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Amphibians of Central America

Gunther Kohler

Caver photos: Front cover, leR: AgaIychk spurrelli (for locality see Fig. 484). Photo: G. Kijhler Right top: Bolitoglossa salvinii (for locality see Fig. 108). Photo: G. Kohler Right bottom: Dendmbates p u d o (for locality see Fig. 4). Photo: G. Kijhler

Photo p. 4: Cruziohyla dcar i fe r (for locality see Fig. 490). Photo: G. K6hler

Back cover: Anotheca spinosa (top; for locality see Fig. 510). Photo: G. Kijhler Hypsiboas rufitelus (middle; for locality see Fig. 566). Photo: J. Sunyer Ecnomiohyla fimbrimembra (bottom; for locality see Fig. 539). Photo: A. Hertz

I(lihler, Gmther Amphibians of Central America Offenbach: Herpeton Verlag, 201 1 ISBN 3-936180-33-4

All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the publisher.

No liability is assumed with respect to the use of information contained herein.

8 2011 Expetan, Verlag Elke Kahler, lbhdz 22, D.63075 menbach, Germany

Eke JGhk, Wenbach

Gunther Kiihler

Amphibians of Central America

930 Color photographs 92 Drawings and diagrams

177 Maps

H~AP~TVN Verlag Elke Kohler

%word ............................................................ 6 bntroduction ...................................................... 10

..... $bmments on the Usage of this Book 11 Classifkation of the Amphibians of

............................................. :Central America 14 ........................................... The Environment 16

..................... Climate in Central America 24

................................................... . hphibia 26 y k .ceecilians (Gymnophiona) ........................ 27

............................. &lamandem (Caudata) 36 F

.......................... : Phga and Toads (Anura) 93 ......................................................... ' Bufonidae 96

........................................ ........ Centrolenidae ; 117 ................................................ Dendrobatidae 128

.......................................................... Terrarana 140 .............................................. Craugastoridae 158

..................................... Eleutherodactylidae 181 ............................................. Strabomantidae 186 ............................................. Hemiphr actidae 192

.............................................................. Hylidae 195 ...................................................... teiuperidae 273

.............................................. [ teptodactylidae 276 ................................................... Microhylidae 282

............................................................... Pipidae 290 .............................................................. Ranidae 291

............................................. Rhinophrynidae 296

The Tadpoles of Central American ...................................................... Anurans 298

Amphibian Declines and ................................................. Ektindiom 315

................................... Acknowledgments 327 ....................................... Literature Cited 329

....................................................... Glossary 370 Addendum .................................................. 373 Index ............................................................. 374

Foreword

Some seven years ago, I had the honor and pleasure to write the foreword for the first edition of Gunther Kohler's Reptiles of Central America. I was equally pleased he asked me to contribute the foreword for the second edition of this landmark publication, which appeared five years later. The fact that only five years separate the publication dates for the first and second editions of this book is a testament to the rapidity with which herpetologists are adding to the knowledge base about reptiles in the highly biodiverse, yet relatively small region of the Central American tropics.

Now, Dr. Kohler has undertaken the daunt- ing task of creating a companion treatment of the region's amphibian fauna. In some ways, completion of this task is more urgent than the one involving his earlier book. Current research initiatives are exposing previously unrecognized diversity among the amphibians of Central America at a pace comparable or perhaps even in excess of the rate a t which this activity is transpiring among reptiles. This is especially the case among salamanders and anurans, most prominently those inhabiting highland regions of Central America. Molecular systematic techniques have become indispensible in this research. At the same time, concurrent research priorities are demonstrating that the impos- ing diversity of the Central American amphibian fauna is under severe threat, both from the well known anthropogenic effects, especially habitat degradation and destruction, and the impact of the lethal disease chytridiomycosis caused by the chytrid fungal pathogen Ba trachochytrium dendrobatidis (Bd). The synergy between these two major causes of amphibian population decline appears to have reached a point a t which new species are being described after they have disappeared or

just as they are about to do so. An example of the former occurrence involves the Costa Rican endemic Atelopus chirnpoensis, the description of which appeared in 2009. This harlequin frog is known from a single specimen collected in 1980 and has not been seen in the intervening 30 years. Thus, this anuran follows the same pattern documented in so many other members of its genus. The latter occurrence is exempli- fied by the Panamanian endemic fringe- limbed treefrog Ecnomiohyla rabborum. This spectacular gliding frog was described in 2008 on the basis of specimens raised in captivity, inasmuch as its type locality lies in the El Cop6 highland area of central Panama, an area in which amphibian populations have been devastated by Bd. Investigators who were able to move in advance of the southward-moving front of chytridiomycosis have documented the dis- astrous impact of the sweep of this disease through the El Cop6 region by comparing the makeup of the amphibian community before and-after the e~idemic. Thev concluded that abundance, species divirsity, and evolutionary history were decimated by the fungal pathogen's effect. Thus, the fungal pathogen is expressing its lethality over the course of both ecological and evolutionary time.

Although a large amount of attention has been directed toward documenting the progress of chytridiomycosis through anuran populations, it is only one aspect of the overall problem of amphibian population decline, albeit an extremely significant one, as discussed by Dr. Kohler in an important chapter in this book. I can draw an example of the impact of habitat degradation and destruction from my own work and that of my colleagues in Honduras. Over the last four and one-half decades, we have documented an impressive amount of species-level herpetofaunal endemism in

Foreword

this country that exceeds the proportion seen in any other country in Central America. In Honduras, the percentage of amphibian species endemism is higher than that for reptiles and recent fieldwork is demonstrating that the increase in recognition of additional endemic species is higher for amphibians than for reptiles as well. Currently, 48 endemic salamanders and anurans are known to occur in Honduras of a total amphibian fauna of 130 species. More than a third of these species are limited to a single mountain range in the country, the Cordillera Nombre de Dios. Present and past fieldwork has centered in three areas within this range in the northern portion of the country, all of which lie within ostensibly protected areas. Two of these three areas -- - have been subjected to serious anthro-

AgaIYChniS amae de Bim,, pogenic environmental degradation Heredia, Costa Riea, 1630 m). Photo: G. K6hIer ing unsustainable logging and agricultural practices. As a consequence, population decline of several endemic amphibian species has occurred. Our most recent fieldwork in the third of these areas has taken place where there has been relative- major causes, i.e., chytridiomycosis, habi- ly little disturbance and where we have tat degradation and destruction, and glob- been fortunate to document robust popula- al climate change. Evidence is accumulat- tions of endemic amphibians whose popu- ing to indicate that these three causes are lations have been decimated in the other synergistically intertwined, in effect creat- two better-studied areas in the Cordillera ing an environmental super-problem that Nombre de Dios. grows more resistant to solution with time. Tropical salamanders also are known to be These environmental agents are bringing involved in the global amphibian crisis and about losses in diversity at many levels of it is hypothesized that global climate biological organization existing both change might be responsible for document- through space and time. It is telling that ed drastic declines in salamander abun- they are occurring at the same time that dance, community structure, and phyloge- new techniques for exposing biological netic diversity, especially evident in one of diversity and evolutionary history are the best studied and most diverse commu- being developed and employed that depend nities along an elevational transect on the upon the continued existence of robust southern slopes of Volcan Tajumulco in the communities of organisms, including Guatemalan department of San Marcos. amphibians. Such losses are not simply On the other hand, little is known about just a matter of a species of frog, salaman- how caecilians have been affected by these der, or caecilian being on Earth one day environmental pressures, as Dr. Kohler and gone the next, as regrettable as that has noted elsewhere in this volume. might be; the results can exacerbate to the

level of irreparable changes in community Mesoamerican amphibian population structure as measured over ecological time decline appears to involve at least three and, beyond this, to losses of pages of phy-

logenetic history as measured over evolutionary time.

Examples of similar situations can be documented throughout Central America. In fact, these stories are more and more rapidly becoming the norm in one of the planet's most important biodiversity hotspots. As a result of these develop- ments, a premium has been placed on cat- aloguing this diversity as we now understand it, with a view to updating this understanding as time advances. Thus, Dr. Kohler7s book comes at an opportune time in providing us a snapshot of what we know about Central American amphibian diversity today. Subsequent editions, if the reception of this book follows that of the companion volume on reptiles, will perform a hugely valuable function of keeping the book's readers updated on what we know about composition and distribution of the region's amphibian fauna. To the extent that the reptile companion volume can be an indication, in the five years that separated the 2003 from the 2008 editions, the number of species in the area of coverage increased from 537 to 557, an increase of 3.7%. My prediction is that if five years separates the present edition of this book and the one that might follow, the percentage increase in species numbers will be higher for amphibians than for reptiles. An extended discussion of these issues impact- ing attempts at amphibian conservation in Central American is presented toward the end of this book.

It is not fortuitous that I have come to pen the words of this foreword or that Gunther has come to write the rest of the book. The stage for this association was set undoubtedly at the 2002 Mesoamerican Workshop for the Global Amphibian Assessment, which occurred at the La Selva Biological Station in Costa Rica. As fellow students of the Central American herpetofauna, we were asked to convene with a number of our herpetological colleagues to provide our professional opinions on the conservation status of the region's amphibian

fauna. Ultimately, this work contributed to the overall assessment of the state of the global amphibian fauna that produced the alarming conclusion that nearly one-third of the world's caecilians, salamanders, and anurans are threatened with extinction. Work subsequent to this workshop has only served to underscore this sorry state of affairs.

Dr. Kohler also contributed to the book I co-edited entitled Conservation of Mesoamerican Amphibians and Reptiles, published earlier this year. He collaborated with his then-student Javier Sunyer on the chapter dealing with the conservation status of the Nicaraguan herpetofauna. Javier and I spent time together, along with others, on a study of the herpetofauna of the Bosawas biosphere reserve in northern Nicaragua, work underwritten in part by Dr. Kohler. Thus, so it goes with our careers intertwining as time moves along.

I am very pleased that Dr. Kohler has asked me to provide the foreword for the present book. Another reason, in addition to those indicated above, is that I expect this book on amphibians will become as much a companion during my own fieldwork as has his reptile book. Those famil- iar with fieldwork, especially in Central America, are aware that space and weight are always at a premium in places in which it is often the case that one's gear must be transported on mules and horses andlor one's own back. The 6 l/2 x 9 114 inch for- mat is very convenient for transport in the field and the contents especially useful for the kinds of identifications necessary to help determine the best use of the usually limited amount of time available.

The organization of this book is similar to that of the Reptiles of Central America in that there are initial sections providing an introduction, comments on the usage of the book, and a discussion of the classification of Central American amphibians. These sections are followed by the longest portion of the book dealing with the various groups

Following these accounts is a long section pmviding a key to the tadpoles of the anu- ram of Central America that possess these free-living larval stages (none of the Terrarana do). Having written keys to tadpoles previously, I know they are not easy to construct, so this will be a particularly useful aspect of Dr. Kohler's book.

The next section is the comprehensive dis- a s i o n of amphibian declines and extinctions. This section is extensively documented and concludes with a list of further readings that will prove helpful to the reader who wishes more information on this complicated and still-evolving topic. The book concludes with a section of cited literature containing more than one thou- sand entries, and, finally, a glossary and an index to all scientific names.

This book is concerned with a group of animals that is being shown to be under increasing threat from synergistically operating environmental problems in a region that is part of one of the largest and most important biodiversity hotspots in the world. Amphibians in Central America

Foreword

of amphibians, beginning with caecilians (and the single family of them in Central America), continuing on with the salamanders (and their single Central American family), and finishing with the lengthy treatment of the 14 families of anurans. The arrangement of these treatments is similar to that in the reptile book, with a description of each family followed by a key to its Central American genera, a section on further pertinent readings, a description of each genus, a key to its species (all checked for accuracy and utility by Dr. giihler's students), further readings, and the accounts of the Central American species in each genus. As with the reptile Book, this part of the amphibian book is accompanied by good-quality photographs and maps to guide the user's Fig. 2. Bolitoglossa striatula (Rio San Juan, attempts at identification. About 87 per- Nicaragua, 30 m). Photo: G. Kohler cent of the regions' 498 species of amphibians are illustrated and all are mapped.

are highly diverse, widely distributed, and given to extensive endemism, but are undergoing population declines on a broad- ening front. In light of this reality, this book will provide an important basis from which to continue the study of not only the biology of these fascinating creatures but also the many threats they face.

I wrote most of this foreword at my home on the eastern edge of the Florida Everglades and finished it in Trujillo, Honduras, a favorite place of mine, in a set- ting with an easy view of the incomparable Bay of Trujillo and beyond it, the Bay Island of Guanaja.

Larry David Wilson 15 October 2010

Introduction

Central America has a large and complex amphibian fauna. It has been postulated that the Neotropics contain the Earth's largest reservoir of amphibian biodiversity (SANTOS et al. 2009). In the past decades, works have been published on selected groups of Central American amphibians (e.g., DUELLMAN 1970, SAVAGE & M. WAKE 1972, CAMPBELL & SAVAGE 2000, DUELLMAN 2001, SAVAGE & M. WAKE 2001) as well as on the herpetofauna or amphibians alone of selected countries or regions (e.g., MERTENS 1952a, LEE 1996, CAMPBELL 1998, MCCRANIE & WILSON 2002, SAVAGE 2002, GUYER & DONNELLY 2005, MCCRANIE et al. 2005, KOHLER et al. 2006). FLORES-VILLELA et al. (1995) published a compilation of identification keys for the herpetofauna of Mexico. As presently understood, the amphibian fauna of Central America consists of 498 species representing 72 genera and 16 families (Table 1). This breathtaking diversity is due to the rich physiography, exceptional vegetational variation, and the vivid geological history of Central America; the geological history of the region was summarized in KOHLER (2008) and is not repeated here.

During the past ten years, we have wit- nessed drastic changes in the taxonomy of amphibians (see summary of changes during the past 20 years by JOHNSON et al. 2010). Some species have undergone repeated changes in the generic allocation during the past few years (i.e., the marine toad has been variously assigned to the genera Bufo, Chaunus, or Rhinella; those bufonid species allocated to Incilius today, have been assigned to Bufo, Cranopsis, or Ollotis just a few years ago). The recent - and most probably ongoing - period of taxonomic change (WIENS 2007, FROST et al. 2008) is due mostly to new research approaches, basically in the field of molec-

Isthmus of

I El Salvador 7

Fig. 3. The zoogeographic region of Central America (shaded).

ular genetics. This book has the purpose to accomplish an up-to-date summary of the diversity and taxonomy of the amphibian fauna of Central America and provide identification keys to the species level. The contents of this book can only serve as a momentary snapshot of the status quo; new discoveries and a better knowledge of the phylogenetics of the taxa treated here undoubtedly will usher in numerous taxonomic changes in the near future.

Revisionary work is needed urgently in several taxonomic groups of Central American amphibians. Examples are the issues of taxonomic identity of the Panamanian populations of frogs referred to the genera Elachistocleis and Hyloxalus, respectively. Some authors have suggested that Relictivomer might be a synonym of Elachistocleis, although without presenting evidence to support this opinion (e.g., ZWEIFEL 1986, LAVILLA et al. 2003); morphologic differences among species currently assigned to either genus are small and molecular genetic data could be useful to clarify this issue. Other clusters of morphologically poorly differentiated species with partly questionable validity of certain taxa include some caecilians (e.g., Oscaecilia elongata and 0. osae), some salamanders (e.g., some taxa

the dunni, rostrata, and morio species Comments on the Usage ofthis exes, respectively) and some frogs some taxa in the Ecnomiohyla mil- Book species group, as well as those in the us coccifer complex, the Craugastor

species group, the Lithobates pip- This book treats the species of amphibians species group, the genera Hypo- known to occur within the geographic lira-

pechus and Crepidophzyne, and also the its of Central America, here defined as the d e m e l y similar Smilisca species, S. sila geographic area between the Isthmus of and S. sordida). These are just a few of the Tehuantepec (level of narrowest portion) 'numerous taxonomic problems that pre- and the Panama-Choc6 border. Subspecies vail among Central American amphibians. are not considered. I have adopted the tax- f hope that this book will stimulate onomic arrangement as proposed by FROST ,wearch and conservation activities on et al. (2006), modified by those in the '*e amphibians in the Central American works of FAIVOVI~H et al. (2005), GRANT et m t r i e s . Only on the basis of sound tax- al. (20061, WILKINSON & NUSSBAUM (2006), ammy and a thorough understanding of SANTOS et al. (2009), and FAIVOVICH et al. the interactions with the environment of (2010). As to which species to include, I the species in question can successful con- have accepted all published species unless remation strategies be developed and there is convincing evidence to the con- "implemented. trary available in print.

During the past decades, dramatic declines in amphibian populations have led to population crashes and extinctions &species worldwide and Central America has been hit particularly hard by this mass extinction phenomenon (WAKE & VREDENBURG 2008, COLLINS & CRUMP 2009). The seriousness and urgency of this Inpic made it mandatory to include a dis- m i o n of the amphibian decline issue in &is book.

The intent of this book is to provide an identification guide to the numerous amphibian species found in Central America, as well as for higher taxa (genera and families), for both scientists and trav- elers. For this purpose, identification keys are provided to all families, genera, and species of amphibians that are known to occur in Central America, even though this is not explicitly stated in the respective key headings. These keys are supported by line drawings, schematic illustrations of characteristics, and numerous color photographs of the majority of

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Introduction

species in life. Some Central American species of amphibians are so distinctive in coloration, as well as body size and shape, that an animal in question often can be identified readily by photographs alone. However, identification keys provide additional information on the morphology andlor coloration of the respective species and should be consulted. Additionally, the distribution maps can be included in the identification process, because many species have very restricted distributions. The keys are dichotomous, meaning that they provide pairs of contrasting alternatives (couplets). Such a key works by using a step-by-step elimination of alternatives until only a single alternative, the solution, remains. It is important to remember when using the identification key that the juveniles of many amphibian species have different coloration and pattern than the adults and might lack characteristics present in the adults. Therefore, in many cases, the keys might not work to defini- tively identify juveniles. The key to the tadpoles also is challenging to employ, since tadpoles are significantly smaller than the adults of the same species. It is most often necessary to use a hand lens or even a microscope to be able to see the dis- tinguishing features sufficiently well. The technical terms and morphological character designations used in the keys can be unfamiliar to non-specialists, but I am optimistic that anyone willing to invest some effort in learning the terms will be able to use the keys with confidence and success. The glossary included in the book will be helpful in this regard, as will the illustrations accompanying the keys. Abbreviations used are CCL (combined fore and hind limb length); C W C (Costa Rican Amphibian Research Center); HFW (hind foot width); HL (head length); HW (head width); P.N. (Parque National); and SVL (snout-vent length; distance from tip of snout to vent; to distal end of vent in salamanders). Numbers of teeth refer to total number (both sides summed), if not otherwise indicated. The term "limb interval", used in the keys to salamanders, indi-

cates the number of costal folds between adpressed fore and hind limbs; the total count of costal folds includes one each in axilla and groin.

For some species-rich taxa (i.e., Bolitoglossa, Oedipina, and Terrarana), I have provided two keys, one that treats the species distributed from southern Mexico to Nicaragua, and the second covering the species known to occur in Nicaragua, Costa Rica, and Panama. Nicaragua is included twice in these keys because zoogeographically, northern Nicaragua belongs to Nuclear Central America (sharing with the northern countries species such as Bolitoglossa mexicana, Ptychohyla hypomykter, Plectrohyla sp., Mesaspis morele tii, Anolis beckeri, Tan tillita lin toni, etc.). Southern Nicaragua, on the other hand, has many elements that are otherwise only found in Lower Central America, such as Oophaga pumilio, Dendrobates aura tus, Lepidoblepharis xanthos tigma, Anolis pen taprion, etc.). Thus, Nicaragua, much of which comprises the Nicaraguan Depression, is somewhat "transitional" between Nuclear Central America, and Lower Central America. It would be arbi- trary to divide the country and make the split, for example, in central Nicaragua.

After the introduction to each major group of Central American reptiles, the following information is provided in the species accounts: the current scientific name with author and year of description (in the case that the author and year of description are enclosed within parentheses, this signifies that the original generic name has been changed to the current one), followed by the citation of the location of the original description of the species and a statement of the type locality; the maximum size (usually SVL), and the known geographic and elevational distribution. Detailed anatomical descriptions are beyond the scope of this book because it is intended for use as a field guide. The recommendations for further reading at the end of each

(near San nam6a Be La Fwgen, Hmdia, Costa Rim). Photo: C4. KBhl~r

The index at (19711, h a clb; (3ms- (19761, ALTIG inhdhgthe (19871, DUELLMAN & CAMPBELL (19921,

, mpple- be an w u s d addition to a field guide descrip- such as thig one, I would be remiss to net

Class3cation of the Amphibians of Central America

ClassXcation of the Amphibians of Central A m h

I ~ m ~ h i b i a (Amphibians)

Bolitoglossa (86) Bradytriton (1) Cryptotriton (6) Dendrotriton (8) Nototriton (14) NyctanoLis (1) Oedipina (29)

Dseudoeurycea (6

Family - Genus (in parentheses is the number of species

that occur in Central America)

claaacation ofthe Amplu'bh of Central America

L : , I 8

~lophryniscus (1 Atelopus (8) Chaunus (2)

Crepidophryne (3) Incilius (22) Rhaebo (1) Rhinella (2

Espadarana (1) i Ameerega (1) Hyalinobatrachium (7: Colostgthus (3)

Sachatamia (2) Dendrobatm (9) Teratohyla (2) Hyloxalus (1)

PhyUobates (2) Phyllodactylus (31

miphractus (

Gastrotheca (2

Agalychnis (6) Anotheca (1)

Bromeliohyla (1) Charadrahyla (1)

Cruziohyla (1) Dendropsophus (5) Duellrnanohyla (7)

Exerodonta (5)

I Diasporus (6 )

Eleuthe,m&&pJm I

- -'dae Leiupenaae eurodema ( I

% I Engystomops (

The Environment

The Environment

The various mountain complexes that form the physiography of Central America can be divided into two major massifs. The northern block includes the Mesa Central of Chiapas, the highlands of central and southern Guatemala, and those of Honduras, El Salvador, and northern Nicaragua. These are separated from the southern highlands by the Nicaraguan Depression. The southern highlands include the Cordilleras de Guanacaste, Central, and Talamanca, as well as the relatively low highlands of Dari6n in eastern Panama. The two large Nicaraguan lakes (Lagos de Managua and Nicaragua) are the largest inland bodies of water in Central America, with surface areas of 1035 km2 and 8157 km2, respectively. The diverse landscape of Central America supports very different regional climatic situations (DUELLMAN 1966, STUART 1966). On the Pac5c versant of Nuclear Central America, the dry season extends from January through April. A subhumid corridor, made up predominantly of dry forest or the remains thereof, stretches along the Pacific versant of Central America from the Isthmus of Tehuantepec through northwestern Costa Rica. Generally, the Caribbean versant experiences an equable, tropical climate with relatively mesic conditions throughout the year. Exceptions are the northern part of the Yucath Peninsula, as well as some interior valleys, which experience a more arid climate. The wettest regions of Central America are found in southeastern Nicaragua and northeastern Costa Rica, which receive an excess of 6000 mm precipitation per year.

' J B*

Rainfomst , " - * - ?

The beauty and majesty of a primary tropical rainforest or an old mature growth can be breathtaking. Tropical rainforest is found in regions with monthly temperatures exceeding 18 "C during all months of the year and an annual rainfall of more than 2000 mm. Rainforest soils are typically poor, due to leaching of soluble nutrients. In the northern parts of Central America, rainforest is found mainly in the Caribbean lowlands. In Costa Riea and Panama, it also covers large areas of the Pacific versant. During the day, temperatures in the tropical rainforest reach 3045°C. During

Fig. L. , iooded rainforest at Rio San Juan, Nicaragua (30 a elevation). Photo: G. Kijhl81'

The h-at

Cloud fontst

eavyr- moist forest characterized by frequent aftemom. mist or clouds at the vegetation level.

I Photo: G. Ktihler

The Environment I streams channel the water on its way to the lowlands. At very high altitudes along 1 exposed ridges, a special sub-type of cloud forest grows, the so-called elfin forest. Subject to low temperatures, high precipitation, and high wind speeds, trees in the elfin forest usually attain less than 1-3 m in height. The fauna of cloud forests tends to be less diverse than that of the lowlands, but there is a high number of endemic species, i.e., species that only occur in one particular area, such as a summit or a mountain chain.

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Dry forest Dry forests are typical of the Pacific ver- I sant of Central America and a few interi- - 7s or valleys of Honduras and Guatemala. ~~ The climate is characterized by low Pre- Fig. 7. Cloud forest at La Never*, Ngbbe cipitation (500-1200 mm) and long dry ~ u g i ~ , Panama, 1700 m. Photo: G.

Fig. 8. Dry forest near Arenal, Motagua Valley, Guatemala, 70 m. Photo: G. Kiihler 1

The Environment

last several months and vary hic location The precipita- seasonal and heavy down- ical during the rainy season.

! dry seaon, when precipitation le, a majority of the trees shed

permitting sunlight and ate the ground layer. Only

r rivers contain water throughout : whereas smaller streams are

* dry during the dry season. mals of this extreme habi-

sll adapted to the prolonged wiods. Dry forests consist of a iand cover of succulents and leistant shrubs. Tree heights

not exceed 20 m, and many of m armed with thorns. A curious

e formation is the presence of which ofken reach heights equal

Marshes, swamps, and seasonal flooded forests Aside from streams, ponds, and lakes, there are also aquatic habitats in Central America that are characterized by frequent or continuous flooding. These habitats include marshes, swamps, and seasonal flooded forests, which constitute important habitats for amphibians, especially frogs. In these wetlands, typically the water is shallow and features grasses, rushes, reeds, typhas, sedges, other herbaceous plants, as well as low-growing shrubs. A common feature of swamps and marshes is water stagnation. In so-called swamp forests, the seasonal flooding replenishes the soils with fertile silt.

- habitat between Hatt~evllle and ~ e l i z e City, Belize, 5 m. Photo: G. Kohler

The Environment - I

Fig. 10. P: pesrubra.

Cerro de la Muerte, Cartago, Costa Ri habitat of Bolitogll~ssa Photo: G. Kiihler

Fig. 11. Lagune near the entrance to P.N. Juan Castro Blanco, Alajuela, Costa Rica, 1915 m; breeding site of Lithobates vibicarius and Isthmohyla pseudopuma. Photo: G. K6hler

- aland ST

' Mesa Central de Chiapas \

Cordillera Nombre de Dios

E 4

Sierra Madre de Chia~

Guatemalan Plateau

Nicaraguan Depression

Cordillera Central

Cordillera de Talamanca

Cordillera de Guanacaste i

Cordillera de Tila*

vergreen rain forest (Caribbean versant)

Evergreen rain forest (Pacific versant)

Cloud forest f l . '

Dry forest

Pine-oak forest

Pine savanna

I B I

The Environment

Climate in Central America

The climate in Central America is highly location dependent, with the yearly cycle largely characterized by variations in precipitation levels. On the Caribbean side, the heaviest rains fall during December and January, while the Pacific side receives virtually no rain during the northern winter (the dry season on the Pacific side lasts from January until April). In general, the Caribbean side of Central America is more humid and receives more precipitation than the Pacific side.

The following climate diagrams illus- trate the yearly climate in selected 3 wo areas. The mean monthly precipitation - is indicated by bars. The mean daily f

j"'@' temperature can be read from the upper g curve and the mean nightly temperature from the lower curve (data from M ~ E R 1996, except for Utila, which was collec- 250

ted personally). 200

160

100

m

0 J F M A M J J A S O N D

- 160 -. -- 10 - - - 1W -- -

5 MI --

J F M A M J J A S O N D

10

6

0


The Environment 5 -



Amphibia

Amphibia

The extant species of amphibians are classified in three orders: (1) Anura (frogs and toads; about 5860 species); (2) Caudata, also called Urodela (salamanders; about 600 species); and Gymnophiona (caecilians; about 180 species). Amphibians inhabit every continent except Antarctica, with the highest diversity found in the tropics and subtropics (DUELLMAN & TRUEB 1986). Amphibians not only inhabit wetlands such as streams, ponds, lakes, and marshes, but they also live in deserts and high mountains. The highest record in the world for amphibians is held by Pleurodema marmorata, which has colo- nized newly formed ponds in recently deglaciated terrain at 5244 m in Peru (SEIMON et al. 2007). Amphibians play an important role in energy flow and nutrient recycling. Most amphibians are oppor- tunistic feeders. They eat just about any live prey they can fit into their mouths.

independent of water bodies for breeding and instead have a life cycle in which the eggs are laid in non-aquatic environments (e.g., in leaf litter, under rotten tree trunks), and the embryos develop within the egg directly into miniatures of the adults (WELLS 2007). Reproduction. in most lowland anurans is seasonal and occurs during the rainy season; some species call but only few (e.g., Smilisca sila, S. sordida) reproduce during the dry season (DONNELLY & GUYER 1994).

Key to the orders of adult a m p h i i 1 a No legs present; a protrusible tentacle pre-

sent on the side of the head between n w ......................... tril and eye Ggmnophiona

b Legs present; no tentacle present on the side of the head between nostril and eye ................................................................... 2

................................ 2 a No tail present Anura b A tail present ............................... Caudata

Some species, however, such as dendrobatids and many Neotropical microhylids Rsading

TAYLOR & GU~CTMAN 1977, DUELLMAN & TRUEB feed primarily on termites and ants. 1986, FEDER & BURGGREN 1992, STEBBINs & Amphibians display an amazing variety of COHEN 1995. MCDIARMID & ALTIG 1999. reproduction modes. Some species are DUELLMAN 1999, WEB 2007, OGIELSBA 2009

'

Fig. 14. Boliloglossa lir~colni (Nenton, Huehuetenango, Guatemala). Photo: J. Sunyer

Gymnophiona

Caecilians (Gymnophiona)

Caecilians are wormlike, limbless amphibians that are highly specialized for burrowing. At first glance, most people would probably mistake a caecilian for a large earthworm. However, they are readily distinguished from the latter by the presence of a well-developed mouth, teeth, and nostrils (M. WAKE 1983, ZHANG & M. WAKE 2009). Caecilians can be differentiated f h m wormlike reptiles such as amphisbae- nians and blind snakes by their scaleless Zmooth skin (GOWER & WILKINSON 2008). Ln caecilians, the body is segmented by annular grooves and folds. They have a rery short tail or no tail at all, and the vent is nearly terminal to terminal. The eyes are reduced in size and covered with skin or bone (M. WAKE 1983). The left lung is rudimentary or absent; both lungs are absent in the South American species Caecilita iwokramae and A tretochoana eiselti (WAKE & DONNELLY 2010). Caecilians have a protrusible, chemosenso- ry tentacle, which is located on the side of the head between the eye and the nostril ~BILLO & WAKE 1987). The animals contin- ually extend and withdraw the tentacles in order to sense the environment (M. WAKE 1983). When protruded, the tentacle picks up chemical particles from the environment. When it is retracted, the tentacle is pulled inside the tentacle aperture and the opening closes over. The chemical particles are then carried to the olfactory epithelium of the vomeronasal organ (SCHMIDT & WAKE 1990). There is no apparent external sexual dimorphism (SAVAGE 2002).

There are about 180 living species of caecilians worldwide, grouped into six families: Caeciliidae (104 species), Ichthy- ophiidae (46 species), Rhinatrematidae (10 species), Scolecomorphidae (6 species), Typhlonectidae (13 species), and Uraeo- typhlidae (7 species) (WILKINSON & NUSS- BAUM 2006). Caecilians are circumtropical

in distribution, but are absent from Madagascar, the eastern East Indies, New Guinea, Australia, and the Pacific islands. Fifteen species of caecilians are known to occur in Central America, all belonging to the family Caeciliidae (SAVAGE & M. WAKE 2001). The species are found in lowland to premontane regions, from sea level to about 1500 m, and rarely as high as 2000 m above sea level (in Dermophis costaricense; SAVAGE & M. WAKE 2001).

Because of their secretive lifestyle, relatively little is known about the natural history of caecilians. These are fossorial species that are rarely encountered on the surface. During the rainy season, they can occasionally be found under fallen logs, leaf litter, and other surface debris. After excep- tionally heavy rains they may be moving through the leaf litter or on the ground. Caecilians feed mostly on earthworms, termites, and soft-bodied larvae or instars of insects (M. WAKE 1983). A specimen of Dermophis mexicanus that was kept alive a t the Research Institute Senckenberg for about two years readily accepted earthworms. The most important predators on caecilians are snakes, especially coral snakes (genus Micrurus), and occasionally birds (M. WAKE 1983).

Caecilians exhibit a diversity of reproductive modes, including oviparity with aquatic larvae, oviparity with direct development, and a t least two modes of viviparity (LOADER et al. 2003). Caecilians have internal fertilization. The protrusible organ of the males, the phallodaeum, is an eversible portion of the cloaca, which is inserted into the female's vent during copulation (M. WAKE 1980). Some species of the family Caeciliidae for which reproductive data are available, such as Dermophis mexicanus and Gymnopis multiplicata, are live-bear- ers (M. WAKE 1983). Development takes

place in the oviduct, and the embryos oviductal wall and on cells from the oviduc- exhaust the supply of yolk before they tal lining (M. WAKE 1977). Other caeciliids emerge fi-om the egg membranes. After the such as the South American Caecilia orien- embryos have hatched in the oviducts, they talis reproduce by orpary with terrestrial feed on rich secretions from the thickened eggs ( F m et al. 2004).

1 annular grooves

Y annular folds

/ eye

I tentacle nostril

Fig. 15. Lateral view of head and anterior body of a caecilian (Demophis mexicanus). Photo: G. Kahler; Drawing: M. Vesely

Fig. 16. Ventral view of the body of a caecilian. Drawing: G. Kohler

Caeciliidae

Caeciliidae Key to genera of caecilians 1 a Tentacle and tentacular foramen posi-

tioned immediately below nostril ............ 2 b Tentacle and tentacular foramen posi-

................ tioned well posterior to nostril 4 2 a No secondary folds; more than 220 primary

folds .............................................................. ... O~caecilia (in park 0. elongata, 0 . omel

b Secondary folds present; fewer than 210 ............................................ primary folds 3

3 a Primary annular grooves complete around body; grooves of primary folds contrastingly darker than ground color; eye not visible

..................................................... externally ......... &&a (in park 0. o c h u e a w )

b Primary annular grooves incomplete at least on anterior body; grooves of primary folds not contrastingly darker than ground color; eye usually visible externally ........... ........................................................ Caecilia

4 a Eye not visible externally (eye covered by bone); a pair of splenial teeth behind den-

..................... tary series of teeth G-oN b Eye visible externally; no pair of splenial

teeth behind dentary series of teeth .......... .................................................... Dezmophis

Further Reading DUNN 1942a, TAYLOR 1968, SAVAGE & M. WAKE 1972, NUSSBAUM 1979, NUSSBAUM & HOOGMOED 1979, MOORE et a1.1983, International Commission on Zoological Nomenclature 1987, DUCEY et al. 1993b, EXBRAYAT & HRAOUI- BLOQUET 1994, LYNCH 1999c, SAVAGE & M. WAKE 2001, GOWER & WEKINSON 2005

Fig. 17. CaeciJia isthmica (Reserva Natural Privada Chucanti, Distrito de Chepigana, Darien, Panama, 797 m). Photo: A. J. Crawford

Only four species of the predominantly South American genus CaeciLia occur in the southern portion of Central America. They are thought to have reached the region relatively recently, following closure of the final segments of the Panamanian seaway about 3.3 million years ago (SAVAGE & M. W m 2001). Species in this genus are moderate to large sized caecilians (total length to well over one meter). They are fossorial species that are found mostly when turning rotten logs, or digging into deep leaf litter or other forest debris.

Ca&a isthmica COPE 1877, Proc. Am. Philos. Soc. 17: 91; type locality: "East side of the isthmus of Daribn", h a m a . Total length to 578 mm. Eastern Panama and western Colombia, details on its altitudinal range are lacking.

Caecilia Jeucocephala TAYLOR 1968, Caecilians of the World: 404; type locality: "Virology Field Station, Rio Riposo, Edo. Valle del Cauca, south of Buenaventura, Colombia." Total length to 455 mm. Eastern Panama to northwestern Ecuador, 50-1000 m elevation.

Caecilia nign'cans BOULENGER 1902, Ann:Mag. Nat. Hist. Ser. 7, 9: 51; type locality: "the Rio Lita, 3000 feetn, northwestern Ecuador. Total length to 1030 mm. Southeastern Dari6n Province, Panama, through the Pacific lowlands of Colombia to extreme northwestern Ecuador, near sea level to 915 m elevation.

Caecilia voJcm' TAnoR 1969, Univ. Kansas Sci. Bull. 48: 315; type locality: "El Valle de Antbn, Cocl6, Panam6 (elev. 550 m)." Total length to 320 mm. Western and central Panama, 550-1180 m elevation.

Caecilia isthmica r Caecilia leucocephala

Caecilia volcani

$ Fig. 18. Caecilia volcani (Santa F6, Veraguas, Panama, 800 m). Photo: M. Ponce

Fig. 19. Caecilia nigricans (Rio Tuira at Rio Mono, DariBn, Panama). Photo: W. E. Duellman (Courtesy of the Biodiversity Instilate, University of Kansas)

Key to Ca& 1 a 150-188 primary folds; 42-62 secondary

............................ folds; total folds 196-252 ........................................ Caecilia nigricans

b 112-147 primary folds; 12-54 secondary ......................... folds; total folds 126-185 2

2 a A small species, total length to 320 mm; 112-124 primary folds; 14-37 secondary folds; total folds 126-156 C a d a wlm'

b Large species, total length to 570 mm; 118-147 primary folds; 12-54 secondary

......................... folds; total folds 143-185 3 3 a 131-147 primary folds; 12-21 secondary

folds; total folds 143-168 C a d a iethmica b 118-131 primary folds; 32-54 secondary

............................ folds; total folds 150-185 .................................. Caecilia leucooephala

Further Reading TAYLOR 1968, SAVAGE & M. WAKE 1972, 2001, SUMMERS 2001, SUMMERS & M. WAKE 2001

Dermophis

The six recognized species of the genus Dermophis are distributed on the Atlantic versant from Veracruz, Mexico, to western Honduras, and on the Pacific versant from Jalisco and Michoach, Mexico, to northwestern Colombia (SAVAGE & M. WAKE 2001). They are moderate-sized (total length to 600 mm) and relatively robust. Diagnostic characters of Dermophis include the positioning of the tentacle about halfway between the eye and nostril, the presence of secondary grooves, and the absence of both an unsegmented terminal shield and terminal vertical keel (SAVAGE & M. WAKE 2001). The eyes are visible in all species of Dermophis.

Just as with other terrestrial caecilians, all Dennophis are fossorial species that are only occasionally encountered on the surface. Specimens have been discovered under rotten logs and decaying plant material. Also, they are often found when digging in loose, friable, organic soil, as on coffee plantations, as well as in the leaf litter in tree buttresses and even under cow manure (M. WAKE pers. comm. September 2010). They feed primarily on earthworms and soft-bodied insects.

M. WAKE (1980) studied the reproduction and population structure of Dermophis mexicanus in western Guatemala. She reported that these animals reach maturity at an age of one year, and most females breed at the beginning of their second year and males at two years of age. Repro- ductive females are usually between 300 and 400 mm in total length (M. WAKE 1983). Fertilization occurred from late April to early June and the gestation period was one year. Two to sixteen young were born from May through June at the beginning of the rainy season. Size at birth was documented to be 108-155 mm total length (M. WAKE 1980). The juveniles feed on maternal nutritive secretions using specialized fetal teeth (M. WAKE 1980).

Females of D. mexicanus reach sexual maturity at a size between 300 and 400 mm in total length (M. WAKE 1980).

Demophis costaricense TAYLOR 1955, Univ. Kansas Sci. Bull. 37: 506; type locality: "Cinchona, [Canthn de Alajuela, Provincia AlajuelaJ Heredia Province Fn error], Costa Rica." Total length to 387 mm. Atlantic slopes of the Cordilleras Tilargn, Central, and Talamanca in Costa Rica, 1000-1300 m elevation.

Dermophis glandulosus TAYLOR 1955, Univ. Kansas Sci. Bull. 37: 509; type IocaIity: "San Isidro del General, [Cantbn de Perez Zeledbn,] San Jose Province", Costa Rica. Total length to 405 mm. Southern Pacific versant of Costa Rica to western Colombia, 400-2000 m elevation. Dermophis balboai TAYLOR 1968 is regarded as a synonym of D. glandulosus according to SAVAGE & WARE (1972,2001).

Dermophis gracilior GUNTEIER 1902, Biol. Centr. Amer., Rept. Batr.: 306; type locality: "Panama, Chiriqui." Total length to 345 mm. Southern PacSc versant of Costa Rica and adjacent western Panama, 50-1200 m elevation.

Demophis mexicanus ( D ~ R I L & BIBRON 1841), Erp. Gen. 8: 284; type locality: "Mexique"; restricted to "Cuatotolapam", Veracruz, Mexico. Total length to 600 mm. On the Atlantic versant of southeastern Mexico and disjunct in eastern Guatemala and northwestern Honduras, on the Pacific versant from Oaxaca, Mexico to central Nicaragua, near sea level to 1500 m elevation. Dermophis eburatus TAYLOR 1968 and Dermophis septentrionalis TAYLOR 1968 are regarded as synonyms of D. mexicanus, according to SAVAGE &WARE 1972, 2001.

Demophis occidentalis TAYLOR 1955, Univ. Kansas Sci. Bull. 37: 503; type locality: "Dominical Road approximately 15 km. WSW of San Isidro del General in open meadow under stones", San Jose Province, Costa Rica. Total length to 235 mm. Southern Pacific versant of Costa Rica, 50-970 m elevation.

Dermophis paniceps (DUNN 1924a), Occas. Pap. Boston Soc. Nat. Hist. 5: 93; type locality: "La Loma, on trail from Chiriqui Grande to Boquete, altitude about 2000 feet, Province of Bocas del Toro, Panam&" Total length to 217 mm. Atlantic slopes of Costa Rica and western Panama, also in parts of central Panama and on Isla Popa, Bocas del Toro, Panama, near sea level to 1200 m elevation.

- ,. 20. Dermophis parviceps (Gua, _..in, CRARC, Costa Rica, 520 m). Photo: B. Kubicki

A Dermophis costaricense Dermophis glandulosus

r Dermophis gracilior Dermophis occidentalis Dermophis parviceps

Fig. 21. Dermophis mexicanus (El Refugio, Ahuachaph, El Salvador, 225 m). Photo: G. Kohler

Key to Dennophis 1 a Annular grooves marked with dark pig-

ment ventrally, in sharp contrast with the otherwise pale venter; 94-112 primary folds; 35-88 secondary folds; total folds 94-112 .................... Demophis mexicanus

b Annular grooves not contrasting in color with overall ventral coloration; number of

............................................ folds variable 2 2 a Head distinctly paler (pink to whitish in

life, pale brown in preservative) than dor- .............. sal body; 11-37 secondary folds 3

b Head not markedly different in color than ......... dorsal body; 37-96 secondary folds 4

3 a 85-102 primary folds; 11-26 secondary folds; total folds 97-126 .............................. ................................... Dermophisparvicepe

b 95-112 primary folds; 29-37 secondary folds; total folds 126-149 ............................ ............................... Demophis ddentaZis

4 a 91-106 primary folds; 37-60 secondary ............................ folds; total folds 132-159

............................... Dermophis glandulo8us b 91-112 primary folds; 65-96 secondary

folds; total folds 159-208 ......................... 5 5 a 91-102 primary folds; 65-78 secondary

folds; total folds 159-176 ............................ ..................................... DermophiB pc i l io r

b 107-112 primary folds; 74-96 secondary folds; total folds 186-208 ............................ .............................. Dermophb crxstankeme

Fig. 22. Dermophis parviceps (Rara Avis, Las Horquetas de Sarapiqui, Heredia, Costa Rica, 700 m). Photo: T. Leenders

Fig. 23. Dermophis occidentalis (Palmar Norte, Puntarenas, Costa Rica, 25 m). Photo: P. Janzen

Further Reading TAYLOR 1955, 1968, 1969, SAVAGE & M. WAKE 1972. T A ~ O R 1973, M. WAKE 1980,1983, 1998,

-

Gymnopis I The genus Gymnopis is restricted to Central America and contains two species, G. multiplicata and G. syntrema. These are moderate-sized caecilians (total length to 480 mm) that have the eye covered by bone and not visible externally. The few specimens of G. rnultiplicata I have

; encountered so far were found during daytime beneath rotten logs (two specimens

, in southern Nicaragua and one in Punta Leon% Costa Rita); an additional speci- Fig, 24, Gpnopis mulwicata (punta L ~ ~ ~ ~ , men in was found during heavy Puntarenas, Costa Rica, 40 m). Photo: G. Klihler rains on the surface in a pasture. These caecilians feed mostly on earthworms and termites, as well as soft-bodied insect larvae (M. WAKE 1983). Gymnopis multiplica-

, ta gives birth to fully developed young.

! Litter size in G. multiplicata is 2 to 10, and the total length of the newborn varies

j between 100 and 120 mm (M. WAKE 1983).

%ymnopis m ultiplicata PETERS 1874, Monatsber. Preuss. Akad. Wiss. Berlin 1874: 616; type locality: 'Veragua", Panama. Total length to 480 mm. Guatemala to western Panama, also on Isla Cristdbal and Isla Escudo de Veraguas, Bocas del Toro, Panama, and on Isla del Caiio, Costa Rica, sea level to 1400 m elevation. Siphonops proximus COPE 1877 and Siphonops simus COPE 1877 are regarded as synonyms of G. multiplicata according to SAVAGE & M. WAKE 1972,2001.

Fig. 25. Gymnopis multiplicata (El Naranjc Atlantic0 Norte, Nicaragua, 150 m).

Photo: G. Kijhler

Gpnopi s syntrema (COPE 18661, Proc. Acad. Nat. Sci. Philadelphia 18: 129; type locality: "the neighboring region of Honduras." Total , length to 305 mm. Central and eastern ' Guatemala, and in the Mayan Mountains of southern Belize, 440-1000 m elevation. Minascaecilia sartoria M. WAKE & C ~ E L L k ~ ? 1983 and Siphonovs oligozona COPE 1877 are itF"lc- '

regarded as &nonbs oi' G. syntrema, accord- 1 ing to NUSSBAUM 1988 and SAVAGE & M. WAKE ,

I I

Fig. 26. Gymnopis syntrema (Guatemala). I

Photo: J. A. Campbell

Caeciliidae

Fig. 27. Oscaecilia ochrocephala (Rio Tuira at Rio Mono, Daribn, Panama).

Photo: W. E. Duellman (Courtesy of the Biodiversity Institute, University of h a s )

Key to Gymnopie 1 a Dorsum gray, annular grooves contrasting

pink or white; 128-132 primary folds; 63-93 secondary folds; total folds 193-206 .................................... G3m.Qo.w-

b Annular grooves not contrasting in color with overall dorsal coloration; 112-133 primary folds; 84-107 secondary folds; total folds 201-250 ........ Ggmno* multiplic~ta

Gymnopis multiplicata rnnopis syntrema

The primary distribution of the genus Oscaecilia is in northwestern South America, with only three species of the nine species in this genus occurring in the southern portion of Central America. The Central American species of this genus are moderate-sized, mostly very slender caecilians (total length to 620 mm) and as with the species of Gymnopis have the eye covered by bone and not visible externally. Oscaecilia elongata and 0 . osae are known from only a few specimens each. As far as is known, the Central American species of this genus are all fossorial forms that are found in humid situations under logs and within leaf litter and other debris; the holotype of 0 . osae was collected at night "fi-om a flooded path in a short-cropped grassy airstrip during a moderate rain. In contrast to the other two Central American species of Oscaecilia, 0. ochrocephala has been collected more frequently and appears to inhabit a diverse array of habitats; it was found as deep in soil as 10 m (when digging building foundations in Panama City), and has been observed on the ground surface at dawn and dusk with some frequency in Garnboa, Panama (M. W m pers. comm. September 2010).

Oscaecilia elongata ( D m 1942a), Bull. Mus. Comp. Zool. 91: 527; type locality: ''Panam&." Total length to 620 mm. Known only from "within the area of Yaviza, Dari6n Province, Panam6'' ( D m 1942) and from Los Algarrobos, Chiriqui (Fig. 29), 50-145 m above sea level.

Oscaecilia ochrocephala (COPE 18661, Proc. Acad. Nat. Sci. Philadelphia 18: 132; type locality: Panama. Total length to 617 mm. Panama and northwestern Colombia, near sea level to 610 m elevation.

Oscaecilia osae WAS & SAVAGE 1992, Copeia 1992: 703; type locality: "airstrip at La Sirena, Peninsula de Oea, Cant6n de Osa,

Further Rsading Puntarenas Province, Costa Rica (approx. 3 M. WAKE 1963, T A ~ O R 1968, SAVAGE & M. WAKE m)." length 382 rnm. GoLf~ 1972, T m o ~ 1973, M. wm & CAMPBELL 1983, region in Pacific southern Costa Rica, near sea NUSSBAUM 1988, M. WARE 1988a,b, SAVAGE & M. level to 40 WAKE 2001

28. Oscaecilia elongata (Lo r i d , Panama, 145 m).

&y to Oscaecilia 1 a Primary annular grooves complete around

body; 7-31 secondary folds present; 112-198 primary folds; body not extremely attenuate (ratio total length 1 diameter at midbody 40-60) ... OscaeciLia o c h m m w

b Primary annular grooves incomplete at least on anterior body; no secondary folds;

r id 226-232 primary folds; body extremely attenuate (ratio total length I diameter at

........................................ midbody 83-91) 2 # a Dermal scales present in posterior annular

grooves (visible in a X-ray image); 232 pri- .......................... mary folds Oscaecilia osae

b No dermal scales present in a ~ u l a r .................. grooves; 226-231 primary folds

Osca& elongata

. Fig. 29. Oscaecilia elongata (Los Algarrobos, ...................................... Chiriqui, Panama, 145 m). Photo: A. Hertz

garrobos, Fig. 30. Oscaecilia osae (La Gamba near rnoto: A. Hertz GoEto, Costa Rica, 60 m). Photo: P. Weish

Rnther Reading 1942a, T A ~ O R 1968, 1969, SAVAGE & M.

WAKE 1972, TAYLOR 1973, NUSSBAUM & WILKINSON 1989, LAHANAS & SAVAGE 1992, Fig. 31. Oscaecilia osae (La Gamba near WAGE & M. WAKE 2001 Golfito, Costa Rica, 60 m). Photo: P. Weish

Plethodontidae

Salamanders (Caudata)

Fig. 32. Bolitoglot 2esrubri . de 1 uerte, C - , uww . . ler

Salamanders comprise a diverse group of amphibians that includes terrestrial, arboreal, fossorial, and aquatic species. They are distributed from southern Canada to central Bolivia in the Americas, and in northern, central and eastern Asia, as well as Europe and northern Africa in the Old World. In Central America (narrowest portion of Isthmus of Tehuantepec to eastern Panama), salamanders are well represented by 151 species in seven genera, with Bolitoglossa being by far the most speciose genus.

Plethodontidae Plethodontid salamanders are characterized (among other traits) by the absence of lungs; therefore, respiration is primarily cutaneous and via the buccopharyngeal membranes. All tropical plethodontids have an extremely long tongue that works just like a true projectile (WAKE & DEBAN 2000). The tongue can be shot with great accuracy at prey (mostly small insects and

spiders) several centimeters away, reaching the target in a few milliseconds. This specialized biomechanical feeding system, combined with excellent vision, enables the salamanders to feed efficiently on prey that might be at some distance from the animal's head (-40% of the salamander's SVL) (WAKE 2005). Tropical plethodontid salamanders have extraordinarily low basal metabolic rates and, therefore, require relatively little food (FEDER 1983).

All Central American salamanders belong to the subfamily Bolitoglossinae within the family Plethodontidae. The species of this family are distributed .from Nova Scotia and extreme southeastern Alaska southward, reaching eastern Brazil and central Bolivia. A major center of diversity of this group is Mesoamerica (Mexico and Central America), with 13 currently recognized genera and about 220 species, which equals nearly half of all salamander species (WIENS et al. 2007, WILSON & JOHNSON 2010). In contrast, only two of the

- : :r. 33. Bolitoglossa dofleini (male; Nueva Concepcion, Senahu, mta Verapaz, Guatemala, 1 3 1000 In). Photo: J. Sunyer

=- r 34. Bolitoglossa mombachoensis (Volcan Mombacho, Granada, Nicaragua, 1100 in 1. A .-. Photo: G. Kohler

37

Fig. 35. Bolitoglossa salvinii (Coatepeque, Quetzaltenango, Guatemala, 530 m).

Photo: G. Klihler

currently recognized genera, with none endemic, and 28 species, are found in South America (WAKE 2005). The success of this group of animals is based partly on their reproductive strategy of direct development (no free-living aquatic larvae). After internal fertilization (the female picks up a spermatophore that the male has deposited), the female produces large and yolky eggs that are laid in a moist and humid, but non-aquatic environment, such as under leaf litter, fallen logs, or a similar moist and protected place. The development of the larvae takes place within the egg membrane, and after 42-169 days, the young salamanders hatch as miniatures of the adults.

Rey to the genera of Central American salamanders 1 a Shiny black with bright red spots on eye-

lids as well as orange-red spots on dorsal surfaces of body, limbs, and tail that become orange, then yellow, and finally cream-colored posteriorly in life (black with discrete pale spots in preservative), spots on head and shoulder regions almost

.... symmetrically arranged (Figs. 158-1601 Ngctanolis ....................................................

...................... b Color pattern not as above 2 2 a Costal grooves 17-20; body extremely elon-

gate (Fig. 161); limbs very short (limb ..... interval usually >7, rarely as low as 6)

...................................................... w - b Costal grooves fewer than 15; body not

extremely elongate; limbs moderate to well developed (limb interval usually c5, rarely as high as 6) .............................................. 3

........... 3 a Sublingual fold absent Balitog10888 b A distinct sublingual fold present (a fleshy

fold running transversely across the mouth below the tongue) ................................... 4

4 a Stocky body with no clearly defined neck; short, slender limbs bearing syndactylous hands and feet; tail laterally compressed .. ................................................. Brad~hftan

... b Combination of characters not as above 5 5 a Larger, more robust species (SVL in adults

usually >40 mm); feet broader (ratio HFWISVL >0.10); legs longer (ratio CCWSVL >0.5); ratio nostril diameter / SVL ~ 0 . 0 5 .......................... Pseudoewycea

b Diminutive, slender species (SVL in adults usually c35 mm, rarely to 39 mm); feet narrower (ratio HFWISVL usually ~0.10); legs shorter (ratio CCWSVL usually c0.5, or if

premaxillary teeth

An interesting antipredator strategy in plethodontid salamanders is their ability to autotomize their tails. After urotomy, the salamander regenerates a new tail, developing vertebrae and all other tissues except the notochord (WAKE & DRESNER 1967). This is unlike the autotomy in lizards where vertebrae are not regenera- ted after tail breakage (VITT & CALDWELL 2009). Whereas no salamander is able to see colors, most plethodontid species have some degree of binocular vision (SAVAGE Fig. 36. Roof of mouth in a salamander. 2002). Drawing: G. Kohler

Plethodontidae

.0.5, then ratio nostril diameter I SVL ........................................................ ~0.05) 6

Tostrils smaller, ratio nostril width / SVL .... :0.019; ratio CCWSVL usually ~ 0 . 4 6 7

b Nostrils larger, ratio nostril width 1 mL usually ~0.018, or if 40.019, then ratio

........................................ CCIJSVL >0.47 8 a Head narrower (ratio HWISVL ~0.16); legs

shorter (ratio CCLISVL usually c0.40, rarely to 0.45) Notokitan

I ...........................

b Head broader (ratio HWISVL >0.15); legs .................... longer (ratio CCUSVL >0.44)

................................ Dendlv~ton (in part) a Legs shorter, ratio CCIJSVL <0.47, or if ~ i ~ . 38. oedipina uniformis (Albergue

'0'479 then ratio nostril diameter ' Monterreal, near P.N. Juan Castro Blanco, <0.019 and ratio tail length I SVL >1.25; feet narrower (ratio HFWISVL 0.06-0.09) A1ajuela, Rita, lgoO m).

Ciyptotriton Photo: G. Kiihler ................................................ b Legs longer, ratio CCIJSVL >0.47; ratio

nostril diameter 1 SVL usually >0.019; ratio tail length I SVL usually c1.25; feet

......... broader (ratio HFW/SVL 0.09-0.12) ................................ Dendlvtn'ton (in part)

lrtber Reading INN 1926, TAYLOR 1944, WAKE & DRESNER 67, DODD & BRODIE 1976, WAKE & LYNCH V6, HANKEN et al. 1980, WAKE et al. 1980, -N 1983, WAKE & ELIAS 1983, ELIAS 1984, b g ~ 1987, &SEN et al. 1989, SESSIONS & EER 1991, WAKE et al. 1992, WAKE & HANKEN 96, BRUCE 2000, GARCIA-PAR~S et al. 2000, b g ~ & DEBAN 2000, BRUCE 2003, CHAN 2003, LTELLER et d. 2004

! rsal view

Fig. 40. Pseudoeurycea b m a t a (San Marcos, Guatemala). Photo: D. B. Wake costal grooves

cloaca

tral view V head length

Fig. 39. Foot of Bolitoglossa heiroreias; a. dorsal Fig. 37. External characteristics of salamanders. view; b. ventral view; note subdigital pads (red

Drawings: G. KiShler circle). Photos: G. Kohler

Bolitoglossa

Bolitoglossa is the largest and most widely distributed genus in the order Caudata and includes about 16% of the recognized salamander species (PARRA-OLEA et al. 2004). While most species of Bolitoglossa occur in Central America and Mexico, the genus ranges throughout most of tropical South America as well ( B m & WAKE 196313). These salamanders are found in environments ranging from lowland rainforests to cool, humid cloud forests and paramo of the Talamanca (to 3620 m elevation; SAVAGE 2002). Hands and feet have diversified greatly in the genus (WAKE & LYNCH 1976). Extensive webbing is usually associated with climbing behavior and semi-arboreal to arboreal life histories (WAKE & LYNCH 1976).

Bolitoglossa differs from the remaining tropical salamanders in lacking a sublingual fold (a fleshy fold running transversely across the mouth below the tongue in all other tropical genera and many North American species with freely projectile tongues), having an extraordinarily long and rapidly projected tongue, the so-called "mushroom tongue," (in association with a unique hyobranchial apparatus) and by the tendency of having tarsal reductions (resulting in broad hands and feet with relatively short, stubby digits) and extensive interdigital webbing (WAKE & ELIAS 1983). Monophyly of Bolitoglossa is supported by morphological and molecular genetic data (PARRA-OLEA et al. 2004).

Seven subgenera of Bolitoglossa are currently recognized (PARRA-OLEA et al. 2004): Bolitoglossa (12 species previously included in the mexicana species group), Eladinea (all South American and some Lower Central American species), Magnadigita (a morphologically heterogeneous assemblage including the species previously included in the d m ; f rankl i~~~, and rostrata species groups), Mayamandra (includes three species previously placed in

the veracrucis species group), Nanotriton (diminutive, short-tailed salamanders with small hands and feet: the two species previously included in the rufescens species group plus the recently described B. nymphal, Oaxakia (includes five species, previously included in the macrinii species group) and Pachymandra (includes two large to very large species with hands and feet that are fully webbed and first caudal vertebrae that have branched transverse processes: B. dofleini and B. alvaradoi).

Bolitoglossa alberchi GARC~A-PAR~S, PARRA- OLEA, BRAME &WAKE 2003, Rev. Esp. Herpetol. 16: 57; type locality: "near Playa Escondida, approximately 14 k m (road) NE Sontecomapan, Veracruz, MBxico, 18"56'N, 95"05W, a t ca. 50 m elevation." Maximum SVL in males 69.6 mm, in females 97.1 mrn; tail 1engthISVL ratio about 1.0. Southern Mexico in the Los Tuxtlas area, in east-central Veracruz, the Los Chimalapas area of extreme eastern Oaxaca, and the Caribbean slopes of western and central Chiapas, near sea level to 1080 m elevation.

Bolitoglossa alvaradoi TAYLOR 1954, Univ. Kansas Sci. Bull. 36: 604; type locality: "Moravia de Chirripo, [Cantbn de Turrialbal, Lim6n Province, Costa Rica." Maximum SVL in males 65 mm, in females 79 mm; tail lengeh/SVL ratio 0.92-0.98. Atlantic slopes of Costa Rica, near sea level to 1120 m elevation.

Bolitoglossa anthraciua BRAME, SAVAGE, WAKE & HANKEN 2001, Copeia 2001: 700; type locality: "north slope of Cerro Pando, Prov. Bocas del Toro, Panam6, elevation 1450 m, . . . Estimated coordinates are 8"58' N, 82"42' W." Maximum SVL in only known male 46 mm, in females 70.8 mm; tail length/SVL ratio 1.01-1.27. Western highlands of Panama, 1100-1450 m elevation.

Bolitoglossa biseriata TANNER 1962, Herpetologica 18: 19; type locality: "Rio Tuira a t mouth of Paya Tributary, Dari6n Province, Panama." Maximum SVL in males 36.7 mm, in females 48.7 mm; tail 1engtWSVL ratio 0.80-1.16. Panama to northwestern Ecuador (see recent record by CISNEROS-HEREDIA 2006), near sea level to 500 m elevation.

Bolitoglossa bramei WAKE, SAVAGE & HANKEN 2007, Copeia 2007: 561; type locality: "Costa Rica, Provincia Puntarenas, trail to Cerro Pando, above Las Tablas, 8"56'N, 82"46W, 2200 m." Maximum SVL in males 39.1 mm, in

1 41.0 mm; tail IengthISVL ratio 91-1.12. Western highlands of Panama and

$

r ~ g . 43. Bolitoglossa mvaradoi (Guayach, CRARC, Costa Rica, 500 m). Photo: B. Kubick

Fig. 44. Bolitoglossa bramei (Las Tablas, Punta- renas, Costa Rica). Photo: M. Garcia Paris

~hg. 41. Bolitoglossa alberchi (Berriozabal, Fig. 45. Bolitoglossa carri (Cerro Cantagallo, i" -epas, Mexico, 800 m). Photo: A. Ramirez V. Francisco MorazBn, Honduras).

Photo: J. R. McCranie

3 I Bolitoglossa biseriata (Bilsa Biological Fig. 46. Bolitoglossa cataguana (P.N. Montaiia

.lull, Esmeraldas, Ecuador, 550 m). de Yoro, Francisco Morazan, Honduras, Photo: H. M. Ortega-Andrade 1850 m). Photo: J. H. Townsend

Plethodontidae

Bolitoglossa celaque MCCRANIE & WILSON 1993, Herpetologica 49: 11; type locality: "near the Rio Arcaqual, eastern side of Cerro Celaque, Cordillera de Celaque (14"32'N, 88"40W), 2480 m elevation, Departamento de Lempira, Honduras." Maximum SVL in males 55.1 mm, in females 62.2 mm; tail length/SVL ratio 0.70-0.99 (males), 0.63-0.87 (females). Southwestern Honduras (Montafia de Celaque, Sierra de Opalaca, Sierra de Montecillos), 1900-2620 m elevation.

Bolitoglossa centenorurn CAMPBELL, SMITH, STREICHER, ACEVEDO & BRODIE 2010, Misc.

Huehuetenango, Guatemala, 3250 m (15.815000" N, 91.478000" W)". Maximum SVL in females 56.4 mm (adult males unknown); tail lengtWSVL ratio tail length

Publ., Mus. Zool., Univ. Michigan 200: 28; type Fig. 47. Bolitoglossa ,,~pacta (l.,.lan B a d , locality: "Cerro Bobic, SSE San Mateo Ix ta th , chiriqui, panama, 3000 m). photo: A. ~ ~ r t ~

0.73-0.94. Only known from the type locality

Bolitoglossa cerroensis (TAYLOR 1952a), Univ. Kansas Sci. Bull. 34: 724; type locality: "Cerro de la Muerte at approximately 7000 feet elevation, Pacific slope, 2 km. below Millville [= Villa Mills] on the Pan-American Highway", Cant6n de Perez Zeledbn, Cartago Province, Costa Rica. Maximum SVL in males 72 mm, in females 76 mm; tail 1engthISVL ratio 0.85-1.08. Cordillera de Talamanca, Costa Rica, 2530-2990 m elevation.

BoLitoglossa colonnea (DUNN 1924c), Field Mus. Nat. Hist. Publ., Zool. Ser. 12: 96; type locality: "La Loma, on trail from Chiriquicito to Boquete, altitude about 2000 feet, Bocas del Toro, Panama." Maximum SVL in males 54 mm, in females 53 mm; tail 1engtWSVL ratio 0.79-0.96. Atlantic versant of Costa Rica and western Panama, and Pacific versant of southern Costa Rica and Cerro Campana, Panama, from near sea level to 1250 m elevation.

Bolitoglossa compacta WAKE, BRAME & DUELLMAN 1973, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 248: 12; type locality: "north slope of Cerro Pando, 1920-1970 m (6298-6462 R) elevation, Provincia de Bocas del Toro, western Panama." Maximum SVL in males 53.4 mm, in females 74.2 mm; tail length/SVL ratio 0.79-1.00. Lower montane areas on or near the Costa Rica-Panama border, 16503000 m elevation.

Bolitoglossa conanti MCCRANIE & WESON 1993, Herpetologica 49: 4; type locality: "Quebrada Grande (15"05'N, 88"55W), 1370 m elevation, Departamento de Coph , Honduras." Maximum SVL in males 53.1 mm, in females 60.7 mm; tail 1engtWSVL ratio 0.75-1.11 (males), 0.72-0.91 (females).

Western Honduras (El Chaguiton and El Portillo in the Departamento de Ocotepeque; Quebrada Grande in the Departamento de Copb, and Parque Nacional Cerro Cusuco) and eastern Guatemala (areas of La Unidn in the Departamento de Zacapa, as well as Cerro Nylon and Cerro Negro Norte in the Sierra de Caral, Departamento Izabal) (S. ROVITO pers. comm. Nov. 2010), 950-2000 m elevation.

Bolitoglossa copia WAKE, HANKEN & IBAREZ 2005, Copeia 2005: 223; type locality: "Panama, Provincia de Cocle, Parque Nacional General de Divisidn Omar Torrijos Herrera, from near the summit of Cerro Pefia Blanca, approximate UTM coordinates 0541522 0957221, 1315 m elevation." Only known specimen has a SVL of 81.5 mm and a tail length , of 83.2 mm (tail length/SVL ratio 1.02). Known I only from type locality.

Bolitoglossa cuchumatana (STUART 1943a1, I

Misc. Publ. Mus. Zool. Univ. Michigan 56: 14; i type locality: "Oak forest about 2 kilometers north of Nebaj, [Departmento] El Quiche, Guatemala." Maximum SVL in males 43.5 mm, in females 57.5 mm; tail length/SVL ratio 0.69-0.87. Cordillera de 10s Cuchumatanes and the area of Chichicastenango, QuichB, Guatemala (S. ROWTO pers. comm. Nov. 2010), 1200-2500 m elevation.

Bolitoglossa cuna WAKE, B m & DUELLMAN 1973, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 248: 2; type locality: 'Camp Sasardi, 12 m (39 ft) elevation, Territorio de San Blas, Panama." Maximum SVL 55.7 mm; tail length/SVL ratio 0.94-0.97. Eastern lowlands of San Blas, Panama, sea level to 350 m elevation.

Bolitoglossa daryorum CAMPBELL, SMITH, STREICHER, ACEVEDO & BRODIE 2010, Misc.

Fig. .x. Bolitoglossa cuchumatana (Laj Chimel, ler QichB, Guatemala, 2266 m).

Photo: S. M. Rovito

--

'Pig. 49. Bolitoglossa cerroensis (Cerro de la Fig. 53. Bolitoglossa cuna (Nusagandi, Kuna = Huerte, Costa Rica). Photo: W. Van Devender Yala, Panama, 350 m). Photo: G. Kiihler

Fig. 54. Bolitoglossa dargonun (holotype). CRARC, Costa Rica, 400 m). Photo: G. Kahler Photo: J. A. Campbell

, , ,. ,,. ,,litog,,,a conanti (El Portillo, Fig. 55. Bolitoglossa decora (holotype). Ocotepeque, Honduras). Photo: G. Kohler Photo: J. R. McCranie

Publ., Mus. Zool., Univ. Michigan 200: 16; type locality: "Biotopo del Quetzal, Plan de Geomaya, 2235 m, Baja Verapaz, Guatemala (15.204732" N, 90.233556" W)". Maximum SVL in males 46.9 mm, in females 60.9 mm; tail 1engtWSVL ratio 0.46-0.80. Western and central portions of the Sierra de las Minas, Baja Verapaz and Progreso, Guatemala, 2400-2760 m elevation.

Bolitoglossa decora MCCRANIE & WILSON 1997, Proc. Biol. Soc. Washington 110: 367; type locality: "along the trail to Cerro Enrnedio near the ~ o n t e icon dido campground (15"05'N, 86"44W), Parque Nacional La Muralla, 1440 m elev., Departamento de Olancho, Honduras." Maximum SVL in males 40.2 mm, in females 62.1 mm; tail 1engtWSVL ratio 0.75-0.77. Only known from the vicinity of the type locality, 1430-1550 m elevation.

Bolitaglossa diaphora MCCRANIE & WILSON 1995, J. Herpetol. 29: 448; type locality: "above the visitors center of Parque Nacional El Cusuco, Cerro Cusuco (15"31'N, 88"12'W), 5.6 Ism WSW Buenos Aires, 1550 m elevation, Sierra de Omoa, Departamento de CortBs, Honduras." Maximum SVL in males 50.4 mm, in females 51.9 mm; tail 1engtWSVL ratio 0.43-0.72. Sierra de Omoa on the Atlantic versant of northwestern Honduras, 1470-2200 m elevation.

Bolitodossa diminuta ROBINSON 1976. Proc. Biol. got. Washington 89: 289; type locality: "Quebrada Valverde, a precipitous stream 8.8 km NE by road from the bridge crossing the Rio Grande de Orosi near Tapanti, Cartago Province, Costa Rica a t an altitude of approximately 1555 m, ... The exact site of the stream was determined from aerophotos to be 9'43'18"N latitude and 83'41'48W longitude." SVL in single known female 31.1 mm; tail 1engtWSVL ratio 1.14. Only known from the type locality.

Bolitoglossa dofleini (WERNER 19031, Abh. Math. Physik. C1. Bayer. Akad. Wiss. 22: 352; type locality: "Guatemala." Maximum SVL in males 70.0 mm, in females 125.9 mm; tail length/SVL ratio 0.79-0.84. Extreme northern Alta Verapaz, Guatemala, and Cayo District in southern Belize to north-central Honduras, 50-1550 m elevation.

Bolitoglossa dunni (SCHMIDT 19331, Field Mus. Nat. Hist. Publ., Zool. Ser. 20: 16; type locality: "mountains west of San Pedro [= San Pedro Sula, Departamento de CortBs], Honduras. Altitude 4500 feet." Maximum SVL in males 59.6 mm, in females 73.2 mm; tail length/SVL ratio 0.90-1.04 (males), 0.80-0.89 (females).

Eastern mountains of the Atlantic versant of Guatemala and western mountains of Honduras, 1020-1600 m elevation.

Bolitoglossa engelhardti (SCHMIDT 1936), Field Mus. Nat. Hist. Publ., Zool. Ser. 20: 156; type locality: "Volcan Atitltm, 7,000 feet above Olas de Moca, Solola, Guatemala." Maximum SVL in males 43.2 mm, in females 46.6 mm; tail len&h/SVL ratio 0.85-1.17. Extreme southeastern Chiapas, Mexico, along the Pacific versant to Volcan Atitlan, south-western Guatemala, 1100-2600 m elevation.

Bolitoglossa epimela W m & B w 1963, Rev. Biol. Tropical 11: 63; type locality: "a point on the Turrialba-Peralta road 6.2 miles northeast of the Rio Reventaz6n bridge (approximately 7 miles east-northeast of Turrialba), a t about 915 meters (3,000 feet) elevation, [Cantbn de Turrialba] Provincia de Cartago, Costa Rica." Maximum SVL in males 42.8 mm, in females 46.0 mm; tail 1engtWSVL ratio 1.08-1.27. Atlantic slope in central Costa Rica (Cartago Province), 775-1550 m elevation.

Bolitoglossa eremia CAMPBELL, SMITH, STREICHER, ACEVEDO & BRODIE 2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 11; type locality: "near La Soledad, on road to Torre de Guatel, 2650 m, Depto. Jalapa, Guatemala (14.530146" N, 90.142417" W)". Maximum SVL in males 42.3 mm, in females 62.1 mm; tail leneth/SVL ratio 0.62-0.83. Miramundo highla&, Departamento de Jalapa, Guatemala, 2360-2660 m elevation.

Bolitoglossa flavimembris (SCHMIDT 19361, Field Mus. Nat. Hist. Publ., Zool. Ser. 20: 158; type locality: "Volcan Tajumulco, at 7,200 feet, on the trail above El Porvenir, San Marcos, Guatemala." Maximum SVL in males 55.4 mm, in females 64.5 mm; tail length/SVL ratio 0.64-1.08. Slopes of the Tacand volcano in Chiapas, Mexico, and mountains along the southwestern Guatemalan Plateau (Atitltm, Tajumulco and Tacana volcanoes), 1500-2500 m elevation.

Bolitoglossa flaviventris (SCHMIDT 1936), Field Mus. Nat. Hist. Publ., Zool. Ser. 20: 148; type locality: "Chicharras, Chiapas", Mexico. Maximum SVL in 95 mm; tail lengtWSVL ratio 0.95-1.01. Pacific versant of Chiapas, Mexico, and western Guatemala, sea level to 1000 m elevation.

Bolitoglossa franklini (SCHMIDT 19361, Field Mus. Nat. Hist. Publ., Zool. Ser. 20: 159; type locality: 'Volcan Tajumulco, at 5,600 feet altitude, on the trail above El Porvenir, [Departmento del San Marcos, Guatemala." Maximum SVL 66 mm; tail length/SVL ratio

Fig. 56. Bolitoglossa diaphora (P.N. Cusuco, Fig. 60. Bolitoglossa engelhardti (Finca La Cortks, Honduras, 1550 m). Photo: G. Kohler Insula, San Marcos, Guatemala, 2057 m).

Photo: S. M. Rovito

Fig. 57. Bolitoglossa diminuta (holotype). M Fig. 61. Bolitoglossa epimela (holotype). Photo: D. B. Wake Photo: D. B. Wake

Fig. 58. BoLitoglossa dofleini (Quebrada Grande, Fig. 62. Bolitoglossa en ~mw Cop&, Honduras, 1300 m). Photo: G. Kahler Jalapa, Guatemala, 26L- ...,.


Fig. 59. Bolitoglossa dunni (EN. Cusuco, CortBs, Fig. 63. Bolitoglossa flavimembris (Finca La Honduras, 1810 m). r Photo: J. E. Kolby Insula, San Marcos, Guatemala, 2057 m).

Photo: S. M. Rovito

45

1.10-121.00 the Sa&e venmt &rm south- mtm Chiapas, Me*, dong &@ F d c ver- smt of &atemala as far as Vol& At&h, 1506-3000 m elmdon.

BdliW- &mnm. wg, SUB- L I?bMlaN 2007, Cop& a002 ':857; type "Cmh Etiea, P r o ~ c h Puntm6nae. La CNCes ~dogical station, a04~35 N, 15ae5mo w2 BNI my lkximum SVL in d e ~ 49.9 mm, in females 54,s aun; tail len@b%VL ratio Q.%l.lb PtilCitie dope of wesbru hia;bl&~tds pf T%mama Bad aaaeent Costa Itiaa, &o knew from the ~ ~ ~ l t e r n a d of the Fila (Joe'tefia in sou.theastepa Costa Eea, 1170-2IZ0 rn slewtioa

3 w i h h grid% mmm, mmoN L W m 1987, Revagfr Biol. T r d d 35: 87; ms localits: "Ria Q&L about 1 Itm NE by road &am the w e I W O ~ ~ ~ J tha Rfo Gkmde de

=ax Ttrpantf, Cert~go Province, Costa litica, at an elwatim of a p p r o h b b 1,280 m d@maUao ... R04'730'W, B3°4T4!8& -0 mP Ivkdmum 6VL in Paales 37.6 mm, ia &ximb% 82.4 mnh: td len&m d o 1.2Z-1.IS. Onls

178: 10; type l d t p *4.B d m w Cri&iBal de La Cams, Chiapas, MQ&O ... at absut 2134 rs (7000 h t ) elm-.'' M w a SVL ia malea 46.8 ma, in &mda 53.6 m; kd bB%$b/8m ~aaio O.ba.%, NO&hmh'd Chiapm (Gem T W u i B , Saa Cristobal de lm Gasas, aad QlgmuLa &Mcts), Mexim, snd Cordillera de 10s Chduumtanm, GuaWJ% lZ0&301)0 m elevation.

-- -.- - lbhxhltrm sm ia malw44.9 mm, is f e m h 63.1 gmq tdl hgtW1$VL ratio O.T.24.84 h males, 0.684.82 in females. @&a de %taptin range at the borders of M Wvdor, GUgtemala, md Hondu~w (dtawgh it ia auh yef known from khe Hod- p& of this m o d a h range), as well as Boa O ~ O &yetzdte~wque, CPuathtmalbl(5. m o pers. eamm. Nm. 20101,l84Q-%806 sn elevation.

B-1- B&& ( S ~ T 19861, Rdd MUB, N&. Mst. Publ., h l . Ser. 20: 16% tme l ~ d w "naou12W above Fin- Sam% w& of Cob- Alta Vmapaz, Qua-? d 5,W feet alMtude,w bhzimm 6VL in maledi 47.4 mm, in f'ePL4enaE88 4 . 7 mm; ;tail le-VL =tie Q.67-0.95. South- Altpl VmpaZ grid . Baja Verapm Deparbnah~, Guatemala f00e2@0 zn &ation.

Fig. 64. Bditoglossa indio (hobtype). Phnta: J. Sun jer

r

~ ~ ~ ; c z h u ~ h ~ e t e n m g a ~ i ~ Gem, tihmi, 8-m A m &% B m 2010, &. Publ., Mw. W., Univ. 1Vliri4&p11200: 80; type l d * : "140 km Fa' jlmtbn of h Mateo Lxfatlin=BarilIae, road to Mmt&n, Ruehuebnga, Oua- at 3781) m devc tion (35.81944" N, 91.5641W Ffn9 Maximma 5VL in males 56.1 nun, m females 56.0 mm; w l e V L rati~ tail k* O.W.94h duehamatan Subwea (sensu CaMBBm Qz V~W~UINT 1 8 Departamento de BuSllueCnatlgo, Guatemala, MBO-2$85 la

elmation. M*1og9(1 MB S-ER, LoTnaT, l3mnz, wm7 m, RCEWEQ $ 0 9 && Bid. 88: 320; I d t . YDos Boeas wo (11~%64.8"N, rn06gt&.4"7W), 2E m elma- iim, Depwtam~nto de Bo Saa 3ml;l, M-e 8% in only ~WWZI fern& @-- mea &8 mm; kil I - a rat& 0.74, ICgeamsnlyti.8m~etgpel*.

Photo: J. Sunyer

Fig. 70. Bolitoglossa hartwegi (Cerro , Chiapas, Tzontehuiz, Chiapas, Mexico). Photo: T. Bille

b

t Fig. 67. BoLitoglossa franklini (Cerro Mozotal, Fig. 71. Bolitoglossa heiroreias (P.N. Monte- Chiapas, Mexico, ca. 3000 m). Photo: L. Gray cristo, Santa Ana, El Salvador, 1904 m).

Photo: T. Leenders

Fig. 68. BoLitoglossa gomezi (Jurutungo, Chiriqui, Panama, 2120 m). Photo: G. Kiihler Quetzal, Baja Verapaz, Guatemala, 1650 m).

Photo: G. Kiihler

Fig. 69. BoLitoglossa gracilis (near Tapanti, ( Fig. 73. BoLitoglossa huehuetenanguensis tago, Costa Rica, 1350 m). Photo: M. F r a ~ ~ ~ , . ~ (holotype). Photo: J. A. Campbell

vation, Isla de Ometepe, Departamento de Rivas, Nicaragua." SVL in only known female specimen 64.7 mm; tail 1engthISVL ratio 0.86. Known only from the type locality.

Bolitoglossa jacksoni ELIAS 1984, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 348: 7; type locality: %as Nubes sector of Finca Chiblac, approximately 12 km NNE of Santa Cruz Barillas, Depto. Huehuetenango, Guatemala, at about 1,400 m elevation." Maximum SVL in females 65 mm (males unknown). Only known fkom the vicinity of the type locality.

Bolitoglossa kaqchikelorum CAMPBELL, SMI'I'H, STREICHER, ACEVEDO & BRODIE 2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 5; type locality: "vicinity of San Juan Sacatepkquez, Guatemala, Guatemala (14.717798' N, 90.650107" W), 1850 mu. Maximum SVL in males 42.0 mm, in females 53.3 mm; tail 1engWSVL ratio 0.70-0.93. Eastern portion of the Guatemalan Plateau, including the highlands bordering the western side of Guatemala City and near Antigua, 1690-1850 m elevation.

Bolitoglossa la CAMPBELL, S m , STREICHER, ACEVEDO & BRODIE 2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 23; type locality: "3.0 km SSE Chichicastenango, valley between Paxot and Caminibal, 2100 m, El Quich6, Guatemala (14.921181" N, 91.103626' W)". Maximum SVL in males 43.6 mm, in females 57.5 mm; tail length/SVL ratio tail length 0.63-0.80. Northern edge of the Guatemalan Plateau and the extreme western portion of the Sierra de C h u a h , 2100-2390 m elevation.

Bolitoglossa Lignicolor (PETERS 18731, Monatsber. Preuss. Akad. Wiss. Berlin 1873: 617; type locality: "Chiriqui", Panama; corrected to "Camron, Provinz Chiriqui", Panama, on the basis of museum records by BAUER et al. (1993: 295). Maximum SVL in males 67.7 mm, in females 81.2 mm; tail length/SVL ratio tail length 0.82-1.04. P a d c versant of southern Costa Rica and adjacent western Panama, as well as on khe Azuero Peninsula and Isla Coiba, Panama, sea level to 880 m elevation.

Bolitoglossa lincolni (STUART 1943a), Misc. Publ. Mus. Zool. Univ. Michigan 56: 9; type locality: "Monte at Salquil Grande, El QuichB, Guatemala. Altitude, 2450 meters." Maximum SVL in males 63.0 mm, in females 86.0 mm; tail length1SVL ratio 0.82. Central Plateau of Chiapas, Mexico, and in several mountainous areas in western Guatemala, including the upper slopes of the Guatemalan Plateau close to the Mexican border (Vulcan Tajumulco, Montanas de Quilco, and the southwestern

an13 eastern Sierra de 10s Cuchumataneal, 15003000 m elevation. I Bolitoglossa longissima MCCRANIE & CRUQ- D~AZ 1996, Caribb. J. Sci. 32: 195; type locality: "along the trail to Pico La Picucha, Sierra de Agalta (14"58W, 85"55W), ca. 10 airline km NNW Catacamas, 1900 m elevation, Departamento de Olancho, Honduras." Maximum SVL in males 48.2 mm, in females 61.1 mm; tail length/SVL ratio 0.77-0.89 (males), 0.81-0.96 (females). Only known from Parque Nacional Sierra de Agalta, in the Departamento de Olancho, east-central Honduras, 1840-2240 m elevation.

Bolitoglossa magnifica WN, WAKE & SAVAGE 2005, Copeia 2005: 228; type locality: "PanamB, from the southeast slope of Volc6n Chiriqui, 7.8 km W and 2.5 km N (airline) of Boquete, Prov. Chiriqui, g047.6'N, 82"30.OSW, 2400 m elevation." Maximum SVL in males 69.6 mm, in females 104.0 mm; tail length/SVL ratio 0.76-0.92 (males), 0.75-0.98 (females). Southern Cordillera de Talarnanca- Bard Massif in southwestern Panama, between Cerro Punta and Boquete, Province Chiriqui, 1250-2450 m elevation.

Bolitoglossa marmorea (TANNER & BRAME 1961), Great Basin Nat. 21: 23; type locality: "Crater of Volch Baru (Chiriqui), elevation 10,500 feet, Chiriqui Province, Panama." Maximum SVL in males 65.5 mm, in females 68.6 mm; tail length/SVL ratio 0.93-1.15. Southern portion of the Cordillera de Talamanca-Bani of eastern Costa Rica and western Panama, 1920-3444 m elevation.

Bolitoglossa medemi BRAME & WAKE 1972, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 219: 2; type locality: Tinca Chibigu', approximately 76"30'W, 6"15'N, on the Rio Arqufa, Departamento de Antioquia, Colombia. . . .

I Elevation approximately 300 m (980 ft)." Maximum SVL in males 46.7 mm, in females 48.2 mm; tail length/SVL ratio 0.75. Central

Fig. 74. Bolitoglossa longissima (Pico La Picucha, Olancho, Honduras).


Lag Fig. 75. Bolitoglossa jacksoni (from type locali- Fig. 79. Bolitoglossa lincolni (Sierra de 10s

Photo: J. Hanken Cuchumatanes, Guatemala). Photo: T. Bille

Fig. 76. Bolitoglossa kaqchikelorum (Cerro Alux, Fig. 80. Bolitoglossa magnifica (Las Tablas, Guatemala, 2107 m). Photo: J. A. Campbell Puntarenas, Costa Rica). Photo: M. Garcia Paris

Fig. 77. Bolitoglossa la (holotype). Fig. 81. Bolitoglossa marmorea (Volch Baru, Photo: J. A. Campbell Chiriqui , Panama, 3400 m). Photo: A. Hertz

i 'I Fig. 78. Bolitoglossa ligrugrucolor (Meseta de Fig. 82. Bolitoglossa medemi (near Belky,

I Chorcha, Chiriqui, Panama, 250 m). Antioquia, Colombia). Photo: P. A. Silverstone photo: G. Kijhler (Courtesy of the Biodiversity Institute, University of Kansas)

49

Plethodontidae

and eastern Panama to the Choc6 region of Colombia, near sea level to 800 m elevation.

Bolitoglossa me.?iana WAKE & LYNCH 1982, Herpetologica 38: 258; type locality: Uvicinity of Santa Rosa Pass, 9 km NE Santa Cruz del QuichC, [Departamento] El Quiche, Guatemala (elevation 2520 m)." Maximum SVL in males 66.0 mm, in females 88.0 mm; tail lengthlSVL ratio 0.84-1.10 (males), 0.79-0.98 (females). Central Guatemala, including the Sierra de Chuads and the Sierra de la Minas, 1550-2730 m elevation.

Bolitoglossa mexicana D&R~L, BIBRON & D-n 1854, Erp. Gen. 9: 95; Type localities: "province d'Oaxaca au Mexique" and Vera- Crux." Maximum SVL in males 64.1 mm, in females 71.6 mm; tail 1engthISVL ratio 0.96-1.10 (males), 0.81-1.09 (females). On the m Atlantic versant of Chiapas, Mexico, through southern and central Belize and northern Guatemala to northeastern Honduras; also an isolated population in the Yucatan Peninsula, near sea level to 1900 m elevation.

Bolitoglossa minutula WAKE, BRAME & BUELLMAN 1973, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 248: 7; type locality: 'north slope of Cerro Pando, 1920 m (6298 ft) elevation, Provineia de Bocas del Toro, western PanamB." Maximum SVL in males 36.0 mm, in females 36.5 mm; tail length/SVL ratio 0.86-1.12. Both slopes of the southern Cordillera de Talamanca of eastern Costa Rica and western Panama, 1670-2660 m elevation.

Bolitoglossa mombachoensis K~HLER & MCCRANIE 1999, Senckenb. Biol. 79: 89; type locality: "Volcln Mombacho (11°49.99'N, 85"58.77W), 1100 m elevation, Departmento de Granada, Nicaragua." Maximum SVL in males 57.5 mm, in females 66.0 mm; tail 1engthISVL ratio 0.74-0.98. Endemic at Volch Mombacho, Departamento de Granada, Nicaragua, 950-1250 m elevation.

Bolitoglossa morio (COPE 1869b), Roc. Acad. Nat. Sci. Philadelphia 21: 103; type locality: "Mountains of Guatemala." Neotype from '3.7 km S Santa Maria Visitaci611, Sololl, Guate- mala at 2000-2200 (14.723073" N, 91.330252" W)" (CAWBELL et al. 2010). Maximum SVL in males 55.0 mm, in females 44.4 mm; tail length/SVL ratio 0.69-0.81. Central portion of the Guatemalan Plateau, 1400-2500 m elevation. Oedipus boeourti B~occm 1882 is regarded as a synonym of B. morio according to CAMPBELL et al. (2010).

Bolitoglossa mulleri (BROCCHI 1883), Miss. Scient. Mex, Amer. Centr., Rech. Zool. 3 (2, livr. 3): 116; type locality: Several localities in

"haute Vera Paz." Maximum SVL in fernalee 80 mm (data for males not available); tail length/SVL ratio 0.69-1.07. Atlantic slopes of eastern Chiapas, Mexico, and northern Guatemala (Sierra de Los Cuchumatanes, Alta Verapaz highlands, and the Sierra de Santa Cruz), 100-1500 m elevation.

Bolitoglossa nigreseens (TAYLOR 1949), Univ. Kansas Sci. Bull. 33: 282; type locality: "Boquete Camp (on highway between Millville [= Villa Mills] and San Isidro del General), [Cantba de Perez Zeledbn, Provincia San Jose,] Costa Rica, elevation 6,000 feet." Maximum SVL in females 94.1 mm (males unknown); tail IengtWSVL ratio 0.79-0.81. Northern and central portions of the Cordillera de Talamanca of Costa Rica, 1650-3000 m elevation.

Bolitoglossa ninadormida CAMPBELL, S ~ T R , STREICHER, ACEVEDO & BRODIE 2010, Misc. Publ., Mus. Zool., Univ. 200: 28; type locality: 'near Todos Santos Cuchumath, Carretera between Chiabal and El Rancho, Huehuetenango, Guatemala, ca. 3200 m (15.469815' N, 91.543401" W)". Maximum SVL in males 41.8 mm, in females 57.1 mm; tail length/SVL ratio tail length 0.66-0.89. Only known from the type locality.

Bolitoglossa nussbaumi CAMPBELL, SMITH, STREIC~R, ACEVEDO & BROBIE 2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 26; type locality: 7.2 km SE Todos Santos, Huehue- tenango, Guatemala, 3259 m (15.484167" N, 90.552222' W)". Maximum SVL in females 53.4 mm (adult males unknown); tail 1engthBVL ratio tail length 0.66-0.89. Only known from the type locality.

Bolitoglossa nympha CAMPBELL, SMITH, STREICHER, ACEVEDO & BRODIE 2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 41; type locality: "E of San Miguelito, along tributary of Itio Bobos, Sierra de Caral, Municipio de Morales, Izabal, Guatemala, between 510-550 m elevation (15.357485" N, 88.719444" W)".

rig. as. ~outoglossa nympna {nolocype). Photo: J. A. Campbell

0 84. Bolitoglossa meliana (Santa Rosa Fig. 88. Bolitoglossa morio (3.7 km S Santa , QuichB, Guatemala). Photo: T. Bille Maria Visitacibn, Solola, Guatemala, 2200 m).


3 - -. t ?'A

I I I

. Bolitoglossa mexicana (P.N. Pico Fig. 89. Bolitoglossa mullen (Coban, Alta Atlhtida, Honduras). Photo: G. Kohler Verapaz, Guatemala, 1380 m). Photo: G. Kijhler

olitoglossa minutula (Jurutungo, Fig. 90. Bolitoglossa nigrescens (Cerro de la anama, 2060 m). Photo: G. Kohler Muerte, Costa Rica). Photo: W. Van Devendw

k. 87. Bolitoglossa mombachoensis (Volcitn Fig. 91. Bolitoglossa ninadormida (holotype). Photo: J. A. Campbell

Plethodontidae

Maximum SVL in males 38.0 nun, in females 43.0 mm; tail length/SVL ratio tail length 0.75-0.95 in males, 0.70-0.86 in males. Sierra de Card in eastern Guatemala near the border with Honduras, 275-1165 m elevation.

Bolitoglossa obscura m N , W m & SAVAGE 2005, Copeia 2005: 233; type locality: "Costa Rica, Prov. Cartago, 8 km S of Tapanti, Parque Nacional Tapanti, fbom Quebrada Valverde, g043'18"N, 83"46'48"W, 1555 m elevation." Only known from a single female with 87.7 mm SVL and a tail lengtWSVL ratio of 0.86. Only known &om the type locality.

Bofitoglossa occidentalis TAYLOR 1941, Univ. Kansas Sci. Bull. 27: 145; type locality: "La Esperanza, Chiapas, Mexico, elevation 500 feet." Maximum SVL in males 39.0 mm, in females 38.1 mm; tail 1engthlSVL ratio 0.55-0.75. Pacific versant from Cerro Bad in extreme eastern Oaxaca, Mexico, to Volch Atit lh in central southern Guatemala; isolated populations in the continental divide region of western Chiapas (Mexico), sea level to 2000 m elevation.

Bolitoglossa odonnelli (STUART 1943a1, Misc. Publ. Mus. Zool. Univ. Michigan 56: 10; type locality: "Cafetal just east of the hacienda at Finca Volch, [Departamento del Alta Verapaz, Guatemala. Altitude, 1200 meters." Maximum SVL in males 69.0 mm, in females 86.0 mm; tail 1engtWSVL ratio 0.98. Atlantic slopes of Alta Verapaz to the Montaiias del Mico, Guatemala, 100-1200 m elevation.

Bolitoglossa omniumsanctorum (STUART 1952), Proc. Biol. Soc. Washington, 65: 4; type locality: Todos Santos, Department of Huehuetenango, Guatemala. Elevation about 2500 meters." Maximum SVL in males 46.2 mm, in females 76.6 mm; tail 1engthISVL ratio 0.57-0.87. Extreme western portion of the Sierra de C h u a d , as well as the Sierra de 10s Cuchumatanes and Montaiias del Cuilco, Guatemala. Resurrected fmm synonymy of B. mono by CAMPBEU et al. (2010).

BoLitoglossa oresbia MCCRANSE, ESPINAL & WILSON 2005, J. Herpetol. 39: 108, Type locality: "Cerro El Zarciadero, U043.662'N, 87"53.925W, 1880 m elevation, Departamento de Comayagua, Honduras." Maximum SVL in females 55.5 mm (males unknown); tail length/SVL ratio 0.82-0.87. Only known kom the type locality.

Bolitoglossa pacaya CAMPBELL, SMITH, STREICHER, ACEVEDO & BRODIE 2010, Misc. Publ., MUS. Zool., Univ. Michigan 200: 8; type locality: "4 k m (airline) SSE San Vicente Pacaya on west slope Volch de Pacaya,

between 2000 and 2050 m, Depto. Escuintla, Guatemala (14.585821" N, 90.609385" W)". Maximum SVL in males 45.8 mm, in females 48.6 mm; tail len&h/SVL ratio 0.66-0.83. Southeastern highlands of Guatemala (VoIcAn Pacaya and the Las Nubes highland block), 1700-2300 m elevation.

Bolitoglossa pesrubra (TAYLOR 1952a1, m v . Kansas Sci. Bull. 34: 707; type locality: "Cerro de la Muerte, [boundary area between Cartago and San Jos6 provinces,] Costa Rica, elev. 10,900 ft.." Maximum SVL in males 57 mm, in females 67 mm; tail 1engtWSVL ratio 0.96-1.04. Cordillera de Talamanca in Costa Rica, 1870-3620 m elevation. I BoLitoglossa phalarosoma WARE & BRAME 1962, Los Angeles Co. Mus. Contrib. Sci. 49: 1; type locality: "Medellin, Departamento de Antioquia, Colombia ... The altitude of Medellin is 5,045 feet (1538 m)." Maximum SVL 58.2 mm. Department of Antioquia, Colombia, and Dari6n Province, Panama, 730-960 m elevation.

Bolitoglossa platydactyla (GRAY in CUVIER 18311, Animal Kingdom (GRIFFITH & PIDGEON, eds.) 9: 107; type locality: "Mexico." Maximum SVL in males 79.0 mm, in females 75.0 mm; tail lengtWSVL ratio 0.93-1.29. Southern Tamaulipas, Mexico, to northwestern Chiapas, Mexico, near sea level to 1300 m elevation.

Bolitoglossa porrasorum MCCRANIE & WILSON 1995b, Herpetologica 51: 132; type locality: "east slope of Pico Pijol (15"10'N, 87'33W1, Montaiia de Pijol northwest of Tegucigalpita, 1860-1900 m elevation, Departamento de Yoro, Honduras." Maximum SVL in males 58.1 mm, in females 65.7 mm; tail length/SVL ratio 0.70-1.09 (males), 0.69-0.88 (females). Montaiias de Pijol and Macuzal as well as Cordillera Nombre de Dios on the Atlantic versant in north-central Honduras, 980-1920 m elevation.

Bolitoglossa psephena C ~ B E L L , SMITH, STREICHEE, ACEVEDO & BRODIE 2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 33; type locality: "Finca Santa Elena, 5.2 km W of Health Center at Agua Escondida, Chimaltenango, Guatemala, 2500 m (14.791487" N, 91.051613O W)". Maximum SVL in single known female specimen 71.5 mm; tail lengtWSVL ratio tail length 0.73. Only known from the type locality.

Bolitoglossa pygmaea Bo-os & WAKE 2009, Zootaxa 1981: 58; type locality: "Fgbrega Massif (coordinates 9O07'00"N, 82O52'40W) at 3100 m elevation, Provincia de Bocas del Toro, Panama." Maximum SVL in males 32.6 mm, in

mg. ya. Bolitoglossa occidentalis (Wca U L ~ U U ~ ~ , Fig. 96. Bolitoglossa pesrubra (Cerro de la mapas , Mexico, 1050 m). Photo: G. K6hler Muerte, Costa Rica, 3300 m). Photo: G. Kohler

. Bolitoglossa odonnelh (B'mca &I r lg. 97. Bok'toglossa phalarmoma (Cordillera Senahfi, Mta Verapaz, Guatemala, Central, Deph. Tolima, Colombia, 1800 m).

Photo: S. M. Rovito Photo: A. Acoska

I

-

wsa ~mnlum9anc Fib. ". 1litOg10ssa plawdactyla (Veracruz, Mexico). Photo: M. Garcia Paris

Photo: 6. A. Campbell

litoglossa oresbia (Cerro Zarciadero, Fig. 99. Bolitoglossa ponasorum (2.5 airline agua, Honduras). Photo: J. R. McCranie km NNE of La Fortuna, Yoro, Honduras).


53

Plethodontidae

, ig. 1 L Y V - w - Y , - I mg(. Chiriqui, pan-a, 1965 m). Photo: A. Hertz

Fig. 101. Bolitoglossa sooyonun (north slope of Cerro de la Muerte, Cartago, Costa Rica, 3000 m). Photo: W. E. Duellman (Courtesy of the Biodiversity Institute, University of Kansas)

females 36.8 mm; tail lengthlSVL ratio 0.65-0.80 (males), 0.544.81 (females). Only known from the type locality.

Bolitoglossa robinsoni BOLAROS & WAKE 2009, Zootaxa 1981: 61; type locality: "Cerro Echandi (top of Cerro 3167 m), Prov. Puntarenas, Costa Rica, latitude 9"01750"N (9.0306), longitude 82"49'20W (-82.8222) (3.4 km max. error, WGS84 datum), elevation 3162 m." Maximum SVL 72.5 mm; tail length/SVL ratio 0.75-0.94. Section of the Cordillera de Talamanca on the continental divide that forms the boundary between Costa Rica and PanamB, and in the FBbrega Massif, which extends northward into Panamti, 2300-3335 m elevation.

Bolitoglossa robusta (COPE 1894), fioc. Acad. Nat. Sci. Philadelphia 46: 194; type locality: Taldas of Volcano Irazu", ca. g057'N, 83"52'W, Cant6n de Cartago, Provincia de Cartago, Costa Rica (HANKEN et al. 2005). Maximum SVL in males 113.9 mm , in females 133.5 mm; tail length 0.67-1.11 (males), 0.79-1.10 (females). Northern Costa Rica, from Volch

Cacao southeastward through the Cordilleras Ti larh and Central and the Talamanca-Bani Massif, to the Fortuna region, Province Chiriqui, Panama, 650-2100 m elevation.

Bolitoglossa rostrata (B~occm 1883), Miss. Scient. Mex. Amer. Centr., Rech. Zool. 3 (2, livr. 3): 112; type locality: "les hauteurs de Tonicapam [= Totonicaph, Departamento de Totonicaph] (Guatemala occidental)." Maximum SVL in males 49 mm, in females 54 mm; tail 1engtWSVL ratio 1.22-1.30. Guatemalan Plateau from the Tecph Ridge westward across the Cumbre Maria T e c b to the Volch Tajumulco region in San Marcos; also known from Cerro Tzontehuitz, Chiapas, and the Sierra de 10s Cuchumatanes, Guatemala (S. ROVITO pers. comm. Nov. 20101, 24003480 m elevation.

Bolitoglossa rufescens (COPE 1869b), Proc. Acad. Nat. Sci. Philadelphia 21: 104; type locality: "Orizava," [= Orizaba, Veracmz,] Mexico. Maximum SVL in males 40.5 mm, in females 45.0 mm; TLEIVL 0.78-0.95 (males), 0.74-0.85 (females). On the Atlantic versant from San Luis Potosi and Veracruz, Mexico, across Guatemala and Belize to northwestern Honduras, sea level to 1500 m elevation.

Bolitoglossa salvinii (GRAY 1868), Ann. Mag. Nab. Hist. Ser. 4, 2: 297; type locality: "Guate- mala." Maximum SVL in males 83.0 mm, in females 96.5 mm; tail length/SVL ratio 0.92-1.03. Pacific versant of southern Guate- mala and El Salvador, 600-1250 m elevation.

Bolitoglossa schizodactyla WAKE & BRAME 1966, Fieldiana Zool. 51: 1; type locality: "El Valle de Anton, 560 meters (1,837 feet), Provincia de Cocle, Panama." Maximum SVL in males 60.9 mm, in females 62.0 mm; tail length/SVL ratio 1.00-1.38. Atlantic versant from southern Costa Rica to central Panama, sea level to 850 m elevation.

Bolitoglossa sombra HANHEN, W m & SAVAGE 2005, Copeia 2005: 234; type locality: 'Costa Rica, Prov. Puntarenas, from aqueduct trail, along Rio Coton, below Las Tablas, g056.6'N, 82"45.0W, 1760 m elevation." Maximum SVL in males 70.0 mm, in females 84.5 mm; tail lengtb/SVL ratio 0.0.68-0.90 (males), 0.79-0.93 (females). Pacific slopes of the Cordillera de Talamanca of eastern Costa Rica and adjacent western Panama, 1500-2300 m elevation.

Bolitoglossa sooprum VIAL 1963, Rev. Biol. Tropical 11: 89; type locality: "Cordillera Talamanca, Cerro de la Muerte (= Cerro Buena Vista) at 9,000 feet (2,745 meters), 12 miles (19.2 km.) southeast of El Empalme, [Cant611 El Guarco,] Prwincia de Cartago,

102. BolitogIossa psepnena fiolotype). Fig. 106. Bolitoglossa rostrata (Sierra de 10s Photo: J. A. Campbell Cuchumatanes, Guatemala). Photo: T. Bille

t

qsa pygz i (from type Fig. 107. BoLitoglossa rufescens (1 km SSE Photo: G. Kijhler Tegucigalpita, Honduras, 40 m).Photo: G. Kiihler

Fig. 108. Bolitoglossa .-1vini.i (Coatepeque, Quetzaltenango, Guatc la, 530 m).

PhcL- G. Kohler

1 " hlhY

. 105. Bolitoglossa robusta (Mto Url, B o w Fig. 109. Bolitoglossa schizodactyla (Cocl6, Toro, Panama, 1080 m). Photo: M. Ponce Panama). Photo: W. Van Devender

55

- - - - - -

Costa Rica." Maximum SVL in males 65 mm, in females 72 mm; tail length/SVL ratio 0.72-1.08. Atlantic and Pacific slopes of the Cordillera de Talamanca of Costa Rica, 2355-3000 m elevation.

Bolitoglossa striatula (NOBLE 19181, Bull. Am. Mus. Nat. Hist. 38: 344; type locality: "Cukra [near Bluefields], eastern Nicaragua." Maximum SVL in males 54.0 mm, in females 65.9 mm; tail length/SVL ratio 0.96-1.22. Atlantic versant from northeastern Honduras to southern Costa Rica, near sea level to 1050 m elevation.

Bolitoglossa stuarti WAKE & BRAME 1969, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 175: 22; type locality: "7.8 mi (by road) SE of Ciudad Cuauht6moc (El Ocotal), Chiapas, Mexico, in Huehuetenango [Department], Guatemala, at about 950 m (3120 feet) elevation." Maximum SVL in males 51.1 mm, in females 57.0 mm; tail length/SVL ratio 0.72-0.80. Slopes drained by the upper tributaries of the Rio Grijalva, Departamento de Huehuetenango, Guatemala, 950-1660 m elevation. ------

Bolitoglossa subpslmata (BOULENGER 18961, Ann. Mag. Nat. Hist. Ser. 6,18: 341; type locality: "La Palma", Costa Rica. Maximum SVL in males 56 mm, in females 60 mm; tail length/SVL ratio 1.04-1.38. Cordilleras de Guanacaste, Tilarh, and Central in Costa Rica, 1245-2900 m elevation.

Bolitoglossa suchitanensis CAMPBELL, SMITR, STREICHER, ACEVEDO & BRODIE 2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 16; type locality: "Volcan Suchitan, above Aldea Suchitan, Municipio Santa Catarina Mita, 1960 m, Jutiapa, Guatemala (14.403209" N, 89.779753" W)". Maximum SVL in males 45.0 mm, in females 55.0 mm; tail length/SVL ratio 0.72-0.76. Volcan Suchith, Departamento de Jutiapa, Guatemala, 1850-1990 m elevation.

Bolitoglossa synoria MCCRANIE & K~HLER 1999, Senckenb. Biol. 78: 226; type locality: "Quebrada La Quebradona (14"23.96'N, 89"07.38W), north slope of Cerro El Pital, 2150 m elevation, Departamento de Ocotepeque, Honduras." Maximum SVL in males 52.0 mm, in females 59.8 mm; tail length/SVL ratio 0.76-0.90 (males), 0.69-0.72 (females). Cerro El Pital in the Department of Ocotepeque in southwestern Honduras, and in adjacent northwestern El Salvador, 2150-2710 m elevation.

Bolitoglossa taylori WAKE, BRAME & MYERS 1970, Am. Mus. Novit. 2430: 3; type locality: "south ridge of Cerro Cituro, Serrania de Pirre,

at an elevation of approximately 1100 meters (3608 feet), Dari6n Province, Republic of Panama." Maximum SVL in males 53.3 mm, in females 64.7 mm; tail length/SVL ratio 1.04-1.28. Eastern Daridn region, Panama, 800-1440 m elevation.

Bolitoglossa tica GARC~A-PAR&, PARRA-OLEA & WAKE 2008, Herpetol. J. 18: 24; type locality: "0.8-0.9 km SSW El Empalme junction on Interamerican Highway 2, Prov. Cartago and San Jos6, Costa Rica; 9.7137, -83.9507; 2360-2370 m elevation." Maximum SVL in males 59.7 mm, in females 53.9 mm; tail 1engthISVL ratio 0.91-1.32. Northern slopes of the Cordillera de Talamanca of Costa Rica, 1745-2500 m elevation.

Bolitoglossa tzultacaj CWBELL, SMITH, STREICHER, ACEVEDO & BRODIE 2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 18; type locality: "17.7 km above Jones on road to Finca Alejandria, Sierra de las Minas, 1800 m, Zacapa, Guatemala (15.116067" N, 89.614982" W)". Only known female specimen (subadult) has a SVL of 37 mm and a tail 1engthISVL ratio 0.64. Only known from the type locality in the eastern Sierra de las Minas.

Bolitoglossa veracrucis TAYLOR 1951, Univ. Kansas Sci. Bull. 34: 189; type locality: "at a point 35 km. SE of Jesiis Carranza (Santa Lucretia), elev. approx. 350 feet, Veracruz, Mexico." Maximum SVL in males 50 rnm, in females 55 mm; tail length/SVL ratio 0.84-0.92. Western portion of the Isthmus de Tehuantepec (in extreme southern Veracruz, extreme northeastern Oaxaca, and western Chiapas), 100-1000 m elevation.

Bolitoglossa xibalba CAMPBELL, SMITH, STREICHER, ACEVEDO & BRODIE 2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 36; type locality: "7.0 km by road SW San Juan Ixcoy, Huehuetenango, Guatemala, at 2750 m elevation (15.570678" N, 91.441228" W)". Maximum SVL in males 50.9 mm, in females 49.7 mm; tail 1engtWSVL ratio tail length 0.53-0.86. Western Guatemalan highlands including the Montaiias del Cuilco, 1980-2760 m elevation.

Bolitoglossa yucatana (PETERS 1882a), Sitzungsber. Ges. Naturforsch. Freunde Berlin 1882: 137; type locality: "Yucatan", Mexico. Maximum SVL 66 mm; tail length/SVL ratio 0.69-0.96. Northern and central portions of Yucath and Quintana Roo, Mexico; also an isolated population in south-central Campeche, Mexico, and one in northeastern Belize, near sea level to 300 m elevation.

'14. llu. aomoglossa striatula (Bartola, Rio Fig. 114. Bolitoglossa taylori (Serrania de Pirre, - in Juan, Nicaragua, 30 m). Photo: G. Kohler Darien, Panama, 950 m). Photo: A. J. Crawford

- . 115. Bolito ,sa tica (Macho Gaff.,

arta ago, Costa nica). Photo: M. Garcia Paris Photo: J. A. Campbell

. 112. Bolitoglossa subpalmata (P.N. Juan Fig. 116. Bolitoglossa xibalba fiolotype). Castro Blanco, Costa Rica, 1900 m). Photo: J. A. Campbell

Photo: G. Kohler

Fig. 113. Bolitoglossa synoria (holotype). Fig. 117. Bolitoglossa yucatana (2 km E Photo: G. Kohler Chichen ItzB, YucatAn, Mexico).

Photo: W. E. Duellman (Courtesy of the Biodiversity Institute, University of Kansas)

3. occidentalfs 3. rufescens

r B . hartwegi

Bo~oglossa 1 B. alberchi B. flaviventris

T B. jacksoni I I B. mexicana

B. mulleri 8 . odonnelli B. platydactyla

,r B. salvinii ./

7 -

/L - . cuchumatana

B. centenorurn B. la

r B. psephena

~olito~lossa' 7

A B. emmia - B. daryorum / ~uatemala r B. kaqchikelorurn

m B. morio B. omniumsanctorum

Bolitoglossa

B. mombachoensis

6. biseriata robusta maanific

I. alvaradai r B. subpalmata

B. pesrubra A B. marmorea

B. taylori

r B. cenoen

Key to Bolitoglos~a of northern Central America (southern Mexico to Nicaragua) 1 a Hands and feet moderately webbed (one or

two phalanges of longest digits free of webbing; Figs. 118a-n, 119a-g); subdigital pads usually present (Fig. 39b) ............... 2

b Hands and feet extensively to fully webbed (usually only the longest digits projecting slightly; Figs. 119h-n); subdigital pads usually absent ........................................ 38

2 a Toes triangular with acutely rounded to pointed tips (Figs. 118a-d); dorsum dark brown without conspicuous pattern ........ 3

b Tips of toes bluntly rounded or truncate (Figs 118e-n); dorsal coloration variable 6

3 a Slightly less webbing of hands and feet (Figs. 118a,b); SVL of adult males 33-56 mm, of adult females 45-65 mm ............. 4

b Slightly more webbing of hands and feet (Figs. 118c,d); SVL of adult males 33-48 mm, of adult females 31-58 mm ............. 5

4 a Venter pale; usually with pale limbs that contrast with darker body coloration; 29-56 m d a ~ teeth (<49 in individuals with a SVL <50 mm); 18-31 vomerine teeth; SVL of males to 55.4 mm, of females

....... to 64.4 mm Bolitoglossa flavimembris b Venter dark with fine reticulation of small

pale spots; limbs not paler than body; 40-70 maxillary teeth; 19-41 vomerine teeth; SVL of males to 43.6 mm, of females to 57.5 mm .......................... BolitogIossa la

5 a Ventral surface of tail tan in life; 25-70 maxillary teeth; 17-41 vomerine teeth; SVL of males to 43.5 mm, of females to 57.5 mm ................... Bolitoglossa cu&umatana

b Ventral surface of tail orange or pink in life; 51-62 maxillary teeth; 23-39 vomerine teeth; SVL of males to 47.6 mm, of females to 45.7 mm ............. Bolitoglh~sa helmrichi

6 a Dorsum with contrasting brick-red, orange-red, greenish or silvery blotches, or with a contrasting coral red (in life) longitudinal band on a black background (Figs.

............................ 14, 67, 79); venter dark 7 b Dorsal pattern not as above; venter vari-

able ........................................................... 8 7 a Adpressed fore and hind limbs touch; SVL

................. to 66 mm Bolitoglossa W b Two costal folds between adpressed limbs;

SVL to 57 mm ........... Bolitoglosrra linlincolni 8 a Dorsal and lateral surfaces of head, body,

and tail uniformly black without any pattern; tips of toes tkncate; SVL to 66 mm in males and 88 mm in females ....................... .................................... Bolitoglossa melitUW Fig. 118. Feet (dorsal view) of Bolitoglo~~a.

b Coloration of dorsal and lateral surfaces of Photos: G. Kohler

head, bods and tail not as above; tips of toes bluntly r-deed; maximum SVL usually r66 mm ...........................,............ 9

9 a Flanks usually with extensive pinkish coloration iB. life con€r&dhg 6 t h the black dorsal and p a y ventral uoloratiog 8546 masiUary tee& 1 3 4 4 vomerine teeth .. U)

b Coloration in life not as above; numbers of maxillary and vomerine teeth variable .. 11

18a Limb interval 1.5-4.0; 354-3 m a d l w teeth in &, & 9 4 2 in feznales; 13-16 oomerine teeth in males, 22-81 ia females; SVL to 46.9 mm in males and 60.9 mm in females ... Bok'&&.lae~a o m d m c t ~ ~ ~ ~ ~

b Limb interval 0.54.0; 35-66 m d l a r y tee* 1644 vomerine tee*, SVL to 42.3 mm in males and 62.1 nun in females ....;.. ..................................... j%&fYgrm arapUa

l la Bold pale spote and blotches on flanks and most of lateral and ventral eurfaees of tail; limb internal 0.0-2.0; 40-56 r n d a r g . teeth; 18-28 mmelliDe tee* SVL to 48.9 mm in malee and 60.9 mm in females .. 12

b Color pattern net as above; limb inCwaI variable; numbers of maxilltiry a d vomerine tees variable; SVL of adults variable ............... .. ..... .. ........................... l3

laa Ground color gray; not more than one phalanx free of webbbing on hands and feet; 4 W 6 madlarg teeth; 18-26 vomezke teeth; SVL to 4.6.9 mm in males and 60.9 ....... mm B females ,&lib&- daqmmm

b Giromd d o t brom; two to slighhiy more thaD two phalanges free of we'bbing on bands and feet; 47-70 m d q teeth, 23-31 vomerine teea, %I% to 40.2 mm in males and 62.1 in females ................... ...................................... 33db&wm deaora

1Sa Less webbbg of hands and feet, l .M.O phdaagm free of webbing ..................... 14

b More webbing of hands and feet, 1.0 pha- laax or less free of webbing ................... 25

14a Anterior three-fonrkhs of the subcaudal surface uniformly buff-yellow (in life; yellowish brown with anly widely scattered fleaka in preservative), this pale doration extending onto the nntrolokral portion of the tail or even onto the venColateral portions of tail and body; m d l a r y teeth aeries extending to lead of posteriol- end of orbit ............................ Bah'@- omsbia

b Ventrc,labd portiorrua of tail and body same color as, or darker than, that of dorsal coloration; maxillary teeth series &ending tp slightly beyond center of orbit .................................................................. 16

15a Dorsal surface of body with pde spotting or blotching, or a pale longitudinal band; dor-

aal surface of anterior portim of tail 6th conspirmous pde 1 e spots or blotche (in Y specimens with pa e longitudiaal band, this band conti6uee onto tail); usudy .................... fewer than 75 m d teeth %-w- '?manti .................................... Dorsum u s d y Without a conspicuous pale pattern, if auch pattern is present &en w i h t conspieuovs pale large spob or blotches on d a d surface of anterior portion of tail and then usually amre than ................................ 76 mardllary teeth ... 16 Number of maxiUary tee+& 8543, if more @an 64 maxillary teeth, then a pldr ofdm- solateral pde stripers usudlp present, or .......... venter orange to ru~ty red ia lifg 17 Number of teeth 64-97; no pair of domolakd pde a&ipes; venter brown ....................... or reddish brown in l& 21 Dorsal @&ace of body with a pair of doreo- lateral pale stripes (h~o~hlpicuous in some specimehl,) ..................... El&- cani Dmal surface of body uniform pale to dark brown, or with lichenous or black blot&es, or with pale rno-g or flecking, but with- ont a pair of dorsolaterd pale stripes .....M Less webbing (Finger III usudly with 2.0-2.1 phalanges on both aides and bgi%veen Toes m-VI freg of webbing Fig. ll8kl .......................... iwtcglm k!molh Mare webbing (Finger III usually 1.5-2.0 phalstnges onboth sides and beFw~ea b s

.................... IXl-Vr free af webbing) : ,.... 19 ... Subcaudal surface b m or daxk brown

............................... l30lbgIloaea Q-

Suboaudd d a c e pale yellm, orang;$ or m$y red in life (pale jelbw to brown in ........................................... preservative^ 20 FYngw II shorter t bm I"mger IV; dorsal surface of body uniform pale to dak brown, or with pale flecks; restricted to Montmla de Celaque (Departments of Lempira and Ocotepeqye) in westera ................... Hoadwras l%M@w#8 eala~pe Finger II same length as or longer t h Finger nJ; dorsal surface of body usualIy with lichenow or black blokches; resstricted to Sierra de Metapbn Range at the h r d m of El Sdvedor, Guatemala, and Hondurtw ................................. Bol&glvssahe@mia~ Dorsal srnrfaca sf feet with small, conspicuous gold or white epots in He; mom webbing (Finger LU wuafly with 0.5-1.5 p b langes on both sides and 1.0-1.5 phalanges between Toes III-VT free of web- Fig. 11811); restricted to northwestern Honduras ..................... B0l.i-m drmui Dorsal sElrfilees of feet without small, con-

Plethodontidae - _ -

spicuous gold or white spots in life; less webbing (Finger I11 usually with 1.5 to slightly over 2.0 phalanges on both sides and between Toes 111-VI free of webbing)

00 .................................................................. YY

22a Shorter limbs (adpressed fore and hind limbs slightly overlapping in males, a limb interval of 0.5 to 1.5 in females); more webbing (Finger I11 usually with 1.5 phalanges on the inside 1.5-2.0 phalanges on the outside and 1.5-2.0 between Toes 111-VI free of webbing); restricted to north-central Honduras ............ Bolitoglossa pomorutu

b Longer limbs (adpressed fore and hind limbs well overlapping in both sexes); less webbing (Finger I11 usually with 2.0 to slightly over two phalanges on both sides and between Toes 111-VI free of webbing);

........... m ............................... restricted to east-central Honduras

Bolitoglmna longisnima 23a Dorsal surfaces of head, body, and tail uni-

formly dark gray without any pattern, lateral and ventral surfaces of head, body, and tail slightly paler (grayish brown in preservative) than dorsum; limb interval 3.54.0 ..................................................... 24

L b Coloration not as above; limb interval <3.5 .................................................................. 26

24a Head narrow, ratio head width / SVL <0.17; SVL in single known female (an adult) 71.5 mm ......................... Bolitoglmna peephena

b Head broad, ratio head width / SVL >0.17; SVL in single known female (a subadult) 37 mm ...................... Bolitoglossa tzultaq'

25a Usually three costal folds between adpressed limbs; uniform dark gray or with dorsal pale band .. Bolitoglorssa engelhardti

b Usually less than three costal folds between adpressed limbs; dorsal pattern variable ................................................... 26

26a Dorsum with a pair of pale dorsolateral stripes (these can be fragmented into mottling); venter dirty white to brown ........ 27

b Dorsal pattern variable but without a pair .................... of pale dorsolateral stripes 32

.............. 27a More webbing of hands and feet 28 b Less webbing of hands and feet ............ 29

28a Limb interval 0.5-2.5 (mean 1.52); 27-81 (mean 58.3) maxillary teeth; 14-28 (mean 19.4) vomerine teeth; SVL to 45.8 mm in males and 48.6 mm in females ........... ...................... Bolitoglossa pacaya (in part)

b Limb interval 0.5-1.5 (mean 1.00); 3 5 4 5 (mean 39.2) maxillary teeth; 22-28 (mean 25.0) vomerine teeth; SVL to 53.3 mm in females (data for males not available)

Bolitoglossa kaqchikelonm (in part) ..........

29a Dorsolateral pale (cream color or reddish brown in life) stripes relatively broad, can be broken into mottling; limb interval 1.0-1.5 (mean 1.10); 50-74 (mean 59.0) maxillary teeth; 21-29 (mean 25.2) vomerine teeth; SVL to 41.8 mm in males and 57.1 mm in females .............................. ........................... Bolitoglon~a ninadom-da

b Dorsolateral pale stripes narrow; limb interval 1.5-2.5; number of maxillary and vomerine teeth variable: adult size variable .................................................................. 30

30a Limb interval 1.5-2.0 (mean 1.63); 60-82 (mean 69.8) maxillary teeth; 20-31 (mean 25.0) vomerine teeth; SVL to 56.4 mm in females (data for males not available) ............................. Bolitogllossa centenom

b Limb interval 2.0-3.0 (usually >2.0); number of maxillary and vomerine teeth vari-

......................... able; adult size variable 31 31a Habitus more slender; limb interval

.... 2.0-2.5 Bolitogllosa nussbaurni (in part) b Habitus more robust; limb interval 2.5-3.0

.................................... BoMoghm mtrata 32a Dorsum with a straight-edged broad pale

(tan, orange brown, or orange in life) middorsal stripe ............................................ 33

b No straight-edged broad pale middorsal stripe ....................................................... 36

33a Pale middorsal stripe tan to orange brown in life; limb interval 0.5-2.5 (mean 1.52); 27-81 (mean 58.3) maxillary teeth; 14-28 (mean 19.4) vomerine teeth; SVL to 45.8

....... mm in males and 48.6 mm in females ...................... Bolitogl088a pacap (in part)

b Pale middorsal stripe orange, copper or metalic brassy colored in life (metalic brassy color lost in preservative); limb interval 0.5-1.5; 35-45 maxillary teeth; 22-28 vomerine teeth; SVL to 53.3 mm in females (data for adult males not available)

34a Pale middorsal stripe copper or metalic brassy colored in life (metalic brassy color lost in preservative); limb interval 2.0-2.5; 49-87 maxillary teeth; SVL to 53.3 mm in females (data for adult males not available) .............. Bolitoglmna huehuetenanguensin

b Pale middorsal stripe orange in life; limb interval 0.5-1.5; 35-45 maxillary teeth; SVL to 50.1 mm in males and 56.0 mm in females .........................................................

Bolitoglma kaqchikelorum (in part) ......... 36a Dorsum uniformly gray or dark brown, no

.......................................... pale stippling 36 b Dorsum dark gray to black with pale (pale

gray, blue, or white in life) stippling that is sometimes clumped ................................ 37

- - .- - Plethodontidae --- 1

36a Flanks gray with white spots or mottling; limb interval 1.0-2.0; SVL to 54.3 mm in

.......................... females (data for adult males not available)

Bolitoglossa suchitanensis b No pale markings on flank; limb interval

.I ..................... 2.0-2.5; SVL to 53.4 mm in females (data for adult males not available) ............... Bolitoglossa nussbaumi (in part)

37a Dorsum and flanks dark gray to black with heavy pale gray or blue stippling that is sometimes clumped; limb interval 0.5-1.5 (mean 1.00); SVL to 53.3 mm in females (data for adult males not available) ........... ...................... Bolitoglossa pacaya (in part)

b Dorsal surfaces of head, body, and limbs dark gray to black with scant to moderate white stippling; flanks as dorsum or with

I few paler (salmon or rost-colored in life) streaks or blotches; limb interval 0.0-1.5 (mean 0.86); SVL to 44.5 mm in males and

..... 55.0 mm in females Bolitoglossa mono 38a Maxillary teeth absent 39

r ...........................

b Maxillary teeth present ......................... 40 ...... 39a Longer limbs (limb interval 1.0 to 2.5)

................................. Bolitogllorra nrfe~ems b Shorter limbs (limb interval 3.0 to 4.0) .....

................................... Bolitoglossa nymph 40a Dorsum of body and tail with a conspicuous

longitudinal dark brown band, bordered by an edging of immaculate white less than a millimeter in width and well-demarcated

I ......... from brilliant yolky yellow (in life) lateral body coloration Bolitoglossa jacksoni

......................... b Coloration not as above 41 41a Dorsum with contrasting dark or pale

blotches or longitudinal stripes (Figs. 41, 64, 66, 85, 89, 93, 98, 108) ...................... 42

b Dorsal color pattern variable, but no con- I trasting dark or pale blotches or longitudi-

nal stripes ............................................... 49 42a Venter immaculate pale ..............................

.............................. Bolitoglossa flavivenim~ b Venter brown to black ............................ 43

43a Dorsal surfaces of body and tail black with a conspicuous narrow vertebral pale (orange-yellow in life) stripe, or completely

............ .........................

black (Fig. 89) Bolitoglossa mulleri b Coloration not as above 44

44a Dorsal surfaces of body and tail with a conspicuous broad, yellow or orange dorsal band (Fig. 98) .... Bolitoglossa pla&damla

......................... b Coloration not as above 45 45a Dorsal pattern usually including at least

three colors, black or dark brown, pale (buff P 12 I 1:) Feet tdor.nl \ 1 e h v ) uf BoI l tog~o i> ,~ to orange-yellow in life) and pale brown to

.................. Photo. C; Koll le~ reddish brown (Figs. 46, 85) 46

b Dorsal pattern consisting of only two colors, black or dark brown and pale (orange-

.......................................... yellow in life) 47 46a Dorsal coloration usually consists of well

demarcated cream longitudinal stripes contrasting with black ground color with a middorsal orange-red (in life; brown in preservative) stripe usually present; ratio tail length / SVL 0.81-1.10; ratio head

................................ length / SVL 0.20-0.23 ................................. Bolitoglossa mesicana

b Dorsal coloration consisting of less contrasting pattern, no middorsal orange-red (in life; brown in preservative) stripe; ratio tail length / SVL 0.74 in single known specimen; ratio head length / SVL 0.24 ............ ........................................ Bolitoglossa indio

47a Longer limbs (limb interval 0.0-2.0 in

......................... ................................... I males, 1.5-3.0 in females); head broad and well demarcated from body

Bolitoglossa alberchi b Shorter limbs (limb interval 3.0-5.0 in both

sexes); head narrower and less demarcated from body ................................................ 48

48a Dorsal coloration consisting of narrow dorsolateral pale stripes, sometimes broken into rows of dots that do not meet across the dorsal midline or produce a reticulated pattern (Fig. 93); snout usually black ........ ................................. Bolitoglossa odonnelli

b Dorsal coloration consisting of dorsolateral pale stripes that join anteriorly to produce a pale-colored snout and that frequently meet across the dorsal midline or produce a reticulated pattern (Fig. 108); snout usually brown ..................... Bolitoglma salvinii

49a Number of maxillary teeth <21; SVL to ................. 39 mm Bolitoglossa m'dentalis

b Number of maxillary teeth 2G122; SVL of adults usually >40 mm .......................... 50

5Oa Large robust species (SVL to 70 mm in males, to 126 m in females); 80-122 maxillary teeth; limb interval 2-3; dorsum with well-defined dark spots, some spots comon- ly centered on costal grooves on dorsolateral

............ portion of body Bolitogllosa doffeini b Smaller species (adult SVL usually

<60 mm); fewer than 80 maxillary teeth, if more than 80, then limb interval less than 2; limb interval variable; dorsal coloration variable ................................................... 51

51a Vomerine teeth arranged in several series, sometimes forming a patch; dorsum gray, brown, or reddish brown with an irregular mottling of cream or pale tan; venter uniform brown or tan; limb interval 2-3 ......... ................................. Bolitoglossa yucatana

b Vomerine teeth arranged in a single linear series; dorsal and ventral coloration variable; limb interval variable ................... 52

52a Limbs shorter (limb interval 2.5-3.5); hands and feet fully webbed; maxillary teeth series extending to center of orbit .... ............................................................... 53

b Limbs longer (limb interval 0.0-2.0); hands and feet extensively webbed; maxillary teeth series extending to beyond center of orbit ......................................................... 55

53a Dorsal coloration consisting of a single pair of non-delimited dorsolateral shadings, moderately darker than dorsal ground color in a dorsal pattern otherwise homogeneously brown with dark brown mottling .................................. Bolitoglossa insularis

b Dorsal coloration consisting of a variable number of well delimited dorsal dark brown stripes ......................................... 54

54a Venter mostly dark brown with pale streaks, spots, or mottling; 14-28 maxillary teeth ............. Bolitoglossa mombachoed

b Venter pale with dark brown streaks (Fig. 127a); 22-37 maxillary teeth ...................... .................................. Bolitoglaea sfxiatula

55a Dorsum brown with irregularly scattered pale spots or irregular shaped blotches; tail 1engtWSVL ratio 0.84-0.92; 12-19

..... vomerine teeth Bolitoglossa veramcis b Dorsum uniform brown or brown with dark

brown mottling; tail 1engtWSVL ratio 0.43-0.73; 19-41 vomerine teeth ........... 56

56a Venter dark, either uniformly or with small white dots that may be visible only under magnification .......................................... 57

b Venter dirty white with a diffuse mottling of brown pigment, this mottling distributed homogeneously or becoming more dense midventrally and forming a rather large midventral patch .................................... 58

57a Slightly less webbing of hands and feet (Fig. 119k); in life, dorsum dark brown with

............................... reddish brown mottling .................................. Bolitoglossa hartwegi

b Slightly more webbing of hands and feet (Fig. 1191); in life, dorsum uniformly bown or pale brown with dark brown mottling ... ..................................... Bolitoglossa stuarti

58a Dark mottling on venter becoming more dense midventrally and forming a dark brown midventral streak (Fig. 122a) ......... ..................................... Bolitoglossa xibalba

b Dark mottling on venter distributed homogeneously (Fig. 122b) .................................... .................................. Bolitoglossa diaphm

Key to Bolitogllosa of southern Central America (Nicaragua to Panama) 1 a Distinct transverse ridge across frontal

region (Fig. 120); hands and feet fully webbed; 0 to 6 maxillary teeth in adults, rows not extending to eye Bolitogloesa domes

b No transverse ridge across frontal region; hands and feet variously webbed; more than 6 maxillaq teeth in adults, usually

............................ reaching to level of eye 2 2 a Overall coloration black with a pale (cream

to or reddish orange) ring around base of tail (Fig. 121); hand and feet moderately webbed; large species (SVL to 114 mm in males, to 134 mm in females) ..................... .................................... Bolitoglorna mbusta

b Coloration not as above, especially, no pale ring around base of tail; webbing of hand and feet variable; maximum SVL variable ................................................................. 3 C

3 a Biseriate color pattern with black lateral bands, buff dorsal band and whitish ventral band (Fig. 42); hand and feet fully webbed ........... ....... Bolitog..os~a bisariata

b Color pattern not conspicuously biseriate; I

hands and feet variously webbed ............ 4 4 a Hands and feet moderately webbed (one or

two phalanges of longest digits free of webbing; Figs. 123a-n); subdigital pads (Fig. 39b) usually present ........................ 5

b Hands and feet extensively to fully webbed (only the longest digits projecting slightly; Fig. 124); usually no subdigital pads ...................................................................... 23

5 a Dorsal, lateral and ventral coloration more or less uniform black; large species (SVL of adults usually >60 mm, up to 134 mm) .. 6

b Dorsal, lateral and ventral coloration not more or less uniform black; smaller species (SVL of adults usually <60 mm), or, if >60 mm, then dorsum brown with red-orange markings forming blotches, a middorsal field, or irregular paired dorsolateral

..................................................... stripes 11 6 a White pigment present along jaws and

throat; webbing of hands and feet more developed (only portions of distal-most phalanges of longest digits free of webbing) ........................................ BoIitqg10~~~ mpia

b No white pigment present along jaws and throat; webbing of hands and feet less developed (two or more phalanges of longest digits free of webbing) ................. 7

7 a Very small teeth that barely protude from the gums; relatively few teeth (35 maxil-

.................... lary teeth, 19 vomerine teeth) .................................... Bolitogloma obscura

b Larger teeth that protude well from the gums; usually more than 35 maxillary

........ teeth, more than 19 vomerine teeth 8 8 a Dorsum with whitish flecks, dorsal surface

of tail with larger whitish patches ............. ............................... Bolitogllosa magnifica

b Dorsum without whitish flecks, dorsal surface of tail without larger whitish patches .................................................................... 9

9 a More than 30 maxillary teeth (per side) in adult males, 39 or more in adult females ............................. Bolitoglossa anthracina

b Fewer than 29 maxillary teeth (per side) in adult males, 39 or fewer in adult females ... ................................................................. 10

10a Number of costal folds between adpressed limbs 2.5 to 4.0; ratio SVWhead width 5.9-6.5 in females; ratio SWwidth across

.................................. shoulders average 7.8 ............................... Bolitoglma n i g n w n i g r e r r o e n s

b Number of costal folds between adpressed

I limbs 0.5 to 2.5; ratio SVWhead width 6.5-6.7 in females; ratio SWwidth across shoulders average 8.2 .................................. .................................... Bolitoglossa omb bra

l la Almost unpigmented; internal black pigmentation of posterior abdomen and stomach showing through semitransparent ven-

very small species (SVL to 32.6 mrn in .................... males, to 36.8 mm in females)

Bolitoglossa p-ea

r tral and lateral body wall (in preservative); 1

.................................. b Pigmentation normal; no indication of

internal black pigmentation of posterior abdomen and stomach showing through ventral and lateral body wall; size variable .................................................................. 12

l2a Dorsum brown with red-orange markings forming blotches, a middorsal field, or irregular paired dorsolateral stripes; SVL to 53 mm in males, to 74 mm in females ................................ Bolitoglma mpacta

b Dorsal coloration not as above; size variable .......................................................... 13

13a Very small species, adults usually <40 mm SVL (maximum SVL 39 mm in males, 41

....................................... mm in females) 14 b Larger species, adults usually >45 mm SVL

.................................................................. 16 14a Number of maxillary teeth 34; number of

vomerine teeth 12 in single known specimen (a male); SVL 31 mm; truncus dorsally brown with a well-demarcated dark brown lateral band .. Bolitoglossa diminuta

I I

b Number of maxillary teeth 31-66; number of vomerine teeth 22-28; average adult size >37 mm SVL, no well-demarcated dark Fig. 123. r w a l , \UUIU~I view) of BoLitoglossa.

brown lateral band ................................. 15 Photos: G. Kohler

two phalanges of longest digits free of web- .......... bing); dorsal coloration not as above

..................................... B o l i ~ o 8 8 8 h e i Maxillary teeth series extending to level of center of eye; more webbing (less than two phalanges of longest digits free of webbing); dorsal and ventral surfaces lavender to purplish brown with yellow markings (streaks, stipples andlor spots), some specimens dark brown with pale markings in preservative, with or without a pair of

........ irregular dorsolateral pale stripes 17 Maxillary teeth series extending three- fourths the length of eye; less webbing (two phalanges of longest digits free of web-

................ bing); coloration not as above 20 Number of maxillary teeth 1945; number of vomerine teeth 19-25; a pair of irregular dorsolateral pale stripes present (Rg. 125); number of costal folds between adpressed limbs 0.5 to 1.0 in males, 1.0-2.0 in females ................................ Bok'toglossa cemxwh Number of maxillary teeth 43-80; number of vomerine teeth 22-38; no pair of irregular dorsolateral pale stripes; number of costal folds between adpressed limbs 0.0 to

............................................................ 2.5 18 Number of maxillary teeth 50-65; hands and feet relatively narrow; dorsum brownish black but with so much pale spotting

..... that the animal appears generally pale ................................. BoliWlossa mbinsoni Number of maxillary teeth 43-80; hands and feet relatively broad; dorsum, flanks, and venter purpelish brown, heavily

............ flecked and spotted with yellow 19 Occurs in the western portion of the Cordillera de Talamanca in Costa Rica; number of costal folds between adpressed limbs 0.0 to 0.5 in males, 0.0-1.0 in females ................................. Bolitogllossa swyonrm Occurs in the eastern portion of the Cordillera de Talamanca in extreme eastern Costa Rica and in western Panama; number of costal folds between adpressed limbs 0.0 to 0.5 in males, 1.5-2.5 in females ................................ Bok'togl0888 marmorea Head well delimited from the neck; 19-34 vomerine teeth in males, 2 2 4 6 in females; number of costal folds between adpressed limbs 0.0 to 1.5 in males, 1.0-2.5 in females

Bolitogllossa tica ......................................... Head not well delimited from the neck; 15-30 vomerine teeth in males, 17-29 in females; number of costal folds between adpressed limbs 0.0 to 3.0 in males, 2.0-3.0 in females ................................................ 21 Maxillary teeth in adults 37-65; Cordilleras de Guanacaste, Tilarhn, and

= 8. alvaradoi

Fig. 124. Feet (dorsal view) of Bolitoglossa. Photos: G. Kijhler

Photo: M. Garcia Paris

Number of costal folds between adpressed limbs 3.0 to 3.5; less webbing (two phalanges of longest digits free of webbing); dorsum bright yellowish to golden tan with brown to black streaks and spots ..... .................................... BoLitogfossa gradis Number of costal folds between adpressed limbs 0.0 to 1.0; more webbing (less than

.................................. Central of Costa Rica ............................. Bolitoglos~a ~ubbpalmata Maxillary teeth in adults 29-57; Cordilleras de Talamanca of Costa Rica

............................. and adjacent Panama 22 SVL to 49.9 mm in males, to 54.5 mm in females; tail lengthlSVL ratio 0.93-1.19

................ (mean 1.04) Bolitogl0888 lgamezi

SVL to 52.8 mm in males, to 57.8 mm in females; tail 1engthISVL ratio 0.96-1.04 ................................. BoliWassa pembra

Ground color solid black with a few white ~atches in two dorsolateral series along the :ides of the head and over the shou1;ders; Fig. 126. BoLitoglossa cuna (dorsal view of head very narrow, ratio HWISVL head). Photo: G. Kohler

...... 0.125-0.135 Bolitoglwa phahwoma b Coloration not as above; head moderately

...................................................... wide to broad, ratio HWISVL usually >0.130 24

24a Anterior portion of dorsal head at level of mideye abruptly paler than posterior portion of dorsal head (Fig. 126); 66-77 maxillary teeth; 33-38 vomerine teeth; number of costal folds between adpressed limbs about 3.5 ........................ Bolitogllos~a curia

b Anterior portion of dorsal head not abrupt- I

ly paler than posterior portion of dorsal head; number of maxillary and vomerine teeth variable; number of costal folds between adpressed limbs variable ........ 26

26a Hands and feet extensively webbed (Figs. 61,114); SVL to 43 mm in males, to 46 mm in females; 3 2 4 3 maxillary teeth; 16-24 vomerine teeth ........................................ 26

b Hands and feet fully webbed (Fig. 124); size variable; number of maxillary and vomerine teeth variable ......................... 27

26a SVL to 43 mm in males, to 46 mm in females; 3 2 4 3 maxillary teeth; 16-24 vomerine teeth ......... Bolitoglo8~a epimela

b SVL to 53.3 mm in males, to 64.7 mm in females; 44-78 maxillary teeth; 1 8 4 5 vomerine teeth ........... Bolitoglos~a taylon

27a Venter mostly dark brown to black, uniform or with pale streaks, spots, or mottling ......................................................... 28

b Venter pale (cream, yellow, or dirty white) with or without dark brown streaks, or with a broad median dark stripe .......... 33

28a Very small species, adults average 34 mm (SVL to 36 mrn in males, to 37 mm in females); 34-55 maxillary teeth; 1 5 4 0 vomerine teeth ....... Bolitoglasssa minutula

b Larger species, adults usually >40 mm SVL; number of maxillary and vomerine teeth variable Anterior ........................... 29 Fig. 127. Vent& surfaces in a. Bolitoglossa

29a Dorsum with contrasting dark or pale striatula; b. B. insularis. Photos: J. Sunyer

- Plethodontidae blotches (Fig. 43); maxillary teeth in adults 42-89; maxillary teeth series extending nearly to level of posterior margin of eye ................................ Bolitoglma alvaradoi

Dorsal color pattern variable (with broad cream to tan field, cream to yellow with paired brown dorsal stripes, paired dorsolateral pale stripes, or dark brown with pale streaks, but no contrasting dark or pale blotches); m d a r y teeth in adults 30-48; maxillary teeth series extending to level of center of eye or to about two-thirds of eye ..................................................... 30 Dorsum very dark, almost black, without markings; number of costal folds between adpressed limbs 0-1.5; maxillary teeth series extending to about two-thirds of eye .................................. Bolitoglh~88 med-

Dorsal coloration variable but not as above; number of costal folds between adpressed limbs 2.5-3.5; maxillary teeth series . extending to level of center of eye ......... 31 A pair of dark edged yellow to cream dorsolateral stripes present ............................. ........................................ Bolitoglma indio Color pattern not as above .................... 32

32a Number of vomerine teeth 14-28; venter ................. dark brown with pale streaks

...................... Bolitogloma mombachoe . b Number of vomerine teeth 22-40; venter

dark brown without pale streaks although ...................... pale mottling can be present

................................. Bolitoglma Ligniwlor 33a Venter yellow with a conspicuous irregu-

larly shaped dark brown central stripe, rarely immaculate; 43-104 maxillary teeth, 27-61 vomerine teeth; terminal pha-

................ langes on most digits bifurcated ......................... Bolitoglosaa achizodactpla

b Venter without an irregularly shaped dark brown central stripe; number of maxillary and vomerine teeth variable; terminal phalanges on digits not bifurcated .............. 34

34a Dorsum more or less homogeneously brown with dark brown mottling and a pair of faint, not well demarcated dark brown stripes; venter with indistinct dark brown

................... speckles Bolitog108sa i r ~ ~ u f t u i ~ b Dorsum cream to yellow with a pair of dis-

tinct, well demarcated dark brown stripes; venter with dark brown streaks (Fig. 127a) .................................. Bolitoglma striatula

Fq: 128. Bolitoglossa lignicolor (8 km NE Rio Fig. 129. Bolitoglossa sL---odactyI~ .-- Cop4 Chiriqui, Panama, 1210 m). Cocl6, Panama, 640 m). Photo: K.-H. Jungfer

Photo: G. K6hler

& WAKE 1963a,b, WAKE & BRAME 1963a, b, c, VIAL 1968, WAKE & BRAME 1969, WAKE et al. 1976, ALE~ERCH & ALBERCH 1981, HANKEN & WAKE 1982, WAKE & LYNCH 1982,

R et il. 2008, BOLAROS &WAKE 2009, CAMPBELL et al. 2010, TOWNSE'ND et al. 2009,2010

69

Plethodontidae

Bradytriton

In 1983, WAKE & ELIAS described the new genus and species Bradytriton silus based on material collected in northwestern Guatemala. This monotypic genus is diag- nosed by having a laterally compressed tail, stocky body with no clearly defined neck, and short, slender limbs bearing syndactylous hands and feet (WAKE & ELIAS 1983). The poor degree of development suggests that Bradytriton does not use its limbs for specialized locomotory activity such as climbing. Most of the known specimens of B. silus were found beneath pieces of wood in a clearing surrounded by mixed hardwood forest that receives over 6000 mm of annual precipitation (ELIAS 1984). It occurs syntopically with Nyctanolis pernix.

~ r a ' d ~ t o n silus WAKE & ELIAS 1983, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 345: 4; type locality: "Finca Chiblac, 15 km NE Barillas, Depto. Huehuetenango, Guatemala, elevation 1,310 m (4,300 ft)." Maximum SVL in males 53.3 mm, in females 53.0 mm; tail 1engthISVL ratio 0.59-0.83. Known from the vicinity of the type locality (including San Jos6 Maxbal) on the eastern slopes of the Sierra de 10s Cuchumatanes, as well as from Finca Ixcashn on the northern slopes of the Cuchumatanes, northwestern Guatemala.

F'urther Reading WAKE & ELIAS 1983, E m 1984

Fig. 130. Bradytriton silus (Finca Ixc 11.0 road km E Yalambojoch, Huehuetenango, Guatemala, 1640 m). Photo: T J. Papenfuss

Cryptotriton

Prior to WAKE & ELIAS (1983), the genus Chiropterotriton was used as a catchall for most of the small Middle American salamanders that were not referred to Bolitoglossa, Oedipina, or Pseudoeurycea. Currently, the Middle American species formerly referred to Chiropterotriton are placed in three genera: Cryptotriton (7 species; northern Oaxaca, Mexico, to northwestern Honduras), Dendrotriton (8 species; southwestern Chiapas, Mexico, to northwestern Honduras), and Nototriton (14 species; eastern Guatemala to central Costa Rica). The genus Chiropterotriton, as recognized today, occurs west and north of the Isthmus of Tehuantepec (GARC~A- Pmfs & WAKE 2000). The species of Cryptotriton are diminutive, slender salamanders that live either in moss mats or arboreal tank bromeliads ( G ~ c ~ A - P A R ~ S & WAKE 2000, MCCRANIE & WILSON 2002, TOWNSEND & WILSON 2008).

etween 1740 and 1780 meters." SVL 31.2 mm; tail IengtWSVL ratio

, Guatemala, 1610-2290 m eleva-

Fig. 131. Cryptotriton monzoni (5.2 km SE La Unidn, Guatemala, 1560 m). Photo: S. M. Rovito

Fig. 132. Cryptotn'ton nasalis (P.N. Cusuco, C O ~ B ~ , ~onduras, 1577 m). Photo: J. Kolby I.

rig. 133. Cryptotriton sierraminensis (from type locality). Photo: S. M. Rovito

r '

Fig. 134. C~yptotn'ton veraepacis (El Quiche, Guatemala). Photo: J. A. Campbt

Fig. 135. Cryptotriton wakei (holotype). Photo: J. A. Campb

1 KeytocqFptotn'ton 1 a Nostrils smaller (ratio nostril width / SVL

<0.020; Fig. 136b) Qyptotriton veraepac.5~ b Nostrils larger (ratio nostril width / SVL

>0.019; Fig. 136a,c) ................................... 2

I 2 a Venter in life yellow with a greenish cast

tive) or clear with punctate melanophores

I (pale beige with gray mottling in preserva- Fig. 136. a. C , t o t d t o n and a few widely spaced iridophores ...... 3 b. C. veraepacis; c. C. wakei. rnocos: G. Kijhler

b Venter in life and preservative dark or pale brown with or without dense brown flecking .................................................................... 4

8 a Venter with punctate melanophores; ratio tail length 1 SVL 1.08-1.21; head broader (ratio HWIHL >0.72) ................................... ....................... Crpptotn'ton alvarezdeltoxvi

b Venter pale beige with gray mottling in preservative; ratio tail length 1 SVL 1.29; head narrower (ratio HW/HL 10.70) ...........

C~totritoIl r i a r r a m i n d ........................ 4 a Legs shorter (ratio CCUSVL 0.42); vomer-

ine teeth 8 in single known specimen; venter dark brown ............................................. .................................. Crpptotntnton rn-'

b Legs longer (ratio CCUSVL 0.43-0.50); vomerine teeth 10-18; venter pale brown, unicolor or heavily suffused with brown or with dense brown flecking ........................ 6

6 a Ratio HUSVL 0.18-0.21; ratio HWISVL 0.13-0.14 ................... Crpptotriton naaalie

b Ratio HUSVL 0.22 in single known specimen; ratio HWISVL 0.15 ..............................

Crpptotriton wakai .......................................

Further Reading LYNCH & WAKE 1978, PAPENFUSS & WAKE 1987, GARC~A-PAR~S & WAKE 2000, MCCRANIE & WILSON 2002, CAMPBELL & SMITH 1998, TOWNSEND & WILSON 2008

B Plethodontidae

ies of diminutive, slender salamanders are distributed from southwestern

mas. Mexico, to northwestern

Fig. 139. Dendrotriton localitv).

chujonun (from type Photo: J. A. Cam~bell

" ieliads, the latter up to ten meters we the ground (MCCRANIE & WILSON !, CAMPBELL et al. 2010). The holotype .

D. kekchiorum was encountered at r t 19:OO h sitting on a fern about 1 m -

. > . . ._ . - .. . - I . ; . ..

Fig. 140. Dendrotriton chujorum (holotype). Photo: J. A. Campbell

137. Dendrotriton bromeliacius (SW of Fig. 141. Dendrotriton kekchiorum (holotype). @e above El Rincon, San Marcos, Guate- Photo: J. A. Campbell b, 2760 m).

U% Photo: S. M. Rovito

- I Fig. 142. Dendrotriton sanctibarbarus (ca. 3

&am. Dendrotriton megarhinus (Cerro Tres km S San Luis 10s Planes, Santa Barbara, s, Chiapas, Mexico). Photo: S. M. Rovito Honduras, 1705 m). Photo: S. M. Rovito

Plethodontidae

Denohtriton bromeliacius SCHMIDT 1936a1, Field Mus. Nat. Hist. Publ., Zool. Ser. 20: 161; type locality: "Volcan Tajumulco, at 8,000 feet altitude, on the trail above El Porvenir,

I - [Departamento de] San Marcos, Guatemala." Maximum SVL 37.9 mm; tail 1engthISVL ratio 1.11-1.35. Southern slopes of VolcAn

the Departamento de San Marcos, southwestern Guatemala, 1700-2700 m elevation.

I Tajumulco and adjacent volcanic highlands in

Dendrotriton chujorum C ~ B E L L , SMITH, STREICKER, ACEVEDO & BRODIE 2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 45; type locality: "14.0 km by road on the road to Nent6n NW of the junction with the San Mateo Ixtatzin to Barillas road, Huehuetenango, Guatemala, at 2780 m elevation (15.892222" N, 91.564167" W)". Maximum SVL in males 31.6 mm, in females 27.6 mm; tail length/SVL ratio 0.97-1.10. Only known from the vicinity of the type locality, 2780-2800 m elevation.

Dendrotn'ton cuchurnatanus (LYNCH & WAKE 1975), Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 265: 6; type locality: "forest along hig&way 9N, 8.5 km (%y road) SW San Juan Ixcoy, Huehuetenago, Guatemala, at about 2860 m elev." Maximum SVL in males 31.0 mm, in females 32.0 mm; tail length/SVL ratio 0.86-0.97. Only known from the type locality.

Dendrotriton kekchiorum CAMPBELL, S m , STREICHER, ACEVEDO & BRODIE 2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 48; type locality: "road between El Chime1 and San Pablo El Baldio, El QuichB, Guatemala, at 2100 m elevation (15.511944" N, 90.791944' W)". Maximum SVL in females 38.0 mm (adult males unknown); tail lengthlSVL ratio 1.01-1.20. Mountains north of the town of Uspantzin, Departamento de El QuichB, 2100-2591 m elevation.

Dendrotriton megarhinus (RABB 19601, Copeia 1960: 304; type locality: "northern slopes of Cerro Tres Picos, about 19 kilometers northeast of Tonala, Chiapas, Mexico. Altitude approximately 7000 feet." Maximum SVL in males 32 mm, in females 26 mm; tail 1engtWSVL ratio 0.92-1.26. Known only from Cerro Tres Picos in extreme southwestern Chiapas, Mexico, 2000-2900 m elevation.

Dendrotn'ton rabbi (LYNCH & WAKE 19751, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 265: 2; type locality: "9.5 km W, 8.5 km S (airline) La Democracia, Huehuetenango, Guatemala, ... between 2100 and 2500 meters." Maximum SVL in males 36.7 mm, in females 37.5 mm; tail length/SVL ratio 0.93-1.14.

Fig. 143. Dendrotriton xolocalcae (Reserva de la Biosfera El Triunfo, Chiapas, Mexico, 2000 m). Photo: S. M. Rovito

Fig. 144. Dendrotn'ton rabbi (Montaiias de Pefia Blanca, Huehuetenango, Guatemala, 2974 m). Photo: S. M. Rovito I Western Guatemala (Montaiias de Cuilco and Sierra de 10s Cuchumatanes), 2100-2700 m elevation.

Dendrotriton sanctibarbarus (MCCRANIE & WILSON 1996), Southwest. Nat. 41: 111; type locality: "Montaiia de Santa Bfirbara (14"55'N, 88"07W), 2,100 m elevation, Departamento de Santa Bhbara, Honduras." Maximum SVL in males 32.5 mm, in females 39.2 mm; tail length/SVL ratio 0.96-1.26 (males), 0.82-1.14 (females). Montaiia de Santa BArbara on the Atlantic versant in western Honduras, I

1830-2745 m elevation.

Dendrotriton xolocalcae (TAYLOR 19411, Univ. , Kansas Sci. Bull. 27: 148; type locality: "Cerro , Ovando, Chiapas, Mexico, between 6,800-8,000 feet elevation." Maximum SVL in males 33.2 mm, in females 34.2 mm; tail length/SVL ratio 1.00-1.11. Only known from the kype locality, 1500-2500 elevation.

eg to Dandrotn'ton A distinct sexual dimorphism in nostril width, nostrils small in males but large in adult females (ratio nostril width / SVL 0.005 in males, 0.015-0.016 in females; Fig. 145a,b); SVL in males to 31.6 mm, in &males to 27.6 mm; limb interval 0.5-1.0; ratio head width / SVL 0.15-0.21 ............... ............................... Den&tntnton Ch~OIurn

6 Nostrils large or small, but no difference " between sexes; adult size variable; limb

interval variable; ratio head width / SVL variable ..................................................... 2 I Nostrils smaller (ratio nostril width / SVL

<0.008) ...................................................... 3 b Nostrils larger (ratio nostril width / SVL

A.009) ...................................................... 6 @ Nostrils larger (ratio nostril width / SVL

0.006-0.007); head narrower (ratio HWBVL 0.15-0.16); hind foot narrower

.................... (ratio HFW/SVL 0.087-0.093) ...................................... DanhMton rabbi b Nostrils smaller (ratio nostril width / SVL

0.003-0.006; Fig. 145c); head broader (ratio HWISVL 0.15-0.21); hind foot broader (ratio HFW/SVL 0.096-0.110) ................. 4 SVL to 33.2 mm in males, to 34.2 mm in females; digits not expanded at the tip; tail IengthJSVL ratio 1.00-1.11 ......................... .............................. Dendrotritan xolocalcde SVL to 38 mm in females (adult males unknown); digits, except the first one, expanded at the tip; tail length/SVL ratio 1.01-1.20 .......... Dendtotn'tan k&&io~l~n

a Maxillary teeth usually >50, rarely as few as 44 in females; SVL to 32.5 rum in males, to 39.2 mm in females ................................. ...................... Dandrotriton a a n c t i b a r h

i) Maxillary teeth 50 or fewer; SVL variable 6 a Nostrils smaller (ratio nostril width / SVL

<0.017); ratio tail length / SVL 0.86-0.97 ..................... Dandrotrjtan cuchumatanus

b Nostrils larger (ratio nostril width / SVL >0.017); ratio tail length / SVL 0.92-1.35 ... 7 Head broader (ratio HWISVL 0.162-0.166); hind foot broader (ratio HFWISVL 0.096-0.105); ratio tail length / SVL

92-1.26 ...... Dendtob'to~~ megarhinus ower (ratio HWISVL

, hind foot narrower (ratio .089-0.094); ratio tail length /

Dandrotxiton bromebcius

, LYNCH & WAKE 1975, COLLINS- BUTH 1990, M c C m & WILSON 1998, MCCRANIE & WILSON 2002,

Fig. 145. Dendrotriton chujorum (a. male; b. female); c.D. kekchiorum (c.). Photos: G. Kohler

A D. sanctibarbams a D. xolocalcae

Nototriton

The genus Nototn'ton comprises 14 species of small terrestrial to arboreal salamanders that are distributed from eastern Guatemala to central Costa Rica at an elevational range of 740 to 2500 m. In Costa Rica, these animals typically inhabit moss mats, sometimes on the ground, but usually on the trunks and branches of trees. The Honduran species N. barbouri and N. lignicola were reported to have been taken during daytime under and inside rotten

Fig. 146. Nototriton abscondens (El Angel, Costa Rica). Photo: M. Franzen

logs and (in the case of N. barboun? in abandoned hummingbird nests; at night, N. barbouri was found to be active on low limbs and branches of small trees and shrubs (MCCRANIE & WILSON 2002). Another Honduran species, N. limnospeo tator, was mostly collected fiom forest leaf litter on the ground, but also from inside a bromeliad on a fallen trunk (MCCRANIE & WILSON 2002). The single known specimen of N. tomarnorum was collected during the daytime fkom leaf litter packed onto a rock ledge alongside a small creek (TOWNSEND et al. 2010).

In Costa Rica, these salamanders readily penetrate among the stems and rhizoids of

Fig. 147. Nototriton barbom. (Montana ae the mosses where the females deposit their Macuzal, Yoro, Honduras, 1780 m). clutches of 1-19 eggs (BRUCE 1998, JOINSH

Photo: J. T. Townsend & GARC~A-PAR~S 1998, MCCRANIE & WILSON 2002). The available evidence suggests that females of Nototriton do not attend ' their clutches (GOOD &. WAKE 1993, BRUCE

I Fig. 149. Eggs of Nototriton picadoi (P.N.

Fig. 148. Nototriton brodiei (from type locality). Tapanti, Cartago, Costa Rica). Photo: J. A. Campbell Photo: W. Van Devender

Plethodontidru

I months; hatching is completed by of the wet season in December

in abandoned hummingbird nests. Fig. 150. Nototriton gamezi (Monteverde, Costa Rica). Photo: M. Franzeu

h t r i ton abscondens (TAYLOR 1948), Proc. 01. Soc. Washington 61: 177; type locality: Ja Bonita (American Cinchona Plantation), w. 5500 ft., Volcdn Poas, Caribbean

and Central of Costa Rica, 1010-2500

Volch Cacao (elevation 1580 m), te National Park, Prov. de Guana- t a Rica." Maximum SVL 34 mm; tail

Fig. 151. Nototriton guanacaste (Cerro Cacao, Guanacaste, Costa Rica, 1540 m).

Photo: G. Kohler

Fig. 152. Nototriton lignicola (P.N. Montaiia de Yoro, Francisco MorazBn, Honduras, 2020 m).

Photo: J. T. Townsend

1engthISVL ratio 1.12-1.33. Volch Orosi and Volch Cacao, Province of Guanacaste, Costa Rica, 1400-1580 m elevation.

Nototriton lignicola MCCRANIE & WILSON 1997b, Proc. Biol. Soc. Washington 110: 369; type locality: "Cerro de Enmedio (15"06'N, 86'44W) along the trail above the Monte Escondido campground, Parque Nacional La Muralla, 1780 m elev., Departamento de Olancho, Honduras." Maximum SVL in males

33.9 mm, in females 32.8 mm; tail length/SVL ratio 0.90-1.06 (males), 0.84-1.01 (females). Only known from the type locality, 1760-1780 m elevation.

Nototriton limnospectator MCCRANIE, WILSON & POLISAR 1998, Herpetologica 54: 455; type locality: "northwestern side of Montaiia de Santa B6rbara southwest of San Luis de 10s Planes (14"56'N, 88"08W), 1910 m elevation, Departamento de Santa Bgrbara, Honduras." Maximum SVL in males 38.2 mm, in females 37.8 mm; tail 1engtWSVL ratio 1.25-1.30 (males), 1.03-1.06 (females). Montafia de Santa Barbara on the Atlantic versant in western Honduras, 1640-1980 m elevation.

Nototriton major GOOD & WAKE 1993, Herpetol. Monogr. 7: 137; type locality: "along Quebrada Platanillo, approximately 6 km east of Moravia de Chirripb, Prov. Cartago, Costa Rica . . . Elevation ca. 1200 m." SVL 37.9 mm in single known male specimen; tail lengtWSVL ratio 1.44. Only known from the type locality.

Nototriton picadoi (STEJNEGER 19111, Proc. U.S. Natl. Mus. 41: 285; type locality: "La Estella, southeast of Cartago, [Cartago Province,] Costa Rica." Maximum SVL 32.0 mm; tail 1engtWSVL ratio 1.20-1.42. North- western portion of the Cordillera de Tala- manca, Costa Rica, 1200-2200 m elevation.

Nototriton richardi (TAYLOR 19491, Univ. Kansas Sci. Bull. 33: 284; type locality: Tsla Bonita (American Cinchona Plantation), Atlantic slope of Volch Pols at an elevation of about 6,500 feetn, Costa Rica. Maximum SVL 24 mm; tail 1engthISVL ratio 1.07-1.48. Atlantic slopes of the Cordillera Central of Costa Rica, 1370-1800 m elevation.

Nototriton saslaya KO- 2002, Herpetologica 58: 205; type locality: "S slope of Cerro Saslaya (13"46.0'N, 85"02.3'W), 1371 m elevation, Regidn Aut6noma Atlantic0 Norte, Nicaragua." Maximum SVL 34.6 mm; tail 1engtWSVL ratio 0.88-1.26. Cerro Sasalya and Cerro El Torro in Parque Nacional Saslaya, north-central Nicaragua, 1280-1370 m elevation.

Nototriton stuarti WAKE & CAMPBELL 2000, Proc. Biol. Soc. Washington 113: 817; type locality: "11.6 km (road) WSW Puerto Santo Tomas, Montaiias del Mico, Depto. Izabal, Guatemala, 88"40W, 15"38'N, 744 m elev." Maximum SVL 32.6 mm; tail length/SVL ratio 0.57-0.62. Known only from the type locality.

Nototriton tapanti GOOD & WAKE 1993, Herpetol. Monogr. 7: 134; type locality: "Rio Quiri (elevation ca. 1300 m), 0.3 km northeast

of the junction of the Tapanti Road and the Tausito Road, near Tapanti, Prov. Cartago, Costa Rica." SVL 23.5 mm in single known female specimen; tail 1engtWSVL ratio L2L Only known fi-om the type locality.

Nototriton tomamom TOWNSEND, BUTLER, WILSON & AUSTIN 2010, Zootaxa 2434: 4; type locality: "2.5 km NNE of La Fortuna (15"25.965'N, 87"18.556W), 1550 m, Refbgio de Vida Silvestre Texiguat, Departamento de Yoro, Honduras." Single known female specimen has a SVL of 26.9 rnm and a tail length/SVL ratio of 0.91. &own only from the type locality.

* N. limnospectator

N. tornamorurn

to Nototriton a Hands and feet syndactylous, fingers and

toes not separate from one another, without subterminal pads; ratio 5th toe length /

........... SVL <0.004; tips of digits pointed 2 b Fingers and toes separate from one anoth-

er for at least one third of its length, subterminal pads usually present; ratio 5th toe length / SVL >0.005; tips of digits rounded or pointed .................................................. 4 b a Nostrils larger (ratio nostril width / SVI >0.017); hind foot narrower (ratic

. HFWISVL <0.039) Nototn'ton tomamom b Nostrils smaller (ratio nostril width / SVL

<0.017); hind foot broader (ratio HFWISVL >0.039) ...................................................... 3

a Legs shorter (ratio CCUSVL 0.31); hind foot narrower (ratio HFWISVL 0.041); Toe IV only outlined by a groove externally; maxillary teeth >28; ratio trunk length / SVL 0.61 ....................... Nototn'ton tapanti

b legs longer (ratio CCUSVL 0.33-0.37); hind foot broader (ratio HFWISVL 0.044-0.050); Toes I and IV only outlined by grooves externally; maxillary teeth <28;

............ ratio trunk length / SVL 0.49-0.57 ...................................... NototEI'ton r idudi

a Maxillary teeth >58; ratio nostril width / ....................... SVL 0.004-0.005 (Fig. 153a)

........................................ Nototriton bnniiei b Maxillary teeth <56; ratio nostril width /

............................................. SVL variable 5 a Nostrils larger (ratio nostril width / SVL

>0.009; Fig. 153b,e) .................................. 6 Nostrils smaller (ratio nostril width 1 SVL ~0.010; Fig. 153c,d) ................................... 9 Hind foot narrower (ratio HFWISVL 0.04-0.06); occurring in Honduras and

................................................ Guatemala 7 Hind foot broader (ratio HFWISVL

......... 0.06-0.07); occurring in Costa Rica 8 Maxillary teeth 36 in single known speci-

Plethodontidae

c. N. guanawsb

Fig. 153. Lateral view of head in species of Nototriton; note different nostril sizes.

Photos: G. Kohler

men; nostrils larger (ratio nostril width / SVL 0.012); head broader (ratio HWISVL

............................. 0.138) Nototriton stuarti b Maxillary teeth 41-54; nostrils smaller

(ratio nostril width / SVL 0.005-0.011); head narrower (ratio HWISVL 0.104-0.132 ...................... NototritM banboun' (in part)

8 a Habitus slender (ratio trunk width / SVL 0.08-0.10); legs shorter (ratio CCLISVL

.................. 0.34); SVL of adults 24-26 mm ........................................ Nototn'ton gamezi

b Habitus robust (ratio trunk width / SVL 0.10-0.11); legs longer (ratio CCLISVL

.................. 0.35); SVL of adults 20-32 mm ........................................ Nototn'tan picadoi

9 a Hind foot narrower, ratio HFWISVL ~0.045; ratio tail length 1 SVL d .07 in

........................ males and d . 0 1 in females N ~ t o ~ t o n lignimla .....................................

Plethodontidae

b Hind foot broader, ratio HFWISVL >0.045 or, if ~0.045, then ratio tail length / SVL

.... >1.19 in males and >1.02 in females 10 10a Legs longer (ratio CCLlSVL 0.42-0.45);

nostrils small (ratio nostril width 1 SVL 0.002-0.003); hind foot broader (ratio HFWISVL 0.075-0.091); maxillary teeth

........................ 17-23; vomerine teeth 3-11 ....................................... Nototn'ton saslaya

b Legs shorter (ratio CCLISVL 0.30-0.39); nostril size variable; hind foot narrower (ratio HFWISVL 0.040-0.072); maxillary teeth 25-55; vomerine teeth 10-26 ....... Photo: M. Franzen

l la Head relatively long and broad (ratio HLISVL 0.22-0.24; ratio HW/SVL 0.015-0.016); habitus robust (ratio trunk width / SVL 0.11-0.12); parotoid gland

..... often prominent Nototfiton guanacaste b Head moderate to short in relative length

and width (ratio HLISVL 0.16-0.21; ratio HWISVL 0.010-0.014); habitus slender (ratio trunk width / SVL usually ~0.11); parotoid gland usually less well developed .................................................................. 12

12a Maxillary teeth 41-55; vomerine teeth 12-26; occurring in Honduras ............... 13 Fig. 155. Nototriton limnospectator (Cerro

b Maxillary teeth 27-38; vomerine teeth h u l Mehbar , Comayagua, Honduras, 8-18; occurring in Costa Rica ................ 14 1650 m). Photo: J. T. Townsend

13a Nostrils smaller (ratio nostril width I SVL

narrow pale (cream to silver) &ripe sips- rating the dorsal and ventral colorations .. ........................... Nototriton limnospectator

b Nostrils larger (ratio nostril width I SVL 0.005-0.011); head broader (ratio HWISVL 0.116-0.128 in males, 0.104-0.132 in females); limb interval 4.0-6.5; no pale lat-

.... era1 stripe Notokiton barbod (in part) 14a Head moderate in relative length and

width (ratio HUSVL 0.18-0.21; ratio HWISVL 0.012-0.014); vomerine teeth 8-11; ratio tail length I SVL 1.10-1.31; SVL

............ to 32.2 mm Notohitan abscondem b Head relatively short and narrow (ratio

HWSVL 0.177; ratio HWISVL 0.011); vomerine teeth 18 in single known specimen; ratio tail length / SVL 1.44; SVL to 37.9 mm .......................... Nototrib w ' o r

0.003); head narrower (ratio HWISVL 0.110-0.118 in males, 0.095-0.098 in females): limb interval 3.0-5.0: usuallv a

Fig, 156. Nototriton richardi (Villa Calas, Alajuela, Costa Rica, 1920 m). Photo: G. Kohler

Further Reading MCCRANIE &WILSON 1992, Goon & WAKE 1993, 1 MCCRANIE 1996b, BRUCE 1998,1999, CAMPBELL & SMITH 1998, JOKUSH & GARC~A-PAR~S 1998, GARC~A-~AR~S & WAKE 2000, MCCRANIE & Fig. 157. Nototriton saslaya (holotype). W n s o ~ 2002,2003, TOWNSEND et al. 2010 Photo: G. Kiihler:

me-nrtidae -- enus Nyctanolis was proposed by & WAKE (1983) to accommodate a

mala and Chiapas, Mexico (ELIAS . It differs from all other tropical

& WAKE 1983). Most of the speci- Fig. 16G. IV~LWUIOL~S pernix San of the type series were collected Jose Maxbal, Huehuetenango, Guatemala,

Photo: S. M. Rovito

ofis pernix & WAIZE IgSa, in

r & ~ A T A (eds.), Adv. Herpetof. Evol. ; type locality: "Finca Chiblac, 10 km

(air) NE Barillas, Huehuetenango, Guatemala, (91°16W, 15"53'N), 1,370 m (4,500 ft) elevation." Maximum SVL in males 68.1 mm, in females 73.6 mm; tail 1engtWSVL ratio 1.19-1.37. Disjunct populations in southem Chiapaa, Mexico, and western and c a d Guatemala? 1200-1610 m elevation. , 8 8 ,

Nyctanolis pernix corn type locality). Photo: J. Hanken

C~ig. 159. Nyctanolis pernix (3.8 k m SE _?urulha, Baja Verapaz, Guatemala).

t Photo: J. A. Campbell (Courtesy of the Biodiversity Institute, University of Kansas)

Nyctanolis pemix

Further Reading ELIAS & WAKE 1983, ELIAS 1984

Plethodontidae

Fig. 161. Oedipina uniformrs (Albergue Montemeal, near P.N. Juan Castro Blanco, Alajuela, Costa

1 Rica, 1900 m). Photo: G. Kohler

I

Oediplna

The species of the genus Oedipina stand out among all other Neotropical salamanders because of their extremely slender, elongate bodies, tiny limbs and very long tails. Whereas the representatives of all other tropical salamanders have 14 trunk vertebrae, this number is 18 or more in the species of Oedipina (GARc~A-PAR~S & WAKE 2000, MCCRANIE et al. 2008). Oedipina is distributed from southern Mexico to northwestern South America with the majority of species (18 of 29) occurring in Costa Rica and Panama; only two species of this genus, 0. complex and 0. parvipes, have invaded South America. The species of Oedipina form a well-supported (by morphological and molecular data) monophyletic group divided into three subgenera: Oedipina (usually with 20-23 trunk vertebrae; western Guatemala to Panama), Oeditriton (with 20-21 trunk vertebrae; Honduras and Nicaragua), and Oedopinola (usually with 18-19 trunk vertebrae; Chiapas, Mexico, to Ecuador). They range from sea level to elevations in excess of 2200 m. These semifossorial or fossorial animals are usually found under

and inside rotten logs, beneath rocks, or in leaf litter and decaying plant material. They have also been reported to live under slabs of bark on fallen trees and in insect burrows within rotten logs (SAVAGE 2002). According to M. RYAN (pers. com. August 2010), individuals of Oedipina alleni and 0. savagei have been observed at night on the upper surface of leaves up to 1.5 m above the ground during the rainy season. If one of these salamander is touched, the usual reaction is body flipping. It is assumed that no parental care of eggs takes place in the species of Oedipina (GOOD & WAKE 1993).

Oedipina alfaroi DUNN 1921, Proc. Biol. SOIL Washington 34: 144; type locality: "Zent, , [Cantdn de Matina, Limdn Province,] Costa Rica." Maximum SVL 60 mm; tail 1engthISVL [ ratio 1.27-1.50. Atlantic versant of Costa Rica . ! and extreme northwestern Panama, near sea - ' level to 850 m elevation.

Oedipina alleni TAYLOR 1954, %v. Sci. Bull. 36: 607; type l o d t y : "Palma. Mortef, , Canth de Osa], Puntarenas Province, C d ~ i c a ~ ~ a x i m u m SYL in males 4 mm, i&j females 58 mm; tail 1engthlSVL ratid' 1.27-1.63. Pacific versant of southwester0

Plethodontidae

Fig. 162. Oedipina alleni (near Parrita, Puntarenas, Costa Rica, near sea level).

Toto: 1,eenders

b~g. 163. Oedipina carablanca (Pocora, Costa m Rica, 250 m). Photo: B. Kubicki

Fig. 164. Oedipina collarj.~ (P.N. Cr. D. Omar Torrijos H., near El Cop6, CoclB, Panama, 709 m). Photo: R. Brenes

Fig. 165. Oedipina complex (Alto de Piedra, Veraguas, Panama, 885 m). Photo: A. Hertz

Plethadontidae

Oedipina fortunensis K~HLER, PONCE & BATISTA 2007, Senckenbergiana Biol. 87: 214; type locality: 'Reserva Forestal Fortuna, near Finca de Gustavo Santamaria, BQ43'52"N, 82"15'35W, 1270 m, Chiriqui, Panama." SVL to 33.5 mm; tail length/SVL ratio 1.20-1.39. Known only from the Fortuna region in western Panama, 1200-1310 m elevation.

Oedipina gephyra McCJUNIE, WILSON & WILLIAMS 1993, Roc. Biol. Soc. Washington 106: 385; type locality: "2.5 airline km NNE La Fortuna (15'26'N. 87"18W), 1690 m elev., Cordillera Nombre de Dios, Departamento de Yoro, Honduras." Maximum SVL in males 52.3 mm, in females 58.1 mm; tail lengthlSVL ratio 1.30-2.23. Atlantic versant in the Cordillera Nombre de Dios of north-central Honduras, 1580-1810 m elevation.

Oedipina gracils TAYLOR 1952a, Univ. h a s m s, B.,, 34: 773: t, locali,: , ~iamantes , one mile sduth of ~ u b ~ i l e s , [Canhjn de Pocod, Provincia de Lim6~1 Costa Rica." Maximum SVL in males 44 mm, in females 47 mm; tail length/SVL ratio 1.39-2.57. Atlantic lowlands of Costa Rica and :xtreme northwestern Panama, sea level to 110 m elevation.

Oedipina grandis BRAE~E & DUELLMAN 1970, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 201: 1; type locality: "northern slopes of Cerro Pando, between 1810 and 1930 m elevation (5937-6330 feet), Rovincia de Bocas del Toro, extreme western Panama near the border with Costa Rica." Maximum SVL in males 68 mm, in females 72 mm; tail length/SVL ratio 1.78-2.23. Cordillera de Talamanca in extreme southern Costa Rica and adjacent western Panama, 1810-1950 m elevatien.

Oedipina ignea STUART 1952, Proc. Biol. Soc. Washington 65: 1; type locality: "along the Rio Las Brisas, just south of Yepocapa, Depart- ment of Chimaltenango, Guatemala. Elevation, about 1450 meters." Maximum SVL in males 61.7 mm, in females 62.1 mm; tail length/SVL ratio 1.87-2.97. Pacific versant of south-central and southeastern Guatemala as well as southwestern Honduras, 1000-2000 m elevation.

Oedipina kasios MCCRANIE, VIEI'EES % WAKE 2008, Zootaxa 1930: 11; locality: "near Quebrada Pinol, 15"07'N, 86"44W, Parque Nacional La Muralla, 1190 m a.s.l., Dept. Olancho, Honduras." Maximum SVL in males 37.5 mm, in females 46.8 mm; tail 1engthISVL ratio 1.6-1.8+. Endemic to Parque Nacional La Muralla in northwestern Olancho, Honduras, 950-1780 m elevation.

Oedipina leptopoda MCCRANIE, VIEITES & WAKE 2008, Zootaxa 1930: 13; locality: "32 km (road) W of Yoro on road to Morazh, 15.267480 N, 87.434820 W, Dept. Yoro, Honduras." Maximum SVL in males 49.9 mm (adult females unknown); tail 1engthISVL ratio unknown. 'The species is known only from three specimens collected many years ago. Field notes are precise with respect to locality, two from west and one from south of Yoro (Fig. 4), but vague with respect to habitat. The estimated elevation for these specimens is from ca. 700 to 1300 m m elevation.": (MCCRANIE et al. 2008: 14). i Oedipina man'tima GARC~A-PAR~S & W m 2000, Copeia 2000: 50; type locality: Tscudo Camp, West Point, Isla Escudo de Veraguas, Prov. Boeas del Toro, PanamB, approximate g06.1'N, 81°4.5W." Maximum SVL in males 46.2 mm, in females 44.3 mm; tail lengthlSVL ratio 1.47-1.89. Isla Escudo deveraguas in the Bocas del Toro Province, Panama; close to?ea level. ; 7 Oedipina nica SUNYER, WAKE, TOWNSEND, TRAVERS, ROVITO, PAPENFUSS, OBANDO & K ( 1 m 2010, Zootaxa 2613: 32; type IocaEty: "El Gobiado, Reserva Natural Cerro Datanli- El Diablo, 13"09'N, 85"52W, 1420 m above sea level (a.s.1.1, Dept. Jinotega, Nicaragua." Maximum SVL in males 44.5 mm, in females 48.5 mm; tail lengthlSVL ratio to 1.92. North- central Nicaragua, 1360-1660 m elevation.

Oedipina pacxcensis TAYLOR 1952a, Univ. 1

Kamas Sci. Bull. 34: 775; type locality: 'about two miles east of San Isidro El General," Costa Rica. Maximum SVL in males 48 mm, in females 51 mm; tail IengtWSVL ratio 1.39-2.57. Pacific slope of southern Costa Rica and adjacent western Panama, sea level to 730 m elevation.

Oedipina p m p e s (PETERS 18791, Monatsber. I Preuss. Akad. Wiss. Berlin 1879: 778; type locality: "Laceres [= CBceresl am Cauca, einem Nebenfluss des Magdalenenstroms in der Provinz Antioquia (Neu Granada)", Colombia. Maximum SVL in males 47.5 mm, in females 57.7 mm; tail length/SVL ratio 1.30-1.77. Central Panama to Choc6 region and the lower Cauca Valley in Antioquia Department of northwestern Colombia, near sea level to 865 m elevation.

Oedipina paucidentata BRAME 1968, Herpetol. 2: 41; type locality: "near El Empalme, (along the Pan American Highway, south of Cartago) in the northern end of the Cordillera de Talamanca, Cantiin de Dota [in error, Cant6n de El Guarcol, Provincia de Cartago, Costa Rica, about 2286 m (7500 8)

b. Oedipina fortunensis (holotype). Fig. 171. Oedipina kasios (from type locality). Photo: G. Kohler Photo: J. R. McCranie

-2. Oedipina gephyra (from type locality). Fig. &, 2. Oedip-a .,topoda (Los Pinos,

C Photo: J. R. McCranie Cort6s, Honduras). Photo: J. R. Collart

I Fig. 173. Oedipina nica (KilamM, Jinotega,

2060 m). Photo: G. Kohler Nicaragua, 1490 m). Photo: G. Kohler

85

gethodontidae

?levation." Maximum SVL in males 52 mm, in ,em of northeastern Honduras, 70-540 m el+ kmales 62 mm; tail 1engtWSVL ratio vation.

3edipinapoelziB~~lll~ 1963,Contrib. Sci. Nat. Copeia 2000: 52; type locality: "Finca Las( list. Mus. Los Angeles Co. 65: 7; type locality: Cruces, 6 km S San Vito de Java, Prov. Punta-

r 'area of the stone quarry in the W t y of El renas, Costa Rica, 8"47'35"N, 82"57'30W, ingel Waterfall (former American Cinchona approximately 1200 m elev.." Maximum SVL 'lantation and Isla Bonito of TAYLOR 1952),3. in males 38.5 mm, in females 39.3 mm; tail 5 miles by road south of Cariblanco (5 miles length/SVL ratio 1.08-1.86. Pacific versant in iorth of Vara Blanca), along the Vara extreme southern Costa Rica and adjacent Blanca-Puerto Vieio Road, Provincia de Panama. 1200-1400 m elevation. Heredia-Provincia "de ~ l a j u e l a boundary, :osta Rica; elevation 5000 feet (1520 meters)"; :orrected by SAVAGE (1974) to "0.5 km S :inchona, near Rio Santiaguito, Cariblanco road, Cant611 de Heredia, Provincia de Heredia: 1350 m", Costa Rica. Maximum SVL in males 63 mm, in females 64 mm; tail 1engthISVL ratio 1.33-2.13. Cordilleras de t'ilarh, Central, and Talamanca in Costa dica, 775-2050 m elevation

Oedipina pseudoum'formis BRAME, 1968, J. Herpetol. 2: 25; type locality: "Cienega Colorado approximately three kilometers by road east of Juan V i W and 6.3 km by road west' of Turrialba, Cant611 de Turrialba, Provincia de Cartago, Costa Rica, elevation 1035 m (3400 R)." Maximum SVL in males and "emales 53 mm; tail length/SVL ratio 1.44-2.03. Northern Nicaragua to central Costa Rica, near sea level to 1253 m elevation. Yowever, SUNYER et al. (2010) pointed out that he Nicaraguan populations refemed to 0. >seudouniformjs represent an undescribed rpecies.

3edipina quadra MCCRANIE, VIEITES & WAKE 2008, Zootaxa 1930: 6; locality: "UNS Tingni Kiamp, 14"55'N, 84"41W, tributary of upper portion of Rio Warunta, 160 m above sea level (a.s.l.), Dept. Gracias A Dios, Honduras." Maximum SVL in males 43.7 mm, in females 55.5 mm; tail 1engtWSVL ratio to 2.4. From south of La Ceiba, Atlfintida, eastward and southeastward to the Rfo Coco drainage sys-

- . 7 ,. - r

- -

Oedipina stenopodia BRODIE & CAMPBELL 1993, Herpetologica 49: 260; type locality: "south slope of Volcfin Santa Maria, Finca El Faro, Departamento de Quezaltenango, Guatemala; ... This locality is located between 1219 and 1432 m elevation at about 14"43'N, 9lo34'W." Maximum SVL 51 mm; tail length/SVL ratio 2.13-2.18. Pacific versant of southwestern Guatemala in the departments of San Marcos and Quezaltenango, 1200-1500 m elevation.

Oedipina stumti BRAME 1968, J. Herpetol. 2: 47; type locality: "Amapala, Isla Tigre, in the Golfo de Fonseca, Departamento de Valle, Honduras." Maximum SVL in males 54.0 mm, in females 63.1 mm; tail length/SVL ratio to 2.30. With certanty only known from the type locality; it has also been reported from the city of Tegucigalpa at 1000 m elevation, but this record has been questioned (MCCRANIE & WILSON 2002).

Oedipina taylon' STUART 1952, Proc. Biol. Soc. Washington 65: 2; type locality: "4 ldometew east of Hacienda La Trinidad (23 air-line kilometers southeast of Chiquimulilla), Depart- ment of Jutiapa, Guatemala. Elevation, about 100 meters." Maximum SVL 69.3 mm; tail length/SVL ratio to 2.01-2.63. Pacific versant from southeastern Guatemala along southern Honduras to central El Salvador, also known fkom the upper Motagua Valley in the vicinity of Dofia Maria in southeastern Guatemala, 140-1140 m elevation.

Fig. 174. Oedipina pseudounifonnis (near Turrialba, Cartago, Costa Rica). Photo: W. E. Duellman (Oo-y ofthe Biodivemity Institute, University of 1

Plethodontidae

si MCC- 2006,J. Herpetol. Oedipina m'formi's K E F E ~ 1868a, Arch. locality: "near Sendero de Naturgesch. 34: 299; type locality: "Costarica."

e Nacional El Cusuco, 15"30'N, Also described as new by KEFER~TEIN (1868b: tW014W, 1800 m elevation, Departamento de 331). Maximum SVL in males 57 mm, in

; €!at&, Honduras." Maximum SVL in males females 55 mm; tail length/SVL ratio 57.5 mm, in females 62.9 mm; tail 1engthlSVL 1.56-2.85. In Costa Rica from V o l h Tenorio ratio 1.90-2.03. Only known from the type south through the cordilleras and Meseta

E U t y Central to the Panaman border, 750-2150 m elevation.

Fig. 178. ueaipina quadra (P.N. Pico Bonito, Costa Rica). Photo: M. Garcia Paris Atlhtida, Honduras, 160 m). Photo: G. Kohler

6. Oedipina parvipes (Burbayar Lodge, Photo: B. Kubicki

Fig. 179. Oedipina taylori (P.N. El Imposible, Ahuachaph, El Salvador, 1120 m).

Photo: V. Henn'quez

177. Oedipina poelzi (Monteverde, Costa rlg. lau. uempma comas1 (r.lu. busuco, Cortes, Photo: M. Franzen Honduras, 1810 m). Photo: J. Kolby

87

Key to the Oedipina of northern Central America (southern Mexico to north-centd Nicaragua) 1 a Costal grooves 21-22; hind foot narrow

(ratio HFWISVL ~0.020); head narrow (ratio HWISVL ~0.070); maxillary teeth 0-13 ................................. Oedipina taylori

b Costal grooves 17-20; hind foot broader (ratio HFWISVL usually >0.020); head broader (ratio HWISVL >0.070); maxillary

................................................. teeth 0-98 2 ............................... 2 a Costal grooves 17-18 3

b Costal grooves 19-20 ............................... 6 3 a Maxillary teeth absent; hind foot broader

(ratio HFWISVL usually >0.030); limb ....... interval usuallyc9 Oedipina elongata

b Maxillary teeth 18-58; hind foot narrower (ratio HFWISVL usually ~0.030); limb interval 9-14 ........................................ 4

4 a Limb interval 12-14; maxillary teeth 18-37; vomerine teeth 9-18 ....................... .................... Oedipina stenopdia (in part)

A 0. carablanca

+ 0. pseudouniformis

b Limb interval 9-11; maxillary teeth .................. 43-58; vomerine teeth 16-23 5

5 a Body jet black in life; a row of pale flecks present ventrolaterally on body in life; less interdigital webbing on longest toe of hind

................... foot (nearly one segment free) .......................................... Oedipina tom&

b Body blackish brown in life; no row of pale flecks present ventrolaterally on body in life; more interdigital webbing on longest toe of hind foot (fully webbed with only the

...................... toe tip protruding; Fig. 182a) ....................................... Oedipina gephyra

6 a Maxillary teeth 80-98; hind foot broader (ratio HFWISVL 0.046-0.050); limb interval usually 9-11 ............ Oedipina co&zfs

b Maxillary teeth 18-57; hind foot narrower (ratio HFWISVL 0.019-0.037); limb interval variable .............................................. 7

7 a Head broader (ratio HWISVL 0.100-0.135); limb interval 9-12 ................................... 8

b Head narrower (ratio HWISVL 0.073-0.102); limb internal 10.5-14 .............................. 9

8 a Not all digits tightly fused; maxillary teeth 36-57; vomerine teeth 17-34; hind foot broader (ratio HFWISVL 0.029-0.037) ..... ......................... Oedipina pseudouniformis

b All digits tightly fused (digits can be dis- cerned only because there are ill-defined grooves on the dorsal surfaces); maxillary teeth 29-40; vomerine teeth 10-17; hind

....... foot narrower (ratio HFWISVL 0.027) ..................................... Oedipina leptopoda

9 a Hind foot broader (ratio HFWISVL 0.026-0.036) ........................................... 10

b Hind foot narrower (ratio HFWISVL 0.019-0.026) ........................................ 11

10a Tail nearly rectangular in cross section; SVL to 43.7 mm in males, to 55.5 mm in females ........................... Oedipina quadz-a

b Tail round to ovoid in cross section; SVL to 61.7 mm in males, to 62.1 mm in females ........................................... Oedipina ignea

lla Head broader (ratio HWISVL 0.085-0.101) ................................................................. 12

b Head narrower (ratio HWISVL 0.074-0.083) ........................................... 13

12a Maxillary teeth 41-48 ........ Oedipina nica ................................ b Maxillary teeth 1 8 3 7

.................... Oedipina stenopadia (in part) 13a Maxillary teeth >20; SVL to 37.5 mm in

males, to 46.8 mm in females .................... .......................................... Oedipina kasios

b Maxillary teeth <20; SVL to 54.0 mm in males, to 63.1 mm in females .................... ......................................... Oedipina stuarti

Key to te Oedipina of southern Central America (southern Nicaragua to Panama)

............................... 1 a Costal grooves 17-18 2

............................... b Costal grooves 19-20 8 2 a Maxillary teeth >19; 13-14 vomerine teeth;

ratio HUSVL 0.165-0.180 .......................... ................................... Oedipina fortunensis

b Maxillary teeth <20, or if r19 then ratio HUSVL ~0.170; number of vomerine teeth variable .................................................... 3

3 a Head broader (ratio HWISVL 0.104-0.119) 4 b Head narrower (ratio HWISVL

.............................................. 0.085-0.106) 6 4 a Maxillary teeth absent; vomerine teeth

........................ 10-12 Oedipina carablanca b Maxillary teeth 7-45; vomerine teeth

11-32 ....................................................... 5 5 a Snout more bluntly rounded; limb interval

7.0-9.5; maxillary teeth 1345; vomerine ................... teeth 17-32 Oedipina complex

b Snout less bluntly rounded; limb interval i 6.0-7.0; maxillary teeth 7-18; vomerine , .................... teeth 11-22 Oedipina savagei 1

6 a Venter pale; maxillary teeth 0-19 .............. Oedipina pamipes .......................................

b Venter dark, with or without pale markings; maxillary teeth 0-8 ......................... 7

7 a Hind foot broader (ratio KFWISVL >0.050); dorsal surfaces pale brown with cream markings on tail and usually head and body including a large occipital pale blotch; venter reticulated with brown around

....................... cream spots Oedipina alleni b Hind foot narrower (ratio HFWISVL

<0.050); dorsum dark brown, almost chocolate on sides, paler along indistinct dorsal

................. band; venter paler than dorsum ...................................... Oedipina maritima

8 a Maxillary teeth absent; vomerine teeth 4-13; dorsal surface of head usually distinctly paler than that of body ................... ........................................... Oedipina alfami

b Maxillary teeth 14-98; vomerine teeth 9-34; dorsal surface of head usually not distinctly paler than that of body although pale blotched can be present on dorsal sur-

............................................... face of head 9 9 a Maxillary teeth 80-98; hind foot broader

(ratio HFWISVL 0.0460.050); limb interval usually 9-11 ............. W p i n a coflaris

b Maxillary teeth 14-70; hind foot narrower (ratio HFWISVL 0.013-0.040); limb inter-

............................................. val variable 10 10a Venter darker (usually black) than dorsum

of body (brown), and separated by a creamy or silvery lateral stripe .......................... 11

Plethodontidae

1 4 n I Fig. 181. Dorsal view of head in a. Oedipma altura; b. 0. cyclocauda; c. 0. paucidentata; d. 0. savage5 e. 0. uniformis. Photos: G. Kohler 1 Fig. 182. Dorsal view of hind foot in a. Oedipina gephyra; b. 0. quadra; c. 0. uniformis; d. 0. taylori.

Photos: G. Kohler

b Venter paler (usually gray to grayish black) than dorsum of body (black to blackish gray), no creamy or silvery lateral stripe ........... 12

l l a Head broader (ratio HWISVL 0.100-0.118); hind foot broader (ratio HFWISVL 0.035-0.040) ...................... Oedipina poelP'

b Head narrower (ratio HW/SVL 0.085-0.100); hind foot narrower (ratio HFWISVL 0.021-0.28) .. Oedipina grandis

12a Head broader (ratio HWISVL 0.100-0.113); hind foot broad (ratio HFWISVL 0.029-0.037); vomerine teeth 17-34 .......... .......................... Oediea pseudomif6rmis

b Head narrower (ratio HWISVL 0.075-0.100); hind foot usually narrower (ratio HFWISVL 0.013-0.035); vomerine teeth 9-26 ............................................... 13

13a Snout long and narrow, tip truncate (Fig. 181a); maxillary teeth 19-27; vomerine teeth 9-17; dorsum grayish black with some small white spots on head, neck, shoulder, and upper limb surfaces; venter black ................................. Oedipina altura

15a Head long and relatively narrow (ratio HWISVL 0.075-0.095; Fig. 181~); maxillary teeth 14-43 ........... Oedipina paucidenteta

b Head short and relatively broad (ratio HWISVL 0.083-0.100; Fig. 181e); maxillary teeth 29-55 ............................................. 16

16a Dorsum and venter grayish black, often with a whitish area in parietal region andlor near limb insertions or joints ......... .................................. Oedipina p a d &

b Coloration usually not as above, usually head same color as dorsum or head in general slightly paler than dorsum and without a whitish area in parietal region .... 17

17a SVL to 44 mm in males, to 47 mm in females; occurs on Atlantic lowlands (sea level to 710 m elevation) Oedipina gracilis

b SVL to 57 mm in males, to 55 mm in females; occurs in uplands (750-2200 m

.................... elevation) Oedipina d r m i e

b Snout short, rounded (Fig. 18lb,c,e); maxil- Re- lary teeth 14-55; vomerine teeth 10-26; Bw 1963, 1968, Bw & DUELLMAN 1970, ............. coloration usually not as above 14 BRODIE & CMBELL 1993, LIPS 1993d,

14a Hind foot broader (ratio HFWlSVL M C C ~ et al. 1993, MCCRANIE 1996c, GOOD & 0.027-0.035); legs moderate (10-15% of W- 1997, GARC~A-PAR~S & WAKE 2000, SVL) .......................... M P ~ dmuda MCCRANIE et al. 2001. MCCRANIE 2006. KOHLER

b Hind foot narrower (ratio HFWISVL et al. 2007, MCCRANIE' et al. 2008, S&R et al. 0.013-0.29); legs tiny (7.8-11.3% of SVL) 15 2010

eudoeuryeea

Fhe genus Pseudoeurycea comprises about 60 species, inhabiting the cloud forests and Is". humid pine-oak forests in northern Mexico bnd Guatemala, usually above 1200 m ele- q d o n (WILSON & JOHNSON 2010). It is a -7: -i - * --". * ?&-;mi; qoup of moderate-sized to very large sala- .

manders with well-developed hands and Fig. 183. Pseudoeurycea brunnata (3 km W of && in which the middle digits me marked- El Rincbn, Quetzaltenango, Guatemala, ca. Itp longer than the outer ones. Some species 2400-2500 m). Photo: J. Hanken

4 Pseudoeulycea can be locally abundant, leaching densities higher than those of any other Guatemalan salamander (ELIAS El&&). Most species have very restricted

1; \pographic ranges. As pointed out by , { ~ C H & WAKE (1989), the genus Pseudo-

i auycea, as currently constituted, appears :: r to be a taxonomic group of convenience and ~2 not necessarily a natural unit. Indeed, 1,

t - nost species assigned to this genus are

r! h a c t e r i z e d mainly by their lack of L Fig. 184. Pseudoeurycea p m a (Cerro Badl, b. derived traits that define other plethodon- Chiapas, Mexico, 1900 m). Photo: T. J. Papenfuss I 1; tid genera. FROST et al. (2006a) placed 1, Wotn'ton in the synomymy of Pseudo- 7- -

SVL 59 mm; tail length/SVL ratio 1.08. Soconus- co area, Chiapas, Mexico, eastward to VolcAn Agua in Guatemala, 2400-3200 m elevation.

Pseudoeupea nigra (WAKE & JOHNSON 19891, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 411: 2; type locality: "12 km (7.5 mi) NW BerriozAbal, Chiapas, Mexico, elevation ca. 1,068 m." Maximum SVL in males 59.7 mm, in females 58.9 mm; tail 1engthISVL ratio 1.35-1.56. Known only from the type locality.

r. auycea. These salamanders have been h d beneath and within logs, under bark,

i; smss, grass clumps, and rocks and were b m e d active in the open on wet nights

i @LIAS 1984).

udoeurycea b m a t a BUMZAREM & SMITH 5, Herpetologica 11: 73; type locality:

de Socoausco, Chiapas, Mexico." um SVL to 60 mm; tail length/ml ratio 03. From Volch TacanA (in extreme

mistern Chiapas, Mexico), discontinuously ee~tward as far as V o l b Chicabal (in s~uth- Fig. 185. Pseudoeurycea goebeli (Lagu

Guatemala), 2400-2800 m elevation. Chicabal, Quetzaltenango, Guatemala, doeuqcea exspectata STUART 1954, Proc. 2722 m). Photo: S. M. Rovito . $06. Washmgton 67: 159; type locality:

roadleaf forest about 3 kin. west of the aldea Miramundo (about 7 air-line kilometers

satheast of Jalapa), Department of Jalapa, Gnatemala. Elevation, 2525 m." Maximum W L 52.7 mm; tail 1engtWSVL ratio 0.85. Only known from the type locality.

I

I

Pseudoeuiyeea pama L- & WAI(E 1989, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 411: 15; type locality: "on a ridge SE Cerro Bad, 21 km W Rizo de Om, Chiapas [error for Oaxaca], Mexico (ca. 1,600 m). The type locality is just within the eastern border of the state of Oaxaca", Mexico. M u m SVL in males 40 mm (females unknown); tail la&dSVL ratio 0.62-1.02. Cerro Bad, extreme eastern Oaxaca and adjacent Chiapas, 1500-2000 m elevation.

Pseudoezqcea rex ( D m 19211, Proc. Biol. SOC. Washington 34: 143; type locality: "Sierra Santa Elena (near Tecpam), Guatemala (alt. 9,500 feet)." Maximum SVL 48 mm; tail length/SVL ratio 1.07. Sierra de 10s Cuchu- matanes and the southwestern highlands of Guatemala; on Volch Tacani it also occurs on m the Mexican side, 2450-4000 m elevation.

Key to Pseudoeuzycea 1 a Pattern consisting of brown dorsal stripe

and lichenous white spots on tail; small ............................... species (SVL to 40 mm)

.................................. Pseudoeuzycea parva b Pattern not as above; larger species (adult

SVL usually >40 mm) .............................. 2 2 a Undersurface of tail with pale marblings

................................ on a dark ground color ................................ Pseudoeuqycea goebeli

b Undersurface of tail pale or dark, but always unicolor without marblings ......... 3

3 a Coloration solid black (in life by day, becoming brown by night) without any markings; legs conspicuously long (limbs overlap for 2.5 to more than 4 costal grooves); ratio tail length 1 SVL 1.35-1.56 ................................... P s e u d o e m nigra

b Coloration not as above; legs not conspic%- ously long (limb interval variable, from non-overlapping to overlapping for more than 2 costal grooves); ratio tail length I SVL ~ 1 . 2 0 ................................................. 4

4 a Adpressed fore and hind limbs overlap ..... ............................ Pseudoeuzycea arpectata

b Adpressed fore and hind limbs fail to overlap to the extent of at least 2 costal grooves -

5 a Coloration in preservative: dorsal surfaces pale brown, sides and belly darker brown ........................... P~8udoeulyCea bluzulata

b Coloration in preservative: dorsal surfaces pinkish gray, sides gray with or without white marbling, belly pale gray .................. ....................................... Pseudaeuzycea mx

Fig. 186. Pseudoeuzycea rex (Sierra de 10s' Cuchumatanes, Guatemala). Photo: T. BiUe

Fig. 187. Pseudoeulycea rex (Caserio Monzo, San Marcos, Guatemala, 2785).

Photo: S. E. Rovito

Further Reading WAKE & JOHNSON 1989, LYNCH & WAKE 1989, 1999, PARRA-OLEA et al. 2001, PARRA-OLEA 2008

~- - - Anura

Frogs and Toads (Anura)

Consisting of almost 6000 recognized species, frogs are among the most diverse groups of vertebrates, distributed almost ubiquitously woldwide; they are only absent from Antarctica, Arctic regions of North America and Eurasia, Greenland and some oceanic islands. They are characterized by usually having long, powerful hind legs, a broad head, a short body, large protuberant eyes, and the absence of a tail. Adult size ranges from less than 1 cm to 32 em. Frogs exhibit an enormous variation in morphlogy, behavior, life history, and reproductive mode. Although many species have the classic amphibian biphasic life history,

interdigital webbing

with an aquatic tadpole going through metamor~hosis to transform to an air- breathing adult, numerous species show moderate to extreme deviations from this theme. For example, many species have direct development within the egg capsule (e.g., species of the families Craugastor- idae, Eleutherodactylidae, and Strabo- mantidae), or the development of young in dermal vacuities on the dorsum (e.g., Pipa, Gastrotheca), in the stomach (e.g., Rheobatrachus), or even in vocal sacs (e.g., Rhinoderma). In Central America, 332 anuran species in 14 families are recognized.

subarticular tubercles accessory palmar tubercles n

outer metatarsal tubercle n

/ plantar tubercles A \

~ ~ ~ g i t a l webbing ,.

- inn; metatarsal tubercle subarticular tubercles

webbing formula: 12-2 314 I l l 1/2-8 111 1 314-3+ IV 3 1B- 1 If2 V

C.

Fig. 188. Morphological characteristics of anurans: a. general external morphology; b. palmar view of hand; c. plantar view of foot (b,c. Plectrohyla s a g o m ) . Drawings: a. G. Kohler; b,c. M. Vesely

Fig 192. Hind f i t of mops pu&zbus; note t m d tubercle (arrow),

Drawing M. Vedy

Fig. W. Eind foot af B b i ~ o p ~ u s dm& note spade-like inaer metatarsal tubenrclas bmws). Drawing: M . B d y

Fig. 190. Forefoot af Daadrobab auratuq note pairs of acute-Eke flaps ~mows) .

Drav&g M, Vedy

Fig. 191. hcilius valliceps (near San Jod El .

Idolo, Sucbitap4quez, Guaternda, 100 rp); note enlarged pardtoid glad@ Earrow4).

PBotc G. KBhler

Fig;. 1193. E&popa&w vm'alo~qe; note Imam- verse fold of akin across head posterior to ayes Carrowl. Photo: G. K~hler

Fig. 194 IF~padeFana pmmIbIepm Wl and H~obatrachiwa f l d 4 b (right); note transparent VB.XI~@~~. Photo: G. mhlet

Fig. 195. Tip of Ping- (lahrd view) is a. C r r t m b r ndei (note digtd groava); b. Leptcvda&~Ius bolidaw. Photos: G. KiMw

Anura

Key to families of h g s 1 a Four toes; two elongate spade-like inner

metatarsal tubercles, each with a free edge .......................... (Fig. 189) Rhino phrynidae

J b Five toes; no elongate spade-like inner

.... metatarsal tubercles, with a fkee edge 2 I a Wholly aquatic species with a distinctly

dorsoventrally flattened body and fully webbed feet; finger tips modified to form

........ three projecting lobes; tongue absent ......................................................... Pipidae

b Semiaquatic or terrestrial species without a distinctly dorsoventrally flattened body;

tongue present 3

1 degree of foot webbing variable; finger tips rlg. lvti, ventral surraces m a. ~eptodactylus not modified to form three projecting lobes; poecilochilus; note ventral disc (arrow).

.......................................... Photo: G. Kiihler - A cons~icuous tarsal tubercle resent at

end of inner tarsal foldA(Fig. 192); upper jaw without teeth; dorsum extreme-

........................................................ ly warty ........... Leiuperidae (in part: Eqgystamops) No conspicuous tarsal tubercle; upper jaw with or without teeth; texture of dorsum variable ..................................................... 4 A prominent deep purple to black colored lumbar gland, usually with pale (white to

...... pale blue in life) central spot (Fig. 672) ............. Leiuperidae (in part: P l e d e m a ) No such colored lumbar gland present ... 5 Well-developed parotoid glands present (Fig. 191) ................................................... 6

............................ Parotoid glands absent 7 Fingers with enlarged discs; pupil vertically elliptical in bright light (in life) or

................................................. preservative ................ Hylidae (in part: PhgIIomedusa) Fingers without enlarged discs; pupil horizontally elliptical in bright light (in life) or

................ preservative Bufonidae (in part) A pair of scute-like flaps on upper surface

........... of tips of fingers and toes (Fig. 190) ............................................. Dendrobatidae

No pair of scute-like flaps on upper surface of tips of 6ngers and toes ......................... 8 A transverse fold of skin across head posterior to eyes (Fig. 193); one or two fleshy ridges across posterior region of roof of mouth ..................................... Mimhylidae No transverse fold of skin across head posterior to eyes; no fleshy ridges across posterior region of roof of mouth ...................... 9 A digital groove completely around tips of digits (Fig. 195a); finger discs usually pre-

............... sent, toe discs always present 10 No digital groove completely around tips of digits (Fig. 195b); no finger discs, usually no toe discs ........................................... 13

108 A circular or V-shaped ventral disc present (Fig. 196); fingers without webbing ............

Terrarana (families Craugastoridae, Eleutherodactylidae, and Strabomantidae)

b No ventral disc; finger webbing present or ...................................................... absent 11

l l a Venter transparent, internal organs at least partially visible externally (Fig. 194); eyes directed forward at about a 45 degree

................................... angle Centrolenidae b Venter not transparent, internal organs

not visible externally; eyes oriented more ................................................... laterally 12

12a Fingers not conspicuously long and unwebbed, or, if so, males with nuptial excrescences; degree of finger webbing variable; no triangular dermal flap on each eyelid; no brood pouch in females; no fleshy

......................... proboscis Hylidae (in part) b Fingers conspicuously long, unwebbed,

lacking nuptial excrescences in males; a triangular dermal flap on each eyelid present or absent; a brood pouch in females present or absent; a fleshy proboscis pre-

.................. sent or absent Hemiphractidae 138 A distinct ventral disc present (Fig. 196);

no toe webbing ................. Leptodactylidae b No ventral disc; toes with substantial web-

.......................................................... bing 14 14a A distinct tympanum present; upper jaw

...................................... with teeth Ranidae b Tympanum absent; upper jaw without

........... teeth Bufonidae (in part: ate lop^)

Further Reading TAYLOR & GUTTMAN 1977, DUELLMAN & TRUEB 1986, FEDER & BURGGREN 1992, STEBBINS & COHEN 1995, MCDLARMID & ALTIG 1999, DUELLMAN 1999, FROST et al. 2006a, WELLS 2007, OGIELSKA 2009

Bufonidae The nearly cosmopolitan true toads (fami-

I

ly Bufonidae) are represented by approximately 470 species in 49 genera, distributed almost in all temperate, subtropical and tropical regions of the world with the exception of Madagascar, the Saharan region of Aftica, some oceanic islands, and the Australo-Papuan region (although Chaunus marinus has been introduced to many tropical regions worldwide including those in Australia). Just as with many other groups of anurans, the bufonids have been subjected to repeated substantial taxonomic revisions in the past 10 years (e.g., ~ i ~ . 197. Amplecting pair of hcilius pe+enes

2006, 2008, FkoST et al. 2009)- (Monteverde, Costa Rica). Photo: M. Franzen What had been grouped traditionally under a single generic name - Bufo - is now classified in several genera, four of which ( Chaunus, Incilius, Rhaebo, and Rhinella) occur in Central America.

In Central America, 39 bufonid species in seven genera are recognized. Synapo- morphies of the Bufonidae include the presence of a Bidder's organ (rudimentary ovaries present in males; absent in some D e n d r o p ~ s c u s ) , parotoid glands, and an edentate maxilla and premaxilla (FORD & CANNU 1993, PRAMUII 2006). Many bufonids have thick glandular skins with pustular warts and concentrations of glandular tissue. When stressed, these toads can secrete appreciable quantities of a vis- cous white poison from their parotoid glands.

Most species of this family are crepuscular or nocturnal, but some are diurnal. The majority of bufonids are terrestrial or semifossorial; exceptions include some fossorial and a few scansorial or semiaquatic species.

Key to the genera of Bufonidae 1 a Finger I shorter than Finger 11; tympanum

.................................. absent or indistinct 2 b Finger I same length or longer than Finger II, tympanum indistinct to prominent .... 6

2 a Inner digits rudimentary, enclosed by fleshy web; fingers (except 111) and toes (except IV) nearly fully webbed .................. ............................................. CrepPdo&pe

b Inner digits well developed; fingers and toes not fully webbed ............................... 3

3 a Skin generally smooth, but often with minute pustules dorsally and with well- developed spicules along anterior flank; arms very long and slender; no cranial

............................................ crests Ateloprte b Skin of dorsum rugose with numerous,

scattered, small to medium-sized tubercles andlor warts; arms of moderate length and

......... less slender; cranial crests present 4 4 a Tympanic annulus absent; vocal slits

........................................... absent in males RhiaeUa (in park R. amlopha) ...................

b Tympanic annulus present, might be concealed by skin; single sinistral vocal slit

....................................... present in males 5 6 a Tympanum distinct, not concealed by skin;

dorsal tubercles pointed; ground color greenish to olive brown; large toads, SVL in adult males 53-89 mm, 76-94 mm in

............................ females MUB (in part) b Tympanum indistinct, concealed by skin;

dorsal tubercles rounded; ground color grayish brown; smaller toads, SVL in adult

Atelophryniscus

In 1989, M c C m et al. described a new genus and species Atelophryniscus chrysophorus from premontane rainforest and cloud forest in north-central Honduras. PRAMKJK & LEHR (2005) demonstrated morphological similarities between A. chrysophorus and Rhinella veraguensis of South America and proposed to not recognize Atelophgmiscus as a valid genus. However, because of the geographic distance separating A. chrysophorus and the northernmost species assigned to the R.

Fig. 198. Amplecting pair of hcilius canaliferus veraguensis group (some 2600 b), m d (San Lorenzo, Suchitep6quez, Guatemala, 210 because no molecular data are available to m). Photo: G. Kiihler support this taxonomic decision, 1 prefer to

recognize the genus Atelopluyniscus as valid until additional evidence has been published.

.... males 3540 mm, 49-60 mm in females Atdo- A t e l o p ~ s C L I s chrysopho*s was Collect- ...........................................

6 a A well-developed i ~ e r tarsal fold present; ed along a stream; during daytime, males parotoid gland very large, several times the area of upper eyelid .................................. 7 were encountered sitting on rocks or bod-

b Inner tarsal fold absent; parotoid gland ders in the stream and, usually small to moderate-sized, less than when disturbed, they would jump into the twice the area of upper eyelid ................. 8 stream and swim to the bottom to conceal

, 7 a Cranial crests present; no dark brown to themselves among debris in the bottom of black lateral longitudinal band from head the pools; at night, males were observed in to at 'east posterior margin of ~arotoid the vegetation along the &ream, 1-2 m gland ......... Chaunue (in part: C. manhue)

b No cranial crests; a dark brown to black above the ground (MCCRANIE & WILSON

lateral longitudinal band from head to at 2002). Am~lectant pairs were found in I ... least posterior margin of parotoid gland m i d - F e b r u ~ during a period of no precip-

.......................................................... L h itation for several weeks (MC~RANIE & I 8 a Pretympanic and parietal crests usually

........................ prominent InciIiue (in part) 1 b Pr&ympanic and parietal crests usually

indistinct ................................................... 9 9 a Nostrils oriented laterally or dorsolateral-

t ly; supratympanic crest prominent and I continuing over tympanum as a narrow

shelf ................ Rhinella (in part: R. alata) b Nostrils oriented dorsally; supratympanic

crest indistinct and not continuing over tympanum as a narrow shelf ...................... .................... Chaunue (in part: C. mW)

BLAIR 1972, HOO&OED 1989, GRWBEAL & DE QUEIROZ 1992, PAULY et al. 2004, MULCAHY et al. Fig- lg9. Atelopfiscus chWsophoms 2006, PRAMUK 2006, PRAMUK 2008, FROST et al. (Quebrada de Oro, Atlhtida, Honduras).

I 2009 - Photo: J. R. McCranie

WILSON 2002). The tadpoles of this species show adaptations to living in fast-flowing streams, such as depressed bodies with flattened venters and an enlarged ventral mouth followed by a large suctorial disc (MCCRANIE et al. 1989).

Atelophryniscus chrysophorus MCCRANIE, WILSON & W m s 1989, Occas. Pap. Mus. Nat. Hist. Univ. Kansas 129: 3; type locality: "Quebrada de Oro (15'383, 86'47W), elevation 880 m, tributary of Rio Viejo, south slope of Cerro BMalo, Cordillera de Nombre de Dios, Depto. Atliintida, Honduras." Males to 40.1 mm, females to 59.6 mm. Central and western portions of the Cordillera Nombre de Dios on the Atlantic versant of north-central Honduras, 750-1760 m elevation.

r Atelophryniscus chrysophorus

C

F'urther Reading MCCRANIE et al. 1989, LAVILLA & DE SA 2001, MCCAANIE & WLLSON 2002, MCCRANIE AND C A S T ~ D A 2005,2007, PRAMuK & L E ~ R 2005

The genus Atelopus is a tropical New World endemic and contains 89 currently recognized species (FROST 2010), collective- ly known as "harlequin frogs," which are found in the uplands and lowlands from Costa Rica south to Bolivia and eastward ' into the Guyanas. The adults of most

'

species typically exhibit gaudy, aposematic ] colorations with yellow, orange, red, green . andlor black elements. On the contrary, ,

metamorphs and juveniles of some species I are cryptically colored (e.g., in juvenile A. , zeteki, olive green with dark brown blotches). In A. zeteki, the cryptic juvenile coloration is apparently correlated with the absence of toxic skin compounds (LINDQUIST & HETHERINGTON 1998a). The ontogenetic change in coloration is apparently accompanied by acquisition of highly toxic skin compounds ( D m et al. 1994, 1997). The compounds in the skin secretions are neurotoxic (alcaloids) and car- diotoxic (steroids) (SAVAGE 2002). - Species of Atelopus are terrestrial, diurnal, and usually associated with fast flowing forest streams of slope and highland regions. Walking is the usual locomotion mode for these frogs when foraging or seeking mates. Hopping is used mainly to attempt to escape potential predators. Males are territorial and the advertisment call and - in case an intruder is not deterred by the call - physical combats are used to maintain the territory. At night, individuals of Atelopus have been reported to sleep in the vegetation up to 1 m above the ground or to hide beneath rocks (LOTTERS 1996). In the Central American species, breeding activities (amplexus and oviposition) take place during the wet season (May to July in A. chiriquensis, July to August in A. senex, September to early December in A. varius). Amplexus is axillary and the egg strings are deposited into streams. The tadpoles have a large ventral suctorial disc that allows them to attach to rocks or other objects in the stream.

Fig. 200. Atelopus certus (6.2 km up Rio Piiia h m Bahia Piiia, DariBn, Panama, 85 m).

Photo: A. J. Crawford

r. 201. Atelovus certus (Cerro S a ~ o , Daribn,

Atelopus species have suffered eclines in the past 10-15 years,

the Central American species CA & L O ~ R S 1997, SAVAGE 2002,

WCA et al. 2005).

i s certus BARBOUR 1923, Occas. Pap. 001. Univ. Michigan 129: 12; type loca-

"from a stream on Mt. Sapo, Provincia ." Males to 32 mm,

to 42 mm. Cerro Sapo in southwestern Province, Panama, 50-1150 m eleva-

chirripoensis SAVAGE & BoLARoS Biol. Tropical 51: 382; type locality:

southeastern side of Norte, ca. 4 km N Cerro Chirrip6

e; Parque Nacional Chinipb, Cordillera anca, Distrito Limbn, Cmtbn Limbn, Lirn6n (ca. g032'N, 83"29'W), a t ca.

0-3500 m above sea level." SVL 42.5 mm in

Fig. 202. Atelopus chin'quiensis (male; Las Tablas, Puntarenas, Costa Rica).


Fig. 203. Atelopus chiriquiensis (female; Las Tablas, Puntarenas, Costa Rica).

Photo: M. Garcia Parj

only known female. Known only from the type locality.

Atelopus chiriquiensis SHREVE 1936, Occas. Pap. Boston Soc. Nat. Hist. 8: 269; type locality: "'from Mr. Lewis' place, 14 km NE El Hato del Volch, 1500 m], Rio Chiriqui, Viejo and branches, Panama Republic." Males to 34 mm, females to 49 mm. Cordillera de Talamanca of Costa Rica and western Panama, 1400-2500 m elevation.

Atelopus glyphus D m 1931, Occas. Pap. Boston Soc. Nat. Hist. 5: 396; type locality: "Pini Range, [near head of Rio Limbn,] DariBn", Panama. Males to 36 mm, females to 48 mm. Eastern Panama, in the Serrania de Pirre, and also the Choc6 of Colombia, 884-1500 m elevation.

Atelopus limosus IBAREZ, JARAMILLO & SOL~S 1995, Caribb. J. Sci. 31: 57; type locality: "en las orillas de la Quebrada San Antonio (09"24'00"N, 79"31'07"0), aproximadamente a

180 m sobre el nivel del mar, cerca del poblado San Juan de Pequenf, Provincia de Panaman, Panama. Males to 30.7 mm, females to 40.2 mm. On the Atlantic versant of the eastern portion of central Panama, near sea level to 730 m elevation.

Atelopus senex TAYLOR 195213, Univ. Kansas Sci. Bull. 35: 630; type locality: "near Desengaiio] pass between V o l h Po& and Volch Barba, western slope Volcan Pods, Pacific drainage, elev. app. 6800 R.", Cantdn de Santa Barbara, Provincia de Heredia, 2073 m elevation, Costa Rica. Males to 32 mm, females to 43 mm. Central Costa Rica in the Cordilleras Central and Talamanca, 1100-2200 m elevation.

Atelopus varius (LICHTENSTEIN & MARTENS 1856), Nomencl. Rept. Amph. Mus. Zool. Berol.: 40; type locality: Veragoan; restricted to '%he Pacific portion of the Provincia Veraguas, western Panama." by LOITERS et al. (1998). Males to 39 mm, females to 48 mm. Both versants of the cordilleras of Costa Rica and western Panama, 1G2150 m elevation.

Hey to Atelopus 1 a Legs short, ratio shank length1SVL ~0.35;

pretympanic and temporal regions, flanks, lateral and upper surface of upper arm, upper surfaces of thigh, sacral region, and area lateral to vent with pale tipped coni . ................................. Atelopus Wpoenaia

b Legs longer, ratio shank length/SVL >0.35; without coni or spiculae in regions listed above ......................................................... 2

2 a Coloration in life brick red with black spots (pale reddish with black spots in preservative) ................................... Atelopus certus

b Coloration not as above ........................... 3 3 a No differentiated glandular areas, skin

smooth throughout; tips of fingers slightly ...................................................... swollen 4

b Skin of rostrocanthal area, dorsum and knee region glandular; tips of fingers tapering .............................................................. 'I

4 a Dorsum dark greenish olive to chocolate- brown in life, without blotches or other markings; males with a lateroventral black line; juveniles lime green in life with dark brown markings ............. Atelopus limhsus

Ai$elopus zeteki DuNN 1933b, Occas. Pap. Boston Soc. Nat. Hist. 8: 71; type locality: "El Valle [del Ant6nIn, Panama. Males to 40 mm, females to 58 mm. East of the main Tabasard ridge in Provincias Cocl6 and Panama, Panama, 250-1315 m elevation.

Fig. 204. Atelopus glyphus (Cana, Daribn, Panama, 1300 m). Photo: A. J. Crawford

b Dorsal coloration not as above; males without a lateroventral black line ................. 5

5 a Dorsum brown (in life?), uniform or with ................................. yellowish white spots

........................................ Atedopus gIpp6us b Dorsal coloration not as above ............... 6

6 a Dorsum golden yellow in life with transverse black bands, with black blotches, or uniform golden yellow ..... Atelopus eeteHi

b Dorsal coloration mostly black or chocolate brown with yellow to red bands and blotches ...................................... Atelopus v d u s

7 a Glandular areas well developed; coloration in life black, bluish gray or gray-green, with green to pale orange spots and blotches, or uniform, some individuals with an orange suffusion ............... Atelopus senex

b Glandular areas weakly developed; coloration in life yellow, green, rust, or black with red or yellow spots, blotches, or lines ............................... Atedopus ~ q r u q r u e n s i 6

Fig. 203. ntelopus limosus (Burbayar, Cuna, Panama). Photo: B. Kubicki

Fig. 210. Atelopus senex (Cemo Chompipe, Heredia, Costa Rica). Photo: W. Van Devender I

Fig. 211. Atelopus van-us (Cerro Negro, Veraguas, Panama, 1090 m). Photo: S. Lotzkat

Fig. 212. Atelopus zeteki (Valle de Antbn, Panama). Photo: G. Kiihler

The taxonomy of the South and Central American species formerly assigned to Bufo (sensu lato) has been subject to repeated name changes, and much controversy exists among recent authors as to content and hierachical level of the recognized genera (e.g., CHAPARRO et al. 2007,

r'ig. 213. Atelopus glyphus. Photo: B. Wilson SAVAGE & B o ~ o S 2009b9 PAUL, et 2009, VAN BOCXLAER et al. 2010). Contrary to the proposal of CHAPARRO et al.12007), I here follow SAVAGE & B O ~ O S (2009b) in recognizing the genus Chaunus for the marinus and granulosus species groups, and Rhinella for the margan'tiferus species group and those species formerly placed in Rhamphophryne. NARVAES & RODRIGUES (2009) revised the Chaunus granulosus group recognizing twelve species (single Central American representative: C. centralis) .

Further Reading STARRETT 1967, DUELLMAN & LYNCH 1969, SAVAGE 1972a, CRUMP 1986, POUNDS & CRUMP 1987, CRUMP 1988, C R ~ & POUNDS 1989, COCROFT et al. 1990, IBAREZ et al. 1995, L~DQUIST 1995, LINDQUIST & ~~ETHERINGTON 1996, LOTTERS 1996, LINDQUIST & SWIHART 1997, LA WCA & LBTTERS 1997, LINDQUIST & HETHERINGTON 1998a,b, LINDQUIST et al. 1999, RAMos et al. 2002, LA W C A et al. 2005, KARRAKER et al. 2006, RICHARDS & KNOWLES 2007

The marine toad ( Chaunus mruinus) is the largest toad of Central America (females can weigh up to 1.5 kg) and is often found in habitats severely altered by human activities. In fact, it is usually more common around human settlements and agricultural areas than in its natural habitat (savanna, secondary growth, and open forest), and closed-canopy rain forest seems to be an effective barrier for this species (ZUG 1983). Adult marine toads are mostly nocturnal, although they begin to leave their daytime shelter at dusk. On the contrary, juveniles are diurnal. Whereas exotic populations of the marine toad have been assumed to be acyclic, the reproductive activities of native populations reflect the seasonality of the local environment; tim- ing seems to assure that metamorphosis takes place during periods of high humidity and prey abundance (ZUG 1983). Females produce paired strings with several thousands of eggs in the shallow water of both permanent and temporary water bodies. The small black tadpoles metamorphose in 60-75 days (SAVAGE 2002). The toxic and noxious properties of eggs and

Bufonidae

tadpoles are efficient for antipredator defense. The juveniles can usually be differentiated from other lowland toads by regular rows of paravertebral tubercles.

Chaunus centralis NARVAES & TREFAUT RODRIGUES 2009, Arquiv. Zool. 40: 46; type localitv: "Chitre. Herrera. Panama." Males to

p5 ,,,, 58.4 hm, femiles to 61.8 mm. Eastern Panama, sea level to 200 m elevation.

Chaunw marinus (LINNAEUS 17581, Syst. Nat. .&>&, I .-d. 10, 1: 211; type locality: "America." Males .-., L . -- , to 145 mm, females to 175 mm. From southern

Texas, USA, through tropical Mexico and b 2U. amus (' kXf2 Cenbal America to northern South he- ; Panama). Ph&W. E. he- introduced to many tropical countries (and

ofthe B i o Q i e * Ins~tuWs u&e~!&b of -f subtropical regions within largely temperate countries, e.g., the USA) world wide, sea level to 3000 m elevation (in Central America up to

1 1600 m elevation).

I Key to aaunus 1 a Parotoid gland very large, several times

the area of upper eyelid; SVL 85-145 mm in adult males. 90-175 mm in adult

I F Rg. 215. Chaunus marinus (Santa Eulalia, Alajuela, Costa Rica, 825 m). Photo: G. Kiihler I

females ........................ Chaunus marinus b Parotoid gland of moderate size, not larger

than the area of upper eyelid; SVL <60 mm in adult males, <66 mm in adult females ........................................................ ...................................... Chaunus aentkah

Chaunus centralis 1 :haunus marinus

Further Reading ZUG & ZUG 1979, ZUG 1983, EASTEAL 1986, NARVAES & RODRIGUES 2009

u ~ u n u s marinus (Bartola, Rio San ragua, 30 m). Photo: G. Kahler

--

Crepidophryne

The species of the genus Crepidophryne are small toads that have been documented a t localities between 1050 to slightly over 2000 m elevation in wet montane forest on the Atlantic versant of Costa Rica and western Panama (SAVAGE 2002). According to VAUGHAN & MENDELSON (20071, three species with disjunct ranges are recognized. Very little is known about the life history of these toads. They are diurnal inhabitants of primary or mature secondary forests (cloud forest or highland oak forest with closed canopy) where they dwell apparently fossorially to semi- fossoriallyin deep layers of leaf litter (SAVAGE 2002). During the months of April, small aggregations of 2-8 individuals of C. epiotica have been observed in the morning hours (VAUGHAN & MENDELSON 2007). The only available information on the reproductive biology of these toads was generated from the dissection of two gravid females of C. epiotica (one collected in March, the other one in October, both with approximately 55 eggs) and one of C. guanacaste (collected in April, number of eggs not reported); the mode of amplexus is unknown and no tadpole has ever been found (VAUGHAN & MENDELSON 2007).

Crepidophryne chompipe VAUGHAN & MENDELSON 2007, Copeia 2007: 308; type locality: "Costa Rica, Heredia [provincel, Cant6n San Rafael, Distrito Los Angeles, Reserva Dantas." Males to 27.3 mm, females to 33.5 mm. Cordillera Central in Costa Rica (several localities near Cascajal, Cerro Chompipe, and Reserva Dantas), 1400-2050 m elevation.

Crepidophtpe epiotica (COPE 18761, J. Acad. Nat. Sci. Philadelphia Ser. 2,8: 97; type locality: %om 5000 feet elevation on Pico Blanco" [= Cerro m m u k , Cant6n de Talamanca, Provincia Limhn], Costa Rica; in error for "Cerro Utyum, 5000 ft, Cant6n de Talamanca, Provincia de Lim6n: 1524 m", according to SAVAGE (1974). Males to 33.3 mm, females to 40.2 mm. Atlantic versant of the Cordillera de Talamanca in Costa Rica and western Panama, 1050-2040 m elevation.

Fig. 217. Crepidophryne chompipe (Cascajal de Coronado, San JosB, Costa Rica, 1740 my.

Photo: G. kohl^

Fig. 218. Crepidophryne chompipe (same as in Fig. 217). Photo: G. Kiihler

Crepidophryne guanacaste VAUGHAN & MENDELSON 2007, Copeia 2007: 308; type locality: 'Costa Rica, Guanacaste [Province], Cant611 Bagaces, Distrito Fortuna, Volchn Miravalles." Males to 21.6 mm, females to 31.6 mm. Volcanes de Miravalles and Rinc6n de la Vieja in the Cordillera de Guanacaste, Costa Rica, 1820-2000 m elevation.

mt" 7 d - e 1 a Finger &ee fm almost half its length (Fig. ...................... aisbs **e-&&*~

b JTinger I completely embedded in p a w e .................................. webbing 21Ba) 2 t a S U on dar& s e w relatively srnmth,

with many iaclistiwt. low, rounded tuber- oles; lateral row of tuherdm ia&stin&, low, rou~ded; lateral a d m of body newly ern&, with few, nrnnded tubercles; scapu= lar texture generally &dm to tlhblt ofsur - TOW- areas, with i n W c t 3 low, romd- ed hberolw, domum and venter nearly un&nn dull brown-gray .......,........... ,,.,. .,., ......................... w&@me&m-=-

b Skin on doma1 sarfacea rough, with dis- Ihet3.y raised, founded or poiaha tubercles; late~al row of tube~cles large sharply poiated; latezal surface of bady with shsrply pointed tubercles; eeapular textme distinct &om monndiag are- lacking distinct tubmkq dozsm d d b m with irregular dark brown markings, venter dull E brown wit& c?mam mar-* e-y on .......................... the pelvic areas and limbs I ................... ..,.. ...... ampid* t&mriplPe

[k& 219. Ventral view of hand in a. Fig. 222. Crepidophgme epiotica (Rio Macho

Fig. 221. Crepidophxyne epiotica Quvenile, 7,7 ram SVL, Rio Changena, Panama, 1650 m).

Photo: A. Hertz

e chompipe and b. C. epiotica. de Orosi, Cartago, Costa Rim, 2000 m). Photos: G. Kiihler Photo: G. KLihler

105

The species of the genus hcilius are distributed from the southern USA south to Panama and then south along the Pacific versant to Ecuador. Most species in this genus have well-developed cranial crests and parotoid glands. As currently understood, the genus Incilius is represented in Central America by 22 species. This group of anurans has experienced repeated taxonomic name changes in the past few years. FROST et al. (2006a) initially moved the members of this group to a separate genus, Cranopsis, only to Gnd out a little later (FROST et al. 2006b) that Cranopsis was unavailable. The latter individuals applied the generic name Ollotis for this taxon. Unfortunately, FROST et al. (2006a,b) neglected to note an available name for this taxon, Incilius Cope 1863 (type species: Bufo coniferus COPE 1862b; designated by KELLOGG 1932:29), and for reason of priority FROST et al. (2009) transferred all species included in Ollotis to Incilius. Note that some herpetologists (e.g., following Pauly et al. 2004) are retaining the use of Bufo as the valid name for this group of toads..

MCCRANJE & CASTAREDA (2007) did not follow the taxonomy for the Ollotis coccifer complex suggested by MENDELSON et al. (2005). Despite the obvious difficulties in indentifying species of this complex based on the information provided by MENDELSON et al. (2005), I prefer to follow this arrangement until a different view based on published evidence is available. Contrary to the claim of L~PEZ et d . (2009), Incilius occidentalis (CAMERANO 1874) and I. per- plexus (TAYLOR 1943) have not been recorded east of the Isthmus of Tehuantepec (J. MENDELSON pers. comm. July 2010) and, therefore, occur outside of the area covered in this book.

Most species of this genus are crepuscular or nocturnal, but some highland species are diurnal. They are usually terrestrial or

Fig. 223. Lncilius aucoinae (GoKto, Puntarenas, Costa Rica, 120 m). Photo: G. Kohler

Fig. 224. Incilius bocourti (San Cristdkal de las Casas, Chiapas, Mexico, 2136 m).

Photo: A. Ramirez Veliizquez I semifossorial, exceptions including some scansorial species such as Incilius com'ferus that can be found several meters above ground level on shrubs or trees (KOHLER 1993). Amplexus is axillary and egglaying takes place in shallow water of ponds, lakes, and water courses with slow current. Most species of the I. valliceps group (i.e., I. aucohae, I. campbelli, I. macrocristatus, I. melanochloris, I. tutelarius) breed in streams during the dry season; I. valliceps is unique within this species group by laying eggs in ponds during the wet season (MENDELSON et al.1999). Eggs are typically laid in two strings. Tadpole aggregations Pschools") of hundreds to thousands of individuals are common in many Incilius species. When disturbed, the groups can break up but are reestablished within a short time. Recently metamorphed froglets of Incilius species are tiny and usually have no cranial crests or have them poorly to partially developed (SAVAGE 2002).

The genus Incilius includes a assemblage of highland species that are diurnal and eemifossorial: the members of the fastidiosus and periglenes groups. The golden toad, I. periglenes, spends most of the year underground except for a few weeks near the end of the dry season (March through June) when they emerge to breed in temporary pools (CRW 1989a, CRW et al. 1992). The only known population of I. periglenes declined rapidly in the 1980s, with the last individual seen in 1989 ( C R ~ et al. 1992). The species is now con-

fncilius aucoinae (O'NEILL & MENDELSON %004), J. Herpetol. 38: 487; type locality: "air- port a t the town of Aguabuena, Distito Sierpe, Osa Peninsula, Puntarenas Province, Costa Bica (approximately B026'N, 83O38W." Males to 67.2 mrn, females to 104.5 mm. On the Paciiic versant from central Costa Rica to weehrn Panama, 25-500 m elevation.

M u s bocourti (BROCCHI 1877c), Bull. Soc. Pbiiomath., Paris, Ser. 7, 1: 186; type locality: 'Totonicapam", Chiapas, Mexico. SVL to 77 inm in females (males smaller). Southwestern Chbpas (Sierra Madre del Sur and central Plateau of Chiapas), Mexico to southern Guatemala, 10003200 m elevation.

hcilius campbelli (MENDELSON 1994), Occas. Pap. Mus. Nat. Hist. Univ. Kansas 166: 4; type kudity "Las Escobas, 5.1 km W Puerto Santo %mBs, 104 m, Montaiias del Mico, Departa- m a t o de Izabal, Guatemala." Males to 62.6 mm, females to 86.2 mm. Northern and east- 81p Guatemala, southern Belize, and extreme western Honduras, 100-1200 m elevation.

M u s canalXem (COPE 18771, Proc. Am. Philos. Soc. 17: 85; type locality: T e s t Tehu8ntepecn, Oaxaca, Mexico. Males to 43

, females to 52 mm. On the Pacific versant

Fig. 225. LnciLius canaliferu, ,A inca Irlanda, Chiapas, Mexico, 1050 m). Photo: G. Kahler

Fig. 226. Incilius coccifer (near San Jose El Idolo, SuchitepBquez, Guatemala, 100 m).

Photo: G. Kiihler

Fig. 227. Incilius coniferus (Fortuna, Chiriqui, Panama, 1030 m). Photo: G. Kohler

rig. am. Incilius campbelli (Morales, Izabal, Gua- temala, 920 m). Photo: C. R. Vhsquez Almazh

lantic versantlof Honduras and from the. lantic coastal regions of Honduras and

Nicaragua, sea level 2080 m elevation.

Incilius coniferus (Cope 1862b), Proc. Acad. Nat. Sci. Philadelphia 14: 158; type locality: "Turbo region, New Granada" = Choc6, Colombia, according to COCW & GOIN (1970). Males to 72 mm, females to 94 mm. North-central Nicaragua to Pacific lowlands of Colombia and northern Ecuador, sea level to 1550 m elevation.

Fig. 229. InciIius fastidiosus (Lm ' as, Punt Incilius fastidiosus (COPE 1876), J. Acad. Nat. renas, Costa Rica). Photo: M. Garcia Paris sei. Philadelphia Ser. 2, 8: 96; type locality:

"2500 feet elevation on the slope of the Pico Blanco ICant6n de Talamanca. Provincia

Fig. 230. Incilius holdridgei (Cerro Chompipe, Heredia, Costa Rica, 2090 m). Photo: G. Kohler

Fig. 231. Incilius ibarrai (San Miguel Ixtahuach, San Marcos, Guatemala, 2050 m).

Photo: M. Acevedo

Fig. 232. Lncilius karenlipsae (holotype). Photo: J. R. Mendelson

Lim~jn], the district of urenn,'Costa Rica ["probably between the Rio Uren and Rio Lari a t 760 m," according to SAVAGE (197411. Males to 52 mm, females to 60 mm. Both slopes of the southern Cordillera de Talamanca of Costa Rica and the Atlantic slope of adjacent Panama, 760-2100 m elevation.

Incilius holdridgei ( T A ~ O R 1952b1, Univ. Kansas Sci. Bull. 35: 607; type locality: "at an elevation of approximately 7500 ft. [2272 m] on Volch Barba, western slope", Cant611 de Santa Barbara, Heredia Province, Costa Rica. Males to 46 mm, females to 53 mm. Volch Barva, Cordillera Central, Costa Rica, 2000-2200 m elevation.

Incilius ibmai (STUART 19541, Roc. Biol. Sm. Washington 67: 162; type locality: "Oak-pine zone a t Aserradero San Lorenzo (about 12 airline kilometers slightly east of north of Jalapa), Department of Jalapa, Guatemala. Elevation, 1725 m." Males to 82.4 mm, females to 78.4 mm. Central and southern Guatemala, to southwestern Honduras (see comments in MENDELSON et al. 20051, 1360-2050 m elevation.

Fig. 233. Incilius luetkenii (San Juan de Dios, Chontales, Nicaragua, 60 m). Photo: J. Sunyer

i hcilius karedpsae MENDELSON & MUW:AHY 2010, Zootaxa 2396: 62; type locality: "Panama: Cocl6.Province: El Cop&, Parque Nacional G. D.

I Omar Torrijos (8"40'N, 80Q37'17"W; approximately 850m)." SVL 89.3 mm in single known male. Only known from the type locality.

Incilius leucomyos (MCCRANIE & WILSON 2000), J. Herpetol. 34: 22; type locality: "Quebrada El Pin01 a t its confluence with Quebrada Las Cantinas (15"09'N, 86'43W),

I b a u e Nacional La MuralIa. 950 m elevation, ~ e p k a m e n t o de Olancho, ~ondurag." Males Fig. 234. InciLius luetkenii (P.N. El Imposible, to 64.4 mm, females to 90.7 Disjunct Pop- Ahuachaph, El SaIvador, 720 m). ulations on the Atlantic versant of north-cen- Photo: G. Kohler tral Honduras, sea level to 1600 m elevation.

Incilius luetkenii (BOULENGER 18911, Ann. Mag. Nat. Hist. Ser. 6, 8: 455; type locality: "Cartago, Costa Rica"; type locality considered erroneous by SAVAGE (1974). Males to 96 mm, fiemales to 107 mm. Pacific lowlands from Chiapas, Mexico to central Costa Rica, and on the Atlantic dope in dry interior valleys of Guatemala and Honduras, and in the upper Rfo San Juan drainage in Costa Rica, sea level to 1300 m elevation.

W u s macrocristatus (FIRSCHEIN & SE~PI~I 1957), Herpetologica 13: 219; type locality %tween La Gloria and Cerro Azul (presumably nearer C e m Azul), Oaxaca", Mexico; the original collector reported the locality as "Cerro Atravesado in view to SE and Cerro Azul to E or slightly S of E ...." Males to 68.7 nun, females to 76.8 mm. From the Los Chimalapas region in south-central Oaxaca, Mexico, south and east along the Atlantic versant as far as the Sierra de 10s Cuchwnat.gee in Guatemala, 200-1600 m elevation.

hcilius marmoreus (WIEGMANN 18331, Isis von Oken 26: 661; type locality: "mexicanischen"; rendered as Vera C d , Veracruz, Mexico by lWmx (1932). SVL to 75 mm. Pacific versant of Mexico from southern Sonora to northern Chiapas; also in the Balsas basin in adichoactin and on the Atlantic versant in Veracruz, sea level to 1100 m elevation.

Jndius melanochlom (COPE 1877), Proc. Am. Mas. SOC. 17: 85; type locality: "Bast Costa &can; specified as "Costa Rim, east or eastern, 4CanMn de Limhn, Provincia de Lim6n" by $AVACE (1974). Males to 73.6 mm, females to 106.7 mm. On te Atlantic versant from southeastern Nicaragua to northern Panama -tly recorded in Bocas del Toro Prwince by J~ARAMILLO et al. 20091, near sea level to 1080 rn elevation.

heiIius periglenes ( S A V ~ 1967 "1966"), Rev. Biol. Tropical 14: 153; type locality: "Costa

Fig. 235. Incilius mat &tatus (Raydn, Chiapas, Mexico, 1888 m). Photo: A. Ramirez Velizquez

rg. haw. LnciLius macrocristatus (Rayon, Chiapas, Mexico, 1888 m). Photo: A. Ramirez Velizquez

I Fig. 237. hcilius melanochloms (Rio San Juan, Nicaragua, 30 m). Photo: G. Kohler

Fig. 238. Incilius periglenes (male, Monteverde, Costa Rica). Photo: M. Franzen

Fig. 239. bcilius porteri (Cerro Uyuca, Fran- cisco Morazh, Honduras). Photo: G. Kiihler

Fig. 240. bcilius tacanensis (Top of Cerro Mozotal, Chiapas, Mexico, 3016).

Photo: S. M. Rovito

Rica: Provincia de Alajuela: Cant6n de San Carlos: Cordillera de Tilarb, 2 miles ENE of Monteverde, Provincia de Puntarenas; 1590 meters." Males to 48 mm, females to 56 mm. This species was known only from the Reserva Biolcigica Monteverde, Costa Rica, 1500-1620 m elevation.

Lnciliusperipatetes (SAVAGE 19721, J. Herpetol. 6: 26; type locality: "Panama: Provincia de Chiriqui: above Boquete on Almirante trail, 1500 m." Males to 67.0 mm, females to 72.3 mm. Known only from two sites: the type locality and Cerro Bollo (both in the western Panama), 1500-1856 m elevation.

Incilius porteri (MENDELSON, WILLIAMS, SHEIL & MULCAHY 20051, Sci. Pap. Nat. Hist. Mus. Univ. Kansas 38: 17; type locality: "6 mi l9.6 km] NE Escuela Panamericana, Cerro Uyuca, 5200 ft [I584 ml, Francisco MorazBn, Honduras." Males to 58.7 mm, females to 73.0 mm. Cerro La Tigra and Cerro Uyuca in the department of Francisco MorazAn, Honduras, 1584-2100 m elevation.

Incilius signifer (MENDELSON, WILLIAMS, SHEll. & MULCAHY 20051, Sci. Pap. Nat. Hist. Mus. Univ. Kansas 38: 20; type locality: "7 mi N (11.2 km) and 2 mi (3.2 km) W of David, Chiriqui, Panama." Males to 64.1 mm, females to 77.0 mm. On the Pacific versant in the vicinity of the city of David, Chiriqui, and then from the Province of Veraguas to the Canal Zone, Panama; sea level to 900 m elevation.

Incilius tacanensis (SMITH, P.W. 1952), Copeia 1952: 176; type locality: "at 1500 meters on Volch de TacanB, Unicin Ju&-ez, Chiapas, Mkxico." SVL to 53 mm. On the Pacific versant of eastern Chiapas (Unicin de Juarez), Mexico, to western Guatemala, 200-2200 m elevation.

Fig. 241. InciIius vallicep~ \ l v l v - ~ h e r , Ag. 242. h d u s valIiceps (Cerro El Toro, Atlan- Toledo, Belize, 7 m). Photo: G. Kiihler tico Norte, Nicaragua, 830 m). Photo: G. Kiihler

Bufonidae

Incilius tutelarius (MENDELSON 19911, Herpetologica 53: 15; type locality: &Colonia Rodulfo Figueroa, 19.0 km NW Rizo de Oro, Oaxaca, Mexico, 1370 m (16"32'N, 94"08'W)." Males to 76.2 mm, females to 103.6 mm. On the Pacific versant from the Chimalapas region in southeastern Oaxaca, Mexico, to extreme western Guatemala, 1000-2000 m elevation.

hcil ius valliceps (WIEGMANN 1833), Isis von Oken, 26: 657; type localities: "mexicanischen"; "Mexicon and "Vera Cruz, Mexico." Males to 76 mm, females to 82 mm. On the Atlantic versant from central Veracruz, Mexico, to northern Costa Rica; and on the Pacific slope .from around the Isthmus ofTehuantepec, Mexico, to

' extreme western El Salvador (see K O ~ E R et al. 2006), sea level to 2000 m elevation. Bufo

i valliceps wilsoni BAYLOR & STUART 1961 is

, regarded as a synonym of I. valliceps according to M E N D ~ Q N (1998).

I Fig. 241. Lncilius tutelarius (Jiquipilas, Chiapas, Mexico, 1214 m). Photo: A. Ramfrez VelAzquez

lncilius valliceps

' I

lncilius coccifer ' 'ncilius ibarrai - 'ncilius porteri r lncilius tutelariws

*-- PL lncilius canalifem

lncilius carnpbelli Incilius bocourti lncilius karenlipsa

lncilius rnacrocristatus r lncilius tacanensis

Key to Incitme 1 a Tympanum absent or indistinct (Figs. 244

and 245) ..................................................... 2 ................. b Tympanum distinct (Fig. 246) 7

2 a Parotoid gland very large, its length almost equal head length ..................................... 3

b Parotoid gland small; hands and feet with fleshy pads ................................................ 4

3 a Parotoid gland at least twice as long as wide and its anterior border closely attached to the posterior border of eye; postocular crest well developed; tarsal fold pronounced ....................... hcilius bwourti

b Parotoid gland less than twice as long as wide and its anterior border not closely attached to the posterior border of eye; postocular crest poorly developed; tarsal fold rudimentary ................. hcilius tacand

4 a Males uniform orange in life (yellow in preservative), females olive-green to black with red spots .............. hcilius peziglenes

b Ground color grayish brown to black with parotoid glands and some warts reddish

........................................................ brown 5 5 a Cranial crest low, obsolescent; parotoid

gland less than 2/3 of area of upper eyelid; venter pale with some dark mottling on chest ............................... Incilius holdridgai

b Cranial crest massive, elevated; parotoid gland 2/3 or more than of area of upper eyelid; venter pale, uniform or heavily mottled with black ................................................. 6

6 a Dorsum without enlarged warts; venter almost uniform pale tan; posterior thigh dark black, contrasting to dorsum; shank with moderate rugose warts ....................... ..................................... ..hcilius penpenpetetes

b Dorsum with regular, longitudinal series of enlarged warts; venter pale, heavily mottled with black; posterior thigh same color as dorsum; shank with huge pointed warts

Incilius fastidiosus ...................................... 7 a Finger I shorter than Finger 11; upper sur-

faces covered with widely spaced dark- tipped, keratinized, pointed warts and spicules; toes with considerable webbing (Fig. 249) ................................................... 8

b Finger I same length or longer than Finger 11; warts and spicules on upper surfaces, if present, not keratinized, pointed; toes with basal webbing (Fig. 250) .......................... 9

8 a Fingers slender and normally developed with little or no webbing between Fingers 1-111 ................................ hcilius conifivlLe

b Fingers short and plumb with extensive .................. webbing between Fingers 1-111

..................................... Inciliu8 ktiledp~at?

Fig. 244. Incilius holdridgei (Cerro Chompipe, Heredia, Costa Rica, 2090 m). Photo: G. Kohler

Fig. 245. Incilius bocourti (San CrisMbal de las Casas, Chiapas, Mexico, 2290 m).

Photo: A. Ramirez Velazquez

Fig. 246. Incilius conifems (Bartola, Rio San Juan, Nicaragua, 30 m). Photo: G. Kijhler

Fig. 247. Incilius melanochlorus (Soltis Center, Alajuela, Costa Rica, 460 m). Photo: G. Kohler

Fig. 248. Dorsal view of head in species of Incilius: a. I. canaliferus; b. I. coccifer; c. I. luetkem ' d. I. valliceps; note variation in shape and size of parotoid glands. Drawings: M. Vesely

Fig. 249. Foot of Lncilius conifem. Drawing: M. Vesely

9 a Parotoid gland twice the size of area of upper eyelid, twice as long as wide (Fig. 248a); parietal crests absent ...................... ..................................... Inclllue canalif$rus

b Parotoid gland signXcantly less than twice the size of area of upper eyelid, much less than twice as long as wide (Figs. 248b-d);

.......... parietal crests present or absent 10 10a ~arotoid gland significantly smaller than

the size of area of upper eyelid (Fig. 248~); a lateral row of tubercles present ......... 11

b Parotoid gland same size or larger than the size of area of upper eyelid (Figs. 248b,e) or, if slightly smaller then without a lateral row of tubercles ...................... 13

l la Parotoid gland round to oval, area 112 or less area of upper eyelid (Fig. 248c); suborbital, and labial crests well-developed ... ........................................ InciZiu~ luetkenii

b Parotoid gland triangular, very narrow and elongate, area about 314 of area of upper eyelid; suborbital, and labial crests weak or absent ....................................... 12

12a No transverse folds between parietal crests; pretympanic and preorbital crests

Fig. 250. Foot of Lncilius aucoinae. Drawing: M. Vesely

absent or poorly developed; throat and ........................ chest pale In&w aucainae

b Transverse folds between parietal crests present; pretympanic and preorbital crests well-developed; throat and chest black (Fig.

........................ 247) hcik'ue melanochlaru8 13a Parietal crest absent . In&~8 marmomue

b Parietal crest present, usually prominent . .................................................................. 14

14a Subarticular tubercles on fingers and toes bifid or double; suborbital and labial crests usually well-developed; usually a pale

....................................... interorbital bar 15 b Subarticular tubercles on fingers and toes

single; suborbital and labial crests usually absent or weakly developed; interorbital

................................ bar, if present, dark 18 1Sa Supratympanic crests large and bulbous

............................................... (Fig. 251a) 16 b Supratympanic crests relatively thin (Fig.

251b) ........................................................ 17 16a Larger toad, SVL in adults 43-82 mm,

mean 62.2 mm (males), 61-94 mm, mean 78.4 mm (females); occurs in central and

rig. 251. Laterm surrace 01 neaa in a. Inciliu porten and b. I. coccifer, note different condil ons of supratympanic crest (arrow).

Photos: G. Kijhler

bital crests higher than postorbital crest at meeting point of these three crests (this is more pronounced in females than in males); parietal crest higher than supraorbital crest, arched and usually gradually

............... sloping downward posteriorly 20 19a Hind foot short (ratio foot length / SVL

0.35-0.42 in males, 0.35-0.41 in females); males with dark throats resulting from underlying darkly pimented interhyoideus

............................. muscle hcilius valliceps b Hind foot longer (ratio foot length / SVL

usually 0.45-0.49 in males, 0.42-0.43 in females; males lacking darkly pimented

.... interhyoideus muscle hcilius ~ ~ m p b e l I i 20a Tympanum smaller (ratio tympanum

length / SVL usually ~0.38); hind leg shorter (ratio hind leg length / SVL about 0.41 in males, 0.38 in females) hcilius tutdan'u~

b Tympanum larger (ratio tympanum length / SVL usually >0.38); hind leg longer (ratio hind leg length / SVL 0.42-0.49 in males,

............................. I 0.42-0.46 in females) 21 21a Males and females with elevated, rather

thin parietal and supraorbital crests; hind leg longer (ratio hind leg length / SVL

... 0.44-0.49 in males, 0.42-0.46 in females) ....................................... hcilius leucomyoe

b Males with low, relatively thick parietal and supraorbital crests (in most females these crests are elevated, rather thin); hind leg shorter (ratio hind leg length 1 SVL

............. 0.42-0.43) hcilim macromktatus

southern Guatemala as well as southwestern Honduras .................... Incilius ibarrai

b Smaller toad, SVL in adults 40-59 mm, mean 49.6 mm (males), 56-73 mm, mean 64.5 mm (females); occurs in south-central Honduras ........................... hdus pod

17a Venter with no, or few, relatively indistinct dark brown markings; cranial crests rela-

........ tively thick and high hcilius d m b Venter boldly marked with a distinct mar-

bled pattern; cranial crests relatively thin and low ............................. h&us Bignifer

18a Tympanum smaller (ratio tympanum length / SVL usually <0.50); parietal and supraorbital crests usually not higher than postorbital crest at meeting point of these three crests, if parietal and supraorbital crests higher than postorbital crest at confluence, then parietal crest not higher than supraorbital irest and usually concave just ~osterior to confluence with supraorbital crest ......................................................... 19

b Tympanum larger (ratio tympanum length / SVL usually >0.50); parietal and supraor-

Further Reading PORTER 1963, SAVAGE 1966, 1972b, NOVAK & ROBINSON 1975, MCDL~RM~I & FOSTER 1981, DUNDEE & ROSSMAN 1987, LEE & SALZBURG 1989, JACOBSON & VANDENBERG 1991, ROSSMAN 1991, FREED 1992, SAVAGE & DONNELLY 1992, BLAIR et al. 1995, LIPS & KREMPELS 1995, FLEET & AUTREY 1997, HAAS & KOHLER 1997, MENDELSON 1997, 1998, A'ITUM & EASON 1999, FRANKLIN & FRANKLIN 1999, MENDELSON et al. 1999, KOHLER et al. 2000, McC- & KOHLER 2000, McCRANIE & WILSON 2000, MULCAHY & MENDELSON 2000, DAYTON et al. 2001, MCGEHEE et al. 2001, MENDELSON 2001, JOHNSON 2002, KOHLER 2003, O'NEILL & MENDELSON 2004, MENDELSON et al. 2005, JARAMILLO et al. 2009, L~PEZ et d. 2009, MENDELSON & MULCAHY 2010

The generic name Rhaebo was revived by -T et al. (2006a) to accommodate nine species of toads one of which, R. haematiticus, occurs in Central America. The species d this genus are distributed from eastern Honduras to Panama and also widely in tropical South America on both sides of the Andes. Species of Rhaebo typically have a relatively smooth dorsal skin lacking obvi- oas "warts", large parotoid glands, and no cranial crests. The large parotoid glands in I& haematiticus are inconspicuous because &ey are not very elevated and their borders blend in with the surrounding skin. Bhaebo haematiticus is a nocturnal inhab- Mt of the forest floor where it can be h m e d foraging for small ants and other e o p o d s in the leaf litter (LIEBERMAN 9.986). During the breeding season (March

July), Rhaebo haematiticus typically nrngregates in rocky pools along the bor- ,& of forest streams and rivers (SCOTT '*a). It is an explosive breeder that lays $ggs in paired strings. After metamorpho- 'e the froglets are about 19 mm SVL and U y identifiable by having a black venter wrinkled with white spots in life; in gmxsemative, the pale spots lose their i&&ensity (SAVAGE 2002).

Fig. 253. Rhaebo haematiticus (P.N. Saslaj Atlantic0 Norte, Nicaragua, 720 m).

Photo: G. Kijhler

Rhaebo haematiticus (COPE 1862b), Proc. Acad. Nat. Sci. Philadelphia 14: 157; type locality: "Region of the Truando, New Granada [= Chocb, Colombia] ." Males to 62 mm, females to 80 mm. From eastern Honduras (Caribbean slopes) and from southern Costa Rica (Pacific versant) south to northern Colombia and northwestern Ecuador (and isolated populations on the eastern slopes of the Cordillera Oriental of Colombia extending into adjacent Venezuela), sea level to 1300 m elevation.

Rhaebo haematiticus Y 3 a

Further Reading CRUZ DfAZ & WUSON 1983, SCOTT 1983a, BARRIO

er ~ M O R ~ S 2001

115

The taxonomic history of Rhinella is deeply intertwined with that of Bufo (sensu lato), with repeated name changes and much controversy among recent authors as to content and hierachical level of this taxonomic group (e.g., CHAPARRO et al. 2007, SAVAGE & BOLAROS 2009b, PAULY et al. 2009, VAN BOCXLAER et al. 2010). Clearly more research is needed in order to clarify the phylogenetic relationships of the heterogeneous assemblage of species currently grouped under the generic names Chaunus and Rhinella. See also comments in the Chaunus account. I follow here CHAPARRO et al. (2007) in placing the former species of Rhamphophryne in Rhinella. GRAYBEAL & CANNATELLA (1995) had already noted that there is no unarn- biguous evidence in support of the monophyly of former Rhamphophzyne.

-ella acrolopha (TRUEB 1971), Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 216: 18; type locality: "Cerro Mali, Darien Province, P d , elevation 1410 la" Males to 38 mm, females to 45 mm. Cordilleras of southeastern Dari6n Province, Panama, to northwestern Colombia, near sea level to 1480 m elevation.

Rhinella data (THQMINOT 1884), Bull. Soc. Philomath., Paris, Ser. 7, 8: 151; type locality: Wsthme de Panama ... Obispo." Males to 43 mm, females to 55 mm. Possibly western Panama to northern Colombia, but the geographic distribution of R. data and its morphological differentiation &om R. acutirostris are poorly understood (HOOGMOED 1989, BARRIO-AMOR~S 19998, ~oFuuL.4 & S~ARIS 1999, B A R R I O - ~ O B ~ S 2004).

Keytomella 1 a Finger I shorter than Finger 11; tympanum

absent or indistinct ..................................... ...................................... Rhinella auvlopha

b Finger I same length or longer than Finger 11; tympanum distinct ................................

Rhinella alata ..............................................

The small toads Rhinella acrolopha and R. - alata both can be found during daytime a amidst leaf litter on the forest floor; gravid females of R. acrolopha were obtained during the months of February, March, and fi July (TRUEB 1971, IBAE-JEZ et al. 1999b). Both species feed mainly on ants and other small arthropods (TOFT 1981). Rhinella pqF data is an explosive breeder that uses &i. k' ponds and areas in the forest Fig. 254. &inella acrolopba (Cerro Mali, reproduce, primarily near the end of the Da,.i&, Photo: W. E. Duellman r a i ~ season and in the early dry .wason ( C O A S ~ of the Biodiversity Institute, University of Kansas) (BAREz et al. 1999b).

r Rhinella acrolopha

Fig. 255. Rhinella da ta (Nusagandi, Cuna, Panama, 350 m). Photo: G. Klihler

Centrolenidae

Centrolenidae Glassfrogs are Neotropical endemics and emstitute important components of the rain and cloud forest assemblages. These approximately 150 species of predominantly green (in life), arboreal frogs are distri- bnted from southern Mexico across Central America and tropical South America (to Peru on the Pacific versant and to south- central Bolivia, southeastern Paraguay, and northeastern Argentina on the Atlantic versant, excepting the lower Amazon basin) (FROST 2010). Nine genera nre currently recognized in the family Centrolenidae (CISNEROS-HEREDIA & I~CDIARMID 2007, GUAYASAMIN et al. 2009, PBosr 20101, five of which have representatives in Central America. Most cen- trolenids are small (<30 mm SVL), delicate frags with large eyes that are directed forward at about a 45 degree angle (hylids usually have the eyes oriented laterally). The common name "g1assf?ogs" is derived h m their partially or completely transparent venters that allow an inspection of tfie viscera in living individuals of these frogs. Most species have well-developed discs and webbing on both fingers and toes (SAVAGE 2002).

&I glassfrogs are nocturnal and can be &md in vegetation along small rivers and b s . During the rainy season, the &&-pitched call of the males can be &ad. .The females of most species deposit #bir eggs on leaves overhanging fast-mov-

&reams, on moss on branches, or on above streams (MCDIARMID 1975, ARRANZA & LYNCH 1991), although

ella granulosa is reported to breed nds occasionally (KUBICKI 2007). ding on the species, the eggs can be on the upper leaf surface, at the tip,

neath the leaf (GUAYASAMIN et al. . During development and with the

of water, the gelatinous egg ell and increase their volume. In

species of glassfrogs, varying degrees care are present, such as in the Hyalinobatrachium. It is the

Ng. 256. SacAatamla llex (Soltls tienter, San Isidro de Pefias Blancas, Alajuela, Costa Rica, 360 m). Photo: G. Kohler

male that attends and guards the clutch, thereby increasing the rate of survivorship of the eggs and unhatched larvae. Predators on glassfrog eggs include wasps and fly larvae (VILLA 1977, MCDIAR~WD 1983). Adult males of H. valerioi attend their egg clutches during night and day, being conspiciously similar in appearance to their egg clutches (Fig. 273): the frog's reticulate green pattern on a yellowish background is thought to be an evolutionary response to visually hunting diurnal predators (MCDIARMID 1983). This resemblance between frogs and eggs is not present in species that desert their clutch during the day, such as H. colymbiphyllum, which is uniform green with small yellow dots. After night falls, the males of this species return to their egg sites.

Depending on local weather conditions, the larvae are ready to hatch after 2-3 weeks. Often clutches develop to a certain point and then hatch during substantial rains when they are washed or fall into the stream. One benefit of hatching during rain is that tadpoles dropping into water are less prone to predation by fish due to

the then turbid water (MCDIARMILI 1983). The tadpoles hide in the decaying leaf litter and detritus that accumulates in the slower parts of the stream. The predominately reddish color of most glassfrog tadpoles is caused by extremely high concentrations of hemoglobin and an extensive system of surface capillaries enhancing cutaneous respiration in this low-oxygen I environment (SAVAGE 2002). The tadpoles F I ~ . ao ,. nana or a. H. chmpoi and b. H. take several months to a year before meta- colymbiphylllum. Photos: B. Kubicki morphosis (KUBICKI 2007).

Key to the glasshge of Central America 1 a Bones white in life; dorsal coloration in

preservative white or cream; dentigerous process of the vomer and vomerine teeth absent; completely transparent ventral parietal peritoneum (Fig. 259a,b) ............. 2

b Bones pale to dark green in life; dorsum

preservative; dentigerous process of the

1 creamy lavender to dark lavender in a.

and transparent posteriorly or completely

24 vomer present, with or without teeth; ven- Fig. 258. Profile of head in a. Sachatamia albo- tral parietal peritoneum white anteriorly maculata and b. Cochranella euknemos.

Drawings: G. Kohler .......................... transparent (Fig. 259cf) 8 2 a Two to six conspicuous, clearly demarcated

large, bright yellow dots present on the usually white (members of some popula- H*Obawum tions lack a completely white-pigmented

b No conspicuous, clearly demarcated large, pericardium) . Hjdhobatrachium valerid bright yellow dots present, although numerous small yellow flecks can be pre- b Do'sum with yellow spots but not Bur- sent ............................................................ 3 rounded by a reticulum of green pigment

and dark punctations in life (whitish or yel- 3 a Substantial webbing between fingers I1 lowish, often with a minute brown or and I11 (Fig. 257a) ....................................... purpelish punctations in preservative); ...................... Hyalinobatmchium chimi.. pericardium white or colorless ................ 7

b Only vestigial webbing between fingers I1 7 a Tympanum absent or very indistinct; peri- and 111 (Fig. 257b) ..................................... 4 cardium white or colorless; nostrils in very

4 a Dorsum with a dark green vertebral stripe; slightly indicated fleshy swellings ............. ............................... pericardium colorless 5 ............... qyalinobatrachium fleischmanni

b Dorsum green with small yellow spots or b Tympanum distinct; pericardium colorless; reticulated with green and yellow in life, no nostrils in strongly protuberant fleshy dark green vertebral stripe; pericardium swellings ......................................................

.......... white or colorless ...................................... 6 Hplinobatrachium c o l ~ p 4 v I I u m 5 a Dark green vertebral stripe bordered on 8 a Prepollex with a free, spiny distal end; dor-

either side by yellow stripes in life (the sum without distinct spots; SVL to 23 mm ...................................... dark stripe indicated by purplish or brown- Teratohyla @OM

ish pigment in preservative) ....................... b Prepollex without a free, spiny distal end; ............. Hyalinobatrachium vhmittatum dorsum with or without distinct spots; SVL b Dark green vertebral stripe not bordered variable ..................................................... 9

on either side by yellow stripes in life ....... m o b a t r a c h i u m talamancae 9 a Skin texture strongly granular; dorsum ................. uniform green or bluish-green without yel-

6 a Dorsum with yellow blotches surrounded low spots (although evenly marked with by a reticulum of green pigment and dark tiny bluish to whitish flecks) but usually punctations in life (a reticulum of black with scattered purple or black spots; a punctations in preservative); pericardium white parietal peritoneal sheath present;

$259. Ventral view of glassfrog species: a. H. vireovittatum; b. H. valeriot wmaculata; e. T. pulverata; f. C. euknemos.

SVL to 29 mm in males, to 32 mm in ; females ................. Cochranella grandoea a b Skin texture not conspicuously granular "though scattered tubercles can be pre-

sent (in C. euknemus); dorsal coloration .......................... variable; SVL variable 10

a Dorsum uniform green in life or with numerous small dark spots, no yellow spots; a humeral spine present in adult males although this spine can be protruding (E. prosoblepon) or be hidden in

............................. musculature (S. ilex) 11 b Dorsum green with yellow or white spots

in life; no humeral spine in adult males 12 lla A protruding, externally visible, humeral

hook present in adult males; dorsum uniform green in life or with numerous small dark spots; webbing between toes 111-IV-V not reaching base of disc on toes 111 and V, not reaching distal subarticular tubercle on toe TV; nostrils not elevated on protuberant ridges; snout subovoid in dor-

..................................................... sal aspect ............................ Espadarana pmsoblepon

b A humera1 spine present in adult males although this spine is mostly hidden in musculature and therefore, not visible externally; dorsum uniform green in life, never with numerous small dark spots; webbing between toes 111-IV-V reaching

. granulosa; d. S. Photos: B. Kubicki

almost to base of disc on toes 111 and V, reaching distal subarticular tubercle on toe IV; nostrils elevated on protuberant ridges; ........... snout semicircular in dorsal aspect ........................................... Sachatamia ilex

1% Snout vertical to rounded in profile (Fig. 258a); no white pigment on digestive tract ......................... Sachatamia albomaculata

b Snout conspicuously sloping in profile (Fig. 258b); white pigment on digestive tract 13

13a Dorsum limegreen with numerous fine white or yellowish spots; pale green bones; a translucent parietal peritoneum (Fig. 259e) ......................... Teratohyla pdvetata

b Dorsum deep bluish-green with numerous clearly demarcated yellow spots; dark green bones; a white parietal peritoneum (Fig. 2590 .............. CochranelIa ezhemoe

F'urther Reading TAYLOR 1949b, 1958a, SAVAGE & STARRETT 1967, STARRETT & SAVAGE 1973, M c D m & ADLER 1974, Villa 1974, DUELLMAN 1977, VILLA 1977, HAYES & STARREIT 1980, VILLA 1984, Ruu- CARRANZA & LYNCH 1991, MCCRANIE & WILSON 1997c, BARREU-RODRIGUEZ 2000, SAVAGE 2002, CISNEROS-HEREDIA & MCDIARMID 2007, KUBICKI 2007, GUAYA~AMIN et al. 2008, 2009, HOFFMANN 2010

Cochranelli

CochraneLla contains seven species, only two of which are distributed in Central America (FROST 2010). In Central America, these frogs inhabit vegetation along streams in humid forests at elevations below 1750 m above sea level. At night, males call from the upper surfaces of leaves, often several meters above ground level. Females deposit eggs on the upper

I sides of leaves overhanging the water, usu- 1 ally near the tip of the leaf. In both Central American species, the eggs are darkly pig- Fig- 260. C ~ c h m e f l a mdcnemos (Nusagan& rnented and the jelly masses can contain Kuna Yda, Panama, 350 m). Photo: G. Kohler

40-70 eggs (KUBICKI 2007).

Codranella euknemos (SAVAGE & STARIWTT 19671, Copeia 1967: 604; type locality: "Costa Rica: Provincia de San Jos6: Cant611 de Coronado: 1.5 km S Alto La Palma, 1500 m." Males to 25 mm, females to 32 mm. Central Costa Rica to western Colombia, 90-1650 m elevation.

CochraneLZa granulosa (TAYLOR 19491, Univ. Kansas Sci. Bull. 33: 265; type locality: "Los Diamantes one mile south of Guapiles, [Cant& de Pococi, Provincia Lim6n,l Costa Rica." Males to 29 mm, females to 32 mm. Atlantic versant from eastern Honduras to central Panama and on the Pacific versant from central Costa Rica to western Panama, 30-1500 m elevation. Fig. 261. Cochranella granulosa (3 km SE

Ayapal, Nicaragua, 200 m). Photo: G. Hiohler

Zochranella euknernos

Further Reading MCCRANIE et al. 1999, SAVAGE 2002, GUAYASAMIN & BONACCORSO 2004, KUBICIU 2007, H o ~ 2010 I

recently proposed genus (GUAWAMIN 9) includes three species, only one Espadarana prosoblepon, occurs

al America. This is a nocturnal that is usually observed on the veg- along fast-moving streams. Males soblepon call from the upper sur- leaves throughout the year with Fig. 262. Espadarana prosoblepon (male, P.N.

ed vocal activity during the rainy Saslaya, Nicaragua, 920 m). Photo: G. Kohler

Fig. 263. Espadarana prosoblepon (female, P.N. Saslaya, Nicaragua, 920 m). Photo: G. Kohler

alinobatrachium I

a prosoblepon (BOETTGER 1892), b Samml. Mus. Senckenb. Natur-

C

The genus Hyalinobatrachium comprises 27 species, seven of which are found in Central America (FROST 2010). Morphological characteristics of the species in this genus include white bones and the absence of vomerine teeth (SAVAGE 2002). Males of this genus usually call at night from the underside of leaves in vege-

> tation overhanging streams. During day-

! time, these small frogs hide well camou- s flaged within streamside vegetation. The b females deposit their eggs, which are green

when laid, on the underside of leaves Espadamna prosoblepon (RUIZ-CARRANZA & LYNCH 1998, SAVAGE

2002). Males in this genus typically guard the eggs (KUBICKI 2007). Hyalinoba-

Further Reading trachium fleischmanni is the glassfrog JACOBSEN 1985, CRUZ-DIAZ & MCCRANIE 1999, with the widest geographical and eleva- SAVAGE 2002, KUBICRI 2007, HOFFMANN 2010 tional range of any centrolenid species.

I

.Hyalinobatrachium aureoguttatum (BARRE& : XODRIGUEZ & RUIZ-CARRANZA 1989), Trianea, 3: .77; type locality: "Colombia, Departamento del "Chocb, vertiente occidental de la cordillera "Occidental, Municipio El Carmen de Atrato, ,,Kin. 23 carretera El Carmen - Quibdo, 5'47' Jatitud N y 76"20W, 1030 m.s.n.m." Males to <23.3 mm, females to 23.9 mm. Eastern cordilleras af the Dari6n Province in Panama to northwestern Ecuador, 45-1570 m elevation.

.Hyalinobatrachium chirripoi (TAYLOR 1958a), Univ. Kansas Sci. Bull. 39: 59; type locality: "Cocales Creek, Suretka, [Cant611 de Talamanca,] L i m b Province C. R." Males to 26 mm, females to 30 mm. Southern Caribbean

-Costa Rica, and central and eastern Panama to western Colombia, and also in eastern Honduras, sea level to 700 m elevation. Hyalinobatrachium cardiacalyptum MCCRANIE & WILSON 1997 is regarded as a synonym of H, chidpoi according to C I S ~ O S - HEREDIA & MCDIARMILI (2007).

Hyalinobatrachium colymbiphyllum (TAYLOR 1949b), Univ. Kansas Sci. Bull. 33: 262; type locality: "American Cinchona Plantation, elev. 5,600 ft., Caribbean drainage of V d c h Po&, [Canun de Alajuela, Provincia de Alajuela,] Costa Rica." Males to 27 mm, females to 29 mm. Costa Rica to Colombia, and also in eastern Honduras, sea level to 1710 m elevation. Hyalinobatrachium crybetes MCCRANIE & WILSON 1997 is regarded as a synonym of H. colymbiphyllum according to CISNEROS- HEREDIA & MCDIARMID (2007).

Hyalinobatrachium fleischmanni (BOETTGER 18931, Ber. Senckenb. Naturforsch. Ges., 1893: 251; type locality: "San Jos6, Costa Rica." Males to 28 mm, females to 32 mm. Guerrero and Veracruz, Mexico, south through Central America to Colombia, sea level to 1750 m elevation.

Hyalinobatrachium talamancae (T. )R 1952b), Univ. h a s Sci. Bull. 35: 781, ~ ~ g e locality: "Moravia [de Chirrip6, Cant611 de Turrialba, Prov. Cartago, Costa Rica], Carib- bean drainage." Males to 22 mm, females to 27 mm. Caribbean versant of Costa Rica to eastern Panama, 400-1600 m elevation.

Hyalinobatrachium valerioi (DUNN 19311, Occas. Pap. Boston Soc. Nat. Hist. 5: 397; type locality: "La Palma, [Canthn Coronado, Prov. San JosB,] Costa Rica, 4,500 feet." Males to 24 mm, females to 26 mm. On the Atlantic versant from northern Costa Rica to northwestern Ecuador, sea level to 1500 m elevation.

Hyalinobatrachium vireovittatum (STARRETT & SAVAGE 19731, Bull. S. California Acad. Sci.

'472: 66; type locality: "Costa Rica: Provincia de :Sari Jose: Canun Perez Zeledon: 0.5 km NE @ombra: a place 16 km SW San Isidro de El ,\General on the road to Dominical, 880 m." ales to 23 mm, females to 25 mm. Definitely *own only Com the central Pacific foothills in #.Costa Rica and in the area of El Valle de Ant6n in west-central Panama (KUBICKI 2007).

1 i630-1100 m elevation. I .-

Hyalinoba frachium

J

A Y. aureogutatum t

I. fleischmanni

4 H. vireovittatum r H. talamancae

Hyalinobafrachium

f. colymbiphyllum - I. chirripoi

Centrolenidae

Fig. 265. Hyalinobatrachium aureoguttatunl (Ecuador). Photo: M. R. Bustamante 1 BiographicaO

I

Fig. 264. Hyalinobatrachium colymbiphyllum Fig. 266. Hyalinobatrachium aureoguttatum (near Parrita, Puntarenas, Costa Rica, 250 m). (Ecuador).

Photo: B. Kubicki Photo: M. R. Bustamante 1 BiographicaB

Fig. 267. Hyalinobatrachium chimipoi (Aguas Fig. 270. Hyalinobatrachium chirripoi (P.N. Zarcas, Limbn, Costa Rica). Photo: G. Kohler Patuca, Olancho, Honduras). Photo: G. Kiihler

Fig. 268. Hyalinobatrachium fleischmanni Fig. 271. Hyalinobatrachium fleischmanni (Salto lab^, Atlantic0 Norte, Nicaragua, (Piedra de Candela, Chiriqui, Panama, 480 m). Photo: G. Kohler 1470 m). Photo: G. Kohler

Fig. 269. Hyalinobatrachium talamancae (La Fig. 272. Hyalinobatrachium vireovittatum Nevera, Ngobe Bugle, Panama, 1600 m). (near Parrita, Puntarenas, Costa Rica, 250 m).

Photo: A. Hertz Photo: B. Kubicki

Fig. 273. Hyalinobatrachium valerioi (male with eggs, Guayacan, CRARC, Costa Rica, 550 m). Photo: G. Kijhle

Fig. 274. Hyalinobatrachium valerioi (male, Fig. 275. Hyalinobatrachium valerioi (female, Guayacin, CRARC, Costa Rica, 550 m). Costa Rica). Photo: G. Kiihler

Photo: G. Kohler

Further Reading GOIN 1964, GREER & WELLS 1980, MCDIARMID 1983, VILLA 1984, IBAREZ & JARAMILLO 1997, JARAMILLO et al. 1997, MCCRANIE & WILSON 1997, bAFmz et al. 1999a, BARRERA-RODRIGUEZ 2000, SERARIS 2001, SAVAGE 2002, KUBICRI 2004a, KUBICKI 2007, HOFFMANN 2010

125

Sachatamia & The generic name Sachatamia was proposed by GUAYASAMIN et d. (2009) for three species of glassfrogs, two of which occur in Central America. Characteristics of frogs in this genus include having green bones (in life), having the ventral parietal peritoneum white anteriorly and transparent posteriorly, and changing of the dorsal coloration from green in life to lavender in preservative. A humeral spine (mostly hidden in musculature) is present in S. ilex I but is absent in S. albomaculata. Fig. 276. Sachatamia albomaculata (Guayach,

CRARC, Costa Rica, 550 m). Photo: G. Kiihler As with most glassfrogs, the species of Sachatamia are typically found at night in the vegetation along fast-moving streams in humid forests. Males usually call eom the upper surface of leaves, up to seven meters above the ground (KUBICKI 2007). The eggs of S. albomaculata are brownish gray with a violet tint and are normally laid on the upper surfaces of leaves overhanging streams (KUBICKI 2007, HOFF- MANN 2010). Occasionally, clutches are discovered on rocky surfaces alongside water- falls (PUSCHENDORF et al. 2004). Clutch size is 16-25 eggs in S. ilex (KUBICKI 2007).

Sachatamz'a albomaculata TAYLOR 1949b), Univ. Kansas Sci. Ed. 33: 267; type locality: 'Zos Diamantea, one mile south of Gu&piles, ECant6n de Poaoci, Provincia Limbn,] Costa Rica." Males to 29 mm, females to 32 2. North-cenkral Honduras to western Colombia, sea level to 1500 m elevation. Recently, this species has been recorded from NW Ecuador (GUAYASAMIN et al. 2006).

Sachatamia ilex (SAVAGE 1964), Copeia 1967: 326; type localits 'Costa Rica: F'rovincia de Lim6n: Cant6n de Limb: Alta Talamanca: 16 km SW Amubri, on Rio Lari, 300 m." Males to 29 mm, females to 34 mm. Eastern Nicaragua to western Panama and western Colombia (Choc6 and Valle del Cauca departments), in Costa Rica also on the Pacific versant, 250-1420 m elevation.

Fig. 277. Sachatamia ilex (Soltis Center, Alajuela, Costa Rica, 460 m). Photo: G. Kiihler

Further Reading SAVAGE 2002, HOFFMANN 2004, F'USCHENDORF et al. 2004, KUBICKI 2007, HOFFMANN 2010

Centrolenidae

Teratohyla - Frogs of the genus Teratohyla occur in the lowlands and premontane regions of Central America and in the Pacific and Amazonian lowlands of South America. Four species are recognized in this genus, two of which are found in Central America. Characteristics of frogs in this genus

, include having pale to dark green bones (in life), the absence of humeral spines, having the ventral parietal peritoneum white anteriorly and transparent posteriorly (T. spinosa) or completely transparent (T. pulverata), and changing of the dorsal coloration from green in life to creamy lavender to dark lavender in preservative. A protruding prepollical spine is present in T. spinosa but absent in T. pulverata.

: The two Central American species of this genus are found along streams in forests,

1 where males call from the upper surfaces of leaves at night. In T. spinosa, the egg

1 masses (clutch size 15-25 eggs) are usual- [ ly attached to the undersides of leaves, / along their margins, whereas in T. pulver- I ata the eggs (clutch size 30-75 eggs) are i laid on the upper surface and toward the i 1 tip of leaves (KUBICKI 2007). During early t stages of development, the eggs of T. spin-

ma are greenish white, whereas those of T. pulverata are pale greenish blue (KUBICKI 2007).

Teratohyla pulverata (PETERS 18731, Monatsber. Preuss. Akad. Wiss. Berlin 1873: 614; type locality: "Chiriqui", Panama. Males to 29 mm, females to 33 mm. On the Atlantic versant from northwestern Honduras, and on the Pacific slope from southern Costa Rica to northwestern Ecuador, sea level to 960 m elevation.

Teratohyla spinosa TAYLOR 1949b, Univ. Kansas Sci. Bull. 33: 259; type locality: "Los Diamantes, one mile south of GuBpiles, [Cant611 de Pococi, Pravincia Lim6n,] Costa Rica." Males to 20 mm, females to 23 mm. Eastern Honduras to western Colombia south to Rio Palenque in Ecuador, near sea level to 800 m elevation.

Fig. 278. Teratohyla spinosa (Guayach, CRARC, Costa Rica, 550 m). Photo: G. Kohler

Fig. 279. Teratohyla pulverata (Cerro El Toro, Atlantic0 Norte, Nicaragua, 480 m).

Photo: G. Kohler

I A T. spinosa T. pulverata

F'urther Reading KUBICKI 2007, SAVAGE 2002, HOFFMANN 2010

127

Dendrobatidae Recently, there has been some controversy regarding the higher taxonomy of dendrobatid frogs (GRANT et al. 2006, SANTOS et al. 2009). The monophyly of Dendrobatidae (sensu NOBLE 1926) has always been strongly supported by previous studies (VENCES et al. 2000, SANTOS et al. 2003). The taxonomic split of Dendrobatidae into two families (i.e., Allobatidae and Dendro- batidae), as proposed by GRANT et al. (2006), was rejected as unnecessary by SANTOS et al. (2009), who returned to the more traditional taxonomy. Also, SANTOS et al. (2009) found Dendrobates sensu lato (including Adelphobates, Dendrobates, Excidobates, Minyobates, Oophaga, and Ranitomeya) to be a well-supported monophyletic group and, therefore, synonymized Adelphoba tes, Excidobates, Minyoba tes, Oophaga, and Ranitomeya with Dendro- bates. Dendrobatidae, as defined by SANTOS et al. (2009), contains 274 species that are mostly distributed in South America. In Central America, this family is represented by 19 species in seven genera. Most dendrobatids are small diurnal, terrestrial, semiarboreal, or semiaquatic species, with the remarkable exception of the nocturnal Ammobates nocturnus MYERS et al. (1991) that reaches a SVL of 62 mm. Species of the genera Dendrobates and Phyllobates typically have aposematic colorations and highly toxic skin secretions; the alkaloids in the skin of dendrobatids are derived from their diet of ants and melyrid beetles (DALY 1995, DALY et al. 1980,1992,1994, DUMBACHER et al. 2004).

Key to genera of Dendrobatidae 1 a Discs of outer fingers (Fingers III and TV)

and toes (Toes IV and V) usually at least two times width of digits; no maxillary

....................................... teeth Dendrobah b Discs of outer fhgers (Fingers 111 and IV)

and toes (Toes IV and V) only slightly wider ....... than digits; maxillary teeth present 2

2 a Toes with extensive webbing; dorsal surfaces chocolate brown with a yellow flash mark atop base of arm in life; SVL of single

......... known specimen 19.5 mm Z & l d ~ 8 b Toes with basal or no webbing; dorsal color

pattern variable; SVL of adults variable 3 3 a Finger I as long as Finger 11; dorsum with

pale spots, no stripes; SVL of single known ..................... specimen 18.9 mm Am-

b Finger I longer than Finger 11; dorsum without pale spots, but usually dorsolateral or lateral stripes present; SVL of adults variable ..................................................... 4

4 a Ventral surfaces of body and thighs black with pale mottling .................. Phyllobates

b Ventral surfaces of body and thighs white, although throat and chest can be black in

............................................... adult males 5 5 a No oblique lateral pale stripe anterior to.

groin, although a dorsolateral pale stripe extends from eye to sacrum, and a ventrolateral pale line from below tympanum to groin in some individuals; no toe webs; SVL of adult males 17-24 mm, of adult females 16-25 mm .................................... AUobates

b Lateral dark band interrupted by an oblique pale stripe, either completely from upper posterior corner of eye to groin or from level of midbody to groin; degree of toe webbing variable; SVL of adults variable 6

6 a Finger I11 swollen in adult males (Fig. 280); lateral pale line, strongly oblique, usually extending forward to eye, sometimes incomplete and interrupted near level of

........... midbody; no toe webs sihrmbneia b Finger I11 not swollen in adult males;

lateral pale line, almost parallel to dorsolateral pale line, extending only a short distance forward from groin, not reaching level of axilla; degree of toe webbing variable .................................... Col&th~8

Further Reading SAVAGE 1968a, MYERS 1987, Z- & ZIMMEFMANN 1988, DALY et al. 1992, BAUER

Fig. 280. Hand of Silverstonia flotatoq note 1994, CLOUGH & SUMMERS 2000, GRANT et al. swollen Finger 111 (arrow). Photo: G. Kijhler 2006, SANTOS et al. 2009

Dentrobatidae

Allobates - - The predominantly South American genus Mobates contains 47 species, only one of which occurs in Central America (FROST 2010). Allobates talamancae is a diurnal denizen of the forest floor usually sitting on top of the leaf litter. Males call throughout the day but tend to be most active at periods of low light intensity (i.e., early morning, late afternoon, and during cloudy periods); the call consists of a series of slightly harsh "peet-peet-peets" (SAVAGE 2002). Males and females are territorial ( b m z et al. 1999b). Mating takes place in the moist leaflitter where the eggs are laid and attended by the male ( I B ~ z et al. 1999b). The males transport 8-29 tadpoles on their backs to water-filled depressions or shallow puddles on the edge of streams where the tadpole complete their development (IBAREz et al. 1999b, SAVAGE 2002).

Fig. 281. Allobates talamancae (near Miramar, Bocas del Toro, Panama, 20 m). Photo: G. Kohler

mobaces taIamancae ('OpE 1876)9 J' Fig. 282. Allobates talamancae (Lagarto Lodge, Nat. Sci. Philadelphia Ser. 2,s: 102; Type localitv: "near old H~~~~~~ [= puerto vieio1, on the Alajuela, Costa Rica, 90 m). Photo: G. Kohler

~*as t coast", [Cantbn -de Limbn, ~=ovincia L i d n , Costa Rica]. Males to 24 mm, females to 25 mm. Southeastern Nicaragua to the Pacific lowlands of Colombia and to northern Ecuador, sea level to 800 m elevation.

-- Ameeregi

The genus Ameerega contains 31 species distributed widely in tropical South America (FROST 2010). The single Panamanian representative of this genus, A. maculata, is known only from the type specimen found over a century ago (MYERS 1982). The holotype of A. maculata was redescribed by MYERS (1982).

Ameerega maculata (PETERS 18731, Monatsber. Preuss. Akad. Wiss. Berlin, 1873: 617; type locality: "Chiriqui", Panama; at the time of the description "Chiriqui" included both Atlantic and PacXc versants of extreme

- western Panama according to MYERS (1982). SVL of single known female 18.9 mm. Known only fkom the type specimen.

Further Reading M e r Reading SAVAGE 1968a, IBAREZ et al. 1999b, SAVAGE 2002 MYERS 1982

The genus Colostethus has its center of diversity in northern South America; of the 19 currently recognized species only three occur in Lower Central America (FROST 2010). Recently, GRANT (2004) resurrected C. panamensis from synonymy with C. inguinalis, and restricted the latter species to Colombia. The species of Colostethus are diurnal denizens of rainforest and cloud forest. Colostethus latinasus is a non- riparian frog usually found on the floor of wet montane forest, including cloud forest, away from streams, whereas C. panamensis is riparian, occurring only in the imme- diate adjacency of forest streams, often along rocky sections (IBAREZ et al. 199913, GRANT 2004). Males and females are territorial; the call of male C. panamensis consists of one to four whistled, moderately high-pitched notes in rapid succession (IBAREZ et al. 1999b). In C. panamensis, eggs are laid in the moist leaf litter, and after hatching, the female carries the tad-

- o s t e u s -he, Bocas del Toro, Panama, 10 m). Photo: G. Kiihler

Fig. 284. Colostethus panamensis (Nusagandi, Kuna Yala, Panama, 350 m). Photo: G. Kohler

poles on the back to a small water-flled depression or quiet backwater area where the tadpoles develop (IBAREZ et al. 1999b).

Colostethus latinasus (COPE 1863), Proc. Acad. Nat. Sci. Philadelphia 15: 48; type locality: "Truando region, New Granada" [= Depto. Chocb, Colombia]; neotype from "Panama, probably on Cerro Pirre above Santa Cruz de Cana, Dari6n Provincen (see GRANT 2004). Males to 24.5 mm, females to 34 mm. Known only from the Serrania de Pirre, Darien Province, Panama, sea level to 1440 m elevation. Colostethus panamensis ( D m 1933b), Occas. Pap. Boston Soc. Nat. Hist. 8: 69; type locality: "El Valle de Anton in the northwestern corner of the Province of Panama", Panama. Males to 27.1 mm, females to 29.2 mm. Panama and adjacent regions in Colombia, sea level to 900 m elevation.

. - Colostethus pratti (BOULENGER 1899), Ann. Mag. Nat. Hist. Ser. 7, 3: 274; type locality: "Santa Ines, N. of Medellin, Departamento Antioquia,] Republic of Colombia; altitude 3800 feet." Males to 24 mm, females to 25 mm. On the Atlantic versant from western Panama (including Isla Col6n) to Colombia (departments of Choc6 and Antioquia), 110-1160 m alevation.

mc -m - d g. 285. Foot of a. Colostethus panamensis d b. C. pratti. Photos: G. Kohler

Dentrobatidae

Dendrobates

&ey to Colost%t&us 1 a Toes with lateral fringes and distinctly

webbed at base (Fig. 285a); SVL of adult males 18.8-27.1 mm, of adult females 24.9-29.2 mm ..... Colostethue p a n a m d

b Toes without lateral fringes and webbing absent between all toes or barely detectable between toes I11 and TV (Fig. 285b); SVL of adults variable .................. 2

O a A dark stripe along posterior surface of thigh; throat immaculate white in females, mottled with gray to dark brown in adult 1 males; no toe webs; SVL of adult males to 24 mm, of adult females to 25 mm ............. ........................................ Colostetbus pratti

b No dark stripe along posterior surface of thigh; throat brown with white spots forming mottled pattern in females, darker in

#. adult males; webbing absent between all I toes or barely detectable between toes 111

and TV; SVL of adult males to 24.5 mm, of .............................. adult females to 34 mm 9- .................................. Colostethus latinasus

SANTOS et al. (2009) synonymized the nominal genera Adelphobates, Excidobates, Minyoba tes, Oophaga, and Ranitomeya with Dendrobates, the latter genus now containing about 50 species of tiny to moderately sized frogs. All species in this genus have more or less distinct aposematic colorations and produce highly toxic lipophilic alkaloid skin toxins (DALY et al. 1992). Studies on the geographic variation of skin toxins in D. pumilio have demonstrated a high variability in the alkaloid profiles among the various populations as well as among individual specimens (SAPORITO et al. 2006, 2007a, MEBS et al. 2008). The alkaloids in dendrobatid frogs are of dietary origin and, therefore, indicative for the availability of prey (mites, ants, and other arthropods) containing these chemical compounds (SAPORITO et al. 2004, 2007b). The Dendrobates species inhabit predominantly lowland rainforests; most are terrestrial or semiarboreal, but at least D. arboreus and D. vincentei are strictly arboreal, living in epiphyte-laden trees (MYERS et al. 1984, JUNGFER et al. 1996). Dendrobates pumilio and D. granuliferus exhibit a considerable diversity in color patterns (SUMMERS et al. 2003, BATISTA & KOHLER 2008). There are examples of distinct populations in close proximity without any obvious physiographic barriers that show no evidence of hybridization; these abrupt transitions have been interpreted as evidence of lack of or minimal gene flow (BATISTA & KOHLER 2008). DALY & MYERS (1967: 973) stated that for D. pumiLio "a static subspecies concept does not seem well suited to the dynamics of the situation". However, the descriptions of the phenotypes by these authors agree well with the observations by BATISTA & KOHLER (2008), indicating that over the course of at least four decades the phenotypic pattern had remained stable. The underlying mechanisms and barriers that prevent mixing of the phenotypes in the natural populations are poorly understood.

Dentrobatidae I Dendroba tes gran uliferus, previously thought to be a characteristic endemic of the Golfo Dulce region on the Pacific versant, was found in sympatry with D. pumilio on the Caribbean coast of southeastern Costa Rica, near the Panamanian border (MYERS et al. 1995). Males call throughout the day, but most intensive after rains and in the early afternoon; the call of D. auratus is a low, slurring buzzing sound lasting 2-4 seconds, that of D. arboreus, D. granuliferus, D. pumilio, and D. speciosus a harsh, insect-like "chirp- chirp-chirp" (MYERS et al. 1984, SAVAGE 2002). Eggs are deposited in the leaf litter; there is no amplexus but during fertilization of the eggs, male and female sit facing away from one another with their rear ends juxtaposed (SAVAGE 2002). In D. speciosus, D. granuliferus, and D. pumilio, tadpoles are transported singly on the back by females to small accumulations of water such as water-filled leaf or stem axils of bromeliads and other plants (SUMMERS et al. 1999). The female then returns to the pools periodically and lays unfertilized eggs that are eaten by the tadpole (WEYGOLDT 1980, JUNGFER 1985, BRUST 1993).

Dendrobates arboreus MYERS, DALY & MAR- 1984, American Mus. Novit. 2783: 5; Type Locality: "cloud forest a t 1120 m elevation on the continental divide above the upper Quebrada de Arena, a t longitude 82"12'31nW, on the border between the provinces of Chiriqui and Bocas del Toro, western Panama." Males to 21.6 mm, females to 22.2 mm. On the Atlantic versant in west- e m Panama, 20-1300 m elevation.

Dendrobates auratus ( G m 1855), Proc. Acad, Nat. Sci. Philadelphia 7: 226; type locality: "Island of Taboga, in the Bay of Panama." Males to 40 mm, females to 42 mm. Southeastern Nicaragua on the Atlantic versant and southern Costa Rica on the Pacific versant to northwestern Colombia, sea level to 800 m elevation.

Dendrobates claudiae JUNGFER, L ~ T ~ E R S & J ~ R G E N ~ 2000, Herpetofauna, Weinstadt, 22: 12; type locality: "Panama: Pronvincia Bocas

3. auratus

Dendrobates 4 1

I 7. granuliferus

I . minutus I. arboreus

Y -: Dendrohafes

pumilio fulguritus claudiae vicentei

Dentrobatidae

Bocas del Toro, Panama, 1090 m). arboreus (Camino Palo Fig. 880. Uendrobates claumae (mainland

opposite of Isla Loma Partida, Bocas del Toro, Photo: K.-H. Jungfer Panama, 20 m). Photo: K.-H. Jungfer

Rg. 287. Dendrobates auratus (Bartola, Rio Fig. 291. Dendrobates claudiae (mainland Sari Juan, Nicaragua, 30 m). Photo: G. K6hler opposite of Isla Loma Partida, Bocas del Toro,

Panama, 20 m).

Fig. 288. Dendrobates auracus wan Antonio ae r ~ g . g . 2 ~ . uendrobaces mgmitus (huuagandi, Damas, Puntarenas, Costa Rica, 60 m). Kuna Yala, Panama, 330 m).

Photo: G. K6hler Photo: K.-H. Jungfer

I

Fig. 289. Dendrobates auratus (Nusaganm, Fig. 293. Dendrobates minutus (Nusagandi, Kuna Yala, Panama, 350 m). Photo: G. Kohler Kuna Yala, Panama, 350 m). Photo: G. Kiihler

133

Dentrobatidae

del Toro: Festland westlich der Island Loma Partida (82°11W/09009'N)." Males to 13.4 mm, females to 13.2 mm. Known only from the vicinity of the type locality, sea level to 140 m elevation.

Dendrobates fuIgurjtus SILVERSTONE 1975, Sci. Bull. Nat. Hist. Mus. Los Angeles Co. 21: 28; type locality: "Playa de Oro, Departa- mento del Choc6, Colombia, 160 m." Males to 15 mm, females to 16.5 mm. Central and eastern cordilleras of Panama to the Pacific lowlands of Colombia as far south as the Fig. 294. Dendrobates grandifems (Rincbn de lower Rio Anchicaya, 160-800 m elevation. Osa, Puntarenas, Costa Rica, 10 m).

Dendrobates granuliferus TAYLOR 1958b, Photo: G. Kiohler Univ. Kansas Sci. Bull. 39: 10; type locality: "on low mountains north of the Rio Diquis, about 3 miles north of Palmar [Nortel, [Cantbn de Osa,] Puntarenas Province Costa Rica." SVL to 25 mm in both sexes. Pacific versant of southern Costa Rica and adjacent western Panama as well as on the Caribbean coast of southern Costa Rica, near the Panamanian border, sea level to 100 m elevation.

Dendrobates minutus S m t ~ v ~ 1935, Occas. Pap. Boston Soc. Nat. Hist. 8: 212; type locality: "Barro Colorado Island, Panama Canal Fig. 295. Dendrobates granuliferus (San Zone", Panama. Males to 15.0 mm, females Antonio de Damas, Puntarenas, Costa Rica, 15.5 mm. Central Panama (El Valle and 60 m). Photo: G. Kohler Cerro Campana) to Colombia (south to Rio Saija), near sea level to 1100 m elevation.

Dendrobates pumilio SCHMIDT 1857, Sitzungsber. Akad. Wiss. Wien Phys. Math. Naturwiss. K1. 24: 12; type locality: "Neugranada." SVL to 24 mm in both sexes. On the Atlantic versant from eastern central Nicaragua to western Panama (including islands in the Bocas del Toro archipelago), sea level to 940 m elevation.

Dendrobates speciosus SCHMIDT 1857, Sitzungsber. Akad. Wiss. Wien Phys. Math. Natunviss. K1. 24: 12; type locality: "Neugranada." SVL to 31 mm in both sexes. On both versants of the Cordillera de Talamanca in western Panama, 1100-1370 m elevation.

Dendrobates vicentei JUNGFER, WEYGOLDT & JURASKE 1996, Herpetofauna, Weinstadt, 18 (103): 18; type locality: "auf dem Ful3weg von El Cop6 nach Rio Blanco del Norte, Paljhohe der Kontinentalscheide etwa 1 km ijstlich des Cerro Blanco, etwa 8"40'N, 80°36W, 912 m NN, Provincia de Cocl6, PanamB." Males to 20.3 mm, females to 19.4 mm. Known only eom the type locality.

r lg. LJO. U ~ I I U ~ V U ~ L G @ ~ U U I I I I V \LUU C/ULLILLL L L ~ ,

Rio San Juan, Nicaragua, 30 m). Photo: G. Kijhler

Fig. 297. Dendrobates pumiLio (Rio Chimurria, Rio San Juan, Nicaragua, 30 m). Photo: G. Kiihler

Dentrobatidae

5 298. Uendrobates pumluo (lsla Dolane, cas del Toro, Panama, 10 m). Photo: G. Kohl€

rlg. au2. Denohbates purnilio (Isla Popa, Bocas del Toro, Panama, 10 m). Photo: G. Kiihler

;. 299. Dendrobates pumilio (Isla Bastimen- Fig. 303. Dendrobates speciosus (Fortuna, ;, Bocas del Toro, Panama). Photo: G. Kohler Chiriqui, Panama, 1100 m). Photo: K.-H. Jungfm I

Fig. 300. L,Jrobates pumilio (Isla Bastimen- Fig. 304. Dendrobates vicentei (Rio BelBn, tos, Bocas del Toro, Panama). Photo: G. Kohler Veraguas, Panama). Photo: K-H. Jungfer

Fig. 301. Dendrobates pumilio (Cerro Brujo, Fig. 305. Uendrobates vicentei (El Cop6, Coc16, Bocas del Toro, Panama). Photo: G. Kiihler Panama, 680 m). Photo: R-H. Jungfer

b Pale dorsal markings white, pale yellow, or golden in life; dorsal surfaces of legs brown to grayish brown without contrasting pale and black markings; pale ventral blotches pale blue or grayish white in life; SVL in adults of both sexes 13.1-13.4 mm ............ .................................. Dendrobates claudiae

7 a Dorsal and ventral surfaces of body and limbs brown or black with contrasting, mostly circular pale (yellow or yellowish green in life) spots; SVL of adults of both

I sexes 20-22 mm .... Dandrobates arbomus Fig. 306. Dendrobates pumilio (Guayachn, b Coloration and pattern of dorsal and ven- CRARC, Costa Rica, 550 m). Photo: G. Kohl1 tral surfaces of body and limbs not as

described above; SVL of adults variable . 8 8 a Dorsum extremely granular; hind limbs

usually bright jade green in life; SVL of .................. Key to Dendzvbatee adults of both sexes 19-22 mm

........................... 1 a SVL of adults 2542 mm ......................... 2 Dandrobates gran* b SVL of adults 13-24 mm ......................... 3 b Dorsum smooth; hind limbs coloration vari-

.................... 2 a Dorsum and venter bluish green, green, or able; SVL of adults variable 9 cream with large black or dark brown 9 a Throat coloration in males not different blotches, bands, or stripes ........................... from that in females; Coloration and pat-

Dandrobates auratus tern of dorsal and ventral surfaces of body ................................... b Dorsum red in life, with or without black and limbs variable; SVL of adults

flecks; venter uniform red 14-24 mm . Dandrobata pumilio (in part) ........................... ................................ Dandrobates s p e C j m b !mroat in males dark, in females pale (Fig.

3 a Dorsum with longitudinal stripes 4 307); dorsal and ventral surfaces of body ........... and limbs olive green, yellowish green, or b Dorsum patternless, or with speckles, bluish green in life, with or without brown

...................................... spots, or blotches 7 or black blotches or reticulations; SVL of 4 a SVL of adults >17 mm; dorsolateral pale adults 12.0-20.3 mm Dendrobates M'eentei

stripe, if present, not continuous from eye to sacrum .. Dandrobates purnilio (in part)

b SVL of adults c17 mm; a continuous dorsolateral pale stripe from eye to sacrum .... 5

5 a Venter with numerous small pale spots (pale blue or white in life) blotches; a distinct broad pale (golden or yellow in life) flash marking on dorsal surface of thigh, covering almost the complete dorsal surface of thigh; no longitudinal median pale stripe; SVL of adults of both sexes 12.0-15.5 mm ......... Dendrobates minutw

b Venter with a few large pale (green, yellowish green, yellow, or pale blue in life) blotches; pale markings on thigh, if present, not broad nor yellow in life; a complete or incomplete longitudinal median pale stripe on anterior portion of dorsum Fig. 307. Dendrobates vicentei: male (left) and present or not; SVL of adults variable .... 6 female (right). Photo: P. Weygoldt

6 a All pale dorsal markings of the same color (green or yellowish green) in life; dorsal surfaces of legs with contrasting pale Further Reading (green or yellowish green) and black mark- SAVAGE 1968a, SILVERSTONE 1975, DONNELLY ings; pale ventral blotches green, yellowish 1989, 1991, JUNGFER et al. 1996, GRAVES 1999, green, or yellow in life; SVL in adults of JUNGFER et al. 2000, SAVAGE 2002, SUMMERS et both sexes 13.5-16.5 mm ............................ al. 2003, LOTTERS et al. 2007, HAUSWALDT et aL ............................... hdrobates fidguritus 2010

Hyloxalus

MYEW (1991), referred several specimens i h m Colombia and Ecuador, and one individual from Panama to "Colostethus chocoensis" [= Hyloxalus chocoensis (BOULENGER 1912)l. MYERS & GRANT (2009) assigned populations from northwestern Ecuador to the new Anomaloglossus con- fusus and restricted the range of Hyloxalus chocoensis to its type locality and one locality in Antioquia, both in Colombia, thereby rendering the single Panamanian specimen (KU 172790 from El Llano-Cartf road, km 11.7, 150 m elevation, Province ' PanamA) formerly referred to as 'Colostethus chocoensis" without an identification. Thus, I have refrained from including a species account under this genus.

Pnrther Reading MYERS 1991, COLOMA 1995, MYERS & GRANT 2009

indeed very similar in their external morphology and coloration, and have been found to be sister species in the molecular phylogeny published by WIDMER et al. (2000). Furthermore, they readily hybridize under captive conditions (J. VARGAS pers. com.). These frogs are diurnal terrestrial denizens in the leaf litter of forested stream valleys where they move about predominantly by short hops inter- spersed with walking (SAVAGE 2002). Males typically call from within holes found between roots or rocks and are difficult to locate (KOHLER unpubl. obs.). The call of both Central American species consists of a low rasping trill lasting 2-6 seconds and repeated after a pause (SAVAGE 2002). Oviposition is preceeded by a complex

As currently understood, the genus PPlylobates contains five species, two of which are found in Lower Central America MYERS et al. 1978, 1995, GRANT et al. Fig. 308. Phyllobates lugubris (Rio Uyama, 2006). The batrachotoxins in the skin Bocas del Toro, Panama, 35 m). Photo: G. Kijhler secretions of these five species are among the most potent naturally occurring poi- ~ o n s (MYERS et al. 1978, DALY et al. 1980). Although the two Central American @es are not as dangerously toxic as is P. terribilis of Colombia, they should still be handled with care because their secretions ean be very irritating to eyes and mucous membranes, and can possibly cause serious systemic poisoning if the toxins enter the body through skin cuts (DALY et al. 1980). WAGE (1968a) considered P. vittatus as a synonym of P. lugubris, but SILVERSTONE (1976) recognized both as valid species. Phyllobates lugubris and P. vittatus are

P. lugubris P. vittatus

Phyllobates _i

Dentrobatidae - - -

courtship; there is no amplexus, instead the animals have been observed sitting with their rear ends touching (SILVERSTONE 1976, LEENDERS 2001). Eggs are deposited in the moist leaf litter and the clutches are not guarded, although males have been observed to moisten them when they are among dry leaves (SAVAGE 2002). After hatching, the tadpoles wriggle onto the back of the male, who carries one or several tadpoles to small puddles on the forest floor, water-filled depressions in logs, and backwater areas near streams (SILVER- STONE 1976, LEENDERS 2001). Metamor- phosis occurs about two months later, and the froglets are 12-13 mm SVL (SAVAGE 2002).

Phyllobates lugubris (SCHMIDT 1857), Sitzungsber. Ahad. Wiss. Wien Phys. Math. Naturwiss. Kl. 24: 12; type locality: "Neugranada"; restricted to "der Weg zwischen Bocca del Toro und dem Vulcan Chiriqui [Panama]. zwischen 5000' und 7000' Hohe" by SCHMIDT (1858). Males to 21 mm, females to 24 mm. Atlantic versant from extreme southeastern Nicaragua to central Panama (including islands in the Bocas del Toro archipelago), sea level to 600 m elevation.

Phyllobates vittatus (COPE 1893), Proc. Am. Philos. Soc. 31: 340; type locality: "Buenos Ayres*, [= Buenos Aires, Cant6n de Buenos Aires, Provincia de Puntarenas], Costa Rica. Males to 26 mm, females to 31 mm. Pacific versant of southern Costa Rica (from near Dominical to the Golfo Dulce region), 20-550 m elevation.

Fig. 309. Phyllobates vittatus (P.N. Corcovado, Puntarenas, Costa Rica). Photo: G. Kohler

1

Key to PhyIobates 1 a Dorsolateral stripe yellow in life, usually

narrow, bordering upper eyelid laterally; upper surfaces of limbs marbled with gold, yellow, or yellowish green in life; SVL of adult males 19-21 mm, of adult females 20-24 mm ................. Phyllobates lzzgrzblis

b Dorsolateral stripe orange-red in life, usually broad, nearly as wide as upper eyelid; upper surfaces of limbs marbled with bluish green in life; SVL of adult males 22-26 mm, of adult females 26-31 mm ..... .................................... Phyllobates vittatus

Further Reading SAVAGE 1968a, SILVERSTONE 1976, MYERS et al. 1978, DALY et al. 1980, DONNELLY et al. 1990a, WLDMER et al. 2000, RYAN 2002

Silverstoneia

GRANT et al. (2006) erected the genus Silverstoneia to accomodate three species formerly referred to as Colostethus. DUNN (193313) and SAVAGE (1968a) considered S. flotator as a synonym of S. nubicola, but IBAS~EZ & SMITH (1995) demonstrated that both are valid species that occur sympatri- cally a t several localities in Panama. These species are diurnal, terrestrial inhabitants of the leaf litter in relatively undisturbed rainforest (IBAREz & SMITH 1995). During the rainy season, individuals can be found a t considerable distance to the nearest streams, whereas in the dry season they tend to aggregate along the vicinity of streams (JAEGER et al. 1976). Males call throughout the day, usually from beneath leaf litter, with highest intensity after showers during the early morning and late afternoon (SAVAGE 2002). The call of S. flotator has been described as a relatively slow, weak "peet-peet-peet" lasting from 19 sec to more than ten minutes, that of S. nubicola as a rapid "peet- peet-peet", resembling a trill that last about two seconds (IBMEZ & SMITH 1995). The eggs are deposited in the leaf litter and after hatching, the male carries two to ten tadpoles on his back to streams where they complete development (SAVAGE 2002).

Dentrobatidae

Silverstoneia flotator @UNN 1931), Occas. Pap. Boston Soc. Nat. Hist. 5: 389; type locality: T3arro Colorado Island, Panama Canal Zone", Panama. Males to 16.5 mm, females to 18.0 mm. On the Atlantic versant from Costa Rica to east-central Panama, and on the Pacific versant from southern Costa Rica to west-central Panama, sea level to 865 m elevation.

Silverstoneia nubicola (DUNN 1924b), Occas. Pap. Mus. Zool. Univ. Michigan 151: 7; type localitv: "rain forest above Boquete on the trail to cGriqui Grande, 4,500 -feetp, Chiriqui, Panama. Males to 20 mm, females to 23 mm. Southern Pacific Costa Rica to western Colombia 200-1600 m elevation. Fig. 310. Silverstoneia flotator (Guayachn,

CRARC, Costa Rica, 550 m). Photo: G. Kijhler

Key to Silmtoneia 1 a Throat gray in adult males; venter white in

adult females in life; oblique lateral pale stripe complete or incomplete; usually no dark bars or spots on dorsal surface of shank; SVL of adult males 14.5-16.5 mm,

......................... of adult females 15-18 mm ................................... Silvmtaneia flotator

b Throat black in adult males; venter yellow in adult females in life; oblique lateral pale stripe complete; usually dark bars or spots on dorsal surface of shank, SVL of adult males 16-20 mm, of adult females 17.5-23 mm ......................... S i l m e i a nzzbicola

Fig. 3 11. Silverstoneia nubicola (Almirante, Bocas del Toro, Panama, 30 m). Photo: A. Hertz Silverstonia flofator

-b 7 ; :

Silverstonia nubicola

1

Fig. 312. Silverstoneia nubicola (P.N. G. D. Omar Torrijos H., near El Cop6, Cocl6, Panama). Photo: R. Brenes

Further Reading SAVAGE 1968a, 1-z & SMITH 1995, SAVAGE 2002

Terrarana New World direct 6 developing ogs)

New World direct developing frogs comprise a diverse assemblage of species that has a confusing taxonomic history. During the last decades of the lgth and the early part of the 2oth centuries, these frogs were assigned to the genus Hylodes, a name proposed by FITZINGER (1826). However, STEJNEGER (1904) determined that Eleutherodactylus was the correct name for the species that formerly had been referred to Hylodes because the latter name has to be applied to a group of frogs related to H. nasus (type species of the genus Hylodes, today in the family Hylodidae of FROST et al. 2006a). This conclusion was supported by G. S. MYERS (1962). Recently, in a series of revisions, the species of Eleutherodactylus (sensu lato) were split into several families and genera (CRAWFORD & SMITH 2005, HEINICKE et al. 2007, HEDGES et al. 2008). HEDGES et al. (2008) included the frog species formerly referred to as Eleutherodactylus (sensu lato) in the taxon Terrarana, an unranked above-family-group name that corresponds to the family Brachycephalidae (sensu lato) of other authors (e.g., VITT & CALDWELL 2009). The Terrarana of HEDGES et al. (2008) includes 883 described species allocated to four families, four subfamilies, 24 genera, 11 subgenera, 33 species series, 56 species groups, and 11 species sub- groups. Thus, Terrarana includes about 28 percent of anurans in the New World tropics and about 13 percent of frogs worldwide (DUELLMAN & LEHR 2009). In Central America, there occur representatives of three families of Terrarana (Craugastor- idae, Eleutherodactylidae, and Strabo- mantidae) with a total of 123 recognized species.

Key to the species of the genera Craugastor, Diaapom, Eleuthdactglus; and Aistimantis of northern Central America (southern Mexico to Nicaragua) 1 a Toe V longer than Toe I11 when each

adpressed to Toe IV; webbing between toes reduced (no webbing or only basal

.............................. webbing; Figs. 314a,b) 2 b Toe V shorter than Toe I11 when each

adpressed to Toe nT; webbing between toes ................................................... variable 15

2 a A pale transverse line demarcating finger and toe discs (Fig. 313); most digits with palmate or spadate covers (Fig. 3150; SVL of adult males 16-21 mm, of adult females 18-24 mm .................. Diaspom diadems

b No pale transverse line demarcating finger and toe discs; disc covers on Fingers 111-IV and on all toes round or truncate (Figs. 315c,d); SVL variable ............................... 3

3 a Disc covers on the outer two fingers (I11 and IV) moderately expanded, rounded (Fig. 314b, 315c) ........................................ 4

Further Reading CRAWFORD 2003, CRAWFORD & SMITH 2005, HEINICKE et al. 2007, HEDGES et al. 2008, DUELLMAN & LEHR 2009

Fig. 313. Diasporus diastema; note pale line demarcating discs (arrows). Photo: G. Kohler

Fig. 314. Foot of a. Diasporus diastema; b. Pristimantis ridens. Drawings: M. Vesely

rig. 315. Finger disc covers of a. Craugastorpodiciferus (unexpanded, even); b. C. jota (slightly expanded); c. Pristin~antis caryophyllaceus (expanded, even, round); d. Craugastor fitzingeri expanded, truncate, slightly emarginate); e. C. monnichorum (expanded, truncate, distinctly

emarginate); f. Diasporus diastema (expanded, spadate); g. Diasporus diastema (expanded, lanceolate). Photos: G. Kohler

Lliasporus diastema (even, triangular).

Fig. 316. I'mger d~sc pads of a. mstunantls caryopnyuaceus (even, ovoid); b. Craugastor fitzingen' (even, broadened); c. C. monnichonun (even, truncate); d. Diasporus diastema (even, ovoid); e.

Photos: G. Kohler

I?&. 317. Lateral view of head showing superciIievy tubercles and enlarged supraocular tubercles ie a Craugastar fitzingeri; b. C. massidigim c. C. sf;eja@iaaus; d. C. rugosn% e. Diasporus diadem4 f Eleuthemdactylus eoqui; g. Plis&nmtis ridem; h, P. ceminus; i. P. cruenb, j. P. museosm. Photos: a.-i. G. Kiihler; j. A. Hertz

141

b Disc covers on the outer two fingers (111 and IV) greatly expanded, truncate, and

......................... emarginate (Figs. 315d,e) 5 4 a No dark supratympanic mark; a large heel

tubercle present (Fig. 336e); subarticular tubercles pungent to pointed; usually W- shaped or H-shaped ridges on anterior portion of dorsum (Fig. 318a); eyes large and protruding in life; SVL of adult males to 25 mm, of adult females to 35 mm .................. ................................ Aistimanti.9 aerasinus

b A distinctive dark supratympanic mark usually present (Fig. 318b); no heel tubercle (Fig. 336a,b); subarticular tubercles round, not projecting; no W-shaped or H- shaped ridges on anterior portion of dorsum; eyes moderately sized and not protruding in life; SVL of adult males to 19 mm, of adult females to 25 mm .................. ..................................... Aietimantis nf ldw

ti a Disc of Toe V wider than that of Toe I1 (Fig. 319a); no vocal slits in males ................... 6

b Disc of Toe V subequal or narrower than that of Toe I1 (Fig. 319b); vocal slits in males present or absent ........................... 8

6 a Total number of vomerine teeth high (6-14); SVL of adult males 20.9-23.7 ram,

................... of adult females 26.6-29.3 mm ................................... Craugador cdmpbelli

I b Total number of vomerine teeth low (0-8); SVL of adult males 24.0-27.3 mm, single known female 39.8 .................................... 7

7 a Proximal subarticular tubercle on Toe V m obtuse, pungent, or pointed; SVL of adult

males 24.0-26.8 mm (females unknown) J ...................................... ..-tor -.- b Proximal subarticular tubercle on Toe V

globular or conical; SVL of adult males 25.1-27.3 mm; single known female 39.8 .. ........................ Craugastor cyanochthebius

8 a Ear opening very large (Fig. 320a; ratio tympanum length I eye length >0.70); supratympanic fold prominent ................ 9

b Ear opening moderate to large (Fig. 320b; ratio tympanum length I eye length <0.70); supratympanic fold present or absent .. 10

9 a Inner tarsal fold absent; dorsum with low, inconspicuous pustules; flank drab, mottled with brown .......... Cmugaetor glaucus

b A strong inner tarsal fold present along entire length of tarsus; dorsum smooth; flank with darker patches .......................... ....................................... Craugarstor taylon'

10a Ear opening moderate (ratio tympanupl length I eye length <0.50 in males, ~0 .35 in females); vocal slits present in males ........ ...................................... ~ b o c o u r t i

b Ear opening large (ratio tympanum length

Fig. 318. Head of a. Pristiznantis cerasinus and b. P. ridens; note W-shaped mark in P. cerasinus (arrow) and dark supratympanic mark in P. ridens (arrow). Photos: G. Kohler

Fig. 319. Foot of a. Craugastor nefEens and b. C. campbelli. Photos: G. Kohler

E ig. azu. Larerw neaa or a. ~raugastor glaucus and b. C. xucanebi. Photos: G. Kiihler

/ eye length >0.50 in males, >0.35 in females); no vocal slits in males ........... 11

lla Overall dorsal coloration pale without distinct dark markings; dorsum completely smooth; an inner tarsal fold present for one-third length of tarsus; occurs in the northeastern portion of the Yucatan Peninsula ............. Craugadmyucatanensis

b Usually with some dark dorsal markings; texture of dorsum variable; inner tarsal fold present or absent; occurs in the base of the Yucatan Peninsula or in the highlands of southern Mexico and Guatemala ..... 12

12a Venter dark brown; ratio tympanum length / eye length >0.50 in females; SVL of single known female 40.2 mm ................ .................................... Craugador siIv1'cola

b Venter dirty white or pale cream color; ratio tympanum length / eye length variable; SVL of adults variable ................. 13

13a Occurs in the lowlands (sea level to 600 m elevation) &om central Veracruz, Mexico, to western El Pet&, Guatemala ................ ., .................................... czmga&m a k d i

b Occurs in the highlands (600-2200 m elevation) of southern Mexico and Guatemala ................................................................. .14

14a Occurs in the central highlands of Guatemala .............. Crazzgwbr xucanebi

b Occurs in the highlands of western Guatemala and adjacent Chiapas, Mexico ...................................... Craugaator

It% Finger discs not expanded, although disc on outer fingers (111 and IV) can be slightly swollen (Figs. 315a,b) ......................... 16

b Finger discs narrowly to broadly expanded (Figs. 315c,d,e, 331) ................................ 26

16a Terminal transverse groove on Fingers I and I1 well-developed (Fig. 322a) ......... 17

b Terminal transverse groove on Fingers I and LI weak or absent (Fig. 32213) ........ 19

17a Three large, well-developed palmar tubercles; first finger longer than second .......... ............................ E I e u t h d ~ I u s l e p m

b Two large palmar tubercles, outer (third) palmar tubercle reduced in size or absent; first finger shorter than second ............ 18

l8a Dorsum dark with pale (red in life) spots ... and blotches; digitai pads not expande'd

............ Eleuthedactylus rubrimamrlatus b Dorsum dark brown with numerous pale

(cream to vellowish in life) s~o ts : digital . * . - ............................... pads slightly expanded

........................ J5'leuthedac&lus pipilans 19a Finger and toes with supernumerary

tubercles ................................................. 20

Fig. 321 doration of thigh in a. Craugasl yucatanensis, b. C. stuarti, and c. C. xucaneor.

Photos: G. Kohler

rig. 322. Tip of Finger I in a. Eleutherodactylus rubrimaculatus; b. Craugastor podiciferus; note terminal transverse groove in a. (arrow).

Photos: G. Kohler

Fig. 323. Foot of Craugastor loki; note supernumerary tubercles and inner metatarsal tubercle (arrow). Drawing: M. Vesely

b Finger and toes without supernumerary tubercles .................................................. 25

20a One or two inner metatarsal tubercles (Fig. 323); no vocal slits in males; nuptial pad present on thumb in males; no thin mem-

..................... branous web between toes 21 b No inner metatarsal tubercles; vocal slits

thumb in males present or absent; basal webbing usually present between Toes

I in males present or absent; nuptial pad on .

11-IV ........................................................ 22 I

21a All fingers and toes bearing ventral pads . ............................................ Craugastor loki

b Innermost fingers, at least thumb, and toes lack ventral pads ...... Craugastor rhodopie

22a A thenar and two palmar tubercles; no vocal slits and no nuptial pad on thumb in

...................... males Craugastor montanus b A thenar and one palmar tubercle; vocal

slits in males present or absent; nuptial pad on thumb in males present or absent . .................................................................. 23

23a No inner tarsal fold; no vomerine teeth; no nuptial pad on thumb in males ................................. hugastor pggmaeus

b A distinct inner tarsal fold present; vomerine teeth present; nuptial pad on thumb in

......................... males present or absent 24 ............ 24a No nuptial pad on thumb in males

................................... Craugastor lauraster b A nuptial pad on thumb in males present

(Fig. 342) ............... Craugastor b&ordii 26a Posterior dorsum with a definite lat-

erosacral system of either V-shaped or non- converging parallel rows of tubercles or ridges; venter with a dark brown reticulum, although this reticulum can be reduced to scattered dark spots (Fig. 325); SVL of adult males 3043 mm, of adult females 50-70 mm ...................................... .......................... Craugastor megamphalus

b Posterior dorsum without a definite laterosacral system of either V-shaped or parallel rows of tubercles or ridges; venter patternless; SVL of adult males 18-22 mm, of

............................. adult females 35-44 mrn .................................... Craugastor aphanus

26a No thin membranous web between toes .... ................... Eleuth-lus plauimstrie

b Toes webbed, at least basally ................ 27 27a A dark eye mask from snout to at least

tympanum present (Fig. 324), if this mask is reduced to contrastingly barred lips, then a distinct tympanic stripe present above and posterior to tympanum; snout slightly protruding with rounded upper edge in profile ........................................ 28

Fig. 324. Craugastor mimus; note dark eye mask along lateral head (Cerro El Toro, Atlantico Norte, Nicaragua. Photo: G. Kijhler

Fig. 325. Craugastor megacephalus (ventral surfaces; Cerro El Toro, Atlantico Norte, Nicaragua. Photo: G. Kohler

b No dark eye mask as described above; .......... snout usually rounded in profile 34

28a Discs on outer two fingers (111 and IV) markedly larger than those on inner fin-

......................................... gers (I and 11) 29 ..................... b Discs on fingers subequal 30

29a Inner tarsal fold present; sole of foot with definite low plantar tubercles (Fig. 326a); dorsum smooth; glandular supratympanic ridge extending at least one-third of body ....................................... Craugaetor noblei

b Inner tarsal fold absent; sole of foot smooth (Fig. 326b,c); dorsum granulate; glandular supratympanic ridge not

................... extending posteriorly to axilla .............................. Craugastor talamancae

30a Toes with moderate webbing, modal formula I11 3 - 4 IV (Fig. 326c); toes with well- developed lateral fleshy fringes continuous with webs (Fig. 326c) .. Crau.gahr m i m ~

b Toes with basal webbing, modal formula III 3+-4 l/2 IV; toes with non-fleshy lateral keels ........................................................ 31

Sla Heel smooth to rugose but without pustular tubercles or calcars; no distinct dark stripe along outer thigh margin; most finger disc covers slightly to lanceolate, with swollen to cuspidate disc pads; no dark seat patch mark ................. Craugagtor latioeps

b Heel with 1-4 more or less distinct small pustular tubercles or 1-2 large calcars; a distinct dark stripe along outer thigh margin; finger disc covers round, with even disc pads; a dark seat patch mark present .. 32

S2a Heel with 1-2 large tubercles (calcars); without paired dark spots at the apex of the triangular dark seat patch mark ......... .................................... C r a m lineatus

b Heel with 1-4 more or less distinct small pustular tubercles; paired dark spots present at the apex of the triangular dark seat

.............................................. patch mark 33

Fig. 326. Foot of a. Craugastor noblei; b. C. talamancae; c. C. mimus. Drawings: M. Vesely

33a Heel pustules weak; row of outer tarsal tubercles very weak or absent; usually no pale vertebral stripe, although a weak, thin ridge similarly colored as adjacent dorsum present in most individuals; SVL of adult males 30-36 mm, of adult females 40-58

............................. mm Craugastor rastralis b Heel pustules well-developed; row of outer

tarsal tubercles distinct; a thin pale vertebral stripe usually present; SVL of adult males 22-27 mm, of adult females 31-39

................................... mm chac 34a Inner tarsal fold absent; tympanum usu-

.... ally hidden or indistinct, rarely distinct QX

b Inner tarsal fold present; tympanum prominent ........................................ 48

36a Body with a pair of prominent dorsolateral ridges ....................................................... 36

b Body without a pair of dorsolateral ridges, although irregular ridges and tubercles can be present ........................................ 38

36a Dorsum smooth to rugose but never with numerous pustules or tubercles (Fig. 328a); no nuptial pads in males ............................ ........................................ Craugastor

b Dorsum strongly granular with numerous ,- large tubercles (Fig. 328b); males with nup-

.................................................. tial pads 37 37a No toe webbing; venter areolate; SVL of

adult males 19-23 mm, of adult females 2 mm ...................... .

b Basal toe webbing; venter almost smooth; 4 SVL of adult males 36-50 mm, of adult

..... females to 65 rnrn Craugarstor matudai 38a Toes with a distinctive lateral fleshy fringe

(that fold ventrally, a t least in preservative) ......................................................... 39

b Toes usually with a well-developed lateral keel, but not with a lateral fleshy fringe (Toe IV with weak fleshy fringes in some large females of C. fecundus; SVL in adult

................... C. fecundus 29.8-37.3 mm) 44 39a Toes extensively webbed (webbing formula

between Toes 111 and IV 1 213 - 3); dorsum wrinkled, a few tubercles laterally and above vent; SVL of adult males 3 4 4 1 mm, of adult females 37-46 mm ......................... ............................. Craugastor chgmozetetes

b Toes moderately or basally webbed (webbing formula between Toes I11 and IV 3 - 4 or 3 - 4 112); dorsum wrinkled or strongly granular with numerous tiny, moderate- sized, or large tubercles; SVL of adult males 21-34 mm, of adult females 2 8 4 7

40a Ratio disc width on Finger I11 / tympanum ..................................... length 0.69-0.93 41

b Ratio disc width on Finger I11 / tympanum length 0.38-0.58 ..................................... 42

41a Upper jaw flared beneath orbit (Fig. 327); snout truncate in profile ............................. ........................... Craqgastor trachydmue

b Upper jaw not flared beneath orbit; snout rounded in profile ........... Craqgastor-'

42a Snout obtuse (sloping) in profile; tympanum indistinct but visible in females (con-

........................... dition in males unknown) C r a e myllam~llan ...............................

b Snout rounded to nearly vertical in profile; tympanum hidden in females, indistinct or hidden in males ...................................... 43

43a Dorsum with tiny tubercles; webbing of toes basal to moderate, modal webbing formula 111 3 4 4 4 1/41 IV ( 4 + 4 ll4)-(2 ll2-2 314) V; SVL of adult males 21-27 mm, of

............................. adult females 28-37 mm .............................. Craugastor epochthidiue

b Dorsum with moderate-sized tubercles; webbing of toes moderate, modal webbing formula 111 3 - 4 IV 4-2 112 V; SVL of adult males 27-33 mm, of adult females 33-47 mm ........................ hugastor 8taddinani

44a Upper lip distinctly flared; tympanum, when visible, large (ratio tympanum length / eye length 0.72-0.85 in males, 0.39-0.52

............. in females); no vocal slits in males ................................. C r a m amoaenaie

b Upper lip margin smooth; upper jaw not flared beneath orbit; tympanum, when visible, large (ratio tympanum length / eye length 0.35-0.63 in males, 0.27-0.47 in

... females); vocal slits present in males 45 #a Dorsum with large tubercles (Fig. 328~);

tympanum distinct in males, indistinct in females; SVL of adult males 28-31 mm, of

............................. adult females 39-41 mm ................................ Craummk adamaatus -

b Dorsum with tiny tubercles; tympanum distinct or indistinct in males, indistinct or hidden in females; SVL of adult males 20-25 mm, of adult females 25-37 mm 46

46a Less toe webbing, webbing basal between

Fig. 327. Craugastor trachydermus; note flarea upper jaw (arrow). Photo: G. Kohler

Toes I-11, r e m a i h g moda formula I1 2+ -3 4/5 I I I 3 + 4 112 IV 4 ll2 -3- V; SVL of adult males 28-31 mm. of adult females I

............. 3 9 4 1 mm ~m'ugamkr s a l ~ m b More toe webbinp; modal webbing formula 1

I 2-2 415 11 2-3 3 4 111 3-4 ~4 IV 4 114 -2 3/4VorI2+-2 415 112-3 314 I I I 3 4 1/2 IV 4 1/2 -3- v ............................................... 47

47a Tympanum indistinct in males, usually hidden, rarely indistinct, in females; ratio tympanum length / eye length 0.36-0.49 in I males; usuallyless webbgg between Toes Fig. 328. Dorsum of a. Craugastor g r e g - b. C. 111-V, modal formula 111 3 4 1/4 IV 4 114 -2 d&; cc. C. adamastus. Photos: G. Kiihler

f rw. 329. Craugastor fitzingen - a. thighs; b. - -- .

Photo: G. Kohler

Fig. 330. Foot of a. Craugastor fitzingeri, b. C. rupinius, and c. C. chingopetaca.

Drawings: a,b M. Vesely, c. L. Czupalla

........... 3/4 V; occurs in northern Honduras .................................... hugas tar i&cundu~

b Tympanum usually distinct, rarely indistinct, in males, indistinct or hidden in females; ratio tympanum length / eye length 0.47-0.63 in males; usually more webbing between Toes 111-V, modal formula 111 3 4 1/2 IV 4 Y2 -3- V; SVL of adult males 20-25 mm, of adult females 25-37

..... mm; occurs in northwestern Honduras ......................................... Craugastor milmBB1

48a A dark seat patch mark present; disc covers on fingers and toes spadate; disc pads

......................... on k g e r s and toes swollen ....................................... Craugar~tor wffellg

b No dark seat patch mark; disc covers on fingers and toes rounded or truncated; disc pads on fingers and toes broadened or truncated ................................................. 49

...... 49a Toe webbing substantial (Fig. 330c) 60 b Toe webbing basal or moderate (Fig.

..................................................... 330a,b) 52 50a Inner tarsal fold ridge-like; discs on

Fingers 111-IV broadly expanded (usually >2.5 times width of digit just proximal to disc); posterior surface of thigh patternless;

........................ vocal slits present in males ............................. hugae tor chingopetam

b Inner tarsal fold flap-like; discs on Fingers 111-IV narrowly expanded ( ~ 2 . 5 times width of digit just proximal to disc); posterior surface of thigh brown with small distinct pale spots or dark gray with beige mottling or spotting; no vocal slits in males

-d

51a Dorsum finely granulate; discs on Fingers 111-lV broader expanded (ca. 2.0 times width of digit just proximal to disc); posterior surface of thigh brown with small distinct pale spots Crazrgastmmatandonenais

b Dorsum tuberculate; discs on Fingers 111-IV narrower expanded (ca. 1.5 times width of digit just proximal to disc); posterior surface of thigh dark gray with beige

.................................... mottling or spotting ................................ Craugastar sandans0mIU

52a Discs on Fingers 111-IV broadly expanded (usually >2.5 times width of digit just proximal to disc); snout subelliptical in dorsal aspect in males, subovoid in females; canthus rostralis angular; vocal slits and nuptial pads present in males; posterior thigh with pale mottling or with discrete and sharply demarcated spots (Fig. 329a); a

.... pale subgular stripe present (Fig. 329b) ................................... hugas tor fi-

b Discs on Fingers 111-IV narrowly expanded ( ~ 2 . 5 times width of digit just proximal to disc); snout usually nearly rounded in dor-

sal aspect; canthus rostralis rounded or angular; vocal slits and nuptial pads present or absent in males; chin and thigh a. coloration variable ................................... 63

.................................. 53a Toe webbing basal 54 b Toe webbing moderate ........................... 58

Ma Vocal slits present in males; canthus rostralis sharply angular; dorsum smooth; venter cream to reddish in life ................... ........................................... C r a w porn

b Vocal slits absent in males; canthus rostralis rounded to moderately angular; dor- fig. 331. Hand of a. Craugastorfitzingeri, and sum smooth, rugose, or tuberculate; b. C. 1aevissimus. Drawings: M. Vesely coloration of venter in life variable ....... 55

55a Discs on Fingers 111-N broader expanded (ca. 2.0 times width of digit just proximal to disc); largest toe discs 1.5 to 2.0 times width of digit just proximal to disc ....... 56 3

b Discs on Fingers 111-IV narrower expanded (~1 .5 times width of digit just proximal to disc); largest toe discs d . 5 times width of digit just proximal to disc .................. 57

56a Dorsum tuberculate; ratio shank length / SVL 0.46-0.57; canthus rostralis rounded; venter yellow or chartreuse, often with red-

of adult females 53-65 mm

P dish in life; SVL of adult males 26-36 mm, Fig. 332. Craugasfnr palenque; note flap-like

......................... inner tarsal fold (arrow). Photo: G. Kohler ....................... C r a u g a e t o r p e e p a m

b Dorsum smooth; ratio shank length I SVL 0.54-0.65; canthus rostralis moderately angular; venter dusky brown, purple, or grayish purple in life; SVL of adult males to disc); canthus rostralis sharply angular;

..... 37-51 mm, of adult females 66-83 mm venter whitish to cream in life; SVL of ...................................... C r a w - adult males 34-38 mm, of adult females

............... 57a Toes with a lateral narrow flange; dorsum 65-67 mm hugestor d w l a rugose; ratio shank length / SVL 0.55-0.64; b Discs on Fingers 111-IV broader expanded canthus rostralis moderately angular; (1.5 to >2.0 times width of digit just proxi- occurs on the Pacific versant of south-cen- mal to disc); canthus rostralis rounded to tral Honduras .............. Craugastar emleni sharply angular; coloration of venter vari-

........................ b Toes with a lateral keel; dorsum granulate; able; adult SVL variable 60 ratio shank length 1 SVL 0.51-0.57; can- 60a Canthus rostralis rounded ..................... 61 thus rostralis rounded; occurs on the b Canthus rostralis moderately to sharply Atlantic versant in east-central Guatemala angular .................................................... 65 ..................................... Craugagtor inachurr 61a Males with single dextral vocal slit (howev-

58a Inner tarsal fold flap-like (Fig. 332); discs er, possibly an anomaly in single known on Fingers 111-IV ca. 1.5 times width of male); number of vomerine teeth 2-3 in digit just proximal to disc; venter whitish males, 4 4 in females; discs on Fingers or yellowish in life; vocal slits absent in 111-IV ~ 1 . 5 times width of digit just proxi- ........ males; nuptial pads present in males ma1 to disc; SVL of adult males 27-35 mm, ................................... Chugastar palanque of adult females 43-51 mm; only known

b Inner tarsal fold ridge-like, or, if flap-like, from two sites in the Montaiia de Celaque, then discs on Fingers 111-IV ca. 2.0 times Departments of Lempira and Ocotepeque, width of digit just proximal to disc; col- western Honduras ..... C r a w anciano oration of venter variable; vocal slits and b vocal in males, if present, bilateral, ... nuptial pads present or absent in males number of vomerine teeth 2-11 in males, .................................................................. 51 5-12 in females; relative width of discs on

59a Discs on Fingers 111-N narrower expand- Fingers 111-IV variable; adult SVL vari- .......................................................... ed ( d . 5 times width of digit just proximal able 62

$2a Discs on Fingers 111-IV broader expanded (ca. 2.0 times width of digit just proximal to disc); vocal slits and nuptial pads present in males ................................................... 63

b Discs on Fingers HI-IV narrower expanded (ca. 1.5 times width of digit just proximal to disc); vocal slits and nuptial pads absent in males ...................................... 64

6&i Largest toe discs c1.5 times width of digit just proximal to disc; venter white, cream, or yellow in life; SVL of adult males 3144

................ mm, of adult females 69-81 mm ............................... Craugestor auzilegulw

b Largest toe discs >1.5 times width of digit just proximal to disc; venter yellowish green in life; SVL of adult males 33-43

................ mm, of adult females 59-62 mm .................................. Cra-peca-

(Ida Ratio tympanum length / eye length . 1.00-1.14 in males; SVL of adult males 18-30 mm (females unknown) ................... ................................. Craugwbr olanchano

b Ratio tympanum length / eye length 0.66-0.71 in males; SVL of adult males 26-45 mm, of adult females 40-74 mm ..... .................................... Crar;rgerrtorranoidee

@5a Discs on Fingers 111-IV narrower expanded (ca. 1.5 times width of digit just proximal to disc); vocal slits and nuptial pads absent in males ...................................... 66

b Discs on Fingers 111-IV broader expanded (> 1.5 times width of digit just proximal to disc); vocal slits and nuptial pads present or absent in males .................................. 67

66a Less toe webbing (modal formula I11 34+ IV); usually with bold lip bars; number of vomerine teeth 5-9 in males, 7-10 in females; SVL of adult males 29-38 mm, of adult females 51-67 mm ............................ .................................. Craugastor charadra

b More toe webbing (modal formula I11 3 - 4 IV); usually lacking bold lip bars; number of vomerine teeth 3-6 in males, 6-7 in females; SVL of adult males 25-28 mm, of adult females 44-57 mm ............................ .............................. Craugastor hepisaimus

67a Vocal slits absent in males; nuptial pads present in males; discs on Fingers 111-IV broader expanded (ca. 1.5 times width of digit just proximal to disc) .................... 68

b Vocal slits present in males; nuptial pads absent in males; discs on Fingers 111-IV narrower expanded (usually ~1.5 times width of digit just proximal to disc) ..... 69

68a Canthus rostralis moderately angulate; SVL of adult males 25-36 mm, of adult females 58-64 mm ..... C r a e n d m

b Canthus rostralis sharply angulate; SVL of adult males 41-57 mm, of adult females

................ 76-110 mm C r a e r P e l m 69a Dorsum rugose; SVL of adult males 28-40

mm, of adult females 58-72 mm ............... ................................... Cra- mpinius

b Dorsum smooth to granulate; SVL of adult males 38-44 mm, of adult females 69-83 mm ........................... C r a w t o r #*us

Figs. 333. Craugastor greggi (Finca Santa Fig. 334. Craugastor charadra (near Julia, San Marcos, Guatemala, 1490 m). Tegucigalpita, Honduras, 40 m).

Photo: J. Sunyer Photo: G. Kiihler

Key to the epeciea of the genera Crau- gastor, ~ a s p o ~ , E l e u t h ~ l u s , Ria- timantis, and S e m a n t i s of southern Central America (Nicaragua to Panama) 1 a Toe V longer than Toe 111 when each

adpressed to Toe IV; no webbing between ............................................................ toes 2

b Toe V shorter than Toe I11 when each adpressed to Toe IV; webbing between toes variable ................................................... 23

2 a A pale transverse line demarcating finger and toe discs on dorsal surface (Fig. 313); discs on outer fingers (111 and IV) subequal or only slightly larger than those on inner fingers (I and 11) (Fig. 314a); some digits with palmate, spadate, lanceolate, or papillate disc covers (Figs. 315f,g); most disc pads broadened or triangular (Figs. 316d,e); Finger I shorter than Finger 11; SVL of adult males 11-22 mm, of adult

................................... females 13-26 mm 3 b No pale transverse line demarcating finger

and toe discs; discs on outer fingers (111 and IV) usually distinctly larger than those on inner fingers (I and 11) (Fig. 314b); digits usually with round or truncate disc covers (Figs. 315c,d); disc pads usually ovoid or broadened (Figs. 316a,b); relative lenghts of Fingers I and I1 variable; SVL variable - . .................................................................... 8

3 a Disc cover lanceolate or papillate (Fig. 315g) .......................................................... 4

b Disc cover palmate or spadate (Fig. 3150 L:

4 a Dorsum shagreened; fingers without thick lateral fkinges; Toe V not partially fused

Fig. 335. Plistimantis pardalis (Cerro Man- posa, Veraguas, Panama); note white blotche on anterior surface of thigh (arrow).

Photo: G. Kohler

with Toe IV; SVL of adult males 14.0-16.0 ......... mm, of adult females 16.5-18.0 rnm

....................................... Diasparue vocator b Dorsum scattered low warts; fingers with

thick lateral fringes; Toe V partially fused with Toe IV; SVL of adult males 10.9-14.8

......... mm, of adult females 13.S16.9 mm ................................... Diaspo~u3 qutqutddiW

6 a Fingers I1 and I11 with palmate disc covers and broadened, non-triangular disc pads; adults with vomerine teeth ........................ .................................... DiaspoRu3 diastema

b Fingers I1 and I11 with spadate disc covers and triangular disc pads; adults with or

........................... without vomerine teeth 6 6 a Venter in most individuals with distinct

black and white blotches; dorsum and dorsal surfaces of arms and legs in some individuals bright red in life ............................ ........................ Diaaponte ventrtpaculatus

b Venter patternless or with a few small black dots; dorsum and dorsal surfaces of arms and legs brown, cream, or yellow in life ............................................................. 7

7 a Posterior surface of thigh unpigmented (yellow in life); adults with vomerine teeth ....................................... Diaspom tigrill0

b Posterior surface of thigh pigmented (brownish, often suffused with red in life); adults without vomerine teeth .................. ............................... D i a q o ~ ~ h y l a e f b d

8 a Surfaces of groin and anterior thigh with large contrasting pale blotches on a dark gray to black background (Fig. 335) ....... 9

b Surfaces of groin and anterior thigh without large contrasting pale blotches on a

............. dark gray to black background 10 9 a Discs on Fingers I11 and IV greatly

expanded, equal in size to tympanum; pale blotches of groin and anterior thigh silvery white in life ............ Ridmantie pard&

b Discs on Fingers I11 and IV moderately expanded, smaller than tympanum; pale blotches of groin and anterior . . thigh orange red in life ..................... A38hmantie altae

10a A well-developed pointed calcar (heel tubercle; Fig. 336e); usually an enlarged tubercle on upper eyelid (Fig. 31%-j) . 11

b Heel smooth (Fig. 336a) or with a one to several small tubercles of about the same size and similar to others scattered over upper surface of hind limb; enlarged tuber-

........ cle on upper eyelid present or not 15 l l a Snout long and pointed in dorsal view (Fig.

337a), ratio snout length / head width usually >1.1, ratio snout width / SVL usually

........................... ~0.37; dorsum smooth 12

Fig. 336. Heel region of a. Craugastor fitzingen'; b. C. podiciferus; c. C. monnichonrm; d. C. jota; e. Pristimantis caxyophyllaceus; note calcar (arrow). Photos: G. Kohler

b Snout of moderate length and rounded to truncate in dorsal view (Fig. 337b), ratio snout length 1 head width ~ 1 . 1 , ratio snout width / SVL usuallv >0.37; dorsum manu- - lar or tuberculate ................................... 13

l2a Tip of Toe V reaching beyond distal subarticular tubercle of Toe IV; tip of Toe I reaching beyond distal subarticular tubercle on Two 11; subarticular tubercles non-projecting; broadened and rounded h g e r discs and toe pads; inner metatarsal tubercle small, elongate, and obscure; outer metatarsal tubercle obscure, small, and rounded; no wartlike white spots on dorsum, arms, and lateral surfaces ................. . . ..................... A.lstzmantis CalyOphgIleoeurr

b Tip of Toe V not reaching distal subarticular tubercle of Toe IV; tip of Toe I not reach-

Fig. 337. Dorsal view of head in a. Pristimantis car~"ophy11aceus; b. P. cruentus.

Photos: G. Kiihler

ing distal subarticular tubercle on Two 11; subarticular tubercles projecting; expanded, round, and even finger discs and toe pads; inner metatarsal tubercle large and elongate; outer metatarsal tubercle large, conical, and projecting; most individuals with wartlike white to yellowish spots on dorsum, arms, and lateral surfaces ............ . . ............................... Arstunantis educatoris

13a Tip of Toe V not reaching distal subarticular tubercle on Toe IV (Fig. 340a); upper eyelid with several small tubercles along edge and several small warts on upper surface of eyelid (Fig. 317h); some subarticular tubercles pungent to pointed; tarsus with 1-2 outer tubercles; hidden surfaces of groin, thigh, and calf regions red in life ..... . ,. ................................ ti8cerasinw

b Tip of Toe V reaching distal subarticular tubercle on Toe IV (Fig. 340b); a well-developed pointed or flap-like tubercle on upper eyelid (Fig. 317ij); subarticular tubercles oval, globular, non-projecting; tarsus with 2 4 outer tubercles; hidden surfaces of groin, thigh, and calf regions not red in life .................................................................. 14

14a Dorsal coloration predominantly green in life (purple in preservative); usually one to several pale spots outlined by dark pigment; black groin spot in preservative; outer tarsal tubercles enlarged and conical ................................ Plietimantis mnseoeus

b Dorsal coloration predominantly tan to black in life (brown to black in preservative); groin usually with one to several pale spots outlined by dark pigment; outer tarsal tubercles moderately sized and . . rounded .................. A.letzmantis mentus

16a Tympanum internal, barely visible; tympanic annulus not evident (Fig. 338a) .. 16

b Tympanum prominent; tympanic annulus distinct (Fig. 33813) ................................. 18

16a A distinctive dark supratympanic mark usually present (Fig. 338b); upper eyelid with one to several supraocular tubercles . . (Fig. 317g) ................... Pnstunantis ridens

b No distinctive dark supratympanic mark usually present; no supraocular tubercles on upper eyelid ....................................... 17

17a Toes short, ratio Toe 4 lmgth I SVL ~0.21; coloration in life dark purplish brown dorsally with or without brick red mottling,

. grading into white along flanks and with dark brown eyes (in preservative dorsally dark brown, venter pale qay) ....................

R l s t m a U t i s m ................................. b Toes longer, ratio Toe 4 length I SVL >0.20; --

coloration in life green with a red head and red orange eyes preseNa- Fig. 338. Lateral view of head in a. Pristi- tive) ............................... P a mantis taeniatus and b. P. ridens.

18a Width of largest finger discs greater than Photos: a. C. Hernandez J.; b. G. Kohler

length of tympanum; toes with prominent lateral fringes; sole of foot with numerous supernumerary plantar tubercles, largest ones nearly as large as basal subarticular

........................... tubercle on Toes IV and V R&zmaJltis taaniatue ................................

b Width of largest finger discs smaller than length of tympanum; toes without prominent lateral Enges, although narrow later-

- a1 Enges can be present; sole of foot without or with only several supernumerary plantar tubercles, largest ones distinctly smaller than basal subarticular tubercle on Toes IV and V ........................................ 19

19a Venter smooth; toes with feeble lateral keels or narrow lateral fringes; Finger I usually longer than Finger 11; canthus rostralis sharp ............................................. 20 Fig. 339. Pristimantis gaigei (Nusagandi,

b Venter granulate or coarsely areolate; toes Kuna Yala, Panama, 350 m). Photo: G. Klihler without lateral keels or fringes; Finger I1 usually longer than Finger I; canthus rostralis weak ............................................. 21

20a Dorsum with a pair of vivid orange, orange red, or gold dorsolateral stripes on a black ground color (Fig. 339); width of larges finger discs <0.5 times tympanum length; on vocal slits absent in males ........................... ......................................

b Coloration not as above; width of larges finger discs >0.5 times tympanum length;

........................ vocal slits present in ma l~s PhEltunantie achatinus ...............................

21a Larger £rogs, SVL of adult males to 28 mm, of adult females 47 mm; disc covers trun- b. cate ..................... Eleuth-1-

b Smaller frogs, SVL of adult males to 37 mm, of adult females 52 mm; disc covers usually rounded ...................................... 22

22a Dorsum smooth are slightly granulate; posterior surface of thigh dark brown with cream spots or mottling; dark supratympanic stripe not extending to arm; nuptial pad absent in breeding males; SVL of adult Fig. 340. Foot of a. Pristimantis cerasinus; b. males to 28 mm, of adult females 47 mm .. P. cruentus. Drawings: M. Vesely

..................... Eleufilmdm&lus antillen& b Dorsum strongly granulate or tuberculate;

posterior surface of thigh pale brown; dark supratympanic stripe usually extening to arm; nuptial pad present in breeding males; SVL of adult males to 25 mm, of adult females 35 mm ................................... ..................... Eleuthemdactylus johnstonel

!Ba Finger discs not expanded, although disc on outer fingers (111 and IV) can be slightly

......... swollen, disc covers often pointed 24 b Finger discs narrowly to broadly expanded;

disc covers usually rounded, truncate, or emarginate ........................................ 36

Ma Sole of foot with supernumerary tubercles; ventral pattern not as described below .. 26

b Sole of foot without supernumerary tubercles; venter with a dark brown reticulum, although this reticulum can be reduced to

.............. scattered dark spots (Fig. 341) 32 8Sa A conspicuous large, triangular calcar on

heel; a prominent supratympanic-suprascapular ridge from posterior corner of eye to suprascapular region; ulnar and outer tarsal folds present .......... hqgastar jota

b Heel smooth to rugose or with 1-3 heel tubercles, no conspicuous large, triangular calcar; no prominent supratympanic- suprascapular ridge from posterior corner of eye to suprascapular region; ulnar and outer tarsal folds absent; SVL of adults variable ................................................... 26

2$a Heel with a 1-3 well-developed conical tubercles; subarticular tubercles on hands low and rounded; venter smooth, at least centrally; ventral disc circular with its posterior margin not reaching level anterior to insertion of hind limbs; no nuptial pads in males; SVL of adult males 21-28 mm, of adult females 2 3 4 0 mm ............................. ................................ craugaetorpodi*

b Heel smooth to rugose or with several very small subequal tubercles; subarticular tubercles on hands projecting; venter coarsely areolate; ventral disc V-shaped with its posterior margin reaching beyond level of insertion of hind limbs; nuptial pads in males present or absent; SVL of males to 26.6 mm, of females to 30 mm 27 Toes I11 and IV usually without supernumerary tubercles; subarticular tubercles of hands and feet low and round, usually obtuse in lateral view; nuptial pads present in males; SVL of adult males 16-26 mm, of

............................. adult females 18-30 mm ............................... Craugagtor undarwoodi Toes 111 and IV with supernumerary tubercles; subarticular tubercles of hands and feet projecting and pointed, usually obtuse

Fig. 341. Craugastor rugosus (ventral surfaces; Refugio El Rodeo, Costa Rica). Photo: G. Kiihler

Fig. 342. Hand of male Craugastor bransfordii; note nuptial pad (arrow) Photo: G. Kahler

in lateral view; nuptial pads present or absent in males; SVL of adults variable 28

28a Finger I1 longer than Finger I; SVL of males to 18 mm, of females to 23 mm; nup-

....................... tial pads absent in males 29 b Fingers I and 11 subequal; SVL of adults

variable; nuptial pads present or absent in ....................................................... males 30

29a Toes with basal webbing; a weak inner ... tarsal fold present C r a m Iaurantar

b No thin membranous web between toes; inner tarsal fold barely evident .................. ................................. craugaetorperaimilie

30a Thenar tubercle much smaller than palmar tubercle. about the same size as subarticular tubercles of Fingers II-IV (Fig. 343a);

nuptial pads absent in males; SVL of adult males 12-18 mm, of adult females 13-22 mm .................. Craugastar stejnegehnus

b Thenar tubercle about the same size or only slightly smaller than palmar tubercle, much larger than subarticular tubercles of Fingers 11-IV (Fig. 343b); nuptial pads present or absent in males; SVL of adults variable ................................................... 31

31a Nuptial pads present in males (Fig. 342); SVL 13-23 Fig. 343. Hand of a. Cravgastor stejnegerianw, females 14-26 mm C r a w t a r and b. C. bransfordii. Drawings: M. Vessljr

b Nuptial pads absent in males; SVL of adult males 15-27 mm, of adult females 16-29 mm ....................... CraugastarP0b'P-m b Upper eyelid without prominently elongat-

32a Dorsum essentially smooth, lacking ed supraocular tubercle, although several suprascapular ridges, but in juveniles char- small superciliary tubercles usually pre- acteristically figured weak suprascapular sent; eyes brown or grayish brown in life ridges present, which disappear during ........................ Strabomantis b&- 0ntogenY; SVL of adult females 7s103 mm 378 A dark eye mask from snout to at least (adult males unknown) Craugtwtargdmue tympanum present, if this mask is reduced

b Dorsum of adults and juveniles with dis- to contrastingly barred lips, then a distinct tinct paired suprascapular ridges; SVL of tympanic stripe present above and posteri- adult females to 70 mm ......................... 33 or to tympanum (Figs. 345 and 346); snout

33a Posterior surface of thigh with pale and slightly protruding with rounded upper dark spotting or mottling; posterior dorsum edge in prome; finger disc non-emarginate

.................................................................. with a definite laterosacral system of 38 either V-shaped or non-converging parallel b No dark eye mask as described above, or, if rows of tubercles or ridges .......................... dark eye mask present, then finger disc

C r a ~ m e g a o 8 p a a l u s emarginate; snout usually rounded in pro- .......................... ............................................................ b Posterior surface of thigh with alternating file 41

black and red (white in juveniles and in 38a Discs on outer two fingers (111 and Tt3 preservative) or brown to black with or markedly larger than those on inner fin- without may areas; posterior dorsum with- gers (I and 11) ......................................... 39 out a d e h b laterosacral system of either V-shaped or parallel rows of tubercles or ridges ....................................................... 34

34a Posterior surface of thigh brightly barred black and red in life (black and white in juveniles and in preservative); dorsum often with pale dorsolateral stripes; dorsum strongly tuberculate with numerous large warts and ridges Craugastor rugasus

b Posterior surface of thigh not brightly barred black and red (or white), although barred with black and tan in some individuals; no pale dorsolateral stripes on dorsum; dorsum smooth with scattered low pustules ....................... Craugaetor opitnus

36a Finger discs not expanded; head very ................ broad, head width >45% SVL 36

b Finger discs expanded; head usually not conspicuously broad, head width ~ 4 0 % SVL .......................................................... 37

36a Upper eyelid with prominently elongated supraocular tubercle; eyes blue or grayish

........... blue in life Sh.abamantis laticcupus

...................... b Discs on fingers subequal 40 39a Inner tarsal fold present; sole of foot with

definite low plantar tubercles (Fig. 326a); dorsum smooth; glandular supratympanic ridge extending at least one-third of body . ........................................ Craugtwtor noblai

b Inner tarsal fold absent; sole of foot smooth (Fig. 326b,c); dorsum granulate; glandular supratympanic ridge not extending posteri-

........ orly to axilla Craugastor talamancae 40a Heel with one or two well-developed cal-

cars; toes with moderate webbing, modal formula I11 3 - 4 IV (Fig. 326c); toes with well-developed lateral fleshy fringes continuous with webs (Fig. 32613; dark seat

.... patch mark forming an inverted V or U ....................................... Cmugastor mimus -

b Heel without tubercles; toes with basal webbing, modal formula 111 3 4 IV (Fig. 344b); toes without lateral fleshy fringes (Fig. 344b); dark seat patch mark usually reduced to a pair of dark spots above and lateral to vent, never forming an inverted V or U ............................. CraugastargaUman'

b Toe discs well developed; posterior surface of thigh with distinct pale spots or pale mottling; nuptial pads present or absent in

. males ....................................................... 46 &a Posterior surface of thigh black with small,

............................... regular yellow spots 46 b Posterior surface of thigh mottled with pale

yellow and brown ................................... 47 46a Vocal slits and nuptial pads present in

males; toe webbing extensive; inner tarsal fold flap-like; posterior thigh black with small pale (yellow in life) spots; SVL in adult males 35-56 mm, females 62-86 ...... ....................................... hugestor 0-

b Vocal slits and nuptial pads absent in males; toe webbing strong; inner tarsal fold very strong; posterior thigh black with large pale (yellow in life) spots; SVL in adult males 32-50 mm, females 56-64 ...... ................................... C r a e evanemm

Fig. 344. Foot of a. Craugastor crassidigitus; b. C. gollmeri; c. C. melanostictus.

Drawings: M. Vesely I 41a Toe webbing substantial (Fig. 344a) ..... 42

b Toe webbing basal or moderate, or no webbing at all (Fig. 344b,c) .......................... 48

42a Posterior surface of thigh patternless, not spots or mottling; discs on Fingers 111-N broadly expanded (usually >2.5 times

I width of digit just proximal to disc) ...... 43 Fig. 345. Craugastormimus (Selva Negra, Mata-

b Posterior surface of thigh bra- or black galpa, Nicaragua, 1300 m). Photo: G. Kiihler with small distinct pale spots or mottling; discs on Fingers 111-IV narrowly expanded (~2.5 times width of digit just proximal to disc) ......................................................... 44

4% Finger and toe disc covers retuse; indistinct accessory palmar tubercles present only on base of Fingers I and 11; supratympanic ridge only slightly curved downward; midgular pale stripe usually absent .......... ............................ C r a m Chingvpetam

b Finger and toe disc covers non-retuse; distinct accessory palmar tubercles present on base of all Fingers; supratympanic ridge distinctly curved downwards; midgular pale stripe usually evident or at least indicated anteriorly .. Craugastor itus us

44a Toe discs weakly developed; posterior sur- m face of thigh with pale mottling; nuptial Fig. a*o. b h u g ~ t o r ~ o l l m e r i (Nusagandi,

.... pads absent in males CrarTgastar taupas Kuna Yala, Panama, 350 m). Photo: G. KWer

47a Venter white in life; fingers with lateral keels; 6-9 vomerine teeth in males, 9-15 in females; occurs ... Crauga&upun6d~

b Venter pale yellow in life; fingers with lateral fringes; 6-7 vomerine teeth in males, 8 in females .... Craugaetar rhyawbatrachus

48a Disc on some Fingers (usually I11 and IV) emarginate (Figs. 315e, 316c) ................ 49

b Discs on Fingers non-emarginate (Fig. 315c, 316a,b) ........................................... 66

49a Toe webbing moderate ........................... 60 b Toe webbing basal or no webbing at all 51

60a Posterior surface of thigh patternless (dark purple in life); a heel tubercle present .......................................... Craugaetorrago

b Posterior surface of thigh chocolate brown with large bright yellow stripes andlor

.... spots; no heel tubercle Craugaetar an& 5la Overall coloration including that of posteri-

or surface of thigh grayish white without any distinct pattern .. C ~ ~ U @ & C W ~

b Coloration not as above ......................... 52 52a Posterior surface of thigh with dark trans-

verse bars that alternate with light areas (Fig. 347); Finger 11 distinctly longer than Finger I (tip of Finger I reaches to distal end of penultimate phalanx on Finger 11) -- . ................................................................. W

b Posterior surface of thigh with mottling, spotting, or small bright yellow spots that may fuse to form vertical stripes; usually Fingers I and I1 subequal or Finger I longer than Finger 11, rarely Finger I1 slightly longer than Finger I ............................... 64

63a No heel tubercle; toe webbing basal, nup- ........................ tial pads absent in males ; ....

.................................. Craugaet~~ tnbamme b A heel tubercle present; no toe webbing;

.................... nuptial pads present in males ........................... C r a m m e l a n - ' a

Ma Finger I longer than Finger II; posterior surface of thigh dark brown with small bright yellow spots that may fuse to form vertical stripes ......... hugas tar w q u m

b Fingers I and I1 subequal or Finger I1 slightly longer than Finger I; posterior surface of thigh very dark with fine pale stippling or dots ............................................ 66

55a Heel with several tubercles; posterior surface of thigh very dark with fine bluish-

............ gray stippling C r a w t o r emcalae b Heel with one tubercle; posterior surface of

thigh very dark with fine white dots; Finger I1 longer than Finger I; a heel tubercle present ........ Craugaetor m-choru~n

Fig. 347. Craugastor melanostictus (Sendem Los Quetzales, Chiriqui, Panama, 2135 m).

Photo: A. Hertz

Ma Toe webbing basal or no webbing at all 57 ........................... b Toe webbing moderate 60

57a Finger I1 longer than Finger I; a moderately-sized heel tubercle present; no toe webbing; SVL in adult males 19-20 mm,

....... females unknown Aistimantie adnus b Finger I1 shorter than Finger I; no heel

tubercle; minimal to basal toe webbhing present; SVL in adult males 26-46 mm, females 38-75 ......................................... 68

68a Toe discs barely enlarged; discs on fingers weakly enlarged, discs on Fingers I11 and IV about 1.25 times the width of the digit; posterior surface of thigh blqtched or. mottled ................ Craugastor fl-

b Toe discs definitely enlarged, discs on Toes 111-V at least 1.5 times the width of the digit; discs on fingers weakly enlarged, discs on Fingers 111 and IV at least 1.5 times the width of the digit; posterior surface of thigh uniform or with obscure pale mottling ................................................... 59

59a No vocal slits in adult males; discs on Fingers 111 and IV about 1.5 times the width of the digit; venter bright pale yellow, orange, or red in life ............................. .................................. Craugaetor axgelicu~

b Vocal slits present in adult males; discs on Fingers 111 and IV almost 2.0 times the width of the digit; venter bright tomato red

........................... in life Cra- 60a Posterior surface of thigh patternless

brown ................... C r a w t o r 1- b Posterior surface of thigh with mottling,

................................ flecking, or spotting 61 61a Discs on Fingers 111-IV broadly expanded

(usually >2.5 times width of digit just prox-

Terraram

imal to disc); snout subelliptical in dorsal contrasting small pale spots on dark brown aspect in males, subovoid in females; can- background; vocal slits and nuptial pads

........ thus rostralis distinctly angular; vocal slits absent in males Craugaetor ranoidw and nuptial ads present in males ....... 62 b Toe discs moderately enlarged, discs on

b Discs on Fingers III-IV narrowly expanded Toes III-V 1.5 or less times the width of the (<2.5 times width of digit just proximal to digit; canthus rostralis rounded or moder- disc); snout usually nearly rounded in dor- ately angulate; posterior surface of thigh sal aspect; canthus rostralis rounded or with pale spotting or mottling; vocal slits moderately angular; vocal slits and nuptial and nuptial pads present or absent in

....................................................... pads present or absent in males ........... 63 males 64 6% Spots on posterior thigh usually discrete 64a Toe discs greatly enlarged, discs on Toes

and sharply demarcated, at least in adults 1114 almost 2.0 times the width of the (Fig. 329a); in life, thigh spots normally yel- digit; canthus rostralis rounded; posterior lowish green in individuals in central and surface of thigh with distinct small yellow eastern Panama, occasionally tan or gray spots; vocal slits and nuptial pads present in western Panama; median white gular in males; SVL of adult males 3045 mm, of

............................. stripe always present (Fig. 329b), although adult females 45-75 mm if throat is generally sparsely pigmented, ................................... Craugaetor catalinae

i the stripe may be only weakly indicated as b Toe discs moderately enlarged, discs on an area devoid of melanophores; SVL of Toes III-V 1.5 or less times the width of the adult males 23-35 mm, of adult females digit; canthus rostralis moderately angu- 36-53 mm ................ Craugaetar f i u late; posterior surface of thigh with a few

b Posterior surface of thigh with many pale rather small light spots or with pale spot- flecks or with larger, coalescing spots that ting or mottling; vocal slits present or are crowded and poorly defined, these absent in males; nuptial pads absent in markings normally pale brown or tan in males; SVL of adult males 24-37 mm, of life; melanophores normally distributed adult females 33-57 mm ........................ 66 across entire with a M a Dorsum granulate; posterior surface of median white line or very faint ill-defined thigh with a few rather small light spots; stripe; SVL of adult males 27-43 mm, of discs on Toes III-V about 1.5 times the adult females 52-74 mm (individuals of width of the digit; vocal slits present in both sexes smaller on the Pearl Islands, males .................... Crauga&~ m d Panama) ................. Craugaetor- b Dorsum smooth; posterior surface of thigh

63a Toe discs greatly enlarged, discs on Toes with pale spotting or mottling; discs on III-V almost 2.0 times the width of the Toes III-V g1.5 times the width of the digit; canthus rostralis rounded; posterior digit; vocal slits absent in males ................ surface of thigh with numerous sharply .............................. Craugmh laevissimw

Fig. 348. Craugastor laevzssmus (&lambe, ~'lg. 349. Craugastor monnichonun (Aha Nicaragua). Photo: J. Sunyer Chiquero, Volch Bani, Chiriqui, Panama,

1820 m). Photo: G. Kohler

Until a few years ago, with over 700 Neotropicd species, the genus Eleu&em- dac@lus sensu lato ranked as the most species-rich of all vertebrate genera ( C R A ~ O R D & S m 2005). Ftecently, this very large, unwieldy genus was split into several pnera, and these new genera were removed from the family Lephdactylidae md asaigned to three separate families: Craugastoridaq Ehutheradacd&dae, md Strabomantidae RAWFOR FORD 80 S~MPPH 2005, ~ C S E S et d. 2008). cmrentIy understood, Crauga8bridae contains two genera, fl.augmtor (11% species) and Bkddatw (2 species); the specie8 of Craugastoridae are distributed from southern Arizona and central Texas (USA) rind south 60 northwestern Ecuador and Colombia, and thmee to eastern Brazil (FaosT 2010).

Recently, CRAWFORD 6t SMun (20051, w d d

on molecular genetic data, elevated the former subgenus Craugwtor to the rank of genua, This genus represents a group .of frogs Wt has its greatest diversiiy i-n Middle America, and with 93 species, this genus is well reprersented in Central America, Many epecies of Craugmtur are notorious for the high degree of polymorphism and polycbomatism, meaning that intraspecific variation 11 coloration, dorsal skin te~ture, patterns of skin tubercles, war& and ridges can be asto&hingly great in some species (SAVAGE & EMERSON 1970). Because of this phenomenon, species diversity can easily be over- or underestimated. In the case ofthe C. ru&rr- losw specties group, early studies s e e d to grossly cwersimpliEy the situation, refer- ring most Mexican and Nuclear Central American populations to a single species (LYNCH 1965d, SAVAGE 1975); currently, a totd of 34 rrpeciaa is recognized in the C.

~ O S U S specia group, several of which have broadly overlapphg geographical ranges (CWBILL & SAVAGE 2000, R m et al, ZOlOb). Because of intraqedic v&- tion and supeffieial resemblance among many h s , species identification is not always an ea&y .Y task this group of frogs. This is especially true with p r e m e d spec- hens that have lost theb life colaration, which in some w e s may be helpful in dis- tingdehing among ~pecies.

Species of Cmugaator are ge11erdly emti- c a y colored inhabitwts of the rainforest. Some species are stream-associated and can be observed sit* on racks or the ground along stream@ at night (e-g., &base of the fitr&geri and ruguluslls gcoupe; M c C m & WILSON 2002). Others are diurnal leaf litter inhabitants and can reach high papulation densities lo& (e.g., members of the br;aasfor& specie8 group; S~Cvrr 1983b). Also diurnal leaf lib ter species, the members of tbe goheri species group may pass unnoticed by a per- son ,walking through the rainforest because of their cryptic domal pattern and coloration, but typically dart across the forest floor in a zigzag pattam sf long leaps when approached too closely (SAVAGE 1987). The species of Craugasbr are food generalists feeding on a wide variety of arthropads and lager species may eat 8.mall fkogs (SAVAGE 2002). Sexual dimorphism in tympanum she, with males having a larger tympanwn k h a n females, is typical far khe majority of species in Craug~tstor Q Y N ~ 2000; Fig. 350). Also, in most species of this genus, females are larger than males (CAMPBELL & SAVAGE 2000, SAVAGE 2002). The males of the majority of species in this genus have the ability to produce vocalizations to attract females during the breeding season ( S C ~ F F E R & FSGEROA-S~mf 1998). However, some species (e.g., those of the goUmd species group) apparently do not emit any kind of calls (%PAGE 19873. As f a as is known? dl spe@es in this genus have direct development, i.e., they lay encapsu- lated eggs in moist terrestrial sites, and

dimorphism in tympanum size. ~ i g . 3550. ~raugastorpodiciferus; note sexual Fig. sol. braugastor &edi (El Ocote,

Chiapas, Mexico). Photo: G. Kijhler Photos: G. Kohler

the embryos undergo direct development entirely within the egg membranes, hatching as small frogs without a free-swimming aquatic tadpole stage (VITT & CALDWELL 2009). Nest construction and female attendance of the clutch has been reported for several species of Craugastor (HAYES 1985, MENDOZA QUIJANO et al. 2002). HAYES (1985) inferred that C. angelicus may have internal fertilization.

Craugastor adamastus (CAMPBELL 1994b1, Herpetologica 50: 404; type locality: "northern slopes of the eastern portion of the Sierra de las Minas, Municipio Los Amates, 0.8 km W Aldea Vista Hermosa, 650 m, ... The type locality is located a t about 15"17W, 8g013W and is drained by the headwaters of the Rio Las Caiias, which flows into Lago de Izabal", east- e m Guatemala. Males to 31.0 mm, females to 40.8 mm. Known only from the type locality, 600-650 m elevation.

Craugastor aEedi (BOULENGER 1898a), Roc. Zool. Soc. London 1898: 481; type locality: "Atoyac, State of Vera Cruz", Mexico. Females to 45.0 mm (data for males not available). Central Veracruz, Mexico, to western El Pet&, Guatemala, sea level to 600 m elevation.

Craugastor amniscola (CAMPBELL & SAVAGE 2000), Herpetol. Monogr. 14: 227; type locality: "Santa Ana Huista, 920 m, Huehuetenango, Guatemala. This site lies in Premontane Dry Forest at about 15"41'N, 91°49W." Males to 38.8 mm, females to 67.4 mm. Middle and upper Rio Gfijalva Basin eom the Sierra de 10s Cuchumatanes and the Montafias de Cuilco in western Guatemala to north of

Fig. 352. Craugastor amniscola (El Zapotal Chiapas, Mexico). Photo: W. E. D u e l L m ~ (Courtesy of the Biodiversity Institute, University of Kansas) r

Fig. 353. Craugastor anciano (above Villa Verde, Lempira, Honduras). Photo: J. R. McCranie

Fig. 354. Craugastor andi (Bajo La Hondura, San Jos6, Costa Rica). Photo: W. Van Devender

159

Ocozocuautla in Chiapa O,~UU-1 - - m elevation.

Craugastor anciano (SAVAGE, MCCRANIE & W n s o ~ 1988), Bull. S. California Acad. Sci. 87: 53; type locality: "El Chagiiit6n (14"3OYN, 88"48'W), elevation 1830 m, 18.8 km SE

Corquin, CordilIera, [sic] de Celaque, Departamento de Ocotepque, Honduras." Males to 33.0 mm, females to 41.2 mm. Only known from two sites in the Montaiia de Celaque, Departments of Lempira and Ocotepeque, western Honduras, 1400-1840 m elevation.

Craugastor andi (SAVAGE 1974), Herpetologica 30: 295; type locality: "Costa Rica: Provincia de San Jose: Cant611 de Coronado: 0.8 km N junc- ture Rio Clam and Rio La Hondura, 1150 m." Males to 55 mm, females to 80 mm. Atlantic slopes of the Cordillera Central and extreme northeastern portions of the Cordillera de Talamanca, Costa Rica, 830-1500 m elevation.

Craugastor angelicus (SAVAGE 19751, Copeia 1975: 274; type locality: "Costa Rica: Proyincia de Alajuela: Cant6n de Alajuela: Salto El Angel, 1380 m." Males to 46 mm, females to 75 mm. Cordilleras de Guanacaste (Cerro Cacao), Tilarh, and Central (northern and eastern dopes of V o l h Po&) in Costa Rica, 600-1700 D elevation.

Craugastor, aphanus (CAMPBELL 1994a), Herpetologica 50: 297; type locality: 92.0 km WSW Puerto Santo Tom&, 786 m, De~artamento de Izabal. Guatemala. . .. This loiality is located on the'eastern slopes of the Montafias del Mico at about 15"41'N, 88"41W. The town of Santo Tom& is called Puerto Santo Tom& de Castilla or Puerto Matias de Gdvez on some maps." Males to 21.6 mm, females to 44.1 mm. Montaiiaa del Mico and the adjacent portion of the Sierra de las Minas, eastern Guatemala, 591-786 m elevation.

Craugastor aurilegulus (SAVAGE, MCCRANIE & WILSON 19881, Bull. S. California Acad. Sci. 87: 50; type locality: "Quebrada de Oro (15"38'N, 86"47W), elevation 780-840 m, tributary of Rio Viejo, south slope of Cerro Bdfalo, Cordillera de Nombre de Dios, Departamento de Atltintida, Honduras." Males to 49.8 mm, females to 80.5 mm. Cordillera Nombre de Dios, Montafia La Muralla, and Montaiia Texiguat, in north-central Honduras, 50-1550 m elevation.

to the Azuero Peninsula, Panama, 16940 elevation.

Craugastor bocourti (BRoccm 1877b), Bull. Soc. Philomath., Paris, Ser. 7,l: 130; type locality: "montagnes de Coban", Alta Verapaz, Guatemala. Females to 59 mm (data for males not available). Mountains of Alta Verapaz and the Sierra de las Minas. Guatemala. 1300-1700 m elevation. P K r . '. 8 - -

Males to 23 mm, f e a e s to 2k mm. 0; the Atlantic versant from eastern Nicaragua to southern Costa Rica, 60-1600 m elevation.

Craugastor brocchi (BOULENGER l882b in BROCCHI 1882), Miss. Scient. Mex. Amer. Centr., Rech. Zool. 3 (2, livr. 2): 60; type locality: "Guatemala." Males to 51.0 mm, females to 83.3 mm. On the Atlantic versant from Chiapas (Parque Nacional Lagunas de Montebello), Mexico, across the northern highlands of Guatemala (northwestern Huehuetenango through the Sierra de 10s Cuchumatanes to the Sierra de lraa Minas), 1200-2000 m elevation. '

Craugastor campbelli (SMITH 20051, Herpetologica 61: 287; type locality: Torres de Guatel, C e m San Gil, Montaikw del Mico, Municipio de Puerto Barrios, Departamento de Izabal, Guatemala, 962 m." Males to 23.7 mm, females to 29.3 mm. Montatias del Mico, Izabal Department, eastern Guatemala, 260-962 m elevation.

Craugastor catalinae (CAMPBELL & SAVAGE 20001, Herpetol. Monogr. 14: 276; type locality: "Las Alturas, Rio Bellavista, approximately 0.5 km N of bridge, 1400 m, Cantdn de Coto Brues [= Coto Brus], Provincia de Puntarenas, Costa Rica. This locality is in Tropical Premontane Wet Forest at about 8"75'N, 82"52W." Males to 44.8 mm, females to 75.3 mm. On the Pacific versant in the Rio Cot611 drainage of southern Costa Rica and adjacent western Panama (Pacific slopes of Volch Bani), 1220-1800 m elevation.

Craugastor chac (SAVAGE 1987), Fieldiana, Zool. N.S., 33: 31; type locality: "12.6 km W Santo TomBs, [Departamento] Izabal, Guatemala, 774 m." Males to 26 mm, females to 39 mm. Atlantic versant from central Belize and central Guatemala to the La Ceiba city area on the northern coast of Honduras, 20-1000 m elevation.

Craugastor azueroensis (SAVAGE 1975), Copeia 1975: 277; type local i~: T ~ ~ ~ : prOvincia de Craugastor charadra (CAMPBELL & SAVAGE 10s Santos: N slope Cerro Cambutal, 480 2000), Herpetol. Mono@. 14: 247; type locality: Males to 40 mm, females to 65 -. Endemic W P W ~ reaches of Webrada San Antonio, SW

I- - Fig. Costa 355. Rica). Craugastor angelicus Photo: (Monteverde, M. fianzen - Fig. CRARC, 359. Costa Craugastor Rica, 550 bransfordii m). Photo: (Guayach, G. Kijhler

Fig. 356. Craugastor aphanus (Guatemala). Fig. 360. Craugastor bmchi (Guatemala). Photo: J. A. Campbell Photo: J. A. Campbell

Fig. 35;. ~raugastcl~ aJrilegulus (QuetLa,, ~d Fig. 36;. v~,,,,,~r CL~,,, (M~-~..iias del Om, Atlhtida, Honduras). Photo: J. R. McCranie Mico, Izabal, Guatemala). Photo: J. A. Campbel

Fig. 358. Craugastor bocourti (Guatemala). Fig. 362. Craugastor chac (Cockscomb bserve , Photo: J. A. Campbell Stann Creek, Belize, 80 m). Photo: G. Kijhler

side Cerro del Nylon, 1310 m, Sierra de Espiritu Santo, Municipio Los Amates, [Departamento] Izabal, Guatemala. This site lies in Premontane Wet Forest a t about 15"12'33"N, 88O53'47W." Males to 37.9 mm, females to 67.0 mm. Highlands between the Rio Motagua in Guatemala and the Rio Chamelec6n in Honduras, 30-1370 m elevation.

Craugastor chingopetaca KOHLER & SUNYER 2006, Senckenb. Biol. 86: 262; type locality: "Chingo Petaca, Rio San Juan, 10°44'50.9"N, 83"50'26.3W, Departamento Rio San Juan, Nicaragua, 40 m elevation." SVL to 37.6 mm. Southeastern Nicaragua, sea level to 280 m elevation.

Craugastor chrysozetetes (MCCRANIE, SAVAGE & WILSON 19891, Proc. Biol. Soc. Washington 102: 483; type locality: "Quebrada de Oro (15"3B7N, 86"47W), elevation 880 m, tributary of Rio Viejo, south slope of Cerro Btifalo, Cordillera de Nombre de Dios, Departamento de Atlhtida, Honduras." Males to 41.3 mm, females to 45.6 mm. Known only from the type locality, 880-1130 m elevation.

Craugastor coffeus (MCCRANIE & K ~ H L E R 1999b), Senckenb. Biol. 79: 83; type locality: "between Laguna del Cerro and Quebrada Grande (15"04.95'N, 88"56.08W), 1000 m elevation, Departamento de Coptin, Honduras." SVL to 33.9 mm. Extreme western Honduras, around 1000 m elevation.

Craugastor crassidigitus (TAYLOR 1952b), Univ. Kansas Sci. Bull. 35: 740; type locality: "Isla Bonita, eastern slope Volcan Pols", Cant611 de Alajuela, Provincia de Alajuela, 1680 m, Costa Rica. Males to 31 mm, females to 48 mm. Northern Costa Rica to extreme northwestern Colombia. sea level to 2000 m elevation.

Craugastor cruzi (MCCRANIE, SAVAGE & WILSON 19891, Proc. Biol. Soc. Washington 102: 485; type locality: "south slope of Cerro Blifalo (15"38'N, 86"47'W), elevation 1520 m, Cordillera de Nombre de Dios, Departamento de Atlhtida, Honduras." Males to 33.2 mm (females unknown). Only known from the type locality.

Craugastor cuaquem (SAVAGE 1980), Bull. S. California Acad. Sci. 79: 14; type locality: "1.75 km east southeast of Monteverde; Provincia Puntarenas, Cant6n de Puntarenas, Costa Rica; 1520 m." Males to 39 mm, females to 48 mm. Known only from the type locality, 1520-1600 m elevation.

Craugastor cyanochthebius MCCRANIE & SMITH 2006, Herpetologica 62: 185; type locality: "San Isidro, on the western slopes of Montaiia del Cerro Azul, Sierra del Espiritu Santo, Departamento de Coptin, Honduras, 1200 m elevation. . . . The locality is a t approximately 15"07'38.2"N, 88"56'13.8W." Males to 27.3 mm, females to 39.8 mm. Western slopes of Montaiia del Cerro Azul, Sierra del Espiritu Santo, Departamento de Coptin, Honduras, 900-1200 m elevation.

Craugastor daryi (FORD & SAVAGE 19841, Occas. Pap. Mus. Nat. Hist. Univ. Kansas 110: 1; type locality: "3.8 km (by road) SE PurulhB, Baja Verapaz, Guatemala, 1585 m." Males to 36 mm, females to 46 mm. Sierra Xucaneb and Sierra de las Minas, central Guatemala, 1500-2380 m elevation.

Craugastor emcelae (LYNCH 1985a1, Herpetologica 41: 443; type locality: "Rio Claro near its junction with Rio Changena, Bocas del Toro, Panama, 910 m." Males to 46.4 mm, females to 70.5 mm. Western central cordillera of Panama, 910-1450 m elevation.

dig. 363. Craugastor daryi (3.8 k m SE PurulhB, Baja Verapaz, Guatemala).


Fig. 364. Craugastor emcelae (north slope Cerro Pando, Bocas del Toro, Panama, 1350 m). Photo: W. E. Duellman (Courtesy of the Biodiversity Institute, University of Kansas)

Fig. 365. Craugastor chrysozetetes (Quebrada Fig. 369. Craugastor charadra (near Teguci de Oro, Atlantida, Honduras). pita, Honduras, 40 m). Photo: G. KO-----


Fig. 366. Craugastor coffeus (holotype). Fig. 370. Craugastor chingopetaca (Rio San Photo: G. Kohler Juan, Nicaragua, 30 m). Photo: J. Sunyer

I

Fig. 367. Craugastor crassidigitus (Guayach, Fig. 371. Craugastor emleni (Los Golondrinos, CRARC, Costa Rica, 550 m). Photo: G. Kiihler Francisco Morazh, Honduras).


Fig. 368. Craugastor cyanochthebius (San ,Fig. 372. Craugastor epochthidius (Quebrada Isidro, Coph, Honduras). Photo: J. R. McCranie de Las Man'as, Olancho, Honduras).

Photo: J. R. McCranie 163

Craugastor escoces (SAVAGE 19751, Copeia 1975: 280; type locality: "Costa Rica: Prwincia de Heredia: Cant611 de San Rafael: south slope of Volcan Barba: E margin Alto de Roble, 0.5 km N Cerro Chompipe, nr. Rio Las Vueltas: 2030 m." Males to 46 mm, females to 72 mm. Volcanoes Barva, I r d , and Turrialba, Costa Rica, 1100-2100 m elevation.

Craugastor evaneseo RYAN, SAVAGE, LIPS & GIERMAKOWSKI 2010, Copeia 2010: 405; type locality: "Panama, Cocl6 Province, Parque ~acional G. D. 0mar Tonijos H. (El CO&, headwaters of Rio Guabal. 8"39.59'N.

I -. . - . . . -

80°35.33W, 769 m.a.s.1. elevation." Males to Fig. 373. Craugastor escoces (Bajo La 50.2 mm, females to 64.3 mm. Eastern portion Hondura, San Jos6, Costa Rica). of the Serrania de TabasarB, west-central

photo: W. van Devender Panama, 80-709 m elevation.

Craugastor fecundus (MCCRANIE & WILSON 1997a), Alytes 14: 162; type locality: "Quebrada de Oro (15"38'N, 86'477171, elevation 880 m, tributary of Rio Viejo, south slope of Cerro Baalo S of La Ceiba, Cordillera Nombre de Dioa, Departamento de Atlhtida, Honduras." Males to 23.5 mm, females to 37.3 mm. Quebrada de Oro and Cerro Calentura in the Cordillera Nombre de Dios in the Departments of Atlhtida and Colbn, northern Honduras, at elevations of 200-1260 m elevation.

Craugastor fitzingen (SCHMIDT 18571, Sitzungsber. Akad. Wiss. Wien Phys. Math. Naturwiss. K1. 24: 12; type locality:

Fig. 374. Crauga--- - ---:o (holotype). "Neugranada"; SAVAGE'S (1974) designation of Photo: M. Ryan LACM 76859, from Barro Colorado Island,

Panama Canal Zone, as neotype was considered invalid by LYNCH & MYERS (1983). Males to 35 mm, females to 53 mm. Northeastern Honduras to northwestern Colombia, near sea

Craugastor emled ( D m 1932), Copeia 1932: level to 1520 m elevation.

97; type locality: "San Juancito, in the cloud Craugastor fleischmanni (BOETTGER 18921, forest at 6600 feet and ... at 6000 feet 11829 &t. Batr, Samml. Mus. Senckenb. m]," Departamento Francisco Morazh, Naturforsch. Ges.: 27; type locality: "San Jod, Honduras. Males to 28.1 mm, females to 49.8 [Cantbn de San Jos6, Provincia San Jose,] mm. On the Pacigc versant of south-central Costa Eta." Males to 47 mm, females to 75 Honduras CRio Choluteca drainage system mm. On the Pacific slopes of the volcanoes between Tegucigalpa and about El Zamorano, Po& and Barva, and on the Atlantic slopes of Departamento Francisco Morazh), 800-2000 the volcanoes Irma and Turrialba, and on both m elevation. slopes of the western portion of the Cordillera

de Talamanca, Costa Rica, 1050-2286 m eleva- Craugastor epochthidius (MCCRANIE & WILSON tion. 1997a), Alytes 14: 169; type locality: YRio Seco (14"55'N, 85'56'W), elevation 1050 m, Ixibu- Craugastor glaueus (LYNCH 1967a1, Trans. tary of Rio Guayape N of Catacamas, Sierra de K m a s Acad. Sci. 70: 177; type locality: "1.6 Agalta, Departamento de Olancho, Honduras." km SW San Cristobal [de las Casas], Chiapas, Males to 27.4 mm, females to 36.7 mm. Sierra M6xico,2100 m." Males to 31.7 mm (maximum de Agalta and Sierra Punta Piedra in the sbe for females not available). Only known departments of Olancho and ColBn, east-cen- h m the type locality. tral and northeastern Honduras, 150-1459 m elevation.

Fig. 375. Craugastor fecundus (Quebrada de Fig. 379. Craugastorglaurms (San Crld6bal de Oro, Atlhtida, Honduras). Photo: J. R. McCranie las Casas, Chiapas, Mexico). Photo: T. Bille

Fig. 376. Craugastor fitzingen (Lagarto Lodge, r'ig. mu. craugastor golunen (nusagami, Alajuela, Costa Rica, 90m). Photo: G. Kijhler Kuna Yala, Panama, 350 m). Photo: G. Kohler

Fig. 377. Craugastor fitzingen' (Lagarto Lodge, Fig. 381. Craugastor greggi (Volch Tacand, Alajuela, Costa Rica, 90m). Photo: G. Kohler Chiapas, Mexico). Photo: L. Gray

1 Kg. 378. Craugastor fitzingeri (near Uvita, Fig. 382. Craugastor gulosus (Cerro Frio, P ~ t a r e n a s , Costa Rica, 10 m). Photo: G. Kohler Bocas del Toro, Panamd). Photo: E. Boza

Craugastor gollmeri (PETERS 1863a1, Monatsber. Preuss. Akad. Wiss. Berlin 1863: 409; type locality: "Caracas"; in error, according to RIVERO (1961); see SAVAGE (1987) who discussed reasons for correcting the type locality to 'Veragua" (= western Panama). Males to 37 mm, females to 57 mm. On the Atlantic versant from northern Costa Rica to eastern-central Panama, near sea level to 1520 m elevation.

Craugastor greggi (BUMZAHEM 19551, Copeia 1955: 118; type locality: "Volcan Tajumulco, near San Marcos, Guatemala." Males to 36 mm, females to 46 mm. Soconusco region and slopes of V o l h Tacand, Chiapas, Mexico, and adjacent western Guatemala a t Volch Tajumulco, 1500-2700 m elevation.

Craugastor gulosus (COPE 18761, J. Acad. Nat. Sci. Philadelphia Ser. 2, 8: 112; type locality: "on a spur of the Pico Blanco, a t 6000 feet elevation", Costa Rica; in error for "Cerro Utyum, 6000 ft, Canthn de Talamanca, Provincia de Lim6n: 1829 mn, according to SAVAGE (1974). Females to 103 mm (adult males unknown). Southern Cordillera de Talamanca of Costa Rita-and western Panama, 1000-1870 m elevation.

Craugastor inachus (CAMPBELL & SAVAGE 2000), Herpetol. Monogr. 14: 232; type locality: "Quebrada Los Chorros, 650 m, Aldea Santa Elena, new San Diego, [Departamento] Zacapa, Guatemala. This locality is located in the Rio Motagua Valley in premontane Dry Forest at about 14"501N, 89'4TW." Females to 60.2 mm (single known male 25.3 mm SVL). Middle Motagua Valley and associated tributaries in east-central Guatemala, and the slopes of the Salam6 Bash in Baja Verapaz, Guatemala, 500-1400 m elevation.

Craugastor jota (LYNCFI 1980a), Trans. Kansas Acad. Sci. 83: 101; type locality: "along &e Rio Changena, Provincia Bocas del Toro, Panamd, 830 m." Females to 28.5 mm (males unknown). Atlantic slopes of the Cordillera de Talamanca in western Panama, 830-1820 m elevation.

Craugastor laevissirnus (WERNER 1896), Verh. Zool. Bot. Ges. Wien 46: 349; type locality: "Hondurasn; considered likely to have come &om the "vicinity of San Pedro Sula, the major town in northwestern Honduras" CMCCRANIE & W ~ O N 2002). Males to 31.8 mm,'females to 60.9 mm. Atlantic and Pacific slopes of Honduras to northern and southwestern Nicaragua, sea level to 2000 m elevation.

Craugastor laticeps ( D m 18531, AM. Sci. Nat. Paris, Ser. 3, 19: 178; type locality: "Yucatan (AmBrique centrale)", Mexico. Males

to 34.8 mm, females to 67.7 mm. On the Atlantic versant from southern Veracruz to Tabasco and Chiapas, Mexico, and southward to western Belize, Guatemala, and northern Honduras, from near sea level to 1600 m elevation.

Craugastor lauraster (SAVAGE, MCCRANIE & ESPINAL 1996), Proc. Biol. Soc. Washington 109: 366; type locality: "above the Quebrada El Piiiol (15"07'N, 86'43W1, Parque Nacional La Muralla, Departamento de Olancho, Honduras, elevation 1200 m," Males to 17.2 mm, females to 22.3 mm. On the Atlantic versant .from central and eastern Honduras to northerncenM Nicaragua, 40-1300 m elevation.

Craugastor lineatus (BROCCHI 1879), Bull. Soc. Philomath., Paris, Ser. 7, 3: 22; type locality: "Attitlan (Mexique)" [= slopes of Volch Atitlh, Guatemala]. Males to 32 mm, females to 47 mm. On the Pacific versant from Guerrero, Mexico, to the southwestern highlands of Guatemala, 300-2000 m elevation.

Craugastor loki (SHANNON & WEItLER 1955), Trans. Kansas Acad. Sci. 58: 370; type locality: "at an altitude of 3500 feet on Volch San Martin, Veracruz", Mexico. SVL to 32.1 mm. On the Atlantic versant from San Luis Potosi and Veracruz, Mexico, to Belize and northwestern Honduras; crossing the Isthmus of Tehuantepec and thence southeast along the Chiapas and Guatemalan versant to El Salvador (LYNCE 20001, sea level to 2100 m elevation.

Craugastor lo&rostris (BOULENGER 1898b), Proc. Zool. Soc. London 1898: 120; type lo&- ty: "Cachab6 [=Ca&abfl ... a small village on the river of that name, on the N.W. Coast, in the Prov. Esmeraldas. ... probably about 500 feet above the sean, Ecuador. LYNCH & MYERS (1983) discussed this locality and noted that it is not the same rl8 San Javier de Cachabi. Males to 39.2 mm, females to 59.6 mm. Extreme eastem Panama (Dari6n Province), throughout western Colombia to the southern

Fig. 383. Craugastor inachus @om near type locality). Photo: J. k Campbell

Fig. 384. Craugastor jota (Rio Changena, Bocas Fig. 388. Craugastor lineatus (Santa Barbara, del Toro, Panama, 1765 m). Photo: A. Hertz S slope of Volcfin Atit lb, Suchitepbquez,

Guatemala, Guatemala). Photo: G. Kohler

IF! I Fig. 385. Craugastor laevissimus (P.N Aff --eico Norte, Nicaragua, 780 m). intla, Guatemala, 300 m). Photo: G. Kohler

Photo: G. Kohler

Izabal, Guatemala). Fig. 386. Craugastor laticeps (Cerro San Gil, Fig. 390. Craugastor longirostris (Cana, DariBn,

Photo: G. Kohler Panama, 1597 m). Photo: A. J. Crawford

ng . 38.1. craugastor lauraster (P.N. Saslaya, Fig. 391. Craugastor megacephalus (Cerro El Atlantic0 Norte, Nicaragua). Photo: G. Kohler Toro, Nicaragua, 830 m). Photo: G. Kohler

part of Guayas Province in Ecuador, sea level the slope of Chiriqui volcano above Boquete, to 1200 m elevation. ChiriauIi Province". Panama. SVL to 65.7 mm.

Craugastor matudai (TAYLOR 1941), Univ. Kansas Sci. Bull. 27: 154; type locality: %It. Ovando, Chiapas", Mexico. Males to 50 mm, females to 65 mm. On the Pacific versant in southern Chiapas (Cerro Ovando), Mexico, and aqjacent Guatemala, 1500-2000 m elevation.

Craugastor megacephalus (COPE 18761, J. Acad. Nat. Sei. Philadelphia Ser. 2,8: 110; type locality: ''Pico Blanco, at 6000 feet elevation", Costa Rica; corrected to "Cerro Utyum, 6000 R, Cant& de Talamanca, Provincia de Lim6n: 1829 m", Costa Rica by SAVAGE (1974). Males to 43 mm, females to 70 mm. Atlantic versant h m extreme southeastern Honduras to western Panama (including the Bocas del Toro islands), sea level to 1230 m elevation.

Craugastor melanostictus (COPE 18761, J. Acad. Nat. Sci. Philadelphia Ser. 2,8: 109; type locality: "7000 feet elevation on the Pico Blanco", Costa Rica; corrected to "Cerro Utyum, 7000 ft, Cant611 de Talamanca, Provincia de Lim6n: 1829 m", Costa Rica by SAVAGE (1974). Males to 43 mm, females to 56 mm. Cordilleras of Costa Rica and westem and central Panama, 1150-2700 m elevation.

Craugastor merendonensis ( S c m 19331, Field Mus. Nat. Hist. Publ., Zool. Ser., 20: 17; type locality: 'lower Santa Ana Canyon, west of San Pedro [= San Pedro Sula, Departamento de Cortds], Honduras. Altitude 500 feet." Males to 48 mm, females to 83 mm. Montaflas del Merend6n on the Sierra de Omoa, west of San Pedro Sula in the Department of Corths, Honduras, 150-200 m elevation.

Craugastor milesi (Scwimrr 1933), Field Mus. Nat. Hist. Publ., Zool. Ser., 20: 18; type locality: "Mountains west of San Pedro [= San Pedro Sula, Departamento de C o ~ s l , Honduras. Altitude 4500 feet." Males to 24.8 mm, females to 37.1 mm. Western and northwestern Honduras (Cerro Azul in Departamento de Cop&, and from MonMas del Cusuco and Merend6n in the Departmento de Cort&s), 1050-1720 m elevation.

Craugastor mimus (TAYLOR 19551, Univ. Kansas Sci. Bull. 37: 517; type locality: "5 km. NNE Tilarh, [Cantbn de Tilarh,] Guanacaste Province, Costa Rica." Males to 37 mm, females to 58 mm. On the Atlantic versant from eastern Honduras to southern Costa Rica, sea level to 1330 m elevation.

Craugastor monnichonun (DUNN 19403, Proc. Acad. Nat. Sci. Philadelphia 92: 107; type locality: "Valley of the Velo, Finca Lerida, on

west& cordillera de Talamanca of Panama, 610-1830 m elevation.

Craugastor montanus (TAYLOE 1942), Univ. Kansas Sci. Bull., 28: 67; type locality: "Mount Ovando, Chiapas, 6,000 ft. elevation," Mexico. SVL to 27 mm. Cerro Ovando and surrounding areas, in the Sierra Madre de Chiapas, Mexico, 1200-1990 m elevation. Eleutherodactylu~ sartori LYNCH 1965a is regarded as a synonym of C. montanus according to HEDGES et al. (2008).

Craugastor myllomyllon (SAVAGE 20001, Rev. Biol. Tropical 48: 993; type locality: "Finca Volcan, Sierra de Xucaneb, Departamento de Alta Verapaz, Guatemala, 875 m." Known only from a single immature female (SVL 30.1 mm). Only known from the type locality.

Craugastor nefiens (SMITE 20051, Herpeto- logica 61: 287; type locality: "La Firmeza, Sierra de Caral, Municipio de Morales; Departamento de Izabal, Guatemala, 840 m. Males to 26.8 mm, single known female 25.8 mm. Sierra de Caral, southeastern Guate- mala, 800-1000 m elevation.

Craugastor noblei (BARBOUR & DUNN 1921), Proc. Biol. Soc. Washington 34: 161; type locality: "Guapiles, [Cantdn Pococi, Provincia Limbn,] Costa Rica." Males to 48 mm, females to 66 mm. On the Atlantic versant h m northeastern Honduras to western Panama, also on the Pacific versant in Costa Rica and west-central Panama, sea level to 1330 m elevation.

Craugastor obesus (BARBOUR 19281, Proc. New England Zool. Club 10: 27; type locali* "Gutierrez, Bocas del Toro Province, Panama (near the Costa Rican hntier)." Males to 56.2 mm, females to 86.3 mm. On the Atlantic versant of southern Costa Rica and western Panama, 400-1450 m elevation.

Fig. 392. Craugastor melanostictus (Pico Picacho, Chiriqui, Panama). Photo: G. Kohler

It R8.393. Crau~astor milesi (Y.N. I;usuco. PIE. 391. t i rau~as tor netrens (rmca ~a Photo: Kolby

- Firmeza, ~ o r i e s , Izabal, Guatemala).

Photo: C. R. Vhsquez Almazhn

Fig. 394. Craugastor mimus (Lagarto Lodge, Fig. 398. Craugastor noblei (Selva Negra, Mata- Alajuela, Costa Rica, 90 m). Photo: G. Kohler galpa, Nicaragua, 1300 m). Photo: G. Kohler

m I b 395. Craugastormonnichom (Ate Cbique- &g. YYY. tiraugastor oDesus ( ~ o ~ q w ,

,, ,Wqui, Panama, 1820 m). Photo: G. Kijhler Panama, 1050 m). Photo: K-H. Jungfer

Fig. 396. Craugastor montanus (Volchn Tacanh, Fig. 400. Craugastor olanchano (Quebrada El 1

Chiapas, Mexico, 900 m).Photo: k Ramirez Velslzquez Pinol, Olancho, Honduras). Photo: J. R. McCranie

169

Craugastor olanchano (MCCRANIE & WILSON 1999b), Proc. Biol. Soc. Washington 112: 518; type locality: "hillside above the Quebrada El Pin01 (15"07'N, 86"44W), Parque Nacional La Muralla, Departamento de Olancho, Honduras, 1200 m elev." Males to 29.8 mm, females unknown. On the Atlantic versant in the northwestern portion of the Department of Olancho, north-central Honduras. 1180-1350 m elevation.

Craugastor omoaensis (MCCRANIE & WILSON 1997a), Alytes 14: 155; type locality: "about 10 Fig. 401. Craugastor opimus (Altos de Cerro airline km WSW San Pedro Sula on road to Azul, PanamB, Panama, 820 m). Pen3 (15"28'N, 88"06W), elevation 1150 m, Photo: A. J. Crawford Sierra de Omoa, Departamento de Cort6s, Honduras." Males to 30.0 mm, females to 38.4 mm. Sierra de Omoa WSW of San Pedro Sula on the Atlantic versant of northwestern Honduras, 760-1150 m elevation.

Craugastor opimus (SAVAGE & MYERS 2002), American Mus. Novit. 3357: 34; type locality: "Quebrada Docord6, about 10 km above junction with Rio San Juan, about 100 m elev., Depto. Choc6, Colombia (approx. 4'34'N, 77"03W)." Females to 69 mrn (adult males unkriown). Central Panama to northwestern Colombia, near sea level to 1040 m elevation.

Fig. 402. Craugastor pechonun (Quebrada Craugmtor paIenque (CAMPBELL & SAVAGE Machin, Col6n, Honduras). Photo: J. R. McCranie 20001, Herpetol. Monogr. 14: 221; type locality: "10.7 km SE Cristbbal Col6n on road to Bonampak, ca. 350 m, Chiapas, Mexico. This drain into the lower Rio Grijalva and K O site lies in Bemontane Wet Forest at about Usumacinta, 350-1700 m elevation. 17"08'N, 91°34W." Males to 43.1 mm, females to ~ 5 . ~ mm. c&bbean slopes of northern Craugastorpersimilis (BARBOUR 1926), Occas. Chiapas, xi^^, and adjacent ~ ~ ~ ~ ~ ~ ~ l ~ , Pap. Boston Soc. Nat. Hist. 5: 193; type locali- where it is horn eom tributaries that ty: "Suretka, Costa Rica, near the boundary of

drain into the lower mo Grijalva, 200400 Bocas del Toro Province, Panamd." Males to

elevation. 17.8 mm, females to 23 mm. On the Atlantic versant in central to southern Costa Rica,

Craugastor pechorum (MCCRANIE & WILSON 40-1200 m elevation. 1999b1, Proc. Biol. Soc. Washington 112: 515; type locality: "from near a dam along a Craugastor phasma (LIPS & SAVAGE 1996), small tributary of the Quebrada de Las Marfa8 Roc. Bid. Sot. Washington 109: 7 e type (15017.64'N, 84021.29~), about 12 airline km locality: "Costa Rica: Puntarenas Province, - L~ coloda, D~~~~~~~~~~ de 0lancho, Cmt6n Coto Brus, Zona Protectors Las ~ ~ ~ d ~ ~ ~ ~ , 680 rn elev." ~ a l ~ ~ to 42.5 -, Tabla, Finca Jaguar, 8"55N 82"44W on the females to 62.1 mm. On the Atlantic versant of 20 km - of La Lucha, the and eastern portions of the elevation." Single known specimen 47.9 mm Honduran departments of Olancho and Colbn, Known from the type locality. respectively, i50-680 m elevation. Craugastor podiciferus (COPE 1876), J. Acad. craugastor pelorus (C-BEL~ & sAVAGE Nat. Sci. Philadelphia Ser. 2,s: 107; type local- 2000), Herpetol. 14: 218; type locality: ity: "from the level of 5000 to 7000 feet ... on "2.7 km N ~ ~ l ~ ~ ~ ~ h i ~ ~ ~ , ca. 400 m, chiapas, the Pico Blanco", Costa Rica; corrected to ~ ~ ~ i ~ ~ , hi^ site lies in premontane wet "Cerro Utyum, 5000-7000 ft, Cant6n de F~~~~~ at about 17027'N, 93003v." ~ a l ~ ~ to Talamanca, Provincia de Limbn: 1524-2134 56.8 =, females to l lO.~ mm. caribbean m", Costa Rica by SAVAGE (1974). Males to 28 slopes of northern Chiapas, Mexico, where it is to 40 Both 'lopes of the known from the upper tributaries that cordilleras of Costa Rica and adjacent western

Panama, 1089-2650 m elevation.

I

Fig. 403. Craugastor persirnilis (Guayacan, Fig. 407. Craugastor podiciferus (Guacuyo, CRARC, Costa Rica, 550 m). Photo: G. Kohler Heredia, Costa Rica, 1850 m). Photo: G. Kohler

Fig. 404. Craugastor podiciferus (Alto Chiquero, Fig. 408. Craugastor podicifem Paso Llano, Chiriqui, Panama, 1820 m). Photo: G. Kiihler Heredia, Costa Rica, 1700 m). Photo: G. Kohler

Fig. 405. Craugastorpodicifem (Volch Barva, Fig. 409. Craugastor polyptychus (P.N. Manza Heredia, Costa Rica, 2000 m). Photo: G. Kiihler nillo, Lim6n, Costa Rica, 30 m). Photo: G. Kijhler

Fig. 404. Craugastorpodzciferus (Alto Chiquero, Fig. 410. Craugastor polyptychus (same as in Chiriqui, Panama, 1820 m). Photo: G. Kiihler Fig. 409). Photo: G. Kohlt

Craugastor polyptychw (COPE 1886), Proc. Am. Philos. Soc. 23: 276; type locality: "Nicaragua." Males to 26.6 mm, females to 28.7 mm. Atlantic slope of southeastern Nicaragua to southeastern Costa Rica and adjacent northwestern Panama, sea level to 260 m elevation.

Craugastor pozo (JORNSON & SAVAGE 19951, J Herpetol. 29: 501; type locality: "12 km NW o- BerriozAbal (elev. ca. 1060 m), Municipality of BerriozAbal, Chiapas, M6xico (ca. 16"53'N lat. 93'23W long.)." Males to 48.3 mm, females tc 80.8 mm. Western foothills and highlands of Chiapas (in an area bordered by Cintalapa, Berriozabal and Presa Nezahualcoyotl), Mexico, 500-1200 m elevation.

Craugastor psephosypharus (CAMPBELL, SAVAGE & MEYER 19941, Herpetologica 50: 413; type locality: '7.8 km WSW Puerto Santo Tom&, 375 m, northeastern slope Montaib~ del Mico, Departamento de Izabal, Guatemala . . . This locality is located on the eastern slopes of the Montaiias del Mico at about 15a41'N, 88O40W. The town of Santo Tomas is called Puerto Santo Tomb de Castilla or Puerto Matias de Gglvez on some maps." Males to 36.3 mm, females to 65.3 mm. Central and eastern Guatemala, and adjacent southern Belize, 150-1170 m elevation.

Craugastor punctariolus (PETERS 1863b), Monatsber. Preuss. Akad. Wiss. Berlin 1863: 462; type locality: "Veraguan, Panama. Males to 50 mm, females to 81 mm. Western and central cordilleras of Panama, 560-1800 m elevation.

Craugastor pygmaew (TAYLOR 1937a), Trans. Kansas Acad. Sci. 39: 352; type locality: '1 mile north of Rodriguez Clara, Vera Cruz", Mexico. SVL to 18 mm. Extreme southern Michoach and M6xico (state), Mkco, south and east along the Pacific versant to western Guatemala; also in Veracruz and eastern Oaxaca, Mexico, on the Atlantic slope; sea level to 2000 m elevation.

Craugastor raniformis (BOULENGER 18961, Ann. Mag. Nat. Hist. Ser. 6,17: 19; type locality: "Buenaventura and Cali", Departamento Valle del Cauca,] Colombia. Males to 43.2 mm, females to 74.0 mm. East-central Panama to western Colombia, near sea level to 1500 m elevation.

Craugastor ranoides (COPE 1886a), Proc. Am. Philos. Soc. 23: 275; type locality: "Nicaragua." Males to 45 mm, females to 74 mm. Eastern Nicaragua on the Atlantic slope and northwestern Costa Rica on the Pacific versant to extreme western Panama (including Isla

Fig. 411. Craugastorpozo (12.5 kilometers North of Berriozabal, Chipas, Mexico, 1100 m).

Photo: T. J. Papenfuss

Fig. 412. Craugastor psephosypharus (11.8 km WSW Puerto Santo Tom& Izabal, Guatemala).

Photo: J. A. Campbell (Courtesy of the Biodiversity Institute, University of Kansas)

Escudo de Veraguas), exclusive of the Golfo Dulce region of southwestern Costa Rica, sea level to 1300 m elevation.

Craugastor ray0 (SAVAGE & DEWEESE 1979), Bull. S. California Acad. Sci. 78: 107; type locality: "second sabana on the trail from Finca El Helechales to Sabanas Esperanza, 5 km, airline, east of Finca El Helechales, Cant611 de Buenos Aires, Provincia Puntarenas, Costa Rica, 1640 m elevation." Males to 45 mm, females to 71 mm. Pacitic slope of the Cordillera de Talamanca of southern Costa Rica, 1480-1820 m elevation.

Craugastor rhodopis (COPE 18671, Proc. Acad. Nat. Sei. Philadelphia 18: 323; type locality: "Vera Cruz, at Orizava [= Orizabal and Cordova I= Cbrdobal", Veracruz, Mexico; restricted to Orizaba, Vera-, Mexico, by lectotype designation (SMITH & TAYLOR 1950). Males to 28.5 mm, females to 45.7 mm. Western V e r a m and adjacent Hidalgo and Puebla in Mexico; also in central and southeastern Chiapas and adjacent Oaxaca, mostly 100-1000 m elevation.

! FSg. 413. Craugasiarpunotbuiolus (El Valle de Fig. 416. Craugastor ranaides (5 lun W Anttin, C d d , Panama, 800 m). Photo: M. Ryan Ahhnte , B o w del Tom, Panam3.

Photo: W. E. Due- (Couftess ofthe B i o d i p e Instihb, U n i U m of ~ ~ S B B )

Fig. 414. Craugastor pygmaats (30 km N San fig. 417. Cmugwtor rh~dopis (rma Rancho ff

Gabriel Itfixtap, Oaxaca, Ikhdco). Alegre* Ghiapaa, Msxioo, ilOO m). Photo: W. E. Duehan Phob: G. &@or I

- ~ g . 415. Craugmtor radio& (Chepigam Fig. 418. Cmugastor dpulw. Dmidn, Panama, l322;m). Photo: M. Ponce Photo: J. A. Campbell

Craugastor rhyacobatrachus (CAMPBELL & SAVAGE 2000), Herpetol. Monogr. 14: 274; type locality: "Quebrada Fortuna, 19 km N San Isidro de El General, 1700 m, Cant6n Perez- Zeledbn, Provincia de San Jose, Costa Rica. This locality is in Tropical Premontane Rainforest at approximately 9"28'25"N, 83O41'42W." Males to 50.0 mm, females to 80.9 mm. Southern slopes of the Cordillera Talamanca-Bani of Costa Rica and western Panama, 950-1800 m e1evation.L

Craugastor rivulus (CAMPBELL & SAVAGE 2000), Herpetol. Monogr. 14: 224; type locality: "Samac-Cham& Trail on Finca Samac, Alta Verapaz, Guatemala. This site lies in the "coffee belt" at about 1250 m." Males to 39.2 mm, females to 63.9 mm. Highlands of northern Guatemala, including the mountains of Alta Verapaz and the eastern portion of the Sierra de Los Cuchumatanaes in Quich6,770-1250 m elevation.

Craugastor rostralis (WERNER 1896), Verh. Zool. Bot. Ges. Wien 46: 350; type locality: "Honduras." Males to 35.7 mm, females to 59.0 m. Extreme eastern Guatemala to western and north-central Honduras, 850-1800 m ele-

Craugastor saltuarius (MCCRANIE & WILSON 1997a), Alytes 14: 165; type locality: "south slope of Cerro Blifalo (15'39' N, 86"48W), elevation 1550 m, Cordillera Nombre de Dios, Departamento de Atlhtica, Honduras." Males to 22.4 mm, single known female 17.8 mm. Western and central portions of the Cordillera Nombre de Dios on the Atlantic versant of northern Honduras. 1550-1800 m elevation.

Craugastor sandersoni (SCHMIDT 19411, Field Mus. Nat. Hist. Publ., Zool. Ser., 22: 485; type locality: "Double Falls, west of Stann Creek, British Honduras [= Belize]." Males to 50.2 mm, females to 85.2 mm. Caribbean slopes of the Maya Mountains in east-central Belize southward to the Montaiias del Mico in eastern Guatemala and westward into the Sierra de Santa Cruz, the eastern portion of the Sierra de las Minas, and the foothills of the northern Alta Verapaz, sea level to 1160 m elevation.

vation. m Craugastor rugosus (PETERS 1873), Monatsber. Preuss. Akad. Wiss. Berlin 1873: 610; type locality: "Chiriqui", Panama. Males to 44 mm, females to 69 k. Paciiic versant of central and southern Costa Rica and adjacent Panama, sea level to 1220 m elevation.

Craugastor rupinius (CAIWBELL & SAVAGE 2000), Herpetol. Monogr. 14: 213; type locality: "Finca El Faro, S slope Volch Santa Maria, 4.0 km N El Palmar, 875 m, [Departamento] Quezaltenango, Guatemala. This site lies in Premontane Wet Forest at about 14"40'30nN, 9lo35'00W." Males to 40.3 mm, females to 72.1 mm. Pacific versant of southeastern Chiapas, Mexico, southeastward through southern Guatemala to eastern El Salvador, 200-2000 m elevation.

Craugastor sabrinus (CAMPBELL & SAVAGE 2000), Herpetol. Monogr. 14: 239; type locality: "Las Escobas, 5.1 km WSW Puerta Santo Tom& 150 m, Monteas del Mico, Izabal, Guatemala. This locality is located on the NE slopes of Cerro las Escobas in Lowland Wet Forest at about 15"38'N, 88"41W." Males to 43.6 mm, females to 82.6 mm. Foothills of eastern Guatemala, including the Montaiias del Mico, the eastern portion of the Sierra de las Minas, and the northern portion of the Sierra de Merend6n; also known from the Maya Mountains of Belize, near sea level to about 900 m elevation.

Craugastor silvicola (LYNCH 1967b), Proc. Biol. 1 Soc. Washington 80: 211; type locality: "12 mi. NNE Zanatepec, Oaxaca, Mkxico, 4900 ft. ele-

r vation." SVL 40.2 in single known female. 1 Only known from the type locality, 1450-1600 m elevation. 3 1

i 1 i

Fig. 419. Craugastor rugosus (Refugio El Rodeo, Costa Rica, 995 m). Photo: G. Kohler

I I L Fig. 420. Craugastor rostralis (P.N. Cusuco, ! Cartes, Honduras, 660 m). Photo: J. Kolby

m

Fig. 421. Craugastor rupinius (El Imposible, Fig. 425. C s a u g a ~ t o ~ stadRZmd ( I airline b Ahuachaph, El Salvador, 720 m). NNE of La Fofttma, Yoro, Honduras).

Photo: G. Kiihler Photo: J. R. McCrmie

I -

Fig. 422. CPauga&r 5abau8 (Gem San GI1, Fig. 426. Craugastor+-w Wends Bani, Mal, Guatemda, 160 m). Photo: G. Kahle~ Puntarenas, Costa Rica, 20 m). Photo: G. Kijhler

I Fig. 423. Craugastor saltuarius (south slope of Fig. 427. Craugastor stuarti (Jaltenango E l Cerro Blifalo, Atlhtida, Honduras). Triunfo], Chiapas, Mexico, 1960 m).

Photo: J. R. McCranie Photo: A. Ramirez Veldzquez

Fig. 424. Craugastor sandersoni (Sierra de Fig. 428. Craugastor tabasarae (La Nevera, Chinajd, Alta Verapaz, Guatemala, 615 m). Ngobe Bugl6, Panama). Photo: G. Kohler

Photo: J. Sunyer

Craugastor stadelmani (Sctmamr 193&BJ, Proc. Biol. Soc. Washington 49: 44; type locality: "Portillo Grande, [Departamento del Yoro, Honduras, at 4800 feet altitude." Males to 33.1 mm, females to 47.4 mm. Western portion of the Cordillera Nombre de Dios southward to Montaiia de Pijol, Yoro, and eastward to northwestern Olancho on the Atlantic versant of northern Honduras. 1125-1900 m elevation. I Craugastor stejnegerianus (COPE 18931, Proc. Am. Philos. Soc. 31: 338; type locality: "Palmar"* Provincia Puntarenas, Costa @& Fig. 430. Craugastor t a m s (Las Mellizas, Males to 18.0 mm, females to 22.4 mm. Pacific Punts Burica, Chiriqui, Panama, 120 m). versant of Costa Rica and western Panama, sea level to 1330 m elevation. Photo: M. Ponce

Craugastor stuarti (LYNCH 1967a). Trans. ~ a n s & Acad. Sci. 70: 180; type locality: "Aldea Paraiso, 13 km. S. La Mesilla (on Guatemala-M6xico border), Depto. Huehuetenango, Guatemala, 2200 m." Males to 30.5 mm, females to 40.5 mm. Departamentos de San Marcos and Solola, Guatemala, and adjacent eastern Chiapas, Mecico, 1300-2300 m elevation.

f?raugastor tabasarae (SAVAGE, HOLLINGS- WORTH, LIPS & JASMW 20041, Herpetologica 60: 521; type locality: "9.7 km NNW El Cop6, on the continental divide at sawmill, La Pintada District, Cocl6 Province, Panama, 600400 m (8"40'04"N, 8Oo35'6"W)." Males to 33.7 mm, females to 54.2 mm. West-central and central Panama, 600-910 m elevation.

Craugastor talamancae ( D m 19311, Occas. Pap. Boston Soc. Nat. Hist. 5: 385; type locality: Wrnirante, Bocas del Tom, Panama." Males to 30 mm, females to 50 mm. On the Atlantic versant from Nicaragua to eastern Panama, 15-646 m elevation.

Craugastor taurus (TAYLOR 1958b), Univ. Kansas Sci. Bull. 39: 17; type locality: "Golfito, [Cantbn de Golfito,] Puntarenas Province", Costa Rica. Males to 44 mm, females to 80 mm. Pacific versant of southern Costa Rica and adjacent western Panama, 25-525 m elevation.

Craugastor Caylori (LYNCH 19661, Trans. Kansas Acad. Sci. 69: 76; type locality: "Mexico, Chiapas, 6.2 km S Ray6n Mescalapa." SVL 26.0 in single known male specimen. Only

Fig. 431. Craugastor trachydermus (from type locality). Photo: J. A. Campbell

known from-the type locality. Fig. 432. Craugastor talamancae (Nusagandi, Craugastor trachydermus (CAMPBELL 1994b), Kuna Yala, Panama, 350 m). Photo: G. Kohler Herpetologica 50: 400; type locality: "Xiacam, Sierra de Santa Cruz, Departamento de Izabal, Guatemala, 900 m. ... This locality is located at approximately 15"33'N, 8go32W on the headwaters of the Rio Sauce." Males to

33.5 mm, females to 46.1 mm. Known only from the type locality

Craugastor underwoodi (BOULENGER 1896b1, Ann. Mag. Nat. Hist. Ser. 6,18: 340; type locality: "La Palma", Cantdn de Coronado, Provincia de San Joe&, Costa Rica, 1600 m elevation. Males to 26 mm, females to 30 mm. Cordilleras de Tilarh, Central, Costefia, and Talamanca, as well as western Panama, 920-1590 m elevation.

Craugastor wcanebi (STUART 19411, Proc. Biol. Soc. Washington 54: 199; type locality: "Cloud forest above F i c a Voldn (49 kilometers [straight line] east of Cobfin), Alta Verapaz, Guatemala; altitude about 1300 meters." Males to 32.8 mm, females to 46.0 mm. Central highlands of Guatemala, including the Sierra de Los Cuchumatanes, the Sierra de Xucaneb, and the Sierra de las Minas, 600-1300 m elevation.

Craugastor yucatanensis (LYNCH 1965131, Herpetologica 20: 249; type locality: "in a cave ... 1.5 km. S, 1 krn. E Pueblo Nuevo X-Can, Quintana Roo, Mexico, at 10 meters elevation." SVL to 36 mm in females (maximum size for males not available). Northeastern portion of the Yucatan Peninsula (east-central Yucatdn and north-central Quintana Roo), Mexico, near sea level to 30 m elevation.

L

C. crassidigitus C. chingopetaca

Craugastor

I C. fitzingen

Fig. 433. Craugastor xucanebi; a. from 3.8 km SE PurulhB, Baja Verapaz, Guatemala; b. from San Jose Maxbal, Barillas, Huehuetenango).

Photos: a. J. A. Campbell (Courtesy of the Biodiversity Institute, University of Kansas);

b. C. R. VBsquez Almazh

C. talarnancae

Craugastor

' 5ransfordii lauraster

Craugastor

2. xucanebi ,. rhodopis 2. chrysozetetes >. ornoaensis

8 C. coffeus

2- I L'.

yucatanensis C. nefrens

f :. glaucus .. taylori

A C. silvicola

:. loki :. rnyllomyllon

r C. rostralis A C. epochthidius

---

a C. saltuarius

r C. pygmaeus

8 C. stadelrnani r C. rnatudai A C. fecundus

2. melanostlctus ,. tabasarae S. monnichorum

Craugastor

S. erncelae S. longirostris

r C. cuaquero C. phasma

I C. rupintus C. sabrinus C. pelorus C. pozo C. emleni C. montanus

sandersoni I c. rivulus 1 C. amniscola % C. olanchano

C. merendonensis C. pechorum

C. Iaeviss~mus

C. inachus

Craugastor

V C. rhyacobafrachus C. chingopetaca

I I C. ranoides E C. escoces I C. punctario/us I -

V C. catalinae I

Further Reading SAVAGE 1975. SAVAGE & DEWEESE 1979, LYNCH 1980b, SAVAGE 1980, SAVAGE & DEWEESE 1981, MIYAMOTO 1983, Ford & SAVAGE 1984, MIYAMOTO & TENNANT 1984, LYNCH 1985a, M~PAMOTO 1986, SAVAGE 1987, SAVAGE et al. 1988, MCCRANIE et al. 1989, C A ~ B E L L 1994, CAMPBELL et al. 1994, JOHNSON & SAVAGE 1995, LYNCH 1996, SAVAGE et al. 1996, MCCRANIE & WILSON 1997a, KOHLER 1998, LYNCH 1998, DONNELLY 1999, HOBEL 1999a, M c C m & KOHLER 1999b, MCCRANIE & WILSON 199913, CAMPBELL & SAVAGE 2000, LYNCH 2000, SAVAGE 2000, MCCRANIE & WILSON 2002, SAVAGE & MYERS 2002, F'USCHENDORF & CRAWFORD 2003, SAVAGE et al. 2004, CRAWFORD & S m 2005, SMITH 2005, KOHLER & SUNYER 2006, HEDGES et al. 2008

Eleutherodactylidae

Eleutherodactylidae The recognition of the family Eleuthero- dactylidae containing two subfamilies (Eleutherodactylinae and Phyzela- phryninae), four genera, and a total of 201 species follows HEDGES et al. (2008) and FROST (2010). This family covers a wide geographic distribution ranging from central Texas (USA) and western Mexico across Central America, as well as widely in northern South ~Ger ica and on the Antilles.

F'urther Reading HEDGES et 81.2008

Fig. 433. Diasporus hylaeformis (Alto Chiquer Chiriqui, Panama, 1820 m). Photo: G. Kohl

Dias~orus

The genus Diasporus was erected by HEDGES et al. (2008) to accommodate nine species of the so-called dink frogs (FROST 2010); the common name refers to the characteristic "dink" or "tink" call the males of species in this genus produce. These small anurans are nocturnal, rather common forest inhabitants that dwell in primary as well as disturbed forests. They are more often heard than seen and can be quite common locally Occasionally, individuals of Diasporus can be observed walking on their stubby legs rather than jump- ing (Scorn 1983~). Food consists of a variety of small arthropods, including ants (TOW 1981). The males start calling shortly after dusk from sites in vegetation from near ground level to several meters up in the trees (WILCZYNSKI & BRENOWITZ 1988, Wncmsm et al. 1993). They continue calling throughout most of the night with peaks of calling activity between 10:OO pm and 2:00 am (SAVAGE 2002). For such a tiny frog, it is an impressively loud call, especially when heard at close range. During daytime, these nocturnal frogs can be found hiding in bromeliads and in leaf litter (LIEBERMAN 1986). Nesting sites of these frogs include bromeliads, leaf peti- oles, and other cavities (TAYLOR 1955, OVA~KA & RAND 2001). The development is

direct, and after little less than four weeks little frogs hatch out of the eggs (Scorn 1983c, OVASKA & RAND 2001).

Diasporus diastema (COPE 18761, J. Acad. Nat. Sci. Philadelphia Ser. 2, 8: 155; type 1ocalit;pr "camp Mary Cmtta , Panama." Males to 2%

n mm, females to 24 mm. Southeastern Honduras to eastern Panama; sea level td 1620 m elevation.

I Diasporu~i h y l a e f d (COW 1876), J. Acad Nat. Sci. Philadelphia Ser. 2,8: 107; type locality: "mountain of Pico Blanco, at 7000 feet elevation", ROV. Lim&n, Costa Rica; corrected to "Cerro Utpm, 7000 R, Cant6n de Tdamanca, Provincia de Lim6n: 2134 m", Costa Rica bigr SAVAGE (1974). Males to 22 mm, females to 26 mm. Cordilleras of Costa Rica and western Panama, 1580-2500 m elevation,

Diasporus quidditus (LYNCH 20011, Rev. Acad. Colomb. Cienc. Exact. Fis. Nat. 25: 294; type locality: "Colombia, Valle del Cauca, municipio Buenaventura, Estaci6n Forestal (de la Universidad de Tolima) 'Bajo Calima', 50 m, 3"59'N 76O57W." Males to 14.8 mm, females to 16.9 mm. Central Panama to northwestern Colombia, sea level to about 600 m elevation.

Diasporus tigrillo (SAVAGE 19971, Amphibia-Reptilia 18: 241; type locality: "Alto Lari a t the confluence of the Rfo Lari and Rio Dipari, about 21 km SW Amubri, Distrito Bratsi, Cant6n de Talama[n]ca, Provincia de L i d n , Costa Rica, (440 m)." Males to 18 mm,

Fig. 434. Diaspom ventrimaculatus (Valle del Silencio, Costa Rica). Photo: R. Vargas

Figs. 435-436. Diaspom ventrimaculatus (Valle del Silencio, Costa Rica). Photo: R. Vargas

Fig. 437. Diasporus diastema (Guayach, CRARC, Costa Rica, 550 m). Photo: G. Kohler

. Kn o localitie valley of the Rio Lmi, in Limb

Wvince, Costa Rica, 400-440 m elevation.

!@iasPoms ventrimaculatus CHAVES, GARCfP WDR~GUEZ, MOM & LEAC 2009, Zootaxa 2088 9; type locality: Valle del Silencio at the easz $&ge of the Cordillera de Talamanca 20 km I t e s t from the Costa Rica-Panama borde @.1116 N, -82.96172 W, 2550 m elevatinn) Males to 23.5 mm, females to 24.7 mm. Kr only from the type localiwdns.-

~Diasporus vqcator (T-' ) 2 ~ . ) 0 l , uw &msas Sci. Bull. 37: 522; type locality: "Agu

. Buena, [Cant611 de Golfito,] Puntarena IPr~vince, Costa Rica." Males to 16 mxx. bmales to 18 mm. On the Pacific versant fiom louthern Costa Rica to central Panama; i- central Panama also on the Atlantic versanl EL&w@@~ to 1220 m elevation. 7, .‘ ;p Q

Fig. 43Y. Uiasporus dastema (tiuayacan, CRARC, Costa Rica, 550 m). Photo: G. Kohler

Fig. 4;-. 3 i a s p o m vocatc- .-Jusagandi, Kuna Fig. 440. Diaspon -1. (Fortuna, Chiriqui, Yala, Panama, 350 m). Photo: G. Kijhler Panama). Photo: G. Kiihler

For more than a century all Neotropical direct developing frogs were placed in a single, enormous genus, Eleutherodactylus sensu lato (e.g., STEJNECER 1904, CRAWFORD & SMITH 2005). After the split- ting of the genus Eleutherodactylus into several families and genera, Eleuthero- dactylua, as currently understood, forms a group of 185 species that has its center of diversity in the West Indies (HBDGES et al. 2008, FROST 2010). The genus is distributed throughout the West Indies, Florida, and southern Texas, USA, Mexico, Belize, and Guatemala (HEDGES et al. 2008); introduced to Honduras, Costa Rica, and Panama (1-z & RAND 1990, m z et al. 1999b, SAVAGE 2002, MCCRANIE et d. 2008). The native Central American species (E. leprus, E. pipilans, and E. mbrimacula tus) belong to the subgenus Sphophus. The other species listed below are assigned to the nominal subgenus ( E l e u t h e r o d a ~ l ~ ~ ) .

The wide distribution and ability to suc- cessfully invade new areas (at least in one species, E. plm'rostris) of members of this genus probably derives from desiccation resistance of s me species (STEWART & MARTIN 1980)[ The three introduced species are ecologicd generalists that can be found in mesic and xeric forests, cutover fields, gardens, and greenhouses (SCHWABTZ & HEINDEBON 1991, KAISER & HARDY 1994). The three species of the sub-

-Rsading genus Sphophus are nocturnal inhabi-

SCOTT 1983c, LYNCH 2001, OVASKA & RaMD 2001, tants of open areas, including altered habi- MCCRANIE et al. 2002, SAVAGE 2002, CHAVEB et tats such as gardens and backyards; a1.2009 they can be locally abundant (Vow et al.

1997). ~ o u g h o u t the rainy season, the males of E. antillensis, E. coqui, and E. johstonei call from ground level up to three meters above the ground from sites such as piIes of debris (e.g., coconut husks), rock walls, shrubs, and trees (SCHWARTZ & HENDEBON 1991, IBAREZ et al. 1999b). The females lay 10 to 30 eggs in moist sites such as under debris or in concealed places

near ground level; one of the parents attends the eggs until they hatch about two weeks later (SAVAGE 2002). With one known exception, all species of Eleuthero- dactylus have direct development of eggs in a non-aquatic environment; the only exception is the Puerto Rican E. jasperi, which gives birth to living young (DREWRY & JONES 1976).

Fig. 441. Eleu therodactylus an tillensis (Panama City, Panama). Photo: B. Barker

I- - d Fig. 442. Eleutherodactylus johnstonei (Mandeville, Jamaica, 650 m).

Photo: K.-H. Jungfer

rclg. 44s. fi~eutherodactylus p~a.rurosms wan Pedro Sula, Cortb, Honduras).


Fig. 444. Eleutherodactylus coqui (Turrialba, Cartago, Costa Rica, 620 m). Photo: G. Kohler

Eleutherodactylus antillensis (REINHARDT & L ~ K E N 18631, Vidensk. Medd. Dansk Naturhist. Foren. Ser. 2,4: 209; type localities: "St. Croix", "St. Thomas", and "St. Jan" and "Vieques", Virgin Islands. Males to 28 mm, females to 47 mm. Islands of the Puerto Rican Bank, including Puerto Rico, the British Virgin Islands, and the U.S. Virgin Islands; introduced to Panama City, Panama, sea level to 1219 m elevation.

Eleutherodactylus coqui THOMAS 1966, Q. J. Florida Acad. Sci., 28: 376; type locality: "11.8 km S Palmer, Area Recreo La Mina, Puerto Rico." Males to 37 mm, females to 52 mm. Originally a Puerto Rican endemic, Eleutherodactylus coqui has invaded the Bahamas, the Dominican Republic, Ecuador, Guam, New Zealand, as well as Florida and Hawaii, USA, recently reported from Turrialba, Costa Rica (GARCIA-RODRIGUEZ et al. 2010), sea level to 1200 m elevation.

Eleutherodactylus johnstonei BARBOUR 1914, Mem. Mus. Comp. Zool. 44: 249; type locality: "St. Georges, [St. George Parish,] Grenada." Males to 25 mm, females to 35 mm. Most of the Lesser Antilles; introduced in many other regions including Jamaica, Venezuela, Colombia, Guyana, Trinidad, and Tobago, as well as Costa Rica and Panama, sea level up to a t least 1300 m elevation.

Eleutherodactylus leprus (COPE 18791, Proc. Am. Philos. Soc. 18: 268; type locality: "Santa Efigenia", Oaxaca, Mexico. Males to 26 mm, females to 29 mm. Atlantic versant of Mexico from central Veracruz eastwards to the Isthmus of Tehuantepec and north-central

- ~ e u t h e r o d a C ~ l u s mbrimacula tus (Finca Irlanda, Chiapas, Mexico, 1050 m).

Photo: G. Kijhler

Chiapas to northern Guatemala; also in northeastern Guatemala and southw88tern Belize; also on the Pacific v6rsat at the Isthmus df Tehuantepec; sea l&l to 800 m elevation.

Eleutherodactyilzs pipihas (TAYWR 19401, Roc. Biol. SOC. Wa8hbgton 53: 95; type lucali- ty: '9 mi. south of M a m t l b , aUemero, MMco (h. 3371." SVL to 31 mm. On the P a d c versant from central Guerrero, Mexico, to weetern Guatemala, 100-1800 m elevation.

E l e u t h ~ l ~ ~ d a ~ l w p l h h i s COP^: 1862b, Proc. Acad. Nat. Sci. Philadelphia 14: 163; type Z d t y : "New ~d~ Island, Bahamasw. SVL to 36 mm. Widespread on Cuba, the Isla de Juventud, Cagmdln Islands, Gaia Ida&, introduced in Florida, sonthem Louisiana, southern &or& and Hawaii (USA), h, Jamaica, Hondurw (see MCCRANE et 81. 2008), and V e r a m (Mexico), sea level to 720 m elevation.

Eleutheroda&tlw rubrimactzlam (TAYLOR & SMITa IS&), Proc. U.S. Natl. M w 96: 583; type locality: "La Esperanzfi, Chiapasw, Mexico. Males and female8 to 22.5 mm. On the P a d c vererant from & m e aciuthembrn Cbiaparr, &fexicos to western Guatemala, sea level to 1000 m elevation.

Fig. 447. Eleutherodactylus pipilans (Sierra Madre de Chiapas, Arriaga, Chiapas, Mexico, 690 m). Photo: A. Ramirez Veliizquez

1 LYNCH 1970a, SCKWARTZ & HENDERSON 1991,

Fig. 446. Eleutherodactylus lepnrs (Xalapa, KAISER & HARDY 1994. KAISER 1997, IBAI?EZ et ~eAcruz, Mexico). Photo: P. ~ e i m e s d . 1999b, KAISER et d.'2002, ~AvA~E.2002

Str abomantidae HEDGES et al. (2008) introduced the new family Strabomantidae to accommodate some 560 species of Neotropical direct- developing frogs distributed from eastern Honduras moss Lower Central America and widely in tropical South America, as well as on the Lesser Antilles. The current taxonomy divides this family into two subfamilies, the Holoadeninae (eix genera with a total of 45 species) and the Strabomantinae (eleven genera with a total of 517 species) (FROST 2010). In Central America occur representatives of two genera of the latter subfamily, Pn'stimantis and Strobmantis. PADIAL et al. (2009) provided molecular evidence that Strabomantidae, as currently recognized, may not be monophyletic. - Readin€! HFINICKE et al. 2007, HEDGES et al. 2008, PADIAL et al. 2009

On the basis of molecular evidence, SIEINICKE et al. (2007) resurrected the genus PTistimantis from synonymy with Eleutherodac@lus. The genus includes 434 recognized species distributed from Honduras southward across Central America and widely in tropical South America; also it is found on Trinidad and Tobago, Grenada, and on the Lesser Antilles (FROST 2010). The majority of species is distributed in tropical South America, whereas only 14 species are known to occur in Central America (D~LLMAN & LEHR 2009).

Most species of;this genus are inhabitants of mesic forests, where they can be observed at night perched on low vegetation; during daytime, species such as P. cerasinus and P. ridens can be found in the leaf litter (LIEBERMAN 1986). Other species,

Fig. 448. Pristimantis achatinus (Cerro ~ u i a Daribn, Panama). Photo: W. E. Duellman (Courtesy of the Biodiversity Institute, Universiw of Kansas)

rn A noteworthy phenomenon is the suggested mimicry relationship between &ti- inantis gaigei and poison-dart frogs of the genus Phylobatea (DUELUUN & TRUEB 1986, TOLEDO & HADDAD 2009). The palat- able Pristimantis gaigei has a color pat- I tern closely resembling the highly toxic potential models Phyllobates aurotaenia and Phyllobates lugubris. Pristimantis gaigsi occurs syntopically with the latter

Fig. 449. histimantis adnus (holotype). species in soubheastern Costa Rica and Photo: A. J. Crawford adjacent northwestern Panama, and with

such as P. moro, have been reportedto use epiphytic bromeliads as daytime hiding places (SAVAGE 2002). Food consists of a variety of small arthropods. As far as is known, all species of Pristimantis deposit eggs on the ground, in crevices of tree trunks, or on leaves of bromeliads (MYERS 1969); females of P. caryophyllaceus have been observed to brood their eggs (SAVAGE 2002). As is the case for almost all species of Terrarana, development in Pristimantis is direct and froglets hatch out of the eggs without a free-swimming aquatic tadpole stage (SAVAGE 2002).

P. aurotaenia in northwestern Colombia (LYNCH 1985b). Unlike most other Pristi- mantis, P. gaigei makes short hops as their principal means of locomotion, as do Phyllobates (SAVAGE 2002). As pointed out by SAVAGE (2002), the possibly mimetic and/or aposematic role for the color pattern in P. gaigei ia not established conclusively and needs further study.

Mtimantis achatinus (BOULENGER 1898b1, Proc. Zool. Soc. London 1898: 120; type locality: "CachabB [= Cachabi] . . . a small village on the river of that name, on the N.W. Coast, in the Prov. Esmeraldas. ... probably about 500 feet above the sean. Ecuador. Males to 36.2

Fig. 450. Pristimantis altat i road km NW S& Ram6n, Alajuela, Costa Rica, 1233 m).

Photo: A. J. Crawforfl

-, females to 46:l mm. Extreme eastern Panama (DariBn Province) to southern Ecuador, sea leveI to 900 m elevation.

Pristimantis adnus CRAWFORD, RYAN & J m 2010, Herpetologica 66 (2):194; type locality: "along a trail approximately 700 m in - elevation at the foot of Cerro Piiia in the F'! . - - . - - - -. - . - - -. . _-_ -- - . ,/ -*,- - Serrania del Sapo, above the Rfo Piiia on the Pacific coast of the Dari6n Province, Republic Fig. 451. Pristimantis caryophylaceus (La Ne- of Panama." Males to 20.0 m, females vera, Ngobe Bugl6, Panama). Photo: G. Kiihler unknown. Known only from the type locality.

Pristimantis altae (DUNN 1942b), Not. Nat. Philadelphia 104: 1; type locality: "Atlantic. slope of Costa Rica, on the pass between Barba and Irazu, below the divide and above the finca of Felix Delgado, approximately 4000 feet", Cant6n de El Guarco, Provincia de. Cartago, 2860 m, Costa Rim. Males to 28.5:; mm, females to 27.0 mm. On the Atlantic versant from northern Costa Rica to extreme! northwestern Panama, 60-1245 m elevation.

Pristimantis caryophyliaceus (BARBOUR 19281, Proc. New England Zool. mub 10: 28; type Fig. 452. Plistimantis caryophyllaceus (female locality: "La Loma on the trail from with eggs; Alto de Piedra, Veraguas, Panama, Chiriquicito to Boquete, Bocas del Toro 910 m). Photo: A. Hertz Province, Panama." Males to 24 mm, females to 26 mm. On the Atlantic versant from northern Costa Rica to westen-central Panama, and along the Pacific vmant from extreme southern Costa Rica to western Panama, sea level to 1900 m elw?tion.

Pristimantis cerasinus (COPE 18761, J. Acad. Nat. Sci. Philadelphia Ser. 2,B: 112; type local-

I ity: *eastern slope of the Pico Blanco", Costa Rica; corrected to "Cerro Utyum, 5000-7000 R, Cant6n de Talamanca, Provincia de Lim6n: 1524-2134 m", Costa Rica by SAVA~E (1974). Males to 25 mm, females to 35 mm. d m-l Northeastern Honduras to eastern Panama Fig. 453. Pristimantis educatoilD &om type

locality, 800 m). Photo: A. J. Crawford

.including islands in the nocas del Toro archipelago) and marginally on the Pacific slope in northwestern Costa Rica, 20-920 m elevation. Eleutherodactylus operosus SAVAGE, MCCRANIE & WILSON 1999 is regarded as a synonym of P. cerasinus according to MCCRANIE &WILSON 2003b.

Pristimantis cruentus . (PETERS 18731, Monatsber. Preuss. Akad. 'Wiss. Berlin 1873: 609; type locality: "Chiriqui", Panama (see comment under Ameerega maculata). Males to

~ i ~ . 454. fiStUdtiS bGx asinLu ,Lbancho ~ ~ t ~ - 28 mm, females to 42 mm. On the Atlantic versant from northern Costa Rica to western-cen-

ralista, Costa Rica, 900 m). Photo: G. Kohler tral Panama, 40-1800 elevation.

Fig. 455. Pristimantis cerasinus (Nusagandi, I

Kuna Yala, Panama, 350 m). Photo: G. Kohler

Fig. 456. Pristimantis cruentus (Albergue Monterreal, Alajuela, Costa Rica, 1850 m).

Photo: G. Kohler

Fig. 457. Pristiman tis gaigei (Changuinola, Bocas del Toro, Panama, 10 m). Photo: M. Ponce

Pristimantis educatoris RYAN, LIPS & GIERMAKOWSKI 2010, J. Herpet. 44: 194; type locality: "forest just below the Continental Divide on Atlantic slope; Parque Nacional G. D. Omar Torrijos H., Cocle', PanamP (So39.95'N: 80°35.55W 709 m elevation; ... approximately 8 km north of El CopiS, Coclh Province, Panama." Males to 20.4 mm, females to 37.7 mm. Santa FiS, Veraguas Province in central Panama to Colombia, with a disjunct population in extreme southeastern Costa Rica on the Atlantic versant of Cerro Uatsi, Bratsi, Limdn Province, 450-1450 m elevation.

Pristimantis gaigei (DUNN 19311, Occas. Pap. Boston Soc. Nat. Hist. 5: 387; type locality: "Fort Randolph, Panama Canal Zone", Panama. Males to 31 mm, females to 44 mm. On the Atlantic versant from extreme southern Costa Rica to eastern Panama, also on the Pacific versant in central Colombia, 20-200 m elevation. I here follow BRANDON- JONES et al. (2007) in the usage of the species' name as originally spelled, and not the emen- ded name ("gaigeae"), as proposed by T A ~ O R (1952b).

Pristimantis moro (SAVAGE 19651, Bull. S. California Acad. Sci. 64: 106; type locality: "Costa Rica: Provincia de San Josh: Cantdn de Coronado: La Hondura, 1245 meters (4085 feet)." Males to 19.5 mm, females to 25 mm. Atlantic versant in central Costa Rica (near La Hondura) and Pacific versant in central Panama; also in western Colombia, 550-1245 m elevation.

Pristimantis museosus (I~ArriEz, JARAMILLO & AROSEMENA 19941, Amphibia-Reptilia 15: 337; type locality: "Culebra (08"52'14"N, 82"25'12W), head waters of Rio Changuinola, Distrito de Bocas del Toro, Provincia de Bocas del Toro, Republics de PanamP, 1000 m." Males to 22.4 mm, females to 38.2 mm. Western and central cordilleras of Panama, 700-1000 m elevation.

Fig. 458. Pristimantis moro (El Cop6, Cocl6, Fig. 462. Pristimantis ridens (near Rio Sereno, Panama, 700 m). Photo: B. Kubicki Chiriqui, Panama, 1210 m). Photo: G. Kohler

Fig. 459. Pristimantis museosus (Cerro Negro, Fig. 463. Pristimantis ridens (9 km NW For- Veraguas, Panama, 770 m). Photo: A. Hertz tuna, Alajuela, Costa Rica). Photo: G. Kohler

Fig. 460. Pristimantis pardalis (Cerro Mariposa, Fig. 464. Pristimantis ridens (P.N. Saslaya, Veraguas, Panama, 870 m). Photo: G. Kijhler Atlantic0 Norte, Nicaragua, 820 m).

Photo: G. Kohler

Fig. 461. Pristimantis pirrensis (Cana, Darikn, Fig. 465. Pristimantis taeniatus (Los Santos, Panama, 1300 m). Photo: A. J. Crawford Santander, Colombia, 1640 m).

Photo: C. Hernandez J. 189

%a Loma, in the cloud forest on trail fr Chiriquicito to Boquete, Bocas del TO Province, Panama? Males to 19 mm, femal to 29 mm. On the Pacific versant in southe*? Costa Rica and on the Atlantic versant fro&& northwe- t~ wSern P- 50-1qQ J&<

elevation :. ,, L

a

J%stimantis g&nais (- & CRAWOW,' 2004), J. Herpetol. 38: 241; type locality: ' ' a io~, the headwaters of the Cana River, appro& mately 100 m below the Pirre Tent Camp by Ancon Expeditions a t Cana, D a r i b National Park, Dan& Province, Panama, 1 2 w m elevation, 07"45'48.6"N, 77O43'19.8Wf" Females to 30.0 mm (males unknown). Know4X only from the vicinity d twBs m. 1250-1550 m elevation.

1 d to 18fB &I @?m&.Qian. I ' r f a m g 8 @OULEpBieTp 1.SWh Ann. Mag, 'W. %.% Bex+ $10: 18% Qp lad-

I I ity: "No- E. N-al, Rio Raa J m Chco, $.X COED*.^^ p.1 mm* caryophY1lalaceus f d w tjg 8g.k 'wecentral P- 9 pardalis

9 pirrensis (--A& &fib Haita Tabmar4 I$&@ C' C~Ibmbk -6 & h@'h&@ b vdh calla * h m t * d acro5a $he noAem

0f W- =a en^ m- thewe b the m5ddle Mrgdaha V&E 8&1400 m d~v&ari.

F'urther Reading SAVAGE 1981b, MIYAMOTO 1984, LYNCH 1985b, k4Fmz et al. 1994, LYNCH & ARDILA-ROBAYO

Pristimantis 1999, IBAREZ & CRAWFORD 2004, CRAWFORD et al. 2010, RYAN et al. 2010

P. cruentus I? moro 1

Strabomantis

The genus Strabomantis was removed from synonymy with Eleutherodactylus by HEINICKE et al. (2007). It includes 17 species of broad-headed frogs distributed in southern Costa Rica and Panama, as well as through the wet tropics of Colom- bia, northern Venezuela, Ecuador, eastern Peru, and western Brazil (FROST 2010). Just as with most of the species in this genus, both Central American representatives are toad-like terrestrial frogs. Strabo-

1896a), Ann. Mag Nat. Hist. Ser. 6,17: 19; type locality: UBuenaventuran, Colombia. Males to 69 mm, females to 94 mm. On the Atlantic versant from extreme southern Costa R i a to western Colombia, 16-350 m elevation.

Strabomantis laticorpus (MYERS & LYNCH 1997), Am. Mus. Nwit. 3196: ;4; type locality: "montane forest on southwest sector of Cerro Tacarcuha mass% 1590 m elevation, Dari6n Province, Republic of Wnama. Approximatsly 8"10W, 77O4OW." Males to 30 mm, females to 40 mm. Cerro Tacarcuna and Cerro MaIf in the? southeastern Daribn Province, Panama, 1460-1630 m elevation. - -7 8 Z . L , 8 . . - rb:.. - & a , , = . '

mantis laticorpus is reported ti be active Further Rsading in the leaf litter by day, but also encoun- MYERS & LYNCH 1997, wz et al. 1999b, tered at the edge of a stream at night sAVAGE 2002, DUELLMAN & LEHR 2009 (MYERS & LYNCH 1997). Strabomantis bufoniformis is a nocturnal terrestrial stream- associated frog that can be found on the ground along small forest streams at night (K~HLER unpubl. obs.). I B ~ Z et al. (1999b) reported males calling from the edges of small streams just after dusk. The call of S. bufoniformis consists of paused short low bark-like notes followed by even shorter knock-like ones ( I B ~ z et al. 1999b).

Fig. 466. Strabomantis bufoniformis (Nusagandi, Kuna Yala, Panama, 350 m). Photo: G. Kijhler

Hemiphractidae +--- - , - WWSSWSP----

3 Fig. 467. He1111phractus fasciatus (Valle de Anton, Cocl6, Panama). Photo: B. Wilson

Hemiphractidae

Recently, GUAYA~AMIN et al. (2008) found ies of water (DUELLMAN & ~MANEss 1980, that the species of the former families WASSERSUG & DUELLMAN 1984, WIENS et al. Amphignathodontidae, Cryptobatrachidae, 2007). and Hemiphractidae form a monophyletic group that they recognized as a single fam- f i y to the species of Hemiphractidae ily, Hemiphractidae. This family is a pre- 1 a A fleshy proboscis; head triangular in dor- dominantly South American group of frogs sal view; no brood pouch in females

.............................. consisting of 93 species in five genera, with Hemi- fzleciatue only three species in two genera occurring b NO fleshy proboscis; head not triangular in in Lower Central America (FROST 2010). dorsal view; a brood pouch present in

...................................................... females 2 Most species in this family have direct a A triangular flap on upper eyelid; development, with eggs brooded by the flank and posterior surfaces of thigh not female in a dermal pouch (e.g., in darker than dorsum Gestrotheca m u t a Gastrotheca) or attached to the dorsal skin b No t f i m d a r dermal flap on upper eyelid; (e.g., in Hemiphractus) (DUELLMAN 1970, flank and posterior surfaces of thigh dark-

......... MENDELSON et al. 2000). In all hemiphrac- er than dorsum GWmtheca nioefhn' tids, eggs are carried on the female's dor- -er Re* sum for some or all of development; eggs TRUEB 1974, DUELLMAN & ~ E S S 1980, hatch directly into froglets or into tadpoles WAS~ERSUG & DUELLMAN 1984, m ~ ~ s o ~ et that are deposited in ponds and other bod- al. 2000, GUAPASAMIN et al. 2008

Gastrotheca c -

Fig. 468. Gastrotheca cornuta (female with eggs; Valle de Antdn, Cocl6, Panama).

Photo: B. Wilson

The 58 species of Gastrotheca are widespread throughout much of South America with only two species occurring in Lower Central America (FROST 2010). Gastrothe- ca cornuta is a nocturnal canopy dweller in tropical rainforest and low montane forest characterized by high humidity m I throughout the year (DUELLMAN 1970, SAVAGE 2002). The males call from high in I the trees, with the call having been ~

I described as a loud "bop" (DUELLMAN 1970). The larvae develop fully enclosed - within a special brood pouch that develops Fig. 470. Gastrotheca cornuta (male; Nusagandi, as a posterior invagination of the female's Kuna Yala, Panama, 320 m).

I dorsal skin; the embryos are partly or com- Photo: E-H. Jungfer pletely encased in large, thin bell-shaped

gills providing a surface for gas exchange (PINO 1980, WASSERSUG & DUELLMAN 1984). In some Gastrotheca species, including G. cornuta, the eggs hatch directly into froglets, whereas in some high Andean species the eggs hatch into tadpoles that are released into ponds (DUELLMAN 1983, WASSERSUG & DUELLMAN 1984, WIENS et al. 2007).

I

Fig. 469. Gastrotheca nicefori (Ridge between ' Rio Jaque and Rfo Imamado, Daribn, Panama).

Photo: C. W. Myers Fig. 471. Gastrotheca cornuta (P.N. Chagres, (Co- of the Biodiversity Institute, University of Kansas) Col6n, Panama, 790 m). Photo: M. Ponce

Hemiphractidae 1

Gastrotheca cornuta (BOULENGER 1898b1, Proc. Zool. Soc. London 1898: 124; type locality: "Cachab6 [=Cachaba . . . a small village on the river of that name, on the N.W. Coast, in the Prov. Esmeraldas. ... probably about 500 feet above the sea", Ecuador. Males to 81.1 mm, females to 76.9 mm. On the Atlantic versant from central Costa Rica to central Panama, and on the Pacific versant from eastern Panama to western Ecuador, sea level to 1000 m elevation. Hyla ceratophrys STEJNEGER 1911 is regarded as a synonym of G. cornuta according to DTPLLMAN (1983).

Gastrotheca nicefon' GAIGE 1933, Occas. Pap. Mus. Zool. Univ. Michigan 263: 1; type locality: "Pensilvania, [Departamento Caldas,] Colombia." Males to 73.5 mrn, females to 82.0 mm. Western Panama to Colombia and Venezuela (there is a large gap between the records in western and eastern Panama), 800-1100 m elevation.

Further Reading Fig. 472. Hemphractus fasciatus (Nusagandi,

DUELLMAN 1980, DUELLMAN & MAN!XSS 1980, Kuna Yala, 320 m)- DUELLMAN 1983, WASSERSUG & DUELLMAN 1984, Photo: K.-H. Jungfer DUELLMAN 1989, WIENS et al. 2007

Gastrotheca comuta Gastrotheca nicefori I

1 I Herniphractus fasciatus i

Hemiphractus

Some of the most distinctive, if not the most bizarre looking frogs belong to the genus Hemphractus, an assemblage of six species with large, triangular heads ornamented with bony processes and fleshy protuberances adorning eyelids and snout (MENDELSON et al. 2000, SHEIL & MENDELSON 2001). The very large mouth contains a pair of sharp, fang-like premaxillary teeth (odontoids) (DUELLMAN 1970). Hemiphractus is a Neotropical endemic with only one species occurring in southeastern Central America, the other species being distributed in tropical South America (TRUEB 1974, SHEIL & ~ N D E L S O N 2001). Hemphractus fasciatus is nocturnal and inhabits the forest floor of dense, tropical rainforest and cloud forest; by day individuals of this species hide in leaf litter; at night, they can be found on low vegetation at a height of less than one meter above the ground (DUELLMAN 1970, KOHLER unpubl. obs.). They feed on a variety of invertebrates (arthropods and gastropods)

Hylidae

Hylidae

Fig. 473. Hemphractus fasciatus (Nusagand, Kuna Yala, Panama, 350 m). Photo: G. Kohler

and vertebrates (e.g., frogs and lizards) that they subdue rapidly and with a powerful bite (DUELLMAN 1970). When handled, H. fasciatus gaped their mouth and readily attempted to bite, which can be quite painful to the one bitten, given the sharp odontoids on fiont of the lower jaw (MYERS 1966). Very little is known about the reproductive biology in Hemphractus, although females have been reported to carry the eggs openly on the back, in a series of depressions in the dorsal skin with the eggs adhering firmly to the female's back; the developing embryos are completely enclosed in large, membranous gills that are embedded in deep pits of the female's back (DUELLMAN 1970). All species of Hemiphractus have direct development with froglets hatching from the eggs (WIENS et al. 2007).

Hemphractus fasciatus PETERS 1862, Monatsber. Preuss. Akad. Wiss. Berlin 1862: 149; type locality: "Pastassa-Thal [Pastaza Valley], an der Ostseite der Anden in Ecuador", [in error, according to TRUEB (1974)l. Males to 55.7 mm, females to 68.7 mm. Central Panama to , northwestern Ecuador, 300-1600 m elevation. Cerathyla panamensis STEJNEGER 1917 is regarded as a synonym of H. fasciatus according to TRUEB (1974).

Further Reading MYERS 1966, TRUEB 1974, MENDELSON et d. 2000, SHEIL & MENDELSON 2001, SHEIL et al. 2001

The family Hylidae (treefrogs) is a very large and heterogeneous assemblage of tiny to very large frogs distributed over much of North, Central, and South America, the West Indies, temperate Eurasia, the Australo-Papuan region, and North Africa (DUELLMAN 1970, 2001). Currently, close to 900 species of treefrogs are recognized, organized in three subfamilies: Hylinae (637 species), Pelodryadinae (196 species, none in Central America), and Phyllomedusinae (59 species) (FROST 2010). In Central America, this family is very well represented with a total of 105 species in 22 genera. Characteristics of hylid frogs include expanded adhesive digital discs, an intercalary cartilage between the penultimate and terminal phalanges, and usually long legs (DUELLMAN 1970, SAVAGE 2002). The members of the two subfamilies occurring in Central America are readily distinguished by the shape of the pupil: the pupil is vertically elliptical in species of the subfamily Phyllomedusinae and horizontally elliptical in species of the subfamily Hylinae (DUELLMAN 1970). Most hylids are nocturnal and arboreal, although some species are terrestrial and a few are fossorial (DUELLMAN 1970).

Key to the subfamiles of Hylidae 1 a Pupil vertically elliptical in bright light (in

life) or preservative; most species with bright green overall dorsal coloration in life (blue in preservative) . .. . Phyllomedusinae

b Pupil horizontally elliptical in bright light (in life) or preservative; dorsal coloration variable ........................................... H y b

Further Reading DUELLMAN 1970, SAVAGE & HEYER 1970, DUELLMAN 2001, MCCRANIE & WILSON 2002, SAVAGE 2002

Phyllomedusinae The subfamily Phyllomedusinae (lear- frogs) is endemic to the Neotropics and contains five genera and 59 species (FROST 2010). The species of this subfamily are distributed from southern Mexico across Central America and also widely in South America (DUELLMAN 1970, 2001). The

I Central American phyllomedusines are

Fig. 474. Agalychnis callidryas (Soltis Center, placed into three genera: Aga l~c f i s (five Alajuela, Costa Rita, 460 m). Photo: K&ler species), Cruziohyla (one species), and ~ h . y l l o i e d u s (one species? (DUELLMAN 1970, 2001). Except for A. a m a e that occurs at moderate elevations, the Central American phyllomedusines are primarily denizens of the lowland rainforest (SAVAGE 2002). During daytime, the Central American species hide in trees, often high up in the forest's canopy (KUBICKI 2004b). The breeding takes place primarily during the rainy season, and the females deposit their eggs on leaves, vines, roots, bark, or wood overhanging water; on hatching, tadpoles fall or are washed by rain into the water body below (SAVAGE 2002, KUBICKI 2004b). Phyllomedusine tadpoles, except those of C. calcarifer, are often observed orienting themselves with the head up near the water surface, moving just the tip of the tail in a rapid motion (SAVAGE 2002, KUBICKI 2004b).

Key to the species of phyllomedusine frogs (genera Agalychi~, Cciuziohyla, and l?hyflomedwsa) 1 a Webbing between fingers and toes, if pre-

sent, only rudimentary (Fig. 47713) .......... 2 b Hands and feet with significant webbing 3

2 a Parotoid glands absent; Toe I shorter than Toe 11; SVL of adult males 30-41 mm, of adult females 39-53 mm Agalychni.9 lemur

b Well-developed parotoid glands present; Toe I longer than Toe II; SVL of adult males 75-86 mm, single known adult female 98 mm ......................... Phyhmedusa venusta

3 a Flanks with contrasting pale and dark vertical bars (Figs. 475, 492) ......................... 4

b Flanks without contrasting pale and dark ............................. vertical bars (Fig. 476) 5

4 a Lower eyelid not reticulate; heel with flap- like calcar; flanks yellow or orange in life

with vertical black bars; iris gray in life; SVL of adult males 50-81 mm, of adult females 60-87 mm .. ~ o 4 y . a cal&

b Lower eyelid reticulate (Fig. 474); heel without flap-like calcar; flanks blue, purple, or brown in life with cream or white vertical bars; iris red in life; SVL of adult males 39-59 mm, of adult females 51-71 mm ...........................

5 a Fingers with reduced webbing, webbing between Fingers 111 and IV not reaching to penultimate subarticular tubercle of Finger 111 (Fig. 477e); flanks blue or purplish blue; dorsum usually with wavy transverse lines; SVL of adult males 34-54 mm, of adult females 52-66 mm ................ ..................................... - saltator

b Fingers with substantial webbing, webbing between Fingers 111 and IV reaching to penultimate subarticular tubercle of Finger I11 (Fig. 477a,c,d); flank coloration variable blue; dorsum without wavy trans-

......... verse lines; SVL of adults variable 6 6 a Fingers with complete or almost complete

webbing, webbing between Fingers I11 and IV reaching well beyond penultimate subarticular tubercle of Finger I11 (Fig. 4778; SVL of adult males 48-75 mm, of adult females 81-87 mm .... &d jmhk spurrelLi

b Fingers without complete webbing, webbing between Fingers I11 and IV reaching to penultimate subarticular tubercle of Finger 111 (Fig 477a,c); SVL of adults variable ............................................................. 7

7 a Flanks, as well as anterior and posterior surfaces of thighs blue in life; iris yellow to orange in life; SVL of adult males 57-74 mm, of adult females 67-84 mm ................ ........................................ Ad* annae - -

b Flanks, as well as anterior and posterior surfaces of thighs pinkish to orange in life; iris maroon to dark red in life; SVL of adult males 51-66 mm, of adult females 71-83 mm ............................ Agalychuis momletii

I Fig. 475. Agalychnis callidwas (Volch Fig. 476. Agalychnis moreletii (El Refugio,

I Mombacho, Granada, Nicaragua, 1000 m). Ahuachaph, El Salvador, 225 m). I Photo: G. Kohler Photo: G. Kohler

I I

Fig. 477. Hand of a. Agalychnis annae; b. A. lemuq c. A. morelee d. A. s p m e f i , e. A. saltator. Drawings: M. Vesely

Agalychnis

Fourteen species are currently assigned to the genus Agalychnis; the most recent change has been the transfer of the species lemur from Hylomantis to Agalychnis (FAIVOVICH et al. 2010); as currently understood, Agalychnis consists of 14 species distributed from southern Sonora, Mexico, across Central America to the humid lowlands of northern South America on both

SAVAGE 2002). Males have been reported to gather in breeding aggregations on vegetation above the breeding site (SCOTT & STARRETT 1974, ROBERTS 1994, KUBICKI 2004b). Amplexus is axillary (DUELLMAN 1970). Prior to egg laying, the amplecting pair descends to the water body where the female remains with ,her posterior body submerged in the water, supposedly filling her urinary bladder with water (SAVAGE 2002). So far as is known, all species of

sides of the Andes, including the Atlantic coastal forests of Bahia and Pernambuco, Brazil (FAIVOVICH et al. 2010). The species of this genus are moderately-sized to large, mostly very photogenic treefrogs. During daytime, and generally during the dry season, these nocturnal frogs seek shelter in the treetops. Males call throughout the year in rainforest regions, but are only active during the rainy season in seasonal forests (SAVAGE 2002). The call of A. cal- Lidryas consists of a single "chock" or a double "chock-chock" (A. callidryas also often emits a series of clucks, especially when present in small aggregations; B. KUBICI~I pers. comm. Oct. 2010), those of A. annae and A. moreletii were described as a single note "wo-or-op", that of A. spurrelli as a single low-pitched "groan", and that of A. lemur as a very short "tick" (DUELLMAN 1963b, 1970, JUNGFER & WEYGOLDT 1994,

I

Fig. 478. Agalychnis moreletii (Ocozocuautla, Chiapas, Mexico, 1000 m). Fig. 479. Agalychnis moreletii (El Occ-,

Photo: A. Ramirez-Velazquez Chiapas, Mexico). Photo: G. Kohler

Hylidae

Fig. 480. Agalychnts amae (Plan de Birri, Fig. 482. Agalychnis annae (La Nevera, Ngobe Heredia, Costa Rica, 1630 m). Photo: G. Kohler BuglB, Panama). Photo: G. Kohler

Fig. 481. Agalychnis callidryas (KilambB, Fig. 483. Agalychnis callidryas (Peninsula Jinotega, Nicaragua, 1015 m). Photo: G. Kohler Burica, Chiriqui, Panama, 65 m).

Photo: A. Hertz

Agalychnis deposit the eggs on the upper surface of leaves (Fig. 478), branches, or vines overhanging lentic water bodies such as ponds, puddles, road-side ditches, swimming pools, garden fountains, and backwaters of slow-moving small streams (PIBURN 1963,1964, JUNGFER & WEYGOLDT 1994). After hatching, which occurs five to seven days after deposition, the tadpoles fall or are washed by rains into the water bodies below (KUBICKI 2004b). Typically, free swimming tadpoles float in midwaters with their bodies directed at about 45" to the water surface, maintaining this position by constant rapid movements of the tail tip (SAVAGE 2002, KUBICKI 2004b). Parachuting ability has been reported for --- = STARRETT 1974, ROBERTS 1994). nas, Costa Rica). A. sdtator and A. ~purrelli (SCOTT & Fig. 484. Agalychnis spurreLLi (Bani, Puntare-

Photo: G. Kohler

Fig. 486. Agalychnis lemur (Guayach, CRARC, Costa Rica, 550 m). Photo: G. Kiihler

Fig. 485. Agalychnis spurrelli (Guayach, I

CRARC, Costa Rica, 550 m). Photo: G. Kohler

Agaly&.uk annae (DUEW 1963b), Rev. Biol. Tropical 11: 1; type locality: "Tapanti, [Cant6n de Pardso,] Cartago Province, Costa Rica, 1200 meters." Males to 73.9 mm, females to 84.2 mm. Cordilleras de TilarBn, Central, and Talamanca, Costa Rica; recently recorded from the Serrm'a de Tabasara in western-central Panama (HERTZ et al. in press), 780-1650 m elevation. I Agalychnis callidryas (COPE 1862a), Proc. Acad. Nat. Sci. Philadelphia 14: 359; type locality: "Panama." Males 56 mm, femalesto ~ i ~ . 487. ~ g a l ~ ~ f i ~ moxletii ( ~ i ~ ~ ~ ~ ~ l ~ d ~ , 71 mm. On the Atlantic versant from southern Chiapas, Mexico, 1050 m). photo: G. ~m~~ Veracruz and northern Oaxaca in Mexico, southeastward to central Panama; on the Pacific versant from southern Costa Rica to Colombia, near sea level to 1325 m elevation. Agalychnis helenae COPE 1885 is regarded as a synonym of A. cdidryas according to SAVAGE & HEYER (1967).

Agalychnis lemur (BOULENQER 1882a), Cat. Batr. Sal. Coll. Brit. Mus. Ed. 2: 425; type locality: "Costa Rica." Males to 40.8 mm, females to 50.6 mm. Costa Rica and Panama, and marginally in Colombia, 440-1600 m elevation.

Agalychnis moreletii (DuI&R~, 1853), Ann. Sci. Nat. Paris, Ser. 3, 19: 169; type locality: 'Vera- Paz", Guatemala. Males to 65.7 mm, females to 82.9 mm. On the Atlantic versant from northeastern Puebla and southcentral Veracruz, Mexico, to northwestern Honduras; and on the Pacific versant from southcentral Guemem, Fig. 488. Agalychnis saltator (La Selva,

Heredia, Costa Rica, 70 m). Photo: G. Kohler

Mexico, to central El Salvadur, 200-2130 m elevation.

Agalychnis sdtator TAYLOR 1955, Univ. Kansas Sci. Bull. 37: 527; type locality: "4 km. NNE of Ti larh [= Finca San Bosco, Cant611 de T i l a h according to SAVAGE 19741, [Provincial Guanacaste, Costa Rica." Males to 46.7 mm, females to 61.5 mm. Atlantic slopes from northeastern Honduras to southeastern Costa Rica, near sea level to 820 m elevation.

Agdychnis spurreIli BOULENGER 1913, Roc. Zool. Soc. London 1913: 1024; type locality: "Peiia Lisa, Condoto, altitude 300 feet," Provincia Choc6, Colombia. Males to 75.6 mm, females to 92.8 mm. Southern Costa Rica on both versants across Panama and the Pacific lowlands of Colombia to northwestern Ecuador, 15-885 m elevation. Phyllomdnsa litookyas DUELLMAN & TRUEB 1967 is regarded as a synonym of A. spurrelli according to ORTEGA-ANDRADE (2008).

. A. spumlli A. annae

Agalychnis

Fig. 489. Agalychnis spurrelli (near Golfito, Puntarenas, Costa Rica, 50 m). Photo: B. Kubicki . A. moreletii

Further Reading I PWURN 1963,1964, SAVAGE & HEYER 1967, PYBURN 1970, SCOTT & STARRETT 1974, PROY 1992, PROY 1993a, JUNGFER & WEYGOLDT 1994, ROBERTS 1994, D'ORGEM & TURNER 1995, GRAY & RAND 1997, PROY 2000, MCCRANIE et al. 2003, HOFFMANN 2005a

Fig. 490. Cruziohyla calcarifer (Guayacan, CRARC, Costa Rica, 550 m). Photo: G. Kahler

w Cruziohyla

The genus Cruziohyla was introduced by FAIVOVICH et al. (2005) for two species of leaf-frogs, one of which, C. calcarifer, occurs in Central America, whereas C. craspedopus is restricted to the Amazonian lowlands in South America. Cruziohyla calcarifer is a large nocturnal treefrog that inhabits the rainforest canopy and only descends to find suitable breeding sites during the rainy season (SAVAGE 2002). Males call from trees surrounding or one to seven meters above the breeding sites (KUBICKI 2004b). The call of C. calcarifer has been described as a soft single note, "rurm" or "whuunk"; several minutes to hours can pass between the calls (SAVAGE 2002, KUBICK~ 2004b). This species does not

utilize permanent or temporary ponds for breeding; rather, the females deposit the eggs on substrate (leaves, aerial roots, wood, or bark) over water-filled cavities of living trees, depressions in the tree trunk or its buttresses or similar habitats on fallen trunks (MARQUIS et al. 1986, ROBERTS 1995, KUBICKI 2004b). The eggs hatch within five to ten days and on hatching, the tadpoles fall or are washed by rain into the water below (SAVAGE 2002). The tadpoles feed on organic material such as algae and decaying vegetable matter in their puddle; due to the low concentration of organic food in this environment, it can take up to a year or more before metamorphosis takes place (KUBICKI 2004b). Under laboratory conditions, tadpoles metamorphosed after a minimum of 211 days (DONNELLY et al. 1987).

Hylidae

Cruzioh~la calcarifer (BOULENGER 19021, Ann. Mag. Nat. Hist. Ser. 7,9: 52; type locality: "Rio Duraneo. 350 feet". Provincia Esmeraldas. 1 ~ c u a d & . . ~ a l e s to 81.0 mm, females to 78.5 mm. On the Atlantic versant f?om eastern Honduras to central Panama. and on the Pacific versant from eastern panama through the Pacific lowlands of Colombia to northwestern Ecuador, sea level to 820 m elevation.

Fig. 491. Cruziohyla calcarifer (Guayacan, CRARC, Costa Rica, 550 m). Photo: G. Kohler

Further Reading MARQUIS et al. 1986, DONNELLY et al. 1987, CALDWELL 1994

I Fig. 492. Cruziohyla calcarifer (La Selva, Heredia, Costa Rica, 70 m). Photo: G. Kohler

203

- . - . -

Phyllomedusa - - - - --.

Only a single species of the predominantly South American genus Phyllomedusa reaches extreme eastern Panama (DUELLMAN 1970, 2001). Phyllomedusa venusta is large nocturnal, arboreal frog that inhabits the tree tops of the rainforest, and only descends to reach its breeding sites during the rainy season (RENJIFO & LUNDBERG 1999). The females deposit the eggs on the upper leaf surface of plants overhanging small ponds, puddles and water-filled depressions within the forest ~ i ~ . 494. p.yllomedusa venusta (Montes de (RENJIFO & LUNDBERG 1999). The five type ~ b ~ , sucre, colombia). Photo: A. Acosta specimens of this species were found on vegetation in a swamp forest "where the species probably breeds" (DUELLMAN 1970: 138).

Phyllomedusa venusta DUELLMAN & TRUEB 1967, Copeia 1967: 128; type locality: "a small swamp approximately 1 km west-southwest of the junction of the Rio Mono and the Rio Tuira, DariBn Province, PanamB, elevation 130 m." Males to 86.3 mm, females to 97.7 mm. Eastern Dar ik Province, Panama, as well as in the middle Magdalena Valley, Colombia, 130-1200 m elevation.

Fig. 495. Phyllomedusa venusta (Montes de Maria, Sucre, Colombia). Photo: A. Acosta

Fig. 493. Phyllomedusa venusta (Cana main camp, P.N. Darien, D d n , Panama, 500 m). Further Reading

Photo: A. J. Crawford DUELLMAN & TRUEB 1967, DUE= 1970,2001

Hylinae The frogs of the subfamily Hylinae are predominantly nocturnal, arboreal species, although a few (e.g., ACRS and Pseudacris) are terrestrial (VIIT & CALDWELL 2009). According to the current taxonomy, 637 species in 38 genera are recognized, distributed disjunctly across temperate Eurasia, northern Africa, and throughout the Americas and West Indies (FROST 2010). Axillary amplexus is typical for hyline frogs, and the females deposit the eggs in a variety of sites ranging from leaf surfaces over water bodies, in tree holes and bromeliads to lentic and flowing water bodies on the ground (SAVAGE 2002). Parental care is uncommon but occurs in gladiator frogs (genus Hypsiboas) and two species of Smilisca (KLUGE 1981, MALONE 2004).

Key to the genera of hyline frogs 1 a Snout with greatly expanded labial shelf

(Fig. 496a); head much longer than wide .. ....................................................... nipl-2-on

b Snout not expanded laterally; ratio head ................... length 1 head width variable 2

2 a Bony occipital, canthal, and frontal spines present in adults (Fig. 497); a distinctive dorsal pattern of black or dark brown

..... blotches outlined with white Anotheca b No bony occipital, canthal, or frontal

........ spines; dorsal pattern not as above 3 3 a Snout protruding in profile (Fig. 496b);

webbing between Toes I and I1 reduced to a fringe along inner margin of Toe I1 or absent, not extending beyond subarticular tubercle of toe I (Fig. 499a) ............ Scinax

b Snout not or only slightly protruding in profile and with rounded upper edge; webbing between Toes I and I1 not greatly reduced relative to webbing between other

.................................... toes (Fig. 499b-d) 4 4 a Skin on dorsal shrface of head co-ossified

with roof of skull to form a definite casque (Fig. 498); finger discs very large; dorsal coloration in life pale green, ashy gray, pale brown, or reddish brown with bold dark reticulations or elongate blotches ..... .................................................... o~teopilw

b Skin on dorsal surface of head not co-ossified with roof of skull to form a definite casque, or, if co-ossified, then scalloped

Fig. 496. Snout profile: a. Trprion petasatus; b. Scinax elaeochroa. Photos: G. Kohler

Fig. 497. Anotheca spinosa (Rancho Naturalists, near Platanillo, Costa Rica, 900 m).

Photos: G. Kijhler

Fig. 498. Osteopilus septentrionalis (Florida, USA). Photos: T. Leenders

Fig. 499. Foot of a. Scinax staufferi; b. Isthmohyla pseudopuma; c. Isthmohyla rivularis; d. Smilisca baudinii. Drawings: M. Vesely

- dermal fringes present on posterior edges of forearms and tarsal segments (Figs. 537, 539); size of finger discs variable; dorsal coloration variable .................................... 5

5 a Skin on dorsum thick, glandular, and tuberculate (Fig. 5001, a particularly dense zone of thickened, rugose skin in occipital and frontal regions; hands and feet large with large discs and extensive webbing; males with paired lateral vocal sacs behind angles of jaws (Fig. 506b); no prepollex in adult males ....................... Th-phalw

b Skin on dorsum not conspicuously thick and glandular, or, if so, then hands and feet not having large discs and extensive webbing and males with single subgular vocal sacs and with prepollex; color of bones in life variable ............................................... 6

6 a A protuberant prepollex (Fig. 501a) present in adult males or in both sexes; vocal sac in adult males absent or, if present, single, subgular; large to very large species, SVL

Fig. 500. Trachycephalus venulosus (Scotland Halfboon, Belize, 10 m). Photo: G. Kohler

Fig. 501. a. Hand of Plectrohyla matudai (Finca Irlanda, Chiapas, Mexico, 1050 m); note protuberant prepollex (arrow); b. lateral head of Plectrohyla sagorum (Mapastepec, Chiapas, Mexico, 2014m).

Photos: a. G. K6hler; b. A. Ramirez-Veldzquez

of adult males and females >40 mm, ofien much larger .............................................. 7

b Males and females without a protuberant prepollex, or, if a prepollex is present, then SVL c40 mm; vocal sac, if present, single or paired, subgular or lateral; size variable, but usually smaller than indicated above . .................................................................. 10

7 a Extensive fleshy fringes, usually with a scalloped edge, along posteroventral margin of lower arms and legs .... Emomiohyla

b No extensive fleshy scalloped fringes along posteroventral margin of lower arms and legs .......................................................... 8

8 a Skin on dorsum smooth, never thick, glandular, or tuberculate; with a middorsal dark line, or, if no such line present, then webbing red in life .................... Hypsiboas

b Skin on dorsum tuberculate, or, if smooth, then skin thick and glandular; never with a middorsal dark line or red webbing in life 9

w, P. h:- - ~ - ; .::.

rr?> &.*..*

La*;;

Fig. 502. Hyloscirtus palmeri (Guayacb, Fig. 504. Dendropsophus subocularis CRARC, Costa Rica, 550 m); note mental gland (Rionegro, Santander, Colombia, 831 m). b row) . Photo: G. Kohler Photo: C. Hernhdez-Jaimes

- Fig. 503. Ventral surfaces of Isthmohyla rig. sus. ~oloratlon or thigh in Isthmohyla insolita. Photo: J. R. McCranie lancasteri (Guayach, Vueltade Queque, Costa

Rica). Photo: G. Kohler

9 a Head not conspicuously large and deep; discs very large, width of disc on Finger I11

........ nearly twice the length of tympanum ........................................ cllaraela

b Head conspicuously large and deep (Fig. 501b); discs moderately large, width of disc on Finger I11 only slightly larger than the length of tympanum; vocal sac absent or, if present, single, median, subgular ............... ................................................... Plecbhyla

10a Bones pale green in life; a circular whitish mental gland (Fig. 502) in adult males and some adult females; vomerine tooth patches acute, angulate, extending posterior to choanae; a low inner tarsal fold present; toe webbing extensive, modal formula I11 1 - 2 IV or I11 1+ - 2 IV; dorsum usually pale or olive green, sometimes pale brown (yellowish white in preservative) ....... I.ipl08cirt~~

b Bones white in life; no circular whitish mental gland; vomerine tooth patches

transverse, not extending posterior to choanae; inner tarsal fold present or absent; toe webbing variable; dorsal col-

...................................... oration variable 11 lla Chin and throat boldly marked with a con-

spicuous pale (yellow to white in life, dirty white in preservative) central area, flanked by solid, dark pigment (Fig. 503); dorsum with numerous wart-like tubercles; SVL of adult males 30-36 mm, of adult females 34-38 mm bthmohyh (in park 1 insolit.)

b Chin and throat not as described above; dorsum with or without numerous wart-

.... like tubercles; SVL of adult variable 12 12a Posterior surface of thigh contrastingly

barred black and yellow in life (black and white or cream in preservative) (Fig. 505); one or more dark blotches in groin, SVL of adult males 27-34 mm, of adult females

.................................................... 31-38 mm ............. bthmohyla (in park I. lancasteri?

b Posterior surface of thigh without dark bars; no dark blotches in groin, although small punctations can be present; SVL of adult variable ......................................... 13

13a Dorsal surfaces of head, body, and limbs with large, tubercular spines; adult males with small black spines on thumb; SVL of adult males 26-35 mm, of adult females 3 1 4 0 mm Iethmo4vla (in part: I. d.ypa)

b Dorsal surfaces of head, body, and limbs smooth, granular, or tuberculate, but without large, tubercular spines; nuptial pads of adult males with or without small black spines; SVL of adults variable ............... 14

14a A distinct pale (white or yellowish) lateral or dorsolateral stripe between levels of axilla and groin, or at least between levels

............................. of midbody and groin 15 b No pale lateral or dorsolateral stripe

between levels of axilla and groin, nor between levels of midbody and groin .... 23

16a A dorsolateral pale stripe extending from posterior corner of eye along upper

edge of tympanum (stripe can be indistinct or missing on head and anterior body) and then to sacral area or groin ................... 16

b A lateral pale stripe extending from lower posterior corner of eye or from axilla to groin ........................................................ 20

ria Posterior surface of thigh unpigmented except for a yellow spot bordered by black . . Dendropophue (in part: D. auhocularie)

b Posterior surface of thigh uniform yellow or pale brown .............................................. 17

17a Dorsolateral white lines broad .............. 18 b Dorsolateral white lines narrow ............ 19

18a A distinct axillary membrane present; webbing of fingers thin; a low inner tarsal fold present; flanks yellow in Life; posterior surface of thigh uniform yellow in life; SVL of adult males 19-21 mm, of adult females 19-22 mm . . ZTa.ld yla (in part: T. pi&)

b Axillary membrane absent; webbing of fingers thick; inner tarsal fold absent; flanks dark brown; posterior surface of thigh pale brown in life; SVL of adult males 33.5-38.7 mm, single known adult female 40.5 mm .. .................. Ist&nohyla (in part: I. graceae)

19a Fingers either essentially lacking webbing; venter peppered with brown flecks; SVL of adult males 30-34 mm, of adult females 34-37 mm ....... Iethmolwla angustilineata

b Fingers with basal webbing; venter immaculate; SVL of adult males to 26 mm, of

... adult females to 28 mm Dandropsophus ........................ (in part. D. mbertmertensi3

b. external. lateral. ~ a i r e

Fig. 506. Vocal sacs in hylid frogs: a. Smilisca baudinii (El Cmcero, Nicaragua); b. Trachy- cephalus venulosus (Guaquira, Venezuela); c. Scinax staufEeri (San Juan de Dios, Nica- ragua); d. Dendropsophus ebraccatus (Green Hills, Belize). Photos: a.4. J. Sunyer; d. G. Kahler

Fig. 507. Nuptial pads in hyline frogs: a. Smilisca b a u W , b. S. sordida; c. Isthmohyla pseudopuma; d. Ptychohyla euthysanota; e. P. hypomykter, f. Isthmohyla picadoi; g. I. zeteki.

Photos: G. Kahler

20a Adult males with a patch of small black spines on base of thumb (Fig. 507d,e); a vertical rostral keel present .... Ryd106gla (in part: P. euthyrranota, P. panch013

b Adult males with a brown non-spinous pad on base of thumb (Fig. 507b,c); no vertical rostral keel ........................................ 21

21a No continuous pale lateral stripe from head to groin, although a pale lateral stripe can be present between midbody and groin; vomerine tooth patches positioned near posterior margin of choanae ....................... .................. lethmohyla (in part: I. de-1

b A continuous pale lateral stripe from head to groin or from axilla to groin present; vomerine tooth patches positioned between choanae ................................................... 22

2% No suborbital pale spot, labial stripe, ifpre- sent, not expanded below eye and termi-

..................... nating anterior to tympanum ................... la (in part. P. 1egk13

b A suborbital pale spot present or pale labial stripe expanded below eye, or, if a suborbital pale spot is absent, then a continuous white stripe from tip of snout along upper lip and below tympanum to groin .............. Duellrnanohyla (in park all species of

I genw except D. soralia) 23a An axillary membrane present, extending

at least one-fourth distance towards elbow; iris red in life; dorsal surfaces of head and body with a lichenous pattern of some shade of green in life (a pale lichenous pattern on a dark brown to dark gray background in preservative); posterior surface of thigh lemon yellow with dusting of brown pigment in life (cream with brown flecking in preservative); SVL of adult males 28-32 mm, of adult females 35-38 mm .... Duellmanohyla (in part: D. saralia)

b Axillary membrane absent, or if present, then iris not red in life and color pattern of dorsal surfaces of head and body and of

posterior surface of thigh not as described ................ above; SVL of adult variable 24

24a Large frogs, SVL of single known adult female 84 mm; fingers about two-thirds webbed (modal formula I11 2 - 1 1/2 IV) with large discs, width of disc on Finger I11 slightly greater than tympanum length; toes nearly fully webbed (modal formula 111 1 - 1 1/2 nn ........................................a*.

.... -la (in part. P. d e n t h u m ) b Combination of characters not as above,

particulary, if large frogs, then with less finger and toe webbing .......................... 25 r

%a Adult males with a patch of small black spines on base of thumb (Fig. 507d,e); prominent ventrolateral gland present or absent in adult males

B ............................ 26 1 2 b Adult males with a brown non-spinous pad

or a patch of very tiny spinules on base of thumb (Fig. 507b,c); no prominent ventro-

................. lateral gland in adult males 27 26a Adult males with a patch of ten black

spines on base of thumb; no ventrolateral gland in adult males; modal toe webbing formula I11 2- - 3 IV; SVL of adult males 22-23 mm, of adult females 24-26 mm .... ............ lethm06gla (in park I. melacaena)

b Adult males with a patch of more than 30 black spines on base of thumb; ventrolateral gland usually well-developed in adult males; modal toe webbing formula 111 1+ - 2 IV to I11 1 - 2- W, SVL of adult males 30-39 mm, of adult females 35-46 mm ptscaohyla (in part: P. &pomy&my P. macrotpmpanum, P. spinipoUe~)

27a Small frogs, SVL of adult males 21-32 mm, of adult females 24-35 mm; overall coloration tan, yellowish brown to yellowish green, often with accumulation of dark pigment in loreal region; finger webs vestigial; toe webbing reduced (modal toe webbing formula I11 1 213 - 3 IV to I11 2 - 3 IV); axillary membrane absent; vomerine

teeth in linear series entirely posterior to choanae; tympanum partly or completely concealed; heavy muscular development in temporal region; adult males with white

................................ nuptial pads (Fig. 507f;g) ...... (inpart: I zelkdd, I.picadrn3

b Combination of characters not as above, particularly, if small h g s with overall coloration tan, yellowish brown to yellowish

- green, then either more finger and toe web- ................. bing and tympanum distinct 28

28a A well-developed ax ihry membrane present, extending along upper arm more than one-half distance toward elbow (Fig. 508); SVL of adult males 1 8 4 5 mm, of adult females 2 3 4 2 mm; posterior surface of thigh patternless yellow or red in life, much paler than dorsal surface of shank;

................. no mottling in groin or flank 29 b Axillary membrane absent, or, if present,

abbreviated, not reaching beyond halfway to elbow; SVL of adults variable; coloration of posterior surface of thigh, shank and groin variable ......................................... 30

29a Usually a dark stripe from nostril along canthus rostralis to eye; u s u d y some narrow markings on dorsum; posterior surface of thigh yellow in life; SVL of adult males

..... 18-25 mm, of adult females 2 3 3 5 mm Dandropsophm (in part: D. ebraccatue; D. m i c m c e ~ m , D. phlebudes)

b No dark stripe from nostril along canthus rostralis to eye; usually no distinct markings on dorsum; posterior surface of thigh red in life; SVL of adult males 33-46 mm,

......................... of adult females 3 8 4 7 mm .................. !f!W+la (in park T. loquax)

30a Toes with reduced webbing, modal formula 111 2 - 2 l/3 W, no finger webbing; a dark brown face mask below cathus rostralis preseqt; upper surfaces of thigh and shank without dark transverse bars; inner tarsal

fold present; a dark brown lateral stripe present from posterior eye through tympanum to groin; SVL of adult males 29-40

....... mm, of adult females 3 1 4 1 mm qvla b Toes usually with more webbing than indi-

cated above, modal formula at least I11 1 314 - 2 213 IV, usually more webbing; degree of finger webbing variable; no dark brown face mask below canthus rostralis present, or, if present, then upper surfaces of thigh and shank with distinct dark transverse bars; a dark brown lateral stripe present or absent; inner tarsal fold present or absent; SVL of adult variable 31

31a A dark brown or black postorbital mark encompassing tympanum present, usually continuing as dark lateral band a t least to level of axilla (Fig. 509); vocal sacs in adult males external, subgular, paired (Fig. 506a); SVL of adult males 40-75 mm, of

............................. adult females 47-90 mm Smilieca (in part: S. baudihii, S. cyanostz'c- ta, S. Paaeota)

b No dark brown or black postorbital mark encompassing tympanum present; vocal sacs in adult males variable; SVL of adults

................................................... variable 32 32a Posterior surface of thigh with discrete

pale (yellow in life, white in preservative) spots; venter with extensive dark pigment or not; finger webbing basal or absent .............................................. ......-&la ............ (in park I.pmctip, I. xanthosticta)

b Posterior surface of thigh patternless or with pale and dark mottling, never with a series of discrete pale spots; venter without extensive dark pigment; finger webbing variable ................................................... 33

33a Venter usually peppered with dark spots (Fig. 590); snout acute in profile; tympanum hidden, length no more than one-third eye length; modal toe webbing formula I11

Fig. 508. Dendropsophus ebraccatus; note axil- Fig. 509. Smilisca baudinii (both from Mazate- lary membrane (arrow). Photo: G. Kohler nango, Guatemala, 360 m). Photos: G. Kohler

1 l/4 -2 213 W, SVL of adult males 2934 ................ mm, of adult females 33-37 mm

3kthuo4pla (in part: I. r iv7 lkb) ................ Venter immaculate or with Erie brown stippling; snout shape variable; tympanum distinct or hidden, length variable; modal toe webbing formula variable; SVL of adult

................................................... variable 34 Finger webbing vestigial or absent; white stripes present on upper lip, forearm, and tarsus; dorsum with two elongate blotches extending fmm head to sacrum, usually anastomosing at one or several points along midline; transverse supracloacal white stripe present; posterior surface of thigh in lilfe and in preservative dark brown; SVL of adult males 32-38 mm, of adult females 40-46 mm ............................. ......................... S$lilieca (in part: S. puma) Combination of characters not as above, particulary, if webbing vestigial, then dorsal coloration not as above ..................... 36 Small species, SVL of adult males 20-30 mm, of adult females 18-36 mm, a mdi- mentary or small (extending along upper arm one-fourth to one-half distance toward elbow) axillary membrane usually present .................................................................. 36 Larger species, SVL of adult males >30 mm, of adult females >36 nm, or, if adult males <31 mm and adult females <37 mm, then without a trace of an axillary membrane ....................................................... 38 Toe webbing reduced, modal formula 111 1+ - 3' IV; SVL of adult males 26-30 mm, of adult females 27-36 mm ..... Brnmelloil~h Toes extensively webbed, modal formula I11 1 -2 lVto I I I 1 - 2 l/2IV;SVLofadults variable .................................................... 37 Snout nearly rounded in dorsal aspect, canthus rostralis rounded to nearly angular; SVL of adult males 27-35 mm, of adult

.................................... femdes 34-42 mm ....... P@&*a (in part: P. salved-) Snout acuminate or pointed in dorsal view (Fig. 544-551); canthus rostralis usually angular; SVL of adult males 2033 mm, of

............................. adult females 18-36 mm .............. ..E.arodanta lin park E. bhcata, E. catracha, E. ch imdqq E. mmidmdal A discontinuous, irregular, white or cream stripe extending from the lower posterior corner of eye to groin, snout acuminate or pointed in dorsal view; canthus rostralis usually angular; axillary membrane extending about half of upper arm; SVL of adult females 33.1-37.5 mm (males unknown) I h m d h t a (in park E. p a r ! 3

b No white or cream stripe from eye to groin; axillary membrane, if present, extending no more than one-fourth of upper arm; snout nearly rounded in dorsal aspect; canthus rostralis rounded to nearly angular;

........................... SVL of adults variable 39 39a Tarsal fold distinct; subarticular tubercles

under Fingers 111 and IV single, rounded; modal toe webbing formula 111 1 - 2' IV to III 1 l/4 - 2 l/4 IV; adult males with paired, internal, subgular vocal sacs; groin and posterior thigh reticulated with dark brown surrounding cream ar pale blue flecks; SVL of adult males 31-54 mm, of

............................. adult females 44-64 mrn .......... SiniIieca (in park 5. rdla, s. EmIdida)

b Tarsal fold weak or absent; subarticular tubercles under Fingers Ill or IV often double or bzd; modal toe webbing formula variable; adult males with single, median or bilobate, internal, subgular vocal sacs; coloration of groin and posterior thigh not as described above; SVL of adult males 2745 mm, of adult females 33-52 mm 40

40a Dorsum lime green with scattered pale green dots in life; flanks with a narrow zone of reticulatian where dorsal and ventral coloration meet; adult males with a patch of tiny black spines on base of thumb; SVL of adult males 29-33 mm, of

............................. adult females 41-51 mm ....... ptscho4rh (in part: P. sauc&mcmCie)

b Dorsum pale tan, dull brown, or reddish brown with or without scattered dark brown blotches in life; flank coloration not as described above; adult males with a brown non-spinom pad on base of thumb;

........................... SVL of adults variable 41 41a Larger species, SVL of adult males 37-45

................ mm, of adult females 41-52 mm .......................................... ..........am ........... (in part: I. Mizcata, I, peeudamma)

b Smaller species, SVL of adult males 27-35 mm, of adult females 3342 mm ........... 42

42a Dorsum and upper surfaces of limbs granulate to tuberculate; modal toe webbing

....................................... formula 111 1-2 IV ........................ lidlrn*. (in park I. ti=)

b Dorsum and upper surfaces of limbs smooth; modal toe webbing formula I11 1 l/2 -2 IV ....................................................... ........ &mhbyIa (in park P. e a l v y i d d )

Fig. 510. Anotheca spinosa (Rancho Naturalists, near Platanillo, Costa Rica, 900 m). Photo: G. Kohler

Anotlieca is a mo110typic genus, and its only species, A, spinosa, is a large, strik- ingly patterned frog with sharply pointed and knob-like projections on the head. It has a patchy geographic distribution and no records are available from Belize, Guatemala, El Salvador, or Nicaragua (MEYER & FOSTER 1996, CAMPBELL 1998, K O ~ R 2001, K ~ E R et al. 2006). Anotheca spinos'a is a diflicult to find, nocturnal, arboreal frog (TAYLOR 1954, DUELLMAN 1970, SESSIONS 19783. Even though t-he males of this species lack vocal slits and a vocal sac, they produce vocalizations (DUELLMAN 1970, 2001). The male calls from water-filled cavities such as treeholes, open bamboo internodes, or

bromeliad leaf axils (JUNGFER 1996). The call was described as a loud "boop-boop- boop* (DUELLMAN 1970, JUNGFER 1996). This species uses bromeliads, wate~fflled tree cavities, and water-filled bamboo internodes for breeding (TAYLOR 1954, ROBINSON 1961, DUEW 1970, J U N G ~ R 1996). Afker egg deposition, the male dis- appears, but the female continue to visit the developing tadpoles a t intervals of about 5 days (range 1-14 days) to deposit unfertilized (nutritive) eggs into the water body, with the eggs mostly being grabbed and consumed by a tadpole as soon as they were extruded &om the female's cloaca (JUNGFER 1996, V m & CALDWEZL 2009). The tadpoles metamorphose aRer 60-136 days at a froglet size of 26-28 mm, and then the female approaches a calling male to breed again (JUNGFER 1996).

ETZ

Bromeliohyla I The genus Bromeliohyla was introduced by FAJYOVICH et al. (2005) for the species of the Hyla bromeliacia group of DUELLMAN (1970, 2001). Two species are assigned to this genus, only one-of which, B. bromelia- I cia, occurs east of the Isthmus of Tehuantepec (DUELLMAN 2001). Bromelio- hyla bromeliacia is a small nocturnal cloud forest frog. that lives in arboreal tank I bromeliadi (MCCRANIE & WILSON 2002). Males call from arboreal bromeliads or infrequently from vegetation such as on Fig. 513. Bromeliohyla bromeliacia (Quebrada

top of leaves of branches outside of C0ph7 1300 m).

bromeliads; the call consists of five or six Photo: G. Kiihler

soft notes repeated at intervals of 45-70 seconds (DUELLMAN -1970, MCCRANIE & WILSON 2002). The female deposits the eggs into water-filled arboreal bromeliads and the tadpoles complete their development within the bromeliads (STUART

I 1943b, DUELLMAN 1970, MCCRANIE &. WILSON 2002). In Honduras, eggs and tadpoles were found in bromeliads from May through August (MCCRANIE & WILSON 2002).

Bromeliohyla bromeliacia (SC-T 19331, Field Mus. Nat. Hist. Publ., Zool. Ser., 20: 19; M e locality: "mountains west of San Pedro G-san pe& Sula, Departamento de Cort6~1, Honduras. Altitude 4500 feet? Males to 29.5 mm, females to 32.7 mm. On the Atlantic versant from northwestern Chiapas, Mexico, across central and eastern Guatemala, the Maya Mountains of Belize to northw qonduras, 350-1790 m elyyation.

Fig. 514. Brorneliohyla bromeuacia (Eel\. Cusuco, CortBs, Honduras, 1355 m).

Photo: J. Kolby

Further Reading DUELLMAN 1970, WTINEZ-CORONEL et al. 1995, MCCRANIE & WILSON 2002

Fig. 515. Charadrahyla chaneque (extreme northwestern Chiapas on Hwy 190, Mexico, ca. 700 m). Photo: L. Gray

Charadrahvla - The genus Charadrahyla was proposed by FANOVICH et al. (2005) to accommodate the species of the Hyla taeniopus group of DUELLMAN (1965, 2001). DUELLMAN (1965) published a review of this group of fiogs, at that time containing only two recognized species, C. taeniopus and C. chaneque. Currently, six species are recognized in the genus Charadrahyla (CAMPBELL et al. 2009). Charadrahyla chaneque fiom the cloud forests of western Chiapas is the only member of the genus known to occur east of the Isthmus of Tehuantepec (CAMPBELL et al. 2009). All members of this genus are relatively large, stream-breeding frogs that are found in cloud forest or humid pine-oak forest in central and southern Mexico at elevations of 1100-2200 m above sea level.

Charadrahyla chaneque (DUELLMAN 1961~1, Herpetologica 17: 1; type localiw. "stream above (6.2 kilometers by road south of) R a m Mescalapa, Chiapas, Mexico (elevation 1890 meters)." Males to 60.7 mm, females to 79.3 mm. Known from only three localities in northern Chiapas and eastern Oaxaca, Mexico, 800-!$200 melevation:

- I F . 2*. -

Further Reading DUELLMAN 1961c, DUELLMAN 1965,1970, PEREZ- HIGAREDA 1981, MENDELSON & CAMPBELL 1999, CAMPBELL et al. 2009

Dendropsophus 1968). Amplexus is axillary and the eggs of this species are laid on or near the surface of the water, where they are usually

The generic name Dendropsophus was resurrected from synonymy with Hyla by FAJYOVICH et al. (2005). As currently understood, this genus contains 91 species, all except five being distributed in South America (FROST 2010). The five Central American species of Dendropsophus are moderately to very small treefrogs that are widespread and common in the humid lowlands. Throughout the rainy season, the males can be heard calling from tall grass, reeds, or sedges at the edge of water, or while perched on emergent vegetation; they produce insect-like calls, those of D. ebraccatus sounding like a prolonged "wreeeek" or "creeeek", those of D. microcephalus, D. phlebodes, D. robertmertensi like "creek-eek-eek-eek", and those of D. subocularis like a short trill (DUELLMAN 1970, SAVAGE 2002). Calling usually begins at dusk and continues until near dawn; males usually sing in large choruses and they will respond to conspecific males and males of other species within a vocalizing group (SCHWARTZ & WELLS 1985, SAVAGE 2002). Preferred breeding areas are temporary ponds, roadside ditches, flooded areas, and marshes (DUELLMAN & FOUQUETTE

attached to vegetation near the water surface (DUELLMAN 2001, SAVAGE 2002). The tadpoles of the Central American ~ e n d r o ~ s o ~ h u s have beaks but lack denti- cles (SAVAGE 2002). In D. ebraccatus, tadpole development <s completed within 4-6 weeks (SAVAGE 2002).

Dendropsophus ebraccatus (COPE 18741, Proc. Acad. Nat. Sci. Philadelphia 26: 69; type locality: "region of Nicaragua." DUNN & STUART (1951:58) noted that the type specimen is labeled "Machuca", and that it surely came from "Nicaragua along the San Juan River." Males to 27.8 mm, females to 36.5 mm. Atlantic versant of southern Mexico across northern Guatemala and Belize to Colombia and northwestern Ecuador, sea level to 1600 m elevation.

Dendropsophus rnicrocephalus (COPE 18861, Proc. Am. Philos. Soc. 23: 281; type locality: "along a mountain stream in the department of Chiriqui", Panama; at the time of the description "Chiriquin included both Atlantic and Pacific versants of extreme western Panama according to MYERS (1982). Males to 27 mm, females to 32 mm. On the Atlantic versant from southern Veracruz and northern Oaxaca, Mexico, to Panama; also widely distributed in South America (Colombia, Vene-

Fig. 516. Dendropsophus ebraccatus (male; La Fig. 517. Dendropsophus ebraccatus Itemale; Selva, Heredia, Costa Rica, 70 m). Bartola, Rio San Juan, Nicaragua, 30 m).

Photo: G. Kohler Photo: G. Kohler

Hylidae

zuela, Brazil, Trinidad, and Tobago), sea level to 1300 m elevation. Hyla underwoodi BOULENGER 1899 is regarded as a synonym of D. microcephalus according to SAVAGE & IIEYE (1970).

Dendropsophus phlebodes (STEJNEGER 19061, Proc. U.S. Natl. Mus. 30: 817; type locality: "San Carlos, Costa Rica." SAVAGE (1974) corrected it to "Boca de Arenal, Cant6n de San Carlos, Provincia de Alajuela; 55 m", Costa Rica. Males to 23.6 mm, females to 27.8 mm. On the Atlantic versant from southeastern Nicaragua, including Isla de Maiz Grande, to eastern Panama; and on the Pacific versant from central Panama to northwestern Colombia, and barely across the continental divide into northwestern Costa Rica, 20-700 m elevation.

rig. 313. ~enaropsophus microcephalus (Volch Mombacho, Nicaragua, 1100 m). Photo: G. Kijhler

Dendropsophus robertmertensi (TAYLOR 1937b), Proc. Biol. Soc. Washington 50: 43; type locality: "near Tapachula, Chiapas", Mexico. Males to 26.4 mm, females to 28.1 mm. Pacific versant from extreme eastern Oaxaca to central El Salvador; also in the Cintalapa Valley in southwestern Chiapas, sea level to 750 m elevation.

Dendropsophus subocularis (DUNN 1934), Am. Fig. 520. Dendro~Oflus micmoeMus (Hopby M ~ ~ . N ~ G ~ . 747: 2: tvDe localitv: u ~ i o Stann Creek Belize, 5 m). Photo: G. Kohler Chucunaque, first creek a6;ve entrance of Rio Tuquasa", Provincia DariBn, Panama. Males to 23.1 mm, females to 26.1 mm. Eastern lowlands and cordilleras in the Dari6n Province of Panama, and in Colombia, sea level to 800 m elevation.

I Fig. 521. Dendropsophus phlebodes (Turrialba, Cartago, Costa Rica, 600 m). Photo: G. Kohler

Fig. 518. Dendropsophus robertmertensi (road I from Hacienda San Cayetano to San Jose El Fig. 522. Dendropsophus subocularis Idolo, Suchitepbquez, Guatemala, 100 m). (Rionegro, Santander, Colombia, 831 m).

Photo: G. Ktihler Photo: C. Hernhdez-Jaimes

Key to hdropsophus 1 a Posterior surface of thigh unpigmented

except for a yellow spot bordered by black; SVL of adult males to 23 mm, of adult females to 26 mm ........................................ ........................ Dendropsophue subocul&

b Posterior surface of thigh uniform yellow; SVL of adults variable ............................. 2

2 a Pale dorsolateral lines present, extending to groin or nearly so; SVL of adult males to

............. 26 mm, of adult females to 28 mm .................. Dendrupsophus r o l l e r t n l ~ ~

b Pale dorsolatera1 lines, if present, not .................................... extending to groin 3

3 a Head broad; scalloped dark mark on dorsal surface of shank; dorsum without marking or marked with brown blotches, or hour-

mark; SVL of males Fig. 523. Eggs of Dendropsophus ebraccatus 23-27 rnm, of adult females 30-35 mm ..... ......................... Dendrupsophus ebraocatm ( G u a ~ a c b p CRARC* Cartago* Rita*

b Head narrow; narrow transverse dashes or 550 m). Photo: G. Kijhler

longitudinal dark line on dorsal surface of shank, dorsum patternless, with brown dashes, longitudinal dark lines, or reticulations; SVL of adult males 18-25 mm, of

......................... adult females 23-31 mm 4 4 a Dorsolateral dark stripe indistinct, present

only above tympanum and insertion of foreleg; dorsum with narrow lines and dashes, sometimes interconnecting; SVL of adult males 19-24 mm, of adult females 23-28 mm ........ Dendrupsophurr pldebodes

b Dorsolateral dark stripe distinct, bordered above by a narrow pale line, extending .from eye at least to point level with elbow when arm folded along side, often reaching to level of groin, and usually bordered below by a narrow white line in life; dorsum patternless or with a pair of narrow longitudinal dark lines, or with interconnecting lines and dashes; SVL of adult males 18-25 mm, of adult females 24-31 mm ............ Dendrupsophus ~~~~CruCephalus - I D. microcephalus

D. robertmertensi 1 Dendropsophus

A D. phlebodes D. subocularis

n -

l?urther Reading DUELLMAN & F O U Q U E ~ 1968, DVELLMAN 1970, SCHWARTZ & WELLS 1984, ROBLE 1985, SCHWARTZ & WELLS 1985, BACKWELL & JENNIONS

- 1993, DUELL~~IAN 2001

wid-

- - -

Duellmanohvla

The genus Duellmanohyla was described by CAMPBELL & SMITH (1992) to accommodate a group of eight small to moderately small Central American hylid frogs with tadpoles having large pendant mouths and a certain tooth combination (212,213, or 313 short tooth rows). All species in this genus have more or less distinct red eyes. Only one species, D. ignicolor, is extralimital, occurring in northern Oaxaca, Mexico (DUELLMAN 2001).

The species of Duellmanohyla are nocturnal, arboreal inhabitants of small, shallow, cascading mountain streams or seeps draining into streams in humid forests (SAVAGE 2002). During the day, they seek shelter in leaf axils of arboreal bromeliads and terrestrial aroids (DUELLMAN 1970, SAVAGE 2002). Males call from dense vegetation several meters away from the stream, from plants a t the edge of or overhanging the stream; the advertisement call of D. uranochroa was described as sounding like a series of bell-like notes ("boop- boop-boop-boop"), that of D. d o c u l i s as a short rattle or "scraape" consisting of three or four short notes, that of D. lythrodes as a weak bell-like "pings' in bursts of five to seven notes, that bf 5. soralia as a single low-pitched "peep", repeated every 20-30 seconds, and that of D. chamulae and D. schmidtonun as a series of short, raucous low-pitched notes (DUELLMAN 1970, MCCRANIE & WILSON 2002, SAVAGE 2002). The tadpoles live in pools of the stream where they adhere to pebbles, stones, and rocks in the relatively quiet water; when disturbed, they seek refuge amidst debris on the bottom of th6 pools (DUELLMAN 1970). Although these species are apparently active throughout the year, breeding activity seems to peak in the early to middle rainy season (May to July); some species such as D. uranochroa are reported to have a second reproductive peak in the late rainy season (DUELLMAN 1970, MCCRANIE & WILSON 2002, SAVAGE 2002).

Fig. 524. Duellmanohyla chamulae (6.2 km S Ray6n Mescalapa, Chiapas, Mexico).


Fig. 525. Duellmanohyla lythrodes (Rio Chan- guinola near Quebrada Guabo, Bocas del Toro, Panama). Photo: C. W. Myers (Courtesy of the Biodiversity Institute, University of Kansas)

Duel2manohyla chamulae (DuEuMAN 1961b1, Univ, Kansas Publ. Mns. Nat. Eat. 13: 364; type localiQ: 'a stream above (6.2 kilometers by r o d 8 o d ) Ray6n Mescalapa, Chiapas,; Mgxico (1690 meters)." Males to 80.5 mm,, females to 31.8 mm. North- sIopes of tha. central highkh of Chiapas, Mexico, from' Jitotol to Soluschiapa, 350-1700 m elevation. Duellmanohyla fgthrodes (SAVAGB 1968b),! Bd. S. California Acad. Sci. 67: 1; type locali~ ! ty: "Cash Rica: Pmvhcia de Lim6n; Cant6n dm \

Lim6n: Alta Talamanca: 21 km SW Amubri a6 confluence of Rio Lari and Rio Dipari, 800 m"; elevation is actually 440 m according to SAVAGE (1974). Males to 32.6 mrn, females unknown. Atlantic versant in southern Costa Rica and adjacent northwestern Panama, 170440 m elevation. Duehanohyla mfioctd.is (TAYLOR 1962b), Univ. Kansas Sci. Bull. 35: 827; type locality:

"Isla Bonita (American Cinchona Plantation), eastern slope of Volch Pois, [CantBn de Alajuela, Provincia de Alajuela,] Costa Rica", 1200 m elevation. Males to 30 mm, females to 39.9 mm. Caribbean and Pacific slopes of the mountains of Costa Rica and adjacent Panama, 460-1580 m elevation. DueLlmanohyla salvavida (MCCRANIE & WILSON 19861, Proc. Biol. Soc. Washington 99: 51; type locality: "Quebrada de Om (15"38'N, 86"47W), elevation 880 m, tributary of Rio Viejo, south slope of Cerro BGalo, Cordillera

females to 34.3 mm. Cordillera de Nombre de

de Nombre de Dios, Departamento de Fig. 526. Duellmanohyla rufioculis ( Atlhtida, Honduras." Males to 28.0 mm, Center,Alajuela, Costa Rica, 460 m).

Photo: G. Kohler Dios and Montafia Macuzal (Departamentos de Atlintida and Yoro), north-central Honduras, 90-1400 m elevation. Duellmanohyla schmidtorum (STUART 19541, Proc. Biol. Soc. Washington 67: 170; type locality: "Finca El Porvenir (17 air-line kilometers due west of San Marcos), Department of San Marcos, Guatemala. Elevation, unknown but El Porvenir terrain includes elevations ranging from about 500 m. to possibly 2000 m." Males to 32.8 mm, females to 38.3 mm. Extreme eastern Oaxaca (Chimalapas) and extreme southwestern Chiapas, Mexico, as well as adjacent Guatemala, 500-2200 m elevation. Fig. 527. Duellmanohyla rufioculis (Rancho

~ ~ ~ l l ~ ~ ~ ~ h ~ l ~ soda (wlu0~ & M ~ c ~ ~ ~ ~ ~ Naturalista, Costa Rica, 900 m).Photo: G. K6hler 1985), Herpetologica 41: 133; type locality: "Quebrada Grande (15"05'N, 8B055W), 1370 m elevation, Sierra de Omoa, Depto. Copin, Honduras." Males to 32.3 mm, females to 37.7 mm. Sierras de Omoa and Espiritu Santo in northwestern Honduras and northeastern Guatemala, 40-1570 m elevation. Duellmanohyla uranochroa (COPE 1876), J. Acad. Nat. Sci. Philadelphia Ser. 2,8: 103; type locality: "near Sipurio", Cant6n de Talamanca, Lim6n Province, Costa Rica. Males to 36.8 mm, females to 39.9 mm. Cordilleras of Costa Rita and western Panama, 70-1740 m elevation. Flg. o m . uueumanonyla soralia (Quebrada

Grande, Honduras, 1300 m). Photo: G. Kohler

D. salvavida D. charnulae

Key to Du-ela 1 a Dorsum with a reticulate pattern of white,

cream, green, or turquoise blotches on a brown, reddish brown or dark grayish background; no white lateral stripe; SVL of adult males 28-32 mm, of adult females 35-38 mm ............. D u h e k &

b Dorsal color pattern not as above; a white lateral stripe usually present; adult SVL

..................................................... variable 2 2 a White labial stripe usually not expanded

below eye; undersurfaces of foot not suffused with dark pigment; dorsum leaf green in life (bluish to purplish in preservative); an inner tarsal fold present; distance from eye to nostril equals eye length ........................ D u w l a lrranochraa

b White labial stripe expanded below eye; undersurfaces of foot heavily suffused with dark pigment; dorsal coloration variable; an inner tarsal fold present or absent; relative eye to nostril distance variable ........ 3

3 a Distance from eye to nostril equals eye length; an inner tarsal fold present; dorsum leaf green in life (bluish to purplish in preservative) .... Dueltnanwfa salvavida

b Distance fkom eye to nostril less than eye length; inner tarsal fold weak or absent; dorsum brown, greenish brown, or leaf green in life (brown, bluish, or purplish in preservative) ............................................ 4

4 a No ventrolateral glands in males; a brown nuptial pad on base of thumb in males; rel-

............. ative size of tympanum variable 6 b Ventrolateral glands present in males; no

nuptial pad on base of thumb in males; .... tympanum length >50% of eye length 6

6 a Tympanum length <50% of eye length; throat and venter white in life; posterior surface of thigh usually suffused with dark pigment; no inner tarsal fold ...................... ............................ ouellman*k docu l iB

b Tympanum length >50% of eye lengt6; throat and venter yellow in life; posterior surface of thigh not suffused with dark pigment; a weak inner tarsal fold ................... ............................ Duel&anaky.fa &hrude#

6 a Dorsum predominately green in life (bluish to purplish in preservative); venter yellow in life; a mental gland present in males .... ........................... Duellmanwla chamulae

b Dorsum brown in life (and in preservative); venter white in life; no mental gland in males ........... -la ~chmidtmum

Fig. 529. Duellmanohyla salvadda (Liberacibn, Atlhtida, Honduras, 1050 m).

Photo: J. H. Townsend

Fig. 580. Duellmanohyla schmidtarum (Finca La Paz, San Marcos, Guatemala).

Photo: W. E. Duellman ( C o h y of the BiodiversiQ Institute, U n i v e n a a )

Fig. 531. Duellmanohyla uranochroa (Palo Seco, Ngobe Bugl6, Panama, 1060 m).

Photo: A. Hertz

F'urther Reading DUELLMAN 1961b, SAVAGE 1968b, DUELLMAN 1970, WILSON & MCCRANIE 1985, 1986, MCCRANIE & WILSON 1990, DUELLMAN 2001

221

- Ecnomionyla -----

Twelve species of moderately small to very large frogs are currently assigned to Ecnomiohyla, a generic name proposed by F ~ r v o v r c ~ et al. (2005). All Central American taxa except one (i.e., E. m i o t p - panurn) are large nocturnal canopy species referred to as fringe-limbed treefrogs; they are rarely seen or collected (SAVAGE 2002). Fringe-limbed treefrogs have large hands and feet and with fleshy fringes along the outer margins of forearm and foot (DUELLMAN 1970). I t was hypothesized that the thick, rugose skin and the co-ossifica- tion of the dorsal head surface in some species may provide some protection against desiccation in the canopy (DUELLMAN 1970). Unlike most frogs, males of most fringe-limbed treefrogs are larger than females (SAVAGE 2002). Very little information on the natural history is available for fringe-limbed treefrogs. Males of E. miliaria have been reported to call from tree holes; one male was sitting above the water line inside the upper edge of the tree hole (MCCRANIE et al. 2003). The call of E. miliaria was described as a loud, booming single note repeated 12-15 times followed by 15-25 minutes of silence (MCCRANIE et al. 2003); SAVAGE (2002) reported the vocalizations of this species to be a single "gurrrgh" repeated at widely separated intervals, similar to the call of E. rabborum ("grrrrrck") as described by MENDELSON et al. (2008). Males are territorial and attend the oviposition site to possibly physically defend it (SAVAGE 2002, MENDELSON et al. 2008). It appears that an individual male may mate with several females using the same tree hole throughout the year (MENDELSON et al. 2008). Females deposit their eggs just above the water line on the exposed wood or bark inside the tree hole; the clutch size being estimated to be about 60-200 eggs (MENDELSON et al. 2008). Some evidence suggests that tadpoles of E. miliaria are (obligate?) egg eaters and are fed by the female with unfertilized nutritive eggs (WILSON et al. 1985, MCCRANIE &

Fig. 532. Ecnomiohyla rabborum (from type locality). Photo: B. Wilson

WILSON 2002, SAVAGE 2002). However, in captivity it was observed that the atten- dant males of E. rabborum spend the day with their bodies half submerged in the water with the tadpoles swimming around the male and eating "small flecks of what appears to be skin coming off his body" (MENDELSON et al. 2008: 755).

Very different from the canopy dwelling fiinge-limbed treefrogs, E. miotympanum is a moderately small species with weakly developed dermal fringes that can be found along small mountain streams during the breeding season, December through April (DUELLMAN 1970). During daytime, these frogs seek shelter in bromeliads and elephant-ear plants (DUELLMAN 1970). The tadpoIes develop in quiet pools of the stream and when disturbed, they seek refuge amidst debris on the bottom of the pools (DUELLMAN 1970). FAIVOVICH et al. (2005) already noted that additional data and analyses might find the species c u - rently included in Ecnomiohyla to represent more than one clade. The obvious differences in habitus and life history between the canopy dwelling fringe-limbed treefrogs on one hand, and E. miotppanum on the other, certainly supports this view.

I Fig. 533. Ecnomiohyla miotympanum (Hidalgo, Fig. 534. Ecnomiohyla milida (Reserva Mexico). Photo: G. Kohler Biologics Rus Rus, Gracias a Dios, Honduras,

180 m). Photo: J. H. Townsend

Ecnomiohyla fimbrimembra (TAYLOR 1948b1, Copeia 1948: 235; type l o c a l i ~ Ysla Bonita ('I'he American Cinchona Plantation), on the Caribbean slope of Volch Poiis, [Heredia Prov.,] Costa Rica; elevation, 5,200 R.." Males to 79 mm, females to 92 mm. Cordilleras de Tilarh, Central, and Talamanca of Costa Rica and western Panama, 500-1900 m elevation.

Ecnomiohyla miliaria (COPE 1886), Proc. Am. Philos. Soc. 23: 272; type locality: "Nicaragua." Males to 110.0 mm, females to 86.2 mm. Atlantic versant from southeastern Honduras to southern Costa Rica, and on the Pacific versant from southern Costa Rica to western and central Panama, near sea level to 1330 m elevation.

Ecnomiohyla minera ( W m , MCCRANIE & WILLIAM^ 19851, Herpetologica 41: 145; type locality: "4.2 km (by road) S PurulhB, 1760 2 elevation, Depto. Baja Verapaz, GuatemaIa. Males to 83.1 mm, females to 80.7 mm. Known from two disjunct populations, in the Municipio de l?urulhii, in the Department Baja Verapaz, and the Sierra de Santa Cruz, Department Izabal, both in Guatemala, 630-1830 m. DUELLMAN (2001: 898) suggested that this nominal taxon might be conspecifk with E. valancifer, see also comments in MCCRANIE &WILSON (2002) and MENDELSON et al. (2008).

Ecnomiohyla miotppanum (COPE 18631, Proc. Acad. Nat. Sci. Philadelphia 15: 47; type locality: "Near Jalapa, Mexico" and "Mirador", Veracruz, Mexico. Males to 38.4 mm, females to 51.0 mm. On the Atlantic versant of the Sierra Madre Oriental from central Nuevo Leon to central Veracruz, Mexico; also disjunct populations in southern Veracruz, north-central Oaxaca, and northern Chiapas, Mexico, 100-2282 m elevation.

Fig. 535. Ecnomiohyla minera (San JosB, SE slope Cerro Verde, Baja Verapaz, Guatemala).

Photo: J. A. Campbell (Courtesy of the Biodiversity Institute, University of Kansas)

Fig. 536. Ecnomiohyla miotympanum (Chiapas, Mexico). Photo: A. Ramirez-Velazquez

Ecnomiohyla rabborum MENDELSON In, SAVAGE, GRIFFITH, Ross, KUBICKI & GAGLIARDQ 2008, J. Herpetol. 42: 751; type locality: "Panama: Cocle: near El V d e de Anton, Rio de Jesus, 990 m. O€i037.9689'N, 08O0O4.6049W." Males to 97.0 mm, females to 100 mm. Mountain~~ in the vicinity of the town of El Valle de Antbn, in both Cocl6 and Panama provinces, Panama, 900-1150 m elevation.

Ecnomiohyla salve& (WILSON, MCCRANIE & WILLIAMS 19851, Herpetologica 41: 141; type locality: "Quebrada Grande (15"05'N, 8s055W, 1370 m elevation, Depto. Copb, Fig. 537. Ecnom'oh~la r a b b o m (Cemo Negro, Honduras." Males to 86.0 mm. females Veraguas, Panama, 540 m). Photo: A. Hertz unknown. Known from Cemo del Mono near La Uni6n in eastern Guatemala close to the border with Honduras, and a t Quebrada Grande, Departamento de Coptin, in extreme western Honduras, 1370-1520 m elevation.

Ecnomiohyla sukia SAVAGE & KUBICKI 2010, Zootaxa 2719: 23; h e locality: "Costa Rica: Provincia de ~ & 6 6 I ~ a n t b n - de Siquirres: Distrito de Siquirres: Guayach: Alto Colorado, 710 m (N 10°02'13.7" W 83'31' 22.4")." Males to 63.3 mm, females to 68.7 mrn. Definitely known from three localities on the Atlantic versant in northeastern Costa Rica (Alto GuayaA, Alto Colorado, and on the Fig. 538. Ecnomiohyla s a l v e (P.N. Cusuco,

'lopes of Volch Tunialba), 400-1000 Cortbs, Honduras, 1355 m). Photo: J. Kolby m elevation.

Ecnomiohyla thysanota (DUELLMAN 1966131, Univ. Kansas Publ. Mus. Nat. Hist. 17: 259; type locality: "Cerro Mali, Dari6n Province, Panam4 (elevation 1265 meters)." SVL of only known female specimen 95.7 mm. Only known from the type locality.

Ecnomiohy1a valancifer (FIRSCHEIN & SMITH 19561, Herpetologica 12: 18; type locality: "Volch San Martin, 4500 ft., Veracruz, Mexico." Males to 77.7 mm, females to 82.0 mm. Sierra de Los Tuxtlas at Volcb San Martin, southern Veramz, Mexico, 500-1380 m elevation. There is one specimen from Toledo District in Belize that has been assigned to this species, but further taxonomic work is required to co~l.firm the identity of this population, which probably belongs to another species.

E. miotympanum

rig. oad. Ecnomiohyla fimbrimembra (Bajo Mono, Chiriqui, Panama, 1820 m). Photo: A. Hertz

Key to E ~ n 0 ~ o h y l 8 prominent and scalloped; texture of dor- 1 a Small green frogs, SVL of adult males sum variable; conditions of prepollex and

27-38 mm, of adult females 37-51 mm; fin- nuptial excrecences variable; skin of dorsal gers about one-third webbed; distinct white surface of head in adults co-ossified with stripe present on ~?argin of upper lip skull or not ................................................ 3 ....................... E ~ o ~ & ' l a m o ~ P a n u m 3 a Dorsum uniform green in life; dorsum

b Large, mostly brownish frogs, SVL in adult smooth to minutely granular; skin of dorsal males 60-110 mm, in females 70-100 mm; surface of head not co-ossified with skull; fingers at least two-thirds webbed; no white heel without tubercles but with an acutely

.................................. stripe present on margin of upper lip ..... 2 scalloped dermal fold ............................ 2 a Webbing between Fingers 11-IV not Ecnomiohyla thpmota

extending beyond penultimate subarticu- b Dorsum brown, reddish brown, or brown lar tubercle on Finger 111; supratympanic with green or darker brown markings or fold continuing posteriorly behind tympa- mottling; dorsum granular or tuberculate; num to terminate above axilla; dermal skin of dorsal surface of head in adults co- fringe along lateral edge of forearm and ossified with skull or not; condition of heel tarsus narrow and not or only weakly scal- variable ..................................................... 4 loped; dorsum smooth to minutely granular 4 a Prepollical projection in adult males (tuberculate in juveniles); prepollex in recurved, terminating in a spine .............. 6

males with terminus and scat- b Prepollex in adult males elongate, flat, tered black nuptial spines; skin of dorsal surface of head in adults co-ossified with rounded, ending blunt, not terminating in

....................................................... skull Ecnomiohyla fhbrimembra a spine 7 .............. Webbing between Fingers II-IV extending 6 a Prepollex in adult males with scattered

well beyond penultimate subarticular small black spines; heel without tubercles;

tubercle on Finger 111; supratympanic fold humera1 projection in adult males present;

not continuing posteriorly behind tympa- hands fully webbed (to base of disc on all ........................ num towards axilla; dermal fringe along digits) Ecnomiohyla minera

lateral edge of fore- and tarsus usually b Prepollex in adult males without scattered

Hylidae

small black spines; heel with one or several pointed tubercles or, if without tubercles, then with a scalloped fringe; humeral projection in adult males absent; hands extensively webbed (to base of disc on some, but not all all digits) ....................................... 6

6 a Heel with one or several pointed tubercles (tipped with black keratin in males); black keratin tipped tubercles over most of flanks and venter; prepollical projection terminating in a sharp spine in adult males: skin of dorsal surface of head co- ossified with skull; SVL of adult males 86-110 mm, of adult females to 86 mm ...a.

Fig. 541. Hand of a. Ecnom.Oh~la hbrimem- E o o h bra; b. E. r ~ b ~ ~ n u n . Drawings: M. Vesely ..................................

b Heel without tubercles but with a scalloped fringe; no black keratin tipped tubercles on flanks and venter; prepollical projection pointed but not terminating in a sharp spine in adult males; skin of dorsal surface of head not co-ossified with skull; SVL of adult males 57-63 mm, of adult females 5&69 mm ................... E~-ahyvla alkia

7 a Prepollex without scattered small black soines: heel with one large tubercles; no hLume;al projection in add; males ...... ......

Em-ohyla valancifm ............................... I b Prepollex with scattered small black

spines; heel without tubercles; humeral projection in adult males present ........... 8 I

8 a Dorsum smooth to granular; skin of d~rsa l Fig. 542. Hand of female Ecnomiohyla minera surface of head not co-assified with skull .. m~ A 24958). Photo: Q Kohler ............................... EuloziIiohyIa rabLnnum

b Dorsum tuberculate, skin of dorsal surface of head co-ossified with skull ...................... .................................... Ecnoznioh yla salvqje

Fig. 543. Ecnomiohyla valancifer (South slope Volch San Martin, Veracruz, Mexico, 1170 m).

Photo: W. E. Duellman (Courtesy of the Biodiversity Mtute, University of Kansas)

Further Reading DUELLMAN 1970, SAVAGE 198lc, Wilson et d. 1985, HAYES et al. 1986, IBAREZ et al. 1991, DUELLMAN 2001, MCCRANIE & WILSON 2002,

Fig. 540. Ecnomiohyla sukia (from type locality). SAVAGE 2002, MCCRANIE et al. 2003, MENDELSON Photo: B. Kubicki et al. 2008

Exerodonta +-.'

FAIVOVICH et al. (2005) resurrected the name Exerodonta from synonymy with Hyla and assigned eleven species of small to moderately small treefrogs to the former taxon. These h g s are denizens of cloud forests and pine-oak forests where they breed in streams and ponds (DUELLMAN 1970, MCCRANIE & WILSON 2002). During the day, frogs of the genus Exerodonta have been recorded hiding in arboreal tank bromeliads and in axils of giant elephant- ear plants, some at considerable distance from the nearest streams of ponds, suggesting these species might sometimes migrate considerable distances to breed (CAMPBELL & BRODIE 1992, MCCRANIE & WILSON 2002). During the breeding season (June to August in E. catracha and E. sumichrasti?, males call at night from low vegetation and on the ground around slow moving streams and small ponds (STARRETT 1960, DUELLMAN 1970, MCCRANIE & WILSON 2002).

The advertisement call of E. catracha was described as a ''muffled two-tone crackle", that of E. bivocata as a series of short notes and occasional long notes, that of E. sumichrasti as a series of quickly repeated notes ("haah-haah-haah") (DUELLMAN 1970, M c C m & WILSON 2002). Tadpoles of E. catracha were collected from slow moving streams and small ponds; after completing metamorphosis, the froglets apparently move uphill from the streams and hide in bromeliads (MCCRANIE & WILSON 2002).

Exerodonta bivocata (DUELLMAN & HOYT 1961), Copeia 1961: 414; type locality: "a small stream above (6.2 kilometers by road south of) Ray6n Mescalapa, Chiapas, M6xic0, elevation 1690 meters." Males to 28.5 mm (females unknown). Northeastern slopes of the central highlands of Chiapas, Mexico, 1500-2100 m elevation.

Exerodonta catracha (PORRAS &WILSON 19871, Copeia 1987: 478; type locality: "Zacate Blanco (14"20'N, 88"15W), a village located 14 km WNW of La Esperanza, 2020 m, Sierra de Opalaca, Departamento de IntibucB, Honduras." Males to 28.7 mm, females to 32.1 mm. Sierra de Opalaca and Sierra de Montecillos, Montafia de La Sierra, in southwestern Honduras, 1800-2160 m elevation.

Exerodonta chimalapa (MENDELSON & CAMPBELL 1994), Proc. Biol. Soc. Washington 107: 399; type locality: "Colonia Rodulfo Figueroa, 19 km NW Rizo de Oro (Chiapas), 1542 m, Oaxaca, Mexico." Males to 24.9 mrn, females to 26.6 mm. Chimalapas region (Sierra Atravesada in southeastern Oaxaca and adjacent Chiapas), Mexico, 850-1540 m elevation.

Exerodonta perkinsi (CAMPBELL & BRODIE 19921, J. Herpetol. 26: 187; type locality: "21.1 km (by road) NNE Barillas, Finca Chiblac, Departamento de Huehuetenango, Guate- mala, ... This site is located at 1080 m elevation on the northern slope of the Sierra de 10s Cuchumatanes at approximateIy 15'53'N, 91°16W." Females to 37.5 mm (males unknown). Known only from the type locality, 615-1080 m elevation.

Exerodonta sumichrasti BROCCHI 1879, Bull. Soc. Philomath., Paris, Ser. 7,3: 20; type locality: "Santa Efigenia pr&s de Tehuantepec", Oaxaca, Mexico. Males to 27.7 mm, females to 33.0 mm. On the Pacific versant from central Guerrero and eastern central Oaxaca to Chiapas, Mexico, 200-2000 m elevation.

Fig. 544. Exerodonta bivocata (6.2 km S Ray6n Mescalapa, Chiapas, Mexico).


I KeytoI5!zmdmte 1 a Axillary membrane absent or rudimentary;

tympanum larger (ratio tympanum length / eye length >0.50); tympanum distinct or indistinct; inner tarsal fold present or

........................................................ absent 2 b Axillary membrane extending at least one-

fourth of upper arm; tpnpanum distinct; tympanum smaller (ratio tympanum length / eye len& ~0.50); inner tarsal fold present ...................................................... 8

2 a Inner tarsal fold absent; dorsum without daxk spots; snout pointed in dorsal view; tympanum indistinct; SVL of adult males 24.8-27.7 mm, of adult females 28.5-33.0 mm ...................... Emdanta Bumichraeti

b Inner tarsal fold present; dorsum usually with dark spots; snout nearly rounded in dorsal view; tympanum distinct or indis- tin& SVL of adult males 24.9-27.4 mm, of adult females 28.6-32.1 mm ....................... ................................... Eremdanta cam

3 a A broad continuous pale stripe along sides of body; tympm~~ll larger (ratio tympanum length / eye length ~0.30); SVL of adult males 26.2-28.5 mm, single known female 26.0 mm .................... Xkemdanta biwcfifa

b No broad pale stripe along sides of body, or, if a stripe present, then discontinuous and irregular; tympanum smaller (ratio tympanum length / eye length c0.30); SVL of ........................................ adults variable 4

4 a A discontinuous, irregular, white or cream stripe extending from the lower posterior corner of eye to groin; axillary membrane extending about half of upper arm; SVL of adult females 33.1-37.5 mrn (males unknown) .................. Exemdmta par-

b No white or cream stripe &om eye to groin; axillary membrane extending about one- fourth of upper arm; SVL of adult males 20-25 mm, of adult females 18-27 mm ................................. lbrodmta c?himsla-

Fig. 545. Exerodonta catracha (Zarciadero, Comayagua, Honduras, 1460 m).


Fig. 546. Exerodonta chimalapa (a. Guate- mala; b. Cerro Baa, Mexico).

Photos: a. J. A. Campbell, b. T. Bille

Fig. 547. Exerodonta perkinsi (from type locality) Photo: J. A. Campbell

Fig. 548. Exerodonta ~umichrasti (Copalita, Sierra de Miahuath, Oasaca, Mexico).

Photo: P. Heimes

Fig. 549. Exerodonta sumichrasti (Villa Flores, Rancho Alegre, Chiapas, Mexico, 970 m).

Photo: A. Ramirez-VelAzquez

Fig. 550. Exerodonta sumiclwasti (Jiquipilas L a Palmita], Chiapas, Mexico, 1557 m).

Photo: A. Ramfrez-Velazquez

Fig. 551. Exerodonta sum3hrasti (Portillo Nejapa, Oaxaca, Mexico). Photo: W. E. Duellman (Courtesy of the Biodiversity Institute, University of Kansas)

Hyla

Hyla bocourti and H. walkeri are the only Central American treefrogs that have remained in the genus Hyla after the major taxonomic revisions proposed by FAIVOVICH et al. (2005). The 34 species that are currently assigned to Hyla are distributed in Europe, eastern Asia, northwestern Africa, as well as and North and Central America (FROST 2010). The two Central American species are nocturnal inhabitants of pine and pine-oak forests (DUELLMAN 1970). Hyla bocourti has been recorded hiding beneath sheaths of banana plants and in bromeliads at daytime (STUART 1948). During the early rainy season (June), males call from temporary grassy ponds, frequently in clearings or meadows devoid of trees; the advertisement call of H. wdkeri consists of series of four to six short, quickly repeated low- pitched notes, the call of H. bocourti is unknown (DUELLMAN 1970,2001). Tadpoles of H. walkeri have been found in grassy ponds where they complete their development (DUELLMAN 1970).

Hyla bowurti (MOCQUARD 1899), Nouv. Arch. Mus. Natl. Hist. Nat. Paris, Ser. 4,l : 341; type locality: "Haute Vera-Paz", Guatemala. Males to 34.6 mm, females to 37.7 mm. Highlands of Alta and Baja Verapaz, Guatemala, 1300- 1536 m elevation.

Hyla walkeri STUART 1954, Proc. Biol. Soc. Washington 67: 165; type locality: "marshy pasture at Aserradero San Loreneo (12 km slightly east of north of Jalapa), Department of Jalapa, Guatemala at 1725 meters." Males to 35.9 mm, females to 37.8 mm. Central Chiapas, Mexico, to the Sierra de 10s Cuchu- matanes, Guatemala, 1450-2900 m elevation.

Further Reading DUELLMAN & HOYT 1961, PORRAS & WILSON 1987, CAMPBELL & BRODIE 1992, MCCRANIE et al. 1993a, MENDELSON & CAMPBELL 1994

Hylidae

Fig. 552. Hyla bocourti (3.8 km W PurulhB, Baja Verapaz, Guatemala).

Photo: J. A. Campbell (Coultesy of the Biodidty Institute, University of I(ansas)

brown and spotted with yellow; feet about three-fourth webbed; SVL of adult males

..... 32-40 mm, of adult females 3 4 4 1 mrn ............................................... Hyla boaourh'

b Dorsum smooth, green in life; posterior surfaces of thigh usually uniformly tan; feet about two-tliiraS webbed; SVL of adult males 29-36 mm, of adult females 31-38 mm ......................................... Ewa wdkeri

E'lg. 563. Hyla walkefi (18 Inn NW Gomitiin, Chiapas, Mexico). Photo: W. E. Duellman (Courtesy of the Biodiversity Institute, University of Kawad

Fig. 555. Thigh pattern in a. Hyla bocourti; b. H. wdkeri. Photos: G. Kijhler

Further Reading DUELLMAN 1970, CAMPBELL & LAWSON 1992, DlJlEUuN 2001

Fig. 554. Hyla walkeri (12 road km on highway to Ocosingo from junction with Hwy 190, Chiapas, Mexico, 2320 m). Photo: S. M. Rovito

Hyloscirtus is an old name proposed by PETERS (1882b) that was resurrected .from synonymy with Hyla by F ~ O V I C H et al. (2005). As currently understood, the genus Hyloscirtus contains 30 species, all except two being distributed in South America (FROST 2010). The two species treated here are unique among Central American hylids by having a whitish or greenish-yellow mental gland present in adult males and usually in adult females (SAVAGE 2002). Both H. colymba and H. paheri are nocturnal stream-breeding treefrogs that inhabit small to moderate-sized fast flowing watercourses with rocky beds in lowland and premontane rainforests (SAVAGE 2002, KOHLER unpubl. obs.). Males of H. colymba have been recorded calling from cracks in rocks, fkom crevices under rocks, and from vegetation overhanging streams, hidden between leaves; the call was described as a metallic "pin-pin-pin" (KUBICKI pers. comm. March 2010, HERTZ pers. comm. September 2010). Males of H. palmeri call from beneath rocks in fast flowing streams emitting paired notes best described as "chi-chi, chi-chi" (K~HLER unpubl. obs.). Occasionally, females of H. colymba have been encountered at night sitting without cover on leaves about 1 m above the ground in the forest some hun- dred meters away from the nearest stream (HERTZ pers. comm. September 2010). Breeding is assumed to be restricted to the drier parts of the year (SAVAGE 2002).

Hyloscirtus colymba ( D m 19311, Occas. Pap. Boston Soc. Nat. Hist. 5: 400; type locality: %a Loma (or Buenavista) on trail from Chiriqui Lagoon to Boquete, about 1500 feet altitudea, Prwincia Bocas del Toro, Panama. MaIes to 37.0 mm, females to 43.3 mm. On the Atlantic versant from southern Costa Rica to central Panama, and also on the Pacific versant in eastern Panama, sea level to 1410 m elevation.

Hyloscirtus palmen' (BOULENGER 19081, Ann. Mag. Nat. Hist. Ser. 8, 2: 515; type locality: "Jimenez", Department of Valle, Colombia. Males to 45.0 mm, females to 50.0 mm. On the Atlantic versant from central Costa Rica to central Panama, and on the Pacific versant from western Colombia to northwestern Ecuador, 100-920 m, in Colombia up to 1600 m elevation.

Fig. 557. Hyloscirtus colymba (El Cop4 CoclB, Panama, 750 m). Photo: B. Kubicki

Fig. 556. Hyloscirtus colymba (Alto de Piedra, Fig. 558. Hyloscirtus palmeri (Guayach, Veraguas, Panama, 910 m). Photo: A. Hertz CRARC, Costa Rica, 550 m). Photo: G. Kohler

Fig. 559. Hyloscirtus p a h e n (Guayach, CRARC, Costa Rica, 550 m). Photo: B. Kubicki

Key to HyIoscirtua 1 a A yellow stripe extending along canthua

rostralis and margin of upper eyelid to level of axilla; a dermal calcar present on heel; SVL of adult males 31-37 mm, of adult females 31-39 mm ............................. .................................... Hylmcidus ml+

b No yellow stripe extending along canthus rostralis and margin of upper eyelid to level of axilla; no dermal calcar on heel; SVL of adult males 36-45 mm, of adult females 36-50 mm ..... E&lmcidus palmme

M e r Reading DUELLMAN 1970, MYERS & DUELLMAN 1982, bArjEz et al. 199913, DUELLMAN 2001, SAVAGE 2002

Hypsiboas

The genus Hypsiboas contains 82 species of moderately sized to very large arboreal nocturnal treefrogs (FROST 2010). Hypsiboas was removed from the synonymy of Hyla by FAIVOVICH et al. (2005). Hypsiboas includes a group of large tan, brownish, or gray frogs, termed "gladiator frogs", formerly referred to as the Hyla boas species group (e.g., DUELW 1970, 2001). Gladiator frogs have large extensively webbed hands and feet and a protuberant prepollex, with terminal spine in males, usually concealed in a retractable fleshy sheath (FAIVOVICH et al. 2005). Fights between males seem to be common in gladiator frogs, and males use the prepollical spines as a weapon to inflict injuries on one another during combat (KLUGE 1981).

Gladiator frogs have a special breeding system in which the males usually construct basin-like nests on the margins of ponds, roadside ditches, or slow-moving streams, the nests being fed with by seepage from these water bodies (KLUGE 1981, HOBEL 2000). However, these frogs show great plasticity in respect of nest site choice depending largely on the ecological factors and frog densities, and utilize natural depressions formed in leaf litter or roots and even puddles and cattle foot- prints as substitutes for male-built nests (HOBEL 1999b). However, clutches of gladiator frogs are never found in open water of ponds or streams, suggesting that a basin - male-built or natural - is critical for reproductive success (HOBEL 2000). During daytime, these frogs retreat to the nearby forest canopy; early in the evening, males begin to call and descend to their nests (SAVAGE 2002). Males of H. rosenbergi have a complex vocal repertoire with at least five different call types (HOBEL 2000). In a Panamanian population of this species, females seemed to inspect the nest of males prior to amplexus, being rather selective, and at least in half the time they

did not approve the nest and left (KLUGE 1981). In case the female found the nests acceptable, she remained and, while in axillary amplexus, spent hours renovating the nest before oviposition, which mostly took place in the early morning hours (2:00 to 5:OO) (KLUGE 1981). The eggs come to form a monolayer floating at the water surface, and hatching occurs 40-66 hours after deposition ( ~ U G E 1981, HOBEL 2000). Flooding of the nests releases the tadpoles into the main body of water where they live on sand or litter banks in streams and ponds. The only Central American Hm~iboas species not belonging to the group of gladiator frogs is H. rufitelus; it was formerly referred to the Hyla albomarginata species group (e.g., DUELLMAN 1970, SAVAGE 2002). It is a moderately-sized treefrog having a green dorsum in life and bright red digital webs. The males call from dense vegetation at the edge or fiom emergent plants in forest ponds or flooded areas in swamp forest (DUELLMAN 1970, SAVAGE 2002, K~HLER unpubl. obs.). The advertisement call consists of a series of

I clucks (DUE- 1970). The eggs are I deposited as a surface film in shallow ! water, and the tadpoles develop in the shal-

low ponds and pools (DUELLMAN 197'0). The juveniles of H. rufiklus differ from the adults in having a distinct pale dorsolateral stripe from tip of snout along upper eyelid to sacral area on each side and in the

I absence of red on the digital webs (see Fig. 1 568).

1 i

Fig. 560. Hypsiboas boans (Laguna Purriche, Darien, Panama, 449 m). Photo: A. J. Crawford

Fig. 561. Hypsiboas crepitans (Ponuga, Veraguas, Panama, 35 m). Photo: A. Hertz

H e b o a s born (LINNAEUS 17581, Syst. Nat. Ed. 10,l: 213; type locality "America." Males to 132.0 mm, females to 117.5 mm. Central and eastern Panama and widely distributed in tropical South America, s'ea level to 300 m elevation.

Hypsiboas erepitans (WED-NEUWIED 18241, Abbild. Naturgesch. Brasil., Heft 8: PI. 47, fig. 1; type locality: 'Tamburil, Jiboya, Arrayd da Conquista" Bahia, Brazil. Males to 58.5 mm, females to 69.9 mm. Azuero Peninsula, central Panama and northern South America (northern Colombia, most of Venezuela and Fig. 562. Hh'siboas Pugnax (Guarinocito near

Magdalena River, Colombia). Photo: A. Acosta

into northern Brazil, the islands of Trinidad and Tobago and throughout much of the Guianas); also in the Atlantic Forest region of Brazil, sea level to 300 m elevation.

Hypsiboas pugnax (SCHMIDT 18571, Sitnmgsber. Akad. Wiss. Wien Phys. Math. Naturwiss. Kl. 24: 11; type locality: "Neugranada." Males to 77.9 mm, females to 80.0 mm. Azuero Peninsula and central Panama; also in northern Colombia and western Venezuela, sea level to 700 m elevation.

Hypsiboas rosenbergi (BOULENGER 1898b1, Proc. Zool. Soc. London 1898: 123; type locali-

... ty: "CachabB [=Cachabfl a small village on the river of that name, on the N.W. Coast, in

... the Prov. Esmeraldas. probably about 500 feet above the sean, Ecuador. Males to 90.0 mm, females to 93.2 mm. On the Atlantic versant disjunctively in southern Costa Rica, Panama, and in the Pacific lowlands of Colombia and the northern slopes of the Cordillera Occidental, to western Ecuador, near sea level to 900 m elevation.

Hypsiboas rufitelus (FOUQUETTE 19611, Fieldiana, Zool. 39: 595; type locality: "Barro Colorado Island, Canal Zone", Panama. Males to 49.2 mm, females to 52.6 mm. Eastern Nicaragua to central Panama on the Caribbean slope, near sea level to 650 m elevation. Venezuelan specimens previously assigned to H. albomarginatus are provision- ally assigned to H. rulitelus.

Key to H9p91'boae 1 a Dorsum green in life (pale tan with dark

flecks in preservative), no middorsal dark line; flanks and posterior surfaces of thighs patternless; webbing red in life; SVL of adult males 39-49 mm, of adult females 46-55 mm ................... Hppaiborre d t e l u 8

b Dorsum tan, cream, or brown in life (and in preservative), usually with a middorsal dark line; fld and posterior surfaces of thighs with dark reticulations or vertical bars; webbing tan or brown in life; SVL of adults greater than indicated above ....... 2

2 a Webbing of fingers rudimentary, webbing between Fingers 111 and IV not reaching penultimate subarticular tubercle on Finger 111 (Fig. 563a); flanks with faint dark bars; SVL of adult males 50-59 mm, of adult females 63-70 mm ......................... .................................... Hjpdboas crep-tam3

b Webbing of fingers substantial, webbing between Fingers I11 and IV reaching at least penultimate subarticular tubercle on Finger 111 (Fig. 563b); flank pattern variable; SVL of adults variable .................... 3

3 a Calcar present; palpebral membrane reticulated; flanks mottled with dark brown; SVL of adult males 101-132 mm, of adult

....... females 90-118 mm &p.ibotts bane b Calcar absent; posterior surfaces of thighs

with narrow vertical bars; flanks with distinct dark bars; palpebral membrane not

......... reticulated; SVL of adults variable 4 4 a Webbing not extending beyond penulti-

mate subarticular tubercle on Finger IV; dark transverse marks on dorsal surfaces of the thighs extending onto anteroventral surfaces; SVL of adult males 69-78 mm, of adult females 66-80 mm Hqrpaibae slpugnax

b Webbing extending almost to base of disc on Finger 1% dark transverse marks on dorsal surfaces of the thighs not extending onto anteroventral surfaces; SVL of adult males 60-90 mm, of adult females 63-93 mm ........................... ~bot t smenbe lg i

Hylidae

Fig. 568. &ad d a. -?ha arepiha% h. a. 4ql~enW. Drawingtx M. Kmely

Isthmohyla

In 2005, FAWOVICH et al. proposed a new genus, Isthmohyla, for 15 species of small to moderate-sized treefrogs restricted to Central America where they occur in moderate to high elevations (370-2770 m above sea level). Typically, Isthmohyla species are nocturnal, arboreal stream breeders; however, a t least three species, I. angustilineata, I. lancasteri, and I. pseudopuma are known to utilize small puddles and other water-filled depressions in the forest and even roadside ditches for breeding; and I. picadoi and I. zeteki are found in arboreal tank bromeliads where they also deposit the eggs (SAVAGE 2002). Individuals of most Isthmohyla species are present along the streams throughout the year (SAVAGE 2002). During daytime, they seek shelter in ground and arboreal tank bromeliads and in the leaf axils of other plants (SAVAGE 2002). At night, males, which are often highly territorial, call from vegetation near or overhanging the stream, as well as from rocks in or along watercourses during most of the year (DUELLMAN 1970, LIPS 1996, SAVAGE 2002). The call of I. debilis and I. tica has been described as a series of cricket-like "chirp- chirp-chirps", that of I. rivularis as a series of high-pitched "cheep-cheep-cheeps", that of I. pictipes as a single low-pitches, pulsed "eeck", and that of I. calypsa and I. lancasteri as a low-pitched, vibrating extended note (SAVAGE 1968b, DUELLMAN 1970). Amplexus is axillary (SAVAGE 2002). These frogs reproduce primarily in the drier months of the year when stream flow is reduced or during the early part of the rainy season (SAVAGE 2002). Some Isthmo- hyla species (e.g., I. calypsa) deposit the eggs in a mass on leaf surfaces of plants directly above the stream; on hatching, tadpoles fall or are washed by rain into the stream below (WILSON et al. 1994, LIP^ 1996). Isthmohyla insolita lays eggs in masses of moss, which are brooded by the male (WILSON et al. 1994). Other Isthmohyla species (e.g., I. angustilineata, I. lancasteri,

Fig. 569. Isthmohyla angustilineata (Pianists Trail, Ngobe Bugl6, Panama, 1800 m).

Photo: M. Ponce

Fig. 570. Isthmohyla calypsa (north slope Cerro Pando, Bocas del Toro, Panama, 1920 m). Photo: W. E. Duellman (Courtesy of the Biodiversity Institute, University of Kansas)

I. pseudopuma) lay the eggs in the water in single long strands (SAVAGE 2002). Females of I. pseudopuma appear to select breeding sites lacking conspecific tadpoles ( C R ~ 1991). The tadpoles of Isthmohyla species inhabit the quieter areas of the stream such as slow-moving portions and backwater pools (LIPS 1996, SAVAGE 2002). In some species, such as I. rivularis, the tadpoles have very large oral discs to adhere to large rocks in the stream (SAVAGE 2002).

Isthmohyla angustilineata ( T A ~ O R 1952b), Univ. Kansas Sci. Bull. 35: 850; type locality: "La Palma, [Provincia San JosB,] Costa Rica." Males to 34.2 mm, females to 36.8 mm. Cordilleras de Tilaran, Central, and Talamanca of Costa Rica and western Panama, 1100-2300 m elevation.

Fig. 572. Isthmohyla debilis (Cerro Tugri, Ngobt Bugl6, Panama, 1570 m). Photo: A. Hert

isthmohyla caiypsa (LIPS 1996), Copeia 1996: 617; type locality: "Finca Jaguar (1900 m), approximately 18 Inn NNE La Lucha, Coto Brus, Puntarenas Province, Costa Rica, 8"55'N, 82"44W." Males to 36.0 mm, females to 41.0 mm. Southern portion of the Cordillera de Talamanca in Costa Rica and Panama, 1500-2100 m elevation.

Isthmohyla debilis (TAYLOR 1952b), Univ. Kansas Sci. Bull. 35: 880; type locality: l s l a Bonita, (American Cinchona Plantation) east- ' ern slope of V6lcan P6as, ICant6n de Alajuela, Provincia de Alajuela,] Costa Rica, 5600 B." [I200 m elevation]. Males to 29.5 mm, females to 31.6 mm. Atlantic versant of the Cordilleras Central and Talamanca in Costa Rica and western Panama, alao on the Pacific versant of southwegtern Panama, 910-1700 m elevation.

Isthmohyla graceae (MYERS & DUELLMAN 1982), Am. Mus. Novit. 2752: 5; type locality: 'hear continental divide southeast of Cerro Colorado, approximately 1650 m. elev., Chiriqui Province, Panama. .. . the type locality lies on a rough eonstructiw road (&om San Felix) at a point about 4 km. airline southeast of Cerro Colorado and about 0.6 km. due south of the continental divide (border of Bocas del Toro)." Males de to Talamanca 38.7 mm, females and to 41.4 mm. - 1

de Fig. 573. Isthmohyla debilis (La Nevera, Ngobe Tabasan5 of Panama, 1120-1710 m elevation. Bugl6, Panama, 1643 m). Photo: A. Hertz

5

Fig. 571. Isthmohyla graceae (Fortuna, Chiri- Fig. 574. Isthmohyla graceae (Cerro Sagui, quf, Panama, 950 m). Photo: IC-H. Jungfer Ngobe Bug14 Panama, 1710 m).Photo: A. Hertz

pig. 575. Isthmohyla graceae (Quebrada de 1 Arena, Chiriqui, Panama). Photo: C. W. Myers 1

(Courtesy of the 3iodiversit.v Institute, University of Kansas) 1 -

Fig. 576. Isthmohyla infucata (Bocas uGl L U L ~ , Fig. 579. Isthmohyla insoLita (2.5 airline km Panama). Photo: E. Boza NNE of La Fortuna, Yoro, Honduras).


1 rig. u I I . r u u u u u u y x a ~ r u u w ~ a r r \rwu vuuugrrru, s. r e vuu.

Bocas del Toro, Panama, 1605 m). Photo: A. Hertz CRARC, ~ca). Photo: G. Kohler

Ishohyla infucata ( D m 1968331, Univ. Kansas Publ. Mus. Nat. Hist. 17: 564; Qp locality: "Rio Changena, Bocas del Toro Province, Panam$, 830 meters." Males to 42.9 mm, females to 45.6 mm. Known only from the vicinity of the type locality, 830-1610 m elevation.

IsthmohJrla insolib (MCCI~ANLE, WILSON & WUUAW 1993b), Copeia 1993: 1057; type locality "2.5 km (airline) NNE La Fortuna (15"26%, 87"18W), 1550 m elevation, Cor- dillera Nombre de Diw Departamento de YWO, Honduras." Males to 36.0 mm, females to 38.0 mm. M y known &om the type locality.

kthmohyla laneasten' (BARBOUR 19281, ROC. New England Zool. Club 10: 31; type locality: "high forest a t Peralta, [Cant611 de Turrialba,

Fig. 578. Lshnohyla iniizcata ( h m type l d t y ) . Provincia Cartago,] Costa Rica, altitude, 500 Photo: W. E. Duellman meters." Males to 33.6 mm, females to 41.1

(Courtesy of the Biodivdty Wtuta, University of -8)

I I

Fig. 581. Isthmohyla lancasteri (Fortuna, Chiriqui, Panama). Photo: G. Kohler tago, Costa Rica, 1000 m). Photo: B. Kubicki

rig. sax. lstnmonyla melacaena w.n. &I uusuco, Fig. 584. Isthmohyla picadoi (Volcdn Barva, CortBs, Honduras). Photo: J. R. McCranie Heredia, Costa Rica, 2260 m) Photo: G. Kohler

mm. Cordillera de Talamanca of Costa Rica I

and western Panama, 90-1450 m elevation.

Isthmohyla melacaena (MCCRANIE & CASTEREDA 2006), Herpetologica 62: 309; type locality: "Parque Nacional El Cusuco Centro de Vistantes, 15"31'N, 88"12W, 1550 m elevation, Departamento de CortBs, Honduras." Males to 22.6 ram, females to 25.9 mm. Only known from the type locality.

Isthmohyla picadoi ( D d 1937), Copeia 1937: 164; type locality: "Volcan Barba, [southwestern slope, Cant6n de Santa Barbara, Provincia de Heredia,] Costa Rica, a little above the farm of Manuel Acosta, 2140 m." Fig. 585. Isthmohyla zeteki (La Nevera, Ngobe Males to 32.8 mm, females to 35.2 mm. Bugl6,Panama) Photo: G. Kahler Cordilleras Central and Talarnanca of Costa Rica and western Panama, 1920-2770 m elevation.

jMhm~&la pl'c@xw (GQPE 1816), J. Ad. Nak. Bu. Phihddphia 8 ~ . 2, & 106; type loidiw %om bhe C m w r x $ at h m 5000 to 7009 feetn, C&a Rita; in error for "Cerro Ulpm, 5000-7000 ft, C a n e de lldamanm, Provida de L i n : 1524-2134 m", a c o o ~ to SAYAGE (1974). Males ta 39.0 mm, &malea to 45.1 mm. Cordjlleras Central and Talamanm in Gosh Ica, 1930-2800 m ekevation.

Fig, 586. Isthmohyla pictim (SE slope of Volch Bama, Heredia, Costa Rica, ca. 2000 m). Photo: E-H. Jungfer

Fig. 587. Isthmohyla pictipes (1 km W Poasito, Alajuela, Costa Rica). Photo: W. E. Duellman (Courtesy of the Biodiversity Xnatitute, University of Kansa~)

Fig. 588. Isthmohyla pseudopuma (female; road fiom Copey to Providencia, San Jos6, Costa Rica, 2540 m). Photo: G. Kohler

lethmo3lpa pseudapuma (G- 1901). Bid. Cenk Amer., Rapt.. m., Part 166: 274; type lo&* "Costa &ca, [prwincia S m Jm4, C&n de Cmnado,] La Paha, alt. 1600 metres." Males to 41.4 mm, female8 to 47.6 mm. Cordilleras de 'Pilar61-1, Central, aad Tdamanca of Cosba Rica and western Panama, 1000-2540 m elevation.

Isthmohyla rivularis ( T ~ R 1952b), Univ. Ransas Sci. Bd. 35: 847; type localiQ: "American Cinohona l?lantati~n EIsla Bonita), eastern slope of V O W Potis, [Cant611 de Alajyla, Provincia de AlajuelaJ Costa Rka, elevabon 5,500 ft." I1209 m elevation]. Wem to 34.0 mm, females to 56.4 mm. C o d l e ~ $ s T i h h , Central anrl Tabmanca of Gosta Em and 4acen t western Fbama, 1210-2840 m elevation.

Ikbhmahyls tica (STARRETT 1966), Bull. S. California Acad. Sd. 65: 23; type l d t y : "a stream, 4500 feet, on Volc6n Turridba, [Cantbn de 'PurridbaJ Cartago Province, Costa Rica" Males to 34 mm, females to 42 rmn. Cordilleras de Tilarbn, Central, and Talamanca of Costa Rica and western Panama, 835-1920 m elevation.

I s h o h y l a xanthostkta (DUE- 1968b), Univ. Kansas Publ. Mw. Nat. a t . 17: 662; type localits "south Fork of the Rio Las Vueltas on the south s l qe of Volch Barba, near the northwest base of Cerro CBompipe, Heredia Province, Costa Rica, elevation 2100 metem." SVL of single known female specimen 29.3 mm. Known only from the type locality.

I . a h y l a eet& (GAIGE 19291, Occas. Pap. M a . Zool. Univ. Michigan 207: .4; type locality: "from a bromdiad in the Caldera Yalley, above Boquete, Chiriqui P r h e , Panama." M a l a to 23.6 mns, females to 26,2 mm. Cordilleras Central and Talamanca of Costa Rica and wesfern Panama, 680-1804 m elevation.

Fig. 589. Isthrnohyla pseudopuma (male; road Fig. 592. Isthmohyla tica (Tapanti, Cartago, &om Copey to Providencia, San JosB, Costa Costa Rica, 1150 m). Photo: K.-H. Jungfer Rica, 2230 m). Photo: G. Kohler

Fig. 590. Isthmohyla rivularis (Jurutungo, Fig. 593. Isthmohyla tica (Finca Bambito, El Chiriqui, Panama, 2070 m; variation in ventral Volch, Chiriqui, Panama). coloration). Photo: G. Kahler Photo: W. E. Duellman

(Courtesy of the Biodiversity Institute, University of Kansas)

Fig. 591. Isthmohyla rivularis (Jurutungo, Fig. 594. Isthmohyla xanthosticta (Rama Sur Chiriquf, Panama, 2070 m). Photo: A. Hertz Rio Las Vueltas, Heredia, Costa Rica).


Hylidae

I . I. pseudopuma ! I. infucata 1

I. lancasteri . I. tica I. calypsa

Key to I d l m e l a 1 a A broad (greatest width varying from one-

half tympanum length to greater than tympanum length) dorsolateral pale (white or yellowish) stripe extending from upper posterior corner of eye to sacral area; venter immaculate; thick basal finger webbing present; posterior surface of thigh pale brown without markings; SVL of adult males 33.5-38.7 mm, single known adult female 40.5 mm ......... Isthm06pla graaeae

b No dorsolateral pale stripe, or, if present, then stripe very narrow and venter with brown flecking; coloration of posterior surface of thigh variable; SVL of adults variable ............................................................ 2

2 a A dorsolateral pale pinstripe from tip of snout along edge of upper eyelid and then along body to level of groin (can be indistinct on head and anterior body), bordered below by a broad dark brown or gray stripe; dorsum uniform green, pale brown, or olive tan in life (some shade of brown or grayish bown in preservatiye), patternless; venter peppered with brown flecks; SVL of adult males 30-34 mm, of adult females 34-37 mm .................. Isthmo4pla mgwtiheata

b No dorsolateral pale stripe from tip of snout to level of groin; flank coloration variable; dorsum with markings; ventral coloration variable; SVL of adults variable .............. 3

3 a Males with 7-10 large black spines on thumb; skin on dorsum smooth; posterior part of dorsum transIucent in life; ventral

surfaces of head and body dirty white; posterior surface of thigh yellowish brown; SVL of adult males 22-23 mm, of adult females 24-26 mm lethm&h melacaena

b Males without large black spines on thumb, or, if spines present, then dorsum with large, tubercular spines; posterior part of dorsum translucent in life or not; coloration of ventral surfaces of head and body variable; coloration of posterior surface of thigh variable; SVL of adults variable ............................................................ 4

4 a Chin and throat boldly marked with a conspicuous pale (yellow to white in life, dirty white in presemative) central area, flanked by solid, dark pigment (Fig. 503); dorsum with numerous wart-like tubercles; SVL of adult males 3 0 3 6 mm, of adult females

................... 34-38 mm Lthmohyla ineolita b Chin and throat not as described above;

dorsum with or without numerous wart- ..... like tubercles; SVL of adults variable 5

6 a Posterior surface of thigh contrastingly barred black and yellow in life (black and white or cream in preservative) (Fig. 505); one or more dark blotches in groin; SVL of adult males 27-34 mm, of adult females 31-38 mm ............... Lthmohy.. heasten'

b Posterior surface of thigh without dark bars; no dark blotches in groin, although small punctations can be present; SVL of

.......................................... adults variable 6

6 a A pale suborbital spot present; a continuous white or golden yellow lateral stripe or an elongate pale blotch on glandular ridge between midlateral region and groin, tympanum small, its length about one-third eye length; SVL of adult males 25-30 mm,

......................... of adult females 27-32 mm ....................................... Isthmohyla d e w

b No pale suborbital spot; no continuous pale lateral stripe or elongate pale blotch present between midlateral region and groin; tympanum size variable; SVL of adults variable ..................................................... 7

7 a Posterior surface of thigh with discrete pale spots (yellow in life, white in preserv-

...................... ative); venter usually dark 8 b Posterior surface of thigh immaculate or

with mottling, but never with discrete pale spots; venter usually pale ........................ 9

8a A bronze stripe on canthus rostralis; large yellow spots on groin and posterior thigh; tympanum length about 50% eye length; single known adult female 29 mm SVL .............................. lethmohyla xanthosticta

b No stripe on canthus rostralis; moderate- sized yellow spots on groin and posterior thigh; tympanum length about 30% eye length; SVL of adult males 31-39 mm, of

............................. adult females 40-45 mm ..................................... Istlunohyla pPct!*p

9a Dorsal surfaces of head, body, and limbs with large, tubercular spines; adult males with small black spines on thumb; SVL of adult males 26-35 mm, of adult females

..................... 31-40 mm Isthmohyla calgpaa b Dorsal surfaces of head, body, and limbs

smooth, granular, or slightly tuberculate, but without large, tubercular spines; nuptial pads of adult males without small

..... black spines; SVL of adults variable 10 10a Toes with extensive webbing, modal formu-

la I11 1-2 IV or I11 1 l/2 -2 112 N (Fig. 595a); moderate webbing between Fingers 111 and IV, modal formula 111 2 l/3 -2 IV 11

b Toes with moderate webbing, modal formula I11 1 l/4 -2 2/3 IV to I11 2 -3 IV (Fin. 595b); webbing between Fingers I11 and fi vestigial, modal formula I11 2 2/13 -2+ IV 13

l la Groin with discrete yellow spots edged by reddish brown; hidden surfaces of groin, anterior and posterior thigh red or maroon in life (pinkish tan in preservative); snout rounded in lateral view; dorsum smooth; SVL of adult males 38-43 mm, of adult females 4 1 4 6 mm .... Isthmohyh infucata

b Groin with yellow mottling of indistinct yellow spots; hidden surfaces of groin, anterior and posterior thigh orange yellow, tan, brown, or orange brown in life (dirty white,

tan, or brown in preservative); snout rounded or truncate in lateral view; texture of dorsum variable; SVL of adults variable ................................................. 12

l2a Snout short and truncate in profile; dorsum and upper surfaces of limbs usually strongly tuberculate; modal toe webbing formula I11 1-2 TV; no discrete yellow spots or a single yellow area in groin, although flank usually mottled brown and yellow, or with irregular pale spots; no dark brown lateral band between axilla and groin in breeding males; SVL of adult males 27-34

............... mm, of adult females 3 3 4 2 mm ........................................... lethmohyh tica

b Snout moderately long and acurninate in lateral view; dorsum and upper surfaces of limbs smooth to granular; modal toe webbing formula III1 112 -2 l/2 IV; groin usually dark brown with indiscrete yellow spots or a single yellow area encroaching on lateral dark band from below; breeding males usually with a distinct dark brown lateral band between axilla and groin; SVL of adult males 37-45 mm, of adult females 41-52 mm ......... I8thmohyla peendopuma

13a Venter usually peppered with dark spots (Fig. 590); dorsum gold orange, yellowish tan, to olive brown in life; modal toe webbing formula I11 1 114 -2 2/3 IV (Fig. 595b); SVL of adult males 29-34 mm, of adult females 33-37 mm .. Isthmohyla rivduie

b Venter imaculate; dorsum grayish brown or tan in life; modal toe webbing formula I11 1 2/3 -3 IV to III 2-3 IV (Fig. 595~);

.......................... SVL of adults variable 14 14a Flanks and ventral surfaces of arms

smooth and not glandularly thickened; eyes directed forward at about a 45" angle; tympanum distinct and directed dorsolaterally; a single odontoid present at the tip of lower jaw; prepollical pad greatly enlarged (Fig. 507g); SVL of adult males

.... 21-24 mm, of adult females 24-27 mm ..................................... lethm*la z e t a b Flanks and ventral surfaces of arms dis-

tinctly granulate to glandularly thickened; eyes directed forward at less than a 45" angle; tympanum indistinct or hidden and directed vertically; a pair of odontoids present at the tip of lower jaw; prepollical pad moderately enlarged (Fig. 5079; SVL of adult males 27-32 mm, of adult females 34-35 mm ................... Isthmohyla picadoi

Further Reading CRUMP 1984,1989b, 1990, CRW & TOWNSEND 1990, AROSEMANA & I B ~ Z 1991, CRW 1991

Osteopilus (West Indian treefiogs) is an Antillean genus of eight moderate-sized to very large species (FROST 2010); one species, 0. septentrionalis, has been introduced to the urban center of Puerto Limbn, Costa Rica (SAVAGE 2002). The skin secretions of 0. septentrionalis are highly irritating to mucous membranes (SAVAGE 2002). This is a nocturnal treefrog that thrives in and around human habitations; as an explosive breeder, it reproduces throughout the rainy season after heavy rains utilizing any permanent or temporary water body that is available such as ponds, roadside ditches, drainage ditches, flooded areas, and even swimming pools (SCHWARZ & HENDERSON 1991, SAVAGE 2002). Males call from vegetation (primarily bushes and trees) near the water body and the advertisement call has been described as rasping snarl (SAVAGE 2002). The female lays up to 130 eggs per clutch in a thin sheet on the water surface (SAVAGE 2002).

Osteopilus septentrionds (DUMI~RIL & BIBXON 1841, Erp. Gen. 8: 538; type locality: "Cuba". Males to 89 mm, females to 165 mm. West Indies (Cuba, Isla de Juventud, Cayman Islands, Bahama Islands); introduced on Puerto Rico, Curacao, St. Maarten and Saba (Netherlands Antilles), Antigua, Hawaii, Virgin Islands, Anguilla, Florida and adjacent coastal Georgia (USA), and Puerto Lim6n, Costa Rica, sea level to 1110 m elevation. See discussion of this species' nomenclature by MI- (1950).

Fig. 596. Osteopilus septentrionalis (Florida, USA). Photo: T. Leenders

Fig. 597. Osteopilus septentrionalis (pet trade). Photo: P. Janzen

Further Rsading MIT~LEMAN 1950, DUELLMAN & CROMBIE 1970, TRUEB & TPLER 1974, SCHWARZ & HENDERSON 1991, MESHAKA 2001, SAVAGE 2002

Fig. 598. Plectrohyla exquisita (P.N. Cusuco, CortBs, Honduras, 1833 m). Photo: J. Kolby

r~ectron yla

The genus Plectrohyla has been recognized for a distinctive group of cloud forest hylids from southeastern Mexico and northern Central America since the early 1940s (e.g., HARTWEG 1941, DUELLMAN & CAMPBELL 1992). Currently, with the inclu- sion of the species formerly assigned to the Hyla bistincta group by FAIVOVICH et al. (2005), 42 species of Plectrohyla are considered valid, 18 of which occur in the area treated by this book (FROST 2010). The Central American Plectrohyla species inhabit cloud forests and pine-oak forests in the highlands (615-3500 m elevation) where they remain along streams throughout most, if not all, of the year (DUELWIAN & CAMPBELL 1992). Daytime retreats of these nocturnal frogs include arboreal tank bromeliads, rock crevices, and moss

mats; individuals of P. exquisita were sleeping on vegetation three to six meters above a stream in early afternoons (MCCRANIE & WILSON 2002). At night, Plectrohyla species have been found along cascading streams where they were cling- ing to rocks and large boulders in the splash zone of cascades or behind water- falls, or perching on branches of bushes or on tree trunks at the edge or above the stream, often at places receiving mist from cascades (DUELLMAN & CAMPBELL 1992, MCCRANIE & WILSON 2002). I encountered juveniles of an undetermined Plectrohyla at Cerro Saslaya, Nicaragua, about 20 meters away from the nearest stream. Plectrohyla species apparently have an extended breeding season, possibly more or less continuous throughout the year (DUELLMAN & CAMPBELL 1992). Males of some species are known to vocalize; the call of P. guatemalensis has been

Hylidae

Tadpoles of Plectrohyla species live in the quieter areas of streams such as pools and small tributaries where they adhere to stones and rocks on the bottom of the pool (DUELLMAN & CAMPBELL 1992). Tadpoles of some Plectrohyla species (e.g., P. hartwegi and P. teuchestes) have greatly enlarged suctorial oral discs (DUELLMAN & CM- BELL 1992).

Fig. 599. Plectrohyla exquisita (P.N. Cusuco, Cortks, Honduras, 1833 m). Photo: J. Kolby

described as a "single low-pitched note

- a that is best described as a grunt which is repeated a t intervals of about two min." (MCCRANIE et al. 1987: 72).

In Plectrohyla, males are larger than females in species having a single knife- like spine, whereas in those species having a blunt prepollex or bifid prepollex, males and females are about equal in size (DUELLMAN & CAMPBELL 1992). Although no combat has been reported in any Plectrohyla species, the presence of scars on bodies and forelimbs has been inter- Fig. 600. Plectrohyla dasypus (P.N. CUSUCO, preted as evidence for rnale-rnale aggres- Codes, Honduras, 1385 m). Photo: J. Kolby sions (DUELLMAN & CAMPBELL 1992). Amplexus is axillary, and the chin and swollen upper lips of the male are pressed against the top of the head of the female; breeding males exhibit glandular swelling of the upper lips and maxillary and premaxillary teeth protruding through the * lips, a condition reminiscent of the condition in plethodontid salamanders in which males have enlarged. premaxillary teeth that pierce the lip and that is used to scratch the females back in order to "inoc- ulate" females with secretions of the male's mental gland (BROCK & VERRELL lgg4)- McCFUN1~ et al. (IgB7) found a Fig. 601. Ampleding pair of Plectrohyla matudai clutch of P. g~atemalensis with 240 eggs urith eggs submerged in a stream (volch that were adherent to the roots of a shrub Tacanl, Chiapas, Mexico, 1750 m). on the bottom of a shallow stream. Photo: A. Ramirez-Vellzquez

Hylidae

Fig. 602. Plectrohyla acanthodes (San Crist6bal de las Casas, Chiapas, Mexico). Photo: l? Heirnes

, .g. 603. Plectrohyla avia (Volck TacanB, Chiapas, Mexico, 1805 m).

Photo: A. Ramirez-Velhquez

Fig. 604. Plectrohyla chrysopleura (La Libera- cibn, Atlktida, Honduras, 1030 m).


Plectrohyla acanthodes DUELLMAN & CAMPBELL 1992, Misc. Publ. Mus. Zool. Univ. Michigan 181: 2; type locality: "a stream 6.2 km (by road) S of Ray& Mescalapa, 1690 m, Chiapas, Mexico (17"10'N, 92'58'W)." Males to 63.2 mm, females to 62.3 mm. Northern slopes of the Meseta Central de Chiapas, Mexico, and the Sierra de 10s Cuchumatanes, western Guatemala, 1540-2250 m elevatim.

Plectrohyla avia S'SULT 1952, Proc. Biol. Soc. Washington 6S: 6; type locality: "Granaja [= Graqja] Lorena (about 10 air-line kilometers northwest of Colomba), Departament of Quesaltemmgo, Guatemala. Elevation, about 1750 meters." Males to 90.4 mm, females are unknown. Extreme southwestern Chiapas, Mexico, southward to northern Pacific Guatemala, 1650-2200 m elevation.

Plectrohyla cfuysopleura WILSON, MCCRANIE & CRUZ-DfAZ 1994, F'roc. Biol. SOC. Washington 107: 68; type locality: "along the Quebrada de Om (15"38'N, 86"47W, 970 m elev., tributary of the Rio Viejo, south slope of Cerro Bfifalo, Cordillera Nombre de Dios, Departamento de Atlhtida, Honduras." Males to 65.6 mm, females to 65.6 mm. Known only fkom the viciniQ of the type locality, 930-1550 m elevation.

Plectrohyla dasypus MCCRANIE & WILSON 1981, Occas. Pap. Mus. Nat. Hist. Univ. Kansas 92: 1; type locality: "Quebrada Cusuco at El Cusuco (15"30'N, 88"13'W), a finca located 5.6 km WSW Buenos Aires (the latter locality about 19 km N Cofkadia), 1580 m, Sierra de Omoa, Departamento de Cort6s, Honduras." Males to 44.0 mm, females to 48.3 mm. Parque Nacional Cusuco, Departamento de CoMs, northwestern Honduras, 1348-2040 m elevation.

Plectrohyla exquisita MCCRANIE & WILSON 1998, J. Herpetol. 32: 96; type locality: "El Cusuco, above the visitors center of Parque Nacional El Cusuco, Cerro Cusuco (15"31'N, 88"12'W), 5.6 km WSW Buenos Aires, 1550 m elevation, Sierra de Omoa, Departamento de Cort6s, Honduras." Males to 80.7 mm, females to 80.7 mm. Endemic to Parque Nacional Cu- suco in the Sierra de Omoa, Departemento de CorMis, northwestern Honduras, 1490-1833 m elevation.

Plectrohyla glandulosa (BOULENGER 1883b1, Ann. Mag. Nat. Hist. Ser. 5,12: 164; type locality: "Guatemala." Males to 49.1 mm, females to 49.7 mm. Sierra de 10s Cuchumatanes and the central and southern highlands of Guatemala, 24004500 m elevation.

Fig. 605. Plectrohyla duysopleura (La Liberacibn, Atlhtida, Honduras, 1030 m).


rig. 605. rrectrohyla dasypus (P.N. Cusuco, CortBs, Honduras, 2040 m).


Fig. 606. Plectrohyla dasypus (P.N. Cusuco, Fig. 608. Plectrohyla guatemalensis (P.N. CortBs, Honduras, 1348 m). Photo: J. Kolby Montaiia de Yoro, Francisco MorazBn,

Honduras, 1850 m). Photo: J. H. Townsend

Plectrohyla guatemalemi~ BROCCHI 187Ta, Bull. Soc. Philomath., Paris, Ser. 7,l: 92; type lodity: Tacicilla [= Patzicia, Departamento Chimaltenango] (Guatemala)." Males to 59.5 mm, females to 54.1 mm. On both versants from southeastern Chiapaa, Mexico, to east- central Honduras, 900-2800 m elevation.

Plectmhyla hartwegi DUEW 1968b, Univ. Hans= Publ. Mus. Nat. Hist. 17: 576; type locality: "Barrejonel (19 kilometers west of Chicomuselo), Chiapas, Mchico, elevation 1000 meters." Males to 75.8, females to 76.7 mm. Chiapas, Mexico, and western Guatemala, 926-2700 m elevation.

Plectrohyla ixil STUART 1942, Occas. Pap. Mus. Zool. Univ. Michigan 456: 4; type locality: "Small stream in a cafetal at Finca San Fig. 609. Plectrohyla hartwe@ (east slope of Francisco (about 25 km. northeast of Nebaj), Cerro Celaque, Lempira, Honduras).


Fig. 610. Pkckrohyla id (Tapalapa, Chispass, Mexico, 1757 m). Photo: A. Ramirez-VelgZquez

Fig. 611. Plectdyla id (San Jose Maxbal, B d a s , Huehuetenango, Guatemala, 1390 m).

Photo: C. R. VAsquez Almazh

Fig. 612. Plectrohyla 1acertos.a (Chiapas, Mexico), Photo: A. Ramhe-Velhquez

[De-toI El Q&h6, &atemala. Altitvde a b u t 1175 met;ar~." Males to 41.6 mu, females to 4622 mm. Two disjunct populations on the Atlantic slope8 of northern Chiapas, Mesioo, and western Guatemala, 1100-2200 m death.

Plaetrohyla lacerbo~a B ~ M & 5 m 1854, Herpetolagica 110: % type locdity: "Etegi6n de ~ 0 c o n u 8 ~ ~ F t 8 8 ~ Mexico? Males to 47.8 mm, females fa 48.1 mm. R n m only from h'0 l&- bl B ~ P CkpSkE (c-0 Tres Ficos and Sboonmm region), Mexico, 1000-2134 m elevation.

Pledmhyia matnsai -G 1941, Occas. Pap. Mus. Zool. Univ. Michigan 437: fi.; type locality: "at an elevation of 1800 meters in the cloud fomt on Mount OPando, District of Sownmco, Chiapas", MexicoCO Males to 46,O mm, females to 49.0 mm. P d c slope^ from Oaxaca and Chiapas, Mexico, to central C f u a M a , 700-2a00 m b a t i o n .

PI~chhpla p h c h i D- & CAMPBELL 1984, Copia 1984: 394; type l d t y : "Rio Sananj6, 3.6 km E of La Uni6n Barrios, Departamento Baja Verapaz, Guatemala, 1,585-1,707 m (lSO1l'N, 90"lQ'Wl." Males to 85.2 mm, females to 49.0 Sderra de las Wnm and the Sierra de Xn~aneb, central and eastern Guatemala, 1&33-1900 m elevation.

PIeofmhyla &odmm M a C m L WILSON 1999r, S e n W . Biol. 78: 232; type locality: "Rio Arcdgual E14"3B'N, 8S04(YW), Cerro Celaque, 2630 m ebvation, Departamento de Lempkh Bondpa&" Males to 48.5 mm, females to 4!3.0 llma Cerro Cekqllie, w&rn h d w a e * d ~ E l P i W @ * r ~ O f El Sdvadar d Hcadr2ras'I, 2M0-2580 m elevation.

P1-rL! pmn&a Baa 1959, Herpa* l~&& 16: 85; type 1 d * "near Coyame, Vemmq Mexico, at about 13aO feePb; in e m , acmrding bo Dmumm C1-WQ). Mdes Q 60.5 mm, fmdw w.p&pm Meseka Central de Chiapas, 2000-2430 elevation.

Pl&m qusectu" Sml?' w, Q c ~ c c fill. Mus. Zool, Wpix nsichigein U: 1; @pe locality: "SCTeam in Barr- L a Ptdmm abut 2 km. north of Fin- Lm A l p (43 Irmn. emt and eligbiily south c$ Cobh btraight line]), IDepa&ameatal Alta Verapaz, Guatemala. Altitude, about 1014 xlpe4ws," W e s to 42.8 mm, b J e s to 46.7 mm, H* highlands of (Siema de las C-tan88 to the highlands of Alh Vmpm and the Siema de,lss Abwl, 615-160 m elevation.

- .:-- ;;.ti

''+--%' ' -+-? , ., .9 ̂

,fl- A

Fig. 613. Plectrohyla psilodema (holotype). Fig. 616. Plectrohyla matudai (r'lnca lrlanaa, Photo: G. Kohler Chiapas, Mexico, 1050 m). Photo: G. Kohler

Fig. 614. Plectrohyla matudai (Volch Tacana, Fig. 617. Plectrohyla quecchi (Finca Los Alpes, Chiapas, Mexico, 1750 m). Senahu, Alta Verapaz, Guatemala, 1000 m).

Photo: A. Ramirez-Veldzquez Photo: C. R. Vasquez Almazdn

Fig. 615. Plectrohyla pokomchi (E slope C e m Fig. 618. Plectrohyla quecchi (3.8 km SE Quisis, Baja Verapaz, Guatemala). Purulhti, Baja Verapaz, Guatemala).

Photo: J. A. Campbell Photo: J. A. Campbell (Courtesy of the Biodiversity Institute, University of Ransasf (Courtesy of the Biodiversity Institute, University of Kansas)

Hylidae * > Y * - + - - ~ ~ W ~ ~ ~ ~ Y ~ ~ K + m-

,'lectrohyla sagonun H A X ~ G 1941, Occas. Pap. Mus. Zool. Univ. Michigan 437: 2; type locality: "at an elevation of 1800 meters in the cloud forest on Mount Ovando, District of Soconusco, Chiapasn, Mexico. Males to 45.5 mm, females to 51.9 mm. On the Pacific slopes of the Sierra Madre del Sur de Chiapas, south- central Chiapas, Mexico, southwest to Guatemala and also in northern El Salvador, 1000-2400 m elevatiofi.

Plectrohyla tecunumani DUELLMAN & CAMPBELL 1984, Copeia 1984: 391; type locality: "a cave 1 km east of Chemal, Departamento Huehuetenango, Guatemala, 3,395 m (15"27'N, 91°31'W)." Males to 61.6 mm, females to 57.3 mm. Sierra de 10s Cuchuma-

Fig. 619. Plectrohyla sagorum (VolcBn Tacana, tams in western Guatemala, 3200-3395 m Chiapas, Mexico, 1810 m). elevation.

Photo: A. Radez-VelBzquez Plectrohyla teuchestes DUELLMAN & CAMPBELL 1992, Misc. Publ. Mus. Zool. Univ. Michigan 181: 15; type locality: "Finca Los Alpes, ca. 1000 m, Departamento Alta Verapaz, Guate- mala (15"22'N, 90°01'W)." Males to 67.1 mm, females to 73.6 mm. Known onlv from the tvpe , .y. ' :;,- !&'

4 .% locality. ,":+'? - .. $.

Fig. 620. Plectrohyla sagom (Rgserva de la Fig. 622. Plectrohyla sp. (subadult; Saalaya, Biosfera El Triunfo, Chiapas, Mexico, Atlantic0 Norte, Nicaragua). Photo: G. Kahler ca. 2000 m). Phatn: S. M. Rovito

- Fig. 621. Plectrohyla teuchestes (Finca Los Fig. 623. Plectrohyla exquisita (P.N. Cusuco, Alpes, Alta Verapaz, Guatemala). CortBs, Honduras). Photo: J. R. McCranie


a f ! chrysopleura F! matudai

=Yb ...................... 1 a Prepollex b' d (Fig. 624ef) 2 . b Prepollex flat, blunt, or pointed, not bifid ........................................... (Fig. 624a-d) 7 2 a Anterior and posterior surfaces of thigh

wikh bold black or dark green and cream in life (black and cream in preservative), usually vertical, markings; SVL of adult males ..... 54-63 mm, in adult females 54-77 mm .................................... PbCtdyh harm?@

b Color pattern of anterior and posterior surfaces of thigh not as described above; SVL

...................................... of adults variable 3 3 a D o r m smooth or nearly so, with only

small wbercles; dorsal surfaoes of head and body uniformly green to greenish olive or greenish brown in life (dark gray to dark brown in preservative); size larger, SVL of adult males 69-81 mm, in adult females 6841 mm ................................................. 4

b Dorsum with numerous large round or conical tubercles; dorsal surfaces of head and body not uniformly green to greenish olive in life; size smaller, SVL of adult males 40-63 mm, in adult females 42-62 mm .. 6

4 a Snout obtuse in profile, moet prominent in males; iris greenish gold or pale brown with black reticulations in life; SVL of adult males 69-81 mm, in adult females 69-81 mm ................ Ple&m&la exquidta

b Snout nearly vertical in profile; iris golden yellow in life; SVL of adult males 72-76

................ mm, in adult females 68-74 mm ................................ Pl- h e & &

6 a Tubercles on dorsum conical, scaktered; dorsum dull olive green or dull gray with large pale olive green spota in life (dull dark brown with or without large faint pale spots); SVL of adult males 48-63 mm, in adult females 54-62 mm ............................. ...............................

b Tubercles on dorsum round, either scattered or densely packed; SVL of addta

..................................................... variable 6 6 a Dorsum with densely packed tubercles;

dorsum dull olive bmwn to dull green with reddish brown markings in He; webbing tan to dull gray in life; vocal slits absent in adult males; SVL of adult males 40-52

................ mm, in adult females 41-54 mm .......................... R* gaatamalaaaie

b Darsum with scattered tubercles; dorsum bright green with darker green or gray tubercles in life; webbing bright red in life; vocal sliOs preaent in adult males; SVL of adult males 48-55 mm, in single known ......... female 49 mm F?b&w&Is pcbnchi

7 a Prepollex flat and distally blunt (Fig. .......................................................... 624a,b) 8

b Prepollex terminally curved with a project- ............................. ing spine (Fig. 624c,d) 14

8 a An outer tarsal fold present; dorsum mostly smooth with few weak tubercles; dorsal surfaces of head and body olive green with irregular olive brown or dark brown marks in males; vocal slits absent in males; SVL of adult males 42-49 rnm, of adult females 39-50 mm ............. Pktmhyla glandulosa

b Outer tarsal fold absent; texture of dorsum variable; dorsal pattern variable; vocal slits present or absent in males; SVL of adults variable ..................................................... 9

9 a Dorsum with numerous bold, irregular dark spots; dorsum strongly tuberculate; maxillary teeth pointed; nuptial excrecences absent in adult males; SVL of adult males to 62 mm, of adult females to 57 mm .............................. Plectrohyla t e c u n m *

b Dorsal pattern not as described above; dorsum smooth or weakly to moderately tuberculate; maxillary teeth pointed or spatulate; nuptial excrecences present in

...... adult males; SVL of adults variable 10 10a Dorsum bronze with discrete black spots,

narrowly outlined with green; dorsum smooth to weakly tuberculate; maxillary teeth spatulate; vocal slits present in adult males; SVL of adult males 3 9 4 4 mm, of

............................. adult females 43-48 mm .................................... Plectdvla dasypw

b Dorsal pattern not as described above; dorsum smooth or weakly to moderately tuberculate; maxillary teeth pointed or spatulate; vocal slits present or absent in

...... adult males; SVL of adults variable 11 lla Dorsum smooth, no tubercles; snout obtuse

in profile; dorsum usually some shade of green in life, with or without brown markings; maxillary teeth pointed; vocal slits absent in adult males; SVL of adult males

..... 4 2 4 9 mm, of adult females 4 3 4 9 mm ............................... Plectrohyla peilodenna

b Dorsal with scattered tuberculate; snout truncate or bluntly rounded in profile; dorsal pattern variable; maxillary teeth pointed or spatulate; vocal slits present or absent in adult males; SVL of adults variable ..................................................... 12

12a Hidden areas of legs, axilla, and groin with prominent yellow marks in life; dorsum weakly to moderately tuberculate; maxillary teeth spatulate; vocal slits present in adult males; SVL of adult males 57-66

................ mm, of adult females 63-66 mm ............................ Plecimhyla dupopleura

Fig. 624. Hand (ventral view) in a. Plectrohyla tecunumani; b. P. psilodema; c. P. matudai, d. P. sagorum; e. P. pokomchi, f P. guatemalensis. Photos: G. Kiihler

b Hidden areas of legs, axilla, and groin without prominent yellow marks in life; texture of dorsum variable; maxillary teeth pointed or spatulate; vocal slits absent in adult

............... males; SVL of adults variable 13 13a Dorsum with irregular brown markings in

preservative; inner tarsal fold flap-like; maxillary teeth pointed; SVL of adult males 54-72 mm, of only known two adult

....... females 48 mm H d y l a lacertosa b Dorsum grayish brown with bluish tan

flecks in preservative; inner tarsal fold not flap-like; maxillary teeth spatulate; SVL of adult males 53-61 mm (females unknown) ............................... n-h p y c n d

14a Dorsal surfaces of head and body uniformly green in lie; dorsum smooth; vocal slits

-1 absent in adult males; large size, SVL of adult males 81-90 mm (femdes unknown) fig. WAS. Dorsal view of head in ........................................ H d y l a avia a. Plectrohyla sagorum; b. P. quecchi; c, P.

b Dorsal surfaces of head and body some d. p. Photos: G. Kijhler shades of brown in life; texture of dorsum variable; vocal slits present in adult males; size smaller, SVL of adult males 32-72

........... mm, in adult females 40-52 mm 16 .................. 16a Vertical rostral keel present 16

Vertical rostra1 keel absent ................... 17 Snout acurninate in dorsal view (Fig. 625a); flanks with small dark flecks (Fig. 626a); dorsum weakly tuberculate; SVL of adult males 34-46 mm, of adult females 40-52 mm ............................ Pldyla agon nun Snout bluntly rounded in dorsal view (Fig. 62513); flanks with large dark spots (Fig. 62613); dorsum moderately tuberculate; SVL of adult males 54-72 mm, of only known adult female 47 mm ........................ ..................................... Pl,?ctm4p.la quecchi Snout acurninate in dorsal view (Fig. 625c); sides of body usually with a broad pale lateral stripe bordered below by a narrow dark stripe (Fig. 626c); SVL of adult males

........... 37-42 mm, adult females to 47 mm ........................................ Pltxh&da id

Snout truncate in dorsal view (Fig. 625e); flanks with a dark lateral line or a series of dark spots (Rg. 626d), no broad pale lateral stripe; SVL of adult males 32-46 mm,

............................... adult females to 49mm ................................... PIdyla matudai

Further Reading BUMZAHEM & SMITH 1954, MCCRANIE & WILSON 1981, DUELLMAN & CAMPBELL 1984, MCCRANIE et ale 1987, DUELL- & CAMPBELL 1992, Fig. 626. Flank coloration in a. Plectrohyla WILSON. MCCRANIE & ~ ~ J z - D ~ A z 1994, sagorum; b. P. quecchi; c. P. S, d. P. matudai. M C C R ~ & WILSON 1998,1999a, 2002

-

Photos: G. Kohler

Fig. 627. Ptychohyla legleri (Alfombra, Puntarenas, Costa Rica, 880 m). Photo: M. Ryan

The genus Ptychohyla contains 13 species of stream-breeding treefrogs that inhabit moderate elevation forests (broad-leafed and mixed pine, 100-2200 m above sea level) from south-central Guerrero and northern Oaxaca, Mexico, across Central America to western Panama with a distributional gap in the Nicaraguan De- pression; nine species are known to occur in Central America (FROST 2010). Adults of this nocturnal treefrog are found on low vegetation, logs, boulders, and on the ground within, above, or near clear, swift streams in the forest (MCCRANIE & WILSON 2002). Males have been reported calling from bushes and thick grasses along streams and seepage areas (MCCRANIE & WILSON 2002). The vocalizations of P.

spinipollex have been described as "a rapidly repeated series of five or six short, high-pitched notes almost like a stuttered whistle" (MCCRANIE & WILSON 1989: 15). DUELLMAN (1970) described the call of P. euthysanota, P. macrotympanum and P. hypomykter as a single soft note ("wraaack"). A clutch of about 135 eggs, presumably from P. spinipollex, was found attached to the undersurface of a large, rather flat rock in a seepage area (MCCRANIE & WILSON 2002). In captivity, females of P. legleri attached the eggs to the roof of a cavity, usually below the waterline (PRoY 1993). The tadpoles of Ptychohyla species develop in the slower- moving shallows or gravel-bottomed pools of the streams where they rest and feed on the bottom (DUELLMAN 1970, SAVAGE 2002; KOHLER unpubl. obs.).

Ptychohyla dendrophasma (CAMPBELL, SMITH & ACEVEW 2000), Herpetologica 56: 251; type locality: "along the Riachuelo Sancapech, Finca San Francisco, near Aldea Yalambojoch, Huehuetenango, Guatemala. This locality is located at about 16O01'56"N and 91°33'29W at 1270 m in the northwestern portion of the Sierra de Los Cuchumatanes, approximately 4.0 km NW of the main ranch-house of Finca San Francisco." Known only from a single female (SVL 84.1 mm). Known only from the type locality.

P@chohyla euthysanota (KELGOGC 19281, Proc. Biol. Soc. Washington 41: 123; type locality: "Los Esesmiles, Departament of Chalatenan- go, El Salvador, C.A. Altitude 6400 feet." Males to 37.3 mm, females to 53.3 mm. Southeastern Oaxaca, Mexico, to Guatemala and western El Salvador, 200-2200 m elevation.

Ptychohyla hypomykter MCCRANIE & WILSON 1993b, Southwest. Nat. 38: 101; type locality: "beside the Quebrada Cafion Oscuro (15"05'N, 8g056W), 1150-1170 m elevation, a stream on Montaiia Cerro Azul below Quebrada Grande, Departamento de CopBn, Honduras." Males to 35.4 mm, females to 39.6 mm. This species ranges from western Guatemala, through most of Honduras to central Nicaragua, 340-2070 m elevation.

Ptychohyla leg& (TAYLOR 1958b), Univ. Kansas Sci. Bull. 39: 33; type locality: "15 km WSW San Isidro del General, San Jose Province . . . in a small swamp", Costa Rica [1.5 km NW Alfombra, Cant6n de Perez Zeledbn, Provincia de San Jose 970 m according to SAVAGE (197411. Males to 36.7 mm, females to 37.0 mm. Pacific slopes of the Cordillera de Talamanca of Costa Rica and extreme western Panama, 880-1600 m elevation.

Ptychohyla m a m o ~ p a n u m (TANNER 19571, Great Basin Nat. 17: 52; type locality: "10 miles east of Chiapa de Corzo, Chiapas", Mexico. Males to 38.8 rnm, females to 44.8 mm. Known only from the road to La Cumbre, Municipio de la Demoeracia, Huehuetenango, Guatemala, 600-2000 m elwation.

Ptychohyla panchoi DUELLMAN & CAMPBELL 1982, Herpetologica 38: 374; type locality: "Aldea Vista Hermosa, Municipio Los Amates, Departamento de Izabal, Guatemala. . . . Vista Hermosa is situated on the northern escarp- ment of the Sierra de las Minas at approximately 15"17'N, 8Q013W, at an elevation of 600 m elwation." Males to 34.2 mm, females to 37.3 mm. Eastern and central Guatemala (Sierra de las Minas, Montafias del Mico, and Sierra Xucaneb), 100-895 m elevation.

Fig. 628. Ptychohyla euthysanota (Finca Irlanda, Chiapas, Mexico, 1050 m). Photo: G. Kohler

Fig. 629. Ptychohyla hypomykter (Posada del Quetzal, Baja Verapaz, Guatemala, 1495 m).

Photo: G. K i e r m

Fig. 630. Ptychohyla hgpomykter (Selva Negra, Matagalpa, Nicaragua, 1300 m). Photo: G. Kiihler

I Fig. 631. Ptychohyla leglen' (Las Cruzes, Punta- renas, Costa Rica, 1220 m). Photo: J. Vargas

257

Hvlidae

Fig. 632. Ptychohyla salvadorensis (Nahua- Fig. 635. ptychohyla macrotympanum terique, M o r a h , El Salvador, 1390 m). (Tapalapa, Chiapas, Mexico, 1741 m).

Photo: T. Leenders Photo: A. Ramirez-Vekizquez

Fig. 633. Ftychohyla macrotympanum Fig. 636. Ptychohyla panchoi (Los Amates, (Berriozaba., Chiapm, Mexico, 1241 m). Aldea Vista Hermosa, Izabal, Guatemala).

Photo: A. Ramirez-Velbquez Photo: J. A. Campbell (Courtesy of the Biodiversity W t u t e , University of I(ansae)

Fig. 634. Ptychohyla macrotympanum (same loca- , 37. . , :hohyla spin :same localil I lity as in Fig. 635). Photo: A. Emhz-VeUzquez as in Fig. 639). Photo: J. H. Townsend

Hylidae

d Fig. 638. Ptychohyla hypomykter (P.N. Saslaya, Atlantico Norte, Nicaragua). Photo: J. Sunyer

Ptychohyla salvadorensis (MERTENS 1952b1, Senckenb. Biol. 33: 169; type locality: "Hacienda San Jose, 800 m H., Metapan- Gebirge, Dept. Santa Ana, El Salvador!' Males to 34.9 mm, females to 41.8 mm. Extreme southeastern Guatemala, northern and central El Salvador, and southwestern to south- ' central Honduras, 700-2050 m elevation.

Ptychohyla sanctaecrucis CAMPBELL & SMITH 1992, Herpetologica 48: 155; type locality: "near Chinamococh, Finca Semuc, Departa- mento de Izabal, Guatemala; . . . This locality is

Fig. 639. Ptychohyla spinipollex (La located at 640 m elevation in the Rio Sauce

Liberacibn, Atlhtida, Honduras, 1030 m). drainage of the Sierra de Santa Cruz at about 15"37'N, 89O29W." Males to 32.5 mm, females

Photo: J. H. to 50.7 mm. Sierra de Santa Cruz in eastern Guatemala, 366-1150 m.

Ptychohyla spinipollex (SCHMIDT 1936131, Proc. Biol. Soc. Washington 49: 45; type locality: "mountains behind Ceiba, Departamento del Atlhtida, Honduras." Males to 39.1 mm, females to 46.1 mm. Cordillera Nombre de Dios in northern Honduras, 160-1580 m elevation.

Fig. 640. Ptychohyla spinipollex (same locality as in Fig. 639). Photo: J. H. Townsend

259

Key to ptschohyla 1 a A distinct lateral white stripe from axilla

to groin; width of disc on Finger I11 distinctly greater than tympanum length; iris red or reddish bronze in life (usually reddish brown in preservative) ..................... 2

b No lateral white stripe from axilla to groin; width of disc on Finger I11 smaller than or only slightly greater than tympanum length; iris not red in life (not reddish brown in preservative) ............................. 5

2 a Venter with scattered brown flecks; adult males with a patch of 50-55 moderately large black spines on base of thumb; dorsal surfaces of head and body grayish tan with green flecks, especially prominent on eyelids; a vertical rostral keel present; modal webbing formula of fingers I11 2+ - 1 IV; SVL of adult males 31-34 mm, of single known female 37 mm &ychohyla p c h o i

b Venter immaculate; adult males with a brown non-spinous pad on base of thumb or with a patch of 125-135 small to moderately small black spines; dorsal surfaces of head and body dark brown, uniform or mottled with small pale, bronze, or green flecks, or pale reddish brown with dark brown reticulations; no vertical rostral keel; modal webbing formula of fingers I11 2+ - 2 IV; SVL of adult males 31-37 mm, of adult females 36-39 mm ......................... 3

3 a Width of disc on Finger I11 distinctly greater than tympanum length; adult males with a brown non-spinous pad on base of thumb; ventrolateral gland absent in adult males; iris red in life; SVL of adult males 31-37 mm, of adult females 36-39 mm ................................ Pip&& yla leglmeR

b Width of disc on Finger I11 about subequal to tympanum length; adult males with a a patch of 125-135 small to moderately small black spines on base of thumb; ventrolateral gland present in adult males; iris reddish bronze in life; SVL of adult males

..... to 37 mm, of adult females to 53 mm 4 4 a Venter immaculate; adult males with a a

patch of 44-143 small to moderately small black spines; iris reddish bronze in life; SVL of adult males to 37 mm, of adult females to 53 mm Rydiohyla e u ~ ~ o t a

b Venter with scattered brown or black flecks; adult males with a a patch of 90-100 small to moderately small black spines; iris coppery bronze in life SVL of adult males to 38 mm, of adult females to 45 mm ......... &ychohyla macro@npanum

5 a Fingers about two-thirds webbed, modal formula I11 2 l/2 - 2 112 IV; toes nearly fully webbed, modal formula I11 1 -1 l/2 IV; a distinct inner tarsal fold present present;

SVL of single known female 84 mm ........... .......................... Pip&ohyla dendrophasma

b Fingers about one-fourth to one-half webbed, modal formula I11 2+ -2 IV to I11 2 l/2 - 2 l/2 IV; toes with less webbing than described above; inner tarsal fold distinct, weak, or absent; SVL of adults <60 mm . 6

6 a Dorsum lime green with scattered pale green dots in life; fingers about one-fourth webbed, modal formula I11 2 l/2 - 2 l/2 IV; flanks with a narrow zone of reticulation where dorsal and ventral coloration meet; adult males with a patch of tiny black spines on base of thumb; a distinct inner tarsal fold present present; SVL of adult males 29-33 mm, of adult females 41-51 mm .................... &ychohyla sanctaecrucis

b Dorsum pale tan, dull brown, or reddish brown with or without scattered dark brwon blotches in life; fingers about one- half webbed, modal formula I11 2+ -2 IV; flank coloration not as described above; nuptial excrecences in males variable; an inner tarsal fold distinct, weak, or absent; SVL of adults variable ............................. 7

7 a Adult males with a non-spinous pad on base of thumb (Fig. 641a); no ventrolateral gland in adult males; SVL of adult males

..... 27-35 mm, of adult females 34-42 mm ............................ &ychohyla salvado&

b Adult males with a patch of black spines on base of thumb (Fig. 641b); ventrolateral gland weakly to prominently developed, or only scattered small glands present between axilla and groin in adult males; SVL of adults variable ............................. 8

8 a Flank without pale spots or mottling; adult males with a patch of 90-100 small to moderately small black spines on base of thumb; inner tarsal fold weak, low and rounded; SVL of adult males to 38 mm, of adult females to 45 mm ..............................

-&la macro@npanum ...................... b Flank with pale spots or mottling; adult

males with a patch of 27-78 moderately large black spines on base of thumb; inner tarsal fold distinct; SVL of adults variable 9

9 a Belly and ventral surfaces of thighs immaculate or with widely scattered flecks; ventrolateral gland prominently developed in adult males; SVL of adult males 30-35 mm, of adult females 3 5 4 0

....................... mm &&+la hypom yktar b Belly and ventral surfaces of thighs usual-

ly peppered throughout with brown flecks; ventrolateral gland not well-developed, only scattered small glands present between axilla and groin in adult males; SVL of adult males 33-39 mm, of adult females 39-46 mm P&.hohyh spini'IIex

Fig. 641. Nuptial pads in a. mchohyla salvadorensis; b. P. euthysanota. Drawings: M . Vesely

Fig. 642. Hand and foot of ptschohyla salvadorensis (left,) and P. euthysanota (right).

Drawings: M. Vesely

Further Readkg DUELLMAN 1963c, 1970, DUELLMAN & CAMPBELL 1982, MCCRANIE & WILSON 1989, CAMPBELL & SMITH 1992, MCCRANIE & WILSON 1993b, CAMPBELL et al. 2000, DUELLMAN 2001

P: salvadorensis

H ylidae

Fig. 643. Scinax boulengeri (La Selva, Heredia, Fig. 644. Scinax elaeochroa (Punta Uva, Costa Rica, 70 m). Photo: G. Kiihler Limbn, Costa Rica, 10 m). Photo: G. Kohler

About 100 species of small to moderate- sized anurans are currently assigned to the genus Scinax, the vast majority of them occurring in South America (FROST 2010). Central America is home to six species of this genus. Some controversy had existed as to the gender of the genus name Scinax (DUELLMAN & WIENS 1992, KOHLER & B O m 1996, SAVAGE 2002). I follow SAVAGE (2002) in regarding Scinax as of masculine gender. All Scinax are nocturnal and arboreal. Scinax staufferi is found in a wide variety of habitats including dry, moist, and wet forests, usually in relatively open situations, avoiding deep forest (LEE 1996). Scinax elaeochroa and S. boulengeri, on the contrary, are more typical lowland rainforest species, inhabiting forest habitats, often found near breeding habitats (SAVAGE 2002, SCOTT 1983d). These two species are prolonged breeders, and males can be heard calling throughout most of the year (DUELLMAN 1970). The breeding activities of S. staufferi are

restricted to the rainy season, given the strong seasonality that prevails where this species occurs (CAMPBELL 1996). During the dry season, individuals of S. s tader i have been found hiding under logs or rocks on the ground, beneath the bark of standing trees, and within bromeliads near water bodies (SMITH 1941, CAMPBELL 1996, LEE 1996). The call of S. elaeochroa has been described sounding like "waak", that of S. boulengeri like "wraak", and that of S. boulengeri Like "ah-ah-ah-ah" (LEE 1996, SAVAGE 2002). Scinax species typically breed in shallow temporary bodies of water throughout most of the wet season (LEE 1996, SAVAGE 2002). After the first heavy rains hit the breeding site, males, especially of the common species, such as S. elaeochroa and S. staufferi, suddenly congregate, and soon numerous males produce a raucous chorus (SAVAGE 2002). Amplexus is axillary in Scinax and eggs are laid as small clumps in the shallow water where the tadpole hatch and complete their development (DUELLMAN 1970, LEE 1996, SAVAGE 2002).

Hylidae

Scinax d tae ~ U N N 1933a), Occas. Pap. Boston Soc. Nat. Hist. 8: 61; type locality: "Summit, Panama Canal Zone", Panama. Males to 26.0 mm, females to 27.8 mm. Subhumid Pacific lowlands of Panama from Concepcibn, Chiriqui Province, to Chepo in the lower Bayano valley, Panama Province, Panama, sea level to 700 m elevation.

Scinax boulengeri (COPE 1887), Bull. U.S. Natl. Mus. 32: 12 ; type locality: "Nicaragua."Males to 48.7 mm, females to 52.8 mm. On the Atlantic versant from northeastern Honduras to central Panama, also on the Pacific versant in Costa Rica and an apparently disjunct population in Colombia, sea level to 700 m elevation.

Scinax elaeochroa (COPE 18761, J. Acad. Nat. Sci. Philadelphia Ser. 2, 8: 105; type locality: "east foot of mountains near Sipurio", CanMn de Talamanca, Provincia Lim6n, Costa Rica. Males to 37.7 mm, females to 40.3 mm. On the Atlantic versant from eastern Nicaragua to northwestern Panama, and on the Pacific versant from the Rio Barranca drainage of central Costa Rica to western Panama; also an isolated population in the Pacific lowlands of Colombia, sea level to 1200 m.

Scinax rostratus (PETERS 1863b), Monatsber. Preuss. Akad. Wiss. Berlin 1863: 466; type locality: "Caracas", Venezuela. Males to 45.7 mm, females to 41.0 mm. Central and eastern Panama, and widely in northern South America (Colombia, northern Venezuela to Guyana and Surinam), in Panama sea level to 300 m elevation.

Scinax mber (LAURENTI 17681, Spec. Med. Exhib. Synops. Rept.: 5; type locality: "America"; neotype from "Paramaribo, Surinam" (DUELLMAN & WIENS 1993)). Males to 35.2 mm, females to 40.8 mm. Central and eastern Panama, and widely in Tropical South America; in Panama sea level to 300 m elevation.

Scinax staufferi (COPE 1865), Proc. Acad. Nat. Sci. Philadelphia 17: 195; type locality: "Orizava [= Orizaba, Veracruz], Mexico." Males to 29.0 mm, females to 31.6 mm. On the Atlantic versant from central Tamaulipas, Mexico, to Belize, Guatemala, Honduras and Nicaragua (including the Corn Islands); on the Pacific versant from Guerrero, Mexico, to northwestern Costa Rica, sea level to 1530 m elevation.

Fig. 645. Scinax altae (Los Algarrobos, Chiriqui, Panama, 125 m). Photo: A. Hertz

Fig. 646. Scinax elaeochroa (Guayach, CRARC, Costa Rica, 550 m). Photo: G. Kohler

Fig. 647. Scinax staufferi (near San Jose El Idalo, Suchitep6quez, Guatemala, 70 m).

Photo: G. Kijhler

Fig. 648. Scinax staufferi (Santa Lucia, Intibuca, Honduras, 370 m). Photo: G. Kohler

Key to Scinax 1 a Posterior surface of thigh with contrasting

dark bars on a pale (greenish yellow to bright yellow in life, white in preservative) background (Figs. 649a,b); supratympanic fold black; dorsum blotched or spotted; SVL of adult males 36-49 mm, of adult females

................................................. 44-52 mm 2 b Posterior surface of thigh uniform or weak-

ly, never contrastingly barred; supratympanic fold pale brown; dorsum with a linear pattern; SVL of adult males 21-38 mm, of

......................... adult females 2 1 4 1 mm 3 2 a Dorsum tuberculate; snout subacuminate

in dorsal view; vocal sac flecked with brown; inner tarsal fold weak; h g e r s lacking webbing; black spots absent in scapular region; groin with a dark spot or several spots; SVL of adult males 36-49 mm, of

............................. adult females 44-52 mm ........................................ Scinax bod-

b Dorsum smooth; snout pointed in dorsal view; vocal sac dark gray; inner tarsal fold absent; fingers with a trace of webbing; two or more elongate black spots in scapular region; groin without a dark spots; SVL of adult males 4 0 4 6 mm, of single known Panamanian female 41 m m ........................ ........................................ Scinaxrostratus

3 a Tympanum length less than half eye length; prevomerine elevations smaller than choanae; dorsum strongly granulate to tuberculate; snout pointed in dorsal view; bones white in life; SVL of adult males 21-27 mm, of adult females 21-32 mm ............................................................ 4

b Tympanum length a t least two-thirds of eye length; prevomerine elevations about the size of the choanae; dorsum smooth to weakly granulate, never tuberculate; snout rounded to acutely rounded in dorsal view; bones green in life; SVL of adult males 26-38 mm, of adult females 3 0 4 1 mm .. 5

4 a Dorsal surface of shank with transverse bars; dorsum brown with irregular dorsolateral stripes and interrupted paravertebral stripes; interorbital bar present; SVL of adult males 21-27 mm, of adult females 21-32 mm ......................... &ax 8ta-

b Dorsal surface of shank with longitudinal stripe; dorsum gray with uninterrupted dorsolateral and paravertebral stripes; interorbital bar absent; SVL of adult males

..... 22-26 mm, of adult females 26-28 mm ................................................. Scinax altae

5 a Width of disc on Finger I11 about one-half tympanum length; posterior surface of thigh with bold dark brown or black reticulations enclosing pale (bright yellow in life,

creamy tan in preservative) spots (Fig. 650); SVL of adult males 31-35 mm, of

...... adult females 37-41 mm L3cinaxrnbe.r b Width of disc on Finger I11 distinctly

greater than tympanum length; posterior surface of thigh uniform pale yellow or with suffuse olive tan mottling; SVL of adult males 26-38 mm, of adult females 3 0 4 0 mm ..................... elaeochroa

S. stauffen'

I

S. altae

'4

Scinax

Hylidae

r'ig. b 4 ~ . ,mign pattern m a. acmax boulengeri and b. S. rostratus.

Photos: a. G. Kiihler; b. A. Hertz

i Fig. 650. Scinax ruber (Wacu~u, Panama,

[ Panama, 130 m).

J Photo: A. J. Crawford

Fig. 651. Scinax rostratus. (Wacuco, PanamB, Panama, 120 m). Photo: A. J. Crawford

Further Rsading RIVERO 1969, DUELLMAN 1970, FOUQUETTE & PY~URN 1972, FOUQUETTE & DELAHOUSSAYE 1977, SCOTT 1983d, DUELLMAN 1986, JUNGFER 1987, DUELLMAN & WIENS 1992, LANGONE 1992, DUELLMAN & WIENS 1993, K o m ~ & B ~ H M E 1996, NIETO 1999, SAVAGE 2002

The genus Smilisca contains eight species of moderate-sized to large anurans, six of which occur in Central America; the two extralimital species, S. dentata and S. fodi- ens, traditionally were assigned to a separate genus, Pternohyla, before FANOVICH et al. (2005) synonymized the latter genus with Smilisca. The Central American species of this genus are common treefrogs that are found in a wide array of habitats sach as dry, moist, and wet forests, secondary growth, and pastures (CAMPBELL 1996, DUELLMAN 1970). Smilisca baudinii and S. phaeota are usually the first anurans to begin calling after the onset of the rainy season, and some males will even call during the day (CAMPBELL 1996, WEBB 1971). The call of S. baudinii has been described sounding like "wonk-wonk- wonk" (CAMPBELL 19961, that of S. phaeota like "wawkn (SAVAGE 2002). Males of S. baudinii usually call from the edge of the water body or from low vegetation near the shore, males often call from the water surface of small rain pools (KOHLER unpubl. obs.).

Amplexus is axillary in all Central American Smilisca species (SAVAGE 2002). Four of the six Central American Smilisca species (i.e., S. baudinii, S. cyanosticta, S. phaeota, and S. puma) utilize temporary pools (ponds, roadside ditches, etc.) for breeding following heavy rains, whereas S. sila and S. sordida breed along streams during the dry season (JAMES 1944, MALONE 2004). During the rainy season, streams utilized by S. sila and S. sordida can be roaring torrents, apparently not suitable for the adults or tadpoles of these species (DUELLMAN 1970). The pond-breeding species of Smilisca exhibit a general- ized reproductive mode with eggs deposited as a surface film in lentic water; the two stream-breeding species on the other hand, construct basins for egg deposition (MALONE 2004). During amplexus, females of S. sila and S. sordida dig a depression in

Hylidae

the substrate over the course of several hours by pushing gravel with the hind feet; these basins are constructed on rocky gravel islands or on adjacent banks, usually with stream water flowing into basins but some basin without a connection with the stream (MALONE 2004). The female then deposits eggs within the depression; the tadpoles are found in pools and backwaters of the stream and use the large oral disc to maintain position (SAVAGE 2002, MALONE 2004).

Fig. 652. SmiLisca baudinii (Mazatenango, Guatemala, 395 m). Photo: G. Kiihler

Smilisca baudinii ( D U ~ I L & BIBRON 18411, Erp. Gen. 8: 564; type locality: "Mexique." Males to 76.0 mm, females to 90.0 mm. On the Atlantic versant from the Rio Grande embay- ment in extreme southern Texas, U.S.A. to Costa Rica, including the Yucatan Peninsula; and on the Pacific versant from southern Sonora to northwestern Panama, sea level to 1925 m elevatian.

Smilisca cyanosticta (SMITH 1953), Herpeto- logica 8: 150; type locality: "Piedras Negras, Pet&, Guatemala." Males to 56.0 mm, females to 70.0 mm. On the Atlantic versant from Oaxaca and southern Veracruz, Mexico, through northern Chiapas, Meico, and northern Guatemala (El Pet6n and northern Alta Verapaz) to Belize, 100-1200 m elevation.

Smilisca phaeota (COPE 1862a), Proc. Acad. Nat. Sci. Philadelphia 14: 358; type locality: "Turbo" [Intendencia de Choc6, Colombia]. Males to 65.0 mm, females to 78.0 mm. On the Atlantic versant from northeastern Honduras to northern Colombia; on the Pacific versant from Costa Rica to western Ecuador, sea level to 1255 m elevation.

SmiLisca puma (COPE 18851, Proc. Am. Philos. Soc. 22: 183; type locality: "Nicaragua." Males to 38.0 mm, females to 46.0 mm. Caribbean lowlands of Costa Rica and adjacent Nicaragua, 15-520 m elevation.

Smilisca sila DUE- & TRUEB 1966, Univ. Kansas Publ. Mus. Nat. Hist. 17: 318; type locality: "a small stream a t north edge of the village of El Volch, Chiriqui Province, PanamB, elevation 1280 meters." Males to 45.0 mm. females to 62.0 mm. On the Pacific versant from southern Costa Rica to eastern Panama, and on the Atlantic versant from cen-

; tral Panama to northern Colombia and the middle Magdalena Valley, sea level to 1350 m , elevation.

I Smilisca sordida (PETERS 1863b), Monatsber. Fig. 653. Smilisca baudinii (Lagarto Lodge, Preuss. Akad. Wiss. Berlin 1863: 460; type Alajuela, Costa Rita, 90 m) photo: G. Kohler locality: veragua", Males to 45.0-

mm, females to 64.0 mm. On the Atlantic ver-

I sant from northeastern Honduras to northwestern Panama, and on the Pacific versant from southern Costa Rica to west-central Panama; also a record from the middle Magdalena Valley, Colombia, sea level to 1525 m elevation.

I Fig 654. Srmlisca puma (Rlo San Juan, Nicaragua, 30 m). Photo: G. Kohler

Hylidae

u

Nicaragua). Photo: G. ~ 6 h l k r

I

Fin. 655. SmiIisca ~haeota (near Pueblo Wiso,

Fig. 659. Smilisca cyanosticta (El Ocote, Chiapas, Mexico). Photo: G. Kohler

Key to S d c a 1 a A dark brown or black postorbital mark

encompassing tympanum present, usually continuing as dark lateral band at least to level of axilla; SVL of adult males 40-75

............. mm, of adult females 47-90 mm 2 b No dark brown or black postorbital mark

Fig. 656. Smilisca phaeota (Cerro El Toro, encompassing tympanum present; SVL of adult males 31-54 mm, of adult females

Atlantic0 Norte, Nicaragua). Photo: G. Kohler 40-62 mm ................................................. 4 2 a A brown blotch on upper lip present (at

level of anterior portion of eye); a series of whitish warts or tubercles along posteroventral margin of arm present; flanks cream with bold mottling in groin; SVL of adult males 47-75 mm, of adult females 56-90 mm ...................... S . baudinii

b No brown blotch on upper lip present (at level of anterior portion of eye); no series of whitish warts or tubercles along posteroventral margin of arm; flank coloration different than above ................................. 3

Fig. 657. fj'milisca slla Rafael Norte, Sari 3 a Groin as well as anterior and posterior Sur- JosB, Costa Rica, 990 m). Photo: G. Kohler faces of thighs with blue spots; SVL of

adult males 50-56 mm, of adult females

Fig. 658. Smilisca sordaa (ruco ae neara, Veraguas, Panama, 900 m). Photo: G. K'ohle

b Groin cream with fine black venation, no blue spots on anterior and posterior surfaces of thighs; SVL of adult males 40-66 mm, of adult females 50-78 mm ................ ........................................ S&cd phaeota

4 a Webbing between fingers essentially absent (Fig. 660a); no series of warts along posteroventral margin of arm; a pale stripe, bordered below by a dark line, from heel to foot; dorsum with a pair of irregularly shaped longitudinal dark brown blotches that fuse medially at several points; a dark brown interorbital bar or

spot usually present; SVL of adult males 32-38 mm, of adult females 4046 mm ..... ............................................. Smilisce puma

b Webbing between fingers substantial, extending to or near to distal subarticular tubercle on finger IV (Fig. 660b-d); a series of warts along posteroventral margin of arm; no pale stripe fi-om heel to foot; dorsal pattern variable but not with a pair of irregularly shaped longitudinal dark a. brown blotches that fuse medially at several points; usually no dark brown interorbital bar or spot ........................................ 5

5 a Less webbing between fingers (Fig. 6604; dorsal surfaces strongly tuberculate; snout short, truncate; blotches on dorsum not having the tendency to form transverse bars; vocal sacs of breeding males gray or brown; SVL of adult males 31-45 mm, of

....... adult females 44-62 mm S&CB aila b More webbing between fingers (Fig. 660d); C. dorsal surfaces smooth; snout longer, slop-

ing. rounded: blotches on dorsum having th;;r'tendencyto form transverse bars; vocd

I

sacs of breeding males white; SVL of adult Fig. 660. Hand of a. Smilisca baudinii; b. S. males 32-54 mm, of adult females 56-64 Puma; c- S. sila; d. S. sordida. mm .................................... Smilieca Bardida Drawings: M. Vesely

Smilisca I 4 -

I :. baudinii

Smilisca

S. sila S. puma

. L

Smilisca I

*

Tlalocohyla

The genus Tlalocohyla was established by FAIVOVICH et al. (2005) for a group of four species of hylid frogs formerly referred to as the Hyla godmani species group (e.g., DUELLMAN 2001). Two representatives of this genus occur in the area covered by this book, the other two species, T. godmani and T. smithi, have a more northerly dis-

I tribution in Mexico (DUELLMAN 2001). ! Tlalocohyla loquax is a widespread noctur.

nal frog that is found in and around temporary and permanent ponds and small lakes (DUELLMAN 1970, MCCRANIE & WILSON 2002). During the dry season, this predominantly arboreal frog has been found in arboreal bromeliads high in the trees (CAMPBELL 1998). Most males of this species that I have encountered were calling from emergent plants or floating vegetation some distance from the shore in the deeper parts of the pond making it impossible to access them without getting wet hip-deep. This is one of the last frogs to arrive at the breeding ponds after the beginning of the rainy season (usually after several heavy rains) and breeding activities peak in the middle of the rainy season in July and early August (CAMPBELL 1998, SAVAGE 2002). The call of T. loquax has been likened to the honking of a goose (DUELLMAN 1970). The females deposit large gelatinous egg masses attached to submerged plants near the water surface; one such clutch contained about 250 eggs (DUELLMAN 1970, CAMPBELL 1998, SAVAGE 2002). The tadpoles are exceedingly wary and inhabit the deeper portions of the water body where they prefer an abundance of aquatic vegetation (DUELLMAN 1970). T2alocohyla picta is a common frog that inhabits a variety of lowland habitats such as savannas, forest edges, and tall forest (DUELLMAN 1970, LEE 1996). During the day and in the dry season, individuals of T. picta have been found in the axils of the leaves of elephant-ear plants (DUELLMAN 1970). After the onset of the rainy season, males congregate at temporary and perma-

Fig. 661. Tlalocohyla loquax (Guayach, CRARC, Costa Rica, 550 m). Photo: G. Kohler

Fig. 662. Tlalocohyla picta (near Chetumal, Quintana Roo, Mexico, 20 m). Photo: G. K6hler

nent ponds and small lakes, forest pools, grassy depressions in pastures, and roadside ditches and call from emergent vegetation (DUELLMAN 1970, LEE 1996). Amplexus is axillary in both T. loquax and T. picta (LEE 1996). The call of T. picta consists of series of high-pitched, insect-like chirping notes, described as "cric-ic-ic-ic" (DUELLMAN 1970, LEE 1996, ROBLE 1997). Clutch size in T. picta was determined to vary between 160 and 956 eggs with an average of 543 eggs per clutch (ROBLE 1997). The females deposit the eggs in small packages at the water surface o r attached to aquatic plants (ROBLE 1997).

Tlalocohyla loquax (GAIGE & STUART 1934), Occas. Pap. Mus. Zool. Univ. Michigan 281: 1; type locality: YIxpuc Aguada, north of La Libertad, El Pethn, Guatemala." Males to 44.7 mm, females to 47.0 mm. On the Atlantic versant from southern Veracruz, Mexico, to Costa Rica, sea level to 1585 m elevation.

Tldocohyla picta (GifNTHER 1901), Biol. Centr. Amer., Rept. Batr., Part 166: 286; type locality: "Mexico, [Veramz,] Jalapa." Males to 21.4 mm, females to 22.1 mm. On the Atlantic versant f?om southern San Luis Potosi, Mexico, southward through northern Guatemala and Belize to northern Honduras, sea level to 1300 m elevation.

Rey to W e l a 1 a No pale lateral or dorsolateral stripe

between levels of axilla and groin, fingers webbing reduced, webbing between Toes III and IV not reaching penultimate tubercle on Toe IV; webbing red in life; posterior surface of thigh red in life; SVL of adult males 36-45 mm, of adult females 39-42 mm ..... ....................................... m*la loquax

b A distinct pale (white or yellowish) lateral or dorsolateral stripe between levels of axilla and groin; fingers moderately webbed, webbing between Toes 111 and ZV reaching penultimate tubercle on Toe IV; webbing yellow or yellowish tan in life; pos-

. terior surface of thigh uniform yellow in life; SVL of adult males 19-21 mm, of adult females 19-22 mm .......... 111aloa&la picta

- - . _ I _ _ n..a..l.l-.__

T: loquax r: picta

1 i t 1

Further Reading DUELLMAN 1970, ROBLF: 1985, LEE 1996, ROBLE 1997, CAMPBELL 1998, M c C m & WILSON 2002, SAVAGE 2002

, ,. 663. , x,,,uA, loquax (near San Ignacio, Cayo, Belize, 240 m). Photo: G. Kijhler

Trach ycephalus

The genus Trachycephalus consists of 12 species of large, robust frogs with a center of diversity in South America; only a single species, T. venulosus (milky treefrog), occurs in Central America (FROST 2010). The latter species was assigned to Phrynohyas for the past decades before FAIVOVICH et al. (2005) placed Pluynohyas in the synonymy of Trachycephalus. The milky treefrog is a widespread, nocturnal, arboreal inhabitant of primary and disturbed habitats. I t passes the dry season sequestered in bromeliads, tree holes and crevices, under bark on standing trees, and in the axils of the leaves of heliconias or bananas (LEE 1996, SAVAGE 2002). At night, they leave their daytime shelters and can be seen perching in the vegetation. Thanks to the large, webbed hands and feet, this species has well-developed parachuting ability (Con 1926). Characteristic for T. venulosus are the thick dermal glands on the dorsum that can secrete a large amount of sticky white, poisonous, water- insoluble mucus, that is highly irritating to mucous membranes and can cause sneez-

ing in humans even without direct contact (DUELLMAN 1956). These secretions have proved to discourage potential predators (LEARY & RAZAFINDRATSPTA 1998) and are also helpful to reduce cutaneous water loss

Trachycephalus venulosus is an explosive breeder and depends on heavy rains to initiate breeding; the males call while sitting or floating in shallow water of temporary ponds (SAVAGE 2002). The advertisement

P call was described as a loud growl, repeated at regular, short intervals (Duellman 1970; LEE 1996). Amplexus is axillary, and a single amplectant pair can produce several egg masses in one night; females lay eggs as a film on the water surface (ZWEIFEL 1964a, LEE 1996). The tadpoles complete metamorphosis in 37 to 47 days (ZWEIFEL 1964a, PYBURN 1967). The tadpoles have very large gills, most likely as , an adaptation to living in warm water with ' a low oxygen level (ZWEIFEL 1964a). Fig. 664. Trachycephalus venulosus (near

Chetumal, Quintana Roo, Mexico, 20 m). Photo: G. Kohler

Trachycephalus venulosus ( L A ~ N T I 17681, Spec. Med. Exhib. Synops. Rept.: 31; type locality: "Indiisn; changed by neotype designation (HEMMING 1958) to "in arbustis et arboribus ad flwnen Teffbn, Brazil. Males to 100.5 mm, females to 113.7 mm. From central Tamau- lipas and southern Sinaloa, Mexico, southward on both versants to central Nicaragua, and on to the Pacific versant through Panama; also widely distributed in tropical South America, sea level to 1610 m elevation.

I Fig. 665. Trachycephalus venulosus (Alami- kamba, Atlantico Norte, Nicaragua, 20 m).

Photo: G. Kohler

Further Reading DUELLMAN 1956, HEMMING 1958, ZWEIFEL 1964a, PYBURN 1967, MCDIARMID 1968, DUELLMAN 1970, 1971a, LEE 1996, CAMPBELL 1998, MANZANILLA et al. 1998, MCCRANIE & WILSON 2002, SAVAGE 2002

271

Triprion petasatus (shovel-headed treefrog) is a bizarre-looking denizen of the Yucatan Peninsula. Earlier, two species (T. petasatus and T. spatulatus) were assigned to the genus Triprion (e.g., DUELLMAN 1970, 2001). S. SMITH et al. (2007) removed T. spatulatus to the genus Diaglena to resolve the detected paraphyly of Triprion with respect to Anotheca. Thus, Triprion is now regarded as monotypic (FROST 2010).

fiprion petasatus is a nocturnal frog that inhabits savannas and low dry and moist forests (DUELLMAN 1970, LEE 1996, CAMPBELL 1998). This species is highly seasonal and apparently remains dormant for the duration of the dry season (LEE 1996, M c C m & WILSON 2002). During the day, they seek shelter in holes of tree and limestone rocks as well as in the recesses of tree trunks and in rock crevices (STUART 1935, LEE 1996, KOHLER 1997). The frogs plug the openings with their casque-like

6 heads, a behavior called "phragmosis", and are almost impossible to extract; the m- ossified dorsal head surface obviously helps to reduce cutaneous water loss (LEE 1996). During the first heavy rains of the wet season, males congregate at temporary ponds, limestone solution pits, and sink holes at night, and call from shrubs and small trees as well as from the edge of the water bodies (LEE 1996, MCCRANIE & WILSON 2002). The call was described as a series of low-pitched notes reminiscent of the quacking of a duck (DUELLMAN 1970, CA~MPBELL 1998). Breeding activity peaks in the early and middle portion of the wet season (months of May to July) (DUELLMAN 1970, MCCRANIE & WILSON 2002). Amplexus is axillar~, and the females deposit the eggs in clumps in the water where the tadpoles complete their development (DUELLMAN 1970, LEE 1996). Tadpoles prefer shaded areas and seek refuge in decaying vegetation on the bottom of the water body (DUELLMAN 1970).

k'lg. 666. Triprion petasatus (near 'I'eabo, Yucath, Mexico, 20 m). Photo: G. Kohler

Trprion petasatus (COPE 18651, Proc. Acad. Nat. Sci. Philadelphia 17: 193; type locality: "vicinity of Mbrida, Yucatan . . . f?om a hole in the rocky wall of the Cenote Pamanche, on the new road to Progreso" 117 km north of Mbrida, Yucatsn, Mexico]. Males to 60.8 mm, females to 74.2 mm. Yucath Peninsula, Mexico, southward to Guatemala and Belize; also at Lago de Yojoa in northwestern Honduras, sea level to 740 m elevation.

I Triprion petasatus

F'urther Reading DUELLMAN & KLAAS 1964, DUELLMAN 1970, '~'RUEB 1970, LEE 1996, KOHLER 1997, CAMPBELL 1998, MCCRANIE & WILSON 2002

Leiuperidae

Leiuperidae

The species now accommodated in the family Leiuperidae (foam nesting frogs and dwarf frogs) traditionally have been included in the family Leptodactylidae. The recognition of Leiuperidae is based on GRANT et al. (2006). As currently understood, this family contains 79 species distributed from southern Mexico across Central America and South America (FROST 2010). Two genera of Leiuperidae with one species in each occur in Central America, the wars &og (Engystomops pustulosus) and the Colombian four-eyed frog (Pleurodema brachyops). The bhgara frog is one the most studied Neotropical -

frogs with one book and dozens of papers Fig- 667. EngJ'stomo~s ~ustdosus (Guaquira,

dedicated to its ecology, life history, and Yaracuy, Venezuela). Photo: J. Sunyer

sexual selection kg., RYAN 1985, RYAN et al. 1996, RON et al. 2005, LG~'~PERT et al. 2007).

Key to Central American species of laemus where it had been placed by LXNCH Leiuperidae (1970b). FUNR et al. (2008) suggested that 1 a A prominent deep purple to black colored the data provided by NASCIMENTO et al.

~ ~ ~ ~ U ~ l ~ & ~ ~ $ h ~ " , ~ f ~ ~ ~ ~ ~ ~ (2005) did not just* the separation of faces of groin, and posterior surface of E n ~ s t o m o ~ s from PhJ'salaemus. Pub- thigh and shank orange; ventral surface lished data suggest that the Panamanian without dark spots H e d a m a bradww~ and South American populations of E. pus-

b No lumbar gland present; hidden surfaces tulosus represent a species distinct from of groin, posterior surface of thigh and the Mexican and Central American popu-

same "lor as adjacent (brown lations north of Panama (RYAN et al. 1996, to grayish brown); ventral surface usually with small to large dark spots .................... WYNN & HEYER 2002, WEIGT et al. 2005, ............................. ~ o p e p u s t u l o ~ u ~ B O H L et al. 2006).

Further Reading GRANT et al. 2006

Where it occurs, Engystomops pustulosus can be quite common. It is a nocturnal and terrestrial species that is found savannas, pastures, and open forests, as well as in and around human settlements, avoiding - both very dry and very wet areas (RAND

Engystomops 1983). It is a leaf litter species hunting various kinds of small arthropods at night.

Eight species are currently recognized in DUE^ (1960) reported on a series of the genus Engystomops, distributed from this species that he obtained by digging southern Mexico across Central America them out of a dry river bank during the dry and South America south to Bolivia (FROST season. For breeding, the frog makes use of 2010). NASCIMENTO et al. (2005) removed smaller bodies of water such as ponds, Engystomops from synonymy with Physa- roadside ditches, road ruts, puddles, pot-

holes, and overflow pools along streams; running streams and permanent large bodies of water are rarely used (RAND 1983). Breeding activities concentrate during the first weeks of the wet season (May to June); males can be observed calling at night while floating on the water with their body inflated (RYAN 1985). The call has two components, a whine and several chug-like sounds (RYAN 1985). The eggs are laid in foam nests, generally under cover, in pools. A detailed account of the construction of the foam nest by this species is provided by HEYER & RAND (1977). The foam can prevent egg desiccation during short periods without rain (RYAN 1985). The eggs hatch within three days or fewer, and the tadpoles leave the foam nest to enter the water below. After four to nine weeks, metamorphosis takes place and the froglets leave the water (RAND 1983, RYAN 1985).

Fig. 668. Engystomops pustulosus (Nusagandi, Kuna Yala, Panama, 350 m). Photo: G. Kohler

Aead. Nat. Sci. Philadelphia 16: 180; type Fig. 669. Enggstomops pustulo~us (6: locality: 'New Grenada, on the River Lorenzo, Suchitep6quez, Guatemala, 210 m, Truandow, Colombia. Males to 34 rmn, females ~ ~ ~ t ~ ~ ~ l ~ ) . Photo: G. Kiihler to 35 mm. On the Atlastic versaat from eouth- ern Vera~uz, Mexico, and on the Pa&c versant imm Oaxaca, Mexico, to e ~ t e r n Panama; also in northern South America, sea level to 1540 m elevation. Pleurodema

Engystornops pustulosus !

Further Reading RAND 1983, CANNATELCA & DUE~LLMAN 1984, RYAN 1985, RYAN et al. 1996, RON et al. 2005, LAMPERT et al. 2007

As currently understood, the genus Pleurodema consists of 14 species with a predominantly South American distribution (FROST 2010). Pleurodema bufonia is the most southern frog, reaching the Straits of Magellan at 53"s latitude (DUELLMAN & VELOSO 1977). Only a single species, P. brachyops, occurs in central and eastern Panama (IBAREZ et al. 1999b). Many species in this genus have a pair of prominent elevated lumbar glands dorsolaterally above the inguinal region. Pleurodema brachyops is a nocturnal and terrestrial species that inhabits open grassy areas, llanos, savannas, xeric shrub forest, and mesic broad-leaf forest (DUELLMAN & VELOSO 1977, HOOGMOED & Go= 1979, M. RYAN pers. com. October

2010). When handled, this fiog inflates its body and elevates the hind parks of the body displaying the conspicuously colored

1 lumbar glands and the orange coloration of the groin and posterior thighs; this suppos- I edly intimidating posture has been interpreted to represent a deimatic behavior typical for animals that cannot flee very fast (MARTINS 1989). This is one of the first fiogs to breed afber the initial rains of the wet season. Thereafter, their distinctive

I call can be heard on any rainy night for the ! rest of the rainv season: toward the end of I "

the season, the aggregations seem to con- / sist almost exclusively of males (HOOG- MOED & GORZULA 1979). For breeding, P. ~ig. 670. Pleurodema brachyops (Cien6ga of bxachyops uses any available temporary Caimanera, Sucre, Colombia). Photo: A. Acosta water body such as shallow pools, puddles. hoofprint8, and tlooded ditches; i d e s tali while floating on the surface of the water and amplexus is d l a r y (HOOGMOED & GORZULA 1979, I B ~ Z et al. 1999b). The eggs are laid in a foam nest floating freely on the surface of the water (HOOGMOED & GORZULA 1979).

P l e d e m a bra&pop% COPE 186% g868", h e . M. Nat, fi. PbhLIphia 20: 312; type bdty: "Wqddena Bveq New Cm& I= Colambia]." SVL to 44 ma. Penfmd8 d0 huwo and central Pamum -Do nozthern South Am& (northern Cohmbia, Venemela to Ou-' BFd" in Panamtt eea Fig. 6'71. PIemdema brachyops (Cien6ga of ld ta k?OU m elmathon.

Caimanera, Sucre, Colombia). Photo: A. Acosta

Further Rsading Fig. 672. Pleurodema brachyops (El Rodeadero DU~UMAN & VELOSO 1977, HOOGMOED & de Gaifa, Magdalena, Colombia). GORZULA 1979, MARTINS 1989, FERRARO & Photo: W. E. Duellman CASAGRAN~A 2009 (Courtesy of the Biodiversity Institute, Univerai@ of Kansas)

Leptodactylidae

Prior to 2006, the Leptodactylidae was a very large unwieldy family of frogs. RWINSKY & MAXSON (1996) suggested that Leptodactylidae (sensu lato) was poly- phyletic and FROST et al. (2006a) parti- tioned the former Leptodactylidae into several reformulated families. Subsequently, GRANT et al. (2006) split the Lepto- dactylidae into the Leptodactylidae (as now recognized) and the Leiuperidae. As currently understood, the family Lepto- dactylidae contains four genera, one of which, Leptodactylus, occurs in Central America (FROST 2010). Most species in this family are moderate-sized frogs, but some species can reach a very large (up to 185 mm SVL) adult size (VITT & CALDWELL 2009).

M e r Reading GORHAM 1966, L m c ~ 1971, RUVINSKY & MAXSON 1996, FROST et al. 2006, GRANT et al. 2006

The genus Leptodactylus consists of 88 recognized species, distributed from southern North America, across Central America and South America, and into the West Indies (FROST 2010). HEYER (1969) rede- fined the genus, and in numerous subsequent papers contributed much to the current taxonomic knowledge of this group of frogs (e.g., HEYER 1970a,b, 1972, 1974, 1978, 1979, 1994, 2002, 2005). Lepto- dactylus are terrestrial or semiaquatic and nocturnal, occupying a variety of habitats including tropical rainforest, marshes, lakes, ponds, and roadside ditches throughout Lowland and Premontane Dry, Moist, and Wet Forest zones (sensu HOLDRIDGE 1967) (SAVAGE 2002). All Leptodactylus build foam nests in which the eggs are laid; nest of L. savagei can contain 2-7 liters of foam (SCOTT 1983e). The foam nests are either floating on the surface of the water

or are placed hidden in vegetation, under debris, or in the mouth of a burrow in streambeds or other seasonal flooded sites (BREDER 1946, SAVAGE 2002). The eggs hatch within two to three days; subsequent rains flood the nest site, and the tadpoles are washed into the water where development is completed. Development is fast, and the tadpoles metamorphose in two to four weeks after hatching (SAVAGE 2002). Several of the Central American species superficially resemble frogs of the genus Lithobates.

Leptodactylus savagei is the largest Central American species in this genus. During the day, this nocturnal frog hides in burrows and under logs; juveniles can be found active at daytime on leaf litter in shady forest situations (SAVAGE 2002). They are important forest floor predators and eat just about any live prey they can fit into their mouths, including arthropods, frogs, lizards, snakes, birds, and mammals (SCOTT 1983e). When caught, these frogs sometimes emit a loud high-pitched scream (SCOTT 1983e). The skin secretions of adult individuals contain noxious compounds that are highly irritating to mucous membranes and little cuts in the skin (VILLA 1969); the mucus compounds can be irritating by diffusion through the air without direct contact, resulting in heavy sneezing and swelling of the eyes (SAVAGE 2002). These secretions can discourage potential predators; R U N et al. (2010) reported on a Bothrops asper that they encountered while it was attempting to consume a specimen of L. savagei. The snake regurgitated the frog and showed signs of poisoning including the loss of the righting reflex; it took more than an hour for the snake to recover. Males call solitar- ily from margins of ponds and streams; the call is a loud "whorup" repeated at intervals of five to ten seconds (BREDER 1946, VILLA 1969). An attracted female is grasped by the male by axillary amplexus and held firmly by the muscular forearms and the stout nuptial spines on thumbs and chest (SAVAGE 2002).

Leptodactylus bolivianw BOULENGER 1 8 9 8 ~ Ann. Mus. Civ. Stor. Nat. Genova, Ser. 2, 19: 131; type locality: "Barraca and Missiones Mosetenes," Bolivia; re6tricted to "Barraca, Rfo Madidi, Bolivia," by leetotype designation (CAPOCACCIA 1957). Males to 95 mm, females to 120 mm. Along the Pacific versant of costa Rica, south through Panama to northern South America, where it has a patchy distribution; also present on the Caribbean slope of Nicaragua and on Providencia and San An&s Islands (Colombia), and San Mipe l and Saboga I~lands (Panama), sea level to 1400 m elevation. Lephikctylm ins- BARBOUR 1906 is regarded as a synonym of L. bolivianus, according to REYER (1970a, 1974) and SAVAGE (2002).

Leptodadyllue h& (BROCCRI 1877~1, Bull. Soc Philomath., Paris, Ser, 7,l: 182; type locality: Tehuantepec (Mexique)." Males to 36 mm, females to 40 mm. Extreme southern Texas, USA, through Mexico and Central America to the Magdalena Valley of northem Colombia, thmngh most of northern Venezuela, a s far as the lower Orinoco basin, and the Llano8 region of both Venezuela and Colombia, sea I d to 1700 m elevation. Much of the literature regarding this species uses the i n m t m e L. labial& (e.g., HEYER 1971), which is now a junior synonym of L. mystacjnus (see HEYElR 1978,2002).

Leptodact-JIm f u s ~ t l ~ (SCHNEIDER 1799), Hist. Amph. Nat.: 130; type locality: implied to be from Surinam; neotype from "Surinam" (HJWIIR 19681). h e m Peninsula to centid Panama, and widely over tropical South America east of the Andes, in Panama sea level to 600 m elevation.

Fig. 674. Leptodactylus boliviii~lus (male, Ponuga, Veraguas, Panama, 35 m).

Photo: G. Kahler

Fig. 675. Leptodadylus bolivianus (male, Ponuga, Veraguas, Panama, 35 m).

Photo: G. K6hler

Fig. 673. Leptodactylus fragilis (road from Fig. 676. Leptodadylus &agjlis (Cem Mariposa, FIacienda Sari Cayetano to Sari Jose El Iddo, Veraguas, Panama, 900 m). Photo: G. Kohler Suchitep6quez, Guatemala, 70 m).

Photo: G. Kohler

Leptodactylus melanonotus (HALLOWELL 1861), Roc. Acad. Nat. Sci. Philadelphia 12: 485; type locality: "Nicaragua." Neotype from "Nicaragua, Zelaya, Bonanzan (HEYER 1970b). Males to 45 mm, females to 55 mm. k o m southern Tamaulipas and southern Sonora, Mexico, across Central America to South America west to the Andes (as far south as Rios Province in Ecuador), sea level to 1440 m elevation.

Leptodactylus poecilochilus (COPE 1862131, Proc. Acad. Nat. Sci. Philadelphia 14: 156; type locality: 'Near Turbo, New Granada" [= Antioquia, Colombia]. Males to 49 mm, females to 50 mm. On the Pacific slope from northwestern Costa Rica to Colombia and

Fig. 677. Leptodactylus savagei (juvenile; San sea level to 1150 Ram6n de La Virgen, Heredia, Costa Rica, Leptodactylus savagei HEYEa 2005, Arq. Zool. 10 m). Photo: G. Kohler Siio Paulo 37: 330; type locality: "Rincon de

Osa, Puntarenas, Costa Rica, OS042'N, 63"29W." Males to 177 mm, females to 185 mm. Northern Honduras on the Atlantic versant and northern Nicarama on the Pacific

Fig. 678. Leptodactylus savagei (Guayacln, CRARC, Costa Rica, 550 m). Photo: G. Kohler

versant south to ~olombia, sea level to 1200 m elevation.

Leptodactylus silvanimbus MCCRANIE, WILSON & PORW 1980, J. Herpetol. 14: 361; type locality: "Belbn Gualcho, Cordillera de Celaque, Depto. Ocotepeque, Depto. Ocotepeque, Honduras, elevation 1700-1900 m (14'29'N, 88"47W)." Males to 53 mm, females to 48 mm. Montaiias de Celaque and Sierra del Merenddn (Department of Ocotepeque) in extreme western Honduras, 1470-2000 m elevation.

Fig. 679. Leptodactylus savagei (Nusagandi, Fig. 680. Leptodactylus silvanimbus (fiom type 1 Kuna Yala, Panama, 350 m). Photo: G. Kahler locality). Photo: J. R. McCranie

I

L. poecilochilus

I

Lepfodactylus ck - 1 C

- L. savagei

Fig. 681. Leptodactylus melanonotus (Lagarto Lodge, Alajuela, Costa Rica, 90 m).

Photo: G. Kohler Leptodactylus I <._I. --,.

I melanonotus

L. fuscus

I -4 Leptodactylus

l b . Fig. 683. Ventral surface of foot in a. Lepto- dactylus fi-agilis (foot and tarsus with prominent tubercles); b. L. poecilochilus (foot and tarsus smooth). Photos: G. Klihler

Key to Leptodactglus 1 a A pale longitudinal stripe present on poste-

rior surface of thigh (Fig. 682); no lateral fringes on toes; males without black keratinized thumb spines; males with paired

....................... lateral external vocal sacs 2 b Posterior surface of thigh uniform, mottled,

spotted, or marbled (Fig. 684b) but without a pale lon&udinal stripe; lateral fringes on toes present or absent; males with black keratinized thumb spines (Fig. 684a);

............... Fig. 682. Posterior surfaces of thigh and shank males with internal vocal sacs 4 in Leptodactylus poecilochilus; note pale Ion@- 2 a Dorsum with six distinct longitudinal tudinal stripe (arrow). Photo: G. Kohler folds; no pale vertebral stripe .....................

................................... Leptodactglus fiLecus b Dorsum with two to four distinct longitudi-

nal folds, or, if with six longitudinal folds, ....... then a pale vertebral stripe present 3

................... 1 ..................................

3 a Ventral surfaces of foot and tarsus with prominent tubercles (Fig. 683a)

Leptda@ylus i b g i k b Ventral surfaces of foot and tarsus smooth,

without prominent white tubercles (Fig. .............. 68313) Leptudactglus poecilochilus

4 a A pair of distinct dorsolateral glandular

I folds; adults large, SVL in adult males 50-177 mm, in adult females 70-185 mm 5

b No pair of distinct dorsolateral glandular folds, although indications of several tuberculate dorsolateral ridges can be present; adults small to moderate-sized, SVL in adult males 30-53 mm, in adult females

................................................. 3548 mm 6 5 a Lower surface of tarsus smooth; distinct

lumbar glands present; lateral fringes on toes absent, although weak ridges present in some individuals; males with or without black keratinized spines on chest; a single black keratinized thumb spine; SVL in adult males 100-177 mm, in adult females

......... 115-185 mm Leptodactglm sa- b Lower surface of tarsus tuberculate; no

lumbar glands; lateral fringes on toes present; males without black keratinized spines on chest; two black keratinized thumb spines; SVL in adult males 50-95

.............. mm, in adult females 70-120 mm ............................ Leptodactglus bolipianus

6 a Well-developed lateral fleshy fringes on toes present; arms not noticeably hypertrophied in adult males; SVL in adult males 30-40 mm, rarely to 45 mm, in adult

.......... ........................ females 3543 mm, rarely to 55 mm

Leptodactglus melanonotus

I b Toes with lateral keels or only weakly developed lateral fleshy fringes; adult

Fig. 684. a. Hand of Leptodactglus melanono- males with hypertrophied arms; SVL in tus; b. Posterior thigh of L. b ~ l i ~ a n u s . adult males 40-53 mm, in adult females

Photos: C: Kiihler 4348 mm ...... Leptodactglus ailvanimbus

--

Fig. 685. Leptodactylus poecilochilus (Meseta de Chorcha, Chiriqui, Panama, 250 m). Photo: G. Kohler

Fig. 686. Leptodactylus fuscus (near Minacu, Fig. 688. Leptodactylus melanonotus (Ponu,-, Goias, Brazil). Photo: W. Van Devender Veraguas, Panama, 35 m). Photo: G. Kohler

FurtherRaading HEYER 1970a,b, 1971,1972,1974, STRAUGHAN & HEYER 1976, HEYER 1978, 1979, HEYER & MAX~ON 1982, Scom 1983e, MCCRANIE et al. 1986, SOLANO 1987, MARTINS 1988, D w o ~ s & ' HEYER 1992, HEYER 1994, HEYER et al. 1996,

Fig. 687. Leptodactylus melanonotus (9 km 2002, HEYER 2002, HEYER & REID 2003, HEYER NW Fortuna, Alajuela, Costa Rica). 2005, HEYER et al. 2005, PONSSA 2008, HEYER et

Photo: G. Kohler al. 2010

Microhylidae The family Microhylidae (narrow-mouthed toads) contains 466 species in 66 genera, which is the largest number of genera of any frog family (FROST 2010). Eight of the nine Central American microhylids are included in the Gastrophryninae, whereas Relictivomer has not been assigned to any of the currently recognized subfamilies. Microhylid frogs occur throughout the tropical and warm temperate regions of North America, Central America, South America, Subsaharan Africa, eastern and southern India, Sri Lanka, southeastern Asia, through to New Guinea and northern Australia (VIW & CALDWELL 2009).

The Central American microhylids are all characterized by having a round to ovoid body with short legs, a very small, narrow head, and a hidden tympanum. The pheno-

typically similar Rhinopluynus can be distinguished easily from the microhylids by the presence of two large spadelike inner metatarsal tubercles (only one inner metatarsal tubercle is present in the Central American microhylids).

All Central American microhylids are secretive fossorial inhabitants of humid forests, which emerge to the surface only during heavy rains for short periods to breed. They have a highly specialized feeding mode, the highly protrusible tongue enabling them to feed on termites and ants underground. The head shape is adapted for breaking into the termite passageways and ant tunnels (SAVAGE 2002). The mucous skin secretions of these frogs apparently efficiently protect them from being attacked by their potential prey (SAVAGE 2002).

Fig. 689. Hypopachus van'olosus (Santa Eulalia, Alajuela, Costa Rica, 825 m). Photo: G. Kahler

282

Key to the genera of Central American Mimhylidae 1 a Inner and outer metatarsal tubercle pre-

sent (Fig. 690) ........................................... 2 b Only an inner metatarsal tubercle present

...... or no metatarsal tubercle (Fig. 691b) 3 2 a Toes with moderate webbing (Fig. 690) .....

................................................. b Toes without webbing (Fig. 691a) ................

.................. GwWplupe (in part: G. usta) 3 a Toes with moderate webbing (Fig. 691b);

venter without distinct markings; dorsal Fig. 690. Foot of Hypopachus variolosus; note surfaces of head, body, and limbs essential- outer metatarsal tubercle (arrow). ly uniform glossy black ..... Nelemoplqme Photo: G. Kiihler

b Toes with basal or no webbing; venter with or without markings; dorsal surfaces of head, body, and b b s essentially not uni-

...................................... form glossy black 4 4 a Venter with large white spots; toes with

...................... basal webbing w d 4 - .......... (in part: G. e l m , G. pictivenkia)

b Venter uniform or with faint flecking; toes without webbing ....................................... 6

6 a Middorsum dark brown, well demarcated from paler (pale reddish brown to yellow in life; pale brown to cream color in preservative) dorsolateral band, bordered below by

............ a dark grayish brown lateral band ................................................ Chiaamoeleie

b Coloration not as above ........................... 6 6 a An isolated yellow spot present in inguinal

region and usually another one in the axilla; inguinal region and posterior surface of thigh without flashy red markings in life; usually a narrow pale stripe on posterior

.................... surface of thigh Eli*& b No discrete yellow spots Present in a d l a r ~ Fig. 691. Foot of a. Gastrophryne usta; b.

or inguinal region; inguinal region and pas- NelBOnoPhryne ate-a, Drawings: M. Vesely terior surface of thigh with flashy red markings in life; no stripe on po&erior surface of thigh ...................... RaZictivrnner

FurtherRsading PARKER 1934, Durn 1949, DE CARVALHO 1954, WALKER 1973, ZWEIFEL 1986, WILD 1995, VAN DER MEIJDEN et al. 2007, & CALDWELL 2009 -

Fig. 692. Relictivomer pearsei (ventral surfaces; Malagana, Bolivar, Colombia, 30 m).


Chiasmocleis is a predominantly South American genus containing 25 recognized species distributed principally in tropical South America, north and east of the Andes (FROST 2010). Only a single species, C. panamensis, enters southern Central America. Most species of this genus show marked sexual dimorphism in the degree of toe webbing (DE CARVALHO 1954, WALKER & DUELLMAN 1974). Specimens of C. panamensis have been found under logs and rocks (RENJIFO & LUNDBERG 1999). The species is reported to be locally common in central Panama where individuals primarily occupy open areas with bushy vegetation; they are only encountered occasionally in the forest ( I B ~ z et al. 1999b). Breeding activities are concentrated during the first half of the rainy season; after heavy rains, the males call from the ground or from partially submerged vegetation in temporary water bodies such as shallow pools and flooded ditches ( I B ~ z et al. 1999b). The eggs are laid in water where the tadpoles develop (IBAREz et al.

diasmocleis panamensis D m , TRAPmo & EVANS 1948, Am. Mus. Novit. 1376: 1; type locality: "Old Panama, Republic of Panama." SVL to 26 mm. Central Panama to Colombia (Caribbean coast and the Uraba Gulf to the Magdalena valley), sea level to 600 m elevation.

Fig. 693. Chiasmocleis panamensis (Gamboa, CoMn, Panama, 29 m). Photo: A. J. Crawford

- *->-*.. ' , ~ , , . , s a A . ~ , ~ - ~ . ~ , . . > . ~ ->...,*..- .**, -, ..- 7

Chiasmocleis panamensis 1 E

Further Reading DUNN et al. 1948, DUNN 1949, NELSON 1972~, WALKER & DUELLMAN 1974

Elachistocleis

The predominantly South American genus Elachistocleis is composed of eleven species, which are distributed from Panama and Colombia southward, east of the Andes, to southern Paraguay, southeastern Bolivia, central and southeastern Brazil, and Uruguay; the genus is also found on Trinidad (CARAMASCHI 2010, TOLEDO 2010, TOLEDO et al. 2010). CARAMASCHI (2010) discussed the taxonomic position of E. ovaLis (SCHNEIDER 1799) and considered this taxon as a nomen dubi- Fig. 695. ~ ~ ~ ~ ~ ~ ~ h r y n ~ elegans ( ~ ~ i ~ ~ f ~ de la

thereby rendering the Panamanian Cruz, Atlhntida, Honduras, 5 m). populations formerly referred to as Photo: J. Sunyer "Elachistocleis ovalid' without an identification. Thus, I have refrained from including a species account under this genus. Elachistocleis are small nocturnal, fossori- a1 frogs. As is typical for most microhylids, GastrODhrvne individuals of Elachistocleis are only encountered during the wet season - usually after heavy rains -when they breed in temporary ponds, often in pastures (TOLEDO 2010). Males have been observed to call with the hind legs in the water and the body out of water, holding onto emergent plant parts with the hands. The amplexus is axillary. The eggs are laid at the water surface attached to floating or emergent vegetation (TOLEDO et al. 2010).

F'urther Reading COCHRAN & GOIN 1970, LAVILLA et al. 2003, TOLEDO 2010, CARAMASCHI 2010, TOLEDO et al. 2010

Five species are currently recognized in the predominantly North American genus Gastrophryne, distributed from the southern USA to Costa Rica (NELSON 1973b). Habitats occupied by the Central American species of this genus include tropical deciduous forest, marshes, and savannas (G. usta), and tropical rainforest (G. elegans and G. pictiventris) (NELSON 1972b). These are nocturnal secretive frogs that are present in the leaf litter of the forest floor at all times of the year (NELSON 1963). Local densities of Gastroplnpe can be as high as 145 individualsha (LIEBERMAN 1986, data for La Selva, Costa Rica) but in other regions it is an uncommon component of the anuran fauna (LEE 1996). Gastrophryne pictiventrs seems to prefer undisturbed forest sites and is less

i frequently found in secondary growth (SAVAGE 2002). All species of Gastrophryne feed on ants, termites, small beetles, and other small insects. Typical for explosive

b*.n; breeders, they arrive at their breeding -/ sites - shallow pools and flooded ditches -

Fig. 694. Elachistocleis sp. (P.N. G. D. Omar after heavy rains, mostly during the Torrijos H., near El Cop6, Cocl6, Panama). months from May to August (STUART 1934,

Photo: R. Brenes CAMPBELL 1998, SAVAGE 2002). The mating

call of G. elegans has been described as a prolonged "baaaaaaa" (CAMPBELL 1998) or a high-pitched, rather nasal "naaaaaaa" (LEE 1996,2000). The males of G. pictiventris have been observed calling while float-

of several males (NELSON 1973c7 SAVAGE 2002). The eggs are laid on the surface of

E ing in the water, with synchronous calling 1

the water and hatch within a few days (MEYER & FOSTER 1996). The tadpole of G. pictiventris has been described and illustrated by DONNELLY et al. (1990b).

I Fig. 696. Gastropluyne pictiventris (Guayachn, CRARC, Costa Rica, 400 m). Photo: B. Kubicki

Gastrophryne elegans (BOULENGER 1882a), Cat. Batr. Sal. Coll. Brit. Mus. Ed. 2: 162; type locality: "Cordovan [= C6rdoba1, Veracruz, Mexico. Males to 26 mm, females to 30 mm. On the Atlantic versant from southern Tamaulipas, Mexico, to Belize and north-central Honduras, with an isolated population in Sian Ka7an Biosphere Reserve (Yucattin Peninsula), sea level to 350 m elevation.

Gastrophgme pietiventris (COPE 18861, Proc. Am. Philos. Soc. 23: 272; type locality: "Nicaragua." Males to 31 mm, females to 38 mm. Atlantic slopes of southeastern Nica- ragua and Costa Rica, sea level to 500 m elevation.

Gastrophgme usta (COPE 1866), Proc. Acad. Nat. Sci. Philadelphia 18: 131; type locality: "Guadalaxara [= Guadalajara, Jaliscol, West Mexicon; apparently in error, according to N m o ~ (1972b). Males to 28.6 mm, females to 30.4 mm. From southern Sinaloa and central Veracruz to Pacific versant Guatemala and El Salvador, sea level to 1000 m elevation.

Fig. 697. Gastrophryne usta (road from Hacienda San Cayetano to San Jose El Idalo, SuchitepBquez, Guatemala, 70 m).

Photo: G. Kohler

Key to Garrtzuphgme 1 a Inner and outer metatarsal tubercle pre-

sent; toes without webbing; venter pale with pepper fine dark spots forming darker

..................... reticulum Gastrophqme wta b Only an inner metatarsal tubercle present;

toes with basal webbing; venter dark with ............ distinct white blotches and spots 2

2 a White blotches on venter separated from adjacent blotches by much less than their diameter and often partially fused with adjacent blotches; a dark brown inguinal spot present above the more or less distinct dark brown lateral band ............................. ................................. Garrtropluyne elegans

b White blotches on venter generally separated from adjacent blotches by at least their least diameter; no dark inguinal spot present above the more or less distinct dark brown lateral band ............................. ........................... ~phgneppctrctrvmtris

Further Reading NELSON 1963, LEGLER 1964, NELSON 1972a,b, 1973b,c DONNELLY et al. 1990b, VALENZLTELA GALVAN et al. 2004

Microhylidae

The genus Hypopachus (sheep frogs) consists of two species whose morphological boundaries are poorly defined (see comments in MCCRANIE & WILSON 2002, SAVAGE 2002). Whereas H. variolosus has a broad mostly lowland distribution from southern Texas (USA) and southern Sonora (Mexico) to Costa Rica, the second species, H. barberi, has been recorded from mountainous regions in Nuclear Central America (MCCRANIE & WILSON 2002, KOHLER et al. 2006). During much of the year, these anurans lead a secretive, primarily subterranean life. When the animals are near the surface (principally during the rainy season), they can be found under rocks and rotten logs, and in debris. Otherwise, they are nocturnal inhabitants of burrows and other cavities, often up to 1 m underground (SAVAGE 2002). They feed mostly on ants, termites and other small arthropods. Typical for an explosive breeder, these frogs emerge after heavy down- pours, mostly during the early rainy season, and primarily use temporary water bodies such as flooded pasture, roadside ditches, and marshes for reproduction (LEE 1996, SAVAGE 2002). The male mating call is a sheep-like bleat ("baaaaaaa") that may last 2-3 seconds (MEYER & FOSTER 1996, CAMPBELL 1998). Males have been reported to call from shallow depressions beneath

Fig. 698. Hypopachus barben' (3.5 km NNW Guajiquiro, La Paz, Honduras, 2205 m).


Fig. 699. Hypopachus variolosus (Morgans Rock, Rivas, Nicaragua). Photo: J. Sunyer

vegetation a t the edge of the water and also while floating on top of the water (CAMPBELL 1998). Synchronization of calling males can result in rhythmic bursts of sound (SAVAGE 2002). Amplexus is axillary, and the eggs (600-800 per clutch) are laid as a loose mass on the surface of the water where they hatch within 12-24 hours; metamorphosis takes place in about three to four weeks (LEE 1996, MEYER & FOSTER 1996, SAVAGE 2002).

Hypopachus barben' (SCHMIDT 19391, Field Mus. Nat. Hist. Publ., Zool. Ser. 24: 1; type locality: "TecpBn, Solo16 [in error for Chimaltenango, according to NELSON 1973a: 151, Guatemala." Males to 40.5 mm, females to 45.3 mm. Central Chiapas, Mexico, to Guatemala, El Salvador, and southwestern Honduras, 1300-2500 m elevation.

Hypopachus variolosus (COPE 1866), Proc. Acad. Nat. Sci. Philadelphia 18: 131; type locality: "Arriba [= Meseta Central according to SAVAGE 19741, Costa Rica." Males to 39 mm, females to 53 mm. On the Atlantic versant from southern Texas, USA, to Costa Rica, and on the Pacific versant from southern Sonora to Costa Rica, sea level to 1610 m elevation.

&Y to J%Ppecbm 1 a Outer metatarsal tubercle small, widely

separated from the inner metatarsal tubercle (inner and outer metatarsal tubercles seoarated from each other bv 0.9-1.7 times Gdth of outer metatarsal "tubercle) (Fig. 700b) .......................... &~o.@?&uB ibanban' 1

b Outer metatarsal tubercle large, narrowly separated from the inner metatarsal tubercle (inner and outer metatarsal tubercles separated from each other by 0.3-0.6 times width of outer metatarsal tubercle) (Fig. 700a) ..................... &pop&118 v d o h ~ ~

Fig. 700. Foot of a. Hypopachus variolosus; b. H. barben'. Drawings: M. Vesely

Hypopachus w Further Reading NELSON 1973a, 1974, MCCRANIE & WILSON 2002, K~RLER et al. 2006

Fig. 701. Nelsonophzyne aterrima (Coco de Siquirres, Lim6n, Costa Rica, 250 m).

Photo: B. Kubicki

The genus Nelsonophyne is restricted in distribution to lower Central America and northwestern South America; it contains only two specim, one of which, N. atenima, occurs in lower Central America. Nel~onophqyne atenima is an uncommon, fossorial, nocturnal h g that has been collected from beneath debris and surface litter on the forest floor and also from a depth of 80 cm in the ground (SAVAGE 2002). The individual of N. aterrima pictured in Figure 701 was unearthed by heavy machinery removing sections of secondary growth during the (ICE) hydroelectric dam project of the Rio Reventaz6n, near Coco de Siquirres, Costa Rica ( B m KUBICM pers. comm. July 2010). Gravid females have been collected during the months of June to August with a reported clutch size of 70-80 eggs in the oviducts (SAVAGE 2002). DONNEUY et al. (1990b) collected tadpoles of N. aterrima from a small puddle at the edge of a logging road in early October,

N e ~ m n o p ~ e a t e h a (GONTHER 19011, Biol. Centr. Amer., Rept. Batr., Part 166: 210; type locality "Costa Rica." Males to 61 mm, females to 67 mm. Costa Rica and Panama to Colombia and northwestern Ecuador, near sea level to 1600 m elevation.

I

PARKER 1934, DE CARVACHO 1954, BOGART & 1 NELSON 1976, DONNELLY et al. 1990, SAVAGE

2002

Relictivomer

Relictivomer is a monotypic genus, and some authors have questioned its validity and suggested that it might be a synonym of Elachistocleis (e.g., FROST 1985, ZWEIFEL 1986). Relictivomer pearsei is a fossorial species that lives in termite nests, feeding on larvae and adults of termites (RENJIFO & LUNDBERG 1999). As with other microhylids, it only emerges to the surface after heavy rains t o reproduce in temporary pools and flooded areas (RENJIFO & LUNDBERG 1999).

Relictivomer pearsei (RUTHVEN 1914), Proc. Biol. Soc. Washington 27: 77; type locality: "vicinity of Fundacion, [Sierra Santa Marta,] Colombia." SVL to 41.5 mm. Caribbean versant of central Panama and northern Colombia, including the Middle Magdalena Valley, to northwestern Venezuela (Maracaibo drainage), near sea level to about 500 m elevation.

Fig. 702-703. Relictivomer pearsei (Los Algarrobos, Chiriqui, Panama, 125 m).

Photos: A. Hertz

Further Reading DE CARVALHO 1954, COCHRAN & GOIN 1970, RENJIFO & LUNDBERG 1999

Pipidae

Pipidae

The family Pipidae contains about 30 species in five genera, all of which are exclusively aquatic. Characteristics of this family include the absence of a tongue, a dorsoventrally flattened body, completely webbed feet, small dorsally placed eyes, highly modified ears for producing and receiving sound underwater, and a lateral line system ( V m & CALDWELL 2009). The family Pipidae has two centers of distribution: tropical South America (one genus) and sub-Saharan Africa (four genera).

The genus Pipa is a Neotropical endemic with seven recognized species (TRUEB & CANNATELLA 1986). Males in this genus lack vocal cords but are able to communi- cate underwater via "clicking" sounds generated through a modified laryngeal apparatus (RABB 1960b). Species of the genus Pipa are known for their elaborate mating behavior, which includes the performance of aquatic turnovers by the amplectant pair, allowing for the eggs to be deposited on the female's back (RABB & RABB 1960, RABB & SNEDIGAR 1960). The fertilized eggs sink into the skin and eventually become embedded completely. In the South American P. pipa, development is direct, and fully formed froglets hatch from the skin pockets (RABB & FLUB 1963). The only

Central American pipid has free-living tadpoles that initiate their development within individual pits in the highly vascular- ized skin of the female's dorsum. The free- living tadpoles have a wide mouth as seen in anterior view.

Pipa myersi i s the only Central American representative of this genus. It was collected from swamps in high-canopy lowland rain forest and also from ponds in secondary growth; the water was described as clear but brown-colored containing abundant leaves (HEATWOLE 1963, TRUEB 1984). The tadpoles P. myersi are dark gray with a silvery white abdominal peritoneum (in life); barbels and keratinous mouthparts are absent; tadpoles were more common near the surface of the water at night than by day (TRUEB 1984).

Pipa myersi TRUEB 1984, Herpetologica 40: 225; type locality: "7 km above the mouth of the Rio Ucurganti, Provincia Daribn, Panama, 30 m." Males to 35.5 mm, females to 44.0 mm. Eastern Panama (Rio Ucurganti and Rio Canglon, Dari6n Province) and northeastern Colombia, about 30 m elevation.

F'urther Reading RABB & RABB 1960, RABB & SNEDIGAR 1960, HEATWOLE 1963, RABB & RABB 1963, GORHAM 1966, TRUEB 1984, TRUEB & CANNATELLA 1986

Fig. 704. Pipa myersi (KU 113662, paratype); a. dorsal view; b. ventral view. Photo: A. Campbell (Courtesy of the Biodiversity Institute, University of Kansas)

Ranidae

Ranidae

The Ranidae (true fiogs) has the widest distribution of any fkog family, with members occurring in North America, South America (south to north-central Bolivia and eastern Brazil), Europe, Asia, Madagascar, Africa, and the East Indies to New Guinea and extreme northern Australia. Typically, true frogs are smooth, moist-skinned frogs, with large, powerful legs and extensively webbed feet. Many ranids are aquatic or live close to water. However, there are also arboreal species of the family Ranidae and even species that can live in brackish water (GORDON et al. 1961). All Central American members of this family belong to a single genus, Lithobates, divided into two species groups, the Lithobates pipens species group (in Central America: L. brownonun, L. forren; L. miadis, and L. taylori) and the Lithobates palmipes species group (the remaining species as listed below).

HILLIS & DE SA (1988) published a revision of the Lithobates palmipes species group. However, our state of taxonomic knowledge regarding the Lithobates pipiens species group is less advanced. Much uncertainty is associated with the nominal species L, macroglossa (BROCCHI 1877~). Some authors considered it to be synonym of L. maculata (e.g., STUART 1963, HOLMAN & BIRKENHOLZ 1963, h~ 1976). S m et al. (1966:171) designated MNHN 6321 as the lectotype of Lithobates macroglossa and stated that this specimen-"clearly represents the species Rana pipiens." MCCRANIE &WILSON (2002) analyzed the morphological variation in frogs of the L. pipiens species complex in Honduras and provided a lengthy discussion of the complex situation there. They concluded that at this point the available data on Honduran material can be interpreted to support

fig. 705. Lithobates vibican'us (subadult; near entrance to P.N. Juan Castro BIanco, AIajuela, Costa Rica, 1900 m). Photo: G. Kiihler

three possible taxonomic scenarios: one, two, or three distinct species with possible hybridization among the forms. Obviously, additional studies including molecular methods are needed in order to clarify these issues. We have even less published information on the status of the Guate- malan populations of the L. pipiens species complex aside from that provided by HrrJJs (1988), who mentioned that an undescribed species of this complex occurs on the Meseta Central of Chiapas and the Central Plateau of Guatemala. Because there is no way to morphologically delimit L. macroglossa from L, forreri and L. brownom and no published evidence is available to support the species status of this form, I prefer not to recognize L. macroglossa as a valid species until a taxonomic revision of the Central American members of the L, pipiens species complex has been published (see also comments in HILLIS & DE SA [1988], who considered L. macmglossa to be a synonym of L. berlan- diem?. The Central American s~ecies of Lithobates are semiaquatic denizens of ponds, slow-moving streams, river over- flows, swamps, and marshes active both at day and at night. They are characteristic inhabitants of the aquatic margin (DUELLMAN 1963a); when disturbed, these frogs jump into the water and swim to the

bottom t o conceal themselves among debris. Males call w i th their paired vocal sacs inflated while floating o n the water surface. The eggs are deposited in globular masses, often a t tached t o submerged vege- tat ion o r bottom debris (HILLIS & DE SA 1988, SAVAGE 2002).

Fig. 706. Lithobates brownorum (near Chetumal, Quintana Roo, Mexico, 10 m).

Photo: G. Kohler

Fig. 707. Lithobates forreri (Zona Protectors El Rodeo, Alajuela, Costa Rica).

Photo: G. Kohler

Fig. 708. Lithobates taylori (Cerro Mariposa, Veraguas, Panama, 900 m). Photo: G. Kohler

Lithobates brownorum (SANDERS 19731, J. Herpetol. 7: 87; type locality: "41 mi. W of Xicalango, Campeche, Mexico." Males to 88 mm, females to 91 mm. Southern Veracruz and northeastern Oaxaca east through the Yucatan Peninsula and the uplands of Chiapas to eastern central Nicaragua, near sea level to 1650 m elevation.

Lithobates f0rFer.i (BOULENGER 1883a1, AM. Mag. Nat. Hist. Ser. 5, 11: 343; type locality: "Presidion, Sinaloa, Mexico. Males to 90 mm, females to 114 mm. On the Pacific versant from southwestern Sonora, Mexico, to Costa Rica, sea level to 2260 m elevation.

Lithobates juliani ( H ~ L I ~ & DE SA 19881, Herpetol. Monogr, 2: 7; type locality: "SW end of Little Quartz Ridge, Maya Mountains, Toledo District, Belize, 16"24'N, 8g06W, elevation 915 m." Males to 70 mm, females unknown. Endemic to the Mayan Mountains in Belize, 450-920 m elevation.

Lithobates maculatus ( B R O C C ~ 1877~1, Bull. Soc. Philomath., Paris, Ser. 7, 1: 178; type locality: "Totonicapam (Mexique)" [= Totonicapan, Guatemala]. Males to 75 mm, females to 113 mm. On the Atlantic versant from Chiapas, Mexico, to central Nicaragua; and on the Pacific versant from eastern Oaxaca, Mexico, to Honduras and El Salvador, 40-3000 m elevation.

Lithobates miadis (BARBOUR & LOVERIDGE 19291, Bull. Mus. Comp. Zool. 69: 143; type locality: "Little Corn Island, forty miles off the Nicaraguan coast", Nicaragua. Males to 90 mm, females to 97 mm. Isla de Maiz Pequefia, Nicaragua.

Lithobates taylon. (SMITH 19591, Herpetologica 15: 214; type locality: "Peralta, [Cantdn de Turrialba, Provincia de Cartago,] Costa Rica." Males to 78 mm, females to 88 mm. Eastern Nicaragua to southeastern Costa Rica, 60-1860 m elevation.

Lithobates vaillanti (BROCCHI 18771, Bull. Soc. Philomath., Paris, Ser. 7, 1: 175; type locality: "sur les rives de la rivi&re de Mullins prhs de Belize ([British] Honduras)." Males to 94 mm, females to 125 mm. Southern Veracruz and southeastern Oaxaca, Mexico, to Panama; also widely in northwestern South America (to southwestern Ecuador), sea level to 990 m elevation.

Lithobates vibicarius (COPE 18941, Proc. Acad. Nat. Sci. Philadelphia 46: 197; type locality: "Rancho Redondo on the divide of the Irazu Rangen and "Isla Nueva near the head of the Rio Sucio, on the Atlantic side", Costa Rica;

restricted by lectotype designation (ZWEIFEL 1964) to "Rancho Redondo, on the divide of the Irazu rangen, Cant6n de Goicoechea, Provincia de San Jose, Costa Rica. Males to 73 mm, females to 92 mm. Cordilleras de Tilarh, Central, and Talamanca of Costa Rica and western Panama, 1500-2700 m elevation.

Lithobates warszewitschii (SCHMIDT 18571, Sitzungsber. Akad. Wiss. Wien Phys. Math. Naturwiss. R1. 24: 11; type locality: "Neugranada." Males to 52 mm, females to 63 mm. On the Atlantic versant from northeastern Honduras to eastern Panama, on both versants in Costa Rica and Panama, sea level to 1740 m elevation.

Fig. 709. Litho,,,.s miadis (Isla de Maiz

Fig. 711. Lithobates vibicarius (adult; near entrance to P.N. Juan Castro Blanco, Alajuela, Costa Rica, 1900 m). Photo: G. Kohler

Fig. 712. Lithobates maculatus (Laguna del Cem, Cop&, Honduras, 770 m). Photo: G. Kiihler

~equeiia, Nicaragua). Photo: J. Sunyer 1

I Fig. 713. Lithobates vaillanti (P.N. Ma~~llnillo, Limb, Costa Rica, 10 m). Photo: G. Kohler

I Fig. 710 (above). Lithobates julim' (Columbia River Forest Reserve. Toledo. Belize).

Photo: J. C. Lee (Courtesy of the Biodiversity Institute, University of Kansas) Figg. 714 (right). Lithobates vaillanti (Bartola, Rio San Juan, Nicaragua, 30 m).Photo: G. Kijhler

Key to Lithobatee 1 a Skin on dorsum denticulate; toe tips

.............................. expanded (Fig. 719a-c) 2 b Skin on dorsum smooth or rugose but not

denticulate; toe tips not expanded (Fig. ......................................................... 719d) 6

2 a Tympanum large (greater than eye diameter); no distinct dark face mask no distinct

............................. pale supralabial stripe ...................................... Lithobates vaillanti

b Tympanum smaller (less than eye diameter); distinct dark face mask present; a dis-

....... tinct pale supralabial stripe present 3 3 a Two metatarsal tubercles; toe pads pre-

sent; webbing on hind feet reduced (Fig. Fig. 715. Lithobates warszewitschii (3 km SE 719b) .......................................................... 4 Ayapal, Atlantic0 Norte, Nicaragua, 200 m). b One metatarsal tubercle; toe pads absent;

Photo: G. Kohler webbing on hind feet nearly complete (Fig. 719c) ......................................................... 5

4 a Posterior surface of thigh with large pale (yellow in life) spots or bars on a dark background (Fig. 718); toe tips bordered laterally by a well-defined marginal grooves; iris gold above and brown below in

.................... life Lithobates warezedtschii

I Fig. 716. Lithobates warszewitschii iRio San

b Posterior surface of thigh uniform orange to bright red in life (uniform pale brown in preservative); no grooves around margins

......................... of toe tips; iris green in life ................................... I Lithobates vibimCBRus

1 5 a Tympanum length almost as large as eye diameter; flanks uniform dark brown; no vocal slits in males ....... Lithobates juliani

b Tympanum length approximately half the diameter of the eve or less: flanks with -

Juan, Nicaragua). Photo: G. Kohler dark blotches; vocd slits present in adult ....................... males Lithobates madatus

6 a Dorsal surfaces of lower hind limbs with irregular longitudinal dark bands; restrict-

..... ed to Isla de Maiz Pequeiia, Nicaragua ........................................ Tithobates miadis

b Dorsal surfaces of lower hind limbs with dark crossbars; occurs on mainland Central America ....................................... 7

Fig. 717. Lithobates warszewitschii Fig. 718. Coloration of posterior thigh in (Guayach, CRARC, Costa Rica, 550 m); note Lithobates warszewitschii (Rio Curinwas, different coloration of male (right) and female BOSAWAS, Atlantico Norte, Nicaragua). (left). Photo: G. Kohler Photo: G. Kohler

7 a Dorsolateral folds usually continuous, pos- dorsal blotches usually 13 or fewer; vocal terior portion not inset medially; dorsal sacs not pigmented in adult males; blotches usually 15 or more; vocal sacs Miillerian ducts present or absent in adult darkly pigmented in adult males; males; occurs on the Caribbean versant .8 Miillerian ducts present in adult males; 5 a ~~~u~~~~ ducts present in adult males .... occurs on the Pacific versant ...................... ............................... Lithobates brownorum ........................................ Lithobah h r i

b Miillerian ducts absent in adult males b Dorsolateral folds usually interrupted, ........................................ Lithobates taylon'

with the posterior portion inset medially;

Fig. 719. Foot of Lithobates vaillanti; b. L. warszewitschii; c. L. maculatus; d. L. taylori. Drawings: M. Vegely

..... -L------ ------.> . . . . a .

L. brownoturn i Y L. forreri

L. rniadis 1

L. rnaculatu: + L. juliani .

6 I

Lithobates a

d *..a

L. warszewitschii L. vibicarius t

i

$

1 B .a

1 Lithobates

Further Reading ZWEIFEL 1964b, SMITH et al. 1966, ZWEIFEL 1967, LEE 1976, HILLIS et al. 1983, HILLIS & FROST 1985, H n m 1988. HILLIS & DE S A 1988. VILLA 1988, 1990, ACOSTA-GALVIS 1999, RAM[REz et al. 1998,

Rhinophrynidae

The Mexican burrowing toad (Rhino- phquus dorsalis) is the only living species of the genus Rhinophrynus and the family Rhinophrynidae. The family was once more widespread, including species ranging as far north as Canada, but these became extinct during the Oligocene (SAVAGE 2002). Rhinophgmus dorsalis is a peculiar frog with a small, cone-shaped head and four short, robust limbs projecting from a large, globular body. Its eyes are relatively small, and the tympanum is not visible. It feeds underground, specialized on a diet of ants and termites. The tongue that is not attached at the front can be protruded well beyond the tips of the jaws - an adaptation for feeding on subterranean insects (TRUEB & GANS 1983).

Rhinophrynus dorsalis is an inhabitant of mostly arid environments with friable soils that permit burrowing, such as savannas, open dry forests, and non-forested areas, as well as cultivated fields, pasture, and gardens (DUELLMAN 1971~). These fossorial frogs remain inactive underground in deep burrows for most of the year and emerge only after and during heavy rains for brief period of breeding. They will breed any time of the year provided it rains strongly enough. This frog is an explosive breeder that uses temporary pools, roadside ditches, and flooded areas created by heavy rainfall for egg laying (JAMES 1966). The males call at night while floating, emitting

' ->".-- . * --- - ----#-~..---- Rhinophrynus dorsalis

a loud, low pitched "wh-o-o-o-a" with a ris- ing inflection a t the end (FOUQUETTE 1969, FOSTER & MCDIARMID 1983). Large choruses of R. dorsalis may be heard for several kilometers (FOSTER & MCDIARMID 1983). Amplexus is inguinal. Both the female and the male are submerged during egg laying; clutch size may range from 2000 to 8000 eggs (STUART 1935, FOSTER & MCDIARMID 1983). The eggs take several days to hatch, and the tadpoles develop over one to three months, depending on food availability (STUART 1961). Adults reach sexual maturity at 60-65 mm (FOUQUETTE 1969).

Rhinophrynus dorsalis D ~ R I L & BIBRON 1841, Erp. Gen. 8: 758; type locality: "Vera- Cmz," Mexico. Males to 75 mm, females to 89 mm. On the Atlantic versant from southern Texas, USA to northeastern Nicaragua; and on the Pacific versant from the Rio Balsas (Mexico) to Costa Rica, sea level to above 700 m elevation.

Fig. 720. Rhinophrpus dorsalis (2 km W Tegucigalpita, Honduras, 40 m).


Fig. 721. Rhinophrynus dorsalis (near Teguci- galpita, Honduras, 40 m). Photo: G. Kohler

Fig. 722. Rhinophzynus dorsalis (near Tegucigalpita, Honduras, 40 m). Photo: G. Kohler

Fig. 723. Rhinophrynus dorsalis (Cintalapa, Chiapas, Mexico, 577 m); male calling while floating. Photo: A. Ramirez Velzizquez

Fhrther Reading STUART 1961, FOUQUETTE & ROSSMAN 1963, G o m 1966, NELSON & NICKERSON 1966, F O U Q U E ~ 1969, DUELLMAN 1971c, FOSTER & MCDIARMID 1983, TRUEB & GANS 1983, GARRETT & BARKER 1987

Tadpoles

The Tadpoles of Central American Anurana

Most species of Central American anurans have fiee-living tadpoles; exceptions are the species in the families Craugastoridae, Eleutherodactylidae, Hemiphractidae, and Strabomantidae that have direct development without aquatic free-swimming tadpoles (MCDIARMID & ALTIG 1999). Available data on the tadpole morphology of the majority of Central American anurans make possible species identifications based on tadpoles. Tadpoles usually remain longer and at greater individual densities at the breeding site than the adults and, therefore, are easier to detect than the adults. Thus, including tadpoles in regional amphibian inventories can increase the detectability threshold and time period of the species' presence at the study site, especially in the so-called explosive breeders. However, tadpoles of several Central American anurans remain unknown (Table 2). The species with unknown tadpoles, mostly bufonids, dendrobatids, and hylids, are either rare or difficult to find, are explosive breeders with short larval periods, or have not been differentiated from congeners (ALTIG 1987). The following key is useful for specimens between GOSNER (1960) Stages 25 and 40; generally, Stage 40 tadpoles are two to two and a half times as long as those at Stage 25 (SAVAGE 2002). In some few cases, it is not possible to distinguish between the tadpoles of two species, and in these cases they are placed in a single portion of a couplet and separated by "or." More than 30 ecomorphologi- cal guilds have been distinguished among tadpoles (ALTIG & JOHNSTON 1989, SAVAGE 2002). These larval adaptations reflect mostly ecological differences related to the type of habitat (lentic versus lotic water), the niche in the water column, and the feeding mode and food items (SAVAGE 2002).

Terminology of tadpole morphology follows MCDIARMID & AZTIG (1999). Figure 724

gives an overview on the tadpole body and tail morphology. The spiracle is the exit for respiratory water and its number, location, and morphological configuration varies among species. The vent tube is the posterior opening of the digestive tract and its position (medial versus dextral) is useful for tadpole identification. The oral disc contains the keratinous and soft tissue structures adjacent to the mouth. The anterior (called "upper" by some authorstposterior (called "lower" by some authors) direction of the oral disc is the same as the head-tail direction. Oral discs can have emarginations (Fig. 725 left side) or not (Fig. 725 right side). The jaw sheaths of the beak are usually ornamented by serrations. Tooth rows consist of keratinous spicules and are found anteriorly and posteriorly to the mouth. The labial tooth row formula describes the number and arrangement of the labial tooth rows. The numbers before the forward slash indicate how many tooth rows are on the anterior labium and the numbers after the forward slash indicate the number of tooth rows on the posterior labium. Tooth rows are always counted in an anterior to posterior direction. Tooth rows in the anterior labium are represented by a letter A followed by the respective row number, whereas rows in the posterior labium are represented with a letter P followed by its respective number. Thus, as examples, the first tooth row on the anteri- or labium is termed A-1, and the second row on the posterior labium is called P-2. The oral disc usually is bordered by rows of marginal papillae. The marginal papillae can occur in one (uniserial), two (biserial), or several (multiserial) rows. Submarginal papillae are situated on the face of the oral disc, particularly common near the lateral ends of the lower tooth rows besides the jaw sheaths and the emarginations. Other abbreviations used in the following tadpole key are BL (body length) and BW (body width).

Tadpoles

\ \ \ Vent tube ventral fin

b. 'urnbelliform oral disc d. tail rnuiculature

Fig. 724. Lateral view of tadpoles: a. Dendrobates auratus (spiracle ventrolateral; tail tip rounded); b. Silverstoneia flotator (spiracle lateral, located near midline of side of body; large umbelliform [funnel-ahaped] oral disc; tail tip long and attenuate); c. Dendropsophus ebraccatus (spiracle lateral, located near midline of side of body; dorsal fin high, extending onto body; tail M a t i n g in a thin flagellum); d. Gthobates taplori (spiracle lateral, located near midline of side of bodx dorsal fin high, extending onto body). Drawings: A Schulze

anterior tooth rows anterior ("upper')

emarginate disc

posterior tooth rt

marginal papillae posterior ("lower") (in two rows)

marginal papillae (in one row)

Fig. 726. Morphological characteristics of the oral disc and mouthparts of tadpoles. Drawing: A. Schulze

Tadpoles

Table. 2. Central American anuran species without published data on tadpole morphology. Species marked with an asterisk have either only hatchlings known, have not been sufficiently well described, or have tadpoles that cannot be allocated to the respective species with confidence.

Ameerega maculata Atelopus chiriquiensis* Atelopus chirripoensis Atelopus glyphus Atelopus limosus Atelopus senex Chaunus centralis Crepidophryne chompipe Crepidophryne epiotica Crepidophryne guanacaste Dendrobates claudiae Dendrobates fulguritus*

Dendrobates vicentei* Dendropsophus robertmertensi Duellmanohyla lythrodes Ecnomiohyla miliaria Ecnomiohyla minera Ecnomiohyla thysanota Ecnomiohyla valancifer Exerodonfa bivocata Exerodonta chimalapa Exerodonta perkinsi Incilius aucoinae lncilius bocourti lncilius campbelli* lncilius canaliferus lncilius karenlipsae lncilius melanochlorus lncilius peripatetes lncilius signifer lncilius tacanensis lsthmohyla infucata lsthmohyla insolita* lsthmohyla melacaena lsthmohyla xanthosticta Lithobates miadis Plectrohyla lacertosa Plectrohyla pycnochila Ptychohyla dendrophasma Rhinella acrolopha Rhinella alata

MYERS 1982 SAVAGE 2002 SAVAGE & BOLAROS 2009 DUELLMAN & LYNCH 1969 IBAFJEZ et al. 1995 SAVAGE 2002 NARVAES & RODRIGUES 2009 VAUGHAN & MENDELSON 2007 VAUGHAN & MENDELSON 2007 VAUGHAN & MENDELSON 2007 JUNGFER et al. 2000 SILVERSTONE 1975 JUNGFER et al. 1996 KOHLER et al. 2006 SAVAGE 2002 SAVAGE 2002 DUELLMAN 2001 DUELLMAN 1970,2001 DUELLMAN 1970, 2001 DUELLMAN 1970, 2001 MENDELSON & CAMPBELL 1994 CAMPBELL & BRODIE 1992 O'NEILL & MENDELSON 2004 ALTIG 1987 MCCRANIE & WILSON 2002 KOHLER et al. 2006 MENDELSON & MULCAHY 2010 O'NEILL & MENDELSON 2004 SAVAGE 1972, SAVAGE & DONNELLY 1 992 MENDELSON et al. 2005 ALTIG 1987 DUELLMAN 1970, 2001 MCCRANIE & WILSON 2002 MCCRANIE & CASTENEDA 2006 SAVAGE 2002 ZWEIFEL 1967 DUELLMAN & CAMPBELL 1992 DUELLMAN & CAMPBELL 1992 CAMPBELL et al. 2000, DUELLMAN 2001 TRUEB 1971 R. IBAREZ pers. comm. Oct. 201 0

Tadpoles

Key to the Tadpoles of Central American Anmans

1 a Keratinized jaw sheaths absent .............. 2 b Keratinized jaw sheaths present ........... 10

2 a Two lateral or ventrolateral spiracles; barbels present or absent .............................. 3

b One ventral spiracle; no barbels .............. 4 3 a Barbels present a t corners of mouth ..........

................................ R b i n o ~ ~ d d ............................... b No barbels Pipa qpmi

4 a Spiracle opens near midbelly; no oral flaps ............................. N h o p l q m e aatenima

b Spiracle opens on lower abdomen, near vent; oral flaps present ............................ 5

5 a Oral flaps papillate or scalloped, medial margins overlapping or abutting on midline (Fig. 726a) ............................................. ....... Yypopachus vaflvaflollaeurr or H. barban'

b Oral flaps not papillate or scalloped, medial margins parallel or divergent (Fig. 726b)

6 a Tail terminating in a thin flagellum (E'ig. ................. 724c) Chbmocl& p a m m d

b Tail not terminating in a thin flagellum 7 7 a Caudal fins transparent with dark brown

spots; upper half of lateral body brown, lower half pale gray or dark brown with pale spots ..................................................... ~ e i E ~ o r R a l i c t m a m a r ~ ~

b Caudal fins uniform brown or mottled with brown and white; lateral surface of body uniform brown or black, or brown, mottled with white ................................................. 8

8 a Tip of tail pointed; caudal h s low, at level of midtail, fin height less than diameter of tail muscle ................... - e d

b Tip of tail rounded; caudal fins high, a t level of midtail, fin height greater than diameter of tail muscle ............................ 9

9 a Caudal fins mottled with brown and white; lateral surface of body brown, mottled with white ................ ~~e pi&vantriS

b Caudal fins uniform brown; lateral surface of body uniform black .................................. .................................. G@@Pkme el-

10a Labial tooth rows absent ....................... 11 b Labial tooth rows present ...................... 16

lla Tail terminating in a thin flagellum ..... 12 b Tail not terminating in a thin flagellum 15

1% Posterior labium with papillae-like protuberances ................................................ 13

b Posterior labium without papillae-like protuberances ............................................... 14

Fig. 726. Oral flaps of a. Hypopachus barberi; b. Gastrophryne usta. Drawings: M. Vesely

Fig. 727. Tadpole of Dendropsophus ebraccatus (GuayacBn, CRARC, Costa Rica, 550 m).

Photos: G. Kiihler

13a Vent tube medial, in preservative, dorsal and lateral surfaces of body dark brown, mottled with tan and a white stripe extending from mouth posteriorly below eye along body; caudal fins pale with gray or black blotches (colorful in life; Fig. 727). ......................... DenmOp~ophus ebraaxtus

b Vent tube dextral, in preservative, dorsal and lateral surfaces of body black with a dorsolateral tan stripe; caudal fins black with irregular tan vertical marks ....................... DandropBophae subocalans

14a Postorbital dark stripe absent or extending onto tail; no dark spots on tail .................. Lhdlvpeophus PLI-QD~~*UB

Tad-

b Postorbital dark stripe restricted to body; dark spots ........ Dandropophw &bod&

16a Byes dorsal; moderate oral disc, protuberant labia, velum covering anterior jaw sheath ........... Iethmela picadai (in part)

b Eyes lateral; large umbelliform oral disc ...... (Figs. 724b, 728) dlilvtmnku~aia flotatnr

l6a Enlarged ventral sudorial disc present 17 b Enlarged ve'ntral sucturial disc absent . 18

178 Tail musculature black with contrasting

I pale tan (pale yellow in life) bands a d spots .......... Awph@mw chyw&mw

b Tail musculature not black with contrasting pale bands and spots ............................ ....................... I A t d b p u ~ c e r t u s o r A . ~ ~ ~

18a Labial tooth rows 011, YO, 210, l/1, or Y2 19 b Labial tooth rows 212 or greater ............ 25

19a Vent tube dextral; papillae present on .................................. upper disc margin 20 b.' L . b Vent tube medid no papillae on upper disc

................... .......................... margin ... 22 Fig. 728. Tadpole of Silverstoneia flotator (Guayach, CRARC, Costa Rica, 550 m).

20a Labial tooth rows l/0 or 210; umbeIliform oral disc ................... eia n&aola

Photos: G' b Labial tmth rowa l / l z m a l l oral disc .................................................................. 21

.................................. .......................

21a Labial tooth rows l/1

... Ietbmo4yla picadai (in part)

b Labial tooth rows Y2 h % ~ ~ a h p l a d ..................................

................................

I ............................. 22a Labial tooth rows 011

Dandrobates qJeci~w b Labial tooth rows l/1 23

23a A-1 short, restricted to area above middle of beak or lateral with large gap above

...... mouth (Fig. 729)) htdhbatesprrmili~ b A-1 extends uninterrupted to level of later-

.............. A 8.1 margins of upper jaw sheath 24

Fig. 729. Tadpole of Dendrobates pumilio 24a Tail tip rounded; beak huge, occupying ..... (Maria Luisa, Limbn, Costa Rica).

most of oral disc D a n d m h &WI

Photos: G. Kiihler b Tail tip long and attenuate; beak large but not occupying moat of oral disc .................. ........................... h t d h b a t e s g r a n m

............................. 26s Labial tooth rows 2/2 26 ............ b Labial tooth rows 2/3 or greater 29

26a Oral disc not funnel-shaped; lahial papillae large; median gap in A-2 tooth row broad, over twice length of either lateral segment; ........... jaw sheaths wide Anotheca Bpinaea

b Oral disc funnel-shaped; labial papillae tins median gap in A-2 tooth row, if present, less than length of either lateral segment; jaw sheathdl narrow to medium-sized .................................................................. 27

Fig. 730. Tadpole of Denohhates auratus (Gua- 27a Oral disc completely bordered by one row of ya&, CRARC, Costa Rica). Photos: G. Kiihler tiny papillae; oral disc without emargina-

Tadpoles

Fig. 731. Oral disc of a. Chaunus marinus; b. Incilius luetkenii; c. I. coccifer. Drawings: M. Vesely

tion on medial portion of posterior labium (Figs. 737a) ................................................... ............ Duellmanohyla r u i i 4 (in part)

b Papillae restricted to lateral and ventral portions of oral disc; oral disc with small emargination on medial portion of posteri- or labium ................................................. 28

28a Labial tooth row formula 2(2)/2(1); all tooth rows shorter than upper jaw sheath; oral disc papillae prominent and conical (Fig. 73712) ...................... Duellmanohyla soralia

b Labial tooth row formula 2/2; anterior tooth rows and P-1 row about as long as upper jaw sheath; oral disc papillae elongate ................... Duellmanohyla salvavida

............................. 29a Labial tooth rows 2/3 30 b Labial tooth rows greater than 2/3 ..... 130

30a Oral disc emarginate (Figs. 731a-c, 735b) 31 b Oral disc non-emarginate ...................... 52

31a Gaps in anterior and posterior labial papillae (Figs. 731a-c) ..................................... 32

b No gaps in posterior labial papillae ...... 43 32a Caudal musculature dark with pale sad-

dles or with cream-colored mottling or blotching on lateral surface of tail musculature; venter lacking or nearly lacking dark pigment; submarginal papillae absent ............................................................... 33

b Tail musculature solid black or brown above, with brown punctuations, or dark dorsally with narrow pale area ventrally; ventral coloration variable; submarginal papillae present or absent ..................... 34

33a Scattered dark pigment on caudal fin ....... .......................................... hcilius d e r

b Reticulate pattern on caudal fin ................ ........... hcilius vaUiceps and I. marmoreus

34a Median gap in A-2 tooth row broad, one- half to almost equal in length to either lateral segment ........................................... 35

b Median gap in A-2 tooth row, if present, narrow, several times shorter than either lateral segment ....................................... 37

.................... 35a Submarginal papillae present ............................................. hcilius porteri

b Submarginal papillae absent ................. 36 36a Ventral surfaces of body lightly pigmented

............................................. hcilius ibd b Ventral surfaces of body darkly pigmented

...................................... hcilius fastidiosus 37a P-3 tooth row noticeably shorter than P-1

and P-2 rows; one or two rows of labial ...................................... papillae present 38

b Posterior tooth rows subequal in length; usually one row of labial papillae present on

.............................................................. 35 38a Labial teeth weakly keratinized, those on

A-1 and P-3 rows noticeably less developed than those on A-2 and P-1 rows; upper jaw sheath weakly keratinized, ser- 3 rated edge with a broad, medial shallow indentation; labial papillae in one row 3 ...... ................................... Rhaebo haematitim 2

b Labial teeth well developed, all tooth rows more or less equally developed; upper jaw sheath well developed (Fig. 731a), without a broad, medial shallow indentation on ser- rated edge; labial papillae at least partial-

............... ly in two rows Chaunus marinus 39a Submarginal papillae present only on pos-

............. tenor disc area hcilius periglenes b Submarginal papillae present only on lat-

eral disc area .......................................... 40 40a Median gap in A-2 tooth row relatively

broad (about 20-30 teeth wide); posterior tooth rows slightly shorter than anterior

............ rows (Fig. 73 1b) hcilius luetkenii b Median gap in A-2 tooth row, if present,

very narrow (one to five teeth wide); all ............... tooth rows subequal in length 41

41a All labial teeth of A-1 about subequal; two to five irregular rows of submarginal papillae present laterally; oral disc ventral in position ......................... hcilius 1eucomyos

b Labial teeth at midline of A-1 smaller, more closely arranged, than teeth at ends of row; two to three submarginal papillae

b Dorsal caudal fin with brown flecks (Fig. 732b) .......................... p4VIIobatea lugthis

49a Two complete rows of labial papillae on

E osterior margin of disc; caudd .fin with ............. rown spots Allobdm )RiarnnaFnn

b One complete row of labial papillae on pas- tenor margin of disc; caudal fin without brown spots ......................................... 60

Mla Body dark-brown or blaclr; tail lightly ..................................... marked with brown

...........................a. Calcmtet&wparrurmaaaie b Dorsal caudal fin without brown blotches

.................................................................. Fig. 732. Tadpole of P6gUobates lugubh (EN. 51 h a d l o , Cwta Rim, 10 m). Photos: G. K6hler 5la Body tan, tail lightly marked wit21 brown;

no scattered very large white guanophores ...... on body and tail Calo~te#iw &i-

present laterally; oral disc anhoventral in position ............................................. 42

4% Tail fins uniformly dark brown, with tiny pigment granules; spiracle with lateral wall shorter than medial wall, aperture ............................ round li~ciliw CuWw

b Body and tail pale with some brownish pigment especially dorsally; scabtered very large white guanophores on body and tail . ...................................... Calastetbw prath'

52a Vent tube medial .................................... 53 .................................... b Vent tube dextral 78

b Tail fins transparent, with scattered, small pigment granules and a few large pigment granules; spiracle walls of nearly equal ....................... length, aperture ovoid ,..:....

~ c a k u / 3 ~ t a e ............................... 43n Oral disc anteroventral; body ovoid; one ....... row of papillae posterior to mouth 44

b Oral disc ventral, body depressed; one or two rows of papillae posterior to mouth 46

48a Dorsal fin low, not extending onto body; jaw sheaths narrow; size moderately small, total length <35 mm in Stage 36 .... ............................ ~ p ~ p ~ ~

b Dorsal fin high, extending onto body (Fig. 735a) ..................................................... 45

46a P-1 tooth row complete; body brown, tail heavily pigmented with large spots or blotches; well-developed lateral line sys-

................................ t en Lithobah iinrai b P-1 tooth row with narrow median gap;

body dark brown, tail with light brown or whitish punctations or flecks; lateral line system indistinct (Fig. 735) m b a - .............. t6,ylmi or L. brownarum (in part)

46a Spiracle ventrolateral ........................... 47 b Spiracle lateral, located near midline of

side of body ........................................ 49 47a Two complete rows of labial papillae on

posterior margin of disc ............................. ................................. n&lobate# vittaha b One complete rows of labial papillae on ........................ posterior margin of disc 48

............... 48a Caudal h without brown flecks ............................... Dandzvbatea minu*

........... 65a Gap in papillae of posterior labium ................................ l?kmdba XIlatcihgDpe b No gap in papillae of posterior labium .54

64a A-2 complete or with narrow gap ......... 66 b Median gap in A-2 broad, longer than, or

nearly as long as each lateral segment. 64 56a Labial papillae complete across anterior

labium; oral disc greatly expanded; deep folds lateral to mouth lkth~~&h p i c t p

b Gap in papillae of anterior labium; oral disc unexpanded; no folds lateral to mouth

66a Spiracle sinistral, but located near midlength of body; body not greatly depressed; eyes round; A-2 tooth row segments, if median gap present, not situated at lateral edges of jaw sheathe; overall coloration not reddish in life ...................... 57

b Spiracle sinistral, but located more posteriorly, at 5341% of body length toward vent; body greatly depressed; eyes usually ares- cent-shaped; A-2 tooth mw segments very short, situated at lateral edges of jaw sheaths; overall coloration reddish in life ............................................. (Fig. 734s) 65

Ma Spiracle venholaterd, jaw sheath without .................. serrations ~~ aahqje

b Spiracle lateral, located near midline of side af body; jaw sheaths with at least small serrations .................................... 68

B8a A broad median gap in P-2 tooth row present, about one-fourth length of each lateral section .............. Ecnrrrmrrrma4ph d a

b No gap in P-2 tooth row ........................ 69

Tadpoles

59a Labial papillae on posterior margin of disc in one row; tail musculature and fins nearly uniform in color or with a few scattered small pale spots ...................................... 60

b Labial papillae on posterior margin of disc in two rows; coloration of tail musculature and k s variable ..................................... 61

60a No submarginal papillae lateral to jaw sheaths; gap in anterior labial papillae broad, distinctly broader than width of posterior jaw sheath (Fig. 7H3a) L@&&h mnrnMmnjhRor h silvammh

b Submarginal papillae present lateral to jaw sheaths; gap in anterior labial papillae narrower, about as broad as width of posterior jaw sheath ............. fndw h&d&

61a AU labial tooth rows complete .................... ............................ Leptdkww bnlmianw

................... b Median gap in A-2 present 62 62a Body and tail musculature uniformly black

................................... Dendmbates amatus b Body and tail with small dark flecks and

spots .................................................... g8 63a No median gap in P-1 .................................

........................ Leptdactylw paeddw b A narrow median gap in P-1 present ........

................................... lkphh&vIw ibmw

Fig. 733. Oral disc of a. Leptodactylus melano- notns; b. L. &agilis. Drawings: M. Vesely

Fig. 734. Tadpole of H-obatrachim fleischmanni (Costa Rica). Photos: G. K6hler

64a No distinct pale spot posterior to oral disc; oral disc terminal in position; gap in A-2 and P-1; P-3 half length of P-1 and P-2 .. ................................. LeptaLkc&lw 8 & ~ *

b Distinct ventral pale spot present posterior to oral disc; oral disc anteroventral in position; P-3 more than half length of P-1 and P-2 (Fig. 733b) ....... lk@&c&.l~ 6.sgi2ie

K C - g&-&& ,S!@&-*Nq? * 66a Spiracle located at 7281% of body length

toward vent (at 71-79% in hatchlings) ..66 b Spiracle located at 6345% of body length

toward vent (at 51-71% in hatchlings) ... 74 66a All parts of the sinus hyobranchialis con-

spicuously voluminous, its wings not sepa-

n rated medially, dominating the ventral buccopharyngeal body part ............................... ................... Taratolvla pulvarata (in part)

b Sinus hyobranchialis narrow and its wings broadly separated medially ................... 67

67a Dorsum with strong pigmentation, dark

(BL:BW = mostly c2.0) 68 with broad Fig. 735. Tadpole of U,thobates taYjoi (near ..........................

La Fortuna, Alajuela, Costa Rica); note emar- b Dors- sparsely pigmented; body slender gination (arrow) of oral disc. Photos: G. Kiihler .............................. (BL:BW mostlp 2.0) 71 68a Interorbital distance narrow, ratio interor-

bital distance / oral disc width c0.45 (Fig. 736a) ................................. S a c h W ik

b Interorbital distance broader, ratio interor- .... bitd distance / oral disc width >0.45 69

Tadpoles

I . Teratohyia poiverata 69a Sinus hyobranchialis with well developed receptacula transversales hyobranchiales (RTH); ratio tail height to tail length c0.17; buccopharyngeal ventrum without spots

................... (Fig. 73613) Taratohy. epinosa b Sinus hyobranchialis with narrow RTH,

broad receptacula venosa disci oralis (RDO); ratio tail height to tail length >0.17; buccopharyggeal ventrum with or without spots ...................................................... 70

70a Body conspicuously flat (ratio BWBH >1.40); lateral part of the buccopharyngeal ventrum with numerous pigment spots ..... ........................... Sachatarnia albomacalata

b Body not conspicuously flat (ratio BW/BH

C ~1.40); lateral part of the buccopharyngeal ventnun without numerous pigment spots .............. Es-mMepan (in part)

7la Oral disc with more than 60 papillae in two rows; center of upper jaw narrowly keratinized and flexible ......................................

H*khchium ladcwioi (in part) ......... b Oral disc with fewer than 60 papillae in

one row; center of upper jaw broadly keratinized ..................................................... 72

7% Sinus hyobranchialis with well developed receptacula; ratio tail height / tail length ~0.17; body slender (ratio BL:BW ~ 2 . 0 ) ..... ................... Taratohpa pulverata (in part)

b Sinus hyobrancbialis narrow; ratio tail height / tail length > 0.17; body broader (ratio BL:BW e2.0) ................................. 73

73a Serrations on lower jaw sheath distinctly longer laterally than medially; sinus hyobranchialis narrow-concave; ratio BW:BH usually e1.35 .................................. .............. Emdanuu, ~ b l e p o n (in part)

b Lower jaw sheath with small, uniform serrations throughout; sinus hyobranchialis narrow plain or convex; r?tio BFBH usu: ally >1.35 @ d n o b a h c b m fl-

I 74a Oral disc with more than 60 papillae in two rows; center of upper jaw narrowly keratinized and flexible ...................................... ......... ~ o b a ~ u m ladcwioi (in part)

..................................... I b Oral disc with fewer than 60 papillae in

one row; center of upper jaw broad keratinized and rigid 76

........................................................ 76a Sinus hyobranchialis well developed (Fig.

7368 76 b Sinus hyobranchialis absent (EYg. 736e) .77

76a Sinus hyobranchialis extremely large, the two wings of the RDO broadly in contact

................. medially Cockanella granulosa b Sinus hyobranchialis large but the two

Fig. 736. Centrolenid tadpoles (dorsal view). wings of the RDO are mostly separated Photos: H. Hoffmann medially (Fig. 7368 t2ochmella euHnemos

d Cachranella euknernos

uR-: Tadpoles

I- Fig. 737. Oral disc of a. Duellmanohyla rufioculis (locality as in Fig. (locality as in Fig. 740); c. D. soralia CP.N. Cusuco, CortBs, Honduras, 700 m).

Photos: a,b. G. Kbhler; c. J. Kolby

Dorsum with few isolated spots; oral disc emarginated or notched, side-lobes of the oral disc smoothly separated from the upper lip ...... 4yaZinobafxachium chimpoi .................................. or H. co&mbip4vIIum Spots a t the dorsum fused to groups, resulting in a netlike pattern on dorsum and dorsal tail, oral disc acutely separated from the upper hp, without emargination between its lateral and caudal part and therefore forming broad lateral lobes ........... Bjdhobatraahium taikmancae or

.......................................... H. vireovittatum 78a Oral disc with continuous labial papillae

(Fig. 737a-c) ........................................... 79 b Oral disc with gap in anterior labial papil-

beyond level of beak margins 80

........................... lae (Fig. 744, 748a-c) 104 Fig. 739. Tadpoie of Duellmanony~a nrfiocuLis 79a Labial tooth rows not extending laterally (Soltis Center, Alajuela, Costa Rica, 460 m).

Photos: G. Kohler ................ b Labial tooth rows extending laterally well

beyond level of beak margins ................ 81 80a Labial tooth row A-1 shorter than A-2 .

e b. I

Fig. 740. Tadpole of Duellmanohyla uranochroa Fig. 738. Tadpole of Cruziohyla calcarifer (Gua- (Rancho Naturalista, near Platanillo, Costa yach, CRARC, Costa Rica). Photos: G. Kijhler Rica, 900 m). Photos: G. Kohler

Tadpoles

b Labial tooth rows A-1 and A-2 about same length (Fig. 737b); body more massive (Fig. 740) uranoohnaa (bt part)

.......... 8la Spiracle sinistral, but ventrolateral .................................. ch5i&yl&l tale

b Spiracle sinistral, located at or slightly ......................................... below midline 83

82a Upper jaw sheath with greatly enlarged pointed serrations to either side of midline ................................................................. 83

b Upper jaw sheath serrati01m subequal in length ..................................................... 84

Ma Upper jaw sheath with one or two greatly enlarged fang-like serrations on either side ........................... J?ktdp%matadai

b Upper jaw sheath with three or four greatly enlarged fang-like serrations on either side ................................... PIech&k id

84a Oral disc greatly expanded into a suctorial structure, as wide as or nearly as wide as body ........................................................ 86

b Oral disc smaller (c 2/3 widtb of body), not ..... expanded into a sudorial structure 90

8Sa No more than three rows of papillae ..... posterior to mouth; median gap in A-2

..................................... b t h ~ l a d&& b Four to eight rows of papillae below poste-

rior tooth rows; median gap in A-2 present or absent ................................................ 86

86a Oral disc lacking lateral folds; upper jaw sheath with a slightly paler colored poste- .......................................

I rior concavity 87

b Oral disc with lateral folds, upper jaw ... sheath without a posterior concavity 89

87a A-2 tooth row with a narrow median gap .................................. -arqrriaite

.......... b No median gap in A-2 tooth row 88 888 Lips extending laterally beyond side of

snout; single row of large, discrete papillae between fringing papillae and A-1 tooth row ........................... n-d~ ht t=~+

b Lips not extending laterally beyond side of snout; distinct smooth, sharply raised fold between fringing papillae and A-1 tooth row ........................ PIech&la te*

89a Tail fin not &ending onto body; lateral folds of disc lacking papillae ...................... ........................................... la tic8

b Tail fin extending onto body; several large papillae in lateral folds of disc .................. ................................... lkthmo4vla rivuhuh

90a Oral disc with two or three rows of labial papillae ................................................... 91

b Oral disc with a single row of labial papillae ................................................... 97

Bla P-1 and P-2 tooth rows of equal length, P-3 row slightly shorter than P-1 and P-2 rows ...................... PI-@od-

b P-1 and P-3 tooth rows of equal length, P-2 row slightly longer than P-1 and P-3

......................................................... rows 92 9% No median gap in A-2 tooth row; papillae

in submarginal rows anteriorly and posteriorly-to jaw sheaths larger than those in labial rows; size moderately large, total length r4O mm in Stage 36 .......... 93

b A-2 tooth row with a narrow median gap; papillae in submarginal rows anteriorly and posteriorly to jaw sheaths, if present, similar-sized to those in labial rows; size moderately small, total length c40 mm in Stage 36 .................................................. 96

.... 93a Jaw sheaths bearing pointed serrations ...........................a PI- ctulg9&m

b Jaw sheaths bearing blunt serraiiom .. 94 94a Nostrils about midway between eyes and

tip of snout ...... PI-la g u a W & b Nostrils about one-third distance &om eyes

to tip of snout ......... J%&V&h pokamclhi 95a Caudal 5 patternless; snout truncate ....

..................................... lahm&yla cabrpaa b Caudal fins with brown mottling; snout ................................................... rounded 96

96a Eyes directed laterally; tail musculature cream with pale brown mottling .................................. &mdmf;e oatr?zcha

b Eyes directed dorsolaterall~r; tail musculature cream with black spots .................... ................................. Isttuno4pla liumwki

97a A-2 tooth row with a narrow median gap; papillae in submarginal rows anteriorly and posteriorly to jaw sheaths, if present, similar~ized to those 'in labial rows; size

Fig. 741. Tadpoles of Plectroh jla dasypus (P.N. Cusuco, Corths, Honduras, 1370 m).

Photo: 3. Kolby

moderately small, total length <40 mm in Stage 36 ........................... Smilisca d d a

b No median gap in A-2 tooth row; papillae in submarginal rows anteriorly and posteriorly to jaw sheaths larger than those in labial rows; size moderately large, total length >40 mm in Stage 36 ................... 98

98a P-1, P-2, and P-3 tooth rows subequal in length ..................................................... 99

b P-3 tooth row slightly shorter than P-1 and P-2 tooth rows .............................. 101

99a Lateral processes of upper jaw sheath absent; iris coppery red in life .................... .................................... m-h-

b Lateral processes of upper jaw sheath short, narrow; iris color variable, usually pale brown to brown in life .................. 100

lOOa Row of discrete, large papillae between fringing papillae and outer tooth rows on anterior and posterior margins ................ ......................................... PIect&cla avia

b Row of discrete, large papillae between fringing papillae and outer tooth row only on posterior margin ................................... ............................. P I e h acanthodee

lOla No submarginal papillae lateral to jaw sheaths ..................... PItwtmhyla q u d

b At least a few submarginal papillae lateral to jaw sheaths ................................ 102

102a Lateral processes on upper jaw sheath weakly developed ... R ~ h sagmum

b Lateral processes on upper jaw sheath absent ......................... .. ................... 103

103a Numerous small submarginal papillae lateral to jaw sheaths ................................ ............................ PIei9mhy.a tecmumani

b No more than two to four large submarginal papillae lateral to jaw sheaths ......... ............................. PI-Ia g h d u l ~ ~ ~

104a Gaps in anterior and posterior labial papillae; P-3 tooth row mounted on short, vertically movable extension .................... ....................................... S c i n a r b o u l ~ .

b No gaps in posterior labial papillae; P-3 tooth row not on extension ................ 106

106a Spiracle ventrolateral ........................ 106 b Spiracle lateral ................................... 112

106a Tail fins high, dorsal &d ventral fins about subequal in height (Fig. 742a-c) 107

b Tail fins lower, dorsal fin lower than ventral fm (Fig. 742d) .............................. 109

107a P-3 tooth row with a median gap ................................... AgallPchnie saltator

b P-3 tooth row complete ..................... 108 108a Body without brown flecks or mottling

.............................. By..ameduaa van&

Tadpoles

a.

Fig. 742. Tadpoles of a. Agalyehnis annae; b. A. , spurrelli; c. A. lemur, d. A. cailidryas (a. from I Santo Domingo de Heredia, Costa Rica, 1180 m; b . 4 . from Guayach, CRARC, Costa Rica, 550 m). Photos: G. K i e r

b Body with brown flecks or mottling (Fig. 742d) ..................... &&hz&

lO9a Body a t least 15% deeper than wide; tail much deeper than body ............................ ..................................... Ag&ch& m a e

b Body only slightly deeper than wide; tail equal to or slightly deeper than body 110

llOa Dorsal tail h not reaching to body (Fig. 742b); well-developed lateral line system ................................. -epurralli

b Dorsal tail fin extending to body (Fig. 742c); lateral line system indistinct ... 111

l l l a Tail fins transparent with scattered gray flecks; dorsal tail fin only slightly lower than ventral h ..... Agdjdmk m d e t i i

b Tail &IS transparent without scattered gray flecks .................. Agdydds lemur

ll7a A narrow gap in A-1 tooth row; no gaps in posterior both rows; tail musculature and fins marked with small brown flecks, tail

..................... fin not extending onto body; oral disc anteroventral @la howurti

b No gap in A-1 tooth row, gaps in posteri- or tooth rows present or absent; coloration of tail musculature and fins variable; tail fin exten%g onto body or not; oral disc ventral or antemventral .................... 118

ll8a Two rows of labial papillae immediately ........................ I lateral to anterior gap 119

b One row of labial papillae immediately lateral to anterior gap ........................ 126

ll9a Two rows of labial papillae on posterior margin ................................................ 120

................................................. b One row of labial papillae on posterior margin 125

................................................. l20a Tail fin not extending onto body; oral disc

ventral 121 b Tail fin extending onto body; oral disc

...................................... anteroventral 123

Fig. 743. Tadpole of Tlalocohyla loquax l21a Ventral surface of body paler than dorsal ........ (Guayach, CRARC, Costa Rica, 550 m). surface h 0 p z t O P Z t ~ . miOgmpanum

Photos: G. Kijhler b Ventral and dorsal surfaces of body about ..................................... the same color 122

122a Tail heavily flecked with brown; body uni- 1128 A narrow median gap in P-2 tooth row; formly dark brown (F'ig. 745) .....................

P-3 tooth row extremely short, much ........................... lktbn&la peeudopuma shorter than P-1 or P-2 tooth rows (Fig. 743b) mmlal- b Tail creamy tan with brown spots; body .......................... mottled bmwn and tan ..............................

b No median gap in P-2 tooth row; P3 sube- ........................ mw . . ta quai in length slightly than P-l 12Sa P;( tooth ............................... or P-2 tooth rows 113 P-2 tooth rows ....................

113a Body noticeably higher than wide; dorsal tail with high arch, considerably high- b P-3 tooth row subequal in length as or er than tail musculature a t midlength of only shghtly shorter than P-1 and P-2

........................................... tooth rows 124 tail ....................................................... 114 b Body as wide as, or, slightly wider than 124a Tip of tail pointed; gap in A-2 tooth row

high; dorsal tail fin without high arch, only broad, over 70% length of either lateral ........................ slightly higher than, or same depth as, segment hdbca baadinii

tail musculature at midlength of tail 117 114a Conspicuous dark dorsolateral stripe on

..................... body extending through eye ......................................a mela pPcta

b No conspicuous dark dorsolateral stripe ................................................ on body 115

......... 116a Caudal 61ns with large brown spots ........................................ &in4x metratus

b Caudal fins with small brown fleaks and reticulations ....................................... 116

116a In life, dorsum tan, venter pale golden white, tail with minute brown or black flecks, iris pale bronze ......... Scinax altae

b In life, dorsum tan, venter white, tail with dark brown reticulations, iris pale gold Fig. 744- Oral disc of A ~ ~ Y c - annae (same ........................................ Sciaar stauifiai individual as in Fig. 742a). Photo: G. Kahler

Tadpoles

Tip of tail rounded (Fig. 746); gap in A-2 tooth row narrow; less than 20% length of either lateral segment I s t h e l a graaeee Median gaps in A-1 and A-2 tooth rows;

......................................... body tuberculate ..................... Em-ohyla fbbrimembra Median gaps only in A-2 tooth row; body not tuberculate .................... Smilieca & One row of labial papillae on posterior margin; tail tapering to a long point . 127 Two rows of labial papillae on posterior

................ margin; tip of tail rounded 128 No median gap in P-1 tooth row .............. ...................................... Scinax elaeochroa P-1 tooth row with a narrow median gap .............................................. ScinaxNhr

P-3 tooth row much shorter than P-1 and ................. P-2 tooth rows S&CB puma

P-3 tooth row subequal in length as or only slightly shorter than P-1 and P-2

........................................... tooth rows 129 Oral disc nearly terminal; P-1 tooth row slightly shorter than anterior tooth rows, others progressively shorter; serrations on jaw sheaths pointed; no pale crescent- shaped mark dorsolaterally on posterior part of body ................ wprianpeta~atue Oral disc anteroventral; posterior tooth rows subequal in length, slightly shorter than A-2 tooth row; serrations on jaw sheaths blunt; pale crescent-shaped mark usually present dorsolaterally on posteri- or part of body (Fig. 749) L & ~ ~ % c B pheoth Vent tube medial; gap in anterior labial papillae; tooth rows 3 4 4 - 6 (Fig. 747); dark brown lateral stripe or streak present anteriorly on tail musculature ......... ........................ l l - a ~ p h a l u ~ venulosue Vent tube dextral, gap in anterior labial papillae present or absent; number of tooth rows variable; dark brown lateral

Fig. 745. Tadpole of Isthmohyla pseudopuma (Albergue Montemeal, Alajuela, Costa Rica, 1820 m'l Photos: G. Kihler

Fig. 746. Tadpole of Isthmohyla graceae (Cemo Sagui, Ngobe Bugl6, Panama, 1710 m).

Photo: A. Hertz

Fig. 747. Oral disc of Trachycephalus venulosus. Drawing: M. Vesely

Fig. 748. Oral disc of a. Agalychnis call iwas, b. A. spurrelli; c. A. lemur (same individuals as in Fig. 742b-d). Photos: G. Kohler

311

stripe or streak absent on tail musculature ................................................... 131

131a Two or three anterior tooth rows ...... 132 b Four or more anterior tooth rows ...... 144

132a No gap in anterior labial papillae ..... 133 b Gap in anterior labial papillae .......... 138

13% Six or seven posterior tooth rows; oral disc greatly expanded, as wide as body ........... ........................... Ekemdanta EIlmiM

b Three to five posterior tooth rows; oral ...... disc smaller, less wide than body 134

Fig. 749. of S&m phaeota (Guaya- 134a E v e posterior tooth rows; tooth rows long, &, CRARC, Costa Rica). Photos: G. EUihler much longer than width of beak, two rows

of labial papillae ........................................ ..... FtycZ~w saZvadozmsis or P. legled

Fig. 75u. adpo pole of Bromeliohyla bromeliacia (P.N. Cusuco, Co&s, Honduras, 1616 m).

Photo: J. Kolby

Fig. 751. Tadpole of Hypsiboas dtelus. Drawing: H. Hoffinann

Fig. 752. Oral disc of Hypsibom rufitelus. Drawing: H. Hoffmann

b Three or four posterior tooth rows; tooth rows short or long; one or two rows of ................................... labial papillae 136

135a Four posterior tooth rows; tooth rows long, much longer than width of beak; disc surface mostly covered by tooth rows

.............. rows; two rows of labial papillae ............................ C m l a chmeqne

b Three posterior tooth rows; too% rows short, only slightly longer than wdth of beak, disc surface with large papillae; one .......................... row of labial papillae 136

136a Usually no median gaps in A-1 and A-3 .................................................. tooth rows ....... Duallmaaela wgnochme (in part)

b Usually median gaps present in A-1 and .................................... A 4 tooth rows 137

137a No median gap in P-1 tooth row; belly mottled; tail heavily blotched with brown .................. DuenmRnallV;la sdlmia-

b P-1 tooth row with a narrow median gap; belly uniformly dark gray; tail with dense brown flecking, givin brown appearance ......................... &olrslacbamd

138a Three posterior tooth rows; oral disc emarginated 1aterallyLithobates taylon' .......................... or L. inmmanm, (in part)

b Four posterior tooth rows; oral disc not ........................ emarginated laterally 139 139a Labial papillae in twa to three rows; body

pale brown with brown flecks; tail fins without flecking (Fig. 750) ........................ . . ......................... BmmaE~la luvmhm

b Labial papillae in a single row, at least on dorsolateral portion of oral disc; body pale tan, gray, or black without brown flecks;

..... tail fins with dark brown flecking 140 14Qa Body black ..... O&opilua s e p t e n t n ' d

b Body tan to gray ................................. 141 141a Labial papillae in two rows on ventro-

lateral and ventral portions of oral disc

Tadpoles

- -- b Labial papillae in a single row on ventro-

lateral and ventral portions of oral disc 142 142a Submarginal papillae present on lateral

disc area ........................ -b rn b Submarginal papillae absent ............. 143

143a Tail shorter, ratio taiI length l body Iength ~ 1 . 7 ........................ -baas-

b Tail longer; ratio tail length / body length >1.7 ... @ p d h mpa'tane or EL pqgnar

144a Oral disc with continuous labial papillae .............................................................. 146

b Oral disc with anterior gap in labial papillae ........................................................ 152

146a Four anterior tooth rows (A-1 can be fi-ag- mented) ............................................... 146

b Five to seven anterior tooth rows ..... 149 146a Cream-colored crescent-shaped mark on

posterior part of body, if present, not bordered posteriorly by a large brown mark .............................................................. 147

b Cream-colored crescent-shaped mark on osterior part of body bordered posterior- $ by a large brown mark .................. 148

147a P-1 tooth row with a narrow median gap ............................ P&cb&la

.............. b No median gap in P-1 tooth row ................................... - l a d m

148s Caudal musculature uniformly flecked with brown; P-1 to P 4 tooth rows about subequal in length to upper tooth rows ... ............................ mo4pla~t4m=ota

b Caudal musculature with brown squarish blotches dorsally on anterior one-half of tail, P-1 to P-4 tooth rows about usually slightly shorter than upper tooth rows .... .................... -&9-la-tlvmpan~

149a Two rows of labial papillae continuously around oral disc; five or six anterior tooth rows (A-1 and A-2 fragmented), seven to nine posterior rows (P-8 and P-9, if present, fragmented) ............................... 150

b Papillae on anterior margin reduced to one row; labial tooth rows 5-717-10 .. 151

150a Six anterior tooth rows, eight or nine posterior rows ...........

b Five anterior tooth rows< seven posterior rows ................. l 3 y c b b y l a 9 a n ~ c i e

l5la TaiI with stronger dark flecking; labial tooth rows 5-718-10 (Fig. 754) ................... ................................... -1- *m'

b Dark flecking on tail less pronounced; labial tooth rows 6-716-10 ........................ .................................. qvl- k?.&nh

162a Median gap in A-2 tooth row broad, more or less one-fourth to one-half length of

Fig. 753. TSidpole of Ptychohyla hypomykter (P.N. Cusuco, Cortks, Honduras, 1370 m).

Photo: J. Kolby

Fig. 754. Hyloscirtus palmen' (Guayadn, CRARC, Costa Rica, 550 m): a,b. Tadpole; c. metamorph. Photos: G. Ghle r

Fig. 755. Tadpole (preserved) of Hyloscirtus cofymba (Alto de Piedra, Veraguas, Panama, 910 m). Photo: G. Klihler

either lateral segment; oral disc emarginated laterally ................................. 153 Median gap in A-2 tooth row narrow; oral disc not emarginated laterally .......... 164 Five or six anterior tooth rows (Fig. 758) ................................. Lithobatas vibicarius Four (occasionally five) anterior tooth rows ......................... Lithobates vaillanti

................ Five or six posterior tooth rows ...................................... Lithobatas juliani

.................. Four posterior tooth rows 155 166a ~ e d i a n gap in P-1 tooth row narrow, less b.

than 10 teeth wide; labial teeth usually present in stages >30 ................................

Lithobates maeuIatus ................................ b Median gap in P-1 tooth row broader, A

more than 15 teeth wide; labial teeth absent (Fig. 7591 Lithobakw ~anizewitschii 1

Fig, 758. Tadpole of Lithobates vibicarius (ne- Albergue Monterreal, Alajuela, Costa Rica, 1915 m). Photo: G. Kishler

Fig. 756. Tadpole of Litfiobates maculatus (P.N. h 1 C&O, C&S, Honduras, 520 m).

Photo: J. Kolby MI- &; -

Fig. 757. Metamorph of Lithobates vibicarius (near Albergue Monterreal, Alajuela, Costa Rica, 1915 m). Photo: G. Kijhler

Further Reading STABRETT 1960, DUE- & LYNCH 1969, DUE- 1970, ~ T I G & BRANDON 1971, ACTIG & CHRISTENSEN 1976, A ~ G 1987, DUELLMAN & CAMPBELL 1992, MCDIARMID & ALTIG 1999, MENDELSON et al. 1999, DUELLMAN 2001, M C C ~ & WILSON 2002, SAVAGE 2002, rlg. .'(by. 'mapole or urnonaces warszewztsmn Horn 2005b, LYNCH 2006, HOFFMANN 2010, (Soltis Center, Alajuela, Costa Rica, 460 m). SANCHEZ 2010 Photo: G. Kiihler

. . . c H a m r y ~ ~ ~ e ~ ~ ~ ~ - - x 2 , m ~ e Amphibian D&ea

Amphibian Declines and Extinctions

Amphibians are severely declining worldwide. Hundreds of species are on the brink of extinction or have already vanished (POUNDS et al. 2006, WAKE & VREDENBURG 2008). Although declines of amphibians have been reported throughout at least the last century (e.g., DICKERSON 1906, CORN & FOGLEMAN 1984, KAGARISE SHERMAN & MORTON 1993), we are witnessing an alarming increase in regional declines and extinctions of frog and salamander species since the late 1980s (RICHARDS et al. 1993, LIPS 1998, 1999, LIPS et al. 2004); populational data for the majority of caecilian species are essentially unavailable and we, therefore, have no information on the population trends in the species in this order of amphibians (GOWER & WILKINSON 2005).

Furthermore, many of the reported declines and disappearances have been labeled mysterious or enigmatic, because they took place in protected areas with no obvious changes in the environment such as habitat destruction (CRW et al. 1992, POUNDS et al. 2006).

Land use change obviously causes the greatest anthropogenic threat to biodiversity (GROOM et al. 2006). Destruction, fragmentation, and severe modification of habitats through agricultural use, human settlements, logging, and burning are direct causes for species declines and extinctions. Fragmentation produces smaller, more isolated populations that have reduced options for dispersal and a

Fig. 760. Habitat degradation, destmction, contamination, and alteration are still the major causes for the loss of amphibian populations worldwide; Serrania de Tabasari, Ngobe Bugl6, Panama. Photo: G. Kiihler

higher risk of local extinction. The contamination of natural habitats with inorganic and organic toxins can diminish the quality of the habitat for many organisms. The introduction of exotic species can have severe negative influences on the native species through competition, predation, hybridization, or transfer of pathogens (GROOM et al. 2006). Overexploitation by humans for food, products (e.g., skin, med- icines), or pets has resulted in major declines in several species (GROOM et al. 2006). Recently, the effect of emerging infectious disease8 has been recognized as a potential major cause for the loss of biodiversity (LIPS et al. 2008). F'inally, global climatic changee, including global warming, will have severe effects on local and regional biodiversity (GROOM et al. 2006). Each of the factors mentioned above can reduce biodiversity; and, in many cases, these factors act synergistically (COLLINS & CRUMP 2009).

rnoLo: m. r ranzen

Extinction is a normal component of life; more than 99 percent of the species that ever inhabited our planet have become extinct. However, E.O. WILSON (1992) pointed out that the destruction of rain forest has increased the extinction rate to between 1000 and 10,000 times the "normal" or "background" extinction rate. Some authors suggested that the current extinction episode might eventually surpass some of the greatest mass extinctions in the history of earth, including the disappearance of the dinosaurs (COLLINS & CRW 2009). Unlike the mass extinctions of the past, the current extinction episode is caused directly and indirectly by human activities (WAKE & VREDENBURG 2008).

It is problematic to quantify extinction phenomena because it is difficult to know for certain that a species has become extinct. Also, the majority of species inhabiting the Earth has not yet been formally

Amphibian Declines

described; estimates vary between 85 to 95 percent still being undescribed (PURm & HECTOR 2000). This high percentage of undescribed species is highlighted by the results of a recent frog survey on Sri Lanka that resulted in the discovery of close to 100 undescribed species, mostly of the genus PElautus (PETH~GODA et al. 2008); a dramatic increase compared to the 50 frog species recognized before these recent explorations (DUTTA & MANAMENDRA- ARACHCHI 1996). However, many Sri Lankan species known from 19th-century museum collections were not recorded in the recent surveys of natural habitats and are probably extinct (MEEOASWMBURA et al. 2002).

For most areas of the world, baseline data on the historical and current distribution or abundance of amphibian species are not available. Possible disappearances of populations might have happened unnoticed and unreported because of the lack of prior studies. COLLINS & CRUMP (2009:45) emphasize the importance to also publish results documenting stable or increasing populations in order to not "paint a biased picture of worldwide changes."

Thanks to the efforts of the Global Amphibian Assessment, about 99 percent of the currently recognized amphibian species were evaluated; alarmingly, 1 percent of those were considered extinct and 32 percent threatened (IUCN categories Critically Endangered, Endangered, and Vulnerable); 23 percent of species were classified "Data Deficient" (STUART et al. 2008).

In the late 1980s, awareness of the amphibian decline crisis xoalesced with announcements of cases in Australia and Central America (BLAUSTEIN 1994). To gain a better understanding of what was going on, surveys of the amphibian species in these two regions were intensified (RICHARDS et al. 1993). Fieldwork conducted in the early 1990s in the Monteverde region of Costa Rica revealed that 20 frog

species (40 percent of the anuran fauna) had disappeared since 1987, including the golden toad (Incilius periglenes; Fig. 7611, Atelopus varius, eight hylids, and several species of glass frogs (VAN DEVENDER 1980, HAYES et al. 1989, POUNDS et al1997).

All Atelopus species have severely declined, and an estimated 67 percent of the known species of this bufonid genus have disappeared from their known localities (LA WCA et al. 2005, POUNDS et al. 2006).

Although no long-term field study has ever been carried out on caecilians (Go- & WILKINSON 2005), anectotal observations by M. W m in Costa Rica and Guatemala indicate that there are cases of caecilian population declines due to land use changes (COLLINS & CRW 2009): a once high-density population of Dermophis mexicanus that occurred at a coffee finca in Guatemala that was been transformed from a shaded into an open growing plantation resulted in a severe reduction in caecilian density; in the time it would have taken M. Wake and colleagues "to find more than 100 individuals 30 years ago, they found 4." (COLLINS & CRUMP 2009:20). The evaluation of the numerous cases of amphibian declines all over the world revealed several common traits:

(1) Declines seem to affect mostly high elevation species; this refers to elevations above 500 m in Central America and above 1000 m in South America (YOUNG et al. 2001, STUART et al. 2004, CRUMP 2005, LPS et al. 2008). In many Neotropical species with a broader vertical distribution it was observed that these were declining at high elevations, but populations of the same species at lower elevations were not (CRW 2005).

(2) Species that depended on water such as streams or ponds for breeding were affected most strongly by the declines and extinctions (LAURANCE et al. 1996a,b, LIPS 1998, STUART et al. 2004). Also, stream- associated species such as the rain frogs of

Amphibian Declines

the Craugastor rugulosus group were among the species that declined severely or disappeared entirely (MCCRANIE & WILSON 2002). Direct developing terrestrial species (no aquatic larvae), such as many species of rain frogs (many species in the families Craugastoridae, Eleuthero- dactylidae, and Strabomantidae) and plethodontid salamanders were not or were less affected (LIPS et al. 2003b).

(3) Most of the species that became extinct had small geographic ranges. Wide-ranging species have many populations scattered across a large area; in case one or a few of these populations become extinct, chances are that sources for immigrants exist so that the area with the lost population can be re-inhabited by the species (BROWN & KODRIC-BROWN 1977). This is not the case in species with a very restricted distribution comprised of a single (in the case of Incilius periglenes) or just a few populations (COLLINS & CRUMP 2009). Therefore, widespread species tend to persist whereas species with very small ranges tend to become extinct when they are affected by declines (LIPS et al. 2003b, COOPER et al. 2008).

In short, those s~ecies that are stream- associated, living at high altitudes, and having a small geographic range are most likely to be affected by declines. After the first announcements of massive frog declines, skepticism was high among biologists (e.g., PECHMANN & WILBUR 1994). How, they asked, is it possible to differenti- ate between normal population fluctuations and long-term trends? Frog populations are known to fluctuate widely in response to climatic factors. Over unfavor- able years, populations can decrease severely and then recover to original densities in one or two good seasons (SEMLITSCH et al. 1996, JANSEN et al. 2009). These extreme population fluctuations are especially typical for aquatic-breeding frogs ( ~ K ~ s H 2001). The difficulty of differ- entiating between normal population fluctuations and long-term downward spirals

that might lead to extinction has several implications of conservation relevance: Not reacting might lead to the loss of populations and species; over-reacting or reacting to false signs can waste money and credi- bility. BLAUSTEIN (1994:94) pointed out that "one must balance the risk of lost credibil- ity, which might seriously compromise future conservation efforts in this and other arenas, against the cost of failing to respond to a potentially serious environmental crisis."

By the mid-1990s, however, most biologists interested in amphibians shared the opinion that the numerous examples of unex- plained cases of amphibian declines and extinctions occurring simultaneously on different continents could not be attributed to normal population fluctuations (BLAUSTEIN & WAKE 1995, LIPS 1998).

After the first announcements of frog declines, speculation was widespread about the possible causes of these "new mysterious amphibian declines," including environmental contamination. climate change, and emerging infectious diseases (POWDS & CRUMP 1994, P o m s et al. 1999, 2006, 2007). The classic causes for loss of biodiversity such as land use change (habitat alterations) and overexploitation could be ruled out as possible causes because many of these new declines took place in protected areas without any obvious changes in the environment (POUNDS et al. 2006).

Some biologists have hypothesized that global warming is responsible for certain amphibian declines; affecting amphibian populations both directly and indirectly (POUNDS et al. 2006). High temperature can cause amphibians to die outright and severe droughts can be too much for the animals to endure physiologically. Indirect effects include changes in abundance of prey or predators, changes in diseases dynamics, or immunosuppression from stress resulting from environmental extremes making them more susceptible to

Amphibian Declines

pathogens (POUNDS et al. 2006, 2007). Some researchers have linked the outbreaks of chytridiomycosis with climate phenomena, proposing the "climate linked epidemic hypothesis" (POUNDS et al. 2006), which predicts amphibian declines in unusually warm years. At first glance, this hypothesis appears to be paradoxical because the causative agent, Batrachochytrium dendrobatidis (Bd), becomes more pathogenic at lower temperatures, and thrives better under moist conditions whereas warm years are compara- tively dry. POUNDS et al. (2006) suggested that an increased cloud cover might result in both nighttime warming (through the reduction of outgoing long-wave radiation) and daytime microhabitat cooling (through blocking radiant heating). These changes in microhabitat climate are considered responsible for a shift toward the optimum for Bd and resulting outbreaks of chytridiomycosis during unusually warm years (POUNDS et al. 2006). CORN (2005: 59) pointed out that "climate change may be a relatively minor cause of amphibian declines, but it may be the biggest future challenge to the persistence of many species." In this connection, ROHR & RAFFEL (2010) provided evidence for a link between climate change and Atelopus declines.

Increased levels of ultraviolet B (UV-B) have been discussed as possible causes for amphibian declines (BLAUSTEIN et al. 1998, BANCROFT et al. 2008). While this might be true for diurnal species in open areas such as Lithobates forreri and L. vibicarius, it is an unlikely cause for decline in nocturnal forest species (CAREY et al. 2004). At present, there is no published evidence suggesting that UV-B has caused amphibian extinctions, and there is conflicting evidence regarding its role in declines (COLLINS & CRUMP 2009).

Environmental contamination has also been suspected as a possible primary cause for the enigmatic amphibian population declines (CORN & VERTUCCI 1992). Studying the effects of toxins in the envi-

ronment and in organisms is difficult. Detection of chemicals requires specialized analyses and an investigator must at least have a strong suspicion about what he or she is looking for (LINDER et al. 2004). Often, the breakdown products of pesticides are more toxic to amphibians than their parent compounds (SPARLING & FELLERS 2007). Sediments can have higher chemical concentrations than the water, making sediment toxicity a potentially important factor (LINDER et al. 2004). Elevated levels of nitrogen, caused by usage of nitrate fertilizers, are known to have a negative effect on development and survivorship of amphibians (ROUSE et al. 1999). An increased frequency of abnormalities (missing or extra legs) in green frogs (Lithobates clamitans) was traced back to elevated levels of nitrogen in the natural habitat: nitrogen fertilizer that enters the aquatic ecosystem causes increased algal growth that in turn supports more snails; the snails are the intermediate host of trematode parasites (flukes; the adult parasites live in the digestive tract of birds), leading to an increased frequency of the infectious stages (cercariae) of this parasite; the cercariae infect tadpoles, embed in the region of the developing hind limbs, thereby forming cysts that can disrupt normal limb development, resulting in frogs with missing or extra hind limbs (SESSIONS 2003). In short: elevated levels of nitrogen - more snails - more parasites - more abnormalities in the frogs. Exposure of the frogs to pesticides were shown to increase the frequency of trematode infection and the resulting abnormalities; pesticide exposure was correlated with a decrease in circulat- ing leukocytes in the frogs and, therefore, a pesticide-mediated immunosuppression was hypothesized (KIESECKER 2002). However, it needs to be emphasized that parasites are only one possible cause for morphological abnormalities in amphibians; other possible causes (e.g., chemical pollutants, perhaps via chromosomal dam- age) have to be considered, too (LANNOO 2008).

Amphibian Declines =Z-T-+SSZS

I Fig. 762. Still around after dramatic population declines and local extinctions: Atelopus varius at Cerro Negro, Veraguas, Panama, 770 m).

Photo: A. Hertz

Fig. 765. Hyloscirtus colymba (Alto de Piedra, Veraguas, Panama, 910 m) with clinical signs of chytridiomycosis; note vasodilation.

Photos: A. Hertz

Pesticides (insecticides, herbicides, fungi- cides) can cause a variety of effects in amphibian populations, ranging from fem-

I inization of males due to endocrine effects

Fig. 763. Rediscovered: Isthmohyla debilis to mortality (HAYES et al. 2002). One of the (Cerro Tugri, Ngobe BuglB, Panama, 1570 rn). most commonly used pesticides, atrazine,

Photo: A. Hertz is a potent endocrine disruptor that can cause castration and feminization in male

m amphibians (HAYES et al. 2002, 2006). Pesticides can be transported over large distances in the atmosphere; they either evaporate or enter the atmosphere directly when planes apply them to fields. The toxins are then transported on prevailing winds. It could be demonstrated that the most severe amphibian population declines are in the Sierra Mountains east of the Central Valley and downwind of the intensely agricultural San Joaquin Valley of California in the United States

Fig. 764. Craugastormilesi was rediscovered (DAVIDSON & ~ A P P 2007, SPARLUVG et d. at P.N. Cusuco, CortBs, Honduras, 1839 m. 2009). Wind-borne chemical deposition

Photo: J. Kolby generally increases with elevation for

Amphibian Declines

mountains a t middle and high latitudes (DALY & WANIA 2005). Furthermore, an analysis carried out by HAGEMAN et al. (2006) revealed that current and historical agricultural activity are primarily responsible for the presence of pesticides in national parks in the western United States. Although more research is obviously needed, the available data indicate that environmental contamination with pesticides reaches levels locally that are suitable to make amphibian populations and species vulnerable to extinction (COLLINS & CRUMP 2009).

Parasite infestations and infedious diseases occur in all animals. As seen above, trematodes can be a cause for morphological abnormalities in amphibians. And pro- tozoa, bacteria, fungi, and viruses have been implicated as causes of amphibian declines (PFENNIG et al. 1991, KIESECKER et al. 2001, JOHNSON & WELLEHAN 2005). No information on infectious diseases as possible causes for declines in caecilians is available (GOWER & WILK~NSON 2005).

Just a few years after the major amphibian die-offs in Australia, it was speculated that a rapidly spreading epidemic disease might have caused the catastrophic declines (LAURANCE et al. 1996a,b). A major breakthrough in the research into the causes of the "mysterious" amphibian declines was the isolation and description of the putative causative pathogen, the chytrid fungus Batrachochytrium dendrobatidis (Bd) (BERGER et al. 1998, LONGCORE et al. 1999, LIPS et al. 2003a). Bd is a basal fungal lineage in the Chytridiomycota which are distinct from other fungi in that they have a motile, flagellate zoospore (FISHER et al. 2009).

At first, however, skepticism about the actual role of this fungus was high (BLAUSTEIN 1994): Is it a ubiquitous organism that has been present always in the amphibian environment worldwide? Is it then only a secondary cause finally killing the amphibians after "something" has had

a detrimental effect on the amphibian's immune system? Or is it a "new", emerging disease that has been introduced into naive amphibian populations? If the latter case is true, then no predisposition would be necessary: once Bd is introduced into an amphibian population, massive die-offs will follow.

There was good reason for the skepticism. There was no prior model for an infectious disease causing such catastrophic declines in any group of animals (COLLINS & CRUMP 20091, or a pathogen that would have such a low host specificy that it would infect all members of an entire class of animals (PASMAN et al. 2004). Also, it is atypical for a pathogen to not just cause a reduction of population size but drive numerous species to extinction!

The answers to the questions above differ among species and geographic locations, and, of course, among authors. However, two factors point to the emerging disease hypothesis as the primary cause of the majority of the "mysterious" amphibian declines (DASZAK et al. 2003, LIPS et al. 2006): (1) the wave-like epidemiological phenology of the amphibian declines in Australia and Lower Central America (LAURANCE et al. 1996a,b, LIPS et al. 2008); and (2) the low genetic variability observed in Bd worldwide (MOREHOUSE et al. 2003). In Costa Rica and Panama, Bd has been isolated from dead or dying anurans in the wild (LIPS et al. 2003a).

LIPS et al. (2003b) analyzed the phenology of the cases of declining amphibians from four sites in Central America and identified ecological traits, with their "aquatic index" - which estimates the degree of associations with stream habitats over a species' life span - being the most significant factor associated with declines at all four localities. Based on this analysis, they predicted the likelihood of decline for species of the then unaffected site of El Cop6 in west-central Panama. The available data also indicated a directional dis-

Amphibian Declines

persal of the pathogen (Bd), similar to that hypothesized by LAURANCE et al. (1996a,b) in Australia. The epidemic wave was documented to move across Lower Central America at a rate of 15 to 43 kilometers per years (LIPS et al. 2008). Based on this trend, LIPS and colleagues (cited in COLLINS & CRUMP 2009) predicted that Bd would invade El Valle de Ant6n in central Panama in 2006, causing the same catastrophic amphibian declines there as it did a t the sites further west previously; and they were right: in the spring of 2006, Bd arrived at El Valle de Ant611 resulting in frogs dying from chytriomycosis (GAGLIARDO et al. 2008).

An investigation of museum holdings revealed that Bd has been present on frogs collected in Puerto Rico at least since 1976 (BURROWES et al. 20041, in Costa Rica at least since the late 1970s or early 1980s (PUSCHENDORF 2003, PUSCHENDORF et al. 2006, LIPS et al. 20081, and on historical specimens of Xenopus laevis from Africa collected in the 1930s (WELDON et al. 2004). Because Bd was prevalent as a stable endemic infection in southern Africa for 23 years before any positive specimen was found outside Africa, it was proposed that Africa is the origin of the amphibian chytrid fungus (WELDON et al. 2004). The available molecular data are all consistent with a single origin of Bd with subsequent regional and international spread (FISHER et al. 2009).

The regions with the highest suitability for Bd include habitats that contain the world's most diverse amphibian faunas (RON 2005). In Australia, Bd has infected 46 frog species of which 13 have declined and 3 are considered to be extinct (COLLINS & STORFER 2003). The wide range of conditions under which Bd has been found and its high detection rate in the Neotropics suggests that Bd may already occupy a significant portion of the major natural regions in Central and South America (RON 2005).

Why do some amphibian species appear to be more susceptible than others to chytridiomycosis, and why are the most severe declines found in the highlands whereas lowland sites are generally unaffected? The answer seems to lie in the physiologi- cal and ecological preferences of the fungus: Bd thrives best under typical cloud forest conditions: high humidity and temperature between 15" and 25" C (COLLINS & CRUMP 2009). Bd can remain viable in sterilized water for up to seven weeks (JOHNSON & SPEARE 2003).

Although evidence has been presented that Bd is the primary cause of amphibian declines and die-offs in many parts of the world (LIPS et al. 2006, POUNDS et al. 20061, many unresolved issues and numerous questions remain, and some authors argue that the case for Bd as the primary agent has not yet been made conclusively (e.g., MCCALLUM 2005, POUNDS et al. 2006). However, other investigators such as CYNTHIA CAREY and KAREN LIPS (among others), are convinced that no other factors need to be invoked and "once the chytrid infects a formerly chytrid-free population of toads, the toads die whether it's a warm year, cold year, dry year, or wet year." (CAREY cited in COLLINS & CRUMP 2009: 141). RYAN et al. (2008) conducted a mark- recapture study of three subpopulations of the stream frog Craugastorpunctariolus at a mid-elevation site in central Panama between 1999 and 2005. Prior to the invasion of Bd, the subpopulations of C. punctariolus were large and showed a stable age-size distribution. Following detection of Bd at this site in late September 2004, all three subpopulations completely disappeared from this site within two months. CRAWFORD et al. (2010) presented long- term field surveys and DNA barcode data of an amphibian community in central Panama; they documented a mass-mortality event caused by Bd resulting in the loss of 41% of total amphibian lineage diversity, including five undescribed species.

Amphibian Declines

Bd therefore belongs to a select group of virulent multihost pathogens that have had profound effects on entire communities and ecosystems. The mechanisms by which chytridiomycosis affects amphibians are poorly understood. Typically, amphibians with chytridiomycosis show progres- sive deterioration in condition eventually leading to death ( B o s c ~ et al. 2001, NICHOLS et al. 2001). Clinical signs of this disease include vasodilation (Fig. 76513) and hyperkeratosis (PESSIER et al. 1999). A disturbed res~iration or water balance due to hyperkeratosis and poisoning by a fungal toxin have been discussed as the main factors that induce mortality in amphibians with clinical chytridiomycosis (PESSIER et al. 1999, CARVER et al. 2010). More research is needed to determine how Bd causes clinical chytridiomycosis and immunological mechanisms by which hosts respond to Bd.

Reports on rediscovered frog species from both Central America (Figs. 762-764) and Australia have been published (BARRIO- ~ ~ 0 ~ 6 s 2004, PUSCHENDORF et a1 2005, LAMP0 et a1 2006, RODR~GUEZ-CONTRERAS et al. 2008, COLLINS & CRUMP 2009, KOLBY & MCCRANIE 2009, MCCRANIE et al. 2010). It is not always clear whether these represent survivors of the disease outbreak (remnant populations), unaffected disjunct populations in remote places, or new colo- nizations (LOTTERS et al. 2005). However, Bd is still present in these areas (RETALLICK et al. 2004, LOTTERS et al. 2005, KOLBY & MCCRANIE 2009).

If Bd is still resent in the environment. how can we explain the reappearances of several frog species that have not been seen for several (up to 25) years? The pep- tides in amphibian skin secretions are known to have antimicrobial properties (CLARKE 2007). Intraspecific variation in skin secretions can be high in frogs, and, in the presence of a pathogen, a strong pres- sure of natural selection will favor those individuals that have peptide secretions that are more effective against the

pathogen. The rediscovered individuals might be immune to Bd or at least able to tolerate its presence (COLLINS & CRUMP 2009). Or has Bd evolved into less deadly strains? Another sr~eculation would be that naturally occurring bacteria or fungi on the amphibian's skin have changed in composition or nature and are outcompet- ing Bd (COLLINS & C R ~ 2009). Whatever is the case in the rediscovered frogs we do not know. The remnant populations of some Australian frogs that suffered severe declines in the past, now persist in the natural habitat with stable Bd-infections (RETALLICK et al. 2004).

During the past 20 years, we have wit- nessed intensive research into the causes of amphibian declines worldwide. Having standadzed methods and protocols for monitoring amphibian populations is essential to ensure that data are comparable among researchers and across sites. A major step toward this goal has been the publication of the "Standard Methods for Amphibians" (HEYER et al. 19941, a com- pendium that recommends and defines techniques for monitoring amphibian populations including suggestions for field design and statistical analyses.

Despite substantial achievements and resolving some of the initial questions, much controversy remains and many additional questions have been raised. We do not yet understand all the details of this phenomenon, but a t least we now have explanations and hypotheses for the "mysterious die-offs" of amphibians. In addition, we have learned from the mod- ern amphibian extinctions that protected areas are no guarantee for the long-term survival of species. On the positive side are reports of rediscoveries of "lost frog species." Do these rediscoveries indicate that the species involved are rebounding from the brink of extinction or that they are still in a period of decline (KOLBY & MCCRANIE 2009)? Will evolutionary processes such as mutations and natural selection work toward the evolution of

Amphibian Declines

amphibian species that are resistant to Bd? Who will win the evolutionary arms race: the amphibians or Batrachochytrium dendrobatidis? Only time will tell.

However, given the complex and multifac- torial causes of the amphibian decline phenomenon, it would not be wise to narrow our focus to exclusively study the effects of Bd. Obviously, Bd plays a key role in many cases of amphibian population declines and extinctions, but this should not dis- tract us from research into other possible causes. As was demonstrated by DASZAK et al. (20051, amphibian population declines a t the Savannah River Site, South Carolina, United States, were more likely caused by an increase in the number of years with insufficient rainfall than by chytrid epidemics.

However, habitat degradation, destruction, contamination, and alteration are still the major causes for the loss of amphibian populations worldwide. As BLAUSTEIN (1994) concluded, the amphibian decline problem is part of the overall crisis in biodiversity. The world is steadily losing an unprece- dented number of species in many different groups of animals and plants (WILSON 1992). Ongoing deforestation, drainage and pollution of wetlands and other anthropogenic effects causes the alteration and destruction of suitable habitat for many organisms in all regions of the world, including amphibians. For 90 percent of the amphibian species classified as threatened, habitat loss and degradation is a major threat (HOFFMAN et al. 2008). BECKER et al. (2007) have emphasized habitat split - the human-induced discon- nection between habitats used by different life history stages of a species - as yet another important threat for amphibians. Amphibians, including their larvae, play an important role in energy flow and nutrient recycling in all major terrestrial and freshwater ecosystems worldwide. Therefore, we have to address questions such as the ecological consequences of the disappearance of a substantial portion of

an entire class of animals for the structure and functioning of ecosystems. While we are studying the possible causes of amphibian decline, frogs and salamanders are continuing to disappear before our eyes. What can be done to stop and even reverse these processes?

The classic strategies including the decla- ration and management of nature reserves are still important components of any long- term conservation program. If a species depends on forest, then this species will disappear if the forest is cut down. Amphibian populations that have survived Bd infections will further suffer as environments deteriorate (POUNDS et al. 2006). However, because climatic changes, toxic exposures, and emerging diseases will not be controlled by reserve boundaries, it is obvious that current laws and policies alone are insufficient to address global threats. Several possible solutions and conservation tactics have been proposed in order to respond to the global decline of amphibians (HOFFMAN et al. 2008). It must be emphasized that properly managed biotic reserves still play a key role in long- term conservation strategies. The empha- sis here lies on "properly managed" because reserves that only exist on the map are obviously useless. Currently, many protected areas in Central America lack adequate personnel and operational funding resulting in ongoing deforestation, unregulated hunting, use of pesticides, and anthropogenic development (ACEVEDO et al. 2010, SUNYER & KOHLER 2010, WILSON & TOWNSEND 2010).

For some species ex-situ programs might be the only option to avoid extinction (e.g., Atelopus zeteki and Nectophrynoides asperginis; ZIPPEL 2002, KRAJICK 2006; however, see criticism by POUNDS et al. 2006). AmphibianArk, an alliance of conservation organizations, has the goal to maintain in captivity breeding groups of species that otherwise would become extinct. The ultimate goal of these ex-situ programs should the restoration of the

Amphibian Declines

species in the wild. These captive programs should be established in the countries where the threatened species naturally occur. Given the very restricted infra- structure and funding resources in most of the tropical regions, support from countries in the temperate regions is urgently needed. This refers not only to funding but also to capacity building. Training in hus- bandry and breeding techniques, as well as hygiene and disease control are essential for these kinds of programs. Critics (e.g., POUNDS et al. 2006) pointed out that the number of species that could be taken care of in such ex-situ programs is very limited, and that reintroductions would be uncertain for most species that have become extinct in the wild, especially for those whose habitat has been destroyed. At least we are "buying time" with these ex-situ programs, hoping this will allow research to progress and finally open more opportunities for survival of these species in their natural habitat.

Environmental education is an extremely powerful conservation tool to raise awareness about the global amphibian crisis and teach local communities about ecological issues such as the effects of pesticides on amphibian populations (including morphological abnormalities and cases of feminization). It is essential to encourage public appreciation for streams, ponds, lakes, and marshes, and the creatures that inhabit them.

Finally, conservation action is much more effective when supported by a proper taxonomic basis (HOFFMAN et al. 2008). The large percentage of species in the IUCN category "Data deficient" indicates that much research into the taxonomy, geographic variation, population genetics, population dynamics, and natural history of numerous amphibian species is urgent-

ly needed (STUART et al. 2010). Long-term monitoring of amphibian populations has only been done at very few places (DASZAK et al. 2005). Finally, investigations on the population dynamics and the Bd infection rates of surviving populations of amphibian species - those that had suffered severe losses or had not been found in decades and also those that were seeming- ly not affected - will yield valuable information on the interactions between the pathogen and the amphibians. It seems that Bd will persist in the habitats of the caecilians, salamanders, and frogs of Central America, and hopefully the amphibians will find ways to survive despite its continued presence. But they will need all the help they can get in order to succeed - at least, they need decent habitat conditions.

WILSON & TOWNSEND (2010) summarized various aspects of the conservation of the Mesoamerican herpetofauna and offered six broad-based recommendations for the challenging conservation work in this region. Obviously, these recommendations are difficult to implement, but as these authors point out, the only realistic approach is "to assess any given situation or problem and design solutions" (WILSON & TOWNSEND 2010: 777).

Further Reading BLAUSTEIN 1994, HEYER et al. 1994, BLAUSTEIN & WAKE 1995, LIPS 1998,1999, SPARLING et al. 2000, YOUNG et al. 2001, SEMLITSCH 2003, LINDER et al. 2004, LIPS et al. 2003b, LIPS et al. 2004, STUART et al. 2004, LANNOO 2005, LANNOO 2008, STUART et al. 2008, WAKE & VREDENBURG 2008, COLLINS & CRW 2009, STUART et al. 2010, WILSON et al. 2010

Amphibian Declines

Amphibians as Bioindicators

Some biologists suggested that amphibians might be useful as bioindicators because of their permeable skin and their biphasic life style, leading to a kind of double jeopardy (BLAUSTEIN & W m 1995, KOHLER 2001). Disappearing frog populations might be interpreted as early warning signals of environmental deterioration. Although this concept was a great market- ing tool and helped fund raising, it was eventually demonstrated that this hypothesis was false (COLLINS & CRUMP 2009). To be a good bioindicator, a group of species should meet certain criteria, including the following ones (HILTY & MERENLENDER 2000, COLLINS & CRUMP 2009):

(1) It should be readily accessible any time when sampling might be done. However, most amphibians species are notoriously seasonal, and detectability is reduced over much of the annual cycle.

(2) Monitoring of the group should be straightforward and cost-effective. However, this is not the case for most amphibian species. Estimating an amphibian species' population size is technically difficult and laborious.

(3) The taxonomy and natural history of the species in the group should be fairly well known. However, for the majority of amphibian species life history data are scarce andfor anecdotal, the taxonomy of most major groups still is poorly understood, and many species are known from only a few individuals. For a large proportion of amphibians, the reproductive biology and activity periods are unknown. We lack basic data on habitat requirements for numerous species that aggregate in aquatic habitats for breeding but spend the rest of the year abroad in terrestrial environments.

(4) Normal population fluctuations should be relatively small so as not to mask declines resulting from environmental problems on the one hand or to trigger false alarms on the other. However, frog populations are known to fluctuate by orders of magnitude in different years, and the population dynamics of most species are unknown (COLLINS & CRUMP 2009). Many amphibian populations have "boom and bust" population cycles in which numbers may vary by a factor of some 20 times between successive years (SKELLY et al. 1999).

(5) Changes in the group's abundance should occur quickly to the same environmental changes that might also influence the remaining biota and the ecosystem as a whole. However, no evidence for this kind of relationship has been published. BOONE & BRIDGES (2003) concluded that - depending on species studied - amphibians can be either more or less sensitive to toxins than other organisms.

Consequently, there is no evidence to suggest that amphibians are better bioindicators than other animals (BEEBEE & GRIFFITH 2005, COLLINS & CRUMP 2009). Any decline or disappearance of an amphibian population is an indication that something is or was affecting that specific population negatively; but this "something" does not necessarily have an influence on the remaining biota (PECHMANN & WILBUR 1994).

Further Reading PECHMANN & WILBUR 1994, HILTY & MERENLENDER 2000, BEEBEE & GRIFFITH 2005, COLLINS & CRUMP 2009

Acknowledgments

Acknowledgments

During the past 20+ years of my work on Central American amphibians and reptiles, I received support and help from many individuals and institutions in Central America, the USA, and Europe; without them, this book project never would have been possible.

I am particularly thankful to Lany David Wilson who patiently read several versions of the various parts of the book manuscript and provided invaluable assistance, advice, and encouragement during the process of writing this book. Also, he contributed the foreword to this book - his help is much appreciated!

I appreciate the support I have received from forestry and wildlife officials in Central America. Collecting and exporta- tion permits were provided by Z. Alain, Edgar Arauz, Yariela Hidalgo, C. Medina, Abdiel Mendoza, Adalberto Montezuma M., Ricardo Rivera, and Aleida Salazar, Autoridad Nacional de Ambiente (ANAM), Panama City, Panama; Tito Perez, Congreso General Kuna, Panama City, Panama; Javier Guevara Sequeira, Rafael Gutierrez Rojas, and Marta Liliana Jimenez Fernandez, Ministerio de Recursos Naturales, Energia y Minas (MINAE), San Jose, Costa Rica; Milton G. Camacho, Mauricio Fonseca Cuevas, Carlos Peres-Roman, Carlos Rivas Leclair, and Sandra Tijerino Mejia, Ministerio del Ambiente y 10s Recursos Naturales (MARENA), Managua, Nicaragua; Angel Barahona, Karla Cantero, Ruben Castro Lobo, Tomas Garcia, Ana Patricia Martinez, Antonio Ortez Turcios, and Carlos Romero, Corporation Hondureiia de Desarollo Forestal (COHDEFOR), Tegucigalpa, Honduras; Patricia Quintana Vda. De Lopez, and Andres Sanchez, Ministerio de Agricultura y Ganaderia, El Salvador; J.I. Galvez Duiiionez, Oscar F.

Lara. L., and Alba Nydia Perez, Consejo Nacional de Areas Protegidas (CONAP), Guatemala City, Guatemala; F. Ramirez Ruiz de Velasco and L. Lozano, Secretaria de Medio Ambiente, Recursos Naturales y Pesca, Mexico D.F., Mexico.

For logistic support for fieldwork, I thank Robert Dilger, Hauke Hoops, Siegfried Kastl, Mausi and Eddi Kiihl, Juan Carlos Martinez Sanchez, Ivan Ortega, Roger Romtin, and Francisco Serrano. For field assistance, I thank Juan Abarca, Manuel E. Acevedo, Abel Batista, Martin Dolling, Jorge Ferrari, Andreas Hertz, Martin Jansen, Elke Kohler, Johannes Kohler, Jorg Kreutz, Brian Kubicki, Sebastian Lotzkat, James Randy McCranie, Fernando Mendoza, Kirsten Nicholson, Michael Obermeier, Marcos Ponce, Ardiel Z. Quintana, Antonio Ramirez Vasquez, Fabian Schmidt, Robert Seipp, Jorge Soto, Pablo Soto, Javier Sunyer, Joseph Vargas, and Jose Manuel Zolotoff. I am especially thankful to these people for their valuable help. Many students and associates of the department of herpetology a t Senckenberg Museum, Frankfurt, Germany, have tested the identification keys included in this book. Their help is greatly appreciated. And I appreciate the support I have received from my assistant a t Sencken- berg, Linda Acker.

I am grateful to numerous colleagues for photos of Central American reptile species-these photographers are acknowl- edged in the captions of their respective photos. All of the excellent pen and ink drawings of anuran hands and feet are the painstaking efforts of Milan Vesely, and Arne Schulze contributed illustrations of tadpoles. Carl Franklin, Andreas Hertz, Johannes Kohler, and Mason Ryan provided unpublished data on maximum sizes of various amphibian species. Brian Kubicki,

Acknowledgments

Heinz Hoffman, and Joseph Vargas were instrumental in obtaining living tadpoles for photographic purposes. Brian Kubicki pointed out to me the sexual dichromatism present in Lithobates warszewitschii.

For the loan of specimens or access to the collections under their care, I thank Linda Ford and Darrel R. Frost, American Museum of Natural History (AMNH), New York City; Ted Daeschler, Academy of Natural Sciences of Philadelphia (ANSP), Philadelphia; Colin J . McCarthy, The Natural History Museum (BMNH), London; Jens Vindum, California Academy of Sciences (CAS), San Francisco; Alan Resetar and Harold Voris, Field Museum of Natural History (FMNH), Chicago; John E. Simmons, Eli Greenbaum, Linda Trueb, and William E. Duellman, University of Kansas Natural History Museum (KU), Lawrence; Jeff A. Seigel, Natural History Museum of Los Angeles County (LACMI, Los Angeles; Douglas A. Rossman and Frank T. Burbrink, Museum of Natural Science, Louisiana State University (LSUMZ), Baton Rouge; Jose Rosado and James Hanken, Museum of Comparative Zoology (MCZ), Harvard; Alain Dubois, Ivan Ineich and Annemarie Ohler, Museum National d7Histoire Naturelle (MNHN), Paris; Eunice E. Echeverria, Museo de Historia Natural de El Salvador (MUHNES), San Salvador; David B. Wake and Carla Cicero, Museum of Vertebrate Zoology (MVZ), Berkeley; Franz Tiedemann, Naturhistorisches Museum Wien (NMW), Wien; Janalee P. Caldwell, Amy D. Estep, and Laurie J. Vitt, Sam Noble Oklahoma Museum of Natural History (OMNH), Norman; James R. Dixon and Katherine Vaughan, Texas Cooperative Wildlife Collection (TCWC), College Station; Federico Bolafios and Gerardo Chaves Cordero, Museo de Zoologia Universidad de Costa Rica (UCR), San Jos6; F. Wayne King and Kenneth Krysko, Florida Museum of Natural History (UF), Gainesville; Christopher A. Phillips, John Petzing, and Steven D. Sroka, Museum of Natural History,

University of Illinois a t Urbana (UIMNH), Urbana; Ronald Nussbaum, Arnold Kluge, and Greg Schneider, University of Michigan Museum of Zoology (UMMZ), Michigan; Ronald Crombie, Steve Gotte, Ronald Heyer and Robert Wilson, Smithsonian Institution, National Museum of Natural History (USNM), Washington DC; Jonathan Campbell, Carl Franklin, and Eric Smith, University of Texas a t Arlington (UTA), Arlington; Victor J . Hellebuyck (VH), San Salvador; Wolfgang Bohme, Zoologisches Forschungsinstitut und Museum Alexander Koenig (ZFMK), Bonn; and Rainer Giinther, Museum f i r Naturkunde der Humboldt-Universitat zu Berlin (ZMB), Berlin; Frank Glaw and Dieter Fuchs, Zoologische Staatssammlung Miinchen (ZSM), Munich.

I owe an immense debt of gratitude to Andreas Hertz, Brian Kubicki, Sebastian Lotzkat, Joseph R. Mendelson 111, Mason Ryan, David B. Wake, Marvalee Wake, and Larry David Wilson, who patiently supported this project and read various portions of the manuscript; all individuals provided valuable comments and advice that significantly improved the text.

Special thanks are due to my father, Alfred A. Schmidt, for his constant support and encouragement.

Finally, I am especially grateful to my beloved wife Elke Kohler and daughter Isabelle for their emotional support, patience, and inspiring encouragement. Elke and Isabelle accompanied me in the field and provided valuable assistance. Elke also merits special thanks for her very careful editing of the manuscript. My indebtedness to her cannot be exaggerated.

Literature cited

Literature cited

ACEVEDO, M., WILSON, L.D., CANO, E.B. & V~QUEZ-ALMAZAN, C. (2010): Diversity and conservation status of the Guatemalan herpetofauna. - Pp. 406-434. In WLLSON, L.D., TOWNSEND, J.H. & JOHNSON, J.D. (eds.): Conservation of Mesoamerican Amphibians and Reptiles. - Eagle Mountain Publishing, LC, Eagle Mountain, Utah. i-xviii + 1-812 PP.

ACOSTA-GALVIS, A.R. (1999): Distribution, varia- cion y estatus taxonomico de las poblaciones del complejo Rana palmipes (Amphibia: Anura: Ranidae) en Colombia. - Revista de la Academia Colombiana de Ciencias Exactas Fisicas y Naturales 23: 215-224.

ALBERCH, P. & ALBERCH, J. (1981): Heterochronic mechanisms of morphological diversification and evolutionary change in the Neotropical salamander, Bolitoglossa occidentalis (Amphibia: Plethodontidae). - Journal of Morphology 167 (2): 249-264.

ALTIG, R. (1987): Key to the anuran tadpoles of Mexico. - Southwestern Naturalist 32 (1): 75-84.

ALTIG, R. & BRANDON, A. 1971. Generic key and synopses for larvae and tadpoles of Mexican amphibians. - Tulane Studies in Zoology and Botany 17: 10-15.

ALTIG, R. &. T. CHRISTENSEN (1976): A bibliogra- phy of tadpole biology. - Smithsonian Herpetological Information Service 31: 1-3.

ALTIG, R. & JOHNSTON, G.F. (1989): Guilds of anuran larvae: relationships among developmental modes, morphologies, and habitats. - Herpetological Monographs 3: 81-109.

ANDERSON, M.T. & WTHIS, A. (1999): Diets of two sympatric Neotropical salamanders, Bolitoglossa mexicana and B. rufescens, with notes on reproduction for B. rufescens. - Journal of Herpetology 33 (4): 601-607.

AROSEMANA, F.A. & IBAREZ, D.R. (1991): Geographic distribution: Hyla angustilineata (NCN). - Herpetologica! Review 22 (4): 133.

ATTUM, 0. & EASON, P. (1999): Bufo campbelli (Campbell's forest toad). - Herpetological Review 30 (2): 106.

BACKWELL, P.R.Y. & JENNIONS, M.D. (1993): Mate choice in the Neotropical frog, Hyla ebracca- ta: sexual selection, mate recognition and signal selection. - Animal Behaviour 45 (6): 1248-1250.

BANCROFT, B.A., BAKER, N.J. & BLAUSTEIN, A.R. (2008): A meta-analysis of the effects of ultraviolet B radiation and other stressors on survival in amphibians. - Conservation Biology 22: 987-996.

BARBOUR, T. (1906): Vertebrata from the savanna of Panama: reptilia and amphibia. - Bulletin of the Museum of Comparative Zoology 46: 224-229.

BARBOUR, T. (1914): A contribution to the zoogeography of the West Indies, with special reference to amphibians and reptiles. - Memoirs of the Museum of Comparative Zoology. Cambridge, Massachusetts 44: 205-359.

BARBOUR, T. (1923): Notes on reptiles and amphibians from Panama. - Occasional Papers of the Museum of Zoology, University of Michigan 129: 1-16.

BARBOUR, T. (1926): New Amphibia. - Occasional Papers of the Boston Society of Natural History 5: 191-194.

BARBOUR, T. (1928): New Central American frogs. - Proceedings of the New England Zoological Club. Cambridge, Massachusetts 10: 25-31.

BARBOUR, T. & DUNN, E.R. (1921): Herpetological novelties. - Proceedings of the Biological Society of Washington 34: 157-162.

BARBOUR, T. & LOVERIDGE, A. (1929): Vertebrates from the Corn Oslands. Reptiles and Amphibians. - Bulletin of Museum of Comparative Zoology, Harvard University 69: 138-146.

BARRERA-RODRIGUEZ, M. (2000): Estudio anatomic0 de cuatro especies de ranitas de cristal del genero Hyalinobatrachium RUIZ & LYNCH 1991 grupo fleischmanni (Amphibia: Centrolenidae). - Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 23: 245-260.

BARRERA-RODRIGUEZ, M. & RUIZ-CARRANZA, P.M. (1989): Una nueva especie del genero Centrolenella NOBLE 1920 (Amphibia: Anura: Centrolenidae) de la Cordillera Occidental de Colombia. - Trianea 3: 77-84.

BARRIO AMOR~S, C.L. (2001): Geographic distribution. Bufo haematiticus. - Herpetological Review 32 (3): 189.

BARRIO-AMOROS, C.L. (2004): Atelopus mucubajiensis still survives in the Andes of Venezuela. - Froglog 66: 2-3.

Literature cited

BATISTA, A. & K~~HLER, G. (2008): Variation in Oophaga pumilio (Amphibia: Anura: Dendrobatidae) in western Panama. -

Salamandra 44 (4): 225-234 BAUER, A.M., GOOD, D.A. & G ~ T H E R , R. (1993):

An annotated type catalogue of the caecilians and salamanders (Amphibia: Gymno- phiona and Caudata) in the Zoological Museum, Berlin. - Mitteilungen aus dem Zoologischen Museum in Berlin 69: 285-306.

BAUER, L. (1994): New names in the family Dendrobatidae (Anura, Amphibia). - Ripa, Netherlands, Fall: 1-6.

BAYLOR, E.R. & STUART, L.C. (1961): A new race of Bufo valliceps from Guatemala. -

Proceedings of the Biological Society of Washington 74: 195-202.

BECKER, C.G., FONSECA, C.R., BAPTISTA HADDAD, C.F., BATISTA, R.F. & INACIO PRADO, P. (2007): Habitat split and the global decline of amphibians. - Science 318: 1775-1777.

BEEBEE, T.J.C. & GRIFFITH, R.A. (2005): The amphibian decline crisis: A watershed for conservation biology? - Biological Conservation 125: 271-285.

BERGER, L., SPEARE, R., DASZAK, P., GREEN, D.E., CUNNINGHAM A.A., GOGGIN, C.L., SLOCOMBE, R., RAGAN, M.A., HYATT, A.D., MCDONALD, K.R., HINES, H.B., LIPS, K.R., MARANTELLI G. & PARKES, H. (1998): Chytridiomycosis causes amphibian mortality associated with population declines in the rain forests of Australia and Central America. - Proceedinzs of the National Academv of Sciences oUf the United States of America 95: 9031-9036.

BILLE, T. (1991): Central American rainforest salamanders, Bolitoglossa mexicana-group. - Newsletter of the Australian Society of Herpetologists 34 (7): 138-141.

BILLE, T. (1998): Bolitoglossa hartwegi, en lun- gelos salamander fra Mexico og Guatemala, med en notits om B. veracrucis-gruppen. - Nordisk Herpetologisk Forening 41 (4): 87-91.

BILLE, T. (2000): Bolitoglossa meliana (NCN): Antipredator posture. - Herpetological Review 31 (2): 97-98.

BILLE, T. & BRINGSOE, H. (1998): Zu Brutpflege und Wachstum bei Bolitoglossa mexicana (Caudata: Plethodontidae): On parental care and growth in Bolitoglossa mexicana (Caudata: Plethodontidae). - Salamandra 34 (3): 219-222.

BILLO, R. & WAKE, M.H. (1987): Tentacle development in Dermophis mexicanus (Amphibia, Gymnophiona) with an hypothe-

sis of tentacle origin. - Journal of Morphology 192 (2): 101-111.

BLAIR, K.B., KILLEBREW, F.C., SMITH, H.M. & CHISZAR, D. (1995): Bufo valliceps (Gulf Coast toad). - Herpetological Review 26 (3): 152.

BLAIR, W.F. (1972): Evolution in the genus Bufo. - University of Texas Press. 459 pp.

BLAUSTEIN, A.R.'(1994): Chicken Little or Nero's fiddle? A perspective on declining amphibian populations. - Herpetologica 50: 85-97.

BLAUSTEIN, A.R., KIESECKER, J.M., CHIVERS, D.P., HOKIT, D.G., MARCO, A., BELDEN, L.K. & HATCH, A. (1998): Effects of ultraviolet radiation on amphibians: field experiments. - American Zoologist 38: 799-812.

BLAUSTEIN, A.R. &WAKE, D.B. (1995): The puzzle of declining amphibian populations. -

Scientific American 272: 56-61. BOETTGER, 0 . (1892): Katalog der Batrachier-

Sammlung im Museum der Sencken- bergischen Naturforschenden Gesellschaft im Frankfurt and Main. - Museum der Senckenbergischen Naturforschenden Gesellschaft. Frankfurt: Knauer. 73 pp.

BOETTGER, 0. (1893): Ein neuer Laubfrosch aus Costa Rica. - Berichte der Sencken- bergischen Naturforschenden Gesellschaft in Frankfurt am Main 1893: 251-252.

BOGART, J.P. & NELSON, C.E. (1976): Evolutionary implications from karyotypic analysis of frogs of the families Microhylidae and Rhinophrynidae. - Herpetologica 32: 199-208.

BOLANOS, F., ROBINSON, D.C. & WAKE, D.B. (1987): A new species of salamander (genus Bolitoglossa) from Costa Rica. - Revista de Biologia Tropical 35: 87-92.

BOLAROS, F. & WAKE, D.B. (2009): Two new species of montane web-footed salamanders (Plethodontidae: Bolitoglossa) from Costa Rica-Panama border region. - Zootaxa 1981: 57-68.

BOONE, M. & BRIDGES, C. (2003): Effects of pesticides on amphibian populations. - Pp. 152-167 in: SEMLITSCH, R.D. (ed.) (2003): Amphibian conservation. - Smithsonian Books. Washington, D.C.

B o s c ~ , J., MARTINEZ-SOLANO, I. & GARCIA-PARIS, M. (2001): Evidence of a chytrid fungus infection involved in the decline of the common midwife toad (Alytes obstetricans) in protected areas of central Spain. - Biological Conservation 97: 331-337.

BOULENGER, G.A. (1882a): Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum. 2nd edition. Taylor & Francis, London.

Literature cited

BOULENGER, G.A. (188213): 21. Hylodes Brocchi. - Pp. 60-61 in BROCCHI, M.P. (1881-1883): Mission Scientifique au Mexique et dans l ' h e r i q u e Centrale, ouvrage publie par ordre du Ministre de L'Instruction Publique. Recherches Zoologiques. Troisibme Partie. 2. Section. Etude des Batraciens de YAmerique Centrale. 122 pp., Paris. Imprimerie Nationale.

BOULENGER, G.A. (1883a): Descriptions of new lizards and frogs collected by Herr A. Forrer in Mexico. - Annals and Magazine of Natural History, Series 5, 11: 342-344.

BOULENGER, G.A. (188313): Description of new species of reptiles and batrachians in the British Museum. - Annals and Magazine of Natural History, Series 5, 12: 161-167.

BOULENGER, G.A. (1891): Notes on American batrachians. - Annals and Magazine of Natural History ser. 5, 11 (65): 342-344.

BOULENGER, G.A. (1896a): Descriptions of new reptiles and batrachians from Colombia. - Annals and Magazine of Natural History ser. 6, 17 (97): 16-21.

BOULENGER, G.A. (1896b): Descriptions of new batrachians collected by Mr. C. F. Under- wood in Costa Rica. - Annals and Magazine of Natural History. Ser. 6,18 (106): 340-342.

BOULENGER, G.A. (1898a): Fourth report on additions to the batrachian collection in the Natural-History Museum. - Proceedings of the Zoological Society of London 1898 (3): 473-482.

BOULENGER, G.A. (1898b): An account of the reptiles and batrachians collected by Mr. W. F. H. Rosenberg in western Ecuador. - Proceedings of the Zoological Society of London 1898: 107-126.

BOULENGER, G.A. (1898~): A list of the reptiles and batrachians collected by the late Prof. L. Balzan in Bolivia. Annali del Museo Civico di Storia Naturale Genova ser. 19: 128-133.

BOULENGER, G.A. (1899): Descriptions of new batrachians in the collection of the British Museum (Natural History). - Annals and Magazine of Natural History. Ser. 7, 3: 273-277.

BOULENGER, G.A. (1902): Descriptions of new batrachians and reptiles from north-western Ecuador. - Annals and Magazine of Natural History ser. 7, 9: 51-57.

BOULENGER, G.A. (1908): Descriptions of new batrachians and reptiles discovered by Mr. M. G. Palmer in south-western Colombia. - Annals and Magazine of Natural History. ser. 8, I1 (12): 515-522.

America, presewed in the British Museum. - Annals and Magazine of Natural History ser. 8, 10 (56): 185-191.

BOULENGER, G. A. (1913): A collection of batrachians and reptiles made by Dr. H.G.F. SPURRELL, F.Z.S., in the Choco, Colombia. - Proceedings of the Zoological Society of London 1913 (4): 1019-1038.

B m , A.H. (1963): A new Costa Rican salamander (genus Oedipina) with a re-examination of 0. collaris and 0 . serpens. - Natural History Museum of Los Angeles County Contributions in Science 65: 1-12.

B m , A.H. (1968): Systematics and evolution of the Mesoamerican salamander genus Oedipina. - Journal of Herpetology 2: 1-64.

B m , A.H. & DUELLMAN, W.E. (1970): A new salamander (genus Oedipina) of the uniformis group from western Panama. - Natural History Museum of Los Angeles County Contributions in Science 201: 1-8.

B m , A.H., Jr., SAVAGE, J.M., WAKE, D.B. & HANKEN, J. (2001): New species of large black salamander, genus Bolitoglossa (Plethodontidae) from western Panama. - Copeia 2001 (3): 700-704.

BRAME, A.H. & WAKE, D.B. (1963a): Redescription of the plethodontid salamander Bolitoglossa lignicolor (Peters), with remarks on the status of B. palustris TAYLOR. - Proceedings of the Biological Society of Washington 76: 289-296.

B m , A.H. & WAKE, D.B. (1963b): The salamanders of South America. - Natural History Museum of Los Angeles County Contributions in Science 69: 1-72.

B m , A.H. & WAKE, D.B. (1972): New species of salamanders (genus Bolitoglossa) from Colombia, Ecuador and Panama. - Natural History Museum of Los Angeles County Contributions in Science 219: 1-34.

BRANDON-JONES, D., DUCKWORTH, J.W., JENKINS, P.D., RYLANDS, A.B. & SARMIENTO, E.E. (2007): The genitive of species-group scientific names formed from personal names. - Zootaxa 1541: 41-48.

BREDER, C.M., jr. (1946): Amphibians and reptiles of the Rio Chucunaque drainage, Darien, Panama, with notes on their life histories and habits. - Bulletin of the American Museum of Natural History 86: 375-436.

B~occm, M.P. (1877a): Description d'un nouveau genre de phaneroglosse hylaeforme (Plectrohyla guatemalensis). - Bulletin de la Soci6t6 Philomathique de Paris, ser. 7, 1: 92-93.

BOULENGER, G.A. (1912): Descriptions of new batrachians from from the Andes of South

BRoccm, M.P. (1877b): Note sur quelques batraciens hylaeformes recueillis au Mexique et

Literature cited

au Guatemala. - Bulletin de la SocietC Philomathique de Paris, ser. 7, 1: 122-132.

BROCCHI, M.P. (1877~): Sur quelques batraciens raniformes et bufoniformes de l'Am6rique Centrale. - Bulletin de la Soci6t6 Philomathique de Paris, ser. 7, 1: 175-197.

BROCCHI, M.P. (1879): Sur divers Batraciens anoures de l'Am6rique Centrale. - Bulletin de la Soci6t6 Philomathique de Paris Ser. 7, 3: 19-24.

BROCCHI, M.P. (1881-1883): Mission Scientifique au Mexique et dans YAmerique Centrale, ouvrage publie par ordre du Ministre de L'Instruction Publique. Recherches Zoolo- giques. Troisieme Partie. 2. Section. Etude des Batraciens de 1'Amerique Centrale. 122 pp., Paris. Imprimerie Nationale.

BROCK, J. & VERRELL, P. (1994) : Courtship behavior of the seal salamander, Desmo- gnathus monticola (Amphibia: Caudata: Plethodontidae). - Journal of Herpetology 28: 411-415.

BRODIE, E.D. & CAMPBELL, J.A. (1993): A new salamander of the genus Oedipina (Cau- data: Plethodontidae) from the Pacific versant of Guatemala. - Herpetologica 49: 259-265.

BRODIE, E.D., Jr., DUCEY, P.K. & BANESS, E.A. (1991): Antipredator skin secretions of some tropical salamanders (Bolitoglossa) are toxic to snake predators. - Biotropica 23 (1): 58-62.

BROWN, J.H. & KODRIC-BROWN, A. (1977): Turnover rates in insular biogeography: Effect of immigration on extinction. - Ecology 58: 445449.

BRUCE, R.C. (1997): Life history attributes of the salamander Bolitoglossa colonnea. - Journal of Herpetology 31: 592-594.

BRUCE, R.C. (1998): Nesting habits, eggs, and hatchlings of the salamander Nototriton picadoi (Plethodontidae: Bolitoglossini). -

Herpetologica 54: 13-18. BRUCE, R.C. (1999): Life history attributes of a

rare Neotropical salamander, Nototriton picadoi (Plethodontidae: Bolitoglossini). -

Herpetological Review 30: 76-78, BRUCE, R.C. (2000): Sexual size dimorphism in

the Plethodontidae. - In: R. C. BRUCE, R.G. JAEGER & L.D. HOUCK (eds.): The Biology of Plethodontid Salamanders. Kluwer Academic/Plenum Publishers, New York, New York, 243-260.

BRUCE, R.C. (2003): Life Histories. Pp. 477-525 in SEVER, D.M. (ed.): Reproductive Biology and Phylogeny of Urodela (Amphibia): Science Publishers, Inc., Enfield, New Hampshire.

BRUST, D. (1993): Maternal brood care by Dendrobates pumilio - a frog that feeds its young. -Journal of Herpetology 27: 96-98.

BUMZAHEM, C.B. (1955): A new species of frog of the genus Eleutherodactylus from Guate- mala. - Copeia 1955: 118-119.

BUMZAHEM, C.B. & SMITH, H.M. (1954): Additional records and descriptions of Mexican frdgs of the genus Plectrohyla. - Herpetologica 10: 61-66.

BUMZAHEM, C.B. & SMITH, H.M. (1955): Additional notes and descriptions of plethodontid salamanders from Mexico. - Herpetologica 11: 73-75.

BURROWES, P.A., JOGLAR, R.L. & GREEN, D.E. (2004): Potential causes for amphibian declines in Puerto Rico. - Herpetologica 60 (2): 141-154.

CALDWELL, J.P. (1994): Natural history and survival of eggs and early larval stages of Agalychnis calcarifer (Anura: Hylidae). Herpetological Natural History 2 (2): 57-66.

CAMERANO, L. (1879): Di alcune specie di anfibii anuri esistenti nelle collezioni del R. Museo Zooligico di Torino. - Atti della Reale Accademia delle Scienze di Torino. Classe di Scienze Fisiche, Matematiche et Naturali 14: 866-897.

CAMPBELL, J.A. (1998): Amphibians and reptiles of northern Guatemala, the Yucatan, and Belize. - University of Oklahoma Press, Norman, Oklahoma, U.S.A., 380 pp.

CAMPBELL, J.A. (1994a): A new species of Eleutherodactylus (Anura: Leptodactylidae) of the biporcatus group from eastern Guatemala. - Herpetologica 50: 296-302.

CAMPBELL, J.A. (1994b): New species of Eleutherodactylus (Anura: Leptodactylidae) of the milesi group from Guatemala. - Herpetologica 50 (4): 398411.

CAMPBELL, J.A. (1998): Amphibians and Reptiles of Northern Guatamala, the Yucatan, and Belize. - University of Oklahoma Press, Norman, Oklahoma.

CAMPBELL, J.A. (2000): The herpetofauna of the mesic upland forests of the Sierra de las Minas and Montafias del Mico, Guatemala. - Pp. 80-92 in: JOHNSON, J.D., WEBB, R.G. & FLORES-VILLELA, O.A. (eds): Mesoamerican Herpetology: Systematics, Zoogeography and Conservation. The University of Texas a t El Paso. El Paso, Texas.

CAMPBELL, J.A., BLANCAS-HERNANDEZ, J.C. & SMITH, E.N. (2009): A new species of stream- breeding treefrog of the genus Charadrahyla (Hylidae) from the Sierra Madre del Sur of Guerrero, Mexico. - Copeia 2009 (2): 287- 295.

Literature cited

CAMPBELL, J.A. & BRODIE, E.D., Jr. (1992): A new species of treefrog (Hylidae) from the Sierra de 10s Cuchumatanes of Guatemala. - Journal of Herpetology 26 (2): 187-190.

CAMPBELL, J.A. & LAWSON, D.P. (1992): Hyla bocourti (Mocquard, 18991, a valid species of frog (Anura: Hylidae) from Guatemala. - Proceedings of the Biological Society of Washington 105: 393-399.

CAMPBELL, J.A. & SAVAGE, J.M. (2000): Taxonomic reconsideration of Middle American frogs of the Eleutherodactylus rugulosus group (Anura: Leptodactylidae): A reconnaissance of subtle nuances among frogs. - Herpetological Monographs 14: 186-292.

CAMPBELL, J.A., SAVAGE, J.M. & MEYER, J.R. (1994): A new species of Eleutherodactylus (Anura: Leptodactylidae) of the rugulosus group from Guatemala and Belize. - Herpetologica 50: 412-419.

CAMPBELL, J.A. & SMITH, E.N. (1992): A new frog of the genus Ptychohyla (Hylidae) from the Sierra de Santa Cruz, Guatemala, and description of a new genus of Middle American stream-breeding treefrogs. - Herpetologica 48 (2): 153-167.

CAMPBELL, J.A. & SMITH, E.N. (1998): New species of Nototriton (Caudata: Pletho- dontidae) from eastern Guatemala. - Scien- tific Papers of the Natural History Museum of the University of Guatemala 6: 1-8.

CAMPBELL, J.A., SMITH, E.N. & ACEVEDO, M.E. (2000): A new species of fringe-limbed frog from the northern Cuchumatanes of Guatemala. - Herpetologica 56: 250-256.

CAMPBELL, J.A., SMITH, E.N., STREICHER, J., ACEVEDO, M:E. & BRODIE, E.D. (2010): New salamanders (Caudata: Plethodontidae) from Guatemala, with miscellaneous notes on known species. - Miscellaneous Publications, Museum of Zoology, University of Michigan 200: 1-60.

CAMPBELL, J.A. & VANNINI, J.P. (1989): The distribution of amphibians and reptiles in Guatemala and Belize. - The Western Foundation of Vertebrate Zoology 4: 1-21.

CANNATELLA, D.C. & DUELLMAN, W.E. (1984): Leptodactylid frogs of the Physalaernus pustulosus group. - Copeia 1984: 902-921.

CAPOCACCIA, L. (1957): Catalogo dei tipi di anfibi del Museo Civico di Storia Naturale di Genova. - Annali del Museo Civico di Storia Naturale di Genova. Serie 3, 69: 208-222.

CAREY, C., BRADFORD, D.F., BRUNNER, J.L., COLLINS, J.P., DAVIDSON, E.W., LONGCORE, J.E., PESSIER, A.P. & SCHOCK, D.M. (2004): Biotic factors in amphibian population declines. - Pp. 153-208 in LINDER, G.L.,

KREST, S.K. & SPARLING, D.W. (eds.): Amphibian decline: An integrated analysis of multiple stressor effects. - SETAC Press, Pensacola, Florida.

CARVER, S.,. BELL, B.D. & WALDMAN, B. (2010): Does chytridiomycosis disrupt amphibian skin function? - Copeia 2010 (3): 487-495.

CHAN, L.M. (2003): Seasonality, microhabitat and cryptic variation in tropical salamander reproductive cycles. - Biological Journal of the Linnean Society 78 (4): 489496.

CHAFARRO, J.C., P ~ u K , J.B. & GLUESENKAMP, A.G. (2007): A new species of arboreal Rhinella (Anura: Bufonidae) from cloud forest of southeastern Peru. - Herpetologica 63: 203-212.

CHAVES, G., GARC~.~-RODR~GUEZ, A., MOM, A. & LEAL, A. (2009): A new species of dink frog (Anura: Eleutherodactylidae: Diasporus) from Cordillera de Talamanca, Costa Rica. - Zootaxa 2088: 1-14.

CISNEROS-HEREDIA, D.F. (2006): Amphibia, Caudata, Plethodontidae: Bolitoglossa equa- toriana and Bolitoglossa biseriata: range extensions, new provincial records from Ecuador, and natural history. - Check List 2 (3): 64-67.

CISNEROS-HEREDIA, D.F. & MCDIARMID, R.W. (2007): Revision of the characters of Centrolenidae (Amphibia: Anura: Athesphatanura), with comments on its taxonomy and the description of new taxa of glassfrogs. - Zootaxa 1572: 1-82.

CLARKE, B.T. (2007): The natural history of amphibian skin secretions, their normal functioning and potential medical applications. - Biological Reviews 72 (3): 365-379.

CLOUGH, M. & SUMMERS, K. (2000): Phylogenetic and biogeography of the poison frogs: evidence from mitochondria1 DNA sequences. - Biological Journal of the Limean Society 70: 515-540.

COCHRAN, D.M. & GOIN, C.J. (1970): Frogs of Colombia. - United States National Museum Bulletin 288: 1-655.

COCROFT, R.B., MCDIARMID, R.W., JASLOW, A.P. & Run-CARRANZA, P.M. (1990): Vocalizations of eight species of Atelopus (Anura: Bufonidae) with comments on communication in the genus. - Copeia 1990 (3): 631-643.

COLLINS, J.P. & GRUMP, M.L. (2009): Extinction in our times. Global amphibian declines. - Oxford University Press, New York, New York. 273 pp.

COLLINS, J.P. & STORFER, A. (2003): Global amphibian declines: Sorting the hypotheses. -Diversity and Distributions 9: 89-98.

Literature cited

COLLINS-RAINBOTH, A. & BUTH, D.G. (1990): A reevaluation of the systematic relationships among species of the genus Dendrotriton (Caudata: Plethodontidae). - Copeia 1990 (4): 955-960.

COLOMA, L.A. (1995): Ecuadorian frogs of the genus Colostethus (Anura: Dendrobatidae). - Miscellaneous Publication. Museum of Natural History, University of Kansas 87: 1-72.

COOPER, N., BIELBY, J., G. THOMAS, G. & PURVIS, A. (2008): Macroecology and extinction risk correlates of frogs. - Global Ecology and Biogeography 17: 2 11-22 1

COPE, E.D. (1862a): Catalogue of the reptiles obtained during the explorations of the Parana, Paraguay, Vermejo, and Uruguay Rivers, by Capt. Thos. J. Page, U.S.N., and of those procured by Lieut. N. Michler, U.S. Top. Eng., Commander of the expedition con- ducting the survey of the Atrato River. - Proceedings of the Academy of Natural Sciences of Philadelphia 14: 346-359.

COPE, E.D. (186213): On some new and little known American Anura. - Proceedings of the Academy of Natural Sciences of Phila- delphia 14: 151-158.

COPE, E.D. (1863): On Trachycephalus, Scaphiopus and other American Batrachia. - Proceedings of the Academy of Natural Sciences of Philadelphia 15: 43-54.

COPE, E.D. (1864): Contribution to the herpetology of tropical America. - Proceedings of the Academy of Natural Sciences of Philadelphia 16: 166-181.

COPE, E.D. (1865): Third contribution to the herpetology of tropical America. - Proceedings of the Academy of Natural Sciences of Philadelphia 17: 185-198.

COPE, E.D. (1866): Fourth contribution to the herpetology of tropical America. - Proceedings of the Academy of Natural Sciences of Philadelphia 18: 123-132.

COPE, E.D. (1867 "1866"): Fifth contribution to the herpetology of tropical America. - Proceedings of the Academy of Natural Sciences of Philadelphia 18: 317-323.

COPE, E.D. (1868): An examination of the Reptilia and Batrachia obtained by the Orton Expedition to Equador and the Upper Amazon, with notes on other species. - Proceedings of the Academy of Natural Sciences of Philadelphia 20: 96-140.

COPE, E.D. (1869a "1868"): Sixth contribution to the herpetology of tropical America. - Proceedings of the Academy of Natural Sciences of Philadelphia 20: 305-313.

COPE, E.D. (1869b): A review of the species of

the Plethodontidae and Desmognathidae. - Proceedings of the Academy of Natural Sciences of Philadelphia 21: 93-118.

COPE, E.D. (1874): Description of some species of reptiles obtained by Dr. John F. Bransford, Assistant Surgeon United States Navy, while attached to the Nicaraguan surveying expedition in 1873. - Proceedings of the Academy a f Natural Sciences of Philadelphia 26: 64-72.

COPE, E.D. (1876 "1875"): On the Batrachia and Reptilia of Costa Rica. - Journal of the Academy of Natural Sciences of Philadelphia (2) 8 (4): 93-154.

COPE, E.D. (1877): Tenth contribution to the herpetology of Tropical America. - Proceedings of the American Philosophical Society 17: 85-98.

COPE, E.D. (1879): Eleventh contribution to the herpetology of tropical America. -

Proceedings of the American Philosophical Society 18: 261-277.

COPE, E.D. (1885 "1884"): Twelfth contribution to the herpetology of tropical America. - Proceedings of the American Philosophical Society 22: 167-194.

COPE, E.D. (1886): Thirteenth contribution to the herpetology of Tropical America. - Proceedings of the American Philosophical Society 23: 271-287.

COPE, E.D. (1887): Catalogue of batrachians and reptiles of Central America and Mexico. - Bulletin of the United States National Museum 32: 1-98.

COPE, E.D. (1893): Second addition to the knowledge of the Batrachia and Reptilia of Costa Rica. - Proceedings of the American Philosophical Society 31: 333-347.

COPE, E.D. (1894): Third addition to a knowledge of the Batrachia and Reptilia of Costa Rica. - Proceedings of the Academy of Natural Sciences of Philadelphia 46: 194-206.

CORN, P.S. (2005): Climate change and amphibians. - Animal Biodiversity and Conservation 28: 59-67.

CORN, P.S. & FOGLEMAN, J.C. (1984): Extinction of montane populations of the northern leopard frog (Rana pipiens) in Colorado. - Journal of Herpetology 18: 147-152.

CORN, P.S. & VERTUCCI, F.A. (1992): Descriptive risk assessment of the effects of acidic deposition on Rocky Mountain amphibians. - Journal of Herpetology 26: 361-369.

COTT, H.B. (1926). Observations on the life- habits of some batrachians and reptiles from the lower Amazon: And a note on some

Literature cited

mammals from Marajo Island. - Proceedings of the Zoological Society of London 1926: 1159-1178

CRAWFORD, A.J. (2003): Huge populations and old species of Costa Rican and Panamanian dirt frogs inferred from mitochondrial and nuclear gene sequences. - Molecular Ecology 12 (10): 2525-2540.

CRA~TORD, A.J., LIPS, K.R. & BERMINGHAM, E. (2010): Epidemic disease decimates amphibian abundance, species diversity, and evolutionary history in the highlands of Panama. - Proceedings of the National Academy of Sciences of the United States of America"l07 (31): 13777-13782.

CRAWFORD, A.J., RYAN, M.J. & JARAMILLO, C.A. (2010): A new species of Pristimantis (Anura: Strabomantidae) from the Pacific coast of the Darien province, Panama, with a molecular analysis of its phylogenetic position. - Herpetologica 66 (2): 192-206.

CRAWFORD, A.J. & SMITH, E.N. (2005): Cenozoic biogeography and evolution in direct-developing frogs of Central America (Lepto- dactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. - Molecular Phylogenetics and Evolution 35: 536-555.

CRW, M.L. (1984): Ontogenetic changes in vul- nerability to predation in tadpoles of Hyla pseudopuma. - Herpetologica 40 (3): 265-271.

CRUAP, M.L. (1986): Homing and site fidelity in a Neotropical frog, Atelopus varius (Bufonidae). - Copeia 1986 (4): 1007-1009.

C R ~ P , M.L. (1988): Aggression in Harlequin Frogs: male-male competition and a possible conflict of interest between the sexes. - Animal Behaviour 36 (4): 1064-1077.

CRUMP, M.L. i1989a): Life history consequences of feeding versus non-feeding in a faculta- tively non-feeding toad larva. - Oecologia 78: 486-489.

CRW, M.L. (1989b): Effect of habitat drying on developmental time and size a t metamorphosis-in Hyla pseudopuma. - Copeia 1989 (3): 794-797.

CRW, M.L. (1990): Possible enhancement of growth in tadpoles through cannibalism. - Copeia 1990 (2): 560-564.

CRW, M.L. (1991): Choice ofloviposition site and egg load assessment by a treefrog. - Herpetologica 47 (3): 308-315.

CRUIIIP, M.L. (2005): Why are some species in decline but others not? - Pp. 7-9 in LANNoo, M. (ed.): Amphibian declines: The conservation status of United States species. - University of California Press. Berkely, California.

C R ~ , M.L., HENSLEY, F.R. & CLARK, K.L. (1992): Apparent decline of the Golden Toad: Underground or extinct? - Copeia 1992 (2): 413420.

CRUMP, M.L. & POUNDS, J.A. (1989): Temporal variation in the dispersion of a tropical anuran. - Copeia 1989 (1): 209-211.

CRUMP, M.L. & TOWNSEND, D.S. (1990): Random mating by size in a neotropical treefrog, Hyla pseudopuma. - Herpetologica 46 (4): 383-386.

CRUZ Dm, G.A. & WILSON, L.D. (1983): Bufo haematiticus Cope: an addition to the anuran fauna of Honduras. - Herpetological Review 14 (1): 31.

CRUZ-DIAZ, G.A. & MCCRANIE: J.R. (1999): Centrolene prosoblepon. - Herpetological Review 30 (1): 49.

DALY, G.L. & W m , F. (2005): Organic contami- nants in the mountains. - Environmental Science and Technology 39: 385-398.

DALY, J. W. (1995): The chemistry of poisons in amphibian skin. - Proceedings of the National Academy of Sciences of the United States of America 92: 9-13.

DALY, J.W. & MYERS, C.W. (1967): Toxicity of panamanian poison frogs (Dendrobates): some biological and chemical aspects. - Science 156 (3777): 970-973.

DALY, J. W., MYERS, C.W., WARNICK, J.E., & ALBUQUERQUE, E.X. (1980): Levels of batrachotoxin and lack of sensitivity to its action in poison-dart frogs (Phyllobates). - Science 208: 1383-1385.

DALY, J.W., GUSOVSKY, F., MYERS, C.W., YOTSU- YAMASHITA, M. & YASUMOTO, T. (1994): First occurrence of tetrodotoxin in a dendrobatid frog (Colostethus inguinalis), with further reports for the bufonid genus Atelopus. - Toxicon 32 (3): 279-285.

DALY, J.W., PADGETT, W.L., SAUNDERS, R.L. & COVER, J.F., Jr. (1997): Absence of tetrodotox- ins in a captive-raised riparian frog, Atelopus varius. - Toxicon, 35 (5): 705-709.

DALY, J.W., SECUNDA, S.I., GARRAFFO, H.M., SPANDE, T.F., WISNIESKI, A., NISHIHARA, C. & COVER, J.F. (1992): Variability in alkaloid profiles in neotropical poison dart frogs (Dendrobatidae): genetic versus environmental determinants. - Toxicon 30: 887-898.

DASZAK, P., SCOTT, D.E., KILPATRICK, A.M., FAGGIONI, C., GIBBONS, J.W. & PORTER, D. (2005): Amphibian population declines a t the Savannah River Site are linked to climate, not chytridiomycosis. - Ecology 86 (12) 3232-3237.

Literature cited

DASZAK, P., CUNNINGHAM, A.A. & HYATT, A.D. (2003): Infectious disease and amphibian population declines. - Diversity and Distributions 9: 141-150.

DAVDSON, C. & KNAPP, R.A. (2007): Multiple stressors and amphibian declines: Dual impacts of pesticides and fish on yellow- legged frogs. - Ecological Applications 14: 1892-1902.

DAYTON, G.H., FITZGERALD, L.A. & GOLDSTEIN, M.I. (2001): Bufo valliceps (Gulf Coast toad). - Herpetological Review 32 (2): 112.

DE CARVALHO, A.L. (1954): A preliminary synopsis of the genera of American microhylid frogs. - Occasional Papers of the Museum of Zoology, University of Michigan 555: 1-19.

DICKERSON, M.C. (1906): The frog book. - Doubleday, Page and Company. New York.

DODD, C.K., JR. & BRODIE, E.D., 111 (1976): Defensive mechanisms of Neotropical salamanders with an experimental analysis of immobility and the effect of temperature on immobility. - Herpetologica 32 (3): 269-290.

DONNELLY, M.A. (1989): Reproductive phenology and age structure of Dendrobates pumilio in northeastern Costa Rica. - Journal of Herpetology 23: 362-367.

DONNELLY, M.A. (1991): Feeding patterns of the strawberry poison frog Dendroba tes pumilio (Anura: Dendrobatidae). - Copeia 23: 723-730.

DONNELLY, M.A. (1999): Reproductive phenology of E1eutherodactylus bransfordii in northeastern Costa Rica. - Journal of Herpetology 33 (4): 624-631.

DONNELLY, M.A. & GUYER, C. (1994): Patterns of reproduction and habitat use in an assemblage of Neotropical hylid frogs. - Oecologia 98: 291-302.

DONNELLY, M.A., GUYER, C., KREMPELS, D.M. & BRAKER, H.E. (1987): The tadpole of Agalychnis calcarifer (Anura: Hylidae). - Copeia 1987 (1): 247-250.

DONNELLY, M.A., GUYER, C. & DE SA, R.0. (1990a): The tadpole of a dart poison frog, Phyllobates lugubris (Anura: Dendro- batidae). - Proceedings of the Biological Society of Washington, 103: 427431.

DONNELLY, M.A., DE SA, R.0. & GUYER, C. (1990b): Description of the tadpoles of Gastrophryne pictiventris and Nelsono- phryne aterrima (Anura: Microhylidae), with a review of morphological variation in free-swimming microhylid larvae. - American Museum Novitates 2976: 1-19.

D'ORGEM, C.A. & TURNER, B.J. (1995): Multiple paternity in the red-eyed treefrog Agalychnis callidryas (Cope). - Molecular Ecology 4: 505-508.

DREWRY, G.E. & JONES, K. (1976): A new ovovi- vaparous frog, EIeutherodactylus jasperi (Amphibia, Anura, Leptodactylidae) from Puerto Rico. - Journal of Herpetology 10: 161-165.

DUBOIS, A. & HEYER, W.R. (1992): Leptodactylus labialis, the valid name for the American white-lipped frog (Amphibia: Leptodactylidae). - Copeia 1992 (2): 548-585.

DUCEY, P.K., BRODIE, E., JR. & BANESS, E.A. (1993a): Salamander tail autotomy and snake predation: role of antipredator behavior and toxicity for three Neotropical Bolitoglossa (Caudata: Plethodontidae). - Biotropica 25 (3): 344-349.

DUCEY, P.K., FORMANOWICZ, JR, D.R., BOYET, L., ~MAILLOUX, J. & NUSSBAUM, R.A. (1993b): Experimental examination of burrowing behavior in caecilians (Amphibia: Gymnophiona): effects of soil compaction on burrowing ability of four species. - Herpetologica 49: 450-457.

DUELLMAN, W.E. (1956): The frogs of the hylid genus Phrynohyas FITZINGER, 1843. - Miscellaneous Publications. Museum of Zoology, University of Michigan 96: 1 4 7

DIJELLMAN, W.E. (1960): A distributional study of the amphibians of the Isthmus of Tehuantepec, Mexico. - University of Kansas Publications Museum of Natural History 13: 19-72.

DUELLMAN, W.E. (1961a): The amphibians and reptiles of Michoacan, Mexico. - University of Kansas Publications Museum Natural History 15: 1-148.

DUELLMAN, W.E. (1961b): Descriptions of two species of frogs, genus Ptychohyla. - University of Kansas Publications Museum Natural History 13 (8): 349-357.

DUELLMAN, W.E. (1961~): Description of a new species of tree frog from MBxico. Studies of American hylid frogs VI. - Herpetologica 17: 1-5.

DUELLMAN, W.E. (1963a): Amphibians and reptiles of the rainforest of southern El PetBn, Guatemala. - University of Kansas Publications Museum Natural History 15: 205-249.

DIJELLMAN, W.E. (1963b): A new species of tree frog, genus Phyllomedusa, from Costa Rica. - Revista de Biologia Tropical 11: 1-23.

DIJELLMAN, W.E. (1963~): A review of the Middle American tree frogs of the genus Ptychohyla. - University of Kansas Publications Museum Natural History 15: 297-349.

DUELLMAN, W.E. (1965): Frogs of the Hyla taeniopus group. - Copeia 1965: 159-168.

Literature cited

DUELLMAN, W.E. (1966a): The Central American herpetofauna: an ecological perspective. - Copeia 1966 (4): 700-719.

DUELLMAN, W.E. (1966b): A new species of fringe-limbed tree frog, genus Hyla, from Darien, Panama. - University of Kansas Publications. Museum of Natural History 17: 257-262.

DUELLMAN, W.E. (1968a): Smilisca phaeota. - Catalogue of American Amphibians and Reptiles 61: 1-2.

DUELLMAN, W.E. (1968b): Descriptions of new hylid frogs from Mexico and Central America. - University of Kansas Publications, Museum of Natural History 17 (13): 559-578.

DUELLMAN, W.E. (1970): The hylid frogs of Middle America. - University of Kansas Museum of Natural History. Lawrence, Kansas.

DUELLMAN, W.E. (1971a): A taxonomic review of the south american hylid frogs, genus Phrynohyas. - Occasional Papers of the Museum of Natural History of the University of Kansas 4: 1-21.

DUELLMAN, W.E. (1971b): The nomenclatural status of the names Hyla boans (Linnaeus) and Hyla maxima (Laurenti) (Anura: Hylidae). - Herpetologica 27: 397-405.

DUELLMAN, W.E. (1971~): The burrowing toad, Rhinophrynus dorsalis, on the Caribbean lowlands of Central America. -Herpetologica 27: 55-56

DUELLMAN, W.E. (1977): Liste der rezenten Amphibien und Reptilien: Hylidae, Centrolenidae, Pseudidae. - Das Tierreich 95: 1-225.

DUELLMAN, W.E. (1980): A new species of marsupial frog (Hylidae: Gastrotheca) from Venezuela. - Occasional Papers of the Museum of Zoology University of Michigan 690: 1-7.

DUELLMAN, W.E. (1983): A new species of marsupial frog (Hylidae: Gastrotheca) from Colombia and Ecuador. - Copeia 1983 (4): 868-874.

DUELLMAN, W.E. (1986): Two new species of Ololygon (Anura: Hylidae) from the Venezuelan Guayana. - Gopeia 1986 (4): 864-870.

DUELLMAN, W.E. (1989): Lista anotada y clave de 10s sapos marsupiales (Anura: Hylidae: Gastrotheca) de Colombia. - Caldasia 16 (76): 105-111.

DUELLMAN, W.E. (ed.): (1999) Patterns of Distribution of Amphibians: a Global Perspective. - The Johns Hopkins University Press. Baltimore. viii + 633.

DUELLMAN, W.E. (2001). The Hylid Frogs of Middle America. 2nd ed. - Society for the Study of Amphibians and Reptiles. Ithaca, New York, USA.

DUELLMAN, W.E. & CROMBIE, R.I. (1970): Hyla septentrionalis D&RIL and BIBRON Cuban treefrog. - Catalogue of American Amphibians and Reptiles 92: 1 4 .

DUELLMAN, W.E. & CAMPBELL, J.A. (1982): A new frog of the genus Ptychohyla (Hylidae) from the Sierra de las Minas, Guatemala. - Herpetologica 38: 374-380.

DUELLMAN, W.E. & CAMPBELL, J.A. (1984): Two new species of Plectrohyla from Guatemala (Anura: Hylidae). - Copeia 1984: 390-397.

DUELLMAN, W.E. & CAMPBELL, J.A. (1992): Hylid frogs of the genus Plectrohyla: systematics and phylogenetic relationships. - Miscellaneous Publications, Museum of Zoology, University of Michigan 181: 1-32.

DUELLMAN, W.E. & FOUQUETTE, M.J. (1968): Middle American frogs of the Hyla microcephala group. - University of Kansas Science Bulletin 55: 329-379.

DUELLMAN, W.E. & HOYT, D.L. (1961): Description of a new species of Hyla from Chiapas, Mexico. - Copeia 1961: 414-417.

DUELLMAN, W.E. & KLAAS, L.T. (1964): The biology of the hylid frog Triprion petasatus. - Copeia 1964 (2): 308-321.

DUELLMAN, W.E. & LEHR, E. (2009): Terrestrial- breeding frogs (Strabomantidae) in Peru. - Natur und Tier Verlag, Miinster 382 pp.

DUELLMAN, W.E. & LYNCH, J.D. (1969): Descriptions of Atelopus tadpoles and their relevance to atelopodid classification. - Herpetologica 25: 231-240.

DUELLMAN, W.E. & MANESS, S.J. (1980): The reproductive behavior of some hylid marsupial frogs. - Journal of Herpetology 14:213-222.

DUELLMAN, W.E. & TRUEB, L. (1966): Neotropical hylid frogs, genus Smilisca. - University of Kansas Publications. Museum of Natural History 17: 281-375.

DUELLMAN, W.E. & TRUEB, L. (1967): Two new species of tree frogs (genus Phyllomedusa) from Panama. - Copeia 1967 (1): 125-131.

DUELLMAN, W.E. & TRUEB, L. (1986): Biology of amphibians. - McGraw-Hill Book Company. New York.

DUELLMAN, W.E. & VELOSO, A. (1977): Phylogeny of Pleurodema (Anura: Leptodactylidae): a biogeographic model. - Occasional Papers of the Museum of Natural History, University of Kansas 64: 1-46.

DUELLMAN, W.E. & WIENS, J.J. (1992): The status of the hylid frog genus Ololygon and the

Literature cited

recognition of Scinax WAGLER, 1830. - Occasional Papers of the Museum of Natural History of the University of Kansas 151: 1-23.

DUELLMAN, W.E. & WIENS, J.J. (1993): Hylid frogs of the genus Scinax Wagler, 1830, in Amazonian Ecuador and Peru. - Occasional Papers of the Museum of Natural History of the University of Kansas 153: 1-57.

DUMBACHER, J.P., WAKO, A,, DERRICKSON, S.R., SAMUELSON, A,, SPANDE, T.F. & DALY, J.W (2004): Melyrid beetles (Choresine): a putative source for the batrachotoxin alkaloids found in poison-dart frogs and toxic passer- ine birds. - Proceedings of the National Academy of Sciences of the United States of America 101: 15857-15860.

DUMERIL, A.H.A. (1853): M6moire sur les batraciens anoures, de la famille des hylaeformes ou rainettes, comprenant la description d'un genre nouveau et de onze especes nouvelles. - Annales Des Sciences Naturelles, Zool., ser. 3, 19: 135-179.

DUMERIL, A.M.C. & BIBRON, G. (1841): Erpktologie g6nerale ou Histoire naturelle complhte des reptiles. Vol. 8: 1-792. Paris: Librarie Enclyclopedique de Roret.

D&Rn, A.M.C., BIBRON, G. & DUMI~RIL, A.H.A. (1854): Erpktologie g6n6rale ou Histoire naturelle complete des reptiles. Vol. 9: 1-440.

DUNDEE, H.A. (1998): Bolitoglossa sooyorum. - Catalogue of American Amphibians and Reptiles 662: 1-2.

DUNDEE, H.A. & ROSSMAN, D.A. (1987): Geographic distribution, Bufo valliceps valliceps (Gulf Coast toad). - Herpetological Review 18 (2): 37.

DUNDEE, H.A. & SCHEIBE, J.S. (2000): Bolito- glossa yucatana (Peters): Yucatan salamander. - Catalogue of American Amphibians and Reptiles 702: 1-3.

DUNN, E.R. (1924a): New amphibians from Panama. - Occasional Papers of the Boston Society of Natural History 5: 93-95.

DWNN, E.R. (192413): Some Panamanian frogs. - Occasional Papers of the Museum of Zoology, University of Michigan 151: 1-16.

DUNN, E.R. (1924~): New salamanders of the genus Oedipus with a synoptical key. - Field Museum of Natural History Publication. Zoological Series 12: 95-100.

DUNN, E.R. (1926): The salamanders of the family Plethodontidae. - Smith College, Northampton, Mass.

DUNN, E.R. (1931): New frogs from Panama and Costa Rica. - Occasional Papers of the Boston Society of Natural History 5: 385401.

DUNN, E.R. (1932): A preoccupied name in Eleutherodactylus. - Copeia 1932: 97.

DUNN, E.R. (1933a): A new Hyla from the Panama Canal Zone. - Occasional Papers of the Boston Society of Natural History 8: 61-64.

DUNN, E.R. (193313): Amphibians and reptiles from El Valle de Anton, Panama. -

Occasional Papers of the Boston Society of Natural History 8: 65-79.

DUNN, E.R. (1934): Two new frogs from Darien. -American Museum Novitates 747: 1-2.

DUNN, E.R. (1937): The amphibian and reptilian fauna of bromeliads in Costa Rica and Panama. - Copeia 1937 (3): 163-167.

DUNN, E.R. (1940): New and noteworthy herpetological material from Panama. - Proceedings of the Academy of Natural Sciences of Philadelphia 92: 105-122.

DUNN, E.R. (1942a): The American Caecilians. - Bulletin of Museum of Comparative Zoology, Harvard University 91: 439-540.

DUNN, E.R. (194213): A new species of frog (Eleutherodactylus) from Costa Rica. - Notulae Naturae of the Academy of Natural Sciences of Philadelphia 104: 1-2

DUNN, E.R. (1949): Notes on South American frogs of the family Microhylidae. - American Museum Novitates 1419: 1-21.

DUNN, E.R., TRAPIDO, H. & EVANS, H. (1948): A new species of the microhylid frog genus Chiasmocleis from Panama. - American Museum Novitates 1376: 1-8.

DUTTA, S.K. & MANAMENDRA-ARACHCHI, K. (1996): The amphibian fauna of Sri Lanka. Wildlife Heritage Trust of Sri Lanka. Colombo, Sri Lanka. 230 pp.

EASTEAL, S. (1986): Bufo marinus. - Catalogue of American Amphibians and Reptiles 395: 1-4.

EHMCKE, J., WISTUBA, J., CLEMEN, G. & SCHLATT, S. (2003): Targeted expression of androgen receptors in tooth-forming tissues of a neotropical salamander (Bolitoglossa schizodactyla) enables region-specific formation of dimorphic types of teeth in the male. - General and Comparative Endocrinology 134 (1): 26-35.

ELIAS, P. (1984): Salamanders of the northwestern highlands of Guatemala. - Natural History Museum of Los Angeles County Contributions in Science 348: 1-20.

ELIAS, P. & WAKE, D.B. (1983): Nyctanolis pernix, a new genus and species of plethodontid salamander from northwestern Guatemala and Chiapas, Mexico. - Pp. 1-12 in: RHODIN, G.J. & MNATA, K (eds.): Advances in Herpetology and Evolutionary Biology:

Literature cited

Essays in Honors of Ernest E. Williams. Museum of Comparative Zoology. Cambridge, Massachusetts.

EXBRAYAT, J.-M. & HRAOUI-BLOQUET, S. (1994): An example of heterochrony: The metamorphosis in Gymnophiona. - Bulletin de la Sociktk Zoologique de France 119: 117-126.

FATVOVICH, J., HADDAD, C.F.B., GARCIA, P.C.O., FROST, D.R., CAMPBELL, J.A. & WHEELER, W.C. (2005): Systematic review of the frog family Hylidae, with special reference to Hylinae: Phylogenetic analysis and taxonomic revision. - Bulletin of the American Museum of Natural History 294: 1-240.

FAIVOVICH, J., HADDAD, C.F.B., BAETA, D., JUNGFER, K.-H., ALVARES, G.F.R., BRANDAO, R.A., S m n , C.A., BARRIENTOS, L.S., BARRIO- AMOROS, C.L., CRUZ, C.A.G. D. & WHEELER, W.C. (2010): The phylogenetic relationships of the charismatic poster frogs, Phyllomedusinae (Anura, Hylidae). -

Cladistics 26 (3): 227-261. FEDER, M.E. (1983): Integrating the ecology and

physiology of plethodontid salamanders. - Herpetologica 39: 291-300.

FEDER, M.E. & BURGGREN, W.W. (eds.) (1992): Environmental physiology of the amphibians. - The University of Chicago Press. Chicago. 646 pp.

FERRARO, D.P. & CASAGRANDA, M.D. (2009): Geographic distribution of the genus Pleurodema in Argentina (Anura: Leiuperidae). - Zootaxa 2024: 33-55.

FIRSCHEW, I.L. & SMITH, H.M. (1956): A new fringe-limbed Hyla (Amphibia: Anura) from a new faunal district of Mexico. -

Herpetologica 12: 17-21. FIRSCHEIN, I.L. & SMITH, H.M. (1957): A high

crested race of toad (Bufo valliceps) and other noteworthy reptiles and amphibians from southern Mexico. - Herpetologica 13: 219-222.

FISHER, M.C., GARNER, T.W.J. & WALKER, S.F. (2009): Global emergence of Batracho- chytrium den trobatidis and amphibian chytridiomycosis in space, time, and host. - Annual Review of Microbiology 63: 291-310.

FITZINGER, L.J.F.J. (1826): Neue Classification der Reptilien nach ihren Natiirlichen Venvandtschaften nebst "einer Venvandt- schafts-Tafel und einem Verzeichnisse der Reptilien-Sammlung des K. K. Zoologisch Museums zu Wien. - J.G. Heubner. Wien.

FLEET, R.R. & AUTREY, B.C. (1997): Bufo valliceps (Gulf coast toad). - Herpetological Review 28 (1): 48.

FLORES-VILLELA, O.A., MENDOZA-QUIJANO, F. & GONZALEZ-PORTER, G. (1995): Recopilacion de claves para la determinacion de anfibios y

reptiles de Mexico. - Publicaciones Especiales del Museo de Zoologia Uni- versidad Nacional de Autonoma de Mexico 10: 1-285.

FORD, L. & CAWATELLA, D. (1993): The major clades of frogs. - Herpetological Monographs 7: 94-117.

FORD, L.S. & SAVAGE, J.M. (1984): A new frog of the genus Eleutherodactylus (Lepto- dactylidae) from Guatemala. - Occasional Papers of the Museum of Natural History of the University of Kansas 110: 1-9.

FOSTER, M.S. & MCDIARMID, R.W. (1983): Rhinophrynus dorsalis (alma de vaca, sapo borracho, Mexican burrowing toad). - In: JANZEN, D.H. (ed.) Costa Rican Natural History. - Chicago & London (Univ. Chicago Press): 419-421.

FOUQUETTE, Jr, M.J. (1961): Status of the frog Hyla albomarginata in Central America. - Fieldiana Zoology 39 (55): 595-601.

FOUQUETTE, Jr, M.J. (1969): Rhinophrynidae, Rhinophrynus, R. dorsalis. - Catalogue of American Amphibians and Reptiles 78: 1-2.

F O U Q ~ T T E , Jr., M.J. & DELAHOUSSAYE, A.J. (1977): Sperm morphology in the Hyla rubra group (Amphibia: Anura: Hylidae), and its bearing on generic status. - Journal of Herpetology 11 (4): 387-396.

FOUQUETTE, Jr, M.J. & PYBURN! W.F. (1972): A new Colombian treefrog of the Hyla rubra complex. - Herpetologica 28: 176-181.

FOUQUETTE, Jr, MM & ROSSMAN, D.A. (1963): Noteworthy records of Mexican amphibians and reptiles in the Florida State Museum and Texas Natural History Museum. - Herpetologica 19 (3): 185-201.

FRANKLIN, C.J. & FRANKLIN, J. (1999): Bufo campbelli (Campbell's forest toad). - Herpetological Review 30 (2): 105.

FRANZEN, M. (1997): Ein Fund von Bolitoglossa gracilis BOLANOS, ROBINSON & WAKE, 1987. - Herpetofauna (19 (111): 23-25.

FREED, P. (1992): An anomalous Gulf Coast toad, Bufo valliceps. - Bulletin of the Chicago Herpetological Society 27 (7): 149.

FROST, D.R. (ed.) (1985): Amphibian Species of the World: A Taxonomic and Geographic Reference. -Allen Press and the Association of Systematic Collections. Lawrence, Kansas. 1-732.

FROST, D.R. (2010): Amphibian Species of the World: an Online Reference. Version 5.4 (8 April, 2010). Electronic Database accessible a t http:llresearch.amnh.orglvzkerpetol- ogylamphibial. American Museum of Natural History, New York, USA.

Literature cited

FROST, D.R., GRANT, T., FAIVOVICH, J. B e , R.H., HAAS, A., HADDAD, C.F.B., DE SA, R.O., CHANNING, A., WILKINSON, M., DONNELLAN, S.C., RAXWORTHY, C.J., CAMPBELL, J.A., BLOTTO, B.L., MOLER, P., DREWES, R.C., NUSSBAUM, R.A., LYNCH, J.D., GREEN D.M. & WHEELER, W.C. (2006a): The amphibian tree of life. - Bulletin of the American Museum of Natural History 297: 1-291.

FROST, D.R., GRANT, T. & MENDELSON, 111. J.R. (2006b): Ollotis Cope, 1875, is the oldest name for the genus current referred to as Cranopsis Cope, 1875. - Copeia 2006: 558.

FROST, D.R., G R . ~ , T., FANOVICH, J. BA;N, R.H., HAAS, A., HADDAD, C.F.B., DE SA, R.O., CHANNING, A,, WILKINSON, M., DONNELLAN, S.C., RAXWORTHY, C.J., CAMPBELL, J.A., BLOTTO, B.L., MOLER, P., DREWES, R.C., NUSSBAUM, R.A., LYNCH, J.D., GREEN D.M. & WHEELER, W.C. (2008): Is the amphibian tree of life really fatally flawed? - Cladistics 24: 385-395.

FROST, D.R., MENDELSON, J.R. I11 & PRAMW, J. (2009): Further notes on the nomenclature of Middle American toads (Bufonidae). - Copeia 2009 (2): 418.

FLWK, W.C., ANGULO, A., CALDWELL, J.P., RYAN, M.J. & CANNATELLA, D.C. (2008): Comparison of morphology and calls of two cryptic species of Physalaemus (Anura: Leiu- peridae). - Herpetologica 64: 290-304.

FUNK W.C., FLETCHER-LAZO G., NOGALES- SORNOSA F. & ALMEIDA-REINOSO D. (2004): First description of a clutch and nest site for the genus Caecilia (Gymnophiona: Caecili- idae). - Herpetological Review 35:128-130.

GAGLIARDO, R., CRUMP. P., GRIFFITH, E., MENDELSON, J., ROSS, H. & ZIPPEL, K. (2008): The principles of rapid response for amphibian conservation, using the programmes in Panama as an example. - International Zoo Yearbook 42: 125-135.

GAIGE, H.T. (1929): Three new tree frogs from Panama and Bolivia. - Occasional Papers of the Museum of Zoology University of Michigan 207: 1-6.

GAIGE, H.T. (1933): A new Gastrotheca from Colombia. - Occasional Papers of the Museum of Zoology University of Michigan 263: 1-3.

GAIGE, H.T. & STUART, L.C. (1934): A new Hyla from Guatemala. - Occasional Papers of the Museum of Zoology University of Michigan 281: 1-3.

GARC~A-PARIS, M., GOOD, D.A., PARRA-OLEA, G. & WAKE, D.B. (2000): Biodiversity of Costa Rican salamanders: implications of high levels of genetic differentiation and phylogeo- graphic structure for species formation. -

Proceedings of the National Academy of Sciences of the United States of America 97: 1640-1647.

GARCIA-PARIS, M., PARRA-OLEA, G., BRAME, A.H., JR., I1 & WAKE, D.B (2003 "2002"): Systematic revision of the Bolitoglossa mexicana species group (Amphibia: Plethodontidae) with description of a new species from Mexico. - Revista Espanola de Herpetologia 16: 43-71.

GARC~A-PARIS, M., PARRA-OLEA, G., BRAME, A.H. & WAKE, D.B. (2002): Systematic revision of the Bolitoglossa mexicana species group (Amphibia: Plethodontidae) with description of a new species from Mexico. - Revista Espafiola de Herpetologia 16: 43-71.

GARC~A-PARIS, M., PARRA-OLEA, G. & WAKE, D.B. (2000): Phylogenetic relationships within the lowland tropical salamanders of the Bolitoglossa mexicana complex (Amphibia: Plethodontidae). - Pp. 199-214 in: Bruce, R.C., Jaeger, R.G. & Houck, L.D. (eds.): The Biology of Plethodontid Salamanders. Kluwer Academic / Plenum Publishers. New York.

GARCIA-PAR~S, M, PARRA-OLEA, G. & WAKE, D.B. (2008): Description of a new species of the Bolitoglossa subpalmata group (Caudata: Plethodontidae) from Costa Rica. -

Herpetological Journal 18: 23-31. GARCIA-PARIS, M. & WAKE, D.B. (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 (1): 42-70.

GARCIA-RODR~GUEZ, A,, CHAVES, G., WAINWRIGHT, M. & VTLLEGAS, A. (2010): Geographic Distribution: Eleutherodactylus coqui (Puerto Rican Coqui). - Herpetological Review 41 (3): 375.

GARRETT, J.M. & BARKER, D.A. (1987): Field guide to reptiles and amphibians of Texas. - Texas Monthly Fieldguide Series, Gulf Publishing Company, Houston, Texas.

GIRARD, C. (1855 "1854"): Abstract of a report to Lieut. James M. Gilliss, U. S. N., upon the reptiles collected during the U. S. N. Astronomical Expedition to Chili. -

Proceedings of the Academy of Natural Sciences of Philadelphia 7: 226-227.

GOIN, C.J. (1964): Distribution and synonymy of Centrolenella fleischmanni in northern South America. - Herpetologica 20: 1-8.

GOOD, D.A. & WAKE, D.B. (1993): Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with descriptions of three new species. - Herpetological Mono- graphs 7: 131-159.

Literature cited

GOOD, D.A. & WAKE, D.B. (1997): Phylogenetic and taxonomic implications of protein variation in the Mesoamerican salamander genus Oedipina (Caudata: Plethodontidae). - Revista de Biologia Tropical 45: 1185-1208.

GORDON M.S., SCHMIDT-NEILSON, K. & KELLY, H.M. (1961): Osmotic Regulation in the Crab-Eating Frog (Rana cancrivora). J. Exp. Biol. 38(3): 659-678

GORHAM, S.W. (1966): Liste der rezenten Amphibien und Reptilien. Ascaphidae, Leiopelmatidae, Pipidae, Discoglossidae, Pelobatidae, Leptodactylidae, Rhinophryn- idae. - Das Tierreich Lieferung 85: I-XVI, 1- 222.

GOSNER, K.L. (1960): A simplified table for stag- ing anuran embryos and larvae with notes on identification. - Herpetologica 16: 183-190.

GOWER, D.J. & WILKINSON, M. (2005): Conservation Biology of caecilian amphibians. - Conservation Biology (19 (1): 45-55.

GOWER, D.J. & WILKINSON, M. (2008): Essay 1.2. Caecilians. - Pp. 19-20 in STUART, S., HOFFMANN, M., CHANSON, J., COX, N., BERRIDGE, R., R,AMANI P. & YOUNG, B. (eds.): Threatened Amphibians of the World. - Lynx Edicions. Barcelona, Spain. 758 pp.

GRANT, T. (2004): On the identities of Colostethus inguinalis (COPE, 1868) and C. panamensis (DUNN, 19331, with comments on C. latinasus (COPE, 1863) (Anura: Dendrobatidae). - American Museum Novi- tates 3444: 1-24.

GRANT, T., FROST, D.R., CALDWELL, J.P., GAGLIARDO, R., HADDAD, C.F.B., KOK, P. J.R., MEANS, D.B., NOONAN, B.P., SCHARGEL, W.E. & WHEELER, W. (2006): Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia, Athesphatanura, Dendroba- tidae). - Bulletin of the American Museum of Natural History 299: 1-262.

GRAVES, B.M. (1999): Die1 activity patterns of the sympatric poison dart frogs, Dendro- bates auratus and D. pumilio, in Costa Rica. - Journal of Herpetology 33 (3): 375-381.

GRAY, J.E. (1831): A synopsis of the species of the class Reptilia. - Pp. 1-110 in Griffith, E. & Pidgeon, E. (eds.): The animal kingdom, vol. 9: The class Reptilia. w i t t ake r , Treacher, and Co. London.

GRAY, J.E. (1868): Notice of two new salamanders from Central America. - Annals and Magazine of Natural History ser. 4, 2: 297-298.

GRAY, L.A. & RAND, A.S. (1997): A daybreak chorus in the frog, Agalychnis callidryas. - Journal of Herpetology 31 (3): 440441.

GRAYBEAL, A. & CANNATELLA, D.C. (1995): A new taxon of Bufonidae from Peru, with descriptions of two new species and a review of the phylogenetic status of supraspecific bufonid taxa. - Herpetologica 51: 105-131.

GRAYBEAL, A. & DE QUEIROZ, K. (1992): Inguinal amplexus in Bufo fastidiosus, with comments on the systematics of bufonid frogs. - Journal of Herpetology 26 (1): 84-87.

GREENBAUM, E. (2004): A new species of Bolitoglossa (Amphibia: Caudata: Pletho- dontidae) from montane forests in Guatemala and El Salvador. - Journal of Herpetology 38 (3): 411-421.

GREER, B.J. & WELLS, K.D. (1980): Terretorial and reproductive behavior of the tropical American frog Centrolenella fleischmami. - Herpetologica 36 (4): 318-326.

GROOM, M., MEFFE, G.K. & CARROLL, C.R. (2006): Principles of conservation biology. 3rd edition. - Sinauer Associates. Sunderland, Massachusetts.

GUAYASAMIN, J.M. & BONACCORSO, E. (2004): A new species of glass frog (Centrolenidae: Cochranella) from the lowlands of northwestern Ecuador, with comments on the Cochranella granulosa group. - Herpeto- logica 60 (4): 485494.

GUAYASAMIN, J.M., CASTROVIEJO-FISHER, S., AYARZAGUENA, J., TRUEB, L. & VILA C. (2008): Phylogenetic relationships of glassfrogs (Centrolenidae) based on mitochondria1 and nuclear genes. - Molecular Phylogenetics and Evolution 48: 574-595.

GUAYASAMIN, J.M., CASTROVIEJO-FISHER, S., TRUEB, L., AYARZAGUENA, J., RADA, M. & Vila C. (2009): Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. - Zootaxa 2100: 1-97.

GUAYASAMIN, J.M., CISNEROS-HEREDIA, D.F., Ymz-Mur;ioz, M.H. & BUSTAMANTE, M.R. (2006): Amphibia, Centrolenidae, Centro- Iene ilex, Centrolene litorale, Centrolene medemi, Cochranella albomaculata, Cochra- nella ametarsia: range extensions and new country records. - Check List. Journal of Species Lists and Distribution 2: 24-25.

G ~ T H E R , A. (1885-1902): Biologia Centrali- Americana. Reptilia and Batrachia. - Taylor and Francis. London. 326 pp.

GUYER, C. & DONNELLY, M.A. (2005): Amphi- bians and Reptiles of La Selva, Costa Rica and the Caribbean Slope: A Comprehensive Guide. - University of California Press, Berkeley.

HAAS, W. & KOHLER, G. (1997): Freiland- beobachtungen, Pflege und Zucht von Bufo

Literature cited

luetkenii BOULENGER, 1891. - Herpetofauna 19 (109): 5-9.

HAGEMAN, K.J., SIMONICH, S.L., CAMPBELL, D.H., WILSON, G.R. & LANDERS, D.H. (2006): Atmospheric deposition of current-use and historic-use pesticides in snow a t national parks in the western United States. - Environmental Science and Technology 40: 3174-3180.

HALLOWELL, E. (1861 "1860"): Report on the Reptilia of the North Pacific exploring expedition, under command of Capt. John Roger, U.S.N. - Proceedings of the Academy of Natural Sciences of Philadelphia 1860: 480-509.

HANKEN, J., LYNCH, J.F. & WAKE, D.B. (1980): Salamander invasion of the tropics. - Natural History Reader in Evolution 89 (12): 46-53.

HANKEN, J. & WAKE, D.B. (1982): Genetic differentiation among plethodontid salamanders (genus Bolitoglossa) in Central and South America: implications for the South American invasion. - Herpetologica 38 (2): 272-287.

HANKEN, J., WAKE, D.B. & SAVAGE, J.M. (2005):A solution to the large black salamander problem (genus Bolitoglossa) in Costa Rica and PanamB. - Copeia 2005: 227-245.

HARTWEG, N. (1941): Notes on the genus Plectrohyla, with descriptions of new species. - Occasional Papers of the Museum of Zoology, University of Michigan 437: 1-10.

HAUSWALDT, J.S., LUDEWIG, A.-K., VENCES, M. & PROHL, H. (2010): Widespread co-occurrence of divergent mitochondria1 haplotype lineages in a Central American species of poison frog (Oophaga pumilio). - Journal of Biogeography 2010, doi:10.1111/j.1365- 2699.2010.02438.x

HAYES, M.P. (1985): Nest structure and attendance in the stream-dwelling frog, Eleutherodactylus angelicus. - Journal of Herpetology (19 (1): 168-169.

HAYES, M.P. & STARRETT, P.H. (1980): Notes on a collection of centrolenid frogs from the Colombian Choc6. - Bulletin Southern California Academy of Sciences 79 (3): 89-96.

HAYES, M.P., POUNDS, J.A. & ROBINSON, D.C. (1986): The fringe-limbed tree frog, Hyla fimbrimembra (Anura: Hylidae): new records from Costa Rica. - Florida Scientist 49 (4): 193-198.

HAYES, M.P., POUNDS, J.A. & TIMMERMAN, W.W. (1989): An annotated list and guide to the amphibians and reptiles of Monteverde, Costa Rica. - Herpetological Circular 17: 1-67.

HAYES, T.B., COLLINS, A., LEE, M., MENDOZA, M., NORIEGA, N., STUART, A.A. & VONK, A. (2002): Hermaphroditic, demasculinized frogs after exposure to the herbicide atrazine a t low ecologically relevant doses. - Proceedings of the National Academy of Sciences of the United States of America 99: 5476-5480.

HAYES, T.B., STUART, A.A., MENDOZA, M., COLLINS, A., NORIEGA, pJ., VONK, A., JOHNSTON, G., LN, R. & DODZO, D. (2006): Characterization of Atrazine-Induced Gonadal Malformations in African Clawed Frogs (Xenopus laevis) and Comparisons with Effects of an Androgen Antagonist (Cyproterone Acetate) and Exogenous Estrogen (17,-Estradiol): Support for the Demasculinization/Feminization. ? Hypothesis. Environmental Health Perspectives 114: 134?141. doi: 10.1289lehp.8067

HEATWOLE, H. (1963): The frog genus Pipa in Panama. - Copeia 1963 (2): 436?438.

HEDGES, S.B., DUELLMAN, W.E. & HEINICKE, M.P. (2008): New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation. - Zootaxa 1737: 1-182.

HEINICKE, M.P., DUELLMAN, W.E. & HEDGES, S.B. (2007): Major Caribbean and Central American frog faunas originated by oceanic dispersal. - Proceedings of the National Academy of Sciences of the United States of America 104: 10092-10097.

HEMMING, F. (ed.) (1958): Opinion 520. Suppression under the Plenary Powers of the specific name tibiatrix LAURENTI, 1768, as published in the combination Hyla tibiatrix, and of the generic name Acrod!ytes FITZINGER, 1843, and interpretation under the same Powers of the nominal species Rana venulosa LAURENTI, 1768 (Class Amphibia). - Opinions and Declarations Rendered By The International Commission on Zoological Nomenclature 19: 169-200.

HERTZ, A., LOTZKAT, S., STADLER, L., HAMAD, N., CARRIZO, A. & KOHLER, G. (in press): Note- worthy amphibians from western Panama. - Herpetological Review.

HEYER, W.R. (1968): The proper name for the type-species of the genus Leptodactylus. - Copeia 1968: 160-162.

HEYER, W.R. (1969): Studies on the genus Leptodactylus (Amphibia, Leptodactylidae) 111. A redefinition of the genus Lepto- dactylus and a description of a new genus of leptodactylid frogs. Contributions in Science. - Natural History Museum of Los Angeles County 155: 1-14.

HEYER, W.R. (1970a "1968"): Studies on the genus Leptodactylus (Amphibia: Lepto-

Literature cited

dactylidae): 2. Diagnosis and distribution of the Leptodactylus of Costa Rica. - Revista de Biologia Tropical 16 (2): 171-205.

HEYER, W.R. (1970b): Studies on the frogs of the genus Leptodactylus (Amphibia: Lepto- dactylidae). VI. Biosystematics of the melanonotus group. Contributions in Science. - Natural History Museum of Los Angeles County 191: 1 4 8 .

HEYER, W.R. (1971): Leptodactylus labialis (COPE) white-lipped frog. - Catalogue of American Amphibians and Reptiles 104: 1-3.

HEYER, W.R. (1972): The status of Leptodactylus pumilio BOULENGER (Amphibia, Lepto- dactylidae) and the description of a new species of Leptodactylus from Ecuador. - Contributions in Science. Natural History Museum of Los Angeles County 231: 1-8.

HEYER, W.R. (1974): Relationships of the mar- moratus species group (Amphibia, Lepto- dactylidae) within the subfamily Lepto- dactylinae. - Natural History Museum of Los Angeles County Contributions in Science 253: 1 4 6 .

HEYER, W.R. (1978): Systematics of the fuscus group of the frog genus Leptodactylus (Amphibia, Leptodactylidae). - Natural History Museum of Los Angeles County Science Bulletin 29: 1-85.

HEYER, W.R. (1979): Systematics of the pentadactylus species group of the frog genus Leptodactylus (Amphibia: Leptodactylidae). - Smithsonian Contributions to Zoology 301: 1-43.

HEYER, W.R. (1994): Variation within the Leptodactylus podicipinus-wagneri complex of frogs (Amphibia: Leptodactylidae). - Smithsonian Contributions to Zoology 546: 1-124.

HEYER, W.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.

HEYER, W.R. (2005): Variation and taxonomic clarification of the large species of the Leptodactylus pentadactylus species group (Amphibia: Leptodactylidae) from Middle America, northern South America, and Amazonia. - Arquivos de Zoologia. Sgo Paulo 37: 269-348.

HEYER, W.R., DE SA, R.O. & HEYER, M.M. (2010): Leptodactylus savagei. - Catalogue of American Amphibians and Reptiles 867: 1-19.

HEE-ER, W.R., DE SA, R.O. & MULLER, S. (2002): Leptodactylus silvanimbus. - Catalogue of

American Amphibians and Reptiles 743: 1-3.

HEYER, W.R., DE SA, R.O. & MULLER, S. (2005): On the enigmatic distribution of the Honduran endemic Leptodactylus silvanimbus (Amphibia: Anura: Leptodactylidae). - Pp. 81-101 in: DONNELLY, M.A., CROTHER, B.I., GUYER, C., WAKE, M.H. & WHITE, M.E. (eds.): Ecology and evolution in the tropics: A herpetological perspective. - University of Chicago Press, Chicago.

HEYER, W.R., DONNELLEY, M.A., MCDIARMID, R.W., HAYEK, L.C. & FOSTER, M.S. (1994): Measuring and Monitoring Biological Diversity. Standard Methods for Amphi- bians. - Smithsonian Institution Press, Washington, D.C.

HEYER, W.R. & MAXSON, L R. (1982): Distributions, relationships, and zoogeography of lowland frogs: The Leptodactylus complex in South America, with special reference to Amazonia. - Pp. 375-388 in: FRANCE, G.T. (ed.): Biological diversification in the tropics. Columbia University Press. New York.

HEYER, W.R. & RAND, A.S. (1977): Foam nest construction in the leptodactylid frogs Leptodactylus pentadactylus and Physa- laemus pustulosus (Amphibia, Anura, Leptodactylidae. - Journal of Herpetology 11: 225-228.

HEYER, W.R. & REID, Y.R. (2003): Does advertisement call variation coincide with genetic variation in the genetically diverse frog taxon currently known as Leptodactylus fuscus (Amphibia: Leptodactylidae)? - Anais da Academia Brasileira de Cigncias 75: 39-54.

HEYER, W.R., DE SA, R., MCCRANIE, J.R. & WILSON, L.D. (1996. Leptodactylus silvanimbus (Amphibia: Anura: Leptodactylidae): natural history notes, advertisement call, and relationships. - Herpetological Natural History 4 (2): 169-174.

HILLIS, D.M. (1988): Systematics of the Rana pipiens complex: puzzle and paradigm. - Annual Review of Systematics and Ecology 19: 39-63.

HILLIS, D.M. & DE SA, R. (1988): Phylogeny and taxonomy of the Rana palmipes group (Salientia: Ranidae). - Herpetological Mono- graphs 2: 1-26.

HILLIS, D.M. & FROST, J.S. (1985): Three new species of leopard frogs (Rana pipiens complex) from the Mexican Plateau. - Occasional Papers of the Museum of Natural History of the University of Kansas 117: 1-14.

H ~ L I S , D.M., FROST, J.S. &WRIGHT, D.A. (1983): Phylogeny and biogeography of the Rana

343

Literature cited

pipiens complex: a biochemical evaluation. - Systematic Zoology 32: 132-143.

HILTY, J. & MERENLENDER, A. (2000): Faunal indicator taxa selection for monitoring ecosystem health. - Biological Conservation 92: 185-197.

HOBEL, G. (1999a): Notes on the natural history and habitat use of Eleutherodactylus fitzingeri (Anura: Leptodactylidae). - Amphi- bia-Reptilia (20 (1): 65-72.

HOBEL, G. (199913): Facultative nest construction in the gladiator frog Hyla rosenbergi (Anura: Hylidae). - Copeia 1999 (3): 796-800.

HOBEL, G. (2000): Reproductive ecology of Hyla rosenbergi in Costa Rica. - Herpetologica 56 (4): 446-454.

HOFFMANN, H. (2004): Description of the previously unknown tadpole of Hyalino- batrachium pulveratum (Anura: Centro- lenidae). - Revista de Biologia Tropical 52 (1): 219-228.

HOFFMANN, H. (2005a): Some ecological notes on A. annae (Anura: Hylidae). - Brenesia 65: 73-77.

HOFFMANN, H. (2005b): The tadpoles of Hyla rufitela (Anura: Hylidae). - Revista de Biologia Tropical 53 (3-4): 561-568.

HOFFMANN, H. (2010): The glass frog tadpoles of Costa Rica (Anura: Centrolenidae): A study of morphology. - Abhandlungen der Sencken- berg Gesellschaft fiir Naturforschung 567: 1-78.

HOFFMANN, M., CHURCH, D., COLLINS, J.P., COX, N., GASCON, C., MENDELSON 111, J.H., MOORE, R.D., STUART, S.N. & ZIPPEL, K.C. (2008): Amphibian conservation - responding to the global decline of amphibians. - Pp. 114-124 in: STUART, S., HOFFMANN, M., CHANSON, J., Cox, N., BERRIDGE, R., RA~MANI P. &YOUNG, B. (eds.): Threatened Amphibians of the World. - Lynx Edicions. Barcelona, Spain. 758 pp.

HOLDRIDGE, L.R. (1967): Life Zone Ecology. - Tropical Science Center, San Jose, Costa Rica.

HOLMAN, J.A. & BIRKENHOLZ, D.E. (1963): On a collection of amphibians and reptiles from Guatemala. - Herpetologica 19 (2): 144-146.

HOOGMOED, M.S. (1989): South American bufonids (Amphibia: Anura: Bufonidae), an enig- ma for taxonomists. Pp. 167-180 in: FONTANET, X. (ed.): Treballs d'Ictiologia i Herpetologia. Volume 2. Societat Catalana d'Ictiologia i Herpetologia, Barcelona.

HOOGMOED, M.S. & GORZULA, S.J. (1979): Checklist of the savanna inhabiting frogs of the El Manteco region with notes on their ecology and the description of a new species

of treefrog (Hylidae, Anura). - Zoologische Mededelingen 54 (13): 183-216.

I B ~ z , D.R., & CRAWFORD, A.J. (2004): A new species of Eleutherodactylus (Anura: Leptodactylidae) from the DariCn Province, PanamB. - Journal of Herpetology 38: 240-244.

IBA~EZ, D.R., & JARAMILLO, C.A. (1997): Hyalinobatrachium chimipoi (Suretka glass frog). - Herpetological Review 28 (1): 48.

I B ~ z , D.R., JARAMILLO, C.A. & AROSEMENA, F.A. (1994): A new species of E1eutherodactylus (Anura: Leptodactylidae) from Panama. - Amphibia-Reptilia 15 (4): 337-341.

I B ~ z , D.R., JARAMILLO, C.A., SOLIS, F.A. & JARAMILLO, F.E. (1991): Hyla fimbrimembra (fringe-limbed treefrog). - Herpetological Review 22 (4): 133-134.

I B ~ z , D.R., JARAMILLO, C.A. & SOLIS, F.A. (1995): Una especie nueva de Atelopus (Amphibia: Bufonidae) de Panama. - Carib- bean Journal of Science 31 (1-2): 57-64.

IBANEZ, R., JARAMILLO, F. & JARAMILLO, C. (1999a): Ampliacion del Bmbito de distribu- cidn y description del renacuajo de la rana de cristal Hyalinobatrachium aureoguttatum (Anura: Centrolenidae). - Revista de la Academia Colombiana de Ciencias Exactas Fisicas y Naturales 23: 293-298.

I B ~ z , D.R., & RAND, S. (1990): Eleuthero- dactylus johnstonei. Panama. - Herpeto- logical Review 21: 37.

IBA-z D., R., R m , A.S. & JARAMILLO A., C.A. (199913): Los Anfibios del Monument0 Natural Barro Colorado, Parque Nacional Soberania y Areas Adyacentes 1 The Amphibians of Barro Colorado Nature Monument, Soberania National Park and Adjacent Areas. - D'Vinni Editorial Ltda., Santa F6 de BogotB, Colombia. 187 pp.

I B ~ E Z D., R. & SMITH, E.M. (1995): Systematic status of Colostethus flotator and C. nubicola (Anura: Dendrobatidae) in Panama. - Copeia 1995: 446456.

International Commission on Zoological Nomenclature ( 1987): Opinion 1462 Caeciliidae Rafinesque-Schmaltz, 1814 (Amphibia, Gymnophiona) and Caeciliidae Kolbe, 1880 (Insects, Psocoptera): a ruling to remove the homonymy. - Bulletin of Zoological Nomenclature 44 (4): 263-264.

JACOBSEN, S.K. (1985): Reproductive behavior and male mating success in two species of glass frogs (Centrolenidae). - Herpetologica 41 (4): 396-404.

JACOBSON, S.K. & VANDENBERG, J.J. (1991): Reproductive ecology of the endangered Golden Toad (Bufo periglenes). - Journal of Herpetology 25 (3): 321-327.

Literature cited

JAEGER, R.G., HAILMAN, J.P. & JAEGER, L.S. (1976): Bimodal die1 activity of a Panamanian dendrobatid frog, Colostethus nubicola, in relation to light. - Herpetologica 32: 77-81.

JAMES, M.S. (1944): Notes on the breeding of Hyla nigripes in Costa Rica. - Copeia 1944: 147-148.

JAMES, P. (1966): The Mexican burrowing toad, Rhinophrynus dorsalis, an addition to the vertebrate fauna of the United States. - Texas Journal of Science 18: 272-276.

JANSEN, M. & KOHLER, G. (2001): ijkologie des Mombacho-Salamanders, Bolitoglossa mombachoensis KOHLER & MCCRANIE, 1999, am Volcan Mombacho, Nicaragua (Caudata: Plethodontidae). - Salamandra 37 (2): 83-98.

JANSEN, M., SCHULZE, A., WERDING, L. & STREIT, B. (2009): Effects of extreme drought in the dry season on an anuran population in the Bolivian Chiquitano region. - Salamandra 45: 233-238.

JARAMILLO, C.A., CRAWFORD, A.J. & I B ~ E Z , R. (2009): Geographic Distribution: Incilius melanochlorus. - Herpetological Review 40 (1): 108.

JARAMILLO, F.E., JARAMILLO, C.A. & I B ~ E Z , R. (1997): Renacuajo de la rana de cristal Hyalinoba trachium colym biphyllum (Anura: Centrolenidae). - Revista de Biologia Tropical 45 (2): 867-870.

JOHNSON, A.J. & WELLEHAN, J.F.X. (2005): Amphibian virology. - Veterinary Clinics of North America: Exotic Animal Practice 8: 53-65.

JOHNSON, J.A. (2002): Bufo valliceps (Gulf Coast toad). - Herpetological Review 33 (3): 219.

JOHNSON, J.D. & SAVAGE, J.M. (1995): A new species of the Eleutherodactylus rugulosus group (Leptodactylidae) from Chiapas, Mexico. - Journal of Herpetology 29 (4): 501-506.

JOHNSON, J.D., WILSON, L.D. & TOWNSEND, J.T. (2010): Recent taxonomic changes. - Pp. 19-27. In Wnsos, L.D., TOWNSEND, J.H. & JOHNSON, J.D. (eds.): Conservation of Mesoamerican Amphibians and Reptiles. - Eagle Mountain Publishing, LC, Eagle Mountain, Utah. i-xviii + 1'-812 pp.

JOHNSON, M.L. & SPEARE, R. (2003): Survival of. Batrachochytrium dendrobatidis in water: quarantine and disease control implications. - Emerging Infectious Diseases 9: 922-925.

JOKUSH, E.L. & GARCIA-PARIS, M. (1998): Nototriton abscondens (Cordilleran moss salamander): reproduction. - Herpetological Review 29: 38.

JUNGFER, K.-H. (1987): Beobachtungen an Ololygon boulengeri (COPE, 1887) und anderen 'Knickzehenlaubfroschen'. -

Herpetofauna 9: 6-12. JUNGFER, K.-H. (1996): Reproduction and

parental care of the coronated treefrog, Anotheca spinosa. - Herpetologica 52 (1): 25-32.

JUNGFER, K.-H. & WEYGOLDT, P. (1994): The reproductive biology of the leaf frog Phyllomedusa lemur BOULENGER, 1882, and a comparison with other members of the Phyllomedusinae. - Revue Franqaise d'Aquariologie 21: 57-64.

JUNGFER, K.-H., WEYGOLDT, P. & JURASKE, N. (1996): Dendroba tes vicentei, ein neuer Pfeilgiftfrosch aus Zentral-Panama. -

Herpetofauna. Weinstadt 18: 17-26. JUNGFER, K.-H., BIRKHAHN, H., KULPMANN, V. &

WASSMANN, K. (1996): Haltung und Fortpflanzung von Dendrobates fulguritus SILVERSTONE, 1975, mit Anmerkungen zur Gattung Minyobates MYERS, 1987. - Herpetofauna 18 (104): 19-27.

JUNGFER, K.-H., LOTTERS, S. & JORGENS, D. (2000): Der kleinste Pfeilgiftfrosch - eine neue Dendrobates-Art aus West-Panama. - Herpetofauna 22 (129): 11-18.

KAGARISE SHERMAN, C. & MORTON, M.L. (1993): Population declines of Yosemite toads in the eastern Sierra Nevada of California. - Journal of Herpetology 27: 186-198.

KAISER, H. (1997): Origins and introductions of the Caribbean frog, Eleutherodactylus johnstonei (Leptodactylidae): management and conservation concerns. - Biodiversity and Conservation 6: 1391-1407.

KAISER, H. & HARDY, Jr, J.D. (1994): Eleuthero- dactylus johnstonei. - Catalogue of American Amphibians and Reptiles 581: 1-5.

KAISER, H., BARRIO AMOR~S, C.L., TRUJILLO, J.D. & LYNCH, J.D. (2002): Expansion of EleutherodactyIus johnstonei in northern South America: Rapid dispersal through human interactions. - Herpetological Review 33 (4): 290-294.

KARRAKER, N.E., RICHARDS, C.L. & ROSS, H.L. (2006): Reproductive ecology of Atelopus zeteki and comparisons to other members of the genus. - Herpetological Review 37 (3): 284-288.

KEFERSTEW, W. (1868a): m e r einige Batrachier aus Costarica. - Archiv fiir Naturgeschichte, Berlin 34: 291-300.

KEFERSTEIN, W. (1868b): Beschreibung einiger neuer Batrachier aus Australia und Costa Rica. - Nachrichten von der Gesellschaft der Wissenschaften zu Gottingen 15: 326-332.

345

Literature cited

KELLOGG, R. (1928): An apparently new Hyla from El Salvador. - Proceedings of the Biological Society of Washington 41: 123-124.

KELLOGG, R. (1932): Mexican tailless amphibians in the United States National Museum. - Bulletin of the United States National Museum 160: 1-224.

KIESECKER, J.M. (2002): Synergism between trematode infection and pesticide exposure: A link to amphibian limb deformities in nature? - Proceedings of the National Academy of Sciences of the United States of America 99: 9900-9904.

KIESECKER, J.M., BLAUSTEIR', A.R. & BELDEN, L.K. (2001): Complex causes of amphibian population declines. - Nature 410: 681-684.

KLUGE, A.G. (1979): The gladiator frogs of Middle America and Colombia - a reevaluation of their systematics (Anura: Hylidae). - Occasional Papers of the Museum of Zoology University of Michigan 688: 1-24.

KLUGE, A.G. (1981): The life history, social organisation and parental behaviour of Hyla rosenbergi BOULENGER, a nest building gladiator frog. - Miscellaneous Publications of the Museum of Zoology University of Michigan 160: 1-170.

KOHLER, G. (1993): Bufo coniferus COPE. - Sauria, Berlin, 15 (3): 1-2.

KOHLER, G. (1997): Triprion petasatus (COPE). - Sauria, Berlin, 19 (1): 1-2.

KOHLER, G. (1998): Eleutherodactylus Iauraster. - Herpetological Review 29 (4): 246.

KOHLER, G. (2001): Anfibios y Reptiles de Nica- ragua. - Herpeton. Offenbach, Germany.

KOHLER, G. (2002): A new species of salamander of the genus Nototriton from Nicaragua (Amphibia: Caudata: Plethodontidae). - Herpetologica 58: 205-210.

KOHLER, G. (2003): Bufo valliceps (Gulf Coast toad). - Herpetological Review 34 (2): 161.

KOHLER, G. (2008): Reptiles of Central America. - Herpeton. Offenbach, Germany.

KOHLER, G. & MCCRANIE, J.R. (1999): A new species of salamander from Volcan Mom- bacho, Nicaragua, formerly referred to Bolitoglossa striatula (Amphibia, Caudata, Plethodontidae). - Senckenbergiana Biolo- gica 79 (1): 89-93.

KOHLER, G., LEHR, E. & MCCRANIE, J.R. (2000): The tadpole of the Central American toad Bufo luetkenii BOULENGER. - Journal of Herpetology 34 (2): 303-306.

KOHLER, G. PONCE, M. & BATISTA, A. (2007): A new species of worm salamander (genus Oedipina) from Fortuna, western central Panama (Amphibia, Caudata, Pletho-

346

dontidae). - Senckenbergiana Biologica 87 (2): 213-217.

KOHLER, G. & SUNYER, J. (2006): A new species of rain frog (genus Craugastor) of the fitzingeri group from Rio San Juan, southeastern Nicaragua (Amphibia, Anura, Leptodactylidae). - Senckenbergiana Biologica 86: 261-266.

KOHLER, G., V E S ~ Y , M. & GREENBAUM, E. (2006): The amphibians and reptiles of El Salvador. - Krieger Publishing Company, 238 S.; Malabar, Florida.

KOHLER, J. & BOHME, W. (1996): Anuran amphibians from the region of Pre-Cambrian rock outcrops (inselbergs) in northeastern Bolivia, with a note on the gender of Scinax Wagler, 1830 (Hylidae). - Revue Franqaise de Aquariologie 23 (3-4): 133-140.

KOLBY, J.E. & MCCRANIE, J.R. (2009): Discovery of a surviving population of the montane streamside frog Craugastor milesi (SCHMIDT). - Herpetological Review 40 (4): 444.

KOLBY, J.E., MCCRANIE, J.R. & ROVITO, S.M. (2009): Geographic distribution. Nototriton brodiei (NCN). - Herpetological Review 40 (4): 444.

~ C K , K. (2006): The lost world of the Kihansi toad. - Science 311: 1230-1232.

KUBICKI, B. (2004a): Rediscovery of Hyalino- batrachium chirripoi (Anura: Centrolenidae) in southeastern Costa Rica. - Revista de Biologia Tropical 52 (1): 215-218.

KUBICKI, B. (2004b): Leaf-frogs of Costa Rica. - Editorial INBio, San Jos6, Costa Rica.

KDICKI, B. (2007) Ranas de vidrio de Costa Rica 1 Glass frogs of Costa Rica. - Editorial INBio, Santo Domingo de Heredia, 304 pp.

LA MARCA, E., LIPS, K.R., LOTTERS, S., PUSCHENDORF, R., IBAP~EZ, R., RUEDA- ALMONACID, J.V., SCHULTE, R., W T Y , C., CASTRO, F., MANZANILLA-PUPPO, J., GARCLA- PEREZ, J.E., TOW, E., B O ~ O S , F., CHAVES, G., POUNDS, J.A. & YOUNG, B. (2005): Catastrophic population declines and extinctions in Neotropical harlequin frogs (Bufonidae: Atelopus). - Biotropica 37 (2): 190-201.

LA W C A , E. & LOTTERS, S. (1997): Monitoring of declines in Venezuelan Atelopus. - Pp. 207-213 in: B o H M E , ~ . , BISHOFF, W. & ZIEGLER, T. (eds.): Herpetologia Bonnensis. - Society European Herpetology. Bonn.

WM, P. N. & SAVAGE, J.M. (1992): A new species of caecilian from the Peninsula de Osa of Costa Rica. - Copeia 1992 (3): 703-708.

Literature cited

LAMPERT, K.P., BERNAL, X.E., RAND, AS., MUELLER, U.G. & RYAN, M.J. (2007): Island populations of Physalaemus pustulosus: history influences genetic diversity and morphology. - Herpetologica 63: 3 11-319.

LAMPO, M., BARRIO-AMOROS, C. & HAN, B. (2006): Batrachochytrium dendrobatidis Infection in the Recently Rediscovered Atelopus mucubajiensis (Anura, Bufonidae), a Critically Endangered Frog from the Venezuelan Andes. - EcoHealth doi: 10.1007/~10393-006-0068-y.

LANGONE, J.A. (1992): Comentarios sobre algunos taxa asignados a1 g6nero Scinax WAGLER, 1830 (Amphibia, Anura, Hylidae) - Boletin de la Asociacidn Herpetologica Argentina. 8: 5-6.

LANNOO, M. (ed.) (2005): Amphibian declines: The conservation status of United States species. - University of California Press. Berkely, California.

LANNOO, M. (2008): Malformed frogs: The col- lapse of aquatic ecosystems. - University of California Press. Berkely, California.

LARSEN, J.H., JR., BENESKI, J.T., Jr. & WAKE, D.B. (1989): Hyolingual feeding systems of the Plethodontidae: comparative kinematics of prey capture by salamanders with free and attached tongues. - Journal of Experimental Zoology 252 (1): 25-33.

LARSON, A. (1983): A molecular phylogenetic perspective on the origins of a lowland tropical salamander fauna. I. Phylogenetic infer- ences from protein comparisons. - Herpetologica 39: 85-99.

LAURANCE, W.F., MCDONALD, K.R. & SPEARE, R. (1996a): Catastrophic declines of Australian rain forest frogs: Support for the epidemic disease hypothesis. - Conservation Biology 10: 1-9.

LAURANCE, W.F., MCDONALD, K.R. & SPEARE, R. (199613): Epidemic disease and the catastrophic decline of rain forest frogs. -Consemation Biology 10 (2): 406413.

LAURENTI, J.N. (1768): Specimen medicum, exhibens synopsin Reptilium emendatum cum experimentis circa venena et antidota Reptilium Austriacorum. 214 pp.

LAVILLA, E.0. & DE SA, R. (2001): Chondro- cranium and visceral skel'eton of Atelopus tricolor and Atelophryniscus chrysophorus tadpoles (Anura, Bufonidae). -

Amvhibia-Revtilia 22 (2): 167-177.

LAZCANO-BARRERO, M.A. (1992): Bolitoglossa mulleri (Brocchi). - Catalogue of American Amphibians and Reptiles 533: 1-2.

LEARY, C. & RAZAFINDRATSITA, V. (1998): Attempted predation on a hylid frog, Phrynohyas venulosa, by an indigo snake, Drymachon corais, and the response of conspecific frogs to distress calls. - Amphibia-Reptilia 19: 442446.

LEE, J.C. (1976): Rana maculata Brocchi, an Addition to the Herpetofauna of Belize. - Herpetologica 32 (2): 211-214.

LEE, J.C. (1996): The Amphibians and Reptiles of the Yucatan Peninsula. - Cornell University Press. Ithaca, New York, USA.

LEE, J.C. (2000): A Field Guide to the Amphibians and Reptiles of the Maya World. - Cornell University Press. Ithaca, New York, USA.

LEE, J.C. & SALZBURG, M.A. (1989): Mating success and pairing patterns in Bufo valliceps (Anura: Bufonidae). - Southwestern Naturalist 34 (1): 155-157.

LEENDERS, T. (2001): A guide to amphibians and reptiles of Costa Rica. - Zona Tropical, Miami.

LEENDERS, T.A.A.M. & WATKINS-COLWELL, G.J. (2003): Morphological and behavioral adaptations in Bolitoglossa colonnea (Caudata, Plethodontidae) in relation to habitat use and daily activity cycle. - Phyllomedusa 2 (2): 101-104.

LEGLER, J.D. (1964): A narrow mouthed toad (Gastrophryne usta) in El Salvador. - Herpetologica 19 (4): 286-287.

LICHTENSTEIN, M.H.C. & VON MARTENS, E.C. (1856): Nomenclator Reptilium et Amphibiorum Musei Zoologici Berolinensis. Akademie der Wissenschaften zu Berlin. Berlin.

LIEBERMAN, S.S. (1986): Ecology of the leaf litter herpetofauna of a Neotropical rain forest La Selva, Costa Rica. - Acta Zoologica Mexicana Nueva Serie 15: 1-72.

LINDER, G.L., KREST, S.K. & SPARLING, D.W. (eds.) (2004): Amphibian decline: An integrated analysis of multiple stressor effects. - SETAC Press, Pensacola, Florida.

LINDQUIST, E.D. (1995): Atelopus zeteki (Pan- amanian golden frog): Pure tonal vocalization. - Herpetological Review 26 (4): 200-201.

LAVILLA, E.O., VAIRA, M. & FER-I, L. (2003): A LINDQUIST, E.D. & HETHERINGTON, T.E. (1996): new species of ~ ~ ~ ~ h i ~ t ~ ~ l ~ i ~ ( A ~ ~ ~ ~ : Field studies on visual and acoustic signal- ~ i ~ ~ ~ h ~ l i d ~ ~ ) from the h d e a n yungas of ing in the "earless" Panamanian Golden Argentina, with comments on the Frog, Atelopus zeteki. - Journal of Elachistocleis ovalisE. bicolor controversy. 30 (3): 347-354. - Amphibia-Reptilia 24: 269-284.

Literature cited

LINDQUIST, E.D. & HETHERINGTON, T.E. (1998a): Tadpoles and juveniles of the Panamanian Golden Frog, Atelopus zeteki (Bufonidae), with information on development of coloration and patterning. - Herpetologica 54 (3): 370-376.

LINDQUIST, E.D. & HETHERINGTON, T.E. (1998b): Semaphoring in an earless frog: the origin of a novel visual signal. - Animal Cognition 1 (2): 83-87.

LINDQUIST, E.D., HETHERINGTON, T.E. & BADGLEY, D.D. (1999): Atelopus zeteki (Panamanian golden frog): Attempted predation. -

Herpetological Review 30 (1): 36. LINDQUIST, E. D. & SWIHART, D. W. (1997):

Atelopus chiriquiensis (Chiriqui Harlequin Frog): Mating behaviour and egg-laying. - Herpetological Review 3 (28): 145-146.

LINNAEUS, C. (1758): Systema naturae per regna tria naturae, secundum classes, ordines, genera, species cum characteribus, differen- tiis, synonymis, locis. Ed 10 Tom 1: 824 pp. Laurentii Salvii. Stockholm.

LPS, K.R. (1993a): Bolitoglossa compacta. - Herpetological Review 24 (3): 107.

LIPS, K.R. (199313): Bolitoglossa minutula. - Herpetological Review 24 (3): 107.

LIPS, K.R. (1993~): Bolitoglossa nigrescens. - Herpetological Review 24 (3): 107.

LIPS, K.R. (1993d): Oedipina grandis. -

Herpetological Review 24 (3): 107. LIPS, K.R. (1996): New treefrog from the

Cordillera de Talamanca of Central America with a discussion of systematic relationships in the Hyla lancasteri group. - Copeia 1996: 615-626.

LIPS, K.R. (1998): Decline of a tropical montane amphibian fauna. - Conservation Biology 12 (1): 106-117.

LPS, K.R. (1999): Mass mortality and population declines of anurans at an upland site in western Panama. - Conservation Biology 13: 117-125.

LPS, K.R., DIFFENDORFER, J., MENDELSON 111, J.R. & SEARS, M.W. (2008): Riding the wave: Reconciling the roles of disease and climate change in amphibian declines. - PloS Biology 6 (3): e72.doi:lO. 13711journa1.pbi0.0060072.

LPS, K.R. & DONNELLY, M.A. (2005): Lessons from the tropics. - Pp. 198-205 in LANNOO, M. (ed.): Amphibian declines: The conservation status of United States species. - University of California Press. Berkely, California.

LIPS, K.R. & KREMPELS, D.M. (1995): Eggs and tadpoles of Bufo fastidiosus COPE, with com-

ments on reproductive behavior. - Copeia 1995 (3): 741-746.

LIPS, K.R., GREEN, D.E. & PAPENDICK, R. (2003a): Chytridiomycosis in wild frogs from southern Costa Rica. - Journal of Herpetology 37: 215-218.

LIPS, K.R., GREEN, D.E., BREM, F., BRENES, R., REEVE, J.D., ALFORD, R.A., V o n ~ s , J., CAREY, C., LIVO, L.; PESSIER, A.P. & COLLINS, P. (2006): Emerging infectious disease and the loss of biodiversity in a Neotropical amphibian community. - Proceedings of the National Academy of Sciences of the United States of America 103: 3165-3170

LIPS, K.R., MENDELSON 111, J.R., MUNOZ-ALONSO, A., CANSECO-MARQUEZ, L. & MULCAHY, D.G. (2004): Amphibian population declines in montane southern Mexico: resurveys of historical localities. Biological Conservation 119: 555-564.

LIPS, K.R., REEVE, J.D. & WITTERS, L.R. (2003b): Ecological traits predicting amphibian population declines in Central America. - conservation Biology 17: 1078-1088.

LIPS, K.R. & SAVAGE, J.M. (1996): A new species of rainfrog, EIeutherodactyIus phasma (Anura: Leptodactylidae), from montane Costa Rica. - Proceedings of the Biological Society of Washington 109: 744-748.

LOADER, S.P., WILKINSON, M., GOWER, D.J. & MSUYA, C.A. (2003): A remarkable young Scolecomorphus vittatus (Amphibia: Gymnophiona: Scolecomorphidae) from the North Pare Mountains, Tanzania. - Journal of Zoology 259: 93-101.

LOTTERS, S. (1996): The Neotropical toad genus Atelopus. Checklist - Biology - Distribution. - Vences und Glaw Verlags GbR, Koln, Germany. 129 pp.

LOTTERS, S., B o r n , W. & G ~ H E R R. (1998): Notes on the type material of the neotropical harlequin frogs Atelopus varius (LICHTEN- STEIN & MARTENS, 1856) and Atelopus cruciger (LICHTENSTEIN & MARTENS, 1856) deposited in the Museum fiir Naturkunde of Berlin (Anura, Bufonidae). - Mitteilungen aus dem Museum fiir Naturkunde in Berlin. Zoologische Reihe 74: 173-184.

LOITERS, S., JUNGFER, K.H., HENKEL, F.W. & SCHMIDT, W. (2007): Pfeilgiftfrosche. Biolo- gie, Haltung, Arten. - Edition Chimaira, Frankfurt am Main, Germany. 668 pp.

LOTTERS, S., LA W C A , E., GAGLIARDO, R., SENARIS, J.C. & VEITH, M. (2005): Harlequin frogs back? Some thoughts and speculations. - Froglog 70:l-3

LONGCORE, J.E, PESSIER, A.P & NICHOLS, D.K. (1999): Batrachochytrium dendrobatidis

Literature cited

gen. et sp. nov., a chytrid pathogenic to amphibians. - Mycologia. 91: 219-27.

L~PEZ, L.O., WOOLRICH PIRA, G.A. & LEMOS ESPINAL, J.A. (2009): La farnilia Bufonidae en MBxico. Universidad Nacional Autonoma de MCxico. Cornision Nacional Para El Conocimiento Y Uso De La Biodiversidad. MBxico D.F. 139 pp.

LYNCH, J.D. (1965a): A review of the eleuthero- dactylid frog genus Microbatrachylus (Leptodactylidae). - Natural History Miscellanea. Chicago 182: 1-12.

LYNCH, J.D. (196513 "1964"): Two new species of Eleutherodactylus from Mexico (Amphibia: Leptodactylidae). - Herpetologica 20: 246-252.

LYNCH, J.D. (1965~): The races of the microhylid frog, Gastrophryne usta, in Mexico. - Transactions of the Kansas Academy of Science 68: 396400.

LYNCH, J. D. (196511): A review of the rugulosus group of Eleutherodactylus in northern Central America. - Herpetologica 21: 102-113.

LYNCH, J.D. (1966): A new species of EIeutherodactylus from Chiapas, Mexico (Amphibia: Leptodactylidae). - Transactions of the Kansas Academy of Science 69 (1): 76-78.

LYNCH, J.D. (1967a): Two new species of Eleutherodactylus from Guatemala and Mexico (Amphibia: Leptodactylidae). - Transactions of the Kansas Academy of Science 70: 177-183.

LYNCH, J.D. (1967b): Two new Eleutherodactylus from western Mexico (Amphibia: Lepto- dactylidae). - Proceedings of the Biological Society of Washington 80: 211-218.

LYNCH, J.D. (1970a): A taxonomic revision of the Leptodactylid frog genus Syrrhophus COPE. - University of Kansas Publications, Museum of Natural History 20 (1): 1 4 5 .

LYNCH, J.D. (1970b): Systematic status of the American leptodactylid frog genera Engy- stomops, Eupemphix, and Physalaemus. - Copeia 1970: 488496.

LYNCH, J.D. (1971): Evolutionary relationships, osteology, and zoogeography of leptodactylid frogs. - Miscellaneous, Publications, University of Kansas, Museum of Natural History 53: 1-238.

LYNCH, J.D. (1980a): A new frog of the genus, Eleutherodactylus from western Panama. - Transactions of the Kansas Academy of Science 83 (3): 101-105.

LYNCH, J.D. (1980b): Systematic status and distribution of some poorly known frogs of the genus Eleutherodactylus from the Chocoan

lowlands of South America. - Herpetologica 36 (2): 175-189.

LYNCH. J.D. (1985a): A new s~ecies of Eleuther odactylus from western ~ i n a m a (Amphibia: Leptodactylidae). - Herpetologica 41 (4):

LYNCH, J.D. (198513): Mimetic and Non-Mimetic Populations of Eleutherodactylus gaigeae (DUNN) in lower Central America and Colombia (Amphibia: Anura, Leptodactylidae). - Studies on Neotropical Fauna and Environment 20 (4): 195-202.

LYNCH, J.D. (1996): Replacement names for three homonyms in the genus Eleu- therodactylus (Anura: Leptodactylidae). - Journal of Herpetology 30 (2): 278-280.

LYNCH, J.D. (1998): New species of Eleuthero- dactylus from the Cordillera Occidental of western Columbia with a synopsis of the distributions of species in western Columbia. - Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 22 (82): 117-148.

LYNCH, J.D. (1999~): Una approximacion a las cuebras ciegas de Colombia (Amphibia: Gymnophiona). - Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 23 (Special Suppl.): 317-337.

LYNCH, J.D. (2000): The relationships of an ensemble of Guatemalan and Mexican frogs (Eleutherodactylus: Leptodactylidae: Amphibia). - Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 24 (90): 129-156.

LYNCH, J.D. (2001): Three new rainfrogs of the Eleutherodactylus diastema group from Colombia and Panama. - Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 25: 287-297.

LYNCH, J.D. (2006): The tadpoles of frogs and toads found in the lowlands of northern Colombia. - Revista de la Academia Colombiana de Ciencias 30 (116): 443-457.

LYNCH, J.D. & ARDILA-ROBAYO, M.C. (1999): The Eleutherodactylus of the taeniatus complex in western Colombia: Taxonomy and distribution. - Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 23 (89): 615-624.

LYNCH, J.D. & MYERS, C.W. (1983): Frogs of the fitzingeri group of EIeutherodactylus in eastern Panama and Chocoan South America (Leptodactylidae). - Bulletin of the American Museum of Natural History 175 (5 ) : 481-572.

LYNCH, J.D. & RENJIFO, J.M. (1990): Two new toads (Bufonidae: Rhamphophryne) from the northern Andes of Colombia. - Journal of Herpetology 24 (4): 364-371.

Literature cited

LYNCH, J.D. & SUAREZ-MAYORGA, A.M. (2001): The distributions of the gladiator frogs (Hyla boans group) in Colombia, with comments on size variation and syrnpatry. - Caldasia 23 (2): 491-507.

LYNCH, J.F. & WAICE, D.B. (1975): Systematics of the Chiropterotriton bromeliacia group (Amphibia: Caudata), with description of two new species from Guatemala. - Natural History Museum of Los Angeles County Contributions in Science 265: 1 4 5 .

LYNCH, J.F. & WAKE, D.B. (1978): A new species of Chiropterotriton (Amphibia: Caudata) from Baja Verapaz, Guatemala, with comments on relationships among Central American members of the genus. - Natural History Museum of Los Angeles County Contributions in Science 294: 1-22.

LYNCH, J.F. & WAKE, D.B. (1989): Two new species of Pseudoeurycea (Amphibia: Caudata) from Oaxaca, Mexico. - Natural History Museum of Los Angeles County Contributions in Science 411: 11-22.

LYNCH, J.F. & WAKE, D.B. (1999): Pseudoeurycea parva Lynch and Wake. - Catalogue of American Amphibians and Reptiles 687: 1-2.

MALONE, J.H. (2004): Reproduction in three species of Smilisca from Costa Rica. - Journal of Herpetology 38 (1): 27-35.

MALONE, J.H. (2006): Ecology of the basin construction reproductive mode in Smilisca sordida (Anura: Hylidae). - Journal of Herpetology 40 (2): 230-239.

MANZANILLA, J., LA MARCA, E., VILLAREAL, 0 . & SANCHEZ (1998): Phrynohyas venulosa (Milki frog, Rana lechosa), antipredator device. - Herpetological Review 29 (1): 39-40.

MARQUIS, R.J., DONNELLY, M.A. & GUYER, C. (1986): Aggregations of calling males of Agalychnis calcarifer BOULENGER (Anura: Hylidae) in a Costa Rican lowland wet forest. - Biotropica 18 (2): 173-175.

MARSH, D.M. (2001): Fluctuations in amphibian populations: A meta-analysis. - Biological Conservation 101: 327-335.

MARTINS, M. (1989): Deimatic behavior in Pleurodema brachyops. - Journal of Herpetology 23 (3): 305-307.

MARTINEZ-CORONEL, M., RAMIREZ-BAUTISTA, A. & VIDAL-LOPEZ, R. (1995): Hyla bromeliacia (treefrog). - Herpetological Review 26 (2): 104-105.

MARTINS, M. (1988): Biologia reprodutiva de Leptodactylus fuscus em Boa Vista, Roraima (Amphibia, Anura). - Revista Brasileira de Biologia. 48: 969-977.

MCCALLUM, H. (2005): Inconclusiveness of chytridiomycosis as the agent in widespread frog declines. - Conservation Biology 19: 1421-1430

MCCRANIE, J.R. (1996a): Geographic distribution: Bolitoglossa porrasorum. - Herpeto- logical Review 27 (1): 28.

MCCRANIE, J.R. (1996b): Geographic distribution: Nototriton barbouri. - Herpetological Review 27 (1): 28.

MCCRANIE, J.R. (1996~): Geographic distribution: Oedipina gephyra. - Herpetological Review 27 (1): 29.

MCCRANIE, J.R. (2006): New species of Oedipina (Amphibia: Caudata) from Parque Nacional El Cusuco, northwestern Honduras. - Journal of Herpetology 40 (3): 291-293.

MCCRANIE, J.R. & F.E. C A S T ~ D A (2005): The herpetofauna of Parque Nacional Pico Bonito, Honduras. - Phyllomedusa 4 (1): 3-16.

MCCRANIE, J.R. & F.E. CASTAREDA (2006): A new species of hylid frog from northwestern Honduras. - Herpetologica 62: 318-323.

MCCRANIE, J.R. & C A S T ~ E D A , F.E. (2007): Guia de Campo de 10s anfibios de Honduras. - Salt Lake City. 304 pp.

MCCRANIE, J.R., COLLART, J.R., CASTEREDA, F.E. & SOLIS, J.M. (2008): Geographic distribution: Eleutherodactylus (Euhyas) plani- rostris. - Herpetological Review 39: 362.

MCCRANIE, J.R. & CRUZ, G.A. (1996): A new species of salamander of the Bolitoglossa dunni group (Caudata: Plethodontidae) from the Sierra de Agalta, Honduras. - Caribbean Journal of Science 32 (2): 195-200.

MCCRANIE, J.R., CRUZ, G.A. & RIVERA, D.M. (2001): Geographic distribution: Oedipina taylori. - Herpetological Review 32: 54.

MCCRANIE, J.R., ESPINAL, M.R. & WILSON, L.D. (2005): New species of montane salamander of the Bolitoglossa dunni group from northern Comayagua, Honduras (Urodela: Plethodontidae). - Journal of Herpetology 39: 108-112.

MCCRANIE, J.R. & KOHLER, G. (1999a): A new species of salamander of the Bolitoglossa dunni group from Cerro El Pital, Honduras and El Salvador (Amphibia, Caudata, Plethodontidae). - Senckenbergiana Biologica 78 (1-2): 225-229.

MCCRANIE, J.R. & KOHLER, G. (1999b): A new species of rainfrog of the Eleutherodactylus gollmeri group from western Honduras (Amphibia, Anura, Leptodactylidae). -

Senckenbergiana Biologica 79 (1): 83-87. MCCRANIE, J.R. & KOHLER, G. (1999~): Errata.

Correction for: J.R. MCCRANIE & G. KOHLER

Literature cited

(1999): A new species of salamander of the Bolitoglossa dunni group from Cerro El Pital, Honduras and El Salvador (Amphibia, Caudata, Plethodontidae). Senckenbergiana Biologica 79 (1): 107.

MCCRANIE, J.R. & KOHLER, G. (2000): Notes on the type series of Bufo valliceps Wiegmann, with the designation of a lectotype. - Southwestern Naturalist 45 (1): 71-74.

MCCRANIE, J.R., NICHOLSON, K.E. & CASTAREDA, F.E. (2002): Geographic distribution. Eleutherodactylus diastema (NCN). - Herpetological Review 33 (3): 220.

MCCRANTE, J.R., SAVAGE, J.M. & WILSON, L.D. (1989): Description of two new species of the Eleutherodactylus milesi group (Amphibia: Anura: Leptodactylidae) from northern Honduras. - Proceedings of the Biological Society of Washington 102 (2): 483490.

MCCRANIE, J. R. & SMITH, E.N. 2006. A new species of Craugastor (Anura: Lepto- dactylidae) of the alfredi group from western Honduras. - Herpetologica 62: 183-190.

MCCRANIE, J.R., TOWNSEND, J.H. & WILSON, L.D. (2003): Hyla miliaria (Anura: Hylidae) in Honduras, with notes on calling site. - Caribbean Journal of Science 39 (3): 398-399.

MCCRANIE, J.R., VALDES ORELLANA, L. & HIMES, J.G. (2010): Rediscovery of two Honduran endemic streamside frogs, Craugastor emleni (DUNN) and Craugastor stadelmani (SCHMIDT). - Froglog 94: 12-15.

MCCRANIE, J.R., VIEITES, D.R. & WAKE, D.B. (2008): Description of a new divergent lineage and three new species of Honduran salamanders of the genus Oedipina (Caudata, Plethodontidae). - Zootaxa 1930: 1-17.

MCCRANIE, J.R., WAKE, D.B. & WILSON, L.D. (1996): The taxonomic status of Bolitoglossa schmidti, with comments on the biology of the Mesoamerican salamander Bolitoglossa dofleini (Caudata: Plethodontidae). - Caribbean Journal of Science 32 (4): 395-398.

MCCRANIE, J.R. & WILSON, L.D. (1981): A new hylid frog of the genus Plectrohyla from a cloud forest in Honduras. - Occasional Papers of the Museum of Natural History of the University of Kansas 92: 1-7.

MCCRANIE, J.R. & WILSON, L.D. (1986): A new species of red-eyed treefrog of the Hyla uranochroa group (Anura: Hylidae) from northern Honduras. - Proceedings of the Biological Society of Washington 99 (1): 51-55.

MCCRANIE, J.R. & WILSON, L.D. (1990): A description of the tadpole of Hyla salvavida

(Anura: Hylidae). - Copeia 1990 (4): 1152-1154.

MCCRANIE, J.R. & WILSON, L.D. (1992): Life history notes: Nototriton barbouri. Repro- duction. - Herpetological Review 23: 115-116.

MCCRANIE, J.R. & WILSON, L.D. (1993a): A review of the Bolitoglossa dunni group (Amphibia: Caudata) from Honduras with the description of three new species. - Herpetologica 49: 1-15.

MCCRANIE, J.R. & WILSON, L.D. (1993b): Taxonomic changes associated with the names Hyla spinipollex SCHMIDT and Ptychohyla merazi Wilson and MCCRANIE (Anura: Hylidae). - Southwestern Naturalist 38 (2): 100-104.

MCCRANIE, J.R. & WILSON, L.D. (1995a): A new species of salamander of the genus Bolitoglossa (Caudata: Plethodontidae) from Parque Nacional El Cusuco, Honduras. - Journal of Herpetology 29 (3): 447452.

MCCRANIE, J.R. & WILSON, L.D. (199513): A new species of salamander of the Bolitoglossa dunni group (Caudata: Plethodontidae) from northern Honduras. - Herpetologica 51 (2): 131-140.

MCCRANIE, J.R. & WILSON, L.D. (1996): A new species of salamander of the genus Nototriton (Caudata: Plethodontidae) from Montana de Santa Barbara, Honduras. - Southwestern Naturalist 41 (2): 111-115.

MCCRANIE, J.R. & WILSON, L.D. (1997a): A review of the Eleutherodactylus milesi-like frogs (Anura, Leptodactylidae) from Honduras with the description of four new species. - Mytes 14 (4): 147-174.

M c C m , J.R. & WILSON, L.D. (1997133: 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 (3): 366-372.

MCCRANIE, J.R. & WILSON, L.D. (1997~): Two new species of centrolenid frogs of the genus Hyalinobatrachium from eastern Honduras. - Journal of Herpetology 31 (1): 10-16.

MCCRANIE, J.R. & WILSON, L.D. (1998): Specific status of the Honduran frogs formerly referred to Plectrohyla teuchestes (Anura: Hylidae). - Journal of Herpetology 32 (1): 96-101.

MCCRANIE, J.R. & WILSON, L.D. (1999a): Description of a new species of Plectrol~yla from Cerro Celaque, Honduras, formerly referred to Plectrohyla glandulosa (Amphibia, Anura, Hylidae). - Sencken- bergiana Biologica 78 (112): 231-236.

Literature cited

MCCRANIE, J.R. & WILSON, L.D. (1999b): Two new species of the Eleutherodactylus rugulosus group (Amphibia: Anura: Lepto- dactylidae) from Honduras. - Proceedings of the Biological Society of Washington 112: 515-522.

MCCRANIE, J.R. & WILSON, L.D. (2000): A new species of high-crested toad of the Bufo valliceps group from north-central Honduras. - Journal of Herpetology 34 (1): 21-31.

MCCRANIE, J.R. & WILSON, L.D. (2002): The Amphibians of Honduras.- Society for the Study of Amphibians and Reptiles. Ithaca, New York, USA. 625 pp.

MCCRANIE, J.R. & WILSON, L.D. (2003a): Nototriton limnospecta tor. - Catalogue of American Amphibians and Reptiles 763: 1-2.

MCCRANIE, J.R. & WILSON, L.D. (2003b): Eleutherodactylus operosus Savage, McCranie, and Wilson, 1999: A synonym of Eleutherodactylus cerasinus (Cope, "1876 [1875]). - Journal of Herpetology 37: 408- 409.

MCCRANIE, J.R., WILSON, L.D. & KOHLER, G. (2005): The Amphibians and Reptiles of the Bay Islands and Cayos Cochinos, Honduras. - Bibliomania, Salt Lake City, Utah.

MCCRANIE, J.R., WILSON, L.D. & POLISAR, J. (1998): Another new montane salamander (Amphibia: Caudata: Plethodontidae) from Parque Nacional Santa Barbara, Honduras. - Herpetologica 54 (4): 455461.

MCCRANIE, J.R., WILSON, L.D. & Porras, L. (1980): A new species of Leptodactylus from the cloud forests of Honduras. - Journal of Herpetology 14 (4): 361-367.

MCCRANIE, J.R., WILSON, L.D. & TOWNSEND, J. H. (2003): Agalychnis callidryas (red-eyed treefrog): Reproduction. - Herpetological Review 34 (1): 43.

MCCRANIE, J.R., WILSON, L.D. & WILLIAMS, K.L. (1986): The tadpole of Leptodactylus silvanimbus, with comments on the relationships of the species. J o u r n a l of Herpetology 20: 560-562.

MCCRANIE, J.R., WILSON, L.D. & WILLIAMS, K.L. (1987): Plectrohyla guatemalensis. Repro- duction. - Herpetological Review 18: 72.

MCCRANIE, J.R., WILSON, L.D. & WILLIAMS, K.L. (1989): A new genus and species of toad (Anura: Bufonidae) with an extraordinary stream-adapted tadpole from northern Honduras. - Occasional Papers of the Museum of Natural History of the University of Kansas 129: 1-18.

MCCRANIE, J.R., WILSON, L.D. & WILLIAMS, K.L. (1993): A new species of Oedipina

(Amphibia: Caudata: Plethodontidae) from northern Honduras. - Proceedings of the Biological Society of Washington 106 (2): 385-389.

MCCRANIE, J.R., WILSON, L.D. & WILLIAMS, K.L. (1993a): Description of the tadpole of Hyla catracha (Anura: Hylidae). - Caribbean Journal of Science 29 (3): 256-258.

MCCRANIE, J.R., WILSON, L.D. & WILLIAMS, K.L. (1993b): New species of tree frogs of the genus Hyla (Anura: Hylidae) from northern Honduras. - Copeia 1993 (4): 1057-1062.

MCCRANIE, J.R., WILSON, L.D. & WILLIAMS, K.L. (1999): Cochranella granulosa. - Herpetological Review 30 (2): 106.

MCDIARMID, R.W. (1968): Populational variation in the frog genus Phrynohyas FITZ~NGER in Middle America. - Contributions in Science of the Natural History Museum of Los Angeles County, 134, 1-25.

MCDIARM~, R.W. (1975): Glass frog romance along a tropical stream. - Terra 13 (4): 14-18.

MCDIARMID, R.W. (1983): Centrolenella fleischmanni (Ranita de Vidrio, Glass Frog). - In: JANZEN, D.H. (ed.) Costa Rican Natural History. - Chicago & London (Univ. Chicago Press): 389-390.

MCDIARMID, R.W. & ADLER, K. (1974): Notes on the territorial and vocal behaviour of Neotropical frogs of the genus Centrolenella. - Herpetologica 30 (1): 75-78.

MCDIARMID, R.W. & ALTIG, R. (eds.) (1999): Tadpoles. The biology of anuran larvae. - The University of Chicago Press. Chicago. 444 pp.

MCDIARMID, R.W. & FOSTER, M.S. (1981): Breeding habits of the toad Bufo coccifer in Costa Rica, with a description of the tadpole. - Southwestern Naturalist 26 (4): 353-363.

MCGEHEE, R.T., REAMS, R. & BROWN, M.E. (2001): Bufo valliceps (Gulf Coast toad): Diet. - Herpetological Review 32 (2): 101-102.

MEBS, D., POGODA, W., BATISTA, A,, PONCE, M., KOELER, G. & KAUERT, G. (2008): Variability of alkaloid profiles in Oophaga pumilio (Amphibia: Anura: Dendrobatidae) from western Panama and southern Nicaragua. - Salamandra 44 (4): 241-247.

MEEGASKUMBURA, M., BOSSWT, F., PETHNAGODA, R., NIANAMENDRA-ARACHCHI, K., BAHIR, M., MILINKOVITCH, M.C. & SCHNEIDER, C.J. (2002): Sri Lanka: An amphibian hot spot. - Sience 298: 379.

MENDELSON 111, J.R. (1994): A new species of toad (Anura: Bufonidae) from the lowlands of eastern Guatemala. - Occasional Papers

Literature cited

of the Museum of Natural History of the University of Kansas 166: 1-21.

MENDELSON 111, J.R. (1997): A new species of toad (Anura: Bufonidae) from the Pacific highlands of Guatemala and southern Mexico, with comments on the status of Bufo valliceps macrocristatus. - Herpetologica 53 (1): 14-30.

MENDELSON 111, J.R. (1998): Geographic variation in Bufo valliceps group (Anura: Bufonidae), a widespread toad in the United States and Middle America. - Scientific Papers of the Natural History Museum of the University of Kansas 8: 1-12.

MENDELSON 111, J.R. (2001): A review of the Guatemalan toad Bufo ibarrai (Anura: Bufonidae), with distributional and taxonomic comments of Bufo valliceps and Bufo coccifer. - Pp. 10-19 in: JOHNSON, J.D., WEBB, R.G. & FLORES-VILLELA, 0 . (eds.): Mesoamerican Herpetology: Systematics, Natural History, and Conservation. The University of Texas a t El Paso. El Paso, Texas.

MENDELSON 111, J.R., BRODIE JR., E.D., MALONE, J.H., ACEVEDO, M.E., BAKER, M.A., SMATRESK, N.J. & CAMPBELL, J.A. (2004): Factors associated with the catastropic decline of a cloud- forest frog fauna in Guatemala. -

International Journal of Tropical Biology. 52 (4): 991-1000.

MENDELSON 111, J.R. & CAMPBELL, J.A. (1994): Two new species of the Hyla sumichrasti group (Amphibia: Anura: Hylidae) from Mexico. - Proceedings of the Biological Society of Washington 107 (2): 398-409.

MENDELSON 111, J.R. & CAMPBELL, J.A. (1999): The taxonomic status of populations referred to Hyla chaneque in southern Mexico, with the description of a new treefrog from Oaxaca. - Journal of Herpetology 33 (1): 80-86.

MENDELSON 111, J.R. & MULCAHY, D.G. (2010): A new species of toad (Bufonidae: Incilius) from central Panama. - Zootaxa 2396: 61-68.

MENDELSON 111, J.R., SAVAGE, J.M., GRIFFITH, E., Ross, H., KUBICKI, B. & GAGLIARDO, R. (2007): Spectacular new gliding species of Ecnomiollyla (Anura: Hylidae) from central Panama. - Journal of Herpetology 42 (4): 750-759.

MENDELSON 111, J.R., DA SILVA, H.R. & MAGLIA, A.M. (2000): Phylogenetic relationships among marsupial frog genera (Anura: Hylidae: Hemiphractinae) based on evidence from morphology and natural history. - Zoological Journal of the Linnean Society 128: 1-24.

MENDELSON 111, J.R., USTACH, P.C. & NIETO MONTES DE OCA, A. (1999): Description of the tadpole of Bufo tutelarius, natural history notes on the Bufo valliceps group, and a key to the tadpoles of the group. - Journal of Herpetology 33 (2): 324-328.

MENDELSON 111, J.R., WILLIAMS, B.L., SHEIL, C.A. & MULCAHY, D.G. (2005): Systematics of the Bufo coccifer complex (Anura: Bufonidae) of Mesoamerica. - Scientific Papers of the Natural History Museum of the University of Kansas 38: 1-27.

MENDOZA QUIJANO, F., SANTOS-BARRERA, G. & PACHECO-RODR~GUEZ, J. (2002): Eleuthero- dactylus fitzingeri (common rain frog) clutch size and parental care. - Herpetological Review 33 (2): 125.

MERTENS, R. (1952a): Die Arnphibien und Reptilien von El Salvador. - Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 487: 1-120.

MERTENS, R. (1952b): Zur Kenntnis der Amphibien-Fauna von El Salvador. -

Senckenbergiana Biologica 33 (113): 169- 171.

~ S H ~ K A Jr, W.E. (2001): The Cuban treefrog in Florida: Life history of a successful coloniz- ing species. - University Press of Florida. Gainesville, Florida.

MEYER, J.R. & FOSTER, C.F. (1996): A guide to the frogs and toads of Belize. - Krieger Publishing Company, Malabar, Florida. 80 PP.

MITTLEMAN, M.B. (1950): Status of the name Hyla septentrionalis. - Herpetologica 6: 24-26.

MNAMOTO, M.M. (1983): Biochemical variation in the frog Eleutherodactylus bransfordii: geographic patterns and cryptic species. - Systematic Zoology 32 (1): 43-51.

MNAMOTO, M.M. (1984): Central American frogs allied to Eleutherodactylus cruentus: allozyme and morphological data. - Journal of Herpetology 18 (3): 256-263.

MNAMOTO, M.M. (1986): Phylogenetic relationships and systematics of the Eleutherodactylus fitzingeri group (Anura, Leptodactylidae). - Copeia 1986 (2): 503-511.

MIYAMOTO, M.M. & TENNANT, M.R. (1984): Phylogenetic relationships of the lower Central American rain frog Eleuthero- dactylus melanostictus. - Copeia 1984 (3): 765-768.

MOORE, T.E., NUSSBAUM, R.A. & MOCKFORD, E.L. (1983): Caeciliidae in Amphibia and Insecta (Psocoptera): proposals to remove the homonymy. Z.N. (S.) 2333. - Bulletin of Zoological Nomenclature 40 (2): 124-128.

Literature cited

MOREHOUSE, E.A., JAMES, T.Y., GANLEY, A.R., VILGALYS, R., BERGER, L., MURPHY, P.J. & LONGCORE, J.E. (2003): Multilocus sequence typing suggests the chytrid pathogen of amphibians is a recently emerged clone. - Molecular Ecology 12: 395-403.

MULLER, M.J. (1996): Handbuch ausgewahlter Klimastationen der Erde. 5. Ed. - Trier (For- schungsstelle Bodenerosion der Universitat Trier): 400 pp.

MUELLER, R. L., MACEY, J.R., JAEKEL, M., WAKE, D.B. & BOORE J.L. (2004): Morphological homoplasy, life history evolution and historical biogeography of plethodontid salamanders inferred from complete mitochondria1 genomes. - Proceedings of the National Academy of Sciences of the United States of America 101: 13820-13825.

MULCAHY, D.G. & MENDELSON 111, J.R. (2000): Phylogeography and speciation of the morphologically variable, widespread species Bufo valliceps, based on molecular evidence from mtDNA. - Molecular Biology and Evolution 17 (2): 173-189.

MULCAHY, D.G., M0rri11, B.H. & MENDELSON 111, J.R. (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.

MYERS, C.W. (1966): The distribution and behavior of a tropical horned frog, Cerathyla panamensis STEJNEGER. - Herpetologiea 22: 68-71.

MYERS, C.W. (1969): Ecological geography of cloud forest in Panama. - American Museum Novitates 2396: 1-52.

MYERS, C.W. (1982): Spotted poison frogs: Descriptions of three new Dendrobates from western Amazonia, and resurrection of a lost species from Chiriqui. - American Museum Novitates 2721: 1-12.

MYERS, C.W. (1987): New generic names for some Neotropical poison frogs (Dendro- batidae). - Avulsos de Zoologia, Museu de Zoologia da Universidade de Sao Paulo. 36 (25): 301-306.

MYERS, C.W., DALY, J.W., GARRAFFO, H.M., WISNIESKI, A. & COVER, J.F. JR (1995): Discovery of the Costa Rican poison frog Dendrobates granuliferus in sympatry with Dendrobates pumilio, and comments on taxonomic use of skin alkaloids. - American Museum Novitates 3144: 1-21.

MYERS, C.W., DALY, J.W. & MALKIN, B. (1978): A dangerously toxic new frog (Phyllobates) used by Embera Indians of Western Colombia, with discussion of blowgun fabri- cation and dart poisoning. - Bulletin of the

American Museum of Natural History 161: 307-366.

MYERS, C.W., DALY, J.W. & MAF~TINEZ, V. (1984): An arboreal poison frog (Dendrobates) from western Panama. - American Museum Novitates 2783: 1-20.

MYERS, C.W. & DUELLMAN, W.E. (1982): A new species of Hyla from Cerro Colorado, and other tree EFog records and geographical notes from western Panama. - American Museum Novitates 2752: 1-32.

MYERS, C.W. & GRANT,T. (2009): AnomaIoglossus confisus, a new Ecuadorian frog formerly masquerading as "Colostethus" chocoensis (Dendrobatoidea: Aromobatidae). -

American Museum Novitates 3659: 1-12. MYERS, C.W. & LYNCH, J.D. (1997.

Eleutherodactylus laticorpus, a peculiar new frog from the Cerro Tacarcuna area, Panamanian-Colombian frontier. -

American Museum Novitates 3196: 1-12. MYERS, C.W. PAOLLLLO-O.,A. & DALY J.W. (1991):

Discovery of a defensively malodorous and nocturnal frog in the family Dendrobatidae: Phylogenetic significance of a new genus and species from Venezuelan Andes. - American Museum Novitates 3002: 1-33.

MYERS, G.S. (1962): The American leptodactylid frog genera Eleutherodactylus, Hylodes (= Elosia), and Caudiverbera (= Calypto- cephalus). - Copeia 1962: 195-202.

NARVAES, P. & TREFAUT RODRIGUES, M. (2009): Taxonomic revision of Rhinella granulosa species group (Amphibia, Anura, Bufonidae), with a description of a new species. -Arquivos de Zoologia 40 (1): 1-73.

NASCIMENTO, L.B., CARAMASCHI, U. & CRUZ, C.A.G. (2005): Taxonomic review of the species groups of the genus Physalameus FITZINGER, 1826 with revalidation of the genera Engystomops JIMENEZ-DE-LA-ESPADA, 1872 and Eupemphix STEINDACHNER, 1836 (Amphibia, Anura, Leptodactylidae). - Arquivos do Museu Nacional63: 297-320.

NELSON, C. (1963): Gastrophryne elegans (BOULENGER): Elegant Narrow-mouthed Toad. - Catalogue of American Amphibians and Reptiles 121: 1-2.

NELSON, C. (1972a): Gastrophryne usta. Two- spaded narrow-mouthed toad. - Catalogue of American Amphibians and Reptiles 123: 1-2.

NELSON, C.E. (197213): Systematic status of the North American microhylid genus Gastrophryne. - Journal of Herpetology 6: 111-137.

NELSON, C.E. (1972~): Distribution and biology of Chiasn~ocleis panamensis (Amphibia: Microhylidae). -Copeia 1972: 895-898.

Literature cited

NELSON, C.E. (1973a): Systematics of the Middle American upland populations of Hypo- pachus (Anura: Microhylidae). - Herpeto- logica 29: 6-17.

NELSON, C.E. (197313): Gastrophryne. - Cata- logue of American Amphibians and Reptiles 134: 1-2.

NELSON, C. E. (1973~): Gastrophryne pictiventris (COPE): Southern Narrow-mouthed Toad. - Catalogue of American Amphibians and Reptiles 135: 1-2.

NELSON, C.E. (1974): Further studies on the systematics of Hypopachus (Anura: Micro- hylidae). - Herpetologica 30: 250-275.

NELSON, C.E. & ALTIG, R. (1972): Tadpoles of the microhylids of Gastrophryne elegans and G. usta. - Herpetologica 28 (4): 381-383.

NELSON, C.E. & NICKERSON, M.A. (1966): Notes on some Mexican and Central American amphibians and reptiles. - Southwestern Naturalist 11 (1): 128-131.

NICHOLS, D.K., LAMIRANDE, E.W., PESSIER, A.P. & LONGCORE, J.E. (2001): Experimental trans- mission of cutaneous chytridiomycosis in dendrobatid frogs. - Journal of Wildlife Diseases 37: 1-11.

NIETO, M.J. (1999): Estudio preliminar de las especies del gknero Scinax (Amphibia: Anura: Hylidae) en Colombia. - Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 23 (Special Suppl.): 339-346.

NOBLE, G.K. (1926): The pectoral girdle of the brachycephalid frogs. - American Museum Novitates 230: 1-14.

NOVAK, R.M. & ROBINSON, D.C. (1975): Observa- tions on the reproduction and ecology of the tropical montane toad, Bufo holdridgei TAYLOR in Costa Rica. - Revista de Biologia Tropical 23: 213-237.

NUSSBAUM, R.A. (1979): The status of Copeotyphlinus syntremus (Amphibia: Gymnophiona). -Journal of Herpetology 13: 121-123.

NUSSBAUM, R.A. (1988): On the status of Copeo typhlin us syn trem us, Gymnopis oligozona, and Minascaecilia sartoria (Gymno- phiona, Caeciliidae): a comedy of errors. - Copeia 1988 (4): 921-928.

NUSSBAUM, R.A. & HOOGM~ED, M.S. (1979): Surinam caecilians, with notes on Rhina- trema bivittatum and the description of a new species of Microcaecilia (Amphibia, Gymnophiona). - Zoologische Mededelingen 54 (14): 217-235.

OGIELSKA, M. (ed.) (2009): Reproduction of amphibians. - Science Publishers. Enfield (NH). 422 pp.

O'NEILL, E.M. & MENDELSON, J.R. (2004): Taxonomy of Costa Rican toads referred to Bufo melanochlorus COPE, with the description of a new species. - Journal of Herpe- tology 38 (4): 487494.

ORTEGA-ANDRADE, H.M. (2008): Agalychnis spurrelli BOULENGER (Anura, Hylidae): variacibn, distribuci6n y sinomia. - PapCis Avulsos de Zoologia 48: 103-117.

OVASKA, K. & Rmn, A.S. (2001): Courtship and reproductive behavior of the frog Eleuthero- dactylus diastema (Anura: Leptodactylidae) in Gamboa, Panama. - Journal of Herpetology 35344-50.

PADIAL, J.M., CASTROVIEJO-FISHER, S. & DE LA RIVA, I. (2009): The phylogenetic relationships of Yunganastes revisited (Anura: Terrarana). - Molecular Phylogenetics and Evolution 52: 911-915

PAPENFUSS, T.J. & WAKE, D.B. (1987): Two new species of pletodontid salamanders (genus Nototriton) from Mexico. - Acta Zoologica Mexicana Nueva Serie 21: 1-16.

PAPENFUSS, T.J., WAKE, D.B. & ADLER, K. (1983): Salamanders of the genus Bolitoglossa from the Sierra Madre del Sur of southern Mexico. - Journal of Herpetology 17: 295-307.

PARKER, H.W. (1934): A Monograph of the Frogs of the Family Microhylidae. - Trustees of the British Museum. London. 208 pp.

PARRA-OLEA, G. (2002): Phylogenetic relationships of the salamander of the genus Pseudoeurycea (Caudata: Plethodontidae). - Molecular Phylogenetics and Evolution 22: 234-246.

PARRA-OLEA, G. & GARCIA-PARIS, M. (1998): Amphibia: Caudata: Plethodontidae: Bolito- glossa hartwegi Wake and Brame. - Cata- logue of American Amphibians and Reptiles 641: 1-2.

PARRA-OLEA, G., GARC~A-PARIS, M. & WAKE, D.B. (1999): Status of some populations of Mexican salamanders. - Revista de Biologia Tropical 47: 217-223.

PARRA-OLEA, G., M. GARC~A-PARIS & WAKE, D.B. (2002): Phylogenetic relationships among the salamanders of the Bolitoglossa macrinii species group (Amphibia: Plethodontidae), with descriptions of two new species from Oaxaca (Mexico). - Journal of Herpetology 36: 356-366.

PARRA-OLEA, G., M. GARCU-PARIS & WAKE, D.B. (2004): Molecular diversification of salamanders of the tropical American genus Bolitoglossa (Caudata: Plethodontidae) and its evolutionary and biogeographic implications. - Biological Journal of the Linnean Society 81: 325-346.

Literature cited

PARRA-OLEA, G., PAPENFUSS, T.J. & WAKE, D.B. (2001): New species of lungless salamanders of the genus Pseudoeurycea (Amphibia: Caudata: Plethodontidae) from Veracruz, Mexico. - Scientific Papers of the Natural History Museum of the University of Kansas 20: 1-9.

PASMANS, F., ZWART, P. & HYATT, A.D. (2004): Chytridiomycosis in the Central American bolitoglossine salamander (Bolitoglossa dofleinil. - Veterinary Record 154: 153.

PAULY, G.B., HLLLIS, D.M. & CANNATELLA, D.C. (2004): The history of a Nearctic coloniza- tion: Molecular phylogenetics and biogeography of the Nearctic toads (Bufo). -Evolution 58: 2517-2535.

PAULY, G.B., HILLIS, D.M. & CANNATELLA, D.C. (2009): Taxonomic freedom and the role of official lists of species names. -Herpeto- logica 65: 115-128.

PECHMANN, J.H.K & WILBUR, H.M. (1994): Putting declining amphibian populations in perspective: Natural fluctuations and human impacts. - Herpetologica 50: 65-84.

PEREZ-KIGAREDA, G. (1981): Geographic distribution. Hyla chaneque. - Herpetological Review 12 (2): 64.

PESSIER, A.P., NICHOLS, D.K., LONGCORE, J.E. & FULLER, M.S. (1999): Cutaneous chytri- dionlycosis in poison dart frogs (Dendro- bates spp.) and White's tree frogs (Litoria caerulea). - Journal of Veterinary Diagnostic Investigation 11: 194-199.

PETERS, W.C.H. (1862): ijber die Batrachier- Gattung Hemiphractus. - Monatsberichte der Koniglich Preussischen Akademie der Wissenschaften zu Berlin 1862: 144-152.

PETERS, W.C.H. (1863a): Uber eine neue Schlangen-Gattung, Styporhynchus, und verschiedene andere Amphibien des zoologischen Museums. - Monatsberichte der Koniglich Preussischen Akademie der Wissenschaften zu Berlin 1863: 399-413.

PETERS, W.C.H. (1863b): Mittheilungen iiber neue Batrachier. - Monatsberichte der Koniglich Preussischen Akademie der Wissenschaften zu Berlin 1863: 445-470.

PETERS, W.C.H. (1873): Uber eine neue Schildkrotenart, Cinosternon Effeldtii und einiger andere neue oder weniger bekannte Amphibien. - Monatsberichte der Preussischen Akademie der Wissenschaften zu Berlin 1873: 301-618.

PETERS, W.C.H. (1874): ijber neue Amphibien ( Gymnopis, Siphonops, Polypeda tes, Rhacophorus, Hyla, Cyclodus, Euprepes, Cyclemmys). - Monatsberichte der Konig- lich Preussischen Akademie der Wissen- schaften zu Berlin 1874: 616-624.

PETERS, W.C.H. (1879): Mittheilung iiber neue Amphibien des Kgl. Zoologischen Museums (Euprepes, Acontias, Typhlops, Zamenis, Spilotes, Oedipus). - Monatsberichte der Koniglich Preussischen Akademie der Wissenschaften zu Berlin 1879: 773-779.

PETERS, W.C.H. (1882a): Herr W. Peters legte eine neue Art der urodelen Batrachier, Oedipus yucatanus, aus Yucatan (CentralameAca) vor. - Sitzungs-Bericht der Gesellschaft naturforschender Freunde zu Berlin 1882:145-148.

PETERS, W.C.H. (1882b): Herr W. Peters schlug vor den Namen der Batrachiergattung Hylonomus in Hyloscirtus zu andern und legte zwei neue Arten von Schlangen, Microsoma notatum und Liophis Ygraecum. - Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1882: 127-129.

PETHTYAGODA, R., MEEGASKUMBURA, M., MANAMENDRA-ARACHCHI, K. & SCHNEIDER, C.J. (2008): Essay 1.2. Amphibian diversity and the case of Sri Lanka's burgeoning inventory. - Pp. 18-19 in STUART, S., HOFFMANN, M., CHANSON, J., COX, N., BERRIDGE, R., RAMANI P. & YOUNG, B. (eds.): Threatened Amphibians of the World. - Lynx Edicions. Barcelona, Spain. 758 pp.

PFENNIG, D.W., LOEB, M.L.G. & COLLINS, J.P. (1991): Pathogens as a factor limiting the spread of cannibalism in tiger salamanders. - Oecologia 88: 161-166.

PINO, E. del(1980): Morphology of the pouch and incubatory integument in marsupial frogs (Hylidae). - Copeia 1980: 10-17.

PONSSA, M.L. (2008): Cladistic analysis and osteological descriptions of the frog species in the Leptodactylus fuscus species group (Anura, Leptodactylidae). -Journal of Zoological Systematics and Evolutionary Research 46: 249-266.

PORRAS, L. & WILSON, L.D. (1987): A new species of Hyla from the highlands of Honduras and El Salvador. - Copeia 1987 (2): 478-482.

PORTER, K. (1963): Distribution and taxonomic situation of seven species of Mexican Bufo. - Herpetologica 19 (4): 229-247.

POUNDS, J.A., BUSTAMANTE, M.R., COLOMA, L.A., CONSUEGRA, J.A., FOGDEN, M.P.L., FOSTER, P.N., LA MARCA, E., MASTERS, K.L., MERINO- VITERI, A., PUSCHENDORF, R., RON, S.R., SANCHEZ-AZOFEIFA, G.A., STILL, C.J. & YOUNG, B.E. (2006): Widespread amphibian extinctions from epidemic disease driven by global warming. - Nature 439: 161-167.

POUNDS, J.A., BUSTAMANTE, M.R., COLOMA, L.A., CONSUEGRA, J.A., FOGDEN, M.P.L., FOSTER, P.N., LA MARCA, E., MASTERS, K.L., MERINO-

Literature cited

VITERI, A., PUSCHENDORF, R., RON, S.R., SANCHEZ-AZOFEIFA, G.A., STILL, C.J. & YOUNG, B.E. (2007): Global warming and amphibian losses: The proximate cause of frog declines? (Reply). - Nature 447: E5-E6.

POUNDS, J.A., CARNAVAL, A.C., PUSCHENDORF, R., HADDAD, C.F.B. & MASTERS, K.L. (2006): Responding to amphibian loss. - Science 314: 1541-1542.

POUNDS, J. A. & CRW, M. L. (1987): Harlequin Frogs along a tropical montane stream: Aggregation and the risk of predation by frog-eating flies. - Biotropica 19 (4): 306-309.

POUNDS, J. A. & CRUMP, M. L. (1989): Temporal variation in the dispersion of a tropical anuran. - Copeia 1989 (1): 209-211.

POUNDS, J. A. & CRUMP, M. L. (1994): Amphibian declines and climate disturbance: The case of the Golden Toad and the Harlequin Frog. - Conservation Biology 8 (1): 72-85.

POUNDS, J.A., FOGDEN, M.P.L. & CAMPBELL, J.H. (1999): Biological response to climate change on a tropical mountain. - Nature 398 (6728): 611-615.

POUNDS, J.A., FOGDEN, M.P.L., SAVAGE, J.M. & GORMAN, G.C. (1997): Tests of null models for amphibian declines on a tropical mountain. - Conservation Biology 11 (6): 1307-1322.

PRAMUK, J.B. (2006): Phylogeny of South American Bufo (Anura: Bufonidae) inferred from combined evidence. - Zoological Journal of the Linnean Society 146: 407452.

P a m , J.B. & LEHR, E. (2005): Taxonomic status of Atelophryniscus chrysophorus McCranie, Wilson, and Williams, 1989 (Anura: Bufonidae) inferred from phylogeny. -Journal of Herpetology 39: 610-618.

PRAMUK, J.B., ROBERTSON, T., SITES, J.W., Jr. & NOONAN, B.P. (2008): Around the world in 10 million years: biogeography of the nearly cosmopolitan true toads (Anura: Bufonidae). - Global Ecology and Biogeography 17: 72-83.

PROHL, H., KOSHY, R.A., MUELLER, U.G., R.wD, A.S. & RYAN, M.J. (2006): Geographic variation of genetic and behavioral traits in northern and southern tungara frogs. - Evolution 60: 1669-1679. .

PROY, C. (1993a): Beobachtungen zur Biologie und Erfahrungen bei der Erhaltung und Nachzucht von Agalychnis annae (DUELL- MAN, 1963). - Herpetofauna 15 (84): 27-35.

PROY, C. (1993b): Erste Terrarienbeobachtungen zur Fortpflanzung von Hyla legleri T A ~ O R , 1958 aus Costa Rica (Anura: Hylidae). - Herpetozoa 6 (3-4): 105-111.

PROY, C. (2000): Unterschiedliche Temperatur- verlaufe und kiinstliche Beregnung zur Stimulierung der Paarung bei zwei Greif- frijschen (Phyllomedusa lemur and Aga- lychnis annae). - Herpetofauna 22 (128): 29-34.

PROY, C.H. (1992): Zur Biologie von Agalychnis saltator TAYLOR, 1955 (Anura: Hylidae). - Herpetozoa 5 (3-4): 99-107.

PURVIS, A. & HECTOR, A. (2000): Getting the measure of biodiversity. -Nature 405: 212-219.

PUSCHENDORF, R. (2003): Atelopus varius (harlequin frog): Fungal infection. - Herpeto- logical Review 34: 355.

PUSCHENDORF, R. & CRAWFORD, A.J. (2003): Eleutherodactylus stejnegerianus (Stej- neger's dirt frog): Urban habitat. - Herpetological Review 34 (3): 230.

PUSCHENDORF, R., B O ~ O S , F. & CHAVES, G. (2006): The amphibian chytrid fungus along an altitudinal transect before the first reported declines in Costa Rica. - Biological Conservation 132: 136-142.

PUSCHENDORF, R., CHAVES, G., CRAWFORD, A.J. & BROOKS, D.R. (2005): Eleutherodactylus ranoides (NCN): Dry forest population, refuge from decline? - Herpetological Review 36 (1): 53.

PUSCHENDORF, R., KUBICKI, B., RYAN, M. & VAUGHAN, C. (2004): Cochranella albomaculata (NCN): Reproduction. - Herpetological Review 35 (1): 52.

URN, W.F. (1963): Observations on the life history of the treefrog, Phyllomedusa callidryas (Cope). - Texas Journal of Science 15: 155-170.

PYBURN, W.F. (1964): Breeding behavior of the leaf-frog, Phyllomedusa callidryas, in southern Veracruz. - The American Philosophical Society Yearbook 1964: 291-294.

PYBURN, W.F. (1967): Breeding and larval development of the hylid frog Phrynohyas spilom- ma in southern Veracruz, Mexico. - Herpetologica 23 (3): 184-194.

PYBuRN, W.F. (1970): Breeding behavior of the leaf-frogs Phyllomedusa callidryas and Phyllomedusa dacnicolor in Mexico. - Copeia 1970 (2): 209-218.

RABB, G.B. (1959): A new frog of the genus Plectrohyla from the Sierra de 10s Tuxtlas, Mexico. - Herpetologica 15: 45-47.

RABB, G.B. (1960a): A new salamander of the genus Chiropterotriton from Chiapas, Mexico, with notes on related species. - Copeia 1960: 304-311.

RABB, G.B. (1960b): On the unique sound pro- duction of the Surinam Toad, Pipa pipa. - Copeia 1960: 368-369.

357

Literature cited

RABB, G.B. & RABB, M.S. (1960): On the mating and egg-laying behavior of the Surinam Toad, Pipa pipa. - Copeia 1960: 271-276.

RABB, G.B. & RABB, M.S. (1963): Additional observations on breeding behavior of the Surinam Toad, Pipa - Copeia 1963: 636-642.

RABB, G.B. & SNEDIGAR, R. (1960): Observations on the breeding and development of the Surinam toad, Pipa pipa. Copeia 1960: 40-44.

RAMIREZ, J., VOGT, R. C. & VILLARREAL-BENITEZ, J. (1998): Population biology of a neotropical frog (Rana vaillanti). - Journal of Herpetology 32: 338-344.

W O S , C.W., PIMENTEL, N. & MARTINEZ-CORTES, V. (2002): Karyotype of the endemic golden frog Atelopus zeteki (Dunn) from Panama. - Caribbean Journal of Science 38 (3-4): 268-270.

RAND, A.S. (1983): Physalaemus pustulosus (rana sapito, t6ngara, foam frog, mud-puddle frog). - In: JANZEN, D.H. (ed.) Costa Rican Natural History - Chicago & London (Univ. Chicago Press): 412-415.

REINHARDT, J.T. & LUTKEN, C.F. (1863 "1862"): Bidrag ti1 det vestindiske Origes og navnli- gen ti1 de dansk-vestindiske Ors herpetolo- gie. - Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn, Serie 2,4: 153-291.

RENJIFO, J.M. & LUNDBERG, M. (1999): Guia de campo Anfibios y reptiles de Urra. -Skannka. Medellin. 96 pp.

RETALLICK, R., MCCALLUM, H. & SPEARE, R. (2004) Endemic infection of the amphibian chytrid fungus in a frog community post- decline. - PLoS Biology 2: 1965-1971.

RICHARDS, C.L. & KNOWLES, L.L. (2007): Tests of phenotypic and genetic concordance and their application to the conservation of Panamanian golden frogs (Anura, Bufonidae). - Molecular Ecology 16: 3119-3133.

RICHARDS, S.J., MCDONALD K.R. & ALFORD R.A. (1993): Declines in populations of Australia's endemic tropical rainforest frogs. -Pacific Conservation Biology 1: 66-77

RIVERO, J.A. (1961): Salienta of Venezuela. - Bulletin of the Museum of Comparative Zoology 126 (1): 1-207.

RIVERO, J.A. (1969): Sobre la Hyla rubra LAURENTI y la Hyla x-signata SPE (Amphi- bia, Salientia). - Memoria de la Sociedad de Ciencias Naturales La Salle 29 (83): 109-118.

ROBERTS, W.E. (1994): Explosive breeding aggregations and parachuting in a neotropical

frog, Agalychnis saltator (Hylidae). - Journal of Herpetology 28 (2): 193-199.

ROBERTS, W.E. (1995): Agalyclinis calcarifer (NCN): Reproduction. - Herpetological Review 26 (4): 200.

R o ~ m s o ~ , D.C. (1961): The identity of the tadpole of Anotheca coronata (STEJNEGER). -

Copeia 1961: 495. ROBINSON, D.C. C1976): A new dwarf salamander

of the genus Bolitoglossa (Plethodontidae) from Costa Rica. - Proceedings of the Biological Society of Washington 89: 289-294.

ROBLE, S.M. (1985): Observations on satellite males in Hyla chrysoscelis, Hyla picta and Pseudacris triseriata. - Journal of Herpetology (19 (3): 432436.

ROBLE, S.M. (1997): Hyla picta (rana arbon'cola). - Pp. 483-486 in GONZALEZ SORIANO, E., DIRZO, R. & VOGT, R.C. (eds.): Historia Natural de Los Tuxtlas. - Universidad Nacional Aut6noma de Mexico, MBxico, D.F., Mexico. 647 pp.

RODRIGUEZ-CONTRERAS, A., SENARIS, J.C., -0, M. & RIVERO, R. (2008): Rediscovery of Atelopus cruciger (Anura: Bufonidae): current status in the Cordillera de La Costa, Venezuela. - Oryx (2008),42:301-304.

ROHR, J.R. & RAFFEL, T.R. (2010): Linking global climate and temperature variability to widespread amphibian declines putatively caused by disease. - Proceedings of the National Academy of Sciences of the United States of America 107 (18): 8269-8274.

RON, S.R. (2005): Predicting the Distribution of the Amphibian Pathogen Batrachochytrium dendrobatidis in the New World. -

Biotropica 37: 209-221. RON, S.R., COLOMA, L.A. & CANNATELLA, D.C.

(2005): A new, cryptic species of Physa- laemus (Anura: Leptodactylidae) from western Ecuador, with comments on the call structure of the P. pustulosus species group. - Herpetologica 61: 178-198.

ROSSMAN, D.A. (1991): Bufo vdliceps (Gulf Coast toad). - Herpetological Review 22 (2): 64.

ROUSE, J.D., BISHOP, C.A. & STRUGER, J . (1999): Nitrogen pollution: An assesment of its threat to amphibian survival. - Environ- mental Health Perspectives 107: 799-803.

RUIZ-CARRANZA, P.M. & LYNCH, J.D. (1991): Ranas Centrolenidae de Colombia I. Propuesta de una nueva clasificaci6n generi- ca. - Lozania (Acta Zooldgica Colombians): 57: 1-30.

RUIZ-CARRANZA, P.M. & LYNCH, J.D. (1998): Ranas Centrolenidae de Colombia XI. Nuevas especies de ranas cristal del genero

Literature cited

Hyalinobatrachium. -Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 22: 571-586.

RUTHVEN, A.G. (1914): Description of a new engystomatid frog of the genus Hypopachus. Proceedings of the Biological Society of Washington 27: 77-80.

RUVINSKY, I., & MAXSON, L.R. (1996): Phylo- genetic relationships among bufonoid frogs (Anura: Neobatrachia) inferred from mitochondrial DNA sequences. Molecular Phylo- genetics and Evolution 5: 533-547.

RYAN, M.J. (1985): The t6ngara frog: A study in sexual selection and communication. - The University of Chicago Press. Chicago. 230 PP.

RYAN, M.J. (2002): Phyllobates vittatus (Golfodulcean poison dart frog). - Herpetological Review 33: 318.

RYAN, M.J., BLEA, N.J., LATELLA, I.M. & KULL, M.A. (2010): Leptodactylus savagei (Smoky jungle frog). Antipredator defense - Herpetological Review 41 (3): 337-338.

RYAN, M.J., LIPS, K.R. & EICHHOLZ, M.W. (2008): Decline and extirpation of an endangered Panamanian stream frog population (Craugastor punctariolus) due to an outbreak of chytridiomycosis. - Biological Conservation 141: 1636-1647.

RYAN, M.J., LIPS, K.R. & GLERMAKOWSKI, J.T. (2010a): New species of Pristimantis (Anura: Terrarana: Stabomantinae) from Lower Central America. - Journal of Herpetology 44 (2): 193-200.

RYAN, M.J., RAND, A.S. & WEIGT, L.A. (1996): Allozyrne and advertisement call variation in the tungara frog, Physalaemus pustulosus. - Evolution 50: 2435-2453.

RYAN, M.J., SAVAGE, J.M., LIPS, K.R. & GIERMAKOWSKI, J.T. (2010b): A new species of the Craugastor rugulosus series (Anura: Craugastoridae) from west-central Panama. - Copeia 2010 (3): 405409.

SANCHEZ, D.A. (2010): Larval development and synapomorphies for species groups of Hyloscirtus PETERS, 1882 (Anura: Hylidae: Cophomantini). - Copeia 2010 (3): 351-363.

SANDERS, 0 . (1973): A new leopard frog (Rana berlandieri brownorum). Journal of Herpetology 7: 87-92. .

SANTOS, J.C., COLOMA, L.A. & CANNATELLA, D.C. (2003): Multiple, recurring origins of apose- matism and diet specialization in poison frogs. - Proceedings of the National Academy of Sciences of the United States of America 100: 12792-12797.

SANTOS, J.C., COLOMA, L.A., SUMMERS, K., CALDWELL, J.P., REE, R. & CANNATELLA, D.C.

(2009): Amazonian amphibian diversity is primarily derived from Late Miocene Andean lineages. - PLoS Biology 7(3)e56: 0001-0014.

SAPORITO, R.A., GARRAFFO, H.M., DONNELLY, M.A., EDWARDS, A.L., LONGINO, J.T. & DALY, J.W. (2004): Formicine ants: An arthropod source for the pumiliotoxin alkaloids of dendrobatid poison frogs. - Proceedings of the National Academy of Sciences of the United States of America 101: 8045-8050.

SAPORITO, R.A., DONNELLY, M.A., GARRAFFO, H.M., SPANDE, T.F. & DALY, J.W. (2006): Geographic and seasonal variation in alkaloid-based chemical defences of Dendrobates pumilio from Bocas del Toro, Panama. - Journal of Chemical Ecology 32: 795-814.

SAPORITO, R.A., DONNELLY, M.A., JAIN, P., GARRAFFO, H.M., SPANDE, T.F. & DALY, J.W. (2007a): Spatial and temporal patterns of alkaloid variation in the poison frog Oophaga pumilio in Costa Rica and Panama over 30 years. - Toxicon 50: 757-778.

SAPORITO, R.A., DONNELLY, M.A., NORTON, R.A., GARRAFFO, H.M., SPANDE, T.F. & DALY, J.W. (2007b): Oribatid mites: a major dietary source for alkaloids in poison frogs. - Proceedings of the National Academy of Sciences of the United States of America 104: 8885-8890.

SAVAGE, J.M. (1965): A new bromeliad frog of the genus Eleutherodactylus from Costa Rica. - Bulletin of the Southern California Academy of Sciences 64: 106110.

SAVAGE, J.M. (1966): An extraordinary new toad (Bufo) from Costa Rica. - Revista de Biologica Tropical 14 (2): 153-167.

SAVAGE, J.M. (1967): A new tree-frog (Centro- lenidae) from Costa Rica. - Copeia 1967 (2): 325-331.

SAVAGE, J.M. (1968a): The dendrobatid frogs of Central America. - Copeia 1968 (4): 745-776.

SAVAGE, J.M. (1968b): A new red-eyed tree-frog (family Hylidae) from Costa Rica, with a review of the Hy1a uranochroa group. - Bulletin Southern California Academy of Sciences 67: 1-20.

SAVAGE, J.M. (1972a): The harlequin frogs, genus Atelopus, of Costa Rica, and western Panama. - Herpetologica 28: 77-94.

SAVAGE, J.M. (197213): The systematic status of Bufo simus 0. SCHMIDT with description of a new toad from western Panama. -Journal of Herpetology 6 (1): 25-33.

SAVAGE, J.M. (1974): On the leptodactylid frog called Eleutherodactylus palmatus (BOULENGER) and the status of Hylodes

Literature cited

fitzingeri 0. SCHMIDT. - Herpetologica 30 (3): 289-299.

SAVAGE, J.M. (1975): Systematics and distribution of the Mexican and Central American stream frogs related to Eleutherodactylus rugulosus. - Copeia 1975: 254-306.

SAVAGE, J.M. (1980): A new frog of the genus Eleutherodactylus (Leptodactylidae) from the Monteverde Forest Preserve, Costa Rica. - Bulletin Southern California Academy of Sciences 79 (1): 13-19.

SAVAGE, J.M. (1981b): The systematic status of Central American frogs confused with Eleutherodactylus cruentus. - Proceedings of the Biological Society of Washington 94 (2): 413-420.

SAVAGE, J.M. (1981~): The tadpole of the Costa Rican fringe-limbed tree-frog, Hyla fimbrimembra. - Proceedings of the Biological Society of Washington 93 (4): 1177-1183.

SAVAGE, J.M. (1987): Systematic and distribution of the Mexican and Central American rainfrogs of the Eleutherodactylus gollmeri group (Amphibia: Leptodactylidae). - Fieldiana Zoology New Series. 33: 1-57.

SAVAGE, J.M. (1997): A new species of rainfrog of the Eleutherodactylus diastema group from the Alta Talamanca region of Costa Rica. - Amphibia-Reptilia 18 (3): 241-247.

SAVAGE, J.M. (2000): A new species of rainfrog of the Eleutherodactylus milesi species group (Anura: Leptodactylidae) from Guatemala. - Revista de Biologia Tropical 48 (4): 993-999.

SAVAGE, J.M. (2002): The Amphibians and Reptiles of Costa Rica: A Herpetofauna between two Continents, between two Seas. - University of Chicago Press. Chicago.

SAVAGE, J.M., & B o ~ ~ s ; F. (2009a): An enigmatic frog of the genus Atelopus (family Bufonidae) from Parque Nacional Chirripo, Cordillera de Talamanca, Costa Rica. - Revista de Biologia Tropical 57: 381-386.

SAVAGE, J.M., & B O ~ O S ; F. (2009b): A checklist of the Amphibians and Reptiles of Costa Rica: Additions and nomenclatural revisions. - Zootaxa 2005:l-23.

SAVAGE, J.M. & DEWEESE, J.E. (1979): A new species of leptodactylid frog, genus Eleutherodactylus from the Cordillera de Talamanca, Costa Rica. - Bulletin Southern California Academy of Sciences 78 (2): 107-115.

SAVAGE, J.M. & DEWEESE, J.E. (1981): The status of the Central American leptodactylid frogs Eleutherodactylus melanostictus (COPE) and Eleutherodactylus pla tyrhynchus ( G ~ T H E R ) . - Proceedings of the Biological Society of Washington 93 (4): 928-942.

SAVAGE, J.M. & DONNELLY, M.A. (1992): The second collection of, and variation in the rare Neotropical toad Bufo peripatetes. - Journal of Herpetology 26 (1): 72-74.

SAVAGE, J.M. & EMERSON, S.B. (1970): Central American frogs allied to Eleutherodactylus bransfordii (COPE): A problem of polymorphism. - Copeia 1970 (4): 623-644.

SAVAGE, J.M. & HEYER, W. R. (1967): Variation and distribution in the tree-frog genus Phyllomedusa in Costa Rica, Central America. - Beitrage zur Neotropischen Fauna 5 (2): 111-131.

SAVAGE, J.M. & HEYER, W. R. (1970 "1969"): The tree-frogs (Family Hylidae) of Costa Rica: diagnosis and distribution. - Revista de Biologia Tropical 16 (1): 1-127.

SAVAGE, J.M., HOLLINGSWORTH, B.D., LPS, K.R. & JASLOW, A.P. (2004): A new species of rainfrog (genus Eleutherodactylus) from the Serrania de Tabasara, west-central Panama, and reanalysis of the fitzingeri species group. - Herpetologica 60: 519-529.

SAVAGE, J.M. & MYERS, C.W. (2002): Frogs of the Eleutherodactylus biporcatus Group (Leptodactylidae) of Central America and northern South America, including rediscovered, resurrected, and new taxa. -American Museum Novitates 3357: 1 4 8 .

SAVAGE, J.M., MCCRANIE, J.R. & ESPINAL, M. (1996): A new species of EIeutherodactylus from Honduras related to Eleutherodactylus bransfordii (Anura: Leptodactylidae). - Proceedings of the Biological Society of Washington 109 (2): 366-372.

SAVAGE, J.M., MCCRANIE, J.R. & WILSON, L.D. (1988): New upland stream frogs of the Eleutherodactylus rugulosus group (Amphi- bia: Anura: Leptodactylidae) from Honduras. - Bulletin Southern California Academy of Sciences 87 (2): 50-56.

SAVAGE, J.M., MCCRANIE, J.R. & WILSON, L.D. (1999): A new species of rainfrog of the Eleutherodactylus cruentus group from eastern Honduras (Amphibia: Anura: Leptodactylidae). - Proceedings of the Biological Society of Washington 112 (4): 787-792.

SAVAGE, J.M. & STARRETT, P.H. (1967): A new fringe-limbed tree-frog (family Centrolenidae) from lower Central America. - Copeia 1967: 604-606.

SAVAGE, J.M. & W m , M.H. (1972): Geographic variation and systematics of the Middle American caecilians, genera Dermophis and Gymnophis. - Copeia 1972 (4): 680-695.

SAVAGE, J.M. & WAKE, M.H. (2001): Reevaluation of the status of taxa of Central American caecilians (Amphibia: Gymno-

Literature cited

phiona), with comments on their origin and evolution. - Copeia 2001 (1): 52-64.

SCHLAEPFER, M.A. & FIGEROA-SANDI, R. (1998): Female reciprocal calling in a Costa Rican leaf-litter frog, Eleutherodactylus podiciferus. - Copeia 1998 (4): 1076-1080.

SCHMIDT, A. & WAKE, M.H. (1990): Olfactory and vomeronasal systems of caecilians (Amphibia: Gymnophiona). - Journal of Morphology 205: 255-268.

SCHMIDT, A.A. & KOHLER, G. (1997): Zur Biologie von Bolitoglossa mem.cana: Freilandbeob- achtungen, Pflege und Nachzucht. -

Salamandra 32 (4): 275-284. SCHMIDT, K.P. (1933): New reptiles and amphib-

ians from Honduras. - Zoological Series of Field Museum of Natural History 20: 15-22.

SCHMIDT, K.P. (1936a): Guatemalan salamanders of the genus Oedipus. - Zoological Series of Field Museum of Natural History 20: 135-166.

SCHMIDT, K.P. (1936b): New amphibians and reptiles from Honduras in the Museum of Comparative Zoology. - Proceedings of the Biological Society of Washington 49: 43-50.

SCHMIDT, K.P. (1939): New Central American frogs of the genus Hypopachus. Field Museum of Natural History Publication. Zoological Series 24: 1-5.

SCHMIDT, K.P. (1941): The amphibians and reptiles of British Honduras. - Zoological Series of Field Museum of Natural History 22 (8): 475-510.

SCHMIDT, 0. (1857): Diagnosen neuer Frosche des zoologischen Cabinets zu Krakau. - Sitzungsberichte der Akademie der Wissenschaften in Wien, Mathematisch- Natunvissenschaftliche Klasse 24: 10-15.

SCHMIDT, 0 . (1858): Deliciae herpetologicae Musei Zoologici Cracoviensis, Beschreibung der im K.K. Museum zu Krakau befindli- chen, von J.V. Warszewitz in Neu-Granada und Bolivia gesammelten Ungeschwantzen Batrachier. - Denkschriften der Akademie der Wissenschaften in Wien, Mathematisch- Natunvissenschaftliche Klasse 14: 237-258.

SCHNEIDER, J.G. (1799): Historia Amphibiorum Naturalis et Literarariae. Fasciculus Primus. Continens Ranas, Calamitas, Bufones, Salamandras et Hydros in Genera et Species Descriptos Notisque suis Distinctos. Jena: Friederici Frommanni.

SCHWARTZ, A. & HENDERSON, R.W. (1991): Amphibians and Reptiles of the West Indies Descriptions, Distributions, and Natural History. University of Florida Press, Gainesville, Florida. 720 pp.

SCHWARTZ, J.J. & WELLS, K.D. (1984): Vocal behavior of the Neotropical treefrog Hyla phlebodes. - Herpetologica 40 (4): 452-463.

SCHWARTZ, J. J. & WELLS, K. D. (1985): Intra- and interspecific vocal behavior of the neotropical treefrog Hyla microcephala. - Copeia 1985 (1): 27-38.

SCOTT Jr, N.J. (1976): The abundance and diversity of the herpetofaunas of tropical forest litter. - Biotropica 8 (1): 41-58.

SCOTT Jr, N.J. (1983a): Bufo haematiticus (sapo, toad). - Pp. 385 in: JANZEN, D.H. (ed.) Costa Rican Natural History - Chicago & London (Univ. Chicago Press).

SCOTT, N.J. (198313): Eleutherodactylus bransfordii (rana). - Pp. 399 in: JANZEN, D.H. (ed.) Costa Rican Natural History - Chicago & London (Univ. Chicago Press).

SCOTT Jr, N.J. (1983~) : Eleutherodactylus diastema (martillito, tink frog). - Pp. 399 in: JANZEN, D.H. (ed.) Costa Rican Natural History - Chicago & London (Univ. Chicago Press).

SCOTT Jr, N.J. (1983d): Hyla boulengeri (ranita de Boulenger, Boulenger's hyla). - Pp. 401 in: JANZEN, D.H. (ed.) Costa Rican Natural History - Chicago & London (Univ. Chicago Press).

SCOTT Jr, N.J. (1983e): Leptodactylus pentadactylus (rana ternero, smoky frog). - Pp. 405406 in: JANZEN, D.H. (ed.) Costa Rican Natural History - Chicago & London (Univ. Chicago Press).

SCOTT, N.J. & STARRETT, A. (1974): An unusual breeding aggregation of frogs, with notes on the ecology of Agalychnis spurrelli (Anura: Hylidae). - Bulletin of the Southern California Academy of Sciences 73 (2): 86-94.

SEIMON, T.A., SEIMON, A., DASZAK, P., HALLOYS, S.R.P., SCHLOEGEL, L.M., AGUILAR, C.A., SOWELL, P., HYATT, A.D., KONECKY, B., & SIMMONS, J.E. (2007): Upward range extension of Andean anurans and chytridiomycosis to extreme elevations in response to tropical deglaciation. - Global Change Biology 13: 288-299.

SEMLITSCH, R.D. (ed.) (2003): Amphibian conservation. - Smithsonian Books. Washington, D.C.

SEMLITSCH, R.D., SCOTT, D.E. PECHMANN, J.H.K. & GIBBONS, J.W. (1996): Structure and dynamics of an amphibian community: evidence from a 16-year study of a natural pond. - Pp. 217-248 In: CODY, M.L. & S~IALLWOOD, J.A. (eds): Long-term studies of Vertebrate communities. - Academic Press, San Diego, California.

Literature cited

SENARIS, J.C. (2001): Una nueva especie de Hyalinobatrachium (Anura: Centrolenidae) de la Cordillera de la Costa, Venezuela. - Memoria de la Sociedad de Ciencias Naturales La Salle. 59 (152): 133-147.

SESSIONS, S.K. (1978): The chromosomes of Anotheca spinosa (Stejneger), family Hylidae. - Herpetologica 34: 70-73.

SESSIONS, S.K. (2003): What is causing deformed amphibians? - Pp. 169-186 in: SEMLITSCH, R.D. (ed.) (2003): Amphibian conservation. - Smithsonian Books. Washington, D.C.

SESSIONS, S.K. & KEZER, J. (1991): Evolutionary cytogenetics of bolitoglossine salamanders (family Plethodontidae). - Pp. 89-130 in: GREEN, D.M. & SESSIONS, S.K. (eds.): Amphibian cytogenetics and evolution. Academic Press, San Diego.

SHANNON, F.A. & WERLER, J.E. (1955): Notes on amphibians of the Los Tuxtlas range of Veracruz, Mexico. - Transactions of the Kansas Academy of Science 58: 360-386.

SHEIL, C.A. & MENDELSON III., J.R. (2001): A new species of Hemiphractus (Anura: Hylidae: Hemiphractinae), and a redescription of H. jol~nsoni. - Herpetologica 57 (2):189-202.

SHEIL, C.A., MENDELSON, III., J.R. & Da Silva, H.R. (2001): Phylogenetic relationships of the species of Neotropical horned frogs, genus Hemiphractus (Anura: Hylidae: Hemiphractinae), based on evidence from morphology. - Herpetologica 57: 203-214.

SHREW, B. (1935): On a new teiid and Amphibia from Panama, Ecuador, and Paraguay. - Occasional Papers of the Boston Society of Natural History 8: 209-218.

SHREW, B. (1936): A new Atelopus from Panama and a new Hemidactylus from Colombia. - Occasional Papers of the Boston Society of Natural History 8: 269-271.

SILVERSTONE, P.A. (1975): A revision of the poison-arrow frogs of the genus Dendrobates Wagler. - Natural History Museum of Los Angeles County Science Bulletin 21: 1-55.

SILVERSTONE, P.A. (1976): A revision of the poison arrow frogs of the genus Phyllobates Bibron in Sagra (Family Dendrobatidae). - Natural History Museum of Los Angeles County Science Bulletin 27: 1-53.

SKELLY, D.K.: WERNER, E .E. & CORTWRIGHT, S.A. (1999): Long-term distributional dynamics of a Michigan amphibian assemblage. - Ecology 80: 2326-2337.

SMITH, E.N. (2005): Two new species of Eleutherodactylus (Anura: Leptodactylidae) of the alfredi group from eastern Guate- mala. - Herpetologica 61: 286-295.

SMITH, H.M. (1959): Herpetozoa from Guatemala. 1. Herpetologica 15 (4): 210-216.

SMITH, H.M., LYNCH, J.D. & REESE, R.W. (1966): Rana maculata Daudin, 1801 (Amphibia): proposed suppression under plenary powers. - Bulletin of Zoological Nomenclature 23: 169-173.

SMITH, H.M. & TAYLOR, E.H. (1950): Type localities of Mexican reptiles and amphibians. - University of Kansas Science Bulletin 33: 313-380.

SMITH, P.W. (1952): A new species of toad from the highlands of Guatemala and Chiapas. - Copeia 1952: 175-177.

SMITH, S.A., ARIF, S. NIETO-MONTES DE OCA, A. & WIENS, J.J. (2007): A phylogenetic hot spot for evolutionary novely in Middle American treefrogs. - Evolution 61: 2075-2085.

SOLANO, H. (1987): Algunos aspectos de la biologia reproductiva del sapito silbador Leptodactylus fuscus (Schneider) (Amphi- bia: Leptodactylidae). - Arnphibia-Reptilia 8: 111-128.

SPARLING, D.W. & FELLERS, G. (2007): Comparative toxicity of chlorpyrifos, diazi- non, malathion and their oxon derivatives to larval Rana boylii. - Pollution 147: 535-539.

SPARLING, D.W., FELLERS, G.M. & MCCONNELL, L.L. (2009): Pesticides and amphibian population declines in California, USA. -

Environmental Toxicology and Chemistry 20 (7): 1591-1595.

SPARLING, D.W., LINDER, D.W. & BISHOP, C.A. (eds.) (2000): Ecotoxology of amphibians and reptiles. - SETAC Press, Pensacola, Florida.

STARRETT, P. (1960): Descriptions of tadpoles of Middle American frogs. - Miscellaneous Publications Museum of Zoology, University of Michigan 110: 5-37.

STARRETT, P (1966): Rediscovery of Hyla pictipes COPE, with description a new montane stream H-yla from Costa Rica. - Bulletin Southern California Academy of Sciences 65: 17-28.

STARRETT, P (1967): Observations on the life history of frogs of the family Atelopodidae. - Herpetologica 23 (3): 195-204.

STARRETT, P.H. & SAVAGE, J.M. (1973): The systematic status and distribution of Costa Rica grassfrogs, Genus Centrolenella (Family Centrolenidae), with description of a new species. - Bulletin of the Southern California Academy of Sciences 72 (2): 57-78.

STEBBLNS, R.C. & COHEN, N.W. (1995): A natural history of amphibians. - Princeton Uni- versity Press. Princeton, New Jersey. 316 pp.

Literature cited

STEINDACHNER, F. (1864): Batrachologische Mittheilungen. - Verhandlungen der kaiser- lich-kijniglichen zoologischen botanischen Gesellschaft in Wien 14: 239-288.

STEJNEGER, L. (1904): The herpetology of Porto Rico. - Report of United States National Museum for 1902: 549-724.

STEJNEGER, L. (1906): A new tree toad from Costa Rica. - Proceedings of the United States National Museum 30: 817-818.

STEJNEGER, L. (1907): A new salamander from Nicaragua. - Proceedings of the United States National Museum 32: 465-466.

STEJNEGER, L. (1911): Descriptions of three new batrachians from Costa Rica and Panama. - Proceedings of the United States National Museum 41: 285-288.

STEJNEGER, L. (1917): A new species of horned tree-toad from Panama. - Proceedings of the Biological Society of Washington 30: 31-34.

STEWART, M.M. & MARTIN, G.E. (1980): Coconut husk piles: a unique habitat for Jamaican terrestrial frogs. - Biotropica 12: 107-116.

STRAUGHAN, I.R. & HEYER, W.R. (1976): A functional analysis of the mating calls of the Neotropical frog genera of the Leptodactylus complex (Amphibia, Leptodactylidae). - Papeis Avulsos de Zoologia 29: 221-245.

STUART, L.C. (1934): A contribution to a knowledge of the herpetological fauna of El Peten, Guatemala. - Occasional Papers of the Museum of Zoology University of Michigan 292: 1-18.

STUART, L.C. (1935): A contribution to a knowledge of the herpetology of a portion of the savanna region of central Peten, Guatemala. - Miscellaneous Publications Museum of Zoology, University of Michigan 29: 1-56.

STUART, L.C. (1941): Two new species of Eleutherodactylus from Guatemala. - Proceedings of the Biological Society of Washington 54: 197-200.

STUART, L.C. (1942): Description of two new species of Plectrohyla BROCCHI with comments on several forms of tadpoles. - Occasional Papers of the Museum of Zoology University of Michigan 455: 1-14.

STVART, L.C. (1943a): Taxonomic and geographic comments on Guatemalan salamanders of the genus Oedipus. - Miscellaneous Publications Museum of Zoology University of Michigan 56: 1-33.

STUART, L.C. (1943b): Comments on the herpetofauna of the Sierra de 10s Cuchumatanes of Guatemala. - Occasional P a ~ e r s of the Museum of Zoology LJniversityLof Michigan 471: 1-28

STUART, L.C. (1948): The amphibians and reptiles of Alta Verapaz, Guatemala. -

Miscellaneous Publications Museum of Zoology University of Michigan 69: 1-109.

STUART, L.C. (1952): Some new amphibians from Guatemala. - Proceedings of the Biological Society of Washington 65: 1-12.

STUART, L.C. (1954): Descriptions of some new amphibians and reptiles from Guatemala. - Proceedings of the Biological Society of Washington 67: 159-178.

STUART, L.C. (1961): Some observations on the natural history of tadpoles of Rhinophrynus dorsalis Dumeril and Bibron. -

Herpetologica 17: 73-79. STUART, L.C. (1963): A checklist of the herpeto-

fauna of Guatemala. - Miscellaneous Publications, Museum of Zoology, University of Michigan. 122: 1-150.

STUART, L.C. (1966): The environment of the Central American cold-blooded fauna. - Copeia 1966 (4): 684-699.

STEWART, M.M. & MARTIN, G.E. (1980): Coconut husk piles: a unique habitat for Jamaican terrestrial frogs. - Biotropica 12: 107-116.

STUART, S.N., CHANSON, J., COX, N. &YOUNG, B.E. (2010): Distributional patterns of the herpetofauna of Mesoamerica, a biodiversity hotspot. - Pp. 3-15. In WILSON, L.D., TOWN- SEND, J.H. & JOHNSON, J.D. (eds.): Con- servation of Mesoamerican Amphibians and Reptiles. - Eagle Mountain Publishing, LC, Eagle Mountain, Utah. i-xviii + 1-812 pp.

STUART, S.N., CHANSON, J., COX, N., YOUNG, B.E., RODRIGUES, A.S.L., FISHMAN, D.L. & WALLER, R. W. (2004): Status and trends of amphibian declines and extinctions worldwide. -

Science 306: 1783-1786. STUART, S.N., HOFFMANN, M., CHANSON, J., COX,

N., BERRIDGE, R., RAMANI P. & YOUNG, B. (2008): Threatened Amphibians of the World. - Lynx Edicions. Barcelona, Spain. 758 pp.

SUMMERS, A.P. (2001): Caecilia volcani TAYLOR. -

Catalogue of American Amphibians and Reptiles 721: 1-2.

SUMMERS, A.P. & WAKE, M.H. (2001): Clarifi- cation regarding the holotype of Caecilia volcani (Amphibia: Gymnophiona). - Copeia 2001 (2): 561-562.

SUMMERS, K., CRONIN, T. W. & KENNEDY, T. (2003): Variation in spectral reflectance among populations of Dendrobates pumilio, the strawberry poison frog, in the Bocas del Toro Archipelago, Panama. - Journal of Biogeography 30: 35-53.

SUMMERS, K., WEIGT, L.A., BOAG, P. & BERMINGHAM, E. (1999): The evolution of

Literature cited

female parental care in poison frogs of the genus Dendrobates: evidence from mitochondrial DNA sequences. - Herpetologica 55:254-270.

SUNYER, J. & KOHLER, G. (2010): Conservation status of the herpetofauna of Nicaragua. - Pp. 488-509. In WILSON, L.D., TOWNSEND, J.H. & JOHNSON, J.D. (eds.): Conservation of Mesoamerican Amphibians and Reptiles. - Eagle Mountain Publishing, LC, Eagle Mountain, Utah. i-xviii + 1-812 pp.

SUNYER, J., LOTZKAT, S., HERTZ, A,, WAKE, D.B., ALEMAN, B.M., ROBLETO, S.J. & KOHLER, G. (2008): Two new species of salamanders (genus Bolitoglossa) from southern Nicaragua (Amphibia, Caudata, Plethodontidae). - Senckenbergiana Biologica 88: 319-328.

SUNYER, J., WAKE, D.B., TOWNSEND, J.H., TRAVERS, S.L., ROVITO, S.M., PAPENFUSS, T.J., OBWO, L.A. & KOHLER, G. (2010): A new species of worm salamander (Caudata: Plethodontidae: Oedipina) in the subgenus Oeditriton from the highlands of northern Nicaragua. - Zootaxa 2613: 29-39.

TANNER, W.W. (1957): Notes on a collection of amphibians and reptiles from southern Mexico, with a description of a new Hyla. - Great Basin Naturalist 17: 52-56.

TANNER, W.W. (1962): A new Bolitoglossa (salamander) from southern Panama. -

Herpetologica 18: 18-19. TANNER, W.W. & Bw, A.H. (1961): Description

of a new species of salamander from Pana- ma. - Great Basin Naturalist 21: 23-26.

TAYLOR, D.H. & GUTTMAN, S.I. (eds.) (1977): The reproductive biology of amphibians. - Plenum Press. New York, 475 pp.

TAYLOR, E.H. (1937a "1936"): New species of Amphibia from Mexico. - Transactions of the Kansas Academy of Science 39: 349-363.

TAYLOR, E.H. (1937b): New species of hylid frogs from Mexico with comments on the rare Hyla bistincta Cope. - Proceedings of the Biological Society of Washington 50: 43-54.

TAYLOR, E.H. (1940): A new Syrrophus from Guerrero, Mexico. - Proceedings of the Biological Society of Washington 53: 95-98.

TAYLOR, E.H. (1941): New Amphibians from the Hobart M. Smith Mexican collections. - University of Kansas Science Bulletin 27: 141-167.

TAYLOR, E.H. (1942): New tailless Amphibia from Mexico. - University of Kansas Science Bulletin 28: 67-89.

TAYLOR, E.H. (1943): Herpetological novelties from Mexico. - University of Kansas Science Bulletin 29 (8): 343-361.

TAYLOR, E.H. (1944): The genera of plethodont salamanders in Mexico, Part 1. - University of Kansas Science Bulletin 30 (12): 189-232.

TAYLOR, E.H. (1948a): New Costa Rican salamanders. - Proceedings of the Biological Society of Washington 61: 177-180.

TAYLOR, E.H. (1948b): Two new hylid frogs from Costa Rica. - Copeia 1948: 233-238.

TAYLOR, E.H. (1949a): New salamanders from Costa Rica. - University of Kansas Science Bulletin 33 (6): 279-288.

TAYLOR, E.H. (1949b): Costa Rican frogs of the genera Cen trolene and Centrolenella. - University of Kansas Science Bulletin 33 (1): 257-270.

TAYLOR, E.H. (1951): A new Veracrucian salamander. - University of Kansas Science Bulletin 34: 189-193.

TAYLOR, E.H. (1952a): The salamanders and caecilians of Costa Rica. - University of Kansas Science Bulletin 34 (12): 695-791.

TAYLOR, E.H. (1952b): The frogs and toads of Costa Rica. - University of Kansas Science Bulletin 35 (5): 577-942.

TAYLOR, E.H. (1954): Additions to the known herpetofauna of Costa Rica with comments on other species, no. 1. - University of Kansas Science Bulletin 36 (9): 597-639.

TAYLOR, E.H. (1955): Additions to the known herpetofauna of Costa Rica with comments on other species, no. 2. - University of Kansas Science Bulletin 37 (13): 499-757.

TAYLOR, E.H. (1958a): Notes on Costa Rican Centrolenidae with descriptions of new forms. - University of Kansas Science Bulletin 39 (2): 41-68.

TAYLOR, E.H. (1958b): Additions to the known herpetofauna of Costa Rica with comments on other species, no. 2. - University of Kansas Science Bulletin 39 (1): 3-40.

TAYLOR, E.H. (1968): The caecilians of the world. A Taxonomic Review. - University of Kansas Press. Lawrence, Kansas.

TAYLOR, E.H. (1969): A new Panamanian caecilian. -University of Kansas Science Bulletin 48: 315-323.

TAYLOR, E.H. (1973): A caecilian miscellany. - University of Kansas Science Bulletin 50: 187-231.

TAYLOR, E.H. & Smith, H.M. (1945): Summary of the collections of amphibians made in MCxico under the Walter Rathbone Bacon traveling scholarship. - Proceedings of the United States National Museum 95: 521-613.

THOMAS, R. (1966): New species of Antillean Eleutherodactylus. - Quarterly Journal of

Literature cited

the Florida Academy of Sciences 28: 375-391.

THOMINOT, A. (1884): Note sur un Batracien d'es- p&ce nouvelle provenant de Panama. - Bulletin de la Soci6tC Philomathique de Paris, ser. 7,8: 151-152.

TOFT, C.A. (1981): Feeding ecology of Pana- manian litter anurans: Patterns in diet and foraging mode. - Herpetologica 15 (2): 139-144.

TOLEDO, L.F. (2010): A new species of Elachistocleis (Anura; Microhylidae) from the Brazilian Amazon. - Zootaxa 2496: 63-68.

TOLEDO, L.F. & HADDAD, C.F.B. (2009): Colors and some morphological traits as defensive mechanisms in anurans. - International Journal of Zoology 2009. Article ID 910892, 12 pages; doi:10.115512009/910892.

TOLEDO, L.F., LOEBMANN, D. & HADDAD, C.F.B. (2010): Revalidation and redescription of Elachistocleis cesarii (Miranda-Ribeiro, 1920) (Anura: Microhylidae). - Zootaxa 2418: 50-60.

TOWNSEND, J.H., BUTLER, J.M., WILSON, L.D., & AUSTIN, J.D. (2009): A new species of salamander in the Bolitoglossa dunni group (Caudata: Plethodontidae: Bolitoglossinae) from Parque Nacional Montaiia de Yoro, Honduras. - Salamandra 45: 95-105.

TOWNSEND, J.H., BUTLER, J.M., WILSON, L.D., & AUSTIN, J.D. (2010): A distinctive new species of moss salamander (Caudata: Plethodontidae: Nototriton) from an imper- illed Honduran endemism hotspot. -

Zootaxa 2434: 1-16. TOWNSEND, J.H., & WILSON, L.D. 2008. Guide to

the amphibians and reptiles of Cusuco National Park, HondurasIGuia de 10s anfibios y reptiles del Parque Nacional El Cusuco, Honduras. - Bibliomania!, Salt Lake City, Utah. xiv + 322 pp.

TRUEB, L. (1970): The evolutionary relationships of casque-headed treefrogs with co-ossified skulls (family Hylidae). - University of Kansas Publications, Museum of Natural History 18: 547-716.

TRUEB, L. (1971): Phylogenetic relationships of certain Neotropical toads with the description of a new genus (Anura: Bufonidae). - Natural History Museum of Los Angeles County Contributions in Science 216: 1-40.

TRUEB, L. (1974): Systematic relationship of neotropical horned frogs, genus Hemiphractus (Anura: Hylidae). - Occasional Papers of the Museum of Natural History of the University of Kansas NO. 29: 1-60.

TRUEB, L. (1984): Description of a new species of Pipa (Anura: Pipidae) from Panama. - Herpetologica 40: 225-234.

TRUEB, L. & CANNATELLA, D.C. (1986): Syste- matics, morphology, and phylogeny of the genus Pipa (Anura: Pipidae). - Herpeto- logica 42 (4): 412-449.

TRUEB, L. & GANS, C. (1983): Feeding specialization of the Mexican burrowing toad, Rhinophrynus dorsalis (Anura: Rhino- phrynidae). - Journal of Zoology, London 199: 189-208.

TRUEB, L. &TYLER, M.J. (1974): Systematics and evolution of the Greater Antillean hylid frogs. - Occasional Papers of the Museum of Natural History, University of Kansas 24: 1-60.

VALENZUELA GALVAN, D., CHAVEZ JUAREZ, J. & ALCARAZ CRUZ, H. (2004): Geographic distribution: Gastrophryne usta. - Herpetological Review 35: 184.

VAN BOCXLAER, I., LOADER, S.P., ROELANTS, K., B I J ~ , S.D., MENEGON, M. & B o s s m , F. (2010): Gradual adaptation toward a range- expansion phenotype initiated the global radiation of toads. - Science 327: 679-682.

VAN DEVENDER, R.W. (1980): Preliminary checklist of the herpetofauna of Monteverde, Puntarenas Province, Costa Rica and vicinity. - Brenesia 17: 319-326.

VAN DER MEIJDEN, A,, VENCES, M., HOEGG, S., BOISTEL, R., CHANNING, A. & MEYER, A. (2007): Nuclear gene phylogeny of narrow- mouthed toads (Family Microhylidae) and a discussion of competing hypotheses concerning their biogeographical origins. -

Molecular Phylogenetics and Evolution 44: 1017-1030

VASQUEZ-ALMAZAN, C.R., ROVITO, S.M., GOOD, D.A. & WAKE D.B. (2009): A new species of Cryptotriton (Caudata: Plethodontidae) from Eastern Guatemala. - Copeia 2009 (2): 313-319.

VAUGHAN, A. & CALVO, W. (2003): Crepidophryne epiotica (NCN): Costa Rica: Heredia. - Herpetological Review 34 (3): 257.

VAUGHAN, A. & MENDELSON 111, J.R. (2007): Taxonomy and ecology of the Central American toads of the genus Crepidophryne (Anura: Bufonidae). - Copeia 2007 (2): 304-314.

VENCES, M., KOSUCH, J., LOTTERS, S., WIDMER,A., JUNGFER, K.-H., KOHLER, J. & VEITH, M. (2000): Phylogeny and classification of poison frogs (Amphibia: Dendrobatidae), based on mitochondria1 16s and 125 ribosomal RNA gene sequences. - Molecular Phylogenetics and Evolution 15: 34-40.

Literature cited

VW, J.L. (1963): A new plethodontid salamander (Bolitoglossa sooyorum) from Costa Rica. - ~ e s s t a de ~iologia Tropical 11: 89-97.

VIAL, J.L. (1968): The ecology of the tropical salamander, Bolitoglossa subpalmata, in Costa Rica. - Revista de Biologia Tropical 15: 13-115.

VILLA, J. (1969): Comportamiento defensivo de la "rana ternero," Leptodactyluspentadacty- lus. - Revista de Biologia Tropical 15: 323-329.

VILLA, J. (1977): A symbiotic relationship between frog (Amphibia, Anura, Centrole- nidae) and fly larvae (Drosophilidae). - Journal of Herpetology 11: 317-322.

VILLA, J.D. (1984): Biology of a neotropical glass frog, Cen trolenella fleischmanni (BOETTGER), with special reference to its frogfly associates. - Milwaukee Public Museum Contributions in Biology and Geology 55: 1-60.

VILLA, J. D. (1988): Rana vibicaria (COPE) Rana montaiiera. - Catalogue of American Amphi- bians and Reptiles 437: 1-2.

VILLA, J. D. (1990): Rana warszewitschii (SCHMIDT). - Catalogue of American Amphi- bians and Reptiles 459: 1-2.

VITT, L.J. & CALDWELL, J.P. (2009): Herpetology. An Introduct ry Biology of Amphibians and Reptiles. 3~~ edition. Elsevier, Academic Press. Burlington, MA. 697 pp.

VOGT, R.C., VILLAREAL BENITEZ, J.-L. & PEREZ- HIGAREDA, G. (1997): Lista anotada de anfibios y reptiles. - Pp. 507-522 in G O N Z ~ E Z SORIANO, E., DIRZO, R. & VOGT, R.C. (eds.): Historia Natural de Los Tuxtlas. - Universidad Nacional Autonoma de Mexico, MBxico, D.F., Mexico. 647 pp.

WAKE, D.B. (1970): The abundance and diversity of tropical salamanders. - American Naturalist 104: 21 1-2 13.

WAKE, D.B. (1987): Adaptive radiation of salamanders in Middle American cloud forests. - Annals of the Missouri Botanical Garden 74: 242-264.

WAKE, D.B. (1998): On the taxonomic status of Nototriton sanctibarbarus MCCRANIE and Wilson (Amphibia: Caudata). -

Southwestern Naturalist 43 (1): 88-91. W m , D.B. (2005): Diversity of Costa Rican

salamanders. - Pp. 65-80 in DONNTELLY, M.A., CROTHER, B.L., GUYER, C., WAKE, M.H. &WHITE, M.E. (eds.): Ecology and Evolution in the Tropics. A Herpetological Perspective. - University of Chicago Press. Chicago.

WAKE, D.B. & BRAME, A.H. (1962): A new species of salamander from Colombia and the status

of Geotriton andicola Posada Arango. - Natural History Museum of Los Angeles County Contributions in Science 49: 1-8.

WAKE, D.B. & BRAI~IE, A.H. (1963a): The status of the plethodontid salamander genera Bolitoglossa and Magnadigita. - Copeia 1963: 382-387.

WAKE, D.B. & BRAME, A.H. (196313): A new species of Costa Rican salamander, genus Bolitoglossa. - Revista de Biologia Tropical 11: 63-73.

WAKE, D.B. & BRAME, A.H. (1966): A new species of lungless salamander (genus Bolitoglossa) from Panama. - Fieldiana Zoology. 51: 1-10.

WAKE, D.B. & BRAI~IE, A.H. (1969): Systematics and evolution of neotropical salamanders of the Bolitoglossa helmrichi group. - Natural History Museum of Los Angeles County Contributions in Science 175: 1-40.

WAKE, D.B., BRAME, A.H. & DUELLMAN, W.E. (1973): New species of salamanders, genus Bolitoglossa, from Panama. - Natural History Museum of Los Angeles County Contributions in Science 248: 1-19.

W m , D.B., BRAME, A.H. & MYERS, C. (1970): Bolitoglossa taylori, a new salamander from cloud forest of the Serrania de Pirre, Eastern Panama. - American Museum Novitates 2430: 1-18.

WAKE, D.B. & C.WBELL, J.A. (2000): A new species of diminutive salamander (Plethodontidae: Nototriton) from the Montaiias del Mico of Guatemala. -

Proceedings of the Biological Society of Washington 113 (3): 815-819.

WAKE, D.B. & DEBAN, S. (2000): Terrestrial feeding in salamanders. - Pp. 95-116 in: SCHWENK, K. (ed.): Feeding: Form, function, and evolution in tetrapod vertebrates. - Academic Press, San Diego.

WAKE, D.B. & DRESNER, I.G. (1967): Functional morphology and evolution of tail autotomy in salamanders. - Journal of Morphology 122: 265-306.

WAKE, D.B. & ELIAS, P. (1983): New genera and a new species of Central American salamanders, with a review of the tropical genera (Amphibia, Caudata, Plethodontidae). - Natural History Museum of Los Angeles County Contributions in Science 345: 1-19.

WAKE, D.B. & HANKEN, J. (1996): Direct development in the lungless salamanders: what are the consequences for developmental biology, evolution and ~hylopenesis? - International Journal of ~e ;e lo~men ta l Biology 40: 859-869.

WAKE, D.B., HANKEN, J. & I B ~ z , R. (2005): A new species of big black Bolitoglossa

Literature cited

(Amphibia: Caudata) from central PanamB. - Copeia 2005: 223-226.

WAKE, D.B. & JOHNSON J.D. (1989): A new genus and species of plethodontid salamander from Chiapas, Mexico. - Natural History Museum of Los Angeles County Contributions in Science 411: 1-10.

WAKE, D.R. & LYNCH, J.F. (1976): The distribution, ecology, and evolutionary history of plethodontid salamanders in tropical America. Natural History Museum of Los Angeles County Science Bulletin 25: 1-65.

WAKE, D.B. & LYNCH, J.F. (1982): Evolutionary relationships among Central American salamanders of the Bolitoglossa fianklini group, with a description of a new species from Guatemala. - Herpetologica 38: 257-272.

WAKE, D.B. & LYNCH, J.F. (1988): The taxonomic status of Bolitoglossa resplendens (Amphi- bia: Caudata). - Herpetologica 38: 257-272.

WAKE, D.B., PAPENFUSS, T.J. & LYNCH, J.F. (1992): Distribution of salamanders along elevational transects in Mexico and Guatemala. - Tulane Studies in Zoology and Botany, Suppl. Publ. 1: 303-319.

WAKE, D.B., SAVAGE, J.M & HANKEN, J. (2007): Montane salamanders from the Costa Rica- Panama border region, with descriptions of two new species of Bolitoglossa. - Copeia 2007 (3): 556-565.

WAKE, D.B. & VREDENBURG, V.T. (2008): Are we in the midst of the sixth mass extinction? A. view from the world of amphibians. - Proceedings of the National Academy of Sciences of the United States of America 105: 11466-11473.

WAKE, D.B., YANG, S.Y. & PAPENFUSS, T.J. (1980): Natural hybridization and its evolutionary implications in Guatemalan plethodontid salamanders of the genus Bolitoglossa. - Herpetologica 36: 335-345.

WAKE, M. H. (1963): Gymnopis multiplicata. - Catalogue of American Amphibians and Reptiles 411: 1-2.

WAKE, M.H. (1977): The reproductive biology of caecilians. - Pp. 73-101 In: TAYLOR, D.H. & GUTTMAN, S.I. (eds.): The reproductive biology of amphibians. - Plenum Press. New York and London. 475 pp.

WAKE, M.H. (1980): Reproduction, growth, and population structure of the Central American caecilian Dermophis mexicanus. - Herpetologica 36: 244-256.

WAKE, M.H. (1983): Gymnopis multiplicata, Dermophis mexicanus, and Dermophis parviceps (soldas, suelda con suelda, dos cabezas, caecilians). - In: JANZEN, D.H. (ed.) Costa Rican Natural History - Chicago &

London (Univ. Chicago Press): 400401. WAKE, M.H. (1988a): Gymnopis, G. multiplicata.

- Catalogue of American Amphibians and Reptiles 411: 1-2.

WAKE, M. H. (1988b): Gymnopis syntremus. - Catalogue of American Amphibians and Reptiles 412: 1.

WAKE, M.H. (1998): Dermophis oaxacae (Mertens). - Catalogue of American Amphi- bians and Reptiles 661: 1-2.

WAKE, M.H. & CAMPBELL, J.A. (1983): A new genus and species of caecilian from the Sierra de las Minas of Guatemala. - Copeia 1983: 857-863.

WAKE, M.H. & DONNELLY, M.A. (2010): A new lungless caecilian (Amphibia: Gymnophiona) from Guyana. -Proceedings of the Royal Society of London B 227 (1683): 915-922. doi: 10.1098/rspb.2009.1662

WALKER, C.F. (1973): A new genus and species of microhylid frog from Ecuador. - Occasional Papers of the Museum of Natural History, University of Kansas 20: 1-7.

WALKER, C.F. & DWELLMAN, W.E. (1974): Description of a new species of microhylid frog, Chiasmocleis, from Ecuador. - Occasional Papers of the Museum of Natural History, University of Kansas 26: 1-6.

WALLS, J. G. (1994): Jewels of the Rainforest: Poison Frogs of the Family Dendrobatidae. - J.F.H. Publications, Neptune City, New Jersey.

WARKENTW, K.M. (2000): Wasp predation and wasp-induced hatching of red-eyed treefrog eggs. - Animal Behavior 60: 503-510.

WARKENTW, K.M., BUCKLEY, C.R. & METCALF, K.A. (2006): Development of red-eyed treefrog eggs affects efficiency and choices of egg-foraging wasps. - Animal Behavior 71: 417425.

WASSERSUG, R.J. & DUELLMAN, W.E. (1984): Oral structures and their development in egg- brooding hylid frog embryos and larvae: evolutionary and ecological implications. - Journal of Morphology 182: 1-37.

WEBB, R.G. (1971): Egg deposition of the Mexican smilisca, Smilisca baudinii. - Journal of Herpetology 5:185-187.

WEIGT, L.A., CRAWFORD, A.J., RAND, A.S. & RYAN, M.J. (2005): Biogeography of the tungara frog, Physalaemus ~ustulosus: a molecular -. - perspective. - Nfolecular Ecology 14: 3857-2876.

WELDON, C., DU PREEZ, L.H., HYATT, A.D, MULLER, R. & SPEARE, R. (2004): Origin of the amphibian chytrid fungus. . - Emerging Infectious Diseases 10: 2100-2105.

Literature cited

WELLS, K.D. (2007): The ecology and behavior of amphibians. - The University of Chicago Press. Chicago. 1148 pp.

WERNER, F. (1896): Beitrage zur Kentniss der Reptilien und Batrachier von Central- amerika und Chile, sowie einiger seltenerer Schlangenarten. - Verhandlungen der kai- serlich-koniglichen zoologischen botanischen Gesellschaft in Wien 1896: 344-365.

WERNER, F. (1903): a e r Reptilien und Batrachier aus Guatemala und China in der zoologischen Staats-Sammlung in Miinchen nebst einem Abhang iiber seltene Formen aus anderen Gebieten. - Abhandlungen der Mathematisch-Physikalischen Classe der Koniglich Bayerischen Akademie der Wissenschaften, Munchen, 22: 342-384.

WEYGOLDT, P. (1980): Complex brood care and reproductive behaviour in captive poison- arrow frogs, Dendrobates pumilio 0. SCHMIDT. - Behavioral Ecology and Sociobiology 7: 329-332.

WEYGOLDT, P. (1987): Evolution of parental care in dart poison frogs (Amphibia: Anura: Dendrobatidae). - Zeitschrift fur Zoologische Systematik und Evolutionsforschung 25 (1): 51-67.

WHITFIELD, S.M. & PIERCE, M.S.F. (2003): Eleutherodactylus noblei (NCN): Repro- duction and parental care. - Herpetological Review 34 (4): 357-358.

WIDMER, A., LOTTERS, S. & JUNGFER, K.-H. (2000): A molecular phylogenetic analysis of the neotropical dart-poison frog genus PhyUobates (Amphibia: Dendrobatidae). - Die Natunvissenschaften. Berlin 87: 559- 562.

WIED-NELJWIED, M.A.P., Prinz zu (1824): Ab- bildungen zur Naturgeschichte Brasiliens. - Heft 8. Weimar: Landes-Industrie-Comptoir.

WIEGMANN, A.F.A. (1833): Herpetologischen Beytrage. I. Uber die mexicanischen Kroten nebst Bemerkungen uber ihnen venvandte Arten anderer Weltgegenden. - Isis von Oken 26: 651-662.

WIENS, J.J (2007): Book review: The amphibian tree of life. - Quaterly Review of Biology 82: 55-56.

WIENS, J.J, PARRA-OLEA, G, GmciA-Pmis, M. & WAKE, D.B. (2007): Phylogenetic history underlies elevational biodiversity patterns in tropical salamanders. - Proceedings of the Royal Society B. doi: 10.1098/rspb.2006.0301: 1-10.

WIENS, J.J, KUCZYNSKI, C.A., DUELLMAN, W.E. & REEDER, T.W. (2007): Loss and re-evolution of complex life cycles in marsupial frogs: does ancestral trait reconstruction mislead? - Evolution 61: 1886-1899.

WILCZYNSKI, W. & BRENOWITZ, E.A. (1988): Acoustic cues mediate intermale spacing in a Neotropical frog. - Animal Behaviour 36: 1054-1063.

WLLCZYNSKI, W., MCCLELLAND, B.E. & RAND, A.S. (1993): Acoustic, auditory, and morphological divergence in three species of neotropical frog. - Journal of Comparative Physiology A Sensory Neural and Behavioral Physiology 172 (4): 425-438.

WILD, E.R. (1995): New genus and species of Amazonian microhylid frog with a phylogenetic analysis of New World genera. - Copeia 1995: 837-849.

WILKINSON, M. & NUSSBAUM, R.A. (2006): Caecilian phylogeny and classification. - Pp. 39-78 in EXBRAYAT, J.-M. (Ed.) Reproductive Biology and Phylogeny of Arnphibia, Volume 3. Gyrnnophiona. - Science Publishers Inc., Enfield, NH.

WILSON, E.O. (1992): The diversity of life. -The Belknap Press of Hanard University Press. Cambridge, Massachusetts.

WILSON, L.D. (1979): New departmental records for reptiles and amphibians from Honduras. - Herpetological Review 10 (1): 25.

WILSON, L.D. & JOHNSON, J.D. (2010): Distributional patterns of the herpetofauna of Mesoamerica, a biodiversity hotspot. - Pp. 30-235. In WILSON, L.D., TOWNSEND, J.H. & JOHNSON, J.D. (eds.): Conservation of Mesoamerican Amphibians and Reptiles. - Eagle Mountain Publishing, LC, Eagle Mountain, Utah. i-xviii + 1-812 pp.

W n s o ~ , L.D. & MCCRANIE, J.R. (1985): A new species of red-eyed Hyla of the uranochroa group (Anura: Hylidae) from the Sierra de Omoa of Honduras. - Herpetologica 41 (2): 133-140.

WnsoN, L.D. & MCCRANIE, J.R. (1989): A new species of Ptychohyla of the euthysanota group from Honduras, with comments on the status of the genus Ptychohyla TAYLOR (Anura: Hylidae). - Herpetologica 45 (1): 10-17.

WILSON, L.D. & MCCRANIE, J.R. (1994): Comments on the occurrence of a salamander and three lizard species in Honduras. - Amphibia-Reptilia 15: 416421.

WILSON, L.D. & MCCRANIE, J.R. (1998): Amphibian population decline in a Honduran National Park. - FrogLog 25: 1-2.

WILSON, L.D. & MCCRANIE, J.R. (1998): The biogeography of the herpetofauna of the subhumid forest of Middle America (Isthmus of Tehuantepec to northwestern Costa Rica). - Royal Ontario Museum Life Sciences Contribution 163: 1-50.

Literature cited

WILSON, L.D., MCCRANIE, J.R. & CRUZ-Dh, G.A. (1994): A new species of Plectrohyla (Anura: Hylidae) from a premontane rainforest in northern Honduras. - Proceedings of the Biological Society of Washington 107 (1): 67-78.

WILSON, L.D., MCCRANIE, J.R. & WILLIAMS, K.L. (1985): Two new species of fringe-limbed hylid frogs from nuclear Middle America. - Herpetologica 41 (2): 141-150.

WILSON, L.D., MCCRANIE, J.R. & WILLIAMS, K.L. (1994): Commentary on the reproductive biology and systematic relationships of Hyla insolita MCCRANIE, WILSON and WILLIAMS. - Caribbean Journal of Science 30 (3-4): 214-221.

WILSON, L.D. & TOWNSEND, J.H. (2010): The herpetofauna of Mesoamerica: Biodiversity significance, conservations status, and future challenges. - Pp. 760-812. In WILSON, L.D., TOWNSEND, J.H. & JOHNSON, J.D. (eds.): Conservation of Mesoamerican Amphibians and Reptiles. - Eagle Mountain Publishing, LC, Eagle Mountain, Utah. i-xviii + 1-812 PP.

WILSON, L.D., TOWNSEND, J.H. & JOHNSON, J.D. (eds.) (2010): Conservation of Mesoamerican Amphibians and Reptiles. - Eagle Mountain Publishing, LC, Eagle Mountain, Utah. i-xviii + 1-812 pp.

WYGODA, M. (1988): Adaptive control of water loss resistance in an arboreal frog. - Herpetologica 44 (2): 251-257.

WYGODA, M.L. (1989): Body temperature in arboreal and nonarboreal anuran amphibians: differential effects of a small change in water vapor density. - Journal of Thermal Biology. 14 (4): 239-242.

WYNN, A. & HEYER, W.R. (2002 "2001"): Do geo- graphically widespread species of tropical amphibians exist? An estimate of genetic relatedness within the neotropical frog Letpodactylus fuscus (Schneider 1799) (Anura Leptodactylidae). - Tropical Zoology 14: 255-285.

YOUNG, B., SEDAGHATKISH, G., ROCA, E. & FUENMAYOR, Q. (1999): El Estatus de la Conservacion de la Herpetofauna de PanamB: Resumen del Primer Taller Internacional sobre la Herpetofauna de Panama. - The Nature Conservancy. Arlington, Virginia.

Yomc, B., LIPS, K.R., REASER, J.K., IBANEZ, R.. SALAS, A.W., CEDENO, J.R., COLOIMA, L.X. RON, S., LA W C A , E., MEYER, J.R., Mu%-OZ. A., B O ~ O S , I?., CHAVEZ, G. & ROMO, D. (2001): Population declines and priorities for amphibian conservation in Latin America. - Conservation Biology 15: 1213-1223.

ZALD~VAR-RNER~N, A., V. LE~N-REGAGNON & A. NIETO-MONTES DE OCA. (2004): Phylogeny of the Mexican coastal leopard frogs of the Rana berlandieri group based on mtDNA sequences. - Molecular Phylogenetics and Evolution 30: 3 8 4 9 .

ZHANG, P. & WARE, M.H. (2009): A mitogenomic perspective on the phylogeny and biogeography of living caecilians (Amphibia: Gymnophiona). - Molecular Phylogenetics and Evolution 53: 479491

Z~MMERMANN, H. & ZIMMERMANN, E. (1988): Etho-Taxonomie und zoogeographische Artengruppenbildung bei Pfeilgiftfroschen (Anura: Dendrobatidae). - Salamandra 24: 125-160.

ZIPPEL, K. (2002): Conserving the Panamanian golden frog: Proyecto Rana Dorada. - Herpetological Review 33 (1): 11-12.

ZUG, G. (1983): Bufo marinus (Sapo Grande, Sapo, Giant Toad, Marine Toad. - In: JANZEN, D.H. (ed.) Costa Rican Natural History - Chicago & London (Univ. Chicago Press): 386-387.

ZUG, G. & ZUG, P. (1979): The marine toad, Bufo marinus: A natural history resum6 of native populations. - Smithsonian Contributions to Zoology 284: 1-58.

ZWEIFEL, R. G. (1964a): Life History of Phrynohyas venulosa (Salientia: Hylidae) in Panama. - Copeia 1964 (1): 201-208.

ZWEIFEL, R.G. (1964b): Distribution and life history of the Central American frog Rana vibicaria. - Copeia 1964 (2): 300-308.

ZWEIFEL, R.G. (1986): A new genus and species of microhylid frog from the Cerro de la Neblina region of Venezuela and a discussion of relationships among New World microhylid genera. - American Museum Novitates 2863: 1-24.

ZWEIFEL, R.G. (1967): Systematic status of the Central American frog, Rana miadis. - Herpetologica 23: 54-56

Glossary

advertisement call species-specific call of a male anuran that serves to attract females or establish a territory

alkaloids complex, often toxic organic bases amplectant the state of being in amplexus or

pertaining to amplexus (Figs. 464,549,522, 601,644)

amplexus the sexual embrace or clasping of a female anuran by a male

annulus (plural annuli) a ring or ring-like structure (Figs. 15,161

anterior pertaining to or towards the front (head) end

anman pertaining to the Anura (frogs and toads) or to members of that order

aposematic of or relating to an antipredator defence, often involving bright contrasting warning colors

arboreal living in trees areolate covered with globular, closely set

prominences, e.g., the texture of the ventral skin in some anurans

arthropods animals with jointed appendages and a chitinous exoskeleton ie.g., insects, spiders, crustaceans)

autochthon evolved in a particular location autotomy ability to drop body parts (e.g., tail)

as a protective measure d a r g pertaining to the region formed by the

posterior surfaces of the insertion of the forelimbs and the adjacent surfaces of the body

barble a fleshy tubular extension of the skin basal located a t or pertaining to the base benthic pertaining to the bottom of a body of

water Biddeis organ ovarian tissue retained in

bufonid males, associated with the functional testis

bitid partially divided into two parts, e.g., the subarticular tubercles or prepollical processes in some species of frogs (Figs. 624e,f)

brood pouch an integumentary pouch in some frogs within which the eggs develop (Fig. 468)

casque a modification of the skull and associated structures of the head to resemble a hel- met (Fig. 467)

caudal of or pertaining to the tail

370

choanae internal nares that open into the mouth cavity (Fig. 36)

cloaca the common chamber that receives the contents of the reproductive, digestive, and excretory systems

cloaca1 of or pertaining to the cloaca or the region where the cloaca opens to the outside

compressed flattened side to side conspecific of the same species convex curved toward the outside co-ossified the condition in which the skin is

fused to the underlying bones (Figs. 496, 497,498)

cosmopolitan occurring worldwide costal of or pertaining to a rib cranium skull cryptic coloration coloration matching the sur-

rounding environment cuspidate having a cusp; used in reference to

disc pad shape in anurans cutaneous of or pertaining to the skin depressed flattened dorsoventrally, i.e., from

top to bottom dermal of or pertaining to the skin dextral of or pertaining to the right side dichromatism the existence of two color

morphs digit a finger or toe dimorphism the existence of two morphologies distal away from the base dorsal the upper surface of the body or other

structure dorsolateral intermediate between dorsal and

lateral, transition from the back to the side ecomorph a morphology shared by relatively

distantly related species adapted for similar ecologic conditions

emarghate of or pertaining to a margin with a single notch or indentation

endemic restricted in distribution to a particular location, limited distribution

granular having a surface texture consisting of or resembling tiny grains

groin pertaining to the region formed by the anterior surfaces of the insertion of the hind limbs and the adjacent surfaces of the body

gular of or pertaining to the ventral surface of the throat

heterogeneous non-uniform homogeneous similar, uniform inguinal of or pertaining to the groin region intercalary placed between other parts invertebrates animals without a vertebral col-

umn iridophore a cell containing silvery reflecting

platelets (synonym: guanophore) juvenile young animal that has not reached

sexual maturity yet keeled with a raised ridge lateral of or pertaining to the side of the body

or a structure lentic of or pertaining to standing or slow-mov-

ing water (e.g., in a lake) lotic of or pertaining to fast-moving water (e.g.,

in a stream) m d a r y of or pertaining to one of the bones

of the upper jaw, usually bearing teeth medial, median of or pertaining to the midline,

in the middle melanophore a cell containing melanin, a dark

brown to black pigment mental of or pertaining to the chin region metamorph a juvenile anuran that has recent-

ly transformed from the tadpole stage (Figs. 754, 757)

monophyletic forming a category descended from a common ancestor

monotypic species a species not divided into subspecies

monotypic genus a genus consisting of only a single species

morphologic concerning the external form nomenclature the branch of systematics that is

concerned with naming organisms nostrils external nasal openings, usually paired nuchal of or pertaining to the dorsal region of

the neck nuptial pad a morphologic structure situated

on the thumb of sexually active male anurans, often as a thick, roughened pad (Figs. 342,507)

occipital of or pertaining to the posterior portion of the head

omnivore an animal that eats both plant and animal food

osmoregulation the process involved in maintaining water regulation in an organism

osteoderm superficial dermal bones underlying the epidermis

oviparous egglaying ovoviviparous live bearing from eggs; develop

ment of fertilized eggs in the womb, with young hatching at or shortly after egg lap- ing

ocelli eye-like markings, markings consisting of a light ring with a dark center or a dark ring with a light center

palmate broad and flat paravertebral of or pertaining to the area

immediately lateral to the dorsal midline parotoid gland a swollen, glandular structure

behind the eye in some amphibians (Fig. 248)

peritoneal sheath the white portion of the parietal peritoneum in some frogs

phylogenetic concerning evolutionary his to^ physiography landscape features plicae fleshy longitudinal folds pollex the first finger, thumb polymorphism the existence of two or more dif-

ferent morphological types within a population

population a group of individuals of the same species living in a particular area and interbreeding among themselves

posterior of or pertaining to the rear or caudal portion of the body or a structure

postocular, postorbital located posterior to the eye

preocular located anterior to the eye prepollex a structure at the outer base of the

thumb (Figs. 501a, 541,542,563,6241 proximal situated near the base or point of

attachment receptacula transversales hyobranchiales

(RTH) posterior portion of the sinus hyobranchialis

receptacula venosa disci oralis (RDO) antero- median portion of the sinus hyobranchialis

reticulate having the appearance of a net or mesh (Figs. 341,325)

revision literally a "new perspective" I on a problem), a comprehensive treatment of a taxonomic group

rostra1 on the anterior tip of the upper jam. rudimentary in an early stage of development rugose having a rough or wrinkled surface tex-

ture scalloped having a series of curved projections scansorial climbing scute (pl. scutes) large flat scale semiaquatic lifestyle partially adapted to an

Glossary

aquatic environment; most semiaquatic species live at the edge of a body of water and spend part of the time in the water

semiarid habitat characterized by conditions close to but less extreme than arid ones

serrate notched or toothed along the edge sexual dichromatism differential coloration of

males and females in a species shagreened having a rough surface texture sinistral of or pertaining to the left side sinus hyobranchialis paired blood sinus sytem

that dominates the buccopharyngeal region some centrolenid in tadpoles

spadate shaped like a spade subcaudal of or pertaining to the ventral sur-

face of the tail subocular below the eye SVL snout-vent length, distance from snout tip

to cloacal opening syrnpatric co-occurring in the same geographic

area synonym two or more names for the same

taxon; in systematics, a senior synonym is the earliest available name for a taxon; a junior synonym is any available name other than the senior synonym and is invalid

syntopic occurring together in the same location

taxonomy study of the systematic classification of organisms

terminal of or pertaining to the distal end of a structure

thenar situated near the base of the thumb tubercle a small bump on the surface tympanum the membrane covering the exter-

nal ear opening, the eardrum (Figs. 244, 245,246

urnbellifom shaped like an umbrella (Figs. 724b, 728)

vent the external cloacal opening venter the lower surface of the body ventral on the belly, towards the belly; of or

pertaining to the lower surface of the body or other structure

ventrolateral a position intermediate between ventral and lateral, in transition from the belly to the lateral surface of the body or other structure

vertebral along the dorsal midline of the body vertebrates animals with a vertebral column versant the slope of a mountain or mountain

range vocal sac an outpocketing of the floor of the

mouth in male anurans that acts as a res- onating chamber during vocalization (Fig. 506, 516, 661, 723)

vocal slit an opening in the floor of the mouth in male anurans leading to the vocal sac

viviparous live-bearing zoogeography study of the distribution of ani-

mals

Addendum

Addendum

The following species was added to the known amphibian fauna of Central America a few days before this book was scheduled to go to the printer. It was not possible at that late date to include it in the main portion of this book.

ROVITO, S. M., VASQUEZ-ALMAZAN, C. R. & PAPENFUSS, T. J. (2010): A new species of Bolitoglossa (Caudata: Plethodontidae) from the Sierra de las Minas, Guatemala. - Journal of Herpetology 44 (4): 516-525.

Bolitoglossa zacapensis ROVITO, VASQUEZ- ALMAZAN & PAPENFUSS 2010, J. Herpetol. 44: 519; type locality: "Finca Planada de Margot, 18.7 km (by road) from CA-9 a t Aldea Las Pozas (km 143.5), Municipio Rio Hondo, Sierra de las Minas, Departamento de Zacapa, Guatemala, 1,660-m elevation, 15"10'24"N, 89"30'26W (WGS84 datum)." Maximum SVL in males 45.3 mm, in females 52.3 mm. Only known from the vicinity of the type locality.

Bolitoglossa zacapensis is a member of the B. rostrata species group. In the identification key to the species of Bolitoglossa from northern Central America (pp. 60fO B. zacapensis will key out to couplet 5 (B. cuchumatana and B. helmrichi). Bolito- glossa zacapensis differs from B. helmrichi in having a uniform pale tan coloration of the ventral surface of the tail (orange or pink in B. helmrichi? and in lacking any mottling on the ventral surfaces (mottling on venter, gular region, and underside of limbs in B. helmrichi?. Bolitoglossa zacapensis differs from B. cuchumatana in the number of maxillary teeth (64-85 maxillary teeth in B. zacapensis versus 25-70 in B. cuchumatana).

Index

Index Bold numbers refer t o figures (photographs and line drawings).

A Adelphobates 128, 13 1 Agalychnis 198ff - annae 7,197,198,199,200,

309,310 - callidryas 196, 197, 199, 200,

309,311 - lemur 197,200,200,309,311 - moreletii 197, 198, 200,200 - saltator 197,200,201 - spurrelli 197, 199,200, 201,

309,311 Allobates 129 - talamancae 129, 129 Arneerega 129 - maculata 129 Anotheca 2 12f - spinosa 205,212f, 212,213 Ammobates 128 - nocturnus 128 A t e l o p ~ c u s 97f - chrysophorus 97f, 97 Atelopus 98ff, 317 - certus 98,98 - chiriquiensis 97, 98, 98 - chirripoensis 98 - glyphus 98,100,102 - limosus 98,101 - senex 97,100,101 - varius 100,101,317,320 - zeteki 97,100, 101,324 AtreMona 27 - eiselti 27 axillary membrane 210

B Batra&ochytrjum 319ff - dendrobatidis 319ff Bolitoglossa 38,40ff - alberchi 40,41 - alvaradoi 40,41,67 - anthracina 40 - biseriata 40,41 - bramei 40,41,66 - carri 41,42 - cataguana 41,42 - celaque 42,43,60 - centenorum 42 - cerroensis 42,43,66,67

- colonnea 42,43,65 - compacta 42,42,66 - conanti 42,43 - copia 42 - cuchumatana 42,43,60 - cuna 42,43,67,68 - daryorum 42,43,60 - decora 43,44 - diaphora 44,45,63,65 - dimin uta 44,45,66 - dofleini 37,44,45 - dunni 44,45,60 - engelhardti 44,45,63 - epimela 44,45 - eremia 44,45,60 - flavimembris 44,45,60 - flaviventris 44,47 - franklini 44,47,60 - gomezi 46,47,66 - gracilis 46, 47, 66 - hartwegi 46,47,63 - heiroreias 46,47 - helmrichi 46,47,60 - huehuetenanguensis 46,47 - indio 46, 46 - insularis 46,46,68 - jacksoni 48,49 - kaqchikelorum 48,49,63 - la 48,49,60 - lignicolor 48,49, 67,69 - lincolni 26, 48, 49,60 - longissima 48,48 - magnifica 48, 49 - marmorea 48,49,66 - medemi 48,49 - meliana 50,51, 60 - mexicana 50,51, 63 - minutula 50,51,67 - mombachoensis 37,50,51,67 - mono 50,51,63 - mulleri 50, 51 - nigrescens 50,51,66 - ninadormida 50,51 - nussbaumi 50 - nympha 50,50 - obscura 52,66 - occidentalis 52, 53 - odonnelli 52,53 - omniumsanctorum 52, 53,60

- oresbia 52, 53 - pacaya 52,63 - pesrubra 20,32, 52,53,66 - phalarosoma 52,53 - platydactyla 52, 53 - p o r r a s o m 52,53,60 - psephena 52,55,63 - pygmaea 52,55,66 - robinsoni 54,55 - robusta 54,55,65 - rostrata 54,55,63 - rufescens 54,55 - salvinii 38,54, 55 - schizodactyla 54, 55,69 - sombra 54,54,66 - sooyorum 54,54,66 - striatula 9,56,57, 67,68 - stuarti 56, 57,63 - subpalmata 56,57,66 - suchitanensis 56,63 - synoria 56,57,60 - taylori 56, 57 - tica 56, 57 - tzultacaj 56 - veracrucis 56,63 - xibalba 56,57,63,65 - yucatana 56,57,63 Bradytn'ton 38,70 - silus 70, 70 Bromeliohyla 2 14 - bromeliacia 214,214, 312 Bufo 96 Bufonidae 96ff

C C a d a 29f - isthmica 29,29, 30 - leucocephala 29,30 - nigricans 29,30,30 - orientalis 28 - volcani 29, 30,30 Caeciliidae 29ff C a d t a - iwokramae 27 Charadrahyla 2 15 - chaneque 215,215 Chaunus 102f - centralis 102,103,103 - granulosus 102,103, 103

This book is a completely up-dated and fully colored illustrated guide to the 598 species of amphibians that are known to occur in the region extending from the Isthmus of Tehuantepec in Mexico to the southern extent of the Panamanian isthmus.

8peclalfeaturer:

The rich illustrated Keys for iden111 --.ion will prove useful to anyone who has a doubt about what amphibian is in hand.

I Color photographs of almost 90 percent of the species in life are provided for the caecilians, salamanders and frogs.

With expertly done line drawings of diagnostic features.

Distribution maps to all species.

Identification key for the anuran tadpoles.

Up to date information on amphlbian declines and extinctions.

and more ...

About the author: Dr. Gunther Kohler is the Head of the Department of Herpetology at the Sencken- berg Research Institute in Frankfurt am Main, Germany. To date, he has produced 26 books, 155 papular articles, and Z70 scientific papers on herpetology, most of

- which pe$ain=to Cenfral Amen'can amfMbian-s- - - - -

= 1

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