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Inventory and Conservation Assessment of the Herpetofauna of the Sierra de Omoa, Honduras, with a Review of the Geophis (Squamata: Colubridae) of Eastern Nuclear Central America

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Inventory and Conservation Assessment of the Herpetofauna of the Sierra de Omoa, Honduras, with a Review of the Geophis (Squamata: Colubridae) of Eastern Nuclear Central America
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TOWNSEND, JOSIAH HAROLD
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2008

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Amphibians ( jstor )
Cloud forests ( jstor )
Colors ( jstor )
Environmental conservation ( jstor )
Forests ( jstor )
Highlands ( jstor )
Museums ( jstor )
Reptiles ( jstor )
Snakes ( jstor )
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City of Miami ( local )

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University of Florida
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University of Florida
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Copyright Josiah Harold Townsend. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
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5/31/2008
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496627197 ( OCLC )

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INVENTORY AND CONSERVATION ASSESSMENT OF THE HERPETOFAUNA OF THE SIERRA DE OMOA, HONDURAS, WITH A REVIEW OF THE Geophis (SQUAMATA: COLUBRIDAE) OF EASTERN NUCLEAR CENTRAL AMERICA By JOSIAH HAROLD TOWNSEND A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF ARTS UNIVERSITY OF FLORIDA 2006

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Copyright 2006 by Josiah Harold Townsend

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As a biotic environment the Honduranean cloud forests offer the biologist a challenge and promise of reward that would be hard to duplicate. But more than this, they hold an infallible aesthetic appeal that is as deep as the mystery of the dim, green woods and as varied as its changing aspects. –Archie F. Carr, Jr. (1953: 20)

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ACKNOWLEDGMENTS I would like to thank my supervisory committee chair, Max A Nickerson, and supervisory committee members, F. Wayne King and J. Richard Stepp, for providing guidance and support throughout the process of completing this thesis. My mentor and friend, Larry David Wilson, has been a trusted advisor and a marvelous field companion, and I have benefited enormously from his experience. My friend, Roberto Downing, was of invaluable assistance in arranging the logistics for my work in Cusuco in Spring 2005 and El Paraiso in the Summer of 2005, and helping me obtain collection and export permits. Conrado Gonzles, Martha Moreno, Ibrahim Padilla, and Carla Crcamo of the Departamento de Areas Protegidas y Vida Silvestre (DAPVS) of the Administracin Forestal del Estado Corporacin Hondurea de Desarrollo Forestal (AFE-COHDEFOR), Honduras, generously provided permission to collect and export scientific specimens. James R. McCranie was instrumental in securing the five-year collecting permit I worked under. I gratefully acknowledge funding from a research assistantship at the Florida Museum of Natural History during the 2004-2005 academic year; and from a Tropical Conservation and Development Graduate Fellowship during the 2005-2006 academic year. David Steadman, Scott Robinson, Leslie Campbell, Shuronna Wilson, Hannah Covert, Wanda Carter, and Victoria Gomez de la Torre were integral in providing and administering these assistantships. Fieldwork in Honduras was supported by Operation Wallacea, the Reptile and Amphibian Conservation Corps (RACC), and the Herpetology iv

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Division of the Florida Museum of Natural History. Varying degrees of field assistance in Honduras was provided by Larry David Wilson, T. Lynette Plenderleith, Brooke L. Talley, Sara M. Hughes, James C. Nifong, Douglas C. Fraser, Charlotte C. Woodhead, Phillip J. Lewis, Jackie B. Grant, Makrn Ramrez, Ren Alvarenga, Danlo Alvarenga, Dan Pupius, Ed Anderson, Emma Sherratt, David Carter, Adrian Symonds, Rachel Freer, Justin Hines, Andrew Stronach, Zia Lewindson, Rauri Allan, Gerry Carter, Louisa Sly, and dozens of volunteers from the Operation Wallacea Honduras Forest Project. Tim Coles provided the opportunity to work with Operation Wallacea, and enthusiastically directed the project. Don Enrique Morales Alegra allowed us to work on his property in El Paraiso, and Doa Telma graciously hosted our stay at El Paraiso. The residents of Buenos Aires and El Paraiso (Depto. Corts) and La Fortuna (Depto. Santa Barbara) were gracious hosts and companions during my time in their communities; muchas gracias por todo. Doa Gladys Fasquelle, of the Pastor Fundacon Hector Rodrigo Pastor Fasquelle, San Pedro Sula, Honduras, was kind enough to store specimens associated with this thesis over the course of late 2004 and early 2005. Loans of specimens under their care were facilitated by Steve Gotte, James Poindexter, and Roy McDiarmid of the National Museum of Natural History, Washington, DC; John Simmons, D. Berger, and C. Linkem of the University of Kansas Natural History Museum, Lawrence, Kansas; J. Vindum and A. Harper, California Academy of Sciences, San Francisco, California; Gunther Khler of the Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main, Germany; Ronald Nussbaum and Greg Schneider of the Museum of Zoology, University of Michigan, Ann Arbor, Michigan; Jimmy McGuire and J. Castillo, University of v

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California Museum of Vertebrate Zoology, Berkeley, California; and J. Seigel, Natural History Museum of Los Angeles County, Los Angeles, California. Steve Gotte facilitated my visit to the National Museum of Natural History, and Jos Rosado facilitated my visit to the Museum of Comparative Zoology, Cambridge, Massachussetts. Dan Janes hosted my stay in Cambridge, Massachussetts, and Steve and Terri Townsend provide room and board during my visit to Washington, DC. James R. McCranie kindly allowed use of material he collected in Rus Rus and Warunta, Honduras. Daniel Pupuis, Brooke Talley, and Phillip Lewis provided use of their photographs from Honduras. Stephanie LaRusso took the time and effort to prepare the head scale illustrations used in this thesis. Finally, and most importantly, I want to thank my wife Deborah for her steadfast support, understanding, and caring during the process of researching and writing this thesis. vi

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TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................iv LIST OF TABLES.............................................................................................................ix LIST OF FIGURES...........................................................................................................xi ABSTRACT.....................................................................................................................xiii INTRODUCTION...............................................................................................................1 METHODS AND MATERIALS.........................................................................................9 Compiling an Inventory of the Sierra de Omoa Herpetofauna.....................................9 Taxonomic Standards.................................................................................................12 Coefficient of Biogeographic Resemblance...............................................................13 Environmental Vulnerability Scores...........................................................................13 Specimen Examination...............................................................................................14 Format for Systematic Accounts................................................................................18 THE SIERRA DE OMOA, HONDURAS: SOCIOECONOMIC AND ECOPHYSIOGRAPHIC SETTING...........................................................................20 Socioeconomic Influences on Conservation in The Sierra De Omoa........................20 Ecophysiography of the Sierra de Omoa....................................................................22 Biodiversity in the Sierra de Omoa............................................................................29 RESULTS OF FIELDWORK IN THE SIERRA DE OMOA DURING 2005.................33 CONSERVATION ASSESSMENT OF THE HERPETOFAUNA OF THE SIERRA DE OMOA..................................................................................................................41 STATUS OF Geophis, species inquirenda, AND A REDESCRIPTION OF Geophis nephodrymus TOWNSEND AND WILSON.............................................................53 Differentiation of Geophis fulvoguttatus from G. species inquirenda.......................53 Identity of Geophis, species inquirenda.....................................................................58 Geophis nephodrymus Townsend and Wilson...........................................................59 REVIEW OF THE Geophis OF EASTERN NUCLEAR CENTRAL AMERICA...........73 vii

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Geophis damiani Wilson, McCranie, and Williams...................................................77 Geophis dunni Schmidt...............................................................................................83 Geophis fulvoguttatus Mertens...................................................................................89 Geophis hoffmanni (Peters)........................................................................................94 Geophis rhodogaster (Cope)....................................................................................105 A KEY TO THE SPECIES OF Geophis FROM EASTERN NUCLEAR CENTRAL AMERICA................................................................................................................111 LITERATURE CITED....................................................................................................113 BIOGRAPHICAL SKETCH...........................................................................................124 viii

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LIST OF TABLES Table page 1 Population growth of the four largest cities in the watershed of the Sierra de Omoa, Depto. Corts, Honduras..............................................................................22 2 Amphibian and reptile species recorded during fieldwork in the Sierra de Omoa, northwestern Honduras, during 2005.......................................................................34 3 Site-by-site breakdown of amphibian and reptiles species collected within Parque Nacional El Cusuco during 2005.................................................................40 4 Inventory and conservation assessment of the amphibians and reptiles of the Sierra de Omoa, Honduras.......................................................................................42 5 Coefficient of Biogeographic Resemblance (CBR) matrix of overall herpetofaunal similarity between Parque Nacional El Cusuco, Parque Nacional Cerro Azul, and Parque Nacional Pico Bonito.........................................................48 6 Coefficient of Biogeographic Resemblance (CBR) matrix of similarity of the highland herpetofauna (found only in PWF and LMWF formations) in Parque Nacional El Cusuco, Parque Nacional Cerro Azul, and Parque Nacional Pico Bonito.......................................................................................................................48 7 Coefficient of Biogeographic Resemblance (CBR) matrix of similarity of herpetofauna endemic to Honduras and found in Parque Nacional El Cusuco, Parque Nacional Cerro Azul, and/or Parque Nacional Pico Bonito.........................48 8 Environmental Vulnerability Scores (EVS) for species of amphibians and reptiles that were recently described or reported from Honduras............................51 9 Amphibians and reptiles of conservation significance from the Sierra de Omoa, ranked by EVS.........................................................................................................51 10 Summary of relevant measurements, scale counts, and variation in scale measurement ratios for the Geophis of eastern Nuclear Central America...............54 11 Morphological and meristic characters for Geophis nephodrymus, part I...............71 12 Morphological and meristic characters for Geophis nephodrymus, part II..............72 ix

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13 Morphological and meristic characters for Geophis damiani, G. dunni, G. fulvoguttatus, and G. rhodogaster............................................................................82 14 Relevant measurements and scale counts for Geophis fulvoguttatus and G. rhodogaster specimens with unmeasured head scales.............................................93 15 Relevant measurements and scale counts for Geophis hoffmanni from localities in eastern Nuclear Central America other than the Rus Rus/Warunta region..........95 16 Variation in ratios for Geophis hoffmanni from localities in eastern Nuclear Central America other than the Rus Rus/Warunta region........................................97 17 Relevant measurements and scale counts for Geophis hoffmanni from localities in eastern Nuclear Central America within Parque Nacional Warunta and Reserva Biolgica Rus Rus, Honduras....................................................................98 18 Variation in ratios for Geophis hoffmanni from localities in eastern Nuclear Central America within Parque Nacional Warunta and Reserva Biologica Rus Rus, Honduras..........................................................................................................99 19 Relevant measurements and scale counts for Geophis hoffmanni from localities in eastern Nuclear Central America within Parque Nacional Warunta, Honduras.100 20 Variation in ratios for Geophis hoffmanni from localities in eastern Nuclear Central America within Parque Nacional Warunta, Honduras..............................101 x

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LIST OF FIGURES Figure page 1 Political map of Central America...............................................................................2 2 Eastern Nuclear Central America (eNCA), with areas above 900 m elevation shaded gray.................................................................................................................4 3 Map of localities and features from the Sierra de Omoa, northwestern Honduras, mentioned in the text..................................................................................................5 4 Biogeographic realms of Central America. Areas above 800 m elevation are shaded gray.................................................................................................................7 5 Denuded Premontane Wet Forest (PWF) near Buenos Aires, Sierra de Omoa, photo taken from about 1100 m elevation................................................................26 6 Mixed Cloud Forest, near Cerro Cusuco, around 1800 m elevation, Parque Nacional El Cusuco..................................................................................................28 7 Hepatic or Mossy Forest, Cerro Cusuco, 2000 m elevation, Parque Nacional El Cusuco......................................................................................................................30 8 The author in Heather Wind Scrub habitat, Cerro Cusuco, 2030 m elevation, Parque Nacional El Cusuco......................................................................................30 9 An undescribed species of Oedipina (JHT 1553) from Cantiles, 1780 m, Parque Nacional El Cusuco, Corts, Honduras....................................................................37 10 An undescribed species of bromeliad-dwelling hylid frog (JHT 1572) from bosque enano, 1990 m elevation, Parque Nacional El Cusuco, Corts, Honduras..37 11 Oedipina elongata (UF 144649) from El Paraiso Valley, 185 m, Corts, Honduras..................................................................................................................38 12 Juvenile Geophis nephodrymus (JHT 1334), demonstrating the patternless variant, from Sendero El Quetzal, 1545 m, Parque Nacional El Cusuco, Corts, Honduras..................................................................................................................61 13 Juvenile Geophis nephodrymus (UF 143024), red patterned variant, from finca de Makrn Ramrez, 1580 m, Parque Nacional El Cusuco, Corts, Honduras.........61 xi

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14 Adult male Geophis nephodrymus (JHT 1342), red patterned variant, from Sendero Las Minas, 1580 m, Parque Nacional El Cusuco, Corts, Honduras. .......63 15 Juvenile Geophis nephodrymus (JHT 1343), pale gray patterned variant, from Sendero Las Minas, 1560 m, Parque Nacional El Cusuco, Corts, Honduras. .......63 16 Geophis nephodrymus from Cantiles, 1780 m, Parque Nacional El Cusuco, Corts, Honduras......................................................................................................64 17 Semidiagramatical illustration of the dorsal, lateral, and ventral aspects of the head of Geophis nephodrymus (UF 143022), from Sendero Las Minas, 1580 m, Parque Nacional El Cusuco, Corts, Honduras........................................................65 18 Distribution of Geophis in eastern Nuclear Central America..................................75 19 Holotype of Geophis dunni (MCZ 31870), an adult female from Matagalpa, Nicaragua.................................................................................................................88 20 Preserved specimens of Geophis fulvoguttatus; adult male (KU 57996) from Hacienda Montecristo, 2200 m, Cordillera de Metapn, Santa Ana, El Salvador, and adult female (KU 214781) from El Portillo de Ocotepeque, 1900 m, Ocotepeque, Honduras.............................................................................................88 xii

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Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Arts INVENTORY AND CONSERVATION ASSESSMENT OF THE HERPETOFAUNA OF THE SIERRA DE OMOA, HONDURAS, WITH A REVIEW OF THE Geophis (SQUAMATA: COLUBRIDAE) OF EASTERN NUCLEAR CENTRAL AMERICA By Josiah Harold Townsend May 2006 Chair: Max A. Nickerson Major Department: Latin American Studies Honduras is one of the poorest and least-developed countries in the Western Hemisphere, and also has one of the fastest growing human populations. One of the most densely populated and rapidly growing parts of Honduras is San Pedro Sula and the Sula Valley, in the watershed of the Sierra de Omoa. The Sierra de Omoa supports both mesic and subhumid forests from near sea level to above 2200 m elevation, including cloud forests within the boundaries of Cusuco National Park. Cloud forests are unique ecosystems that provide a variety of ecosystem services for surrounding lowlands and are threatened by deforestation and climatic change. The Sierra de Omoa also supports a variety of endemic plants and invertebrates as well as highland birds and mammals of international conservation significance. Herpetofauna show a high degree of endemism in cloud forests, and this has been well documented in Central America. Many endemic species are fossorial or semi-fossorial snakes. Honduras has more than a dozen cloud forests, and over 78 endemic amphibians and reptiles. The Sierra de Omoa is home to a xiii

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variety of endemic herpetofaunal species, including the snake Geophis nephodrymus. There is also a small series of distinctively patterned Geophis known from the Sierra de Omoa whose taxonomic status remains uncertain. Fieldwork was undertaken at five sites in the Sierra de Omoa during 2005, and 83 species were recorded from localities between 5 and 650 m in El Paraiso and between 1450 and 2200 m in Cusuco National Park. Noteworthy records include material representing at least three undescribed species, two new country records, four published distribution records, and the addition of 20 species to the list known from the park. One-hundred twenty-six species of amphibians and reptiles are known from the Sierra de Omoa, and 50 species are known from Cusuco National Park. There are 13 species endemic to the Sierra de Omoa, 16 more endemic to eastern Nuclear Central America, and a further 16 species are restricted to Nuclear Central America. The highland herpetofauna of Cusuco National Park is similar to that of Cerro Azul National Park, but in terms of endemic species the Sierra de Omoa is shown to be very distinctive. Twenty-five species are considered to be of high conservation significance. Cusuco National Park is vulnerable to anthropogenic degradation largely due to its proximity to the Sula Valley. All specimens of Geophis from the Sierra de Omoa are shown to be conspecific with G. nephodrymus based on characters of coloration and scale arrangement and counts. A total of 58 specimens of six species of Geophis known from eastern Nuclear Central America were examined, and detailed descriptions and species accounts are provided for each species. A key to the Geophis from eastern Nuclear Central America is provided. xiv

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INTRODUCTION Neotropical cloud forests are widely recognized as unique and valuable ecosystems that are rapidly disappearing as the frontier of agricultural expansion presses into the highest reaches of the Latin American highlands (Brown and Kappelle 2001, Bubb et al. 2004, Doumenge et al. 1995). The rate of deforestation in tropical highlands exceeds that of all other tropical forest biomes (FAO 1993, Doumenge et al. 1995), and tropical montane cloud forests have been acknowledged as highly threatened ecosystems that require increased attention and conservation efforts in order to ensure their protection (Aldrich et al. 2000). Tropical montane cloud forests (TMCFs) may be characterized as possessing three principal types of values: first, providing hydrological services, including capture of water from precipitation and wind-driven clouds, sequestration and transportation of that water, and regulation of soil erosion; second, providing habitat for specialized, often endemic, species of plants that form unique and fragile vegetation communities, and that may have added value for local people as medicine or food; and third, harboring a remarkably high degree of faunal specialization, exemplified by abundant examples of site-specific endemic taxa (Doumenge et al. 1995). These sensitive ecosystems can have their climates altered by deforestation of surrounding lowlands (Lawton et al. 2001, Pounds et al. 1999), and TMCFs may also serve as indicators of global climatic change, due to their sensitivity to variation in temperature, precipitation, and cloud cover (Foster 2001, Nair et al. 2003). Biological specialization and endemism are apparent across a 1

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2 variety of taxonomic groups in TMCF, especially plants (Cardels et al. 2006, Kper et al. 2004), invertebrates (Brehm et al. 2005, Olson 1994), birds (Long 1995; Poulsen and Krabbe 1997), and herpetofauna (Duellman 1999, Pounds and Fogden 2000). Central America (Figure 1) is a region whose cloud forest-inhabiting herpetofauna demonstrates a high degree of localized endemism, and is reasonably well studied (Campbell 1983, Janzen and Khler 2002, Pounds and Fogden 2000, Wilson and McCranie 2004a). Honduras is home to over a dozen isolated cloud forests that support a unique and diverse herpetofauna, with at least 78 species endemic to the country (Wilson and McCranie 2004a, 2004b). Seventeen of the reptiles endemic to Honduras are snakes, many of which are restricted to one or two cloud forest localities. Twelve endemic Honduran snakes are also semi-fossorial or fossorial, and TMCF-restricted semi-fossorial Figure 1. Political map of Central America.

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3 or fossorial snakes tend to be poorly understood, particularly in terms of how they are affected by environmental degradation or climatic change. The Sierra de Omoa is one mountain range in Honduras that supports a relatively large area of TMCF, as well as many endemic species of amphibians and reptiles, and was the focal region for my study (Figure 2, 3). Located in northwestern Honduras in the departments of Corts and Santa Barbara, the Sierra de Omoa is an isolated mountain range bounded by the alluvial Sula Valley to the east, the subhumid Ro Chamelecn valley to the southeast and south, a north-south subhumid valley to the west, the Ro Motagua alluvial plain to the west and northwest, and the Baha de Omoa and its associated narrow coastal plain to the north. At least nine species of amphibians and reptiles are found only in the Sierra de Omoa, and at least seven more that are limited to the highlands along the Honduras-Guatemala border (McCranie 2004a, Townsend and Wilson 2006, Wilson and McCranie 2004a). The high degree of endemism in the Sierra de Omoa warrants a more intensive and extensive assessment of the composition, distribution, and conservation status of the herpetofauna of those mountains, in order to provide a well-supported foundation for future conservation efforts. The 46 species of tropical earth snakes, genus Geophis (Squamata: Colubridae), are small, semifossorial or fossorial snakes primarily distributed in the highlands of Mexico, Central America, and northern South America (Downs 1967). Four species of Geophis are known to occur in Honduras. Two of those, G. damiani and G. nephodrymus, are endemic to isolated TMCF sites in the Cordillera Nombre de Dios and Sierra de Omoa, respectively.

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4 Figure 2. Eastern Nuclear Central America (eNCA), with areas above 900 m elevation shaded gray. Highland areas discussed in the text are labeled as follows: 1 = Cerro Montecristo and vicinity, El Salvador, Guatemala, Honduras; 2 = Parque Nacional Cerro Azul and vicinity, Honduras; 3 = Sierra de Omoa, Honduras; 4 = Cordillera Nombre de Dios, Honduras; 4a = Cerro Texiguat, Honduras; 4b = Parque Nacional Pico Bonito, Honduras; 5 = Matagalpan highlands, Nicaragua. My 2004 fieldwork in the highland forests of Parque Nacional El Cusuco in the Sierra de Omoa produced the holotype of G. nephodrymus, a single patternless specimen with 17 dorsal scale rows and fused supraocular and postocular scales (Townsend and Wilson 2006). Four other specimens of Geophis (herein referred to as Geophis, species inquirenda) were also collected. These other four specimens were similar in having: 17 dorsal scale rows Distinct supraocular and postocular scales Red partial bands and laterally-placed blotches on the dorsal surface Similar dorsal ground coloration and ventral coloration to G. nephodrymus

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5 120 ventral scales (compared to 136 in the holotype of G. nephodrymus, and 135 for G. fulvoguttatus, a similarly pattered species known from El Salvador and southwestern and western Honduras) 25 subcaudal scales (24 in G. nephodrymus, 24 in G. fulvoguttatus) First infralabial scales that are not, or are barely, in contact with each other between the mental scale and anterior chinshields (first infralabial contact suture 40% of their total length in G. nephodrymus, 43% in G. fulvoguttatus) Figure 3. Map of localities and features from the Sierra de Omoa, northwestern Honduras, mentioned in the text. Light gray areas are between 600 m and 1400 m elevation, roughly corresponding to the PWF formation; dark gray areas are above 1400 m elevation, roughly corresponding to the LMWF formation; red areas are heavily urbanized; 1 = Ro Chamelecn; 2 = Baha de Omoa; 3 = Ro Cuyamel / Quebrada La Riudosa (Motagua drainage); 4 = Ro Cuyamelito (Motagua drainage); 5 = Ro Cusuco; 6 = Ro Cuyamel; 7 = Ro Naco; 8 = border between Depto. Corts and Depto. Santa Barbara; 9 = Cofradia; 10 = Buenos Aires; 11 = Centro de Visitantes, Parque Nacional El Cusuco; 12 = Guanales camp; 13 = La Fortuna camp; 14 = Cantiles camp; 15 = Cerro Cusuco; 16 = El Paraiso; 17 = El Paraiso Valley Ecological Reserve; 18 = San Pedro Sula; 19 = Choloma.

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6 Three possible taxonomic resolutions pertain to specific allocation of Geophis, species inquirenda: These specimens represent a range extension and previously undocumented morphological variation for G. fulvoguttatus. Despite variation in the numbers of ventrals and subcaudals between G. fulvoguttatus and Geophis, species inquirenda, the possibility of their conspecificity should be addressed for the following reasons: 1) similarities in the dorsal color pattern between the two species; 2) G. fulvoguttatus is reported to occur in Parque Nacional Cerro Azul, which shares a number of other cloud forest endemics with Parque Nacional El Cusuco. These specimens are conspecific with G. nephodrymus and represents a patterned variety of that species. In this case, the fusion of the supraocular and postocular seen in the holotype would be demonstrated to be an individual anomaly rather than a diagnosable character, and the concept of G. nephodrymus would need to be expanded to include new variation in scutellation and color pattern. These specimens represent a second, undescribed species of Geophis from Parque Nacional El Cusuco, distinguishable from G. nephodrymus at least by the separation of the supraocular and postocular and the presence of a dorsal pattern consisting of red lateral blotches and partial bands. Determining the taxonomic status of Geophis, species inquirenda will require the acquisition and subsequent comparison of additional examples of Geophis from Parque Nacional El Cusuco, which will also clarify the morphological variation present within G. nephodrymus. To facilitate precise comparison between the Geophis from the Sierra de Omoa and other species of Geophis from the region, a comprehensive review of morphological and molecular variation within and between the relevant species is required. This study will focus on morphological aspects of variation, in light of the current lack of available tissue samples or sequence data for most species of Geophis. Nuclear Central America (Figure 4) is that portion of the Middle American land mass centered around the highlands of southern Mexico, Guatemala, El Salvador, Honduras, and northwestern Nicaragua and bounded by the Isthmus of Tehuantepec in Mexico to the west and the Nicaraguan depression to the south (Schuchert 1935, Johnson

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7 1989, Campbell 1999). Eight of the ten Geophis found in Nuclear Central America are endemic to the region. Six species of Geophis (G. damiani, G. dunni, G. fulvoguttatus, G. hoffmanni, G. nephodrymus, and G. rhodogaster) occur in the region referred to as eastern Nuclear Central America, which includes the Sierra de Omoa. Eastern Nuclear Central America (eNCA) is the physiographic region defined by Campbell (1999: 116) as the highlands “south of the lower Ro Motagua Valley and east of a line in Guatemala connecting Zacapa, Chiquimula, Concepcon Las Minas, and the Guatemalan-El Salvador border at the Pacific Coast,” including “the Sierra de Merendn and the eastern Chiquimulan highlands in Guatemala, all of the Honduran and El Salvadoran highlands, and the highlands of Nicaragua north of Lago de Nicaragua” (Figure 2). This definition is expanded for the purposes of this thesis to include the lowland areas bounded within the Figure 4. Biogeographic realms of Central America. Areas above 800 m elevation are shaded gray.

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8 region. Four of the six species reported from eNCA are endemic to the region. Only two of the Geophis species from eNCA are presently known from more than six specimens, and those two species also occur outside the region. Three of the endemic species were described based on single specimens, G. damiani, G. dunni, and G. nephodrymus, and only one of these (G. damiani) is known from any additional material. Despite being a region of distinct evolutionary significance for the genus Geophis, the interand intraspecific variation, intrageneric relationships, biogeography, and natural history of the eNCA species remain poorly understood. The goals of this study are to A) inventory and assess the conservation status of the herpetofauna of the Sierra de Omoa, an isolated mountain range of considerable conservation significance in northwestern Honduras, B) determine the taxonomic status of Geophis, species inquirenda, an enigmatic series of snakes from the Sierra de Omoa, and C) comprehensively review the systematics and morphology of the six species of Geophis that are known to occur in eNCA. This study will be undertaken with the objective of providing worthwhile information not only to the herpetological research community but to conservation planners and practitioners in Central America as well.

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METHODS AND MATERIALS Compiling an Inventory of the Sierra de Omoa Herpetofauna Creating a comprehensive list of the amphibian and reptiles species found in the Sierra de Omoa was completed using two avenues of investigation. First, a list of previously documented herpetofaunal species from the Sierra de Omoa and associated foothills was developed using the available literature and a search of online collection databases (www.herpnet.org). Second, a set of intensive field surveys were designed and implemented to catalog the herpetofaunal diversity present at sites in a variety of habitats and disturbance regimes over a broad range of elevations. Beginning in early 2004, I served as Senior Herpetologist for the Operation Wallacea Honduras Forests Project, and was tasked with assessing herpetological diversity and conservation status in Parque Nacional El Cusuco and El Paraiso Ecological Reserve. I recruited and directed a team of field assistants, allowing me to survey five sites simultaneously utilizing work-intensive methods such as drift fences and quadrat sampling. Between 26 February and 5 March 2005, I visited Parque Nacional El Cusuco to search the vicinity of the type locality of Geophis nephodrymus and localities that produced the series of Geophis, species inquirenda for additional material. I also continued cataloguing the herpetological diversity of the park and identified potential drift fence sites for the summer fieldwork. Searches were conducted during the day and night and consisted of actively turning leaf litter, logs, and other debris. Specimens were also acquired from local workers in cafetales (coffee farms) around the communities of 9

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10 Buenos Aires and Baaderos, between 1100 and 1300 m elevation to the east of Parque Nacional El Cusuco. All specimens not released were preserved with 10% formalin and transferred to 70% ethanol for storage within one week. From 22 June to 23 August 2005, the vicinities of four sites were surveyed within the boundaries of Parque Nacional El Cusuco (Figure 3): the Centro de Visitantes (15.6’N, 88.9’W; 1550 m elevation), base camp for the Operation Wallacea Cusuco Project, which provided access to the Ro Cusuco and nearby trails that transect a variety of forest types between 1450 and >2000 m elevation; Guanales (15.9’N, 88.3’W; 1220 m elevation), a campsite at the bottom of a steep canyon on the southern edge of the park in the eastern headwaters of the Ro Naco, nearby trails that transect a variety of forest types between 1220 and 1500 m elevation; Cantiles (15.8’N, 88.5’W; 1780 m elevation), a campsite in the central highlands of the park on its northern drainage slope; and La Fortuna (15.9’N, 88.2’W; 1300 m elevation), a campsite on the southern drainage slope of the park above a tributary of the Quebrada de Rudosa. The fifth site, El Paraiso, is also in the Sierra de Omoa but lies outside the boundaries of Parque Nacional El Cusuco on the narrow northern coastal plain (Figure 3). The name El Paraiso is used in reference to the El Paraiso Valley Ecological Reserve (15’N, 88’W; 5 m elevation) and its vicinity, a privately owned and maintained reserve protecting an entire river valley in the northern foothills of the Sierra de Omoa from near sea level to over 1000 m elevation. Periodic visits to the community of Buenos Aires, where the road to the Centro de Visitantes of Parque Nacional El Cusuco originates, were also made in an attempt to record amphibian and

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11 reptile species in disturbed areas at lower elevations than are found within Parque Nacional El Cusuco. Drift fences were either straight, “L” shaped, or “T” shaped, and were constructed using 10 m lengths of metal mesh window screening that was 0.5 m wide, with 7.6 liter pitfall buckets spaced 3 m apart. Nineteen drift fence arrays with 63 pitfall traps were installed and operated at four sites in the Sierra de Omoa during June, July, and August 2005, for periods ranging from five to nine weeks at each site. Twelve drift fence arrays (38 pitfalls) were put in place in the vicinities of the Centro de Visitantes, Guanales, and Cantiles, Parque Nacional El Cusuco, at 1300 m, 1310 m, 1320 m, 1350 m, 1555 m, 1560 m, 1570 m, 1580 m, 1585 m, 1780 m, and 1800 m elevation. Some of the drift fence arrays were placed in the same drift fence sites that produced the holotype of Geophis nephodrymus and the specimens of Geophis, species inquirenda in 2004. Seven drift fence arrays (25 pitfalls) were installed at El Paraiso, in reclaimed cacao and banana plantations and secondary forest between 100 and 185 m elevation. At each site, drift fence arrays were checked at least once a day, typically before 0900 h. All fieldwork in Honduras was conducted under collecting and research permit DAPVS, as resolved in Resolucin GG–MP04 issued by the Departamento de Areas Protegidas y Vida Silvestre (DAPVS) of the Administracin Forestal del Estado Corporacin Hondurea de Desarrollo Forestal (AFE-COHDEFOR). Specimens were exported under permits issued in September 2004 (provided by Hector O. Cardona, AFE-COHDEFOR), March 2005 (provided by Conrado Gonzales and Ibrahim Padilla, AFE-COHDEFOR), and August 2005 (provided by Conrado Gonzales, Martha Moreno, and Ibrahim Padilla, AFE-COHDEFOR). Copies of all permits were

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12 deposited along with specimens in the Florida Museum of Natural History and the National Museum of Natural History. Taxonomic Standards I used the most up-to-date accepted nomenclature available. Recent taxonomic changes are summarized here. At the generic level, the beta anoles, previously referred to as Norops (sensu Guyer and Savage [1986, 1992]), are placed within the super-genus Anolis following Poe (2004) and in anticipation of further taxonomic revision, Bromeliohyla takes the place of Hyla for B. bromeliacia (Faivovich et al. 2005), Chaunus is used in place of Bufo for C. marinus (Frost et al. 2006), Cranopsis replaces Bufo for C. campbelli and C. valliceps (Frost et al. 2006), Craugastor is used instead of Eleutherodactylus for Central American species (Crawford and Smith 2005), Dendropsophus replaces Hyla for D. microcephalus (Faivovich et al. 2005), Lithobates replaces Rana for Central American species (Frost et al. 2006), Mastigodryas is used in favor of Dryadophis (Dixon and Tipton 2004), and Tlalocohyla is used in place of Hyla for T. loquax (Faivovich et al. 2005). At the species level, Leptodactylus fragilis is the correct name for frogs previously called L. labialis (Heyer 2002), Rana brownorum is used for Honduran populations formerly referred to as R. berlandieri (Zaldvar-Rivern et al. 2004, Frost 2004, McCranie et al. 2006), Trachemys venusta is used for Honduran turtles formerly known as T. scripta (Siedel 2002, McCranie et al. 2006), Drymarchon melanurus is used instead of D. corais (Wster et al. 2001), Atropoides mexicanus replaces A. nummifer (Campbell and Lamar 2004), and Crotalus simus is used in favor of C. durissus (Campbell and Lamar 2004, Savage et al. 2005).

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13 Coefficient of Biogeographic Resemblance Duellman’s (1990) Coefficient of Biogeographic Resemblance (CBR) algorithm was used as a method for comparing the herpetofaunas of different geographic or ecophysiographic areas. The formula for this algorithm is CBR = 2C/(N1 + N2), where C is the number of species in common to both formations, N1 is the number of species in the first area, and N2 is the number of species in the second area. The CBR value produced is a decimal 1.0, indicating increasing biogeographic resemblance as the value approaches 1.0, and with 1.0 meaning the herpetofaunas being compared are identical in size and composition. Comparative herpetological data from other national parks in Honduras follow McCranie (2004a) and McCranie and Castaeda (2005). Environmental Vulnerability Scores Wilson and McCranie (1992) developed a simple theoretical gauge for evaluating the vulnerability of the Honduran amphibian fauna, and subsequently used this Environmental Vulnerability Gauge to produce Environmental Vulnerability Scores (EVS) to aid in conservation analysis of various aspects of the Honduran herpetofauna (Wilson and McCranie 2003, 2004c). I used the EVS provided by Wilson and McCranie (2004c) to aid in assessing the conservation status of the herpetofauna of the Sierra de Omoa, and follow the methodology used in that paper to generate EVS for newly described or reported species not evaluated by Wilson and McCranie (2004c). The EVS is generated by taking the total of three rankings: 1) extent of geographic range, 2a) degree of specialization of reproductive mode for amphibians or 2b) the degree of persecution by humans for reptiles, and 3) extent of ecological distribution in Honduras. Extent of geographic range in Honduras is determined on a scale of 1 to 5, where: 1 = widespread in and outside of Honduras; 2 = peripheral in Honduras, widespread outside

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14 Honduras; 3 = restricted to NCA; 4 = restricted to Honduras; and 5 = only known from the vicinity of the type locality in Honduras. Degree of specialization of reproductive mode for amphibians is scored 1 through 5, where: 1 = both eggs and tadpoles in large or small bodies of lentic or lotic water; 2 = eggs in foam nests, tadpoles in small bodies of lentic or lotic water; 3 = tadpoles occur in small bodies of lentic or lotic water, eggs outside of water; 4 = eggs laid in moist situations on land or moist arboreal situations, direct development; and 5 = eggs and tadpoles in water-retaining arboreal bromeliads or water-filled tree cavities. Degree of persecution by humans for reptiles is scored 1 through 6, with 1 = fossorial, typically escaping human notice; 2 = semifossorial, or nocturnal arboreal or aquatic, nonvenomous and usually non-mimicking, sometimes escaping human notice; 3 = terrestrial and/or arboreal or aquatic, generally ignored by humans; 4 = terrestrial and/or arboreal or aquatic, thought to be harmful (often mistakenly) and may be killed on sight; 5 = venomous species or mimics thereof, killed when encountered; and 6 = species exploited by humans for meat, eggs, or skin. Extent of ecological distribution in Honduras is determined using known distribution of each species within eight different forest formations, with 1 = occurs in eight formations; 2 = occurs in seven formations; 3 = occurs in six formations; 4 = occurs in five formations; 5 = occurs in four formations; 6 = occurs in three formations; 7 = occurs in two formations; and 8 = occurs in one formation. Species with EVS scores from 3 are considered low vulnerability, EVS scores from 10 indicate medium vulnerability, and scores from 14 are high vulnerability. Specimen Examination All measurements are metric unless otherwise noted. A dash (–) is used to separate ranges, while a slash mark (/) is used to delineate characters that differ from the left to the

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15 right side of an individual specimen. Scale nomenclature follows that of Downs (1967). Museum collection names and acronyms used in the text are the following: Academy of Natural Sciences of Philadelphia, Pennsylvania (ANSP), California Academy of Sciences, San Francisco, California (CAS), Florida Museum of Natural History, University of Florida, Gainesville, Florida (UF), Josiah H. Townsend field series (JHT), Museo de Historia Natural de El Salvador, San Salvador, El Salvador (MUHNES), Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (MCZ), Natural History Museum of Los Angeles County, Los Angeles, California (LACM), Museum of Vertebrate Zoology, University of California, Berkeley, California (MVZ), Museum of Zoology, University of Michigan, Ann Arbor, Michigan (UMMZ), National Museum of Natural History, Washington, DC (USNM), The Natural History Museum, London (BMNH), Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main, Germany (SMF), University of Kansas Natural History Museum, Lawrence, Kansas (KU), Universidad Centroamericana, Managua, Nicaragua (UCA), University of Southern California, Los Angeles, California (UCLA), Universidad Nacional Autnoma de Honduras, Tegucigalpa, Honduras (UNAH), and Zologisches Museum, Berlin, Germany (ZMB). Museum collections housing specimens of Geophis from eNCA were identified by a combination of literature records and a search of online museum collection databases (www.herpnet.org). All known specimens were either requested on loan from their respective institution or examined during a visit to that institution. If a specimen was unavailable for examination, then any available data was taken from the literature and are cited as such.

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16 I visited the USNM on 26 and 27 April 2005, and the MCZ on 20 December 2005, in order to examine holotype specimens that could not be lent out. All other specimens examined were requested and received on loan from the following museums: KU, LACM, MVZ, SMF, UMMZ, and USNM. A total of 58 specimens of six species were examined for this study. A series of 53 characters were recorded for each specimen examined. Supralabial and infralabial scales were counted on both sides. All measurements were taken on the left side at their maximum lengths to the nearest 0.1 mm using a stereomicroscope and dial calipers. Specimens with cephalic damage or that were obviously desiccated were not included in the comparison of measurement ratios to ensure accurate ratios. These 53 characters included sex, number of ventral scales, number of paired subcaudal scales, total segmental counts (ventrals + subcaudals), left supralabial scale count, right supralabial scale count, left infralabial scale count, right infralabial scale counts, snout-vent length, tail length, total length, rostral length from above, rostral-frontal distance, internasal breadth, internasal length, prefrontals common suture length, internasals common suture length, prefrontal length, snout length (as measured from the anterior edge of the eye to the tip of the snout), frontal length, frontal breadth, supraocular length, supraocular breadth, postocular length, postocular height, loreal length, loreal depth, frontal-supraocular contact length, prefrontal-supraocular contact length, parietal length, parietal breadth, head length (measured from tip of snout to posterior margin of parietal), postnasal length, prenasal length, postnasal + prenasal length, eye vertical diameter, eye horizontal diameter, eye-lip distance, sixth supralabial lip exposure, fifth supralabial lip exposure, fourth supralabial lip exposure, third

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17 supralabial lip exposure, second supralabial lip exposure, first supralabial lip exposure, mental breadth, mental length, first infralabial length, first infralabial common suture length, anterior chinshield length, anterior chinshield breadth, posterior chinshield length, and posterior chinshield common suture length. I followed the methodology used by Nieto-Montes de Oca (2003) in describing Geophis juarezi to produce a set of 29 scale-measurement ratios and facilitate production of accurate and detailed descriptions of each species, as well as contribute to a standardized dataset for morphological comparison among species of Geophis. One ratio was added to the 28 used by Nieto-Montes de Oca (2003) to evaluate the consistency of contact between the first pair of infralabials anterior to the first set of chinshields. These ratios are presented in Tables 10 and are summarized in the descriptions in each Systematic Account. The 29 ratios, each assigned an individual number, are: (1) rostral length from above/rostral-frontal distance; (2) internasal breadth/internasal length; (3) internasal length/prefrontals common suture length; (4) internasals common suture length/prefrontals common suture length; (5) prefrontal length/snout length; (6) prefrontals common suture length/frontal length; (7) frontal breadth/frontal length; (8) supraocular length/loreal length; (9) frontal-supraocular contact length/supraocular length; (10) frontal-supraocular contact length/prefrontal-supraocular contact length; (11) parietal length/tip of snout-posterior margin of parietal distance; (12) parietals common suture length/frontal length; (13) parietal length/parietal breadth; (14) postocular height/ postocular length; (15) postnasal length/prenasal length; (16) (prenasal + postnasal) length/loreal length; (17) loreal length/loreal depth; (18) snout length/loreal length; (19) loreal length/eye horizontal diameter; (20) snout length/ eye horizontal diameter; (21) eye

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18 vertical diameter/eye-lip distance; (22) fifth supralabial exposure/fourth supralabial exposure; (23) third supralabial exposure/second supralabial exposure; (24) mental breadth/mental length; (25) anterior chinshield length/anterior chinshield breadth; (26) anterior chinshield length/posterior chinshield length; (27) posterior chinshield common suture length/posterior chinshield length; (28) tail length/total length; and (29) first infralabial common suture length/first infralabial length. Format for Systematic Accounts All data for each species that I examined or obtained from the literature are presented in Systematic Accounts, the format for which is described below. Information on the type material and type locality of each species is derived from the original literature. A brief synonymy is provided and includes only those name combinations used in reference to material from eNCA. Species Group placement for each species follows that of Downs (1967), or the original description if the species was described after 1967. The Diagnosis provides characteristics that will clearly differentiate each species from all other species in eNCA, except in the case of G. nephodrymus where the diagnosis is written to distinguish G. nephodrymus from all other species of Geophis, updating the diagnosis provided by Townsend and Wilson (2006). The Description follows a common format for consistency, and all information contained in the Description was measured or recorded by me unless otherwise noted (see Specimen Examination for details of characters used in the descriptions). Coloration was recorded and characterized by me, unless otherwise noted. Capitalization of the names and corresponding numbers of colors given in descriptions indicate the colors corresponding to Smithe (1975). The Distribution provides a summary of the entire known range of the species, and is derived from a combination of the available literature, museum data accompanying each

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19 specimen, and field notes of the author. Elevations are characterized as “low” (sea level to 600 m), “moderate” (600 m), “intermediate” (1501 m), or “high” (>2700 m), following Stuart (1963) and McCranie and Wilson (2002). The Natural History Comments section features information from the available literature, museum data, and my field observations and notes. Forest formation definitions follow those of Holdridge (1967), as modified and used by Meyer and Wilson (1971, 1973), Wilson and Meyer (1985), and McCranie and Wilson (2002). The Remarks section is used to discuss any information that is pertinent but not appropriate for other sections. Any literature pertinent to the species within eNCA is listed in References, as are published Illustrations of the same relevance. Finally, a Specimens Examined section lists all specimens seen by the author and their locality data. In cases in which there are extant specimens that I did not examine, the specimen number and source are listed as Referred Specimens.

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THE SIERRA DE OMOA, HONDURAS: SOCIOECONOMIC AND ECOPHYSIOGRAPHIC SETTING Socioeconomic Influences on Conservation in The Sierra De Omoa The Republic of Honduras, the second largest country in Central America (112,000 km 2 ), is situated in the central part of the Middle American Isthmus, with Guatemala on its western border and El Salvador and Nicaragua on its southern border (Figure 1). In many ways, Honduras epitomizes the socioeconomic challenges faced by Latin American countries in the 21 st century. Honduras is one of the poorest nations in the Western Hemisphere, with the third lowest GNIPPP (Gross National Income adjusted for Purchasing Power Parities) in the hemisphere, behind Haiti and Bolivia, and 44% of the population earning less than $2 US per day (Population Reference Bureau 2005). Honduras currently has an estimated population of 7.2 million with an annual rate of increase of 2.8% (tied with Guatemala for highest rate of population increase in the Western Hemisphere), and it is projected that the population of the country will increase by 104% to nearly 15 million by the year 2050 (Population Reference Bureau 2005). This rapid growth is surprising in light of Honduras’ 3.2% infant mortality rate, which is clearly tempered by the third highest fertility rate in the Western Hemisphere (Population Reference Bureau 2005). The economy is primarily export-based, and exports made up 42% of the 2000 Gross Domestic Product (EarthTrends 2003). The United States is Honduras’ biggest trading partner, accounting for 54.4% of the Honduras export market and providing 37.5% of Honduran imports (Central Intelligence Agency 2005). The 20

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21 Honduran export profile has changed little over the past century, and is strongly based in primary agricultural production with little or no value added prior to export. Most of the land area under cultivation is dedicated to the production of bananas, citrus fruit, coffee, and other export crops (Merrill 1995). A substantial portion of the economic activity in Honduras occurs along the northern coast, and the hub for much of that activity is the sprawling city of San Pedro Sula. San Pedro Sula is the second largest city in Honduras and the capital of Departamento de Corts, and is considered the industrial and economic heart of the country. This expansive city of more than half a million people sits in the lowlands at the eastern edge of the Sierra de Omoa (Figure 3), and has experienced rapid growth due in large part to migration from the southern and central Honduras (Merrill 1995, Table 1). The trend of rural-to-urban migration has fed rapid growth not only in San Pedro Sula, but also in the nearby cities and towns in the Chamelecn and Ula valleys and along the north coast between Puerto Corts and the Guatemalan border (Table 1). Neighboring cities, like Choloma, have witnessed phenomenal urban growth in the previous two decades, but this growth has not been matched by development of urban infrastructure and has created densely populated favelas (shantytowns) where residents have limited access to clean water or health care. Much of the migration to this region is in search of employment in the growing consumer merchandise production sector, largely based in the garment industry, as well as the deterioration of environmental conditions that has dramatically decreased productivity in southern Honduras, particularly around the city of Choluteca.

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22 Table 1. Population growth of the four largest cities in the watershed of the Sierra de Omoa, Depto. Corts, Honduras. Source: www.world-gazetteer.com. 1988 2001 2006 % increase 1988 San Pedro Sula 287,390 437,798 502,939 175 Choloma 39,054 105,899 148,885 381.2 Puerto Corts 31,586 43,845 49,050 155.3 La Lima 28,703 41,490 47,087 164 The rapid expansion of the human population within the watershed areas of the Sierra de Omoa has put unprecedented pressure on the natural resources of the mountain range. These mountains serve as a catchment for vast quantities of water deposited there by both vertical and horizontal precipitation from wind-driven clouds, which is eventually delivered to the densely populated lowlands by way of over a dozen river systems. Horizontal, or occult, precipitation is moisture that is intercepted by vegetation when canopy-level clouds pass through forest, and it is indeed the persistent presence of canopy-level clouds that led to the term “cloud forest” (Cavelier and Goldstein 1989, Cavelier et al. 1996). Watershed function is recognized as one of the most important values added by the presence of healthy TMCF’s, along with providing the setting for a variety of unique and specialized ecosystems, flora, and fauna (Brown and Kappelle 2001, Doumenge et al. 1995, Stadtmller 1987). Unfortunately, deforestation, primarily driven by small-scale coffee cultivation, has continued inside Parque Nacional El Cusuco at ever-higher elevations, continually degrading the water capture and erosion control functions of the Sierra de Omoa (pers. observation; Lennkh 2005). Ecophysiography of the Sierra de Omoa Most of the Honduran landmass makes up the eastern part of the Nuclear Central American highlands, and the mountainous interior of the country is often referred to as

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23 the serrana (Figs. 2, 4; Wilson and Meyer 1985, McCranie and Wilson 2002). The humid Caribbean lowlands extend along the entire length of the northern coast in a narrow plain, extending onto the windward slopes of Sierra de Omoa and Cordillera Nombre de Dios and inland along major river valleys, eventually widening in the eastern part of country in the lowland region known as the Mosquitia (Figure 2; Wilson and Townsend 2006). In sharp contrast to the humid Caribbean lowlands are the semiarid interior valleys, found in the upper reaches of the alluvial plains of the Agun, Chamelecn, Choluteca, and Comayagua. These valleys divide the serrana into fragmented areas of cloud forest-supporting mountains, which are analogous to “islands” and “archipelagos” of cool mesic habitat separated from each other by hot subhumid habitats (Figure 3). In the northwestern corner of Honduras, the Sierra de Omoa is effectively isolated by the Caribbean Sea and three broad and hot valleys, the Motagua, Chamelecn and Ula, creating an effective “island” of mesic upland habitat (Figure 2). Climatically, the Sierra de Omoa can be divided into the subhumid interior lowlands and foothills, the humid windward lowlands and foothills, and the mesic highlands, each with characteristic climatic regimes, forest formations, and biotic communities. Subhumid areas in the Chamelecn and Ula valleys, to the south and east of the Sierra de Omoa, are primarily Lowland Dry Forest (LDF) that has been converted to agriculture or urbanized. Lowland Dry Forest (LDF) is typified by having between 1000 and 2000 mm precipitation annually, a mean annual temperature in excess of 24C, and occurring from about 100 to 600 m elevation (McCranie and Wilson 2002; Wilson and McCranie 1998). Agdelo C. (1987) listed the following plant species as characteristic of

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24 LDF in Honduras: Astronium graveolens, Bursera simaruba, Calycophyllum candidissimus, Ceiba petandra, Cecropia spp., Chlorophora tinctoria, Entorolobium cyclocarpum, Guaiacum sanctum, Gyrocarpus americanus, Hymenaea coubaril, Melicocca bijuga, Pithecolobium dulce, Swietenia humilis, Tabebuia rosea, and Tamarindus indica. The alluvial plains of the Chamelecn and Ula rivers combine to form the Sula Plain or Sula Valley, an area that is one of the most densely populated and heavily degraded in Honduras. Much of forests between around 600 to 1500 m elevation on the interior slopes of the Sierra de Omoa are characterized as upland pine-oak forest, falling within the Premontane Moist Forest formation (PMF). Most of the forest in this formation has been converted to agriculture, and nearly all of the forest that does remain is subject to frequent fires, both of natural and human origin (McCranie and Wilson 2002). This formation is subject to the Intermediate Dry climate, which occurs from about 600 to 1500 m elevation, has a mean annual precipitation of 1000 to 2000 mm, and a mean annual temperature ranging from 18 to 24C (Wilson and Meyer 1985, McCranie and Wilson 2002). The upland pine-oak forests of the Sierra de Omoa are dominated by Pinus oocarpa, a widespread species found from northwestern Mexico to northwestern Nicaragua (Agdelo C. 1987, Farjon et al. 1997). Two characteristic species of oaks found in upland pine-oak forest are Quercus oleoides, which forms stands known as encinales, and Q. peduncularis, found in stands called robledales (Agdelo C. 1987). Additional tree species present in upland pine-oak forests include Arbutus xalapensis, Brysonima crassifolia, Curatella americana, Dodonaea viscosa, Genipa caruto, Lysiloma seemannii, and Piscidia grandifolia (Agdelo C. 1987).

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25 The Caribbean, or windward, slope of the Sierra de Omoa is subject to a much wetter climate then the interior slopes, due in large part to their exposure to prevailing onshore winds (Wilson and McCranie 2004a). Lower elevations on the Caribbean slope experience the Lowland Wet climate, receiving more than 2500 mm annual precipitation and having an average annual temperature of greater than 24C (Wilson and Meyer 1985, McCranie and Wilson 2002). Forests below about 600 m in the northern foothills of the Sierra de Omoa fall within the Lowland Moist Forest (LMF) formation (McCranie and Wilson 2002, Wilson and Townsend 2006). The Caribbean coastal plain of the Sierra de Omoa is divided between two biogeographic regions: the Lower Motagua Valley and the West Caribbean Lowlands (McCranie and Wilson 2002, Wilson and Townsend 2006). Agdelo C. (1987) stated that the following plant species are most typical of the LMF formation in Honduras: Andina inermis, Astronium graveolens, Castilla elastica, Cedrela adorata, Cordia alliodora, Luehea seemannii, Pithecolobium arboreum, Roystonea spp., Terminalia oblonga, Vochysia ferrugina, and V. hondurensis. The El Paraiso Valley Ecological Preserve provides access to secondary LMF, much of which from reclaimed cacao plantations that have been left undisturbed since 1992 (Kelly 2004). Between around 600 and 1500 m elevation on the Caribbean slopes exist forests in the Premontane Wet Forest (PWF) formation, although much of the PWF has been converted to agriculture, principally coffee above around 1100 m elevation (Figure 5). This region experiences the Intermediate Wet climatic regime, characterized by 2000 mm or greater annual precipitation, and a mean annual temperature of 18C (Wilson and Meyer 1985, McCranie and Wilson 2002). Outside of Parque Nacional El Cusuco, most of the remaining intact PWF in the Sierra de Omoa is patchily distributed on steep slopes

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26 Figure 5. Denuded Premontane Wet Forest (PWF) near Buenos Aires, Sierra de Omoa, photo taken from about 1100 m elevation. Photograph by B. L. Talley. and difficult to access ridges (Figure 5), including some primary forest protected within the El Paraiso Valley Ecological Reserve. The vegetation found in PWF is dominated by broadleaf trees, with a high-diversity species composition that is generally intermediate to that of LMF and LMWF. At upper elevations, PWF can be considered transitional cloud forest (Meja V. 2001). Agdelo C. (1987) listed the following plant species as characteristic of PWF: Ampelocera hottlei, Brosimum alicastrum, Calophyllum brasiliense, Cedrela adorata, Cordia alliodora, C. gerascanthus, Castilla elastica, C. tunu, Cynometra retusa, Dalbergia tucurensis, Dialium guianense, Hieronyma alchorneoides, Huertea cubensis, Magnolia yocoronte, Mauria sessiliflora, Pterocarpus officinalis, Sterculia mexicana, Swietenia macrophylla, Symphonia globulifera, Tabebuia

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27 guayacan, Terminalia amazonia, Vatairea lundellii, Virola guatemalensis, V. koschnyi, Vochysia hondurensis, and Zollernia tango. Most, if not all, of the remaining intact highland forests in the Sierra de Omoa are contained within the boundaries of Parque Nacional El Cusuco, established in 1987 to initially protect areas above 1800 m elevation, and later expanded to include areas as low as 1300 m (Wilson and McCranie 2004b, Townsend et al. 2006a). Forests within the park experience the Intermediate Wet climate in areas below 1500 m elevation, and the Highland Wet climate in areas above 1500 m elevation, characterized by 1500 mm or greater annual precipitation and a mean annual temperature of <18C (Wilson and Meyer 1985, Espinal et al. 2001, Wilson and McCranie 2004b). All forests within the park are characterized as falling within either the PWF (below about 1500 m elevation) or Lower Montane Wet Forest (LMWF; for areas above 1500 m elevation) formations. The name “cloud forest” is commonly used in reference to the LMWF. The cloud forests in Sierra de Omoa can further be delineated into at least four types, following the characterization of Honduran cloud forests provided by Meja V. (2001; original forest type names in parentheses). Mesic forests below around 1800 m elevation are primarily Pinus and Liquidambar Transitional Cloud Forest (Bosque transicional de conferas y Liquidambar styraciflua), a habitat that has also be referred to as pinabetal, in reference to a local name for Pinus maximinoii, the pinabete (Carr 1949, Meja V. 2001). The pine P. oocarpa may also be present around and below 1500 m elevation, as are broadleaf trees like Carpinus caroliniana, Clethra macrophylla, and L. styraciflua (Kelly 2004). Some forest found below 1800 m elevation on the windward slopes of the park is categorized as High-Diversity Broadleaf Transitional Cloud Forest

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28 (Bosque transicional de diversas especies latifoliados). There are two forest types present from 1800 to around 2000 m elevation: High-Diversity Broadleaf Cloud Forest (Bosque latifoliados muy diverso), which lacks Pinus in the overstory and is dominated by broadleaf trees, including Billia hippocastanum, Clusia masoniana, Persea vesicula, and Quercus cortesii (Kelly, 2004); and Mixed Cloud Forest (Bosque mixto, Figure 6), which also includes P. maximinoi and Podocarpus oleifolius in the overstory. All these forest types have a high diversity and density of bromeliads, primarily in the genera Bromelia and Tillandsia, and other epiphytes like orchids and epiphytic cacti. The understory includes at least six species of tree ferns (Cyatheales), some over 10 m tall, and seven species of palms (Kelly 2004). Figure 6. Mixed Cloud Forest, near Cerro Cusuco, around 1800 m elevation, Parque Nacional El Cusuco. Photograph by D. Pupius.

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29 On the peaks and upper slopes of Cerro Cusuco and Cerro Jilinco, the highest elevations in the Sierra de Omoa between around 2000 and 2242 m elevation, is the forest that residents of nearby communities refer to as the bosque enano, or “dwarf forest.” This is not the Bosque enano described by Meja V. (2001), but is instead classified as Hepatic or Mossy Forest (Bosque heptico o musgoso, Figure 7) and Heather Wind Scrub (Filos secos dominados por la familia Ericaceae, Figure 8), both of which are typified by short vegetation (canopy >10 m high) and an overwhelming abundance of epiphytic plants and fungi (Carr 1950, Meja V. 2001). Elsewhere in this study, these two closely associated forest types are referred to collectively by their local name, bosque enano. These unique elfin forests only occurs in a few sites in Honduras, the most substantial of which is on Pico La Picucha in the Sierra de Agalta, with small areas of similar forest on the tops of Cerro Celaque, Cerro Santa Barbara, and Cerro Azul Membar (Hazlett 1980, Meja V. 2001). These highly adapted mountaintop habitats are extremely vulnerable to degradation due to the slow rate at which they recover from disturbance, and may also be especially sensitive to climatic change. Biodiversity in the Sierra de Omoa Honduras demonstrates remarkable diversity in terms of the richness of its endemic biota. The country has endemic species representing almost all major taxonomic groups, including plants (263 species), invertebrates (more than a dozen species), amphibians (44 species), reptiles (32 species), mammals (two species), and one species of bird (Anderson and Ashe 2000, Cerrato et al. 2002, Nelson S. 2001, Wilson and McCranie 2004a, 2004b). Diversity of endemic invertebrates is likely much higher than is currently known

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30 Figure 7. Hepatic or Mossy Forest, Cerro Cusuco, 2000 m elevation, Parque Nacional El Cusuco. Part of the bosque enano survey site. Photograph by B. L. Talley. Figure 8. The author in Heather Wind Scrub habitat, Cerro Cusuco, 2030 m elevation, Parque Nacional El Cusuco. Part of the bosque enano survey site. Photograph by D. Pupius.

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31 (Anderson and Ashe 2000), given the relatively high levels of diversity exhibited by two better studied groups: plants and herpetofauna. Some of the most notable of the taxa endemic to Honduras are the monotypic family and genus Heptanthus hazlettii (Heptanthaceae), a strange plant known only from a single site near Cerro Texiguat and now thought to be extinct (Nelson S. 2001), and the endemic snake genus Omoadiphas, consisting of two species, O. aurula from the Sierra de Omoa and O. texiguatensis from Cerro Texiguat (Khler et al. 2001; McCranie and Castaeda 2004). These distinctive examples are but three of hundreds of species endemic to Honduras, and a majority of those species are inhabitants of highland or cloud forest (J. H. Townsend, unpubl. data). The Sierra de Omoa itself has a number of endemic plants, including the palms Chamaedorea moliniana and C. frondosa, the shrubs Rondeletia megalantha and Viburnum molinae, and the vine Matelea urophylla (Cerrato et al. 2002, Nelson S. 2001). These mountains also encompass the entire known range of a variety of invertebrates. One species of jewel scarab beetle, Chrysina cusuquensis, is endemic to the highlands of the Sierra De Omoa, and another species, C. spectabilis, previously thought to be extinct, was recently rediscovered in Parque Nacional El Cusuco (Cave 2000). Additionally, there is the scorpion Diplocentrus lourencoi (McWest 1997), and six species of water mites that are endemic to the Sierra de Omoa (Wiles 2005). The forests of the Sierra de Omoa also provide a refuge for a number of species that, while not endemic to Honduras, are of international conservation significance, including Baird’s tapir (Tapirus bairdii), ocelot (Leopardus pardalis), margay (L. weidi), resplendent quetzal (Pharomachrus mocinno), and highland guan (Penelopina nigra). But it is the herpetofauna that demonstrates the greatest degree of endemism of any group in the Sierra de Omoa, and fieldwork there

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32 continues to produce new taxa. The extent of uniqueness exhibited by the amphibians and reptiles of the Sierra de Omoa warrants a more intensive and extensive assessment of their conservation status, based on recent fieldwork and supplemented by published reports and museum records.

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RESULTS OF FIELDWORK IN THE SIERRA DE OMOA DURING 2005 Between 26 February and 5 March 2005, I traveled to Parque Nacional El Cusuco and searched the vicinity of the Centro de Visitantes, the type locality of Geophis nephodrymus and the localities that produced the series of Geophis, species inquirenda. On 2 March 2005, two juvenile specimens of Geophis, species inquirenda were found during the day beneath a pile of plant debris on a small farm near the Centro de Visitantes. Additionally, two species of salamanders, five species of anurans, seven species of lizards, and three other species of snakes were recorded (Table 2). Two of these species, the lizards Sceloporus variabilis and Sphenomorphus incertus, were additions to the herpetofauna of Parque Nacional El Cusuco (Townsend 2005, Townsend et al. 2006a). I returned to Honduras from 22 June to 23 August 2005, as Senior Herpetologist for the Operation Wallacea Honduras Forests Conservation Project. Assisted by a team of herpetologists recruited in early 2005, I directed surveys at five sites in the Sierra de Omoa, four of which were within the boundaries of Parque Nacional El Cusuco. The fieldwork added nine specimens of Geophis, plus one patternless specimen (JHT 1327) that was collected in 2004 by Operation Wallacea herpetologists and retrieved from the museum of the Fundacon Hector Rodrigo Pastor Fasquelle in San Pedro Sula, Honduras. Taxonomic resolution of Geophis, species inquirenda incorporating the new material is provided in the chapter “Status of Geophis, species inquirenda, and a redescription of 33

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34 Table 2. Amphibian and reptile species recorded during fieldwork in the Sierra de Omoa, northwestern Honduras, during 2005. PN El Cusuco 1220 m Buenos Aires 922 m El Paraiso 0 m Species Feb-Mar 05 Jun-Aug 05 Feb-Mar 05 Jun-Aug 05 Jul-Aug 05 Caecilidae: 1 Dermophis mexicanus X Plethodontidae: 7 Bolitoglossa conanti X X X Bolitoglossa diaphora X X Bolitoglossa dunni X Bolitoglossa rufescens X Cryptotriton nasalis X Oedipina elongata X Oedipina sp. X Bufonidae: 3 Chaunus marinus X X Cranopsis campbelli X Cranopsis valliceps X X Centrolenidae: 1 Hyalinobatrachium fleischmanni X Hylidae: 7 Bromeliohyla bromeliacia X Duellmanohyla soralia X X X “Hyla” sp. X Plectrohyla dasypus X X Plectrohyla exquisita X X Ptychohyla hypomykter X X X Smilisca baudinii X X X Leptodactylidae: 4 Craugastor chac X Craugastor charadra X X Craugastor rostralis X X X Leptodactylus melanonotus X Ranidae: 2 Lithobates brownorum X Lithobates maculatus X X Emydidae: 1 Trachemys venusta X Kinosternidae: 1 Kinosternon leucostomum X Anguidae: 2 Celestus montanus X Mesaspis moreletii X Corytophanidae: 3 Basiliscus vittatus X Corytophanes cristatus X Laemanctus longipes X

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35 Table 2. Continued. PN El Cusuco 1220 m Buenos Aires 922 m El Paraiso 0 m Species Feb-Mar 05 Jun-Aug 05 Feb-Mar 05 Jun-Aug 05 Jul-Aug 05 Gekkonidae: 2 Hemidactylus frenatus X Sphaerodactylus millepunctatus X Iguanidae: 2 Ctenosaura similis X Iguana iguana X Phrynosomatidae: 2 Sceloporus malachiticus X X Sceloporus variabilis X X Polychrotidae: 12 Anolis amplisquamosus X Anolis capito X X Anolis cusuco X X Anolis johnmeyeri X Anolis lemurinus X Anolis ocelloscapularis X Anolis petersii X Anolis rodriguezii X Anolis sericeus X Anolis sp. X X X Anolis tropidonotus X X Anolis uniformis X Scincidae: 2 Sphenomorphus cherriei X X Sphenomorphus incertus X X Teiidae: 3 Ameiva festiva X Ameiva undulata X Cnemidophorus lemniscatus X Xanthusidae: 1 Lepidophyma flavimaculatum X Colubridae: 23 Adelphicos quadrivirgatum X X X X Clelia clelia X Drymarchon melanurus X Drymobius chloroticus X X X Drymobius margaritiferus X Geophis nephodrymus X X Imantodes cenchoa X X X Lampropeltis triangulum X X Leptophis ahaetulla X X Leptophis mexicanus X Mastigodryas dorsalis X Mastigodryas melanolomus X Ninia sebae X X Oxyrhopus petula X Pseustes poecilonotus X Rhadinaea anachoreta X Rhadinaea montecristi X Scaphiodontophis annulatus X Sibon nebulatus X Spilotes pullatus X Tantilla schistosa X X X Tantillita lintoni X

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36 Table 2. Continued. PN El Cusuco 1220 m Buenos Aires 922 m El Paraiso 0 m Species Feb-Mar 05 Jun-Aug 05 Feb-Mar 05 Jun-Aug 05 Jul-Aug 05 Tropidodispas sartorii X Elapidae: 1 Micrurus diastema X Viperidae: 4 Atropoides mexicanus X X Bothriechis marchi X Bothrops asper X X Cerrophidion godmani X Total (83 species) 41 species 17 species 51 species Geophis nephodrymus Townsend and Wilson.” Other specimens collected in 2004 and stored with Fundacon Fasquelle included Ninia pavimentata, a new country record for Honduras (Townsend et al. 2005a), and five species (Craugastor charadra, Anolis ocelloscapularis, Adelphicos quadrivigatum, Omoadiphas aurula, and Scaphiodontophis annulatus) that were additions to the known herpetofauna of Parque Nacional El Cusuco (Townsend et al. 2005b, 2006). Fieldwork within Parque Nacional El Cusuco during June, July and August 2005 was highly productive, with 41 species recorded (Table 2), 11 more than were reported from the park by Wilson and McCranie (2004). In addition to the nine specimens of Geophis, five species of salamanders, ten anurans, 11 lizards, and 15 species of snakes were recorded. At least two specimens collected Parque Nacional El Cusuco during 2005 are undescribed and new to science (Oedipina sp. [Figure 9] and Hyla sp. [Figure 10]), and an additional eight (Smilisca baudinii, Anolis capito, A. petersii, Lampropeltis triangulum, Pseustes poecilonotus, Rhadinaea montecristi, Atropoides mexicanus, and Bothrops asper) were added to the park’s herpetofauna (Talley et al. 2005, Townsend and Plenderleith 2005, Townsend et al. 2006a, Wilson et al., 2006).

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37 The result of 2005 fieldwork in Parque Nacional El Cusuco was to increase the number of species recorded in the park by two-thirds; from the total of 30 species reported by Wilson and McCranie (2004b) to 50 species (Townsend et al. 2006a). Results between survey sites differ as a result of the duration of the site visit, available habitat at each site, and intensity of survey at the site (number of herpetologists, extent of use of passive but intensive methods such as drift fences). In 2005, 23 species were Figure 9. An undescribed species of Oedipina (JHT 1553) from Cantiles, 1780 m, Parque Nacional El Cusuco, Corts, Honduras. Photograph by B. L. Talley. Figure 10. An undescribed species of bromeliad-dwelling hylid frog (JHT 1572) from bosque enano, 1990 m elevation, Parque Nacional El Cusuco, Corts, Honduras. Photograph by B. L. Talley.

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38 recorded from the Centro de Visitantes area, five species from Bosque Enano, 24 species from Guanales, 13 species from La Fortuna, and 14 species from Cantiles (Table 3). We did not conduct a systematic survey of the vicinity of the community of Buenos Aires, but 17 species were recorded there at elevations intermediate to those in Parque Nacional El Cusuco and El Paraiso (Table 2). The first Honduran record of the snake Ninia pavimentata was collected in the vicinity of Buenos Aires (Townsend et al. 2005a). Fieldwork at El Paraiso between 29 June and 16 August 2005 produced 51 species, including one species of caecilian, two species of salamanders, 10 species of anurans, two species of turtles, 17 species of lizards, and 19 species of snakes (Table 2). One snake, Rhadinaea anachoreta, was added to the Honduran herpetofauna (Townsend et al., 2005c), and one species of salamander, Oedipina elongata (Figure 11), represented the first verified record for Honduras (Townsend et al., 2006b). Figure 11. Oedipina elongata (UF 144649) from El Paraiso Valley, 185 m, Corts, Honduras. Photograph by J. C. Nifong. Only ten species were collected at both El Paraiso and inside Parque Nacional El Cusuco, two anurans, Smilisca baudinii and Craugastor charadra, one lizard, Anolis capito, and seven snakes, Adelphicos quadrivigatum, Atropoides mexicanus, Bothrops

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39 asper, Imantodes cenchoa, Lampropeltis triangulum, Leptophis ahaetulla, and Tantilla schistosa; only three species, Smilisca baudinii, Adelphicos quadrivigatum, and Tantilla schistosa, were recorded from all three areas.

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40 Table 3. Site-by-site breakdown of amphibian and reptiles species collected within Parque Nacional El Cusuco during 2005. Centro de Visitantes Bosque Enano Guanales La Fortuna Cantiles Species Feb-Mar 05 Jun-Aug 05 Jun-Aug 05 Feb-Mar 05 Jun-Aug 05 Jul 05 Jul-Aug 05 Plethodontidae: 5 Bolitoglossa conanti X X X X X Bolitoglossa diaphora X X X Bolitoglossa dunni X Cryptotriton nasalis X X “Oedipina” sp. X Hylidae: 7 Bromeliohyla bromeliacia X X Duellmanohyla soralia X X X X “Hyla” sp. X Plectrohyla dasypus X X X X Plectrohyla exquisita X X X Ptychohyla hypomykter X X X X Smilisca baudinii X Leptodactylidae: 2 Craugastor charadra X Craugastor rostralis X X X X X Anguidae: 2 Celestus montanus X Mesaspis moreletii X X Phrynosomatidae: 2 Sceloporus malachiticus X X X X X Sceloporus variabilis X Polychrotidae: 7 Anolis amplisquamosus X X Anolis capito X Anolis cusuco X X X Anolis johnmeyeri X X X X Anolis ocelloscapularis X X Anolis petersii X X Anolis sp X X X X Scincidae: 1 Sphenomorphus incertus X X X Colubridae: 12 Adelphicos quadrivirgatum X X Drymobius chloroticus X X Geophis nephodrymus X X X Imantodes cenchoa X Lampropeltis triangulum X Leptophis ahaetulla X Mastigodryas dorsalis X Pseustes poecilonotus X Rhadinaea montecristi X Tantilla schistosa X X Elapidae: 1 Micrurus diastema X X Viperidae: 4 Atropoides mexicanus X Bothriechis marchi X X X Bothrops asper X Cerrophidion godmani X X Site subtotals 14 20 5 2 24 13 13 TOTAL: 43 23 5 24 13 13

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CONSERVATION ASSESSMENT OF THE HERPETOFAUNA OF THE SIERRA DE OMOA The herpetofauna of the Sierra de Omoa has been well documented, owing in large part to its proximity to San Pedro Sula, the second largest city in Honduras and a major international point of entry for the country. Numerous endemic species of amphibians and reptiles have been described from the mountain range over the past 70+ years (Khler et al. 2001, McCranie and Wilson 2002, Schmidt 1933, Townsend and Wilson 2006). The herpetofauna of Parque Nacional El Cusuco, which protects the remaining cloud forests in the Sierra de Omoa, is also well documented (Wilson and McCranie 2004b, Townsend et al. 2006a). Fieldwork summarized in the previous chapter, supplemented by literature records and museum data, confirms 126 species of amphibians and reptiles known to occur in the Sierra de Omoa (Table 4) and 50 species in Parque Nacional El Cusuco (Townsend et al. 2006a). Including the undescribed species of “Hyla”, Oedipina, and Rhadinaea mentioned in the previous chapter, Honduras has a total of 352 species of amphibians and reptiles reported from within its borders, of which six have marine distributions (McCranie 2004a, 2004b, 2004c; McCranie and Castaeda 2004a; McCranie and Wilson 2002; McCranie et al. 2002, 2003a, 2003b, 2005; Townsend et al. 2005a, 2005c; Wilson and McCranie 2002; Wilson et al. 2003). The species known from Parque Nacional El Cusuco represent 14.5% of the 346 species recorded from mainland and insular habitats in Honduras, and the species recorded from the Sierra de Omoa represent 36.4% of the 41

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42 Table 4. Inventory and conservation assessment of the amphibians and reptiles of the Sierra de Omoa, Honduras Distribution: SO = endemic to Sierra de Omoa; eNCA = endemic to eastern Nuclear Central America; NCA = endemic to Nuclear Central America; WS = widespread, found beyond Nuclear Central America. Source: BA = species recorded at Buenos Aires, 900–1350 m elevation, during 2004 and 2005; CU = species recorded in Parque Nacional El Cusuco, 1220 m elevation, during 2004 and 2005; EP = species recorded in El Paraiso Valley Ecological Reserve, 1 m elevation, during 2005; OR = other records, taken from the literature, museum records, and other sources. Methods for deriving EVS score and conservation status are described in text; EVS scores from 3 are considered low vulnerability, from 10 indicate medium vulnerability, and from 14 indicate high vulnerability; S =at least some populations stable; D = all populations believed to be declining; E = extinct at all known localities in the Sierra de Omoa; N = no or insufficient data. Species Distribution Source EVS score Status Caecilidae: 1 Dermophis mexicanus WS EP 12 S Plethodontidae: 8 Bolitoglossa conanti eNCA BA, CU 14 S Bolitoglossa diaphora SO CU 16 S Bolitoglossa dofleini NCA BA 14 D Bolitoglossa dunni eNCA CU 14 S Bolitoglossa rufescens eNCA EP 12 S Cryptotriton nasalis SO CU 15 S Oedipina elongata eNCA EP 15 N Oedipina sp. SO CU 16 N Bufonidae: 3 Chaunus marinus WS EP, BA 5 S Cranopsis campbelli NCA EP 10 S Cranopsis valliceps WS EP, BA 5 S Centrolenidae: 1 Hyalinobatrachium fleischmanni WS EP 9 S Hylidae: 12 Agalychnis callidryas WS OR 10 S Bromeliohyla bromeliacia NCA CU 15 S Dendropsophus microcephalus WS OR 5 S Duellmanohyla soralia eNCA BA, CU 10 D “Hyla” sp. SO CU 18 N Phrynohyas venulosa WS OR 5 S Plectrohyla dasypus SO CU 13 D Plectrohyla exquisita SO CU 13 S Ptychohyla hypomykter NCA BA, CU 9 D Scinax staufferi WS OR 5 S Smilisca baudinii WS EP, BA, CU 4 S Tlalocohyla loquax WS OR 6 S Leptodactylidae: 11 Craugastor chac NCA EP 14 S Craugastor charadra eNCA EP, CU 13 S Craugastor laevissimus eNCA OR 9 S Craugastor laticeps WS OR 14 N Craugastor merendonensis SO OR 17 D Craugastor milesi eNCA OR 15 E Craugastor omoaensis SO OR 16 E Craugastor rostralis eNCA BA, CU 14 S

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43 Table 4. Continued. Species Distribution Source EVS score Status Leptodactylus fragilis WS OR 6 S Leptodactylus melanonotus WS EP 6 S Physalaemus pustulosus WS OR 6 S Microhylidae: 1 Hypopachus variolosus WS OR 6 S Ranidae: 3 Lithobates brownorum WS EP 3 S Lithobates maculatus NCA EP, BA 6 S Lithobates vallanti WS OR 7 N Rhinophrynidae: 1 Rhinophrynus dorsalis WS OR 9 N Kinosternidae: 1 Kinosternon leucostomum WS EP 9 S Emydidae: 1 Trachemys venusta WS EP 12 D Geoemydidae: 1 Rhinoclemmys pulcherrima WS OR 9 N Anguidae: 2 Celestus montanus eNCA CU 14 S Mesaspis moreletii NCA CU 13 S Corytophanidae: 4 Basiliscus vittatus WS EP, BA 7 S Corytophanes cristatus WS EP 11 S Corytophanes hernandesii WS OR 12 N Laemanctus longipes WS EP 9 S Gekkonidae: 5 Coleonyx mitratus WS OR 10 N Hemidactylus frenatus* WS EP Sphaerodactylus dunni NCA OR 14 N Sphaerodactylus millepunctatus WS EP 7 S Thecadactylus rapicauda WS OR 10 N Gymnophthalmidae: 1 Gymnophthalmus speciosus WS OR 8 N Iguanidae: 2 Ctenosaura similis WS EP 11 S Iguana iguana WS EP 12 S Phrynosomatidae: 2 Sceloporus malachiticus WS CU 8 S Sceloporus variabilis WS BA, CU 7 S Polychrotidae: 12 Anolis amplisquamosus SO CU 16 D Anolis biporcatus WS OR 10 N Anolis capito WS EP, CU 11 S Anolis cusuco SO CU 16 S Anolis johnmeyeri eNCA CU 15 S Anolis lemurinus WS EP 9 S Anolis ocelloscapularis eNCA CU 15 S Anolis petersii NCA CU 13 S Anolis rodriguezii NCA EP 10 S Anolis sericeus WS EP 7 S Anolis tropidonotus NCA BA 5 S Anolis uniformis NCA EP 11 S Scincidae: 2 Sphenomorphus cherriei WS EP, BA 7 S Sphenomorphus incertus NCA CU 12 S Teiidae: 4 Ameiva festiva WS EP 10 S Ameiva undulata WS EP 7 S Aspidoscelis deppii WS OR 8 N Cnemidophorus lemniscatus WS EP 12 S

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44 Table 4. Continued. Species Source EVS score Status Xanthusidae: 1 Lepidophyma flavimaculatum WS EP, BA Distribution 11 S Colubridae: 38 Adelphicos quadrivirgatum WS EP, BA, CU 8 S Clelia clelia WS EP 11 S Coniophanes bipunctatus WS OR 11 N Coniophanes fissidens WS OR 9 N OR 9 N Drymarchon melanurus WS BA 9 N Conophis lineatus WS Drymobius chloroticus WS BA, CU 11 S Drymobius margaritiferus WS EP 7 S Geophis nephodrymus SO CU 14 S Imantodes cenchoa WS EP, CU 6 S Lampropeltis triangulum WS EP, CU 9 S Leptodeira annulata WS OR 8 N Leptodeira septentrionalis WS OR 9 N Leptophis ahaetulla WS EP, CU 8 S Leptophis mexicanus WS EP, BA 8 S Mastigodryas dorsalis NCA CU 12 D Mastigodryas melanolomus WS EP 9 S Ninia diademata WS BA 8 S Ninia pavimentata NCA BA 12 N Ninia espinali eNCA OR 12 N Ninia sebae WS EP, BA, CU 4 S Omoadiphas aurula SO CU 15 N Oxybelis aeneus WS OR 9 N Oxyrhopus petula WS EP 13 N Pseudelaphe flavirufa WS OR 12 N Pseustes poecilonotus WS CU 12 S Rhadinaea anachoreta eNCA EP 12 N Rhadinaea montecristi eNCA CU 12 N Rhadinaea sp. SO OR 14 N Scaphiodontophis annulatus WS EP, BA 12 S Senticolis triaspis WS OR 10 N Sibon nebulatus WS EP 8 S Spilotes pullatus WS EP 9 S Stenorrhina degenhardtii WS OR 10 S Tantilla schistosa WS EP, BA, CU 10 S Tantillita lintoni WS EP 13 S Tretanorhinus nigroluteus WS OR 8 N Tropidodipsas sartorii NCA EP 12 S Xenodon rabdocephalus WS OR 12 N Elapidae: 2 Micrurus diastema NCA BA, CU 12 S Micrurus nigrocinctus WS OR 9 N Viperidae: 7 Atropoides mexicanus NCA EP, BA, CU 12 S Bothriechis marchi eNCA CU 16 D Bothriechis schlegelii WS OR 12 N Bothrops asper WS EP, CU 12 S Cerrophidion godmani WS CU 12 S Crotalus simus WS OR 12 N Porthidium nasutum WS OR 12 N Species total: 126 * Hemidactylus frenatus is an introduced species, and is not included in conservation analyses as this species is primarily found around human habitation.

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45 non-marine herpetofauna. Of the species known from the Sierra de Omoa, 45 (35.7%) are restricted in distribution to NCA and 13 (10.3%) are endemic to the Sierra de Omoa (Table 4). The high level of endemism present in the Sierra de Omoa makes this region, and particularly Parque Nacional El Cusuco, critically important in conservation value. Since Wilson and McCranie (2004b) provided the first summary of the herpetofauna of Parque Nacional El Cusuco, the number of documented species of amphibians and reptiles known from the park has increased from 30 to 50 (Townsend et al. 2006a). More important in terms of conservation implications is the fact that four previously undescribed species, one salamander, one frog, and two snakes, have been discovered in the park since Wilson and McCranie (2004b), all of which are considered endemic to Parque Nacional El Cusuco (Townsend et al. 2006a). There are 13 species of amphibians and reptiles that are endemic to the Sierra de Omoa: three salamanders (Bolitoglossa diaphora, Cryptotriton nasalis, and Oedipina sp.), five anurans (Craugastor merendonensis, C. omoaensis, “Hyla” sp., Plectrohyla dasypus, and P. exquisita), three snakes (Geophis nephodrymus, Omoadiphas aurula, and Rhadinaea sp.), and two lizards (Anolis amplisquamosus and A. cusuco). Eleven of these 13 are found in Parque Nacional El Cusuco. Craugastor merendonensis and C. omoaensis are “single-site endemics” currently known only from localities in the Sierra de Omoa that are outside of the park’s boundaries. In addition to the endemic species of the Sierra de Omoa, 16 other species found there are endemic to the next broader biogeographical division, eNCA: four salamanders (Bolitoglossa conanti, B. dunni, B. rufescens, and Oedipina elongata), five anurans (Craugastor charadra, C. laeviventris, C. milesi, C. rostralis, and Duellmanohyla soralia), four snakes (Bothriechis marchi,

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46 Ninia espinali, Rhadinaea anachoreta, and R. montecristi), and three lizards (Anolis johnmeyeri, A. ocelloscapularis, and Celestus montanus). Another 16 species found in the Sierra de Omoa are endemic to Nuclear Central America: one salamander (Bolitoglossa dofleini), five anurans (Bufo campbelli, Bromeliohyla bromeliacia, Craugastor chac, Ptychohyla hypomykter, Rana maculata), three snakes (Atropoides mexicanus, Mastigodryas dorsalis, Tropidodipsas sartorii), and seven lizards (Anolis petersii, A. rodriguezii, A. tropidonotus, A. uniformis, Mesaspis moreletii, Sphaerodactylus dunni, Sphenomorphus incertus). The cloud forest herpetofauna of the Sierra de Omoa, confined to Parque Nacional El Cusuco, shares a marked similarity with that found in the vicinity of Parque Nacional Cerro Azul, ca. 75 km to the southwest (Figure 2; Wilson and McCranie 2004a, 2004b; McCranie 2004). This string of mountains along the western border of Honduras has been referred to collectively as the Cordillera de Merendn, and in Guatemala, portions of this same mountain range are also referred to as the Sierra de Caral and Sierra de Espritu Santo. These two parks are grouped together as the cloud forest area referred to as the “Northwestern Highlands (NWH)” by Wilson and McCranie (2003, 2004a). Subhumid interior valleys, where most of the natural vegetation has been converted to agriculture, now separate these two highland areas, effectively isolating the specialized highland species found in these “islands” of cloud forest habitat. Coefficient of Biogeographic Resemblance (CBR) matrices were developed to better evaluate the similarity between the herpetofaunas of Parque Nacional El Cusuco and Parque Nacional Cerro Azul and their separate and combined effectiveness at offering protection to species of conservation importance. Parque Nacional El Cusuco

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47 and Parque Nacional Cerro Azul were included in the evaluation, as well as Parque Nacional Pico Bonito, a park near the Caribbean coast in north-central Honduras that protects a highland area that demonstrates remarkably high endemism in amphibians and reptiles. Lists of herpetofauna of each park are taken from Townsend et al. (2006; Parque Nacional El Cusuco), McCranie (2004; Parque Nacional Cerro Azul), and McCranie and Castaeda (2005; Parque Nacional Pico Bonito). While all three parks protect areas within the Premontane Wet Forest (PWF) and Lower Montane Wet Forest (LMWF) formations, Parque Nacional Pico Bonito also has areas of Lowland Moist Forest (LMF) and, therefore, has lowland-restricted species included in its herpetofauna. To address this variation in ecophysiography between the three parks and to evaluate each park’s contribution to protecting species endemic to Honduras, sets of CBR values were generated for each park using its entire herpetofauna (Table 5), only highland species, herein considered to be species that occur only in or above the PWF formation and not in the LMF formation (Table 6), and only species that are endemic to Honduras (Table 7). A strong biogeographical relationship between Parque Nacional El Cusuco and Parque Nacional Cerro Azul is evident, as previously indicated by Wilson and McCranie (2003, 2004a). However, this relationship is not as evident when the entire herpetofauna reported from each park is used in the analysis (Table 5). Parque Nacional El Cusuco and Parque Nacional Cerro Azul have 28 species in common (56% of Cusuco herpetofauna) and a CBR value of 0.45, demonstrating a high degree of similarity, but Parque Nacional Cerro Azul and Parque Nacional Pico Bonito also have 25 species in common and a CBR value of 0.34, while Parque Nacional El Cusuco and Parque Nacional Pico Bonito have 13 species in common (26% of Cusuco herpetofauna). The similarity observed in the

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48 Table 5. Coefficient of Biogeographic Resemblance (CBR) matrix of overall herpetofaunal similarity between Parque Nacional El Cusuco, Parque Nacional Cerro Azul, and Parque Nacional Pico Bonito. Cusuco Cerro Azul Pico Bonito Cusuco 50 28 13 Cerro Azul 0.45 74 25 Pico Bonito 0.21 0.34 73 N = number of species in each park; N = number of species in common between two parks; N = CBR value. Table 6. Coefficient of Biogeographic Resemblance (CBR) matrix of similarity of the highland herpetofauna (found only in PWF and LMWF formations) in Parque Nacional El Cusuco, Parque Nacional Cerro Azul, and Parque Nacional Pico Bonito. Cusuco Cerro Azul Pico Bonito Cusuco 31 15 3 Cerro Azul 0.48 31 3 Pico Bonito 0.13 0.13 17 N = number of species in each park; N = number of species in common between two parks; N = CBR value. Table 7. Coefficient of Biogeographic Resemblance (CBR) matrix of similarity of herpetofauna endemic to Honduras and found in Parque Nacional El Cusuco, Parque Nacional Cerro Azul, and/or Parque Nacional Pico Bonito. Cusuco Cerro Azul Pico Bonito Cusuco 17 4 1 Cerro Azul 0.35 6 1 Pico Bonito 0.06 0.08 19 N = number of species in each park; N = number of species in common between two parks; N = CBR value. herpetofaunal diversity of all three parks is attributed to the inclusion of widespread, primarily lowland inhabiting species in the CBR analysis presented in Table 5. By including widespread lowland species this set of comparisons is rendered the least informative in evaluating the distinctiveness or similarity of the three parks’ herpetofaunas, due to the absence of lowland habitat and therefore paucity of lowland inhabiting species in two of the parks, particularly Parque Nacional El Cusuco. When species that inhabit the LMF formation are excluded from the analysis and only the highland herpetofaunas are compared, the relationships between the three parks

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49 are more clearly elucidated (Table 6). Parque Nacional El Cusuco and Parque Nacional Cerro Azul continue to exhibit a high degree of similarity, with 15 highland species in common (48.4 % of Cusuco highland herpetofauna) and a CBR value of 0.48. At this stage Parque Nacional Pico Bonito is shown to be notably distinctive from the Cusuco/Cerro Azul herpetofaunas, having only three highland species in common with each park (Sceloporus malachiticus and Drymobius chloroticus with both, Bothriechis marchi with Cusuco, and Craugastor laticeps with Cerro Azul). Considering only Honduran endemic species, the similarity between Parque Nacional El Cusuco and Parque Nacional Cerro Azul is again very apparent, but added emphasis on the importance and distinctiveness of the herpetofauna of Parque Nacional El Cusuco becomes warranted (Table 7). These two parks have four Honduran endemic species in common (Bolitoglossa conanti, Craugastor milesi, Anolis johnmeyeri, and A. ocelloscapularis) and a CBR value of 0.35. But Parque Nacional Cerro Azul only protects a total of six Honduran endemic species, meaning that two-thirds of Honduran endemic species in Parque Nacional Cerro Azul are shared with Parque Nacional El Cusuco, a park with 17 Honduran endemic species making up 34% of its overall herpetofauna (Table 7). Of those 17 species, four previously mentioned species are also found in Parque Nacional Cerro Azul and only one (Bothriechis marchi) is found in Parque Nacional Pico Bonito, leaving eleven species endemic to Honduras (Bolitoglossa diaphora, Cryptotriton nasalis, Oedipina sp., Hyla sp., Plectrohyla dasypus, P. exquisita, Geophis nephodrymus, Omoadiphas aurula, and Rhadinaea sp., Anolis amplisquamosus and A. cusuco) and one species endemic to eNCA (Celestus montanus) that are only known to occur within one protected area, Parque Nacional El Cusuco (Table 4).

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50 Environmental Vulnerability Scores (EVS) were used to evaluate the susceptibility of the herpetofauna of the Sierra de Omoa to human induced decline. For recently discovered and recently reported species whose EVS had not previously been evaluated, new EVS were generated (Table 8). Of the 126 species of amphibians and reptiles found in the Sierra de Omoa, 51 are categorized as low vulnerability, 50 are categorized as medium vulnerability, and 25 are categorized as high vulnerability (Table 4). The 25 species of amphibians and reptiles classed as “high vulnerability” are considered to be the species of most conservation significance (Table 9). Eighteen of those 25 are restricted to highland and/or cloud forest habitat, and 11 of the most vulnerable species in the Sierra de Omoa are endemic to that mountain range (Table 9). At least two species of anurans, one endemic to the Sierra de Omoa (Craugastor omoaensis) and one endemic to Honduras (C. milesi), have not been collected in the last 20+ years despite repeated efforts and are considered extinct (McCranie and Wilson 2002; Wilson and McCranie 2004b, 2004c; Townsend et al. 2006a ). Another species, C. merendonensis, has not been collected since 1968 and might also be extinct, despite the preservation of its type locality as a water production site for local communities (McCranie and Wilson 2002). Moreover, eight species found in the Sierra de Omoa are considered to have declining populations: one salamander (Bolitoglossa dofleini), three anurans (Duellmanohyla soralia, Plectrohyla dasypus, Ptychohyla hypomykter), one turtle (Trachemys venusta), one lizard (Anolis amplisquamosus), and two snakes (Mastigodryas dorsalis and Bothriechis marchi) (Table 4; Wilson and McCranie 2004b, Townsend et al. 2006a ).

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51 Table 8. Environmental Vulnerability Scores (EVS) for species of amphibians and reptiles that were recently described or reported from Honduras. Species Geographic Distribution Ecological Distribution Reproductive Mode/Human Persecution Total = EVS score Oedipina sp. 4 8 4 16 Hyla sp. 5 8 5 18 Geophis nephodrymus 5 8 1 14 Ninia pavimentata 3 8 1 12 Rhadinaea anachoreta 3 8 1 12 Rhadinaea sp. 5 8 1 14 Table 9. Amphibians and reptiles of conservation significance from the Sierra de Omoa, ranked by EVS. Cus = Endemic to localities inside Parque Nacional El Cusuco; SO = Endemic to the Sierra de Omoa; eNCA = Restricted to eastern Nuclear Central America; NCA = Restricted to Nuclear Central America; WS = Widespread, distribution exceeds Nuclear Central America. Species Distribution Elevational range (m) EVS Status Comments “Hyla” sp. Cus 1990 18 N Breeds in bromeliads Craugastor merendonensis SO 150–200 17 D Probably extinct Bolitoglossa diaphora SO 1470–2200 16 S Oedipina sp. Cus 1780 16 N Craugastor omoaensis SO 760–1150 16 E Extinct Anolis amplisquamosus Cus 1530–2200 16 D Anolis cusuco Cus 1300–1935 16 S Bothriechis marchi eNCA 500 16 D Endemic to Honduras Cryptotriton nasalis SO 1220–2200 15 S Oedipina elongata eNCA 10–770 15 N Bromeliohyla bromeliacia NCA 1250 15 S Breeds in bromeliads Craugastor milesi eNCA 1050–1720 15 E Extinct Anolis johnmeyeri eNCA 1340 15 S Endemic to Honduras Anolis ocelloscapularis eNCA 1150 15 S Endemic to Honduras Omoadiphas aurula SO 1250–1800 15 N Endemic genus Bolitoglossa conanti eNCA 1370–2000 14 S Bolitoglossa dofleini NCA 650–1370 14 D Bolitoglossa dunni eNCA 1200–1600 14 S Craugastor chac NCA 20–1000 14 S Craugastor laticeps WS 650–1500 14 N Craugastor rostralis eNCA 1050–1800 14 S Celestus montanus eNCA 915–1780 14 S Sphaerodactylus dunni NCA 60 14 N Endemic to Honduras Rhadinaea sp. Cus 1550 14 N Geophis nephodrymus Cus 1545–1780 14 S One newly recorded species, an undescribed treefrog (Figure 10) that, prior to the large scale generic revision of Faivovich et al. (2005), would have been assigned to the genus Hyla, has an EVS of 18, one of the three highest scores reported from Honduras (Wilson and McCranie 2004b). Only one species, Ctenosaura bakeri, scored higher (19), and one

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52 species, C. oedirhina, scored the same. Both species are spiny-tailed iguanas that are endemic to the islands of Utila and Roatn, respectively, and are subject to both hunting pressure and intensive habitat degradation (McCranie et al. 2005). The single specimen of Hyla was collected as it jumped from an arboreal bromeliad in bosque enano at 1990 m elevation, and material being used to describe this frog was also obtained from bromeliads in Parque Nacional El Cusuco (J.R. McCranie, pers. comm.). Being in close proximity to one of the largest and fastest growing cities in Honduras, San Pedro Sula, has put the Sierra de Omoa at an ever increasing risk of environmental degradation in the wake of the country’s rapidly-expanding human population, which is currently growing at the fastest rate in the Western Hemisphere (Population Reference Bureau 2005). Clearly, the Sierra de Omoa and Parque Nacional El Cusuco represent an important area in terms of its herpetofauna, and in particular its suite of endemic species. Despite legal protection, Parque Nacional El Cusuco is under continuing colonization and deforestation pressure. In 2005, I saw numerous freshly cleared agricultural plots on the southern edge of the park to the west and north of the village of La Fortuna. Even more extensive clearing of highland and cloud forests has occurred in the northern part of the park, particularly in the areas of Nueva Esperanza and Agua Mansa (Lennkh 2005). Uncatalogued biodiversity clearly exists in the Sierra de Omoa (Anderson and Ashe 2000), and there is no way to know what species may already have disappeared with the loss of their habitat. Hopefully, the data presented here will help to bring additional attention to the significant biological diversity residing in the Sierra de Omoa, and to addressing the anthropogenic pressures that threaten the long-term survival of that diversity.

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STATUS OF Geophis, species inquirenda, AND A REDESCRIPTION OF Geophis nephodrymus TOWNSEND AND WILSON Differentiation of Geophis fulvoguttatus from G. species inquirenda The first of three potential taxonomic resolutions presented is that Geophis, species inquirenda represents a range extension and previously undocumented morphological variation for G. fulvoguttatus. This hypothesis is generated based on: a) similarities in the dorsal color pattern between G. fulvoguttatus and G. species inquirenda, and b) G. fulvoguttatus is reported to occur in Parque Nacional Cerro Azul, which shares a number of other cloud forest endemics with Parque Nacional El Cusuco (see previous chapter). After examination of four of the six known specimens of G. fulvoguttatus and their subsequent comparison to specimens of Geophis, species inquirenda, this hypothesis is demonstrated to be untenable, due to the extent of the difference in variation in ventral and segmental counts, color pattern, and a number of head scale characteristics between the two taxa (Table 10). Geophis fulvoguttatus is distinguished from Geophis, species inquirenda by usually having more ventrals (135 versus 120 in Geophis, species inquirenda), having higher total segmental counts (171 versus 149 in Geophis, species inquirenda), internasal common suture more than half the length of prefrontal common suture (internasal common suture about half or less than that of prefrontal common suture in Geophis, species inquirenda; ratio 4), supraocular 47% the length of the loreal 53

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54 Table 10. Summary of relevant measurements, scale counts, and variation in scale measurement ratios for the Geophis of eastern Nuclear Central America The data presented below is formatted as follows: snout-vent length (SVL) and tail length (TL) in millimeters; measurements, ventral and subcaudal scale counts, and scale measurement ratios appear with the range first, followed by the (sample size), then by the (mean, standard deviation), when no variation is present only the datum and (sample size) are given; supralabial, infralabial, supraocular, and postocular counts are given by left side/right side, with the range of variation on each side, and the (typical condition) demonstrates the counts present when discounting variation that occurs in <10% of the specimens examined. Characters G. damiani G. dunni G. fulvoguttatus G. hoffmanni G. nephodrymus G. rhodogaster SVL 256 (2) (261.5, 7.78) 309 (1) 281 (3) (310.3, 37.8) 73 (29) (133.5, 29.1) 92 (17) (165.2, 54.9) 248 (4) (284.8, 41.9) TL 47 (2) (53.5, 9.19) 58 (1) 38 (4) (45, 6.68) 10 (23) (22.1, 5.75) 12 (14) (25.9, 8.8) 55 (3) (61.3, 6.51) Ventrals 136 (2) (139.5, 4.95) 140 (1) 137 (3) (146.3, 10.06) 114 (30) (122.4, 4.24) 120 (17) (129.3, 5.95) 138 (4) (140, 1.63) Subcaudals 34 (2) (37.5, 4.95) 36 (1) 24 (5) (33, 5.1) 23 (23) (27.4, 2.5) 22 (14) (27, 3.38) 34 (3) (39.3, 4.73) Ventrals + subcaudals 177 (2) 176 (1) 171 (3) (177.3, 5.1) 144 (23) (149.5, 3.68) 149 (14) (155.4, 3.77) 176 (3) (179.3, 2.89) Supralabials 6/6 6/6 6/6 4/5 (5/5) 5/6 (6/6) 6/6 Infralabials 6/6 8/8 6/6 (6/6) 5/5 (6/6) 6/6 (6/6) 6/6 (6/6) Supraoculars 1/1 1/1 0/1 (1/1) 1/1 0/1 (1/1) 0/0 Postoculars 1/1 1/1 1/1 0–1/0–1 0/1 (1/1) 1/1 1 0.27.37 (2) (0.32, 0.07) 0.28 (1) 0.23.27 (2) (0.25, 0.3) 0.21.44 (25) (0.31, 0.06) 0.23.5 (13) (0.36, 0.08) 0.33 (2) 2 1.22.33 (2) (1.28, 0.08) 1.4 (1) 1.08.4 (2) (1.24, 0.23) 1.17.0 (25) (1.46, 0.2) 1.38.0 (13) (1.51, 0.19) 1.39.55 (2) (1.47, 0.11) 3 0.73.0 (2) (0.87, 0.19) 0.5 (1) 0.71.93 (2) (0.82, 0.16) 0.4.71 (25) (0.5, 0.08) 0.33.0 (13) (0.65, 0.15) 1.0.1 (2) (1.05, 0.07)

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55 Table 10. Continued. Characters G. damiani G. dunni G. fulvoguttatus G. hoffmanni G. nephodrymus G. rhodogaster 4 0.4.5 (2) (0.45, 0.07) 0.35 (1) 0.57.64 (2), (0.61, 0.05) 0.2.5 (25) (0.31, 0.06) 0.25.54 (13) (0.43, 0.09) 0.46.50 (2) (0.48, 0.03) 5 0.67.81 (2) (0.74, 0.1) 0.72 (1) 0.77.79 (2) (0.78, 0.01) 0.62.84 (25) (0.73, 0.05) 0.68.83 (13) (0.74, 0.05) 0.61.63 (2) (0.62, 0.01) 6 0.46.52 (2) (0.49, 0.04) 0.57 (1) 0.41.44 (2) (0.43, 0.02) 0.35.57 (25) (0.47, 0.06) 0.29.57 (13) (0.4, 0.07) 0.26.38 (2) (0.32, 0.08) 7 1.03.08 (2) (1.06, 0.04) 0.94 (1) 1.0.06 (2) (1.03, 0.04) 0.95.11 (25) (1.0, 0.06) 0.96.16 (13) (1.09, 0.06) 1.0.26 (2) (1.13, 0.18) 8 0.56.6 (2) (0.58, 0.03) 0.48 (1) 0.57.71 (2) (0.59, 0.17) 0.31.55 (24) (0.41, 0.06) 0.09.5 (11) (0.34, 0.12) – 9 0.55.67 (2) (0.61, 0.08) 0.64 (1) 0.56.60 (2) (0.58, 0.03) 0.4.0 (24) (0.66, 0.18) 0.25.0 (11) (0.61, 0.25) – 10 1.25.0 (2) (1.63, 0.53) 2.33 (1) 2.0.5 (2) (2.25, 0.35) 1.0.0 (25) (2.63, 1.17) 0.5.0 (11) (1.55, 0.9) – 11 0.5.52 (2) (0.51, 0.01) 0.45 (1) 0.51 (2) 0.41.51 (25) (0.46, 0.02) 0.45.54 (13) (0.51, 0.03) 0.48.49 (2) (0.48, 0.01) 12 0.78.87 (2) (0.83, 0.06) 0.71 (1) 0.75.97 (2) (0.86, 0.16) 0.48.79 (25) (0.66, 0.09) 0.62.84 (13) (0.73, 0.07) 0.62.68 (2) (0.65, 0.04) 13 1.58.61 (2) (1.59, 0.02) 1.58 (1) 1.71.89 (2) (1.8, 0.13) 1.24.71 (25) (1.36, 0.1) 1.38.75 (13) (1.48, 0.1) 1.39.4 (2) (1.39, 0.01) 14 1.0 (2) 1.8 (1) 1.14.0 (2) (1.57, 0.61) 0.67.0 (23) (1.12, 0.3) 0.89.67 (11) (1.16, 0.4) 1.56.57 (2) (1.56, 0.01) 15 1.2.22 (2) (1.21, 0.01) 1.08 (1) 1.0.08 (2) (1.04, 0.06) 0.71.33 (25) (1.03, 0.13) 0.86.2 (13) (0.99, 0.11) 0.39.91 (2) (0.65, 0.37) 16 1.06.12 (2) (1.09, 0.04) 0.78 (1) 1.26.5 (2) (1.38, 0.17) 0.67.0 (25) (0.79, 0.1) 0.9.5 (13) (1.2, 0.14) 0.88.0 (2) (0.94, 0.08) 17 1.78 (2) 1.77 (1) 1.4.73 (2) (1.57, 0.23) 1.57.75 (25) (2.13, 0.26) 1.14.0 (13) (1.68, 0.19) 1.42.82 (2) (1.62, 0.28) 18 1.88.0 (2) (1.94, 0.08) 1.87 (1) 2.26.36 (2) (2.31, 0.07) 1.58.36 (25) (1.91, 0.2) 2.0.71 (13) (2.35, 0.2) 1.9.12 (2) (2.01, 0.16) 19 1.6.0 (2) (1.8, 0.28) 1.92 (1) 1.75.11 (2) (1.93, 0.25) 1.57.0 (25) (2.24, 0.31) 1.6.4 (12) (1.97, 0.25) 1.7.0 (2) (1.85, 0.21) 20 3.2.75 (2) (3.47, 0.39) 3.58 (1) 4.13.78 (2) (4.46, 0.46) 3.17.2 (25) (4.29, 0.48) 3.8.2 (12) (4.6, 0.38) 3.6.8 (2) (3.7, 0.14) 21 1.0 .25 (2) (1.13, 0.18) 0.85 (1) 0.64.75 (2) (0.69, 0.08) 0.57.2 (25) (0.81, 0.15) 0.6.88 (12) (0.74, 0.09) 0.73.8 (2) (0.76, 0.05)

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56 Table 10. Continued. Characters G. damiani G. dunni G. fulvoguttatus G. hoffmanni G. nephodrymus G. rhodogaster 22 1.17.2 (2) (1.19, 0.02) 1.27 (1) 1.23.0 (2) (1.62, 0.54) 1.45.83 (25) (1.96, 0.32) 1.11.14 (12) (1.51, 0.28) 1.19.88 (2) (1.54, 0.49) 23 1.25.5 (2) (1.38, 0.18) 1.4 (1) 1.14.57 (2) (1.36, 0.3) 1.2.0 (24) (1.45, 0.21) 1.2.0 (12) (1.51, 0.23) 1.33.5 (2) (1.42, 0.12) 24 2.0.17 (2) (2.09, 0.12) 1.71 (1) 1.86 .33 (2) (2.1, 0.33) 1.25.67 (25) (1.86, 0.25) 1.17.0 (13) (1.54, 0.24) 1.88.89 (2) (1.88, 0.01) 25 1.73 .75 (2) (1.74, 0.01) 1.62 (1) 1.63.85 (2) (1.74, 0.16) 1.43.25 (25) (1.74, 0.23) 1.33.5 (13) (2.01, 0.3) 1.71.85 (2) (1.78, 0.1) 26 1.19.31 (2) (1.25, 0.08) 1.24 (1) 1.13.33 (2) (1.23, 0.14) 0.75.79 (25) (1.06, 0.21) 1.0.67 (13) (1.2, 0.17) 1.0.33 (2) (1.17, 0.23) 27 0.0 (2) 0 (1) 0.17.3 (2) (0.24, 0.09) 0.13.5 (25) (0.33, 0.09) 0.0.5 (13) (0.17, 0.18) 0.13.56 (2) (0.35, 0.3) Tail length/ total length 0.155.183 (2) (0.169, 0.02) 0.158 (1) 0.113.161 (3) (0.129, 0.03) 0.115.174 (24) (0.142, 0.02) 0.115.164 (11) (0.134, 0.02) 0.146.215 (4) (0.177, 0.03) 29 0.57, 0.67 (2) (0.62, 0.07) 0.67 (1) 0.5.75 (2) (0.63, 0.18) 0.25.75 (25) (0.55, 0.16) 0.0.6 (13) (0.27, 0.19) 0.5.67 (2) (0.58, 0.12) (supraocular 9% length of loreal in Geophis, species inquirenda; ratio 8), parietals somewhat more elongate (parietal length/parietal breadth 1.69.89 versus 1.38–1.75 in Geophis, species inquirenda; ratio 13), first infralabials broadly in contact between the mental and anterior chinshields (first infralabials narrowly in contact or not in contact in Geophis, species inquirenda; 29), a grayish brown anterior dorsal coloration that gradually becomes brownish black toward the posterior part of the body (dark gray anteriorly, becoming grayish black posteriorly in Geophis, species inquirenda), red-orange middorsal and dorsolateral blotches that rarely reach the level of the second dorsal scale row and never reach the level of the first dorsal scale row (red middorsal, dorsolateral and/or lateral blotches that regularly enter the first dorsal scale row in Geophis, species inquirenda), and chin scales that are immaculate yellowish white or

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57 white (versus chin scales gray or gray with some cream speckling on the posterior portions in Geophis, species inquirenda). The range in ventral and subcaudal counts demonstrated by the small series of G. fulvoguttatus as well as by Geophis, species inquirenda is consistent with that of other species of the dubius group, with no other species in the group having greater than a 20 scale intraspecific difference in ventral scale counts (Downs 1967). The total segmental count (ventrals plus subcaudals) of KU 183881 (181) is the upper limit of the range, with KU 57996 at the lower end of the segmental range (171) for G. fulvoguttatus. If Geophis, species inquirenda were to be considered conspecific with G. fulvoguttatus, then G. fulvoguttatus would demonstrate an exceptionally wide range in ventral counts (120; range of 37). Within the genus Geophis, this range would only be exceeded by G. semidoliatus (131; 48 scales) from Veracruz, Mexico, and approached by the highly variable G. brachycephalus (119; 34 scales) from Costa Rica and Panama, both of which occur in high densities with large numbers of specimens in museum collections. In light of the aforementioned evidence, the likelihood that an inclusive concept of G. fulvoguttatus that demonstrates an exceptional ventral count range coupled with diagnosable variation in other characters should be regarded as improbable, particularly given the other possible taxonomic resolutions. McCranie and Wilson (1991) reported on a specimen (KU 214782) consisting of a tail and the most posterior portion of the body of a snake, which they assigned to Geophis fulvoguttatus because it had 17 dorsal scale rows at the vent, 36 paired subcaudals, and red-orange middorsal spots and paired lateral blotches on the dorsal surface. This specimen was regurgitated by a Micrurus diastema collected above Quebrada Grande,

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58 1680 m elevation, Depto. Copn, Honduras. The record represents the only locality for G. fulvoguttatus from the Caribbean versant, which is a locality that is shared by a number of regional endemic species also found in the Sierra de Omoa (Wilson and McCranie 2004a, 2004b; McCranie 2004). This suggests the possibility that KU 214782 may actually be conspecific with Geophis, species inquirenda, since G. fulvoguttatus and Geophis, species inquirenda share similarities in coloration and overlapping subcaudal counts. The subcaudal count places KU 214782 above the known range of Geophis, species inquirenda (22) and at the upper limit of the known range of G. fulvoguttatus (24). An examination of KU 214782 revealed a color pattern that appears more consistent with that of G. fulvoguttatus than that of Geophis, species inquirenda, and, currently, assignment of KU 214782 to G. fulvoguttatus appears justified. Identity of Geophis, species inquirenda A single specimen of Geophis (JHT 1327) was collected at the end of the 2004 field season and stored at the Fundacon Hector Rodrigo Pastor Fasquelle in San Pedro Sula, Honduras, until it was retrieved in June 2005. This patternless specimen appeared virtually identical to the holotype of G. nephodrymus, but has distinct and separate supraocular and postocular scales like the specimens of Geophis, species inquirenda. JHT 1327 has two more ventral scales then the holotype of G. nephodrymus, and four more than the highest count for Geophis, species inquirenda, meaning the ventral counts now overlap. Another specimen, JHT 1379, collected on 15 July 2005 near the Centro de Visitantes of Parque Nacional El Cusuco, has fused supraoculars and postoculars on both sides, like the holotype of G. nephodrymus, but exhibited the color pattern of typical Geophis, species inquirenda. This evidence clearly supports acceptance of the second proposed resolution of the taxonomic status of Geophis, species inquirenda, that:

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59 Geophis, species inquirenda is conspecific with G. nephodrymus and represents a patterned variety of that species The fusion of the supraocular and postocular seen in the holotype of G. nephodrymus represents individual variation rather than a diagnosable character The species concept of G. nephodrymus needs to be expanded to include new variation Subsequent collections of Geophis from Parque Nacional El Cusuco have validated the inclusion of patterned snakes formerly referred to as Geophis, species inquirenda within a more inclusive concept of G. nephodrymus, and the full range of meristic and morphological variation within G. nephodrymus sensu lato is presented in the following systematic account and in Tables 11 and 12. Geophis nephodrymus Townsend and Wilson Figs. 12; Tables 11, 12 Geophis nephodrymus Townsend and Wilson, 2006: 151. Holotype: Florida Museum of Natural History (UF) 142577, an adult female collected 11 July 2004 by S. M. Hughes and J. H. Townsend. Type-locality: Sendero Las Minas (15.525’N, 88.705’W), 1570 m elevation, Parque Nacional El Cusuco, Corts, Honduras. Species group: dubius (Townsend and Wilson 2006). Distribution: Known from intermediate (1545 to1780 m) elevations in Parque Nacional El Cusuco, Sierra de Omoa, Departamento de Corts, Honduras (Figure 18). Diagnosis: Geophis nephodrymus has 17 dorsal scale rows throughout the body, distinguishing it from all members of the championi group (G. championi, G. downsi, G. godmani, and G. ruthveni) and semidoliatus group (G. cancellatus, G. juliai, G.

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60 laticinctus, and G. semidoliatus), and some members of the chalybeus (G. dugesii, G. nigrocinctus, and G. tarascae), omiltemanus (G. incomptus and G. maculiferus), and sieboldi (G. bellus, G. betaniensis, G. brachycephalus, G. damiani, G. hoffmanni, G. laticollaris, G. nigroalbus, G. petersii, G. russatus, G. ruthveni, G. sallaei, G. talamancae, and G. zeledoni) groups that have only 15 dorsal scale rows throughout the body. Of the remaining members of the chalybeus group, G. bicolor differs from G. nephodrymus in having 2 postocular scales (only 1 postocular in G. nephodrymus), and G. chalybeus has more ventrals (154) and subcaudals (38) than does G. nephodrymus (120 and 22, respectively). Geophis nephodrymus is separated from the members of the sieboldi group that possess 17 dorsal scale rows (G. dunni, G. nasalis, G. pyburni, and G. sieboldi), and from some members of the dubius group (G. carinosus, G. juarezi, and G. rostralis) by having smooth dorsal scales throughout the length of the body (dorsal scales keeled on at least the posterior part of the body in those species). Geophis nephodrymus differs from most of the members of the latifrontalis group (G. latifrontalis and G. mutitorques) and the members of the omiltemanus group that have 17 dorsal scale rows (G. isthmicus and G. omiltemanus) by lacking an anterior temporal (anterior temporal present in latifrontalis group [except G. blanchardi] and omiltemanus group). Geophis blanchardi differs from G. nephodrymus by having more ventrals (150 versus 120 in G. nephodrymus) and usually 7 infralabials (usually 6 infralabials in G. nephodrymus). Of the remaining species in the dubius group, G. nephodrymus can be differentiated from G. dubius by having internasals that are distinct from the prefrontals (internasals

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61 Figure 12. Juvenile Geophis nephodrymus (JHT 1334), demonstrating the patternless variant, from Sendero El Quetzal, 1545 m, Parque Nacional El Cusuco, Corts, Honduras. Photograph by B. L. Talley. Figure 13. Juvenile Geophis nephodrymus (UF 143024), red patterned variant, from finca de Makrn Ramrez, 1580 m, Parque Nacional El Cusuco, Corts, Honduras. Photograph by J. H. Townsend.

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62 usually fused to prefrontals in G. dubius), from G. anocularis, G. duellmani, and G. rhodogaster by having one supraocular and one postocular (supraocular and postocular absent in G. anocularis and G. duellmani; supraocular absent in G. rhodogaster), from G. immaculatus by having a dark coloration on chin and lateral edges of ventrals (chin and ventral scales immaculate in G. immaculatus), and from G. fulvoguttatus by usually having fewer ventrals (120) and a lower segmental count (149)(G. fulvoguttatus with 135 ventrals, segmental count 171). Description: Dorsal scales in 17 rows, smooth throughout length of body; no apical pits; ventrals 120, 122 in six males, 128 in six females, and 120–134 in five unsexed juveniles; subcaudals 22, 22 in six males, 24 in four females (with complete tails), and 23 in four unsexed juveniles (with complete tails); segmental count 149, 151 in six males, 157 in four females (with complete tails), and 149 in four unsexed juveniles (with complete tails). Snout-vent length 105 mm in six males, 130 mm in four females (with complete tails), 92 mm four unsexed juveniles (with complete tails); tail lengths 20–37 mm in six males, 22 mm in four females, and 12 mm in four juveniles; tail length/total length ratio 0.119.164 in six males, 0.119.145 in four females, and 0.115.131 in four unsexed juveniles. Head indistinct or slightly distinct from neck; snout bluntly rounded from above, rounded in profile, extending beyond the anterior end of the lower jaw; lower jaw reaches to the level of the middle of the first supralabial; rostral wider than high, extending at least somewhat posteriorly between internasals, portion of rostral visible in dorsal view 0.23.5 times the length of its distance from the frontal, with posterior termination at

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63 Figure 14. Adult male Geophis nephodrymus (JHT 1342), red patterned variant, from Sendero Las Minas, 1580 m, Parque Nacional El Cusuco, Corts, Honduras. Photograph by P. J. Lewis. Figure 15. Juvenile Geophis nephodrymus (JHT 1343), pale gray patterned variant, from Sendero Las Minas, 1560 m, Parque Nacional El Cusuco, Corts, Honduras. Photograph by B. L. Talley.

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64 Figure 16. Geophis nephodrymus from Cantiles, 1780 m, Parque Nacional El Cusuco, Corts, Honduras. Photograph by P. Wood. anterior portion of nostril; internasals 1.33.0 times as wide as they are long, angulate anteriorly, contacting the prenasal and postnasal laterally, their length 0.33.0 times as long as prefrontal common suture, their common suture 0.18.54 times as long as prefrontal common suture; prefrontals contacting postnasal, loreal, and orbit laterally, their length 0.67.88 times snout length, their common suture 0.29.57 times the frontal length; frontal 0.96.16 times as wide as is long, angulate anteriorly, in very narrow to moderate contact with supraoculars (length of frontal contact with supraocular 0.25.0 times supraocular length, 0.5.0 times length of prefrontal contact with supraocular-postocular), supraocular usually distinct (supraocular and postocular fused on both sides in UF 142577 and JHT 1379), moderately sized, triangular, narrowly contacting postocular, somewhat smaller than eye, 0.09.54 times as long as loreal; parietals 1.38.75 times as long as broad, their length 0.45.54 times the length of the head, their common suture 0.58.84 times as long as frontal; single postocular usually distinct from supraocular, 0.5.0 times as high as long, slightly smaller than supraocular; nasal divided; postnasal length 0.57.2 times that of prenasal length;

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65 Figure 17. Semidiagramatical illustration of the dorsal, lateral, and ventral aspects of the head of Geophis nephodrymus (UF 143022), from Sendero Las Minas, 1580 m, Parque Nacional El Cusuco, Corts, Honduras (artwork by S. LaRusso). combined length of prenasal and postnasal 0.85.38 times the length of the loreal; loreal 1.14.0 times as long as high, contained 2.0.71 times on snout length; 1.6.6 times as long as eye horizontal diameter, dorsal margin straight; eye small, contained 3.8.2 times in snout length, its vertical diameter 0.6.88 times its distance from the lip. Supralabials usually 6/6 (5/6 in JHT 1382 due to fusion of third and fourth left supralabials); first supralabial reduced in size, contacting rostral and prenasal; second supralabial contacting prenasal, postnasal, and loreal; third supralabial contacting loreal; third and fourth supralabials entering orbit; fourth supralabial contacting postorbital portion of postocular; fifth supralabial largest, contacting postocular and parietal; lip exposure of third supralabial 1.2.0 times that of second supralabial; lip exposure of fifth supralabial 1.11.14 times that of fourth supralabial; anterior temporal absent; one posterior temporal, separating sixth supralabial from parietal; posterior temporals sometimes fused with one nuchal on each side, separated from each other by three to five

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66 nuchals; mental 1.17.0 times as broad as long, rounded anteriorly, separated from chinshields by first infralabials; infralabials 6/6 (6/7 in JHT 1334 due to division of third right infralabial); first infralabials may be broadly in contact between mental and anterior chinshields, narrowly in contact, or separated by the mental and not in contact; first, second, and third infralabial contacting the anterior chinshield; fourth infralabial contacting posterior chinshield; anterior chinshields 1.33.5 times as long as broad, 1.0–1.67 times as long as posterior chinshields; posterior chinshields in narrow contact or not in contact and separated by first medial gular scale; two or three medial gular scales present. Coloration: This species shows a high degree of variation in coloration and color pattern. There are certain consistencies in the coloration of G. nephodrymus, such as having a gray dorsal ground coloration that becomes darker towards the posterior part of the body, a pale gray to yellowish cream venter with gray mottled lateral edges on the ventral scales, gray infralabials and gray mottled chinshields, and an essentially uniform dark gray tail, with pale mottling sometimes present on the subcaudals. There is a gradient in dorsal color pattern ranging from patternless (four of 17 specimens) to extensively marked with bands, laterally offset partial bands, and lateral blotches that range from pale grayish cream to brick red in color. Below are color notes for six specimens of G. nephodrymus, representing the broad spectrum of variation in dorsal pattern within the species. Color notes of JHT 1382, a patternless adult male: dorsal surface of head bluish gray, infralabials gray, chinshields mostly gray with some pale gray mottling posteriorly; dorsal surface of body blackish gray, becoming somewhat paler and more brownish

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67 laterally; dorsum also becoming darker and more brownish posteriorly; ventral scales pale gray with at least outer one-third of scale at each side mottled dark gray with some scales being almost completely mottled gray; dorsal and ventral surfaces of tail dark blackish gray. JHT 1337, a female, exhibits the most reduced color pattern of any of the specimens that were examined: dorsal surface of body brownish gray, becoming darker posteriorly; there is a series of about seven small, laterally offset pairs of spots on the posterior one-third of the body, orangish cream in color, all contained within the first three dorsal scale rows, all less than two dorsal scales in length, with poorly defined edges that mottle into the dorsal ground color. Color notes of USNM 561825, an adult female exhibiting reduced dorsal color pattern, are as follows: dorsal surface of head gray, ventral surface gray with some cream mottling on the posterior portions of the chinshields, infralabials gray; dorsum dark gray with about nine pairs of poorly delineated, laterally offset grayish pink dorsolateral spots with mottled edges, most apparent on the posterior portion of the body; dorsal ground color becoming somewhat darker posteriorly; ventral surface whitish with dark smudging near the lateral edges of the ventrals; dorsal surface of tail grayish black, lacking any pattern, ventral surface of tail gray with whitish mottling, becoming uniform grayish black on posterior half of tail. Another specimen exhibiting a reduced color, JHT 1378, has 18 laterally offset pairs of pale red spots, round in shape with irregular edges, all contained within the first six dorsal scale rows, most contained within the second and fifth dorsal scale rows, and has an otherwise typical dorsal and ventral coloration.

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68 UF 143022, a typically patterned adult male, has: dorsal surface of head dark gray, somewhat darker dorsally than laterally; chin and infralabials gray, with some cream speckling visible in the central areas of the chin and throat; dorsal scales grayish black, becoming somewhat darker posteriorly; dorsum marked with about five brick red crossbands and about 10 pairs of brick red partial bands that are laterally offset along the longitudinal axis, becoming more distinctive towards the posterior portion of the body, regularly entering the first dorsal scale row, typically 1 dorsal scales wide, separated by 4 dorsal scales, pigment only entering the middorsal scale row as crossbands, partial bands laterally or dorsolaterally positioned; ventral surface whitish with gray smudging on the lateral edges of the ventral scales, ventral color becoming slightly darker and pinkish-orange posteriorly; dorsal surface of tail grayish black, becoming black toward the terminus; one dorsal crossband present at level of vent, three pairs of offset partial bands and two small crossbands present on dorsal surface of tail; ventral surface of tail with some pinkish-cream mottling on a dark gray ground color, becoming uniform grayish black towards terminus. One of the more unusual specimens of G. nephodrymus in terms of color pattern is JHT 1343 (Figure 15), a heavily patterned juvenile with pale grayish cream (rather than red) markings. Dorsal and ventral coloration is typical, the dorsum is marked with seven complete bands that extend from the vertebral row to the first or second dorsal scale rows, 11 sets of laterally offset partial bands that reach laterally to between the fifth and seventh dorsal scale rows, and three elongate lateral blotches that do not extend laterally beyond the third dorsal scale row but are four to eight dorsal scales in length. JHT 1343 is

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69 one of the most heavily marked examples of G. nephodrymus, and the only one exhibiting pale grayish cream dorsal markings. Natural History Comments: Of the 17 known specimens, 14 were obtained from pitfall traps associated with drift fence arrays, and all specimens were from disturbed and undisturbed Pinus and Liquidambar Transitional Cloud Forest and undisturbed High-Diversity Broadleaf Cloud Forest (forest type definitions follow Meja V. 2001), in the Lower Montane Wet Forest (LMWF) formation. Other species of reptiles and amphibians collected in or in the immediate vicinity of pitfall traps include Bolitoglossa conanti, B. diaphora, Cerrophidion godmani, Drymobius chloroticus, Tantilla cf. schistosa, Anolis cusuco, Anolis johnmeyeri, and Sphenomorphus incertus. Two juveniles (UF 143024) collected on 2 March 2005 were found mid-morning under a pile of plant debris on a small agricultural plot cut from Pinus and Liquidambar Transitional Cloud Forest. One juvenile (JHT 1334) was collected at about 2200 h on 25 June 2005 as it crawled across Sendero El Quetzal, near the Centro de Visitantes. Remarks: Geophis nephodrymus (JHT 1327; snout-vent length 259 mm) is notably smaller (maximum snout-vent length 259 mm) than the other highland-inhabiting species of Geophis in eNCA, and demonstrates the least contact between the first pair of infralabials of any species from eNCA (Table 10). JHT 1378 has a peculiar deformity taking the place of the left eye, which is replaced with an irregular, pitted, opaque lens the covers that area of the left orbit and left supraocular and postocular scales. References: Townsend and Wilson (2006), Townsend et al. (2006).

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70 Illustrations: Townsend and Wilson (2006; black and white photograph and illustration of the dorsal and lateral aspects of the head of the holotype), Townsend et al. (2006; color photograph). Specimens examined: 17; HONDURAS: Corts: Sendero El Danto, Parque Nacional El Cusuco, Sierra de Omoa, 1555 m (UF 143023, USNM 561825, JHT 1327); Sendero El Quetzal, Parque Nacional El Cusuco, Sierra de Omoa, 1545 m (JHT 1334); Sendero Las Minas, Parque Nacional El Cusuco, Sierra de Omoa, 1560 m (UF 142577 [holotype], UF 143022, USNM 561824, JHT 1337, 1342, 1378, 1382, 1536); finca de Makrn Ramrez, Parque Nacional El Cusuco, Sierra de Omoa, 1580 m (UF 143024). Other records: 1; HONDURAS: Corts: Cantiles, Parque Nacional El Cusuco, Sierra de Omoa, 1780 m (Figure 16).

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71 Table 11. Morphological and meristic characters for Geophis nephodrymus, part I. Honduras: Corts: Parque Nacional El Cusuco, 1545 m Characters holotype UF 142577 UF 143022 UF 143023 UF 143024 juv. UF 143025 juv. USNM 561824 juv. USNM 561825 JHT 1379 SVL 223 187 130 97 92 119 198 219 TL 30 34 22 13 12 18 28 32 Ventrals 136 124 128 134 126 120 132 135 Subcaudals 24 31 32 25 23 29 25 25 Ventrals + subcaudals 160 155 160 159 149 149 157 160 Supralabials 6/6 6/6 6/6 6/6 6/6 6/6 6/6 6/6 Infralabials 6/6 6/6 6/6 6/6 6/6 6/6 6/6 6/6 Supraoculars 1/1 1/1 1/1 1/1 1/1 1/1 Postoculars Fused 1/1 1/1 1/1 1/1 1/1 1/1 Fused 1 0.31 0.5 0.3 0.4 0.44 – 0.43 0.31 2 1.5 1.33 1.5 1.5 1.5 – 1.5 1.43 3 0.55 1 0.67 0.67 0.33 – 0.67 0.7 4 0.18 0.5 0.5 0.5 0.5 – 0.44 0.4 5 0.88 0.71 0.74 0.78 0.68 – 0.73 0.7 6 0.42 0.35 0.29 0.32 0.32 – 0.39 0.43 7 0.96 1.13 1.09 1.11 1.11 – 1.13 1.09 8 0.54 0.25 0.25 0.38 0.43 – 0.25 – 9 0.28 1, 0.67 1 0.33 0.67 – 0.67 – 10 2 3 2 0.5 2 – 1 0.5 11 0.53 0.51 0.54 0.53 0.53 – 0.51 0.53 12 0.58 0.7 0.62 0.74 0.63 – 0.78 0.78 13 1.52 1.38 1.38 1.47 1.44 – 1.52 1.52 14 – 1 1 1 1 – 1.67 – 15 0.57 1 1 0.86 1 – 0.89 0.89 16 0.85 1.17 1.25 1.38 1.29 – 1.17 1.17 17 1.98 1.71 1.6 2 1.75 – 1.71 1.71 18 2 2 2.38 2.25 2.71 – 2.17 2.5 19 2.6 2.4 2 2 1.75 – 2.4 2 20 5.2 4.8 4.75 4.5 4.75 – 5.2 5 21 0.63 0.63 0.8 0.8 0.8 – 0.63 0.88 22 1.71 1.38 1.4 1.33 1.6 – 1.33 1.11 23 2 1.5 1.5 1.25 1.67 – 2 1.6 24 1.57 1.6 2 1.5 1.67 – 1.8 1.43 25 1.89 2.5 2 2 2.2 – 1.9 1.89 26 1.13 1.67 1.27 1.17 1.1 – 1.27 1.21 27 0.07 0 0.09 0.17 0.1 – 0.07 0 28 0.119 0.154 0.145 0.118 0.115 0.131 0.124 0.127 29 0.4 0 0.17 0.17 0.33 – 0.1 0.2

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72 Table 12. Morphological and meristic characters for Geophis nephodrymus, part II. Honduras: Corts: Parque Nacional El Cusuco, 1545 m Characters JHT 1327 JHT 1334 JHT 1337 JHT 1342 JHT 1343 juv. JHT 1378 JHT 1382 JHT 1536 juv. JHT 1537 SVL 259 105 136 223 109 226 189 113 184 TL 35 20 12(I) 30 16 25(I) 37 12(I) 35 Ventrals 138 123 138 123 131 133 122 131 124 Subcaudals 22 31 16(I) 31 25 17(I) 29 24 29 Ventrals + subcaudals 160 154 154 154 156 150(I) 151 155 153 Supralabials 6/6 6/6 6/6 6/6 6/6 6/6 5/6 6/6 6/6 Infralabials 6/6 6/7 6/6 6/6 6/6 6/6 6/6 6/6 6/6 Supraoculars 1/1 1/1 1/1 1/1 1/1 0/1 1/1 1/1 1/1 Postoculars 1/1 1/1 1/1 1/1 1/1 0/1 1/1 1/1 1/1 1 0.38 – 0.23 0.38 0.27 0.28 0.43 – 0.35 2 1.38 – 1.4 1.83 1.4 1.38 2 – 1.43 3 0.67 – 0.63 0.54 0.71 0.73 0.5 – 0.58 4 0.33 – 0.5 0.36 0.43 0.54 0.3 – 0.25 5 0.83 – 0.67 0.78 0.79 0.68 0.74 – 0.73 6 0.44 – 0.42 0.42 0.37 0.42 0.4 – 0.57 7 0.96 – 1.05 1.08 1.16 1 1.16 – 1.14 8 0.42 – 0.4 0.42 0.5 – 0.09 – 0.3 9 0.6 – 0.5 0.4 0.25 – 1 – 0.67 10 3 – 1 2 1 – – – 1 11 0.49 – 0.46 0.49 0.53 0.45 0.53 – 0.51 12 0.7 – 0.84 0.65 0.79 0.65 0.76 – 0.81 13 1.5 – 1.44 1.4 1.38 1.52 1.52 – 1.75 14 1.25 – 1 1.33 2 – 0.5 – 1 15 0.9 – 1.2 0.89 1 1 1.14 – 1.14 16 1.5 – 0.9 1.08 1.13 1.15 1.18 – 1.2 17 1.71 – 1.67 1.71 1.14 1.86 1.57 – 1.67 18 2.42 – 2.1 2.25 2.38 2.38 2.45 – 2.6 19 2 – 2 2 1.6 – 1.83 – 1.67 20 4.83 – 4.2 4.5 3.8 – 4.5 – 4.33 21 0.6 – 0.71 0.75 0.71 – 0.67 – 0.86 22 1.27 – 1.67 1.57 1.8 2.14 – – 1.57 23 1.5 – 1.2 1.4 1.67 1.6 – – 1.2 24 1.33 – 1.2 1.8 1.17 1.43 – 1.6 25 2 – 2.33 1.81 2.33 1.78 2 – 1.33 26 1.18 – 1.27 1 1.08 1.14 1.05 – 27 0.47 – 0 0.3 0.23 0.29 0 – 0.5 28 0.119 0.160 – 0.119 0.128 – 0.164 – 0.160 29 0.4 – 0.5 0.6 0.33 0.25 0 – 0.5

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REVIEW OF THE Geophis OF EASTERN NUCLEAR CENTRAL AMERICA The genus Geophis is one of the most species-rich genera of colubrid snakes in the Western Hemisphere, being surpassed in the number of its constituent members only by Atractus (about 95 species; Williams et al., in preparation) and Tantilla (59 species; Canseco-Mrquez et al. 2002, in press; Greenbaum et al. 2004; Savage 2002; Sawaya and Sazima 2003; Stafford 2004; Wilson 1999; Wilson and Campbell 2000). There are 46 taxonomically recognized species in the genus Geophis distributed from southwestern Chihuahua and southern Tamaulipas, Mexico, through Central America to northwestern Colombia (Wilson and Townsend, unpublished data). These small snakes are fossorial or semi-fossorial in nature, and inhabit a wide range of habitats from lowland rainforest and dry forest to upland pine-oak forest and cloud forest (Downs 1967). Despite the wide geographic distribution and specific diversity within the genus, Geophis remains poorly understood and many species are known only from a single specimen or small series of specimens (Downs 1967; Myers 2003). The center for diversity of the genus Geophis clearly lies within the country of Mexico, where 32 species of Geophis occur, 26 of which are endemic to that country (Wilson and Townsend, unpublished data). Secondary evolutionary radiations within Geophis appear in the highlands of Nuclear Central America, as well as the highlands of Costa Rica and Panama. Eight of the nine species of Geophis found in Costa Rica and Panama have distributions restricted to those countries (Wilson and Townsend, unpublished data). There are six species known to occur in eastern Nuclear Central 73

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74 America (eNCA): G. damiani, G. dunni, G. fulvoguttatus, G. hoffmanni, G. nephodrymus, and G. rhodogaster (Figure 18). Downs (1967), in the most comprehensive work on the genus to date, partitioned the species of Geophis into seven species groups: the chalybeus group, championi group, dubius group, latifrontalis group, omiltemanus group, semidoliatus group, and sieboldi group. Only two of these groups are represented in eNCA, the dubius group and sieboldi group. Regarding the species groups with distributions that are extralimital to eNCA, the chalybeus, latifrontalis, and omiltemanus groups are endemic to Mexico; the semidoliatus group has members found in Mexico and Guatemala; and the championi group is restricted to Costa Rica and Panama. The characters used by Downs (1967) to define the species groups of Geophis do not represent shared derived characters and the groups were assembled on the basis of a non-uniform set of morphological characteristics, and, therefore, the assignment of members to these groups should not be necessarily viewed as a reflection of phylogenetic relationships among species, but rather as a matter of taxonomic convenience (Nieto-Montes de Oca 2003, Townsend and Wilson 2006). The Geophis dubius group is comprised of as many as 10 species: G. anocularis, G. carinosus, G. dubius, G. duellmani, G. fulvoguttatus, G. immaculatus, G. juarezi, G. nephodrymus, G. rhodogaster, and G. rostralis (Campbell et al. 1983, Downs 1967, Nieto-Montes de Oca 2003, Smith and Holland 1969, Townsend and Wilson 2006). The taxonomic status of G. rostralis is disputed, with some authors (Bogert and Porter 1966, Campbell et al. 1983, Townsend and Wilson 2006) choosing to recognize G. rostralis as a full species, whereas others (Downs 1967, Prez-Higareda and Smith 1988, Smith 1959) regard the name as a synonym of G. dubius. In this study, I continue to tentatively

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75 Figure 18. Distribution of Geophis in eastern Nuclear Central America. recognize G. rostralis as a valid species in the dubius group until its taxonomic situation is satisfactorily clarified. Species in the dubius group are distributed from 900 m elevation from Puebla and Veracruz, Mexico, to northwestern Honduras on the Caribbean versant and from Oaxaca, Mexico, to northwestern El Salvador and western Honduras on the Pacific versant (Downs 1967, Smith 1995). Members of the dubius group are characterized by having 17 dorsal scale rows with dorsal scales that may be smooth or keeled; ventrals numbering 114 in males and 118 in females; subcaudals numbering 31 in males and 24 in females; a tail that is 15.3% of total length in males and 11.3–22% of total length in females; a long snout that is bluntly pointed, a prominent rostral whose visible length is one-quarter or more its distance from the frontal, large anteriorly rounded internasals; short prefrontals (internasals and prefrontals usually fused in G. dubius); a frontal with an angulate anterior edge, short parietals; small triangular

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76 supraoculars that may be distinct or absent (as in the cases of G. anocularis, G. duellmani, and G. rhodogaster); small eyes; enlarged postnasals; short loreals; and no anterior temporals (Downs 1967, Nieto-Montes de Oca 2003, Townsend and Wilson 2006). The Geophis sieboldi group is the most speciose of the groups, and consists of 15 recognized species: G. betaniensis, G. brachycephalus, G. damiani, G. dunni, G. hoffmanni, G. laticollaris, G. nasalis, G. nigroalbus, G. petersii, G. pyburni, G. russatus, G. sallei, G. sieboldi, G. talamancae, and G. zeledoni (Downs 1967, Lips and Savage 1994, Wilson et al. 1998). Species in the sieboldi group are distributed from near sea level to 2164 m elevation, from Michoacan, Mexico, to Valle, Colombia, a range that encompasses much of the entire geographic distribution of genus. Members of the sieboldi group are characterized by having 15 or 17 dorsal scale rows, dorsal scales that may be smooth or keeled, a long snout that extends well beyond the lower jaw and appears rounded when viewed dorsally, a rostral that does not extend posteriorly between the internasals, internasals that are less than 61% the length of the prefrontal common suture, postnasals that are shorter than they are wide, prefrontals and loreals that are elongate, postoculars that form the posterior half of the dorsal margin of the orbit, no anterior temporals, a rounded mental, maxillae that extend forward to the middle of the second supralabial and have 8 subequal teeth, a toothless anterior tip of the maxilla, and a capitate hemipenis with a naked basal pocket on the asulcate side (Downs 1967, Lips and Savage 1994, Wilson et al. 1998). Systematic accounts: Below are species accounts for five of the Geophis found in eNCA, with the account for the sixth species, G. nephodrymus, having been presented in

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77 the previous chapter. The format followed below has been outlined in the chapter Methods and Materials. Geophis damiani Wilson, McCranie, and Williams Table 13 Geophis damiani Wilson, McCranie, and 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 (15’N, 87’W), Yoro, Honduras. Species group: sieboldi (Wilson et al. 1998). Distribution: Known only from the vicinity of the type locality at intermediate (1550 to 1750 m) elevations on the slopes of Cerro Texiguat, Honduras (Figure 18). Diagnosis: Geophis damiani is distinguished from the species of Geophis found in eNCA (except G. hoffmanni) by having 15 dorsal scale rows present throughout the length of the body (17 dorsal scale rows in G. dunni, G. fulvoguttatus, G. nephodrymus, and G. rhodogaster). Geophis damiani can be differentiated from G. hoffmanni by having a dark dorsal ground coloration with a red nuchal band and red partial bands and laterally offset red blotches (uniform dark dorsal coloration with a pale nuchal band in juveniles in G. hoffmanni), 136 ventral scales (114 in G. hoffmanni), 34 subcaudal scales (23 in G. hoffmanni), six supralabials (five or fewer in G. hoffmanni), and posterior chinshields in moderately to broadly separated (posterior chinshields in moderate contact in G. hoffmanni).

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78 Description: Dorsal scales in 15 rows, smooth throughout length of body; no apical pits apparent; ventrals 136, 136 in one adult male and 143 in one adult female; subcaudals 34, 41 in one adult male and 34 in one adult female; segmental count 177 in both specimens. In one male, snout-vent length 267 mm, tail length 60 mm, tail length/total length ratio 0.183; in one adult female, snout-vent length 256 mm, tail length 47 mm, tail length/total length ratio 0.155. Head slightly distinct from neck; snout bluntly rounded from above, rounded in profile, extending beyond the anterior end of the lower jaw; lower jaw reaches to the level of the posterior portion of the rostral; rostral wider than high, extending posteriorly in dorsal view between internasals, portion of rostral visible in dorsal view 0.27.37 times the length of its distance from the frontal, 1.0.17 times as long as internasal common suture, with posterior termination at anterior edge of nostril; internasals 1.22–1.33 times as wide as they are long, angular anteriorly, contacting the prenasal and postnasal laterally, their length 0.73.0 times as long as prefrontal common suture, their common suture 0.4.5 times as long as prefrontal common suture; prefrontals contacting postnasal, loreal, and orbit laterally, their length 0.67.81 times snout length, their common suture 0.46.52 times the frontal length; frontal 6-sided, 1.03.08 times as wide as is long, angulate anteriorly, in moderate to broad contact with supraoculars (length of frontal-supraocular contact 0.5.67 times supraocular length, 1.25.0 times prefrontal-supraocular contact length), supraocular moderate to large in size, somewhat triangular, in contact with postocular, roughly equal or slightly smaller than size of eye, 0.5.6 times as long as loreal; parietals 1.52.64 times as long as broad, their length

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79 0.5.52 times the length of the head, their common suture 0.78.87 times as long as frontal; single postocular, 1.0.2 times as high as long, smaller than supraocular; nasal divided; postnasal length 1.11.33 times that of prenasal length; combined length of prenasal and postnasal 1.06.2 times the length of the loreal; loreal 1.67.88 times as long as deep, contained 1.88.13 times on snout length; 1.5.0 times as long as eye horizontal diameter, dorsal margin straight; eye small, contained 3.2.75 times in snout length, its vertical diameter 1.0.25 times its distance from the lip. Supralabials 6/6; first supralabial contacting rostral, prenasal and postnasal; second supralabial contacting postnasal and loreal; third supralabial contacting loreal; third and fourth supralabials entering orbit; fourth supralabial contacting postocular; fifth supralabial the largest, contacting the postocular and parietal; lip exposure of third supralabial 1.25.57 times that of second supralabial; lip exposure of fifth supralabial 1.15.42 times that of fourth supralabial; anterior temporal absent; one posterior temporal, separating sixth supralabial from parietal; posterior temporals separated from each other by four or five nuchals; left posterior temporal divided on USNM 559598 to form a small posterior temporal and a sixth enlarged nuchal; mental 2.0.17 times as broad as long, rounded anteriorly, separated from chinshields by first infralabials; infralabials 6/6; first infralabials broadly in contact between mental and anterior chinshields; first, second, and third infralabial contacting the anterior chinshield; fourth infralabial contacting posterior chinshield; anterior chinshields 1.73–1.83 times as long as broad, 1.12.37 times as long as posterior chinshields; posterior chinshields not in contact, moderately to widely separated by first medial gular scale; three medial gular scales present.

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80 Hemipenis: Wilson et al. (1998:412) described the hemipenis of the holotype (USNM 498356) as “bilobed, distal portion of organ strongly capitate, calyculate, spinulate; sulcus spermaticus intermediate between centrifugal and centrolineal (Myers and Campbell 1981), bifurcation at point of capitation, each branch reaching apex; naked basal pocket on asulcate side; central portion of organ with large spines in oblique rows.” Dentition: The maxilla bears 10 subequal teeth, has a toothless anterior tip, and extends anteriorly to the middle portion of the second supralabial (Wilson et al. 1998). Coloration: Color notes in life of the holotype (from Wilson et al. 1998) as follows: “dorsal portions of body and tail Blackish Neutral Gray (82), with 24 Flame Scarlet (15) crossbands or laterally offset pairs of half crossbands on body; first nine bands complete, next twelve divided (or almost so) and the halves offset from one another along the longitudinal axis, and final three complete; seven similar markings (both crossbands and laterally offset markings) on tail that become increasingly faint towards its tip; head Blackish Neutral Gray (82); each ventral scale Glaucous (79) on anterior portion and white on posterior portion; underside of tail Glaucous (79); iris Jet Black (89).” The first dorsal crossband in the holotype forms a nuchal band. Coloration for USNM 559598 after about 19 months in preservation was: dorsal ground color brownish gray, becoming somewhat paler laterally; no nuchal band present as in holotype; dorsal surface with 25 well-defined pale reddish crossbands (no nuchal band) and two incomplete bands near the tail; dorsal surface of tail with four pairs of laterally offset spots; ventral surface cream colored with dark pigment from dorsal scales forming partial stripes on each ventral scale, becoming complete stripes along posterior edge of each scale towards posterior portion of the body; infralabials and chinshields with

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81 pale gray mottling; subcaudals dark gray with pale posterior edges on each scale, becoming darker posteriorly. Natural History Comments: Geophis damiani is known from disturbed cloud forest in the Lower Montane Wet Forest formation (Wilson et al. 1998, McCranie and Castaeda 2004b). The holotype was collected from under a mahogany plank on a steep slope, and the second specimen was active in the leaf litter in moderately disturbed forest at 1950 h, about one hour after dark. Wilson et al. (1998) remarked that several specimens of the plethodontid salamander Oedipina gephyra, another species endemic to the Cordillera Nombre de Dios, were also collected under mahogany planks in the vicinity of the type locality. Remarks: UF 142543 consists of an egg and an early stage snake embryo, attributable to Geophis damiani on the basis of its coloration when the egg was cut open in the field (J. R. McCranie, pers. comm.). This coloration has since faded in preservative, but remnants of the pattern are still discernable. References: Wilson et al. (1998), McCranie and Castaeda (2004b). Illustrations: Wilson et al. (1998: black and white photograph and illustration of the dorsal and lateral aspects of the head of the holotype); Khler (2001a: color photograph; 2003: color photograph); Campbell and Lamar (2004: color photograph of holotype). Specimens examined: 3; HONDURAS: Yoro: about 2.5 km NNE La Fortuna, Reserva de Vida Silvestre Cerro Texiguat, 1750 m (USNM 498356), 1680 m (USNM 559598), 1550 m (UF 142543; egg with partially developed snake).

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82 Table 13. Morphological and meristic characters for Geophis damiani, G. dunni, G. fulvoguttatus, and G. rhodogaster. Geophis damiani Geophis dunni Geophis fulvoguttatus Geophis rhodogaster Honduras: Texiguat Nicaragua: Matagalpa El Salvador: Montecristo El Salvador: Montecristo Characters USNM 498356 USNM 559598 MCZ 31870 KU 57996 KU 214781 KU 57998 KU 57999 SVL 267 256 309 281 353 249 321 TL 60 47 58 54 45 61 55 Ventrals 136 143 140 137 145 140 140 Subcaudals 41 34 36 34 35 41 34 Ventrals + subcaudals 177 177 176 171 180 181 176 Supralabials 6/6 6/6 6/6 6/6 6/6 6/6 6/6 Infralabials 6/6 6/6 8/8 6/6 7/7 6/6 6/6 Supraoculars 1/1 1/1 1/1 1/1 1/1 0/0 0/0 Postoculars 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1 0.27 0.37 0.28 0.27 0.23 0.33 0.33 2 1.22 1.33 1.4 1.4 1.08 1.55 1.39 3 0.73 1.0 0.5 0.71 0.93 1.1 1 4 0.4 0.5 0.35 0.57 0.64 0.5 0.46 5 0.81 0.67 0.72 0.79 0.77 0.61 0.63 6 0.52 0.46 0.57 0.44 0.41 0.26 0.38 7 1.03 1.08 0.94 1.06 1.0 1 1.26 8 0.56 0.6 0.48 0.71 0.47 – – 9 0.55 0.67 0.64 0.6 0.56 – – 10 1.25 2 2.33 2 2.5 – – 11 0.5 0.52 0.45 0.51 0.51 0.48 0.49 12 0.78 0.87 0.71 0.75 0.97 0.62 0.68 13 1.61 1.58 1.58 1.71 1.89 1.39 1.4 14 1 1 1.8 2.0 1.14 1.57 1.56 15 1.2 1.22 1.08 1 1.08 0.39 0.91 16 1.12 1.06 0.78 1.5 1.26 0.88 1 17 1.78 1.78 1.77 1.4 1.73 1.42 1.82 18 2.0 1.88 1.87 2.36 2.26 2.12 1.9 19 1.6 2.0 1.92 1.75 2.11 1.7 2 20 3.2 3.75 3.58 4.13 4.78 3.6 3.8 21 1.0 1.25 0.85 0.75 0.64 0.8 0.73 22 1.17 1.2 1.27 2 1.23 1.88 1.19 23 1.5 1.25 1.4 1.14 1.57 1.5 1.33 24 2.17 2.0 1.71 1.86 2.33 1.89 1.88 25 1.75 1.73 1.62 1.85 1.63 1.85 1.71 26 1.31 1.19 1.24 1.33 1.13 1.33 1 27 0 0 0 0.17 0.30 0.56 0.13 28 0.183 0.155 0.158 0.161 0.113 0.197 0.146 29 0.57 0.67 0.67 0.5 0.75 0.5 0.67

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83 Geophis dunni Schmidt Figure 19; Table 13 Geophis dunni Schmidt, 1932, Copeia 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. Species group: sieboldi (Downs 1967). Distribution: Known only from the type locality, from moderate elevations in the Matagalpan highlands (Figure 18). Diagnosis: Geophis dunni can be distinguished from the two other members of the sieboldi group that occur in eNCA (G. damiani and G. hoffmanni) by having 17 dorsal scale rows throughout the length of its body (15 dorsal scale rows in G. damiani and G. hoffmanni). From the three remaining species with 17 dorsal scale rows (G. fulvoguttatus, G. nephodrymus, and G. rhodogaster), G. dunni can be distinguished by having distinctly keeled dorsal scales on most of the body (smooth dorsal scales in G. fulvoguttatus, G. nephodrymus, and G. rhodogaster), eight infralabials (six in G. nephodrymus, six or seven in G. fulvoguttatus and G. rhodogaster), dorsal coloration pale yellow with dark brown saddles (dorsal coloration dark brown or blackish gray with red or yellow middorsal blotches in G. fulvoguttatus; gray to blackish gray with no pattern or with red or pale gray dorsolateral blotches, bands, or partial bands in G. nephodrymus; dark reddish brown or brown in G. rhodogaster).

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84 Description: Dorsal scales in 17 rows, keeled throughout length of body, except for neck; apical pits present; ventrals 140 in single adult female; subcaudals 36; segmental count 176. In one female, snout-vent length 309 mm, tail length 58 mm, and tail length/total length ratio 0.158. Head indistinct from neck; snout rounded from above, bluntly rounded in profile, extending beyond the anterior end of the lower jaw; lower jaw reaches to the level of the posterior one-third of the rostral; rostral wider than high, extending posteriorly in dorsal view between internasals, portion of rostral visible in dorsal view 0.28 times the length of its distance from the frontal, with posterior termination at midpoint of nostril; internasals 1.4 times as wide as they are long, angular anteriorly, contacting the prenasal and postnasal laterally, their length 0.5 times as long as prefrontal common suture, their common suture 0.35 times as long as prefrontal common suture; prefrontals contacting postnasal, loreal, and orbit laterally, their length 0.72 times snout length, their common suture 0.57 times the frontal length; frontal 6-sided, 0.94 times as wide as is long, angulate anteriorly, in broad contact with supraoculars (length of frontal-supraocular contact 0.64 times supraocular length, 2.33 times prefrontal-supraocular contact length), supraocular large in size, appearing triangular, in contact with postocular, slightly smaller than size of eye, 0.48 times as long as loreal; parietals 1.58 times as long as broad, their length 0.45 times the length of the head, their common suture 0.71 times as long as frontal; single postocular, 1.8 times as high as long, smaller than supraocular; nasal divided; postnasal length 1.08 times that of prenasal length; combined length of prenasal and postnasal 0.78 times the length of the loreal; loreal 1.77 times as long as deep,

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85 contained 1.87 times on snout length, 1.92 times as long as eye horizontal diameter, dorsal margin straight; eye small, contained 3.58 times in snout length, its vertical diameter 0.85 times its distance from the lip. Supralabials 6/6; first supralabial contacting rostral, prenasal and postnasal; second supralabial contacting postnasal and loreal; third supralabial contacting loreal; third and fourth supralabials entering orbit; fourth supralabial contacting postocular; fifth supralabial the largest, contacting the postocular and parietal; lip exposure of third supralabial 1.4 times that of second supralabial; lip exposure of fifth supralabial 1.27 times that of fourth supralabial; anterior temporal absent; one posterior temporal, separating sixth supralabial from parietal; posterior temporals separated from each other by five nuchals; mental 1.71 times as broad as long, rounded anteriorly, separated from chinshields by first infralabials; infralabials 8/8; first infralabials broadly in contact between mental and anterior chinshields; first, second, and third infralabial contacting the anterior chinshield; fourth infralabial contacting both anterior and posterior chinshield; infralabials 2-4 small, squarish; fifth infralabial largest; anterior chinshields 1.62 times as long as broad, 1.24 times as long as posterior chinshields; posterior chinshields not in contact, separated by first medial gular, almost meeting at a point near their contact with anterior chinshields; three medial gular scales present. Hemipenis: Unknown. Dentition: Downs (1967:154) described the maxilla as extending “anteriorly to the level of supralabial 2; anterior extension about equal to that of palatine; maxilla dorsoventrally compressed, bears 12 curved teeth, subequal in length; anterior tip of

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86 maxilla pointed, toothless; posterior end of maxilla curves ventrally, tapers to blunt point; anterior end of ectopterygoid single, not expanded; postorbital bone present.” Coloration: Coloration of holotype after 96 years in preservation: dorsal surface of head brownish gray, coloration extending posteriorly to the anterior one-fourth to one-third of the parietals; remainder of parietals markedly paler, mottled brown and yellow; first, second, and third supralabials brownish gray, becoming somewhat paler towards ventral edge; fourth supralabial brownish gray on anterior one-third, mottled brown and yellow on posterior portion; ventral surface of head immaculate pale yellow, except mental, first infralabials, and anterior edge of anterior chinshields and second through fifth infralabials brownish gray; dorsal surface of body with pale yellow ground color; a wide brown band beginning 2 dorsal scales posterior to the parietals, 7 dorsal scales long, and extending laterally to the upper half of the second dorsal scale row; dorsum with 24 brown saddles and irregular blotches that extend laterally to the second or sometimes the first dorsal scale row; yellow scales between brown saddles and blotches with varying degrees of brown mottling, usually consisting to a brown spot at the apex of each scale; ventral surface immaculate pale yellow; dorsal surface of tail with three brown saddles and two elongate brown blotches; subcaudals immaculate anteriorly, with brown mottling on edges of scales increasing posteriorly, posterior one-fourth of tail mottled brown and yellow. Natural History Comments: Literally the only thing known about the natural history of G. dunni is that on one occasion, sometime during or before 1909, a single individual was eaten by a Micrurus nigrocinctus.

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87 Remarks: This species is one of the most enigmatic and poorly known snakes of the genus Geophis, or any genus for that matter. Known only from the type specimen, which was discovered in the stomach of a Micrurus nigrocinctus collected during or before 1909 in a locality recorded only as “Matagalpa, Nicaragua” (MCZ Catalogue, original entry, checked by JHT on 20 December 2005), no additional material representing this distinctive snake has been secured since its description. The name provided as the type locality could refer to both a city and a department in northwestern Nicaragua, leaving basic but informative data (such as the elevation and habitat) a mystery. Downs (1967: 153) reported that the type specimen came from 705 m elevation, but there is no known basis for using that elevation. It is unlikely that “Matagalpa” refers to the immediate vicinity of the city of Matagalpa, which is surrounded by denuded pine-oak forest. Since pine-oak forest is a widespread habitat in eNCA, the lack of any additional specimens of G. dunni leaves me skeptical that this species occurs in pine-oak forest at or near the elevation provided by Downs (1967). References: Schmidt (1932), Downs (1967). Illustrations: Downs (1967; black and white illustration of the dorsal and lateral aspects of the head of the holotype). Specimens examined: 1; NICARAGUA: Matagalpa (MCZ 31870).

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88 Figure 19. Holotype of Geophis dunni (MCZ 31870), an adult female from Matagalpa, Nicaragua. Photograph by J. H. Townsend. Figure 20. Preserved specimens of Geophis fulvoguttatus; adult male (KU 57996) from Hacienda Montecristo, 2200 m, Cordillera de Metapn, Santa Ana, El Salvador, and adult female (KU 214781) from El Portillo de Ocotepeque, 1900 m, Ocotepeque, Honduras.

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89 Geophis fulvoguttatus Mertens Figure 20; Table 13, 14 Geophis fulvoguttatus Mertens, 1952b: 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 Metapn, Santa Ana, El Salvador. Species group: dubius (Downs 1967). Distribution: Known from intermediate (1680 to 2200 m) elevations in Santa Ana, El Salvador, and Ocotepeque and Copn, Honduras (Figure 18). Diagnosis: Geophis fulvoguttatus can be distinguished from G. damiani and G. hoffmanni by having 17 dorsal scale rows (15 dorsal scale rows in the aforementioned species). Of the remaining species with 17 dorsal scale rows (G. dunni, G. nephodrymus, and G. rhodogaster), G. fulvoguttatus can be distinguished from G. dunni by having smooth dorsal scales, six or seven infralabials, and a dark brown or gray dorsal coloration with irregular red or yellow dorsal blotches (dorsal scales strongly keeled, eight infralabials, and pale yellow dorsal color with dark brown saddles in G. dunni); from G. nephodrymus by having 135 ventral scales and segmental counts ranging from 171–181 (120 ventral scales and segmental counts ranging from 149 in G. nephodrymus); and from G. rhodogaster by having a supraocular on at least one side and the frontal not entering the orbit (no supraoculars, frontal entering the orbit on both sides in G. rhodogaster). Description: Dorsal scales in 17 rows, smooth throughout length of body; no apical pits; ventrals 135 (137 in one undamaged male and 145 in two adult

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90 females; 135 reported by Khler et al., 2006); subcaudals 24 (34–36 in two males and 24 in two adult females); segmental count 171 (171 in undamaged male and 180 in two adult females). Snout-vent length 110 mm in two males, 297 mm in two females; tail length 20 mm in two males, 38 mm in two females; tail length/total length ratio 0.161.181 in two adult males, 0.113 in two adult females. Head not distinct from neck; snout rounded from above, rounded in profile, extending beyond the anterior end of the lower jaw; lower jaw reaches to the level of the posterior portion of the rostral; rostral wider than high, extending posteriorly in dorsal view between internasals, portion of rostral visible in dorsal view 0.23.27 times the length of its distance from the frontal, 0.67.75 times as long as internasal common suture, with posterior termination in line with middle of prenasal; internasals 1.08.4 times as wide as they are long, angulate to somewhat rounded anteriorly, contacting the prenasal and postnasal laterally, their length 0.71.93 times as long as prefrontal common suture, their common suture 0.57.64 times as long as prefrontal common suture; prefrontals contacting postnasal, loreal, and orbit laterally, their length 0.77.79 times snout length, their common suture 0.41.44 times the frontal length; frontal 6-sided, 1.0.06 times as wide as is long, acutely angulate anteriorly, in moderate to broad contact with supraoculars (length of frontal-supraocular contact 0.56–0.60 times supraocular length, 2.0.5 times prefrontal-supraocular contact length), supraocular moderate in size, triangular, in contact with postocular, roughly smaller than eye, 0.47–0.71 times as long as loreal; parietals 1.69.89 times as long as broad, their length 0.50–0.51 times the length of the head, their common suture 0.75.97 times as long as frontal

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91 (KU 183881 lacks supraocular on left side, frontal entering orbit; supraocular on right side reduced in size and shifted posteriorly around orbit); single postocular, 1.33.0 times as high as long, smaller than supraocular; nasal divided; postnasal length 1.08 times that of prenasal length; combined length of prenasal and postnasal 1.15.5 times the length of the loreal; loreal 1.4.7 times as long as wide, contained 2.15.36 times on snout length; 1.75.11 times as long as eye horizontal diameter, dorsal margin straight; eye small, contained 4.13.78 times in snout length, its vertical diameter 0.64–0.75 times its distance from the lip. Supralabials 6/6; first supralabial contacting rostral, prenasal and postnasal; second supralabial contacting postnasal and loreal; third supralabial contacting loreal; third and fourth supralabials entering orbit; fourth supralabial contacting postocular; fifth supralabial the largest, contacting the postocular, and parietal; lip exposure of third supralabial 1.13.57 times that of second supralabial; lip exposure of fifth supralabial 1.23.0 times that of fourth supralabial; anterior temporal absent; one posterior temporal, separating sixth supralabial from parietal; posterior temporals separated from each other by 5 nuchals; mental 1.86.33 times as broad as long, angulate anteriorly, separated from chinshields by first infralabials; infralabials 6/6 or 7/7; first infralabials broadly in contact between mental and anterior chinshields; first, second, third and, in one case, the fourth infralabial contacting the anterior chinshield; fourth infralabial contacting posterior chinshield; anterior chinshields 1.6.85 times as long as broad, 1–1.58 times as long as posterior chinshields; posterior chinshields narrowly to broadly in contact; three to four medial gular scales present.

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92 Hemipenis: Downs (1967:89) described the hemipenis of KU 57996 as follows: bearing a single large spine and numerous small spines on the basal portion; a naked basal pocket present on the asulcus side, flanked by ridges; central part of hemipenis bearing 35 long, slender spines arranged in oblique rows; distal portion of hemipenis capitate, with the capitation obscured by gradation between spines on central portion of hemipenis and spines on proximal edge of spinulate capitulum; calyces discernable at apex only; apex bilobed; sulcus spermaticus bifurcate, with each branch reaching the apex of one lobe. Dentition: Downs (1967:89) described the maxilla as extending “anteriorly to suture between first and second supralabials; anterior extension greater than that of palatine; maxilla curved in lateral view, slenderest anteriorly; 10 maxillary teeth, increasing in length posteriorly; first tooth at anterior tip of maxilla; posterior end of maxilla laterally compressed into moderate flange; anterior end of ectopterygoid bifurcate, one branch short and blunt, second branch long, compressed, blade-like; no postorbital bone.” Coloration: McCranie and Wilson (1991:113) described the color in life of an adult female (KU 214781) as follows: dorsal surface very dark gray with 26 red-orange middorsal spots more prominent posteriorly and with 2 dorsal and 4 alternating lateral spots on the tail; dorsal surface of head gray, paling laterally onto supralabials; ventral surface white with dark gray on the lateral edges of the ventral scales; subcaudal surface white infused with brownish gray. In the examined specimens, the fourth, fifth, and sixth supralabials are heavily pigmented with ventral coloration, with dorsal pigment only visible near the dorsal edges

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93 Table 14. Relevant measurements and scale counts for Geophis fulvoguttatus and G. rhodogaster specimens with unmeasured head scales. Geophis fulvoguttatus Geophis rhodogaster El Salvador: Montecristo Honduras: Copn El Salvador: Montecristo Characters SMF 43248 KU 183881 KU 214782 (tail only) SMF 77413 KU 57997 SVL 110* 297 – 248 321 TL 20 38 43 68 57+ Ventrals 145-150* 157 – 138 142 Subcaudals 36 24 36 43 34+ Ventrals + subcaudals 171-176* 181 – 181 176+ Supralabials 6/6 6/6 – 6/6 6/6 Infralabials 6/6 6/6 – 7/7 6/7 Supraoculars 1/1 0/1 – 0/0 0/0 Postoculars 1/1 1/1 – 1/1 1/1 Tail length/ total length (29) 0.181* 0.113 – 0.215 0.151 of the scales. The 28 middorsal spots or blotches only extend to the third dorsal scale row on three occasions, only contact the second dorsal scale row in two cases, and never reach to the first dorsal scale row. The blotches appear as either irregular amorphous middorsal markings, or as single or laterally offset pairs of circular spots. Natural History Comments: Specimens of this species have been found under logs in cloud forest and pine-oak forest (Downs 1967, McCranie and Wilson 1991). Remarks: The holotype (SMF 43248) is a subadult male with badly damaged posterior half (Mertens, 1952a, b), and was not examined for this study. References: Mertens (1952a, b), Downs (1967), McCranie and Wilson (1991), Khler et al. (2006). Illustrations: Mertens (1952b; black and white photograph of the holotype), Downs (1967; black and white illustration of the dorsal and lateral aspects of the head of the holotype); Khler et al. (2006; color photograph of preserved specimen).

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94 Specimens examined: 4; EL SALVADOR: Santa Ana: Hacienda Montecristo, Cordillera de Metapn, 2200 m (KU 57996); Metapn, 2200 m (KU 183881). HONDURAS: Copn: above Quebrada Grande, 1680 m (KU 214782); Ocotepeque: El Portillo de Ocotepeque, 1900 m (KU 214781). Referred specimens: 2; EL SALVADOR: Santa Ana: Hacienda Montecristo, Cordillera de Metapn, 2200 m (SMF 43248 [holotype], Mertens 1952a; MUHNES 489, Khler et al. 2006). Geophis hoffmanni (Peters) Tables 15 Colobognathus Hoffmanni Peters, 1859: 276. Geophis hoffmanni: Boulenger, 1894: 319. Geophis bartholomewi Brattstrom and Howell, 1954: 120. Syntypes: Eight: Zoologisches Museum, Berlin (ZMB) 1868 (five specimens), 4003, 4106 (two specimens), and The Natural History Museum, London (BMNH) 1946.1.6.54. Downs (1967: 155) designated ZMB 1870, an adult female, the lectotype. Type-locality: “Costa Rica.” One syntype (BMNH 1946.1.6.54) is from “Porto Caballo,” Costa Rica (Downs, 1967:155). Species group: sieboldi (Downs 1967). Distribution: Known from low and moderate (18 to 670 m) elevations in central and eastern Honduras south through eastern Nicaragua, Costa Rica, and central Panama (Figure 18). Diagnosis: Geophis hoffmanni is distinguished from all other Geophis found in eNCA (except G. damiani) by having 15 dorsal scale rows present throughout the length

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95 Table 15. Relevant measurements and scale counts for Geophis hoffmanni from localities in eastern Nuclear Central America other than the Rus Rus/Warunta region. Honduras Nicaragua Olancho: Coln: Quebrada Machn Ro Tinto El Paraiso: Arenales Matagalpa Jinotega: Bosaws Characters USNM 561035 USNM 561036 USNM 561037 USNM 337522 LACM 20483 CAS 91202 UMMZ 117653 juv SMF 78594 SMF 78595 SVL 158 100 160 161 148 133 96 137 160 TL 26 14 27 29 25+ 13+ 15+ 19+ 23 Ventrals 124 122 123 121 122 124 126 120 122 Subcaudals 28 27 28 29 25(I) 14(I) 24(I) 22(I) 25 Segments 152 149 151 150 147(I) 138(I) 150(I) 142(I) 147 Supralabials 5/5 5/5 5/5 5/5 5/5 5/5 5/5 5/5 5/5 Infralabials 6/6 6/6 6/6 6/6 6/6 6/6 6/6 6/6 6/6 Supraoculars 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 Postoculars 1/1 1/1 1/1 1/1 1/1 1/1 0/0 1/1 1/1 of the body (17 dorsal scale rows in G. dunni, G. fulvoguttatus, G. nephodrymus, and G. rhodogaster). Geophis hoffmanni can be differentiated from G. damiani by having a uniform dark dorsal coloration with a pale nuchal band in juveniles (dark dorsal ground coloration with a red nuchal band and red partial bands and laterally offset red blotches in G. damiani), 114 ventral scales (136 in G. damiani), 23 subcaudal scales (34 in G. damiani), five or fewer supralabials (six in G. damiani), and posterior chinshields in moderate contact (posterior chinshields separated by first medial gualr scale in G. damiani). Description: Dorsal scales in 15 rows, smooth throughout most of length of body and keeled or faintly keeled scales in the region above the vent; paired apical pits apparent near the vent in some individuals; ventrals 114 (114 in eight males, 116 in 18 females, and 120 in four juveniles); subcaudals 23 (27 in eight males, 23 in 14 females with complete tails, and 24 in three juveniles with complete tails); segmental count 144 (145 in eight males, 144 in 14

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96 females with complete tails, and 144 in three juveniles with complete tails). Snout-vent length ranges from 98 mm in eight males, 88 mm in 14 females (with complete tails), 73 mm in three unsexed snakes (with complete tails); tail length from 19 mm in eight males, 15 mm in 14 females, 10 mm in three unsexed snakes, tail length/total length ratio 0.152.174 in eight males, 0.115.146 in 14 females, 0.118.141 in three unsexed snakes. Head somewhat distinct from neck; snout somewhat pointed from above, somewhat pointed in profile, extending beyond the anterior end of the lower jaw; lower jaw reaches to the level of the anterior edge of the first infralabial; rostral higher than wide, not extending appreciably between internasals in dorsal view, portion of rostral visible in dorsal view 0.23.44 times the length of its distance from the frontal, with posterior termination at anterior edge of nostril; internasals 1.17.0 times as wide as they are long, rounded anteriorly, contacting the prenasal and postnasal laterally, their length 0.4.67 times as long as prefrontal common suture, their common suture 0.22–0.4 times as long as prefrontal common suture; prefrontals contacting postnasal, loreal, and orbit laterally, their length 0.62.84 times snout length, their common suture 0.4–0.57 times the frontal length; frontal 6-sided, 0.91.11 times as wide as is long, angulate anteriorly, in broad contact with supraoculars (length of frontal-supraocular contact 0.5–1.0 times supraocular length, 1.0.0 times prefrontal-supraocular contact length); supraocular moderate in size, usually appearing triangular, usually in contact with postocular (supraocular and postocular fused on both sides in USNM 561046), usually smaller than size of eye, 0.31.55 times as long as loreal; parietals 1.24.44 times as

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29 0.5 0.67 0.5 0.67 0.5 – 0.67 0.67 0.67 97Table 16. Variation in ratios for Geophis hoffmanni from localities in eastern Nuclear Central America other than the Rus Rus/Warunta region. Honduras Nicaragua Coln: Quebrada Machn Olancho: Ro Tinto El Paraiso: Arenales Matagalpa Jinotega: Bosaws Ratio USNM 561035 USNM 561036 USNM 561037 USNM 337522 LACM 20483 CAS 91202 UMMZ 117653 juv SMF 78594 SMF 78595 1 0.29 0.33 0.36 0.33 0.4 – 0.38 0.21 0.23 2 1.4 1.5 1.6 1.6 1.4 – 1.25 1.4 2 3 0.5 0.5 0.45 4.2 0.55 – 0.57 0.71 0.4 4 0.3 0.25 0.27 0.25 0.33 – 0.29 0.29 0.2 5 0.67 0.75 0.75 0.71 0.62 – 0.68 0.77 0.76 6 0.5 0.4 0.48 0.57 0.45 – 0.35 0.37 0.53 7 1.1 1 0.96 1 1 – 0.95 1 1.11 8 0.43 0.4 0.39 0.43 0.36 – 0.33 0.42 0.45 9 0.67 0.5 1 0.67 0.5 – 0.5 0.4 0.6 10 4 2 5 4 2 – 2 2 3 11 0.46 0.47 0.47 0.48 0.46 – 0.41 0.51 0.47 12 0.75 0.5 0.48 0.67 0.65 – 0.5 0.79 0.79 13 1.33 1.35 1.38 1.38 1.42 – 1.29 1.71 1.47 14 1 1 0.67 1 1.33 – – 2 1.33 15 1.33 1 1 1.17 1.2 – 1 1 1 16 0.79 0.9 0.85 0.79 0.91 – 0.67 0.67 0.91 17 2.33 2.5 2.17 2.33 1.83 – 2.4 2.4 1.83 18 1.71 2 1.85 1.71 2.36 – 1.58 1.83 2.27 19 2.8 2.5 2.17 2.33 2.2 – 3 2.4 2.2 20 4.8 5 4 4 5.2 – 4.75 4.4 5 21 0.57 0.67 0.86 0.75 0.83 – 0.6 0.67 0.71 22 2.13 2 2.13 2 2.5 – 2.14 1.88 1.7 23 1.25 1.5 1.2 1.67 1.5 – 1.66 1.5 1.25 24 1.75 2 2.67 1.75 1.25 – 2 2 2 25 1.67 1.6 1.83 1.83 1.8 – 1.6 2 1.67 26 1 1.43 0.83 1 0.75 – 1 1 0.91 27 0.2 0.43 0.33 0.27 0.42 – 0.38 0.4 0.36 28 0.141 0.123 0.144 0.153 0.145 0.089* – – 0.126

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98 Table 17. Relevant measurements and scale counts for Geophis hoffmanni from localities in eastern Nuclear Central America within Parque Nacional Warunta and Reserva Biolgica Rus Rus, Honduras. Cabaceras de Ro Rus Rus Bodega del Ro Tapalws Cerro Wahatingni Norte Kilpa Tingni Hiltara Characters USNM 561039 juv. USNM 561038 USNM 561040 USNM 562885 juv. USNM 562879 USNM 562880 USNM 562881 USNM 562882 USNM 562883 USNM 562884 SVL 73 – 134 110 99 154 146 151 136 159 TL 12 – 24 16 16 24 22 18(I) 19 26 Ventrals 121 119 120 131 120 125 126 127 128 126 Subcaudals 30 25 27 27 28 26 25 20(I) 24 27 Ventrals + subcaudals 151 144 147 158 148 151 151 147(I) 152 153 Supralabials 5/5 5/5 5/5 5/5 5/5 4/5 5/5 5/5 5/5 5/5 Infralabials 6/6 6/6 6/6 6/6 6/6 6/7 6/7 6/5 6/6 6/6 Supraoculars 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 Postoculars 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 Tail/total length 0.141 – 0.152 0.127 0.139 0.135 0.131 – 0.123 0.141 long as broad, their length 0.43.49 times the length of the head, their common suture 0.48.79 times as long as frontal; single postocular (in UMMZ 117653, left postocular fused to parietal with parietal contacting orbit; right postocular fused to the fourth supralabial, which interrupts parietal-orbit contact on the right side), 0.67.33 times as high as long, smaller than (occasionally roughly equal in size to) supraocular; nasal divided; postnasal length 0.71.33 times that of prenasal length; combined length of prenasal and postnasal 0.69.0 times the length of the loreal; loreal 1.57.75 times as long as deep, contained 1.71.5 times on snout length, 1.57.0 times as long as eye horizontal diameter, dorsal margin straight; eye small, contained 3.17.2 times in snout length, its vertical diameter 0.57.2 times its distance from the lip. Supralabials 5/5 (4/5 in USNM 562880); first supralabial contacting rostral, prenasal, and postnasal; second supralabial contacting postnasal and loreal; third

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99Table 18. Variation in ratios for Geophis hoffmanni from localities in eastern Nuclear Central America within Parque Nacional Warunta and Reserva Biolgica Rus Rus, Honduras. Bodega del Ro Tapalws Kilpa Tingni Kiamp Hiltara Kiamp Ratio USNM 561038 USNM 561040 USNM 562879 USNM 562880 USNM 562881 USNM 562882 USNM 562883 USNM 562884 1 0.31 0.29 0.27 0.29 0.25 0.23 0.23 0.44 2 1.75 1.4 1.25 1.4 1.75 1.75 1.5 1.2 3 0.4 0.42 0.5 0.5 0.44 0.44 0.5 0.5 4 0.3 0.25 0.38 0.3 0.22 0.33 0.25 0.3 5 0.78 0.79 0.84 0.79 0.71 0.73 0.77 0.67 6 0.5 0.57 0.44 0.53 0.43 0.45 0.4 0.5 7 1.1 1 1 1 1 1.05 1.05 1 8 0.42 0.31 0.36 0.39 0.36 0.42 0.45 0.36 9 1 0.75 0.5 0.6 0.6 0.6 0.8 0.8 10 5 3 2 1 1.5 3 2 4 11 0.46 0.46 0.49 0.47 0.46 0.47 0.44 0.44 12 0.75 0.67 0.72 0.74 0.67 0.6 0.6 0.6 13 1.24 1.35 1.26 1.35 1.24 1.3 1.33 1.39 14 1 1 1 1.25 1.33 1 1.33 1.33 15 1 1.17 0.71 1 1 1 1.2 1.17 16 0.92 0.69 0.73 0.69 0.71 0.92 0.91 0.71 17 2 2.17 2.75 1.86 2 2 1.83 2.33 18 1.92 1.85 1.73 1.85 1.71 1.83 2 1.71 19 2.4 2.17 1.83 2.17 2.33 2 2.2 2.8 20 4.6 4 3.17 4 4 3.67 4.4 4.8 21 0.83 0.83 1.2 0.75 0.86 0.86 0.83 0.73 22 1.78 1.78 1.75 1.73 1.6 1.56 2.14 1.89 23 1.4 1.75 1.25 – 1.4 2 1.33 1.75 24 1.5 1.75 2 2 2 1.75 1.75 1.75 25 2 1.67 2 1.67 1.5 1.67 2.25 1.6 26 1.11 1 1.14 1.25 1 1.11 1.79 1 27 0.33 0.4 0.29 0.13 0.3 0.22 0.14 0.38 28 – 0.152 0.139 0.135 0.131 – 0.123 0.141 29 0.4 0.33 0.33 0.33 0.67 0.25 0.67 0.5

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100 Table 19. Relevant measurements and scale counts for Geophis hoffmanni from localities in eastern Nuclear Central America within Parque Nacional Warunta, Honduras. Urus Tingni Kiamp Warunta Tingni Kiamp Characters USNM 561041 USNM 561930 USNM 561932 USNM 561042 USNM 561043 USNM 561044 juv. USNM 561045 USNM 561046 USNM 561047 USNM 561048 USNM 561931 SVL 163 119 132 161 98 75 157 136 154 174 88 TL 24 25 27 21 19 10 31 27 29 24 15 Ventrals 123 116 118 126 114 120 117 126 118 130 116 Subcaudals 25 31 32 25 29 24 29 31 31 23 28 Ventrals + subcaudals 148 147 150 151 145 144 146 157 149 153 144 Supralabials 5/5 5/5 5/5 5/5 5/5 5/5 5/5 5/5 5/5 5/5 5/5 Infralabials 6/6 6/6 6/6 6/6 5/6 6/6 6/6 6/6 6/6 6/6 6/6 Supraoculars 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 Postoculars 1/1 1/1 1/1 1/1 1/1 1/1 1/1 Fused on both sides 1/1 1/1 1/1 Tail/total length 0.128 0.174 0.170 0.115 0.162 0.118 0.165 0.166 0.158 0.121 0.146 supralabial contacting loreal (loreal does not reach orbit and the third supralabials contact prefrontals in SMF 78595 [both sides] and USNM 561047 [left side]); third and fourth supralabials entering orbit; fourth supralabial contacting postocular; fifth supralabial much larger than other supralabials, contacting the postocular and parietal; lip exposure of third supralabial 1.2.0 times that of second supralabial; lip exposure of fifth supralabial 1.45.83 times that of fourth supralabial; anterior temporal absent; one posterior temporal, posterior temporals usually separated from each other by five nuchals; mental 1.5.67 times as broad as long, rounded anteriorly, separated from chinshields by first infralabials; infralabials typically 6/6, but can range from 5; first infralabials broadly in contact between mental and anterior chinshields; first, second, and third infralabial contacting the anterior chinshield; third and fourth infralabial contacting posterior chinshield; anterior chinshields 1.43.25 times as long as broad, 0.83.79

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101Table 20. Variation in ratios for Geophis hoffmanni from localities in eastern Nuclear Central America within Parque Nacional Warunta, Honduras Urus Tingni Kiamp Warunta Tingni Kiamp Ratio USNM 561041 USNM 561930 USNM 561932 USNM 561042 USNM 561045 USNM 561046 USNM 561047 USNM 561048 1 0.36 0.33 0.27 0.29 0.31 0.33 0.33 0.29 2 1.4 1.4 1.17 1.4 1.4 1.4 1.33 1.4 3 0.5 0.63 0.67 0.5 0.42 0.45 0.55 0.5 4 0.4 0.38 0.33 0.4 0.33 0.36 0.36 0.4 5 0.69 0.73 0.67 0.75 0.74 0.73 0.69 0.75 6 0.45 0.4 0.43 0.48 0.57 0.5 0.5 0.53 7 0.91 0.95 0.91 0.95 1 0.95 1 1.11 8 0.46 0.55 0.46 0.31 0.4 – 0.55 0.46 9 0.83 0.5 0.5 1 0.5 – 0.67 0.67 10 2.5 3 1.5 4 1 1.5 2 2 11 0.43 0.45 0.46 0.46 0.44 0.47 0.45 0.46 12 0.59 0.6 0.71 0.52 0.71 0.64 0.68 0.79 13 1.37 1.32 1.35 1.37 1.35 1.42 1.27 1.44 14 1.33 0.75 1 1.33 0.75 – 1.33 0.67 15 1 0.86 1 1 1 1 1.17 0.83 16 0.69 0.82 0.77 0.75 0.73 0.85 1 0.69 17 2.17 2.2 2.17 2 2.14 1.86 1.57 2.17 18 2 2 1.85 2 1.8 2 2.36 1.85 19 2.17 2.2 2.17 2 2.14 1.86 1.57 2.17 20 4.33 4.4 4 4 4.5 4.33 3.71 4 21 0.75 0.71 0.86 0.75 0.88 1.17 0.88 0.86 22 2 2.83 2.43 1.56 1.45 2 2.13 2 23 1.4 1.25 1.25 1.4 1.4 1.2 1.5 1.5 24 1.75 1.75 2 2 1.8 1.75 1.8 1.75 25 1.67 2 1.43 1.43 1.71 2.2 1.57 1.5 26 0.83 1 1 1 1.09 1 1.1 1 27 0.5 0.3 0.4 0.4 0.36 0.36 0.2 0.33 28 0.128 0.174 0.170 0.115 0.165 0.166 0.158 0.121 29 0.5 0.5 0.75 0.33 0.75 0.67 0.5 0.75

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102 times as long as posterior chinshields; posterior chinshields in moderate contact (posterior chinshield common suture 13% of posterior chinshield length); two to four medial gular scales present. Hemipenis: Based on examination of three adults males (USNM 561045), the hemipenis of G. hoffmanni can be characterized as elongate and unicapitate, with a slightly bilobed apex; distal portion with somewhat papilate calyces; a bifurcate, centrifugal sulcus spermaticus, with both branches extending to the apex; central section of hemipenis with numerous spines, and a weakly-developed basal naked pocket on the sulcal side; proximal section of organ bearing numerous spicules. Dentition: Downs (1967: 157) described the maxilla of G. hoffmanni as extending “anteriorly to the suture between supralabials 2 and 3; anterior extension about equal to that of palatine; maxilla dorsoventrally compressed, bears 8 subequal teeth; anterior tip of maxilla pointed, toothless; posterior end of maxilla curves ventrally, tapers to blunt point; anterior end of ectopterygoid single, not expanded; postorbital bone present.” Coloration: Dorsal surface of head pale to dark gray, often markedly paler from posterior edge of parietals forward; internasals sometimes with pale gray patches; a white to gray nuchal collar present in smaller individuals, becoming less distinct then disappearing with age, extending posteriorly two or three dorsal scales from posterior edge of parietals, can cover half to nearly all of fifth supralabial; dorsal surface of body and tail bluish gray to blackish gray, darker in smaller specimens, first dorsal scale row can be white or cream on lower half or less of each scale; rostral cream to pale gray; infralabials and chinshields can be white, speckled with gray, or pale gray; ventral scales

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103 white or cream, immaculate or with some dark gray mottling on the lateral edges of ventrals; subcaudal surface dark gray, with pale anterior edges of subcaudal scales. Natural History Comments: Geophis hoffmanni is a semifossorial, diurnal inhabitant of undisturbed evergreen broadleaf forest in the Mosquitia. Specimens were collected under and inside of rotten logs, under rocks, and in decaying root masses. This species is locally very abundant in certain areas of Parque Nacional Warunta, Honduras (McCranie et al. 2006). Remarks: Geophis bartholomewi was described by Brattstrom and Howell (1954) based on a single female specimen (CAS 91202; originally UCLA 6163) from “Arenal, elevation 1,200 feet, 25 km. east of Jalapa, Nuevo Segovia, Nicaragua.” The community of Arenales (visited by JHT in 2001) became part of Depto. El Paraiso, Honduras, following the final demarcation of the border with Nicaragua in 1960 (International Court of Justice 1960). Brattstrom and Howell (1954) differentiated G. bartholomewi from G. hoffmanni on the basis of having a shorter tail (14 subcaudals), smooth scales throughout the body, and an immaculate ventral scales. Downs (1967: 158), in justifying his synonymization of G. bartholomewi with G. hoffmanni, accurately pointed out that CAS 91202 has an incomplete tail and therefore an inaccurate subcaudal count, and that it is common for G. hoffmanni to have smooth scales (particularly in females) and immaculate ventral scales. Smith and Smith (1964) reported on a specimen of G. hoffmanni from eastern Olancho, Honduras (UI 53021; a female with 122 ventrals and 25 subcaudals. An additional specimen from Arenales (LACM 20483) was also available to Downs, a female with 25 subcaudals on an incomplete tail. Based on this additional material, Downs (1967: 158) maintained that “there remains the possibility (as suggested by

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104 Brattstrom and Howell) that bartholomewi is worthy of subspecific status, based on lower ventral and subcaudal counts.” Since the publication of Downs (1967), the number of specimens of G. hoffmanni available from northern Nicaragua and Honduras has increased considerably, largely through the collecting efforts of Tomas Manzanres and James McCranie in the Rus Rus and Warunta protected areas of the Honduran Mosquitia. Based on the material now available from eNCA and the data presented above and in Tables 15, a more detailed evaluation of the status of G. bartholomewi is now possible. While preliminary examination of the new data for G. hoffmanni does demonstrate that specimens from eNCA appear to have lower ventral and subcaudal counts, a more detailed study of material from Costa Rica and Panama is needed to facilitate direct comparison among all populations assigned to the name G. hoffmanni. Of the referred specimens listed below, CAS 91202 could not be loaned because of its status as the holotype of G. bartholomewi (J. Vindum, pers. comm.) and UIMNH 53021 was unavailable for examination due to construction at the museum in which it is housed (C. Phillips, pers. comm.). References: Peters (1859), Brattstrom and Howell (1954), Smith and Smith (1964), Downs (1967), McCranie et al. (2006). Illustrations: Peters (1859: black and white illustration of the dorsal and lateral aspects of the head and maxilla of the holotype), Downs (1967; black and white illustration of the dorsal and lateral aspects of the head), Khler (1999: black and white photograph and black and white illustrations of the lateral aspects of two heads showing variation in loreals; 2001b: color photograph and black and white illustrations of the

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105 lateral aspects of two heads showing variation in loreals; 2003: color photograph and black and white illustration of the dorsal and lateral aspects of the head), McCranie et al. (2006: color photograph). Specimens examined: 29; HONDURAS: Coln: Quebrada Machn, Reserva de la Biosfera Ro Pltano, 540 m (USNM 561035); El Paraiso: Arenales (LACM 20483); Gracias a Dios: Bodega de Ro Tapalws, Reserva Biologica Rus Rus, 200 m (USNM 561038, 561040); Cabaceras de Ro Rus Rus, Reserva Biologica Rus Rus (USNM 561039); Cerro Wahatingni Norte, Reserva Biologica Rus Rus, 190 m (USNM 562885); Hiltara Kiamp, Parque Nacional Ro Warunta, 150 m (USNM 562881); Kilpa Tingni Kiamp, Parque Nacional Ro Warunta, 150 m (USNM 562879); Urus Tingni Kiamp, Parque Nacional Ro Warunta, 160 m (USNM 561041, 561930, 561932); Warunta Tingni Kiamp, Parque Nacional Ro Warunta, 150 m (USNM 561042, 561931); Olancho: Campo Nuevo de Ro Tinto, 500 m (USNM 337522). NICARAGUA: Jinotega: vicinity of Pueblo Wiso, Reserva Biosfera Bosaws, 170 m (SMF 78594); Matagalpa: Hacienda la Cumplida (UMMZ 117653). Referred specimens: 3; HONDURAS: Coln: Mata de Maz (UNAH 5358; Wilson and Meyer 1985); El Paraiso: Arenales (CAS 91202; Brattstrom and Howell 1954); Olancho: 1.6 km E of Dulce Nombre de Culm (UIMNH 53021; Smith and Smith 1964). Geophis rhodogaster (Cope) Table 12, 13 Colophrys rhodogaster Cope, 1868: 130. Geophis rhodogaster: Bocourt, 1883: 531. Geophis fulvoguttatus: Khler, 1996: 36 (in error).

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106 Syntypes: Three: Academy of Natural Sciences of Philadelphia (ANSP) 3316 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.” Species group: dubius (Downs 1967). Distribution: Found in the mountains of southern Chiapas, Mexico, east through southern Guatemala to the Cordillera Metapn, Santa Ana, El Salvador, intermediate to high (1500 to 2744 m) elevations. This species probably also occurs in Honduras on the northeastern slope of Cerro Montecristo and northern slope Cerro El Pital along the border with El Salvador (Figure 18). Diagnosis: Geophis rhodogaster can be distinguished from G. damiani and G. hoffmanni by having 17 dorsal scale rows (15 dorsal scale rows in the aforementioned species). Geophis rhodogaster is differentiated from the remaining species with 17 dorsal scale rows (G. dunni, G. fulvoguttatus, and G. nephodrymus) by always lacking a supraocular scale and having the frontal contacting the orbit on both sides of the head (supraocular always present on at least one side of the head in G. dunni and G. fulvoguttatus, and always present on both sides or fused to the postocular in G. nephodrymus; frontal not contacting orbit on both sides in all three species). Description: Dorsal scales in 17 rows, smooth throughout length of body; ventrals 138, 138 in one adult male and 140 in three adult females; subcaudals 34, 43 in one adult male and 34 in three adult females; segmental count 176 in four specimens. Downs (1967: 94), based on specimens from throughout the range, gave a ventral range of 131 for males and 136 for females, and a subcaudal

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107 range of 39 in males and 29 in females. Smith (1995) reported the lowest recorded ventral count for G. rhodogaster, 129, from a male from Chiapas, Mexico. Snout-vent length, tail length, and tail length/total length ratio range of one adult male: 248 mm, 68 mm, 0.215; and three adult females: 249 mm, 55 mm, 0.146–0.197. Head indistinct from neck; snout rounded from above, rounded in profile, extending beyond the anterior end of the lower jaw; lower jaw reaches to the level of the posterior portion of the rostral; rostral generally as wider as high, extending posteriorly in dorsal view between internasals, portion of rostral visible in dorsal view 0.33 times the length of its distance from the frontal, with posterior termination at the level of the anterior edge of nostril; internasals 1.39.55 times as wide as they are long, rounded anteriorly, contacting the prenasal and postnasal laterally, their length 1.0.1 times as long as prefrontal common suture, their common suture 0.46.5 times as long as prefrontal common suture; prefrontals contacting postnasal, loreal, and orbit laterally, their length 0.61.63 times snout length, their common suture 0.26.38 times the frontal length; frontal 6-sided, 1.0.26 times as wide as is long, sharply angulate anteriorly, contacting orbit; supraocular absent; parietals 1.39.4 times as long as broad, their length 0.48.49 times the length of the head, their common suture 0.62.68 times as long as frontal; single postocular, 1.56.57 times as high as long; nasal divided; postnasal length 0.39.91 times that of prenasal length (in KU 57998 the postnasal on both sides is divided to form a small postnasal and a large, squarish preloreal); combined length of prenasal and postnasal 0.88.0 times the length of the loreal; loreal 1.42.82 times as long as deep, contained 1.9.12 times on snout length; loreal 1.7.0 times as

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108 long as eye horizontal diameter, dorsal margin straight; eye small, contained 3.6.8 times in snout length, its vertical diameter 0.73.8 times its distance from the lip. Supralabials 6/6; first supralabial contacting rostral, prenasal and postnasal; second supralabial contacting postnasal and loreal; third supralabial contacting loreal; third and fourth supralabials entering orbit; fourth supralabial contacting postocular; fifth supralabial the largest, contacting the postocular, and parietal; lip exposure of third supralabial 1.33.5 times that of second supralabial; lip exposure of fifth supralabial 1.19.88 times that of fourth supralabial; anterior temporal absent; one posterior temporal, separating sixth supralabial from parietal; posterior temporals separated from each other by seven nuchals; mental 1.88.89 times as broad as long, angulate anteriorly, separated from chinshields by first infralabials; infralabials 6 or 7 (6/6 in two females, 6/7 in one female, and 7/7 in one male); first infralabials broadly in contact between mental and anterior chinshields; first, second, and third infralabial contacting the anterior chinshield; fourth infralabial contacting posterior chinshield; anterior chinshields 1.71.85 times as long as broad, 1.0.33 times as long as posterior chinshields; posterior chinshields barely to broadly in contact; 2 medial gular scales present. Hemipenis: Downs (1967: 94) described the hemipenis as long and slender, extending posteriorly to subcaudals 11; basal portion of organ bearing many minute spines and, in distal parts, two or three large spines; basal naked pocket present on the asulcate side; central part of the organ bearing around 35 medium spines and hooks; distal portion of hemipenis capitate, with the capitation obscured by gradation between spines on central portion of hemipenis and spines on proximal edge of capitulum; capitulum spinulate proximally, calyculate and papillate distally; strongly bilobed, with

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109 each lobe extending two subcaudals in length; sulcus spermaticus bifurcates around subcaudal eight or nine. Dentition: Downs (1967: 94) described the maxilla of G. rhodogaster as extending anteriorly to the suture between supralabials 1 and 2; anterior extension greater than that of palatine; maxilla narrower anteriorly than posteriorly, bearing 14 teeth that increase in length posteriorly; anterior tip of maxilla bearing first tooth; posterior end of maxilla compressed and and expanded into a nearly horizontal flange; posterior flange bearing final few teeth; anterior end of ectopterygoid bifurcate, with one branch short and blunt and the other long and blade-like; no postorbital bone present. Coloration: Based on one male and three female specimens in preservation: dorsal surface of head slate gray, lower half of rostral, postnasals, and loreals cream, first and second supralabials cream, third through sixth supralabials with anterior one-third or less slate gray with cream on posterior portions and mottling at the contact zone between dorsal and ventral coloration, immaculate cream infralabials and ventral surface of head; dorsal surface of body slate gray; first dorsal scale row cream; second dorsal scale row cream with dark gray posterior edges to mostly gray with pale anterior one-third; in KU 57997, the third dorsal scale row has pale anterior one-quarter to one-third; ventral surface of body immaculate cream; dorsal surface of tail slate gray, becoming somewhat darker posteriorly; ventral surface of tail cream with some scattered dark flecks or spots. Khler et al. (2006: 167) presented a color photograph of a live specimen and reported the dorsal and lateral coloration in life as dark grayish to reddish brown, with yellow spots in the second dorsal scale row, and the ventral surfaces as immaculate yellow or yellowish white.

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110 Natural History Comments: This species is found under ground cover in a variety of disturbed and undisturbed habitats, including cloud forest, pine-oak forest, cultivated hillsides and fields, and dry grassy areas (Stuart 1951, Downs 1967, Khler et al. 2006). References: Cope (1868), Downs (1967), Smith (1995), Khler et al. (2006). Illustrations: Downs (1967; black and white illustration of the dorsal and lateral aspects of the head of the holotype), Khler et al. (2006; color photograph). Specimens examined: 4; EL SALVADOR: Santa Ana: Hacienda Montecristo, Cordillera de Metapn, 1780 m (SMF 77413), 2200 m (KU 57997). Referred specimens: 5; EL SALVADOR: Santa Ana: Hacienda Montecristo, Cordillera de Metapn, 2200 m (KU 58000); Montecristo (MUHNES 490, 1141; Khler et al. 2006); Chalatenango: Cerro El Pital, 2100 m (KU 291579). GUATEMALA: Jalapa: 8.5 km ENE Mataquescuintla, 2630 m (MVZ 160733).

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111 reddish to grayish brown; ventral coloration yellow to red-orange; found in northern APPENDIX A KEY TO THE SPECIES OF Geophis FROM EASTERN NUCLEAR CENTRAL AMERICA In order to assist in the identification of specimens of Geophis originating in eNCA, the full range of morphological and meristic data recorded for each species is summarized in Table 10, and a dichotomous key is presented below. 1.a. Dorsal scales in 15 rows at midbody.............................................................................2 b. Dorsal scales in 17 rows at midbody............................................................................3 2. a. Dorsal coloration consists of dark gray ground color with red crossbands and halfbands throughout the length of the body; ventral scales white with heavy infusion of gray, sometimes forming stripes on the posterior edges of each scale; maximum total length 327 mm; apparently restricted to Cerro Texguat, Cordillera Nombre de Dios, north central Honduras, 1550 m elevation.............G. damiani b. Dorsal coloration uniform grayish brown; ventral scales white with no or very little gray pigment on the lateral edges of each scale; maximum total length 190 mm; found in the eastern Caribbean lowlands and foothills of Honduras south to Panama, and on the Pacific versant of Costa Rica, 150 m elevation..............G. hoffmanni 3. a. Dorsal scales prominently keeled; dorsal coloration yellow with brown crossbands; eight infralabials, known only from Matagalpa, Nicaragua............................G. dunni b. Dorsal scales smooth; dorsal ground coloration dark brown or gray..............................4 4. a. Supraocular present on at least one side of head..........................................................5 b. Supraocular absent, frontal contacting orbit; postocular present; dorsal coloration

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112 El Salvador, central Guatemala, and southwestern Chiapas, Mexico, 1500 m elevation................................................................................................G. rhodogaster 5. a. Ventral + subcaudals 171, red-orange or yellow middorsal blotches rarely contacting and never reaching below the 2nd dorsal scale row; maximum total length 443 mm; found in northwestern Santa Ana, El Salvador, and Ocotepeque and Copn, Honduras, 1680 m elevation....................................................... G. fulvoguttatus b. Ventral + subcaudals 149, red or gray dorsolateral and lateral blotches and partial bands regularly entering the 1st dorsal scale row; maximum total length 294 mm; found in Parque Nacional El Cusuco, Corts, Honduras, 1545 m elevation.................................................................................. G. nephodrymus

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118 McCranie, J. R. 2004c. Anomalepis mexicanus Jan (Serpentes, Anomalepididae) in Honduras. Herpetological Bulletin 89: 21. McCranie, J. R. and F. E. Castaeda. 2004a. A new species of snake of the genus Omoadiphas (Reptilia: Squamata: Colubridae) from the Cordillera Nombre de Dios in northern Honduras. Proceedings of the Biological Society of Washington 117: 311. McCranie, J. R. and F. E. Castaeda. 2004b. Notes on the second specimens of Geophis damiani Wilson, McCranie and Williams and Rhadinaea tolpanorum Holm and Cruz D. (Colubridae). Herpetological Review 35: 341. McCranie, J. R. and F. E. Castaeda. 2005. The herpetofauna of Parque Nacional Pico Bonito, Honduras. Phyllomedusa 4: 3. McCranie, J. R., M. R. Espinal and L. D. Wilson. 2005. New species of montane salamander of the Bolitoglossa dunni group from northern Comayagua, Honduras (Urodela: Plethodontidae). Journal of Herpetology 39: 108. McCranie, J. R., K. E. Nicholson, and F. E. Castaeda. 2002. Geographic distribution. Eleutherodactylus diastema. Herpetological Review 33: 220. McCranie, J. R., J. H. Townsend, and L. D. Wilson. 2003a. Three snakes new to the herpetofauna of Honduras. Herpetological Review 34: 391. McCranie, J. R., J. H. Townsend, and L. D. Wilson. 2003b. Hyla miliaria (Anura: Hylidae) in Honduras, with notes on calling site. Caribbean Journal of Science 39: 398. McCranie, J. R., J. H. Townsend, and L. D. Wilson. 2006. The Amphibians and Reptiles of the Honduran Mosquitia. Krieger Publishing, Malabar, Florida. x + 291 pp. McCranie, J. R. and L. D. Wilson. 1991. Geophis fulvoguttatus Mertens and Micrurus browni Schmidt and Smith: additions to the snake fauna of Honduras. Amphibia Reptilia 12: 112. McCranie, J. R., and L. D. Wilson. 2002. The Amphibians of Honduras. Society for the Study of Amphibians and Reptiles Contributions to Herpetology, Volume 19. Ithaca, New York. x + 625 p. McCranie, J. R., L. D. Wilson, and G. Khler. 2005. Amphibians and Reptiles of the Bay Islands and Cayos Cochinos, Honduras. Bibliomania, Salt Lake City, Utah. xiv + 210 p. McWest, K. J. 1997. Description of the male of Diplocentrus lourencoi (Scorpiones: Diplocentridae). Journal of Arachnology 25: 251.

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120 Prez-Higareda, G., H. M. Smith, and M. A. Lpez-Luna. 2001. A new Geophis (Reptilia: Serpentes) from southern Veracruz, Mexico. Bulletin of the Maryland Herpetological Society 37: 42. Poe, S. 2004. Phylogeny of anoles. Herpetological Monographs 18: 37. Population Reference Bureau. 2005. 2005 World Population Data Sheet (http://www.prb.org/) viewed on 12 December 2004. Poulsen, B. O., and N. Krabbe. 1997. Avian rarity in ten cloud-forest communities in the Andes of Ecuador: implications for conservation. Biodiversity and Conservation 6: 1365. Pounds, J. A., M. P. Fogden, and J. H. Campbell. 1999. Biological response to climate change on a tropical mountain. Nature 398: 611. Pounds, J. A., and M. P. Fogden. 2000. Amphibians and reptiles of Monteverde, pp. 537–540. In N. M. Nadkarni and N. T. Wheelwright (eds.). Monteverde. Ecology and Conservation of a Tropical Cloud Forest. Oxford Univ. Press, New York. xxiii + 573 p. Savage, J. M. 2002. The Amphibians and Reptiles of Costa Rica: A Herpetofauna between Two Continents, between Two Seas. University of Chicago Press. Chicago, Illinois. xx + 934 p. Savage, J. M., J. A. Campbell, and W. W. Lamar. 2005. On names for Neotropical rattlesnakes (Reptilia: Serpentes: Viperidae: Crotalus). Herpetological Review 36: 369. Sawaya, R. J., and I. Sazima. 2003. A new species of Tantilla (Serpentes: Colubridae) from southeastern Brazil. Herpetologica 59: 119. Schmidt, K. P. 1932. Stomach contents of some South American coral snakes, with the description of a new species of Geophis. Copeia 1932: 6. Schmidt, K. P. 1933. New reptiles and amphibians from Honduras. Zoological Series of Field Museum of Natural History 20: 15. Schuchert, C. 1935. Historical Geology of the Antillean-Caribbean Region. John Wiley and Sons, New York. 811 p. Seidel, M. E. 2002. Taxonomic observations on extant species and subspecies of slider turtles, genus Trachemys. Journal of Herpetology 36: 285. Smith, E. N. 1995. Geophis rhodogaster (Colubridae), an addition to the snake fauna of Mexico. Southwestern Naturalist 40: 123.

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121 Smith, H. M. 1959. New and noteworthy reptiles from Oaxaca, Mexico. Transactions of the Kansas Academy of Science 62: 265. Smith, H. M., and R. L. Holland. 1969. Two new snakes of the genus Geophis from Mexico. Transactions of the Kansas Academy of Science 72: 47. Smith, H. M., and P. W. Smith. 1964. Range extension of a snake (Geophis) in Central America. Herpetologica 20: 72. Smithe, F. B. 1975. Naturalist’s Color Guide. Part I. Color Guide. American Museum Natural History, New York. Solorzano L., A. 2004. Serpientes de Costa Rica: Distribucin, Taxonoma, e Historia Natural. Snakes of Costa Rica: Distribution, Taxonomy, and Natural History. Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica. 791 p. Stadtmller, T. 1987. Cloud forests in the humid tropics: a bibliographic review. United Nations University, Tokyo, Japan, and Centro Agronmico Tropical de Investigacin y Enseanza, Turrialba, Costa Rica. 81 p. Stafford, P. 2004. A new species of Tantilla (Serpentes: Colubridae) of the taeniata group from southern Belize. Journal of Herpetology 38: 43. Stuart, L. C. 1951. The herpetofauna of the Guatemalan plateau, with special reference to its distribution in the southwestern highlands. Contributions of the Laboratory of Vertebrate Biology, University of Michigan 49: 1. Stuart, L. C. 1963. A checklist of the herpetofauna of Guatemala. Miscellaneous Publications of the Museum of Zoology, University of Michigan 122: 1. Talley, B. L., D. C. Fraser, L. D. Wilson, and J. H. Townsend. 2005. Natural History Notes. Bothrops asper (Barba amarilla, Terciopelo). Maximum elevation. Herpetological Bulletin 94: 29. Townsend, J. H. 2005. Geographic Distribution. Sphenomorphus incertus (Stuart’s Forest Skink). Herpetological Review 36: 337. Townsend, J. H., and T. L. Plenderleith. 2005. Geographic Distribution. Anolis (Norops) petersii (Peters’ Anole). Herpetological Review 36: 466. Townsend, J. H., L. D. Wilson, T. L. Plenderleith, B. L. Talley, and J. C. Nifong. 2005a. Ninia pavimentata (Squamata: Colubridae): an addition to the snake fauna of Honduras. Caribbean Journal of Science 41: 869. Townsend, J. H., S. M. Hughes, and T. L. Plenderleith. 2005b. Geographic Distribution. Anolis (Norops) ocelloscapularis (NCN). Herpetological Review 36: 466.

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122 Townsend, J. H., J. C. Nifong, and L. D. Wilson. 2005c. First record of the colubrid snake Rhadinaea anachoreta Smith and Campbell from Honduras. Herpetological Bulletin 94: 2. Townsend, J. H., and L. D. Wilson. 2006. A new species of snake of the Geophis dubius group (Reptilia: Squamata: Colubridae) from the Sierra de Omoa of northwestern Honduras. Proceedings of the Biological Society of Washington 119: 150. Townsend, J. H., L. D. Wilson, B. L. Talley, D. C. Fraser, T. L. Plenderleith, and Sara M. Hughes. 2006a. Additions to the herpetofauna of Parque Nacional El Cusuco, Honduras. Herpetological Bulletin. In press. Townsend, J. H., J. C. Nifong, and R. Downing M. 2006b. Oedipina elongata (Schmidt, 1936) in Honduras. Salamandra 42(1): 61. Wiles, P. R. 2005. First collection of water mites from Honduras: descriptions of six new species. Journal of Natural History 39: 253. Williams, K. L., L. D. Wilson, and J. H. Townsend. In preparation. Atractus Wagler. Catalogue of American Amphibians and Reptiles. Wilson, L. D. 1999. Checklist and key to the species of the genus Tantilla (Serpentes: Colubridae), with some distributional commentary. Smithsonian Herpetological Information Service 122:1. Wilson, L. D., and J. A. Campbell. 2000. A new species of the calamarina group of the colubrid snake genus Tantilla from Guerrero, Mexico. Proceedings of the Biological Society of Washington 113: 820. Wilson, L. D. and J. R. McCranie. 1992. Status of amphibian populations in Honduras. Unpublished Report to the Declining Amphibian Populations Task Force, 15 August 1992. 14 p. Wilson, L. D. and J. R. McCranie. 1998. The biogeography of the herpetofauna of the subhumid forests of Middle America (Isthmus of Tehuantepec to northwestern Costa Rica). Royal Ontario Museum of Life Sciences, Contribution 163: 1. Wilson, L. D., J. R. McCranie, and K. L. Williams. 1998. A new species of Geophis of the sieboldi group (Reptilia: Squamata: Colubridae) from northern Honduras. Proceedings of the Biological Society of Washington 111: 410. Wilson, L. D. and J. R. McCranie. 2002. Update on the list of reptiles known from Honduras. Herpetological Review 33: 90. Wilson, L. D. and J. R. McCranie. 2003. Herpetofaunal indicator species as measures of environmental stability in Honduras. Caribbean Journal of Science 39: 50.

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123 Wilson, L. D. and J. R. McCranie. 2004a. The herpetofauna of the cloud forests of Honduras. Amphibian and Reptile Conservation 3: 34. Wilson, L. D. and J. R. McCranie. 2004b. The herpetofauna of Parque Nacional El Cusuco, Honduras. Herpetological Bulletin 87: 13. Wilson, L. D. and J. R. McCranie. 2004c. The conservation status of the herpetofauna of Honduras. Amphibian and Reptile Conservation 3: 6. Wilson, L. D., J. R. McCranie, S. W. Gotte, and J. H. Townsend. 2003. Distributional comments on some members of the herpetofauna of the Mosquitia, Honduras. Herpetological Bulletin 84: 15. Wilson, L. D., and J. H. Townsend. 2006. The herpetofauna of the rainforests of Honduras. Caribbean Journal of Science. 42(1). In press. Wilson, L. D., J. H. Townsend, T. L. Plenderleith, B. L. Talley, and D. C. Fraser. 2006. Squamata, Colubridae, Rhadinaea montecristi: distribution extension, geographic distribution map. Check List 2: 8. Wster, W., J. L. Yrausquin, and A. Mijares-Urrutia. 2001. A new species of indigo snake from north-western Venezuela (Serpentes: Colubridae: Drymarchon). Herpetological Journal 11: 157. Zaldvar-Rivern, A., V. Len-Regagnon, and A. Nieto-Montes de Oca. 2004. Phylogeny of the Mexican coastal leopard frogs of the Rana berlandieri group based on mtDNA sequences. Molecular Phylogenetics and Evolution 30: 38.

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BIOGRAPHICAL SKETCH Josiah H. Townsend was born in 1978 in Enid, Oklahoma. After living in New Orleans and Sulphur, Louisiana; Pittsburgh, Pennsylvania; and Hauppauge, New York, he moved to Miami, Florida. There he attended Miami Palmetto Senior High School and upon graduation entered Miami Dade Community College. During his first year at MDCC he met Deborah Talia Vergara, whom he would marry in August of 2004. While at MDCC, and under the tutelage of his professor Dr. Larry Wilson, Townsend began to explore his interest in herpetology and field biology in the forests of eastern Honduras. He transferred to the University of Florida in 2000. In 2004, he received his Bachelor of Science Degree in Wildlife Ecology and Conservation with a minor in Zoology. While at the University of Florida, he volunteered and worked at the Florida Museum of Natural History in the Herpetology Division and the Specimen Preparation Lab, and continued to work alongside Wilson and Randy McCranie in Honduras, conducting herpetological research in the Mosquitia. During this time Townsend also participated in various research projects relating to the native and introduced herpetofauna of Florida. After receiving his bachelor’s degree, Townsend entered the Master of Arts in Latin American Studies Program at the University of Florida, with a concentration in Tropical Conservation and Development. After graduation, Townsend will continue his studies in the Interdisciplinary Ecology PhD program at the School of Natural Resources and Environment, University of Florida. 124