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Rare Snakes—Five New Species from Eastern Panama: Reviews Of PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3391, 47 pp., 20 ®gures, 2 maps, 3 tables February 19, 2003 Rare SnakesÐFive New Species from Eastern Panama: Reviews of Northern Atractus and Southern Geophis (Colubridae: Dipsadinae) CHARLES W. MYERS1 CONTENTS Abstract ....................................................................... 2 Resumen ....................................................................... 2 Introduction .................................................................... 3 Gorgas Memorial Laboratory and the Panama Snake Census ....................... 4 Methods of Study ............................................................. 5 Genus Atractus ................................................................. 8 Key to Panamanian Atractus ................................................... 9 Atractus clarki Dunn and Bailey .............................................. 10 Atractus darienensis, new species .............................................. 16 Atractus depressiocellus, new species .......................................... 20 Atractus hostilitractus, new species ............................................ 22 Atractus imperfectus, new species ............................................. 25 Genus Geophis ................................................................ 28 Key to Panamanian Geophis .................................................. 28 Records Excluded from Central Panama ........................................ 29 Geophis bellus, new species .................................................. 30 Geophis brachycephalus (Cope) ............................................... 37 Geophis, species inquirenda .................................................. 38 Geophis hoffmanni (W. Peters) ................................................ 40 Museum Abbreviations and Acknowledgments .................................... 43 References .................................................................... 44 1 Curator Emeritus, Division of Vertebrate Zoology (Herpetology), American Museum of Natural History. Copyright q American Museum of Natural History 2003 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3391 ABSTRACT The South American genus Atractus barely enters political North America on the eastern half of the Isthmus of Panama, where it is extraordinarily rare. Collected over a period of 39 years, the ®ve Panamanian specimens of Atractus known to the author represent ®ve species! Four new species are described: A. darienensis, A. hostilitractus, A. imperfectus, and A. de- pressiocellus. The ®fth species is A. clarki Dunn and Bailey, for which a second specimen is reported from the Colombia ChocoÂ. The noncapitate hemipenis of A. clarki may be primitive in being calyculate and deeply bilobed. The morphologically convergent Geophis is primarily a Middle American genusÐMexico to western Panama, with two or three outlying species in the western Andes of Colombia (G. betaniensis, G. nigroalbus, and probably G. hoffmanni). The genus is unrecorded from eastern Panama, and a few old records for central Panama seem to have been based on erroneous specimen data. Nonetheless, the genus does occur in east-central Panama, based on two spec- imens of G. hoffmanni (W. Peters) and on a specimen each of Geophis bellus, new species, and G. brachycephalus (Cope)Ðthe latter representing a disjunct population separated by about 340 km from those in the Boquete area of western Panama. Geophis bellus is a tiny snake differing from sympatric G. brachycephalus and South Amer- ican G. nigroalbus in characters of size, color, and hemipenis. Geophis brachycephalus may be a composite species in western Panama. Unicolored specimens from the Atlantic versant seem to differ from those in the polymorphic Boquete population in hemipenial and other characters, and they are set aside as a species inquirenda. The ®rst specimen of Geophis hoffmanni is reported from Colombia, but it lacks precise data. Atractus depressiocellus, A. imperfectus, Geophis bellus, G. brachycephalus, and G. hoff- manni are at least broadly sympatric on the ``Piedras-Pacora Ridge''Ðthe continental divideÐ between the upper drainages of the RõÂo Chagres and RõÂo Pacora, some 30 km northeast of Panama City. This relatively low upland likely is a premontane forest refuge, where some very rare snakes may be making a last stand prior to extinction. RESUMEN El geÂnero Atractus esta ampliamente distribuido en SudameÂrica y tiene su limite norte en la regioÂn oriental del Istmo de PanamaÂ, adonde es extraordinariamente raro. ¡Los 5 uÂnicos ejemplares panamenÄos conocidos y estudiados por el autor, colectados a lo largo de 39 anÄos, corresponden a 5 especies! Cuatro son especies nuevas que se describen en este trabajo: A. darienensis, A. hostilitractus, A. imperfectus,yA. depressiocellus. La quinta especie es A. clarki Dunn & Bailey, de la cual se reporta un segundo ejemplar del Choco colombiano. Los hemipenes no capitados de A. clarki podrõÂan ser primitivos por ser caliculados y profunda- mente bilobulados. Geophis, que es morfoloÂgicamente convergente con Atractus, es primariamente un geÂnero mesoamericanoÐse distribuye desde MeÂxico hasta el oeste de PanamaÂ, con dos o tres especies aisladas en los Andes occidentales de Colombia (G. betaniensis, G. nigroalbus, y probable- mente G. hoffmanni). Geophis no ha sido registrado hasta ahora en el este de PanamaÂ, y unos pocos registros viejos del centro de Panama parecen haber sido basados en especõÂmenes de procedencia incierta. Sin embargo, el geÂnero ocurre en el centro-este de PanamaÂ, como lo indican dos especõÂmenes de Geophis hoffmanni (W. Peters), un espeÂcimen de G. bellus (especie nueva), y un espeÂcimen de G. brachycephalus. Este uÂltimo representa una poblacioÂn disyunta, distante aproximadamente 340 km de las poblaciones mas cercanas en el oeste de PanamaÂ. Geophis bellus es una pequenÄita culebra que se diferencia de la especie simpatrida brachy- cephalus y de la sudamericana nigroalbus en caracteres de tamanÄo, coloracioÂn y hemipenes. La especie polimoÂr®ca G. brachycephalus podrõÂa ser una especie compuesta en el oeste de PanamaÂ. Ejemplares monocromaÂticos de la pendiente atlaÂntica parecen diferir en caracterõÂsticas de los hemipenes y de otros caracteres, y son mantenidos como especie inquirenda. Se reporta el primer espeÂcimen de Geophis hoffmanni de Colombia, aunque faltan datos precisos. Atractus depressiocellus, A. imperfectus, Geophis bellus, G. brachycephalus,yG. hoffmanni son ampliamente simpatridas en la ``cresta Piedras-Pacora'', la divisoria de los altos drenajes de los rõÂos Chagres y Pacora, unos 30 km al nordeste de la ciudad de PanamaÂ. Esta serranõÂa 2003 MYERS: SNAKES FROM PANAMA 3 baja es probablemente un refugio de bosque montano, adonde algunas serpientes muy raras podrõÂan presentar una uÂltima resistencia a la extincioÂn. INTRODUCTION albiceps is Barro Colorado Island, which This paper is about some rare snakes on supports one of the world's best known trop- the eastern half of the Isthmus of Panama. ical herpetofaunas (Myers and Rand, 1969; Biologists familiar with the great diversity of Rand and Myers, 1990). Perhaps Tantilla al- serpents in tropical rain forests know that rar- biceps is extinct, at least on Barro Colorado ity or the appearance of rarity is compounded Island. Perhaps biologists on the island have by several factors, including: (1) many, in- failed to differentiate an occasional specimen deed most, wet-forest snakes seem to have from the more common and similar appear- low population densities relative to temperate ing Enuliophis sclateri. Such explanations species; (2) many are hard to ®nd because of are unveri®able and unsatisfactory. secretive habits; and (3) some are less likely Five or six species in the two genera of to be encountered because of small geo- concern in this paper seem to be, like the graphic ranges and/or specialized microhab- example above, really rare, although to be itats. sure they come from less well-trodden With time, however, suf®cient specimens ground. The terrestrial or semifossorial accumulate to allow approximation of geo- snakes of the genus Atractus reach the north- graphic and ecologic distributions and as- ern limits of the generic range in east-central sessment of morphological variation. But Panama. To my knowledge, only ®ve Pana- there are exceptions to this generalizationÐ manian specimens of Atractus have been col- there are snakes seemingly rare in an abso- lectedÐin the years 1936, 1938, 1966, 1967, lute sense, so rare as to resist veri®able ex- 1974Ðwith each specimen a different spe- planation of their rareness. For example, the cies, of which one was named in 1939 and little Tantilla albiceps is known to science the other four are described herein. only from a single specimen that was found The second genus of concern, the semi- in central Panama three-quarters of a century fossorial Geophis, ranges throughout much ago (Barbour, 1925)2; the type locality of T. of Middle America and occurs also in the western Andes of northern South America. 2 PeÂrez-Santos and MartõÂnez (1997: 451) erroneously Geophis is well known from Costa Rica and reported Tantilla albiceps from Isla Coiba off the Paci®c western Panama, but published records for coast of western Panama; this record is based on a spec- imen of the widespread Enulius ¯avitorques. The col- central Panama are based on erroneous lo- lector, Ignacio De la Riva, kindly provided
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