<%BANNER%>

Integrative Taxonomy Reveals the Chortís Block of Central America as an Underestimated Hotspot for Amphibian Diversity

Permanent Link: http://ufdc.ufl.edu/UFE0043737/00001

Material Information

Title: Integrative Taxonomy Reveals the Chortís Block of Central America as an Underestimated Hotspot for Amphibian Diversity
Physical Description: 1 online resource (350 p.)
Language: english
Creator: Townsend, Josiah H
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2011

Subjects

Subjects / Keywords: 16s -- amphibia -- anura -- barcode -- caudata -- chortis -- coi -- evolution -- extension -- honduras -- mesoamerica -- nicaragua -- nototriton -- phylogenetics -- plethodontidae -- systematics -- taxonomy
Interdisciplinary Ecology -- Dissertations, Academic -- UF
Genre: Interdisciplinary Ecology thesis, Ph.D.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: Central America has a remarkably complex geological history, has served as the biological dispersal route between North and South America, and is the site of extensive in situ evolution. Nuclear Central America is recognized as a region of high biodiversity, and the eastern portion of Nuclear Central America (the Chortís Block) has largely been overlooked as a biodiversity hotspot. In this dissertation, I use an integrative systematic approach to examine evolutionary patterns in a group with high diversity of endemic species that is also of global conservation priority: amphibians. I carried out 17 expeditions to over 60 localities in the Chortís Block for this dissertation, results from which are synthesized with existing data on regional herpetofaunal diversity, distribution, and conservation status. The Chortís herpetofauna is characterized by a high degree of endemism (35% of all species are endemic) and equally high extinction risk (41% threatened, including 96% of endemic species). Endemism is highest among salamanders (86% of species are endemic). A total of 456 amphibian samples representing 52 species were sequenced for two genes (16S and COI) and analyzed using distance-based and phylogenetic methods. I identify at least 36 unnamed "candidate" across eight families of amphibians, while confirming that four taxonomically-uncertain salamander populations represented new allopatric populations of endangered species previously assumed to be single-site endemics. These results reveal a regional amphibian fauna, already recognized as being of global conservation priority, with species diversity underestimated by 26% (47% in salamanders), highlighting the critical role molecular systematics plays in endangered species conservation and management. To directly address the systematics of one group with multiple candidate species, I present an integrative taxonomic revision of the genus Nototriton using data from three genes, external morphology, and osteology. I restrict the taxon N. barbouri to populations from the Sierra de Sulaco, describe three new species from the Chortís Highlands, and a review of the remaining regional species. In light of these findings, I present my vision for using systematic biology as the basis for integrating research, education, and outreach in support of biodiversity conservation in the Chortís Block.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Statement of Responsibility: by Josiah H Townsend.
Thesis: Thesis (Ph.D.)--University of Florida, 2011.
Local: Adviser: Austin, James.
Electronic Access: RESTRICTED TO UF STUDENTS, STAFF, FACULTY, AND ON-CAMPUS USE UNTIL 2014-12-31

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2011
System ID: UFE0043737:00001

Permanent Link: http://ufdc.ufl.edu/UFE0043737/00001

Material Information

Title: Integrative Taxonomy Reveals the Chortís Block of Central America as an Underestimated Hotspot for Amphibian Diversity
Physical Description: 1 online resource (350 p.)
Language: english
Creator: Townsend, Josiah H
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2011

Subjects

Subjects / Keywords: 16s -- amphibia -- anura -- barcode -- caudata -- chortis -- coi -- evolution -- extension -- honduras -- mesoamerica -- nicaragua -- nototriton -- phylogenetics -- plethodontidae -- systematics -- taxonomy
Interdisciplinary Ecology -- Dissertations, Academic -- UF
Genre: Interdisciplinary Ecology thesis, Ph.D.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: Central America has a remarkably complex geological history, has served as the biological dispersal route between North and South America, and is the site of extensive in situ evolution. Nuclear Central America is recognized as a region of high biodiversity, and the eastern portion of Nuclear Central America (the Chortís Block) has largely been overlooked as a biodiversity hotspot. In this dissertation, I use an integrative systematic approach to examine evolutionary patterns in a group with high diversity of endemic species that is also of global conservation priority: amphibians. I carried out 17 expeditions to over 60 localities in the Chortís Block for this dissertation, results from which are synthesized with existing data on regional herpetofaunal diversity, distribution, and conservation status. The Chortís herpetofauna is characterized by a high degree of endemism (35% of all species are endemic) and equally high extinction risk (41% threatened, including 96% of endemic species). Endemism is highest among salamanders (86% of species are endemic). A total of 456 amphibian samples representing 52 species were sequenced for two genes (16S and COI) and analyzed using distance-based and phylogenetic methods. I identify at least 36 unnamed "candidate" across eight families of amphibians, while confirming that four taxonomically-uncertain salamander populations represented new allopatric populations of endangered species previously assumed to be single-site endemics. These results reveal a regional amphibian fauna, already recognized as being of global conservation priority, with species diversity underestimated by 26% (47% in salamanders), highlighting the critical role molecular systematics plays in endangered species conservation and management. To directly address the systematics of one group with multiple candidate species, I present an integrative taxonomic revision of the genus Nototriton using data from three genes, external morphology, and osteology. I restrict the taxon N. barbouri to populations from the Sierra de Sulaco, describe three new species from the Chortís Highlands, and a review of the remaining regional species. In light of these findings, I present my vision for using systematic biology as the basis for integrating research, education, and outreach in support of biodiversity conservation in the Chortís Block.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Statement of Responsibility: by Josiah H Townsend.
Thesis: Thesis (Ph.D.)--University of Florida, 2011.
Local: Adviser: Austin, James.
Electronic Access: RESTRICTED TO UF STUDENTS, STAFF, FACULTY, AND ON-CAMPUS USE UNTIL 2014-12-31

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2011
System ID: UFE0043737:00001


This item has the following downloads:


Full Text

PAGE 1

1 INTEGRATIVE TAXONOMY REVEALS THE CHORT S BLOCK OF CENTRAL AMERICA AS AN UNDERESTIMATED HOTSPOT OF AMPHIBIAN DIVERSITY By JOSIAH HAROLD TOWNSEND A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2011

PAGE 2

2 2011 Josiah Harold Townsend

PAGE 3

3 Para Ileana And i n loving memory of my grandfather, Vernon Lynn Boyd

PAGE 4

4 ACKNOWLEDGMENTS I want to begin b y expressing my gratitude for the support shown by my advisory committee during the course of preparing this dissertation. My Chair, James Austin, supported me both academically and pers onally throughout this process; his leadership and steadfast s upport was instrumental in my success I want to thank my committee members, Rob Fletcher, Mike Miyamoto, Rick Stepp, and Larry David Wilson, for their guidance, advice, and support through the process of researching and writing this dissertation. Writing a dissertation must be both a singularly selfish act and one that relies on the extended patience and support of loved ones, lab mates, and colleagues T his certainly was the case for me. To the one person who is literally all three of those things, my wif e Ileana Luque Montes, I cannot adequately express the amazement with which I have watched you begin your own graduate studies over the past four months; maintaining a rigorous field schedule and supervising undergraduates, while I was of minimal help and fully focused on writing the following document. I simply would not have completed this without you r remarkable dedication and support and my gratitude will be best expressed by giving you the same degree of support as you pursue your own goals I am bles sed with the devoted support and seemingly endless patience and understanding of a wonderful and inspirational family, my parents Steve and Terri Boyd Townsend my sister Katielynn, and my grandparents Vernon Lynn and Norma Jean Boyd and Willa Parker Towns end, who are largely responsible for any past and future successes I may have. In many ways, I am who I am today thanks to the mentorship of Larry David Wilson. As a mentor, colleague, and friend, Larry introduced me to Honduras in 1999

PAGE 5

5 and guided my dev eloping research interests there throughout my community college and undergraduate years becoming my colleague and supporter as I progressed through my graduate studies. I owe no small debt of gratitude to my friends and colleagues in the Austin Lab and in particular for actively assisting with my labwork and providing countless impromptu instructional sessions I want to thank Jason Butler, John Hargrove, Nathan Johnson, Emily Saarinen, Matt Shirley, and Aria (Johnson) St. Louis I also b enefited from the dedicated work of a series of undergraduate researchers and volunteers: Teresa Burlingame, Dania Gutierrez, Jaclyn Irwin, Rachel Shapiro, Vicki Villanova and Lauryn Walter My work for the past six years has benefited greatly from the support of the Section of Protected Areas and Wildlife, Instituto Nacional de Conservacin y Desarrollo Forestal, reas Protegidas y Vida Silvestre, and particularly Iris Acosta, Carla Crcamo, Sad Lainez O. Andrs Alegria Ramn Alvarez L. Wilson Zniga D., Sonia Ma rtnez Moreno, and Wendy Aronne (Instituto Nacional de Conservacin y Desarrollo Forestal, reas Protegidas y Vida Silvestre [ICF]). F ieldwork was carried out under a series of research permits issued by ICF most recently Resolucin DE MP 086 201 0 and Dic tamen DVS ICF 045 2010 In Honduras, my work would not have been possible without the hard work and support variously given by Ileana Luque Montes (UNAH), Melissa Medina Flores ( UNAH, UNA), Luis A. Herrera (UNAH), A llan J. Fuentes and E duardo J. Zavala (P ROLANSATE), Alcalde Adolfo Pagoada Saybe (Municipalidad de Arizona), Alcadesa Teresa Espinosa Aguilar (Municipalidad de Marale), Oliver Komar, Jose Mora, Jorge Ivn Restrepo, and Fredy Membre o (Centro Zamorano de

PAGE 6

6 Biodiversidad), E frain Aguilar (San Jos d e Texguat) A lfonso Contreras (Mezapita) A lionso Portillo (Jilamito Nuevo), J Dubn ( La Liberacin ), Mario Orellana Leiva (El Playon), Leonel Erazo Chvez (El Cedral), Rafael Ulloa (Municipalidad de Gualaco) Eduardo Rico (ICF Gualaco), Carlos Perdomo ( Aldea Global), Paul House (UNAH, Herbario TEFH), Robert Dale (Los Naranjos), and Alicia Ward (Santa Brbara) I would like to acknowledge the following people for their work in the field in support of this project: Carlos Andino, Ben K. Atkinson, James D. Austin, Christopher Begley, Mark Bonta, Jason M. Butler, Brian Campesano, Csar A. Cerrato, Gabriela Diaz, Anne Donnelly, Matthew Donnelly, Yensi Flores, Sergio C. Gonzalez, Levi Gray, Vladlen Henriquez Luis A. Herrera, Paul House, Robert Hyman, Lorraine Ketzler, Ileana Luque, Christina Martin, David Medina, Melissa Medina Flores, Mayron Mckewy Meja, Aaron Mendoza, Wendy Naira, Ciro Navarro, Lenin Obando, Sandy Pereira, Onn A. Reyes, John Slapcinsky, Mario Sols, Fito Steiner, Nathaniel Stewart, Alexande r Stubbs, Katielynn Townsend, Steve Townsend, Scott L. Travers Rony Valle, Hermes Vega, Alicia Ward and Christopher Wolf I am very grateful for the support of Amy Driscoll, Dan Mulcahy, Andrea Ormos and the SI Barcoding Project (Smithsonian Institutio n Laboratory of Analytical Biology), eposition of voucher specimens and timely acquisition of catalog numbers was facilitated by : Jim Mc Guire, Ted Papenfuss, Sean Rovito, Carol Spencer, and David Wake (MVZ), J ose Rosado (MCZ), and Steve Gotte, Jeremy Jacobs, John Poindexter and Robert Wilson (USNM). Jason Butler, Ileana

PAGE 7

7 Luque, Javier Sunyer, and Scott Travers kindly contributed photograph s for use in this dissertation. Various portions of this dissertation was funded by Critical Ecosystem Partnership Fund (CEPF), a Summer Research Grant from the Working Forests in the Tropics IGERT Program (National Science Foundation DGE 0221599) at the University of Florida, a grant to K irsten E. Nicholson (Central Michigan University; National Science Foundation DEB During my dissertation studies, I was supported by a 2007 09 NSF GK 12 Fell owship from the UF SPICE Program, and as much as any experience during my graduate education, my participation in this program was truly formative and help to shape the conceptual framework within which I propose to carry out research. I am especially grat eful to Doug Levey and Suzan Smith, my partner teacher Nate Stewart, and fellow fellows Jackson Frechette, Rachel Naumann, and Tom Tidyman, for helping to make SPICE a truly great experience Following SPICE, my work with the UFTeach Program as an instruct or for their core Research Methods class was key during the final two years of my dissertation, and I thank Alan Dorsey, Dimple F lesner, Griff Jones, Linda Jones, Katrina Short, Gloria Weber, and our RM students for making these last two spring semesters s o enjoyable Finally, I want to take this opportunity to dedicate my work in the past and future to the memory of three friends that I lost while completing this degree. I met Thad Owens as a student in my Herpetology class at UF and we quickly became fri end s as he did with just about everyone who knew him. Thad was one of the most honest, unreserved, and enthusiastic people I will ever have the pleasure of knowing, and rarely a day

PAGE 8

8 passes since he was lost in May 2009 that I do not think about him or fin d inspiration in his memory. Wade Wassenberg was a good friend since high s chool, and was one of the most loyal friends anyone could hope t o have After losing tra ck of him for years, I learned that Wade had j oined the US Army and served in the elite 1st Ba t ta lion of the 75th Ranger Regiment surviving four tours of duty in Afghanistan and two more in Iraq from 2 002 to 2008. After retir ing to rejoin life with his beautiful young family he was suddenly diagnosed with brain cancer and passed soon thereafter at the VA Hospital in Gainesville walking distance from my office I will always regret that while my friend suffered and passed literally minutes away from my home, I was unaware and in the field in Honduras Finally, Don Ma rio Guiffaro of Olancho, Honduras, was an influential and remarkable man whose l ege ndary life story included being a feared pistolero a frontier gold miner, a jaguar hunter, and in his later years, and outstanding advocate and conservationist in the Patu illegal logging and drug trafficking in the Mosquitia was ultimately answered by violence, and he was gunned down in front of his son and friends. The memories of, and examples set, by these three frie nd s, to each of whom I would not have hesitated to entrust with my life, and in some cases did just that, will always be my compass and my motivation as I continue through life

PAGE 9

9 TABLE OF CONTENTS page ACKNOW LEDGMENTS ................................ ................................ ................................ .. 4 LIST OF TABLES ................................ ................................ ................................ .......... 14 LIST OF FIGURES ................................ ................................ ................................ ........ 15 ABSTRACT ................................ ................................ ................................ ................... 18 CHAPTER 1 INTRODUCTION ................................ ................................ ................................ .... 20 2 BIOGEOGRAPHIC DELIMITATION OF THE CHORTS BLOCK: GEOLOGICAL HISTORY, PHYSIOGRAPHY, AND ECOLOGICAL ASSOC IATIONS .................... 25 Geological History ................................ ................................ ................................ ... 27 Contemporary Ecophysiography ................................ ................................ ............. 34 Characterizing Ecological Associations: Holdridge Forest Formations ................... 46 Characterizing Ecological Associations: Updating and Operationalizing the Carr (1950) System for Classifying Honduran Ec osystems ................................ ......... 50 Lowland associated Habitats ................................ ................................ ............ 51 Habitats Shared Between Lowlands and the Chorts Highlands ...................... 57 Habitats of the Chorts Highlands ................................ ................................ ..... 61 Introduction to Biodiversity of the Chorts Block ................................ ...................... 70 3 TAXONOMIC DIVERSITY, DISTRIBUTIONAL PATTERNS, AND CONSERVATION STATUS OF THE CHORTS BLOCK HERPETOFAUNA ......... 72 Methods and Materials ................................ ................................ ............................ 74 Field Sampling ................................ ................................ ................................ .. 74 Taxonomic Scope and Standards ................................ ................................ .... 74 Evaluating Conservation Status ................................ ................................ ....... 77 Results ................................ ................................ ................................ .................... 78 Composition of the Herpetofauna ................................ ................................ ..... 78 Patterns of Distribution and Endemism within the Cho rts Block ...................... 79 Distribution of Chorts Highland Endemics ................................ ....................... 79 Conservation Status ................................ ................................ ......................... 80 Sampling Results by Locality ................................ ................................ .................. 80 Parque Nacional Celaque ................................ ................................ ................. 81 Parque Nacional Cerro Azul Copn ................................ ................................ 82 Parque Nacional Cerro Azul Membar ................................ ............................. 83 Parque Nacional Cusuco ................................ ................................ .................. 86 Parque Nacional La Tigra ................................ ................................ ................. 87 Parque Nacional Montaa de Botaderos ................................ .......................... 87

PAGE 10

10 Parque Nacional Montaa de Comayagua ................................ ....................... 90 Parque Nacional Montaa de Santa Brbara ................................ ................... 91 Parque Nacional Montaa de Yoro ................................ ................................ .. 94 Parque Nacional Pico Bonito ................................ ................................ ............ 95 Parque Nacional Pico Pijol ................................ ................................ ............... 95 Parque Nacional Sierra de Agalta ................................ ................................ .... 96 Refugio de Vida Silvestre Texiguat ................................ ................................ .. 97 Reserva Biolgica Cerro Uyuca ................................ ................................ ....... 98 Reserva Biolgica Guajiquiro ................................ ................................ ......... 100 Reserva Biolgica Gisayote ................................ ................................ ......... 101 Reserva Biolgica Yerbabuena ................................ ................................ ...... 102 Reserva de la Biosfera Bosawas ................................ ................................ .... 102 Jardn Botnico Lancetilla ................................ ................................ .............. 104 rea de Uso Multple Isla del Tigre ................................ ................................ 106 Non Protected Areas ................................ ................................ ...................... 107 Cerro El Zarciadero ................................ ................................ .................. 107 Highlands surround the Meseta de La Esperanza ................................ ... 107 Los Naranjos ................................ ................................ ............................ 108 Montaa de Jacaleapa ................................ ................................ ............. 109 Montaa Macuzal ................................ ................................ ..................... 109 Saguay ................................ ................................ ................................ ..... 112 San Jos de Texguat ................................ ................................ .............. 112 Selva Negra ................................ ................................ ............................. 113 Yeguare Valley ................................ ................................ ......................... 113 Discussion ................................ ................................ ................................ ............ 116 Baseline Herpetological Inventory of Parqu e Nacional Montaa de Yoro ............. 116 A New Species of Anole ................................ ................................ ................. 118 Salamanders of Uncertain Taxonomic Assignment ................................ ........ 119 Hotspot within a Hotspot: the Special Case of Refugio de Vida Silvestre Texguat ................................ ................................ ................................ ............. 120 Discovery of Plectrohyla chrysospleura ................................ .......................... 122 Underestimated Salamander Diversity? ................................ ......................... 124 Highly Endemic and Highly Endangered ................................ ........................ 125 Cr yptozoic Snake Diversity ................................ ................................ ................... 126 A New Species of Centipede Snake (genus Tantilla ) from La Liberacin ...... 127 A Large New Species of Bli ndsnake ( Typhlops tycherus ) .............................. 128 Geophis damiani at La Liberacin ................................ ................................ .. 130 Noteworthy Ninia ................................ ................................ ............................ 131 Unidentified Salamander Populations ................................ ................................ ... 133 4 THE CHORTIS BLOCK IS AN UNDERESTIMATED HOTSPOT OF AMPHIBIAN DIVERSITY AND ENDEMISM ................................ ......................... 166 Methods and Materials ................................ ................................ .......................... 1 72 Sampling and Sample Identification ................................ ............................... 172 Extraction, Amplification, and Seque ncing ................................ ..................... 173 Sequence Evaluation and Alignment ................................ .............................. 174

PAGE 11

11 Distance Based Barcode Metrics ................................ ................................ ... 175 Phylogenetic Analysis ................................ ................................ .................... 176 Species Delimitation and Candidate Species ................................ ................. 176 Results ................................ ................................ ................................ .................. 177 Broad Results for Distance based Analyses ................................ .................. 177 Identification of Potential Candidate Species of Anura ................................ ... 179 BLASTN Results for Unassigned Salamander Sequences ............................ 187 Distance Based Analyses of Caudata ................................ ............................ 187 Phylogenetic Anal ysis of Caudata ................................ ................................ .. 191 Discussion ................................ ................................ ................................ ............ 196 Candidates for Further Taxonomic Study Among Anurans ............................ 196 Incilius coccifer/ibarrai/porteri ................................ ................................ ... 197 Incilius coniferus ................................ ................................ ...................... 198 Rhaebo haematiticus ................................ ................................ ............... 200 Craugastor aurilegulus ................................ ................................ ............. 200 Craugastor laevissimus ................................ ................................ ............ 200 Craugastor sp. inquirenda 1 & 2 ................................ .............................. 201 Diasporus diastema ................................ ................................ ................. 201 Plectrohyla cf. guatemalensis ................................ ................................ .. 201 Ptychohyla hypomykter ................................ ................................ ............ 202 Ptychohyla spinipollex ................................ ................................ .............. 202 Smilisca baudinii ................................ ................................ ...................... 203 Leptodactylus fragilis ................................ ................................ ............... 204 Lithobates brownorum X forreri ................................ ................................ 204 Lithobates brownorum ................................ ................................ ............. 205 Lithobates forreri ................................ ................................ ...................... 205 Lithobates maculatus ................................ ................................ ............... 208 Lithobates taylori ................................ ................................ ...................... 209 Lithobates warszewitschii ................................ ................................ ......... 209 Pristimantis ridens ................................ ................................ .................... 209 Candidate Species and Allopat ric Populations of Salamanders ..................... 210 Bolitoglossa (Magnadigita) sp. inquirenda 1 ................................ ............ 210 Bolitoglossa (Magnadigita) oresbia / sp. inqu irenda 2 .............................. 211 Bolitoglossa (Magnadigita) celaque ................................ ......................... 214 Bolitoglossa (Magnadigita) conanti ................................ .......................... 214 Bolitoglossa (Magnadigita) porrasorum ................................ ................... 215 Bolitoglossa (Nanotriton) rufescens/nympha ................................ ............ 215 Nototriton b arbouri ................................ ................................ ................... 217 Nototriton sp. inquirenda 1 ................................ ................................ ....... 217 Nototriton sp. inquirenda 2 ................................ ................................ ....... 218 Nototriton sp. inquirenda 3 ................................ ................................ ....... 218 Nototriton lignicola / sp. inquirenda 4 ................................ ....................... 218 Nototriton limnospectator / sp. inquiren da 5 ................................ ............. 219 Oedipina kasios / sp. inquirenda 1 ................................ ........................... 219 Oedipina nica / sp. inquirenda 2 ................................ ............................... 220 Oedipina koehleri / sp. inquirenda 3 ................................ ......................... 220

PAGE 12

12 Oedipina gephyra ................................ ................................ ..................... 222 Amphibian Endemism and Conservation Prioritie s in the Chorts Block ............... 223 Iterative Taxonomic Approaches to Biodiversity Inventory ................................ .... 224 5 CRYPTIC DIVERSITY AND REVISIONARY SYST EMATICS OF CHORTS HIGHLAND MOSS SALAMANDERS (CAUDATA: PLETHODONTIDAE) ............. 250 Salamanders as Models for Evolutionary Study ................................ ................... 250 Sala mander Diversity in the Chorts Block ................................ ............................ 252 Methods and Materials ................................ ................................ .......................... 256 Sampling ................................ ................................ ................................ ........ 256 DNA Extraction, PCR Amplification, and Sequencing ................................ .... 257 Sequence Alignment and Model Selection ................................ ..................... 258 Sequence Analy ses ................................ ................................ ........................ 259 Bayesian Inference and Maximum Likelihood Phylogenetic Analyses ........... 260 Comparative Morphology ................................ ................................ ............... 260 Results ................................ ................................ ................................ .................. 261 Discussion ................................ ................................ ................................ ............ 266 DNA Barcode Identification of Chorts Highland Moss Salamander s ............. 266 Influence of Substitution Saturation in Cytochrome B Dataset ....................... 267 Phylogenetic Systematics and Candidate Species ................................ ......... 268 Systematics ................................ ................................ ................................ .......... 269 A Divergent New Lineage from Refugio de Vida Silvestre Texguat ............... 269 Nototriton tomamorum Townsend, Butler, Wilson, & Austin 2010a .......... 270 A New Species from the Sierra de Agalta ................................ ...................... 275 Noto triton picucha Townsend, Medina Flores, Murillo, and Austin 2011 275 Restriction of the taxon Nototriton barbouri (Schmidt, 1936) .......................... 281 Nototriton barbouri (Schmidt 1936) ................................ .......................... 282 Description of unassigned populations from the Cordillera Nombre de Dios .. 284 Nototr iton sp. A, sp. nov. ................................ ................................ .......... 284 Nototriton sp. B, sp. nov. ................................ ................................ .......... 285 Review of the Remaining Species of Nototriton from the Chorts Highlands .. 286 Nototriton brodiei Campbell & Smith 1998 ................................ ............... 287 Nototriton lignicola McCranie & Wilson 1997 ................................ ........... 287 Nototriton limnospectator McCranie, Wilson, & Polisar 1998 ................... 289 Nototriton stuarti Wake & Campbell 2000 ................................ ................ 290 6 INTEGRATING RESEARCH, EDUCATION, AND OUTREACH IN SUPPORT OF CONSERVATION IN THE CHORTS HIGHLANDS ................................ ........ 294 Taxonomic Inventories in Promotion of Education and Extension ........................ 295 Opportunities for Training and Education ................................ .............................. 29 8 Pilot Project: Parque Nacional Montaa de Yoro ................................ .................. 299 Concluding Statement ................................ ................................ ........................... 303 APPENDIX: TAXONOMIC REVIEW OF CAUDATA FROM THE CHORTS BLOCK .. 304

PAGE 13

13 REFERENCES ................................ ................................ ................................ ............ 326 BIOGRAPHICAL SKETCH ................................ ................................ .......................... 350

PAGE 14

14 LIST OF TABLES Table page 3 1 Summary of fieldwork undertaken in the Chorts Block, 2006 2011. ............... 135 3 2 Conservation status and physiographic distribution of the native non marine herpet ofauna of the Chorts Highlands ................................ ............................. 137 3 3 Composition of the Chorts Block herpetofauna ................................ .............. 159 3 4 Broad distributional patterns of herpetofaunal diversity in the Chorts Block ... 160 3 5 Endemic and conservation priority herpetofaunal d iversity from the Chorts Block ................................ ................................ ................................ ................. 160 3 6 Distribution by mountain ranges of the endemic amphibians and reptiles of the Chorts H ighlands ................................ ................................ ....................... 161 4 1 Voucher information for samples used in this study ................................ ......... 228 4 2 Average nucleotide compositions of various taxonomic groups ....................... 244 4 3 Potential candidate specie s identified through this study ................................ 245 4 4 Results of BLASTN searches of the NCBI databas e for 16S consensus sequences representing 10 populations of taxon omically unassigned salamanders ................................ ................................ ................................ ..... 249 5 1 Samples used in sequence divergence and phylogenetic analyses ................. 291 5 2 Models of nucleotide substitution chosen for phylogenetic analyses of Chorts Highland taxa using Akaike Information Criterion values ................................ 292 5 3 W ithin and between species sequence divergence (uncorrected p distance) for Chorts Highland moss salamanders ................................ ........................... 292 5 4 Morphological and morphometric comparison of species of Nototriton ............ 293

PAGE 15

15 LIST OF FIGURES Figure page 1 1 Map showing present da y location of the Chorts Block ................................ ..... 21 2 1 Political divisions of the Chorts Block ................................ ................................ 26 2 2 Tectonic plate reconstructions showing the relative position and movement of the Chorts Block during the Cen ozoic ................................ ............................... 30 2 3 Physiograph ic regions of the Chorts Block ................................ ........................ 35 2 4 M ap showing m ount ain ranges of the Chorts Block ................................ ........... 36 2 5 Mountai n ranges of the Chorts Block I ................................ ............................... 37 2 6 Mountain ranges of the Chorts Block II ................................ .............................. 39 2 7 Ecological asso ciations of the Chorts Block I ................................ .................... 52 2 8 Ecological assoc iations of the Chorts Block II ................................ ................... 58 2 9 Ecological associ ations of th e Chorts Highlands ................................ ............... 64 3 1 Map showing s ampling localities in the Chorts Block ................................ ........ 75 3 2 Sampling in the Chorts Block I ................................ ................................ ........... 84 3 3 S ampling in the Chorts Block II ................................ ................................ .......... 88 3 4 Sa mpling in the Chorts Block III ................................ ................................ ......... 92 3 5 S ampling in the Chorts Block IV ................................ ................................ ........ 98 3 6 Sampling in the Chorts Block V ................................ ................................ ....... 104 3 7 S ampling in the Chorts Block VI ................................ ................................ ...... 110 3 8 Sa mpling in the Chorts Block VII ................................ ................................ ..... 114 3 9 Exemplar paratypes and habitats from Parque Nacional Monta a de Yoro ..... 117 3 10 La Liberacin de Texguat ................................ ................................ ................ 121 3 11 Plectrohyla chrysopleura (Hylidae) from La Liberacin ................................ .... 123 3 12 New species of Tantilla (Colubridae) and Typhlops (Typhlopidae) ................... 129

PAGE 16

16 3 13 Noteworthy cryptozoic snakes ................................ ................................ .......... 132 4 1 Radial phylogram showing coverage of higher level taxonomic groups ........... 178 4 2 Maximum likelihood phylogram of COI data showing generic and subgeneric rela tionships of salamander s amples ................................ ............................... 180 4 3 COI (left) and 16S (right) neighbor joining trees for Bufonidae ......................... 182 4 4 COI (left) and 16S (right) neighbor joini ng trees for Craugastoridae, Eleutherodactylidae, Leptodactylidae, and Strabomantidae ............................. 183 4 5 COI (left) and 16S (right) neighbor joining trees for Hylidae ............................. 185 4 6 COI (left) and 16S (right) neighbor joining trees for Ranidae ........................... 186 4 7 COI (left) and 16S (right) neighbor joining trees for Bolitoglossa ...................... 188 4 8 COI (left) and 16S (right) neighbor joining trees for Cryptotriton Dendrotriton Nototriton and Oedipina ................................ ................................ ................... 190 4 9 Maximum likelihood phylo gram for the genus Bolitoglossa .............................. 192 4 10 Maximum likelihood phylogram for the genera Nototriton Oedipina Dendrotriton and Cryptotriton ................................ ................................ .......... 194 4 11 Candidate Species I: Bufonidae and Craugastoridae ................................ ....... 199 4 12 Candidate Species II: Eleutherodactylidae and Hylidae ................................ ... 203 4 13 Candidate Species III: Ranidae and Strabomantidae ................................ ....... 206 4 14 Candidate Species IV: Bolitoglossa ................................ ................................ .. 213 4 15 Candidate Sp ecies V: Bolitoglossa cf. porrasorum ................................ ........... 216 4 16 Candidate Species VI: Oedipina ................................ ................................ ....... 221 4 17 Candidate Species VII: Oedipina cf. gephyr a ( = O. petiola ) ............................. 221 5 1 Distribution of Nototriton in the Chorts Highlands ................................ ............ 255 5 2 Comparison of tochondrial genes used to delimit species boundaries in Nototriton ................................ ........................... 262 5 3 Bayesian phylograms showing discordance between combined mtDNA phylogenies due to saturation at the third codon posit ion for cytochrome b ..... 265 5 4 Nototriton tomamorum ................................ ................................ ...................... 271

PAGE 17

17 5 5 Nototriton picucha ................................ ................................ ............................. 277 5 6 Nototriton barbouri sensu stricto ................................ ................................ ....... 283 5 7 Nototriton sp. B ................................ ................................ ................................ 286 5 8 Nototriton lignicola ................................ ................................ ............................ 288 5 9 Nototriton limnospectator ................................ ................................ .................. 289 6 1 Print media coverage of the discovery of new endemic species during 2008 and 2009. ................................ ................................ ................................ ......... 297 6 2 Examples of public outreach and dissemination of res ults from taxonomic inventories ................................ ................................ ................................ ........ 300

PAGE 18

18 Abstract of Dissertation Presented to the Graduate School of the University of Flori da, in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy INTEGRATIVE TAXONOMY REVEALS THE CHORTS BLOCK OF CENTRAL AMERICA AS AN UNDERESTIMATED HOTSPOT OF AMPHIBIAN DIVERSITY By Josiah Harold Townsend December 2011 Chair: James D. Austin Major: Interdisciplinary Ecology Central America has a remarkably complex geological history has served as the biological dispersal route between North and South America and is the site of extensive in situ evolution Nucle ar Central America is recognized as a region of high biodiversity, and the eastern portion of Nuclear Central America (the Chorts Block) has largely been overlo oked as a biodiversity hotspot In this dissertation, I use an integrative systematic approach to examine evolutionary patterns in a group with high diversity of endemic species that is also of global conservation priority: amphibians. I carried out 17 expeditions to over 60 localities in the Chorts Block for this dissertation results from which a re synthesized with existing data on regional herpetofaunal diversity, distrib ution and conservation status The Chorts herpetofauna is characterized by a high degree of endemism (35% of all species are endemic) and equally high extinction risk ( 41% thre atened including 96% of endemic species ) Endemism is highest among salamande rs (86% of species are endemic) A total of 456 amphibian samples representing 52 species were sequenced for two genes (16S and COI) and analyzed using distance based and phyloge neti c methods. I i dentify at least 36 unnamed across eight families of amphibians, while confirming that four

PAGE 19

19 taxonomically uncertain salamander populations represented new allopatric populations of endangered species previously assumed to be s ingle site endemics. These results reveal a regional amphibian fauna, already recognized as being of g lobal conservation priority, with species diversity underestimated by 26% (47% in salamanders), highlighting the critical role molecular systematics plays in endangered species conservation and management. To directly address the systematics a poorly known group with multiple candidate species, I present an integrative taxonomic revision of the genus Nototriton using data from three genes, external morpholo gy, and osteology I restrict the taxon N. barbouri to populations from the Sierra de Sulaco, describe three new species from the Chorts Highlands and a review of the remaining regional species. In light of these findings, I present my vision for using s ystematic biology as the basis for integrating research, education, and outreach in support of biodiversity conservation in the Chorts Block.

PAGE 20

20 CHAPTER 1 INTRODUCTION The formation of the Central American land bridge, with subsequent interchange of previou sly isolated organisms of Laurasian and Gondwanian origin, has been a source of inquiry for biogeographers since the founding of the discipline (Wallace 1876; Stehli and Webb 1985). Central America has had a remarkably complex geological history, owing in tectonic plates: the Caribbean, Cocos, Nazca, North American, and South American plates ( Figure 1 1; Bird 2003). Central America has not only served as the dispersal route betw een North and South America, but extreme topographical and ecological heterogeneity has also fuelled significant in situ diversification, particularly associated with the disjunct highland areas of Nuclear Central America and southern Central America (Savage 1966, 1983; Wake 1987). Whereas Nuclear Central America has long been accepted as a region of high biodiversity, some observers have further recognized the western and eastern portions of this highland block as distinct biogeographic entit ies (Johnson 1989; Campbell 1999; Townsend 2006, 2009). Eastern Nuclear Central America corresponding to the Chorts Block tectonic formation ( Figure 1), has been shown to have a distinctive component of endemic biodiversity, particularly in amphibians an d reptiles (Wilson & Johnson 2010), however molecular characterization of evolutionary patterns of diversification in this region has been limited to a few studies of a restricted taxonomic breadth and broader geographic focus (e.g. Castoe et al. 2009).

PAGE 21

21 Figure 1 1. Map showing present day location of the Chorts Block. Relative present day positions of four major tectonic plates and selected elements of the Central American Isthmus are indicated, with the contemporary Chorts Block highlighted in green (map generated using ODSN Plate Tectonic Reconstruction Service 1 and modified by author ). My principal goal is to broadly initiate molecular systematics research and studies in evolutionary biogeography in the Chorts Block ( Figure 1 1) a distinctive bio geographic region with a complex and unique geomorphological history and highly variable contemporary ecophysiographic landscape This region has a largely unrealized biogeographic importance as the first fragment of proto Central America to contact the No rth American plate; furthermore, it is characterized by a high degree of in situ endemism and evolutionary diversification in its extant biota. As a group with the highest documented endemism and highest risk for extinction, amphibians and 1 http://www.odsn.de/odsn/services/paleomap/paleomap.html

PAGE 22

22 particularly sa lamanders, make excellent taxa upon which to base studies of systematics and evolution. I begin in Chapter 2 by providing a synthesis of relevant characteristics of the Chorts Block in order to support a working definition for use in this dissertation a nd beyond. I begin by summarizing the geological histo ry of the Chorts Block, providing a comprehensive account of the contemporary physiographic and ecological landscape, and finally present ing tem for characterizing Honduran animal habitats. In Chapter 3, I provide an assessment of the diversity, conservation status, and distribution of amphibians and reptiles in the Chorts Block. This assessment is based on over 12,560 person hours of fieldwo rk conducted since 2006 (over 2,577 person hours logged myself) and existing data on distribution and conservation status of amphibians and reptiles from the region. The Chorts Block herpetofauna is characterized by a high degree of endemism that is face d with an equally high degree of extinction risk. Thirty five percent of the Chorts Block herpetofauna is endemic and highest threat categories Critically Endangere d, Endangered, and Vulnerable ), including an alarming 96% of endemic species. Endemism is by far highest among the salamande rs (86% of species are endemic); w hi le 75% of salamander species are considered conservation priority by the IUCN (2011). T his figur e likely is underestimated significantly As a result of the fieldwork detailed in Chapter 3, a comprehensive sample set of taxa and localities was assembled for Chorts Block amphibians, with particularly good

PAGE 23

23 representation of salamanders (Caudata). In Chapter 4, take an integrative taxonomic approach involving genetic distance based analysis (i.e., DNA barcoding) and model based phylogenetic analysis to identify and characterize cryptic amphibian diversity in the Chorts Block A nalysis of 456 individua l samples representing 52 named species in eight families of amphibians indicate species (20 anurans and 16 salamanders) in the Chorts Block while confirming that four taxonomically uncertain salamander popu lations actually represented new allopatric populations of endangered species previously assumed to be single site endemics. These results reveal a regional amphibian fauna, already recognized as being of global conservation priority, to be underestimated by 26% (47% in salamanders) in terms of species diversity, highlighting the critical role molecular systematics plays in endangered species conservation and management. In Chapter 5, I further utilize an integrative approach to delineate and estimate spec ies level phylogenetic relationships among and endemic radiation of highly cryptic moss salamanders (genus Nototriton ) in the Chorts Highlands. Using data from three mitochond rial DNA loci I examine the utility of each gene in a distance based DNA barcod ing approach fo r determining species diversity. The evolutionary relationships of diagnosed lineages are inferred using model based phylogenetic analyse s. My results indicate that species level diversity in Nototriton is underestimated in the Chorts Bl ock Three undescribed species of Nototriton were identified: one from the Sierra de Agalta in Departamento de Olancho, and one each from Pico Bonito and Texguat in the Cordillera Nombre de Dios. Finally, the taxonomy of this clade of Nototriton is reviewed and revised, with a restriction of the taxon N. barbouri (Schmidt) to populations from the

PAGE 24

24 Sierra de Sulaco, formal description of the three new species, and a review of the remaining species. Finally, Chapter 6 presents my long term vision for scientific exploration, research, education, and outreach in support of herpetofaunal conservation in Honduras, the country containing the bulk of the Chorts Highlands and the majority of the Chorts e dissertation, reviewing the results within a context of the ir regional and global significance.

PAGE 25

25 CHAPTER 2 BIOGEOGRAPHIC DELIMITATION OF THE CHORTS BLOCK: GEOLOGICAL HISTORY, PHYSIOGRAPHY, AND ECOLOGICAL ASSOCIATIONS My enthusiasm for the topic of Cho rts Block biogeography grew from the realization that patterns of endemism observed over the course of my studies in Honduras and northern Nicaragua seemed to be largely a reflection of emerging patterns in tectonic plate dynamics that have shap ed the com plex geomorphological 2010a) in order to set this region apart, in recognition of its disti nctiveness from proximal regions. This region is geographically analogous to the Chorts Block, an allochthonous 1 geological formation that today forms the only modern continental portion of the Caribbean Tectonic Plate and the largest terrestrial segment of the contemporary Central American land bridge (Rogers 2003; Marshall 2007). The Chorts Block has a challengingly complex history, and has recently been the subject of increased focus, and sometimes contentious debate, within the geological research com munity (James 2007; Mann et al. 2007; Ortega Gutirrez et al. 2007; Silva Romo 2008; Morn Zenteno et al. 2009). Politically, the contemporary region I refer to as the Chorts Block includes all of the country of Honduras, the northern portion of El Salvad or, eastern Guatemala, and northern Nicaragua ( Figure 2 1). As a working delimitation of the biogeographic region, I consider the Chorts Block to be nearly concept of an Eastern Nuclear Central A merican biogeographic province. I include the 1 Allochthonous re f erencing an independent geological structure that has moved f rom its site of origin.

PAGE 26

26 Figure 2 1 Political divisions of the Chorts Block. C ountries are shaded differently and labeled in all capital letters, while departments ar e labeled in sentence case in a smaller font size.

PAGE 27

27 associated coastal plains, with the western extent of the Chorts Block at the edge of the Ro Motagua Valley (the eastern edge of the Polochic Motagua fault complex) to a north south line roughly running through Zacapa, Chiquimula, Concepcon Las Minas, and the Guatemalan El Salvador b order at the Pacific Coast It extends eastward to include all of Honduras and El Salvador, and s outh to a line roughly between Lago Xolotln (= Lago de Managua) and Lago Colcibolca (= Lago de Nicaragua) in northern Nicaragua ( Figure 2 1). Without includin Nuclear Central America is homologous with the highlands of the Chorts Block, referred to collectively as the Chorts Highlands (Marshall 2007) or the serrana (Carr 1950) An integrative definition of the Cho rts Block Biogeographic Province as presented here, combine s an ecological and biodiversity based framework as used to delineate Eastern Nuclear Central America, with that of a physiographically and tecto nically defined Chorts Block. Geological Histor y The history of the Chorts Block is in large part characterized by its eastward movement along a series of strike slip faults on the southern margin of the North American Plate ( Figure 2 2; Dengo 1969; Donnelly et al. 1990; Gordon 1992; Rogers et al. 200 7). As recently as the K T Boundary (65 million years before present [ mybp ] ; Figure 2 2), the Chorts Block was located somewhere south of modern south central Mxico, having moved along a west to east trajectory around 200 km into its current position as the principal surface area of the Central American land bridge and the modern territory of Honduras, El Salvador, eastern Guatemala and northern Nicaragua ( Figure 2 1; Rogers 2003; Ortega Gutirrez et al. 2007).

PAGE 28

28 Origin and Cenozoic d evelopment of the Chor ts Block. The Chorts Block represents the only exposed Precambrian and early Paleozoic continental crust on the contemporary Caribbean Plate (DeMets et al. 2007). The oldest exposed geological formation of the Chorts Block is Precambrian in age and of R odinian 2 derivation, originating during the Grenville orogeny (1 017 20 mybp or 1400 mybp, depending on dating methods) contemporaneously with the Appalachian and Adirondack mountains of eastern North America and the Llano Plateau of Texas and northeaste rn Mxico (Mandon 1996; Gordon et al. 2010). Today this ancient formation is visible as a 60 km long series of exposed outcrops along the Jocotn Ceiba Fault in the Departamento de Yoro, Honduras (Gordon et al. 2010). While it is generally accepted among geologists that the Chorts Block originated approximately 1,100 km west of its current position and became detached during the Eocene, sliding and rotating along the Motagua Polochic Fault Complex at the southern margin of the stationary North American Pl ate, there are al. 2007). The first hypothesis places the Chorts Block along the southwestern margin of the North American Plate and physically contiguous with Mxico an d is supported by the existence of similarly aged Precambrian and Paleozoic rock formations, potential aligned fault systems in present day Honduras and southwestern Mxico and geological evidence that the Chorts Block rotated 30 40 counterclockwise whi le sliding eastward ( Figure 2 2; Gose 1985, Silva Romo 2008). The second hypothesis 800 km south of Mxico with it moving northeastwardly while rotating 40 clockwise (Keppie & Moran Zenteno 2005). A 2 Rodinia was a paleo approximately 2,000 mybp and 750 mybp

PAGE 29

29 t hird, less accepted hypothesis holds that the Chorts Block has remained in virtually the same position relative to the North American Plate, and that structures and evidence to the contrary have essentially been misinterpreted (James 2007). The relativel y dramatic Cenozoic tectonic history of the Chorts Block was dominated by what can only be described as catastrophic, prolonged, and repeated volcanism as the block slid and rotated its way eastward. The beginning of this extended period was the mid Eocen e (approx imately 55 mybp), following initial detachment of the Chorts Block from its parent structure (Jordan et al. 2008). A second flare up took place during the mid Oligocene (around 40 mybp), and the third, and largest, flare up took place during the early to middle Miocene (Jordan et al. 2008). M iocene Ignimbrite Flare up and subsequent u plift. The Mesozoic history of the Chorts Block features an approximately 10 million year period of intense explosive volcanism along the margins of the Chorts Blo ck and Central American Volcanic Front, considered the second largest ignimbrite 1 event in the known geological history of Earth (Jordan et al. 2008). During the mid Miocene over 5,000 km 3 of ignimbrites up to 2,000 m thick were deposited on top of the low relief surface of the southern an d western Chorts Block and tens of thousands of square k ilometers were repeatedly covered in thick layers of ash (Williams & McBirney 1969; Rogers et al. 2002; Jordan et al. 2008). The most intense period of the ignimbrit e flare up lasted from around 20 mybp to 15 mybp, with activity ceasing approximately 10.5 mybp (Gordon & Muehlberger 1994). This period is well documented by a series of deep sea sediment cores from sites in the western Caribbean Sea (Jordan et al. 2008). The site of a fissure like volcano that was 3 Ignimbrite hard rock formed from the deposition of super heated pyroclastic flow

PAGE 30

30 Figure 2 2. Tectonic plate reconstructions showing the relative position and movement of the Chorts Block (shaded green) during the Cenozoic F ollow s the generally accepted tectonic model of Hay et al. (199 9; alternative models reviewed in Rogers et al. 2007) with b lack lines represent tectonic boundaries and gray shading delineates modern day shorelines and landmasses (map s generated using ODSN Plate Tectonic Reconstruction Service 2 and modified by author ) 4 http://www .odsn.de/odsn/services/paleomap/paleomap.html

PAGE 31

31 up is represented today by the Padre Miguel Group geological formation in southwestern Honduras and peripherally in El Salvador and Guatemala (Rogers 2003). Under these circumstances, it would seem unlikely that extant terrestrial organisms and ecosystems on the Chorts Block are survivors of this extreme volcanism, and more likely that the extant biota represent s the product of post volcani c colonization and diversification. Following the Miocene flare up an d up and until approximately 3.8 mybp, the Chorts Block went through a period of rapid uplift driven by the detachment and subsequent subduction of a portion of the Cocos Plate, which induced upwelling in the mantle that raised the Chorts Block up to 1,1 00 m (Rogers et al. 2002). The c ontemporary Chorts Block. The Chorts Block presently continues its eastward movement along the strike slip faults of the Motagua Polochic Fault Zone and Swan Island Fault Zone, interacting in the continental context with the Maya Block of the North American Plate to the north and the Chorotega Block to the south, albeit interrupted by the Nicaraguan Depression ( Figure 1 1, 2 2; Rogers 2003, Marshall 2007). Marshall (2007) defined 15 physiographic provinces in Central Amer ica, four of which (the Chorts Highlands, Chorts Volcanic Front, Chorts Fore Arc, and Mosquitia Coast Lowlands) are geomorphological associates of the Chorts Block. The Chorts Highlands Province consists of a large dissected plateau that forms the gr eater part of the Chorts Block and includes the majority of the territory of the countries of Honduras and El Salvador, as well as western Guatemala and northern Nicaragua (Marshall 2007). The Chorts Highlands Province is subdivided into four regions: th e Western Rifted Highlands, the Central Chorts Plateau, the Eastern

PAGE 32

32 Dissected Highland s and the Honduran Borderlands. The Chorts Volcanic Front Province is an active volcanic front that borders the southern margins of the Chorts Highlands Province, and includes two regions: the Guatemalan Cordillera, which borders the western margin of the Chorts Highlands Province; and the Salvadoran Cordillera, which borders the Median Trough, an elongate graben that extends along the boundary faults at the margin of the active Nicaraguan Volcanic Front (Marshall Pacific Ocean. The Chorts Fore Arc Province encompasses the Pacific coastal plain of the Chorts Volcanic Front Province, and is similarly subdivided into the Guatemalan Coastal Plain and Salvadoran Coastal Plain (Marshall 2007). The Mosquito Coast Lowlands Province is the wide alluvial pla in along the eastern Caribbean slope of Honduras and Nicaragua, a region also referred to as La Mosquitia The Mosquito Coast Lowlands Province is dominated by a massive paleo Coco/Patuca river delta built up during the glacial cycles of the Pliocene Pleis tocene (Marshall 2007). The aforementioned Chorts Highlands Province and its four constituent subregions are of princip al i nterest for this dissertation, and these subregions are described in detail below. The Western Rifted Highlands region of southeas tern Guatemala, southwestern Honduras, and northern El Salvador is a west to east oriented plateau, generally exceeding 1,000 m elevation, which is interrupted by a series of independent, north to south oriented rift valleys featuring flat, xeric valley fl oors (Marshall 2007). The rift valleys, or grabens, of the Western Rifted Highlands include the contemporary

PAGE 33

33 Comayagua Valley and Otoro Valley. This region corresponds to the Padre Miguel Group, a 1,000 2,000 m thick layer of mid Miocene ignimbrites laid d own during the super volcanic eruptions along the margin of the Chorts Highlands and Chorts Volcanic Front. Those super eruptions essentially reset the landscape, allowing the development of new meandering river drainages as the rift valleys began spread ing following the end of the Miocene ignimbrite flare up around 10.5 mybp (Gordon & Muehlberger 1994; Rogers et al. 2002). The Central Chorts Plateau region of the Honduran interior represents the most tectonically stable portion of the Chorts Highland s, forming an essentially level plateau with little dissection or embedding by rivers (Marshall 2007). The Central Chorts Plateau lies atop of Paleozoic bedrock overlain with layered Cretaceous aged sedimentary deposits (Rogers et al. 2002; Marshall 2007) The Eastern Dissected Highlands region of eastern Honduras and Nicaragua includes the lower elevation, higher relief mountains bordering the Mosquito Coast Lowlands, and features three large, deeply embedded river drainages that drain a large portion of the Chorts Highlands Province (Marshall 2007). The Honduran Borderlands region lies along the northern margin of the Chorts Highlands, and is characterized by five major west to east trending faults that border major mountain ranges, including the Cordi llera Nombre de Dios in northern Honduras (Rogers 2003, Marshall 2007). One large graben valley, the Sula Graben, extends north to south from the Caribbean coast to the north end of Lago de Yojoa, and today contains the lower courses of two of the largest watersheds in the Chorts Block: the Ro Chamelecn and the Ro Ulua.

PAGE 34

34 Contemporary Ecophysiography Carr (1950), in his pioneering classification of Honduran ecological associations, recognized three princip al ecophysiographic components that make up the Ch orts Block: the Caribbean versant lowlands, the Pacific versant lowlands, and the mountainous interior region known as the serrana ( Figure 2 ecophysiographic regions correspond well with the geologically based physiographic provinces of Marsha ll (2007 ; described in the previous section ), with the serrana being Subsequently, from this point forward I use the name serrana Carr (1950) fu rther subdivided the Chorts Highlands into the Northern Cordillera, the Southern Cordillera, and the Pacific Colinas. This arrangement was also used by Wilson & Meyer (1985), McCranie & Wilson (2002), and others. Meja Ordez & House (2002) introduced a modified arrangement, based on their comprehensive evaluation of the ecosystems of Honduras using the UNESCO system of Physiognomic Ecological Classification of Plant F ormations of the Earth (Mueller Dombois & Ellenberg 1974), which recognized a Cordillera del Norte Cordillera Central and Cordillera del Sur This arrangement is preferable and is used here as the basis for describing the ecophysiography of the Chorts Block, which I divide into three principal regions: the Caribbean Lowlands, the Pacific L owlands, and the Chorts Highlands, which itself is subsequently subdivided into the Northern, Central, and Southern Cordilleras. Northern Cordillera of the Serrana As first defined by Meja Ordez & House (2002) and expanded here, the Northern Cordil lera consists of the following mountain ranges and groups of ranges:

PAGE 35

35 Figure 2 3 P hysiographic regions of the Chorts Block A fter Carr (1950) The Cordillera (or Sierra) Nombre de Dios ( Figure 2 4) stretches west to east across the departments of Atl ntida, Coln, and Yoro, Honduras, and includes the cloud forest protected areas Refugio de Vida Silvestre (RVS) Texguat (maximum elevation 2,208 m) at the western end and Parque Nacional (PN) Pico Bonito (2,435 m) and PN Nombre de Dios (1,725 m) in the ce ntral portion, with a few scattered low peaks extending to the east, terminating with PN Capiro y Calentura (1,235 m) near Trujillo. Based on my preliminary observations, I consider the Sierra de Mico Quemado, a north to south oriented range in western Yor o, to be the western terminus of the Cordillera Nombre de Dios. This range, which includes Zona de Reserva Ecolgica (ZRE) Montaa de Mico Quemado y Las Guanchias, was considered part of the Central Cordillera by Meja Ordez & House (2002). The Sierra d e Omoa ( Figure 2 4) in the departments of Corts and Santa Brbara, Honduras, with the cloud forest protected area PN Cusuco (2,242 m), as well as the rea de Produccin de Agua Merendn (1,749 m). The Sierra de Espritu Santo ( Figure 2 4) in the departme nts of Copn and Santa Brbara, in Honduras and Izabal and Zacapa in Guatemala, which includes the cloud forest reserve PN Cerro Azul Copn (2,285 m), as well as unprotected highland forests at Ro Amarillo (1,479 m) in Copn, Honduras, and Cerro del Mono (1,653 m) in Zacapa, Guatemala. While these mountain ranges appear disjunct and isolated ( Figure 2 4), particularly with respect to the Sula Graben, their herpetofauna l composition and patterns of

PAGE 36

36 Figure 2 4 Map showing m ountain ranges of the Chort s Block G reen outlines correspond to the Northern Cordillera, blue to the Central Cordillera, red to the Southern Cordillera, and black to ranges extralimital to this study. 1 = Sierra Nombre de Dios, 2 = Sierra de Omoa, 3 = Sierra de Espritu Santo, 4 = Sierra de Joconal, 5 = Montaa de Santa Brbara, 6 = Montaas de Membar, 7 = Sierra de Montecillos, 8 = Sierra de Comayagua, 9 = Sierra de Sulaco, 10 = Cordillera de La Flor La Muralla, 11 = Sierra de Agalta, 12 = Sierra de Botaderos, 13 = Sierra Punta Pi edra, 14 = Montaas de Patuca, 15 = Sierra de Montecristo, 16 = Sierra del Merendn, 17 = Sierra de Celaque, 18 = Sierra de Erandique, 19 = Sierra de Puca Opalaca, 20 = Montaa de la Sierra, 21 = Sierra de Lepaterique, 22 = Sierra de Dipilto, 23 = Montaa de Coln, 24 = Cordillera Dariense, 25 = Salvadoran Cordillera, 26 = Cordillera de Las Marabios.

PAGE 37

37 Figure 2 5 Mountain ranges of the Chorts Block I. A) Cordillera Nombre de Dios, seen from offshore looking south; Cerro Bfalo is the tallest mountain on the left (east) side, and Pico Bonito is the sharp peak on the right (west) side. B) Sierra de Omoa, seen from near Buenos Aires de Baaderos looking east northeast. C) Southern slopes of Cerro Azul de Copn in the Sierra de Espritu Santo, seen from Queb rada Grande. D) Montaa de Santa Brbara, seen from the road west of Peas Blancas looking south southwest. E) Montaas de Membar, seen from the road south of Santa Elena looking south. F) Sierra de Comayagua, seen from road south of San Jernimo looking south. (Photos J.H. Townsend). distribution reflect a greater shared evolutionary history among these mountains than any of them share with other ranges (as evidenced in Chapter 4)

PAGE 38

38 Central Cordillera of the Serrana I follow Meja Ordez & House (20 02) in recognizing a Central Cordillera made up of the remaining Caribbean versant serrana The Sierra de Joconal (1,688 m; Figure 2 4) extends to the roughly west to east from eastern Departament o de Copn (municipalities Nueva Arcadia and San Nicols), across Departamento de Santa Brbara and into western Departamento de Corts (municipality of Villanueva). Montaa de Santa Brbara (2,744 m; Figure 2 4) is an isolated karstic massif rising from the southern terminus of the Ula Chamelecn Plain to the west of Lago de Yojoa, its unique ecological communities wholly contained within the boundaries of PN Montaa de Santa Brbara. T h e Montaas de Membar (2,080 m; Figure 2 4), also called the Monta as de Yule, are a rugged set of peaks on the eastern side of Lago de Yojoa on the border between the departments of Corts and Comayagua, primarily contained within PN Cerro Azul Membar. Meja Ordez & House (2002) apparently included this group of moun tai ns in the Sierra de Montecillos; however I consider it to be a separate formation. The Sierra de Montecillos ( Figure 2 4) is one of two roughly parallel mountain ranges that are oriented northwest southeast and form the margins of the mountain range, which straddles the border between the departments of Comayagua and Intibuc, make up RB Montecillos (2,459 m). The Sierra de Comayagua ( Figure 2 4) runs roughly parallel to th e Sierra de Montecillos, separated by the dry intermontane Comayagua Valley, and extends over 130 km north to south along the border between the departments of Comayagua and Francisco Morazn. The highest portions of this range are found within PN Montaa de Comayagua (2,407 m) and RVS Corralitos (2,117 m). The Sierra de Sulaco ( Figure 2 4) runs roughly west to east in the southwestern part of the Departamento de Yoro, and includes the highlands of PN Pico Pijol (2,282 m) at the western end of the range and Montaa Macuzal (1,945 m) at the eastern end. The Cordillera de La Flor La Muralla ( Figure 2 4) stretches across northern Departamento de Francisco Morazn, southern Departamento de Yoro, and into western Departamento de Olancho, with highland forest pro tected areas PN Montaa de Yoro (2,245 m), RVS La Muralla (2,064 m), Reserva Forestal Anthropolgica (RFA) Montaa de la Flor (1,637 m), RB El Cipresal (1,930 m), and RB Misoco (2,153 m).

PAGE 39

39 Figure 2 6 Mountain ranges of the Chorts Block II. A) Looking west along the spine of the Sierra de Sulaco, taken from the top of Montaa Macuzal, with Pico Pijol being the largest peak in the distance. B) Southeastern reaches of the highest peak in the Sierra de Botaderos. C) The Sierra de Celaque viewed from the ea st, with the tallest peak in the Chorts Block, Cerro de la Minas, visible as the peak in the middle of the photograph. (Photos J.H. Townsend).

PAGE 40

40 The Sierra de Agalta ( Figure 2 4) in central Olancho is a long, relatively narrow and steep range that include s, from west to east, the protected areas Monumento Natural Boquern (1,261 m), PN Sierra de Agalta (2,335 m), Reserva Anthropolgica El Carbn (1,817 m), and PN Sierra de Ro Tinto (1,925 m). The Sierra de Botaderos ( Figure 2 4) is located in northern Ol ancho along the border with the department of Coln, and includes Cerro Ulloa (1,735 m) and Cerro Azul (1,433 m) within the highland reserve PN Montaa de Botaderos, as well as the lower mountains of the Sierra de La Esperanza. The Sierra Punta Piedra ( Fig ure 2 4) is a relatively low elevation range in the departments of Coln and Gracias a Dios, and includes Montaa Punta Piedra (1,500 m), Cerro Antlope (1,075 m), Cerro Mirador (1,200 m), and Cerro Baltimr (1,082 m). These mountains are found in Reserva de Hombre y la Biosfera Ro Pltano. The Montaas de Patuca (1,155 m; Figure 2 4) in Olancho are located between the Ro Guayape, which flows directly southwest to meet the Ro Guayambre and form the Ro Patuca, and a lower course of the Ro Patuca that f lows northeast to the Caribbean Sea. The southeastern portion of this range is found within PN Patuca. Southern Cordillera of the Serrana The Southern Cordillera is a geomorphologically linked series of mountain ranges extending from the vicinity of the El Salvador Guatemala Honduras border region east southeast into northern Nicaragua. The Sierra de Montecristo ( Figure 2 4) has its highest elevations at the point where El Salvador, Guatemala, and Honduras meet, in a tri nationally managed protected are a called Montecristo Trifinio (2,419 m). Most of this range is found in Guatemala, where it extends northward into the department of Chiquimula. The Sierra del Merendn ( Figure 2 4) is a north south oriented range that extends from Guatemala (Chiquimula, Zacapa) across Honduras (Copn, Ocotepeque, Lempira) and into El Salvador (Chalatenango), and includes the following cloud forest areas: RVS Erapuca (2,380 m), Cerro El Pital (2,730 m), Cerro Sumpul (2,167 m), and Reserva Biolgica (RB) Gisayote (2,310 m) The Sierra de Celaque ( Figure 2 4) is a north south oriented range located in Lempira and easternmost Ocotepeque, and contains the highest elevations in the Chorts Highlands in PN Celaque (including peaks of 2,849 m, 2,825 m, and 2,804 m elevation) and RB Volcn Pacayita (2,516 m).

PAGE 41

41 The Sierra de Erandique (2,134 m; Figure 2 4) is a north south oriented range in southeastern Lempira, extending from the municipality of La Campa at the northern end to the municipality of Piraera in the south. The Sierra de Puca Opalaca ( Figure 2 4) is located in Intibuc, northeastern Lempira, and extreme southern Santa Brbara, and includes cloud forest areas found in RC Cordillera de Opalaca (2,390 m), RVS Puca (2,234 m), RVS Mixcure (2,312 m), and RVS Montana Verde (2, 127 m). The Montaa de la Sierra ( Figure 2 4) is found in the department of La Paz and extreme southern Intibuc, and includes a number of peaks and high plateaus, including a number within RB Guajiquiro (2,265 m), RB El Chiflador (1,811 m), RB El Pacayal (1,955 m), RB Mogola (1,648 m), RB Sabanetas (2,047 m), RB San Pablo (1,741 m), and RB San Pedro (1,719 m). The Sierra de Lepaterique ( Figure 2 4) is the roughly U shaped range that circles the southern side of the upper Choluteca valley, which is also t he valley containing the Honduran capital, Tegucigalpa. This range includes PN La Tigra (2,290 m), RB Yerba Buena (2,243 m), RB Cerro Uyuca (2,006 m), RB El Chile (2,190 m), and RB Monserrat Yuscarn (1,825 m). The Sierra de Dipilto ( Figure 2 4) extends ov er 300 km west to east from PN La Botija (1,710 m) in Choluteca, to the Cordillera Entre Ros in PN Patuca, straddling the Honduras Nicaragua border and including Reserva Natural (RN) Cerro Mogotn (2,106 m), the highest point in Nicaragua. The Montaa d e Coln ( Figure 2 4) is a low (maximum elevation 941 m), isolated karstic range located in southeastern Olancho and adjacent Gracias a Dios, and is found primarily within the Reserva de Biosfera Tawahka Asangni. The Cordillera Dariense ( Figure 2 4) is a co llection of cloud forested peaks and highland areas in northern Nicaragua, in the departments of Jinotega, Matagalpa, and Regin Autnoma Atlntico Norte, including Reserva Natural (RN) Apante (1,442 m), RN Cerro Musn (1,438 m), RN Dantal El Diablo (1,68 0 m), RN Kilamb (1,755 m), RN Peas Blancas (1,744 m), RN Saslaya (1,658 m), and RN Volcn Yali (1,709 m). Intermontane Valleys and Plains The Chorts Highlands can be characterized as well for its valleys as it can be for its mountains; the isolated mo untains form isolated areas of hot, dry habitat. Aspects of the physiography, ecological associations, and biogeography of these subhumid valleys were the subject of study by S tuart (1954),

PAGE 42

42 Johannessen ( 1963 ), Wilson & McCranie (1998), Sasa & Bolaos (2004), and Townsend & Wilson (2010b). The Middle Motagua Valley is among the driest areas in Central America, along with the Middle Agun Valley in Honduras. This valley lies betwe en the Sierra de las Minas (extralimital to the Chorts Highlands) and the Sierra Espritu Santo. The Sula Valley in northwestern Honduras is formed from combined drainages of two large watersheds, the Ro Chamelecn and Ro Ula, which have courses that flow closely together in their lower reaches into the Caribbean Sea. The Sula Valley is a north to south oriented graben valley that has been spreading since the late Miocene. The Otoro Valley is a moderately high elevation subhumid graben valley (lowest elevations 500 600 m) lying in a narrow upper portion of the Sula Valley, possessing an ecological character distinctive from that of the broader middle Sula Valley. The Comayagua Valley is a relatively high subhumid graben valley (lowest elevations 580 68 0 m) that forms a principal portion of the Honduran Depression, lying between the Sierra de Montecillos to the west and the Sierra de Comayagua to the east. This valley, like the Otoro Valley to the west, is actually a narrow upper portion of the Sula Vall ey, distinctive enough in character to warrant recognition. The Middle Agun Valley is a west to east oriented fault valley that lies in the rain shadow of the Cordillera Nombre de Dios in Yoro and is one of the driest areas in the Chorts Block. The Sir ia Talanga Valley is a high plain (lowest elevations 620 720 m) in central Francisco Morazn that is the headwaters of two of the largest watersheds in the Chorts Highlands, the Ro Guayambre/Ro Patuca and the Ro Ula. The Olancho Valley (or Guayape Gu ayambre Valley ) is a large valley in central Olancho surrounded by several mountain ranges, including the Sierra de Agalta, Cordillera de La Flor La Muralla, and Montaas de Patuca, to form the headwaters of the Ro Patuca. The Agalta Valley also referre (Wilson & McCranie 1998, McCranie & Wilson 2002, Townsend & Wilson 2010b), is found between the Sierra de Botaderos and Sierra de Agalta in central Olancho. The lowest elevations of the relatively high su bhumid intermontane valley, formed by the Ro Grande (a river whose name changes to Ro Sico, Tinto, and Negro downstream), are 550 650 m. The Middle Lempa Valley lies on a west to east orientation in central El Salvador between the Southern Cordillera of the Serrana and the Salvadoran Cordillera.

PAGE 43

43 The Upper Segovia Valley in Nicaragua is a subhumid region in the headwaters of the Ro Coco (the upper reaches are also called the Ro Segovia, and the lower river is called the Wangki by the indigenous Miskit u). The Choluteca Valley is the only major subhumid intermontane valley on the Pacific versant, and is initially oriented south to north from the headwaters of the Ro Choluteca in the Sierra de Lepaterique, before curving around the north side of the mo untains protected within PN La Tigra and turning south and then southwest on its path to the Pacific Ocean. The Meseta de La Esperanza is the highest plain in the Chorts Highlands, extending 12 km in length across central Intibuc at elevations ranging b etween 1800 and 2000 m. The Meseta de Siguatepeque is located in Comayagua between the Sierra de Comayagua, Sierra de Montecillos, and Montaas de Membar at around 1,100 m elevation. The Meseta de Santa Rosa is a wide plain on a plateau at about 1,100 m elevation in western Copn. Caribbean Lowlands Corresponding to the Mosquito Coast Lowlands Physiographic Province of Marshall (2007), the major ecophysiographic regions of the Caribbean lowlands include (McCranie & Wilson 2002, Wilson & Townsend 2006): the Motagua Plain (lower alluvial plain of the Ro Motagua, east of the river and northwest and west of the Sierra de Omoa and Sierra de Espritu Santo), the Ula Chamelecn Plain (large alluvial plain formed by Chamelecn and Ula rivers, which drain clos e to half of the physical territory of Honduras), the Nombre de Dios Piedmont (the narrow strip of coastal plain backed by the Cordillera Nombre de Dios), the Agun Negro Plain and the wide expanse of the Mosquitia (broad alluvial plain essentially lying between the Sico Paulaya watershed in Honduras and the Ro Grande de Matagalpa watershed in Nicaragua). Two climatic regimes are present in the Caribbean Lowlands (McCranie and Wilson 2002, Wilson & Townsend 2006): the Lowland Wet climate is found on the C aribbean coastal plain from sea level to about 600 m elevation, with mean annual

PAGE 44

44 precipitation exceeding 2000 mm and mean annual temperature exceeding 24C. Important protected areas for Caribbean Lowlands ecosystems include: Parque Nacional (PN) Cuyamel O moa, PN Jeannette Kawas, Refugio de Vida Silvestre (RVS) Cuero y Salado, PN Punto Izopo, Jardn Botnico Lancetilla, RVS Laguna de Guaymoreto, Reserva de Hombre y la Biosfera Ro Pltano, PN Patuca, PN Warunta, Reserva Biolgica (RB) Rus Rus, RB Laguna de Karataska, PN Ro Kruta, Reserva de Biosfera Tawahka Asangni, Reserva de Biosfera Bosawas, Reserva Natural (RN) Cabo Viejo Tela Sulumas, RN Laguna Bismuna Raya, and RN Laguna Pahara. Pacific Lowlands Corresponding to the Salvadoran Coastal Plain of the Chorts Fore Arc Physiographic Province of Marshall (2007), the Pacific ve rsant lowlands consist of a relatively broad coastal plain extending from the western to the southern limits of the Chorts Highlands province, becoming narrowest around the Gulf of Fonseca. These lowlands constitute a single ecophysiographic region with a relatively homogenized biota (Wilson & McCranie 1998; Sasa & Bolaos 2004; Townsend & Wilson 2010b). The Pacific Lowlands are subject to the Lowland Dry climate regime (Wilson & Mey er 1985), found from sea level to about 600 m elevation, with mean annual precipitation below 2000 mm and mean annual temperature exceeding 24C. Important protected areas for Pacific Lowlands ecosystems include: rea Protegida con Recursos Manejados Barra de Santiago, Parque Privada Walter T. Deininger, rea de Proteccin y Restauracin (APR) Nancuchiname, rea de Manejo Laguna El Jocotal, APR Conchagua, rea de Manejo de Habitt de Especie (AMHE) Baha de Chismuyo, AMHE Baha de San Lorenzo, AMHE Las Igua nas Punta Condega, AMHE Los

PAGE 45

45 Delgaditos, AMHE El Jicarito, AMHE La Berbera, AMHE San Bernardo, and RN Delta de Estero Real. Salvadoran Cordillera The Salvadoran Cordillera is not geomorphologically part of the the Chorts Highlands, instead constituting the Chorts Volcanic Front and being dominated by more recent Pliocene and Quaternary volcanic deposits (Marshall 2007). I include the Salvadoran Cordillera for the sake of completeness in this discussion given its position across the Pacific Lowlands of El Salvador and putative inclusion in the Eastern Nuclear Central America biogeographic province of Campbell (1999; as expanded upon by Townsend 2006). This west to east oriented range is a continuation of the Guatemalan Cordillera, and is made up of sever al dozen volcanic cones and peaks, including Santa Ana (2,365 m), San Vincente (2,182 m), and San Miguel (2,130 m). Cordillera Los Marabios Like the Salvadoran Cordillera, this range not geomorphologically part of the Chorts Highlands, and is represente d by a string of northwest to southeast oriented Quaternary (and in some cases active) volcanic cones arising from the Pacific Lowlands of northwestern Nicaragua. Volcanoes in this cordillera include Cosigina (858 m), San Cristbal (1,745 m), Casita (1,40 5 m), Telica (1,060 m), Cerro Negro (726 m), El Hoyo (1,079 m), and Momotombo (1,279 m). Watersheds Major river systems on the Caribbean versant of the Chorts Block include: Motagua (485 km), Chamelecn (200 km), Ula (300 km), Len (60 km), Agun (225 km), Sico or Tinto or Negro or Grande (215 km), Pltano (85 km), Sikre (70 km), Patuca (500 km), Warunta (85 km), Mocorn (92 km), Nac unta (65 km), Kruta (125 km), Coco or Segovia or Wangki (550 km), Wawa (160 km), Kukalaya (140 km), Prinzapolka

PAGE 46

46 (330 km), and Grande de Matagalpa or Awaltara (430 km). Major Pacific versant watersheds include Estero Real (137 km), Negro (85 km), Choluteca (250 km), Nacaome (90 km), Goascorn (115 km), Lempa (422 km) and Paz (134 km). Lakes and coastal lagoons There are very few large inland water bodies in the Chorts Block, the most notable of which is Lago de Yojoa (700 m elevation) in central Honduras. The two other large bodies of freshwater, Embalse El Cajn (285 m) in Honduras and Lago de Apans (970 m) in Nicaragua, a re both reservoirs created by hydroelectric dams. There are a number of large coastal lagoons and lagoon complexes on the Caribbean coast, including Los Micos, Guaymoreto, Ibans, Brus, Tilbalakan, Laguntara, Warunta, Tansn, Karataska, Kohunta, Bismuna, Pa hara, Karat, Huouhnta, and Laguna de Las Perlas. I consider Lago de Izabal in Guatemala and Lago Xolotln (= Lago de Managua) in Nicaragua extralimital to the Chorts Block and do not include them. Islands The principle islands associated with the Chort s Highlands include the Honduran Islas de la Baha (Utila, Roatn, Guanaja, and Cayos Cochinos), the Cayos Miskitos of Nicaragua, and Isla El Tigre and other small islands of the Gulf of Fonseca. Characterizing Ecological Associations: Holdridge Forest Fo rmations Forest formations described below follow the system developed by Holdridge (1967), as applied to Honduras in previous works (Meyer & Wilson 1971, 1973; Wilson & Meyer 1985; Wilson & McCranie 1998; Wilson et al. 2001; McCranie & Wilson 2002). The w idely used Holdridge (1967) system uses climatic, edaphic, and atmospheric conditions to define and determine the distribution of terrestrial ecosystems. The Chorts Block is typified by a wide range of climatic and elevational regimes, resulting in nine H oldridge forest formations being recognized within the region. I am using this system,

PAGE 47

47 described below, to partially define Chorts Block ecosystems, supplemented with other published reports, gray literature, and my own observations. Lowland Moist For est Commonly referred to as lowland rainforest, the Lowland Moist Forest (LMF) formation is defined by a high mean annual temperature (>24C), high mean annual precipitation (>2000 mm; with no month of the year having precipitation <50 mm), and by being f ound from sea level to about 600 m elevation. In the Chorts Block, the LMF formation is restricted to the Caribbean versant, the majority of remaining intact forest being found in the vast region of eastern Honduras and Nicaragua known as La Mosquitia In addition to lowland rainforest, the pine savannas in La Mosquitia open woodlands intersected by veins of gallery forest, are found within the Lowland Moist F o rest formation. Intact LMF is ch aracterized by having a heteroge neous canopy dominated by evergr een broadleaf trees regularly reaching 30 40 m in height (Agdelo C. 1987). Lowland Dry Forest The Lowland Dry Forest (LDF) formation includes habitat commonly referred to as scrub forest, and is defined by high mean annual temperature (>24C), moderate b ut seasonally variable annual precipitation (1000 2000 mm; with at least 3 4 months having precipitation <50 mm), and an elevational range of sea level to about 600 m elevation. In the Chorts Block, the LDF formation is found on the Pacific versant and in several interior valleys. Intact LDF is characterized by having a heterogeneous canopy dominated by deciduous trees typically reaching around 25 m in height (Agdelo C. 1987). Lowland Arid Forest. The Lowland Arid Forest (LAF) formation, commonly called thorn forest, is defined by high mean annual temperature (>24C), low annual

PAGE 48

48 precipitation (500 1000 mm; with at least 3 4 months having precipitation <50 mm), and an elevation range of sea level to approximately 600 m elevation. This formation has one of the most limited extents in the Chorts Block, known only from the Middle Agun Valley and the Upper Motagua Valley. Intact LAF is characterized by having a low, heterogeneous canopy dominated by deciduous trees typically reaching around 10 m in height, wi th vegetation dominated by xeric adapted plants such as cacti (Agdelo C. 1987). Premontane Wet Forest The Premontane Wet Forest (PWF) formation, sometimes called highland rainforest (McCranie & Wilson 2002), is defined by a moderate mean annual temperat ure (18 24C), high annual precipitation (>2000 mm), and an elevational range of approximately 600 to 1500 m elevation. The PWF formation bridges the LMF with higher elevation montane forests, and thus contains characteristics of both. Intact PWF is chara cterized by having a closed canopy dominated by evergreen broadleaf trees reaching 25 30 m in height, sometimes reaching 40 m (Agdelo C. 1987). Premontane Moist Forest The Premontane Moist Forest (PMF) formation, commonly referred to as upland pine oak forest, is defined by a moderate mean annual temperature (18 24C), moderate annual precipitation (1000 2000 mm), and an elevational range of about 600 to 1850 m elevation. The PMF formation is relatively widespread in the Chorts Highlands, particularly on interior slopes. Various habitat types are found within the PMF, and are defined below following the classification system of Carr (1950).

PAGE 49

49 Premontane Dry Forest The Premontane Dry Forest (PDF) formation, which can b is d efined by moderate mean annual temperature (18 24C), low annual precipitation (500 1000 mm), and an elevation range of approximately 600 to 1250 m elevation. This habitat is generally limited to the upper periphery of some xeric interior valleys that oth erwise support LDF or LAF, two formations with which PDF shares its typical characteristics. Lower Montane Wet Forest. The Lower Montane Wet Forest (LMWF) formation, is defined by having a low mean annual temperature (12 18C), high annual precipitation (>2000 mm), and an elevation range of approximately 1500 to 2700 m elevation (note: habitat typical of this formation can also occur at lower elevations, particularly in the Cordillera Nombre de Dios). In the Chorts Highlands, LMWF is primarily distribute d on the Caribbean versant, being replaced by Lower Montane Moist Forest (below) in the somewhat drier Pacific versant highlands. Intact LMWF is characterized by having a closed canopy dominated by evergreen broadleaf trees reaching 50 m in height (Agdelo C. 1987). Lower Montane Moist Forest The Lower Montane Moist Forest (LMMF) formation, also referred to as cloud forest or montane forest, is defined by having a low mean annual temperature (12 18C), moderate annual precipitation (1000 2000 mm), and an elevation range of approximately 1500 to 2700 m elevation. In the Chorts Highlands, LMMF is distributed in highland areas, typically on the Pacific versant and on the leeward slopes of some of the interior most Caribbean versant peaks. The LMMF formation contains both pine and broadleaf dominated habitats (better characterized using the classification system of Carr 1950).

PAGE 50

50 Montane Rainforest The Montane Rainforest (MRF) formation is defined by having a very low mean annual temperature (6 12C), high an nual precipitation (>2000 mm), and an elevation range of above approximately 2700 m elevation. This formation is the most geographically limited formation in the Chorts Highlands, being restricted to the highest slopes of Cerro Celaque (Honduras; maximum elevation 2,849 m), Cerro Santa Brbara (Honduras; maximum elevation 2,744 m), and Cerro El Pital (El Salvador and Honduras; maximum elevation 2,730 m). Characterizing Ecological Associations: Updating and Operationalizing the Carr (1950) System for Class ifying Honduran Ecosystems Based on four years of first hand observation made during his time as a professor at Escuela Agrcola Panamericana (Zamorano), Carr (1950) presented a preliminary erest was in k was system for classifying animal habitats in Honduras and the greater Chorts Bl ock. As opposed to the Holdridge (1967) system, the Carr (1950) system is descriptive and is designed with the intent of being modified and contextualized by specialists to fit their particular study system or taxonomic group. I developed such an operation al system, which I will refer to as the Carr Classification System for Honduran Ecological Associations or simply the Carr System, based on a synthesis of the available literature

PAGE 51

51 and my own observations from 1999 2011. This system is discussed below, and the habitat names defined here are used throughout this dissertation. Lowland associated H abitats Selva or Lowland Broadleaf Rainforest Found primarily within the Lowland Moist Forest (LMF) formation in the Caribbean Lowlands, selva is characterized by a tall, multi layered, closed canopy that is dominated by evergreen broadleaf trees, with upper canopy trees normally reaching 30 40 m (Agdelo C. 1987, Meja Ordez & House 2002) and reaching 60 m in some areas (Carr 1950, Wilson & Meyer 1985). Large expan ses of selva are found within Reserva de Hombre y la Biosfera Ro Pltano, PN Patuca, PN Warunta, Reserva Biolgica (RB) Rus Rus, Reserva de Biosfera Tawahka Asangni, and Reserva de Biosfera Bosawas. Meja Ordez & House (2002) listed the following tree s pecies as typical of Honduran selva : Brosimun alicastrum, Bursera simarouba, Calophyllum brasiliense, Cedrela odorata, Coccoloba anisophylla, Cordia alliodora, Ficus colubrinae, Ficus insipida, Ficus tonduzii, Guarea grandifolia, Hernandia stenura, Licani a platypus, Luehea candida, Nectandra sp. Ocroma pyranidale, Pithecoellobium donnel smithii, Pouteria campechiana, Pouteria sapota, Rinorea guatemalensis, Symphonia globulifera, Swietenia macrophilla, Tabebuia chrysantha, Terminalia amazonia, Virola koshn y i and Vochysia hondurensis ; the relatively open understory is made up of palms ( Acoelorrhaphe wrightii Chamaedorea spp., Bactris spp., and Geonoma spp.), woody plants ( Cespedesia macrophylla, Isertia haenkeana, Piper spp., Cephaelis spp., Psychotria spp. ), and herbaceous plants ( Adiantum spp., Polypodium spp., Begonia spp., Selaginella spp., Philodendron spp., and Syngonium spp.).

PAGE 52

52 Figure 2 7 Ecological associations of the Chorts Block I. A) Selva or Lowland Broadleaf Forest; Ro Tapalws, Reserva Bi olgica Rus Rus, Depto. Gracias a Dios, 180 m; Lowland Moist Forest formation. B) Mosquitia Pine Savanna; between Rus Rus and Awasbila, with the Montaas de Coln in the background, Depto. Gracias a Dios, 200 m; Lowland Moist Forest formation. C) Coastal S crub; Caribbean coast near Kaukira, Depto. Gracias a Dios; Lowland Moist Forest formation. D) Mangrove Swamp, near mouth of the Ro Kruta, Depto. Gracias a Dios; Lowland Moist Forest formation. E) Seasonal Deciduous Forest; near Teocintecito, Depto. Olanch o, 690 m; Premontane Dry Forest formation. F) Freshwater Swamp in Seasonal Deciduous Forest; seepage bog in the upper Valle de Agalta, northeast of Saguay, Depto. Olancho, 570 m; Premontane Dry Forest formation. Photos J.H. Townsend

PAGE 53

53 Mosquitia Pine Savan nas Although wholly classified as a rainforest area by Wilson & Townsend (2006) due to being found within the Lowland Moist Forest formation, the Mosquitia of eastern Honduras and Nicaragua supports large areas of Pinus caribaea savanna that bear a strong er resemblance to subhumid ecosystems than to rainforests (Parsons 1955, Zamora Villalobos 2000, Townsend & Wilson 2010b). The Mosquitia pine savannas resemble an open woodland dominated by P. caribaea with a mix of broadleaf trees and shrubs including Ag arista mexicana var pinetorum, Amaioua corymbosa, Arthrostemma ciliatum, Arundinella deppeana, Byrsonima crassifolia, B. verbasifolia, Calea integrifolia, Cecropia peltata, Cephaelis tomentosa, Chamaecrista nictitans, Clethra calocephala, Clidemia sericea, Cococypsellum sp., Cuphea pinetorum, Davilla kunthii, Guazuma ulmifolia, Gnaphalium semiamplexicaule, Lasianthaeas fruticosa, Lobelia laxiflora, Miconia albicans, M. glaberrima, Myrica cerifera, Psychotria suerrensis, Quercus oleoides, Salvia sp. Vernoni a agyropappa, Vigna vexillata and Xylopia frutescens, with an open understory of fire tolerant grasses (Poaceae) and sedges (Cyperaceae), particularly Paspalum pectinatum, Blechnum serrulatum, Rhynchospora rugosa, Rhynchospora bulbosa, Scleria cyperina, a nd Setaria geniculata (Parsons 1955; Zamora Villalobos 2000; Meja Ordez & House 2002 ). Herpetofaunal diversity present in the Mosquitia pine savannas is almost completely congruent with that of the subhumid forests of the intermontane valleys (Townsend & Wilson 2010 b ), and phylogenetic analyses of subhumid specialized taxa support conspecific relationships among Pacific and pine savanna populations ( Incilius coccifer Mendelson et al. 2005; Porthidium ophryomegas Castoe et al. 2005). Townsend & Wilson ( 2010b: 702 ) presented two princip al and as yet untested,

PAGE 54

54 hypotheses for explaining the apparent continuing connectivity between the Pacific lowlands, subhumid intermontane valleys, and Mosquitia pina savannas: in Nicaragua between the southern end of the Nuclear Middle American highlands and Lago de Nicaragua, allowing for dispersal of subhumid species from the Pacific Lowlands of Honduras, El Salvador and Nicaragua to the pine savannas of the Nicaraguan Mosqui tia, which is contiguous with pine savannas extending to the northern coast of Honduras. 2) Utilization of open areas along large rivers (Patuca, Coco, and Grande de Matagalpa) that have their upper reaches in subhumid areas as routes for dispersal. These rivers originate in subhumid intermontane valleys in the Chorts Highlands, flowing through extensive areas of broadleaf rainforest and on through pine savannas of La Mosquitia. Secondary connectivity may also occur through coastal strand habitat, which c reate s a network among individual river drainages at or near their mouths. Broadleaf Swamp Forest These swamp forests are found along poorly drained margins and backwater areas of large rivers, with notable expanses of Broadleaf Swamp Forest found along the Ro Patuca in Reserva de Hombre y la Biosfera Ro Pltano as well as along Ro Kruta and in PN Jeannette Kawas. While riverine swamp forests are restricted to the LMF in the Caribbean Lowlands, there is also broadleaf swamp forest at the northern and southern ends of Lago de Yojoa (700 m elevation), at the lower edge of the Premontane Wet Forest (PWF) formation. Meja Ordez & House (2002) listed the following species as being found in permanently inundated broadleaf swamp forest: the trees Crias caul iflora, Pachira aquatica, Pterocarpus hayesii, and Pterocarpus officinalis and the palms Roystonea dunlapiana, R. regia var

PAGE 55

55 hondurensis, Acoelorrhaphe wrightii, and Desmoncus orthacantus The following are found in seasonally inundated broadleaf swamp for est: the trees Castilla elastica, Coccoloba sp., Combretum cacoucia, Symphonia globulifera, and Vochysia ferruginea ; and in the flooded forests at the northern end of Lago de Yojoa are found the tree Erythrina fusca and an understory including Calathea spp ., Costus spp., Heliconia spp., Hymenocallis litoralis, Maranta spp., Thalia geniculata, Smilax spp., Philodendron spp ., and Syngonium spp. Palm Swamp A variety of palm dominated swamp forests occur in coastal areas. Tique palm swamps are dominated by th e tique ( Acoelorrhaphe wrightii ), or paurotis palm, in association with Annona glabra, Chrysobalanus icaco, Coccoloba uvifera, Conocarpus erectus, Dalbergia ecastaphylla, Dalbergia monetaria, Davilla kunthii, Morinda citrifolia, Doliocarpus guianensis, Eug enia aeruginea, Henriettea succosa, Miconia glaberrima, Miconia albicans, Montrichardia arborescens, Myrmecophila wendlandii, Palicourea tripilla, Symphonia globulifera, Terminalia bucidoides, Thrinax parviflora, Tococa guianensis, Clidemia sericea, Acroco mia mexicana, Bursera simaruba, Casearia sylvestris, Chrysophyllum mexicanum, Cordia alliodora, C. curassavica, Hibiscus tiliaceus, and Ochroma pyramidala ( Meja Ordez & House 2002) Seasonally inundated mixed broadleaf palm swamps near the Ro Kruta and Cabo Gracias a Dios can have a 40 50 m high canopy dominated by the palm Roystonea dunlapiana and the understory palm Acoelorrhaphe wrightii as well as Mimosa schomburki, Psychotria spp., Alibertia edulis, Spondias mombim, Pachira aquatica, Desmoncus ort hacantus, Bactris sp., Ficus sp., Calophyllum brasiliense, Coccoloba schiedeana., Hirtella racemosa, Xylopia frutesce ns, Dialium guianensise,

PAGE 56

56 Virola koschnyi, Annona glabra, Grias integrifolia, Dalbergia ecastaphyllum, and Trophis racemosa In defining the Huiscoyol Swamp as a habitat, Carr (1950: 587) described thick stands of slender Bactris palms (called huiscoyol glass Bactris swamp. Coastal Scrub A heterogen e ous habitat association found above the beach line along the Caribbean coast, Islas de la Baha, Cayos Cochinos, and Cayos Miskitos, Coastal Scrub includes low coastal strand forest (typi cal plants include: Cannavalia maritima, C. rosea, Euphorbia buxifolia, Ipomoea pescaprae, Sesuvium portulacastrum, Sporobolus virginicus, Chrysobalanus icaco, Coccoloba uvifera, Citharexylum caudatum, Hybiscus tiliaceus and Phyllanthus acidus ) and its ass ociated grass covered dune system (typical ground cover includes Andropogon brevifolius, Aristida sp ., Eleocharis sp ., Eragrostis sp ., Fimbristylis spadicea, and Paspalum sp ). This association is found in relatively long, undisturbed extensions along esse ntially the entire Caribbean coast of the Chorts Highlands ( Meja Ordez & House 2002 ). Carr (1950) described the somewhat peculiar and specialized coastal scrub habitat found on Isla El Tigre in the Gulf of Fonseca and a few exposed hillsides facing the gulf as a distinctive habitat association: Sea Breeze Scrub Forest. While this forest lies within the Lowland Dry Forest (LDF) formation, the forest receives most of its moisture in the form of occult precipitation brought in on Pacific winds. Common plan t species include Bursera simarouba, Cresentia alata, Enterolobium saman, Spondias purpurea, Prosopis juliflora, Acacia spp. Heamatoxilon brasilett i and Zizyphum sp.

PAGE 57

57 Mangrove Swamp These estuarine swamp forests are found along both the Caribbean and Pac ific coasts. Mangrove swamp communities on both coasts typically include the mangroves Avicennia germinan s and Rhizophora mangle with Caribbean swamps including salt tolerant species such as Laguncularia racemosa Acrostichum aureum Cecropia spp., and Co ccoloba uvifera and Pacific mangroves similarly accompanied by Sesuvium portulacastrum, Sporobolus virginicus, Acrostichum aureum, Cecropia spp., Coccoloba uvifera, Conocarpus erectus, and Laguncularia racemosa ( Meja Ordez & House 2002). Habitats Share d Between L owlands and the Chorts Highlands Vegas and Gallery Forest This association is found on rich alluvial soils along stream and river courses, and in areas of low relief around the confluence of two streams (a vega 1950) habitat classes Dry Gullies and Fence Rows and Hondonadas in this category, recognizing that all of the constituent associations are essentially arteries of mesic habitat, often through comparatively xeric areas. Frequent plants of Caribbean Lowland vegas and gallery forests include Carapa guianensis, Hirtella racemosa, Xylopia frutescens, Dentropanax arboreus, Dialium guianense, Ficus sp. Licania platipus, Ochroma lagopus, Pterocarpus rohrii, Symphonia globulifera, Vochysia hondurensis, Schizolobiu m parahybum, Cecropia obtusifolia, Hyeronima alcornoides, Lacmellea panamensis, Prioria copaifera, Enterolobium schomburki Apeiba membranaceae, Casearia sylvestris, Cedrela macrophilla, Dendropanax arboreus, Vismia macrophylla, Xylopia frutescens, and Zue lania guidonia (Meja Ordez & House 2002). Carr (1950: 588) observed that these types of alluvial forest appeared to serve as

PAGE 58

58 Figure 2 8 Ecological associations of the Chorts Block II. A) Thorn Scrub Forest, upper Valle de Agun, northwest of Coyoles Central, Depto. Yoro, 250 m; Lowland Arid Forest formation. B) Pantano or Freshwater Marsh, northern end of Lago de Yojoa, with the Montaas de Membar in the backgr ound, Depto. Corts, 640 m. C) Infrequently burned Ocotal near Guaymas, Depto. Francisco Morazn, 1,450 m; Premontane Moist Forest formation. D) Frequently burned Ocotal Cerro de las Cruces, Depto. Olancho, 1,260 m; Premontane Moist Forest formation. E) Broadleaf Transitional Forest, La Liberacin, Refugio de Vida Silvestre Texguat, Depto. Atlntida, 1,030 m; Premontane Wet Forest formation. F) Mixed Transitional Forest, Montaa de Jacaleapa Depto. Olancho, 1,120 m; Premontane Wet Forest formation. Phot os J.H. Townsend

PAGE 59

59

PAGE 60

60 hts are still grounds generating biogeographic hypotheses testable with modern molecular methods. Seasonal Deciduous Forest Called Monsoon forest by Carr (1950) and commonly referred to as tropical dry forest, this habitat has a distribution limited to th e Lowland Dry Forest and Premontane Dry Forest formations in the intermontane valleys and on the Pacific Lowlands. Meja Ordez & House (2002) reported the following species as frequent in Seasonal Deciduous Forest: Enterolobium cyclocarpun, Bursera simar ouba, Ceiba pentandra, Cordia alliodora, Lysiloma auritum, Lysiloma seemanii, Samanea sama n, Swetenia macrophylla Cochlospermum vitifolium, Gyrocarpus americana, Apeiba membranacea, Alvaradoa amorphoides, Calycophylum candidissimum, Tabebuia neoch rysanta, Samanea saman, Spondias mombin, Lonchocarus minimiflorus, and Guazuma ulmifolia. Thorn Scrub Forest This low (<4 m in canopy height) habitat is d ominated by spine bearing shrub like trees such as Pachycereus sp ., Hylocereun spp ., Mammillaria spp and Op untia spp ., and dry tolerant shrubs and herbaceous plants like Ananas sp ., Argyreia esp eciosa, Cnidoscolus tubulosus, D igitaia insularis, Epidendrum xipheses, Evolvulus sp Gonolobus sp ., Acacia farnesiana, Albizzia neopoides, Combretum fruticosum, Diphysa ribinoides, Jacquinia macricarpa, Karwinskia calderonii, Lepidagastris alopecuroidea, Loeselia sp ., Melanthera nivea, Thouviinidium decandrum, and Watheria americana (Meja Ordez & House 2002). Thorn Scrub Forest is

PAGE 61

61 restricted to the Lowland Arid Forest and Lowland Dry Forest formations in the intermontane valleys of the Chorts Highlands. Pantano or Freshwater Marsh Freshwater marshes dominated by Typha domingensis, Phragmites australis, and/ or Thalia geniculata form an expansive habitat association in some areas, especially the wide alluvial plains of La Mosquitia (Carr 1950; Zamora Villalobos 2000; Meja Ordez & House 2002). Other grasses present include Andropogon brevifolius, Aristida sp., Eleocharis sp ., Eragrostis sp ., Fimbristylis spadicea y Paspalum sp (Meja Ordez & House 2002). In La Mosquitia, the vast pantanos have a similar character to that of the Florida Everglades, including being dotted with islands of pine ( Pinus caribaea ) or paurotis palms ( Acoelorrhaphe wrightii ) with extensiv e areas of habitat found around the Laguna de Karataska, the Ro Kruta, and the lower Ro Coco. Habitats of the Chorts Highlands Ocotal The ubiquitous Mesoamerican pine oak forests are found throughout moderate elevation areas of the serrana and are dom inated by ocote pine ( Pinus oocarpa ), with representation from P. pseudostrobus and other pines at higher elevations. Ocotales are extensively distributed in the serrana in the Premontane Moist Forest formation, and peripherally in the Premontane Dry Fore st and Lower Montane Moist Forest formations, roughly between 800 and 1,600 m elevation. Ocotal is subject to regular burning by humans, in some areas annually, and, therefore, the biotic composition is limited to species able to tolerate or escape frequen t fires. Besides pines, the various species of oaks ( Quercus spp ) and the hardwood trees and herbaceous plants Acacia farnesiana, Brahea salvadorensis, Byrsonima crassifolia, Clethra occidentalis, Myrica cerifera, Enterolobium cyclocarpun, Eritrina berter oana,

PAGE 62

62 Ficus spp. Lysiloma auritum, Mimosa tenuiflora, Psidum guianense, and Tabebuia chrysantha are typically found in ocotales (Meja Ordez & House 2002). Carr (1950) identified a number of subdivisions of the Ocotal habitat, including shaded ocotal p ark ocotal ocotal steppe and ocotal pedregal based on edaphic and climatic variation as well as burn frequency. Mixed Transitional Cloud Forest This transitional forest between ocotales and cloud forest, which Carr (1950) called High Ocotal associatio n is essentially a humid ocotal with higher concentrations of bromeliads and other epiphytes, as well as a denser and more diverse understory. High ocotales are typically found within the Premontane Moist Forest and Lower Montane Moist Forest formations, between around 1,000 1,500 m elevation on the Caribbean versant and 1,200 1,800 m on the Pacific versant. Trees of mixed transitional forest include the pines Pinus oocarpa, P. pseudostrobus, and P. tecunumanii, the hardwoods Arbutus xalapensis, Clethra ma crophylla, Ficus aurea, Heliocarpus apendiculatus, Oreopanax lachnocephalus, Oreopanax xalapensis, and Quercus cortesii and an understory that includes Buddleia americana, Conostegia sp ., Miconia sp ., Psychotria macrophylla, Vernonia arborescens, Calyptra nthes hondurensis, Lobelia laxiflora, Piper launosum, and Verbesina sp. (Meja Ordez & House 2002). In addition to High Ocotales Carr (1950) identified two other types of transitional forest, Pinabetales Diquidambales Liquidambales ), which I re cognize as distinctive localized associations within Pinus and Liquidambar Transitional Cloud Forest. Pinabetales are ridge line groves dominated by the pinabete pine ( Pinus pseudostrobus ) having epiphyte and understory communities similar to those of the High Ocotal association, but also incorporating representative

PAGE 63

63 from higher elevation hardwood forests. Pinabetales are typically found between 1200 1600 m within the Premontane Moist Forest and Lower Montane Moist Forest formations. Besides P. pseudostrob us, the pines P. maximinoi and P. tecunumanii can also be present, as might plants otherwise characteristic of both ocotales and mixed cloud forest. Liquidambales have a similar composition to Pinabetales but are dominated by sweet gums ( Liquidambar styra ciflua ) and are more typical of leeward slopes than exposed ridges. Broadleaf Transitional Cloud Forest Also referred to as Premontane rainforest, this high diversity forest blends diversity from both selva below and the cloud forest above. It is found p rimarily in the Premontane Wet Forest formation from between 500 m and 1,500 m in elevation, primarily on the windward slopes of mountains along the Caribbean versant. Broadleaf Cloud Forest Lower Montane Wet Forest (LMWF) and Lower Montane Moist Forest (LMMF) formations, characteristically found between around 1,500 2,300 m elevation on the Caribbean versant and 1,800 2,600 m elevation on the Pacific versant. This forest characteristically has a hi gh diversity of large canopy trees, with canopy heights regularly reaching 40 50 m. Typical vegetation of Broadleaf Cloud Forest in LMWF includes the trees Alnus arguta, A. jorullensis, Cornus sp Prunus sp ., Olmediella betschieriana, Abies guatemalensis, Taxus globosa, Podocarpus oleifolius Acalypha firmula, Bocona glauci folia, Cleyera theaeoides, Wein mannia pinnata, W. tuerckheimii, Daphnopsis strigillosa, Fuchsia paniculata, F. splendens, Hedyosmun mexicanum, Hoffmannia lineolata, Miconia glaberina, Que rcus cortesii, Q. lancifolia, Q. laurina,

PAGE 64

64 Figure 2 9 Ecological associations of the Chorts Highlands. A) Broadleaf Cloud Forest, above Quebrada Varsovia, Montaas de Membar, Depto. Comayagua, 1,620 m; Lower Montane Wet Forest formation. B) Broadleaf C loud Forest, canyon across top of Montaas de Santa Brbara, Depto. Santa Brbara, 2,190 m; Lower Montane Wet Forest formation. C) Broadleaf Cloud Forest, Montaa de Botaderos, Depto. Olancho, 1,715 m; Lower Montane Wet Forest formation. D) Mixed Cloud For est, Quebrada Cataguana, Montaas de Yoro, Depto. Francisco Morazn 1,860 m; Lower Montane Wet Forest formation. E) Mixed Cloud Forest, Sierra de Omoa, Depto. Corts 1,660 m; Lower Montane Wet Forest formation. F) Mixed Cloud Forest, Sierra de Celaque, D epto. Lempira 2,130 m; Lower Montane Moist Forest formation. G) Ro Arcagual in Mixed Cloud Forest, Sierra de Celaque, Depto. Lempira 2,580 m; Lower Montane Wet Forest formation. H) Hepatic Forest, Sierra de Celaque, Depto. Lempira 2,780 m; Montane Rain forest formation. I) Heather Wind Scrub, Montaa Macuzal, Depto. Yoro 1,730 m; Lower Montane Wet Forest formation. Photos J.H. Townsend

PAGE 65

65

PAGE 66

66 Rondeletia buddleioides, R. laniflora, Rubus eriocarpus, and Saurauia kegeliana the herbaceous plants Senecio j urgensenii, Smilax spinosa, Ternstroemia megaloptycha Begonia convallariodora, B. fusea, B. oaxacana, Cibotium regale, Deppea grandiflora, Lobelia nubicola, L. tatea, Parathesis hondurensis, and Peperomia spp., and the ferns Adiantum piretii, Asplenium ha rpeodes, A. olivaceum, A. pterocarpus, Blechnum lehmannii, and Elaphoglossum eximium (Meja Ordez & House 2002). From Broadleaf Cloud Forest in LMMF on the Pacific versant, Meja Ordez & House (2002) listed the following species as common in PN La Tigr a in the Sierra de Lepaterique: Mauria sessiflora, Ilex chiapensis, Ilex williamsii, Oreopanax xalapensis, Carpinus caroliniana var tropicalis, Weinmannia balbisina, Hieronyma guatemalensis, Hieronyma poasana, Quercus cortesii, Q. lanciflia, Q. laurrina, Q bumelioides, Homalium racemosum, Olmediella betschieriana, Calatola laevigata, Nectandra heydeana, Ocotea veraguensis, Phoebe helicterifolia, Magnolia hondurensis, Miconia argentea, Guarea pittieri, Trophis chorizantha, Ardisia paschalis, Chamaedorea pin natifrons, Clusia rosea, Lophosoria quadripionnata, and Cyathea mexicana. Mixed Cloud Forest Called Bosque mixto by Meja Valdivieso (2001), Mixed Cloud Forest is typically found within the Lower Montane Wet Forest and Lower Montane Moist Forest formation s, between above about 1,500 m elevation on the Caribbean versant and 1,800 m on the Pacific versant up to around 2,500 m on both versants. Trees of Mixed Cloud Forest include the pines Pinus pseudostrobus P. tecunumanii and P. ayacahuite with a high di versity of oaks ( Quercus brumeliodes, Q. cortesii, Q. rugosa, Q. sapotifolia, and Q. acutifolia ) and other hardwoods including

PAGE 67

67 Arbutus xalapensis, Bernoulia flamea, Brunellia mexicana, Clusia spp ., Cornus discifolia, Cyrilla racemiflora, Dendropanax arbore us, Dendropanax hondurensis, Hedyosmun mexicanum, Magnolia sp ., Li quidambar styraciflua, Myrica cerifera, Ocotea sp ., Oreopanax caspitatus, O. xalapensis, O. lachnocephalus, Picramnia teapensis, Symplocos vernicosa, Toxicodendron striatum, Viasmia baccifer a, and Weinmannia pinnata (Meja Ordez & House 2002). At elevations exceeding 2,300 m, particularly on Cerro Celaque and Montaa de Santa Brbara, Laurasian trees at the southernmost extent of their range, such as firs ( Abies guatemalensis ) and yews ( Tax us globosa ), are found syntopically with Gondwanan trees at their northern distributional limit (e.g. Podocarpus oleifolius ). Coniferous Cloud Forest This is a rarely encountered association that is characterized by essentially pure stands of pines ( Pinu s hartwegii, P. maximinoi and P. ayacahuite ) as well as other conifers ( Cupressus lusitanica and Taxus globosa ) and is recorded from only a few drier, open ridges above 2,400 m on Cerro Celaque and Montaa de Santa Brbara (Meja Valdivieso 2001). It is c alled Bosque de conferas by Meja Valdivieso (2001), who described these stands as appearing to be subject to natural fires every 3 5 years. I encountered Coniferous Cloud Forest meeting this description between 1900 2100 m elevation on the southeastern s ide of Montaa de Yoro and within this recently burned h igh pine forest collected herpe t o faunal species otherwise considered endemic to nearby Broadleaf Cloud Forest. Montane Mixed Forest The habitat of the uppermost reaches of LMWF (>2,600 m) and the Mo ntane Rainforest formation, Bosque mixto montano alto Meja Valdivieso (2001) is dominated by the primitive conifers Abies guatemalensis Taxus globosa and

PAGE 68

68 Podocarpus oleifolius the pines Pinus ayacahuite, P. hartwegii P. maximinoi, and P. tecunumanii and the broadleaf trees Alnus arguta, Cornus sp ., Prunus sp ., Olmediella betschieriana, Oreopanax lempirana, Acalypha firmula, Alnus jorullensis, Bocona glaucif olia, Cleyera theaeoides, Weinm annia pinnata, W. tuerckheimii, Daphnopsis strigillosa, Fuchsia p aniculata, F. splendens, Hedyosmun mexicanum, Hoffmannia lineolata, Miconia glaberina, Quercus cortesii, Q. lancifolia, Q. laurina, Rondeletia buddleioides, R. laniflora, Rubus eriocarpus, and Saurauia kegeliana (Meja Ordez & House 2002). This habitat i s limited to high elevation on Cerro Celaque, Cerro El Pital, and Montaa de Santa Brbara. Hepatic Forest A type of mountain top dwarf forest, Hepatic, or Mossy, Forest ( Bosque heptico o musgoso of Meja Valdivieso 2001), that appears to be restricted t o the wet upper slopes of the tallest peaks, including at least Cerro La Picucha (2,100 2,200 m) in the Sierra de Agalta, Cerro Celaque (2,700 m), and Cerro Jilinco (2,200 m) and Cerro Cusuco (1,990 m) in the Sierra de Omoa (Meja Valdivieso 2001, Townsend & Wilson 2008). The canopy does not exceed 10 m in height and is typically shorter, with trees taking on a twisted appearance. Nearly all available surface area is covered, even layered, in epiphytic plants and fungi, up to 50% of which can be liverworts ( Marchantiophyta; Meja Valdivieso 2001 ). In some cases, this luxuriant epiphytic community creates a living exoskeleton that remains long after the death and decay of the tree within. Hepatic Forest is often found in association with Heather Wind Scrub, a nd appears to be a transitional habitat between the exposed scrub and the cloud forest below.

PAGE 69

69 Heather Wind Scrub Found on exposed portions of the highest peaks, this is a wind swept association that Carr (1950) termed Pea Wind Scrub and is variously ref erred to as elfin forest or dwar f forest (Townsend & Wilson 2006, 2008 ), names that reflect the somehow mystic character of these mountain 582) own description of this habitat art fully captures this character: It is a seeming ly incongruous combination of dwarfed and twisted microphyllous and sclerophyllous trees and shrubs, Ericaceae, Myrtiaceae, Myrsinaceae, and the like, implying xeric conditions, but with an astounding array of mosses, filmy ferns, selaginellas, and similar delicate hygrophyllous epiphytes. Although at first glance this is an altogether ill assorted looking flora, the incongruity is only apparent, since each of the two floristic elements is in its own way adapted to withstand drastic reversals in its water e conomy. On these peas the wind blows almost constantly, often violently, and while it usually brings in abundance of moisture, it imposes a heavy penalty when the supply fails for even a short period. The wind pruned trees meet the situation by conservati on of their moisture, while their cryptogamic guests yield freely to desiccation, lapsing into dormancy almost on a moment's notice, and without permanent injury. As indicated by the name, this habitat is dominated by plants in the family Ericaceae with a 2 array of bromeliads and other epiphytes. This habitat association is known from Cerro La Picucha (2,200+ m) in the Sierra de Agalta, Cerro Azul Membar (1,950 m+), Cerro Cela que (various exposed ridges above 2,700 m), and Cerro Jilinco (2,240 m) and Cerro Cusuco (2,010 m) in the Sierra de Omoa (Hazlett 1980, Meja Valdivi eso 2001, Townsend & Wilson 2006 ). Heather Wind Scrub likely is found in at least small patches on the tops of most exposed peaks above around 1,900 m elevation. Elfin Forest This term is herein reserved for use to describe the unique forest association found on Cerro La Picucha in the highest portions of the Sierra de Agalta. Delineated as Bosque enano by Mej a Valdivieso (2001), this habitat superficially

PAGE 70

70 however, in place of Ericaceae, this true dwarf forest is made up of twisted, epiphyte covered, bonsai like versions of the trees Billia hippocastanum Podocarpus oleifolius and Pinus hartwegii with exposed ground covered in clubmosses ( Huperzia and Lycopodium ) and dense patches of ground dwelling tank bromeliads. Introduction to Biodiversity of the Chorts Block Given the wi de variation in physiographic and ecological attributes that are evident in the Chorts Block, it comes as little surprise that the region also supports a rich and diverse biota. While the Caribbean and Pacific lowlands have biotas that are typically chara cterized as being composed of relatively widespread species from both the west and south, the Chorts Highlands and associated piedmont is an area of considerable endemic biodiversity. While investigation of the endemic fauna of the Chorts Highlands has been limited compared with other areas of the Neotropics, particularly southern Central America, the dedicated work of a small group of specialists led to the documentation of endemism across a variety of taxonomic groups. Botanical data may support this better than other groups, with over 263 described endemic species found in Honduras alone (Nelson S. 2001, 2008). Areas of elevated plant endemism include mesic highland forests and xeric intermontane valleys of the Chorts Highlands, and, in particular, t he piedmont of the Cordillera Nombre de Dios (Nelson S. 2008). Despite the high degree of localized plant endemism seen across the Chorts Highlands, to date there has been no published analysis of biogeographic patterns among these taxa. Over 166 native f reshwater fishes have been documented from Honduras, including three described and at least six undescribed endemic species (Martin 1972, Matamoros et al. 2009, Matamoros & Schaefer 2010). Among terrestrial vertebrates, birds (one species;

PAGE 71

71 Monroe 1968) and mammals (three species; Goodwin 1942, Reid 2009) stand out in having very few described endemic species; however, at least for mammals, this low level of endemism is almost certainly an artifact of a lack of focused sampling in the molecular age, particul arly for small mammals, by systematic mammalogists in the Chorts Highlands. It is the herpetofauna, the amphibians and reptiles, which provide s the best opportunity for elucidating patterns of evolutionary diversification in the Chorts Block. Systematic herpetologists have been active in the Chorts Block since at least the early in Townsend & Wilson (2010 a ) I detailed the history of herpetological exploration and research in Honduras My own work in the Chorts Block began as a student of Dr. and had progressed to the point where, when I began this dissertation in 2006, I had identified that the Chorts Block, and particularly the isolated cloud forests of the Chorts Highlands, as being in serious need of intensive study using molecular based approaches. However, taking an approach that was both broad and comprehensive in terms of taxonomic coverage required that virtually all the areas of the Chorts Block, principally those that were known to support endemic species, wou ld be sampled or resampled. This sampling was carried out from 2006 2011, and the results are presented herein

PAGE 72

72 CHAPTER 3 TAXONOMIC DIVERSITY, DISTRIBUTIONAL PATTERNS, AND CONSERVATION STATUS OF THE CHORTS BLOCK HERPETOFAUNA The con stituent countries of t he Chort s Block (El Salvador, Guatemala, Honduras, and Nicaragua) each have their own rich histories of herpetolo gical investigation ( Mertens 1952, Stuart 1963, Meyer 1969, Villa 1972, Wilson et al. 2010). M ost research has expectedly taken a geopolitical ly delimited approach that is often arbitrary and is seldom congruent with biogeographic boundaries. While this approach is practical and largely necessary due to a variety of considerations (e.g., visas, research and export permits, logistics) it can als o come at the expense of a biogeographically meaningful approach to research or of a unified strategy for conservation in transboundary areas T he Chorts Block is an exemplar of this issue, where national boundaries effectively divide efforts to document and conserve isolated cloud forest areas in five mountain ranges that serve to physically delineate national borders In some cases, border regions are avoided due to security related issues. Honduran frontier zones have become increasingly favored by tr ansnational narcotics traffickers, particularly over the past decade as the influence of Mexican drug cartels has expanded into Central America (e.g., Archibold & Cave 2011). In an even more extreme case, the highest mountain range in Nicaragua (Sierra de Dipilto, maximum elevation 2,107 m), which also forms the border with Honduras, was considered a strategic vantage point that was heavily contested, and subsequently landmined, during the Contra United Nations Mine Action Serv ice 1998 ). Despite the apparent biogeographic importance of this mountain range and its potential for supporting endemic species, little to no biological inventory work has been carried out in the Sierra de Dipilto. Beyond this mountain range, a large area of the Chorts

PAGE 73

73 Highlands in northern Nicaragua has also been heavily undersampled as a result of being the principal zone of conflict in the Contra Sandinista War, and has only recently begun to be sampled in a concerted fashion (Sunyer et al. 2009, Trave rs et al. 2011). P olitical boundaries have placed the majority of the endemic rich Chorts Highlands within the borders of one country, Honduras, with significant extensions into three neighboring countries most notably Nicaragua. S ubsequently no concer ted attempt has been made to date to present a unified understanding of the biogeographic logical diversification A consequence of research constrained by political boundaries is that systematic cataloguing of regional diversity has been handicapped As a result, a number of endemic species are known from localities in one country but unconfirmed as occurring across the border in ecophysiographically contiguous areas, as in the cases, for example, of Cryptotriton monzoni (endemic to the Sierra de Espritu Santo in Guatemala) and Anolis johnmeyeri (endemic to the Sierra de Espritu Santo and Sierra de Omoa in Honduras). In at least one case, it has been suggested that two species of Cryptotriton described from opposite sides of the Sierra de Omoa ( called the Sierra de Caral in Guatemala), C. nasalis and C. wakei actually represent the same species (McCranie & Wilson 2002). Minimizing the constraints of these largely political and logistical issues is critical to accurately analy zi ng and interpret ing regional patterns of biogeography, endemism, and conservation priority. My goal in this chapter is to synthesize the available data for the Chorts Block herpetofauna drawing first from a considerable regional literature base This av ailable data are then augmented by the results of sampling efforts from 2006

PAGE 74

7 4 2011 in Honduras and Nicaragua, and presented to provide the basis from which to further investigate herpetofaunal diversity and assess research and conservation priorities moving forward. Methods and Materials This section serves not only to define the methodologies used in Chapter 2, but also to provide some of the broader methods and definitions used throughout this dissertation Field Sampling Between June 2006 and Ap ril 2011 I made 17 field trips totaling over 2,577 person hours of effort ( over 12,560 person hours were logged by expedition participants) over the course of 275 field days in Honduras and Nicaragua sampling over 60 localities in the Chorts Block (Table 3 1; Fi gure 3 1 ). Tissue samples were taken from freshly euthanized vouchers and stored in SED buffer (250 mM EDTA/20% DMSO/saturated NaCl ; Seutin et al. 1991; Williams 1997 ). Voucher specimens were preserved in 10% formalin solution and later transferred to 70% ethanol for permanent storage. Vouchers were deposited in the Florida Museum of Natural History, University of Florida (UF), the Museum of Vertebrate Zoology, University of California, Berkeley (MVZ), and the National Museum of Natural History, Smithsonian Institution (USNM). Taxonomic Scope and Standards I recognize a Chorts Block herpetofauna inclusive of species that occur south or east of the Ro Motagua and north of a latitudinal line across the northernmost edge of Lago Xolotln (= Lago de Managua). I have included taxa from the Islas de la Baha and Cayos Miskitos, as they are continental islands of Chorts Block origin, but not the

PAGE 75

75 Figure 3 1 Map showing s ampling localities in the Chorts Block R ed cr osses = localities sampled 2006 2011, blue cros ses = localities sampled 1999 2005, green diamonds = localities sampled by collaborators for inclusion here

PAGE 76

76 offshore islands of Belize, the smaller cays far offshore from eastern Honduras and Nicaragua, or the Islas del Cisne as they neither geolog ical nor biogeographically related directly to the Chorts Block I have excluded marine taxa (sea turtle families Cheloniidae and Dermochelyidae and the sea snake Pelamis platura ) and introduced species, as well as taxa known only from the Salvadoran Cord illera or southwestern Salvadoran coastal plain as they are not considered herein to be part of the Chorts Block. The taxonomic nomenclature used in this dissertation generally follows that used by Wilson & Johnson (2010). A number of additions and chan ges have occurred since that publication that pertain directly to composition of the Chorts Block herpetofauna, including description or resurrection of the following taxa : Bolitoglossa nympha (Campbell et al. 2010), Cerrophidion sp. (Jadin et al., submit ted), Ctenosaura praeocularis (Hasbn & Khler 2009), Anolis beckeri (Khler 2010), A. unilobatus (Khler & Vesely 2010), Epictia magnamaculata ( Adalsteinnsson et al. 2009), Dendrophidion clarkii (McCranie 2011 a ), Leptodeira rhombifera (Daza et al. 2009), Mastigodryas alternatus (McCranie 2011 a ), Omoadiphas cannula (McCranie & Cruz Daz 2010), Tantilla sp. (Townsend et al., submitted), and Tantilla psittaca (McCranie 2011 b ) In addition, four new species of salamanders have recently been described, in part based on results of this dissertation reported in later chapters, and are included in this chapter for the sake of completeness: Nototriton picucha (Townsend et al. 2011 a ), Oedipina koehleri (Sunyer et al. 2011), O. nica (Sunyer et al. 2010), and O. petiol a (McCranie & Townsend 2011). I follow Myers (2011) the species of the Rhadinaea godmani group as representing a separate genus, Rhadinella

PAGE 77

77 Evaluating Conservation Status I used three different measures to assess the conservation status of the herpetofau na of the Chorts Block : IUCN Red List categorization, Environmental Vulnerability Scores (Wilson & McCranie 2003), and Conservation Status Scores (Wilson & Townsend 2010). IUCN Red List categorizations were taken from one of three sources: the IUCN Red L ist of Thre atened Species (v. 2011.1; www. iucnredlist.org) for amphibians, marine turtles, crocodilians, and Ctenosaura ; Townsend & Wilson (2010) for Honduran reptiles, and Sunyer & Khler (2010) for Nicaraguan reptiles. Species not previously evaluated by the IUCN or other authors were assessed using the stan dard criteria of the IUCN (2001 ). Environmental Vulnerability Scores (EVS) are primarily from Townsend & Wilson (2010 a ). Species not assessed in that study were calculated using the methodology develop ed and refined by Wilson & McCranie (1992, 2003, 2004 a ), which is calculated 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 f or reptiles, and 3) extent of ecological distribution in Honduras; EVS scores from 10 13 indicate medium vulnerability, and scores from 14 19 are high vulnerability ( Wilson & McCranie 2003). Conservation Status Scores (CSS) were developed by Wilson & Town send (2010) to provide a simple measure for assessing conservation status of amphibi ans and reptiles across Mesoa merica. The CSS represents sum of individual scores for 1) numbers of countries, 2) physiographic regions, and 3) vegetation zones occupied by a given species of amphibia n or reptile. The country score range s from 1 to 8 ( the number

PAGE 78

78 of countries of Central American plus Mxico ) the physiographic region score ranges from 1 to 21 and the vegetation zone score from 1 to 15 (Wilson & Townsend 2010) Given this, the CSS can range from 3 (the most restricted endemic species, inhabiting a single vegetative zone in a single physiographic region in a single country) to the theoretical maximum of 44 (for a species found literally everywhere in Mxico and Central America). Results Composition of the Herpetofauna The native, non marine herpetofauna of the Chorts Block is comprised of 382 species in 134 genera and 41 families including 145 species of amphibians and 237 reptiles (Table s 3 2, 3 3). The Chort s Block herpetofauna comprises about approximately 2.3% of global herpetofauna l species diversity (16,301 species as of 6 October 2011; Table 3 3; AmphibiaWeb 2011, Uetz et al. 2011) This includes 2.1% of global amphibians, with 1% of caecilians, 7% of s alamanders, and 1.6% of anurans, along with 2.5% of global reptiles with 3.1% of turtles, 8.3% of crocodilians, and 2.5% of squamates ( 2.3% of lizards, and 4.2% of snakes). There are 41 families of amphibians and reptiles in the Chorts Block (Table 3 3), comprising 30.4% of the fami lies of the world and 65.1% of the families of Mesoamerica (Table 3 3). Within Mesoamerica (considere d in this discussion to include Mxico and Central America), the Chorts Block herpetofauna contains approximately 20.3% of t he regional herpetofauna (Wilson & Johnson 2010). This includes 19.8% of Mesoamerican amphibian species, with 12.5% of caecilians, 17.8% of salamanders, and 21.1% of amphibians, and 20.6% of reptiles, including 18.2% of turtles, two of three crocodilian sp ecies, and 20.6% of squamates (Table 3 3).

PAGE 79

79 Patterns of Distribution and Endemism within the Chorts Block Within the Chorts Block, species were considered to occur within one of eight physiographic regions: the Caribbean Lowlands, Caribbean Versant Inter montane Valleys, Northern Cordillera of the Chorts Highlands, Central Cordillera of the Chorts Highlands, Southern Cordillera of the Chorts Highlands, Pacific Lowlands, Pacific Versant Intermontane Valleys, and Islas de la Baha (Table s 3 2, 3 4 ). The C aribbean Lowlands holds the highest diversity, with 188 species, with the Northern Cordillera being the most diverse portion of the Chorts Highlands with 163 species (Table 3 4). The majority of amphibians (51%) that occur in the Chorts Block are endemi c, i.e., restricted in distribution to within the Chorts Block, with a lesser share of reptiles (24%) being endemic (Table 3 5 ). The salamanders have a particularly high degree of endemism, with 37 of 43 species (86%) being Chorts Block endemics, and 36 of those 37 being endemic to the Chorts Highlands province (Tables 3 2, 3 4) A large share of named anuran species (37%) is also endemic (Table 3 5 ). Distribution of Chorts Highland Endemics The Northern Cordillera is the most endemic rich p ortion of t he Chorts Highlands with 72 species, versus 42 and 35 in the Central and Southern Cordilleras, respectively (Table 3 6 ). The Sierra de Omoa is clearly the mountain range supporting the highest number of Chorts Block endemic species (35 ) and each of the four mountain ranges of the Northern Cordillera support more endemic species than any other mountain range in the Chorts Highlands (Table 3 6 ). In the Central Cordillera, the Sierra de Sulaco and Cordillera de La Flor La Muralla tie for the lead with 13 e ndemic species, with the Sierra de Agalta having 11 (Table 3 6 ). In the Southern Cordillera, the Sierra de l Merendn

PAGE 80

80 supports the most endemic species with 12, followed by the Cordillera Dariense with 9 (Table 3 6 ). Conservation Status A startling portion of the Chorts Block herpetofauna is threatened at the local, regional, and global levels (Table s 3 2 3 5 ). At least 41% of the entire herpetofauna is at endangered at a globally significant level (listed in one of the top three threat categories on the IUCN Red List: Critically Endangered, Endangered, or Vulnerable), including an alarming 74% of salamanders (Table 3 5 ). Although not surprising given their typically narrow geographic distributions, 96% of species endemic to the Chorts Block are also list ed in the top three IUCN Red List categories, with 48% of endemic species listed in the highest risk category: Critically Endangered (Table 3 5 ). Regionally, the Conservation Status Score (CSS) was used to gauge the relative degree of threat facing members of the Chorts Block herpetofauna. Two hundred and ten species (55% of Chorts Block herpetofauna) have CSS between 3 and 11, placing them in the Very High conservation s ignificance the highest risk category employed at the Mesoamerican sca le by Wilson & Townsend (2010). Within the Chorts Block, Environmental Vulnerability Score ( EVS ) was used to provide a local scale measure of the relative degree of threat facing herpetofaunal species. Results of the EVS indicated an elevated degree of su sceptibility to degradation for the Chorts Block herpetofauna with 146 species (38%) having EVS from 14 and Sampling Results by Locality Below is a summary of results pr esented by locality, from herpetofaunal sampling in the Chorts Block from June 2006 to April 2011. For each general locality, typically a

PAGE 81

81 protected area, the location and geographical extent are provided, along with details of the legal protection (if any ), a list of amphibian and reptile species recorded, the sites and dates visited, and a summary of findings. Parque Nacional Celaque Location and Extent: West central Departamento de Lempira and eastern Departamento de Ocotepeque, Honduras; 26,267 ha in e xtent, highest elevation 2,849 m. Status: National Park (legally declared in 1987; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetofaunal results: Bolitoglossa celaque Plectrohyla psiloderma Site visit summary: Campamento Don T oms (2 050 m), vicinity of Campamento Naranjo (2 560 2 850 m), Departamento de Lempira, core zone of PN Celaque. 21 28 June 2008. Comments: PN Celaque has a large core zone, and the physiographic structure of the mountain benefits conservation efforts. Steep sides to this mesa like mountain protect a large, relatively flat area above 2 500 m elevation that is almost completely intact. Upon this mesa originates the primary water source for the nearby city of Gracias, and protection of this and other impor tant watersheds provides added benefit Bolitoglossa celaque was found to be locally abundant around the Ro Arcagual Only one of four species of Plectrohyla reported from Celaque (Mc Cranie & Wilson 2002) was found during my visit ( P. psiloderma ) and additional targeted searches for the other three species ( P. guatemalensis P. hartwegi and P. matudai ) should be carried out

PAGE 82

82 immediately in an attempt to determine their status. Additi onal fieldwork in Celaque would almost certainly produce additional new herpetofaunal species. Parque Nacional Cerro Azul Copn Location and Extent: Western Departamento de Copn, Honduras, along the border with Guatemala; 11,766 ha, highest elevation 2,2 85 m. Status: National Park (legally declared in 1987; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetofaunal results: Caudata: Bolitoglossa nympha B. dofleini ; Anura: Dendropsophus microcephalus Incilius valliceps Lithobates b rownorum Lithobates maculatus, Ptychohyla hypomykter Smilisca baudinii ; Squamata: Anolis ocelloscapularis Lepidophyma flavimaculatum Sceloporus schmidti Adelphicos quadrivirgatum Mastigodryas dorsalis Ninia diademata N. sebae Typhlops stadelmani Site visit summary: Quebrada Grande, Departamento de Copn, 1 280 1 400 m elevation, in the buffer zone of PN Cerro Azul, on the south southeastern side of Cerro Azul. 26 30 July 2008. Summary findings : PN Cerro Azul is both a reserve with high importance with regard to its amphibian diversity and one at high risk due to a lack of management and increasing human population and fo res t clearing. The highest portion of the southeastern flank of Cerro Azul appears steep enough to deter campesinos from clearing most of the remaining forest. Unfortunately, there appear to be no streams flowing through this portion of forest due to the extreme topographical grade. Riparian areas in the vicinity of Quebrada Grande have been completely cleared and converted to livest ock pastures. Below this area, a larger creek (Que brada Caon Oscuro) forms

PAGE 83

83 from smaller streams running through pastures and rapidly descends into a deep canyon, inside which remains some riparian forest. The water source for Quebrada Grande is a spring j ust above town, with a very small (<1 ha) patch of forest remaining converted to agriculture. Additional work is urgently needed to determine the status of remaining forest both at higher elevations and of the side of Cerro Azul opposite to Quebrada Grande. Parque Nacional Cerro Azul Membar Location and Extent: East of Lago de Yojoa in extreme southern Departamento de Corts and northern Departamento de Comayagua; 17,872 ha, highest elevation 2,090 m. Status: National Park (legally declared in 1987; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetofaunal results: Caudata: Bolitoglossa mexicana, B. oresbia Nototriton limnospectator ; Anura: Craugastor laevissimus C. laticeps, Incilius ibarrai, I. porteri, I. valliceps, Lithobates maculatus, Ptychohyla hypomykter Smilisca baudinii Squamata: Anolis lemurinus A. cf. limifrons Sceloporus variabilis Sphaerodactylus millepunctatus Thecadactylus rapicau da Atropoides mexicanus, Bothriechis schlegelii Imantodes cenchoa Site visit summary: Departamento de Corts, 700 1 400 m elevation, 3 5 December 2006, 14 July 2007, 4 5 April 2008, 14 17 April 2008, 4 12 June 2008 25 27 No vember 2009 ; Cerro Azul, 800 1 740 m elevation, 18 April 2008, 5 12 July 2008; San Jos de los Planes, 930 1 ,290 m elevation, 1 3 June 2008.

PAGE 84

84 Figure 3 2 Sampling in the Chorts Block I. A) Campamento Don Toms, 2,080 m, Parque Nacional (PN) Celaque (A D, June 2008). B) The author in Montane Hepatic Forest, 2,770 m, PN Celaque. C) Campamento Los Naranjos, 2,570 m, PN Celaque. D) I. Luque searches for Bolitoglossa celaque in ground dwelling bromeliads; Montane Mixed Forest, 2,630 m, PN Celaque. E) Community of Quebrada Grande, 1,320 m, with PN Cerro Azul (July 2008). F) I. Luque preparing to descend over 350 m to Quebrada Grande, 1,690 m (July 2008). G) Hiking along Quebrada Varsovia, 1,670 m, PN Cerro Azul Membar (August 2008). Figure 3 2B I. Luque, all o ther photos J.H. Townsend.

PAGE 85

85

PAGE 86

86 Summary findings: Highland forest areas of this park previously had not been sampled, and my first two expeditions to cloud forest elevations in July 2008 yielded significant results, the discovery of Bolitoglossa oresbi a considered the most Critically Endangered salamander in Honduras whose previous known distribution consisted of a 1 ha patch of forest on top of the otherwise deforested Cerro El Zarciadero, and Nototriton limnospectator previously known only to occur across Lago de Yojoa in Parque Nacional Montaa de Santa Barbara (Townsend et al. 2011b) Parque Nacional Cusuco Location and Extent: Northwestern Depto. Corts and adjacent northern Depto. Santa Brbara, Honduras, near the Guatemalan border; 17,704 ha, h ighest elevation 2,242 m. Status: National Park (legally declared in 1987; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetofaunal results: Caudata: Bolitoglossa conanti B. diaphora ; Anura: Duellmanohyla soralia Plectrohyla dasypu s, P. exquisita, Ptychohyla hypomykter ; Squamata: Anolis amplisquamosus, A. cusuco, A. johnmeyeri Mesaspis moreletii, Sceloporus schmidti, Cerrophidion sp. Ninia diademata, N. espinali N. sebae Site visit summary: Vicinity of Centro de Visitantes (1 5 20 1 600 m), buffer zone of PN Cusuco; Quebrada Cantiles (1 820 m), core zone of PN Cusuco, 11 17 March 2006, 30 August 3 September 2008, 28 30 November 2009. Summary findings: Details of herpetofaunal work through 2006 are fo und in Townsend & Wilson (200 8); work in 2008 and 2009 did not appreciably add to this knowledge base.

PAGE 87

87 Parque Nacional La Tigra Location and Extent: South central Departamento de Francisco Morazn; 24,340 ha, highest elevation 2,290 m. Status: National Park (legally declared in 1980; Decreto 976 80); Conservation International Key Biodiversity Area. Herpetofaunal results: Anura: Incilius porteri, Lithobates maculatus ; Squamata: Anolis laeviventris, A. tropidonotus. Site visit summary: 500 1 700 m), core zone of PN La Tigra, 6 8 December 2006. Comments: The streamside frog Craugastor emleni has been reported to have undergone extreme declines, and was feared extinct by McCranie & Wilson (2002). This species was recently rediscovered from a small stream at 1,600 m elevation in Parque Nacional La Tigra (McCranie et al. 2010) and additional fieldwork should be undertaken at this easily accessible cloud forest to determin e the extent of the distribution of C. emleni Parque Nacional Montaa de Botaderos Loc ation and Extent: Northern Departa mento de Olancho, and peripherally in adjacent Departamento de Coln; 64,227 ha, maximum elevation 1,724 m. Status: National Park (legally declared in 2011; Acuedro 002 2011 ). Herpetofaunal results: Caudata: Nototriton sp .; Anura: Craugastor lauraster, C. noblei, Craugastor pechorum, Lithobates maculatus, Pristimantis ridens, Ptychohyla hypomykter; Squamata: Anolis capito, A. tropidonotus, Imantodes cenchoa, Ninia maculata, Stenorrhina degenhardtii

PAGE 88

88 Figure 3 3 Sampling i (PN) Cusuco, 1,550 m elevation; the day before the post coup presidential election, with red and white Partido Liberal party members meeting in seclusion (November 2009). B) Looking down from P N La Tigra, with El Rosario in the foreground and the Choluteca Valley beyond (B C, December 2006). C) S. Townsend at Quebrada de la Cascada, 1,850 m, PN La Tigra. D) R. Ulloa on trail leading to cloud forest, 1,640 m, PN Monta a de Botaderos (D G, April 2 m, PN Monta a de Botaderos. F) Our supplies and team near Alao, 1,100 m, on expedition into PN Monta a de Botaderos. G) M. Medina (left) recovering from nighttime sampling efforts while O. Re yes (right) prepares botanical samples at our base camp, 1,300 m, PN Monta a de Botaderos. Photos J.H. Townsend.

PAGE 89

89

PAGE 90

90 Site visit summary: ghest ridge in the range, 1,160 1,730 m elevation, 16 19 Apr il 2011. Summary findings: (Flag #93) to the Municipalidad de Gualaco in Departamento de Olancho, which included the first ever biological ex pedition to the highest portion of this newly decla red park (approved by the Honduran Congress the same week we were there). Parque Nacional Montaa de Comayagua Location and Extent: Southeastern Departamento de Comayagua and adjacent Departamento de Francisco Morazn; 29,767 ha, maximum elevation 2,410 m Status: National Park (legally declared in 1987; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetofaunal results: Caudata: Bolitoglossa mexicana ; Anura: Craugastor laevissimus Ptychohyla hypomykter Squamata: Anolis sminthus A. tropidonotus Sceloporus malachiticus. Site visit summary: La Oki, Departamento de Comayagua, 1 680 2 040 m elevation, edge of core zone for PN Montaa de Comayagua, 21 24 January 2008; Ro Negro, Departamento de Comayagua, 1 100 1 560 m elevation, buffe r zone of PN Montaa de Comayagua, 19 21 April 2008, 15 20 May 2008, 15 22 July 2008. Comments: Accessing areas of cloud forest above 1 500 m elevation has proven very challenging; intact cloud forest occurs as low as 1 200 m elevation along large streams and rivers. The community of Ro Negro is a model of community based conservation and sustainable development, and is a prime candidate to receive funding to support these areas. Areas of cloud forest above 2 000 m elevation need to be

PAGE 91

91 sampled for the pres ence of salamanders, as no salamanders have been found above 1 300 m elevation despite the presence of endemic highland species in surrounding mountain ranges. Parque Nacional Montaa de Santa Brbara Location and Extent: Southeastern Departamento de San ta Brbara; 13,202 ha, maximum elevation 2,744 m. Status: National Park (legally declared in 1987; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetofaunal results: Caudata: Bolitoglossa mexicana, Dendrotriton sanctibarbarus ; Anura: Craugastor laevissimus C. laticeps Lithobates maculatus, Ptychohyla hypomykter, Squamata: Anolis rubribarbaris Sceloporus malachiticus, Typhlops tycherus. Site visit summary: El Cedral, in the buffer zone of Parque Nacional Montaa de Santa Barbara on t he western slope of the mountain, 1 600 1 720 m elevation, 28 29 January 2008; Las Quebradas, in the buffer zone of Parque Nacional Montaa de Santa B rbara on the western slope of the mountain, 1 ,4 00 1 ,45 0 m elevation, 6 7 June 2010; core zone of Parque N acional Montaa de Santa Barbara west to east canyon across the crest of the mountain, 1,450 2,180 m elevation, 6 9 November 2010 Summary findings: The physiography of PN Montaa de Santa Brbara appears to be helping ensure the long term persistence o zone. The steep side and jagged karst composition deter deforestation above 2 000 m, and both endemic species known from within the park appear to have stable populations. My work in PN Montaa de Santa Brbara result ed in two notable discoveries: a remarkable new large species blindsnake ( Typhlops tycherus ; Townsend

PAGE 92

92 Figure 3 4 Sampling in the Chorts Block III. A) I. Luque and M. Medina in Rio Negro, hiking towards Quebrada El Gaviln, 1,200 m, Parque Nacional (PN) Monta a de Comayagua (April 2008). B) Quebrada El Gaviln, 1,260 m, PN Monta a de Comayagua (May 2008). C) J. Austin (foreground) and field team ascending near vertical trail above El Playn, 1,480 m, en route to PN Monta a de Santa Brbara (C E, November 2010). D) J. Austin at basecamp in canyon across top of PN Monta a de Santa Brbara, 2,200 m. E) L. Herrera setting infrared camera trap for mammal survey in PN Monta a de Santa Brbara, 2,190 m. F) G) Mesic ravine through Coniferous Cloud Forest, 1,930 m PN Monta a de Yoro (September 2008). Photos J.H. Townsend.

PAGE 93

93

PAGE 94

94 et al. 2008 a ), and the documentation of variation and distributional records of Anolis rubribarbaris a lizard species previously known from only a single poorly preserved specimen (Townse nd et al. 2008 b ) An expedition into a canyon across the top of Cerro Santa Brbara in November 2010 encountered the bromeliad salamander Dendrotriton sanctibarbarus to be locally abundant particularly around large, bromeliad laden branches that had come to rest on the forest floor. Parque Nacional Montaa de Yoro Location and Extent: Northern Departamento de Francisco Morazn and adjacent southern Departamento de Yoro; 11,766 ha, highest elevation 2,285 m. Status: National Park (legally declared in 1987; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetofaunal results: Caudata: Bolitoglossa cataguana, Nototriton lignicola, Oedipina kasios, Anura: Incilius porteri, I. valliceps, Lithobates maculatus, Ninia sebae, Plectrohyla guatemale nsis, Squamata: Anolis morazani, A. tropidonotus, A. yoroensis, Mesaspis moreletii, Sceloporus malachiticus, Cerrophidion sp. Rhadinella godmani. Site visit summary: Ro Maralito, outside Marale (610 630 m), seat of municipality; Los Planes (1 450 1 500 m), buffer zone community; Cataguana (1 780 2 020 m), northwestern core zone of PN Montaa de Yoro, 8 14 June 2006, 8 15 March 2007; Montaa de la Sierra ( 1,920 m), southeastern core zone of PN Montaa de Yoro, 21 25 September 2008. Comments: Results from this protected area are discussed in detail below.

PAGE 95

95 Parque Nacional Pico Bonito Location and Extent: Eastern Departa mento de Atlntida and adjacent Departa mento de Yoro; 107,107 ha, maximum elevation 2,480 m. Status: National Park (legally declared in 19 87; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetofaunal results: Anura: Craugastor aurilegulus, C. noblei, Duellmanohyla salvavida, Pristimantis ridens, Ptychohyla spinipollex; Squamata: Ameiva festiva, Anolis lemurinus, A. trop idonotus, A. zeus, Basiliscus vittatus, Corytophanes cristatus, Incilius leucomyos, Mabuya unimarginata, Plestiodon sumichrasti, Sphenomorphus cherriei, Thecadactylus rapicauda, Leptophis ahaetulla, Oxybelis aeneus. Site visit summary: Cangrejal, vicinity December 2009; Quebrada de Oro, 930 1,380 m elevation, 21 28 May 2010; CURLA Station, 150 m elevation, 10 12 August 2010 ; Pico Bonito Lodge, 120 m, 7 April 2011 Comments: My fieldwork in Parque Nacional Pico B onito has thus far been limited to the lowlands, supported by two fieldtrips undert aken by Csar Cerrato (UNAH) in 2010. Parque Nacional Pico Pijol Location and Extent: Southwestern Departa mento de Yoro; 11,669 ha, maximum elevation 2,282 m. Status: Nati onal Park (legally declared in 1987; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetofaunal results: Anura: Craugastor laevissimus, C. rostralis, Lithobates maculatus, Ptychohyla hypomykter Smilisca baudinii ; Squamata: Anolis yoro ensis, Oxybelis fulgidus.

PAGE 96

96 Site visit summary: Road above El Porvenir de Morazn, northeastern buffer zone of PN Pico Pijol in the vicinity of Cerro Las Pajarillos, 1 380 1 520 m elevation, 23 25 July 2008; 17 20 September 2008 Summary findings: Extensive shade coffee production dominates the northern side of the buffer zone of PN Pico Pijol. A single frog, Craugastor laevissimus was collected along a path through converted cloud forest; however, a search of a medium sized upper tributary of Quebrada Las Payas surrounded by moderately disturbed forest failed to find any of the normally stream side Craugastor. No individuals of Bolitoglossa porrasorum were collected, despite targeted searching for this species. A single Ptychohyla hypomykter and two groups of tadpoles were the only amphib ians observed along the stream. Parque Nacional Sierra de Agalta Location and Extent: East central Departamento de Olancho; 51,793 ha, maximum elevation 2,354 m. Status: National Park (legally declared in 1987; Decreto 87 87 ); Conservation International Key Biodiversity Area. Herpetological results: Caudata: Bolitoglossa mexicana B. longissima Nototriton sp.; Squamata: Anolis cf. sminthus Atropoides indomitus Cerrophidion sp Site visit summary: Trail to and vicinity of Cerro La Picucha, 615 2,230 m elevation. Summary findings: Fieldwork led by Melissa Medina Flores (Universidad Nacional Autnoma de Honduras) resulting in collection of the first record of Atropoides indomitus from the Sierra de Agalta and two specimens of an unknown species of Nototriton (chapters 4 and 5; Townsend et al. 2011a).

PAGE 97

97 Refugio de Vida Silvestre Texiguat Location and Extent: Southwestern Departamento de Atlntida and adjacent northwestern Departamento de Yoro; 15,736 ha, maximum elevation 2,20 8 m. Status: Wildlife Refuge (legally declared in 1987; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetological results: Caudata: Bolitoglossa nympha, B. cf. porrasorum, Nototriton cf. barbouri, N. tomamorum, Oedipina gephyra; Anu ra: Craugastor aurilegulus, Craugastor cf. rostralis, Duellmanohyla salvavida, Hyalinobatrachium fleischmanni, Incilius leucomyos, I. valliceps, Leptodactylus fragilis Plectrohyla chrysopleura, Ptychohyla spinipollex, Smilisca baudinii, Teratohyla pulvera ta ; Squamata: Anolis beckeri, A. kreutzi, A. loveridgei, A. yoroensis, A. zeus, Corytophanes cristatus, Laemanctus longipes, Lepidophyma flavimaculatum, Sceloporus malachiticus, Sphenomorphus cherriei, Adelphicos quadrivirgatum, Atropoides mexicanus, Bothr iechis marchi, B. schlegelii, Bothrops asper. Dendrophidion percarinatum, Drymobius chloroticus, Geophis damiani, Hydromorphus concolor, Imantodes cenchoa, Leptodeira septentrionalis, Micrurus nigrocinctus, Ninia pavimentata, Ninia sebae, Pliocercus elapoi des, Scaphiodontophis annulatus, Sibon dimidiatus, Sibon nebulatus, Stenorrhina degenhardtii, Tantilla sp. Tropidodipsas sartorii. Site visit summary: 2.5 km NNE of La Fortuna, buffer zone of RVS Texiguat, 1 500 1 890 m elevation, 8 11 April 2008; 10 21 June 2010, 25 July 1 August 2010 Comments: RVS Texiguat might be simultaneously the single protected area of the most biodiversity significance and facing the strongest threat to its long term survival in the Honduran protected areas system. The reserve is discussed in detail later in this chapter.

PAGE 98

98 Figure 3 5 Sampling in the Chorts Block IV. A) Cerro de Pajarillos, 1,460 m, Parque Nacional (PN) Pico Pijol (July 2008). B) J. Slapcinsky (left) and J. Butler (right) at basecamp, La Fortuna, 1,650 m, Refu gio de Vida Silvestre Texguat (B C, April 2008). C) Type locality of Isthmohyla insolita and Nototriton tomamorum La Fortuna, 1,550 m, Refugio de Vida Silvestre Texguat. D) Cattle pond with explosive breeding aggregation of Exerodonta catracha and Hypop achus barberi dark masses in water are H. barberi eggs; 2,210 m, Reserva Biolgica (RB) Guajiquiro (D E, May 2008). E) Male Exerodonta catracha guarding egg clutches in pond from Figure 3 5D. F) Degraded Mixed Cloud Forest, 2,100 m, RB G isayote; Anolis h eteropholidotus and Mesaspis moreletii were abundant in the pasture in the foreground (F G, June 2008). G) Camping under the communications tower at the top RB G isayote, 2,270 m. Photos J.H. Townsend.

PAGE 99

99

PAGE 100

100 Reserva Biolgica Cerro Uyuca Location and Exte nt: Southeastern Departamento de Francisco Morazn, 772 ha, maximum elevation 2,006 m. Status: Biological Reserve (legally declared in 1987; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetological results: Anura: Incilius porteri, Lithobates brownorum X forreri (this population is c onsidered to be interspecific hybrids by McCranie & Wilson 2002) L. maculatus Ptychohyla salvadorensis, Rhinella marina, Tlalocohyla loquax ; Squamata: Anolis laeviventris A. tropidonotus Sceloporus m alachiticus Site visit summary: Vicinity of Cabot Biological Station, 1 620 1 700 m elevation, 17 18 March 2007, 17 19 July 2007. Summary findings: Sampling in Reserva Biolgica Cerro Uyuca took place in the vicinity of the biological station managed by the Centro Zamorano de Biodiversidad, which includes access to a small reservoir supporting a dense population of Lithobates brownorum / forreri Reserva Biolgica Guajiquiro Location and Extent: Departamento de La Paz; 17,165 ha, maximum elevation 2,330 m. Status: Biological Reserve (legally declared in 1987; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetofaunal results: Caudata: Bolitoglossa celaque ; Anura: Exerodonta catracha, Hypopachus barberi, Incilius ibarrai, Lithobates maculatus ; Squamata: Anolis crassulus Drymobius chloroticus Ninia espinali Thamnophis fulvus

PAGE 101

101 Site visit summary: Guajiquiro and surrounding countryside, 1 900 2 240 m elevation, 23 25 May 2008 14 15 August 2008 Summary findings: Highland areas above about 2000 m elevation in the vicinity of the town of Guajiquiro are a matrix of both traditional and modern agriculture and maintained patches of old growth cloud forest. In some places, proximity of existing patches and make up of the intervening agricu ltural areas is such as to allow species such as Bolitoglossa cf. celaque and Exerodonta catracha to persist and, in some cases, apparently thrive. Cattle ponds in small pastures were readily being used for breeding by the anurans E. catracha and Hypopachu s barberi (Ketzler et al. 2011, Luque Montes et al. 2011), and B. cf. celaque was found in a forest patch surrounding a communications tower. Additional highland areas with forest patches in the area need to be visited to fully and accurately assess the co nservation needs of the relatively large but biogeographically connected highland area. Reserva Biolgica Gisayote Location and Extent: Departamento de Ocotepeque; 12,677 ha, maximum elevation 2,330 m. Status: Biological Reserve (legally declared in 198 7; Decreto 87 87); Conservation International Key Biodiversity Area. Herpetofaunal results: Caudata: Bolitoglossa conanti ; Anura: Hypopachus barberi ; Squamata: Anolis heteropholidotus Mesaspis moreletii Sceloporus malachiticus Site visit summary: Commu nication towers southeast of El Portillo de Ocotepeque, core zone of RB Gisayote, 2 080 2 230 m elevation, 18 21 June 2008.

PAGE 102

102 Summary findings: The core zone of RB Gisayote has an access road running along the top of the reserve through high pastures and r emnant cloud forest to a set of communications towers. This reserve appears to support a surprising amount of forest given that it lies in a region that is otherwise heavily impacted by human activity. The status of Leptodactylus silvanimbus is unclear, it has been documented previously inhabiting open fields and pastures with flooded areas in the vicinity. Reserva Biolgica Yerbabuena Location and Extent: Southwestern Departamento de Francisco Morazn 3,522 ha, maximum elevation 2,243 m. Status: Biologic al Reserve (legally declared in 1987; Decreto 87 87); Conservation International Key Biodiversity Area Herpetological results: Caudata: Bolit o glossa carri ; Anura: Incilius ibarrai; Squamata: Anolis sminthus A. tropidonotus Sceloporus malachiticus Site visit summary: Finca La Alondra, cafetal in region of Cerro Cantagallo, and other fincas in the area, 1 750 2 020 m elevation, 16 17 July 2007. Summary findings: Despite the fact that most of the forest around Cerro Cantagallo is disturbed and much of tha t is fully converted to agriculture, Bolitoglossa carri can still be found relatively easily inside bromeliads in remaining forest patches. Additional work should be done to determine the extent of remaining forest on the highes t portions of Cerro Cantagal lo. Reserva de la Biosfera Bosawas Location and Extent: Departamento de Jinotega and Regin Autnom a Atlntico Norte; 1,992,000 ha

PAGE 103

103 Status: UNESCO Man and the Biosphere Reserve (legally declared in 1991 and revised and expanded in 1997 and 2001; Decreto 4 4 91, Decreto 32 96, and Ley 407). Herpetological results: Gymnophiona: Gymnopis multiplicata; Caudata: Bolitoglossa striatula; Anura: Agalychnis callidryas, Cochranella granulosa, Craugastor fitzingeri, C. lauraster, C. megacephalus, C. mimus, C. noblei, Dendropsophus microcephalus, Diasporus diastema, Incilius valliceps, Leptodactylus melanonotus, savagei, Lithobates vallanti, Pristimantis cerasinus, P. ridens, Rhaebo haematiticus, Rhinella marina, Smilisca baudinii, S. phaeota, S. sordida, Teratohyla pul verata; Squamata: Ameiva festiva, Anolis capito, A. limifrons, A. lionotus, A. quaggulus, Basiliscus plumifrons, B. vittatus, Corytophanes cristatus, Iguana iguana, Sphaerodactylus millepunctatus, Sphenomorphus cherriei, Thecadactylus rapicauda, Adelphicos quadrivirgatum, Boa constrictor, Bothrops asper, Drymarchon melanurus, Imantodes cenchoa, Leptodeira septentrionalis, Mastigodryas melanolomus, Micrurus nigrocinctus, Ninia sebae, Oxybelis aeneus, O. brevirostris, Porthidium nasutum, Pseustes poecilonotus Sibon longifrenis, S. nebulatus, Tretanorhinus nigroluteus; Testudines: Kinosternon leucostomum, Rhinoclemmys annulata, R. funerea, Trachemys venusta. Site visit summary: Muru Ta, 12 13 June 2007, 180 m elevation; Muru Lak, 13 18 June 2007, 190 m elevati on; Kalum Kitang, 18 21 June 2007, 180 m elevation; Aran Dak, 11 and 22 June 2007, 150 m elevation; Raiti, 23 June 2007, 140 m elevation. Summary findings: Results from this expedition, the first in a series, to the core zone of Bosawas were reported in S unyer et al. (2009) and Travers et al. (2011).

PAGE 104

104 Figure 3 6 Sampling in the Chorts Block V. A) Cerro Yaluk, Departamento de Olancho, Honduras, seen from the Ro Coco (A D, June 2007). B) Aran Dak, most remote Mayangna community on Ro Lakus, 150 m, Res erva de la Bisfera Bosawas, Nicaragua. C) L. Wilson (seated) being poled down the upper Ro Lakus by Miskitu parataxonomist S. Charley (standing); core zone of Reserva de la Bisfera Bosawas, 170 m. D) Preparing samples; (left to right) L. Wilson, S.Trave rs (seated), Miskitu parataxonomists R. Picado and S. Charley, J. Sunyer (seated), and Mayangna parataxonomist J. Lopez; Muru Lak, 180 m, core zone of Reserva de la Bisfera Bosawas. E) I. Luque ascending the volcanic peak of Isla El Tigre, with a view of the Golfo de Fonseca and the volcanoes of eastern El Salvador; 550 m (August 2010). F) Campsite at the summit of Cerro Zarciadero, 1,890 m, Honduras (July 2007). Photos J.H. Townsend.

PAGE 105

105

PAGE 106

106 Jardn Bot nico Lancetilla Location and e xtent: Western Departame nto de Atlntida ; 2,255 ha, maximum elevation 710 m Status: Nationa l Park (legally declared in 1990 ; Decreto 48 90 ); Conservation International Key Biodiversity Area. Herpetological results: Anura: Craugastor aurilegulus Leptodactylus melanonotus Lithob ates brownorum L. vallanti Smilisca baudinii ; Squamata: Ameiva festiva Anolis lemurinus A. cf. zeus Lepidophyma flavimaculatum Sphaerodactylus millepunctatus Sphenomorphus cherriei Clelia clelia Coniophanes imperialis Site visit summary: 7 9 Jun e 2010, 22 24 July 2010. Summary findings: Besides the extensive grounds of the botanical gardens, Lancetilla also protects the entire watershed of the Ro Lancetilla. This includes 1,281 ha of virgin lowland rainforest, one of the best preserved fragments of lowland rainforest along the northern coast of Honduras. rea de Uso Multple Isla del Tigre Location and Extent: Island in the Golfo de Fonseca on the Pacific Coast, Departamento de Valle; 601 ha, maximum elevation 783 m. Status: Multiple Use Area (legally declared in 1999; Decreto 5 99 E). Herpetological results: Anura: Incilius coccifer ; I. porteri ; Squamata: Sceloporus squamosus Site visit summary: Amapala and trail to summit, 5 783 m elevation, 16 17 August 2010.

PAGE 107

107 Non P rotected Areas Cerro E l Zarciadero Location: Northern Departamento de Comayagua. Herpetological results: Caudata: Bolitoglossa oresbia ; Anura: Exerodonta catracha, Incilius porteri ; Squamata: Drymobius margaritiferus Site visit summary: Cerro El Zarciadero, 1,845 1,890 m ele vation, 14 15 July 2007. Comments: The small patch of forest on top of Cerro El Zarciadero remains intact, and is essentially protected as private property around a set of communications towers. A family lives at the entrance to the tower complex and the husband is paid to guard and maintain the grounds. The residents were already familiar with Bolitoglossa oresbia and the idea that they are protecting the only known habitat of this animal, which was found active on vegetation on the only night we spent on top of the mountain. Exerodonta catracha was also found actively calling and breeding in and around a very small water basin and stream just below the road and towers. The situation with the remaining forest patch, tiny as it is, appears stable for the ti me, since it provides a buffer area around the communications towers at the top of the mountain. The discovery of Bolitoglossa oresbia in PN Cerro Azul Membar also lessens the urgency with which earlier conservation efforts surrounding El Zarciadero were being pursued. Highlands surround the Meseta de La Esperanza Location: Central Departamento de Intibuc, mountainous areas surrounding the Meseta de La Esperanza; 1,700 2,100 m elevation. Herpetological results: Caudata: Bolitoglossa celaque ; Anura: Exero donta catracha Incilius ibarrai Lithobates brownorum X forreri (this population is c onsidered

PAGE 108

108 to be made up of interspecific hybrids [McCranie & Wilson 2002]) ; Squamata: Anolis crassulus, A. sminthus, Sceloporus malachiticus S. variabilis Site visit s ummary: San Pedro La Loma, 1 960 2 020 m elevation, 21 24 January 2008 16 17 August 2008 ; Cerro El Peln, 2 065 m elevation, 30 June 2008; Zacate Blanco, 1 950 2 100 m elevation, 29 30 June 2008 18 19 August 2008. Summary findings: As is the case in RB G uajiquiro, highland areas above around 2 000 m elevation in the vicinity of La Esperanza are a matrix of both traditional and modern agriculture and patches of remnant cloud forest. In some places, proximity of existing patches and make up of the interveni ng agricultural areas are such as to allow species such as Bolitoglossa cf. celaque and Exerodonta catracha to persist and, in some cases, apparently thrive. Additional highland areas with forest patches in the area need to be visited to fully and accurate ly assess the conservation needs of the relatively large but biogeographically connected highland area. Los Naranjos Location: Near the border of Departamento de Corts and Departamento de Santa Brbara, between Lago de Yojoa to the east and Parque Naciona l Montaa de Santa Brbara to the west; 700 730 m elevation. Herpetological results: Caudata: Bolitoglossa mexicana ; Anura: Craugastor laevissimus, Dendropsophus microcephalus Engystomops pustulosus Hyalinobatrachium fleischmanni Hypopachus variolosus Incilius valliceps, Lithobates brownorum, L. maculata, Rhinella marina, Smilisca baudinii ; Squamata: Ameiva undulata Anolis laeviventris A. lemurinus A. tropidonotus, A. unilobatus, A. cf. zeus, Sphaerodactylus millepunctatus, Sphenomorphus cherriei, N inia diademata, N. sebae Tantilla taeniata

PAGE 109

109 Site visit summary: 13 15 August 2007 30 31 January 2008, 5 April 2008, 11 12 April 2008, 20 22 May 2008, 7 June 2010 Summary findings: Collections in this area were primarily made on the Plowden family farm now called Compaia Agrcola El Paraso This large property includes numerous springs, shade coffee plots, and shaded horticultural plots. Montaa de Jacaleapa Location: Central Departamento de Olancho, vicinity of Nahoan community of El Norte, 980 1,1 80 m elevation. Herpetological results: Anura: Craugastor laevissimus, C. lauraster, C. noblei, Lithobates maculatus, Pristimantis ridens, Ptychohyla hypomykter ; Squamata: Anolis sp. A. tropidonotus Sceloporus malachiticus Sphenomorphus cherriei Site visit summary: 11 12 April 2011. Summary findings: This isolated patch of mesic highland forest had not previously been surveyed by biologists, and supports over 500 ha of intact premontane rainforest. Montaa Macuzal Location: Southern Departamento de Yoro, west of Yorito. Herpetological results: Caudata: Bolitoglossa porrasorum Nototriton barbouri ; Anura: Craugastor rostralis ; Squamata: Anolis laeviventris, A. pijolensis A. yoroensis Sceloporus malachiticus. Site visit summary: El Panal, southwest side of Montaa de Macuzal, 1500 1800 m elevation, Departamento de Yoro, 31 January 3 February 2008, 6 7 April 2008. Comments: While the flanks of Montaa de Macuzal are converted to shade coffee cultivation or completely cleared of all natural vegetation the area above 1780 m

PAGE 110

110 Figure 3 7 Sampling Forest surrounded by converted corn fields, 2,230 m, Sierra de Opalaca west of La Esperanza (July 2008). B) Small reservoir supporting populatio n of Lithobates brownorum X forreri 2,010 m, Cerro San Pedro, east of La Esperanza (January 2008). C) Converted cloud forest, Zacate Blanco, highest point 2,250 m (July 2008). D) L. Wilson and I. Luque searching for Bolitoglossa cf. celaque in bromeliads that had been cut and left for cattle feed, 2,010 m, Cerro San Pedro. E) Intact bromeliad cover, 2,020 m, Cerro San Pedro. F) L. Wilson at Pozo Azul, a karstic spring near Los Naranjos, 680 m. G) Monta a Macuzal, viewed from the northeast, showing strip of remnant Broadleaf Cloud Forest at highest elevations (maximum 1,905 m). H) Monta a Macuzal, viewed from the southwest near El Panal, 1,480 m, with yellow wildflowers covered the completely deforested slopes; access to the cloud forest visible in the saddl e (January 2008). I) N. Stewart (front) and J. Butler (back) crossing upper deforested slope of Monta a Macuzal (April 2008). Photos J.H. Townsend.

PAGE 111

111

PAGE 112

112 elevation still supports an approximately 5 10 ha patch of moderately disturbed cloud forest on karsti c soils, and Bolitoglossa porrasorum and Nototriton barbouri are persisting, and in the case of the former species, abundantly so. A stream flowing out of the northern corner of Montaa de Macuzal near the community of El Portillo should be checked for sig ns of Duellmanohyla salvavida and/or Plectrohyla guatemalensis ( McCranie & Wilson 2002) as well as other priority species. Saguay Location: Upper Valle de Agalta, Departamento de Olancho. Herpetological results: Anura: Hypopachus variolosus, Incilius val liceps, Leptodactylus fragilis, Lithobates maculatus Ptychohyla hypomykter, Smilisca baudinii, Tlalocohyla loquax ; Squamata: Anolis capito A. quaggulus A. wermuthi Sceloporus malachiticus ; Testudines: Kinosternon scorpioides Site visit summary: Teoci ntecito ravine, 720 m elevation, and a seepage bog outside Saguay, 580 m elevation, 10 April 2011. Summary findings: Survey sites in remnant dry forest patches in the foothills around the upper Ro Grande (also called the Sico, Tinto, and Negro at differe nt portions downstream) watershed. One locality was a relatively large seepage bog, covered with a t hink mat of floating vegetation that arises from the middle of the dry valley floor. San Jos de Texguat Location: Southern side of the upper Ro Len val ley in the foothills of Texguat, 150 200 m elevation. Herpetological results: Caudata: Bolitoglossa nympha ; Anura: Craugastor aurilegulus Duellmanohyla salvavida Hyalinobatrachium fleischmanni, Leptodactylus

PAGE 113

113 fragilis ; Squamata: Anolis cf. zeus Iguana iguana Lepidophyma flavimaculatum Atropoides mexicanus, Hydromorphus concolor Site visit summary: 1 0 1 1 November 2010 Summary findings: Sampling was carried out in a deep, protected ravine just south of the town. Craugastor aurilegulus and Duellmano hyla salvavida were found in abundance around the small, spring fed creek. A juvenile Hydromorphus concolor was found in a rocky fissure at the spring source. Selva Negra Location: Between cities of Jinotega and Matagalpa, Departamento de Matagalpa, 1,2 00 1,300 m elevation. Herpetological results: Anura: Agalychnis callidryas, Craugastor lauraster, Incilius valliceps, Lithobates cf. taylori Ptychohyla hypomykter, Smilisca baudinii, Tlalocohyla loquax ; Squamata: Anolis capito A. quaggulus A. wermuthi Sceloporus malachiticus Site visit summary: 13 15 August 2007. Summary findings: Collections were made in the premontane rainforest above the Selva Negra coffee farm and lodge. Yeguare Valley Location: Valley associated with an upper tributary of the Ro Choluteca, 810 m elevation Herpetological results: Anura: Engystomops pustulosus ; Squamata: Basiliscus vittatus Gonatodes albogularis Porthidium ophryomegas ; Testudines: Kinosternon scorpioides Trachemys venusta Site visit summary: 16 17 March 20 07; 3 4 June 2010.

PAGE 114

114 Figure 3 8 Sampling in the Chorts Block VII. A) O. Reyes (left) and M. Bonta (right) hiking through the Nahoan community of El Norte en route to Monta a de Jacaleapa, 1,100 m (A E, April 2011). B) Premontane rainforest stream, 1,120 m, Monta a de Jacaleapa. C) The author with a giant (10+ m) teocinte cycad ( Dioon mejiae ) near Saguay, 630 m, Valle de Agalta. D) La Puzunca, upper reach of the Ro Grande, 560 m, Valle de Agalta; distinctive green plants on far hillsides are teocinte cyca ds. E) Giant paddle cactus ( Nopalea cf. hondurensis ), 580 m, Valle de Agalta. F) Artificial lagoon, 1,220 m, Selva Negra, Nicaragua (F H, August 2007). K. Townsend (front) and S. Travers (back) climb a trail through premontane rainforest, 1,460 m, Selva Ne gra. H) Premontane rainforest creek, 1,420 m, Selva Negra. Photos J.H. Townsend.

PAGE 115

115

PAGE 116

116 Summary findings: Collections made in agricultural areas and remnant premontane dry forest in and around the campus of Escuela Agrcola Panamericana Zamorano Discussion Baseline Herpetological Inventory of Parque Nacional Montaa de Yoro In June 2006, March 2007, and September 2008 I led the initial herpetological survey work carried out in Parque Nacional Montaa de Yoro a relatively large yet herpetologically unknown prot ected area in central Honduras Parque Nacional Montaa de Yoro was established in 1987 and sits along the boundary between the Honduran departments of Francisco Morazn and Yoro, with about two thirds of its area in the municipality of Marale in nort hernmost Francisco Morazn ( COHECO 2003 ). With a total area of over 154 km 2 more than 47 km 2 of cloud forest supporting area above 1 800 m, Parque Nacional Montaa de Yoro potentially contains one of the largest areas of cloud forest remaining in Honduras ( COHECO 2003 ). Next to nothing has been are mentioned in the park management plan, with most of those being relatively widespread species that commonly are found around h uman habitation or in habitats below cloud forest ( Townsend & Wilson 2009 ). Additionally, Parque Nacional Montaa de Yoro protects some of the highest portions of the Montaas de la Flor, which is home to the last traditional communities of indigenous Tolu panes that maintain the Tol language and culture (Chapman 1992). Within t people spread across 55 communities, with the population consisting of Tolupan, mestizos and mixed Tolupan mestizo farmers (COHECO 2003). Most of the se people inhabit the

PAGE 117

117 Figure 3 9 Exemplar paratypes and habitats from Parque Nacional Monta a de Yoro. A) Adult male paratype (UF 150000) of A nolis morazani sp. nov. B) A dult female paratype (MCZ 185612) of A morazani sp. nov. C) Dewlap of UF 150000. D) A dult female paratype (MVZ 258030) of Bolitoglossa cataguana B. sp. inquirenda 1 n Chapter 4). E) A rboreal tank bromeliads in disturbed mixed cloud forest, Cataguana, 1,880 m F) Quebrada Cataguana, 1,820 m. G) R emote homestead at Cataguana, 1,910 m Photos J.H. Townsend. nearly completely deforested buffer zone of Parque Nacional Montaa de Yoro and as the buffer zone population grows increasing pressure is being placed on the remaining cloud forest in the nuclear zone. Parque Nacional Montaa d e Yoro has become so heavily impacted by the encroaching agricultural frontier that at one point it was

PAGE 118

118 recommended that the park be re designated as a biological reserve and that its borders be reduced to the remaining extent of intact cloud forest ( Vreug denhil et al. 2002). My work in Parque Nacional Montaa de Yoro was limited to the vicinity of two sites, Cataguana on the northwestern side of the core zone, and the highlands above Guaymas (sometimes referred to as Montaa de la Sierra ; however this is not to be confused with the Montaa s de la Sierra of Departamento de La Paz). Despite the relatively limited amount of fieldwork carried out, the expeditions were no netheless fruitful and resulted in the discovery of a new species of anole ( Anolis morazan i ; Townsend & Wilson 2009), and salamanders from three genera that could not be unambiguously assigned to a named species. A New Species of Anole Fieldwork in Parque Nacional Montaa de Yoro uncovered a relatively common anole lizard assignable to the Ano lis crassulus group, characterized by having moderately to strongly enlarged medial dorsal scales, no more than two scales separating the supraorbital semicircles, four to seven rows of loreals, suboculars, and supralabials in contact under the central por tion of the orbit, enlarged postanals in males, and heterogeneous flank squamation (McCran ie et al. 1992; Khler et al. 1999). The majority of the species in the A. crassulus group are endemic to the Chorts Highlands, including A. amplisquamosus A. heter opholidotus A. muralla A. rubribarbaris A. sminthus and A. wermuthi ( Khler 2008) In addition, one widespread species, A. crassulus is found across the highlands of Nuclear Central America from Chiapas, Mxico to El Salvador and southwestern Hondura s. The population from Parque Nacional Montaa de Yoro was demonstrated to be morphologically distinctive

PAGE 119

119 from all other species in the A. crassulus group based on hemipenial structure and scalation, and I subsequently described it as the new species Anoli s morazani (Townsend & Wilson 2009). Salamanders of Uncertain Taxonomic Assignment Populations of at least three different salamanders were discovered in Parque Nacional Montaa de Yoro A series of Bolitoglossa assignable to the B. dunni species group b ased on having well developed subdigital pads and bluntly rounded, free toe tips but with coloration differing from other described species, was collected at Cataguana in 2006 and 2 007 and above Guaymas in 2008. Three specimens of Nototriton were also coll ected at Cataguana from within ground cover, and a small Oedipina was collected in a deep mesic ravine through coniferous cloud forest. Taxonomic assignment of these three populations, referred to as Bolitoglossa sp. inquirenda 1 Nototriton sp. inquirenda 5 and Oedipina sp. inquirenda 1 are addressed in Chapter 4. Parque Nacional Montaa de Yoro appears to be both overlooked and undervalued by national and international conservation planners due to perceived lack of documented conservation priority s pecies ; however this apparent lack of diversity is a result of undersampling and is biodiversity. Prior to initiation of survey work in Parque Nacional Montaa de Yoro Parque Nacional Montaa de Yoro was surrounded by literally dozens of cloud forest areas many of which have well documented and diverse endemic biotas (Wilson & McCranie 2004 b ). Given the biodiversity present in other cloud forest areas in Ho nduras, it is highly probable that additional undescribed species await discovery in Parque Nacional Montaa de Yoro Future work in Parque Nacional Montaa de Yoro should continue to shed light on

PAGE 120

120 these relationships and provide a clearer picture of highl and biogeography in eastern Nuclear Central America. Hotspot within a Hotspot: the Special Case of Refugio de Vida Silvestre Texguat The Refugio de Vida Silvestre (RVS) Texguat was established in 1987 and consists of approximately 33,267 ha of premontane and lower montane rainforest straddling the border of the Honduran departments of Atlntida and Yoro (CIPF 2009). RVS Texguat is administered by the Instituto Nacional de Conservacin y Desarrollo Forestal, reas Protegidas y Vida Silvestre (ICF) as part of the Sistema Nacional de reas Protegidas de Honduras (SINAPH), with management authority for the reserve in the hands of the non governmental organization Fundacin para la Proteccin de Lancetilla, Punta Sal y Texguat (PROLANSATE). Most herpetologica l surveys in RVS Texguat have been conducted since 1991 on the leeward side of the park in Departamento de Yoro, at elevations above 1,500 m in the vicinity of a coffee farm known locally as La Fortuna (Holm & Cruz 1994; McCranie et al. 1993 ; McCranie & C astaeda 2004a, b; Townsend et al. 2010a; Wilson et al. 1994, 1998). Situated at the western end of the Cordillera Nombre de Dios, the leeward side of RVS Texguat and its counterpart at the eastern end of the Cordillera, Parque Nacional (PN) Pico Bonito, are the most significant areas of herpetofaunal diversity in a country whose national level of endemism has already been demonstrated to be the highest in Central America (Wilson & Johnson 2010). Our preliminary results from the windward side of RVS Texg uat indicate that this area represents a unique opportunity for

PAGE 121

121 Figure 3 10 La Liberacin de Texguat. A) Valley of the Ro Jilamito, with the remote encampment known as La Liberacin de Texguat partially visible as the lower clearing in the center of the picture beyond the emergent palm; the entire expanse from La Liberacin to the farthest peak (Cerro Texguat, 2,208 m) is undisturbed premonta ne and lower montane rainforest. B) T he initial leg of the expedition to La Liberacin requires a 2 hour mule ride through the lowlands (L.D. Wilson on mount). C) T he mule ride is followed by a 6 8 hour hike into the mountain, with around two thirds the trail within this deep, high grade trench (M. Medina Flores following our guides and pack animals). Photos J. H. Townsend.

PAGE 122

122 conserving the remarkable diversity that has already been documented on the leeward side of RVS Texguat, and for expanding the known herpetof aunal diversity of the reserve. Discovery of Plectrohyla chrysospleura Plectrohyla chrysopleura (Anu ra: Hylidae) is a large, critically endangered spikethumb frog only known from the vicinity of its type locality in Parque Nacional Pico Bonito (Cruz et al. 2004). The type locality, Quebrada de Oro, is a premontane rainforest locality that has been under relatively intensive study since 1980 and is among the most remarkable sites of amphibian endemism in Central America (McCranie & Wilson 2002). Unfortunately, this locality also has the distinction of being one of the best documented cases of catastrophic amphibian decline in Central America (McCranie & Wilson 2002; McCranie & Castaeda 2005, 2007; Townsend & Wilson 2010). It is the type locality for six species of amphibians ( Craugastor aurilegulus C. chrysozetetes C. fecundus Duellmanohyla salvavida Plectrohyla chrysopleura and Rhinella chrysophora ), and is just below the type locality (Cerro Bfalo) of two other species ( Craugastor cruzi and C. saltuarius ). Almost all of these species are considered to be in decline, with even some considered to be extinct or close to extinction (McCranie & Castaeda 2007). Like other species from Quebrada de Oro that have declined or disappeared, P. chrysopleura is apparently extirpated from that locality, and given that Quebrada de Oro is the only locality where th is species has been found despite consistent focused work in the area (McCranie and Castaeda 2005, 2007), it raises the concern that the species might be near extinction, if not already extinct. The last time P. chrysopleura was documented as extant was i n May 1996, when two metamorphs and two tadpoles

PAGE 123

123 Figure 3 11 Plectrohyla chrysopleura (Hylidae) from La Liberacin. A) Adult female Plectrohyla chrysopleura ( USNM 573993 ) from La Liberacin, 1,030 m elevation, Refugio de Vida Silvestre Texiguat, Hondu ras. B) Adult male P. chrysopleura ( USNM 573995 ) from Cerro El Chino, 1,420 m elevation, Refugio de Vida Silvestre Texiguat, Honduras. C) Juvenile P. chrysopleura ( USNM 573994 ) from La Liberacin D) Recently metamorphosed P. chrysopleura ( not collected ) f rom La Liberacin. E) Ro Jilamito at La Liberacin, 1,02 0 m E) Tributary of Ro Jilamito, La Liberacin, 1,03 0 m Photos J.H. Townsend.

PAGE 124

124 were collected along Quebrada de Oro (McCranie and Wilson 2002, McCranie and Castaeda 2005). At that time, one of two tadpoles collected had deformed mouthparts (McCranie and Wilson, 2002). Based on these data and additional considerations, Cruz et al. (2004), IUCN (2011 ), and Townsend and Wilson (2010) all judged P. chrysopleura to be Critically Endangered, based on IUCN criteria (IUCN, 2001). For P. chrysopleura the particular red list status was Critically Endangered (A2ace, B1ab[iii,v]+2ab[iii,v]) or suspected over the last 10 years based on direct ob servation, a decline in area of occupancy, extent of occurrence and quality of habitat, and the suspected impact and susceptibility to decline from chytridiomycosis. In June and July 2010, a series of Plectrohyla chrysopleura was collected around La Liberacin and Cerro El Chino on the windward side of RVS Texguat, providing the species, approximately 55 km west southwest of the type local ity (Townsend et al. 2011 c ). Of the six endemic species of premontane forest amphibians described from the Quebrada de Oro area, we now know that two of them ( Duellmanohyla salvavida and Plectrohyla chrysopleura ) occur at La Liberacin. Given the robust na ture of the populations of these two treefrogs and the intactness of the premontane rainforest in this area, I remain hopeful that perhaps others of the endemic anurans that have undergone decline at Quebrada de Oro will be discovered s till resident at La Liberacin. Underestimated Salamander Diversity? Three species of salamanders were previo u sly known from RVS Texguat, Bolitoglossa porrasorum sensu lato Nototriton barbouri s.l. and Oedipina gephyra s.l. each of which is considered endemic to two or mo re isolated localities in northern

PAGE 125

125 Honduras (McCranie & Wilson 2002). Bolitoglossa porrasorum is known from the type locality at Pico Pijol, Montaa Macuzal, Texguat, and Pico Bonito; however, only the Pico Bonito population has been included previo u sly i n phylogenetic studies (Parra Olea et al. 2004). Nototriton barbouri is considered to have the same distribution as B. porrasorum whereas O. gephyra is known from its type locality at La Fortuna and from a single specimen from Pico Bonito (McCranie & Wils on 2002). Both N. barbouri and O. gephrya have been shown to demonstrate species level divergence between the Texguat and Pico Bonito populations (Garca Pars & Wake 2000), and I supplement this existing data with newly collected samples representing bot h putative taxa in Chapter 4. In April 2008, a single specimen of Nototriton possessing the distinctive morphological traits of enlarged nostrils, syndactylous feet, and a contrasting pale venter was collected within leaf litter packed into a crevice in th e side of a small mesic canyon near La Fortuna (1,550 m elevation). This species, described as N. tomamorum by Townsend et al. (2010a), is analyzed genetically in Chapter 4 and reviewed morphologically in Chapter 5. Highly Endemic and Highly Endangered Th e leeward side of RVS Texguat (herein referred to as Yoro Texguat) is highly imperiled due to continued illegal logging of valuable hardwoods and forest clearing for subsistence agriculture, which I witnessed firsthand in April 2008 (Townsend et al. 2010 a). Despite the rapid advancement of deforestation in Yoro, little exploration of the virtually unknown Atlntida side of the refuge has been conducted to date. I led three expeditions to the windward side of RVS Texguat (Atlntida Texguat) during 2010; these visits were limited to 12 days (10 21 June 2010) and seven days (26 July 2 August 2010) in the vicinity of La Liberacin (15.53N/87.29W; camp was established

PAGE 126

126 at 1,030 m elevation), and three days (10 12 November 2010) in the vicinity of San Jos de Texguat (15.52N/87.45W), yet we recorded 46 species (Townsend et al., submitted A) compared to 40 species amassed during the approximately two decade span of limited study of Yoro Texguat (Wilson & McCranie 2004 b ; Townsend et al. 2010a). This pattern suggests that herpetofaunal diversity and endemism demonstrated on the Yoro side will likely be exceeded on the windward side. The 2010 work in this area, along with past work on the leeward slope of RVS Texguat (summarized in Townsend et al. 2010a) and in Parque Nacional Pico Bonito (summarized in McCranie & Castaeda 2005), demonstrates that 29 of the 93 endemic species (close to one third) of amphibians and reptiles reported from Honduras by Townsend & Wilson ( 2010) occur within these three areas of t he same cordillera (Townsend et al., In press A). At least 10 species of amphibians and reptiles are Texguat restricted endemics, and RVS Texguat is home to 14 more species endemic to the Chorts Block (Townsend et al., In press A). Given the preliminar y successes of efforts towards documenting and promoting the conservation of herpetofaunal diversity in RVS Texguat, I believe that the serious problems facing the reserve can be addressed through continued efforts in the coming years to provide enduring protection for this vitally important component of the Honduran patrimony. Cryptozoic Snake Diversity Cryptozoic snake taxa include some of the most species rich groups of snakes, and their remarkable diversity is exceeded by their ability to remain conce aled and evade detection, even in relatively well studied field sites (Myers 2003; Stafford 2004; Townsend 2009). Genera such as Geophis Ninia and Tantilla are all cryptozoic in

PAGE 127

127 nature, typically small in size and often inhabit the interface between leaf litter and soil. Many of these secretive species are known only from one or a handful of specimens, yet are distinctive enough in terms of external morphology to be promptly recognized as distinct species (e.g., Campbell 1998; Stafford 2004; Townsend 2009 ). A New Species of Centipede S nake (genus Tantilla ) from La Liberacin The genus Tantilla currently consists of 6 3 described species (Townsend et al. In press B ), broadly distributed in all three of the major geographic sections of the Western Hemisphere (North America, Mesoamerica, and South America; Wilson 1999) including at least seven species in the Chorts Block (McCranie 2011b ) During survey work in July 2010 around La Liberacin on the windward side of RVS Texguat, we collected a single specimen of Tantilla possessing a set of distinctive morphological characteristics that stand out as unique within the genus. Based on having pale middorsal and/or lateral stripes and pale markings on the nape, we posit that this snake is assignable to the T. taen iata species group (sensu Wilson & Meyer 1971, Campbell 1998, Wilson & McCranie 1999), but represents a previously unknown taxon, which is described in a manuscript submitted for publication in August 2011. The new species differs significantly from all co ngeners on the basis of its dorsal body pattern, consisting of a pale middorsal stripe composed of narrow spots and confined to the middorsal row and a lateral coloration of pale spots on each of rows 1, 2, and 4. In addition, it possesses a middorsally in terrupted pale nuchal band and dark brown pigment on the lateral edges of the ventrals. The new species appears to have no close affinities within the T. taeniata group, and elucidation of its phylogenetic relationships will have to await a group or genus wide assessment, which, given the difficulties in securing fresh material of these taxa, might not occur in the near future.

PAGE 128

128 A Large New Species of B lindsnake ( Typhlops tycherus ) Blindsnakes of the genus Typhlops (Squamata: Typhlopidae) have a cosmopoli tan and primarily tropical distribution (McDiarmid et al. 1999), with the majority of diversity in the Western Hemisphere confined to the islands of the West Indies (Dixon & Hendricks 1979; Thomas & Hedges 2007). Four native species of Typhlops previously were known to occur in Mesoamerica ( Dixon & Hendricks 1979; Khler 200 8 ): T. costaricensis (Honduras to Costa Rica), T. microstomus (Yucatan Peninsula), T. stadelmani (western and northern Honduras), and T. tenuis (Veracruz, Mxico to Guatemala). A fifth typhlopid, the parthenogenetic Ramphotyphlops braminus has been introduced to scattered localities throughout Mesoamerica, primarily a round urban areas (Khler, 2008 ). Two species of blindsnakes ( T. costaricensis and T. stadelmani ) are known to occur in the Chorts Highlands. Both of these species occur in Honduras; Typhlops costaricensis is known from localities in the mesic lowlands of Departamento de Gracias a Dios and from mid elevation pine oak forest localities between Tegucigalpa and Parque Nacion al La Tigra in Departamento de Francisco Morazn ( McCranie et al. 200 6 ; Wilson et al. 1988), and T. stadelmani which is reported from mid elevation localities in the departments of A tlantda, Copn, and Yoro ( McCranie & Castaeda 2005; McCranie & Wilson 2 001). Typhlops stadelmani was, until recently, considered a junior synonym of T. tenuis (Dixon & Hendricks 1979), and was resurrected by McCranie & Wilson (2001) after collection of a large series that verified the distinctiveness of T. stadelmani from T. tenuis based primarily on differences in the number of scales between rostral and tail tip and in color.

PAGE 129

129 Figure 3 12 New species of Tantilla (Colubridae) and Typhlops (Typhlopidae). A) Dorsal, lateral, and central views of the head of adult male holo type (USNM 574000) of Tantilla sp. nov. 1,150 m elevation, La Liberacin de Texiguat B) sulcate and asulcate views of the hemipenis of USNM 574000 C) dorsal and lateral coloration of USNM 574000 D) subadult Typhlops tycherus sp. nov. from El Cedral, Mo nta a de Santa Brbara, 1,550 m. Photos J.H. Townsend.

PAGE 130

130 In January 2008, a single specimen of Typhlops was found freshly killed on an unpaved road between two small communities near the lower edge of intact cloud forest in Parque Nacional Montaa de Sant a Brbara, Honduras. This specimen differed from all other Middle American blindsnakes in its large size, in having 22 22 22 scales around the body, and by having a dark brownish gray dorsum with a well defined pale yellowish gray to immaculate white ventr al coloration. This specimen formed the basis for describing the species Typhlops tycherus which was confirmed in April 2011 by collec tion of a second specimen from the vicinity of the type locality by M Medina Flores (UNAH) Geophis damiani at La Libera cin The genus Geophis (Squamata: Colubridae: Dipsadinae) is currently comprised of 47 species of secretive semifossorial snakes distributed across nearly every terrestrial habitat from Mxico to northern South America (Downs 1967, Wilson & Townsend 2007, Townsend 2009). Six members of the genus are known to occur in the Chorts Block: Geophis damiani G. dunni G. fulvoguttatus G. hoffmanni G. nephodrymus and G. rhodogaster four of which ( G. damiani, G. dunni, G. fulvoguttatus and G. nephodrymus ) are e ndemic (Townsend 2009). Geophis damiani is one of the least known snakes in the Chorts Block and was only known from two adult specimens ( Wilson et al. 1998 McCranie & Castaeda 2004 ). On 29 July 2010, an adult male Geophis damiani ( USNM 573999 ) was col lected during survey work around La Liberacin, representing the third adult specimen and second adult male specimen known for this taxon. The snake was found at 22h30 while active in the bottom of a deep, trench like trail at 1,075 m elevation in moderate ly disturbed Premontane Wet Forest The snake had apparently fallen into and become trapped inside the trench,

PAGE 131

131 which was about 2 m deep at the collection site and over 3.5 m deep in some places This locality lies around 10 km NNW of the previously reporte d localities, and the elevation of the new record is 475 m below the previously known lowest elevational distribution attributed to this species, 1,550 m (based on UF 142543, an egg and embryo), and 605 m below the lowest verified elevation for the species (1,680 m; USNM 573999 ). All previously reported localities for G. damiani are in the Lower Montane Wet Forest formation In Townsend et al. (2010 d ), we provided morphological data and color notes for USNM 573999 and compared it to the two previously kno wn specimens of G. damiani Noteworthy Ninia Ninia pavimentata Ninia pavimentata is a small semifossorial snake reported from pine oak and cloud forest areas in central Guatemala, as well as from a single locality in northwestern Honduras (Smith & Campbel l 1996, Townsend et al. 2005). This taxon was previously considered a synonym of N. maculata (Peters 1861), until Smith & Campbell (1996) resurrected N. pavimentata (Bocourt 1883) to species level on the basis of non overlapping segmental count ranges and other morphological characteristics. The distributions of N. maculata and N. pavimentata are presently known to be separated by a roughly 315 airline km gap in Honduras, with N. pavimentata reaching its easternmost known distribution in the Sierra de Omoa, outside of Parque Nacional Cusuco, Departamento de Corts ( 15 .5 1 N, 88 .18 W ), 1 250 m elevation, and N. maculata its northernmost known locality at Quebrada Machn, Reserva de la Bisfera Ro Pltano, Departamento de Coln (15 .32 N, 85 .28 W), 540 m elevat ion (McCranie et al. 2001; Townsend et al. 2005). On 10 April 2008, J. Butler and I collected a female Ninia pavimentata (UF 152810 ) from under a rock at the edge

PAGE 132

132 Figure 3 13 Noteworthy cryptozoic snakes. A) Third adult specimen, and second male, Geop his damiani (USNM 573999 ), 1,075 m elevation, La Liberacin de Texiguat B) Ninia espinali a Chorts Highlands endemic recorded for the first time in Cerro Azul Membar C) Ninia pavimentata recorded for the first time in Refugio de Vida Silvestre Texgu at Photos J.H. Townsend. of a fragment of cleared cloud forest in Refugi o de Vida Silvestre Texiguat ( 15.44 N, 87 .30 W), 1 715 m elevation, Departamento de Yoro, Honduras (Townsend et al. 2009 b ). This habitat is similar to that of most N. pavimentata s pecimens, which are known from 1 120 1 825 m elevation in pine oak or cloud forest (Smith & Campbell 1996), as well as to the disturbed habitat (a shade cafetal ) where the other Honduran specimen originated. Examination of the specimen showed it to fall w ith the known

PAGE 133

133 range of variation in N. pavimentata (Smith & Campbell 1996; Townsend et al. 2005; Townsend et al. 2009 b ), providing further evidence for the specific status of this taxon. This record extends the known range of Ninia pavimentata approximatel y 65 airline km eas t from its known distribution. The new locality also narrows the geographic gap between N. pavimentata and its most similar congener and presumed sister taxon, N. maculata to approximately 250 airline km. Ninia espinali Ninia espinali is known to occur in western Honduras and northern El Salvador, where it inhabits highland rainforests from 1 580 2 270 m elevation (Khler 2008). On 9 July 2008, I. Luque and I collected an adult female N. espinali on a steep slope approximately 200 m ab ove the Ro de Varsovia ( 1 660 m elevation ) in Parque Nacional Cerro Azul Membar, extending the known range of this species approximately 85 km SSE of localities in Parque Nacional Cusuco, Deptartamento de Corts, and approximately 75 km N of localities in the vicinity of Guajiquiro, Departamento de La Paz (Luque Montes & Townsend 2009) The snake was active at night in an exposed root mass on a steep slope near the top of a ridge. Unidentified Salamander Populations As indicated a bove, fieldwork in Parque Nacional Montaa de Yoro and Refugio de Vida Silvestre Texguat resulted in collection of samples representing five populati ons of salamanders that could not be assigned unambiguously to a known species Besides these five entitie s, at least five other populations of salamanders were sa mpled that can not be unequivocally assigned to a known taxon. From Parque Nacional Cerro Azul Membar, a series of Nototriton and a single distinctively colored Bolitoglossa require additional attent ion, as do samples representing at least two forms of Oedipina from northern Nicaragua. In 2010 and 2011, fieldwork in Departamento de

PAGE 134

134 Olancho resulted in discovery of two previously unknown allopatric populations of Nototriton one from the Sierra de Agal ta and a second from the Sierra de Botaderos. T hese 10 populations of salamanders, as well as other allopatric populations of questionable taxonomic assignment (e.g., Bolitoglossa porrasorum and Nototriton barbouri from Refugio de Vida Silvestre Texguat), are the subject of molecular investigation and taxonomic evaluation in Chapter 4.

PAGE 135

135 Table 3 1 Summary of fieldwork undertaken in the Chorts Block, 2006 2011 DATES LOCALITIES PARTICIPANTS 3 Jun 18 Jun 2006 PN Montaa de Yoro: Cataguana JHT, Larry Davi d Wilson 2 10 Dec 2006 PN Cerro Azul Membar : Los Pinos Brian Campesa no, Lorraine Ketzler, Scott Travers, JHT, Steve Townsend PN La Tigra 8 20 Mar 2007 PN Montaa de Yoro: Cataguana Jason Butler, Lorraine Ketzler, Scott Travers, JHT, Larry Dav id Wilson, et al. RB Cerro Uyuca Jason Butler, Lorraine Ketzler, Scott Travers, JHT 7 29 Jun 2007 Biosfera Bosawas (7 localities) Lenin Obando, Javier Sunyer, Scott Travers, JHT, Larry David Wilson, et al. 13 21 Jul 2007 PN Cerro Azul Membar : Lo s Pinos Lorraine Ketzler, JHT, Larry David Wilson RB Cerro Uyuca RB Yerbabuena Cerro Zarciadero 11 23 Aug 2007 Leon Scott Travers, JHT, Katielynn Townsend Selva Negra Scott Travers, JHT, Katielynn Townsend 20 Jan 5 Feb 2008 PN Mo ntaa de Comayagua: La Ok Lorraine Ketzler, Ileana Luque Montes, JHT, Larry David Wilson PN Montaa de Santa Brbara: El Cedral Marcala San Pedro La Loma Los Naranjos Ileana Luque Montes, JHT, Larry David Wilson Montaa Macuzal I leana Luque Montes, JHT, Larry David Wilson Yeguare Valley Ileana Luque Montes, JHT, Larry David Wilson 4 20 Apr 2008 PN Cerro Azul Membar : Los Pinos Carlos Andino, Csar Cerrato Gabriela Diaz, Lorraine Ketzler, Ileana Luque Montes, Melissa Medina F lores, Aaron Mendoza, Wendy Naira, JHT, Larry David Wilson PN Cerro Azul Membar : Aldea Cerro Azul Csar Cerrato Ileana Luque Montes, Melissa Medina Flores, JHT, Larry David Wilson PN Montaa de Comayagua: Ro Negro Ileana Luque Montes, Melissa Medina Flores, JHT, Larry David Wilson RVS Texguat: La Fortuna Jason Butler, Lorraine Ketzler, Nathaniel Stewart, JHT, Larry David Wilson Montaa Macuzal 14 28 May 2008 PN Montaa de Comayagua: Ro Negro James Austin, Lorraine Ketzler, JHT, Larry Da vid Wilson RB Guajiquiro Csar Cerrato Lorraine Ketzler, Ileana Luque Montes, JHT, Larry David Wilson Los Naranjos James Austin, Lorraine Ketzler, JHT, Larry David Wilson 12 Jun 29 Jul 2008 PN Celaque Lorraine Ketzler, Ileana Luque Montes, JHT, Larry David Wilson PN Cerro Azul Copn: Quebrada Grande Ileana Luque Montes, JHT, Larry David Wilson PN Pico Pijol: Quebrada Las Payas PN Montaa de Comayagua: Ro Negro PN Cerro Azul Membar : Aldea Cerro Azul/Varsovia Ileana Luque Montes, JHT RB Gisayote Lorraine Ketzler, Ileana Luque Montes, Melissa Medina Flores, JHT, Larry David Wilson Copn Ruinas Lorraine Ketzler, Ileana Luque Montes, Melissa Medina Flores, JHT, Larry David Wilson La Esperanza area (3 localities) Lorraine Ketzler, Ileana Luque Montes, JHT, Larry David Wilson 14 Aug 1 Oct 2008 PN Cerro Azul Membar : Aldea Cerro Azul/Varsovia Ileana Luque Montes, JHT PN Cusuco Csar Cerrato Ileana Luque Montes, Melissa Medina Flores, JHT, Larry David Wilson

PAGE 136

136 Table 3 1. Continued TRIP DATES LOCALITIES PARTICIPANTS PN Montaa de Yoro: above Guaymas Ileana Luque Montes, JHT, Larry David Wilson PN Pico Pijol: Pino Alto Csar Cerrato Ileana Luque Montes, JHT, Larry David Wilson RB El Pital Csar Cerrato Melissa Medina Flores, Larry David Wilson RB Gisayote RB Mixicuri Erandique La Esperanza area 10 20 April 2009 PN Montaa de Comayagua: Ro Negro Sergio Gonzalez, Christina Martin, Mario Solis, JHT, Rony Valle, Christopher Wolf 25 Nov 6 Dec 200 9 PN Cerro Azul Membar : Los Pinos Csar Cerrato Vladlen Henriquez, JHT PN Cusuco PN Pico Bonito 1 26 Jun 2010 RVS Texguat: La Liberacin Benjamin Atkinson, Csar Cerrato, Luis Herrera, Mayron McKewy Meja, JHT, Larry David Wilson, et al JB Lancetilla Benjamin Atkinson, Csar Cerrato, Luis Herrera, Mayron McKewy Meja, Ciro Navarro, JHT 20 Jul 21 Aug 2010 PN Cerro Azul Membar : Los Pinos Anne Donnelly, Matthew Donnelly, Ileana Luque Montes, JHT RVS Texguat: La Liberacin Levi Gr ay, Luis Herrera, Melissa Medina Flores, Alexander Stubbs, JHT, etc. JB Lancetilla Roatn Anne Donnelly, Matthew Donnelly, Yensi Flores, Ileana Luque Montes, Melissa Medina Flores, Sandy Pereira, JHT Utila Anne Donnelly, Matthew Donnelly, Ilean a Luque Montes, Melissa Medina Flores, Sandy Pereira, JHT 5 16 Nov 2010 PN Cerro Azul Membar : Los Pinos James Austin, Luis Herrera, Melissa Medina Flores, JHT PN Montaa de Santa Brbara James Austin, Luis Herrera, JHT, Alicia Ward, et al. San Jos de Texguat James Austin, Luis Herrera, JHT 7 22 April 2011 PN Montaa de Botaderos Christopher Begley, Mark Bonta, Robert Hyman, David Medina, Melissa Medina Flores, Onn Reyes, Fito Steiner, JHT PN Pico Bonito Robert Hyman, David Medina, Melissa M edina Flores, Fito Steiner, JHT RB Colibr Esmeralda Robert Hyman, Fito Steiner, JHT Montaa deJacaleapa Mark Bonta, Onn Reyes, JHT Rio Grande, Valle de Agalta Christopher Begley, Mark Bonta, Robert Hyman, David Medina, Melissa Medina Flores, Onn R eyes, Fito Steiner, JHT

PAGE 137

137 Table 3 2 Conservation status and physiographic distribution of the native non marine herpetofauna of the Chorts Highlands. Data primarily sourc ed from Wilson & Johnson (2010) and supplemented by Acevedo et al. (2010), McCrani e (2011 a ), Sunyer & Khler (2010), Townsend & Wilson (2010 a ), and my own observations. Taxa followed by an asterisk (*) indicated new taxa described or being described through the course of this dissertation; details of taxonomic assignment of these specie s are provided in Chapter 3. IUCN Red List status follows is from the IUCN (2011; www.iucnredlist.org ) when available, with other sources indicated by footnotes Elevational Distributions are range wide and not co nfined to the Chorts Highlands. Conservation Status Scores (CSS) are from Wilson & Townsend (2010 a ), and Environmental Vulnerability Scores (EVS) are sources cited above. Species are allocated to one of two General Distribution categories: C B = endemic to the Chorts Block WS = widespread outside Chorts Block Physiographic distribution is related to the Chorts Highl ands ( Chapter 1 provides details of each province), and includes: CL = Caribbean Lowlands physiographic province, CV = Caribbean versant in termontane valleys, NC = Northern Cordillera of the Serrana CC = Central Cordillera of the Serrana SC = Southern Cordillera of the Serrana PL = Pacific Lowlands physiographic province, PV = Pacific versant intermontane valleys, IB = Islas de la Baha ; a plus (+) indicates the taxon is found within the province Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB GYMNOPHIONA Caeciliaidae (2) Dermophis mexicanus Vuln erable A2ac 1 20 12 0 1500 WS + + + Gymnopis multiplicata Least Concern 1 14 12 0 1400 WS + + CAUDATA Plethodontidae (43) Bolitoglossa carri Critically Endangered B1ab(iii)+2ab(iii) 1 3 17 1840 2070 CB + Bo litoglossa cataguana* Critically Endangered B1ab(iii)+2ab(iii) 2 3 16 1800 2080 CB + Bolitoglossa celaque Endangered B1ab(iii) 1 3 16 1900 2820 CB + Bolitoglossa conanti Endangered B1ab(iii) 1 5 14 950 2010 CB + +

PAGE 138

138 Table 3 2 Con tinued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Bolitoglossa decora Critically Endangered B1ab(iii)+2ab(iii) 1 3 17 1430 1550 CB + Bolitoglossa diaphora Critically Endangered B 2ab(iii) 1 3 16 1470 2200 CB + Bolitoglossa dofleini Near Threatened 1 10 14 100 1550 WS + + + Bolitoglossa dunni Endangered B1ab(iii) 1 5 14 1020 1600 CB + Bolitoglossa heiroreias En d angered B1ab(iii) 1 5 15 1840 2300 CB + Bolitoglossa longissima Critically Endangered B1ab (iii) 1 3 17 1840 2240 CB + Bolitoglossa mexicana Least Concern 1 14 9 0 1900 WS + + + + Bolitoglossa nympha Near Threatened 3 6 3 12 3 30 1400 CB + + + Bolitoglossa oresbia Critica ll y Endangered B1ab(iii)+2ab(iii) 1 3 17 1560 1880 CB + Bolitoglossa porrasorum Endangered B1ab(iii) 1 4 15 980 1920 CB + + Bolitoglossa striatula Least Concern 1 10 14 2 1055 WS + Bolitoglossa synoria Critically Endangered B1ab(ii i) 1 4 15 2150 2715 CB + Cryptotriton monzoni Critically Endangered B1ab(iii) 1 3 17 4 1570 CB + Cryptotriton nasalis Endangered B1ab(iii) 1 5 15 1220 2200 CB +

PAGE 139

139 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Cryptotriton wakei Critically Endangered B1ab(iii) 1 3 17 4 1150 CB + Dendrotriton sanctibarbarus Endangered B1ab(iii) 2 [Vulnerable D2 1 ] 3 16 1830 2744 CB + Nototriton barbour i Endangered B1ab(iii) 1 3 15 1530 1920 CB + Nototriton brodiei Critically Endangered B1ab(iii) 1 3 17 875 1140 CB + Nototriton lignicola Critically Endangered B1ab(iii) 1 3 17 1760 2020 CB + Nototriton limnospectator Endangere d B1ab(iii) 1 3 16 1640 1980 CB + Nototriton picucha* Critically Endangered B1ab(iii) 3 3 3 17 3 1890 1920 CB + Nototriton saslaya Vulnerable D2 1 3 17 5 1280 1500 CB + Nototriton stuarti Data Deficient 1 3 17 4 744 CB + Nototri ton tomamorum* Critically Endangered B1ab(iii)+2ab(iii) 2 3 3 17 3 1550 CB + Nototriton sp A (Pico Bonito)* Critically Endangered B1ab(iii) 3 3 3 17 3 1210 1540 CB + Nototriton sp B (Texiguat)* Critically Endangered B1ab(iii)+2ab(iii) 3 3 3 17 3 142 0 1800 CB +

PAGE 140

140 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Nototriton sp. C (Botaderos)* Critically Endangered B1ab(iii) 3 3 3 17 3 1700 1735 CB + Oedipina elongata Least Concern 1 9 15 10 770 WS + + Oedipina gephyra Critically Endangered B1ab(iii)+ 2ab(iii) 3 [Endangered B1ab(iii) 1 ] 3 16 1580 1810 CB + Oedipina ignea Data Deficient 1 7 14 1000 2000 WS + Oedipina kasios Endangered B1ab (iii) 2 4 15 950 1920 CB + Oedipina koehleri* Endangered B1ab(iii) 3 3 3 15 3 628 945 CB + Oedipina leptopoda Endangered B1ab(iii) 2 3 15 700 1300 CB + Oedipina nica* Endangered B1ab(iii) 3 3 3 15 3 1360 1660 CB + Oedipina quad ra Vulnerable B1ab(iii) 2 3 15 70 540 CB + + + Oedipina petiola* Critically Endangered B1ab(iii) 3 3 3 17 3 1580 CB + Oedipina stuarti Data Deficient 1 6 15 0 1000 CB + + Oedipina taylori Least Concern 1 8 15 140 1140 WS + Oedipi na tomasi Critically Endangered B2ab(iii) 1 3 16 1800 CB + ANURA Bufonidae (10) Incilius campbelli Near Threatened 1 11 10 70 1200 WS + +

PAGE 141

141 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Dis tribution (m) General Distribution CL CV NC CC SC PL PV IB Incilius coccifer Least Concern 1 15 6 0 1350 WS + + + + + Incilius ibarrai Endangered B1ab(iii) 1 7 11 1500 1730 WS + + + Incilius leucomyos Endangered B1ab(iii) 1 6 11 0 1600 CB + + + Incilius luetkenii Least Concern 1 16 7 0 1300 WS + + + + + Incilius porteri Endangered B1ab(iii) 3 [Data Deficient 1 ] 3 13 1524 1890 CB + + Incilius valliceps Least Concern 1 26 5 0 2000 WS + + + + + + Rhaebo haematiticus Least Conc ern 1 12 11 0 1300 WS + Rhinella chrysophora Endangered A2ac & B1ab(iii,v) 1 4 12 750 1760 CB + Rhinella marina Least Concern 1 35 5 0 2000 WS + + + + + + + + Centrolenidae (8) Cochranella granulosa Least Concern 1 11 12 0 1500 WS + Espadarana prosoblepon Least Concern 1 14 12 0 1900 WS + Hyalinobatrachium chirripoi Near Threatened 1 9 12 0 700 WS + Hyalinobatrachium colymbiphyllum Least Concern 1 10 12 0 1710 WS + Hyalinobatrachium fleischmanni Least Concern 1 27 9 0 1730 WS + + + + + Sachatamia albomaculata Least Concern 1 12 12 0 1500 WS + + + Teratohyla pulverata Least Concern 1 11 12 0 950 WS + + + Teratohyla spinosa Least Concern 1 9 13 0 560 WS + Craugastoridae (29 ) Craugastor anciano Critically Endangered B1ab(iii,v)+2ab(iii,v) 1 4 15 1400 1840 CB + Craugastor aurilegulus Endangered B1ab(iii,v)+2ab(iii,v) 1 6 14 50 1550 CB + + Craugastor bransfordii Least Concern 1 10 11 5 20 1535 WS +

PAGE 142

142 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Craugastor chac Near Threatened 1 8 14 0 1000 WS + Craugastor charadra Endangered B1ab(iii, v) 1 7 13 30 137 0 CB + + Craugastor chrysozetetes EXTINCT 1 3 17 880 1130 CB + Craugastor coffeus Critically Endangered B1ab(iii) + 2ab(iii) 1 3 17 1000 CB + Craugastor cruzi Critically Endangered A2ace, B1ab(iii,v) +2ab(iii,v) 1 3 17 1520 CB + Craugastor cyanochthebius Critically Endangered B1ab( iii)+2ab(iii)2 [Near Threatened 1 ] 3 17 900 1200 CB + Craugastor emleni Critically Endangered A2ace, B2ab(v) 1 3 14 800 2000 CB + Craugastor epochthidius Critically Endangered A3ce 1 5 15 150 1450 CB + Craugastor fecundus Critically Endangered A2ace 1 5 15 200 1260 CB + Craugastor fitzingeri Least Concern1 14 13 1 1520 WS + + Craugastor laevissimus Endangered A2ace 1 11 8 0 2000 CB + + + + + Craugastor lat iceps Near Threatened 1 13 14 10 1600 WS + + Craugastor lauraster Endangered B1ab(iii) 1 6 14 40 1200 CB + + + Craugastor megacephalus Least Concern 1 10 14 1 1200 WS + Craugastor merendonensis Critically Endangered A2ace, B1ab(v)+ 2 ab(v) 1 3 17 150 200 CB +

PAGE 143

143 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Craugastor milesi Critically Endangered A2ae 1 4 15 1050 1720 CB + Craugastor mi mus Least Concern 1 9 13 15 700 WS + Craugastor nefrens Data Deficient 1 3 17 4 800 1000 CB + Craugastor noblei Least Concern 1 11 13 4 1200 WS + + + Craugastor olanchano Critically Endangered A2ace 1 3 16 1180 1350 CB Craugast or omoaensis Critica lly Endangered A2ace, B1ab(iii) 1 3 16 760 1150 CB + Craugastor pechorum Endangered B1ab(iii) 1 5 15 150 680 CB + Craugastor rhodopis Least Concern 1 16 14 0 1370 WS + Craugastor rostralis Near Threatened 1 6 14 850 1800 WS + + Craugastor saltuarius Critically Endangered A2ace 1 3 16 1550 1800 CB + Craugastor stadelmani Critically Endangered A2ace 1 4 15 1125 1900 CB + + Eleutherodactylidae (1) Diasporus diastema Least Concer n 1 11 14 0 1620 WS + Hylidae (35) Agalychnis callidryas Least Concern 1 17 10 0 1200 WS + + + + + Agalychnis moreletii Critically Endangered A3e 1 19 13 200 2130 WS + + + Agalychnis saltator Least Concern 1 9 13 0 819 WS + Anotheca spinosa Least Concern 1 11 15 95 2068 WS + Bromeliohyla bromeliacia Endangered A2ace 1 8 15 900 1790 WS + Cruziohyla calcarifer Least Concern 1 9 12 30 820 WS + Dendropsophus ebraccatus Least Concern 1 16 11 0 1320 WS + Dendropsophus microcephalus Least Concern 1 19 5 0 1200 WS + + + + + + +

PAGE 144

144 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Duellmanohyla salvavida Criticall y Endangered B2ab(iii,v) 1 5 12 90 1400 CB + Duellmanohyla soralia Critically Endangered B2ab(iii,v) 1 7 10 40 1570 CB + Ecnomiohyla miliaria Vulnerable; B1ab(iii) 1 9 15 0 1330 WS + Ecnomiohyla salvaje Critically Endangered B1ab( iii) 1 5 16 1370 1520 CB + Exerodonta catracha Endangered B1ab(iii)+2ab(iii) 1 3 13 1800 2160 CB + + Isthmohyla insolita Critically Endangered B1ab(iii)+2ab(iii) 1 3 16 1550 CB + Isthmohyla melacaena Critically Endangered B2ab(iv) 2 [Near Threatened 1 ] 3 16 1550 CB + Plectrohyla chrysopleura Critically Endangered A2ace, B1ab(iii,v)+ 2ab(iii,v) 1 4 13 930 1550 CB + Plectrohyla dasypus Critically Endangered A2ace, B1ab(iii,v) +2ab(iii,v) 1 3 13 1410 1990 CB + Plectrohyla exquisita Critically Endangered A3e 1 3 13 1490 1680 CB +

PAGE 145

145 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Plectrohyla guatemalensis Critically Endang ered A3e 1 11 9 900 2800 WS + + + Plectrohyla hartwegi Critically Endangered A3e 1 9 12 925 2700 WS + Plectrohyla matudai Vulnerable B1ab(iii) 1 8 10 700 2300 WS + + Plectrohyla psiloderma Endangered B1ab(iii) 1 4 12 2400 2530 CB + Ptychohyla euthysanota Near Threatened 1 10 11 4 200 2200 WS + Ptychohyla hypomykter Critically Endangered A3e 1 11 9 340 2070 CB + + + Ptychohyla salvadorensis Endangered B1ab(iii) 1 8 11 700 2050 CB + Ptychohyla spinipollex End angered B1ab(iii)+2ab(iii) 1 6 11 160 1580 CB + Scinax boulengeri Least Concern 1 11 11 0 700 WS + Scinax staufferi Least Concern 1 25 5 0 1530 WS + + + + + + + + Smilisca baudinii Least Concern 1 30 4 0 1925 WS + + + + + + + + Smilisca ph aeota Least Concern 1 11 10 0 1116 WS + + Smilisca sordida Least Concern 1 13 11 0 1525 WS + Tlalocohyla loquax Least Concern 1 18 6 0 1585 WS + + + + + Tlalocohyla picta Least Concern 1 15 9 0 1300 WS + + Trachycephalus venulosus Least Concern 1 25 5 0 1610 WS + + + Triprion petasatus Least Concern 1 12 12 0 740 WS + Leiuperidae (1) Least Concern 1 Engytomops pustulosus Least Concern 1 24 6 0 1540 WS + + + + + Leptodactylidae (4) Least Concern 1 Leptodactylus fragilis Least Concern 1 25 6 0 1700 WS + + + + + + + Leptodactylus melanonotus Least Concern 1 25 6 0 1440 WS + + + + + + Leptodactylus savagei Least Concern 1 15 11 0 1200 WS +

PAGE 146

146 Table 3 2 Continued. Taxon IUCN Red Lis t Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Leptodactylus silvanimbus Critically Endangered B2ab(iii,v) 1 4 13 1470 2000 CB + Microhylidae (3) Gastrophryne elegans Least Concern 1 13 11 0 1500 WS + Hypopachus barberi Vulnerable B1ab(iii) 1 10 11 1300 2500 WS + Hypopachus variolosus Least Concern 1 31 6 0 2200 WS + + + + + Ranidae (6) Lithobates brownorum Least Concern 1 18 3 0 1200 WS + + + + Lithob ates forreri Least Concern 1 20 8 0 1960 WS + + Lithobates maculatus Least Concern 1 16 6 40 3000 WS + + + + Lithobates vaillanti Least Concern 1 21 7 0 990 WS + + + + Lithobates warszewitschii Near Threatened 1 13 11 0 2500 WS + L ithobates sp. nov. Endangered B1ab(iii)+2ab(iii) 3 CB + Rhinophrynidae (1) Rhinophrynus dorsalis Least Concern 1 18 9 0 700 WS + + Strabomantidae (2) Least Concern 1 Pristimantis cerasinus Least Concern 1 9 14 19 680 WS + + Pristimantis ridens Least Concern 1 12 12 0 1600 WS + + + REPTILIA TESTUDINES Chelydridae (2) Chelydra acutirostris Data Deficient 3 12 13 0 1164 WS + + Chelydra rossignonii Vulne rable A2d 1 11 14 0 660 WS + + Emydidae (1) Trachemys venusta Near Threatened 2 26 12 0 650 WS + + Geoemydidae (4) Rhinoclemmys annulata Near Threatened 2 [Lower risk/least concern 1 ] 11 13 2 920 WS

PAGE 147

147 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Rhinoclemmys areolata Near Threatened 1 12 12 0 600 WS + Rhinoclemmys funerea Near Threatened 2 [Lower risk/least conce rn 1 ] 7 16 2 600 WS Rhinoclemmys pulcherrima Near Threatened 2 19 9 0 1480 WS + + + Kinosternidae (3) Kinosternon leucostomum Least Concern 2 21 9 0 1500 WS + + Kinosternon scorpioides Least Concern 2 24 9 0 1500 WS + + Staurotypus triporcatus Near Threatened 2 [Lower risk/least concern 1 ] 10 15 0 300 WS + CROCODILIA Alligatoridae (1) Caiman crocodilus Least Concern 2 [Lower risk/least concern 1 ] 15 16 0 300 WS + + Crocod ylidae (1) Crocodylus acutus Vulnerable A1ac 1 22 13 0 650 WS + + + + SQUAMATA: LIZARDS Anguidae (6) Abronia montecristoi Endangered B2ab(iii) 2 [Endangered B1+2c 1 ] 4 15 1370 CB + + Abronia salvadoren sis Endangered B2ab(iii) 2 3 16 2020 2125 CB + Celestus bivittatus Near Threatened 2 7 13 1510 1980 CB + Celestus montanus Endangered B1ab(iii) 2 5 14 915 1372 CB +

PAGE 148

148 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevat ional Distribution (m) General Distribution CL CV NC CC SC PL PV IB Celestus scansorius Endangered B2ab(iii) 2 [Near Threatened 1 ] 3 14 1550 1590 CB + + Mesaspis moreletii Least Concern 2 13 13 1450 3060 WS + + + Corytophanidae (7) Basiliscus plumifrons Least Concern 2 11 13 0 780 WS + Basiliscus vittatus Least Concern 2 25 7 0 1500 WS + + + + + + + + Corytophanes cristatus Least Concern 2 20 11 0 1640 WS + + Corytophanes hernandesii Least Concern 2 14 12 0 1400 WS + Corytophanes percarinatus Vulnerable B1ab(iii) 2 11 14 200 2200 CB + Laemanctus longipes Least Concern 2 17 9 0 1200 WS + + Laemanctus serratus Least Concern 1 17 12 0 1600 WS + Eublepharidae (1) Coleonyx mi tratus Least Concern 2 16 10 0 1435 WS + + + + + Gymnophthalmidae (1) Gymnophthalmus speciosus Least Concern 2 24 8 0 1320 WS + + + + Helodermatidae (1) Heloderma horridum Near Threatened 4 [Least Concern 1 ] 11 14 4 100 15 30 WS + Iguanidae (8) Ctenosaura bakeri Critically Endangered B1ab(i,ii,iii,v)+ 2ab(i,ii,iii,v) 1 3 19 0 5 CB + Ctenosaura flavidorsalis Endangered B1ab(iii,v)+2ab(iii,v) 1 8 13 370 750 CB + + Ctenosaura melanosterna Critically Endangered B1ab(iii,v) 1 5 17 0 300 CB + +

PAGE 149

149 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Ctenosaura oedirhina Critically Endangered B1ab(iii) 1 3 18 0 20 CB + Ctenosaura palearis Endangered B1ab(i,ii,iii,iv,v)+ 2ab(i,ii,iii,iv,v) 1 3 17 4 150 700 CB + Ctenosaura praeocularis Endangered B1ab(iii,v) 3 [Data Deficient 1 ] 3 3 17 3 800 1000 CB + Ctenosaura quinquecarinata Endangered B1 ab(iii,v)+2ab(iii,v) 1 5 16 5 0 250 WS + + Ctenosaura similis Least Concern 1 22 11 0 1320 WS + + + + + + + + Iguana iguana Least Concern 2 26 12 0 1000 WS + + + + + + + + Phrynosomatidae (3) Sceloporus malachiticus Least Concern 2 16 8 540 3800 WS + + + Sceloporus variabilis Least Concern 2 25 7 0 1500 WS + + + + Sceloporus squamosus Least Concern 2 14 10 0 2500 WS + + + Phyllodactylidae (3) Phyllodactylus palmeus Endangered B2ab(iii) 2 3 15 0 30 CB + Phyllodactylus tuberculosus Least Concern 2 23 10 0 1230 WS + + + Thecadactylus rapicauda Least Concern 2 21 10 0 1052 WS + + + + Polychrotidae: (38) Anolis allisoni Least Concern 2 7 13 0 30 WS + Anolis amplisquamosus Endangered B2ab(iii, v) 1 3 16 1530 2200 CB + Anolis beckeri Least Concern 2 22 11 0 1780 WS + + Anolis bicaorum Endangered B2ab(iii) 2 3 16 0 20 CB + Anolis biporcatus Least Concern 2 23 10 0 2000 WS + + + + Anolis capito Leas t Concern 2 19 11 0 1250 WS + + + +

PAGE 150

150 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Anolis carpenteri Least Concern 1 6 13 5 4 682 WS + Anolis crassulus Least Concern 2 11 13 1200 3200 WS + Anolis cupreus Least Concern 2 13 9 0 1435 WS + Anolis cusuco Endangered B1ab(iii) 1 3 16 1550 1935 CB + Anolis heteropholidotus Endangered B2ab(iii) 2 4 14 1860 2200 CB + Anolis johnmeyeri En dangered B1ab(iii) 2 3 15 1340 1825 CB + Anolis kreutzi Critically Endangered B1ab(iii)+2ab(iii) 2 3 16 1670 1690 CB + Anolis laeviventris Least Concern 2 17 9 500 2000 WS + + + Anolis lemurinus Least Concern 2 26 9 0 2000 WS + + + + Anolis limifrons Least Concern 2 13 12 0 1340 WS + Anolis lionotus Least Concern 1 5 13 20 1200 WS + Anolis loveridgei Endangered B1ab(iii) 1 6 14 550 1600 CB + Anolis morazani* Critically Endangered B1ab(iii) 2 3 16 1780 2150 C B + Anolis muralla Vulnerable D2 1 ; Critically Endangered B1ab(iii)+2ab(iii) 2 3 15 1440 1740 CB + Anolis ocelloscapularis Endangered B1ab(iii) 2 4 15 1150 1450 CB + Anolis petersii Vulnerable B1ab(iii) 2 14 13 200 2130 WS + Anolis pijolensis Critically Endangered B1ab(iii) 2 4 14 1180 2050 CB +

PAGE 151

151 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Anolis purpurgularis Endangered B1ab(iii)+ 2ab(iii) 2 3 15 1550 2040 CB + Anolis quaggulus Least Concern 2 8 12 0 1350 WS + Anolis roatanensis Endangered B1ab(iii)+2ab(iii) 2 3 15 0 30 CB + Anolis rodriguezii Least Concern 2 16 10 0 2000 WS + + Anolis rubribarbaris Cr itically Endangered B1ab(iii) 2 3 16 1700 CB + Anolis sminthus Endangered B1ab(iii)+2ab(iii) 2 [Data Deficient 1 ] 4 15 1450 2200 CB + Anolis tropidonotus Least Concern 2 20 5 0 1900 WS + + + + + + Anolis uniformis Least Concern 2 13 11 0 1 370 WS + Anolis unilobatus Least Concern 2 26 7 0 1200 WS + + + + Anolis utilensis Critically Endangered B1ab(iii) 2 3 16 0 5 CB + Anolis wampuensis Endangered B2ab(iii) 2 3 16 95 110 CB + Anolis wermuthi Vulnerable B1ab(ii i) 5 3 15 5 1230 1660 CB + Anolis yoroensis Endangered B1ab(iii)+2ab(iii) 2 4 14 1180 1600 CB + Anolis zeus Endangered B1ab(iii) 2 5 14 90 900 CB + + Polychrus gutturosus Least Concern 2 12 12 6 700 WS + Scincidae (6) Mabuya unimarginata Least Concern 2 27 7 0 1800 WS + + + + + + + Mesoscincus managuae Least Concern 2 10 12 0 920 WS + Plestiodon sumichrasti Least Concern 2 16 11 0 1000 WS + + Sphenomorphus assatus Least Concern 2 16 13 0 2500 WS +

PAGE 152

152 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Sphenomorphus cherriei Least Concern 2 23 7 0 1860 WS + + + + + + Sphenomorphus incertus Vulnerable B1ab(iii) 2 6 12 1350 1670 WS + Sphaerodactylidae (5) Gonatodes albogularis Least Concern 2 20 10 0 1000 WS + + + + Sphaerodactylus dunni Vulnerable B1ab(iii) 2 [Least Concern 1 ] 4 14 60 230 CB + + Sphaerodactylus glaucus Least Conc ern 2 14 13 0 1000 WS + Sphaerodactylus millepunctatus Least Concern 2 19 7 0 1000 WS + + + + + + Sphaerodactylus rosaurae Endangered B2ab(iii) 2 3 15 0 20 CB + Teiidae (5) Ameiva festiva Least Concern 2 20 10 0 1400 WS + + + + Ameiva undulata Least Concern 2 27 7 0 1800 WS + + + Aspidoscelis deppii Least Concern 1 22 8 0 1200 WS + + + + Aspidoscelis motaguae Least Concern 1 12 9 175 1200 WS + + Cnemidophorus lemniscatus Least Concern 2 9 12 0 1000 WS + Xantusiidae (2) Lepidophyma flavimaculatum Least Concern 2 19 11 0 1400 WS + + + Lepidophyma mayae Vulnerable B1ab(iii)2 7 13 100 800 + SQUAMATA: SNAKES Anomalepididae (1) Anomalepis mexic anus Data Deficient 2 8 11 5 500 WS + Boidae (2) Boa constrictor Least Concern 2 32 8 0 1500 WS + + + + + + + + Corallus annulatus Least Concern 2 9 11 0 400 WS + Colubridae (110) Adelphicos quadrivirgatum Leas t Concern 2 [Data Deficient 1 ] 19 8 0 1740 WS + + + + + + Amastridium sapperi Least Concern 2 18 12 100 1600 WS + Chironius grandisquamis Least Concern 2 12 12 0 1600 WS + + +

PAGE 153

153 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevati onal Distribution (m) General Distribution CL CV NC CC SC PL PV IB Clelia clelia Least Concern 2 18 11 0 1000 WS + + + + Coniophanes bipunctatus Least Concern 2 18 11 0 1000 WS + + Coniophanes fissidens Least Concern 2 28 9 0 2200 WS + + + + + Coniophanes imperialis Least Concern 1 19 11 0 2000 WS + + + + + Coniophanes piceivittis Least Concern 1 23 11 0 1305 WS + + + Conophis lineatus Least Concern 1 22 9 0 1500 WS + + + + Crisantophis nevermanni Least Concern 2 11 14 0 138 5 WS + Dendrophidion clarkii Least Concern 2 17 12 30 1500 WS + + + Dendrophidion percarinatum Least Concern 2 13 12 4 1200 WS + + + Dendrophidion vinitor Least Concern 1 18 13 15 1500 WS + + Dipsas bicolor Least Concern 2 8 11 4 1100 WS + + Drymarchon melanurus Least Concern 1 35 9 0 2500 WS + + + + + + + + Drymobius chloroticus Vulnerable; B1ab(iii) 2 [Least Concern 1 ] 18 11 500 2500 WS + + + Drymobius margaritiferus Least Concern 2 33 7 0 2000 WS + + + + + + + Dr ymobius melanotropis Least Concern 1 8 14 0 1400 WS + + Enuliophis sclateri Least Concern 2 10 11 0 1235 WS + Enulius bifoveatus Critically Endangered B1ab(iii) 2 3 15 0 10 CB + Enulius flavitorques Least Concern 2 26 6 0 3000 WS + + + + + Enulius roatanensis Endangered B1ab(iii)+2ab(iii) 2 3 15 0 10 CB + Erythrolamprus mimus Least Concern 2 12 12 70 1400 WS + + Ficimia publia Least Concern 2 18 11 0 1000 WS + Geophis damiani Critically Endangered B1ab(ii i)+2ab(iii) 2 3 15 1075 1750 CB + Geophis dunni Data Deficient 1 3 16 5 900 CB +

PAGE 154

154 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Geophis fulvoguttatus Enda ngered B2ab(iii) 2 6 12 1680 2200 CB + + Geophis hoffmanni Least Concern 2 14 12 18 670 WS + + + + Geophis nephodrymus Endangered B2ab(iii) 2 3 14 1560 1580 C + Geophis rhodogaster Endangered B1ab(iii) 2 [Least Concern 1 ] 8 12 1480 2600 WS + Hydromorphus concolor Least Concern 2 16 9 1 1500 WS + + + Imantodes cen ch oa Least Concern 2 30 6 0 2063 WS + + + + Imantodes gemmistratus Least Concern 2 26 10 2 1435 WS + + + Imantodes inornatus Least Concern 1 12 10 5 1450 WS + + + Lampropeltis triangulum Least Concern 2 36 9 0 2500 WS + + + + + + + Leptodeira nigrofasciata Least Concern 2 18 10 0 1300 WS + + + Leptodeira rhombifera Least Concern 3 28 8 0 2000 WS + + + + + + Leptodeira septentrionalis Least Concern 2 33 9 0 2000 WS + + + + + Leptodrymus pulcherrimus Least Concern 2 14 10 10 1300 WS + + + + + Leptophis ahaetulla Least Concern 2 24 8 0 1680 WS + + + + Leptophis depressirostris Least Concern 5 9 13 5 4 1120 WS + Leptophis mexi canus Least Concern 2 28 8 0 1700 WS + + + + + + + Leptophis modestus Endangered B1ab(iii) 2 8 15 1500 2500 WS + + + Leptophis nebulosus Least Concern 2 12 14 0 1600 WS + Masticophis mentovarius Least Concern 2 32 11 0 2500 WS + + + + + + Mastigodryas alternatus Least Concern 3 N/E 14 6 20 3006 WS + Mastigodryas dorsalis Vulnerable B1ab(iii) 2 10 12 635 1900 WS + + + Mastigodryas melanolomus Least Concern 1 N/E 9 6 0 10406 WS + + + + Ninia diademata Least Concern 1 19 8 0 2200 WS + + + + Ninia espinali Endangered B1ab(iii) 2 [Near Threatened 1 ] 5 12 1590 2242 CB + + +

PAGE 155

155 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Ninia macula ta Least Concern 2 13 12 36 1800 WS + Ninia pavimentata Endangered B1ab(iii)2 5 12 1300 1500 WS + Ninia sebae Least Concern 2 28 4 0 2200 WS + + + + + + + Nothopsis rugosus Least Concern 1 11 12 2 830 WS + Omoadiphas aurula Enda ngered B2ab(iii) 2 3 15 1250 1900 CB + Omoadiphas cannula Critically Endangered B1ab(iii)+2ab(iii) 3 3 3 14 3 1250 CB + Omoadiphas texiguatensis Critically Endangered B1ab(iii)+2ab(iii) 2 [Data Deficient 1 ] 3 14 1690 CB + Oxybelis aen eus Least Concern 2 35 9 0 2500 WS + + + + + + + Oxybelis brevirostris Least Concern 2 12 13 4 800 WS + Oxybelis fulgidus Least Concern 2 27 10 0 1600 WS + + + + + Oxybelis wilsoni Endangered B1ab(iii)+2ab(iii) 2 3 15 0 95 CB + Oxyrho pus petola Least Concern 2 18 13 0 800 WS + + + Pliocercus elapoides Least Concern 1 25 10 0 2000 WS + + + + + Pliocercus euryzonus Least Concern 1 13 14 0 1250 WS + Pseudelaphe flavirufa Least Concern 1 21 12 0 1200 WS + + + Pseuste s poecilonotus Least Concern 1 22 12 0 1330 WS + + + + Rhadinella anachoreta Endangered B1ab(iii) 2 9 12 500 1180 CB + + Rhadinella decorata Least Concern 2 20 11 0 1400 WS + + Rhadinella godmani Vulnerable B1ab(iii) 2 13 9 1200 2200 W S + + + Rhadinella kinkelini Least Concern 1 ; Vulnerable B1ab(iii) 2 10 12 1370 2085 CB + + +

PAGE 156

156 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Rhadinella lachr ymans Least Concern1; Vulnerable B1ab(iii) 2 12 13 500 3000 WS + Rhadinella montecristi Endangered B1ab(iii) 2 7 12 1370 2620 CB + + Rhadinella pegosalyta Critically Endangered B1ab(iii)+2ab(iii) 2 3 14 CB + Rhadinella rogerromani Vulnerable B1ab(iii) 5 3 16 5 1450 CB + Rhadinella tolpanorum Endangered B1ab(iii)+2ab(iii) 2 3 15 1690 1900 CB + Rhinobothryum bovallii Least Concern 1 9 15 4 550 WS + Scaphiodontophis annulatus Least Concern 2 19 12 0 1400 WS + + + + Scaphiodontophis venustissimus Least Concern 2 11 11 2 830 WS + Scolecophis atrocinctus Least Concern 2 14 14 100 1530 WS + + Senticolis triaspis Least Concern 2 30 10 10 2500 WS + + + Sibon annulatus Least Concern 2 10 12 2 1300 WS + Sibon anthracops Least Concern 2 12 14 4 915 WS + + Sibon carri Endangered B1ab(iii) 2 8 12 30 800 CB + + Sibon dimidiatus Least Concern 1 18 11 0 1600 WS + + + + Sibon longifrenis Least Concern 2 9 11 60 750 WS + S ibon manzanaresi Critically Endangered B1ab(iii)+2ab(iii) 2 3 15 250 300 CB + Sibon miskitus Critically Endangered B1ab(iii)+2ab(iii) 2 3 15 150 CB + Sibon nebulatus Least Concern 2 27 8 0 1690 WS + + + + Spilotes pullatus Least Co ncern 2 30 9 0 1500 WS + + + + + + Stenorrhina degenhardtii Least Concern 2 23 10 0 1900 WS + + + + + + Stenorrhina freminvillei Least Concern 1 22 11 0 2000 WS + + + +

PAGE 157

157 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distrib ution (m) General Distribution CL CV NC CC SC PL PV IB Storeria dekayi Least Concern 1 13 9 0 1900 WS + + + Tantilla armillata Least Concern 2 14 9 0 1435 WS + + + + Tantilla impensa Least Concern 1 10 12 300 1600 CB + Tantilla lempira Endangered B1ab(iii)+2ab(iii) 2 4 13 1450 1730 CB + Tantilla sp. nov. Critically Endangered B1ab(iii)+2ab(iii) 3 3 3 16 3 1150 CB + Tantilla psittaca Vulnerable B1ab(iii) 5 3 12 3 5 420 CB + Tantilla schistosa Least Concern 2 24 10 40 1680 WS + + + Tantilla taeniata Least Concern 2 13 10 0 1280 WS + + + + Tantilla tritaeniata Critically Endangered B1ab(iii)+2ab(iii) 2 3 15 0 CB + Tantillita lintoni Least Concern 1 15 13 0 550 WS + Thamnophis fulvus Least Conc ern 1 10 14 1680 3500 WS + Thamnophis marcianus Least Concern 2 22 13 0 1400 WS + Thamnophis proximus Least Concern 1 27 9 0 2500 WS + + + + + Tretanorhinus nigroluteus Least Concern 2 18 8 0 1200 WS + + + Trimorphodon quadruplex Least Concern 2 14 10 0 2000 WS + + Tropidodipsas fischeri Least Concern 1 12 12 1000 3000 WS + Tropidodipsas sartorii Least Concern 2 26 12 0 2000 WS + + + + Urotheca decipiens Least Concern 2 10 11 15 1500 WS + Urotheca guenthe ri Least Concern 1 13 12 25 1600 WS + Xenodon rabdocephalus Least Concern 2 24 12 0 1300 WS + + + + + Elapidae (6) Micrurus alleni Least Concern 2 11 15 1 1620 WS + Micrurus browni Least Concern 3 15 13 0 2200 WS + Micrurus diastema Least Concern 2 19 12 50 600 WS + + + Micrurus mipartitus Least Concern 5 6 15 5 2 1160 WS + Micrurus nigrocinctus Least Concern 2 21 9 0 1600 WS + + + +

PAGE 158

158 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevatio nal Distribution (m) General Distribution CL CV NC CC SC PL PV IB Micrurus ruatanus Critically Endangered B1ab(iii) 1 3 17 0 20 CB + Leptotyphlopidae (2) Epictia ater Least Concern 6 33 6 0 1600 WS + + + + Epictia magnamaculata Endangered B1ab(iii)+2ab(iii) 5 9 0 25 CB + Loxocemidae (1) Loxocemus bicolor Least Concern 2 16 11 0 750 WS + + + Typhlopidae (3) Typhlops costaricensis Least Concern 2 11 11 540 1500 WS + + Typhlops stad elmani Endangered B1ab(iii) 2 6 12 320 1370 CB + + + Typhlops tycherus* Data Deficient 2 3 14 1550 CB + Ungaliophiidae (1) Ungaliophis continentalis Vulnerable B1ab(iii) 2 11 12 990 2300 WS + + Viperidae (12) Agkistrodon bilineatus Near Threatened 1 17 15 0 1500 WS + Atropoides indomitus Endangered B1ab(iii) 2 33 17 670 1200 CB + Atropoides mexicanus Least Concern 2 17 12 0 1600 WS + + + + Atropoides occiduus Vulnerable 4 7 15 4 100 1600 WS + Bothriechis marchi Endangered B1ab(iii, iv) 2 5 16 500 1840 CB + + Bothriechis schlegelii Least Concern 2 21 12 0 1530 WS + + + + Bothriechis thalassinus Vulnerable B1ab(iii)+2ab(iii) 2 5 15 1370 1750 CB + + Bothrops asper Lea st Concern 2 25 12 0 1300 WS + + + + + Cerrophidion sp. Vulnerable B1ab(iii) 2 15 12 1300 2875 WS + + +

PAGE 159

159 Table 3 2 Continued. Taxon IUCN Red List Status CSS EVS Elevational Distribution (m) General Distribution CL CV NC CC SC PL PV IB Crotalus simus Least Concern 2 21 12 500 2600 WS + + + + + Porthidium nasutum Least Concern 1 18 12 0 1100 WS + + + + Porthidium ophryomegas Least Concern 2 15 9 0 1400 WS + + + + Sources for Red List Status: 1 IUCN Red List (2011) 2 Townsend & Wilson (2010) 3 Evaluated for this study 4 Acevedo et al. (2010) 5 Sunyer & Khler (2010) 6 McCranie (2011 a ) Table 3 3. Composition of the Chorts Block herpetofauna, compared with regional (Mesoamerica = MA; Wilson & Johnson 2010) and global (worldwide = WW; Amph ibiaWeb 2011, Uetz et al. 2011). FAMILIES GENERA SPECIES CB MA WW CB MA CB MA WW Gymnophiona 1 1 9 2 4 2 16 188 Caudata 1 4 10 5 17 43 241 614 Anura 11 16 38 33 68 100 474 6,086 AMPHIBIA 13 21 57 40 89 145 731 6,888 Crocodylia 2 2 3 2 2 2 3 24 Te studines 4 9 14 5 19 10 55 323 Squamata 22 31 61 87 179 225 1,090 8,883 REPTILIA 28 42 78 94 200 237 1,148 9,413 TOTALS 41 63 135 134 289 382 1,879 16,301

PAGE 160

160 Table 3 4 Broad distributional patterns of herpetofaunal diversity in the Chorts Block CB = Chorts Block restricted, WS = widespread, CL = Caribbean Lowlands, CV = Caribbean Intermontane Valleys, NC = Northern Cordillera, CC = Central Cordillera, SC = Southern Cordillera, PL = Pacific Lowlands, PV = Pacific Intermontane Valleys, IB = Islas de l a Bahia. Distribution Physiographic Regions CB WS CL CV NC CC SC PL PV IB Gymnophiona 0 2 2 2 0 0 0 1 0 0 Caudata 37 6 6 1 19 17 9 2 2 0 Anura 37 63 51 18 50 37 30 12 16 6 AMPHIBIA 74 7 1 59 21 69 54 39 15 18 6 Crocodylia 0 2 2 2 0 0 0 2 0 1 Testudines 0 10 6 6 0 0 0 2 1 0 Squamata: Sauria 26 61 36 31 32 21 20 18 20 13 Squamata: Serpentes 32 107 85 64 62 60 41 27 36 14 REPTILIA 57 1 80 129 103 94 81 61 49 57 28 TOTALS 131 25 1 188 124 163 135 100 64 75 34 Table 3 5 E ndemic and co nservation priority herpetofaunal diversity from the Chorts Block Families Genera Species Endemic Species (% of endemic species) Critically Endangered Species Endangered Species Vulnerable Species Total % of Species that are CR/EN/VU Total % of Endemic Species that are CR/EN/VU Gymnophiona 1 2 2 0 0 0 1 50% Caudata 1 5 43 3 6 (8 4 %) 19 12 2 74 % 87 % Anura 11 33 100 3 7 (37 %) 26 15 3 44% 100% AMPHIBIA 13 40 145 73 (5 1 %) 45 27 6 54 % 94 % Crocodylia 2 2 2 0 0 0 1 50% Testudines 4 5 10 0 0 0 1 10% Sq uamata: Sauria 13 26 87 32 (37 %) 9 23 6 44 % 97 % Squamata: Serpentes 9 61 139 32 (23 %) 10 18 9 27% 90 % REPTILIA 28 94 237 64 (27 %) 19 41 17 33 % 96 % TOTALS 41 134 382 134 (35 %) 64 68 23 41 % 96 %

PAGE 161

161 Table 3 6 Distribution by mountain ranges of the endemic amphibians and reptiles of the Chorts Highlands (exclusive of Caribbean and Pacific Lowlands and islands); mountain ranges are defined in Chapter 2. Northern Cordillera Central Cordillera Southern Cordillera Taxon Nombre de Dios : Texguat Nombre de Di os : Pico Bonito Omoa Espritu Santo Joconal Santa Brbara Membar Montecillos Comayagua Sulaco La Flor La Muralla Agalta Botaderos Punta Piedra Patuca Montecristo Merendn Celaque Erandique Puca Opalaca De la Sierra Lepaterique Dipilto Coln Dariense CA UDATA Plethodontidae ( 36 ) Bolitoglossa carri + Bolitoglossa cataguana* + Bolitoglossa celaque + + + + Bolitoglossa conan ti + + + Bolitoglossa decora + Bolitoglossa diaphora + Bolitoglossa dunni + + Bolitoglossa heiroreias + Bolitoglossa lon gissima + Bolitoglossa nympha + + + + + Bolitoglossa oresbia + Bolitoglossa porrasorum + + + Bolitoglossa synoria + Crypto triton monzoni + Cryptotriton nasalis + Cryptotriton wakei + Dendrotriton sanctibarbarus + Nototriton barbouri + Notot riton brodiei + Nototriton lignicola + Nototriton limnospectator + + Nototriton picucha* + Nototriton saslaya + Nototri ton stuarti + Nototriton tomamorum* +

PAGE 162

162 Table 3 6 Continued. Northern Cordillera Central Cordillera Southern Cordillera Taxon Nombre de Dios : Texguat Nombre de Dios : Pico Bonito Omoa Espritu Santo Joconal Santa Brbara Membar Montecillos Comayagua Sulaco La Flor La Muralla Agalta Botaderos Punta Piedra Patuca Montecristo Merendn Celaque Erandique Puca Opalaca De la Sierra Lepaterique Dipilto Coln Dariense Nototriton sp A (Pico Bonito)* + Nototriton sp B (Texiguat)* + Nototriton sp. C (Botaderos)* + Oedipina gephyra + Oedipina kasios + Oedipina koehleri* + Oedipina leptopoda + + Oedipina nica* + Oedipina quadra + + Oedipina petiola* + Oedipina tomasi + ANURA (37) Bufonidae ( 3 ) Incilius leucomyos + + + Incilius porteri + + Rhinella chrysophora + + Craugastoridae (20 ) Craugastor anciano + Craugastor aurilegulus + + Craugastor charadra + + Craugastor chrysozetetes + Craugastor coffeus + + Craugastor cruzi + Craugastor cyanochthebius + Craugastor emleni + Craugastor epochthidius + + Craugastor fecundus + Craugastor laevissimus + + + + + + +

PAGE 163

163 Table 3 6 Continued. Northern Cordillera Central Cordillera Southern Cordillera Taxon Nombre de Dios : Texguat Nombre de Dios : Pico Bonito Omoa Espritu Santo Joconal Santa Brbara Membar Montecillos Comayagua Sulaco La Flor La Muralla Agalta Botaderos Punta Piedra Patuca Montecristo Merendn Celaque Erandique Puca Opalaca De la Sierra Lepaterique Dipilto Coln Dariense Craugastor lauraster + + + + + + + + Craugastor merendonensis + Craugastor milesi + + Craugastor nefrens + Craugastor olanchano + + Craugastor omoaensis + Craugastor pechorum + + Craugastor saltuarius + + Craugastor stadelmani + + + + HYLIDAE (13) Duellmanohyla salvavida + + Duellmanohyla soralia + + Ecnomiohyla salvaje + + Exerodonta catracha + + + + Isthmohyla insolita + Isthmohyla melacaena + Plectrohyla chrysopleura + + Plectrohyla dasypus + Plectrohyla exquisita + Plectrohyla psiloderma + + Ptychohyla hypomykter + + + + + + + + + + + + Ptychohyla salvadorensis + + + + + Ptychohyla spinipollex + + Leptodactylidae ( 1 ) Leptodactylus silvanimbus + + Ranidae ( 1 ) Lithobates sp. nov. + + + + SQUAMATA: LIZARDS (32) Anguidae ( 5 )

PAGE 164

164 Table 3 6 Continued. Northern Cordillera Central Cordillera Southern Cordille ra Taxon Nombre de Dios : Texguat Nombre de Dios : Pico Bonito Omoa Espritu Santo Joconal Santa Brbara Membar Montecillos Comayagua Sulaco La Flor La Muralla Agalta Botaderos Punta Piedra Patuca Montecristo Merendn Celaque Erandique Puca Opalaca De l a Sierra Lepaterique Dipilto Coln Dariense Abronia montecristoi + + + Abronia salvadorensis + Celestus bivittatus + + + Celestus montanus + + Celes tus scansorius + + Corytophanidae ( 1 ) Corytophanes percarinatus + + Polychrotidae: ( 17 ) Anolis amplisquamosus + Anolis cusuco + Anolis heteropholidotus + + Anolis johnmeyeri + + Anolis kreutzi + Anolis loveridgei + + + Anolis morazani* + Anolis muralla + Anolis ocelloscapularis + + Anolis pijolensis + Anolis purpurgularis + + Anolis rubribarbaris + Anolis sminthus + + + + Anolis wampuensis + Anolis wermuthi + + Anolis yoroensis + + + Anolis zeus + + + Sphaerodactylidae ( 1 ) Sphaerodactylus dunni + SQUAMATA: SNAKES (27) Colubridae (21 ) Geophis damiani +

PAGE 165

165 Tabl e 3 6 Continued. Northern Cordillera Central Cordillera Southern Cordillera Taxon Nombre de Dios : Texguat Nombre de Dios : Pico Bonito Omoa Espritu Santo Joconal Santa Brbara Membar Montecillos Comayagua Sulaco La Flor La Muralla Agalta Botaderos Punta P iedra Patuca Montecristo Merendn Celaque Erandique Puca Opalaca De la Sierra Lepaterique Dipilto Coln Dariense Geophis dunni + Geophis fulvoguttatus + + Geophis nephodrymus + Ninia espinali + + + + Omoadiphas aurula + Omoadiphas cannula + Omoadiphas texiguatensis + Rhadinella anachoreta + + Rhadinella montecristi + + + + + Rhadinella pegosalyta + Rhadinella rogerromani + Rhadinella tolpanorum + Sibon manzanaresi + Sibon miskitus + Tantilla impensa + + Tantilla lempira + Tantilla psittaca + Tantilla sp. nov. + Typhlopidae ( 2 ) Typhlops stadelmani Typhlops tycherus* Viperidae ( 4 ) Atropoides indomitus + Bothriechis marchi + + + + + + + + Bothriechis thalassinus + + + + Cerrophidion sp. + + + + + + + + + + RANGE TOTALS 26 22 35 22 1 8 9 2 3 13 13 11 5 4 3 7 12 7 3 5 6 9 3 3 10 CORDILLERA TOTALS 72 42 3 7

PAGE 166

166 CHAPTER 4 THE CHORT IS BLOCK IS AN UNDERESTIMATED HOTSPOT OF AMPHIBIAN DIVERSITY AND ENDEMISM Over the past 20 years, amphibians have become emblematic of the myriad of environmental issues that have led to the recognized global loss of biodiversity (Wake & Vredenburg 2008) The global phenomenon of amphibian declines and disappearances, termed the Global Amphibian Crisis, began coming to light at the First World Herpetology Congress in 1989 as amphibian researchers met and related similar stories of unexplained declines from opposite sides of the globe often from seemingly pristine habitats ( Stuart et al. 2004; Mendelson 2011 ). Since that time, a mphibian declines have been traced to a variety of factors, including habitat loss and fragmentation (Cushman 2006), emerging disea ses such as c hytridiomycosis (Berger et al. 1998; Skerratt et al. 2007) and Ranavirus (Gray et al. 2009), and environmental contamination (Reylea & Diecks 2008). Current rates of extinction in amphibians may approach 45,000 times the background rate (Wake & Vredenburg 2008; Honeycutt et al. 2010), indicating that amphibians as a group may be facing the worst threat to their survival as a group in their 365 million year existence. Furthermore, extinction risk is not evenly distributed globally, with species found in tropical areas, and particular tropical stream s facing the highest degree of threat (Dudgeon et al. 2006). Paradoxically the past 20 years has also seen the discovery and taxonomic description of amphibian species increase at a remarkable pace (Khler et al. 2005), with number of recognized species rising 52% from 1992 ( 4,533; Duellman 1993) to October 2011 ( 6,885; AmphibiaWeb 2011), meaning about one quarter of all known amphibians have been described in the past two decades A principal reason for this increase is the advent and widespread implementation of molecular approaches in

PAGE 167

167 systematic biology allowing for rapid evaluation of large numbers of samples and identification of ca ndidate species (e.g. Vieites et al. 2009; Crawford et al. 2010) This rapid increase in amphibian discovery and description was documented in Khler et al. (2005), who se meta analysis demonstrated that most recently described species were not simply subdivisions of described species and were as or more distinctive (a s measured by DNA sequence divergence) as species described in previous eras of taxonomic activity, therefore refuting the notion that the recent rise in new amphibian discoveries is an ar tifact of taxonomic inflation. Rather, the increase is being fuelled by wider implementation of molecular methods in biodiversity inventory intensified efforts to sample poorly known areas, particularly in tropical regions and a rise in the number of working taxonomists from developing countries, i.e. those countries tha t contain the majority of undescribed diversity (Joppa et al. 2011 ) The juxtaposition of increasing loss of diversity and habitats with the accelerated documentation of new diversity has a number of important implications. First, this suggests that amphi regions may be going extinct without ever becoming known to the scientific community (Khler et al. 2005; Crawford et al. 2010). As pointed out in a number of recent works (e.g., Janzen et al. 2009; Monaghan et al. 2009; Honeycutt et al. 2010 ) this potential loss underscores the need for rapid taxonomic assessments across a wide taxonomic b readth. The use of a single or multiple DNA fragments (i.e. DNA barcoding) ha s been proposed, and widely implemented, as a st an dardized method for rapidly assessing taxonomic diversity ( Hebert et al. 2003 ).

PAGE 168

1 68 DNA barcoding initially received a mixed reception among specialists and potential practitioners ( Moritz & Cicero 2004 ), largely fuelled by a backlash to the effort of a limit ed number of advocates ( Tautz et al. 2002, 2003 ) for the development of a single scale use in the assignment of unknown samples to named taxa and the enhancement of taxonomic discovery ( Lipscomb et al. 2003; Seberg et al. 2003 contention and misunderstanding, given its inherent implication that each species possesses a fixed identification marker analogous to the Universal Product Code (UPC) on the side of a cerea l box rather than a perpetually evolving sequence of biological molecules (Moritz & Cicero 2004). A single fragment of the mitochondrial gene cytochrome oxidase subunit I (COI) ~650 base pair (bp) in length, has been intended for use as a universal metaz oan DNA marker, with the goal of generating a comprehensive reference dataset for use in DNA barcode based biodiversity inventory projects ( Hebert et al. 2003; Smith et al. 2008 ). While COI has been demonstrated as effective for species delimitation in mam mals ( Clare et al. 2007 ), birds ( Kerr et al. 2007 ), fishes ( Ward et al. 2009 ), and a variety of invertebrates ( Virgilio et al. 2010 ), been somewhat controversial. In some groups of amphibians, COI shows a misleadingly high degree of intraspecific variation that can overlap with interspecific divergence and mask species boundaries (Vences et al. 2005a, 2005b) while in other groups is has been used effectively ( Smith et al. 2008; Xia et al. 2011), demonstrat ing a clear gap between intraspecific and interspecific sequence divergence, i.e. the A principal problem limiting the universal implementation of COI in

PAGE 169

169 amphibians is in fact its high degree of variation, with variability in the priming r egions across multiple groups leading to high rates of failure in PCR amplification (Vences et al. 2005b) which can be addressed by the use of degenerate primers (Meyer 2003) or by designing group specific primer sets (Smith et al. 2008), however taking t hese steps also diminishes promoted as an advantage of DNA barcoding. The proposed alternative to COI is use of a ~530 bp fragment of the large subunit rRNA gene ( 16S ber of be n e fits compared with COI, including 1) conserved priming regions that amplify successfully across amphibian groups using a single set of primers (Mueller 200 6 ) 2) reduced frequency of overlap between intraspecific and interspecific divergence (Ve nces et al. 2005a, 2005b) and 3) widespread use of 16S in amphibian studies prior to widespread implementation of DNA barcoding methods, providing a relatively large and well explored reference dataset for comparison of newly acquired sequence data ( e.g. Garca Pars & Wake 2000 ). An alternative to selecting a single marker for use in amphibian barcoding studies is to take a two gene barcoding approach for which I see the following advantages: 1. Distance based analyses of COI and 16S data can be directly c ompared for congruence in species delimitation, reducing concerns over erroneous species identification based on use of a single marker. 2. Expansion of available reference data by allowing comparisons to sequences only available for one marker or the other. 3. Taken together, COI and 16S represent fragments of two classes (protein coding and rRNA) of mitochondrial genes with differential rates of evolution, a preferable combination of characteristics for use in phylogenetic analysis. As the benefits and limita tions of DNA barcoding have become more clearly defined (Hebert & Gregory 2005; Hajibabaei et al. 2007), the approach has gained acceptance among systematic biologists as a useful, even powerful, addition to the

PAGE 170

170 methods used for the exploration and descrip tion of biological diversity (e.g. Vieites et al. 2009). Rather than consider DNA barcoding as an alternative to traditional taxonomic methods, I view it as a means of providing verifiable taxonomic assignment of samples and for identifying samples in need of more rigorous evaluation; the molecular analog to the process of providing preliminary taxonomic assignments to samples morphologically using a dichotomous key. In this capacity, sequence data is evaluated using distance based methods with the sole int ention of providing a means of DNA based taxonomic assignment and identification of potential candidate species, rather than generating hypotheses of evolutionary relationships, which is the goal of phylogenetics. Consequently, distance based neighbor join ing (NJ) trees used in DNA barcoding are simply graphical representations of genetic distances used to quickly assess the presence of reciporically mon o phyletic sequence clusters. In cases where further analysis appears warranted, the sequence data used in the barcoding study should be supplemented with data from additional loci and subject to more rigorous phylogenetic analysis. S cientists engaged in rapid biodiversity inventories using DNA barcoding should also be committed to following up that work or at least tangibly supporting the follow up to the work with focused work in descriptive systematics The taxonomic description of new species is considered a top priority as systematic biology moves through the era of elevation biodiversity loss, and is g iven the same level of importance as the biodiversity inventory work itself by Systematics Agenda 2000 (1994) and the IUCN Global Amphibian Assessment (Parra Olea et al. 2007). A n accurate and functioning taxonomy provides the critical basis for not only c onservation and management planning, but also

PAGE 171

171 communicating about biodiversity conservation with the general public ( Parra Olea et al. 2007; Honeycutt et al. 2010 ). While the relative ease with which new species can be identified using molecular methods ha s led to the dramatic rise in known amphibian diversity, the proverbially painstaking and specialized work required to address alpha taxonomy leads to a lag between the molecular identification of potential candidate species and the formal description of n ewly identified taxa (Wheeler 2004) The amphibian fauna of the Chorts Block has largely mirrored the global patterns of both decline and discovery (Townsend & Wilson 2010a) More than half (54%) of Chorts Block amphibians are in one of the three highest IUCN threat categories (Critically Endanger ed, Endangered, and Vulnerable), and include an alarming 74% of salamanders (Chapter 3, Table 3 5 ). Honduras the country making up the majority of the geographic territory of the Chorts Block has 49 endemic am phibian s 46 of which were discovered and described since 1976 (Townsend & Wilson 2010a). While the Chorts Block has been demonstrated to have a regionally distinctive component of endemic biodiversity, particularly in amphibians ( Campbell 1999; Wilson & Johnson 2010), molecular characterization of Chorts Block amphibian diversity has been restricted to a few studies of wider geographic focus and limited within region sampling ( Garca Pars & Wake 2000; Parra Olea et al. 2004; Hillis & Wilcox 2005; Frost et al. 2006 ). Given these considerations, I followed an iterative process to inventory and characterize phylogenetic diversity in Chorts Block amphibians. I present p distance based results from COI and 16S sequencing of 456 samples representing 52 named species in nine amphibian families and use these results to identify samples using the

PAGE 172

172 existing taxonomy and to delimit potential candidate species for future study Among Chorts Block amphibians, salamanders exhibit both the highest degree of endemism and extinction risk, and, based on the current taxonomy are the most comprehensively sampled group among my data. I provide tentative taxonomic assignments for ten unidentified populations of salamanders (discussed in Chapter 3), identifying both new cand idate species and new populations and morphotypes of known species. I then employ maximum likelihood based phylogenetic analysis to evaluate evolutionary relationships among Chorts Block salamanders. Implications for taxonomy, biogeography, and conservati on are discussed. Methods and Materials Sampling and Sample Identification Specimens were collected in the field between 2006 and 2011 (Chapter 3 for details) with fresh tissue samples typically being preserved in SED buffer (20% DMSO, 0.25 M EDTA, pH 7. 5, NaCl 2 saturated; Seutin et al. 19 91; Williams 2007), or 95% ETOH Vouchers were deposited in the Fl orida Museum of Natural History (FLMNH) at the University of Florida (UF), Museum of Vertebrate Zoology, University of California, Berkeley (MVZ), Sencken berg Museum, Frankfurt am Main Germany ( SMF ) and National Museum of Natural History, Smithsonian Institution (USNM). In addition, tissue samples were deposited at the FLMNH Genetics Resources Repository in May 2009. Species assignments for samples are ba sed on morphological identification, and, in the case of endemic species with limited distributions, collecting locality was also used to help sp. inquirenda sp. inq. samples of uncertain taxonom ic assignment, determined a priori

PAGE 173

173 Extraction, Amplification, and Sequencing For salamanders Template DNA was extracted from muscle tissue using the Gentra PureGene Tissue Kit (QIAGEN, Valencia, CA instructions. Concentration a nd purity of DNA extract was estimated using a NanoDrop 1000 spectrophotometer (Thermo Fisher Scientific, Waltham, MA ). Regions of two mitochondrial genes were selected for amplification and sequencing: 16S large subunit RNA (16S) and cytochrome oxidase s ubunit I (COI) The locus 16S is widely used in studies of amphibian phylogeny, and is considered the a Vieites et al. 2009); COI is widely 03), but has not been widely used in amphibians (Smith et al. 2008 Xia et al. 2011 ). Fragments of the mitochondrial genes 16S an d COI were amplified using the primers 16Sar L and 16Sbr H for 16S (Palumbi et al. 1991) and dgLCO 1490 and dgHCO 2198 for COI (Meyer 2003). PCR reactions were typically 20 L in total volume containing ~25ng of DNA template, 4 L 5X PCR buffer (final concentration 1X), 1.2 L of 25 mM MgCl 2 (final concentration 1.5 mM), 0.09 L of 10 mM dNTPs (0.045 mM) 0.8 L of each 10 M prim er ( 0.04 M ), 0.01U of GoTaq Flexi polymerase ( 1U/ L ; Promega, Madison, WI, USA), and 11.91 L H 2 O. Amplification profile for 16S consisted of an initial denaturation for 3 minutes at 94C, 35 cycles of denaturation at 94C for 45 seconds, annealing at 50 C for 45 seconds, and extension at 72C for 45 seconds, with a final elongation at 72C for 5 minutes. COI used an initial denaturation for 1.5 minutes, 37 cycles of 94C for 40 seconds, 45C for 40 seconds and 72C for 40 seconds, and a final elongation at 72C for 5 minutes. PCR products were verified using electropho resis

PAGE 174

174 on a 1.5 % agarose gel stained with ethidium bromide. Unincorporated nucleotides were removed from PCR product using 1 uL of ExoSAP IT (USB, Santa Clara, CA, USA) per 10 L of PCR produc t. Product was cycle sequenced forward and reverse stands using the BigDye Terminator v3.1 Cycle Sequencing kit (Applied Biosystems, Inc., Carlsbad, CA [ABI]); sequencing reactions typically included 0.8 1.2 L of PCR product, 1.1 L 5X ABI sequencing buff er, 0.22 L primer, and the balance in ddH 2 O for a total volume of 5 L. The sequenced product was cleaned using spin column filtration through Sephadex (G 50; GE Healthcare, London, UK ). Cleaned product was electrophoresed on an ABI 3130xl Genetic Analyze r. A nuran samples were amplified and sequenced at the Smithsonian Institution Laboratory of Analytical Biology (Suitland, Maryland) following standardized DNA Barcode of Life (BOLD) protocols (Hebert et al. 2003, Borisenko et al. 2009). For these samples, Template DNA was obtained using phenol chloroform extraction implemented by an AutoGen Gene prep 965 (AutoGen, Holiston, MA) automated DNA isolation robot. Template DNA was then amplified for COI using the primers LCO 1490 and HCO 2198 (Folmer et al. 1994) and for 16S using the primers 16Sar L and 16Sbr H (Palumbi et al. 1991). Unincorporated nucleotides were removed from PCR product using 2 uL of ExoSAP IT per sample Product was cycle sequenced using BigDye Terminator v3.1 Cycle Sequencing kit ( ABI), clea ned using spin column filtration through Sephadex and electrophoresed on an ABI 3730xl DNA Analyzer. Sequence Evaluation and Alignment Sequences were manually assembled and trimmed using CLC Combined Workbench 3 (CLCbio, Denmark). DNA barcode identifica tion was carried out by

PAGE 175

175 comparing generated sequences to those in the National Center for Biological Information (NCBI) database (http://www.ncbi.nlm.nih.gov/). Representatives from all sampled species and unassigned samples were searched against the NCBI database using BLASTN (Zhang et al. 2000) in order to identify reference sequences for inclusion in this analysis; a complete list of samples used in this analysis is presented in Table 4 1 A large number of reference sequences were included from a recent study of Panamanian amphibians (Crawford et al. 2010) and referenced sequences for Ranidae were from Hillis & Wilcox (2005) Samples representing ten populations of salamanders of uncertain taxonomic assignment were searched using B LASTN against the NCBI database. F or use in BLASTN searching consensus sequences were generated in CLC Combined Workbench 3 for each set of non divergent 16S samples representing a single species Finally, my n ewly generated sequences and reference sequences identified in BLAS TN searches were aligned under the default parameters using ClustalW (Thompson et al. 1994) as implemented by the pro gram MEGA5 (Tamura et al. 2011). Alignments were then visually inspected and adjusted. No stop codons were present in the COI sequences, in dicating that these all were amplified from mitochondrial COI and not nuclear pseudogenes (Song et al. 2008 Buhay 2009 ). Distance Based Barcode Metrics Data analysis was undertaken at various scales. First, to provide a graphical view of the family and g eneric level taxonomic discrimination of Chorts Block am phibians all newly generated sequences (Caudata + Anura) were used to construct neighbor joining (NJ) tree s in MEGA5 under the uncorrected (p distance) model using default settings and with nodal su pport calculated with 1,000 nonparametric bootstrap replicates. Then, n ewly generated data were combined with BLASTN reference sequences and

PAGE 176

176 separated into six datasets representing distantly related monophyletic clades (Frost et al. 2006, Pyron & Wiens 20 11): Bolitoglossa other Plethodontidae Bufonidae, Hyl idae, Leptodactylidae, and Strabomantidae (these four groups do not form a monophyletic group in combination). For ea ch dataset, NJ trees were generated for both 16S and COI under the uncorrected (p distance) method described above with 5,000 nonparametric bootstrap replicates Sequence divergences were estimated using uncorrected p distances, using the default paramete rs in MEGA5. Phylogenetic Analysis E volutionary model based hypotheses of phylogenetic relationships were estimated for the salamander data to provide a more rigorous ev al u a tion of candidate species identified in p distance based results The 16S and COI alignments were concatenated into a single dataset and r edundant haplotypes were removed to streamline analysis The combined dataset then was subjected to maximum likelihood (ML) phylogenetic analysis using the program RAxML v7.2.8 (Stamatakis 2006), pe rfor med with 1,000 bootstrap replicates under the GTR GAMMA substitution model (RAxML has a limited number of substitution models that can be implemented ), with data partitioned by 16S, COI codon position 1, COI codon position 2, and COI codon position 3. Species Delimitation and Candidate Species Degrees of divergence between two sequences or groups of sequences (based on uncorrected p distance) are herein < 1 2.0%; COI: 1 .0 > 6 .0% ). To label samples of uncertain taxonomic assignment identified a posteriori I

PAGE 177

177 use the terminology Confirmed Candidate Species, Unconfirmed Candidate Species and Deep Conspecific Lineage, as proposed by Vieites et al. (2009). Unconfirmed Candidate Species (UCS) are lineages identified as being deeply divergent genealogical lineages that have not yet been studied for differences in morphology or other characters, but may represent distinct, unnamed species. Confirmed Candidate Species (CCS) are unnamed but moderately to deeply divergent genealogical lineages that are supported by morphological characters or other evidence of distinctiveness. Deep Conspecific Lineages (DCL) are shallow to deeply divergent genealogical lineages th at lack clear morphological or other differences with described sister species, yet represent putative evolutionary species that warrant further investigation In general discussion, I refer to all three categories as Potential Candidate Species. Results Broad Results for Distance based Analyses Newly generated s equence fragments of the mitochondrial genes COI and 16S were analyzed for a total of 437 individual samples representing 52 nominal species, 17 genera, and nine families of amphibians, as well as ten populations of salamanders with undetermined taxonomic assignments ( Figure 4 1, Table 4 1 ; previously discussed in Chapter 3 ). For my 437 samples, 391 COI seque nces (10.5% fa ilure rate) and 434 16S sequenc es (0.7% failure rate) were successfully obtain ed Average sequence lengths and nucleotide compositions are summarized in Table 4 2 In each of the six taxonomic ally delimited datasets 16S datasets included a greater number of reference sequences than COI datasets, due to widespread use of 16S as the

PAGE 178

178 Figure 4 1 Radial phylogram showing coverage of higher level taxonomic groups F amily level for anurans, genus /subgenus level for salamanders Phylogram generated us ing neighbor joining method with uncorrected p distances and1,000 bootstrap pseudoreplicates Stars indicate potential candidate species identified in this study. Analysis of both 16S and COI datasets revealed at least 36 new Potential Candidate Species i n eight of nine sampled amphibian families (Table 4 3), including 13 Confirmed Candidate Species (CCS), 17 Unconfirmed Candidate Species (UCS), and six Deep Conspecific Lineages (DCL). If both CCS and UCS are assumed to represent distinct undescribed speci es, then a 24.4% increase in diversity among the nine targeted families is projected. However, this study focused on just 52 named species of the 123 represented in those families, and these results represent a 57.7% increase among ingroup taxa

PAGE 179

179 M ean unco rrected p distance between Anura and Caudata was 0.244 for 16S and 0.238 for COI, and mean genetic distance within Anura was 0.206 for COI and 0.146 for 16S, and within Caudata was 0.155 for COI and 0.084 for 16S. For Anuran families with more than one spe cies level lineage sampled, intra family mean genetic distance for COI was 0.122 for Bufonidae, 0.113 for Craugastoridae, 0.154 for Hylidae, and 0.130 for Ranidae, and for 16S was 0.051 for Bufonidae, 0.104 for Craugastoridae, 0.068 for Hylidae, and 0.064 for Ranidae. Between anuran families, mean genetic distance for COI ranged from 0.199 (Bufonidae /Leptodactylidae) to 0.269 (Eleutherodactylidae/Ranidae), and for 16S from 0.128 (Hylidae/Leptodactylidae) to 0.278 (Craugastoridae/Ranidae). All samples of Ca udata represe nt a single family, Plethodontidae, which accounted for 36% of all COI and 38% of all 16S sequences analyzed (Table 4 1 ), far more representation than any other family Analysis of the Caudata dataset recovered the same deeply diverged higher order clades (family, genus, subgenus) as published phylogenetic hypotheses (Garca Pars & Wake 2000, Parra Olea et al. 2004, Wiens et al. 2007, McCranie et al. 2008 ) however relationships within these clades are poorly resolved ( Figure 4 2). Identifica tion of Potential Candidate Species of Anura Twenty new P otential C andidate S pecies of anurans were identified through uncorrected p distance analysis of the COI and 16S data (Table 4 3 ) A comprehensive comparative dataset for 16S is available for Central American Bufonidae, based largely on the work of Mendelson et al. (2005), Mendelson & Mulcahy (2010), Mulcahy & Mendelson (200 0), and Mulcahy et al. (2006). Reference sequences for COI were limited to Panamanian samples assigned to Incilius coniferus and Rhaebo haematiticus (Crawford et al. 2010), and two samples of I. nebulifer the sister species to I. valliceps

PAGE 180

180 Figure 4 2 Maximum likelihood phylogram of COI data showing generic and subgeneric relationships of salamander samples.

PAGE 181

181 Samples from this study were shown to be conspecific with the nominal species I coccifer (16S= 0.2 % ) I. ibarrai (16S= 0.3 %; COI = 0.5 % ) I. leucomyos (16S= 0.2 %; COI = 0.7 % ) I. luetkenii (16S= 0.4 % ) I. porteri (16S= 0.5 %; COI = 0.8 % ) and I. valliceps (16S= 0.8 %; COI = 0.7 % ) Two Un confirmed Candidate Species were identified among the Bufonidae data: I. cf. coniferus and R. cf. haematiticus ( Figure 4 3) The majority of Craugastoridae samples represented taxa that lacked comparative reference sequences; only one sample of C raugastor fitzingeri (N060) was conf irmed by COI and 16S data ( Figure 4 4 ) Most clades correspond with a priori taxonomic assignments. Three Unconfirmed Candidate Species were identified: a divergent population of C. cf. aurilegulus ; and two from northern Nicaragu a, C. sp. inquirenda 1 and C. sp. inquirenda 2 ( Figure 4 4 ) Reference sequences for the genus Diasporus (Eleutherodactylidae) were available for four putative species (Crawford et al. 2010). Four samples of Diasporus cf. diastema from northern Nicaragua form a well supported clade based on 16S and the single COI sequence is also deeply divergent (16S= 7.1 %; COI = 15.7 % ) from the closest available reference sequences (Fig 4 4 ), representing an Unconfirmed Candidate Species awaiting comparative morphological evaluation. While a large number of congeneric reference sequences are available for 16S, and to a lesser extend COI, all newly generated samples of Hylidae represented either the first sequence data available for a putative species for both COI and 16S (the case for at least 10 taxa), or a P otential C andidate S pecies assigned to another taxon. Purported congeners tended to cluster together, except in the cases of Ptychohyla salvadorensis two species of Duellmanohyla and two species of Exerodonta in th e 16S

PAGE 182

182 Figure 4 3 COI (left) and 16S (right) neighbor joining trees for Bufonidae ** = boostrap support 99, = boostrap support 90; candidate species indicated with red stars.

PAGE 183

183 Figure 4 4 COI (left) and 16S (right) neighbor joining trees for Craugastoridae, Eleutherodactylidae, Leptodactylidae, and Strabomantidae ** = boostrap support 99, = boostrap support 90; candidate species indicated with red stars.

PAGE 184

184 NJ tree ( Figure 4 5 ). At least five Unconfirmed Candidate Species were identified i n the 16S NJ tree and supported by the COI NJ tree ( Figure 4 5, Table 4 3 ): a Plectrohyla two Ptychohyla and two Smilisca Samples representing a single putative species, Leptodactylus fragilis (Leptodactylidae), were the only members of this family incl uded in this study. Four samples from Honduras and Nicaragua are deeply divergent (16S= 3.6 %; COI =15.7% ) from that of Panamanian samples assigned to L. fragilis ( Figure 4 4 ). O nly two 16S reference sequences provided taxonomic matches: Lithobates macroglos sa and L. taylori ( Figure 4 6). Two GenBank samples are referred to L. macroglossa ; one sample clusters with samples from Caribbean versant localities in Honduras that were identified as L. brownorum ; the second L. macroglossa sample forms a weakly support ed sister lineage to the L. taylori clade ( Figure 4 6 ) I tentatively consider the latter sequence to represent L. macroglossa while the widespread clade represents L. brownorum Two series of samples, one from Cerro Uyuca and one from San Pedro La Loma, that were considered hybrids between L. brownorum and L. forreri by McCranie & Wilson (2002) form well supported clades for both COI and 16S ( Figure 4 6) The 16S data places these as sister to a sample representing an undescribed highland leopard frog fro m Costa Rica ( Lithobates sp. 5 sensu Hillis & Wilcox 2005). Among samples thought to represent the highland and piedmont stream inhabiting L. maculatus considerable genetic diversity is seen in both COI and 16S data, and this taxon apparently masks at lea st four Unconfirmed Candidate Species in the Chorts Highlands ( Figure 4 6 ). Samples assigned to L. warszewitschii from northern Nicaragua

PAGE 185

185 Figure 4 5 COI (left) and 16S (right) neighbor joining trees for Hylidae ** = bootstrap support 99, = boost rap support 90; candidate species indicated with red stars.

PAGE 186

186 Figure 4 6 COI (left) and 16S (right) neighbor joining trees for Ranidae ** = boostrap support 99, = boostrap support 90; candidate species indicated with red stars.

PAGE 187

187 also a deeply di vergent clade from L. warszewitschii samples from Panam ( Figure 4 6 ). Within the family Strabomantidae, samples from northern Nicaragua assigned to Pristimantis ridens are shown to represent an Unconfirmed Candidate Species, deeply divergent (16S= 4.8 %; COI = 9.1 % ) from Panamanian samples referred to P. ridens but in themselves represent three candidate species ( Figure 4 4 ). BLASTN Results for Unassigned Salamander Sequences Two of ten unassigned salamander populations matched available 16S references sequ ences from NCBI: Nototriton sp. inquirenda 4 with N. lignicola and Oedipina sp. inquirenda 1 with O. kasios (Table 4 4 ). Both unidentified samples were from mixed cloud forest in Parque Nacional (PN) Montaa de Yoro and matched with species previously con sidered endemic to nearby PN La Muralla. A third population from the same localities in PN Montaa de Yoro, Bolitoglossa sp. inquirenda 1, also matches its counterpart from PN La Muralla, B. decora (Table 4 4 ), however these species exhibit markedly differ ent external morphology, discussed in further detail below. The remaining seven salamander populations do not closely match with any sequences in NCBI database ( Distance Based Analyses o f Caudata A relatively comprehensive 16S re ference dataset is available for Mexican and Central American salamanders (Garca Pars & Wake 2000, Parra Olea et al. 2004, Wiens et al. 2007, Adams et al. 2009) ; however only five COI sequences (two Bolitoglossa two Oedipina and one Nototriton ) were a vailable through NCBI. Intra generic divergence was lowest for Nototriton (16S =3.5%, n= 25 ; COI =9.7%, n=16), compared with Bolitoglossa ( 16S =5.5%, n= 192 ; COI =12 7 % n=11 4 ) Cryptotriton

PAGE 188

188 Figure 4 7 COI (left) and 16S (right) neighbor joining trees for B olitoglossa ** = boostrap support 99, = boostrap support 90; candidate species indicated with red stars.

PAGE 189

189 (16S =5.1%, n= 5 ) and Oedipina ( 16S =7.5%, n= 50 ; COI = 1 3 5 % n= 10) Between salamander genera, mean genetic divergence for COI ranged from 20 5 % ( Nototriton / Oedipina and Cryptotriton / Nototriton ) to 21 9 % ( Cryptotriton / Oedipina ), and for 16S ranged from 11 .3% ( Nototriton / Oedipina ) to 20 2 % ( Bolitoglossa / Cryptotriton ). New samples were recovered as conspecific with the following nominal species (intr aspecific divergence indicated parenthetically ): Cryptotriton veraepacis (16S =0.0 % ; COI=0.0% ) Nototriton limnospectator (16S =0.3 %; COI= 0.2 % ) N. lignicola (16S =0.2 % ) Oedipina gephyra (16S =0.1 %; COI = 0.0 % ) O. kasios (16S = 0.1 % ) Bolitoglossa celaque ( 16S = 1.5%; COI = 1 9 % ) B. conanti ( 16S =1.6%; COI = 1 9 % ) B. diaphora (16S =0.0 %; COI =0.1% ) B. dofleini (16S =0.3 %; COI =0.1% ) B. dunn i (16S =0.3 % ) B. heiroreias (16S =0.0 % ) B. longissima (16S =0.3 %; COI =1.2% ) B. mexicana (16S= 2.1 %; COI =1.2% ) B. nympha ( 16S= 3.0 %; COI = 1.4%), and B. synoria (16S= 0.1 %; COI =0.0% ) Sixteen P otential C andidate S pecies of salamanders were identified in the 16S and COI data ( Table 4 2 ) Populations assigned to B. porrasorum and N. barbouri from Texguat and Pico Bonito are divergent from each other at an interspecific level, and are deeply divergent from samples representing the terra tipica of each nominal taxa ( Figures 4 7, 4 8 ). Samples from the Cordillera de Agalta ( Nototriton sp. inq. 2 ) and Sierra de Botaderos ( Nototriton sp. inq. 3 ) representing the first known samples of Nototriton from either mountain range, are sister clades ( intra clade divergence 0.0 % for 16S and 0.2 0.6% for COI; inter clade divergence 1.7 % for 16S and 0.29% for COI) and are both Confirmed Candidate Species (CCS ). Other CCS include a divergent sample

PAGE 190

190 Figure 4 8. COI (left) and 16S (right) neighbor joining trees for Cryptotriton Dendrotriton Nototriton and Oedipina stars. from Texguat ( Nototriton sp. inq. 1 ), Oedipina cf. gephyra from Pico Bonito, Oedipina sp. inq. 2 from the Cordillera de Dariense in Nicaragua, Oedipina sp. inq. 3 from the southeastern piedmont of the Chorts Highlands in Nicaragua, and Bolitogloss a cf. rufescens from the northern foothills of the Sierra de Omoa. Clusters of geographically

PAGE 191

191 discrete moderately divergent subclades ( within subclade divergence 16S =0.0 0.8 %; COI = 0.0 0.3 % ; be tween subclade divergence 16S =0.6 1.6 %; COI =1.7 3.6% ) were also present in samples assigned to B. celaque and B. conanti ( Figure 4 7), which have not previously been differentiated morphologically (McCranie & Wilson 2002) and therefore are considered Deeply Conspecific Line ages pending further evaluation of morphologi cal variation Phylogenetic Analysis of Caudata After removing identical redundant haplotypes, data from 16S and COI were concatenated into a single dataset consisting of 276 individual samples, with a total alig ned sequence length of 1,178 bp Maximum li kelihood (ML) analysis recovered a topology largely congruent with previous analyses of available reference data ( Figure 4 9; Garca Pars and Wake 2000, Parra Olea et al. 2004, Wiens et al. 2007, McCranie et al. 2008). For Bolitoglossa clades representi ng the subgenera Bolitoglossa Eladinea Magnadigita Nanotriton and Pachymandra were recovered as monophyletic ( Figure 4 9). A clade corresponding to the Bolitoglossa dunni species group was also recovered, containing the nominal taxa B. carri B. catagu ana B. celaque B. conanti B. decora B. diaphora B. dunni B. flavimembris B. heiroreias B. longissima B. morio B. oresbia B. porrasorum B. cf. porrasorum (Pico Bonito), B. cf. porrasorum (Texguat) and B. synoria ( Figure 4.9). Within the B. du nni group, four phylogenetic divisions are apparent, each corresponding to a discretely distributed group of species: Clade A ( B. celaque, B. heiroreias, B. synoria ; across the ignimbrite highlands of the Southern Cordillera), Clade B ( B. carri, B. conanti B. diaphora, B. dunni, B. oresbia ; from the

PAGE 192

192 Figure 4 9 Maximum likelihood phylogram for the genus Bolitoglossa C ombined 16S/COI dataset, partitioned by gene with 1,000 bootstrap replicates.

PAGE 193

193 Sierra de Omoa southward to the Sierra del Merendn, then eastward to the Sierra de Lepaterique and north again to the Montaas de Membar), Clade C ( B. cataguana, B. decora, B. longissima, B. porrasorum, B. cf. porrasorum [Pico Bonito], B. cf. porrasorum [Texguat]; the highlands of northern and north central Honduras east to the Sierra de Agalta), and Clade D ( B. flavimembris, B. morio ; Guatemala). Taxonomic discrepencies in distance based analyses ( Figure 4 7) were confirmed by ML analysis ( Figure 4 9). Samples assigned to B. celaque from three mountain rang es in the Southern Cordillera are recovered as an eastern clade (La Paz + Intibuca) and a western clade (Celaque), with the La Paz samples forming a shallow sub clade within the eastern clade ( Figure 4 9). Samples assigned to B. conanti also form a clade o f geographically discrete, shallow ly divergent, monophyletic groups corresponding to the Sierra de Espritu Santo, Sierra del Merendn, Sierra de Omoa, and Cerro del Mono, with the B. conanti complex clade recovered as sister to B. carri itself endemic to the Sierra de Lepaterique ( Figure 4 9). Samples from the B rufescens complex ( sensu McCranie & Wilson 2002) are assignable to either B. nympha or to an unnamed lineage ( Figure 4 9). Clades corresponding to the putative genera Nototriton Oedipina and Cr yptotriton were supported in the ML analysis ( Figure 4 10). For Nototriton clades representing the N. barbouri group, N. picadoi group, and N. richardi group were recovered as monophyletic ( Figure 4 10). As with Bolitoglossa phylogenetic results were con gruent with distance based results in terms of delimiting species level lineages ( Figures 4 8, 4 10). Nototriton sp. inq. 1 was shown to be a phylogenetically distinctive and deeply divergent lineage, sister to the entire N. barbouri group. Together, the N ototriton sp. inq.

PAGE 194

194 Figure 4 10 Maximum likelihood phylogram for the genera Nototriton Oedipina Dendrotriton and Cryptotriton C ombined 16S/COI dataset, partitioned by gene with 1,000 bootstrap replicates.

PAGE 195

195 1 and N. barbouri group clades represent an radiation endemic to the Chorts Highlands. Inclusion of material from the type locality of N. barbouri in phylogenetic a nalysis revealed the taxon to be widely paraphyletic with respect to populations from Pico Bonito and Texguat ( Figure 4 10). Notot riton barbouri sensu stricto from the vicinity of the type locality is the sister species to N. limnospectator while the Pico Bonito and Texiguat populations (themselves forming two distinct monophyletic lineages) form a clade with N. brodiei from the hig hlands along the Guatemala/Honduras border. Nototriton sp. inquirenda 4 and N. sp. inquirenda 5 from central Honduras are shown to be conspecific with N. limnospectator and N. lignicola respectively. In reference to the N. picadoi group, N. saslaya was re covered as the sister lineage to the remaining species of the group and represents the only representative of the N. picadoi group found in the Chorts Block For Oedipina clades corresponding to the subgenera Oedipina Oedipinola and Oeditriton were re covered as monophyletic ( Figure 4 10). Samples of Oedipina from five localities in northern Nicaragua were included, and represent two unnamed lineages, O. sp. inquirenda 2 and O. sp. inquirenda 3 ; one in the Oeditriton clade and one in the Oedipina clade. Oedipina sp. inquirenda 2 found at three isolated mountain localities in northern Nicaragua, is a well supported sister lineage to the other highland dwelling species of Oeditriton O. kasios with the lowland form O. quadra remaining as sister to the tw o highland lineages The second Nicaraguan candidate species, O. sp. inquirenda 3 is sister to a clade containing O. cyclocauda and O. pseudouniformis ( Figure 4 10). The nominal species O. gephyra from Parque Nacional Pico Bonito and

PAGE 196

196 Reserva de la Vida S ilvestre Texiguat, contains two monophyletic lineages (one at each locality), and is itself a monophyletic group. Oedipina tomasi is the sister taxon to the Oedipina gephyra complex, forming a monophyletic clade that is sister to the remaining Oedipinola A sample from a previously unknown allopatric population of Oedipina from central Honduras, O. sp. inquirenda 1 is shown to be conspecific with O. kasios ( Figure 4 10). Discussion Candidates for Further Taxonomic Study Among Anurans Undescribed diversity in the Chorts Highlands is both geographically and phylogenetically widespread. While a disproportionately large share of potential candidate species are salamanders (n=16; 44%), unexpected diversity is present in all families included in this study wit h the exception of Microhylidae ( containing three widespread species that occur in the Chorts Block) Not surprisingly, the majority of Potential Candidate Species have distributions limited to one or a few localities in the Chorts Highlands, however e ve n lowland anuran taxa heretofore considered to be widespread throughout Central America (Khler 2011), such as Diasporus diastema Leptodactylus fragilis Pristimantis ridens Lithobates warszewitschii Rhaebo haematiticus and Smilisca baudinii demonstra te deep divergence indicative of species level diversification (Table 4 3) Below I present a brief account of each Potential Candidate Species, as well as unassigned species identified through this study providing details of distribution and taxonomy re lative to nominal taxa

PAGE 197

197 Incilius coccifer/ibarrai/porteri The toads of the Incilius coccifer species complex include three apparently parapatric species in Honduras, although the distribution and taxonomic status of these species are disputed by authorit ies (Mendelson et al. 2005; McCranie 2009). The taxonomy of these populations has long been recognized as inadequate in terms of characterizing the existing diversity (McCranie & Wilson 2002), and Mendelson et al. (2005) revised the Incilius coccifer compl ex, including resurrection of the taxon I. ibarrai ( Figure 4 11A) and description of a new taxon, I. porteri ( Figure 4 11B) to accommodate premontane and highland populations of I. coccifer complex toads in the Chorts Highlands. A leading authority on Ho nduran amphibians, James McCranie, steadfastly has refused to recognize these taxa as valid (McCranie & Castaeda 2007; I. coccifer complex by Mendelson et al (2005), a thorough review o f this complex is needed that includes language field guide the Honduran amphibian fauna. McCranie provided the following principal reasons for not recognizing the validity of I. ibarrai and I. porteri : 1. Mendelson presented morphological analyses of the I. coccifer complex) detailed ie & Castaeda 2007:125), and the lack of address given to those concerns in Mendelson et al (2005) monograph. 2. The single sample of Honduran I. coccifer sensu stricto included by Mendelson et al (2005) is from the mesic portion of eastern Honduras, whi ch McCranie (by inference) considers distinctive from populations of the Pacific lowlands and therefore not representative of typical I. coccifer 3. Mendelson et al. (McCranie 2009:4).

PAGE 198

198 Results from my 16S and COI data support recognition of both I. ibarrai and I. porteri as valid species ( Figure 4 3 ), and also demonstrate that populations of I. coccifer from eastern Honduras are indeed conspecific with those from the Pacific lo wlands and apparently have a continuous distribution through subhumid areas around the southern terminus of the Nuclear Central American highlands in northern Nicaragua. The distribution of these taxa is more complicated and intertwined than was initially recognized by Mendelson et al (2005), and includes at least one area of sympatry or near sympatry between I. ibarrai and I. porteri on the southern side of Parque Nacional Cerro Azul Membar in Departamento de Comayagua (CAC044, I. porteri from Ro Bonit o, 1,570 m elevation; and IRL002 and IRL 004 05, I. ibarrai from San Jose de Los Planes, 1,290 m elevation, approximately 6 airline km east northeast of Ro Bonito and JHT 2205 ( Figure 4 11A) from Cerro Zarciadero, 1,835 m, approximately 6 km south of Ro Bonito ). Even more curious, samples representing both I. coccifer and I. porteri were collected on Isla del Tigre in the Golfo de Fonseca, with I. coccifer (JHT3301) know from 190 m elevation and I. porteri (JHT 3302) known from 520 m elevation. Clearly, f urther sampling and investigation is needed to determine identify areas of sympatry, and explore ecological or behavior al mechanisms isolating these closely related species. Incilius coniferus The toads referred to the taxon Incilius coniferus are distributed from northern Nicaragua (Travers et al. 2011) south to the Pacific versant of Colombia and northern Ecuador (Savage 2002). A single sample from Bosawas ( Figure 4 11C) is included in a clade with the holotype of I. karenlipsae (Mendelson & Mulcahy 2010) and eight samples referred to as I. coniferus ( Figure 4 3). In comparisons with the available COI

PAGE 199

199 Figure 4 11 Candidate Species I: Bufonidae and Craugastoridae. A) Incilius ibarrai Cerro Zarciadero, 1 800 m ( Photo J.H. Townsend [JHT]) B) I. porteri Cerro Uyuca 1,640 m C) I. cf. coniferus Biosfera Bosawas, 180 m (Photo Scott Travers) D) Rhaebo cf. haematiticus Biosfera Bosawas, 180 m (Photo Javier Sunyer) E) Craugastor cf. aurilegulus adult fe male, San Jos de Texguat, 200 m ( Photo JHT) F) C cf. aurilegulus adult male, same data as E ( JHT) ; G) C. laevissimus Ro Negro de Comayagua, 1,220 m (Photo JHT). H) C. laevissimus Cerro Azul Membar, 800 m (Photo JHT)

PAGE 200

200 data, the Bosawas sam ple (N416) is a strongly supported, distant sister to eight other samples from Panam ( Figure 4 3 ). Rhaebo haematiticus This species is considered a relatively common and widespread inhabitant of lowland and premontane rainforest from eastern Honduras to northern South America (McCranie et al. 2006, Savage 2002). The Nicaraguan samples of Rhaebo haematiticus ( Figure 4 11D) forms a well clade to Panamanian reference samples for both COI and 16S ( Figure 4 3 ). Craugastor aurilegulus This species is currently recognized as endemic to the Cordillera Nombre de Dios. Sampl es of C. aurilegulus from the vicinity of the type locality (Pico Bonito) and the vicinity of Texguat ( Figures 4 11E, 4 11F) clades for both COI and 16S ( Figure 4 4) and accordingly the Texguat population is assigned Deep Conspecific Lineage status and warrants further investigation to clarify its taxonomic status Craugastor laevissimus These data confirm the species C. laevissimus as a widespread member of the C. rugulosus group with a distribution congruent with tha t defined by Campbell & Savage (2000). Samples from the vicinity of the type locality in the Sierra de Omoa (JHT1824) and from throughout the serrana in Honduras ( Figures 4 11G, 4 11H; JHT series samples) and into northern Nicaragua (N series samples) de monstrate relatively low divergence between samples and no outwardly clean pattern of haplotype distribution ( Figure 4 4)

PAGE 201

201 Craugastor sp. inquirenda 1 & 2 Two Unconfirmed Candidate Species ( Figure 4 12A) from northern Nicaragua form a well supported (boot divergent clade with C. bransfordii samples from Panam ( Figure 4 4 ). It may be that one of these populations will be assigned to C. lauraster a small species known from eastern Honduras and northern Nicaragua (Khler 2011), followin g investigation of morphological affinities of these samples. Diasporus diastema Only a single putative species of Eleuthodactylidae, D. diastema is considered to naturally occur in northern Central America (Khler 2011, McCranie et al. 2006). The type l ocality of D. diastema is at the extreme eastern edge of its distribution in the Panama Canal Zone, and samples from northern Nicaragua ( Figure 4 12B) clearly represent a monophyletic lineage divergent from Panamanian samples ( Figure 4 4 ). Further investig ation is needed, preferably with inclusion of samples in the dataset from eastern Honduras, southern Nicaragua, and Costa Rica. Plectrohyla cf. guatemalensis The taxon P. guatemalensis has long been recognized as a potential composite taxon masking undesc ribed diversi ty (Duellman & Campbell 1992: 8, McCranie & Wilson 2002: 301), and the ability to study the phylogenetic relationships of populations assigned to this taxon has suffered from declines and disappearances of this cloud forest treefrog from many localities where it previously was abundant. I have only encountered one population of treefrogs assigned to P. guatemalensis (type locality: Patzi cia, Departamento de Chimaltenango, Guatemala; Stuart 1963) : cloud forest above 1,750 m in Parque N acional Mo ntaa de Yoro ( Figures 4 12C, 4 12D) In both

PAGE 202

202 the 16S and COI NJ trees, this population is recovered as a divergent clade and below the intra generic level does not demostrate any particular relationship to the single reference availabl e sequence ( Figure 4 4 ). Populations assigned to this species are know n from across the Ch orts Highlands (Duellman 2001), and a lack of representative sampling will likely continue to hinder phylogen y based taxonomic evaluation Ptychohyla hypomykter Samples from across th e serrana in Honduras and Nicaragua, including the type locality at Quebrada Grande appear to represent a single widespread species. However single reference sequence assigned to this taxon (ENS8486 from Izabal, Guatemala; Faivovich et al. 2005) is not r ecovered as part of the P. hypomykter clade nor does it cluster closely with any other species of Ptychohyla representing an Unconfirmed Candidate Species ( Figure 4 5 ). Ptychohyla spinipollex This species is presently considered endemic to the vicinitie s of Texguat and Pico Bonito in the Cordillera Nombre de Dios. A single 16S reference sequence from Quebrada de Oro in Parque Nacional Pico B onito was surprising divergent from a series of samples from La Liberacin in Refugio de Vida Silvestre Texguat, well into the range normally applied to intraspecific relationships ( Figure 4 5 ). Given the large genetic distance between the Pico Bonito sample and the 18 samples from Texguat, it is somewhat surp rising to see that the extreme variability in color patt ern variation exhibited by P. cf. spinipollex from Texguat ( Figure 4 12E H ) was not reflected by genet ic variation in 16S and COI.

PAGE 203

203 Figure 4 12 Candidate Species II: Eleutherodactylidae and Hylidae. A) Craugastor sp. inquirenda 1 Cerro Saslaya, 1,000 1,600 m (Photo Javier Sunyer) B) Diasporus cf. diastema Biosfera Bosawas, 180 m (Photo Javier Sunyer). C) Plectrohyla cf. guatemalensis, adult fe male, Monta a de Yoro, 1 820 m (Photo J.H. Townsend [JHT]). D) P. cf. guatemalensis, adult male, Monta a de Yoro, 1820 m (Photo JHT). E G) Ptychohyla cf. spinipollex La Liberacin de Texguat, 1,080 1,120 m. (Photo JHT).

PAGE 204

204 Smilisca baudinii This large treefrog is Mxico to Panam, and is often abundant around areas of human disturbance. It is likely that the disregard shown this taxon in the past, despite indications that there is somewhat considerable geographically linked morphological variation across its distribution (Duellman 2001), has le d to an underestimation of species level diversity Both 16S and COI data support this notion, with two potential candidate species (one in Honduras, one in northern Nicaragua) that are relatively deeply divergent from available reference sequences assigna ble to S. baudinii ( Figure 4 5 ). Leptodactylus fragilis This species is currently considered to have a range extending from Texas to Samples assigned to L. fragilis f rom Honduras and Nicaragua form a clade that is deeply divergent with respect to Panamanian samples referred to L. fragilis ( Figure 4 4) Given that the type locality is in Mxico but remains unsampled, I will continue to refer to Honduran and Nicaraguan s amples as L. fragilis and consider Panamanian samples to be an Unconfirmed Candidate Species pending further investigation. Lithobates brownorum X forreri An unresolved taxonomic problem in the Chorts Highlands relates to the status of populations of Li thobates that occur at high elevation ponds and wetlands in the Southern Cordillera of the Chorts Highlands McCranie and Wilson (2002: 478 479) considered these frogs to represent morphological intermediates that possess a blend of diagnostic features fr om both L. brownorum on the Caribbean versant and L. forreri on the Pacific versant, and judged these intermediate populations to represent hybrids

PAGE 205

205 between the lowland forms L. brownorum and L. forreri ( Figures 4 13A, 4 13B) These high elevation pond bree ding frogs apparently have an extended larval period, with tadpoles regularly reaching or exceeding the size of adult males ( Figure 4 13B). A total of 15 samples were analyzed eight from Cerro Uyuca in Departamento de Francisco Morazn and seven from San Pedro La Loma in Departamento de Intibuc ( Tabl e 4 1 ). All samples are supported as a single clade representing a P otential C andidate S pecies ( Figure 4 6) Analysis of 16S places this candidate species as sister to an undescribed species from the central highlands of Costa Rica ( Rana sp. 5 sensu Hillis & Wilcox 2005). The addition of data from the mitochondrial gene 12S would allow for these Lithobates sp. inquirenda samples to be directly compared to a geographically comprehensive dataset of Mexican Litho bates (Zaldvar Rivern et al. 2004) in addition to that of Hillis & Wilcox (2005), and should further elucidate the relationships of these Honduran highland populations. Lithobates brownorum a series of unresolved taxonomic problems related to Central American frogs of the L. berlandieri group. The identity of lowland forms from the Caribbean versant of Honduras and northern Nicaragua, variously referred to as L. berlandieri (McCranie & Wilso n 2002) and L. brownorum (after Zaldvar Rivern et al. 2004), is unconfirmed due to a lack of comparative sequence data overlapping this dataset and both those of Zaldvar Rivern et al. (2004) and Hillis & Wilcox (2005) Lithobates forreri The status of the lowland frogs from the Pacific versant is even less clearly defined, and they have been assigned to L. forreri from Mxico to Costa Rica (McCranie

PAGE 206

206 Figure 4 13 Candidate Species III: Ranidae and Strabomantidae. A) Lithobates sp. 1 brownorum X forr eri Cerro Uyuca, 1,640 m ( Photo Jason Butler). B) Lithobates sp. 1 brownorum X forreri comparing tadpole and adult male, Cerro Uyuca, 1,640 m (Photo Jason Butler). C) Lithobates cf. maculatus, adult male, Cerro Saslaya, 1,000 1,600 m (Photo Ja vier Sunyer). D) L. cf. maculatus, adult male, Cerro Saslaya, 1,000 1,600 m (Photo Javier Sunyer). E) L. cf. maculatus, adult fe male, Monta a de Yoro, 1 820 m (Photo J.H. Townsend). F) Lithobates cf. warszewitschii Biosfera Bosawas, 180 m (Photo Sco tt Travers). G) Pristimantis cf. ridens Biosfera Bosawas, 180 m (Photo Javier Sunyer). H) P.cf. ridens Cerro Saslaya, 180 m (Photo Javier Sunyer).

PAGE 207

207

PAGE 208

208 & Wilson 2002; Savage 2002). Based on phylogenetic analysis of 12S mtDNA data, Zaldvar Rivern et al. (2004:47) demonstrated that the taxon L. forreri is referable only to frogs from southern Sonora, Mxico, south along the Pacific coastal plain not farther than northern Jalisco, and in Mxico alone there are no fewer than four well supported species m asked under the name L. forreri The relationship between Pacific lowland populations assigned to L. forreri from Guatemala, El Salvador, Honduras, Nicaragua, and Costa Rica, as well as the taxonomic assignment of various populations, remains unresolved. Lithobates maculatus The putative species Lithobates maculatus currently is considered to be found at premontane and lower montane elevations (and peripherally in the lowlands) throughout Nuclear Central America (IUCN 2011; Khler 2011). Both 16S and COI data indicate the existence of at least four discretely distributed P otential C andidate S pecies among the Chorts Highland samples ( Figure 4 6 ): one from northeastern Nicaragua, including Cerro Saslaya and adjacent lowland rainforest areas ( Figure 4 13C, 4 13D) ; one from highland areas in north central Nicaragua, including Cerro Kilamb and Peas Blancas; one from the southern side of the Texguat cloud forest and the northern side of Pico Pijol in north central Honduras; and one from the rest of the Hondu ran serrana ( Figure 4 13E) serrana is composed of four geographically discrete, reciprocally monophyletic subclades ( Figure 4 6 ). Two 16S sequence fragments assigned to L. maculatus were available for comparison: KU 195258, from 19 km NW Rizo de Oro, Oaxaca, Mxico and USNM 559483, from Quebrada de Oro in Parque Nacional Pico Bonito, Honduras (Table 4 1 ). The Quebrada de Oro sample appears to represent a fifth P otential C andidate Species ( Figure 4 6 ). The Oaxaca sample may or

PAGE 209

209 may not be typical L. maculatus whose type locality is Totonicapn, Guatemala, some 350 km southwest of the Oaxacan locality. Lithobates taylori The boundaries between the aforementioned L. brownorum and another Caribbean lowland leopard frog, L. taylori lack c lear definition (McCranie & Wilson 2002, Savage 2002, Hillis & Wilcox 2005). While L. taylori is generally considered to occur from somewhere in the central portion of Nicaragua and extend southward into Costa Rica and presumably Panam (Khler 2011), the boundary between it and L. brownorum to the north (if a boundary exists) is undefined, and the status of populations of Lithobates from Panam also is unclear (Hillis & Wilcox 2005). Interestingly, all Lithobates berlandieri group samples from highland are as in northern Nicaragua cluster with a single sample from a lowland site in eastern Nicaragua ( Figure 4 6 ) that is assigned to L. taylori by Hillis & Wilcox (2005), and I tentatively refer those samples to L. taylori pending additional sampling throughout eastern Nicaragua. Lithobates warszewitschii This species is considered a relatively common and widespread inhabitant of lowland and premontane rainforest from eastern Honduras south to central Panam (McCranie et al. 2006, Savage 2002). Previously publis hed barcoding data (Crawford et al. 2010) indicates that there are potentia l l y two cryptic species concealed under L. warszewitschii in Panam, and samples from northeastern Nicaragua ( Figure 4 13F) form a third clade for both 16S and COI ( Figure 4 6 ). P ristimantis ridens This species is considered a relatively common and widespread inhabitant of lowland and premontane rainforests from northern Honduras to northern South America

PAGE 210

210 (Wang et al. 2008). Samples from low and moderate elevation in northern Nicar agua ( Figures 4 13G, 4 13H) form a clade in both 16S and COI analyses, deeply divergent from samples from Panama ( Figure 4 2 ). The Panamanian samples themselves appear to represent three candidate species (Crawford et al. 2010). Candidate Species and Allo patric Populations of Salamanders I began this study with samples representing ten populations of salamanders that could not be assigned unequivocally to a known species, due either to morphological distinctiveness or from being found in a new locality tha t was geographically and ecologically isolated from congeners. Four of these populations ( Bolitoglossa sp. inquirenda 1 Nototriton sp. inquirenda 1 Oedipina sp. inquirenda 1 and Oedipina sp. inquirenda 2 ), do not correspond to any known species level ta xa, and have subsequently been described as distinct taxa (Sunyer et al. 2010, 2011; Townsend et al. 2009 a 2010 a ). One population Nototriton sp. inquirenda 2 represent s an undescribed species from the Sierra de Agalta in eastern Honduras and is further evaluated in Chapter 5 Another population, Nototriton sp. inquirenda 3 from Montaa de Botaderos, also represents an undescribed species. In the cases of Bolitoglossa sp. inquirenda 2 Nototriton sp. inquirenda 4 N. sp. inquirenda 5 and Oedipina sp. in quirenda 1 these samples were found to represent new allopatric populations of species previously thought to be endemic to a single highland area (Townsend et al. 2011 b ). These cases, as well as a summary of information related to potential candidate spec ies and other resulting taxonomic considerations are detailed below. Bolitoglossa (Magnadigita) sp. inquirenda 1 In June 2006, during the first herpetological expedition into the core zone of PN Montaa de Yoro, two specimens of an unknown salamander diagn osable to the

PAGE 211

211 Bolitoglossa ( Magnadigita ) dunni species group were collected in the vicinity of Cataguana (1 780 2 020 m elevation) in northern Departamento de Francisco Morazn. On a return trip in March 2007, five additional samples were collected at the same locality. BLASTN searches of 16S returned a high degree of similarity to B. decora from PN La Muralla (Table 4 4 ). No 16S sequence divergence was observed between samples series from two localities in Montaa de Yoro and the single available sequence for B. decora typically an indication that samples are conspecific. However, the Montaa de Yoro series differs from B. decora in coloration, head width, and digit length, as well as being allopatric and found wi thin a different elevational band This evi dence, supported by subsequent sequencing and comparative analysis of a widely used fragment of the mitochondrial gene cytochrome b (cyt b ), led Townsend et al. (2009 a ) to recognize the PN Montaa de Yoro population as a young but distinct species, Bolitog lossa cataguana ( Figures 4 14A, 4 14B) Bolitoglossa (Magnadigita) oresbia / sp. inquirenda 2 On 17 18 July 2007, I visit ed Cerro El Zarciadero in central Honduras, the type locality of Bolitoglossa oresbia finding a single subadult B. oresbia (UF 156333 ) active on vegetation at night. This species was described recently on the basis of three specimens collected from an irregular patch of remnant cloud forest (Lower Montane Wet Forest formation) less than 1 hectare in total extent on the peak of Cerro El Zarciadero (McCranie et al. 2005). This patch is the only remaining forest near the peak of this mountain, and is adjacent to a set of communications towers and surrounded by agricultural clearings given over primarily to corn and other staple crops. In th e original description, the authors indicated that B. oresbia

PAGE 212

212 and vulnerable distribution, this species is classified as Critically E nd angered on the IUCN Red List and could be considered one of the most threatened amphibians on earth. From 6 13 July 2008, we surveyed a ridge above Quebrada Varsovia (14.79913N, 87.89128W), on the southwestern side of Parque Naci onal Cerro Azul Membar i n Departamento de Comayagua. Seven specimens of Bolitoglossa were collected, One adult salamander ( Bolitoglossa sp. inquirenda 2; UF 156532) with a distinctive golden yellow coloration covering most of its body ( Figure 4 14C) was collected in moist leaf li tter during the morning of 12 July 2008 along the trail leading to the aforementioned campsite (14.79618N, 87.89527W), 1 560 m elevation. This golden individual was so distinctive in terms of color pattern that it was assumed to represent a distinct taxo n from B. oresbia ; however both distance based (sequence divergence between Zarciadero and Membar samples 0.0 0.2 % for 16S and 0.1% for COI) and phylogenetic analyses confirm that this individual is conspecific with typical B. oresbia ( Figures 4 7). O ne a dult male (UF 156529 ) and four small juven iles (UF 156526 27, 156530 31) were found while active on vegetation at night along a trail that follows a steep hillside through undisturbed Lower Montane Wet Forest, 1 640 1 680 m elevation An adult female (UF 156528 ; Figure 4 14D ) was collected along the same trail during the daytime, when it was dislodged from a standing, rotten tree trunk approximately 1 m above the ground. Two of the juveniles were red orange in color and resembled the golden morph of B. or esbia ( Figure 4 14E), while the adults ( Figure 4 14D) and other two juveniles ( Figure 4 14F) demonstrate coloration more typical of B. oresbia These new localities are only around 8 airline km north of Cerro Zarciadero,

PAGE 213

213 Figure 4 14 Candidate Species IV: Bolitoglossa A) Bolitoglossa sp. inquirenda 1 ( = B. cataguana ) adult female holotype, Monta a de Yoro, 1,820 m; B) B. sp. inquirenda 2 ( = B. oresbia ) subadult male, Cerro Azul Membar, 1,560 m; C) B. sp. inquirenda 2 ( = B. oresbia ) subadult male, Cerro Azul Membar, 1,640 m; D) B. sp. inquirenda 2 ( = B. oresbia ) adult female, Cerro Azul Membar, 1,640 m; E) B. sp. inquirenda 2 ( = B. oresbia ) juvenile, Cerro Azul Membar, 1,640 m; F) B. sp. inquirenda 2 ( = B. oresbia ) juvenile, Cerro Azul Memba r, 1,640 m; G) B. celaque adult female, Cerro Celaque, 2,560 m. Photos J.H. Townsend.

PAGE 214

214 and the intervening area, while wholly converted to agriculture, is no lower than 1 300 m elevation at any given point. Bolitoglossa (Magnadigita) celaque As current ly recognized, Bolitoglossa celaque is a highland salamander found across three mountainous areas of the Southern Cordillera of the Chorts Highlands includ ing the Sierra de Celaque ( Figure 4 14G; the type locality), the Sierra de Puca Opalaca, and the Mo ntaa de la Sierra. Both 16S and COI data indicate that samples from these three ranges each form clades ( Figure 4 7 ). Although divergence among these three clades is moderate ( 1.7 3.6% ) the clear phylogeographic structure (within mountain range divergenc 0.8 associated with these samples indicates they may represent a complex of relatively young species, and should be the focus of further study using finer scale population level approaches, environmental niche modeling, and c omparative morphology to clarify the systematic status of populations assigned to B. celaque Bolitoglossa (Magnadigita) conanti This highland salamander is presently considered to occur in the Sierra de Omoa, Sierra de Espritu Santo, and Sierra del Meren dn, and is characterized by a high degree of variability in coloration within populations (McCranie & Wilson 1993, 2002). Both 16S and COI data revealed clades corresponding to geographically isolated mountains ranges ( Figure 4 7 ). As with B. celaque div ergence among these clades is moderate (2.5 3.0% for COI) and within mountain range is very low ( 0.0 0.2 % for 16S and 0.0% for COI) indicating relatively recent diversification and should be the focus of further study using multiple avenues of finer scal e investigation.

PAGE 215

215 Bolitoglossa (Magnadigita) porrasorum As with the two other species of Chorts Highland Magnadigita currently thought to occur at more than one isolated cloud forest locality, B. porrasorum populations are not recovered as conspecific in d istance based analyses of 16S or COI data ( Figure 4 7) or ML analysis of the combined dataset ( Figure 4 9) Populations of B. porrasorum from the Sierra de Sulaco (where the type locality is found) and Texguat and Pico Bonito in the Cordillera Nombre de D ios each form geographically discrete clades although the lineages assigned to B. porrasorum demonstrate greater genetic distances than those of the B. celaque and B. conanti complexes and are potentially paraphyletic or polyphyletic with respect to B. ca taguana B. decora and B. longissima ( Figure 4 7 ). Samples from Texguat demonstrate a remarkable degree of color polymorphism ( Figure 4 15), however all samples from this locality appear to represent a single taxon ( Figure 4 7). Bolitoglossa (Nanotriton) rufescens/nympha Salamanders of the Bolitoglossa rufescens complex in the Chorts Highlands have a confusing taxonomic history (McCranie & Wilson 2002). Recently, Campbell et al. (2010 ) described a new species from this complex, B. nympha from a localit y on the western side of the Sierra de Caral in eastern Guatemala. Unfortunately, this description did little to resolve the systematic relationships of the B. rufescens complex in the Chorts Highlands, as it was based on 12S and cyt b sequence data from a single sample. The use of only a single sample precludes characterization of diversity and distribution of B. nympha and the choice of 12S in favor of the previously ubiquitous (at least for neotropical salamanders) 16S minimizes the ability of other re searchers to directly ) results with the comprehensive existing

PAGE 216

216 Figure 4 15 Candidate Species V: Bolitoglossa cf. porrasorum A F) variation in color pattern in conspecific samples of Bolitoglossa cf. porrasorum La Libe racin de Texguat, 1,080 1 420 m Photos J.H. Townsend. dataset available at NCBI. Comparative morphological analyses of other species and populations in the B. rufescens complex, particularly the multiple known populations from across the border (albe it in some cases in the same mountains) in Honduras are also not available. Based on the type locality and description of B. nympha I am

PAGE 217

217 inferring that four samples from two localities, one in the Sierra de Omoa and one in the Sierra de Espritu Santo, ar e conspecific with B. nympha ( Figure 4 7 ). Surprisingly, of five samples collected syntopically in El Paraiso Valley on the northern slope of the Sierra de Omoa, three appear to be conspecific with B. nympha while two others represent a deeply divergent (d ivergence between syntopic samples 5.5 % for 16S and 17.6% for COI) cryptic species that is recovered in the 16S NJ tree as the sister to a sample assigned to B. rufescens (MVZ 194254) from Chiapas, Mxico. Nototriton barbouri All analyses support that two populations currently assigned to N. barbouri from the Cordillera Nombre de Dios are paraphyletic with respect to a sample from the vicinity of the type locality of N. barbouri ( Figures 4 8, 4 10 ). Furthermore, those two populations, one from Texguat and one from Pico Bonito (McCranie 1996a) represent two Potential C andidate S pecies, forming a clade with N. brodiei from the Sierra de Omoa and Sierra de Caral ( Figure 4 8 ) A systematic revision of this taxon is presented in Chapter 5. Nototriton sp. inqu irenda 1 During a trip in April 2008 to the leeward side of RVS Texiguat in the vicinity of La Fortuna, a single specimen of Nototriton was collected that possesses a series of morphological characteristics unique among Honduran congeners. This single sala mander had greatly large nares, syndactylous hands and feet with pointed toe tips, and a pale ventral surface with light mottling, characteristics typical of species in the N. richardi group (Costa Rica) or the genus Cryptotriton Data from 16S and COI ind icate that this species forms a sister clade to the remaining species of Chorts Highlands

PAGE 218

218 Nototriton ( Figure 4 8 ), and this species was described as N. tomamorum by Townsend et al. (2010 a ). Nototriton sp. inquirenda 2 Two specimens of Nototriton were col lected in July 2010 from cloud forest in PN Sierra de Agalta, the first record of this genus from eastern Honduras. Both distance and model based analyses of COI and 16S data supports these samples as conspecific and representative an unnamed species ( Figu re 4 8) which is further analyzed and formally described in Chapter 5 (Townsend et al. 2011 a ). Nototriton sp. inquirenda 3 Four specimens of Nototriton were collected in April 2011 along the highest ridge (1,700+ m elevation) in the Sierra de Botaderos i n northeastern Olancho. Both distance and model based analyses of COI and 16S data indicate that these samples are conspecific and represent a new candidate species that is sister to Nototriton sp. inq. 2 ( Figure 4 8). This candidate species is being descr ibed outside of this dissertation Nototriton lignicola / sp. inquirenda 4 On 10 June 2006, we collected a juvenile Nototriton (UF 156544) from inside a fractured rock alo ngside Quebrada Cataguana (15.01N, 87.10W ), 1 820 m elevation, Parque Nacional Mon taa de Yoro, in northern Departamento de Francisco Morazn. On 14 March 2007, we collected two adult Nototriton (UF 156542 43 ) from inside rotten logs on Cerro El Filn above Quebrada Cat aguana 2 020 m elevation. These samples are shown to be conspecific with N. lignicola ( Figure 4 8 ), a Critically Endangered species previously known only from 13 specimens collected inside two rotten logs at Cerro de Enmedio, Par que Nacional La Muralla 1 760 1 780 m elevation (McCranie & Wilson 2002 ).

PAGE 219

219 Nototriton limnosp ectator / sp. inquirenda 5 Nototriton limnospectator is an Endangered moss salamander previously known only from forest above 1 600 m elevation in Parque Nacional Mon taa de Santa Brbara an isolated karstic mountain to the west of L ago de Yojoa During 2 008, four specimens of Nototriton we re collected from three sites in Parque Naci onal Cerro Azul Membar including two i ndividuals (UF 156539 40 ) found on 7 and 9 July 2008 alongside juvenile Bolitoglossa oresbia (UF 156526 27, 156530 31) while active on l ow vegetation at night. Another individual (UF 156541) was collected on 27 August 2008 from near a campsite farther up the Quebrada Varsovia (14.80N, 87.89 W), 1 710 m elevation, while it was active at night on a root buttress. All three of these localiti es are in the Lower Montane Wet Forest formation. In addition, a juvenile N. limnospectator (UF collec tion ) was collected from underneath a moss mat at night on 7 June 2008 along Sendero Bosque Nublado, 1 105 m elevation, above Centro de Visitantes de Parq ue Nacional Cerro Az W), Departamento de Corts, a locality that lies in the Premontane Wet Forest formation. These samples were shown to be conspecific with reference samples of N. limnospectator from Parque Nacional Mon taa de Santa Brbara ( Figure 4 8). Th e new localities are approximately 20 25 km east of the nearest locality in Parque Nacional Montaa de Santa Brbara (McCranie & Wilson 2002), from which they are isolated by Lago de Yojoa the largest freshwater la ke in Honduras. Oedipina kasios / sp. inquirenda 1 On 25 September 2008, a single specimen of Oedipina ( Figure 4 16A; UF 156500 ) was collected in a small mesic ravine on Montaa de la Sierra (14.95N, 87.061W ), 1 920 m elevation, in coniferous cloud fores t on the southeastern side of Parque

PAGE 220

220 Nacional Montaa de Yoro. This specimen was shown to be conspecific with Oedipina kasios Figure 4 8) a species recently described from Parque Nacional La Muralla as a member of a divergent, Chorts Highlands endemic cl ade (subgenus Oeditriton ; McCranie et al. 2008). This species occurs from 950 1 780 m elevation, and has been found in the same logs as Nototriton lignicola (McCranie et al. 2008). The Critically Endangered amphibians Bolitoglossa cataguana and Plectrohyla guatemalensis were also collected within and along the outer margins of this and similar ravines. The forest in this part of PN Montaa de Yoro is typified by regularly burned pine oak forest with an open grassy understory interrupted by deep, narrow seep age ravines that support dense mesic vegetation providing refuge for species like O. kasios during fire events. Oedipina nica / sp. inquirenda 2 Nicaraguan worm salamanders ( Oedipina ) have long been a source of taxonomic uncertainty, the byproduct of cons erved morphology and sparse sampling. During 2008 2009, a series of 18 Oedipina were collected from three highland localities in northern Nicaragua ( Figure 4 16B) These samples could not be assigned to a taxon based on comparisons from BLASTN searches, NJ tree clustering, or sequence divergence, and appeared to represent an undescribed species of the subgenus Oeditriton ( Figure 4 8) Sequencing and subsequent analysis of cyt b data as well as comparative morphology confirmed that these samples represent a new species, which we recently described as Oedipina nica ( S unyer et al. 2010). Oedipina koehleri / sp. inquirenda 3 In his pioneering monograph of Oedipina Brame (1968) described the taxon O. pseudouniformis from central Costa Rica, and included eight paratypes collected in July Hacienda La Cumplida, 1.5 km north of Matagalpa, 731 m elevation

PAGE 221

221 Figure 4 16 Candidate Species VI: Oedipina A) Oedipina sp. inquirenda 1 ( = O kasios ) Monta a de Yoro, 1,920 m (Photo J.H. Townsend). B) O. sp. inquirenda 2 ( = O nica ) Macizos de Peas Blancas 1,515 m (Photo Scott Travers). C) O. sp. inquirenda 3 ( = O koehleri ) Cerro Musn, 724 m (Photo Javier Sunyer) Figure 4 17 Candidate Species VII: Oedipina cf. gephyra ( = O petiola ) A) do rsal view of right hind foot of O cf. gephyra ( = O petiola ) Cerro Bfalo, 1,580 m; B) dorsal view of right hind foot of O. gephyra Texguat, 1,820 m; C) ventral view of right hind foot of O.cf. gephyra ( = O petiola ) ; D) ventral view of right hind foo t of O. gephyra Texguat, 1,820 m. Photo s J.H. Townsend.

PAGE 222

222 Departamento de Matagalpa, Nicaragua. Premontane forests in this area have been severely degraded since 1957, and remaining patches of mesic forest are restricted to the most marginal upper port ions of the surrounding peaks. Khler et al. (2004) later assigned two specimens from pristine forest between approximately 600 and 945 m elevation in Parque Nacional Cerro Saslaya, Regin Autnoma Atlntico Norte, Nicaragua, to O. pseudouniformis recogni zing that these samples appeared conspecific with those from Hacienda La Cumplida. A nalysis of recently collected sample s from PN Cerro Saslaya and an isolated mountain to the south, Cerro Musn ( Figure 4 16C) revealed them to represent a single undescrib ed species ( Figure 4 8) We described this species as O. koehleri based on evidenc e from phylogenetic analyses, ma c r oeco logical models, and morphology (Sunyer et al. 2011) Oedipina gephyra The endemic species Oedipina gephyra was described from the leewar d side of Reserva de Vida Silvestre Texguat in the western portion of the Cordillera Nombre de Dios, Honduras (McCranie et al. 1993), and soon after was reported from Parque Nacional Pico Bonito in the central portion of the Cordillera Nombre de Dios (McC ranie 1996 b ). A single representative from each of the two populations assigned to O. gephyra was included in the phylogenetic analyses of Garca Pars & Wake (2000), who reported interspecific level sequence divergence between the two samples ( Garca Par s & Wake 2000: 70), suggesting that more than one species may be concealed under O. gephyra Multiple attempts to secure addition al material from PN Pico Bonito were unsuccessful, however the addition of two new samples of O. gephyra from the vicinity of the type locality demonstrate s that the two populations are reciprocally monophyletic ( Figure 4 8 ), support ing recognition of two species. Following the

PAGE 223

223 phylogenetic results, McCranie & Townsend (2011) also found diagnostic differe nces in foot morphology ( Figure 4 17) and subsequently described the Pico Bonito population as O. petiola Amphibian Endemism and Conservation Priorities in the Chorts Block The Chorts Block is already known to possess a diverse endemic amphibian fauna, with 74 regionally ende mic species with distributions restricted to within the region (Chapter 3, Table 3 5) The identification of as many as 36 new species from a subset of about 36% of named Chorts Block diversity is largely underestimate d. One third of the Potential Candidate Species (12/36) appear to be restricted to localities in the Northern Cordillera of the Chorts Highlands, a region already known to suppor t the highest level of amphibian endemism in the Chorts Block (Chapter 3, Ta ble 3 6 ). It comes as little surprise to me that six Potential Candidate Species are from the vicinity of Texguat at the western end of the Cordillera Nombre de Dios (Table 4 3) a mountain range identified as one of most significant endemism hotspots in Mesoamerica ( Chapter 3; Townsend et al. 2010a, 2011c, Submitted A ). Four more species are from the vicinity of Pico Bonito, and another two from the Sierra de Omoa (Table 4 3). Given that established protected areas e n compass most of the remaining highland forest in each of these three mountains ranges, tangible opportunities for conserving extant populations of both named and un named endemic species already exist and c ould be strengthened by the description of additional endemic diversity from within each If it is assumed that at least the CCS and UCS represent distinct species, then, including the six candidate species that have already been formally described, a total of 30 new endemic amphibian spec ies are identified in the Chort s Block. Of these 30, at

PAGE 224

224 least 12 would immediately qualify as Critically Endangered based on IUCN Red List criteria ( IUCN 2001 ), and at least an additional nine species would be classified as Endangered (Table 4 3). A somewhat surprising finding was the id entification of Potential Candidate Species from sampled populations assumed a priori to represent widespread lowland taxa such as Diasporus diastema Lithobates warszewitschii Pristimantis ridens Rhaebo haematiticus and Smilisca baudinii Should these populations be confirmed a s distinct species, the implication that there exists a previously unknown endemic amphibian fauna inhabiting the Caribbean lowlands of northeastern Nicaragua (and presumably eastern Honduras) would rather dramatically alter the u nderstanding of patterns influencing Chorts Block biogeography and diversification Given the large number of species that are presently considered to meet the northern limit of their distribution in this area (as was the case with the aforementioned cand idate species), this revelation would warrant a comprehensive re evaluation of conservation and management objectives in the transboundary Mosquitia region which is currently aimed at the maintenance of ecological corridors principally to support populat ions of large mammals such as jaguar, tapir, and white lipped peccaries (CIPF 2009). Accomplishment of a rapid amphibian inventory, emphasizing the collection of molecular data, is urgently needed to assess the extent of endemic cryptic diversity currently overlooked in the Caribbean lowlands of the Chorts Block. Iterative Taxonomic Approaches to Biodiversity Inventory In this chapter, I have provided the first comprehensive assessment of amphibian evolutionary diversity for the Chorts Block of Central A merica a region already recognized for its elevated endemism. I have deliberately employed an iterative,

PAGE 225

225 heuristic approach to the systematic evaluation of regional diversity, with two principal aims in mind. First, to initiate development of a comprehens ive molecular reference dataset upon which future work in the region can be directed and compared. Doing so contributes to the goals set out by both the Systematics Agenda 2000 (1994) and the IUCN Global Amphibian Assessment (2007) chief of which are the e xecution of rapid phylogeny of all species to serve as the basis for classification and as a framework for use by researchers in the life sciences. Second, the use of this ap proach is advantageous in that it allows for, what I consider to be, the appropriate application of DNA barcoding data and methods in taxonomy. After establishing the ecophysiographic setting in Chapter 2 and providing a comprehensive faunal trea t ment of presented and analyzed a comprehensive two gene DNA barcoding dataset for Chorts Block amphibians. As part of an iterative process, the distance based analyses used in association with DNA barcoding provide a vital step that can be used to guide how and where to focus analytical resources moving forward. In this study the Class Caudata (the salamanders) represents both the most taxonomically comprehensive dataset among groups of amphibians analyzed, and presents bot h the most Potential Candidate Species (Table 4 3) and the highest degree of extinction risk (Chapter 3, Table 3 5). As the next step, I used the two gene barcoding dataset as the basis for model based maximum likelihood (ML) phylogenetic analysis in order to generate a phylogenetic hypothesis that can be used to more rigorously establish species limits than can be done using distance based methods alone, and, to address a principal deficiency of

PAGE 226

226 distance based analysis, estimate evolutionary relationships at a deeper level than that of species and species delimitation. Utilization of the two gene approach to DNA barcoding in this case al lows for the initial dataset to be more informative phylogenetically and therefore of more utility at later stages in the process of taxonomic evaluation. In the case of Chorts Block salamanders, DNA barcoding analyses indicated that there are at least 12 CCS and USC and another four DCL that are in need of further systematic study (Table 4 3). Phylogenetic analysis support s each of these Potential Candidate Species as a monophyletic group ( Figures 4 9, 4 10), and revealed further details of evolutionary relationships within and among amphibian clades One group in particular, the moss salamanders ( Nototriton ), exhibited a r elatively high degree of cryptic diversification, and will serve as the model group for use in my subsequent systematic evaluation. Phylogenetic analyses recovered three geographically circumscribed clades ( Figure 4 10): a Northern Cordillera clade consist ing of N. brodiei from the Sierra de Omoa and Sierra de Caral, N. sp. 2 CCS from Pico Bonito, and N. sp. 3 CCS from Texguat; a central clade, consisting of N. barbouri sensu stricto from the Sierra de Sulaco and N. limnospectator from Montaa de Santa Br ba ra and the Montaas de Membar; and an eastern clade, consisting of N. sp. inquirenda 2 (= N. picucha ; Townsend et al. 2011a) and N. sp. inquirenda 3 from the Sierra de Botaderos. Nototriton sp. inquirenda 1 (= N. tomamorum ; Townsend et al. 2010a), which p ossesses distinctive morphological traits, was recovered as the sister lineage to the rest of the Chorts Block endemic clade corresponding to the N. barbouri species group (sensu Garca Pars & Wake 2000 ). The Cordillera de La Flor La Muralla endemic N. l ignicola

PAGE 227

227 is recovered as sist er to the rest of the in group. Chapter 5 deals with the molecular and morphological systematics and taxonomic revision associated with this group, providing what is, in my estimation, the most critically overlooked yet definit ively fundamental activity for contemporary systematic biologists to engage in: the formal taxonomic treatment of candidate taxon identified in molecular studies. Conservation planners, ecologists, legislators, and the general public have little use for m etrics based on un named, un described diversity, and I do not foresee an entity such as Nototriton sp. inq. 1 conservation strategy or public outreach campaign. Despite t he best intentions and most dili gent labor of biologists, in the end it is the greater share of society, made up of non scientists which will determine the success or failure of this to preserve our shared biodiversity resources. Taxonomy provides the most basic means for w hich the specialist can identify and communicate information about biological diversity, and to carry out molecular inventories of biodiversity without promoting the direct taxonomic evaluation of the results is, in my view, a disservice to the study organ isms themselves, our field of study, and to society as a whole.

PAGE 228

228 Table 4 1. series; IRL = Ileana R Luque Montes field series; AJC = Andrew J. Crawford field series; AMNH = American Museum of Natural History; JSF = John Frost tissue collection; KRL = Karen R. Lips field series; KU = University of Kansas Natural History Museum; MVUP = Museo de Vertebratos de la Universidad de Panam SIUC = Herpet ology Collection, Southern Illinois University at Carbondale; USNM = National Museum of Natural History, Smithsonian Institution. Taxon Voucher Locality 16S COI Data source CAUDATA Plethodontidae (29) Bolitoglossa sp. inq. 1 JHT2087 Hondura s: Cataguana + This study (= B. cataguana ) JHT2113 Honduras: Cataguana + This study JHT2114 Honduras: Cataguana + This study JHT2115 Honduras: Cataguana + + This study JHT2125 Honduras: Cataguana + This study JHT2964 Honduras: above Guayma s + This study JHT2965 Honduras: above Guaymas + + This study Bolitoglossa celaque JHT2743 Honduras: Cerro Celaque + + This study JHT2744 Honduras: Cerro Celaque + + This study JHT2745 Honduras: Cerro Celaque + + This study JHT2747 Honduras: Ce rro Celaque + + This study JHT2748 Honduras: Cerro Celaque + + This study JHT2749 Honduras: Cerro Celaque + This study JHT2750 Honduras: Cerro Celaque + + This study JHT2751 Honduras: Cerro Celaque + + This study JHT2752 Honduras: Cerro Celaque + + This study JHT2753 Honduras: Cerro Celaque + + This study JHT2754 Honduras: Cerro Celaque + + This study JHT2755 Honduras: Cerro Celaque + + This study JHT2756 Honduras: Cerro Celaque + + This study JHT2757 Honduras: Cerro Celaque + + This s tudy JHT2903 Honduras: Intibuc + + This study JHT2904 Honduras: Intibuc + + This study JHT2910 Honduras: Intibuc + + This study JHT2911 Honduras: Intibuc + + This study JHT2912 Honduras: Intibuc + + This study JHT2618 Honduras: Guajiquiro + + This study JHT2867 Honduras: Guajiquiro + + This study JHT2868 Honduras: Guajiquiro + + This study JHT2869 Honduras: Guajiquiro + + This study JHT2306 Honduras: Intibuc + This study JHT2307 Honduras: Intibuc + + This study JHT2877 Hondu ras: Intibuc + + This study

PAGE 229

229 Table 4 1. Continued. JHT2878 Honduras: Intibuc + + This study JHT2879 Honduras: Intibuc + + This study JHT2880 Honduras: Intibuc + + This study JHT2887 Honduras: Intibuc + + This study JHT2888 Honduras: Intibuc + + This study JHT2889 Honduras: Intibuc + + This study JHT2890 Honduras: Intibuc + + This study Bolitoglossa colonnea CH 6526 Panam: El Cop FJ784318 FJ766578 Crawford et al. (2010) Bolitoglossa conanti JHT2922 Honduras: Cusuco + + This study JHT2923 Honduras: Cusuco + + This study JHT2924 Honduras: Cusuco + + This study JHT2925 Honduras: Cusuco + + This study JHT2725 Honduras: Ocotepeque + + This study JHT2741 Honduras: Ocotepeque + + This study JHT2742 Honduras: Ocotepeque + + Thi s study UTA A58141 Guatemala: Cerro El Mono + + This study Bolitoglossa diaphora JHT2917 Honduras: Cusuco + + This study JHT2918 Honduras: Cusuco + This study JHT2919 Honduras: Cusuco + + This study JHT2920 Honduras: Cusuco + + This study JHT 2921 Honduras: Cusuco + + This study JHT2989 Honduras: Cusuco + + This study JHT2990 Honduras: Cusuco + + This study JHT2991 Honduras: Cusuco + + This study Bolitoglossa dofleini JHT2430 Honduras: Texguat (Yoro) + + This study JHT2854 Honduras: Quebrada Grande + + This study UTA A56787 Guatemala + + This study Bolitoglossa dunni UTA A51488 Guatemala: Sierra de Caral + + This study Bolitoglossa heiroreias UTA A54712 Guatemala: Chiquimula + + This study Bolitoglossa longissima JHT3194 Hondur as: Cerro La Picucha + This study JHT3195 Honduras: Cerro La Picucha + + This study JHT3196 Honduras: Cerro La Picucha + + This study JHT3197 Honduras: Cerro La Picucha + + This study JHT3198 Honduras: Cerro La Picucha + This study JHT3199 Ho nduras: Cerro La Picucha + This study JHT3200 Honduras: Cerro La Picucha + This study JHT3201 Honduras: Cerro La Picucha + This study Bolitoglossa mexicana JHT2390 Honduras: Los Naranjos + This study JHT2572 Honduras: Comayagua + + This st udy JHT2573 Honduras: Comayagua + This study JHT2960 Honduras: above Guaymas + + This study JHT2512 Honduras: Azul Membar + This study JHT3182 Sierra de Agalta + This study

PAGE 230

230 Table 4 1. Continued. Bolitoglossa nympha 1 JHT1766 Honduras: El Pa raiso Valley + This study JHT1793 Honduras: El Paraiso Valley + + This study JHT1823 Honduras: El Paraiso Valley + + This study JHT2803 Honduras: Quebrada Grande + + This study Bolitoglossa oresbia JHT2225 Honduras: Cerro Zarciadero + + This stud y (includes Bolitoglossa sp. inq. 2) IRL071 Honduras: Azul Membar + + This study IRL072 Honduras: Azul Membar + + This study IRL074 Honduras: Azul Membar + + This study IRL075 Honduras: Azul Membar + + This study IRL079 Honduras: Azul Membar + + This study IRL080 Honduras: Azul Membar + This study IRL081 Honduras: Azul Membar + + This study Bolitoglossa porrasorum JHT2359 Honduras: Macuzal + + This study JHT2360 Honduras: Macuzal + + This study JHT2371 Honduras: Macuzal + + This study JHT2372 Honduras: Macuzal + + This study JHT2373 Honduras: Macuzal + + This study JHT2374 Honduras: Macuzal + This study JHT2419 Honduras: Macuzal + This study Bolitoglossa cf. porrasorum JHT2431 Honduras: Texguat (Yoro) + + This study JHT2432 Honduras: Texguat (Yoro) + This study JHT2449 Honduras: Texguat (Yoro) + + This study JHT2450 Honduras: Texguat (Yoro) + + This study JHT2453 Honduras: Texguat (Yoro) + + This study JHT2454 Honduras: Texguat (Yoro) + This study JHT2455 Honduras: Texguat (Yoro) + + This study JHT3104 Honduras: Texguat (Atlntida) + + This study JHT3105 Honduras: Texguat (Atlntida) + + This study JHT3106 Honduras: Texguat (Atlntida) + + This study JHT3143 Honduras: Texguat (Atlntid a) + + This study JHT3144 Honduras: Texguat (Atlntida) + + This study JHT3145 Honduras: Texguat (Atlntida) + + This study JHT3146 Honduras: Texguat (Atlntida) + This study JHT3147 Honduras: Texguat (Atlntida) + + This study JHT3148 Hond uras: Texguat (Atlntida) + + This study JHT3149 Honduras: Texguat (Atlntida) + + This study JHT3150 Honduras: Texguat (Atlntida) + This study JHT3151 Honduras: Texguat (Atlntida) + + This study JHT3152 Honduras: Texguat (Atlntida) + + T his study JHT3153 Honduras: Texguat (Atlntida) + + This study JHT3171 Honduras: Texguat (Atlntida) + + This study JHT3172 Honduras: Texguat (Atlntida) + + This study JHT3229 Honduras: Texguat (Atlntida) + + This study

PAGE 231

231 Table 4 1. Continued. JHT3230 Honduras: Texguat (Atlntida) + This study JHT3231 Honduras: Texguat (Atlntida) + + This study JHT3250 Honduras: Texguat (Atlntida) + + This study JHT3251 Honduras: Texguat (Atlntida) + + This study JHT3252 Honduras: Texguat (A tlntida) + + This study JHT3253 Honduras: Texguat (Atlntida) + + This study JHT3254 Honduras: Texguat (Atlntida) + + This study JHT3255 Honduras: Texguat (Atlntida) + + This study JHT3256 Honduras: Texguat (Atlntida) + + This study JHT32 57 Honduras: Texguat (Atlntida) + + This study JHT3264 Honduras: Texguat (Atlntida) + + This study JHT3265 Honduras: Texguat (Atlntida) + + This study JHT3266 Honduras: Texguat (Atlntida) + + This study Bolitoglossa cf. rufescens JHT1767 Hon duras: El Paraiso Valley + + This study JHT1797 Honduras: El Paraiso Valley + + This study Bolitoglossa schizodactyla USNM 572791 Panam: El Cop FJ784482 FJ766579 Crawford et al. (2010) Bolitoglossa striatula N145 Nicaragua: Bosawas + + This study B olitoglossa synoria JHT2900 Honduras: Cerro El Pital + + This study JHT2901 Honduras: Cerro El Pital + + This study Cryptotriton alvarezdeltoroi MVZ 158942 Mxico : Chiapas AF199196 Garca Pars & Wake (2000) Cryptotriton sierraminensis MVZ 160907 Gu atemala : Sierra Las Minas AF199198 Garca Pars & Wake (2000) Cryptotriton veraepacis GAR059 Guatemala : Baja Verapaz + + This study (E.N. Smith, UTA) UTA A51396 Guatemala : Baja Verapaz + + This study (E.N. Smith, UTA) MVZ 215913 Guatemala : Baja Vera paz AF199197 Garca Pars & Wake (2000) Dendrotriton rabbi UTA A51086 Guatemala: Uspantn AF199232 Garca Pars & Wake (2000) Nototriton abscondens UCR 12071 Costa Rica: Cascada La Paz AF199199 Garca Pars & Wake (2000) Nototriton barbouri JHT24 20 Honduras: Macuzal + + This study ( N. sp. A) USNM 339712 Honduras: Pico Bonito AF199201 Garca Pars & Wake (2000) ( N. sp. B) JHT3159 Honduras: Texguat (Atlntida) + + This study Nototriton brodiei UTA A5 1490 Guatemala: Sierra de Caral AF199202 + 16S: Garca Pars & Wake (2000); COI: This study (E.N. Smith, UTA) MVZ(FN252407) Honduras: Sierra de Omoa + This study (S.M. Rovito, MVZ) Nototriton gamezi MVZ 207122 Costa Rica: Monteverde AF199200 Garca Pars & Wake (2000) Nototriton guanacaste MVZ 207106 Costa Rica: Volcn Cacao AF199203 Garca Pars & Wake (2000) Nototriton lignicola (= N. sp. inq. 4) USNM 497540 Honduras: La Muralla AF199204 Garca Pars & Wake (2000) JHT2122 Honduras: Cataguana + + This study Nototriton limnospectator None Honduras: Santa Brbara + S.M. Rovito (MVZ) (= N. sp. inq. 5) IRL035 Honduras: Azul Membar + This study IRL070 Honduras: Azul Membar + + This study IRL076 Honduras: Azul Membar + + This study IRL 088 Honduras: Azul Membar + + This study

PAGE 232

232 Table 4 1. Continued. Nototriton picadoi MVZ 225899 Costa Rica: Tapant AF199205 Garca Pars & Wake (2000) Nototriton richardi UCR 12057 Costa Rica: Cascajal de Las Nubes AF199206 Garca Pars & Wake (2000) Nototriton saslaya N650 Nicaragua: Cerro Saslaya + This study SMF collection Nicaragua: Cerro Saslaya + This study Nototriton sp. inq. 1 (= N. tomamorum ) JHT2437 Honduras: Texguat (Yoro) + + This study Nototriton sp. inq. 2 JHT3192 Honduras: Ce rro La Picucha + + This study (= N. picucha ) JHT3193 Honduras: Cerro La Picucha + + This study Nototriton sp. inq. 3 JHT3398 Honduras: Sierra de Botaderos + + This study JHT3399 Honduras: Sierra de Botaderos + + This study JHT3400 Honduras: Sierra de Botaderos + + This study JHT3401 Honduras: Sierra de Botaderos + + This study Oedipina alleni MVZ 190857 Costa Rica: Sirena AF199207 Garca Pars & Wake (2000) MVZ 225903 Costa Rica: Damas AF199208 Garca Pars & Wake (2000) Oedipina carablanca None Costa Rica: Limn FJ196862 McCranie et al. (2008) Oedipina collaris SIUC H 08896 Panam: El Cop FJ196863 Crawford et al. (2010) Oedipina complex DBW5105 Panam: Barro Colorado AF199213 Garca Pars & Wake (2000) DBW5787 Panam: Cerro Campan a AF199212 Garca Pars & Wake (2000) Oedipina cyclocauda MVZ 138916 Costa Rica: La Selva AF199214 Garca Pars & Wake (2000) MVZ 293747 Costa Rica: La Selva AF199215 Garca Pars & Wake (2000) Oedipina elongata UTA A56809 Guatemala + + This stu dy (E.N. Smith, UTA) UTA A56810 Guatemala + This study (E.N. Smith, UTA) Oedipina gephyra JHT2443 Honduras: Texguat (Yoro) + + This study JHT2451 Honduras: Texguat (Yoro) + + This study USNM 530582 Honduras: Texguat (Yoro) AF199218 Garca Pa rs & Wake (2000) Oedipina cf. gephyra (= O. petiola ) USNM 343462 Honduras: Pico Bonito AF199217 Garca Pars & Wake (2000) Oedipina gracilis MVZ 210398 Costa Rica: La Selva AF199219 Garca Pars & Wake (2000) Oedipina grandis MVZ 225904 Costa Rica: Cerro Pando AF199220 Garca Pars & Wake (2000) Oedipina cf. ignea USNM 530586 Honduras: Cerro El Pital AF199231 Garca Pars & Wake (2000) Oedipina kasios MVZ 232825 Honduras: La Muralla FJ196866 McCranie et al. (2008) (=Oedipina sp. inq. 1) JHT 2974 Honduras: above Guaymas + + This study Oedipina parvipes AJC1786 Panam: El Cop FJ784316 FJ766760 Crawford et al. (2010) MVZ 210404 Panam: Nusagandi AF199210 Garca Pars & Wake (2000) MVZ 210405 Panam: Ro Frijoles AF199211 Garca Pars & Wake (2000) Oedipina poelzi MVZ 163703 Costa Rica: Vara Blanca AF199224 Garca Pars & Wake (2000) MVZ 207128 Costa Rica: Monteverde AF199225 Garca Pars & Wake (2000) MVZ 206398 Costa Rica: Braulio Carrillo AF199223 Garca Pars & Wake (2000) Oedipina quadra MVZ 232824 Honduras: Warunta FJ196865 McCranie et al. (2008) Oedipina savagei UCR LG961327 Costa Rica: Cerro Zapote AF199209 Garca Pars & Wake (2000) Oedipina sp. SMF 78738 Nicaragua + Garca Pars & Wake (2000) JHT2974 Hondur as: above Guaymas + + This study

PAGE 233

233 Table 4 1. Continued. Oedipina sp. inq. 2 (= O. nica) N567 Nicaragua: Cerro Kilamb + + This study N569 Nicaragua: Cerro Kilamb + This study N570 Nicaragua: Cerro Kilamb + This study N964 Nicaragua: Cerro Kila mb + + This study N965 Nicaragua: Cerro Kilamb + + This study N1029 Nicaragua: Peas Blancas + This study N1030 Nicaragua: Peas Blancas + This study Oedipina sp. inq. 3 (= O. koehleri) N614 Nicaragua: Cerro Saslaya + + This study JS782 Nica ragua: Cerro Musn + This study Oedipina stenopodia MVZ 163649 Guatemala: San Rafael AF199228 Garca Pars & Wake (2000) Oedipina taylori UGSC 1134 Guatemala: Zacapa HM068304 Sunyer et al. (2010) Oedipina tomasi JHT1553 Honduras: Sierra de Omoa + This study MVZ 258037 Honduras: Sierra de Omoa + Oedipina uniformis MVZ 190853 Costa Rica: Cinega Colorado AF199229 Garca Pars & Wake (2000) MVZ 203751 Costa Rica: Tapant AF199230 Garca Pars & Wake (2000) ANURA Bufonidae ( 14 ) Crepidophryne chompipe UCR 16075 Costa Rica HM563859 Mendelson et al. (In press) Incilius coccifer JHT3301 Honduras: Isla El Tigre + + This study JS1016 Nicaragua: Ometepe + This study JS1058 Nicaragua: Ometepe + This study JS1150 Nicaragua : Atlntico Norte + This study SDSNH AEH 013 Nicaragua: Ometepe AY927857 Mendelson et al. (2005) KU 290030 El Salvador: Morazn AY927856 Mendelson et al. (2005) USNM 547980 Honduras: Rus Rus AY929301 Mendelson et al. (2005) Incilius coniferu s MVUP 1820 Panam: El Cop FJ784379 FJ766768 Crawford et al. (2010) USNM 572086 Panam: El Cop FJ784382 FJ766767 Crawford et al. (2010) USNM 572087 Panam: El Cop FJ784444 FJ766766 Crawford et al. (2010) USNM 572092 Panam: El Cop FJ784586 FJ7667 65 Crawford et al. (2010) toe 140 Panam: El Cop FJ784595 FJ766764 Crawford et al. (2010) toe 141 Ocon Panam: El Cop FJ784597 FJ766732 Crawford et al. (2010) toe 144 Ocon Panam: El Cop FJ784599 FJ766762 Crawford et al. (2010) toe 151 Ocon Pana m: El Cop FJ784601 FJ766761 Crawford et al. (2010) Incilius cf. coniferus N416 Nicaragua: Bosawas + + This study Incilius cycladen UTA JRM 4607 Mexico: Guerrero AY927858 Mendelson et al. (2005) Incilius ibarrai JHT2205 Honduras: Cerro Zarciadero + + This study JHT2604 Honduras: Guajiquiro + + This study JHT2605 Honduras: Guajiquiro + + This study JHT2763 Honduras: Intibuc + + This study JHT2765 Honduras: Intibuc + + This study JHT2874 Honduras: Intibuc + This study JHT2906 Honduras: Intibuc + + This study IRL002 Honduras: Comayagua + + This study

PAGE 234

234 Table 4 1. Continued. IRL004 Honduras: Comayagua + + This study IRL005 Honduras: Comayagua + + This study Incilius karenlipsae UTA A 59522 Panam: El Cop GU552454 Mendelson & Mulc ahy (2010) Incilius leucomyos JHT3034 Honduras: Texguat (Atlntida) + + This study JHT3242 Honduras: Texguat (Atlntida) + This study CAC013 Honduras: Pico Bonito + + This study CAC059 Honduras: Pico Bonito + + This study Incilius luetkenii JS 853 Nicaragua: Las Nubes + + This study Incilius porteri JHT2228 Honduras: Cerro Uyuca + + This study JHT3302 Honduras: Isla El Tigre + + This study CAC044 Honduras: Comayagua + + This study JHT2249 Honduras: Francisco Morazn HM563882 Mendelson e t al. (In press) Incilius valliceps JHT2013 Honduras: Marale + + This study JHT2271 Nicaragua: Selva Negra + + This study JHT2428 Honduras: Texguat (Yoro) + + This study JHT2519 Honduras: Azul Membar + + This study JHT2807 Honduras: Quebrada Gr ande + + This study JHT3175 Honduras: Texiguat (Atlntida) + This study IRL047 Honduras: Azul Membar + + This study LK005 Honduras: Cerro Santa Brbara + + This study N089 Nicaragua: Bosawas + + This study USNM 509524 Honduras: Pico Bonito AY0 08231 Mulcahy & Mendelson (2000) Rhaebo haematiticus MVUP 1842 Panam: El Cop FJ784439 FJ766816 Crawford et al. (2010) USNM 572094 Panam: El Cop FJ784404 FJ766818 Crawford et al. (2010) USNM 572095 Panam: El Cop FJ784426 FJ766817 Crawford et al (2010) USNM 572096 Panam: El Cop FJ784452 FJ766815 Crawford et al. (2010) USNM 572097 Panam: El Cop FJ784546 FJ766814 Crawford et al. (2010) USNM 572098 Panam: El Cop FJ784560 FJ766813 Crawford et al. (2010) Toe 120 Panam: El Cop FJ784593 FJ766812 Crawford et al. (2010) Rhaebo cf. haematiticus N063 Nicaragua: Bosawas + + This study N137 Nicaragua: Bosawas + + This study N594 Nicaragua: Bosawas + + This study Rhinella marina MVUP 1802 Panam: El Cop FJ784357 FJ766819 Crawford et al. (2010) Craugastoridae ( 11 ) Craugastor angelicus MVZ 149762 Costa Rica: Volcn Barba EU186681 Hedges et al. (2008) Craugastor aurilegulus JHT2993 Honduras: Pico Bonito + + This study JHT3015 Honduras: Lancetilla + + This study JHT3016 Hondur as: Lancetilla + + This study JHT3017 Honduras: Lancetilla + + This study JHT3018 Honduras: Lancetilla + + This study JHT3241 Honduras: Texguat (Atlntida) + + This study C007 Honduras: Pico Bonito + + This study C024 Honduras: Pico Bonito + + T his study

PAGE 235

235 Table 4 1. Continued. Craugastor cf. azueroensis KRL0680 Panam: El Cop FJ784332 FJ766637 Crawford et al. (2010 USNM 572219 Panam: El Cop FJ784393 FJ766636 Crawford et al. (2010 USNM 572278 Panam: El Cop FJ784324 FJ766675 Crawford et a l. (2010 USNM 572279 Panam: El Cop FJ784325 FJ766674 Crawford et al. (2010 Craugastor bransfordii USNM 572220 Panam: El Cop FJ784339 FJ766631 Crawford et al. (2010) MVUP 1803 Panam: El Cop FJ784358 FJ766630 Crawford et al. (2010) USNM 572221 P anam: El Cop FJ784376 FJ766629 Crawford et al. (2010) MVUP 1841 Panam: El Cop FJ784427 FJ766628 Crawford et al. (2010) USNM 572222 Panam: El Cop FJ784481 FJ766627 Crawford et al. (2010) USNM 572223 Panam: El Cop FJ784496 FJ766626 Crawford et al. (2010) Craugastor charadra JHT1820 Honduras: El Paraiso Valley + + This study JHT1826 Honduras: El Paraiso Valley + + This study Craugastor fitzingeri N060 Nicaragua: Bosawas + + This study KRL 0693 Panam: El Cop FJ784337 Crawford et al. (20 10) USNM 572256 Panam: El Cop FJ784344 Crawford et al. (2010) MVUP 1798 Panam: El Cop FJ784356 Crawford et al. (2010) Craugastor laevissimus JHT1824 Honduras: El Paraiso Valley + This study JHT2489 Honduras: Azul Membar + + This study JHT2501 Honduras: Azul Membar + + This study JHT2510 Honduras: Azul Membar + + This study JHT2517 Honduras: Azul Membar + + This study JHT2529 Honduras: Comayagua + This study JHT2539 Honduras: Comayagua + This study JHT2552 Honduras: Coma yagua + This study JHT2559 Honduras: Comayagua + This study JHT2779 Honduras: Pico Pijol + This study JHT2978 Honduras: Azul Membar + + This study JHT2979 Honduras: Azul Membar + + This study JHT3000 Honduras: Cerro Santa Brbara + + This study JHT3004 Honduras: Los Naranjos + + This study IRL023 Honduras: Azul Membar + + This study N556 Nicaragua: Cerro Kilamb + This study N557 Nicaragua: Cerro Kilamb + This study N639 Nicaragua: Cerro Saslaya + This study N950 Nicara gua: Cerro Kilamb + This study N961 Nicaragua: Cerro Kilamb + This study Craugastor punctariolus MVUP 1784 Panam: El Cop FJ784333 FJ766673 Crawford et al. (2010) USNM 572281 Panam: El Cop FJ784411 FJ766672 Crawford et al. (2010) USNM 57228 2 Panam: El Cop FJ784417 FJ766671 Crawford et al. (2010) USNM 572283 Panam: El Cop FJ784418 FJ766670 Crawford et al. (2010) Craugastor sandersoni UTA A49803 Guatemala: Sierra de Santa Cruz EF493712 Heinicke et al. (2007) Craugastor tabasarae KRL0 706 Panam: El Cop FJ784342 Crawford et al. (2010) KRL1373 Panam: El Cop FJ784512 Crawford et al. (2010)

PAGE 236

236 Table 4 1. Continued. KRL1387 Panam: El Cop FJ784515 Crawford et al. (2010) Craugastor underwoodi UCR 16315 EF562362 Streicher et a l. (2009) USNM 561403 EF562361 Streicher et al. (2009) Eleutherodactylidae (3) Diasporus aff. diastema MVUP 1783 Panam: El Cop FJ784338 Crawford et al. (2010) USNM 572442 Panam: El Cop FJ784425 Crawford et al. (2010) MVUP 1830 Panam : El Cop FJ784395 Crawford et al. (2010) Diasporus cf. diastema N546 Nicaragua: Cerro Kilamb + + This study N928 Nicaragua: Cerro Kilamb + + This study N934 Nicaragua: Cerro Kilamb + + This study Hylidae (15) Duellmanohyla soralia JHT15 85 Honduras: Sierra de Omoa + + This study UTA A 50812 Guatemala: Sierra de Caral AY843584 Faivovich et al. (2005) Exerodonta catracha JHT2209 Honduras: Cerro Zarciadero + + This study JHT2211 Honduras: Cerro Zarciadero + + This study JHT2315 Hond uras: Intibuc + This study JHT2316 Honduras: Intibuc + + This study JHT2647 Honduras: Guajiquiro + + This study JHT2648 Honduras: Guajiquiro + + This study JHT2649 Honduras: Guajiquiro + + This study JHT2650 Honduras: Guajiquiro + + This stud y JHT2651 Honduras: Guajiquiro + + This study JHT2652 Honduras: Guajiquiro + This study JHT2653 Honduras: Guajiquiro + + This study JHT2654 Honduras: Guajiquiro This study JHT2655 Honduras: Guajiquiro + + This study JHT2656 Honduras: Guaj iquiro + + This study JHT2657 Honduras: Guajiquiro + This study JHT2658 Honduras: Guajiquiro + + This study JHT2659 Honduras: Guajiquiro + + This study JHT2660 Honduras: Guajiquiro + + This study JHT2661 Honduras: Guajiquiro + + This study JH T2861 Honduras: Guajiquiro + + This study JHT2862 Honduras: Guajiquiro + + This study JHT2863 Honduras: Guajiquiro This study JHT2864 Honduras: Guajiquiro + + This study JHT2870 Honduras: Guajiquiro + + This study JHT2871 Honduras: Guajiquiro + + This study JHT2872 Honduras: Guajiquiro + This study JHT2896 Honduras: Intibuc + + This study JHT2897 Honduras: Intibuc + + This study JHT2898 Honduras: Intibuc + + This study JHT2899 Honduras: Intibuc + + This study

PAGE 237

237 Table 4 1. Contin ued. JHT2907 Honduras: Intibuc + This study Plectrohyla chrysopleura JHT3077 Honduras: Texguat (Atlntida) + + This study JHT3081 Honduras: Texguat (Atlntida) + + This study JHT3142 Honduras: Texguat (Atlntida) + + This study JHT3166 Hond uras: Texguat (Atlntida) + + This study JHT3167 Honduras: Texguat (Atlntida) + + This study JHT3169 Honduras: Texguat (Atlntida) + + This study Plectrohyla dasypus JHT1624 Honduras: Cusuco + + This study JHT2988 Honduras: Cusuco + + This stud y Plectrohyla exquisita JHT1600 Honduras: Cusuco + + This study JHT1601 Honduras: Cusuco + + This study JHT1623 Honduras: Cusuco + + This study Plectrohyla guatemalensis JHT2058 Honduras: Cataguana + + This study JHT2059 Honduras: Cataguana + + T his study JHT2060 Honduras: Cataguana + + This study Plectrohyla psiloderma JHT2746 Honduras: Cerro Celaque + + This study Ptychohyla euthysanota JS1244 Guatemala: Los Tarrales + This study UTA A 54786 Mxico: Chiapas AY843744 Faivovich et al. ( 2005) Ptychohyla hypomykter JHT1622 Honduras: Cusuco + + This study JHT2272 Nicaragua: Selva Negra + This study JHT2273 Nicaragua: Selva Negra + + This study JHT2274 Nicaragua: Selva Negra + + This study JHT2276 Nicaragua: Selva Negra + + This study JHT2300 Honduras: Comayagua + + This study JHT2494 Honduras: Azul Membar + + This study JHT2530 Honduras: Comayagua + This study JHT2540 Honduras: Comayagua + + This study JHT2541 Honduras: Comayagua + + This study JHT2548 Honduras: Co mayagua + + This study JHT2855 Honduras: Quebrada Grande + + This study JHT2916 Honduras: Cusuco + + This study JHT2930 Honduras: Pico Pijol + This study JHT2931 Honduras: Pico Pijol + + This study JHT2933 Honduras: Pico Pijol + + This study JHT2934 Honduras: Pico Pijol + + This study JHT2935 Honduras: Pico Pijol + + This study JHT2936 Honduras: Pico Pijol + + This study JHT2945 Honduras: Pico Pijol + + This study JHT2946 Honduras: Pico Pijol + + This study JHT2949 Honduras: Pico Pij ol + + This study JHT2950 Honduras: Pico Pijol + + This study JHT2951 Honduras: Pico Pijol + + This study JHT2954 Honduras: Pico Pijol + This study

PAGE 238

238 Table 4 1. Continued. JHT2998 Honduras: Cerro Santa Brbara + + This study JHT2999 Honduras: Ce rro Santa Brbara + + This study IRL067 Honduras: Azul Membar + + This study IRL091 Honduras: Azul Membar + + This study IRL092 Honduras: Azul Membar + + This study IRL093 Honduras: Azul Membar + + This study IRL094 Honduras: Azul Membar + + This study IRL095 Honduras: Azul Membar + + This study N293 Nicaragua: Cerro Saslaya + + This study N528 Nicaragua: Cerro Kilamb + + This study N547 Nicaragua: Cerro Kilamb + + This study N993 Nicaragua: Cerro Peas Blancas + + This study N 1000 Nicaragua: Cerro Peas Blancas + + This study N1004 Nicaragua: Cerro Peas Blancas + + This study N1028 Nicaragua: Cerro Peas Blancas + + This study Ptychohyla cf. hypomykter ENS 8486 Guatemala: Izabal AY843745 Faivovich et al. (2005) Ptychohyl a salvadorensis JHT2262 Honduras: Cerro Uyuca + + This study Ptychohyla spinipollex USNM 514381 Honduras: Pico Bonito AY843748 Faivovich et al. (2005) Ptychohyla cf. spinipollex JHT3041 Honduras: Texguat (Atlntida) + + This study JHT3042 Honduras : Texguat (Atlntida) + + This study JHT3055 Honduras: Texguat (Atlntida) + + This study JHT3056 Honduras: Texguat (Atlntida) + This study JHT3057 Honduras: Texguat (Atlntida) + + This study JHT3058 Honduras: Texguat (Atlntida) + + This study JHT3059 Honduras: Texguat (Atlntida) + + This study JHT3070 Honduras: Texguat (Atlntida) + + This study JHT3071 Honduras: Texguat (Atlntida) + + This study JHT3072 Honduras: Texguat (Atlntida) + + This study JHT3073 Honduras: Texgu at (Atlntida) + + This study JHT3114 Honduras: Texguat (Atlntida) + This study JHT3115 Honduras: Texguat (Atlntida) + + This study JHT3116 Honduras: Texguat (Atlntida) + + This study JHT3154 Honduras: Texguat (Atlntida) + + This study JHT3170 Honduras: Texguat (Atlntida) + + This study JHT3233 Honduras: Texguat (Atlntida) + + This study JHT3234 Honduras: Texguat (Atlntida) + + This study JHT3235 Honduras: Texguat (Atlntida) + + This study JHT3236 Honduras: Texguat (Atl ntida) + + This study Smilisca cf. baudinii JHT2277 Nicaragua: Selva Negra + + This study JHT2592 Honduras: Los Naranjos + + This study JHT2593 Honduras: Los Naranjos + This study JHT2938 Honduras: Pico Pijol + + This study N781 Nicaragua: Bosaw as + + This study

PAGE 239

239 Table 4 1. Continued. N1019 Nicaragua: Peas Blancas + + This study Smilisca phaeota USNM 572702 Panam: El Cop FJ784413 FJ766835 Crawford et al. (2010) USNM 572703 Panam: El Cop FJ784433 FJ766834 Crawford et al. (2010) Smilisca sila USNM 572707 Panam: El Cop FJ784578 FJ766837 Crawford et al. (2010) USNM 572708 Panam: El Cop FJ784579 FJ766836 Crawford et al. (2010) Smilisca sordida N030 Nicaragua: Bosawas + + This study N031 Nicaragua: Bosawas + + This study N114 Nica ragua: Bosawas + + This study N115 Nicaragua: Bosawas + This study N819 Nicaragua: Bosawas + + This study Tlalocohyla loquax JHT2247 Honduras: Cerro Uyuca + + This study JHT2268 Nicaragua: Selva Negra + + This study JHT2269 Nicaragua: Selva Neg ra + This study Leptodactylidae (1) Leptodactylus fragilis JHT3238 Honduras: Texguat (Atlntida) + + This study N462 Nicaragua: Bosawas + + This study N653 Nicaragua: Bosawas + + This study N841 Nicaragua: Bosawas + + This study Leptodacty lus cf. fragilis USNM 572722 Panam: El Cop FJ784331 FJ766745 Crawford et al. (2010) USNM 572725 Panam: El Cop FJ784416 FJ766744 Crawford et al. (2010) MVUP 1836 Panam: El Cop FJ784437 FJ766743 Crawford et al. (2010) USNM 572727 Panam: El Cop FJ784453 FJ766742 Crawford et al. (2010) Leptodactylus savagei MVUP 1828 Panam: El Cop FJ784394 FJ766748 Crawford et al. (2010) Leptodactylus poecilochilus KRL 0118 Panam: El Cop FJ784321 Crawford et al. (2010) Microhylidae (1) Hypopachus ba rberi JHT2630 Honduras: Guajiquiro + + This study JHT2634 Honduras: Guajiquiro + + This study Ranidae (4) Lithobates areolata KU 204370 USA: Kansas AY779229 Hillis & Wilcox (2005) Lithobates berlandieri JSF1136 USA: Texas AY779235 Hillis & Wi lcox (2005) Lithobates brownorum JHT2329 Honduras: Los Naranjos + This study JHT2349 Honduras: Los Naranjos + This study JHT2350 Honduras: Los Naranjos + This study JHT2351 Honduras: Los Naranjos + This study JHT2466 Honduras: Los Naranjo s + This study JHT2525 Honduras: Comayagua + This study JHT2560 Honduras: Comayagua + This study JHT2561 Honduras: Comayagua + This study JHT2562 Honduras: Comayagua + This study JHT2563 Honduras: Comayagua + This study JHT2564 Hond uras: Comayagua + This study JHT2565 Honduras: Comayagua + This study JHT2566 Honduras: Comayagua + This study

PAGE 240

240 Table 4 1. Continued. JHT2567 Honduras: Comayagua + This study JHT2569 Honduras: Comayagua + This study JHT2760 Honduras: In tibuc + This study JHT2761 Honduras: Intibuc + This study JHT2814 Honduras: Quebrada Grande + This study JHT2815 Honduras: Quebrada Grande + This study JHT2818 Honduras: Quebrada Grande + This study JHT2955 Honduras: above Guaymas + This study JHT2956 Honduras: above Guaymas + This study JHT2957 Honduras: above Guaymas + This study JHT2958 Honduras: above Guaymas + This study IRL051 Honduras: Azul Membar + This study LK002 Honduras: Cerro Santa Brbara + This stu dy Lithobates brownorum X forreri JHT2138 Honduras: Cerro Uyuca + + This study JHT2139 Honduras: Cerro Uyuca + + This study JHT2140 Honduras: Cerro Uyuca + This study JHT2141 Honduras: Cerro Uyuca + + This study JHT2142 Honduras: Cerro Uyuca + + This study JHT2143 Honduras: Cerro Uyuca + + This study JHT2144 Honduras: Cerro Uyuca + + This study JHT2145 Honduras: Cerro Uyuca + + This study JHT2153 Honduras: Cerro Uyuca + + This study JHT2308 Honduras: Intibuc + + This study JHT2310 Ho nduras: Intibuc + + This study JHT2314 Honduras: Intibuc + + This study JHT2327 Honduras: Intibuc + + This study JHT2328 Honduras: Intibuc + + This study JHT2772 Honduras: Intibuc + + This study JHT2883 Honduras: Intibuc + + This study Lit hobates capito TNHC 60195 USA: Florida AY779231 Hillis & Wilcox (2005) Lithobates catesbeiana DMH 84 R2 USA: Kansas AY779206 Hillis & Wilcox (2005) None No data GBX12841 Hillis & Wilcox (2005) Lithobates chiricahuensis KU 194442 Mxico : Durango A Y779225 Hillis & Wilcox (2005) KU 194419 USA: Arizona AY779226 Hillis & Wilcox (2005) Lithobates clamitans JSF1118 USA Missouri AY779204 Hillis & Wilcox (2005) Lithobates forreri KU 194581 Mxico : Sinaloa AY779233 Hillis & Wilcox (2005) Lithob ates grylio MVZ 175945 USA: Florida AY779201 Hillis & Wilcox (2005) Lithobates heckscheri MVZ 164908 USA: Florida AY779205 Hillis & Wilcox (2005) KU 195138 Mxico : Chiapas AY779242 Hillis & Wilcox (2005) UTA A 17185 Guate mala: Solol AY779243 Hillis & Wilcox (2005) Lithobates maculatus KU 195258 Mxico : Oaxaca AY779207 Hillis & Wilcox (2005) USNM 559483 Honduras: Pico Bonito DQ283303 Frost et al. (2006) Lithobates maculatus None Mxico B:AAE6665 B OLD Database

PAGE 241

241 Table 4 1. Continued. Lithobates maculatus complex JHT2007 Honduras: Marale + + This study JHT2028 Honduras: Los Planes + + This study JHT2029 Honduras: Los Planes + + This study JHT2117 Honduras: Cataguana + + This study JHT2136 Hon duras: Cerro Uyuca + + This study JHT2137 Honduras: Cerro Uyuca + + This study JHT2204 Honduras: Cerro Zarciadero + + This study JHT2244 Honduras: Cerro Uyuca + + This study JHT2439 Honduras: Texguat (Yoro) + + This study JHT2492 Honduras: Azul Membar + + This study JHT2528 Honduras: Comayagua + + This study JHT2571 Honduras: Comayagua + + This study JHT2594 Honduras: Los Naranjos + + This study JHT2617 Honduras: Guajiquiro + + This study JHT2851 Honduras: Quebrada Grande + + This stud y JHT2852 Honduras: Quebrada Grande + + This study JHT2853 Honduras: Quebrada Grande + + This study JHT2939 Honduras: Pico Pijol + + This study JHT2940 Honduras: Pico Pijol + + This study IRL015 Honduras: Azul Membar + + This study IRL016 Hond uras: Azul Membar + + This study IRL017 Honduras: Azul Membar + + This study IRL018 Honduras: Azul Membar + + This study IRL019 Honduras: Azul Membar + + This study IRL020 Honduras: Azul Membar + This study IRL052 Honduras: Azul Membar + + This study IRL053 Honduras: Azul Membar + + This study IRL057 Honduras: Azul Membar + + This study IRL058 Honduras: Azul Membar + + This study IRL087 Honduras: Azul Membar + This study N285 Nicaragua: Cerro Saslaya + + This study N371 N icaragua: Bosawas + + This study N610 Nicaragua: Cerro Saslaya + + This study N638 Nicaragua: Cerro Saslaya + This study N649 Nicaragua: Cerro Saslaya + + This study N920 Nicaragua: Cerro Kilamb + + This study N1009 Nicaragua: Peas Blancas + + This study N1021 Nicaragua: Peas Blancas + + This study Lithobates magnaocularis KU 194592 Mxico : Sonora AY779239 Hillis & Wilcox (2005) Lithobates montezumae KU 195251 Mxico : Morelos AY779223 Hillis & Wilcox (2005) Lithobates neovolcanica KU 194536 Mxico : Michoacan AY779236 Hillis & Wilcox (2005) Lithobates okaloosae None (toe clip) USA: Florida AY779203 Hillis & Wilcox (2005) Lithobates omiltemana KU 195179 Mxico : Guerrero AY779238 Hillis & Wilcox (2005)

PAGE 242

242 Table 4 1. Continued. Lit hobates palmipes KU 202896 Ecuador: Napo AY779211 Hillis & Wilcox (2005) Lithobates palustris KU 204425 USA: Indiana AY779228 Hillis & Wilcox (2005) Lithobates pipiens JSF1119 USA: Ohio AY779221 Hillis & Wilcox (2005) None No data GBX12841 Hill is & Wilcox (2005) Lithobates septentrionalis TNHC (no #) Canada: Ontario AY779200 Hillis & Wilcox (2005) Lithobates sevosa TNHC 60194 USA: Mississippi AY779230 Hillis & Wilcox (2005) Lithobates spectabilis KU 195186 Mxico : Hidalgo AY779232 Hilli s & Wilcox (2005) Lithobates sphenocephalus JSF845 USA: Kansas AY779251 Hillis & Wilcox (2005) USC7448 USA: Florida AY779252 Hillis & Wilcox (2005) Lithobates subaquavocalis TNHC (no #) USA: Arizona AY779227 Hillis & Wilcox (2005) Lithobates syl vatica MVZ 137426 USA: New York AY779198 Hillis & Wilcox (2005) Lithobates taylori JHT2263 Nicaragua: Selva Negra + + This study JHT2264 Nicaragua: Selva Negra + + This study JHT2265 Nicaragua: Selva Negra + This study N918 Nicaragua: Cerro Kila mb + + This study N919 Nicaragua: Cerro Kilamb + + This study N997 Nicaragua: Peas Blancas + + This study N1020 Nicaragua: Peas Blancas + + This study TCWC 55963 Nicaragua: 2.5 mi NW Rama AY779244 Hillis & Wilcox (2005) Lithobates tlaloci KU 194436 Mxico : Districto Federal AY779234 Hillis & Wilcox (2005) Lithobates vibicarius MVZ 149033 Costa Rica: San Jos AY779208 Hillis & Wilcox (2005) Lithobates warszewitschii USNM 572770 Panam: El Cop FJ784454 FJ766752 Crawford et al. (2010) U SNM 572779 Panam: El Cop FJ784552 FJ766751 Crawford et al. (2010) USNM 572780 Panam: El Cop FJ784558 FJ766750 Crawford et al. (2010) Lithobates JSF1127 Panam: Cocl AY779209 Hillis & Wilcox (2005) Lithobates aff. warszewitschii USNM 572787 Panam: El Cop FJ784384 FJ766749 Crawford et al. (2010) Lithobates cf. warszewitschii N334 Nicaragua: Bosawas + This study N599 Nicaragua: Bosawas + + This study Lithobates spp. Anura None Mxico B:AAB6680 BOLD Database None Mxico B:AAB6680 BOLD Database None Mxico B:AAB6680 BOLD Database None Mxico B:AAB6680 BOLD Database None Mxico B:AAB6680 BOLD Database None Mxico B:AAB6680 BOLD Database None Mxico B:AAB6680 BOLD Database None Mxico B:AAB6 680 BOLD Database None Mxico B:AAB6679 BOLD Database None Mxico B:AAB6679 BOLD Database None Mxico B:AAB6678 BOLD Database None Mxico B:AAB6681 BOLD Database None Mxico B:AAB6682 BOLD Database None Mxico B:AAB6682 BOLD Databas e None Mxico B:AAB6682 BOLD Database

PAGE 243

243 Table 4 1. Continued. None Mxico B:AAE6666 BOLD Database None Mxico B:AAK8798 BOLD Database None Mxico B:AAC6120 BOLD Database None Mxico B:AAC6120 BOLD Database Lithobates sp. 1 QCAZ 13219 Ec uador: Esmeraldas AY779213 Hillis & Wilcox (2005) Lithobates sp. 2 KU 2044420 Mxico : San Luis Potos AY779224 Hillis & Wilcox (2005) Lithobates sp. 3 KU 194559 Mxico : Michoacan AY779250 Hillis & Wilcox (2005) Lithobates sp. 4 AMNH 124167 Panama: Chiriqu AY779245 Hillis & Wilcox (2005) Lithobates sp. 5 LACM 146764 Costa Rica: Heredia AY779246 Hillis & Wilcox (2005) Lithobates sp. 6 LACM 146810 Costa Rica: Puntarenas AY779247 Hillis & Wilcox (2005) Lithobates sp. 7 KU 194492 Mxico : Jalis co AY779241 Hillis & Wilcox (2005) Lithobates sp. 8 KU 195346 Mxico : Puebla AY779248 Hillis & Wilcox (2005) Strabomantidae (2) Pristimantis caryophyllaceus MVUP 1925 Panam: El Cop FJ784473 FJ766776 Crawford et al. (2010) USNM 572343 Panam : El Cop FJ784375 FJ766772 Crawford et al. (2010) Pristimantis cerasinus N121 Nicaragua: Bosawas + This study Pristimantis ridens A USNM 572416 Panam: El Cop FJ784388 FJ766808 Crawford et al. (2010) USNM 572417 Panam: El Cop FJ784389 FJ766807 C rawford et al. (2010) Pristimantis ridens B MVUP 1829 Panam: El Cop FJ784398 FJ766806 Crawford et al. (2010) USNM 572415 Panam: El Cop FJ784399 FJ766805 Crawford et al. (2010) Pristimantis cf. ridens N090 Nicaragua: Bosawas + + This study N143 N icaragua: Bosawas + + This study N144 Nicaragua: Bosawas + + This study N201 Nicaragua: Bosawas + + This study N263 Nicaragua: Cerro Saslaya + + This study N541 Nicaragua: Cerro Kilamb + This study N998 Nicaragua: Cerro Peas Blancas + + This study N1008 Nicaragua: Cerro Peas Blancas + + This study

PAGE 244

244 Table 4 2. Average nucleotide compositions of various taxonomic groups Taxon # of sequences Avg. Length A G C T COI Amphibia 39 0 636.3 23.9% 18.0% 27.6% 30.5% Anura 256 629.3 23.3% 1 8.1% 28.0% 30.6% Caudata 14 4 649.5 25.1% 17.9% 26.7% 30.2% 16S Amphibia 436 537.0 31.0% 20.1% 23.2% 25.7% Anura 277 553.6 30.0% 20.2% 23.8% 26.1% Caudata 269 512.5 33.2% 19.9% 22.3% 24.7%

PAGE 245

245 Table 4 3 Potential candidate species identified thr ough this study; UCS = Unconfirmed Candidate Species, CCS = Confirmed Candidate Species; DCL = Deep Conspecific Lineage ; for IUCN Red List categories, CR = Critically Enda ngered EN = Endangered, and VU = Vulnerable. Interspecific divergences are measured at the genus level. A priori taxonomic assignment Current status Potential IUCN Red List Intraspecific Divergence Interspecific Divergence Notes Plethodontidae (16) Bolitoglossa sp. inq. 1 Bolitoglossa cataguana CR 16S =0.0% COI =0.7% 16S =2.3 8 .7%, COI =11.6 20.9% (0.0% from B. decora for 16S) Described by Townsend et al. (2009a) Bolitoglossa sp. inq. 2 Bolitoglossa oresbia CR 16S =0.2% COI =0.1% 16S =3.0 9.7% COI =10.0 22.1% Conspecific with B. oresbia and to represent a range extension (Tow nsend et al. 2011b) Bolitoglossa celaque Bolitoglossa sp. 1 DCL EN 16S =0.8% COI =0.3% 16S =1.8 10.0% COI =4.5 21.7% (within B. celaque complex: 16S =1.0 1.5%, COI =1.7 3.0%) Population from Sierra de Puca Opalaca; under study by author Bolitoglossa ce laque Bolitoglossa sp. 2 DCL EN 16S =0.3% COI =0.2% 16S =2.0 10.3% COI =5.2 21.5% (within B. celaque complex: 16S =1.3 1.5%, COI =1.7 3.6%) Population from Montaas de la Sierra; under study by author Bolitoglossa conanti Bolitoglossa sp. 3 DCL EN 16S = 0.0% COI =0.0% 16S =3.1 9.3% COI =9.6 21.4% (within B. conanti complex: 16S =0.6 1.6%, COI =2.7 3.1%) Population from Sierra de Omoa; under study by author Bolitoglossa conanti Bolitoglossa sp. 4 DCL EN 16S =0.0% COI =0.0% 16S =3.5 9.4% COI =9.6 20.4% (within B. conanti complex: 16S =0.8 1.6%, COI =2.5 3.7%) Population from Sierra del Merendn; under study by author Bolitoglossa porrasorum Bolitoglossa sp. 5 C CS CR 16S =0.2% COI =0.7% 16S =2.0 8.6% COI =9.2 23.3% Population from Texguat; under study by author Bolitoglossa porrasorum Bolitoglossa sp. 6 C CS CR 16S =N/A COI = N/A 16S =2.0 8.1% COI = N/A Population from Pico Bonito; under study by author Bolitoglossa rufescens Bolitoglossa sp. 7 CCS 16S =0.2% 16S =5.5 10.3% S yntopic in Sierra de Omoa with B.

PAGE 246

246 COI =0.2% COI =17.6 22.7% nympha ; under study by author Nototriton sp. inq. 1 Nototriton tomamorum CR 16S = N/A COI = N/A 16S =3.3 5.4% COI =11.3 14.1% Described by Townsend et al. (2010a) Nototriton sp. inq. 2 Nototriton picucha CR 16S =0.0% CO I =0.2% 16S =1.2 5.2% COI = 2.9 12.3% Described in Chapter 5 and by Townsend et al. (2011a) Nototriton sp. inq. 3 Nototriton sp. 1 CCS CR 16S =0.0% COI =0.6% 16S =1.7 5.8% COI =2.9 12.5% Population from Sierra de Botaderos Nototriton sp. inq. 4 Nototrit on limnospectator N/A 16S =0.3% COI =0.2% 16S =1.2 5.2% COI =7.8 12.6% Conspecific with N. limnospectator and to represent a range extension (Townsend et al. 2011b) Nototriton sp. inq. 5 Nototriton lignicola N/A 16S =0.2% COI = N/A 16S =2.4 5.9% COI =10.0 13.9% Conspecific with N. lignicola and to represent a range extension (Townsend et al. 2011b) Nototriton barbouri Nototriton sp. 2 CCS CR 16S =N/A COI = N/A 16S =1.2 6.0% COI = N/A Population from Pico Bonito; described in Chapter 5 (as N. sp. A) Notot riton barbouri Nototriton sp. 3 CCS CR 16S =N/A COI = N/A 16S =1.7 5.8% COI =5.9% 13.6% Population from Texguat; described in Chapter 5 (as N. sp. B) Oedipina sp. inq. 1 Oedipina kasios N/A 16S =0.1% COI = N/A 16S =2.6 11.3% COI =6.6 15.7% Conspecific wi th O. kasios and to represent a range extension (Townsend et al. 2011b) Oedipina sp. inq. 2 Oedipina nica EN 16S =0.1% COI =0.0% 16S =2.6 10.2% COI =6.6 16.4% Described by Sunyer et al. (2010) Oedipina sp. inq. 3 Oedipina koehleri EN 16S =0.0% COI = N/A% 16S =3.1 8.4% COI =14.4 16.2% Described by Sunyer et al. (2011) Oedipina gephyra Oedipina petiola CR 16S = N/A COI = N/A 16S =3.7 10.5% COI = N/A Described by McCranie & Townsend (2011) Bufonidae (2) Incilius coniferus Incilius sp. DCL DD 16S = N/ A COI = N/A 16S =3.6 9.3% COI =11.2 17.5% (from Panamanian I. coniferus : 16S =0.7%, COI =3.5%) Population from northern Nicaragua; under study by Sunyer, Townsend, and Travers Rhaebo haematiticus Rhaebo sp. UCS DD 16S =0.0% COI = 0.2% 16S =10.3 12.4% COI =16.1 19.5% (from Panamanian R. haematiticus : 16S =2.2%, COI =5.4%) Population from northern Nicaragua; under study by Sunyer, Townsend, and Travers Craugastoridae (3) Craugastor aurilegulus Craugastor sp. 1 DCL EN 16S =0.1% 16S =4.9 21 .0% Population from vicinity of Texguat;

PAGE 247

247 COI =0.6% COI =11.2 25.6% (from Pico Bonito C. aurilegulus : 16S =1.1%, COI =3.9%) may represent DCL Craugastor sp. inq. 1 Craugastor sp. 2 UCS DD 16S = N/A COI = N/A 16S =7.5 21.1% COI =16.8 26.6% Population from northern Nicaragua; field identification Diasporus diastema Craugastor sp. inq. 2 Craugastor sp. 3 UCS DD 16S =0.0% COI =0.0% 16S =7.5 21.3% COI =11.4 26.7% Population from northern Nicaragua Eleutherodactylidae (1) Diasporus diastema Diasporus sp. UCS DD 16S = N/A COI = N/A 16S =7.1 21.6% COI =15.7 27.2% Population from northern Nicaragua Hylidae (5) Plectrohyla guatemalensis Plectrohyla sp. UCS CR 16S =0.0% COI =0.1% 16S =2.6 13.8% COI =8.8 23.8% Population from Cordillera de La Flor La Muralla P tychohyla hypomykter Ptychohyla sp. 1 UCS DD 16S =N/A COI = N/A 16S =5.5 14.7% COI = N/A Population from eastern Guatemala Ptychohyla spinipollex Ptychohyla sp. 2 UCS CR 16S =0.2% COI =0.5% 16S =4.8 13.4% COI =15.1 22.6% Population from vicinity of Tex guat; may represent DCL Smilisca baudinii Smilisca sp. 1 UCS LC 16S =0.0% COI =0.0% 16S =1.6 15.0 % COI =6.6 22.7% Populations from northern Nicaragua Smilisca baudinii Smilisca sp. 2 UCS LC 16S =0.0% COI =0.1% 16S =1.6 14.4% COI =6.6 22.7% Populations f rom Honduras Leptodactylidae (1) Leptodactylus fragilis Leptodactylus sp. UCS LC 16S =0.0% COI =0.3% 16S =3.6 21.4% COI =15.7 26.2% Populations from Panama (Crawford et al. 2010); Chorts Block samples considered conspecific with nominal form Ranid ae (7) Lithobates brownorum X forreri Lithobates sp. 1 CCS EN 16S =0.8% COI =0.5% 16S =2.2 16.5% COI =9.7 22.4% Populations from Southern Cordillera of Serrana ; being described by author Lithobates maculatus Lithobates sp. 2 UCS EN 16S =0.0% COI = 0.1% 16S =1.6 12.2% COI =2.4 20.5% Populations from highlands in north central Nicaragua; under study by author Lithobates maculatus Lithobates sp. 3 UCS EN 16S =0.4% COI =0.5% 16S =1.6 12.8% COI =2.4 21.1% Populations from low to moderate elevations in northeastern Nicaragua; under study by author Lithobates maculatus Lithobates sp. 4 UCS EN 16S =0.0% COI =0.0% 16S =1.8 12.8% COI =5.7 20.7% Populations from Texguat and Sierra de Sulaco; under study by author

PAGE 248

248 Lithobates maculatus Lithobates sp. 5 UCS E N 16S =0.6% COI =1.2% 16S =2.9 12.3% COI =7.2 21.5% Populations from throughout Honduran Serrana ; under study by author Lithobates maculatus Lithobates sp. 6 UCS EN 16S =N/A COI =N/A 16S =2.6 12.6% COI = N/A Population from vicinity of Pico Bonito; under study by author Lithobates warszewitschii Lithobates sp. 7 UCS DD 16S =0.6% COI = N/A 16S =8.2 17.2 COI =10.1 22.8% Populations from northern Nicaragua Strabomantidae (1) Pristimantis ridens Pristimantis sp. UCS DD 16S =0.3% COI =0.1% 16S =4.8 22.2% COI =9.1 27.0% Populations from northern Nicaragua

PAGE 249

249 Table 4 4 Results of BLASTN searches of the NCBI database for 16S consensus sequences representing 10 populations of taxonomically unassigned salamanders. Sample Matching sequence (GenBank acces sion #) Max ident Max score Query coverage Bolitoglossa sp. inq. 1 1. Bolitoglossa decora (GU725449) 99% 904 100% Cataguana 2. Bolitoglossa morio (GU725452) 97% 830 100% 3. Bolitoglossa porrasorum (AF526151) 96% 824 100% 4. Bolitoglossa morio (AF2184 95) 96% 819 100% 5. Bolitoglossa flavimembris (GU725449) 96% 808 100% Bolitoglossa sp. inq. 2 1. Bolitoglossa flavimembris (GU725449) 96% 843 100% IRL 081 2. Bolitoglossa diaphora (GU725447) 95% 833 100% 3. Bolitoglossa morio (AF526144) 95% 821 100% 4. Bolitoglossa morio (GU725452) 95% 815 100% 5. Bolitoglossa dunni (GU725446) 95% 811 100% Nototriton sp. inq. 1 1. Nototriton lignicola (AF199204) 95% 780 100% Texiguat 2. Nototriton barbouri (AF199201) 94% 760 100% 3. Nototriton richardi (AF1992 06) 94% 758 100% 4. Nototriton abscondens (AF199199) 94% 758 100% 5. Nototriton brodiei (AF199202) 94% 754 100% Nototriton sp. inq. 2 1. Nototriton lignicola (AF199204) 97% 837 100% Agalta 2. Nototriton brodiei (AF199202) 96% 804 100% 3. Nototriton barbouri (AF199201) 96% 804 100% 4. Nototriton richardi (AF199206) 95% 785 100% 5. Nototriton saslaya (GU981761) 95% 776 100% Nototriton sp. inq. 3 1. Nototriton lignicola (AF199204) 96% 813 99% Botaderos 2. Nototriton barbouri (AF199201) 96% 802 99 % 3. Nototriton richardi (AF199206) 95% 784 99% 4. Nototriton brodiei (AF199202) 95% 780 99% 5. Nototriton saslaya (GU981761) 94% 758 99% Nototriton sp. inq. 4* 1. Nototriton lignicola (AF199204) 99% 898 100% Cataguana 2. Nototriton brodiei (AF1992 02) 96% 798 100% 3. Nototriton barbouri (AF199201) 96% 798 100% 4. Nototriton richardi (AF199206) 95% 780 100% 5. Nototriton saslaya (GU981761) 94% 743 100% Nototriton sp. inq. 5 1. Nototriton lignicola (AF199204) 96% 821 100% Membar 2. Nototriton brodiei (AF199202) 96% 804 100% 3. Nototriton barbouri (AF199201) 96% 804 100% 4. Nototriton richardi (AF199206) 96% 797 100% 5. Nototriton saslaya (GU981761) 94% 765 100% Oedipina sp. inq. 1* 1. Oedipina kasios (FJ196866) 99% 893 100% Mta de Yoro 2. Oedipina kasios (FJ196867) 99% 893 100% 3. Oedipina quadra (FJ196865) 95% 769 99% 4. Oedipina uniformis (AF199230) 91% 682 100% 5. Oedipina grandis (FJ196864) 91% 676 100% Oedipina sp. inq. 2 1. Oedipina kasios (FJ196866) 97% 837 99% nica 2. Oe dipina kasios (FJ196867) 97% 835 99% 3. Oedipina quadra (FJ196865) 94% 765 100% 4. Oedipina poelzi (AF199223) 92% 691 99% 5. Oedipina grandis (FJ196864) 91% 680 99% Oedipina sp. inq. 3 1. Oedipina cyclocauda ( AF199214 ) 96% 811 100% 2. Oedipina pse udouniformis ( AF199227 ) 96% 808 100% 3. Oedipina cyclocauda ( AF199215 ) 96% 806 100% 4. Oedipina uniformis (AF199230 ) 95% 782 100% 5. Oedipina uniformis ( AF199229 ) 95% 776 100%

PAGE 250

250 CHAPTER 5 CRYPTIC DIVERSITY AND REVISIONARY SYSTEMATICS OF CHORTS HIGH LAND MOSS SALAMANDERS (CAUDATA: PLETHODONTIDAE) Inconspicuous form, obscure or secretive behavior, and challenging habitat associations are princip al reasons why cryptozoic 1 diversity historically has been overlooked and understudied (Pfenninger & Schwenk 2007). Molecular phylogenetic analyses, facilitated thro ugh techological advances like PCR ( Mullis 1990 ), Sanger sequencing ( Sanger et al. 1977 ) and next generation DNA sequencing ( Schuster 2008 ), have led to the discovery of numerous genetically divergen t but morphologically cryptic lineages (Bickford et al. 2007). Many tropical lungless salamanders (Amphibia: Caudata: Plethodontidae) are characterized by their small size, conserved morphology, and secretive habits, and present some of the most challengin g subjects for systematic study (e.g. Wake & Elias 1983; Highton 1995; Jockusch et al. 2001). As revealed in the preceding chapters, lungless salamanders (Caudata: Plethodontidae) represent the most diverse (in both named and candidate species) and the mos t endangered group of vertebrates in the Chorts Block. In this chapter, I evaluate cryptic diversity and provide a taxonomic revision of an endemic clade of cryptozoic salamanders: the genus Nototriton or moss salamanders. Salamanders as Models for Evol utionary Study Wake (2009) provided an excellent review of the advances in the biological sciences that have result ed in taxon based research focusing on salamanders. In brief (Wake 2009:333): The clade [Caudata] is widespread and diverse, yet sufficientl y small that one can keep all of the species in mind. This facilitates research from 1 Cryptozoic Refers to organisms of small size, cryptic coloration, and that exhibit secretive behavior

PAGE 251

251 diverse perspectives: systematics and phylogenetics, morphology, development, ecology, neurobiology, behavior, and physiology. Different avenues of research offer unique p erspectives on how a relatively old vertebrate clade has diversified. An integrated, hierarchically organized, multidimensional program of research on a taxon illuminates many general principles and processes. Among these are the nature of species and homo logy, adaptation and adaptive radiations, size and shape in relation to issues in organismal integration, ontogeny and development in relation to phylogeny, the ubiquity of homoplasy, ecological niche conservation, species formation, biodiversity, and cons ervation. Specific to the use of salamanders as a study group in the Chorts Block, I recognize a number of advantages. The majority of living salamanders (613 species; AmphibiaWeb, 2 September 2011) are members of the family Plethodontidae (417 species, 6 8% of all living Caudata), and the majority of plethodontids are members of Neotropical restricted genera (265 Neotropical species, 63.6% of all Plethodontidae, 43% of all living Caudata), indicating that an ancient order underwent extraordinary Cenozoic d iversification in tropical America. The Chorts Block is demonstrated to have a diverse endemic salamander fauna, with the majority of endemism found in the highlands (McCranie & Wilson 2002; Wilson & McCranie 2004 b ). In another highland area of high ecoph ysiographic heterogeneity, the comparatively well studied Appalachian Mountains of eastern North America, plethodontids are highly diverse and demonstrate a wide variety of evolutionary patterns and processes, including both adaptive and non adaptive radia tions, parallel evolution, and ecological niche conservatism accompanied wit h morphological conservatism (e.g. Tilley & Mahoney 1996; Kozak & Wiens 2006, 2010; Kozak et al. 2006; Crespi et al. 2010). Among the taxa known from the Chorts Highlands, a numbe r of taxonomic problems exist, and numerous areas of presumably suitable habitat remain unsampled. The Chorts Highlands salamander fauna is highly threatened by loss of habitat,

PAGE 252

252 potentially due to climate change ( as demonstrated in the Appalachian Mountai ns by Milanovich et al. 2010), and by both direct and indirect effects from wider declines in highland amphibian communities (Rovito et al. 2009). Of the 36 species of salamanders recorded from the Chorts Highlands (reviewed in Appendix 1 ), 11 are Critica lly Endangered, 12 are Endangered, 2 are Vulnerable, 1 is Near Threatened, 6 are Least Concern, and 4 are Data Deficient (AmphibiaWeb 2011, IUCN 2011), meaning that close to 70% of all known salamander species in the Chorts Highlands are in the ee highest threat categories (over 80% when poorly known, Data Deficient species are included) Salamander Diversity in the Chorts Block Salamanders originated and initially diversified in Laurasia (Zhang and Wake 2009), with a single family, Plethodonti dae, dispersing into the Neotropics beginning in the late Cretaceous Period (Hanken and Wake 1982; Vieites et al. 2007). Plethodontid salamanders are hypothesized to have dispersed from north to south as the Central American land bridge formed, possibly co lonizing the Chorts Block prior to the divergence of Cryptotriton from remaining neotropical plet hodontids (Appendix 3 of Wiens et al. 2007). Neotropical salamanders can be characterized by their high degree of diversification and widespread morpholog ica l conservatism and homoplasy (Wake 1991; Parra Olea & Wake 2001). Diverse radiations of plethodontids are well documented in western Nuclear Central America (southern Mxico and Guatemala; Wake 1987) and southern Central America (Costa Rica and western Pan ama; Wake 2005); however, much of this cryptic diversity has been revealed only recently by molecular studies (e.g., Garcia Pars et al. 2000; Hanken et al. 2005). Perhaps the best

PAGE 253

25 3 example of this taxonomic uncertainty is seen in the suite of small salaman ders currently divided into the genera Chiropterotriton Cryptotriton Dendrotriton and Nototriton (Garca Pars & Wake 2000). As recently as 1983, these morphologically conserved genera were considered part of a single wide ranging genus, Chiropterotrito n (Wake & Elias 1983). The application of molecular techniques coupled with increased sampling and rigorous studies of external and internal morphology have led to the current taxonomy, which continues to be refined as new populations and taxa are discover ed and described (e.g., McCranie et al. 2008 ; Townsend et al 2010 a ). Even among well studied taxa and regions, such as Bolitoglossa in Costa Rica, morphologically conserved taxa have presented long standing problems that can best be resolved using an inte grative approach that maximizes use of both molecular and morphological evidence (e.g., Hanken et al. 2005). Like western Nuclear and southern Central America, the Chorts Highlands are a site of remarkable plethodontid diversity (McCranie and Wilson 2002) The Bolitoglossa ( Magnadigita ) dunni species group, a Chorts Highlands endemic radiation of 12 nominal species (McCranie et al. 2005), has the highest estimated rate of diversification of any major clade of Mesoamerican salamanders (Wiens et al. 2007). Prior to initiation of this study, there were 13 recognized species of moss salamanders, genus Nototriton distributed across Central American highland forests (Garca Pars & Wake 2000). These salamanders are difficult to differentiate, given their small adult size (all species < 40 mm standard length) and highly conserved morphology (Good & Wake 1993; Garca Pars & Wake 2000). Their utilization of moss mats, deep leaf litter, and rotten logs as habitat makes them particularly difficult to

PAGE 254

254 detect (Good & Wake 1993). The genus is subdivided into three well supported species groups: the N. barbouri group, the N. picadoi group, and the N. richardi group (Papenfuss & Wake 1987, Sav age 2002). Each morphologically defined species group corresponds to a clade su pported by data from the mitochondrial genes 16S and cytochrome b (Garca Pars & Wake 2000, Wiens et al. 2007). The Nototriton barbouri group is the only clade of Nototriton restricted to the Chorts Highlands (Garca Pars & Wake 2000). This group conta ins five nominal species inhabit ing highland forests in eastern Guatemala and northern and central Honduras. The five putative species of the N. barbouri group are each endemic to isolated cloud forest localities in the Chorts Highlands ( Figure 5 1): N. b arbouri (Schmidt 1936), N. brodiei Campbell & Smith 1998, N. lignicola McCranie & Wilson 1996 [ 1997 ] N. limnospectator McCranie, Wilson & Polisar 1998 and N. stuarti Wake & Campbell 2000. Another species of Nototriton found in the Chorts Highlands Notot riton saslaya is extralimital to the Nototriton barbouri group representing the northernmost representative of the N. picadoi group, endemic to a single highland forest area at the southern edge of the Nuclear Central American piedmont in north central N icaragua. Nototriton saslaya aside, the remaining five species of Chorts Highland Nototriton all occur in small areas of highland forest in a belt from easternmost Guatemala to north central Honduras corresponding to the Northern and Central Cordilleras of the Chorts Highlands ( Figure 5 1). While the salamander fauna of Nuclear Central America has been the focus of considerable study (Dunn 1924; Schmidt 1933, 1936; Wake 1987; McCranie & Wilson 1993, 1997; Campbell & Smith 1998), new species continue to be discovered and

PAGE 255

255 Figure 5 1. Distribution of Nototriton in the Chorts Highlands. S haded areas >1000 m elevation; open circle = N. picucha ; open triangles = N. brodiei ; closed triangles = N. limnospectator ; open square = N. sp. A; solid square = N. bar bouri ; open star = N. tomamorum N. sp. B; solid star = N. stuarti ; open diamonds = N. lignicola ; solid diamond = N. saslaya ; N. sp A. and N. sp. B refer to populations currently assigned to N. barbouri described, often from areas that are considered rel atively well known in terms of their amphibian diversity (Campbell et al. 2010; McCranie et al. 2005, 2008; Rovito et al. 2010; Townsend et al. 2009 a 2010 a ; Vsquez Almazn et al. 2009). This can be attributed in part to the small size and cryptic nature of many tropical salamanders, which in the case of Nototriton results in the majority of species being known from 10 or fewer specimens (Good & Wake 1993; McCranie & Wilson 2002). Indeed, the five species of the N. barbouri group have been represented pr eviously in published phylogenetic studies by only seven samples from four putative taxa (Garca Pars & Wake 2000; Wiens et al. 2007; Adams et al. 2009).

PAGE 256

256 Intensive sampling in the Chorts Highlands over the past six years has led to the collection of addi tional samples of nominal taxa and the discovery of new populations of Nototriton from isolated localities (Chapters 3 and 4). In order to evaluate the evolutionary relationships among Chorts Highland Nototriton in light of their currently accepted taxono my, I employed two strategies for analysis of a three gene mtDNA dataset, which supplements the 16S/COI dataset used in Chapter 4 with the a fragment of the gene cytochrome b (cyt b ). First, I used a DNA barcoding approach to identify potential species bou ndaries and candidate species by evaluating intra and interspecific sequence divergence and the effectiveness of each gene for species delimitation using uncorrected distance based methods. Second, I inferred phylogenetic relationships using Bayesian and maximum likelihood (ML) analysis with evolutionary models partitioned by gene and codon. Finally, I applied the distance based and phylogenetic results, supplemented by comparative data from external morphology and osteology, to redefine N. barbouri and de scribe three new species of Nototriton endemic to the Chorts Highlands of Honduras Methods and Materials Sampling Taxa and samples used in this study, along with their associated locality data and museum and GenBank accession numbers, are presented in Ta ble 5 1. All nominal taxa in the N. barbouri group were included with the exception of N. stuarti which is only known from a single specimen col lected in 1991 (Wake & Campbell 2000). Nototriton saslaya the only representative of the N. picadoi group fou nd in the Chorts Highlands, was used as an outgroup. Institutional abbreviations follow those standardized by the American Society of Ichthyologists and Herpetologists

PAGE 257

257 (http://www.asih.org/codons.pdf), except for IRL (field series of Ileana R. Luque Monte s) used for a sample donated to the Florida Museum of Natural History in May 2009 that remains uncataloged. Forest formations follow Holdridge (1967) as applie d by McCranie & Wilson (2002) In this chapter, I present genetic evidence for two undescribed s pecies heretofore considered synonymous with N. barbouri. However, I do not include the morphological diagnoses and descriptions associated with those species, as they have not yet been published at the time of completing this dissertation. DNA Extraction PCR Amplification, and Sequencing Template DNA was extracted from muscle tissue using the Qiagen PureGene genes 16S large subunit RNA (16S), cytochrome b (cyt b ), and cytochrome oxidase subunit I (COI) were amplified using the primers 16Sar L and 16Sbr H for 16S (Palumbi et al 1991), MVZ15L and MVZ16H for cyt b (Moritz et al 1992) and dgLCO 149 0 and dgHCO 2198 for COI (Meyer 2003). PCR reactions were typically 20 L in total volume, containing ~25ng of DNA template, 4 L 5X PCR buffer, 1.2 L MgCl 2 (25mM), 0.09 L dNTPs (10 mM), 0.8 L of each primer (10 M), 0.2 L GoTaq Flexi polymerase (Promega, Madison, WI, USA), and 11.91 L H 2 O. Amplification profiles were as fol lows: for 16S, an initial denaturation for 3 minutes at 94C, 35 cycles of denaturation at 94C for 45 seconds, annealing at 50C for 45 seconds, and extension at 72C for 45 seconds, with a final elongation at 72C for 5 minutes; for cyt b an initial den aturation for 3 minutes at 94C, followed by 38 cycles of 94C for 30 seconds, 48C for 1 minute, and 1 minute for 45 seconds, final elongation at 72C for 5 minutes; and for COI,

PAGE 258

258 denaturation for 1.5 minutes, 37 cycles of 94C for 40 seconds, 45C for 40 seconds and 72C for 40 seconds, with a final elongation at 72C for 5 minutes. All PCR products were verified using electrophoresis on a 1.5% agarose gel stained with ethidium bromide. Unincorporated nucleotides were removed from PCR products using 1 uL o f ExoSAP IT (USB, Santa Clara, CA, USA) per 10uL of PCR product. I cycle sequenced both forward and reverse stands using the BigDye Terminator 3.1 Cycle Sequencing kit, followed by spin column filtration through Sephadex before electrophoresing the product s on an ABI 3130xl (Applied Biosystems, Inc). Sequence Alignment and Model Selection A dataset containing all available sequences of Nototriton was generated from a combination of our newly generated sequence data (Table 5 1) and published data available f rom NCBI (http://www.ncbi.nlm.nih.gov/). Sequences were aligned using ClustalW (Thompson et al 1994) within the program package MEGA5 (Tamura et al 2011) using the default parameters. The dataset was partitioned by gene (16S, which codes for RNA) and by codon position (1st, 2nd, 3rd) for cyt b and COI (both protein coding genes) and selected the best fit model of nucleotide evolution for each gene and each partition with the program jModeltest v0.1 (Posada 2008), which uses PhyML 3.0 (Guindon & Gascuel 2 003) to estimate models under a likelihood framework. The number of substitution schemes was set to three to limit the number of models tested to 24, corresponding to the number of different models that can be implemented in MrBaye s 3.1.2 (Huelsenbeck & Ro nquist 2001). Models selected for each partition are summarized in Table 5 2

PAGE 259

259 Sequence Analyses A DNA barcoding approach was utilized in order to determine whether monophyletic clusters of samples corresponding to named taxa, as well as to identify candid ate species for further taxonomic evaluation (Vences et al 2005; Smith et al 2008). Uncorrected (p distance) pairwise sequence divergence was calculated for all samples and for each gene, and uncorrected neighbor joining (NJ) distance trees (10,000 boots trap pseudoreplicates) were estimated to provide graphical representation of intra and interspecific variation. Sequence divergence estimation and NJ analyses were performed in MEGA5 (Tamura et al. 2011). Two of the three target genes are protein coding, making them potentially susceptible to substitution saturation at the third codon position that, when excessive, can rende r the locus phylogenetically un i nformative (Halanych & Robinson 1999; Farias et al 2004; Parra Olea et al 2004). Substitution satura tion was evaluated using an entropy based index calculated in the program package DAMBE 5.2.34 (Xia & Xie 2001), which uses two simulated topologies (one perfectly symmetrical and one extremely asymmetrical) based on the supplied sequence data to calculate an Index of Substitution Saturation ( I SS ). High substitution saturation is indicated by an I SS significantly greater than or not significantly different than the Critical I SS ( I SS.C ) for both the perfectly symmetrical topology and an extremely asymmetrica l topology (Xia et al 2003; Xia & Lemey 2009) determined with two tailed t test. Prior to measuring substitution saturation, the proportion of invariant sites was estimated for each gene in DAMBE 5.2.34 (Xia & Xie 2001) using a goodness of fit test with a Poisson+Invariant distribution, and the proportion of invariant sites was used as a parameter to optimize estimation of substitution saturation.

PAGE 260

260 Bayesian Inference and Maximum Likelihood Phylogenetic Analyses Bayesian inference (BI) was performed using Mr Baye s 3.1.2 (Huelsenbeck & Ronquist 2001), and consisted of two parallel runs of four Markov chains (three heated, one cold) run for 10 x 10 6 generations and sampled every 1,000 generations, with a random starting tree and the first 2 x 10 6 generations dis carded as burnin. Maximum likelihood analysis was carried out in RAxML v7.2.8 (Stamatakis 2006), with 1000 bootstrap pseudoreplicates under the default GTR GAMMA substitution model (the simplest model implemented in RAxML); the dataset was partitioned by g ene for 16S and by codon position for cyt b and COI Comparative Morphology Morphological measurements were taken with precision digital calipers and a stereo microscope with an optical micrometer, with measurements rounded to the nearest 0.1 mm. Abbrevia tions used for morphological measurements are as follows: snout to posterior edge of vent, SL; axilla groin length, AG; trunk width at midbody, TW; head length from tip of snout to gular fold, HL; head width taken at maximum, HW; tail length, TL; hind limb length, HLL; forelimb length, FLL; combined forelimb and hind limb lengths, CLL; hind foot length, HFL; hind foot width, HFW; and nares length, NL. To allow for comparative studies across taxa, most measurements are standardized by SL; morphological compa risons of the aforementioned characters are presented in Table 5 4 Comparative data for other taxa not examined by the authors is taken from Good & Wake (1993), Campbell & Smith (1998), Lynch & Wake (1978), McCranie et al (1998), Ehmcke & Clemen (2000), Khler (2002), McCranie & Wilson (2002), Savage (2002), and Wake & Campbell (2000).

PAGE 261

261 Results Best fit nucleotide substitution models varied by gene and codon position, supporting use of a gene and codon based partitioning strategy (Table 5 2). For the 489 b p of 16S, 10.8% of nucleotides were variable. Variability increased to 27.4% and 27.8% for COI (658 total bp) and cyt b (702 total bp) respectively. Distance based analyses of each gene yielded unambiguous results in terms of delimiting species level linea ges and clusters, displaying a bimodal pattern of pairwise sequence divergence graphically and easily interpreted using distance scaled NJ trees ( Figure 5 2). The smallest range of intraspeci fic sequence divergence in our data was for COI (0.002 0.003%), followed by 16S (0.0 0.6%) and cyt b (0.0 1.9%). Interspecific divergence ranged from 1.2 6.4% for 16S, the most conservative gene, to 6.1 13.9% for COI and 5.2 14.7% for cyt b (Table 5 3). Fo r all three genes, the bimodal distribution of sequence divergence allows for simple identification between the ranges of intra and interspecific divergence ( Figure 5 2), with 16S having the narrowest gap (0.6%), followed by cyt b (3.3%) and COI (5.8%). Substitution saturation was not a factor for 16S, with the I SS value (0.2197) being highly significantly less than the critical I SS+C value for both the symmetrical ( I SS+C(SYM) = 0.7112, P = 0.0000) and asymmetrical ( I SS+C(ASYM) = 0.4960, P = 0.0000) topologies. Likewise, COI did not exhibit evidence of substitution saturation, with an I SS value (0.3667) significantly less than the critical I SS+C value for both the symmetrical and asymmetrical topologies ( I SS+C (SYM) = 0.7320, P = 0.0000; I SS+C (ASYM) = 0.5482, P = 0.0000 ). For cyt b however, I SS (0.7722) was significantly larger than I SS+C for both the symmetrical ( I SS+C(SYM) = 0.7337, P = 0.0005) and asymmetrical ( I SS+C(ASYM) = 0.5254,

PAGE 262

262 Figure 5 2. Comparison of delimit s pecies boundaries in Nototriton U nrooted neighbor joining trees (10,000 bootstrap replicates) scaled by uncorrected p distance, with shaded area representing gap between upper limit of intraspecific variation and lower limit of interspecific variation. No te that the trees for COI and cyt b are scaled to 0.01, while the 16S tree is scaled to 0.005 due to lower overall divergence. Numbers following taxon labels refer to those in Table 5 1.

PAGE 263

263

PAGE 264

264 P = 0.0000) topologies, indicating that excessive substitution satur ation was present and the data therefore were poorly suited for use phylogenetic study (Xia et al 2003; Xia & Lemey 2009). To further investigate, I tested for substitution saturation at each codon position on cyt b and found the third position to have a higher but not significantly different I SS value (0.6981) on the perfectly symmetrical topology ( I SS+C(SYM) = 0.6797, P = 0.4553) and to be significantly higher than the asymmetrical topology ( I SS+C(ASYM) = 0.4554, P = 0.0000), while the 1st and 2nd codon posit ions did not show evidence of saturation (1st codon position: I SS = 0.3721; I SS+C (SYM) = 0.6797, P = 0.0000; I SS+C (ASYM) = 0.5482, P = 0.0509; 2nd codon position: I SS = 0.3438; I SS+C (SYM) = 0.6797, P = 0.0000; I SS+C (ASYM) = 0.5482, P = 0.0094). Phylogenetic analyses were performed both on the complete combined mitochondrial dataset, as well the combined dataset with the 3rd codon position of cyt b excluded in order to account for saturation ( Figure 5 3). Both analyses recovered a 00/100; pp= 1.0/1.0) consisting of N. brodiei and two unnamed candidate species ( N sp. A and B) currently referred to as N. barbouri sensu lato Also strongly supported by both analyses is the sister relationship between N. barbouri sensu stricto and N. li mnospectator (bs=82/87; pp= 1.0/1.0), a N. barbouri group. The two datasets yielded different topologies in terms of the placement of N. tomamorum and the N. sp. inquirenda 2 with respect to the remaining members of the N. barbouri g roup. The phylogeny excluding 3rd position recovers a topology congruent with recently published phylogenies of Nototriton (Townsend et al. 2010 a ) supplemented by additional sampling, including two samples from a newly discovered

PAGE 265

265 Figure 5 3. Bayesian phy lograms showing discordance between combined mtDNA phylogenies due to saturation at the third codon position for cytochrome b N ote the alternative placement of Nototriton tomamorum and N. picucha sp. nov. as sister to the remaining N. barbouri group. Boot strap scores from maximum likelihood analyses shown above branches; posterior probability values from Bayesian inference shown below branches. population in eastern Honduras. The new species is recovered as sister to the remaining N. barbouri group (bs=82 ; pp= 1.0), with N. tomamorum as sister to t he entire clade (bs=100; pp= 1.0), including the new species ( Figure 5 3). The phylogeny including 3rd position recovers N. tomamorum as sister to N. lignicola (bs=72; pp= 0.84), and instead, places the new species as sister to a N. barbouri group (bs=100; pp= 1.0).

PAGE 266

266 Discussion DNA Barcode Identification of Chorts Highland Moss Salamanders Sequence divergence data from 16S, cyt b and COI all exhibit a bimodal distribution of uncorrected pairwise comparisons, each de indicative of clear different iation between intra and inter specific variation ( Figure 5 2 ). Results for all three loci were congruent, identifying the same sample groupings and unambiguously separating putative species level lineages and revealing three potential candidate species: a newly discovered population from the Sierra de Agalta, N. barbouri sensu lato from Parque Nacional Pico Bonito ( N sp. A), and N. barbouri sensu lato Reserva de Vida Silvestre Texguat ( N sp. B). Samples representing two recently discovered allopatric populations of Nototriton were also included in this study: one from the vicinity of Cataguana in Parque Nacional Montaa de Yoro and the other from Parque Nacional Cerro Azul Membar on the eastern side of Lago de Yojoa. All three gene loci clearly indicated that these samples were conspecific with N. lignicola and N. limnospectator respectively, and therefore confirm that these samples represent new populations of two species previously consider ed to be highly threatened single site endemics (Townsend et al. 2011 b ). In the case of Chorts Highland Nototriton analysis of mtDNA sequence data using a barcoding approach is shown here to be an effective means of identifying potential candidate speci es and newly discovered populations of known taxa, but in and of itself is not grounds for proposing new taxa or making other taxonomic changes. Rather, I view the barcoding approach as an effective and increasingly necessary first step in an integrative p rocess that leads to correct taxonomic allocation of newly collected samples, particularly in difficult and cryptic groups like Nototriton

PAGE 267

267 Influence of Substitution Saturation in Cytochrome B Dataset Our analyses of sequence data from cyt b revealed exce ssive substitution saturation at the third codon position, a phenomenon that has been shown to lead to bias in phylogenetic inference in studies of deep relationships among salamanders (Zhang & Wake 2009), including plethodontids (Mueller et al 2004; Ma ce y 2005; Vieites et al 2011 ). This study provides an example of excessive saturation biasing phylogenetic results at the interspecific level in salamanders, and its influence appears to be limited, at least in this case, to two species level lineages. That the saturation problem has not been previously identified in Nototriton can be attributed to use of a longer fragment of cyt b (702 bp) than had been used in previous phylogenetic analyses of the gen us (385 bp; Garca Pars & Wake 2000; Townsend et al 20 10 a ). When our dataset is trimmed to 385 bp to match the length of most cyt b sequences available from NCBI (http://www.ncbi.nlm.nih.gov/) for Nototriton substitution saturation is not a factor at any codon position, with all I SS values being significantl y less than the critical I SS+C value for both the symmetrical topologies (1st codon position: I SS = 0.2396, I SS+C(SYM) =0.6639, P = 0.0000; 2nd codon position: I SS = 0.0.989, I SS+C(SYM) = 0.6630, P = 0.0000; 3rd codon position: I SS = 0.3061, I SS+C(SYM) = 0.6690, P = 0.0000 ) and asymmetrical topologies (1st codon position: I SS = I SS+C(ASYM) = 0.4113, P = 0.0037; 2nd codon position: I SS+C(ASYM) = 0.4101, P = 0.0004; 3rd codon position: I SS+C(ASYM) = 0.4575, P = 0.0000). Phylogenetic analysis using the 385 bp cyt b fragment (not shown) pr oduced a topology completely congruent with that of the dataset that excluded the 3rd codon position of cyt b (468 bp), suggesting that the 385 bp fragment is short enough to mitigate bias from 3rd position saturation

PAGE 268

268 Phylogenetic Systematics and Candidat e Species As indicated here, paraphyly exists among populations currently referred to as Nototriton barbouri (Schmidt 1936). Populations from Reserva de Vida Silvestre Texguat and Parque Nacional Pico Bonito in northern Honduras referred to N. barbouri (M cCranie & Wilson 1995; McCranie 1996 a; McCranie & Wilson 2002; McCranie & Castaeda 2007) are paraphyletic with respect to a sample from Montaa de Macuzal, the presumed type locality of N. barbouri (McCranie and Wilson 2002: 145). Based on our phylogeneti c results ( Figure 5 3 ), N. barbouri sensu stricto is the sister species to N. limnospectator which Montaa de Santa Brbara eastw ard through the Sierra de Sulaco to Montaa de Macuzal Garca Pars & Wake (2000: 49) suggested that samples of N. barbouri sensu lato from Parque Nacional Pico Bonito (USNM 339712, 497552) and Reserva de Vida Silvestre Texguat (USNM 509333) might represent distinct taxa, an assertion well supported by our res ults. Populations of N. barbouri sensu lato from Parque Nacional Pico Bonito ( N sp. A) and Reserva de Vida Silvestre Texguat ( N sp. B) form a well supported clade with N. brodiei of eastern Guatemala and northwestern Honduras, N. barbouri group ( Figure 5 3 ). Somewhat surprisingly given their geographic distributions, and contrary to the previous cyt b based phylogenetic hypothesis (Garca Pars & Wake 2000), both combined phylogenies strongly support N sp. B from north central Honduras, as sister to a N. brodiei / N sp. A clade (bs=100; pp= 1.0). These results provide phylogenetic support for recognition of both N sp. A and N sp. B as species level taxa; however one known

PAGE 269

269 population of N. barbouri sensu lato remains to be evaluated phylogenetically (Pico Pijol in Departamento de Yoro). Two samples from a newly discovered population of Nototriton in Parque Nacional Sierra de Agalta in eastern Honduras are shown to be conspecific with each other and a relativel y divergent lineage within the group ( Figure 5 3 ). This undescribed species appears to be a typical member of the N. barbouri group in terms of external morphology, with relatively small nostrils and free, well differentiated toes, which leads us to conclu de that the phylogeny excluding 3rd codon position on cyt b which includes the new species as part of an ingroup that is sister to N. tomamorum most accurately represents the evolutionary history of these salamanders. The alternative phylogenetic hypothe sis, obtained using the concatenated mtDNA dataset with cyt b fragment including the saturated 3rd codon position, implies that the distinctive morphology of N. tomamorum such as enlarged nostrils and syndactylous feet, are derived characters that evolved as the result of the divergence of N. tomamorum and N. lignicola from a common ancestor. Future analyses that include additional samples and nuclear gene loci should provide some measure of resolution of the phylogenetic position of N. tomamorum Systema tic s A Divergent New Lineage from Refugio de Vida Silvestre Texguat During my first visit to Refugio de Vida Silvestre Texguat in April 2008, malacologist John Slapcinsky collected a small salamander in l eaf litter packed into a crevice along the stream below the La Fortuna campsite. This single specimen possesses a series of morphological characteristics unique among Honduran salamanders Comparison of morphological and mitochondrial DNA sequence data

PAGE 270

270 confirms that this specimen represents a distinctive and undescribed species and a relict lineage sister to the clade corresponding to the Nototriton barbouri group (Figure 5 4) Nototriton tomamorum Townsend, Butler, Wilson, & Austin 201 0 a Figure 5 4 Nototriton sp. inq. 1: Chapter 3. Holotype. UF 155477, a female from 2.5 km north northeast of La Fortuna 1,550 m elevation, Refugio de Vida Silvestre Texguat, Departamento de Yoro, Honduras. GenBank accession numbers GU9717 31 (16S), GU97173 2 (cyt b ), JN37740 7 (COI). Diagnosis. A small member of the genus N ototriton (SVL=26.9 mm; Table 5 4 ) based on having 13 costal grooves (>16 costal grooves in Oedipina ), hands and feet longer than broad (hands and feet broader than long in Bolitoglossa ), and nares that are smaller than most Cryptotriton and Dendrotriton ( Figure 5 4 A, B; 0.018 NL/SVL; 0.020 0.029 NL/SVL in Cryptotriton [except some individuals of C. veraepacis ] and Dendrotriton ). Cryptotriton veraepacis has nares ranging from 0.017 0.027 NL/SVL (mean 0.022), and can be differentiated from N. tomamorum by ha ving a uniformly dark gray ventral surface (ventral surface pale with gray flecks in N. tomamorum ; Figure 5 4B ). Generic placement in Nototriton is also strongly supported by sequence data from the mitochondrial genes 16S COI, and cyt b (Table 5 3; Figure s 5 2, 5 3 ). This new species is distinguished from all other Nototriton except N. richardi and N. tapanti by having syndactylous hands and feet ( Figure 5 4D, E ; hands and feet with free, differentiated toes in all other species) and relatively large nar es ( Figure 5 3C B; 0.018 NL/SVL versus 0.010 0.016 in N. picadoi 0.003 0.014 in N. abscondens 0.012 in N. stuarti 0.005

PAGE 271

271 Figure 5 4 Nototriton tomamorum. A) Dorsal and ventr al views of the holotype of N tomamorum ( UF 155377 ), standard length 2 6 .9 m m. B) Dorsal view of the head of the holotype of N tomamorum C) Lateral view of the head of the holotype of N tomamorum D) Dorsal view and E) ventral view of right hind foot of the holotype of N tomamorum showing the lack of separation or differentia tion in the toes, and absence of subdigital pads. Photos J.H. Townsend.

PAGE 272

272 0.011 in N. barbouri 0.006 0.009 in N. lignicola 0.004 0.009 in N. guanacaste 0.004 0.005 in N. brodiei 0.003 in N. limnospectator 0.003 in N. major and 0.002 0.003 in N. sasl aya ). Nototriton tomamorum can be further differentiated from members of the N. barbouri group by having a broader head (0.145 HW/SVL versus 0.138 in N. stuarti 0.104 0.132 in N. barbouri 0.120 in N. brodiei 0.103 0.118 in N. lignicola and 0.095 0.118 in N. limnospectator ) and fewer maxillary teeth (26, versus 36 in N. stuarti 41 54 in N. barbouri 42 55 in N. limnospectator 46 54 in N. lignicola and 60 62 in N. brodiei ), from members of the N. picadoi group by having a relatively shorter tail (0.911 TL/SVL, versus 1.441 in N. major 1.123 1.344 in N. picadoi 1.013 1.365 in N. abscondens 1.210 1.337 in N. guanacaste and 1.10 1.30 in N. gamezi ) and narrower feet (0.037 HFW/SVL, versus 0.058 0.071 in N. abscondens 0.059 in N. major 0.060 0.070 in N picadoi and 0.066 0.072 in N. guanacaste ), and from N. saslaya by having shorter forelimbs (0.160 FLL/SVL, versus 0.194 0.210 in N. saslaya ) and hind limbs (0.197 HLL/SVL, versus 0.217 0.244 in N. saslaya ) and narrower feet (0.037 HFW/SVL, versus 0.075 0.091 in N. saslaya ). The new species also differs from N. richardi and N. tapanti in having a pale ventral surface mottled with gray chromatophores (ventral surface brown with dark flecks in N. richardi and dark brown in N. tapanti ), by having a tail tha t is shorter than the snout vent length (0.91 TL/SVL, versus 1.072 1.482 in N. richardi and 1.205 in N. tapanti ), longer forelimbs (0.160 FLL/SVL, versus 0.140 0.146 in N. richardi and 0.147 in N. tapanti ), longer hind limbs (0.197 HLL/SVL, versus 0.174 0. 187 in N. richardi and 0.174 in N. tapanti ), and narrower feet (0.037 HFW/SVL, versus 0.044 0.050 in N. richardi and 0.041 in N. tapanti ). This new species is also well differentiated from all other species of Nototriton

PAGE 273

273 based on mitochondrial sequence dat a, and is 3.5 5.7 % divergent on 16S 11.5 13.9 % divergent on COI, and 9.5 1 3 .1% divergent on cyt b from other Chorts Highland Nototriton (Table 5 3) Description of holotype. Nototriton tomamorum is known only from a single, presumably female (mental gla nd and cloacal papillae absent) specimen, preserved with its mouth open and tongue extended, and is a relatively small member of the genus (SVL=26.9 mm, total length=51.4 mm) with a slender body and reduced limbs. Its head is rounded and slightly broader t han the body, and the nostrils are relatively large (NL/SVL=0.018), and the snout is rounded and of moderate length ( Figure 5 4B, C ). The nasolabial protuberances are apparent but not well developed, and barely extend below the upper lip line. The eyes are relatively large and protuberant, and the parotoid glands appear large but not well defined. The teeth are exceedingly small; there are approximately 26 maxillaries, 4 premaxillaries set slightly forward from the maxillary teeth, and 11 vomerines; vomerin e teeth are arranged in two short medially positioned arches. The limbs are short (CLL/SVL=0.36), with the adpressed limbs being separated by approximately 5.5 costal grooves. The hands and feet are narrow and have poorly developed, poorly differentiated d igits that are fused and lack subdigital pads ( Figure 5 4D, E ). The free tips of digits III on the hands and III and IV on the feet are pointed, and digits I, II and IV on the hands and I, II, and V on the feet are very short and essentially completely fus ed, being demarcated by shallow grooves on the dorsal side of the feet. The relative length of the digits is I
PAGE 274

274 most apparent on the ventral side, and is slightly compressed laterally (tail depth 1.17 times tail width at level of basal constriction). Measurements of holotype (in mm). SVL 26.9; AG 15.2; TW 3.6; HL 4.8; HW 3.9; TL 24.5; HLL 5.3; FLL 4.3; CLL 9.6; HFL 1.7; HFW 1.2; NL 0.5; eye length 1.6; eye width 1.2; interorbital distance 1.4; anterior rim of orbit to snout 1.1; distance separating internal nares 0.8; distance separating external nares 1.8; snout projection beyond mandible 0.6; tip of snout to axill a 7.7; distance from axilla to groin 15.2; snout to anterior edge of vent 24.9; tail depth at basal constriction 2.8; tail width at basal constriction 2.4. Coloration of holotype. Dorsal surfaces of head, body, and tail medium grayish brown, with profuse pale chromatophores laterally, becoming less abundant dorsolaterally. The head has some pale mottling on the top of the snout, and two irregular lines of pale chromatophores extending from the lateral region above the forelimbs onto the posterior portion o f the head and parotoid glands. There is a very thin, pale middorsal stripe, with a herringbone pattern with lines extending from the middorsal stripe posteriorly. There is an indistinct dark dorsolateral stripe starting about one third the way down the tr unk and extending onto the proximal one third of the tail. Ventral surface of head, body, and tail cream, mottled with dark gray chromatophores, becoming somewhat more profuse toward the distal end of the tail. Etymology. tomamorum Tomams are the highest level of deities recognized in the belief system of the indigenous Tolupan of Honduras, of which there are four: Tomam Pones Popawai (Grandfather Tomam), his wife Tomam Pones Namawai (mother of all that exists), and

PAGE 275

275 their children Tomam the Elder and Tomam the Younger (Chapman 1992). This name is given in recognition that the Tolupan are the traditional inhabitants of this area and that this new species is known only from the Cordillera Nombre de Dios, or Distribution and Natural History The single known specimen was collected during the daytime from leaf litter packed onto a rock ledge alongside a small cree k at about 1 550 m elevation in the Lower Monta ne Wet Forest formation (Townsend et al. 2010a) This species is presumably endemic to the vicinity of the type locality, and is sympatric with a congener, N. sp. A and three other plethodontids: Bolitoglossa dofleini Bolitoglossa cf. porrasorum and Oedipina gephyra Conservation Status. Critically Endangered B1ab(iii) +2ab(iii) A New Species from the Sierra de Agalta Nototriton picucha Townsend, Medina Flores, Murillo, and Austin 2011 a Figure 5 5 Nototrito n sp. inq. 2: Chapter 3. Holotype: USNM 578299 ( Figures 5 5 A 5 5 B ) a female, from the northwestern slope of Cerro La Picucha (14.9733N, 85.9279W), 1 890 m, Parque Nacional Sierra de Agalta, Departamento de Olancho, Honduras, collected 17 July 2010 by M Medina F., J. L. Muri llo, I. Zniga, and J. M. Soto. Original field number JHT 3192; Genbank accession numbers: JN377388 (16S), JN377392 (cyt b), JN377404 (COI). Paratype : USNM 578298 (Figure 5 5 C ) same collection data as holotype, except from 1 920 m elevation. Genbank accession numbers: JN377389 (16S), JN377393 (cyt b), JN377405 (COI).

PAGE 276

276 Diagnosis. A member of the genus Nototriton (SL=27.9 mm) possessing 13 costal grooves (>16 costal grooves in Oedipina ), hands and feet longer than broad (hands and feet broader than long in Bolitoglossa ), and small nares (0.007 NL/SL; 0.017 0.029 NL/SL in Cryptotriton and Dendrotriton ). Placement of the new species in the Nototriton barbouri species group is supported by phylogenetic analysis of the mitochondrial genes 1 6S and cyt b ( Figure 5 2). In addition to molecular characteristics, the new species is distinguished from other members of the N. barbouri group by having a broader head (HW/SL N. barbouri N. brodiei N. lignicola and N. stuarti by having relative 0.179, versus 0.142 0.174 in N. barbouri 0.148 0.151 in N. brodiei 0.137 0.160 in N. lignicola and 0.172 in N. stuarti ) and relativ 0.204 0.218, versus 0.153 0.200 in N. barbouri 0.166 0.180 in N. brodiei 0.158 0.181 in N. lignicola and 0.178 in N. stuarti ), and from N. limnospectator by havi ng narrower hind feet 0.043, v ersus 0.048 0 0.007, versus 0.003). Nototriton tomamorum can be distinguished from N picucha by having enlarged nares (NL/SL= 0.018, versus 0.007 0.008 in N picucha ), syndactylous feet lacking subdigital pads (toes well differ entiated with subdigital pads in N picucha ), and fewer max illary and vomerine teeth (MT=26 and VT =11, versus 41 and 16 19 and N picucha ). Nototriton saslaya a member of the N. picadoi group, is known from a single highland forest site in northern Nicara gua, and differs from the new species in having fewer maxillary teeth (17 23, versus 41 in N. picucha ) and vomerines (3 11, versus 16 19 in N. picucha ), relatively longer front limbs (FLL/SL 0.194 0.210, versus 0.179 0.191 in N. picucha ) and relatively wid er hind feet (HFW/SL 0.075 0.091, versus 0.042 0.043 in N.

PAGE 277

277 Figure 5 5 Nototriton picucha A) Dorsolateral and ventrolateral views of the holotype of N picucha (USNM 578299), standard length 27.9 mm (photo J.H. Townsend). B) X ray radiograph of the h olotype of N picucha (photo J.H. Townsend). C) Paratype of N picucha (USNM 578298) i n life, standard length 25.7 mm ( photo M elissa Medina Flores ) picucha ) Comparative morphological data for Chorts Highland Nototriton are summarized in Table 5 4. Description of holotype. A relatively small female (SL=27.9 mm, total length=59.8 mm) with a slender body and reduced limbs. The head is rounded, slightly broader than the body; nostrils are relatively small (NL/SL=0.007), and the snout is acutely rounded and of moderate length. The nasolabial groove is apparent, but nasolabial protuberances are not well developed and barely visible. The eyes are relatively large and protuberant, and the parotoid glands are apparent, but relatively flat and not well define d. There are 41 maxillary teeth, 2 slightly enlarged premaxillary teeth

PAGE 278

278 in line with the maxillaries, and 19 vomerine teeth. The limbs are short (CLL/SL=0.38), with the adpressed limbs being separated by approximately 3.5 costal grooves. The hands and feet are narrow with well developed digits that bear subdigital pads. The relative length of the digits is I
PAGE 279

279 pale chromatophores sep arating dark reddish brown dorsal and dorsolateral ground color from dark blackish gray ventral ground color; venter with some sparse whitish gray spotting primarily on the chin and around ventral midline; yellowish brown coloration extends down from the d orsolateral surfaces around the basal portion of the tail to surround the vent with yellowish brown mottling. Proximal one half to two thirds of forelimbs and two thirds to three quarters of hind limbs mottled yellowish brown dorsally and ventrally. Ventra l surface of head, body, and tail cream, mottled with dark gray chromatophores, becoming somewhat more profuse toward the distal end of the tail. See Figure 5 4 b for color photographs of the dorsal and ventral surfaces of the holotype. Variation in paraty pe. The f emale paratype (Figure 5 5 c; USNM 578298, SL= 25.7, tail missing) agrees almost completely with the holotype in most characteristics and standardized ratios, with the only notable differences being more premaxillaries (6, same size as and in line w ith the maxillaries), fewer vomerines (16), and in having a limb interval of approximately 4.5. Coloration is also similar to that seen in the holotype. Osteology Based on examination of digital radiographs for both the holotype ( Figure 5 5 b ) and paratype Nototriton picucha is a typical member of the genus (Wake & Elias 1983) possessing a single cervical vertebra, 14 trunk vertebrae (the anterior 13 of which bear ribs), 2 caudosacral vertebrae; a complete skull roof formed through contact of the parietal bones; frontal processes of premaxilla fused at point of origin and separate immediately dorsoposterior to origin; septomaxillary bones absent; preorbital vomerine processes well developed into elongate arches bearing a single row of

PAGE 280

280 numerous teeth; colume lla absent; no tarsal spur evident; phalangeal formulae 1 2 3 2 and 1 2 3 3 2; penultimate phalanges reduced, exceeded in length by terminal phalanges on digits II, III, and IV of the forelimbs and digits II, III, IV, and V of hind limbs; terminal phalange s tapered at distal tip, or slightly expanded in digit III of forelimbs and digit III of the hindlimbs; mesopodial elements not mineralized. Etymology. The specific epithet references the type locality and tallest mountain in the Sierra de Agalta, Cerro La Picucha, whose high elevation elfin forests make it one of the natural wonders of Honduras. The type series was collected on the northwestern slope of Cerro La Picucha, above the highest basecamp used to access the summit. Natural history. The type spe cimens of Nototriton picucha were collected at 1 890 and 1 920 meters in the Lower Montane Wet Forest formation on the north northwest slope of Cerro La Picucha (2 354 m) in the Sierra de Agalta. The holotype was collected near the highest base camp used f or accessing the elfin forest atop Cerro La Picucha, and the paratype was found farther up the ridgeline trail to the peak before the transition to elfin forest. Both specimens were collected while active underneath leaf litter on the ground between 21:00 and 00:00 hours on a rainy night. Nototriton picucha is sympatric with another endemic plethodontid salamander, Bolitoglossa longissima which was also found along the same trail. Conservation Status. Critically Endangered B1ab(iii). Remarks. Parque Naci onal Sierra de Agalta contains the easternmost fragment of cloud forest (Lower Montane Wet Forest formation) in Nuclear Central America, being located in Dept. Olancho approximately 180 km to the northeast of Tegucigalpa Honduras The park was established in 1987 to protect an area of approximately of

PAGE 281

281 73,924 ha under the management of the National Institute of Conservation and Forest Development, Protected Areas and Wildlife (ICF). There are 75 species of amphibians and reptiles recorded from Parque Nacion al Sierra de Agalta (Castaeda 2006; McCranie & Cruz 2010); however to date only three species ( Bolitoglossa longissima Nototriton picucha and Omoadiphas cannula ) are considered to have geographic distributions completely restricted to this mountain rang e. Restriction of the taxon Nototriton barbouri (Schmidt, 1936) The precise type locality of Nototriton barbouri is a matter of some uncertainty. The taxon was described by Schmidt (1936: 43) based on material collected by R.E. bromeliads, and from altitudes between 5000 [= 1,524 m] feet (the type), and 6000 [= only locality with that name and local access to elevations above 1,800 m is El Portillo (15.095323N, 87.343902W), a small community sitting at around 1,300 m elevation on the northern side of Montaa Macuzal (maximum elevation approximately 1,945 m ; Chapter 3, Figures 3 7G, 3 7H ), an isolated karstic mountain supporting a small fragment of broadleaf clo ud forest on its summit McCranie and Wilson (1995: 137 ) suggested that Bolitoglossa porrasorum coll ected from Portillo Grande by Stadelman and I agree that Montaa Macuzal indeed represents the only known suitable habitat for cloud forest salamanders in the vicinity of El Portillo, and therefore consider it to be the type locality of N. barbouri

PAGE 282

282 A single specimen (AMNH 54949) from a locality near Lago de Yojoa ( north slopes of Cerro Azul Membar, 860 m elevation [McCranie & Wilson 2002:576] ) is also assigned to this taxon. Given th e confirmation that populations of Nototriton from nearby Cerro Azul Membar represent N. limnospectator (Townsend et al. 2011b; Chapter 4), I consider this single specimen to be represent ative of N. limnospectator and not a distjunct and lower elevation locality for N. barbouri sensu stricto Nototriton barbouri (Schmidt 1936) Oedipus barbouri Schmidt 1936: 43 Holotype. MCZ 21247, an adult male from Portillo Grande, 1,524 m elevation, Departamento de Yoro, Honduras; collecte d 9 May 1934 by R.E. Stadelm an. Paratypes. All female s from the vicinity of the holotype, 1,524 1,828 m elevation; MCZ 21248 50, FMNH 21866 67. G enetype. UF 156538 an adult female from Montaa Macuzal (15.079455N, 87.352985 W), 1 760 m elevation, above El Panal to the west of Yor ito, Departamento de Yoro, Honduras; collected 9 April 2008, by J.M. Butler, L. Ketzler, J. Slapcinsky, N.M. Stewart, J.H. Townsend, and L.D. Wilson; original field number JHT 2420; GenBank accession numbers GU971733 (16S), GU971734 (cyt b ), JN377401 (COI) Referred specimens. 15; ANSP 28981 85, 28200, Montaa Macuzal S of Pueblo Viejo; USNM 339700 08, from the eastern slope of Pico Pijol (15.175N, 87.558W), 1,920 m elevation, Parque Nacional Pico Pijol, Departamento de Yoro, Honduras. Distribution and n atural history I consider this taxon to be restricted to the Sierra de Sulaco in southwestern Departamento de Yoro, where it is known to occur in the vicinity of the type locality in an unprotected patch of remant cloud forest on the top of Montaa Macuza l (maximum elevation approximately 1,945 m) at the eastern

PAGE 283

283 Figure 5 6 Nototriton barbouri sensu stricto. A and B ) Genetype of Nototriton barbouri sensu stricto ( UF 156538), from Monta a Macuzal, Departamento de Yoro, Honduras Photos J.H. Townsend. t erminus of the range and in the cloud forests of PN Pico Pijol (maximum elevation approximately 2,282 m) at the western end of the range. Nototriton barbouri is known to occur in the Lower Montane Wet Forest formation from 1,520 1,920 m elevation Conserva tion status. Endangered B1ab(iii).

PAGE 284

284 Description of unassigned populations from the Cordillera Nombre de Dios Restriction of the taxon Nototriton barbouri to populations inhabiting the highlands of southern Yoro (Montaa de Pijol and Montaa de Macuzal) leav es two allopatric populations from the Cordillera Nombre de Dios without assignment to an existing taxon. With no referable name present in the synonymy of N. barbouri for these populations, putatively endemic to highland forests at opposite ends of the Co rdillera Nombre de Dios in Parque Nacional Pico Bonito and Refugio de Vida Silvestre Texiguat ( Figure 5 1 ), I herein describe these two cryptic species as new taxa. Together, they form a clade with N. brodiei a species residing in the Sierra de Omoa and C ordillera de Merendn in northwestern Honduras and adjacent Guatemala ( Figure 5 1 ). Nototriton sp. A sp. nov. Nototriton barbouri : McCranie 1996 a : 28 Holotype. USNM 497552, an adult female from the south slope of Cerro Bfalo ( 15.66 N, 86.79 W ), 1 540 m elevation, Parque Nacional Pico Bonito, Departamento de Atlntida, Honduras; collected 30 May 1996 by S. Gotte and J.R. McCranie; original field number LDW 10724; GenBank accession number AF199137 (cyt b ). Paratype. USNM 339712, an adult female from Quebra da de Oro (15.64 N 86.80 W), 1 210 m elevation, Parque Nacional Pico Bonito, Departamento de Atlntida, Honduras; collected 13 February 1995 by J.R. McCranie & J.C. Rindfleish; original field number LDW 10378; GenBank accession numbers AF199201 (16S), AF1 99136 (cyt b ). Measurements of holotype (in mm). SL 33.7 ; AG 19.5 ; TW 4 .9; HL 6.0; HW 4.1 ; TL 39.4 ; HLL 6.4 ; FLL 6 .0; CLL 12.4 ; FFW 1.0; HFW 1.9 ; NL 0.3 ; eye length 1.7; eye width 1.2 ; interorbital distance 1. 3 ; anterior rim of orbit to snout 1. 5 ; distance separating internal nares 1.1; distance separating external nares 1. 6

PAGE 285

285 Distribution and natural h istory This species is known only from the vicinity of Cerro Bfalo in Parque Nacional Pico Bonito, Honduras, 1,210 1,540 m elevation. Conservation status Critically Endangered B1ab(iii). Nototriton sp. B, sp. nov. Holotype. USNM 578300, an adult male from Cerro El Chino above La Liberacin (15.525394N, 87.278672W), 1 420 m elevation, Refugio de Vida Silvestre Texiguat, Departamento de Atlntida, Honduras ; collected 19 June 2010 by E. Aguilar, B.K. Atkinson, C.A. Cerrato M., A. Contreras, A. Portillo, J.H. Townsend, and L.D. Wilson; original field number JHT 3159; GenBank accession numbers JN377387 (16S), JN377391 (cyt b ), JN377403 (COI). Paratypes. USNM 339709 10, about 2.5 km (airline) NNE of La Fortuna Refugio de Vida Silvestre Texiguat, Departamento de Yoro, 1 690 m elevation; USNM 339711, about 2.5 km (airline) NNE of La Fortuna Refugio de Vida Silvestre Texiguat, Departamento de Yoro, 1 800 m eleva tion; USNM 509333, about 2.5 km (airline) NNE of La Fortuna Refugio de Vida Silvestre Texiguat, Departamento de Yoro, 1 600 m elevation. Measurements of holotype (in mm). SL 31.9; AG 19.1; TW 3.8; HL 6.9; HW 4.2; TL 41.0; HLL 7.2; FLL 6.4; CLL 13.6; FFW 1.1; HFW 1.5; NL 0.2; eye length 1.8; eye width 0.9; interorbital distance 1.6; anterior rim of orbit to snout 1.5; distance separating internal nares 1.4; distance separating external nares 1.8; tip of snout to axilla 9.6. Distribution and n atural histo ry. This species is known only from highland forest within Refugio de Vida Silvestre Texguat, 1,420 1,800 m. The holotype was collected as it crawled out of an arboreal bromeliad approximately 1.5 m above the ground on a rainy night.

PAGE 286

286 Figure 5 7 Notot riton sp. B. A) Holotype of Nototriton sp. B (USNM 578300) in life B) X ray radiograph of the holotype of Nototriton sp. B. C) Immediate vicinity of the type locality of Nototriton sp. B, Cerro El Chino, 1,420 m, above La Liberacin in Refugio de Vida Silv estre Texguat. Photos J.H. Townsend. Conservation status. Critically Endangered B1ab(iii)+2ab(iii). Review of the Remaining Species of Nototriton from the Chorts Highlands The additions of Nototriton picucha N. tomamorum N. sp. A N. sp. B and N. sp C (identified in Chapter 4 and being described elsewhere) bring s the total number of species in the genus to 18 with 10 of these species being endemic to the Chorts

PAGE 287

287 Highlands This is a more than 27 % increase in the number of species in this cryptozoic genus and a nearly two fold increase in the number of species known from the Chorts Highlands Nototriton brodiei Campbell & Smith 1998 Holotype. UTA A 50000, an adult female from Cerro Pozo de Agua, 1,125 m elevation, Sierra de Caral, Municipi o de Mora les, Izabal, Guatemala. Distribution and Natural History. Premontane forests of the Cordillera de Merendn along the Guatemala/Honduras border, from 875 1,140 m elevation. This species is known from the vicinity of the type locality in the Sierra de Cara l, Guatemala, and the northwestern slope of the Sierra de Omo a, Honduras (Campbell & Smith 1998; Kolby et al. 2009). Conservation Status. Critically Endangered B1ab(iii) (Acevedo et al. 2008); the discovery of this species in a relatively well protected a rea of Honduras should necessitate re Remarks. This species was recently reported from the Sierra de Omoa in Honduras ( Kolby et al. 2009 ). Referred Specimens. GUATEMALA: IZABAL: Sierra de Caral, UTA A 50001 UTA A 5 1490. HONDURAS: CORTS: Parque Nacional Cusuco, MVZ 258034 36. Nototriton lignicola McCranie & Wilson 1997 Nototriton barbouri 2000: 102 Holotype. USNM 497539, an adult male from above the Monte Escondido, 1,780 m elevation, Ce rro de Enmedio (1506'N, 8644'W), Parque Nacional La Muralla, Olancho, Honduras.

PAGE 288

288 Figure 5 8 Nototriton lignicola A and B ) N lignicola ( UF 156543), from Cataguana, 2,020 m, Parque Nacional Monta a de Yoro, Departamento de Francisco Morazn, Honduras. C) Cloud forest habitat at Cataguana. D) Juvenile N. lignicola ( UF 156543) from Cataguana, 1,820 m, Parque Nacional Monta a de Yoro, Departamento de Francisco Morazn, Honduras. Photos J.H. Townsend. Distribution and Natural History. Lower Montane Wet Forest of northern Departamento de Francisco Morazn and northwestern Departamento de Olancho, Honduras, from 1,780 2,020 m elevation. Conservation Status. Critically Endangered B1ab(iii). Referred Specimens. HONDURAS: FRANCISCO MORAZN: Cataguana, UF 156 542 44, OLANCHO: Cerro de Enmedio, USNM 497539 51, 509335 (cleared and stained).

PAGE 289

289 Figure 5 9 A ) Nototriton limnospectator ( UF 156539 ), from above Ro Varsovia, 1,640 m, Parque Nacional Cerro Azul Membar, Depa rtamento de Comayagua, Honduras (Photo J.H. Townsend). B) Juvenile N. limnospectator ( IRL 035) from Cataguana, from above Los Pinos, 900 m, Parque Nacional Cerro Azul Membar, Departamento de Corts, Honduras (Photo I. Luque) Nototriton limnospectator McCranie, Wilson, & Polisar 1998 Holotype. UF 98460, an adult female from southwest of San Lus de los Planes (1456'N, 8808'W), 1910 m elevation, northwestern side of Montaa de Santa Brbara, Parque Nacional Montaa de Santa Brbara, Santa Brbara, Honduras. Distribution and Natural History. T his species is found in Premontane Wet Forest Lower Montane Wet Forest and Montane Wet Forest on both sides of Lago de Yojoa in central Honduras. Conservation Status. Endangered B1ab(iii)

PAGE 290

290 Referred Specimens COMAYAGUA: UF 156539 41, IRL 035 CORTS: El Volcn, AMNH 54949. SANTA BARBARA: El Ocotillo, MVZ 225866, USNM 509334 (CS); Montaa de Santa Brbara, SW of San Lus de Los Planes, UF 98460 66. Nototriton stuarti Wake & Campbell 2000 Holotype. UTA A 33686, an adult male from 11.6 road km west southwes t of Puerto Santo Toms, 744 m elevation, Montaas del Mico (1538'N, 8840'W), Izabal, Guatemala. Distribution and Natural History. This species is known only from a single specimen from p remontane forest in the Montaas del Mico in eastern Guatemala. Conservation Status. Data Deficient (IUCN 2011)

PAGE 291

291 Table 5 1 Samples used in sequence divergence and phylogenetic analyses, with GenBank accession and museum voucher numbers; CR = Costa Rica, GT = Guatemala, HN = Honduras, NI = Nicaragua Taxon Locality GenBank voucher GenBank Accession Numbers 16S cytb COI N. barbouri HN: Yoro: Macuzal UF156538 GU971733 GU971734 JN377401 N. brodiei (1) GT: Izabal: Sierra de Caral UTA A 51490 AF199202 AF199139 JN377402 N. brodiei (2) HN: Corts: Cusuco MVZ258035 JN377384 N. lignicola (1) HN: Olancho: La Mura lla USNM497540 AF199204 AF199141 N. lignicola (2) HN: Olancho: La Mura lla USNM497550 AF199142 N. lignicola (3) HN: Francisco Morazn: Cataguana UF156543 GU971735 GU971736 JN377408 N. limnospectato r (1) HN: Santa Brbara: El Ocotillo MVZ225866 AF199143 N. limnospectator (2) HN: Santa Brbara: San Lus Planes MVZ263852 JN377383 N. limnospectator (3) HN: Comayagua: Azul Membar UF156539 GU971737 GU971738 JN377397 N. limnospectator (4) HN: Comayagua: Azul Membar UF156540 GU971739 GU971740 JN377398 N. limnospectator (5) HN: Comayagua: Azul Membar UF156541 JN377386 JN377396 JN377399 N. limnospectator (6) HN: Corts: Azul Membar UF IRL035 JN377385 JN377395 JN377400 N. picucha sp. nov. (1) HN: Olancho: Sierra de Agalta USNM 578299 JN377388 JN377392 JN377404 N. picucha sp. nov. (2) HN: Olancho: Sierra de Agalta USNM 578298 JN377389 JN377393 JN377405 N. sp. A (1) HN: Atlntida: Quebrada de Oro USNM339712 AF199201 AF199136 N. sp. A (2) HN: Atlntida: Cerro Bfalo USNM497552 AF199137 N. sp. B (1) HN: Atlntida: Texguat USNM 578300 JN377387 JN377391 JN377403 N. sp. B (2) HN: Yoro: Texguat USNM509333 AF199138 N. tomamorum HN: Yoro: Texguat UF155377 GU971731 GU971732 JN377407 N. s aslaya (1) NI: Atlantco Norte: Saslaya MVZ230241 GU981761 N. saslaya (2) NI: Atlantco Norte: Saslaya UF156352 JN377390 JN377394 JN377406

PAGE 292

292 Table 5 2 Models of nucleotide substitution chosen for phylogenetic analyses of Chorts Highland taxa usin g Akaike Information Criterion values Partition Model AIC lnL 16S GTR+G 2335.9224 1126.9612 cyt b (1 st ) HKY+G 1380.1242 651.0621 cyt b (2 nd ) HKY+G 1011.5928 466.7964 cyt b (3 rd ) GTR+G 2527.2855 1220.6427 COI (1 st ) GTR+G 2603.4728 1270.7364 COI (2 nd ) K80+I 1010.9648 481.4824 COI (3 rd ) HKY 641.4734 294.7367 Table 5 3 Within and between species sequence divergence (uncorrected p distance) for Chorts Highland moss salamanders Intraspecific Interspecific Taxon 16S cyt b COI 16S cyt b COI N. barbouri 0.012 0.055 0.071 0.123 0.080 0.121 N. sp. A 0.000 0.014 0.059 0.052 0.142 N. sp. B 0.014 0.018 0.057 0.052 0.144 0.061 0.126 N. brodiei 0.000 0.014 0.061 0.057 0.123 0.061 0.133 N. lignicola 0.002 0.005 0.023 0.064 0. 082 0.120 0.105 0.139 N. limnospectator 0.000 0.006 0.000 0.019 0.002 0.003 0.012 0.059 0.071 0.147 0.080 0.126 N. picucha 0.000 0.008 0.002 0.012 0.049 0.093 0.123 0.085 0.121 N. saslaya 0.006 0.047 0.064 0.104 0.147 0.107 0.136 N. tomamorum 0.035 0.057 0.095 0.131 0.115 0.139

PAGE 293

293 Table 5 4 Morphological and morphometric comparison of species of Nototriton ; see Material and Methods section for explanation of abbreviations; SVL is given in mm C omparative morphological data for species other than N. picucha are from Campbell & Smith (1998), McCranie et al. (1998), Wake & Campbell (2000) Khler (2002), McCranie & W ilson (2002), and Townsend et al. (2010) Taxon SL HW/SL TL/SL HLL/SL FLL/SL HFW/SL NL/SL MT VT N. barbouri 30.2 39.9 0.104 0.132 1.031 1.398 0.153 0.200 0.142 0.174 0.037 0.060 0.005 0.011 41 54 12 23 N. brodiei 33.2 34.5 0.120 1.420 1.440 0.166 0.180 0.148 0.151 0.040 0.060 0.004 0.005 60 62 23 24 N. lignicola 28.3 33.9 0.103 0.118 0.840 1.059 0.158 0.181 0.137 0.160 0.029 0.040 0.006 0.009 46 54 16 24 N. limnospectator 33.0 38.2 0.095 0.118 1.027 1.297 0.164 0.211 0.156 0.183 0.048 0.061 0.003 42 55 16 26 N. picucha 25.7 27.9 0.140 0.148 1.143 0.204 0.218 0.179 0.191 0.042 0.043 0.007 0.008 41 16 19 N. saslaya 28.1 34.6 0.133 0.155 0.883 1.255 0.217 0.244 0.194 0.210 0.075 0.091 0.002 0.003 17 22 3 11 N. stuarti 32.6 0.138 1.264 0.178 0.172 0.049 0.012 36 20 N. tomamorum 26.9 0.145 0.911 0.197 0.160 0.037 0.018 26 11

PAGE 294

294 CHAPTER 6 INTEGRATING RESEARCH, EDUCATION, AND OUTREACH IN SUPPORT OF CONSERVATION IN THE CHORTS HIGHLANDS This dissertation represents the first step in developing an accurate picture of evolutionary diversification and phylogenetic distinctiveness for the Chorts Block. The known to have a high degree of endemism and to be facing a comp arably high risk of extinction, is herein demonstrated to be greatly underestimated in terms of taxonomic diversity. Throughout my research in Honduras and Nicaragua I encountered a series of significant challenges to the continued documentation of Chorts Block biodiversity, and more importantly to the conservation of the ecosystems that support Chorts Block biodiversity First, outside of the herpetofaunal research that has taken place since 1967 (Townsend & Wilson 2010), there has been no concerted effort to inventory and document other major biotic components (e.g. mammals, birds, plants) using modern, molecular based approach es (but see Matamoros et al. [2009] for details of efforts in fre shwater ichthyology) The majority of endemic herpetofaunal species in the Chorts Block are found in the h ighlands owing to the extreme ecophysiographic heterogeneity in its 50+ isolated highland forests and interceding dry valleys. Expl oration of these geographically limited highly threatened mountain top forests continues apace among the best known groups, as evidenced by the continued regular discovery of new amphibians (Chapte r 4) and reptiles (Chapter 3). However, integrative taxonomy inventories ar e desp e rately needed for other taxa. Second, opportunities for recruitment, education, and practical training of aspiring Honduran systemati sts and conservation biologists, those representing the next generation of in the effort to d ocument and conserve their national

PAGE 295

295 biodiversity resources are increasingly rare The countries of the Chorts Block are among the poorest and most underdeveloped in the Americas, and opportunities for higher education in systematics and related fields ar e limited to a few small, under resourced programs in public universities Finally, regional awareness of the unique nature of the endemic Chorts Block biota is low. To date, little concerted outreach has been targeted at the largest group of stakeholder s in regional conservation: the public. My experience has been that Hondura n s generally are interested in and supportive of conservation of their n ational environmental patrimony. H owever lacking access to information about endemic biodiversity limits the ability of individuals and groups to lead or even support conservation efforts. Below I outline my vision for promoting biodiversity conservation through education, training, and outreach opportunities generated through a strategic program to inventory the biodiversity of the Chorts Block. Given that it contains the majority of the endemism rich Chorts Highlands and that geopolitical considerations warrant an initial focus at the national level, I am presenting this strategy as I would see it implemen ted in Honduras Taxonomic Inventor ies in Promotion of Education and Extension As I became aware of these three principal obstacles to biodiversity conservation in the Chorts Block, I sought avenues to address these issues through my own research initiat ives. B iodiversity inventory and systematics research address the first challenge, cataloging the biolo gical diversity of the region; however activities associated with carrying out taxonomic inventory work also offer an excellent opportunity to provide ed ucation and training to multiple groups My experience in Honduras has

PAGE 296

296 demonstrated that a virtually untapped talent pool of u niversity students, parataxonomists, park guards, and conservation practitioners are available for participation in all levels of inventory related research. Taxonomic inventory projects also present frequent opportun it ies for scientists to engage with the public across a variety of platforms. In my case, I have had frequent opportunities to engage the Honduran public through the me dia, promoting local and regional biodiversity awareness and endemic species conservation through newspaper articles and television interviews ( Figure 6 1). I was able to provide both planned and impromptu presentations to municipal governmental leaders, c ommunity groups schools and residents in rural areas, often within the buffer zones of the highland protected areas where I was working ( Figure 6 2). Activities such as these should be ct, and field based systematists have the unique opportunity to simultaneously carry out research, engage the public, and contribute to the development of Honduran capacity in systematic biology. As evidenced in this dissertation, Honduras is a country of remarkable herpetofaunal diversity, having the highest degree of amphibian endemism (i.e., all Central American countries: 36.2%, compared to 27.1% for Guatemala and 26.6% for Costa Rica (Wilson & Johnson 2010). With the discovery of more than 30 potentially new species (Chapter 4), it appears that this high degree of endemism is actually greatly underestimated, and numerous undiscovered species undoubtedly await disc overy across Honduras.

PAGE 297

297 Figure 6 1. Print media coverage of the discovery of new endemic species during 2008 and 2009. As I have emphasized throughout this dissertation, m uch of the diversity is attributed to the high number of isolated highland forest s that are home to suites of endemic species Across Honduras, more than a dozen of these cloud forest remain virtually unexplored biologically, and likely conceal veritable treasure troves of new species from a variety of taxonomic groups As a consequence of the unique biogeography of the Chorts Block Honduras remains a largely unrealized center for biodiversity in the Americas, overshadowed by more thoroughly studied Central American countries.

PAGE 298

298 Opportunities for Training and Education Curre ntly, the School of Biology at the National Autonomous University of Honduras ( Universidad Nacional Autnoma de Honduras or UNAH) offers only a single degree, the Licenciatura en Biologa a degree more comprehensive than a traditional Bachelor of Science degree but lacking the professional preparation and opportunitie s for research typical of a Master of Science program (in 2011 12 UNAH has plans to launch an Master of Science in Tropical Botany). Biology majors complete a capstone research design course (BI 025 Seminario de Investigacin ) as their final class, after which they must complete an 800 hour professional practicum which can take on a wide variety of forms. The primary focus of BI 025 is for students to develop a research proposal, and some high ly motivated students develop proposals to carry out under direction of a supervising scientist in fulfillment of their 800 hour requirement. Despite potential synergies between the requirements of BI 025 and the professional practicum, UNAH faculty do not typically encourage students to attempt linking the two and it is exceedingly challenging for students to propose and find support for independent, or even faculty directed, research projects. As a result of my time spent in Honduras, I have had the oppo rtunity to work closely with numerous UNAH biology students relying on them in innumerable ways while promoting their professional development through participation in remote b iodiversity surveys. In further support of a few dedicated students, I served i n the supervisory role for their final year professional practica, assisting in developing proposals for and carrying out rapid herpetological inventories that, among other things, led directly to the discovery of a new species of salamander (Townsend et a l. 2011a). I see p roviding opportunities of this nature for UNAH students to gain practical

PAGE 299

299 experience is critical to strengthening the national biological community and the promotion of UNAH student participation also serves to enhance the Honduran publ perception of biodiversity conservation and its practitioners The excitement and pursuit of scientific discovery, centered on the collection, identification, and further study of new and poorly known species, was and will continue to be the central t heme to my efforts to give students the opportunity to be fully involved in leading edge systematic research with real world applications in the realm of Honduran biodiversity exploration and conservation Pilot Project: Parque Nacional Montaa de Yoro O ne of my principal goals in taking an approach that combines biodiversity inventory work and integrative systematics is to refine our understanding of the patterns of diversification exhibited in the Chorts Highlands. My initiation of inventory work in Pa rque Nacional Montaa de Yoro (Chapter 3) provided an opportunity to do so, and at the same time develop and practice the educational and outreach related elements described above. Parque Nacional Montaa de Yoro was virtually unknown to biologists prior t o my initial trip there in 2006. As a result of limited baseline biodiversity data, this large area (~50km 2 ) of cloud forest was among the lowest priorities for national level conservation planning and at one point was recommended for a reduction in its pr otected area status (Vreugdenhil et al. 2002 ; COHECO 2003 ) During a single rapid inventory in 2006, we found new species of salamander (Townsend et al. 2009 a ), anole lizard (Townsend & Wilson 2009) and spikethumb treefrog ( Plectrohyla cf. guatemalensis ; Chapter 4) with all three qualif y as Critically Endangered based on IUCN Red List criteria due to severe threat to their limited distributions In contrast, we also found two species of salamanders previously considered to be endemic to Parque

PAGE 300

300 Figure 6 2 Examples of public outreach and dissemination of results from taxonomic inventories. A) Television interview about the endemic biodiversity in Honduras (Photo James Austin) B) Presenting the results of work in La Liberacin de Texguat to the Alcalde (mayor) of the municipality that receives its water from the reserve (Photo Ileana Luque). C) Community members attending a public presentation of results from La Liberacin de Texguat (Photo J.H. Townsend) D) Ileana Luque and I presenting educationa l materials on endemic species to children in Jilamito Nuevo, in the buffer zone of Refugio de Vida Silvestre Texguat (Photo Yensi Flores)

PAGE 301

301

PAGE 302

302 Nacional La Muralla ( Nototriton lignicola and Oedipina kasios ) in Parque Nacional Montaa de Yoro (Chapter 4; Townsend et al. 2011b). Three expeditions were carried out to Parque Nacional Montaa de Yoro in 2006, 2007, and 2008, each in close coordination with national (Honduran National Department of Protected Areas and Wildlife [ICF]), municipal (Municipality o f Marale, collaborating with the alcalde (mayor) and the office of rural development), park administrative (logistics arranged in close collaboration the director of the park), and community management (were accompanied by the local guardabosque or park r anger and work e d out of local homes ) levels. The guardabosque s were not simply viewed as local escorts into the park, rather a deliberate attempt was made to engage them in an active interchange of ecological knowledge, sharing scientific knowledge about biodiversity in exchange for locally derived traditional knowledge of the cloud forest and its inhabitants. In addition to the participation by guardabosques expeditions into Montaa de Yoro included (along with myself and Dr. Larry David Wilson), two gr aduate students and one undergraduate from North American universities, two biology students from the National Autonomous University of Honduras (UNAH), and two Peace Corps volunteers. None of the participants had previous experience in remote field based research in the tropics, but all were able to fully participate and gain experience in a variety of methods, ranging from expedition planning and wilderness survival to preparation of tissue samples and specimens. Following the discoveries in Parque Naci onal Montaa de Yoro a priority became the public dissemination of information about this virtually unknown area. Scientific

PAGE 303

303 results were published in peer reviewed outlets as is standard practice in the field. In addition, I wrote (with Ileana Luque Mont es) a new s paper article that, with the assistance of the Zamorano Biodiversity Center, received front page coverage in the two largest Honduran newspapers ( Figure 6 1). Additional undiscovered species almost certainly occur in this reserve, but intensive s urvey work is needed immediately to forests. Concluding Statement In this dissertation, I have sought to usher in a new era of biodiversity research in the Chorts Block region of Central America. By utilizing an integrat ive framework built around biodiversity inventory and molecular systematics, I have presented a viable model for addressing critical unmet goals for biodiversity conservation through the taxonomic and mo lecular inventory of an endemism hotspot, which simultaneously promotes training for the next generation of Honduran (and Honduras focused) biologists and raising public awareness of endemic rich areas and related conservation issues through extension and outreach. I am anticipating the opportunity to fully pursue implementation of this model with even greater intensity and at a broader scale in the coming years, with the sincere hope that this initiative will catalyze efforts to bring the Chorts Block bio diversity hotspot to the forefront of regional and global conservation diversification.

PAGE 304

304 APPENDIX TAXONOMIC REVIEW OF CAUDATA FROM THE CHO RTS BLOCK Below I present a review of the salamander taxa known t o occur with the Chorts summarized in orde r to provide the context for newly generated data. Dwarf salamanders (genus Cryptotriton Garca Pars & Wake 2000) The genus Cryptotriton contains the six species formerly referred to as the Nototriton nasalis group (sensu Papenfuss & Wake 1987), but reco gnized by Garca Pars & Wake (2000) to represent a cryptic clade distinct from Nototriton An undescribed seventh species was identified from the Sierra de Las Minas in Guatemala by Garca Pars & Wake (2000). Three putative species of Cryptotriton are re ported to occur in the Chorts Highlands; however the status of C. wakei is disputed, with McCranie & Wilson (2002: 139) making the case that this species is a junior synonym of C. nasalis given a lack of morphological distinction between the two and the proximity of their known distributions. However, genetic material referable to C. wakei is lacking, and until the status of this taxon can be assessed phylogenetically its taxonomic validity will remain uncertain. Cryptotriton monzoni (Campbell & Smith 1998) Nototriton monzoni Campbell & Smith 1998: 6. Cryptotriton monzoni : Garca Pars & Wake 2000: 58. elevation (14

PAGE 305

305 Known distribution. Known only from a single sp ecimen from the type locality in easternmost Guatemala. Available genetic data. None. Cryptotriton nasalis (Dunn 1924) Oedipus nasalis Dunn 1924: 97. Chiropterotriton nasalis : Meyer & Wilson 1971: 7. Nototriton nasalis : Wake & Elias 1983: 11. Cryptotr iton nasalis : Garca Pars & Wake 2000: 58. Known distribution. Restricted to the Sierra de Omoa in northwestern Honduras, from 1,220 2,200 m elevation. Available genetic data. Cyt b (1 sample; Garca Pars & Wake 2000). Cryptotriton wakei (Campbell & Smith 1998) Nototriton wakei Campbell & Smith 1998: 5. Cryptotriton wakei : Garca Pars & Wake 2000: 58. e Caral, Municipio de Morales, Izabal, Guatemala, 1150 m elevation (15 Known distribution. Known only from a single specimen from the type locality in easternmost Guatemala, 1,150 m elevation. Available genetic data. None.

PAGE 306

306 Bromeliad salam anders (genus Dendrotriton Wake & Elias 1983) The genus Dendrotriton was proposed to accommodate the species of the Chiropterotriton bromeliacia group (sensu Lynch & Wake 1975) of Chiropterotriton beta (Wake & Elias 1983). Of the six species of Dendrotrito n the single Chorts Highlands species stands as a biogeographic outlier, with the five remaining species restricted to the highlands of western Guatemala and adjacent Chiapas, Mxico (Wake 1998). Bromeliad dwelling salamanders have been known from Cerro Santa Brbara, and isolated karstic mountain in central Honduras, since first collected by XXX in XXX. These salamanders were referred to as Chiropterotriton (and later Nototriton ) nasalis (Meyer & Wilson 1971; Wake & Elias 1983), a species otherwise known only from the Sierra de Omoa in northwestern Honduras, until being recognized as a distinct species genus Nototriton based on its previous assignment to N. nasalis a member o f the N. nasalis group (Papenfuss & Wake 1987; later referred to the genus Cryptotriton ; Garca Pars & Wake 2000); however, Wake (1998) placed the species in the genus Dendrotriton based on external and internal morphology. Dendrotriton sanctibarbarus ( Nototriton sanctibarbarus Dendrotriton sanctibarbarus : Wake 1998: 88. Departamento de Santa Brbara, Hon

PAGE 307

307 Known distribution. Restricted to Montaa de Santa Brbara, 1,800 2,744 m elevation. Available genetic data. None. Moss salamanders (genus Nototriton Wake & Elias 1983) The genus Nototriton included 13 species of miniature salamanders that are d isjunctly distributed across highland forests in Guatemala, Honduras, Nicaragua, and Costa Rica (AmphibiaWeb, 2 September 2011). This genus was one of two named to accommodate the species of Chiropterotriton beta (Wake & Elias 1983). Nototriton sensu Wake & Elias 1983 itself was later shown to be paraphyletic, resulting in description of the genus Cryptotriton by Garca Pars & Wake (2000). There are three species groups currently recognized within the genus Nototriton (Papenfuss & Wake 1987; Savage 2002): the N. barbouri group, the N. picadoi group, and the N. richardi group. These three groups exhibit two discrete patterns of geographic distribution. The N. barbouri group (five species) is restricted to the Chorts Highlands of Honduras and Guatemala, and the N. picadoi group (six species) and N. richardi group (two species) are found in the highlands of Costa Rica, with a single species in the N. picadoi group, N. saslaya endemic to a cloud forest in northern Nicaragua (Khler 2002). This genus is dealt w ith in detail in Chapters 4 and 5. Nototriton barbouri (Schmidt 1936) Oedipus barbouri Schmidt 1936: 43. ?Pseudoeurycea barbouri: Taylor 1944: 209.

PAGE 308

308 Chiropterotriton barbouri : Meyer 1969: 106. Nototriton barbouri : Wake & Elias 1983: 11. Type locality Known distribution. Populations assigned to this taxon are known from the vicinity of Montaa Macuzal and Pico Pijol in Yoro, from Tex guat along the southwestern border between Atlntida and Yoro, and from Pico Bonito in Atlntida; all localities 1,550 1,990 m elevation. A single specimen (AMNH 54949) from a locality near Lago de Yojoa at 860 m elevation is also assigned to this taxon. Available genetic data. 16S (1 sample; Garca Pars & Wake 2000), cyt b (3 samples; Garca Pars & Wake 2000). Nototriton brodiei Campbell & Smith 1998 Nototriton brodiei Campbell & Smith 1998: 3. de Caral, Municipio de Morales, Izabal, Guatemala, 1125 m elevation (15 Known distribution. This species is known only from the Sierra de Caral in easternmost G uatemala, 875 1,140 m elevation. Available genetic data. 16S (1 sample; Garca Pars & Wake 2000), cyt b (1 sample; Garca Pars & Wake 2000). Nototriton lignicola McCranie & Wilson 1997 Nototriton lignicola McCranie & Wilson 1997: 369.

PAGE 309

309 Nototriton barbouri 44'W) along the trail above the Monte Escondido campground, Parque Nacional La Muralla, 1780 m elev., Departamento de Known distribution. Known only from the vicinity of the type locality in the northwestern corner of Olancho in north c entral Honduras, 1,760 1,780 elevation. Available genetic data. 16S (1 sample; Garca Pars & Wake 2000), cyt b (2 samples; Garca Pars & Wake 2000). Nototriton limnospectator McCranie, Wilson, & Polisar 1998 Nototriton limnospectator McCranie et al. 199 8: 455. northwestern side of Montaa de Santa Brbara, southwest of San Lus de los Planes (1456'N, 8808'W), 1910 m elevation, Departamento de Santa Known distribution. Apparently restricted to Montaa de Santa B rbara, 1,640 1,980 m elevation. Available genetic data. Cyt b (1 sample; Garca Pars & Wake 2000). Nototriton saslaya Khler 2002 Nototriton saslaya Khler 2002: 205. elevation, Reg

PAGE 310

310 Known distribution. Known only from the vicinity of the type locality in north central Nicaragua, 1,280 1,370 m elevation. Available genetic data. None. Nototriton stuarti Wake & Campbell 2000 Nototriton stuarti Wake & Campbell 2000: 817. Known distribution. Known only from a single specimen from the type locality in easternmost Guate mala. Available genetic data. None. Worm salamanders (genus Oedipina Keferstein 1868) The taxon Oedipina accommodates some of the mostly morphologically divergent Neotropical salamanders, a monophyletic clade that is sister to Nototriton (Garca Pars & W ake 2000). These elongate, attenuate species are the only Neotropical salamanders to have lengthened their bodies as a result of an increased number of vertebrae, and along with having greatly reduced limbs and elongate tails (which can be more than two ti mes the body length in many species) are well adapted for life inside root masses, leaf litter, rotten logs, and other fossorial microhabitats (Brame 1968). Oedipina represents an excellent example of a non adaptive radiation (highly conserved morphology a cross three distinct clades) that is adaptive at the clade level (the drastic change in body form in order to exploit fossorial microhabitats). Eleven species of

PAGE 311

311 Oedipina are known from the Chorts Highlands, representing all three subgenera: Oedipina Oed itriton and Oedopinola (Garca Pars & Wake 2000, McCranie et al. 2008). Subgenus Oedipina This subgenus is represented by six putative species in the Chorts Highlands, two of which have distributions that extend outside the region, leaving the remaini ng four as endemics. Oedipina cyclocauda Taylor 1952 Oedipina cyclocauda Taylor 1952: 764. Known distribution. Caribbean lowlands of Costa Rica and southern Nicaragua, 60 500 m ele vation. Available genetic data. 16S (2 samples; Garca Pars & Wake 2000), cyt b (2 samples; Garca Pars & Wake 2000). Oedipina ignea Stuart 1952 Oedipina ignea Stuart 1952: 1. t of Known distribution. Pacific versant of south central Guatemala to southwestern Honduras, 1,340 1,750 m elevation.

PAGE 312

312 O. sp. Pars & Wake 2000) cyt b (1 sample; Garca Pars & Wake 2000). Oedipina leptopoda McCranie, Vieites, & Wake 2008 Oedipina cyclocauda (in part): Brame 1968: 30. Oedipina leptopoda McCranie et al. 2008: 13. 480 N, Known distribution. Known from three localities in Departamento de Yoro, Honduras, 700 1,300 m elevation. O. sp. b (1 sample; Garca Pars & Wake 2000). Oedipina pseudouniform is Brame 1968 Oedipina uniformis (in part): Taylor 1952: 350. Oedipina pseudouniformis Brame 1968: 25. Juan Vias and 6.3 km by road west of Turrialba, Canton de Turrialba, Provincia de Known distribution. The type series includes a series of eight specimens from Hacienda La Cumplida, Matagalpa, Nicaragua, 731 m elevation, and is otherwise known from 19 1,253 m elevation in no rthern and central Costa Rica. Available genetic data. 16S (1 sample; Garca Pars & Wake 2000), cyt b (3 samples; Garca Pars & Wake 2000).

PAGE 313

313 Oedipina stuarti Brame 1968 Oedipina stuarti Brame 1968: 47. e Fonseca, Departamento de Known distribution. This enigmatic taxon is known from three specimens, two from Isla El Tigre in the Pacific Ocean, and the third reportedly from Tegucigalpa, 975 m elevation. Available genetic data. None. Oedipina taylori Stuart 1952 Oedipina taylori Stuart 1952: 2. line kilometers southeast of Chiquimulilla, Department of Jutiapa, Guatemala. Elevation, about 100 inal parenthesis supposed to be placed?] Known distribution. Pacific versant of southeastern Guatemala, El Salvador, and Honduras, 140 1,140 m elevation. This species also is known from an isolated locality in the upper Ro Motagua Valley in eastern Guatem ala. Available genetic data. None. Subgenus Oeditriton Oedipina kasios McCranie, Vieites, & Wake 2008 Oedipina cyclocauda : Espinal et al. 2001: 103. Oedipina kasios McCranie et al. 2008: 11.

PAGE 314

314 Nacional La Known distribution. Known only from the vicinity of the type locality in the northwestern corner of Olancho in north central Honduras, 950 1,780 m elevation. Available genetic data. 16S (2 sam ples; McCranie et al. 2008), cyt b (2 samples; McCranie et al. 2008). Oedipina quadra McCranie, Vieites, & Wake 2008 Oedipina cyclocauda (in part): Brame 1968: 30. Oedipina quadra McCranie et al. 2008: 6. Known distribution. Considered relatively widespread in the lowlands and peripheral areas in north central and eastern Honduras, 70 540 m elev ation. Available genetic data. 16S (1 sample; McCranie et al. 2008), cyt b (1 sample; McCranie et al. 2008). Subgenus Oedopinola Oedipina elongata (Schmidt 1936) Oedipus elongatus Schmidt 1936: 165. Oedipina elongata : Taylor 1944: 226. Type locality.

PAGE 315

315 Known distribution. Caribbean versant from near the Isthmus of Tehuantepec, Mxico to northwestern Honduras, 10 770 m elevation. Available genetic data. 16S (1 sample; Garca Pars & Wake 2000), cyt b (1 sample; Garca Pars & Wake 2000). Oedipina gephyra McCranie, Wilson, & Williams 1993 Oedipina gephyra McCranie et al. 1993: 385. Cordillera Nom Known distribution. Known from the vicinity of the type locality around Cerro Texguat at the western end of the Cordillera Nombre de Dios and from Cerro Bfalo in the central portion of the same range in north ern Honduras, 1,580 1,810 m elevation. Available genetic data. 16S (2 samples; Garca Pars & Wake 2000), cyt b (2 samples; Garca Pars & Wake 2000). Oedipina tomasi McCranie 2006 Oedipina sp.: Townsend et al. 2006: 31. Oedipina tomasi McCranie 2006: 291. Known distribution. Known only from the vicinity of the type locality in the Sierra de Omoa in northwestern Ho nduras, 1,780 1,800 m elevation. Available genetic data. None.

PAGE 316

316 Mushroom tongued salamanders (genus Bolitoglossa Dumril, Bibron, & Dumril 1854) The most species rich genus of salamanders at 117 described species and counting (AmphibiaWeb, 2 September 20 11), Bolitoglossa also has the broadest distribution of any salamander genus, ranging from San Luis Potos in central Mxico south through Middle America and into South America to the Amazon Basin and Bolivian highlands (Parra Olea et al. 2004). There are 15 species of Bolitoglossa in the Chorts Highlands, which are spread across four phylogenetically delimited subgenera (after Parra Olea et al. 2004): Bolitoglossa Magnadigita Nanotriton, and Pachymandra The subgenera Bolitoglossa and Nanotriton both ar e found generally in the lowlands and peripherally in the highlands, and do not contain any species heretofore considered endemic to the Chorts Highlands. Pachymandra is represented by a single, giant species in the Chorts Highlands, B. dofleini which i nhabits premontane elevations and peripheral areas. Magnadigita represents a diverse highland restricted radiation that is largely endemic to the Chorts Highlands and includes the morphologically defined B. dunni species group (McCranie & Wilson 1993). Subgenus Bolitoglossa Dumril, Bibron, & Dumril 1854 This diverse and widespread subgenus includes two species in the Chorts Highlands, both of which are found in the lowlands formations peripheral to the highlands of the serrana with one species, B. m exicana widely occurring at premontane elevations.

PAGE 317

317 Bolitoglossa mexicana Dumril, Bibron, & Dumril 1854 Bolitoglossa mexicana Dumril, Bibron, & Dumril 1854: 93. Known distribution. Atlantic versant from southern Verac ruz, Mxico to eastern Honduras, near sea level to 1,400 m elevation. Available genetic data. 16S (17 samples; Garca Pars et al. 2000a ), cyt b (6 samples; Garca Pars et al. 2000a ). Bolitoglossa striatula (Noble 1918) Oedipus striatulus Noble 1918: 344. Bolitoglossa striatula : Taylor 1941: 147. Known distribution. Atlantic versant from eastern Honduras to central Costa Rica, near sea level to 1,050 m elevation. Available genetic data. 16S (1 sample; Garc a Pars et al. 2000a ), cyt b (1 sample; Garca Pars et al. 2000a ). Subgenus Magnadigita Taylor 1944 This subgenus includes the constituent members of the B. dunni species group, the B. franklini species group, and the B. rostrata species group (Parra Ol ea et al. 2004). Only the B. dunni group is found in the region, with 11 putative species endemic to the Chorts Highlands.

PAGE 318

318 Bolitoglossa carri McCranie & Wilson 1993 Bolitoglossa carri McCranie & Wilson 1993: 9. B. (Magnadigita) carri : Parra Olea et al. 2004: 336. Known distribution. Known only from the vicinity of the type locality in the mountains west of Tegucigalpa, H onduras, 1,840 2,070 m elevation. Available genetic data. 16S (2 samples; Parra Olea et al. 2004), cyt b (2 samples; Parra Olea et al. 2004). Bolitoglossa celaque McCranie & Wilson 1993 Bolitoglossa dunni (in part): Hidalgo 1983: 6. Bolitoglossa celaqu e McCranie & Wilson 1993: 11. B. (Magnadigita) celaque : Parra Olea et al. 2004: 336. Kno wn distribution. Highland forests across the departments of Lempira, Intibuc, and La Paz in southwestern Honduras, 1,900 2,620 m elevation. Available genetic data. 16S (2 samples; Parra Olea et al. 2004), cyt b (2 samples; Parra Olea et al. 2004). Boli toglossa conanti McCranie & Wilson 1993 Oedipus morio (in part): Dunn 1926: 387.

PAGE 319

319 Oedipus dunni (in part). Schmidt 1933: 16. Magnadigita dunni (in part): Taylor 1944: 218. Bolitoglossa dunni (in part): Wake & Brame 1963: 386. Bolitoglossa conanti McCrani e & Wilson 1993: 4. B. (Magnadigita) conanti : Parra Olea et al. 2004: 336. Type locality. Known distribution. Several isolated localities in western Honduras along th e Guatemalan and Salvadoran borders, 1,370 2,000 m elevation. Available genetic data. 16S (1 sample; Parra Olea et al. 2004), cyt b (1 sample; Parra Olea et al. 2004). Bolitoglossa decora McCranie & Wilson 1997 Bolitoglossa decora McCranie & Wilson 1997: 367. B. (Magnadigita) decora : Parra Olea et al. 2004: 336. Escondido campground (1505'N, 8644'W), Parque Nacional La Muralla, 1440 m elev., Departamento de Olancho, Hon Known distribution. Known only from the vicinity of the type locality in the northwestern corner of Olancho in north central Honduras, 1,430 1,550 m elevation. Available genetic data. 16S (1 sample; Parra Olea et al. 2004), cyt b (1 sample; Parra O lea et al. 2004).

PAGE 320

320 Bolitoglossa diaphora McCranie & Wilson 1995 Bolitoglossa sp. 1: McCranie & Wilson 1994: 147. Bolitoglossa diaphora McCranie & Wilson 1995: 448. B. (Magnadigita) diaphora : Parra Olea et al. 2004: 336. center of Parque Nacional El Cusuco, Cerro Known distribution. Restricted to the Sierra de Omoa in northwestern Honduras, from 1,470 2, 200 m elevation. Available genetic data. 16S (1 sample; Parra Olea et al. 2004), cyt b (1 sample; Parra Olea et al. 2004). Bolitoglossa dunni (Schmidt 1933) Oedipus morio (in part): Dunn 1926: 387. Oedipus dunni Schmidt 1933: 16 (note: the type series w as later shown to contain more than one species by McCranie & Wilson 1993). Magnadigita dunni (in part): Taylor 1944: 218. Bolitoglossa dunni (in part): Wake & Brame 1963: 386. B. ( Magnadigita ) dunni : Parra Olea et al. 2004: 336. ns west of San Pedro, Honduras. Altitude 4500 feet [= Known distribution. Several isolated localities in western Honduras along the Guatemalan border, 1,200 1,600 m elevation.

PAGE 321

321 Available genetic data. 16S (1 sample; Parra Olea et al. 2004), cyt b (1 sample; Parra Olea et al. 2004). Bolitoglossa heiroreias Greenbaum 2004 Magnadigita engelhardti : Mertens, 1952: 20. Bolitoglossa dunni (in part): Wake and Lynch, 1976: 13. Bolitoglossa engelhardti (in part): Villa et al., 1988: 3. Bolitoglossa conanti (in part): McCranie and Wilson, 1993: 8. Bolitoglossa cf. conanti: Leenders and Watkins Colwell, 2004: 5. Bolitoglossa sp. 3 : Parra Olea et al., 2004: 336. Bolitoglossa heiroreias Greenbaum 2004: 412. e of Cerro Montecristo, Depto. Known distribution. Known from the vicinity of Cerro Montecristo in El Salvador, Guatemala, and Honduras, 1,840 2,300 m elevation. Available gen etic data. 16S (2 samples; Parra Olea et al. 2004), cyt b (2 samples; Parra Olea et al. 2004). Bolitoglossa longissima McCranie & Cruz 1996 Bolitoglossa longissima McCranie & Cruz 1996: 195. B. (Magnadigita) longissima : Parra Olea et al. 2004: 336.

PAGE 322

322 Type Known distribution. Known only from the vicinity of the type locality in the Sierra d e Agalta in eastern Honduras, 1,840 2,240 m elevation. Available genetic data. 16S (1 sample; Parra Olea et al. 2004), cyt b (1 sample; Parra Olea et al. 2004). Bolitoglossa oresbia McCranie, Espinal, & Wilson 2005 Bolitoglossa oresbia McCranie et al. 20 05: 108. Known distribution. Known only from the vicinity of the type locality, a 1 hectare patch of forest on top of an isolated peak i n central Honduras, 1,880 m elevation. Available genetic data. None. Bolitoglossa porrasorum McCranie & Wilson 1995 Bolitoglossa dunni (in part): Meyer 1969: 95. Bolitoglossa conanti (in part): McCranie & Wilson 1993: 8. Bolitoglossa sp.: Holm & Cruz 1 994: 20. Bolitoglossa sp. 2: McCranie & Wilson 1994: 147. Bolitoglossa porrasorum McCranie & Wilson 1995: 132. B. (Magnadigita) porrasorum : Parra Olea et al. 2004: 336.

PAGE 323

323 northwe st of Tegucigalpita, 1860 Known distribution. Populations assigned to this taxon are known from the vicinity of Montaa Macuzal and Pico Pijol in Yoro, from Texguat along the border between Atlntida and Yoro, and from the vicinity of Pico Bonito in Atlntida; 980 1,920 m elevation Available genetic data. 16S (1 sample; Parra Olea et al. 2004), cyt b (1 sample; Parra Olea et al. 2004). Bolitoglossa synoria McCranie & Khler 1999 Bolitoglossa celaque (in p art): McCranie & Wilson 1993: 13. Bolitoglossa synoria McCranie & Khler 1999: 226. B. (Magnadigita) synoria : Parra Olea et al. 2004: 336. Cerro El Pital, 2150 m elevation, De Known distribution. Known only from the vicinity of the type locality on Cerro El Pital along the border between Honduras and El Salvador, 2,150 2,713 m elevation. Available genetic data. 16S (1 sample; Parra Olea et al. 2004), cyt b (1 sample; Parra Olea et al. 2004). Subgenus Nanotriton Parra Olea, Garca Pars, & Wake 2004 This subgenus accommodates two small species, one of which is currently known to occur in the Chorts Highlands. The second species, B. occident alis has been

PAGE 324

324 reported previously from a disjunct locality in Honduras, but a recent review of this specimen showed to represent a poorly preserved juvenile B. mexicana (T. Papenfuss, pers. comm.). Bolitoglossa rufescens (Cope 1869) complex Oedipus ruf escens Cope 1869: 104. Spelerpes (Oedipus) rufescens : Peters 1873: 617. Geotriton rufescens : Smith 1877: 76. Spelerpes rufescens : Boulenger 1882: 71. Bolitoglossa rufescens : Taylor 1941: 145. B. (Nanotriton) rufescens : Parra Olea et al. 2004: 335. T Mxico Known distribution. On the Caribbean versant from central Mxico to central Honduras, from near sea level to over 1,400 m elevation, although typically in the lowlands. Available genetic data. 16S (1 sample; Parr a Olea et al. 2004), cyt b (1 sample; Parra Olea et al. 2004). Subgenus Pachymandra Parra Olea, Garca Pars, & Wake 2004 This subgenus was erected to contain two very large species of terrestrial salamanders, one of which occurs in the Chorts Highlands. Bolitoglossa dofleini (Werner 1903)

PAGE 325

325 Spelerpes dofleini Werner 1903: 352. Oedipus schmidti Dunn 1924: 96. Bolitoglossa doffleini [ lapsus ]: Taylor 1944: 219. B. ( Pachymandra ) dofleini : Parra Olea et al. 2004: 337. Known dis tribution. The Caribbean versant of eastern Guatemala, southern Belize, and northwestern and north central Honduras, from 650 1,370 m elevation. Available genetic data. 16S (1 sample; Parra Olea et al. 2004), cyt b (1 sample; Parra Olea et al. 2004).

PAGE 326

326 REF ERENCES Adalsteinnsson SA, Branch WR, Trape S, Vitt LJ, Hedges SB (2009) Molecular phylogeny, classification, and biogeography of snakes of the family Leptotyphlopidae (Reptilia, Squamata). Zootaxa 2244 1 50. Adams DC, Berns CM, Kozak KH, Wiens JJ ( 2009 ) Are rates of species diversification correlated with rates of morphological evolution? Proceedings of the Royal Society B 276, 2729 2738. Agdelo C N (1987) Ecosistemas Terrestres de Honduras Asociacin Hondurea de Ecologa, Tegucigalpa, Honduras. Amp hibiaWeb ( 2011 ) Information on amphibian biology and conservation. [web application]. Berkeley, California: AmphibiaWeb. Available: (Accessed: Oct 6, 2011: http://amphibiaweb.org/ ). Archibold RC, Cave D (2011) Drug wars push deeper into Central America. New York Times 3/31/ 2011 ( accessed 12 September 2011: http://www.nytimes.com/2011/03/24/world/americas/24drugs.html? _r=1&pagewante d=print ). Berger L, Speare R, Daszak P, Green DE, Cunningham AA, Goggin CL, Slocombe R, Ragan MA, Hyatt AD, McDonald KR, Hines HB, Lips KR, Marantelli G, Parkes H (1998). Chytridiomycosis causes amphibian mortality associated with populati on declines in the rain forests of Australia and Central America Proceedings of the National Academy of Science, USA 95 9031 9036. Bickford D, Lohman DJ, Sohdi NS, Ng PKL, Meier R, Winker K, Ingram KK, Das I (2007) Cryptic species as a window on diversi ty and conservation. Trends in Ecology and Evolution 22, 148 155. Bird P (2003) An updated digital model of plate boundaries. Geochemicals, Geophysics, and Geosystems 4 1027 (doi:10.1029/2001GC000252). Bocourt MF (1883) tudes sur les Reptiles. In: Miss ion Scientifique au Mexique et dans ( 1870 1909) Troisime Partie. 1 re Section (eds Dumril AHA, Bocourt MF, Mocquard F) Imprimerie Nati onale, Paris. Borisenko AV, Sones JE, Hebert PDN (2009) The front end logistics of DNA barcoding: challenges and prospects. Molecular Ecology Resources 9 (supplement 1), 27 34. Brame Jr. AH (1968) Systematics and evolution of the Mesoamerican salamande r genus Oedipina Journal of Herpetology 2 1 64.

PAGE 327

327 Centrale. Bulletin de la Socit Philomatique de Paris ( series 7) 1, 175 197. oming problematic in molecular systematic and DNA barcoding studies. Journal of Crustacean Biology 29 96 110. Campbell JA (1998) Comments on the identities of certain Tantilla (Squamata: Colubridae) from Guatemala, with the descriptions of two new specie s. Scientific Papers, Natural History Museum, University of Kansas 7 1 14. Campbell JA (1999) Distribution patterns of amphibians in Middle America. In: Patterns of Distribution of Amphibians: a global perspective (ed Duellman WE), pp. 111 210. Johns H opkins Univers ity Press, Baltimore, Maryland. Campbell JA, Savage J M (2000) Taxonomic reconsideration of Middle American frogs of the Eleutherodactylus rugulosus group (Anura: Leptodactylidae): a reconnaissance of subtle nuances among frogs. Herpetologica l Monographs 14 186 292. Campbell JA, Smith EN. (1998) New species of Nototriton (Caudata: Plethodontidae) from eastern Guatemala. Scientific Papers of the Natural History Museum of the University of Kansas 6 1 8. Campbell JA, Smith EN, Streicher J, Acevedo ME, Brodie Jr E D (2010) New salamanders (Caudata: Plethodontidae) from Guatemala, with miscellaneous notes on known species. Miscellaneous Publications of the Museum of Zoology, University of Michigan 200 i v, 1 60. Carr AF (1950) Outline for a classification of animal habitats in Honduras. Bulletin of the American Museum of Natural History 94 567 594. Castaeda FE (2006) Herpetofauna del Parque Nacional Sierra de Agalta, Honduras International Resources Group, United States Agency for Interna tional Development, Washington, D.C. Castoe TC, Sasa M, Parkinson CL (2005) Modeling nucleotide evolution at the mesoscale: The phylogeny of the Neotropical pitvipers of the Porthidium group (Viperidae: Crotalinae). Molecular Phylogenetics & Evolution 37 881 898. Castoe TC, et al (2009) Comparative phylogeography of pitvipers suggests a consensus of ancient Middle American highland biogeography. Journal of Biogeography 36 88 103. CIPF ( Centro de Informacin y Patrimonio For estal) (2009) Anuario Estad stico Forestal 2008 Instituto Nacional de Conservacin y Desarrollo Forestal, reas Protegidas y Vida S ilvestre, Comayagela, Honduras

PAGE 328

328 Chapman A (1992) Masters of Animals. Oral Traditions of the Tolupan Indians, Honduras Gordon and Breach, Philadelphia Pennsylvania. Clare EL, Lim BK, Engstrom MD, Eger JL, Herbert PDN (2007) DNA barcoding of Neotropical bats: species identification and discovery within Guyana. Molecular Ecology Notes 7 184 190. COHECO (2003) Plan de Manejo Parque Nacional Montaa de Yoro. Consultor a Hondurea en Ecodesarrollo (COHECO), AFE COHDEFOR, DAPVS, Regon Forestal Francisco Morazn, and Regon Forestal Yoro. Cope ED (1866) Fourth contribution to the herpetology of tropical America. Proceedings of the Academy Natural Scienc es Philadelphia 18 123 132. Crawford AJ, Lips KR, Bermingham E (2010) Epidemic disease decimates amphibian abundance, species diversity, and evolutionary history in the highlands of central Panama. Proceedings of the National Academy of Sciences USA 107 13777 13782. Crespi EJ, Browne RA, Rissler LJ (2010) Taxonomic revisin of Desmognathus wrighti (Caudata: Plethodontidae). Herpetologica 66 283 295. Cruz G Wilson LD, Casteeda F (2004) Plectrohyla chrysopleura In: IUCN Red List of Threatened Spec ies. Version 2010.2 ( www.iucnredlist.org ) Downloaded on 20 July 2010 Cushman SA (2006) Effects of habitat loss and fragmentation on amphibians: a review and prospectus. Biological Conservation 128 231 240. Da udin FM (1803) Histoire Naturelle, Gnrale et Particulire des Reptiles; Ouvrage naturelle rdig par C. S. Sonnini, membre de plusieurs Socits savantes. Tome Sixime F. Dufart, Paris. Daza JM, Smith EN, Pez VP, Parkinson CL ( 2009 ) Complex evolution in the Neotropics: the origin and diversification of the widespread genus Leptodeira (Serpentes: Colubridae). Molecular Phylogenetics and Evolution 53 653 667. DeMets C Mattioli G Jansma P Rogers R Tenorio C Turner HL (2007) Present motion and deformation of the Caribbean plate: c onstraints from new GPS geodetic measurements from Honduras and Nicaragua. In: Geologic and Tectonic Development of the Caribbean Plate in Northern Centra l America (ed Mann P), pp. 21 36 Geological Society of America Special Paper 428 Boulder, Colorado, USA. Dix on JR, Hendricks FS (1979) The wormsnakes (family Typhlopidae) of the Neotropics, exclusive of the Antilles. Zoologische Verhande lingen 173 1 39.

PAGE 329

329 Donnelly TW Horne G, Finch R, Lpez Ramos E (1990) Northern Central America: the Maya and Chorts blocks. In: The Geology of North America, Volume 11: The Caribbean Regi on (eds Dengo G, Case JE ) pp. 37 76. Geological Society of America, Boulder, Colorado, USA. Downs FL (1967) Intrageneric relationships among colubrid snakes of the genus Geophis Wagler. Miscellaneous Publications of the Museum Zoology, University of Michigan 131 1 193. Dudgeon D, et al. (2006) Freshwater biodiversity: importance, threats, status and conservation challenges. Biological Review 81 163 182. Duellman WE (1993) Amphibian Species of the World: Additions and Corrections. University of Kansas Museum of Natural History Special Publication 21. Duellman WE (2 001) The Hylid Frogs of Middle America Society for the Study of Amphibians and Reptiles, Contributions in Herpetology 18. Duellman WE, Campbell JA (1992) Hylid frogs of the genus Plectrohyla : systematics and phylogenetic relationships. Miscellaneous Publ ications of the Museum of Zoology, University of Michigan 181 1 32. Dunn ER (1924) New salamanders of the genus Oedipus with a synoptical key. Field Museum of Natural History Zoological Series 12, 95 100. Dunn ER (1926) The Salamanders of the Family Ple thodontidae Smith College, Northhampton, Massachussetts. Ehmcke J, Clemen G (2000) Teeth and their sex dependent dimorphic shape in three species of Costa Rican plethodontid salamanders (Amphibia: Urodela). Annals of Anatomy 182, 403 414. Faivovich J, et al. (2005) Systematic review of the frog family Hylidae, with special reference to Hylinae: phylogenetic analysis and taxonomic revision. Bulletin of t he American Museum of Natural History 294 1 240. Farias IP, Ort G, Sampaio I, Schneider H, Meyer A (2004) The cytochrome b gene as a phylogenetic marker: the limits of resolution for analyzing relationships among cichlid fishes. Journal of Molecular Evolution 53, 89 103. Frost DR, et al. ( 2006 ) The amphibian tree of life. Bulletin of t he American Mu seum of Natural History 297 1 370. Garca Pars M, Wake DB (2000) Molecular phylogenetic analysis of relationships of the tropical salamander genera Oedipina and Nototriton with descriptions of a new genus and three new species. Copeia 2000, 42 70.

PAGE 330

330 Gar ca Pars M Parra Olea G Wake DB (2000) Phylogenetic relationships within the lowland tropical salamanders of the Bolitoglossa mexicana com plex (Amphibia: Plethodontidae). In: The Biology of Plethodontid Salamanders (eds Bruce RC, Jaeger RG, Houck, LD), pp. 199 214 Kluwer Academic/Plenum Publ., New York. Garca Pars M, Parra Olea G, Brame Jr. AH Wake DB (2002) Systematic revision of the Bolitoglossa mexicana species group (Amphibia: Plethodontidae) with description of a new species from Mxico. Revis ta Espaola de Herpetologa 16 43 71. Good DA, Wake, DB (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton with descriptions of three new species. Herpetological Monographs 7, 131 159. Goodwin GG (1942) Mammals of Honduras Bulletin of the American Museum of Natural History 79 107 195. Gordon M (1992) Northern Cent ral America (the Chorts block). In: Jurassic of the Circum Pacific Region: World and Regional Geology v. 3 (ed Westermann G) p. 107 113. Cambridge University Press. Gordon M, Muehlberger W (1994) Rotation of the Chortis block causes dextral slip on the Guayape fault. Tectonics 13 858 872. Gordon M et al (2010) Dating tectonic events on the Chorts Block. Geological Society of America Abstracts with Progra ms 42 197. Gose WA (1985) Paleomagnetic results from Honduras and their bearing on Caribbean tectonics. Tectonics 4 565 585. Gray MJ, Miller DL, Hoverman JT (2009) Ecology and pathology of amphibian ranaviruses. Diseases of Aquatic Organisms 87 243 266. Guindon S, Gascuel O (2003) A simple, fast, and accurate algorithm to estimate large phylogenies by maximum likelihood. Systematic Biology 52, 696 704. Hajibabaei M, Singer GAC, Hebert PDN, Hickey DA (2007) DNA barcoding: how it complements taxonom y, molecular phylogenetics, and population genetics. Trends in Genetics 23 167 172. Halanych KM, Robinson TJ (1999) Multiple substitutions affect the phylogenetic utility of cytochrome b and 12S rDNA data: examining a rapid radiation in Leporid (Lagomor pha) evolution. Journal of Molecular Evolution 48, 369 379. Hanken J, Wake DB, Savage JM (2005) A solution to the large black salamander problem (genus Bolitoglossa ) in Costa Rica and Panam. Copeia 2005 227 245.

PAGE 331

331 Hanken J Wake DB ( 1982) Genetic differ entiation among plethodontid salamanders (genus Bolitoglossa) in Central and South America: implications for the South American invasion. Herpetologica 38 272 287. Hasbn CR, Khler G (2009) New species of Ctenosaura (Squamata, Iguanidae) from sou theast ern Honduras. Journal of Herpetology 43 192 204. Hay WW, DeConto R, Wold CN, Wilson KM, Voigt S, Schulz M, Wold Rossby A, Dullo WC, Ronov AB, Balukhovsky AN, Soeding E (1999) Alternative global cretaceous paleogeography In: The Evolution of C retaceous Ocean/Climate Systems (eds Barrera E, Johnson C), pp. 1 47. Geological Societ y of America Special Paper 332, Boulder, Colorado, USA. Hazlett DL (1980) A botanical description of Cerro Azul Membar, Honduras. Brenesia 18 201 206. Hebert PDN, Cywinska A, Ball SL, deWaard JR (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society of London, B 270 313 321. Hebert PDN Gregory TR (2005). The promise of DNA barcoding for taxonomy. Systematic Biology 54 852 859. Heyer R ( 2002) Leptodactylus fragilis the valid name for the Middle American and northern South American white lipped frog (Amphibia: Leptodactylidae). Proceedings of the Biological Society of Washington 115 321 322. Highton R (1995) Speciation in eastern North American salamanders of the genus Plethodon Annual Review of Ecology and Systematics 26, 579 600. Hillis DM, de S R (1988) Phylogeny and taxonomy of the Rana palmipes group (Salientia: Ranidae). Herpetol ogical Monogr aphs 2 1 26. Hillis DM, Wilcox TP (2005) Phylogeny of the New World true frogs. Molecular Phylogenetics and Evolution 34, 299 314. Holdridge LR (1967) Life Zone Ecology. Revised Edition Tropical Science Center, San Jos, Costa Rica. Holm P A Cruz D GA (1994) A new species of Rhadinaea (Colubridae) from a clou d forest in northern Honduras. Herpetologica 50 15 23. Honeycutt RL, Hillis DM, Bickman JW (2010) Biodiversity discovery and its importance to conservation. In: Molecular Approaches in Natural Resource Conservation and Management (eds DeWoody JA, et al.), pp. 1 35. Cambridge University Press, New York.

PAGE 332

332 Huelsenbeck JP, Ronquist FR (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics (Oxford) 17, 754 755. IUCN ( 2001 ) IUCN Red List Categories and Criteria: Versi on 3.1. IUCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge, United Kingdom. ( http://www.iucnredlist.org/info/categories_criteria2001 ). IUCN (2011) The IUCN Red List of Threatened Species ( http://www.iucnredlist. org/, accessed 1 September 2011). James KH (2007) Structural geology: from local elements to regional synthesis. In: Central America: Geology, Resources, and Hazards (eds Bundschuh J, Alvarado GE), pp. 277 321 Taylor and Francis, Oxford, United Kingdom. Janzen D et al (2009) Integration of DNA barcoding into an ongoing inventory of complex tropical diversity. Molecular Ecology Resources 9 (supplement 1), 1 26. Jockusch EL, Yanev KP, Wake DB (2001 ) Molecular phylogenetic analysis of slender salamanders, genus Batrachoseps (Amphibia: Plethodontidae), from central coastal California with descriptions of four new species. Herpetological Monographs 15, 54 99. Johannessen CL ( 1963 ) Savannas of Interio r Honduras University of California Press Berkeley 160 p. Johnson JD (1989) A biogeographic analysis of the herpetofauna of northwestern Nuclear Central America. Milwaukee Public Museum Contributions in Biology and Geology 76 1 66. Joppa LN, Rober ts DL, Pimm SL (2011) The population ecology and social behavior of taxonomists. Trends in Ecology and Evolution 26 551 553. Jordan BR, Sigurdsson H, Carey SN (2008) Ignimbrites in Central America and Associated Caribbean Sea Tephra: Correlation and Petr ogenesis VDM Verlag Dr. Mller Keppie JD, Morn Zenteno DJ (2005) Tectonic implications of alternative Cenozoic reconstructions for southern Mexico and the Chorts Block. International Geology Review 47 473 491. Kerr KCR, Stoeckle MY, Dove CJ, Weigt LA, Francis CM, Hebert PDN (2007) Comprehensive DNA barcode coverage of Nother American birds. Molecular Ecology Notes 7 535 543.

PAGE 333

333 Ketzler LP, Luque Montes IR Cerrato M CA Wilson LD Townsend JH (2011) Natural History Notes. Thamnophis fulvus (Central America Highland Garter Snake). Diet. Herpetological Review 42 103. Khler G (2002) A new species of salamander of the genus Nototriton from Nicaragua (Amphibia: Caudata: Plethodontidae). Herpetologica 58, 205 210. Khler G (2008) Reptiles of Central A merica. 2nd Edition Herpeton, Offenbach, Germany. Khler G (2010) A revision of the Central American species related to Anolis pentaprion with the resurrection of A. beckeri and the description of a new species. Zootaxa 2354 1 18. Khler G (2011) Amp hibians of Central America Herpeton, Offenbach, Germany. Khler G, Vesel M (2010) A revision of the Anolis sericeus complex with the resurrection of A. wellbornae and the description of a new species (Squamata: Polychrotidae). Herpetologica 66 186 207 Khler G McCranie JR Wilson LD (1999) Two new species of anoles of the Norops crassulus group from Honduras (Reptilia: Sauria: Polychrotidae). Amphibia Reptilia 20 279 298. Khler G, Quintana AZ, Buitrago F, Diether H (2004) New and noteworthy reco rds of amphibians and reptiles from Nicaragua. Salamandra 40 15 24. Khler J Vieites DR, Bonett RM, Hita Garca F, Glaw F, Steinke D, Vences M (2005) N ew amphibians and global conservation : b oost in species discoveries in a hig hly endangered vertebrate group BioScience 55 693 696. Kolby JE, McCranie JR, Rovito SM (2009) Geographic Distribution: Nototriton brodiei (NCN). Herpetological Review 40 444. Kozak KH, Wiens JJ (2006) Does niche conservatism promote speciation? A case study in North Americ an salamanders. Evolution 60 2604 2621. Kozak KH, Wiens JJ (2010) Niche conservatism drives elevational diversity patterns in Appalachian salamanders. The American Naturalist 176 40 54. Kozak KH, Weisrock DW, Larson A (2006) Rapid lineage accumulatio n in a non adaptive radiation: phylogenetic analysis of diversification rates in eastern North American woodland salamanders (Plethodontidae: Plethodon ). Proceedings of the Royal Society B 273 539 546.

PAGE 334

334 Lipscomb D, Platnich N, Wheeler Q (2003) The intel lectual content of taxonomy: a comment on DNA taxonomy Trends in Ecology and Evolution 18 6 66. Luque Montes IR Townsend JH (2009) Geographic Distribution. Ninia espinali Herpetological Review 40 457. Luque Montes IR, Ketz ler LP Cerrato M CA Wilson LD, Townsend JH (2011) Geographic Distribution. Hypopachus barberi (Montane Sheep Frog). Herpetological Review 42 384 385. Lynch JF, Wake DB (1975) Systematics of the Chiropterotriton bromeliacia group (Amphibia: Caudata), wi th description of two new species from Guatemala. Contributions in Science, Natural History Museum of Los Angeles County 265 1 45. Lynch JF, Wake DB (1978) A new species of Chiropterotriton (Amphibia: Caudata) from Baja Verapaz, Guatemala, with comments on relationships among Central American members of the genus. Contributions in Science, Natural History Museum of Los Angeles County 294, 1 22. Macey JR (2005) Plethodontid salamander mitochondrial genomics: a parsimony evaluation of character conflict an d implications for historical biogeography. Cladistics 21, 194 202. Mant on WI (1996) The Grenville of Honduras. Geological Society of America Abstracts with Programs 28 493. Mann P, Rogers RD, Gahagan L (2007) Overview of plate tectonic history and its unresolved tectonic problems. In: Central America: Geology, Resources, and Hazards (eds Bundschuh J, Alvarado GE), pp. 206 241 Taylor and Francis, Oxford, United Kingdom. Marshall JS ( 2007 ) The geomorphology and physiographic provinces of Central Americ a. In: Central America: Geology, Resources, and Hazards (eds Bundschuh J, Alvarado GE), pp. 75 122. Taylor and Francis, Oxford, United Kingdom. Martin, M. (1972) A biogeographic analysis of the freshwater fishes of Honduras Unpublished Ph.D. Dissertatio n. University of Southern California. Matamoros WA, Schaefer JF, Kreiser BR (2009) Annotated checklist of the freshwater fishes of continental and insular Honduras. Zootaxa 2307 1 38. Matamoros WA, Schaefer JF (2010) A new species of Profundulus (Cypri nodontiformes: Profundulidae) from the Honduran central highlands. Journal of Fish Biology 76 1498 1507.

PAGE 335

335 McCranie JR (1996 a ) Geographic Distibution. Nototriton barbouri (NCN). Herpetological Review 27, 28. McCranie JR (1996 b ) Geographic Distibution. O edipina gephyra (NCN). Herpetological Review 27, 29. McCranie JR (2006) A new species of Oedipina (Amphibia: Urodela) from Parque Nacional El Cusuco, northwestern Honduras. Journal of Herpetology 40 291 293. McCranie JR (2009) Amphibians and Reptiles o f Honduras Listas Zoolgicas Actualizadas UCR ( http://museo.biologia.ucr.ac.cr/Listas/LZAPublicaciones.htm ). Museo de Zoologa UCR. San Pedro, Costa Rica. McCranie J R ( 2011a ) The Snakes of Honduras. Systematics, Distribution, and Conservation. Society for the Study of Amphibians and Reptiles, Contributions in Herpetology. McCranie JR ( 2011 b) A new species of Tantilla of the taeniata species group (Reptilia, Squamata, Colubridae, Colubrinae) from nort heastern Honduras. Zootaxa 3037 37 44. McCranie JR, Castaeda FE (2004a) A new species of snake of the genus Omoadiphas (Reptilia: Squamata: Colubridae) from the Cordillera Nombre de Dios in northern Honduras. Proceeding s of the B iological Society of Washington 117 311 316. McCranie JR, Castaeda FE (2004b): Notes on the second specimens of Geophis damiani Wilson, McCranie, and Williams and Rhadinaea tolpanorum Holm & C ruz D. (Colubridae). Herpetological Review 35 341. McCranie JR, Castaeda FE (2005) The herpetofauna of Parque Nacional Pico Bonito, Honduras. Phyllomedusa 4 3 16. McCranie JR, Castaeda FE ( 2007) Gua de Campo de los Anfibios de Honduras Bibliomania!, Salt Lake City, Utah. McCranie JR, Cruz Diaz GA (2010) A third new species of snake of the genus Omoadiphas (Reptilia, Squamata, Colubridae, Dipsadinae) from Honduras. Zootaxa 2 690 53 58. McCranie JR, Townsend JH ( 2011 ) Description of a new species of worm salamander (Caudata, Plethodontidae, Oedipin a ) in the subgenus Oedopinola from the central portion of the Cordillera Nombre de Dios, Honduras. Zootaxa 2990 59 68. McCranie JR, Wilson LD (1993) A review of the Bolitoglossa dunni group (Amphibia: Caudata) from Honduras with the description of three new species. Herpetologica 49, 1 15.

PAGE 336

336 McCranie JR, Wilson LD ( 1995a ) Two new species of colubrid species of the genus Ninia from Central America. Journal of Herpetology 29 224 232. McCranie JR, Wilson LD ( 1995 b ) A new species of salamander of the Bolito glossa dunni group (Caudata: Plethodontidae) from northern Honduras. Herpetologica 51, 131 140. McCranie JR, Wilson LD (1996 [1997]) Two new species of salamanders (Caudata: Plethodontidae) of the genera Bolitoglossa and Nototriton from Parque Nacional La Muralla, Honduras. Proceedings of the Biological Society of Washington 110 366 372. McCranie JR, Wilson LD ( 1997) Two new species of salamanders (Caudata: Plethodontidae) of the genera Bolitoglossa and Nototriton from Parque Nacional La Muralla, Hondu ras. Proceedings of the Biological Society of Washington 110, 366 372. McCranie JR, Wilson LD (2001) Taxonomic status of Typhlops stadelmani Schm idt (Serpentes: Typhlopidae). Copeia 2001 820 822. McCranie JR, Wilson LD ( 2002) The Amphibians of Honduras Society for the Study of Amphibians and Reptiles, Contributions in Herpetology. McCranie JR, Wilson LD Williams KL (1992) A new species of anole of the Norops crassulus group (Sauria: Polychridae) from northwestern Honduras. Caribbean Journal of Scienc e 28 208 215. McCranie JR Wilson LD, Williams KL (1993 ) A new species of Oedipina (Amphibia: Caudata: Plethodontidae) from northern Honduras. Proceedings of the Biological Socity of Washington 106 385 389. McCranie JR, Wilson LD, Polisar J (1998) Ano ther new montane salamander (Amphibia: Caudata: Plethodontidae) from Parque Nacional Santa Barbara, Honduras. Herpetologica 54 455 461. McCranie JR, Wilson LD, Gotte SW (2001) Three new country records for Honduran snakes. Herpetological Review 32 62 63 McCranie JR, Espinal MR, Wilson LD (2005) New species of montane salamander of the Bolitoglossa dunni group from northern Comayagua, Honduras (Urodela: Plethodontidae). Journal of Herpetology 39 108 112. McCranie JR, Townsend JH, Wilson LD (2006) The Amphibians and Reptiles of the Honduran Mosquitia Krieger Publishing Company, Mal abar, Florida

PAGE 337

337 McCranie JR, Vieites DR, Wake DB (2008) Description of a new divergent lineage and three new species of Honduran salamanders of the genus Oedipina (Caudata, Pl ethodontidae). Zootaxa 1930, 1 17. McCranie JR, Valds Orellana L, Himes JG (2010) Rediscovery of two Honduran endemic streamside frogs, Craugastor emleni (Dunn) and Craugastor stadelmani (Schmidt). Froglog 94 12 15. McDiarmid, R. W., J. A. Campbell, and T. A. Tour. 1999. Snake Species of the World. A Taxonomic and Geographic Reference. Vol. 1 Washington. Meja Ordez & House (2002) Mapa de Ecosistemas Vegetales de Honduras Preparado para el Proyecto P.A.A.R., Tegucigalpa M.D.C. Meja Valdivieso DA (2001) Honduras. In: Bosques Nublados del Neotrpico (eds Kappelle M, Brown AD), pp. 243 282. Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica. Mendelson III J ( 2011 ) Shifted baselines, forensic taxono limbed treefrog: the changing role of biologists in an era of amphibian declines and extinctions. Herpetological Review 42 21 25. Mendelson III J, Mulcahy DG (2010) A new species of toad ( Bufonidae : Incilius ) from Central Panama. Z ootaxa 2396 61 68. Mendelson III J, Williams BL, Sheil CA, Mulcahy DG (2005) Systematics of the Bufo coccifer complex (Anura: Bufonidae) of Mesoamerica Scientific Papers, Natural History Museum of the University of Kansas 38 1 27. Mertens R (1952) Di e amphibien und reptilien von El Salvador. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 487 1 120. Meyer C (2003) Molecular systematics of cowries (Gastropoda: Cypraeidae) and diversification patterns in the tropics. Biological Journ al of the Linnean Society 79, 401 459. Meyer, JR (1969) A Biogeographic Study of the Amphibians and Reptiles of Honduras Unpublished PhD dissertation, University of Southern California. Meyer JR Wilson LD (1971) A distributional checklist of the amphi bians of Honduras. Los Angeles County Museum Contributions in Science 218 1 47. Meyer JR, Wilson LD (1973) A distributional checklist of the turtles, crocodilians, and lizards of Honduras. Natural History Museum of Los Angeles County Contributions in Sc ience 244 1 39.

PAGE 338

338 Milanovich JR, Peterman WE, Nibbelink NP, Maerz JC (2010 ) Projected loss of a salamander diversity hotspot as a consequence of projected global climate change. PLoS One 5 e12189. Monaghan MT, et al. (2009) Accelerated species inventory on Madagascar using Coalescent based models of species delineation. Systematic Biology 58 298 311 Monroe Jr BL (1968) A distributional survey of the birds of Honduras. Amer ican Ornit hological Union, Ornithological Monograph 7 1 458. Morn Zenteno DJ, et al. (2009) Reassessment of the Paleogene position of the Chorts Block relative to southern Mexico: hierarchical ranking of data and features. Revista Mexicana de Ciencias Geolgicas 26 177 188. Moritz C, Cisero C (2004) DNA barcoding: promises and pitfalls. PLoS Biology 2: 1529 1531. Moritz C, Schneider CJ, Wake DB (1992) Evolutionary relationships within the Ensatina eschscholtzii complex confirm the ring species interpretation. Systematic Biology 41, 273 291. Mueller RL, Macey JR, Jaekel M, Wa ke DB, Boore JL (2004) Morphological homoplasy, life history evolution, and historical biogeography of plethodontid salamanders inferred from complete mitochondrial genomes. Proceedings of the National Academy of Science, USA 101, 13820 13825. Mueller RL (2006) Evolutionary rates, divergence dates, and the performance of mitochondrial genes in Bayesian phylogenetic ana lysis. Systematic Biology 55 289 300 Mueller Dombois D, Ellenberg H (1974) Aims and Methods in Vegetation E cology J. Wiley & S ons, New York, USA. Mulcahy DG, Mendelson III JR (2000) Phylogeography and speciation of the morphologically variable, widespread species Bufo valliceps based on molecular evidence from mtDNA. Molecular Phylogenetics and Evolution 17 173 189. Mulcahy DG, Morri ll BH, Mendelson III JR (2006) Historical biogeography of lowland species of toads ( Bufo ) across the Trans Mexican Neovolcanic Belt and the Isthmus of Tehuantepec. Journal of Biogeography 33 1889 1904. Mullis K. (1990) The unusual o rigin of the Polymera se Chain Re action. Scientific American 262 56

PAGE 339

339 Myers CW (2003) Rare snakes Five new species from eastern Panama: Reviews of northern Atractus and southern Geophis (Colubridae: Dipsadinae). American Museum Novitates 3391 1 47. Myers CW (2011) A new ge nus and new tribe for Enicognathus melanauchen Jan, 1863, a neglected South American snake (Colubridae: Xenodontinae), with taxonomic notes on some Dipsadinae. American Museum Novitates 3715 1 33. Nelson S CH (2001) Plantas descritas originalmente de Hon duras y sus nomenclaturas equivalentes actuales. Ceiba 42 1 71. Nelson S CH (2008) Catlogo de las Plantas Vasculares de Honduras. Espermatofitas. Secretria de Recursos Naturales y Ambiente/Guaymuras, Tegucigalpa, Honduras. Ortega Gutirrez F et al. (2007) The Maya Chorts boundary: a tectonostratigraphic approach. International Geological Review 49 996 1024. Palumbi SR, Martin AP, Romano S, McMillan WO, Stice L, Grabowski, G (1991) The uide to PCR Special Publication of the Departm ent of Zoology, University of Hawaii, Honolulu. Papenfuss TJ, Wake DB ( 1987 ) Two new species of plethodontid salamanders (genus Nototriton ) from Mxico Acta Zoologica Mexicana 21, 1 16. Parra Olea G, Wake DB (2001) Extreme morphological and ecological homoplasy in tropical salamanders. Proceedings of the National Academy of Sciences USA 98 7888 7891. Parra Olea G, Garca Pars M, Wake DB (2004) Molecular diversification of salamanders of the tropical American genus Bolitoglossa (Caudata: Plethodonti dae) and its evolutionary and biogeographical implications. Biological Journal of the Linnean Society 81, 325 346. Parra Olea G, et al. (2007) Systematics and Conservation. In: Amphibian Conservation Action Plan (eds Gascon C, et al.), pp. 45 48. IUCN/SS C Amphibian Specialist Group, Grand Switzerland. Parsons JJ (1955) The Miskito pine savanna of Nicaragua and Honduras. Annals of the Association of American Geographers 45 36 63. Pfenninger M, Schwenk K (2007) Cryptic animal species are homogeneously d istributed among taxa and biogeographical regions. BMC Evolutionary Biology 7, 121. Posada D (2008) jModelTest: phylogenetic model averaging. Molecular Biology and Evolution 25, 1253 1256.

PAGE 340

340 Pyron RA, Wiens JJ (2011) A large scale phylogeny of Amphibia with over 2,800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution 61 543 583. Reid F (2009) Field Guide to the Mammals of Central America and Southeast Mxico Se cond Edition. Oxford Uni versity Press. Relyea RA, Diecks N (2008) An unforeseen chain of events: Lethal effects of pesticides at sublethal concentrations. Ecological Applications 18 1728 1742. Rogers RD (2003) Jurassic Recent tectonic and stratigraphic history of the Chorts block of Honduras and Nicaragua (northern Central America). U npu blished Ph.D. dissertation, University of Texas at Austin. Rogers R, Karason H, van der Hilst, R (2002) Epeirogenic uplift above a detached slab in northern Central America. Geology 30 1031 1034. Rogers R, Mann P, Emmet PA (2007) Tectonic terranes of the Chortis Block based on integration of regional aeromagnetic and geological data. In: Geologic and Tectonic Development of the Caribbean Plate in Northern Central America (ed Mann P), pp. 65 88. Geological Society of America Special Paper 428 Boulder, Colorado, USA. Rovito SM, Parra Olea G Vsquez Almazn CR Papenfuss TJ, Wake DB (2009) Dramatic declines in neotropical salamander populations are an important part of the global amphibi an c risis. Proceedings of the National Academy of Sciences USA 106 3231 3236. Rovito SM, Vsquez Almazn CR, Papenfuss TJ (2010) A new species of Bolitoglossa (Caudata: Plethodontidae) from the Sierra de las Minas, Guatemala. Journal of Herpetology 44, 516 525. Sanger F, Nicklen S, Coulson AR (1977) DNA sequencing with chain terminating inhibitors. Proceedings of the National Academy of Sciences of the USA 74 5463 5467. Sasa M, Bolaos F (2004) Biodiversity and conservation of Mesoamerican dry forest he rpetofauna. In: Biodiversity Conservation in Costa Rica: Learning the Lessons in a Seasonal Dry Forest (eds Frankie GW, Mata A, Vinson SB), pp. 177 193. University of California Press. Savage JM (1966) The origins and history of the Central American herp etofauna: dispersal or vicariance? Copeia 1966 719 766. Savage JM ( 1983) The enigma of the Central American herpetofauna: dispersals or vicariance? Annals of the Missouri Botanical Garden 69 464 547.

PAGE 341

341 Savage JM ( 2002) The Amphibians and Reptiles of Cos ta Rica: A Herpetofauna Between Two Continents, Between Two Seas University of Chicago Press. Chicago, Illinois. Schmidt KP (1933) New reptiles and amphibians from Honduras. Zoological Series of Field Museum of Natural History 20, 1 22. Schmidt KP ( 1 936 ) New amphibians and reptiles from Honduras in the Museum of Comparative Zoology. Proceedings of the Biological Society of Washington 49, 43 50. Schuster, SC (2008) Next Nature Methods 5 16 18. Sebe rg O, et al. (2003) Shortcuts in systematics? A commentary on DNA based taxonomy. Trends in Ecology and Evolution 18 63 65. Seutin G White BN, Boag PT (1991) Preservation of avian blood and tissue samples for DNA analyses. Canadian Jou rnal of Zoology 6 9 82 90 Silva Romo G (2008) Guayape Papalutla fault system: A continuous Cretaceous structure from southern Mxico to the Chorts block? Tectonic implications. Geology 36 75 78. Skerratt LF, Berger L, Speare R, Cashins S, McDonald KR, Phillott AD, Hin es HB, Kenyon N (2007) Spread of chytridiomycosis has caused the rapid global decline and extinction of frogs. EcoHealth 4 125 134. Smith BE, Campbell JA (1996) The systematic status of Guatemalan populations of snakes allied with Ninia maculata (Reptil ia: Colubridae). Proceedings of the Biological Society of Washington 109 749 754. Smith MA, Poyarkov Jr NA, Herbert PDN (2008) CO1 DNA barcoding amphibians: take the chance, meet the challenge. Molecular Ecology Resources 8, 235 246. Song H, Buhay JE, Whiting MF, Crandall KA (2008) Many species in one: DNA barcoding overestimates the number of species when nuclear mitochondrial pseudogenes are coamplified. Proceedings of the National Academy of Sciences of the USA 105 13486 13491. Stafford PJ (2004) A new species of Tantilla (Serpentes: Colubridae) of the taeniata group from Southern Belize. Journal of Herpetology 38 43 52. Stamatakis A (2006) RAxML VI HPC: Maximum likelihood based phylogenetic analyses with thousands of taxa and mixed models. Bioinf ormatics 22, 2688 2690.

PAGE 342

342 Stehli F G, Webb SD (1985) The Great American Biotic Interchange Plenum Press, New York. Stuart LC (1954) A description of a subhumid corridor across northern Central America, with comments on its he rpetofaunal indicators. Contri butions of the Laborat ory of Vertebrate Zoology, University of Michigan 65 1 26. Stuart LC ( 1963 ) A checklist of the herpetofauna of Guatemala. Miscellaneous Publications of the Museum of Zoology, University of Michigan 122 1 150. Stuart SN, Chanson J S, Cox NA, Young BE, Rodrigues ASL, Fischman DL, Waller RW ( 2004 ) Status and trends of amphibian declines and extinctions worldwide. Science 306 1783 1786. Sunyer J, Khler G ( 2010 ) Conservation status of the herpetofauna of Nicaragua. In: Conservation o f Mesoamerican Amphibians and Reptiles (eds Wilson LD, To wnsend JH, Johnson, JD), pp. 488 509. Eagle Mountain Publishing LC, Eagle Mountain, Utah. Sunyer J Townsend JH, Wilson LD, Travers SL Obando L Paiz G and Griffith DM ( 2009) Three new country re cords of reptiles from Nicaragua. Salamandra 45 186 190. Sunyer J Wake DB Townsend JH Travers SL Rovito SM Papenfuss TJ, Obando LA ( 2010) A new species of worm salamander (Caudata: Plethodontidae: Oedipina ) in the subgenus Oeditriton from the highla nds of northern Nicaragua. Zootaxa 2613 29 39. Sunyer J Townsend JH, Wake DB Travers SL Gonzalez SG Obando LA Quintana AZ (2011) A cryptic new species of Oedipina (Caudata: Plethodontidae) from premontane elevations in northern Nicaragua, with commen ts on the systematic status of the Nicaraguan paratypes of O. pseudouniformis Brame, 1968. Breviora 526 1 16. Systematics Agenda 2000 (1994). Sy stematics Agenda 2000: Charting the Biosphere New York: Department of Ornithology, American Museum of Natural History. Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S (2011) MEGA5: molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution 28 2731 2739 T autz D, et al. (2002) DNA points the way ahead in taxonomy. Nature 418 479. Tautz D, Arctander P, Minelli A, Thomas RH, Vogler AP (2003) A plea for DNA taxonomy. Trends in Ecology and Evolution 18 70 74.

PAGE 343

343 Taylor EH (1944) The genera of plethodont salam anders in Mxico Part I. University of Kansas Science Bulletin 30, 189 232. Thomas R Hedges SB (2007) Eleven new species of snakes of the genus Typhlops (Serpentes: Typhlopidae) from Hispaniola and Cuba. Zootaxa 1400 1 26. Thompson JD Higgins DG, Gib son TJ ( 1994 ) CLUSTALW: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position specific gap penalties and weight matrix choice Nucleic Acids Research 22, 4673 4680 Til ley SG, Ma hon e y MJ (1996) Patterns of genetic differentiation in salamanders of the Desmognathus ochrophaeus complex (Amphibia: Plethodontidae). Herpetological Monographs 10 1 42. Townsend JH (2006) 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 Florida, Gainesville, Florida, USA. i xiv, 1 124. Townsend JH ( 2009) Morphological variation in Geophis nephodrymus (Squam ata: Colubridae), with comments on the conservation of Geophis in eastern Nuclear Central America. Herpetologica 65 292 302. Townsend JH, Wilson LD ( 2008) Guide to the Amphibians and Reptiles of Cusuco National Park, Honduras / Gua de los Anfibios y Re ptiles de Parque Nacional Cusuco, Honduras Bibliomania!, Salt Lake City, Utah. Townsend JH, Wilson LD ( 2006 ) Denizens of the dwarf forest: herpetofauna of the elfin forests of Cusuco National Park, Honduras. Iguana 13 242 251. Townsend JH, Wilson LD ( 2009 ) New species of cloud forest Anolis (Squamata: Polychrotidae) of the crassulus group from Parque Nacional Montaa de Yoro, Honduras. Copeia 2009 62 70. Townsend JH, Wilson LD (2010 a ) Conservation of the Honduran herpet ofauna: issues and imperative. In: Conservation of Mesoamerican Amphibians and Reptiles (eds Wilson LD, Townsend JH, Johnson, JD), pp. 460 487. Eagle Mountain Publishing LC, Eagle Mountain, Utah. Townsend JH, Wilson LD ( 2010b ) Biogeography and conservation of the Honduran subhumid fore st herpetofauna. In: Conservation of Mesoamerican Amphibians and Reptiles (eds Wilson LD, Townsend JH, Johnson, JD), pp. 686 705 Eagle Mountain Publishing LC, Eagle Mountain, Utah.

PAGE 344

344 Townsend JH, Wilson LD, Plenderleith TL Talley BL Nifong JC (2005) Nini a pavimentata (Squamata: Colubridae): an addition to the snake fauna of Honduras. Caribbean Journal of Science 41 869 870. Townsend JH, Wilson LD Ketzler LP Luque Montes IR ( 2008a ) The largest blindsnake in Mesoamerica: a new species of Typhlops (Squam ata: Typhlopidae) from an isolated karstic mountain in Honduras. Zootaxa 1932 18 26. Townsend JH, Wilson LD Luque Montes IR Ketzler LP ( 2008b ) Redescription of Anolis rubribarbaris (Khler, McCranie, & Wilson 1999), a poorly known Mesoamerican cloud f orest anole (Squamata: Polychrotidae). Zootaxa 1918 39 44. Townsend JH, Butler JM, Wilson LD, Austin JD ( 2009 a ) A new species of salamander in the Bolitoglossa dunni group (Caudata: Plethodontidae: Bolitoglossinae) from Parque Nacional Montaa de Yoro. Salamandra 45, 95 105. Townsend JH, Butler JM, Wilson LD Ketzler LP Slapcinsky J Stewart NM ( 2009 b) Significant range extension for the Central American Colubrid snake Ninia pavimentata (Bocourt 1883) Herpetological Bulletin 106 15 17. Townsend JH Butler JM, Wilson LD, Austin JD ( 2010 a ) A distinctive new species of moss salamander (Caudata: Plethodontidae: Nototriton ) from an imperiled Honduran endemism hotspot. Zootaxa 2434, 1 16. Townsend JH Herrera B LA Medina Flores M Gray LN Stubbs AL W ilson LD ( 2010b ) Notes on the second male specimen of the cryptozoic snake Geophis damiani Wilson, McCranie, & Williams 1998 (Squamata: Colubridae: Dipsadinae). Herpetology Notes 3 305 308. Townsend JH Medina Flores M Murillo JL Austin JD ( 2011a ) Cry ptic diversity in Chorts Highland moss salamanders (Caudata: Plethodontidae: Nototriton ) revealed using mtDNA barcodes and phylogenetics, with a new species from eastern Honduras Systematics and Biodiversity 9 275 287. Townsend JH, Luque Montes IR, Wil son LD ( 2011 b ) Newly discovered populations of four threatened endemic salamanders (Caudata: Plethodontidae) from the highlands of Honduras. Salamandra 47, 50 54. Townsend JH, Wilson LD Cerrato M CA Atkinson BK Herrera B LA McKewy Meja M ( 2011c ) Disc overy of the critically endangered treefrog Plectrohyla chrysopleura in Refugio de Vida Silvestre Texiguat, Honduras. Herpetological Bulletin 115 22 25. Townsend JH, Wilson LD, Medina Flores M, Herrera B LA, Atkinson BK, Stubbs AL, Cerrato M CA, McKewy M eja M, Gray LN, Austin JD ( In press A) Uncovering a hotspot for premontane endemism on the windward side of Refugio de Vida Silvestre Texguat Honduras. Salamandra

PAGE 345

345 Townsend JH, Wilson LD, Medina Flores M, Herrera B LA ( In press B) A new species of centip ede snake in the Tantilla taeniata group (Squamata: Colubridae) from premontane forest in Refugio de Vida Silvestre Texguat, Honduras. Journal of Herpetology Travers SL Townsend JH Sunyer J Obando LA Wilson LD 2011. New and noteworthy records of amp hibians and reptiles from Reserva de la Bisfera Bosawas, Nicaragua. Herpetological Review 42 399 403. Uetz P, et al. (2011) The Reptile Database [website]. (Accessed 6 October 2011: http://www.reptile datab ase.org ). U nited N ations Mine Action Service (1998) Nicaragua Landmine Situation Assessment Mission Report United Nations, 15 December 1998. 6 p. Vsquez Almazn CR, Rovito SM, Good DA, Wake DB (2009) A new species of Cryptotriton (Caudata: Plethodon tidae) from eastern Guatemala. Copeia 2009, 313 319. Vences M, Thomas M, Bonett RM, Vieites DR (2005 a ) Deciphering amphibian diversity through DNA barcoding: chances and challenges. Philosophical Transactions of the Royal Society B 360, 1859 1868. Venc es M, Thomas M, Meijden Avd, Chiari Y, Vieites DR (2005b) Comparative performance of the 16S rRNA gene in DNA barcoding of amphibians. Frontiers in Zoology 2 5. Vieites DR, Min MS, Wake DB (2007) Rapid diversification and dispersal during periods of glo bal warming by plethodontid salamanders. Proceedings of the National Academy of Science, USA 104 19903 19907. Vieites DR, et al. (2009) by an integrative amphibian inventory. Proceedings of the National Academy of Science, USA 106 8267 8272. Vieites DR, Nieto Romn S, Wake MH, Wake DB (2011) A multigenic perspective on phylogenetic relationships in the largest family of salamanders, the Plethodontidae. Molecular Phylogenetic and Evolution 59, 623 635. Villa J (1972) Anfibios de Nicaragua. Instituto Geogrfico Nacional & Banco Central de Nicaragua. Virgilio M, Backeljau T, Nevado B, De Meyer M (2010) Comparative performance of DNA barcoding across insect orders. BMC Bioinformatics 11 206.

PAGE 346

346 Vreugdenhil D, House P Cerrato C, Martnez RA Pereira AC ( 2002) Racionalizacion del S istema de reas P rotegida s del Honduras. Volume I: Main S tudy Tegucigalpa, Honduras: PROBAB World Bank, UNDP, GEF. Wake DB (1987) Adaptive radiation of salamanders i n Middle American cloud forests. Annals of the Missouri Botanical Garden 74, 242 264. Wake DB (19 91 ) Declining amphibian populations. Science 253 860. Wake DB (1998) On the taxonomic status of Nototriton sanctibarbarus McCranie and Wilson (Amphibia: Cau data). The Southwestern Naturalist 43 88 106. Wake DB (2009) What salamanders have taught us about evolution. Annual Review of Ecology, Evolution, and Systematics 40 333 352. Wake DB, Campbell JA (2000) A new species of diminutive salamander (Amphibia : Caudata: Plethodontidae: Nototriton ) from the Montaas del Mico of Guatemala. Proceedings of the Biological Society of Washington 113, 815 819. Wake DB, Elias P (1983) New genera and a new species of Central American salamanders, with a review of the t ropical genera (Amphibia, Caudata, Plethodontidae). Contributions in Science, Natural History Museum of Los Angeles County 345, 1 19. Wake DB, Vredenburg VT (2008) Are we in the midst of the sixth mass extinction? A view from the world of amphibians. Proc eedings of the National Academy of Science, USA 105 11466 11473. Wallace, AR (1876) The Geographic Distribution of Animals. With a Study of the Relations of Living and Extinct Faunas as Elucidating the Past Changes of the Two volumes N ew York: Harper and Brothers. Wang, IJ, Crawford, AJ, Bermingham E (2008) Phylogeography of the pygmy rain f rog ( Pristimantis ridens ) across the lowland wet forests of Isthmian Central America. Molecular Phylogenetics and Evolution 47 992 1004. Ward RD, Hanner R, Hebert PDN (2009) The campaign to DNA barcode all fishes, FISH BOL. Journal of Fish Biology 74 329 356. Wheeler, QD (2004) Taxonomic triage and the poverty of phylogeny. Philosophical Transactions of the Royal Society B London 359 57 583. Wiens JJ, Parra Olea G, Garca Pars M, Wake DB (2007) Phylogenetic history underlies elevational biodiversity patterns in tropical salamanders. Proceedings of the Royal Society B 274, 919 928.

PAGE 347

347 Williams H, McBirney AR ( 1969 ) Volcanic history of Honduras University of California Publications in Geological Sciences 85 1 101. Williams ST (2007) Safe and legal shipment of tissue samples: does it a ffect DNA quality? Journal of Molluscan Studies 73 416 418. Wilson LD ( 1999 ) Checklist and key to the spec ies of the genus Tantilla (Serpentes: Colubridae), with some distributional commentary. Smithsonian Herpetological Information Service 122 1 34. Wilson LD, Johnson JD (2010) Distributional patterns of the herpetofauna of Mesoamerica, a biodiversity hotsp ot. In: Conservation of Mesoamerican Amphibians and Reptiles (eds Wilson LD, Townsend JH, Johnson, JD), pp. 30 235. Eagle Mountain Publishing LC, Eagle Mountain, Utah. Wilson LD, McCranie JR (1998) The biogeography of the herpetofauna of the subhumid fore sts of Middle America (Isthmus of Tehuantepec to northwestern Costa Rica). Royal Ontario Museum of Life Sciences Contribution 163 1 50. Wilson LD, McCranie JR (1999) The systematic status of Honduran populations of the Tantilla taeniata group (Serpentes: Colubridae), with notes on other populations. Amphibia Reptilia 20 326 329. Wilson LD, McCranie JR (2003) Herpetofaunal indicator species as measures of environmental stability in Honduras. Caribbean Journal of Science 39 50 67. Wilson LD, McCranie JR (2004 a ) The conservation status of the herpetofauna of Honduras. Amphibian and Reptile Conservation 3 6 33. Wilson LD, McCranie JR (2004 b ) The herpetofauna of t he cloud forests of Honduras. Amph ibian and Reptile Conservation 3 34 48. Wilson LD, Meyer JR (1971) A revision of the taeniata group of the colubrid snake genus Tantilla Herpetologica 27 11 40. Wilson LD, Meyer JR (1985) The Snakes of Honduras. 2nd Edition Milwaukee Public Museum, Milwaukee, Wisconsin. Wilson LD, Townsend JH (2006) The her petofauna of the rainforests of Honduras. Caribbean Journal of Science 42 88 113. Wilson LD, Townsend JH (2007) Checklist and key to the snakes of the genus Geophis (Squamata: Colubridae), with commentary on distribution and conservation status. Zootaxa 1395 1 31.

PAGE 348

348 Wilson LD, Townsend JH ( 2010) The herpetofauna of Mesoamerica: b iodiversity significance, conservation status, and future challenges. In: Conservation of Mesoamerican Amphibians and Reptiles (eds Wilson LD, Townsend JH, Johnson, JD), pp. 760 812 Eagle Mountain Publishing LC, Eagle Mountain, Utah. Wilson LD, Cruz Daz GA, Villeda E, Flores S ( 1988 ) Typhlops costaricensis Jimnez & Savage: an addition to the snake fauna of Honduras. The Southwestern Naturalist 33 499 500. Wilson LD, McCran ie JR Cruz D GA (1994) A new species of Plectrohyla (Anura: Hylidae) from a premontane rainforest in northern Honduras. Proceedings of the Biolo gical Society of Washington 107 67 78. Wilson LD McCranie JR, Williams KL (1998) A new species of Geophis of the sieboldi group (Reptilia: Serpentes: Colubridae) from no rthern Honduras. Proceedings of the B iological Society of Washington 111 410 417. McCranie JR, Wilson LD, Gotte S (2001) Three new country records for Honduran snakes. Herpetological Review 32 62 63. Wilson LD, McCranie JR, Espinal MR (2001 b ) The ecogeography of the Honduran herpetofauna and the design of biotic reserves. In: Mesoamerican Herpetology: Systematics, Zoogeography, and Conservation (eds Johnson JD, Webb RG, Flores Villela OA), pp. 109 158. Centennial Museum, University of Texas, El Paso, Special Publication 1. Wilson LD, Townsend JH, Johnson JD (2010) Conservation of Mesoamerican Amphibians and Reptiles (eds ) Eagle Mountain Publish ing LC, Eagle Mountain, Utah. Xia Y, et al (2011) COI is better than 16S rRNA for DNA barcoding Asiatic salamanders (Amphibia: Caudata: Hynobiidae). Molecular Ecology Resources DOI: 10.1111/j.1755 0998.2011.03055.x. Xia X, Xie Z (2001) DAMBE: Data Ana lysis In Molecular Biology And Evolution. Journal Of Heredity 92, 371 373. Xia X, Xie Z, Salemi M, Chen L, Wang Y (2003) An index of substitution saturation and its applications. Molecular Phylogenetics and Evolution 26, 1 7. Xia X, Lemey P (2009) Asse ssing substitution saturation with DAMBE. In: T he Phylogenetic Handbook: A Practical Approach to DNA and Protein Phylogeny, 2nd edition (eds Lemey P, Salemi M, Vandamme AM), pp. 615 63 0. Cambridge University Press.

PAGE 349

349 Zaldvar Rivern A, Len Regagnon V, Ni eto Montes de Oca A (2004) Phylogeny of the Mexican coastal leopard frogs of the Rana berlandieri group based on mtDNA sequences. Molecular Phylogenetics and Evolution 30 38 49. Zamora Villalobos N (2000) Arboles de la Mosquitia Hondurea CATIE, Serie Tcnica, Manual Tcnico 43, Turrialba, Costa Rica. Zhang Z, Schwartz S, Wagner L, Miller W (2000) A greedy algorithm for aligning DNA sequences. Journal of Computational Biology 7 203 214. Zhang P, Wake DB (2009) Higher level salamander relationships and divergence dates inferred from complete mitochondrial genomes. Molecular Phylogenetics and Evolution 53, 492 508. Zimku s BM, Schink S (2010) Light at the end of the tunnel: insights into the molecular systematics of East African puddle frogs (Anura: P hrynobatrachidae). Systematics and Biodiversity 8 39 47.

PAGE 350

350 BIOGRAPHICAL SKETCH Josiah (Joe) Townsend received his Ph.D. in Interdisciplinary Ecology from the University of Florida in Fall 2011, and had previously earned his Master of Arts in Latin Americ an Studies and Bachelor of Science in Wildlife Ecology and Conservation, both from the University of Florida. Joe was born in Enid, Oklahoma in 1978, the son of Steve Townsend of Ferriday Louisiana and Terri Boyd Townsend or Medford, Oklahoma After livin g in Medford for his first year, Joe and his family moved to Kenner, Louisiana, and then to Sulfur, Louisiana, where they were joined 198 1 by his sister, Katielynn Boyd Townsend. They soon relocated to Bethel Park, a suburb of Pittsburgh, Pennsylvania, whe re Joe attended Logan Elementary School It was here that Joe spent his childhood and develop ed his fascination with biological diversity in the halls of the Carnegie Museum of Natural History and the creeks and woods of south western Pennsylvania They rel ocated briefly to Pembroke Pines, Florida in 5th grade then to Hauppauge, New York from 6th through 8th grade and finally to Miami, Florida in 1992 where they moved the same week that Hurricane Andrew struck. In Miami, Joe attended Southwood Middle Scho ol and Miami Palmetto Senior High School, where he graduate in 1996. Joe attended Miam i Dade Community College after high s chool while helping to manage a youth hockey league. He began his studies at the University of Florida in 2000. He married Ileana Ro sario Luque Montes o n the 4th of July 2009 at his parent s house in Maryland, and they lived happily ever after.