description, Hallowell ("1860" temporals, and genials. Contact between post- [1861?1, p. 484) stated "a single preanal; .. .," which nasal and preocular was recorded as was contact is highly unusual in Tantilla. This statement is an error, between mental and anterior genials. An ocular however, as all four syntypes have a divided anal plate. micrometer was used to determine length and 1977 VAN DEVENDER AND COLE: COLUBRID SNAKE 3 width of dorsal head scales, eye size, snout length m. elev., LACM 122002, 122003, UMMZ and distance between tip of snout and anterior 135259(6), 135260, 131399, 132191, 132192, edge of naris. 133892(2), 138224 and AMNH 111325-111327; Body length (snout-vent), tail length and the 1.2 km. E Los Angeles de Tilaran, 510 m. elev., ratio of tail length to total length were obtained CRE 3730; 2 km. E Los Angeles de Tilaran, 520 for all animals with complete tails. Presence or m. elev., AMNH 111323, 111324 and UMMZ absence of the left oviduct (Clark, 1970) was 132189; Finca La Pacifica, 5.5 km. NW Canias, checked in four adult females. Reproductive 50 m. elev., CRE 2217. state and diet were determined by dissection. Coloration characters were noted for all speci- RESULTS mens. Maxillary teeth were counted in 18 snakes, palatine and pterygoid teeth in two, and mandib- ECOLOGY ular teeth in two. Skull characters were examined on three dry preparations (UMMZ nos. 132192, Of these 25 specimens all but one was col- 135260, 139892) and in one cleared and stained lected in or under rotting logs in pastures be- specimen (UMMZ no. 132189-OP 6384). tween 0.6 and 2.0 km. (airline) east of Los We examined hemipenes using the methods Angeles de Tilaran, Guanacaste Province, Costa and terminology of Dowling and Savage (1960) Rica, elevation 480 to 520 m. (fig. IA); most of and of Myers (1974). Everted organs were the snakes were 3 to 5 cm. below the surface in studied either without special preparation or moist, sandy soil. The exceptional specimen after injection of colored liquid latex (Dowling (CRE 2217, fig. IB) was under a small stone in and Savage, 1960). tropical deciduous forest at Finca La Pacifica, Chromosomes of bone marrow (from crushed 5.5 km. (airline) northwest of Cafias, Guanacaste vertebrae, as suggested by Dr. Laurence M. Province, elevation 50 m.; it was on the surface Hardy) and testicular cells were prepared for of dry, packed soil near the entrance to a small examination by means of the colchicine, hypo- burrow. tonic citrate, flame-dried procedure (Patton, Tantilla vermiformis is not evident at these 1967), slightly modified for reptiles (Cole and sites throughout the year. Despite considerable Leavens, "1971" [1972]). We employ the effort, only two individuals (CRE 2217 and chromosome terminology used by Cole (1970). UMMZ 138224) were found during the dry sea- son (mid-November to mid-May).