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Zootaxa 1395: 1–31 (2007) www.mapress.com/ zootaxa/

ISSN 1175-5326 (print edition)

Copyright © 2007 · Magnolia Press

ISSN 1175-5334 (online edition)

ZOOTAXA

A checklist and key to the snakes of the genus Geophis (Squamata: Colubridae: Dipsadinae), with commentary on distribution and conservation LARRY DAVID WILSON1 & JOSIAH H. TOWNSEND2 1

Department of Biology, Miami-Dade College, Kendall Campus, Miami, Florida 33176-3393. E-mail: [email protected] School of Natural Resources and Environment, University of Florida, and Division of Herpetology, Florida Museum of Natural History, Gainesville, Florida 32611-7800. E-mail: [email protected] 2

Table of contents Abstract ...............................................................................................................................................................................2 Resumen ..............................................................................................................................................................................2 Introduction .........................................................................................................................................................................2 Acknowledgments ...............................................................................................................................................................3 Checklist ..............................................................................................................................................................................4 Geophis Wagler ...................................................................................................................................................................4 1. Geophis anocularis Dunn ........................................................................................................................................4 2. Geophis bellus Myers ...............................................................................................................................................5 3. Geophis betaniensis Restrepo & Wright ..................................................................................................................5 4. Geophis bicolor Günther ..........................................................................................................................................5 5. Geophis blanchardi Taylor & Smith ........................................................................................................................6 6. Geophis brachycephalus (Cope) ..............................................................................................................................6 7. Geophis cancellatus Smith ........................................................................................................................ ...............7 8. Geophis carinosus Stuart .........................................................................................................................................7 9. Geophis chalybeus Wagler .......................................................................................................................................7 10. Geophis championi Boulenger ...............................................................................................................................8 11. Geophis damiani Wilson, McCranie, & Williams .................................................................................................8 12. Geophis downsi Savage .........................................................................................................................................8 13. Geophis dubius (Peters) .........................................................................................................................................9 14. Geophis duellmani Smith & Holland .....................................................................................................................9 15. Geophis dugesii Bocourt ........................................................................................................................................9 16. Geophis dunni Schmidt ........................................................................................................................................10 17. Geophis fulvoguttatus Mertens ............................................................................................................................10 18. Geophis godmani Boulenger ................................................................................................................................10 19. Geophis hoffmanni (Peters) ..................................................................................................................................10 20. Geophis immaculatus Downs ...............................................................................................................................11 21. Geophis incomptus Duellman ..............................................................................................................................11 22. Geophis isthmicus (Boulenger) ............................................................................................................................11 23. Geophis juarezi Nieto-Montes de Oca .................................................................................................................12 24. Geophis juliai Pérez-Higareda, Smith, & López-Luna ........................................................................................12 25. Geophis laticinctus Smith & Williams .................................................................................................................12 26. Geophis laticollaris Smith, Lynch, & Altig .........................................................................................................13 27. Geophis latifrontalis Garman ...............................................................................................................................13 28. Geophis maculiferus Taylor .................................................................................................................................13 29. Geophis mutitorques (Cope) ................................................................................................................................14

Accepted by S. Carranza: 29 Nov. 2006; published: 22 Jan. 2007

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30. Geophis nasalis (Cope) ........................................................................................................................................14 31. Geophis nephodrymus Townsend & Wilson ........................................................................................................14 32. Geophis nigroalbus Boulenger .............................................................................................................................15 33. Geophis nigrocinctus Duellman ...........................................................................................................................15 34. Geophis omiltemanus (Günther) ..........................................................................................................................15 35. Geophis petersii Boulenger ..................................................................................................................................16 36. Geophis pyburni Campbell & Murphy ................................................................................................................16 37. Geophis rhodogaster (Cope) ................................................................................................................................16 38. Geophis rostralis (Jan) .........................................................................................................................................17 39. Geophis russatus Smith & Williams ....................................................................................................................17 40. Geophis ruthveni Werner ....................................................................................................................... ..............17 41. Geophis sallaei Boulenger ...................................................................................................................................18 42. Geophis semidoliatus (Duméril, Bibron, and Duméril) .......................................................................................18 43. Geophis sieboldi (Jan) ..........................................................................................................................................18 44. Geophis talamancae Lips and Savage .................................................................................................................19 45. Geophis tarascae Hartweg ...................................................................................................................................19 46. Geophis zeledoni Taylor .......................................................................................................................................19 Key to the species of Geophis ...........................................................................................................................................19 Distributional commentary ................................................................................................................................................22 Conservation status of the members of the genus Geophis ...............................................................................................24 Literature cited ..................................................................................................................................................................28

Abstract Geophis Wagler is the fourth largest genus of colubrid snakes in the Western Hemisphere, with 46 species currently recognized. Since the last major treatment of this genus, there have been 12 new species described, three taxa elevated to the species level, and two taxa previously recognized at the species level synonymized with other taxa. We present a checklist for the genus Geophis, summarizing information on taxonomic history, type material and locality, distribution, and species group assignment for 46 species. We also provide an identification key for the 46 species, followed by a brief analysis of distributional patterns and conservation status. Key words: Reptilia, Squamata, Colubridae, Dipsadinae, Geophis, checklist, Mexico, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, Panama, Colombia

Resumen Geophis Wagler es el cuarto género más grande de serpientes de la familia Colubridae en el Hemisferio Occidental, con 46 especies reconocidas actualmente. Desde el último trabajo realizado para este género, 12 nuevas especies han sido descritas, tres taxones se han elevado al nivel de especie, y dos taxones previmante reconocidos como especies se han sinonimizado con otros taxones. Presentamos una lista para el género Geophis, resumiendo la información en la historia taxonómica, material y localidad tipo, distribución, y la asignación a los grupos para 46 especies. También proporcionamos una clave de identificación para las 46 especies, seguidas de un breve análisis de los patrones de distribución y estado de conservación.

Introduction The genus Geophis Wagler 1830 is one of the four most speciose genera of colubrid snakes occurring in the Western Hemisphere, with the other three being Atractus, Rhadinaea, and Tantilla. Köhler (2003) stated that there are approximately 70 described species of Atractus and just over 40 described species of Rhadinaea. Wilson (1999) listed 53 species of Tantilla; three have been described since that paper appeared, and two have been resurrected. We herein include 46 described species of Geophis. Downs (1967) represents the last genus-

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wide treatment of Geophis, in which 33 species were recognized. Since Downs’ (1967) revision, 12 new species have been described, three taxa have been recovered from synonymy to be recognized at the species level, and two of the taxa recognized at the species level by Downs (1967) have been synonymized with other taxa. These 46 species are placed in seven species groups, the recognition of which was established by Downs (1967) in his generic revision. It should be noted, however, that these groups are phenetic in nature, the genus not having been subjected to an explicit phylogenetic analysis, so the presumptive monophyly of these groups has not been tested (Nieto-Montes de Oca 2003). They are recognized as provisional (following Nieto-Montes de Oca 2003) and for the sake of convenience alone. Given the substantial number of changes to the taxonomy of the genus Geophis, we consider it timely to update the information on its content and to provide a new key to the members of this genus, as currently recognized. To these ends, we offer the following paper. Twelve new species have been described in the genus since Downs’ (1967) revision, including: Geophis bellus Myers 2003 Geophis betaniensis Restrepo & Wright 1987 Geophis damiani Wilson, McCranie, & Williams 1998 Geophis downsi Savage 1981 Geophis duellmani Smith & Holland 1969 Geophis juarezi Nieto-Montes de Oca 2003 Geophis juliai Pérez-Higareda, Smith, & López-Luna 2001 Geophis laticollaris Smith, Lynch, & Altig 1965 Geophis nephodrymus Townsend & Wilson 2006 Geophis pyburni Campbell & Murphy 1977 Geophis russatus Smith & Williams 1966 Geophis talamancae Lips & Savage 1994 Two species synonymizations have occurred, as indicated below: Geophis aquilonaris Legler 1959 = Geophis dugesii aquilonaris (Webb 1977: 551) Geophis semiannulatus Smith 1941 = Geophis latifrontalis semiannulatus (Dixon & Thomas 1974: 273) Three species not recognized by Downs (1967) have been resurrected subsequently, as follows: Geophis anocularis Dunn 1920, from synonymy of Geophis dubius (Peters 1861), after Campbell et al. (1983) Geophis nigroalbus Boulenger, 1908, from synonymy of Geophis brachycephalus (Cope 1871), after Myers (2003) Geophis rostralis (Jan [in Jan and Sordelli, 1865]), from synonymy of Geophis dubius (Peters 1861); status in doubt; recognized as distinct from G. dubius by Bogert and Porter (1966) and Campbell et al. (1983), but not by Smith (1959), Downs (1967), Pérez-Higareda and Smith (1988), and Smith and Pérez-Higareda (1991).

Acknowledgments We would like to express our gratitude to Kenneth L. Williams for assistance with literature and for sharing sections of his book on the snake species of the world, and to Colin J. McCarthy (BMNH) for providing inforA CHECKLIST AND KEY TO THE SNAKES OF GEOPHIS

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mation on the type specimen of Geophis nigroalbus.

Checklist Forty-six taxa appear in this checklist. It should be emphasized that G. rostralis is of disputed status (see above). The changes from the information provided by Downs (1967) summarized here are to the date 1 May 2006. The format of the species accounts includes the following sections: species synonymy (including current name and its synonyms); holotype (if known); type locality; distribution (elevational and geographic); species group affiliation; systematic references (containing the most recent and/or comprehensive treatments); remarks (containing systematic information of pertinence, if necessary).

Geophis Wagler Geophis Wagler 1830: 342. Catostoma Wagler 1830: 194. Rabdosoma Duméril, 1853: 440. Colobognathus Peters, 1859: 275. Geophidium Peters, 1861: 923. Colophrys Cope, 1868: 130. Parageophis Bocourt, 1883: 435. Dirosema Boulenger, 1894: 298.

Type-species. Catostoma chalybeum Wagler, 1830, by monotypy (see Remarks). Type-locality. Mexico. Distribution. Geophis occur from 13–2744 m elevation from the Sierra Madre Occidental in southwestern Chihuahua and the Sierra Madre Oriental in southern Tamaulipas, Mexico, through the central highlands and Atlantic lowlands of Central America to the highlands of northern and western Colombia. These snakes occur in vegetation ranging from lowland wet and dry forest, through upland pine-oak forest and wet forest, to highland cloud forest. Content. 46 species. Systematic references. Downs (1967), Smith et al. (1990). Remarks. The name Catostoma Wagler 1830 was first used for this genus, but was quickly substituted with the name Geophis by Wagler (1830) in the same work, in order to avoid confusion with the fish genus Catostomus Lesueur 1817. Smith et al. (1990) further clarified the nomenclatural priority of the name Geophis over Catostoma.

1. Geophis anocularis Dunn Geophis anocularis Dunn 1920: 127. Catostoma anoculare: Amaral 1929: 190. Geophis dubius: Downs 1967: 84.

Holotype. National Museum of Natural History (USNM) 46556, adult male collected 26 July 1894 by E. W. Nelson and E. A. Goldman. Type-locality. Totontepec, Oaxaca, Mexico. Distribution. Known only from 1850-1900 m elevation in the vicinity of the type locality in the Sierra Mije, Oaxaca, Mexico.

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Species group. dubius (Downs 1967; Campbell et al. 1983). Systematic references. Dunn (1920), Downs (1967), Campbell et al. (1983). Remarks. Synonymized with G. dubius by Downs (1967: 84). Resurrected by Campbell et al. (1983), based on the collection of 21 additional specimens from Totontepec, Oaxaca, Mexico.

2. Geophis bellus Myers Geophis bellus Myers 2003: 30.

Holotype. University of Kansas Natural History Museum & Biodiversity Research Center (KU) 110703, adult male, collected 13 December 1964 by C. W. Myers. Type-locality. Near community of Altos de Pacora, a few km northeast of Cerro Jefe (9º15’N, 79º22’W), 700 m elevation, Prov. Panamá, Panama. Distribution. Known only from the type locality. Species group. sieboldi (Myers 2003). Systematic references. Myers (2003).

3. Geophis betaniensis Restrepo & Wright Geophis betaniensis Restrepo & Wright 1987: 191.

Holotype. Universidad del Valle (UV-C) 7360, adult female, collected 18 February 1984 by J. H. Restrepo T. Type-locality. Eastern slope of the Cordillera Occidental, at Corregimiento de Betania (4º23’N, 76º21’W), 1680 m elevation, Municipio de Bolivar, Valle, Colombia. Distribution. Known only from the type locality in the Cordillera Occidental, Valle, Colombia. Species group. sieboldi (Lips & Savage 1994). Systematic references. Restrepo & Wright (1987), Lips & Savage (1994).

4. Geophis bicolor Günther Geophis bicolor Günther 1868: 416. Geophis chalybeus: Bocourt 1883: 530. Dirosema bicolor: Boulenger 1894: 298. R(habdosoma) bicolor: Cope 1885: 385. Catasoma bicolor: Amaral 1929:191.

Syntypes. The Natural History Museum, London (BMNH) 1946.1.1.63-66, collected by H. Doorman. Downs (1967: 55) designated BMNH 1946.1.1.64, an adult male, as the lectotype. Type-locality. “In the neighbourhood of the city of Mexico.” Distribution. Known from the type locality and three localities in Jalisco, Mexico, from 1800–2600 m elevation. Species group. chalybeus (Downs 1967). Systematic references. Downs (1967), Dixon (1968), Smith & Flores Villela (1993). Remarks. Downs (1967: 55) designated BMNH 1946.1.1.64 as the lectotype, as this specimen is the only one of the four syntypes mentioned specifically in the description and was the only adult specimen in a good state of preservation. A CHECKLIST AND KEY TO THE SNAKES OF GEOPHIS

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5. Geophis blanchardi Taylor & Smith Geophis blanchardi Taylor & Smith 1939: 245.

Holotype. Field Museum of Natural History (FMNH) 100037, an adult female collected 14 August 1936 by E. H. Taylor. Type-locality. Around two miles south of Acultzingo, Veracruz, Mexico. Distribution. Known from the vicinity of the type locality in the Veracruz, Mexico around 2300 m elevation, and adjacent parts of Puebla. Also recorded from the Sierra Mazateca in Oaxaca, from 2240–2510 m elevation. Species group. latifrontalis (Downs 1967). Systematic references. Taylor & Smith (1939), Downs (1967). Remarks. The presence of G. blanchardi in the area of Puebla adjacent to the type locality was recently confirmed (Canseco-Márquez and Austin 2005), and this species was also recently reported from a cloud forest locality in northern Oaxaca around 70 km south of the type locality (Canseco-Márquez et al. 2004).

6. Geophis brachycephalus (Cope) Colobognathus brachycephalus Cope 1871: 211. Colobognathus dolichocephalus: Cope 1871: 211. Geophis moestus: Günther 1872: 15. Catastoma chalybaeum: Günther 1872: 16. Catostoma brachycephalum: Cope 1876: 147. Catostoma dolichocephalum: Cope 1876: 147. E(lapoidis) dolichocephalus: Cope 1885: 386. E(lapoidis) brachycephalus: Cope 1885: 386. Rhabdosoma moestum: Cope 1887: 85. Geophis chalybaea quadrangularis: Günther 1893: 87. Geophis dolichocephalus: Günther 1893: 87. Geophis moesta: Günther 1893: 90. Dirosema brachycephalum: Boulenger 1894: 299. Geophis hoffmanni: Boulenger 1894: 319. Catostoma brachycephalum: Amaral 1929: 191. Geophis brachycephalus: Dunn 1942: 4. Geophis dolichocephala: Taylor 1951: 43. Geophis brachycephala: Taylor 1951: 46. Geophis bakeri: Taylor 1954: 689.

Holotype. Academy of Natural Sciences of Philadelphia (ANSP) 3337, juvenile male collected by Van Patten. Type-locality. Costa Rica. Distribution. Known from the Cordillera Central, Cordillera de Talamanca, as well as the northeastern Caribbean and southwestern Pacific lowlands of Costa Rica, and western Panama east to the Piedras-Pacora Ridge in central Panama, 13-2115 m elevation. Species group. sieboldi (Downs 1967). Systematic references. Cope (1871, 1876, 1885), Günther (1872, 1893), Boulenger (1894), Taylor (1951, 1954), Downs (1967), Myers (2003). Remarks. The specific name dolichocephalus actually has priority over brachycephalus in Cope’s (1871) original description. Dunn (1942) selected brachycephalus as the species name without comment, a decision accepted by Downs (1967:152), based at least in part on inaccuracies in Cope’s (1871) description of dolichocephalus. The high degree of color pattern and scale variation in this species has, at least in part, contributed to the confused taxonomic history of this species. Myers (2003) clarified the distribution of G.

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brachycephalus in Panama and reported on a series listed as “Geophis, species inquirenda [G. brachycephalus, auctorum],” that may represent a separate, undescribed taxon.

7. Geophis cancellatus Smith Geophis cancellatus Smith 1941a: 1.

Holotype. National Museum of Natural History (USNM) 46440, female collected February 1896 by Nelson and Goldman. Type-locality. Chicharras, Chiapas, Mexico, ca. 1035 m elevation. Distribution. Known only from two localities on the Pacific versant of Chiapas, Mexico. Species group. semidoliatus (Downs 1967). Systematic references. Smith (1941a), Downs (1967).

8. Geophis carinosus Stuart Geophis carinosus Stuart 1941: 3.

Holotype. University of Michigan Museum of Zoology (UMMZ) 89082, adult male collected 2 August 1940 by L.C. Stuart. Type-locality. Finca San Francisco, 27 km northeast of Nebaj, El Quiché, Guatemala, ca. 1175 m elevation. Distribution. Reported from the Sierra de los Tuxtlas, Veracruz, Mexico, and the Sierra de los Cuchamatanes in El Quiché and Huehuetenango, Guatemala, and adjacent Chiapas, Mexico. Species group. dubius (Downs 1967). Systematic references. Stuart (1941), Downs (1967).

9. Geophis chalybeus Wagler Catastoma chalybeum Wagler 1830: 194. Geophis chalybaeus: Wagler 1830: 342. R(habdosoma) guttulatum Cope 1885: 385. E(lapoidis) chalybaeus: Cope 1885: 386. Geophis chalybaea: Günther 1893: 87. Geophis bicolor: Günther 1893: 91. Dirosema bicolor: Boulenger, 1894: 298. Atractus chalybaeus: Cope 1900: 1232. Catasoma bicolor: Amaral 1929:191. Catostoma chalybaeum: Amaral 1929: 191. Catostoma chalybeum: Slevin 1939: 404.

Holotype. unknown (Downs 1967: 36–40). Type-locality. “Mexico.” The type locality of R. guttulatum Cope is “Vera Cruz.” Distribution. Known only from Mirador, Veracruz, Mexico, ca. 1150 m elevation. Species group. chalybeus (Downs 1967). Systematic references. Wagler (1830), Peters (1859), Cope (1885), Günther (1893), Boulenger (1894), Cope (1900), Amaral (1929), Slevin (1939), Smith (1941a), Downs (1967).

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Remarks. Smith (1941a) placed Rhabdosoma guttulatum Cope in synonymy with G. chalybeus, as R. guttulatum agrees with the descriptions of G. chalybeus supplied by Wagler (1830) and Peters (1859). Downs (1967: 36–40) tentatively accepted this concept of G. chalybeus while detailing the confusing taxonomic history of this species.

10. Geophis championi Boulenger Geophis championi Boulenger 1894: 321. Geophis chalybaea: Günther 1893: 87. Catastoma championi: Amaral 1929: 191. Geophis brachycephalus: Dunn 1942: 4. Holotype. The Natural History Museum, London (BMNH) 1946.1.1.77, collected by J.G. Champion and F.D. Godman. Type-locality. Chiriqui, Panama. Distribution. The type locality simply lists “Chiriqui, Panama”, but Downs (1967) reported an additional specimen (CAS 78977) from Boquete, 1370 m elevation, eastern slope of Volcan Chiriqui, Chiriqui, Panama. Species group. championi (Downs 1967). Systematic references. Boulenger (1894), Dunn (1942), Downs (1967).

11. Geophis damiani Wilson, McCranie, & Williams Geophis damiani Wilson, McCranie, & Williams 1998: 410.

Holotype. National Museum of Natural History (USNM) 498356, adult male collected 26 July 1995 by D. Almendarez, J.R. McCranie, K.L. Williams, and L.D. Wilson. Type-locality. 2.5 km north-northeast La Fortuna (15º26’N, 87º18’W), Yoro, Honduras. Distribution. Known only from the vicinity of the type locality on the slopes of Cerro Texiguat, Honduras. Species group. sieboldi (Wilson et al. 1998). Systematic references. Wilson et al. (1998), McCranie & Castañeda (2004), Townsend (2006).

12. Geophis downsi Savage Geophis downsi Savage 1981: 549.

Holotype. Museum of Natural History of Los Angeles County (LACM) 130254, adult male collected 14-16 July 1972 by N. J. Scott. Type-locality. Las Cruces Field Station and Botanical Garden, 4 km south of San Vito de Jaba, Canton Corredores, Puntarenas, Costa Rica, 1200 m elevation. Distribution. Known only from the vicinity of the type locality in extreme eastern Puntarenas, Costa Rica, from 1100-1200 m elevation. Species group. championi (Savage 1981). Systematic references. Savage (1981).

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13. Geophis dubius (Peters) Geophidium dubium Peters 1861: 923. Geophis dubius: Bocourt 1883: 532. Geophis fuscus: Fischer 1886: 11. Atractus dubius: Cope 1900: 1230. Catostoma dubium: Amaral 1929: 191.

Holotype. Zoologisches Museum, Berlin (ZMB) 4064, an adult female of unknown origin. Type-locality. Unknown. Distribution. At least the pine-oak woodlands of central and southern Oaxaca, Mexico, from 2100–2650 m elevation. Species group. dubius (Downs 1967). Systematic references. Peters (1861), Jan (1865), Fischer (1886), Bogert & Porter (1966b), Downs (1967), Campbell et al. (1983), Smith & Pérez-Higareda (1991).

14. Geophis duellmani Smith & Holland Geophis duellmani Smith & Holland 1969: 47.

Holotype. University of Kansas Natural History Museum & Biodiversity Research Center (KU) 87447, an adult female collected 19 June 1964 by R. L. Holland. Type-locality. Two miles south of Vista Hermosa, 1750 m elevation, Oaxaca, Mexico. Distribution. Known only from the vicinity of the type locality in Oaxaca, Mexico. Species group. dubius (Smith & Holland 1969). Systematic references. Smith & Holland (1969), Smith & Flores Villela (1993).

15. Geophis dugesii Bocourt Geophis dugesii Bocourt 1883: 573. E(lapoidis) dugesi: Cope 1885: 386. Geophis dugesi: Cope 1885: 386. Catastoma dugesii: Amaral 1929: 191. Geophis aquilonaris Legler 1959: 329. Geophis dugesi: Duellman 1961: 96. Geophis dugesii aquilonaris: Webb 1977: 551. Geophis dugesii dugesii: Webb 1977: 551. Geophis dugesi aquilonaris: Vazquez-Diaz & Quintero-Diaz 1999: 235.

Holotype. Museum National d’Histoire Naturelle, Paris (MNHN) 83-278, male collected by O. Navarro and given to A. Dugés. Type-locality. Tangancicuaro, Mexico. Distribution. Known from Aguascalientes, Jalisco, Michoacan, Sinaloa, Durango, and Chihuahua, 15002100 m elevation. Species group. chalybeus (Downs 1967). Systematic references. Bocourt (1883), Smith (1939), Legler (1959), Downs (1967) Webb (1977), Rodriguez Torres & Vásquez Díaz (1990). Remarks. Downs (1967: 62-63) tentatively assigned two specimens from southern Sinaloa to Geophis

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dugesii. Webb (1977: 551) reduced G. aquilonaris to a subspecies of G. dugesii based on a collection of specimens from Durango and Sonora.

16. Geophis dunni Schmidt Geophis dunni Schmidt 1932: 8.

Holotype. Museum of Comparative Zoology (MCZ) 31870, adult female removed from the stomach of a coral snake, Micrurus nigrocinctus (MCZ 17087), collected by W.B. Richardson. Type-locality. Matagalpa, Nicaragua. Distribution. Known only from the type locality. Species group. sieboldi (Downs 1967). Systematic references. Schmidt (1932), Downs (1967), Townsend (2006).

17. Geophis fulvoguttatus Mertens Geophis fulvoguttatus Mertens 1952a: 134.

Holotype. Senckenberg Forschungsinstitut und Naturmuseum (SMF) 43248, juvenile male collected August 1951 by A Zilch. Type-locality. Hacienda Monte Cristo, 2200 m elevation, Cordillera Metapán, Santa Ana, El Salvador. Distribution. Known from 1680-2200 m elevation in Santa Ana, El Salvador, and Ocotepeque and Copán, Honduras. Species group. dubius (Downs 1967). Systematic references. Mertens (1952a,b), Downs (1967), McCranie & Wilson (1991), Townsend (2006).

18. Geophis godmani Boulenger Geophis godmani Boulenger 1894: 322. Catastoma godmani: Amaral 1929: 192.

Syntypes. Two, The Natural History Museum, London (BMNH) 1946.1.6.40-41, collected by F.D. Godman and O. Salvin. Downs (1967: 72) designated BMNH 1946.1.6.41, an adult female, the lectotype. Type-locality. Irazú, Costa Rica. Distribution. Central Costa Rica and western Panama, 1000-2100 m elevation. Species group. championi (Downs 1967). Systematic references. Boulenger (1894), Taylor (1951), Downs (1967), Myers (2003).

19. Geophis hoffmanni (Peters) Colobognathus Hoffmanni Peters 1859: 276. E(lapoides) Hoffmanni: Jan 1862: 21. Geophis hoffmanni: Boulenger 1894: 319. Catastoma hoffmanni: Amaral 1929: 192. Geophis hoffmani: Dunn 1942: 4. Lapsus. Geophis bartholomewi: Brattstrom & Howell 1954: 120.

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Geophis acutirostris: Taylor 1954: 691.

Syntypes. Nine, Zoologisches Museum, Berlin (ZMB) 1868–1870 (five specimens), 4003, 4106 (two specimens), and The Natural History Museum, London (BMNH) 1946.1.6.54, collected by Dr. Hoffmann. Downs (1967: 155) designated ZMB 1870, an adult female, as the lectotype. Type-locality. “Costa Rica.” Distribution. Central and eastern Honduras south through eastern Nicaragua, Costa Rica, and central Panama, and possibly into Colombia, 18-670 m elevation. Species group. sieboldi (Downs 1967). Systematic references.— Peters (1859), Brattstrom and Howell (1954), Taylor (1954), Downs (1967), Savage (2002), Townsend (2006). Remarks. Downs (1967: 158) placed G. acutirostris Taylor in the synonymy of G. hoffmanni, an arrangement later supported by Savage (2002). In the same work (Downs (1967: 158–159), Geophis bartholomewi Brattstrom & Howell was also synonymized with G. hoffmanni.

20. Geophis immaculatus Downs Geophis immaculatus Downs 1967: 90.

Holotype. University of Michigan Museum of Zoology (UMMZ) 107297, adult female, collected by L.C. Stuart. Type-locality. Finca Lorena, ca. 1700 m elevation, Quetzaltenango, Guatemala. Distribution. Known only from the vicinity of the type locality, and from Chiapas, Mexico. Species group. dubius (Downs 1967). Systematic references. Downs (1967). Remarks. This species was recently reported from a cloud forest in central Chiapas, Mexico (Espinosa et al. 1999).

21. Geophis incomptus Duellman Geophis incomptus Duellman 1959: 3.

Holotype. University of Michigan Museum of Zoology (UMMZ) 118840, adult male collected 18 July 1958 by W.E. Duellman and J. Wellman. Type-locality. Dos Aguas, 2100 m elevation, Michoacán, México. Distribution. Known only from the type locality, Sierra de Coalcoman, Michoacan, Mexico. Species group. omiltemanus (Downs 1967). Systematic references. Duellman (1959), Downs (1967).

22. Geophis isthmicus (Boulenger) Atractus isthmicus Boulenger 1894: 307. Rabdosoma zebrinum: Bocourt 1883: 539. Rhegnops zebrinus: Cope 1885b: 178. Geophis zebrina: Gunther 1893: 94. Geophis isthmicus: Dunn 1928: 1. Atractus zebrinus: Amaral 1929: 190. A CHECKLIST AND KEY TO THE SNAKES OF GEOPHIS

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Holotype. Museum National d’Histoire Naturelle, Paris (MNHN) 1984, a juvenile female. Type-locality. “Tehuantepec (Mexique occidental).” Distribution. Unknown, only known from a single specimen from the imprecise type locality. Species group. omiltemanus (Downs 1967). Systematic references. Bocourt (1883), Boulenger (1894), Dunn (1928), and Downs (1967). Remarks. Downs (1967: 122) remarked “the generic allocation of this species remains tentative; it does, however, bear at least superficial resemblances to the members of the omiltemanus group.”

23. Geophis juarezi Nieto-Montes de Oca Geophis juarezi Nieto-Montes de Oca 2003: 574. Geophis carinosus Downs 1967:81.

Holotype. Museo de Zoología of the Facultad de Ciencias, Universidad Nacional Autónoma de México (MZFC) 2236. Adult female, purchased from local collectors in April 1986. Type-locality. Near Metates (17º39’03”N, 96º21’26”W), 900 m elevation, municipality of Santiago Comaltepec, Sierra de Juárez, Oaxaca, Mexico. Distribution. Known only from the vicinity of the type locality, in the Sierra de Juárez, Oaxaca, Mexico. Species group. dubius (Nieto-Montes de Oca 2003). Systematic references. Downs (1967), Nieto-Montes de Oca (2003). Remarks. Nieto-Montes de Oca (2003) described G. juarezi in part based on material previously assigned to G. carinosus by Downs (1967).

24. Geophis juliai Pérez-Higareda, Smith, & López-Luna Geophis juliai Pérez-Higareda, Smith & López-Luna 2001: 42. Geophis semidoliatus: Pérez-Higareda et al. 1987: 16.

Holotype. Estación de Biología Tropical Los Tuxtlas (UNAM-LT) 2775, adult male collected 10 August 1984. Type-locality. Estación de Biología Tropical Los Tuxtlas, 150 m elevation, Veracruz, Mexico. Distribution. Known only from mesic forest in the Los Tuxtlas region of southern Veracruz, Mexico, 150600 m elevation. Species group. semidoliatus (Pérez-Higareda et al. 2001). Systematic references. Pérez-Higareda et al. (2001).

25. Geophis laticinctus Smith & Williams Geophis laticinctus Smith & Williams 1963: 24. Geophis laticinctus albiventris: Smith & Holland 1969: 51.

Holotype. University of Illinois Museum of Natural History (UIMNH) 51352, adult female collected 23 December 1961 by T. MacDougall. Type-locality. Near Colonia Francisco I. Madero, ca. 1800 m elevation, municipality of Jitotol, Chiapas, Mexico. Distribution. Known from 730-1800 m elevation in Chiapas, Mexico.

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Species group. semidoliatus (Downs 1967). Systematic references. Smith & Williams (1963), Smith & Holland (1969), Johnson et al. (1976).

26. Geophis laticollaris Smith, Lynch, & Altig Geophis laticollaris Smith, Lynch, & Altig 1965: 2. Geophis sallaei: Downs 1967: 168.

Holotype. University of Illinois Museum of Natural History (UIMNH) 57170, adult female collected 29 June 1964 by R. Altig. Type-locality. Three miles south of Putla, Oaxaca, Mexico. Distribution. Known only from the type locality in the northern Sierra Madre del Sur, Oaxaca, Mexico. Species group. Unassigned. Geophis laticollaris is apparently closely related to G. sallaei, which is in the sieboldi group (Downs 1967). Systematic references. Smith, Lynch, and Altig (1965), Down (1967), Smith & Chiszar (1992). Remarks. Geophis laticollaris was synonymized with G. sallaei by Downs (1967), but later resurrected by Smith & Chiszar (1992) after examining materials acquired subsequent to publication of Downs’ work.

27. Geophis latifrontalis Garman Geophis latifrontalis Garman 1884: 103. Atractus latifrontalis: Cope 1896: 1023. Geophis semiannulatus: Smith 1941b: 49. Geophis latifrontalis semiannulatus: Dixon & Thomas 1974: 273.

Holotype. Museum of Comparative Zoology (MCZ) 4538, an adult female collected by E. Palmer. Type-locality. From “fifty miles south of San Luis Potosi, Mexico.” Distribution. Hidalgo, Querétaro, San Luis Potosi, and Tamaulipas, Mexico, 1100– 2600 m elevation. Species group. latifrontalis (Downs 1967). Systematic references. Garman (1884), Smith (1941b), Downs (1967), Dixon & Thomas (1974). Remarks. Dixon & Thomas (1974: 273) synonymized G. semiannulatus with G. latifrontalis after collecting a series of specimens from the area between the known ranges of the two taxa.

28. Geophis maculiferus Taylor Geophis maculiferus Taylor 1941: 119.

Holotype. University of Illinois Museum of Natural History (UIMNH) 25078, a juvenile male collected 14 August 1940 by E.H. Taylor. Type-locality. On a road near Cício (= Tzitzio fide Duellman, 1961: 97), ca 1630 m elevation, 17 km south of the Mexico-Guadalajara highway, Michoacán, México. Distribution. Known only from the type locality in the Cordillera Volcanica, Michoacán, México. Species group. omiltemanus (Downs 1967). Systematic references. Taylor (1941), Downs (1967).

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29. Geophis mutitorques (Cope) Rhabdosoma mutitorques Cope 1885: 384. Rhabdosoma longiceps: Cope 1886: 189. Geophis mutitorques: Günther 1893: 93. Atractus longiceps: Boulenger 1894: 305. Geophis longiceps: Dunn 1928:1. Geophis multitorques: Taylor 1949: 194. Lapsus.

Syntypes. Nine, Academy of Natural Sciences of Philadelphia (ANSP) 11324, 14758–65, collected by S. Bernad. Downs (1967: 105) designated ANSP 14762 the lectotype. Type-locality. From “high land about Zacualtipan.” Distribution. Found in cloud forest in San Luis Potosi, Hidalgo, Puebla, and Veracruz, Mexico, above 1500 m elevation. Species group. latifrontalis (Downs 1967). Systematic references. Cope (1885), Smith (1941b), Bogert & Porter (1966b), Downs (1967). Remarks. Downs (1967: 108) synonymized Rhabdosoma longiceps Cope with G. mutitorques.

30. Geophis nasalis (Cope) Catostoma nasale Cope 1868: 131. Elapoides Sieboldi: Müller 1882: 142. R(habdosoma) nasale: Cope 1885: 385. Catostoma chalybeum: Slevin 1939: 404. Geophis nasalis: Smith 1946b: 4.

Syntypes. Unclear, at least four: Academy of Natural Sciences of Philadelphia (ANSP) 3319-21 and National Museum of Natural History (USNM) 12425A, collected by Van Patten. Downs (1967: 160) designated ANSP 3320, an adult female, the lectotype. Type-locality. Near Guatemala City. Distribution. Pacific versant of Chiapas, Mexico to eastern Guatemala, 600-1500 m elevation. Species group. sieboldi (Downs 1967). Systematic references. Cope (1868), Slevin (1939), Smith (1941b), Downs (1967).

31. Geophis nephodrymus Townsend & Wilson Geophis nephodrymus Townsend & Wilson, 2006: 151.

Holotype. Florida Museum of Natural History (UF) 142577, a female collected 11 July 2004 by S. M. Hughes and J. H. Townsend. Type-locality. Sendero Las Minas (15º29.525’N, 88º12.705’W), 1580 m elevation, Parque Nacional El Cusuco, Cortés, Honduras. Distribution. Known only from the vicinity of the type locality in the Sierra de Omoa, Cortés, Honduras. Species group. dubius (Townsend & Wilson 2006). Systematic references. Townsend (2006), Townsend & Wilson (2006).

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32. Geophis nigroalbus Boulenger Geophis nigro-albus Boulenger 1908: 552. Catastoma nigroalbum: Amaral 1929: 192. Geophis brachycephalus: Downs 1967: 146. Geophis negroalbus: Myers 2003: 32. Lapsus.

Holotype. The Natural History Museum, London (BMNH) 1946.1.6.50 (formerly BMNH 1909.4.30.71), juvenile, collected by N.G. Palmer, purchased from Rosenberg. Type-locality. Near Pavas, Valle, Colombia. Distribution. Reported from 900-1700 m elevation in Antioquia, Santander, and Valle de Cauca, Colombia. Species group. sieboldi (Downs 1967). Systematic references. Boulenger (1908), Downs (1967), Restrepo & Wright (1987), Myers (2003). Remarks. Downs (1967) placed G. nigroalbus in synonymy with G. brachycephalus, but in a footnote on p. 146 stated: “A cursory look at a specimen recently obtained in eastern Panama by Charles Myers leads me to doubt that my inclusion of G. nigroalbus Boulenger in the synonymy of G. brachycephalus is justified.” Restrepo & Wright (1987:193) applied the name G. nigroalbus to more recent material from Colombia based on this footnote. The specimen that Downs (1967:146) referred to was the then-undescribed Geophis bellus Myers, and later Myers (2003: 34) tentatively assigned two additional Colombian specimens previously referred to as G. brachycephalus to G. nigroalbus, noting “I have seen too few Colombian specimens to be able to fully characterize Geophis nigroalbus, or even to conclude that but a single species is represented.”

33. Geophis nigrocinctus Duellman Geophis nigrocinctus Duellman 1959: 1.

Holotype. University of Michigan Museum of Zoology (UMMZ) 118841, male collected 18 June 1958 by J. Wellman. Type-locality. Dos Aguas, Michoacán, México, 2100 m elevation. Distribution. Known from the type locality, Sierra de Coalcoman, Michoacán, Mexico, and from the Sierra de Manantlán, Jalisco, Mexico, from 1900 m elevation. Species group. chalybeus (Downs 1967). Systematic references. Duellman (1959), Downs (1967). Remarks. Flores-Villela et al. (1995) recorded G. nigrocinctus from a pine forest in Jalisco, Mexico.

34. Geophis omiltemanus (Günther) Geophis omiltemana Günther 1893: 92. Dirosema omiltemanum: Boulenger 1894: 299. Atractus omiltemanus: Cope 1896: 1024. Catastoma omiltemanum: Amaral 1929: 192. Geophis omiltemana: Smith 1941b: 1. Geophis omiltemanus: Downs 1967: 124.

Syntypes. Three, The Natural History Museum, London (BMNH) 1946.1.6.35-37 collected by H.H. Smith. Downs (1967:124) designated 1946.1.6.37, an adult male, the lectotype. Type-locality. Omilteme, Guerrero, Mexico, ca. 2438 m elevation. A CHECKLIST AND KEY TO THE SNAKES OF GEOPHIS

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Distribution. Known from the vicinity of the type locality, in the Sierra Madre del Sur, Guerrero, Mexico, and from western Oaxaca. Species group. omiltemanus (Downs 1967). Systematic references. Günther (1893), Downs (1967), Hardy (1976). Remarks. Campbell (1988:28) reported this species from San Andrés Chicahuaxtla, Oaxaca, Mexico, based on a personal communication from C. H. Bogert.

35. Geophis petersii Boulenger Geophis petersii Boulenger 1894: 321. Catastoma petersii: Amaral 1929: 192. Geophis petersi: Duellman 1961: 98.

Syntypes. Two, The Natural History Museum, London (BMNH) 1946.1.6.31-32, collected by Doorman. Downs (1967: 164) designated BMNH 1946.1.6.31, a male, the lectotype. Type-locality. City of Mexico, Mexico. Distribution. From the type locality and Michoacan, Mexico, 1800-2000 m elevation. Species group. sieboldi (Downs 1967). Systematic references. Boulenger (1894), Hartweg (1959), Downs (1967).

36. Geophis pyburni Campbell & Murphy Geophis pyburni Campbell & Murphy 1977: 397.

Holotype. University of Texas at Arlington Collection of Vertebrates (UTA) R-4404, female, collected 8 July 1974 by J. A. Campbell. Type-locality. Rancho La Pastilla, ca. 2.5 km west of Dos Aguas, Sierra de Coalcomán, Michoacán, Mexico, 2164 m elevation. Distribution. Known only from the type locality in the Sierra de Coalcomán, Michoacán, Mexico. Species group. sieboldi (Campbell & Murphy 1977; by inference). Systematic references. Campbell & Murphy (1977).

37. Geophis rhodogaster (Cope) Colophrys rhodogaster Cope 1868: 130. Geophis rhodogaster: Bocourt 1883: 531. Geophis chalybaea: Günther 1893: 87. Catastoma rhodogaster: Amaral 1929: 192. Geophis fulvoguttatus: Köhler 1996: 36 (in error).

Syntypes. Three: Academy of Natural Sciences of Philadelphia (ANSP) 3316-17 and National Museum of Natural History (USNM) 12425, collected by Van Patten. Downs (1967: 92) designated ANSP 3317, an adult female, the lectotype. Type-locality. “…elevated country in the neighborhood of the city of Guatemala.” Distribution. The mountains of southwestern Chiapas, Mexico, east through southern Guatemala to the Cordillera Metapán, Santa Ana, El Salvador, 1500-2744 m elevation.

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Species group. dubius (Downs 1967). Systematic references. Cope (1868), Downs (1967), Smith (1995), Townsend (2006). Remarks. Günther (1893) reported G. rhodogaster from Costa Rica. Boulenger (1894) apparently followed Günther (1893) and included Costa Rica in the range of G. rhodogaster. Taylor (1951) also included G. rhodogaster in his review of Costa Rica snakes, but noted “I regard this record as somewhat doubtful.” Köhler (1996) gave measurements and scutellation data for a specimen of G. rhodogaster (SMF 77413) under the name G. fulvoguttatus.

38. Geophis rostralis (Jan) E(lapoides) rostralis Jan (in Jan & Sordelli) 1865: plate 2, figure 2. Geophis rostralis, Bocourt, 1883: 533. R(habdosoma) rostrale, Cope 1885: 385. Catostoma rostrale, Amaral, 1929: 192. Geophis dubius, Downs 1967: 84.

Holotype. Zoologisches Museum, Berlin (ZMB) 6407 (Downs 1967). Type-locality. Unknown. Distribution. Apparently restricted to upper elevations of the Sierra Madre del Sur, Oaxaca, Mexico. Species group. dubius (Downs 1967). Systematic references. Bocourt (1883), Webb & Fulger (1957), Smith (1959), Bogert & Porter (1966b), Downs (1967), Smith & Pérez-Higareda (1991). Remarks. There is disagreement about the specific status of this taxon, which was originally described based on a color plate with no indication of a type specimen or locality. Various authors have recognized G. rostralis as a distinct species (Bogert and Porter 1966; Campbell et al. 1983), while others (Smith 1959; Downs 1967; Pérez-Higareda & Smith 1988; Smith & Pérez-Higareda; 1991) maintain that this species is conspecific with G. dubius.

39. Geophis russatus Smith & Williams Geophis sallaei russatus Smith & Williams 1966: 90. Geophis russatus: Downs 1967: 138.

Holotype. University of Illinois Museum of Natural History (UIMNH) 61409, adult male collected 25 May 1965 by T. MacDougall. Type-locality. La Concepción, near Putla, Oaxaca, México. Distribution. Known only from the type locality in the Sierra de Miahuatlán, Oaxaca, México. Species group. sieboldi (Downs 1967). Systematic references. Smith & Williams (1966), Downs (1967). Remarks. Downs (1967: 138) elevated the name russatus to species level in a footnote.

40. Geophis ruthveni Werner Geophis ruthveni Werner 1925: 60. Catastoma ruthveni: Amaral 1929: 193.

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Holotype. Naturhistorischen Museums Wien (NHMW) 16508, male. Type-locality. “Sarapigui, Brasilien.” Probably refers to Sarapiquí, Heredia, Costa Rica (See Remarks). Distribution. North-central Costa Rica, 85-1600 m elevation. Species group. championi (Downs 1967). Systematic references. Werner (1925), Savage (1960), Downs (1967). Remarks. Downs (1967: 78) agreed with Savage (1960) and Dunn (in Savage, 1960: 31) that the type locality of “Sarapigui,” Brazil is in error and means Sarapiquí, Costa Rica.

41. Geophis sallaei Boulenger Geophis sallaei Boulenger 1894: 318. Catastoma sallaei: Amaral 1929: 193. Geophis sallaei russatus: Smith & Williams 1966: 90. Geophis laticollaris: Downs 1967: 168. Geophis sallei: Smith & Chiszar 1992: 16.

Syntypes. Three, The Natural History Museum, London (BMNH) 1946.1.6.26-27, 1946.1.6.49, collected by M. Sallé. Downs (1967:168) designated BMNH 1946.1.6.27, a female, the lectotype. Type-locality. Mexico. Distribution. Sierra Madre del Sur, Oaxaca, Mexico, ca 1400 m elevation. Species group. sieboldi (Downs 1967). Systematic references. Boulenger (1894), Downs (1967), Smith & Chiszar (1992).

42. Geophis semidoliatus (Duméril, Bibron, and Duméril) Rabdosoma semi-doliatum Duméril, Bibron & Duméril 1854: 93. Catastoma semidoliatum: Cope 1860: 339. E(lapoides) semidoliatus: Jan 1862: 21. G(eophis) semidoliatus: Peters 1859: 276. Geophis (Parageophis) semidoliatus: Bocourt 1883: 534. Geophis semidoliata: Günther 1893: 90. Atractus semidoliatus: Cope 1900: 1230.

Syntypes. Originally five, Museum National d’Histoire Naturelle, Paris (MNHN) 3313 (1–3), 4522, fifth syntype apparently lost. Downs (1967: 133) designated MNHN 4522, an adult female, the lectotype. Type-locality. Mexico. Distribution. Central and southern Veracruz, Mexico, 500-1400 m elevation. Species group. semidoliatus (Downs 1967). Systematic references. Duméril et al. (1854), Downs (1967).

43. Geophis sieboldi (Jan) Elapoides Sieboldi Jan 1862: 21. C(atastoma) sieboldii: Cope 1868: 131. Ninia sieboldi: Garman 1884: 96. E(lapoidis) sieboldii: Cope 1885: 386. Geophis sieboldi: Smith 1941a: 4.

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Syntypes. Originally four syntypes, two deposited in a museum collection in Milan, one in Vienna, and one in Munich. None are known to be extant (Downs 1967: 171). Type-locality. “Mexico” and “Guatemala.” Distribution. Unclear, two specimens referred to G. sieboldi are reported from Almolonga, Guerrero, and Coalcoman, Michoacan, Mexico (Downs 1967). Species group. sieboldi (Downs 1967). Systematic references. Jan (1862), Jan & Sordelli (1865), Garman (1884), Smith (1941a), Downs (1967).

44. Geophis talamancae Lips and Savage Geophis talamancae Lips & Savage 1994: 410.

Holotype. Costa Rica Expeditions (CRE) 5343, adult female collected 1 September 1992 by K.R. Lips. Type-locality. Finca Jaguar, 1800 m elevation, Zona Protectora Las Tablas, Cantón Coto Brus, Puntarenas, Costa Rica. Distribution. Known only from the type locality. Species group. sieboldi (Lips & Savage 1994). Systematic references. Lips & Savage (1994).

45. Geophis tarascae Hartweg Geophis tarascae Hartweg 1959:1.

Holotype. University of Michigan Museum of Zoology (UMMZ) 99151, male collected 18 July 1947 by N. Hartweg. Type-locality. The “Parque Nacional” outside of Uruapan, Michoacán, Mexico, ca. 1676 m elevation. Distribution. Known from 1400-1700 m elevation in Jalisco and Michoacán, Mexico Species group. chalybeus (Downs 1967). Systematic references. Downs (1967), Dixon (1968), Medica et al. (1975).

46. Geophis zeledoni Taylor Geophis zeledoni Taylor 1954: 693.

Holotype. University of Kansas Natural History Museum & Biodiversity Research Center (KU) 31992, collected 24 July 1952 by E.H. Taylor. Type-locality. Finca Zeledón, ca. 1829 m elevation, between Volcán Barba and Volcán Poás, Costa Rica. Distribution. Known from the Cordillera Central, Costa Rica, ca. 1829-2100 m elevation. Species group. sieboldi (Downs 1967). Systematic references. Taylor (1954), Downs (1967).

Key to the species of Geophis This key to identification is original to this paper, but is based on information in Downs (1967), as well as a number of other published sources otherwise cited in this paper.

1 -

Anterior temporal present ............................................................................................................................ 2 Anterior temporal absent ............................................................................................................................. 7

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2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 -

Dorsal scales in 15 rows throughout body................................................................................................... 3 Dorsal scales in 17 rows throughout body................................................................................................... 4 Ventrals immaculate................................................................................................................ G. maculiferus Ventrals each divided into dark anterior portion and pale posterior portion ............................ G. incomptus Postoculars 2 ................................................................................................................................................ 5 Postocular single .......................................................................................................................................... 6 Dorsum pinkish yellow with series of irregular, transverse brown spots; venter pale with numerous reddish brown spots .........................................................................................................................G. isthmicus Dorsum reddish brown with narrow pale crossbands; venter immaculate creamy white .... G. omiltemanus Venter of body white to cream anteriorly grading posteriorly to grayish brown with pale posterior borders on ventrals............................................................................................................................... G. latifrontalis Venter of body black, checkered with red or yellow blotches in juvenile .............................. G. mutitorques Dorsal scales in 17 rows throughout body................................................................................................... 8 Dorsal scales in 15 rows throughout body................................................................................................. 23 Postoculars absent........................................................................................................................................ 9 Postoculars 1 or 2....................................................................................................................................... 10 Dorsum slate black ...................................................................................................................G. anocularis Dorsum red or white with large black dorsal saddles ................................................................ G. duellmani Postoculars 2 .................................................................................................................................. G. bicolor Postocular single ........................................................................................................................................ 11 Dorsal scales smooth throughout body or smooth except above vent, where faintly keeled .................... 12 Dorsal scales distinctly keeled on portions of body ranging from posterior portion of body to all of body except nape ................................................................................................................................................ 18 Venter immaculate white, cream, or yellow, except perhaps for darkly pigmented lateral edges ............. 13 Venter checkered or banded....................................................................................................................... 17 Internasal scales usually fused with prefrontal scales .................................................................... G. dubius Internasal scales not fused with prefrontal scales...................................................................................... 14 No supraocular scale present, frontal scale enters orbit......................................................... G. rhodogaster Supraocular scale usually present, usually distinct from postocular, frontal scale does not enter orbit.... 15 Dorsum brown, with scales in lateral rows spotted cream ........................................................ G. chalybeus Dorsum gray, brown, or blackish with pale gray to brick-red dorsal or lateral blotches or partial bands. 16 135–157 ventral scales, 171–181 ventrals + subcaudals ...................................................... G. fulvoguttatus 120–138 ventral scales, 149–160 ventrals + subcaudals ..................................................... G. nephodrymus Venter checkered with yellowish orange and black..................................................................G. blanchardi Ventrals pale brown anteriorly and yellowish white posteriorly .......................................... G. immaculatus Dorsal scales keeled except on neck.......................................................................................................... 19 Dorsal scales keeled only on the posterior portion of body....................................................................... 20 Dorsum yellowish with dark brown crossbands ...............................................................................G. dunni Dorsum dark brown or gray without markings...............................................................................G. nasalis Ventral scales brownish black anteriorly, yellowish white posteriorly...................................... G. carinosus Ventral surface immaculate........................................................................................................................ 21 Ventrals 124 or fewer...................................................................................................................... G. juarezi Ventrals 126 or more.................................................................................................................................. 22 Ventrals 132 or fewer; dorsum uniform gray................................................................................G. rostralis Ventrals 132 or more; dorsum brown, scales of first row with yellow centers............................. G. sieboldi Postoculars 2 ............................................................................................................................ G. betaniensis Postocular single or absent ........................................................................................................................ 24

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24 Dorsal scales weakly to distinctly keeled on portions of body ranging from posterior one-half to all except neck............................................................................................................................................................ 25 - Dorsal scales smooth throughout body or smooth except above vent....................................................... 33 25 Postocular scale absent ...................................................................................................................G. downsi - Postocular scale single ............................................................................................................................... 26 26 Dorsal body scales weakly to distinctly keeled except on neck ................................................................ 27 - Dorsal body scales weakly to distinctly keeled only on posterior portion ................................................ 29 27 Venter with dark pigment usually present along anterior edges of at least some ventrals; dorsum often with pale lateral blotches, crossbands, or stripes ......................................................................G. brachycephalus - Venter yellowish white, although some ventrals may have brownish pigment along anterior edges; dorsum without pale lateral blotches, crossbands, or stripes.................................................................................. 28 28 Pale nuchal collar absent................................................................................................................. G. sallaei - Broad white nuchal collar present ........................................................................................... G. laticollaris 29 Dorsal and ventral surfaces uniformly dark..................................................................................... G. bellus - Coloration not as above ............................................................................................................................. 30 30 Anterior edge of each ventral dark brown or black, posterior edge white....................................G. ruthveni - Ventral coloration not as above.................................................................................................................. 31 31 Supraocular and postocular scales separated from one another by anterior projection of parietal scale ...... ..................................................................................................................................................G. nigroalbus - Supraocular and postocular scales in contact with one another................................................................. 32 32 Dorsum reddish brown with irregular black crossbars .................................................................G. russatus - Dorsum uniform dark gray to black........................................................................................ G. talamancae 33 Supralabials 5............................................................................................................................................. 34 - Supralabials 6............................................................................................................................................. 35 34 Dorsum with dark dorsal crossbands separated by pale interspaces .....................................G. semidoliatus - Dorsum dark brown or grayish black ........................................................................................G. hoffmanni 35 Venter black with scattered, irregular pale blotches or mottled areas ....................................................... 36 - Venter not colored as above ....................................................................................................................... 37 36 Dorsum blackish with well-defined yellowish-red crossbands ................................................ G. laticinctus - Dorsum uniform gray or black, sometimes with pale lateral stripe and narrow light bands........G. zeledoni 37 Dorsum patterned....................................................................................................................................... 38 - Dorsum patternless .................................................................................................................................... 44 38 Dorsum of body dark with pale markings ................................................................................................. 39 - Dorsum of body pale with dark markings ................................................................................................. 40 39 Dorsum of body dark gray with reddish orange crossbands or lateral blotches ...........................G. damiani - Dorsum brown or black, with narrow pale crossbands on anterior portion of body ..................... G. dugesii 40 Venter immaculate white or cream ............................................................................................................ 41 - Venter not as above .................................................................................................................................... 42 41 Dorsum cream with reddish brown crossbands .......................................................................G. cancellatus - Dorsum red with black crossbands ................................................................................................... G. juliai 42 Venter black with scattered pale blotches ................................................................................. G. laticinctus - Venter white with black spots .................................................................................................................... 43 43 Dorsum with narrow dark crossbands on a gray-brown ground color .................................. G. nigrocinctus - Dorsum with irregular dark markings on head and neck, remainder of body dark brown or black ............. ..................................................................................................................................................... G. tarascae 44 Supraocular absent, parietal scale in contact with prefrontal scale ............................................. G. godmani - Supraocular scale present, parietal scale not in contact with prefrontal scale........................................... 45

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45 46 -

Venter banded, anterior edge brown, posterior edge white....................................................... G. championi Venter immaculate white to creamy white................................................................................................. 46 Ventrals plus subcaudals 171 or fewer.......................................................................................... G. pyburni Ventrals plus subcaudals 175 or more............................................................................................ G. petersii

Distributional commentary The genus Geophis is distributed from the Sierra Madre Occidental in southwestern Chihuahua and the Sierra Madre Oriental in southern Tamaulipas, Mexico, through Central America to northern and western Colombia. Thus, eight countries are occupied by members of the genus, including Mexico, six of the seven countries of Central America, and the South American country of Colombia (Table 1). Geophis, therefore, is largely a Middle American genus, save for two species distributed in Colombia (see below). Mexico is home to 32 species (Table 1), 26 of which are endemic (G. anocularis, G. bicolor, G. blanchardi, G. chalybeus, G. dubius, G. duellmani, G. dugesii, G. incomptus, G. isthmicus, G. juarezi, G. juliai, G. laticinctus, G. laticollaris, G. latifrontalis, G. maculiferus, G. mutitorques, G. nigrocinctus, G. omiltemanus, G. petersii, G. pyburni, G. rostralis, G. russatus, G. sallaei, G. semidoliatus, G. sieboldi, and G. tarascae), four of which are shared with Guatemala (G. cancellatus, G. carinosus, G. immaculatus, and G. nasalis), and one of which is shared with Guatemala and El Salvador (G. rhodogaster). Eighteen species are distributed in Central America (Table 1). Six species occur in Guatemala, four of which are shared with Mexico (see paragraph above), one of which is shared with Mexico and El Salvador (see paragraph above) and one of which is shared with El Salvador and Honduras (G. fulvoguttatus). Only two species are distributed in El Salvador, both of which are shared with Guatemala (one of these is also shared with Mexico; see paragraph above) and one of which also occurs in Honduras (see above). Four species are known from Honduras, one of which is shared with Guatemala and El Salvador (G. fulvoguttatus), another of which is shared with Nicaragua, Costa Rica, and Panama (G. hoffmanni), and two of which are endemic (G. damiani and G. nephodrymus). Two species are known from Nicaragua, one of which is shared with Honduras, Costa Rica, and Panama (G. hoffmanni) and the other of which is endemic (G. dunni). Costa Rica hosts seven species, four of which are endemic (G. downsi, G. ruthveni, G. talamancae, and G. zeledoni), two of which are shared with Panama (G. godmani and G. brachycephalus), and one of which is shared with Honduras, Nicaragua, and Panama (G. hoffmanni). Panama is home to five species, two of which are endemic (G. bellus and G. championi), two of which are shared with Costa Rica (see above), and one of which is shared with Honduras, Nicaragua, and Costa Rica (see above). Colombia has two species, both of which are endemic (G. betaniensis and G. nigroalbus). These are the only two species in the genus extralimital to Middle America (see above). Mexico is the clear center of diversity in the genus Geophis, with 31 of 46 species (67.4%) occurring there. In addition, of the 37 species endemic to a given country, 26 (70.3%) are found only in Mexico. Moreover, of the seven recognized species groups in the genus, six contain species found in Mexico (chalybeus, dubius, latifrontalis, omiltemanus, semidoliatus, and sieboldi groups) and four of these groups (chalybeus, latifrontalis, omiltemanus, and semidoliatus groups) are restricted in distribution to Mexico. With respect to elevation (Table 1), members of the genus Geophis are distributed from 13 m (G. brachycephalus) to 2744 m (G. rhodogaster). Seven species are distributed at low elevations, and only one species (G. juliai) is restricted to these elevations (Table 1). Twenty species are distributed at moderate elevations, with 11 (55%) restricted to these elevations (Table 1). Finally, 31 species occur at intermediate elevations, with 24 (77.4%) restricted to such elevations (Table 1). In summary, the greatest number of species occurs at intermediate elevations, with the next largest number occupying moderate elevations; the genus is poorly represented at low elevations.

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TABLE 1. Distribution of Geophis by country and elevation. Mex = Mexico, Guat = Guatemala, ES = El Salvador, Hon = Honduras, Nica = Nicaragua, CR = Costa Rica, Pan = Panama, Col = Colombia, Low = Low (0-600 m), Mod = Moderate (601-1500 m), Int = Intermediate (1501-2700 m).

Country Species

Mex

G. anocularis

X

Guat

ES

Hon

Nica

Elevation CR

Pan

Col

Low

Mod

Int X

G. bellus

X

G. betaniensis

X X

X

G. bicolor

X

X

G. blanchardi

X

X

G. brachycephalus

X

X

X

X

X

G. cancellatus

X

X

X

X

X

G. carinosus

X

X

X

X

X

G. chalybeus

X

X

G. championi

X

G. damiani

X

X

X

G. downsi

X

X

G. dubius

X

X

G. duellmani

X

X

G. dugesii

X

X

G. dunni

X

G. fulvoguttatus

X

X

X

X

X

G. godmani G. hoffmanni

X

X

X

X

X

X

X

X X

X

X

G. immaculatus

X

X

X

G. incomptus

X

G. isthmicus

X

G. juarezi

X

G. juliai

X

G. laticinctus

X

G. laticollaris

X

X

G. latifrontalis

X

X

G. maculiferus

X

X

G. mutitorques

X

X

G. nasalis

X

X X X X

X

X

X

G. nephodrymus

X

G. nigroalbus

X X

X

G. nigrocinctus

X

X

G. omiltemanus

X

X

G. petersii

X

X

G. pyburni

X

X

G. rhodogaster

X

G. rostralis

X

X

X

X

X X

to be continued.

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TABLE 1 (continued). Country Species

Mex

G. russatus

X

Guat

ES

Hon

Nica

Pan

Col

X

G. sallaei

X

G. semidoliatus

X

G. sieboldi

X

Low

Mod

X

X

X

X X

X

X

X

X

G. zeledoni

X 31

Int

X

G. talamancae

Totals

CR

X

G. ruthveni

G. tarascae

Elevation

6

2

4

2

7

X 5

2

7

20

31

Conservation status of the members of the genus Geophis As noted above, the genus Geophis is one of the largest colubrid snake genera in the Western Hemisphere. It is also essentially a Middle American endemic, with only two species known to occur outside this area. Its members tend to have restricted ranges, creating substantial endemism at the species level (see previous section). Only nine species of 46 (19.6%) have ranges extending beyond the limits of a single country. The prevailing distributional pattern, therefore, is one of the limitation of most species (80.4% of 46) to a single country, with only a few outside of these limits. It is for these reasons that Geophis is of considerable conservation interest. It is the purpose of this section of the paper to examine the conservation status of the members of the genus. In carrying out this assessment, we have used a combination of the patterns of occupancy of physiographic regions (Table 2) and those for vegetational zones (Table 3) to establish a set of categories of vulnerability to environmental damage. In doing so, we make the assumption that the more restricted the physiographic and/or vegetational distribution, the more vulnerable the species and vice versa. Given this assumption, it is possible to erect three categories from least to most vulnerable. These categories and their species composition are as follows: Category 1. Least vulnerable (occurrence in 3-6 physiographic regions and 3-5 vegetation formations) G. brachycephalus—3/5 G. carinosus—5/4 G. dugesii—3/3

G. dubius—4/3 G. hoffmanni—6/4 G. semidoliatus—4/4

Category 2. Intermediately vulnerable (occurrence in 1-2 physiographic regions and 1–6 vegetation formations) G. cancellatus—1/3 G. laticinctus—2/4 G. nasalis—1/6 G. sieboldi—1/3

G. rhodogaster—2/2 G. russatus—2/1 G. ruthveni—2/2

Category 3. Most vulnerable (occurrence in 1 physiographic region and 1-2 vegetation formations) G. anocularis—1/1

G. juarezi—1/1

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G. bellus—1/1 G. betaniensis—1/1 G. bicolor—1/1 G. blanchardi—1/1 G. chalybeus—1/1

G. juliai—1/2 G. laticollaris—1/1 G. latifrontalis—1/1 G. maculiferus—1/1 G. mutitorques—1/2

TABLE 2. Distribution of species of Geophis in physiographic regions. Abbreviations of regions are: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama; CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to southwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras; HN = eastern nuclear Central American highlands; LT = Sierra de Los Tuxtlas; MC = Meseta Central; NP = Caribbean lowlands from Nicaragua to Panama; OCC = Sierra Madre Occidental; ORI = Sierra Madre Oriental; SC = Pacific lowlands from Sinaloa to western Chiapas; SUR = Sierra Madre de Sur; TT = Gulf lowlands from Tamaulipas to Tabasco; YP = Yucatan Platform. Middle American highlands Species

O O RI C C

M C

Geophis anocularis

S U R

L T

C G

H N

C E R P P

Atlantic coast lowlands T T

Y G P H

N P

Pacific coast lowlands S C

C G U

G C R

Northern Andes C P

X

Geophis bellus

X

Geophis betaniensis

X

Geophis bicolor Geophis blanchardi

X X

Geophis brachycephalus

X

Geophis cancellatus Geophis carinosus

X

X

X

X

X

Geophis damiani

X

Geophis downsi

X X

X

Geophis duellmani

X

X

X X

X

X

Geophis dunni

X

Geophis fulvoguttatus

X

Geophis godmani

X

Geophis hoffmanni

X

Geophis immaculatus Geophis incomptus

X

X

X

X

X

X X

Geophis isthmicus Geophis juarezi

X

X

Geophis championi

Geophis dugesii

X

X

Geophis chalybeus

Geophis dubius

X

X X to be continued...

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TABLE 2 (continued). Middle American highlands Species

O O RI C C

M C

S U R

Geophis juliai

L T

H N

C E R P P

T T

Y G P H

N P

Pacific coast lowlands S C

C G U

G C R

X

Geophis laticollaris

X

X X

Geophis mutitorques

X

Geophis nasalis

X

Geophis nephodrymus

X

Geophis nigroalbus

X

Geophis nigrocinctus

X

Geophis omiltemanus

X

Geophis petersi

X

Geophis pyburni

X

Geophis rhodogaster

X

Geophis rostralis

X

X X

X

Geophis ruthveni

X

X

Geophis sallaei Geophis semidoliatus

X X

X

X

X

Geophis sieboldi

X

Geophis talamancae

X

Geophis tarascae

X

Geophis zeledoni Totals

G. championi—1/1 G. damiani—1/1 G. downsi—1/1 G. duellmani—1/1 G. dunni—1/1 G. pyburni—1/1 G. rostralis—1/1 G. sallaei—1/2 G. talamancae—1/1 G. tarascae—1/1 G. zeledoni—1/1

C P

X

Geophis maculiferus

Geophis russatus

Northern Andes

X

Geophis laticinctus

Geophis latifrontalis

C G

Atlantic coast lowlands

X 6

1

10 10 3

6

6

8

0

3

1

1

3

5

0

1

3 2

G. nephodrymus—1/1 G. nigroalbus—1/1 G. nigrocinctus—1/1 G. omiltemanus—1/1 G. petersii—1/1 G. fulvoguttatus—1/2 G. godmani—1/2 G. immaculatus—1/1 G. incomptus—1/1 G. isthmicus—1/1

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Thus, there are 6 category 1 species (13.1% of 46 species), 7 category 2 species (15.2%), and 33 category 3 species (71.7%). In general, most species of Geophis are narrowly distributed, increasing their susceptibility to habitat fragmentation and destruction, as well as other types of deleterious environmental impact. The typical member of the genus is limited in distribution to one country, one physiographic region, and one or two vegetation formations. TABLE 3. Distribution of Species of Geophis in Vegetation Formations. Abbreviations of formations are: TWM = Tropical Wet Forest and Tropical Moist Forest; TD = Tropical Dry Forest; TVD = Tropical Very Dry Forest; SRW = Subtropical Rainforest and Subtropical Wet Forest; SM = Subtropical Moist Forest; SD = Subtropical Dry Forest; LMW = Lower Montane Wet Forest; LMM = Lower Montane Moist Forest; LMD = Lower Montane Dry Forest; MW = Montane Wet Forest; MM = Montane Moist Forest. Vegetation Formations Species

TWM TD

TVD

SRW SM

SD

Geophis anocularis Geophis bellus

LMW LMM LMD MW

MM

X

Totals 1

X

1

Geophis betaniensis

X

Geophis bicolor

1 X

Geophis blanchardi

1

X

Geophis brachycephalus

X

X

Geophis cancellatus

X

X

Geophis carinosus

X

X

X

X

1 X

5

X X

3

X

4

Geophis chalybeus

X

1

Geophis championi

X

1

Geophis damiani

X

Geophis downsi

X

Geophis dubius

1 X

X

Geophis duellmani Geophis dugesi

X

X

Geophis dunni X X

X

X

Geophis immaculatus

X

3

X

X

4

X

1

X X

1 2

X

X

Geophis laticinctus

X

X

Geophis laticollaris

X

X

X

Geophis latifrontalis Geophis maculiferus Geophis mutitorques X

X

4 1

X

X

1 1

Geophis juliai

X

2 2

X

Geophis juarezi

Geophis nasalis

1

X

Geophis incomptus Geophis isthmicus

3

1 X

Geophis godmani

X

X X

Geophis fulvoguttatus Geophis hoffmanni

1

1 X

1

X

X

2

X

X

6 to be continued.

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TABLE 3 (continued). Vegetation Formations Species

TWM TD

TVD

SRW SM

SD

LMW LMM LMD MW

MM

Totals

Geophis nephodrymus

X

1

Geophis nigroalbus

X

1

Geophis nigrocinctus

X

1

Geophis omiltemanus

X

1

Geophis petersi

X

1

Geophis pyburni

X

1

Geophis rhodogaster

X

X

Geophis rostralis

X

Geophis russatus Geophis ruthveni

X X X

1

X X

X

Geophis sieboldi

2 X

2

X

4

X

X

Geophis talamancae

X

3

X

Geophis tarascae

1 X

Geophis zeledoni Totals

1

X

Geophis sallaei Geophis semidoliatus

2

1

X 10

5

0

15

12

2

20

1 18

0

0

1



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Cope, E.D. (1868) An examination of the reptilia and batrachia obtained by the Orton expedition to Ecuador; and the upper Amazon; with notes on other species. Proceedings of the Academy of Natural Sciences, Philadelphia, 20, 96– 140. Cope, E.D. (1871) Ninth contribution to the herpetology of tropical America. Proceedings of the Academy of Natural Sciences, Philadelphia, 23, 200–224. Cope, E.D. (1876) On the batrachia and reptilia of Costa Rica. Journal of the Academy of Natural Sciences, Philadelphia, 8, 93–154. Cope, E.D. (1885) A contribution to the herpetology of Mexico. Proceedings of the American Philosophical Society, 22, 379–404. Cope, E.D. (1900) The crocodilians, lizards, and snakes of North America. Annual Report of the United States National Museum, 1898, 151–1294. Dixon, J.R. (1968) Notes on the snake genus Geophis, from Nevado de Colima, Jalisco, Mexico. Southwestern Naturalist, 13, 452–454. Dixon, J.R. & Thomas, R.A. 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(1865) Livraison 12, In: Jan, G. and Sordelli, F. Iconographie Générale des Ophidiens. Tome Premier (Livrais 1 à 17). J. B. Bailliére et Fils, Paris. 11 pp. + 17 livr., 9 pp. +17 livr., 19 pp. + 16 livr. Johnson, J.D., Ely, C.A. & Webb, R.G. (1976) Biogeographical and taxonomic notes on some herpetozoa from the northern highlands of Chiapas, Mexico. Transactions of the Kansas Academy of Sciences, 79, 131–140. Köhler, G. (1996) Notes on a collection of reptiles from El Salvador collected between 1951–1956. Senckenbergiana Biologica, 76, 29–38. Legler, J.M. (1959) A new snake of the genus Geophis, from Chihuahua, Mexico. University of Kansas Publications, Museum of Natural History, 11, 327–334. Lips, K.R., & Savage, J.M. (1994) A new fossorial snake of the genus Geophis (Reptilia: Serpentes: Colubridae) from the Cordillera de Talamanca of Costa Rica. Proceedings of the Biological Society of Washington, 107, 410–416. McCranie, J.R. & Castañeda, F.E. (2004) Notes on the second specimens of Geophis damiani Wilson, McCranie and Williams and Rhadinaea tolpanorum Holm and Cruz D. (Colubridae). Herpetological Review 35, 341. A CHECKLIST AND KEY TO THE SNAKES OF GEOPHIS

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McCranie, J.R. & Wilson, L.D. (1991) Geophis fulvoguttatus Mertens and Micrurus browni Schmidt and Smith: additions to the snake fauna of Honduras. Amphibia Reptilia 12, 112–114. Medica, P.A., Arndt, R.G. & Dixon, J.R. (1975) Additional records of reptiles from Jalisco, Mexico. Great Basin Naturalist 35, 317–318. Mertens, R. (1952a) Weitere neue Repltilien aus El Salvador. Zoologischer Anzeiger, 149, 133–138. Mertens, R. (1952b) Die Amphibien und Reptilien von El Salvador, auf Grund der Reisen von R. Mertens und A. Zilch. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 487, 1–120. Müller, F. (1882 [1881]). Erster Nachtrag zum Katalog der herpetologischen Sammlung des Basler Museum. Verhandlungen der Naturforschenden Gesellschaft in Basel, 7, 120–165. Myers, C.W. (2003) Rare snakes—Five new species from eastern Panama: Reviews of northern Atractus and southern Geophis (Colubridae: Dipsadinae). American Museum Novitates, 3391, 1–47. Nieto-Montes de Oca, A. (2003) A new species of the Geophis dubius group (Squamata: Colubridae) from the Sierra de Juárez of Oaxaca, Mexico. Herpetologica, 59, 572–585. Pérez-Higareda, G., Vogt, R.C. & Flores-Villela, O. (1987) Lista Anotada de los Anfibios y Reptiles de la Region de Las Tuxtlas, Veracruz. Instituto de Biologia, Universidad Nacional Autónoma de México. 23 pp. Pérez-Higareda, G., Smith, H.M. & López-Luna, M.A. (2001) A new Geophis (Reptilia: Serpentes) from southern Veracruz, Mexico. Bulletin of the Maryland Herpetological Society 37, 42–48. Peters, W. (1859) Die von Hrn. Dr. Hoffmann in Costa Rica gesammelten Schlangen. Monatsberichte der Königlichen Preuss. Akademie der Wissenschaften zu Berlin, 1859, 275–278. Peters, W. (1861) Mittheilung über neue Schlangen des königlichen zoologischen Museums. Monatsberichte der Königlichen Preuss. Akademie der Wissenschaften zu Berlin, 1861, 922–925. Restrepo T., J.H., & Wright, J.W. (1987) A new species of the colubrid snake genus Geophis from Colombia. Journal of Herpetology, 21, 191–196. Rodriguez Torres, J. & Vásquez Díaz, J. (1990) Nuevo registro de distribucion de Geophis dugesi aquilonaris para el Estado de Jalisco, Mexico. Boletín de la Sociedad Herpetológica Mexicana, 2(1), 9–12. Sasa, M. (1993) Distribution and reproduction of the gray earth snake Geophis brachycephalus (Serpentes: Colubridae) in Costa Rica. Revista de Biología Tropical, 41, 295–297. Savage, J.M. (1960) A revision of the Ecuadorian snakes of the Colubrid genus Atractus. Miscellaneous Publications of the Museum of Zoology, University of Michigan, 112, 1–86. Savage, J.M. (1981) A new species of the secretive colubrid snake genus Geophis from Costa Rica. Copeia,1981, 549– 553. Schmidt, K.P. (1932) Stomach contents of some South American coral snakes, with the description of a new species of Geophis. Copeia 1932, 6–9. Slevin, J.R. (1939) Notes on a collection of reptiles and amphibians from Guatemala. I. Snakes. Proceedings of the California Academy of Sciences, 23, 393–414. Smith, E.N. (1995) Geophis rhodogaster (Colubridae), an addition to the snake fauna of Mexico. Southwestern Naturalist, 40, 123–124. Smith, H.M. (1939) Notes on Mexican reptiles and amphibians. Field Museum of Natural History Zoology Series 24(4), 15–35. Smith, H.M. (1941a) Notes on Mexican snakes of the genus Geophis. Smithsonian Miscellaneous Collections, 99(19):1– 6. Smith, H.M. (1941b) A new Geophis from Mexico. Proceedings of the New England Zoölogical Club, 18, 49–55. Smith, H.M. (1959) New and noteworthy reptiles from Oaxaca, Mexico. Transactions of the Kansas Academy of Sciences, 62, 265–272. Smith, H.M. & Chiszar, D. (1992) A second locality for Geophis sallei (Reptilia Serpentes). Bulletin of the Maryland Herpetological Society, 28, 16–18. Smith, H.M. & Flores-Villela, O. (1993) Variation in two species (Geophis bicolor, Geophis duellmani) of Mexican earth snakes (Geophis). Bulletin of the Maryland Herpetological Society, 29, 20–23. Smith, H.M. & Holland, R.L. (1969) Two new snakes of the genus Geophis from Mexico. Transactions of the Kansas Academy of Sciences, 72, 47–53. Smith, H.M., J. D. Lynch, and R. Altig, (1965) New and noteworthy herpetozoa from southern Mexico. Natural History Miscellanea, Chicago Academy of Sciences,180, 1–4. Smith, H.M. & Pérez-Higareda, G. (1991) Clara evidencia de la coespecificidad de Geophis dubius y Geophis rostralis (Reptilia: Serpentes). Boletín de la Sociedad Herpetológica Mexicana, 3, 39–40. Smith, H.M. & Williams, K.L. (1963) New and noteworthy amphibians and reptiles from southern Mexico. Herpetologica 19, 22–27. Smith, H.M. & Williams, K.L. (1966) A new snake (Geophis) from México. Journal of the Ohio Herpetological Society, 5, 90–92. Smith, H.M., Williams, K.L., Wallach, V., & Chiszar, D. (1990) Nomenclatural priority of Geophis Wagler, 1830, over

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A CHECKLIST AND KEY TO THE SNAKES OF GEOPHIS

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