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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 d c a r i f e 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, 2011 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 ............................................................ bntroduction ...................................................... $bmments on the Usage of this Book ..... Classifkation of the Amphibians of :Central America ............................................. The Environment ........................................... Climate in Central America .....................
y. h p h i b i a ................................................... k
6 10 11 14 16 24 26
.ceecilians (Gymnophiona) ........................ 27 &lamandem (Caudata) ............................. 36
F
: Phga and Toads (Anura) .......................... Bufonidae ......................................................... Centrolenidae ........;........................................ Dendrobatidae ................................................ Terrarana .......................................................... '
Craugastoridae .............................................. Eleutherodactylidae ..................................... Strabomantidae ............................................. Hemiphractidae ............................................. Hylidae .............................................................. teiuperidae ...................................................... [ teptodactylidae .............................................. Microhylidae ................................................... Pipidae ............................................................... Ranidae .............................................................. Rhinophrynidae .............................................
93 96
117 128 140 158 181 186 192 195 273 276 282 290 291 296
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 daunting 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 imposing 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 exemplified by the Panamanian endemic fringelimbed 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 disastrous 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 anthropogenic environmental degradation 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 relatively little disturbance and where we have been fortunate to document robust populations of endemic amphibians whose populations have been decimated in the other two better-studied areas in the Cordillera Nombre de Dios. Tropical salamanders also are known to be involved in the global amphibian crisis and it is hypothesized that global climate change might be responsible for documented drastic declines in salamander abundance, community structure, and phylogenetic diversity, especially evident in one of the best studied and most diverse communities along an elevational transect on the southern slopes of Volcan Tajumulco in the Guatemalan department of San Marcos. On the other hand, little is known about how caecilians have been affected by these environmental pressures, as Dr. Kohler has noted elsewhere in this volume. Mesoamerican amphibian population decline appears to involve at least three
---
AgaIYChniSamae de Bim,, Heredia, CostaRiea, 1630 m). Photo:G.K6hIer
major causes, i.e., chytridiomycosis, habitat degradation and destruction, and global climate change. Evidence is accumulating to indicate that these three causes are synergistically intertwined, in effect creating an environmental super-problem that grows more resistant to solution with time. These environmental agents are bringing about losses in diversity at many levels of biological organization existing both through space and time. It is telling that they are occurring at the same time that new techniques for exposing biological diversity and evolutionary history are being developed and employed that depend upon the continued existence of robust communities of organisms, including amphibians. Such losses are not simply just a matter of a species of frog, salamander, or caecilian being on Earth one day and gone the next, as regrettable as that might be; the results can exacerbate to the level of irreparable changes in community structure as measured over ecological time 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 developments, a premium has been placed on cataloguing this diversity as we now understand it, with a view to updating this understanding as time advances. Thus, Dr. Kohler7sbook 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 impacting 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 familiar 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 format 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
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 attempts at identification. About 87 percent of the regions' 498 species of amphibians are illustrated and all are mapped. Following these accounts is a long section pmviding a key to the tadpoles of the anuram 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 disa 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 thousand 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
Fig. 2. Bolitoglossa striatula (Rio San Juan, Photo: G.Kohler Nicaragua, 30 m).
are highly diverse, widely distributed, and given to extensive endemism, but are undergoing population declines on a broadening 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 setting 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 Isthmus of that the Neotropics contain the Earth's El Salvador 7 largest reservoir of amphibian biodiversity (SANTOSet al. 2009). In the past decades, works have been published on selected groups of Central American amphibians (e.g., DUELLMAN 1970, SAVAGE & M. WAKE1972, CAMPBELL & SAVAGE 2000, DUELLMAN 2001, SAVAGE & M. WAKE Fig. 3. The zoogeographic region of Central 2001) as well as on the herpetofauna or America (shaded). amphibians alone of selected countries or regions (e.g., MERTENS 1952a, LEE 1996, & WILSON CAMPBELL 1998, MCCRANIE 2002, SAVAGE 2002, GUYER& DONNELLYular genetics. This book has the purpose to et al. 2005, KOHLER et al. accomplish an up-to-date summary of the 2005, MCCRANIE et al. (1995) pub- diversity and taxonomy of the amphibian 2006). FLORES-VILLELA lished a compilation of identification keys fauna of Central America and provide for the herpetofauna of Mexico. As identification keys to the species level. The presently understood, the amphibian contents of this book can only serve as a fauna of Central America consists of 498 momentary snapshot of the status quo; species representing 72 genera and 16 new discoveries and a better knowledge of families (Table 1). This breathtaking the phylogenetics of the taxa treated here diversity is due to the rich physiography, undoubtedly will usher in numerous taxoexceptional vegetational variation, and nomic changes in the near future. the vivid geological history of Central America; the geological history of the Revisionary work is needed urgently in region was summarized in KOHLER(2008) several taxonomic groups of Central American amphibians. Examples are the and is not repeated here. issues of taxonomic identity of the During the past ten years, we have wit- Panamanian populations of frogs referred nessed drastic changes in the taxonomy of to the genera Elachistocleis and amphibians (see summary of changes dur- Hyloxalus, respectively. Some authors ing the past 20 years by JOHNSON et al. have suggested that Relictivomer might 2010). Some species have undergone be a synonym of Elachistocleis, although repeated changes in the generic allocation without presenting evidence to support 1986, LAVILLAet during the past few years (i.e., the marine this opinion (e.g., ZWEIFEL toad has been variously assigned to the al. 2003); morphologic differences among genera Bufo, Chaunus, or Rhinella; those species currently assigned to either genus bufonid species allocated to Incilius today, are small and molecular genetic data have been assigned to Bufo, Cranopsis, or could be useful to clarify this issue. Other Ollotis just a few years ago). The recent - clusters of morphologically poorly differand most probably ongoing - period of tax- entiated species with partly questionable onomic change (WIENS2007, FROSTet al. validity of certain taxa include some cae2008) is due mostly to new research cilians (e.g., Oscaecilia elongata and 0. approaches, basically in the field of molec- osae), some salamanders (e.g., some taxa
I
the dunni, rostrata, and morio species
exes, respectively) and some frogs some taxa in the Ecnomiohyla milspecies group, as well as those in the us coccifer complex, the Craugastor species group, the Lithobates pipspecies group, the genera Hypopechus and Crepidophzyne, and also the d e m e l y similar Smilisca species, S. sila and S. sordida).These are just a few of the 'numerous taxonomic problems that prevail among Central American amphibians. f hope that this book will stimulate ,wearch and conservation activities on '*e amphibians in the Central American m t r i e s . Only on the basis of sound taxammy and a thorough understanding of the interactions with the environment of the species in question can successful conremation strategies be developed and "implemented. 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 dism i o n of the amphibian decline issue in &is book.
Comments on the Usage ofthis Book This book treats the species of amphibians known to occur within the geographic liraits of Central America, here defined as the geographic area between the Isthmus of Tehuantepec (level of narrowest portion) and the Panama-Choc6 border. Subspecies are not considered. I have adopted the taxonomic arrangement as proposed by FROST et al. (2006), modified by those in the works of FAIVOVI~H et al. (2005),GRANT et al. (20061, WILKINSON & NUSSBAUM (2006), SANTOS et al. (2009), and FAIVOVICH et al. (2010). As to which species to include, I have accepted all published species unless there is convincing evidence to the contrary available in print. 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 travelers. 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 definitively 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 distinguishing 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, Tantillita 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 arbitrary 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
(nearSan nam6a Be La Fwgen, Hmdia, Costa Rim). Photo: C4. KBhl~r
The index at inhdhgthe
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Class3cation of the Amphibians of Central America
ClassXcation of the Amphibians of Central A m h I
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Bolitoglossa (86) Bradytriton (1) Cryptotriton (6) Dendrotriton (8) Nototriton (14) NyctanoLis (1) Oedipina (29) Dseudoeurycea (6
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Family Genus (in parentheses is the number of species that occur in Central America)
claaacation ofthe Amplu'bh of Central America L
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~lophryniscus(1 Atelopus (8) Chaunus (2) Crepidophryne (3) Incilius (22) Rhaebo (1) Rhinella (2
Espadarana (1) Hyalinobatrachium (7: Sachatamia (2) Teratohyla (2)
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miphractus ( Gastrotheca (2
Ameerega (1) Colostgthus (3) Dendrobatm (9) Hyloxalus (1) PhyUobates (2) Phyllodactylus (31
I Agalychnis (6) Anotheca (1) Bromeliohyla (1) Charadrahyla (1) Cruziohyla (1) Dendropsophus (5) Duellrnanohyla (7)
Diasporus ( 6 ) Eleuthe,m&&pJm I
Exerodonta (5)
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The Environment
The Environment '
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- * Rainfomst The beauty and majesty of a primary tropical rainforest or an old mature , "-
The various mountain complexes that form the physiography of Central America can be divided into two major growth can be breathtaking. Tropical massifs. The northern block includes the rainforest is found in regions with monthMesa Central of Chiapas, the highlands ly temperatures exceeding 18 "C during of central and southern Guatemala, and all months of the year and an annual those of Honduras, El Salvador, and rainfall of more than 2000 mm. northern Nicaragua. These are separated Rainforest soils are typically poor, due to from the southern highlands by the leaching of soluble nutrients. In the Nicaraguan Depression. The southern northern parts of Central America, rainhighlands include the Cordilleras de forest is found mainly in the Caribbean Guanacaste, Central, and Talamanca, as lowlands. In Costa Riea and Panama, it well as the relatively low highlands of also covers large areas of the Pacific verDari6n in eastern Panama. The two large sant. During the day, temperatures in the Nicaraguan lakes (Lagos de Managua tropical rainforest reach 3045°C. During 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. ?
Fig. L. ,iooded rainforest at Rio San Juan, Nicaragua (30 a elevation). Photo: G.Kijhl81'
The h-at Cloud fontst eavyraftemom.
moist forest characterized by frequent mist or clouds at the vegetation level.
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The Environment streams channel the water on its way to the lowlands. At very high altitudes along 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. Dry forest Dry forests are typical of the Pacific versant of Central America and a few interior valleys of Honduras and Guatemala. The climate is characterized by low Precipitation (500-1200 mm) and long dry
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Fig. 8. Dry forest near Arenal, Motagua Valley, Guatemala, 70 m.
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1
The Environment last several months and vary hic location The precipitaseasonal and heavy downical 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 habisll 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.
e Belize, 5 m. habitat between Hatt~evllleand ~ e l i z City,
Photo: G. Kohler
The Environment
Fig. 10. P: pesrubra.
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habitat of Bolitogll~ssa Photo: G. Kiihler
Fig. 11. Lagune near the entrance to P.N. Juan Castro Blanco, Alajuela, Costa Rica, 1915 m; Photo: G. K6hler breeding site of Lithobates vibicarius and Isthmohyla pseudopuma.
aland ST
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Pine-oak forest Pine savanna
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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 illustrate the yearly climate in selected 3 wo areas. The mean monthly precipitation is indicated by bars. The mean daily temperature can be read from the upper j"'@' g curve and the mean nightly temperature from the lower curve (data from M ~ E R 250 1996, except for Utila, which was collected personally). 200
f
160 100
m 0
J F M A M J J A S O N D
-. 1W -MI -160
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-- 10
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10
-
6
5
0
J F M A M J J A S O N D
J F M A M J J A S O N D
The Environment 5-
J F M A M J J A S O N D
J F M A M J J A S O N D
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 & TRUEB tropics and subtropics (DUELLMAN 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 colonized 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 opportunistic feeders. They eat just about any live prey they can fit into their mouths. Some species, however, such as dendrobatids and many Neotropical microhylids feed primarily on termites and ants. Amphibians display an amazing variety of reproduction modes. Some species are
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 (WELLS2007). 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 & GUYER1994).
Key to the orders of adult a m p h i i
1 a No legs present; a protrusible tentacle present 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
Rsading
TAYLOR& GU~CTMAN 1977, DUELLMAN & TRUEB 1986, FEDER & BURGGREN 1992, STEBBINs & COHEN 1995. MCDIARMID& ALTIG 1999. DUELLMAN 1999,WEB 2007, OGIELSBA2009
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. WAKE1983, ZHANG& M. WAKE 2009). Caecilians can be differentiated f h m wormlike reptiles such as amphisbaenians 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 r e r y 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. WAKE1983). The left lung is rudimentary or absent; both lungs are absent in the South American species Caecilita iwokramae and A tretochoana eiselti (WAKE & DONNELLY2010). Caecilians have a protrusible, chemosensory tentacle, which is located on the side of the head between the eye and the nostril ~BILLO & WAKE1987). The animals continually 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& WAKE1990).There is no apparent external sexual dimorphism (SAVAGE 2002).
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. WAKE2001). 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 exceptionally 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. WAKE1983). 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. WAKE1983).
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 There are about 180 living species of cae- the males, the phallodaeum, is an eversible cilians worldwide, grouped into six fami- portion of the cloaca, which is inserted into lies: Caeciliidae (104 species), Ichthy- the female's vent during copulation (M. ophiidae (46 species), Rhinatrematidae (10 WAKE1980). Some species of the family species), Scolecomorphidae (6 species), Caeciliidae for which reproductive data are Typhlonectidae (13 species), and Uraeo- available, such as Dermophis mexicanus typhlidae (7 species) (WILKINSON & NUSS- and Gymnopis multiplicata, are live-bearers (M. WAKE1983). Development takes BAUM 2006). Caecilians are circumtropical
place in the oviduct, and the embryos exhaust the supply of yolk before they emerge fi-om the egg membranes. After the embryos have hatched in the oviducts, they feed on rich secretions from the thickened
1
annular grooves
Y
annular folds
oviductal wall and on cells from the oviductal lining (M. WAKE1977). Other caeciliids such as the South American Caecilia orientalis reproduce by orpary with terrestrial eggs ( F m et al. 2004).
/
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 positioned immediately below nostril ............ 2 b Tentacle and tentacular foramen posi-
2 a
b
3a
b
4 a
b
tioned well posterior to nostril ................ 4 No secondary folds; more than 220 primary folds .............................................................. ... O~caecilia(inpark 0. elongata, 0 . omel Secondary folds present; fewer than 210 primary folds ............................................ 3 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 ) Primary annular grooves incomplete at least on anterior body; grooves of primary folds not contrastingly darker than ground color; eye usually visible externally ........... ........................................................ Caecilia Eye not visible externally (eye covered by bone); a pair of splenial teeth behind dentary series of teeth ..................... G-oN Eye visible externally; no pair of splenial teeth behind dentary series of teeth ..........
.................................................... Dezmophis
Further Reading
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 COPE1877, 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.
DUNN1942a, TAYLOR1968, SAVAGE & M. WAKE
Caecilia nign'cans BOULENGER 1902,Ann:Mag. 1979, NUSSBAUM & HOOGMOED Nat. Hist. Ser. 7, 9: 51; type locality: "the Rio 1972, NUSSBAUM 1979, MOORE et a1.1983, International Lita, 3000 feetn, northwestern Ecuador. Total Commission on Zoological Nomenclature 1987, length to 1030 mm. Southeastern Dari6n DUCEYet al. 1993b, EXBRAYAT & HRAOUI- Province, Panama, through the Pacific lowBLOQUET 1994, LYNCH1999c, SAVAGE & M. WAKE lands of Colombia to extreme northwestern 2005 2001, GOWER& WEKINSON 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. 17. CaeciJia isthmica (Reserva Natural Privada Chucanti, Distrito de Chepigana, Darien, Panama, 797 m). Photo: A. J. Crawford
Dermophis
$
Fig. 18. Caecilia volcani (Santa F6, Veraguas, Photo: M. Ponce Panama, 800 m).
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 TAYLOR1968, SAVAGE & M. WAKE 1972, 2001, SUMMERS 2001, SUMMERS & M. WAKE 2001
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 north& M. WAKE western Colombia (SAVAGE 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. WAKE2001). 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. WAKEpers. 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. Reproductive 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. WAKE1980). 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 TAYLOR1955, 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 TAYLOR1968 i s regarded as a synonym of D. glandulosus according to SAVAGE & WARE(1972,2001).
-., 20. Dermophis parviceps (Gua, _..in, CRARC, Costa Rica, 520 m). Photo: B. Kubicki
Dermophis gracilior GUNTEIER1902, 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 TAYLOR1968 and Dermophis septentrionalis TAYLOR1968 are regarded as synonyms of D. mexicanus, according to SAVAGE &WARE1972, 2001. Demophis occidentalis TAYLOR1955, 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 (DUNN1924a), 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.
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-
b 2 a
b 3 a
ment ventrally, in sharp contrast with the otherwise pale venter; 94-112 primary folds; 35-88 secondary folds; total folds 94-112 .................... Demophis mexicanus Annular grooves not contrasting in color with overall ventral coloration; number of folds variable ............................................ 2 Head distinctly paler (pink to whitish in life, pale brown in preservative) than dorsal body; 11-37 secondary folds .............. 3 Head not markedly different in color than dorsal body; 37-96 secondary folds ......... 4 85-102 primary folds; 11-26 secondary folds; total folds 97-126 ..............................
Fig. 22. Dermophis parviceps (Rara Avis, Las Horquetas de Sarapiqui, Heredia, Costa Rica, 700 m). Photo: T. Leenders
................................... Dermophisparvicepe b 95-112 primary folds; 29-37 secondary ............................ ............................... Demophis ddentaZis
folds; total folds 126-149
4 a 91-106 primary folds; 37-60 secondary
............................ ............................... Dermophis glandulo8us
folds; total folds 132-159
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 p c i l i o r b 107-112 primary folds; 74-96 secondary folds; total folds 186-208 ............................
Fig. 23. Dermophis occidentalis (Palmar Norte, Puntarenas, Costa Rica, 25 m). Photo: P. Janzen
.............................. Dermophb crxstankeme Further Reading
TAYLOR1955, 1968, 1969, SAVAGE& M. WAKE 1972. T A ~ O1973, R M. WAKE1980,1983, 1998,
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Gymnopis
; ,
,
!
j
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 speciwas found during heavy men in rains on the surface in a pasture. These caecilians feed mostly on earthworms and termites, as well as soft-bodied insect larvae (M. WAKE1983). Gymnopis multiplicata gives birth to fully developed young. Litter size in G. multiplicata is 2 to 10, and the total length of the newborn varies between 100 and 120 mm (M. WAKE1983).
Fig, 24, Gpnopis mulwicata (puntaL ~ Puntarenas, Costa Rica, 40 m). Photo: G. Klihler
%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
G p n o p i s syntrema (COPE18661, 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 itF"lc1983 and Siphonovs oligozona COPE1877 are regarded as & n o n b s oi' G.syntrema, according to NUSSBAUM 1988 and SAVAGE & M. WAKE ,
k ~ ?
1
I
I Fig. 26. Gymnopis syntrema (Guatemala). Photo: J. A. Campbell
I
'
~
Caeciliidae
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 Fig. 27. Oscaecilia ochrocephala (Rio Tuira at holotype of 0 . osae was collected at night Rio Mono, Daribn, Panama). Photo: W. E. Duellman "fi-om a flooded path in a short-cropped (Courtesyof the Biodiversity Institute,University of h a s ) grassy airstrip during a moderate rain. In contrast to the other two Central American species of Oscaecilia, 0. ochrocephala has been collected more frequentKey to Gymnopie 1 a Dorsum gray, annular grooves contrasting ly and appears to inhabit a diverse array pink or white; 128-132 primary folds; of habitats; it was found as deep in soil as 63-93 secondary folds; total folds 193-206 10 m (when digging building foundations ....................................G 3 m . Q o . w in Panama City), and has been observed b Annular grooves not contrasting in color on the ground surface at dawn and dusk with overall dorsal coloration; 112-133 primary folds; 84-107 secondary folds; total with some frequency in Garnboa, Panama folds 201-250 ........ Ggmno* multiplic~ta (M. W m pers. comm. September 2010).
Gymnopis multiplicata rnnopis syntrema
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.
Further Rsading
M. WAKE1963,T A ~ O1968, R SAVAGE & M. WAKE 1972, T m o 1973, ~ M. wm & CAMPBELL 1983,
NUSSBAUM 1988, M. WARE 1988a,b, SAVAGE & M. WAKE 2001
& SAVAGE1992, Oscaecilia osae WAS Copeia 1992: 703; type locality: "airstrip at La Sirena, Peninsula de Oea, Cant6n de Osa, Puntarenas Province, Costa Rica (approx. 3 m)." length 382 rnm. GoLf~ region in Pacific southern Costa Rica, near sea level to 40
&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; 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 primary folds .......................... Oscaecilia osae b No dermal scales present in a ~ u l a r grooves; 226-231 primary folds .................. Fig. 29. Oscaecilia elongata (Los Algarrobos, ...................................... Osca& elongata Chiriqui, Panama, 145 m). Photo: A. Hertz
. r id
28. Oscaecilia elongata (Lo garrobos, rnoto: A. Hertz r i d , Panama, 145 m).
Fig. 30. Oscaecilia osae (La Gamba near Photo: P. Weish GoEto, Costa Rica, 60 m).
Rnther Reading
1942a, T A ~ O1968, R 1969, SAVAGE & M. WAKE 1972, TAYLOR 1973, NUSSBAUM& WILKINSON1989, LAHANAS& SAVAGE 1992, WAGE & M. WAKE2001
Fig. 31. Oscaecilia osae (La Gamba near Photo: P. Weish Golfito, Costa Rica, 60 m).
Plethodontidae
Salamanders (Caudata)
Fig.32. Bolitoglot
2esrubri .
de 1
uerte, C
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.
-
,
uww
.
.
ler
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) (WAKE2005). Tropical plethodontid salamanders have extraordinarily low basal metabolic rates and, therefore, 1983). require relatively little food (FEDER
All Central American salamanders belong to the subfamily Bolitoglossinae within the family Plethodontidae. The species of this family are distributed .from Nova Scotia Plethodontidae Plethodontid salamanders are character- and extreme southeastern Alaska southized (among other traits) by the absence of ward, reaching eastern Brazil and central lungs; therefore, respiration is primarily Bolivia. A major center of diversity of this cutaneous and via the buccopharyngeal group is Mesoamerica (Mexico and Central membranes. All tropical plethodontids America), with 13 currently recognized have an extremely long tongue that works genera and about 220 species, which just like a true projectile (WAKE& DEBAN equals nearly half of all salamander 2000). The tongue can be shot with great species (WIENSet al. 2007, WILSON& 2010).In contrast, only two of the accuracy at prey (mostly small insects and JOHNSON
-
::r. 33. Bolitoglossa dofleini (male; Nueva 13
1000 In).
Concepcion, Senahu, mta Verapaz, Guatemala, Photo: J. Sunyer
=-.-. r 34. Bolitoglossa mombachoensis (Volcan Mombacho, Granada, Nicaragua,
A
1100 in 1. Photo: G. Kohler
37
Rey to the genera of Central American
salamanders
1 a Shiny black with bright red spots on eyelids 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 elongate (Fig. 161); limbs very short (limb interval usually >7,rarely as low as 6) .....
......................................................
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 (WAKE2005). 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.
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 regenerated after tail breakage (VITT & CALDWELL 2009). Whereas no salamander is able to see colors, most plethodontid species have some degree of binocular vision (SAVAGEFig. 36. Roof of mouth in a salamander. 2002). Drawing: G. Kohler
Plethodontidae
b
a b a
.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 Nostrils larger, ratio nostril width 1 mL usually ~0.018,or if 40.019, then ratio CCIJSVL >0.47 ........................................ 8 Head narrower (ratio HWISVL ~0.16); legs shorter (ratio CCLISVL usually c0.40, rarely to 0.45) ...........................Notokitan Head broader (ratio HWISVL >0.15); legs longer (ratio CCUSVL >0.44) .................... ................................ Dendlv~ton(in part) Legs shorter, ratio CCIJSVL <0.47, or if '0'479 then ratio nostril diameter <0.019 and ratio tail length I SVL >1.25; feet narrower (ratio HFWISVL 0.06-0.09)
'
................................................ Ciyptotriton b Legs longer, ratio CCIJSVL >0.47; ratio nostril diameter 1 SVL usually >0.019;
I ~ i38.~oedipina . uniformis (Albergue Monterreal, near P.N. Juan Castro Blanco, A1ajuela, Rita, lgoO m). Photo: G. Kiihler
ratio tail length I SVL usually c1.25; feet broader (ratio HFW/SVL 0.09-0.12) ......... ................................ Dendlvtn'ton (in part)
lrtber Reading
1926, TAYLOR1944, WAKE & DRESNER 67, DODD& BRODIE1976, WAKE& LYNCH V6, HANKEN et al. 1980, WAKEet al. 1980, -N 1983, WAKE& ELIAS1983, ELIAS1984, b g ~ 1987, &SEN et al. 1989, SESSIONS& EER 1991, WAKE et al. 1992, WAKE& HANKEN 96, BRUCE2000, GARCIA-PAR~S et al. 2000, b g & ~ DEBAN 2000, BRUCE2003, CHAN2003, LTELLER et d. 2004 INN
Fig. 40. Pseudoeurycea b m a t a (San Marcos, Guatemala). Photo: D. B. Wake
costal grooves
rsal view
!
cloaca
tral view
V head length
Fig. 37. External characteristics of salamanders. Drawings: G. KiShler
Fig.39. Foot of Bolitoglossa heiroreias; a. dorsal view; b. ventral view; note subdigital pads (red circle). Photos: G. Kohler
the veracrucis species group), Nanotriton (diminutive, short-tailed salamanders with small hands and feet: the two species preBolitoglossa is the largest and most widely viously included in the rufescens species distributed genus in the order Caudata group plus the recently described B. and includes about 16% of the recognized nymphal, Oaxakia (includes five species, et al. previously included in the macrinii species salamander species (PARRA-OLEA 2004). While most species of Bolitoglossa group) and Pachymandra (includes two occur in Central America and Mexico, the large to very large species with hands and genus ranges throughout most of tropical feet that are fully webbed and first caudal South America as well ( B m & WAKE vertebrae that have branched transverse 196313). These salamanders are found in processes: B. dofleini and B. alvaradoi). environments ranging from lowland rainforests to cool, humid cloud forests and Bolitoglossa alberchi GARC~A-PAR~S, PARRAparamo of the Talamanca (to 3620 m eleOLEA,BRAME &WAKE2003, Rev. Esp. Herpetol. vation; SAVAGE 2002). Hands and feet have 16: 57; type locality: "near Playa Escondida, diversified greatly in the genus (WAKE & 14 k m (road) NE approximately Sontecomapan, Veracruz, MBxico, 18"56'N, LYNCH 1976).Extensive webbing is usually 95"05W,a t ca. 50 m elevation." Maximum SVL associated with climbing behavior and in males 69.6 mm, in females 97.1 mrn; tail semi-arboreal to arboreal life histories 1engthISVL ratio about 1.0. Southern Mexico (WAKE& LYNCH 1976). in the Los Tuxtlas area, in east-central
Bolitoglossa
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& ELIAS1983). Monophyly of Bolitoglossa is supported by morphological and molecular genetic data (PARRA-OLEA et al. 2004).
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 TAYLOR1954, 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. SAVAGE, WAKE Bolitoglossa anthraciua BRAME, & 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, i n 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 Seven subgenera of Bolitoglossa are cura t mouth of Paya Tributary, Dari6n Province, rently recognized (PARRA-OLEA et al. 2004): Panama." Maximum SVL in males 36.7 mm, in Bolitoglossa (12 species previously includfemales 48.7 mm; tail 1engtWSVL ratio 0.80-1.16. Panama to northwestern Ecuador ed in the mexicana species group), 2006), (see recent record by CISNEROS-HEREDIA Eladinea (all South American and some near sea level to 500 m elevation. Lower Central American species), Bolitoglossa bramei WAKE,SAVAGE & HANKEN Magnadigita (a morphologically heteroge2007, Copeia 2007: 561; type locality: "Costa neous assemblage including the species Provincia Puntarenas, trail to Cerro previously included in the d m ; f r a n k l i ~ ~ ~Rica, , Pando, above Las Tablas, 8"56'N, 82"46W, and rostrata species groups), Mayamandra 2200 m." Maximum SVL in males 39.1 mm, in (includes three species previously placed 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, Puntarenas, Costa Rica). Photo: M. Garcia Paris
~hg. 41. Bolitoglossa alberchi (Berriozabal,
"i
-epas, Mexico, 800 m). Photo: A.Ramirez V.
Fig. 45.Bolitoglossa carri (Cerro Cantagallo, Francisco MorazBn, Honduras). Photo: J. R. McCranie
3 I Bolitoglossa biseriata (Bilsa Biological .lull, Esmeraldas, Ecuador, 550 m). Photo: H. M. Ortega-Andrade
Fig. 46. Bolitoglossa cataguana (P.N. Montaiia de Yoro, Francisco Morazan, Honduras, 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& BRODIE2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 28; type locality: "Cerro Bobic, SSE San Mateo I x t a t h , 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 0.73-0.94. Only known from the type locality Bolitoglossa cerroensis (TAYLOR1952a), 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 Coph, elevation, Departamento de 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).
Fig. 47. Bolitoglossa ,,~pacta (l.,.lan B a d , photo: A. ~ chiriqui, panama, 3000 m).
~
r
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. ROVITOpers. 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 only from type locality. Bolitoglossa cuchumatana (STUART1943a1, Misc. Publ. Mus. Zool. Univ. Michigan 56: 14; 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. ROWTOpers. 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.
,I I
i
ler
Fig. .x. Bolitoglossa cuchumatana (Laj Chimel, QichB, Guatemala, 2266 m). Photo: S. M. Rovito
'Pig.49. Bolitoglossa cerroensis (Cerro de la Huerte, Costa Rica). Photo: W. Van Devender
Fig. 53. Bolitoglossa cuna (Nusagandi, Kuna Photo: G. Kiihler Yala, Panama, 350 m).
CRARC, Costa Rica, 400 m). Photo: G. Kahler
Fig. 54. Bolitoglossa dargonun (holotype). Photo: J.A. Campbell
=
, ,.,
--
., ,,litog,,,a conanti (El Portillo, Ocotepeque, Honduras). Photo: G. Kohler
Fig. 55. Bolitoglossa decora (holotype). 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 & WILSON1997, 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 a n 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 (WERNER19031, 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. 19331, Field Mus. Bolitoglossa dunni (SCHMIDT 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& BRODIE2010, 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 (SCHMIDT19361, 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 (SCHMIDT1936),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. 19361, Field Bolitoglossa franklini (SCHMIDT 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
M
Fig. 56. Bolitoglossa diaphora (P.N. Cusuco, Cortks, Honduras, 1550 m). Photo: G. Kohler
Fig. 60. Bolitoglossa engelhardti (Finca La Insula, San Marcos, Guatemala, 2057 m). Photo: S. M. Rovito
Fig. 57. Bolitoglossa diminuta (holotype). Photo: D. B. Wake
Fig. 61. Bolitoglossa epimela (holotype). Photo: D. B. Wake
Fig. 58. BoLitoglossa dofleini (Quebrada Grande, Cop&, Honduras, 1300 m). Photo: G. Kahler
Fig. 62. Bolitoglossa en Jalapa, Guatemala, 26L-
..,.
~mw
Photo: J. A. Campbell
r
Fig. 59. Bolitoglossa dunni (EN.Cusuco, CortBs, Honduras, 1810 m). Photo: J. E. Kolby
Fig. 63. Bolitoglossa flavimembris (Finca La Insula, San Marcos, Guatemala, 2057 m). Photo: S. M. Rovito
45
1.10-121.00 the Sa&e venmt &rm southmtmChiapas,Me*, dong &@ F d c versmt of &atemala as far as Vol& At&h, 1506-3000 m elmdon. BdliW&mnmwg, . SUBL I?bMlaN "Cmh 2007, Cop& a002 ':857; type Etiea, P r o ~ c hPuntm6nae. L a CNCes ~ d o g i c astation, l a 0 4 ~ 3N, 5 15ae5mow2BNI my lkximum SVL in d e 49~.9 mm, in females 54,s aun; tail len@b%VL ratio Q.%l.lbPtilCitie dope of wesbru hia;bl&~tds pf T%mamaBad aaaeent Costa Itiaa, &o knew from the ~ ~ ~ l t ae rdn 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; localits: "Ria Q&L about 1 Itm NE by road &am the w e I W O ~ tha ~ ~ Rfo J Gkmde de =ax Ttrpantf, Cert~goProvince, 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 l e n & m d o 1.2Z-1.IS.Onls
ms
...
w
178: 10; type l d t p *4.B d m 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 &mda53.6 m; kd bB%$b/8m ~aaioO.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, G u a W J % lZ0&301)0 m elevation.
--
-.--
r
O
&yetzdte~wque,CPuathtmalbl(5. m o pers. eamm. Nm.20101,l84Q-%806 sn elevation. B-1B&& ( S ~ 19861, T Rdd MUB,N&. Mst. Publ., h l . Ser. 20: 16% tme l ~ d "naou12W w above Fin- Sam% w& of Cob- Alta Vmapaz, Qua-? d 5,W feet alMtude,wbhzimm 6VL in maledi 47.4 mm, in f'ePL4enaE88 4 . 7 mm; ;taille-VL =tie Q.67-0.95. SouthAltpl VmpaZ grid . Baja Verapm Deparbnah~, Guatemala f00e2@0 zn &ation. ~ ~ ~ hu~ ; h ~ ce t e nz m g Gem, a~i~ 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, Ouaat 3781)m d e v c tion (35.81944"N, 91.5641W Ffn9Maximma 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~UINT1 8 Departamento de BuSllueCnatlgo, Guatemala, MBO-2$85 la elmation. M*1og9(1 MB S-ER, L o T n a T , l3mnz, wm7m,RCEWEQ $ 0 9 && I d t . YDos Boeas Bid.88: 320; wo (11~%64.8"N,rn06gt&.4"7W), 2E m elmaiim, 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
mm,
lbhxhltrm s m ia malw44.9 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(dtawghit ia auh yef known from khe Hodp& of this m o d a h range), as well as Boa O ~
&.
Fig.64. Bditoglossa indio (hobtype). Phnta: J. Sunjer
...
, Chiapas,
Fig. 70. Bolitoglossa hartwegi (Cerro Tzontehuiz, Chiapas, Mexico). Photo: T. Bille
b
t
Fig. 67. BoLitoglossa franklini (Cerro Mozotal, Chiapas, Mexico, ca. 3000 m). Photo: L. Gray
Fig. 68. BoLitoglossa gomezi (Jurutungo, Chiriqui, Panama, 2120 m). Photo: G. Kiihler
Fig. 71. Bolitoglossa heiroreias (P.N. Montecristo, Santa Ana, El Salvador, 1904 m). Photo: T. Leenders
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 ELIAS1984, 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. SMI'I'H, Bolitoglossa kaqchikelorum CAMPBELL, STREICHER, ACEVEDO & BRODIE2010, 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. S m , STREICHER, Bolitoglossa la CAMPBELL, ACEVEDO & BRODIE2010, 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 BAUERet 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 (STUART1943a), 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. Bolitoglossa longissima MCCRANIE& CRUQD ~ A1996, Z 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.
I
WAKE & Bolitoglossa magnifica WN, 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 TalarnancaBard 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& WAKE1972, 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. ... 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). Photo: J. R. McCranie
I
Lag Fig. 75. Bolitoglossa jacksoni (from type localiPhoto: J. Hanken
Fig. 79. Bolitoglossa lincolni (Sierra de 10s Cuchumatanes, Guatemala). Photo: T. Bille
Fig. 76. Bolitoglossa kaqchikelorum (CerroAlux, Photo: J. A. Campbell Guatemala, 2107 m).
Fig. 80. Bolitoglossa magnifica (Las Tablas, Puntarenas, Costa Rica). Photo: M. Garcia Paris
Fig. 77. Bolitoglossa la (holotype). Photo: J. A. Campbell
Fig. 81. Bolitoglossa marmorea (Volch Baru, Chiriqui ,Panama, 3400 m). Photo: A. Hertz
' I i
I
Fig. 78. Bolitoglossa ligrugrucolor (Meseta de Chorcha, Chiriqui, Panama, 250 m). photo: G. Kijhler
Fig. 82. Bolitoglossa medemi (near Belky, Antioquia, Colombia). Photo: P. A. Silverstone (Courtesy of the Biodiversity Institute, University of Kansas)
49
m
Plethodontidae and eastern Panama to the Choc6 region of Colombia, near sea level to 800 m elevation. Bolitoglossa me.?iana WAKE & LYNCH1982, 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 VeraCrux." 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 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 ~ H L E R& 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 (COPE1869b), Roc. Acad. Nat. Sci. Philadelphia 21: 103; type locality: "Mountains of Guatemala." Neotype from '3.7 km S Santa Maria Visitaci611, Sololl, Guatemala 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 ~ o c c m1882 is regarded as a synonym of B. morio according to CAMPBELL et al. (2010). Bolitoglossa mulleri (BROCCHI1883), 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~TR, STREICHER, ACEVEDO & BRODIE2010, Misc. 200: 28; type Publ., Mus. Zool., Univ. 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, S T R E I C ~ ACEVEDO R, & BROBIE2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 26; type locality: 7.2 km SE Todos Santos, Huehuetenango, 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 & BRODIE2010, 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. ~outoglossanympna {nolocype). Photo: J. A. Campbell
0 Fig. 88. Bolitoglossa morio (3.7 km S Santa Maria Visitacibn, Solola, Guatemala, 2200 m). Photo: J.A. Campbell
84. Bolitoglossa meliana (Santa Rosa Photo: T. Bille ,QuichB, Guatemala). 3-
-.
t ?'A
I
I I
. Bolitoglossa mexicana (P.N. Pico Atlhtida, Honduras). Photo: G. Kohler
Fig. 89. Bolitoglossa mullen (Coban, Alta Verapaz, Guatemala, 1380 m). Photo: G. Kijhler
olitoglossa minutula (Jurutungo, anama, 2060 m). Photo: G. Kohler
Fig. 90. Bolitoglossa nigrescens (Cerro de la 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 k m 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 TAYLOR1941, 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 B a d in extreme eastern Oaxaca, Mexico, to Volch Atitlh 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 CAMPBEUet al. (2010).
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 (TAYLOR1952a1, 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. 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 (GRAYin 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. & WILSON Bolitoglossa porrasorum MCCRANIE 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 oresbia MCCRANSE, ESPINAL& WILSON2005, 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 psephena C ~ B E L LSMITH, STREICHEE,ACEVEDO & BRODIE2010, 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 pacaya CAMPBELL,SMITH, STREICHER, ACEVEDO & BRODIE2010, Misc. Publ., MUS.Zool., Univ. Michigan 200: 8; type locality: "4 k m (airline) SSE San Vicente Pacaya on west slope Volch de Pacaya,
& WAKE2009, Bolitoglossa pygmaea Bo-os 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
I
mg. ya. Bolitoglossa occidentalis ( W c a U L ~ U U ~ ~ , Fig. 96. Bolitoglossa pesrubra (Cerro de la Photo: G. K6hler Muerte, Costa Rica, 3300 m). Photo: G.Kohler m a p a s , Mexico, 1050 m).
.Bolitoglossa odonnelh (B'mca &I Senahfi,Mta Verapaz, Guatemala, Photo: S. M. Rovito
r lg. 97. Bok'toglossa phalarmoma (Cordillera Central,Deph. Tolima, Colombia, 1800 m). Photo: A. Acoska
I
wsa ~mnlum9anc
".
-
Fib. 1litOg10ssaplawdactyla (Veracruz, Mexico). Photo: M. Garcia Paris
Photo: 6.A. Campbell
litoglossa oresbia (Cerro Zarciadero, agua, Honduras). Photo: J. R. McCranie
Fig. 99. Bolitoglossa ponasorum (2.5 airline km NNE of La Fortuna, Yoro, Honduras). Photo: J. R. McCranie
53
Plethodontidae Cacao southeastward through the Cordilleras T i l a r h and Central and the Talamanca-Bani Massif, to the Fortuna region, Province Chiriqui, Panama, 650-2100 m elevation.
, ig. 1
Chiriqui, pan-a,
LYV-w-Y,-I
1965 m).
mg(. Photo: A. Hertz
Bolitoglossa rostrata (B~occm1883), 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. ROVITOpers. 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.
Fig. 101. Bolitoglossa sooyonun (north slope of Cerro de la Muerte, Cartago, Costa Rica, Photo: W. E. Duellman 3000 m). (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. & WAKE 2009, Bolitoglossa robinsoni BOLAROS 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 k m 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 (COPE1894), 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
Bolitoglossa salvinii (GRAY1868), Ann. Mag. Nab. Hist. Ser. 4, 2: 297; type locality: "Guatemala." Maximum SVL in males 83.0 mm, in females 96.5 mm; tail length/SVL ratio 0.92-1.03. Pacific versant of southern Guatemala 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,
t
102.BolitogIossa psepnena fiolotype). Photo: J. A. Campbell
qsa pygz
(from type Photo: G. Kijhler
i
Fig. 106. Bolitoglossa rostrata (Sierra de 10s Cuchumatanes, Guatemala). Photo:T. Bille
Fig. 107. BoLitoglossa rufescens (1 k m SSE 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 Toro, Panama, 1080 m).
Photo: M.Ponce
Fig. 109. Bolitoglossa schizodactyla (Cocl6, 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. 19181, Bull. Am. Bolitoglossa striatula (NOBLE 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. SMITR, Bolitoglossa suchitanensis CAMPBELL, STREICHER, ACEVEDO & BRODIE2010, 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. PARRA-OLEA & Bolitoglossa tica GARC~A-PAR&, WAKE2008, 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 1engthISVLratio 0.91-1.32. Northern slopes of the Cordillera de Talamanca of Costa Rica, 1745-2500 m elevation. Bolitoglossa tzultacaj CWBELL, SMITH, STREICHER, ACEVEDO & BRODIE2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 18; type locality: "17.7 k m 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 TAYLOR1951, 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 & BRODIE2010, Misc. Publ., Mus. Zool., Univ. Michigan 200: 36; type locality: "7.0 k m 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
- in Juan, Nicaragua, 30 m).
Photo: G. Kohler
Fig. 114.Bolitoglossa taylori (Serrania de Pirre, 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 Castro Blanco, Costa Rica, 1900 m).
Fig. 116.Bolitoglossa xibalba fiolotype). Photo: J. A. Campbell
Photo: G. Kohler
Fig. 113.Bolitoglossa synoria (holotype).
Photo: G. Kohler
Fig. 117. Bolitoglossa yucatana (2 km E Chichen ItzB, YucatAn, Mexico). Photo: W. E. Duellman (Courtesy of the Biodiversity Institute, University of Kansas)
3. occidentalfs 3. rufescens
r B . hartwegi
B. emmia B. daryorum r B. kaqchikelorurn m B. morio B. omniumsanctorum
A
/
-
~uatemala
Bolitoglossa
B. alberchi B. flaviventris T B. jacksoni I I B. mexicana B. mulleri 8 . odonnelli B. platydactyla ,r B. salvinii 1
Bo~oglossa
./
--
/L
7
. cuchumatana B. centenorurn B. la r B. psephena
~olito~lossa' 7
B. mombachoensis
6. biseriata robusta maanific
I. alvaradai r B. subpalmata B. pesrubra A B. marmorea B. taylori
r B. cenoen
Key to Bolitoglos~aof 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~sahelmrichi 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 variable ...........................................................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 b Coloration of dorsal and lateral surfaces of
Fig. 118. Feet (dorsal view) of Bolitoglo~~a. Photos: G. Kohler
aal surface of anterior portim of tail 6th head, bods and tail not as above; tips of conspirmous pde 1 e spots or blotche (in toes bluntly r-deed; maximum SVL ususpecimens with pa e longitudiaal band, 9 ally r66 mm ...........................,............ this band conti6uee onto tail); u s u d y 9 a Flanks usually with extensive pinkish fewer than 75 m d teeth .................... coloration iB.life con€r&dhg 6 t h the black '?manti .................................... dorsal and p a y ventral uoloratiog 8 5 4 6 Dorsum u s d y Without a conspicuous masiUary tee& 1 3 4 4vomerine teeth U) pale pattern, if auch pattern is present b Coloration in life not as above; numbers of &en w i h t conspieuovs pale large spob maxillary and vomerine teeth variable .. 11 or blotches on d a d surface of anterior 18a Limb interval 1.5-4.0; 354-3 m a d l w portion of tail and then usually amre than teeth in &, & 9 4 2 in feznales; 13-16 76 mardllary teeth ................................ 16 oomerine teeth in males, 22-81 ia females; Number of maxiUary tee+& 8543, if more SVL to 46.9 mm in males and 60.9 mm in females ...Bok'&&.lae~a o m d m c t ~ ~ @an ~ ~64~maxillary teeth, then a pldr ofdmsolateral pde stripers usudlp present, or b Limb interval 0.54.0; 35-66 m d l a r y 17 venter orange to ru~tyred ia lifg tee* 1644 vomerine tee*, SVL to 42.3 64-97; no pair teeth Number of mm in males and 62.1 nun in females ....;.. of domolakd pde a&ipes;venter brown ..................................... j%&fYgrm arapUa 21 or reddish brown in l& ....................... lla Bold pale spote and blotches on flanks and Dorsal @&ace of body with a pair of doreomost of lateral and ventral eurfaees of tail; lateral pale stripes (h~o~hlpicuous insome limb internal 0.0-2.0; 40-56 r n d a r g . specimehl,) .....................-&lE cani teeth; 18-28 mmelliDe tee* SVL to 48.9 Dmal surface of body uniform pale to dark mm in malee and 60.9 mm in females 12 brown, or with lichenous or black blot&es, b Color pattern net as above; limb inCwaI or with pale rno-g or flecking,but withvariable; numbers of maxilltiry a d ont a pair of dorsolaterd pale stripes .....M vomerine tees variable; SVL of adults Less webbing (Finger III usudly with variable ............... .................................. l3 2.0-2.1 phalanges on both aides and laa Ground color gray; not more than one phabgi%veenToes m-VI freg of webbing Fig. lanx free of webbbing on hands and feet; ll8kl ..........................iwtcglm k!molh 4 W 6 madlarg teeth; 18-26 vomezke Mare webbing (Finger III usually 1.5-2.0 teeth; SVL to 4.6.9 mm in males and 60.9 phalstnges onboth sides and beFw~eab s mm B females .......,&lib&- daqmmm IXl-Vr free af webbing) .................... :,.... 19 b Giromd d o t brom; two to slighhiy more Subcaudal surface b m or daxk brown thaD two phalanges free of we'bbing on ...............................l30lbgIloaea Q bands and feet; 47-70 m d q teeth, 23-31 vomerine t e e a ,%I% to 40.2 mm in Suboaudd d a c e pale yellm, orang;$ or males and 62.1 in females ................... m$yred in life (pale jelbw to brown in ......................................33db&wm deaora preservative^ ...........................................20 1Sa Less webbbg of hands and feet, l.M.O FYngw II shorter t b m I"mger IV; dorsal phdaagm free of webbing ..................... 14 surface of body uniform pale to dak brown, or with pale flecks; restricted to b More webbing of hands and feet, 1.0 phaMontmla de Celaque (Departments of laax or less free of webbing ................... 25 Lempira and Ocotepeqye) in westera 14a Anterior three-fonrkhs of the subcaudal Hoadwras l%M@w#8 eala~pe surface uniformly buff-yellow (in life; yelFinger II same length as or longer t h lowish brown with anly widely scattered Finger nJ; dorsal surface of body usualIy fleaka in preservative), this pale doration with lichenow or black blokches; resstricted extending onto the nntrolokral portion of to Sierra de Metapbn Range at the h r d m the tail or even onto the venColateral porof El Sdvedor, Guatemala, and Hondurtw tions of tail and body; m d l a r y teeth .................................Bol&glvssahe@mia~ aeries extending to lead of posteriol- end of orbit ............................ Bah'@omsbia Dorsal srnrfaca sf feet with small, conspicuous gold or white epots in He; mom webb Ventrc,labd portiorrua of tail and body bing (Finger LU wuafly with 0.5-1.5 p b same color as, or darker than, that of dorlanges on both sides and 1.0-1.5 phalanges sal coloration; maxillary teeth series between Toes III-VT free of webFig. &ending tp slightly beyond center of orbit 11811); restricted to northwestern .................................................................. 16 Honduras .....................B0l.i-m drmui 15a Dorsal surface of body with pde spotting or Dorsal sElrfilees of feet without small, conblotching, or a pale longitudinal band; dor-
Y
..
%-w-
..
..........
..
..
...
...................
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
m
L
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); restricted to east-central Honduras ...........
...............................Bolitoglmna longisnima
23a Dorsal surfaces of head, body, and tail uniformly 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 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 (inpart)
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~aninadom-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); num-
ber of maxillary and vomerine teeth variable; 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 (inpart) 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 (inpart) 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
2.0-2.5; SVL to 53.4 mm in females (data for adult males not available) ..................... ...............Bolitoglossa nussbaumi (inpart) 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) ...........
. I
......................Bolitoglossa pacaya (inpart) b Dorsal surfaces of head, body, and limbs
I 38a b 39a b
I I
dark gray to black with scant to moderate white stippling; flanks as dorsum or with 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 Maxillary teeth absent ...........................39 Maxillary teeth present .........................40 Longer limbs (limb interval 1.0 to 2.5) ...... .................................Bolitogllorra nrfe~ems Shorter limbs (limb interval 3.0 to 4.0) .....
...................................Bolitoglossa n y m p h
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 from brilliant yolky yellow (in life) lateral body coloration ......... Bolitoglossajacksoni 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 contrasting dark or pale blotches or longitudinal stripes ...............................................49 42a Venter immaculate pale ..............................
..............................Bolitoglossa flavivenim~ b Venter brown to black ............................43 43a Dorsal surfaces of body and tail black with
b 44a
b 45a P 12 I 1:)
Feet tdor.nl
\ 1ehv)
uf B o I l t o g ~ o i > , ~ Photo. C; K o l l l e ~
a conspicuous narrow vertebral pale (orange-yellow in life) stripe, or completely black (Fig. 89) ............Bolitoglossa mulleri Coloration not as above .........................44 Dorsal surfaces of body and tail with a conspicuous broad, yellow or orange dorsal band (Fig. 98) .... Bolitoglossa pla&damla Coloration not as above .........................45 Dorsal pattern usually including at least three colors, black or dark brown, pale (buff to orange-yellow in life) and pale brown to reddish brown (Figs. 46, 85) ..................46
r
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 ................................. Bolitoglossa mesicana orbit ......................................................... 55 Dorsal coloration consisting of less con- 53a Dorsal coloration consisting of a single pair trasting pattern, no middorsal orange-red of non-delimited dorsolateral shadings, (in life; brown in preservative) stripe; ratio moderately darker than dorsal ground tail length / SVL 0.74 in single known speccolor in a dorsal pattern otherwise homogeimen; ratio head length / SVL 0.24 ............ neously brown with dark brown mottling ........................................ Bolitoglossa indio .................................. Bolitoglossa insularis Longer limbs (limb interval 0.0-2.0 in b Dorsal coloration consisting of a variable males, 1.5-3.0 in females); head broad and number of well delimited dorsal dark well demarcated from body ......................... brown stripes ......................................... 54 ................................... Bolitoglossa alberchi 54a Venter mostly dark brown with pale Shorter limbs (limb interval 3.0-5.0 in both streaks, spots, or mottling; 14-28 maxillary sexes); head narrower and less demarcated teeth ............. Bolitoglossa mombachoed from body ................................................ 48 b Venter pale with dark brown streaks (Fig. Dorsal coloration consisting of narrow dor127a); 22-37 maxillary teeth ...................... solateral pale stripes, sometimes broken ..................................Bolitoglaea sfxiatula into rows of dots that do not meet across 55a Dorsum brown with irregularly scattered the dorsal midline or produce a reticulated pale spots or irregular shaped blotches; tail pattern (Fig. 93); snout usually black ........ 1engtWSVL ratio 0.84-0.92; 12-19 .................................Bolitoglossa odonnelli vomerine teeth .....Bolitoglossa veramcis Dorsal coloration consisting of dorsolateral b Dorsum uniform brown or brown with dark pale stripes that join anteriorly to produce brown mottling; tail 1engtWSVL ratio a pale-colored snout and that frequently 0.43-0.73; 19-41 vomerine teeth ........... 56 meet across the dorsal midline or produce a reticulated pattern (Fig. 108); snout usual- 56a Venter dark, either uniformly or with small white dots that may be visible only under ly brown ..................... Bolitoglma salvinii magnification .......................................... 57 Number of maxillary teeth <21; SVL to b Venter dirty white with a diffuse mottling 39 mm ................. Bolitoglossa m'dentalis of brown pigment, this mottling distributed Number of maxillary teeth 2G122; SVL of homogeneously or becoming more dense adults usually >40 mm .......................... 50 midventrally and forming a rather large Large robust species (SVL to 70 mm in midventral patch ....................................58 males, to 126 m in females); 80-122 maxil57a Slightly less webbing of hands and feet lary teeth; limb interval 2-3; dorsum with (Fig. 119k);in life, dorsum dark brown with well-defined dark spots, some spots comonreddish brown mottling ............................... ly centered on costal grooves on dorsolateral .................................. Bolitoglossa hartwegi portion of body ............ Bolitogllosa doffeini b Slightly more webbing of hands and feet Smaller species (adult SVL usually (Fig. 1191); in life, dorsum uniformly bown <60 mm); fewer than 80 maxillary teeth, if or pale brown with dark brown mottling ... more than 80, then limb interval less than ..................................... Bolitoglossa stuarti 2; limb interval variable; dorsal coloration variable ................................................... 51 58a Dark mottling on venter becoming more dense midventrally and forming a dark Vomerine teeth arranged in several series, brown midventral streak (Fig. 122a) ......... sometimes forming a patch; dorsum gray, ..................................... Bolitoglossa xibalba brown, or reddish brown with an irregular b Dark mottling on venter distributed homomottling of cream or pale tan; venter unigeneously (Fig. 122b).................................... form brown or tan; limb interval 2-3 ......... .................................. Bolitoglossa diaphm ................................. Bolitoglossa yucatana
b Dorsal pattern consisting of only two col-
ors, black or dark brown and pale (orangeyellow 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 ................................
b
47a
I
b
48a
b
49a
b 5Oa
b
51a
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 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~abisariata b Color pattern not conspicuously biseriate; 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 maxillary teeth, 19 vomerine teeth) .................... ....................................Bolitogloma obscura
C I
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 sur-
face 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
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 l a Almost unpigmented; internal black pigmentation of posterior abdomen and stomach showing through semitransparent ventral and lateral body wall (in preservative); very small species (SVL to 32.6 mrn in males, to 36.8 mm in females) .................... .................................. Bolitoglossa p-ea 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 m m 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 b Number of maxillary teeth 31-66; number of vomerine teeth 22-28; average adult size >37 mm SVL, no well-demarcated dark brown lateral band ................................. 15
I r 1
I
I
Fig. 123. r w a l , \UUIU~I view) of BoLitoglossa. Photos: G. Kohler
=
8. alvaradoi
Fig. 124. Feet (dorsal view) of Bolitoglossa.
Photos: G. Kijhler
two phalanges of longest digits free of webbing); dorsal coloration not as above .......... ..................................... B o l i ~ o 8 8 8h 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 threefourths the length of eye; less webbing (two phalanges of longest digits free of webbing); coloration not as above ................ 20 Number of maxillary teeth 1 9 4 5 ; 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
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
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 i n adults 37-65; Cordilleras de Guanacaste, Tilarhn, and
.................................. .............................Bolitoglos~a~ubbpalmata
Central of Costa Rica
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) ................ Bolitogl0888lgamezi SVL to 52.8 mm in males, to 57.8 mm in females; tail 1engthISVL ratio 0.96-1.04
................................. BoliWassa pembra
b 24a
b
26a
b 26a
b 27a
b 28a
b 29a
Ground color solid black with a few white ~ a t c h ein s two dorsolateral series along the :ides of the head and over the shou1;ders; head very narrow, ratio HWISVL 0.125-0.135 ...... Bolitoglwa phahwoma Coloration not as above; head moderately wide to broad, ratio HWISVL usually >0.130 ...................................................... 24 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~acuria Anterior portion of dorsal head not abruptly paler than posterior portion of dorsal head; number of maxillary and vomerine teeth variable; number of costal folds between adpressed limbs variable ........ 26 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 Hands and feet fully webbed (Fig. 124);size variable; number of maxillary and vomerine teeth variable .........................27 SVL to 43 mm in males, to 46 mm in females; 3 2 4 3 maxillary teeth; 16-24 vomerine teeth ......... Bolitoglo8~aepimela SVL to 53.3 mm in males, to 64.7 mm in females; 44-78 maxillary teeth; 1 8 4 5 vomerine teeth ...........Bolitoglos~ataylon Venter mostly dark brown to black, uniform or with pale streaks, spots, or mot28 tling ......................................................... Venter pale (cream, yellow, or dirty white) with or without dark brown streaks, or with a broad median dark stripe ..........33 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 Larger species, adults usually >40 mm SVL; number of maxillary and vomerine teeth variable Anterior ........................... 29 Dorsum with contrasting dark or pale
Fig. 126. BoLitoglossa cuna (dorsal view of head). Photo: G. Kohler
I
Fig. 127. Vent& surfaces in a. Bolitoglossa Photos: J. Sunyer striatula; b. B. insularis.
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~88med-
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 ....................32
Color pattern not as above
Fq:128. Bolitoglossa lignicolor (8 km N E Rio Chiriqui, Panama, 1210 m). Photo: G. K6hler
32a Number of vomerine teeth 14-28; venter dark brown with pale streaks ................. ......................Bolitogloma m o m b a c h o e . 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 irregularly shaped dark brown central stripe, rarely immaculate; 43-104 maxillary teeth, 27-61 vomerine teeth; terminal phalanges 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 distinct, well demarcated dark brown stripes; venter with dark brown streaks (Fig. 127a) .................................. Bolitoglma striatula
Fig. 129. Bolitoglossa sL---odactyI~ .--Cop4 Cocl6, Panama, 640 m). Photo: K.-H. Jungfer
& WAKE 1963a,b, WAKE & BRAME 1963a, b, c, VIAL1968, WAKE& BRAME1969, WAKEet al. & ALBERCH1981, HANKEN & WAKE 1982,WAKE& LYNCH 1982, 1976,ALE~ERCH
R
et il. 2008, BOLAROS&WAKE 2009, CAMPBELL et al. 2010, TOWNSE'ND et al. 2009,2010
69
Plethodontidae Bradytriton In 1983, WAKE& ELIASdescribed the new genus and species Bradytriton silus based on material collected in northwestern Guatemala. This monotypic genus is diagnosed 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 Fig. 130. Bradytriton silus (Finca Ixc hardwood forest that receives over 6000 11.0 road km E Yalambojoch, Huehuetenango, Photo: T J. Papenfuss mm of annual precipitation (ELIAS1984).It Guatemala, 1640 m). occurs syntopically with Nyctanolis pernix. ~ r a ' d ~ t silus o n WAKE & ELIAS1983,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& ELIAS1983, E m 1984
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~APmfs & 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 WAKE2000, MCCRANIE& WILSON2002, TOWNSEND & WILSON2008).
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, Photo: J. Kolby C O ~ B~onduras, ~, 1577 m).
etween 1740 and 1780 meters." SVL 31.2 mm; tail IengtWSVL ratio
, Guatemala,
1610-2290 m eleva-
rig. 133. Cryptotriton sierraminensis (from Photo: S. M. Rovito type locality).
r '
Fig. 134. C~yptotn'tonveraepacis (El Quiche, Guatemala). Photo: J. A. Campbt
I .
Fig. 135. Cryptotriton wakei (holotype). Photo: J. A. Campb
1
KeytocqFptotn'ton
I
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 2 a Venter in life yellow with a greenish cast (pale beige with gray mottling in preservative) or clear with punctate melanophores 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); vomerine teeth 8 in single known specimen;venter dark brown ............................................. .................................. Crpptotntnton '-nr 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 ..............................
I
....................................... Crpptotriton wakai
Further Reading
LYNCH & WAKE1978, PAPENFUSS& WAKE1987, 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 chujonun (from type Photo: J. A. Cam~bell localitv). "
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). Photo: J. A. Campbell
@eabove El Rincon, San Marcos, Guate-
b, 2760 m).
U%
Photo: S. M. Rovito
I &am. Dendrotriton megarhinus (Cerro Tres s,Chiapas, Mexico). Photo: S. M. Rovito
-
Fig. 142. Dendrotriton sanctibarbarus (ca. 3 k m S San Luis 10s Planes, Santa Barbara, 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, [Departamento de] San Marcos, Guatemala." Maximum SVL 37.9 mm; tail 1engthISVLratio 1.11-1.35. Southern slopes of VolcAn Tajumulco and adjacent volcanic highlands in the Departamento de San Marcos, southwestern Guatemala, 1700-2700 m elevation. Dendrotriton chujorum C ~ B E L LSMITH, , STREICKER, ACEVEDO & BRODIE2010, 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 Ixtatzin to Barillas road, Mateo 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.
I I
Fig. 143. Dendrotriton xolocalcae (Reserva de la Biosfera El Triunfo, Chiapas, Mexico, Photo: S. M. Rovito 2000 m).
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 & BRODIE2010, 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.
Fig. 144. Dendrotn'ton rabbi (Montaiias de Pefia Blanca, Huehuetenango, Guatemala, Photo: S. M. Rovito 2974 m).
Western Guatemala (Montaiias de Cuilco and Sierra de 10s Cuchumatanes), 2100-2700 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.
Dendrotriton sanctibarbarus (MCCRANIE& WILSON1996), 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, 1830-2745 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.
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.
I
I
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 INostrils 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 IengthJSVLratio 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, COLLINSBUTH1990, M c C m & WILSON 1998, MCCRANIE& WILSON2002,
Fig. 145. Dendrotriton chujorum (a. male; b. female); c.D. kekchiorum (c.). Photos: G. Kohler
A D. sanctibarbams
a
D. xolocalcae
Nototriton
Fig. 146. Nototriton abscondens (El Angel, Costa Rica). Photo: M. Franzen
Fig. 147. Nototriton barbom. (Montana ae Macuzal, Yoro, Honduras, 1780 m). Photo: J. T. Townsend
'
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 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 the mosses where the females deposit their clutches of 1-19 eggs (BRUCE 1998, JOINSH & 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. 148. Nototriton brodiei (from type locality). Photo: J. A. Campbell
Fig. 149. Eggs of Nototriton picadoi (P.N. Tapanti, Cartago, Costa Rica). Photo:W. Van Devender
Plethodontidru
I months; hatching is completed by of the wet season in December inabandoned hummingbird nests.
Fig. 150. Nototriton gamezi (Monteverde, Costa Rica). Photo: M. Franzeu
h t r i t o n abscondens (TAYLOR1948), 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
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.
Volch Cacao (elevation 1580 m), t e National Park, Prov. de Guanat a Rica." Maximum SVL 34 mm; tail
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.
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& AUSTIN2010, 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.
Nototriton major GOOD & WAKE 1993, Herpetol. Monogr. 7: 137; type locality: "along Quebrada Platanillo, approximately 6 k m 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. Northwestern portion of the Cordillera de Talamanca, Costa Rica, 1200-2200 m elevation. 19491, Univ. Nototriton richardi (TAYLOR 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 KO2002, 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 1engtWSVLratio 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& WAKE1993, Herpetol. Monogr. 7: 134; type locality: "Rio Quiri (elevation ca. 1300 m),0.3 km northeast
*
N. limnospectator
N. tornamorurn
Plethodontidae
to Nototriton
a Hands and feet syndactylous, fingers and
b
b
a
. b
a
b
a
b a
toes not separate from one another, without subterminal pads; ratio 5th toe length / SVL <0.004; tips of digits pointed ...........2 Fingers and toes separate from one another 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 Nostrils larger (ratio nostril width / SVI >0.017); hind foot narrower (ratic HFWISVL <0.039) Nototn'ton t o m a m o m Nostrils smaller (ratio nostril width / SVL <0.017);hind foot broader (ratio HFWISVL >0.039) ...................................................... 3 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 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 i d u d i Maxillary teeth >58; ratio nostril width / SVL 0.004-0.005 (Fig. 153a) ....................... ........................................ Nototriton bnniiei Maxillary teeth <56; ratio nostril width / SVL variable ............................................. 5 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-
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' (inpart) 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 . 0 7 in males and d . 0 1 in females ........................
on .....................................N ~ t o ~ t 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 ....... l l a 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 b Maxillary teeth 27-38; vomerine teeth 8-18; occurring in Costa Rica ................14 13a Nostrils smaller (ratio nostril width I SVL 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 narrow pale (cream to silver) &ripe sipsrating 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 latera1 stripe ....Notokiton barbod (inpart) 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 Further Reading
Photo: M. Franzen
Fig. 155. Nototriton limnospectator (Cerro h u l M e h b a r , Comayagua, Honduras, Photo: J. T. Townsend 1650 m).
Fig, 156. Nototriton richardi (Villa Calas, Alajuela, Costa Rica, 1920 m). Photo: G. Kohler
MCCRANIE &WILSON 1992, Goon & WAKE1993, 1 MCCRANIE 1996b, BRUCE1998,1999, CAMPBELL & SMITH 1998, JOKUSH & GARC~A-PAR~S 1998, Fig. 157. Nototriton saslaya (holotype). GARC~A-~A &RWAKE ~S 2000, MCCRANIE& Photo: G. Kiihler: W n s o 2002,2003, ~ TOWNSEND et al. 2010
-me-nrtidae
enus Nyctanolis was proposed by & WAKE (1983) to accommodate a mala and Chiapas, Mexico (ELIAS differs from all other tropical
. It
& WAKE 1983). Most of the speciof the type series were collected
San Fig. 16G. IV~LWUIOL~S pernix Jose Maxbal, Huehuetenango, Guatemala, Photo: S. M. Rovito
IgSa,
& WAIZE in & ~ A T (eds.), A Adv. Herpetof. Evol. ; type locality: "Finca Chiblac, 10 km
ofis pernix
(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. , , 88
Nyctanolis pemix
Nyctanolis pernix corn type locality). Photo: J. Hanken
Further Reading ELIAS& WAKE1983, ELIAS1984
C~ig. 159. Nyctanolis pernix (3.8 k m SE
t
_?urulha,Baja Verapaz, Guatemala). Photo: J. A. Campbell (Courtesy of the Biodiversity Institute,University of Kansas)
r
Plethodontidae
1
Fig. 161. Oedipina uniformrs (Albergue Montemeal, near P.N. Juan Castro Blanco, Alajuela, Costa Rica, 1900 m). Photo: G. Kohler
I
Oediplna
and inside rotten logs, beneath rocks, or in leaf litter and decaying plant material. The species of the genus Oedipina stand They have also been reported to live under out among all other Neotropical salaman- slabs of bark on fallen trees and in insect ders because of their extremely slender, burrows within rotten logs (SAVAGE 2002). elongate bodies, tiny limbs and very long According to M. RYAN(pers. com. August tails. Whereas the representatives of all 2010), individuals of Oedipina alleni and other tropical salamanders have 14 trunk 0.savagei have been observed at night on vertebrae, this number is 18 or more in the the upper surface of leaves up to 1.5 m species of Oedipina (GARc~A-PAR~S & WAKE above the ground during the rainy season. 2000, MCCRANIE et al. 2008). Oedipina is If one of these salamander is touched, the distributed from southern Mexico to north- usual reaction is body flipping. It is western South America with the majority assumed that no parental care of eggs of species (18 of 29) occurring in Costa Rica takes place in the species of Oedipina and Panama; only two species of this (GOOD & WAKE1993). genus, 0. complex and 0. parvipes, have invaded South America. The species of Oedipina alfaroi DUNN 1921, Proc. Biol. SOIL Oedipina form a well-supported (by morWashington 34: 144; type locality: "Zent, , phological and molecular data) mono[Cantdn de Matina, Limdn Province,] Costa phyletic group divided into three subgenRica."Maximum SVL 60 mm; tail 1engthISVL [ era: Oedipina (usually with 20-23 trunk ratio 1.27-1.50. Atlantic versant of Costa Rica . ! and extreme northwestern Panama, near sea vertebrae; western Guatemala to Panama), level to 850 m elevation. Oeditriton (with 20-21 trunk vertebrae; Honduras and Nicaragua), and Oedopinola Oedipina alleni TAYLOR1954, %v. Sci. Bull. 36: 607; type l o d t y : "Palma.Mortef, , (usually with 18-19 trunk vertebrae; Canth de Osa], Puntarenas Province, C Chiapas, Mexico, to Ecuador). ~ i c ~aa ~ x i m u mSYL in males 4 mm, i&j They range from sea level to elevations in females 58 mm; tail 1engthlSVL ratid' excess of 2200 m. These semifossorial or 1.27-1.63. Pacific versant of southwester0 fossorial animals are usually found under
'
d
Plethodontidae
Fig. 162. Oedipina alleni (near Parrita,
Puntarenas, Costa Rica, near sea level). Toto: 1,eenders
b~g.163. Oedipina carablanca (Pocora, Costa Photo: B. Kubicki Rica, 250 m).
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, Photo: A. Hertz Veraguas, Panama, 885 m).
m
m
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 k m 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 leptopoda MCCRANIE,VIEITES & WAKE2008, 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 Oedipina gracils TAYLOR 1952a, Univ. h a s 46.2 mm, in females 44.3 mm;tail lengthlSVL s, B,. 34: 773: ,t locali,: ratio 1.47-1.89. Isla Escudo deveraguas in the ~ i a m a n t e s , one mile sduth of ~ u b ~ i l e s , Bocas del Toro Province, Panama; close to?ea [Canhjn de Pocod, Provincia de L i m 6 ~ 1Costa level. ; Rica." Maximum SVL in males 44 mm, in Oedipina nica SUNYER,WAKE, TOWNSEND, females 47 mm; tail length/SVL ratio TRAVERS,ROVITO, PAPENFUSS,OBANDO& 1.39-2.57. Atlantic lowlands of Costa Rica and K ( 1 m 2010, Zootaxa 2613: 32; type IocaEty: :xtreme northwestern Panama, sea level to "El Gobiado, Reserva Natural Cerro Datanli110 m elevation. El Diablo, 13"09'N, 85"52W, 1420 m above sea Oedipina grandis BRAE~E & DUELLMAN 1970, level (a.s.1.1, Dept. Jinotega, Nicaragua." Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. Maximum SVL in males 44.5 mm, in females 201: 1; type locality: "northern slopes of Cerro 48.5 mm; tail lengthlSVL ratio to 1.92. NorthPando, between 1810 and 1930 m elevation central Nicaragua, 1360-1660 m elevation. (5937-6330 feet), Rovincia de Bocas del Toro, Oedipina pacxcensis TAYLOR1952a, Univ. 1 extreme western Panama near the border with Kamas Sci. Bull. 34: 775; type locality: 'about Costa Rica." Maximum SVL in males 68 mm, two miles east of San Isidro El General," Costa in females 72 mm; tail length/SVL ratio Rica. Maximum SVL in males 48 mm, in 1.78-2.23. Cordillera de Talamanca in extreme females 51 mm; tail IengtWSVL ratio southern Costa Rica and adjacent western 1.39-2.57. Pacific slope of southern Costa Rica Panama, 1810-1950 m elevatien. and adjacent western Panama, sea level to 730 Oedipina ignea STUART 1952, Proc. Biol. Soc. m elevation. Washington 65: 1;type locality: "along the Rio Oedipina p m p e s (PETERS 18791, Monatsber. I Las Brisas, just south of Yepocapa, DepartPreuss. Akad. Wiss. Berlin 1879: 778; type ment of Chimaltenango, Guatemala. locality: "Laceres [= CBceresl am Cauca, einem Elevation, about 1450 meters." Maximum SVL Nebenfluss des Magdalenenstroms in der in males 61.7 mm, in females 62.1 mm; tail Provinz Antioquia (Neu Granada)", Colombia. length/SVL ratio 1.87-2.97. Pacific versant of Maximum SVL in males 47.5 mm, in females south-central and southeastern Guatemala as 57.7 mm; tail length/SVL ratio 1.30-1.77. well as southwestern Honduras, 1000-2000 m Central Panama to Choc6 region and the lower elevation. Cauca Valley in Antioquia Department of Oedipina kasios MCCRANIE, VIEI'EES% WAKE northwestern Colombia, near sea level to 865 2008, Zootaxa 1930: 11; locality: "near m elevation. Quebrada Pinol, 15"07'N, 86"44W, Parque Oedipina paucidentata BRAME 1968, Nacional La Muralla, 1190 m a.s.l., Dept. Herpetol. 2: 41; type locality: "near El Olancho, Honduras." Maximum SVL in males Empalme, (along the Pan American Highway, 37.5 mm, in females 46.8 mm; tail 1engthISVL south of Cartago) in the northern end of the ratio 1.6-1.8+. Endemic to Parque Nacional La Cordillera de Talamanca, Cantiin de Dota [in Muralla in northwestern Olancho, Honduras, error, Cant6n de El Guarcol, Provincia de 950-1780 m elevation. Cartago, Costa Rica, about 2286 m (7500 8 )
,
7
b.Oedipina fortunensis (holotype). Photo: G. Kohler
Fig. 171. Oedipina kasios (from type locality). Photo: J. R. McCranie
-2. Oedipina gephyra (from type locality). Photo: J. R. McCranie
Fig. &, 2. Oedip-a .,topoda (Los Pinos, Cort6s, Honduras). Photo: J. R. Collart
C
I 2060 m).
Photo: G. Kohler
Fig. 173. Oedipina nica (KilamM, Jinotega, Nicaragua, 1490 m). Photo: G.Kohler
85
gethodontidae ?levation."Maximum SVL in males 52 mm, in kmales 62 mm; tail 1engtWSVL ratio
,em of northeastern Honduras, 70-540 m el+ vation.
3edipinapoelziB~~lll~ 1963,Contrib. Sci. Nat. list. Mus. Los Angeles Co. 65: 7; type locality: 'area of the stone quarry in the W t y of El ingel Waterfall (former American Cinchona 'lantation and Isla Bonito of TAYLOR1952),3. 5 miles by road south of Cariblanco (5 miles iorth of Vara Blanca), along the Vara Blanca-Puerto Vieio Road, Provincia de Heredia-Provincia "de ~ l a j u e l aboundary, :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
Copeia 2000: 52; type locality: "Finca Las( Cruces, 6 km S San Vito de Java, Prov. Puntarenas, Costa Rica, 8"47'35"N, 82"57'30W, approximately 1200 m elev.." Maximum SVL in males 38.5 mm, in females 39.3 mm; tail length/SVL ratio 1.08-1.86. Pacific versant in extreme southern Costa Rica and adjacent Panama. 1200-1400 m elevation. 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 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 (3400R)." 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. et al. (2010) pointed out that Yowever, SUNYER 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-
-
. r
7
,.
--
r
Oedipina stumti BRAME1968, 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 WILSON2002). 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), Department 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
;
E
si MCC2006,J. Herpetol. locality: "near Sendero de e Nacional El Cusuco, 15"30'N, tW014W, 1800 m elevation, Departamento de €!at&Honduras." , Maximum SVL in males 57.5 mm, in females 62.9 mm; tail 1engthlSVL ratio 1.90-2.03. Only known from the type U t y
Costa Rica).
Photo: M. Garcia Paris
Oedipina m'formi's K E F E 1868a, ~ Arch. Naturgesch. 34: 299; type locality: "Costarica." Also described as new by KEFER~TEIN(1868b: 331). Maximum SVL in males 57 mm, in females 55 mm; tail length/SVL ratio 1.56-2.85. In Costa Rica from V o l h Tenorio south through the cordilleras and Meseta Central to the Panaman border, 750-2150 m elevation.
Fig. 178. ueaipina quadra (P.N. Pico Bonito, 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 Photo: M. Franzen
rlg. lau. uempma comas1 (r.lu.busuco, Cortes, 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
b
2 a b
3a b
4 a
(ratio HFWISVL ~0.020);head narrow (ratio HWISVL ~0.070);maxillary teeth 0-13 ................................. Oedipina taylori Costal grooves 17-20; hind foot broader (ratio HFWISVL usually >0.020); head broader (ratio HWISVL >0.070); maxillary teeth 0-98 .................................................2 Costal grooves 17-18 ...............................3 Costal grooves 19-20 ...............................6 Maxillary teeth absent; hind foot broader (ratio HFWISVL usually >0.030); limb interval usuallyc9 .......Oedipina elongata Maxillary teeth 18-58; hind foot narrower (ratio HFWISVL usually ~0.030);limb 4 interval 9-14 ........................................ Limb interval 12-14; maxillary teeth 18-37; vomerine teeth 9-18 ....................... ....................Oedipina stenopdia (inpart)
A
+ 0.pseudouniformis
0.carablanca
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) ...................
..........................................
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;
.......................... ................................... Oedipina fortunensis
ratio HUSVL 0.165-0.180
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) ..... ......................... Oedipinapseudouniformis b All digits tightly fused (digits can be discerned 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 (inpart) 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
b Maxillary teeth <20, or if r19 then ratio 3a b 4 a
b
5a b 6 a
HUSVL ~0.170;number of vomerine teeth variable .................................................... 3 Head broader (ratio HWISVL 0.104-0.119) 4 Head narrower (ratio HWISVL 0.085-0.106) .............................................. 6 Maxillary teeth absent; vomerine teeth 10-12 ........................Oedipina carablanca Maxillary teeth 7-45; vomerine teeth 11-32 ....................................................... 5 Snout more bluntly rounded; limb interval 7.0-9.5; maxillary teeth 1345; vomerine teeth 17-32 ................... Oedipina complex Snout less bluntly rounded; limb interval 6.0-7.0; maxillary teeth 7-18; vomerine teeth 11-22 .................... Oedipina savagei 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 surface 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 interval variable ............................................. 10 10a Venter darker (usually black) than dorsum of body (brown),and separated by a creamy or silvery lateral stripe .......................... 11
,
i 1
Plethodontidae
1
4
n
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)
1
I
Fig. 181. Dorsal view of head in a. Oedipma altura; b. 0.cyclocauda; c. 0.paucidentata; d. 0. Photos: G. Kohler savage5 e. 0. uniformis.
15a Head long and relatively narrow (ratio maxillary HWISVL 0.075-0.095; Fig. 181~); than dorsum of body (black to blackish gray), teeth 14-43 ........... Oedipina paucidenteta no creamy or silvery lateral stripe ........... 12 b Head short and relatively broad (ratio l l a Head broader (ratio HWISVL 0.100-0.118); HWISVL 0.083-0.100; Fig. 181e);maxillary hind foot broader (ratio HFWISVL teeth 29-55 ............................................. 16 0.035-0.040) ...................... Oedipina poelP' b Head narrower (ratio HW/SVL 16a Dorsum and venter grayish black, often with a whitish area in parietal region 0.085-0.100); hind foot narrower (ratio andlor near limb insertions or joints ......... HFWISVL 0.021-0.28) .. Oedipina grandis .................................. Oedipina p a d & 12a Head broader (ratio HWISVL 0.100-0.113); b Coloration usually not as above, usually hind foot broad (ratio HFWISVL head same color as dorsum or head in gen0.029-0.037); vomerine teeth 17-34 .......... .......................... O e d i e a pseudomif6rmis eral slightly paler than dorsum and without a whitish area in parietal region .... 17 b Head narrower (ratio HWISVL 0.075-0.100); hind foot usually narrower 17a SVL to 44 mm in males, to 47 mm in females; occurs on Atlantic lowlands (sea (ratio HFWISVL 0.013-0.035); vomerine level to 710 m elevation) Oedipina gracilis teeth 9-26 ............................................... 13 b SVL to 57 mm in males, to 55 mm in 13a Snout long and narrow, tip truncate (Fig. females; occurs in uplands (750-2200 m 181a); maxillary teeth 19-27; vomerine elevation) .................... Oedipina d r m i e teeth 9-17; dorsum grayish black with some small white spots on head, neck, shoulder, and upper limb surfaces; venter black ................................. Oedipina altura b Snout short, rounded (Fig. 18lb,c,e); maxilRelary teeth 14-55; vomerine teeth 10-26; 1963, 1968, Bw & DUELLMAN 1970, coloration usually not as above ............. 14 Bw 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 W1997, GARC~A-PAR~S & WAKE 2000, SVL) .......................... M P d m~ u d a 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
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r!
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Fhe genus Pseudoeurycea comprises about 60 species, inhabiting the cloud forests and humid pine-oak forests in northern Mexico bnd Guatemala, usually above 1200 m eled o n (WILSON& JOHNSON 2010). It is a q o u p of moderate-sized to very large salamanders with well-developed hands and && in which the middle digits me markedItp longer than the outer ones. Some species 4 Pseudoeulycea can be locally abundant, leaching densities higher than those of any other Guatemalan salamander (ELIAS El&&).Most species have very restricted \pographic ranges. As pointed out by { ~ C & H WAKE (1989), the genus Pseudoauycea, as currently constituted, appears to be a taxonomic group of convenience and not necessarily a natural unit. Indeed, nost species assigned to this genus are h a c t e r i z e d mainly by their lack of derived traits that define other plethodontid genera. FROST et al. (2006a) placed Wotn'ton in the synomymy of Pseudoauycea. These salamanders have been h d beneath and within logs, under bark, smss, grass clumps, and rocks and were b m e d active in the open on wet nights @LIAS
doeuqcea exspectata STUART 1954, Proc. 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 1engtWSVLratio 0.85. Only known from the type locality.
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-i * --". * . Fig. 183. Pseudoeurycea brunnata (3 k m W of El Rincbn, Quetzaltenango, Guatemala, ca. Photo: J. Hanken 2400-2500 m).
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L
Fig. 184. Pseudoeurycea p m a (Cerro Badl, Chiapas, Mexico, 1900 m). Photo: T.J. Papenfuss
7--
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~twardas far as V o l b Chicabal (ins ~ u t h Guatemala), 2400-2800 m elevation.
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Fig. 185. Pseudoeurycea goebeli (Lagu Chicabal, Quetzaltenango, Guatemala, 2722 m). Photo: S. M. Rovito SVL 59 mm; tail length/SVL ratio 1.08. Soconusco 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.
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Pseudoeuiyeea pama L& WAI(E1989, 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 isjust 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 Cuchumatanes and the southwestern highlands of Guatemala; on Volch Tacani it also occurs on the Mexican side, 2450-4000 m elevation.
Fig. 186. Pseudoeuzycea rex (Sierra de 10s' Cuchumatanes, Guatemala). Photo: T. BiUe
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~8udoeulyCeabluzulata b Coloration in preservative: dorsal surfaces pinkish gray, sides gray with or without white marbling, belly pale gray .................. ....................................... Pseudaeuzycea mx
Fig. 187. Pseudoeulycea rex (Caserio Monzo, San Marcos, Guatemala, 2785). Photo: S. E. Rovito
Further Reading
WAKE& JOHNSON 1989, LYNCH& WAKE 1989, et al. 2001, PARRA-OLEA 2008 1999, PARRA-OLEA
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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 1cm 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,
with an aquatic tadpole going through metamor~hosis to transform to an airbreathing 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 Craugastoridae, Eleutherodactylidae, and Strabomantidae), 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.
accessory palmar tubercles
subarticular tubercles
n
outer metatarsal tubercle
/
n
A
plantar tubercles
~
interdigital webbing
~
,.~
\
g
webbing i t
inn;
metatarsal tubercle
subarticular tubercles
webbing formula: 12-2 314 I l l 1/2-8 111 1 314-3+ IV 3 1 B - 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
a
l
mops pu&zbus; note t m d tubercle (arrow), Drawing M.V e d y Fig 192. Hind f i t of
Fig. W .Eind foot af B b i ~ o p ~ dums& note spade-like inaer metatarsal tubenrclas bmws). Drawing: M . B d y
Fig;. 1193. E&popa&w vm'alo~qe;note Imamverse fold of akin across head posterior to ayes Carrowl.
Photo: G. K~hler
Fig.190. Forefoot af Daadrobab auratuq note pairs of acute-Eke flaps ~ m o w s ) . Drav&g
M,Vedy
Fig. 194 IF~padeFanapmmIbIepm Wl and
H~obatrachiwaf l transparent VB.XI~@~~.
. Fig. 191. hcilius valliceps (near San Jod El Idolo, Sucbitap4quez, Guaternda, 100 rp); note enlarged pardtoid glad@Earrow4). PBotc G. KBhler
d 4 b (right); note
Photo: G. mhlet
Fig. 195.Tip of Ping- (lahrd view) is a. C r r t m b r n d e i (note digtd groava); b. Leptcvda&~Iusbolidaw. Photos: G.KiMw
Anura Key to families of h g s 1 a Four toes; two elongate spade-like inner
J
metatarsal tubercles, each with a free edge (Fig. 189) .......................... Rhinophrynidae 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; 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). tongue present .......................................... 3 Photo: G. Kiihler - A cons~icuoustarsal tubercle resent at end of inner tarsal foldA(Fig.192); 108 A circular or V-shaped ventral disc present upper jaw without teeth; dorsum extreme(Fig. 196);fingers without webbing ............ ly warty ........................................................ Terrarana (families Craugastoridae, ........... Leiuperidae (in part: Eqgystamops) Eleutherodactylidae, and Strabomantidae) No conspicuous tarsal tubercle; upper jaw b No ventral disc; finger webbing present or with or without teeth; texture of dorsum absent ...................................................... 11 variable .....................................................4 l l a Venter transparent, internal organs at A prominent deep purple to black colored least partially visible externally (Fig. 194); lumbar gland, usually with pale (white to eyes directed forward at about a 45 degree pale blue in life) central spot (Fig. 672) ...... angle ................................... Centrolenidae ............. Leiuperidae (in part: P l e d e m a ) b Venter not transparent, internal organs No such colored lumbar gland present ... 5 not visible externally; eyes oriented more Well-developed parotoid glands present laterally ................................................... 12 (Fig. 191) ................................................... 6 12a Fingers not conspicuously long and Parotoid glands absent ............................ 7 unwebbed, or, if so, males with nuptial excrescences; degree of finger webbing Fingers with enlarged discs; pupil verticalvariable; no triangular dermal flap on each ly elliptical in bright light (in life) or eyelid; no brood pouch in females; no fleshy preservative ................................................. proboscis ......................... Hylidae (inpart) Hylidae (in part: PhgIIomedusa) b Fingers conspicuously long, unwebbed, Fingers without enlarged discs; pupil horilacking nuptial excrescences in males; a zontally elliptical in bright light (in life) or triangular dermal flap on each eyelid prepreservative ................ Bufonidae (in part) sent or absent; a brood pouch in females A pair of scute-like flaps on upper surface present or absent; a fleshy proboscis preof tips of fingers and toes (Fig. 190) ........... sent or absent .................. Hemiphractidae ............................................. Dendrobatidae 138 A distinct ventral disc present (Fig. 196); No pair of scute-like flaps on upper surface no toe webbing ................. Leptodactylidae of tips of 6ngers and toes ......................... 8 b No ventral disc; toes with substantial webA transverse fold of skin across head postebing .......................................................... 14 rior to eyes (Fig. 193); one or two fleshy ridges across posterior region of roof of 14a A distinct tympanum present; upper jaw with teeth ...................................... Ranidae mouth ..................................... Mimhylidae b Tympanum absent; upper jaw without No transverse fold of skin across head posteeth ........... Bufonidae (in part: ate lop^) terior to eyes; no fleshy ridges across posterior region of roof of mouth ...................... 9 A digital groove completely around tips of Further Reading digits (Fig. 195a); finger discs usually pre- TAYLOR& GUTTMAN 1977, DUELLMAN & TRUEB sent, toe discs always present ............... 10 1986, FEDER& BURGGREN 1992, STEBBINS & No digital groove completely around tips of COHEN 1995, MCDLARMID& ALTIG 1999, 1999, FROSTet al. 2006a, WELLS digits (Fig. 195b); no finger discs, usually DUELLMAN 2009 no toe discs ........................................... 13 2007, OGIELSKA
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Bufonidae
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The nearly cosmopolitan true toads (family 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., ~ i197. ~ 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 Key to the genera of Bufonidae Rhinella) occur in Central America. 1 a Finger I shorter than Finger 11; tympanum absent or indistinct
..................................2
In Central America, 39 bufonid species in b Finger I same length or longer than Finger seven genera are recognized. SynapoII,tympanum indistinct to prominent ....6 morphies of the Bufonidae include the 2 a Inner digits rudimentary, enclosed by presence of a Bidder's organ (rudimentary fleshy web; fingers (except 111) and toes (except IV)nearly fully webbed .................. ovaries present in males; absent in some ............................................. CrepPdo&pe D e n d r o p ~ s c u s ) parotoid , glands, and an edentate maxilla and premaxilla (FORD b Inner digits well developed; fingers and toes not fully webbed ............................... 3 & CANNU 1993, PRAMUII 2006). Many 3 a Skin generally smooth, but often with bufonids have thick glandular skins with minute pustules dorsally and with wellpustular warts and concentrations of glandeveloped spicules along anterior flank; dular tissue. When stressed, these toads arms very long and slender; no cranial crests ............................................Ateloprte can secrete appreciable quantities of a visb Skin of dorsum rugose with numerous, cous white poison from their parotoid scattered, small to medium-sized tubercles glands. andlor warts; arms of moderate length and less slender; cranial crests present .........4 Most species of this family are crepuscular 4 a Tympanic annulus absent; vocal slits or nocturnal, but some are diurnal. The absent in males ........................................... majority of bufonids are terrestrial or ...................RhiaeUa (inpark R. amlopha) semifossorial; exceptions include some fosb Tympanic annulus present, might be consorial and a few scansorial or semiaquatic cealed by skin; single sinistral vocal slit present in males .......................................5 species. 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 (inpart) b Tympanum indistinct, concealed by skin; dorsal tubercles rounded; ground color grayish brown; smaller toads, SVL in adult
Atelophryniscus
Fig. 198.Amplecting pair of hcilius canaliferus (San Lorenzo, Suchitep6quez, Guatemala, 210 Photo: G. Kiihler m).
males 3540 mm, 49-60 mm in females
........................................... Atdo-
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I
1
t
I
6 a A well-developed i ~ e tarsal r fold present; parotoid gland very large, several times the area of upper eyelid .................................. 7 b Inner tarsal fold absent; parotoid gland usually small to moderate-sized, less than twice the area of upper eyelid .................8 7 a Cranial crests present; no dark brown to black lateral longitudinal band from head to at 'east posterior margin of ~arotoid gland ......... Chaunue (in part: C.manhue) b No cranial crests; a dark brown to black lateral longitudinal band from head to at least posterior margin of parotoid gland ...
.......................................................... h
A t e l o p ~ s C L chrysopho*s Is
was Collect-
ed along a stream; during daytime, males were encountered sitting on rocks or bodthe stream and, ders in when disturbed, they would jump into the stream and swim to the bottom to conceal themselves among debris in the bottom of the pools; at night, males were observed in the vegetation along the &ream, 1-2 m above the ground (MCCRANIE & WILSON 2002). Am~lectantpairs were found in m i d - F e b r u ~during a period of no precipitation for several weeks (MC~RANIE &
8 a Pretympanic and parietal crests usually prominent ........................InciIiue (in part) b Pr&ympanic and parietal crests usually indistinct ................................................... 9 9 a Nostrils oriented laterally or dorsolaterally; supratympanic crest prominent and 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 (inpart: C.m W )
BLAIR1972, HOO&OED I
....
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. veraguensis group (some 2600 b), md because no molecular data are available to support this taxonomic decision, 1 prefer to recognize the genus Atelopluyniscus as valid until additional evidence has been published.
1989, GRWBEAL & DE QUEIROZ 1992,PAULY et al. 2004,MULCAHY et al. 2006,PRAMUK 2006,PRAMUK 2008,FROST et al. 2009
-
Fig- lg9.Atelopfiscus chWsophoms (Quebrada de Oro, Atlhtida, Honduras). Photo: J. R. McCranie
WILSON2002). 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).
The genus Atelopus is a tropical New World endemic and contains 89 currently 2010), collectiverecognized species (FROST ly known as "harlequin frogs," which are found in the uplands and lowlands from Atelophryniscus chrysophorus MCCRANIE, Costa Rica south to Bolivia and eastward WILSON & W m s 1989, Occas. Pap. Mus. into the Guyanas. The adults of most Nat. Hist. Univ. Kansas 129: 3; type locality: species typically exhibit gaudy, aposematic "Quebrada de Oro (15'383, 86'47W), elevation 880 m, tributary of Rio Viejo, south slope colorations with yellow, orange, red, green of Cerro BMalo, Cordillera de Nombre de Dios, andlor black elements. On the contrary, Depto. Atliintida, Honduras." Males to 40.1 metamorphs and juveniles of some species mm,females to 59.6 mm.Central and western are cryptically colored (e.g., in juvenile A. portions of the Cordillera Nombre de Dios on zeteki, olive green with dark brown blotchthe Atlantic versant of north-central Honduras, 750-1760 m elevation. es). In A. zeteki, the cryptic juvenile coloration is apparently correlated with the absence of toxic skin compounds (LINDQUIST & HETHERINGTON 1998a). The r Atelophryniscus chrysophorus 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 cardiotoxic (steroids) (SAVAGE 2002).
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C
F'urther Reading
MCCRANIE et al. 1989, LAVILLA & DE SA 2001, MCCAANIE & WLLSON2002, MCCRANIE AND C A S T ~ D2005,2007, A PRAMuK & L E ~ R2005
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 1m 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.
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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 ~ oDaribn, ,
Atelopus species have suffered eclines in the past 10-15 years, the Central American species CA & L O ~ R1997, S SAVAGE 2002, WCA et al. 2005). i s certus BARBOUR1923, 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 elevachirripoensis SAVAGE& BoLARoS Biol. Tropical 51: 382; type locality: southeastern side of Norte, ca. 4 k m 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). Photo: M. Garcia Paris
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 SHREVE1936, 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.
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.
Atelopus senex TAYLOR195213, 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 LOITERSet 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
Fig. 204. Atelopus glyphus (Cana, Daribn, Photo: A. J. Crawford Panama, 1300 m).
b Dorsal coloration not as above; males without a lateroventral black line ................. 5 1 a Legs short, ratio shank length1SVL ~ 0 . 3 5 ; pretympanic and temporal regions, flanks, 5 a Dorsum brown (in life?), uniform or with lateral and upper surface of upper arm, yellowish white spots ................................. upper surfaces of thigh, sacral region, and ........................................Atedopus gIpp6us area lateral to vent with pale tipped coni . b Dorsal coloration not as above ............... 6 ................................. Atelopus Wpoenaia 6 a Dorsum golden yellow in life with transb Legs longer, ratio shank length/SVL >0.35; verse black bands, with black blotches, or without coni or spiculae in regions listed uniform golden yellow .....Atelopus eeteHi above ......................................................... 2 b Dorsal coloration mostly black or chocolate 2 a Coloration in life brick red with black spots brown with yellow to red bands and blotch(pale reddish with black spots in preservaes ...................................... Atelopus v d u s tive) ................................... Atelopus certus 7 a Glandular areas well developed; coloration b Coloration not as above ........................... 3 in life black, bluish gray or gray-green, 3 a No differentiated glandular areas, skin with green to pale orange spots and blotchsmooth throughout; tips of fingers slightly es, or uniform, some individuals with an ...................................................... swollen 4 orange suffusion ............... Atelopus senex b Skin of rostrocanthal area, dorsum and b Glandular areas weakly developed; colknee region glandular; tips of fingers taperoration in life yellow, green, rust, or black ing .............................................................. 'I with red or yellow spots, blotches, or lines ...............................Atedopus ~ q r u q r u e n s i 6 4 a Dorsum dark greenish olive to chocolatebrown in life, without blotches or other markings; males with a lateroventral black line; juveniles lime green in life with dark brown markings ............. Atelopus limhsus
Fig.203.ntelopus limosus (Burbayar, Cuna, Panama). Photo: B. Kubicki
Fig. 210.Atelopus senex (Cemo Chompipe, Heredia, Costa Rica). Photo: W. Van Devender
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
I
r'ig. 213. Atelopus glyphus.
Photo: B. Wilson
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, SAVAGE & B o ~ o S2009b9 PAUL,et 2009, VANBOCXLAER et al. 2010). Contrary to the proposal of CHAPARRO et al.12007), I here follow SAVAGE & B O ~ O(2009b) S 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).
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; timFurther Reading STARRETT 1967, DUELLMAN& LYNCH 1969, ing seems to assure that metamorphosis SAVAGE 1972a, CRUMP1986, POUNDS & CRUMP takes place during periods of high humidi1987, CRUMP 1988, C R & ~POUNDS 1989, COCROFT et al. 1990, IBAREZ et al. 1995, ty and prey abundance (ZUG 1983). L~DQUIST 1995, LINDQUIST & ~~ETHERINGTON Females produce paired strings with sever1996, LOTTERS 1996, LINDQUIST & SWIHARTal thousands of eggs in the shallow water 1997, LA WCA & LBTTERS 1997, LINDQUIST & of both permanent and temporary water HETHERINGTON 1998a,b, LINDQUIST et al. 1999, bodies. The small black tadpoles metamorRAMos et al. 2002, LA W C A et al. 2005, 2002). The KARRAKERet al. 2006, RICHARDS& KNOWLES phose in 60-75 days (SAVAGE toxic and noxious properties of eggs and 2007
I F
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 RODRIGUES2009, Arquiv. Zool. 40: 46; type localitv: "Chitre. Herrera. Panama." Males to 58.4 hm, femiles to 61.8 mm. Eastern Panama, sea level to 200 m elevation. Chaunw marinus (LINNAEUS 17581,Syst. Nat. 10, 1: 211;type locality: "America." Males to 145 mm, females to 175 mm. From southern Texas, USA, through tropical Mexico and Cenbal America to northern South h e - ; introduced to many tropical countries (and subtropical regions within largely temperate countries, e.g., the USA) world wide, sea level to 3000 m elevation (inCentral America up to 1600 m elevation).
p5 ,,,,
.&>&, .-., I -d.
. b 2U. amus(' Panama).
L
--
,
kXf2
Ph&W. E. he-
ofthe B i o Q i e * Ins~tuWs u&e~!&b of -f
1 I Key to aaunus
1 a Parotoid gland very large, several times the area of upper eyelid; SVL 85-145 m m in adult males. 90-175 mm in adult 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
Rg. 215.Chaunus marinus (Santa Eulalia, Alajuela, Costa Rica, 825 m). Photo: G.Kiihler
I
1
Chaunus centralis :haunusmarinus
Further Reading
ZUG & ZUG 1979, ZUG 1983, EASTEAL1986, NARVAES & RODRIGUES 2009
u ~ u n umarinus s (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 Fig. 217. Crepidophryne chompipe (Cascajal de the life history of these toads. They are Coronado, San JosB, Costa Rica, 1740 my. diurnal inhabitants of primary or mature Photo: G. kohl^ secondary forests (cloud forest or highland oak forest with closed canopy) where they dwell apparently fossorially to semifossoriallyin 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 Fig. 218. Crepidophryne chompipe (same as in Photo: G. Kiihler approximately 55 eggs) and one of C. gua- Fig. 217). nacaste (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 (COPE18761, 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.
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.
m t"7 d - e 1a 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 tuberoles; 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 ofsurTOWareas,with i n W c t 3low, romded hberolw, domum and venter nearly un&nn dull brown-gray .......,...........,.,.
......................... w&@me&m-=-
.,
b Skin on doma1 sarfacea rough, with disIhet3.y raised, founded or poiaha tubercles; late~alrow of tube~cleslarge sharply poiated; latezal surface of bady with shsrply pointed tubercles;eeapular textme distinct &om monndiag are- lacking distinct t u b m k q dozsm d d b m with irregular dark brown markings, venter dull E brown wit& c?mammar-* e-y on the pelvic areas and limbs .......................... I ampid* t&mriplPe
.....................,........
Fig. 221. Crepidophxyne epiotica Quvenile, 7,7 ram SVL, Rio Changena, Panama, 1650 m). Photo: A. Hertz
[k& 219. Ventral view of hand in a. e chompipe and b. C. epiotica. Photos: G. Kiihler
Fig. 222. Crepidophgme epiotica (Rio Macho de Orosi, Cartago, Costa Rim, 2000 m). 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. FROSTet al. (2006a) initially moved the members of this group to a separate genus, Cranopsis, only to Gnd out a little later (FROSTet al. 2006b) that Cranopsis was unavailable. The latter individuals applied the generic name Ollotis for this taxon. Unfortunately, FROSTet al. (2006a,b) neglected to note an available name for this taxon, Incilius Cope 1863 (type species: Bufo coniferus COPE1862b; designated by KELLOGG 1932:29),and for reason of priority FROSTet 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..
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 MCCRANJE & CASTAREDA (2007) did not fol- ground level on shrubs or trees (KOHLER low the taxonomy for the Ollotis coccifer 1993).Amplexus is axillary and egglaying complex suggested by MENDELSONet al. takes place in shallow water of ponds, (2005). Despite the obvious difficulties in lakes, and water courses with slow current. indentifying species of this complex based Most species of the I. valliceps group (i.e., on the information provided by MENDELSONI. aucohae, I. campbelli, I. macrocristatus, et al. (2005),I prefer to follow this arrange- I. melanochloris, I. tutelarius) breed in ment until a different view based on pub- streams during the dry season; I. valliceps lished evidence is available. Contrary to is unique within this species group by laythe claim of L ~ P E et Z d . (2009), Incilius ing eggs in ponds during the wet season et al.1999). Eggs are typically occidentalis (CAMERANO 1874) and I. per- (MENDELSON plexus (TAYLOR 1943)have not been record- laid in two strings. Tadpole aggregations ed east of the Isthmus of Tehuantepec (J. Pschools") of hundreds to thousands of MENDELSON pers. comm. July 2010) and, individuals are common in many Incilius therefore, occur outside of the area covered species. When disturbed, the groups can break up but are reestablished within a in this book. short time. Recently metamorphed froglets Most species of this genus are crepuscular of Incilius species are tiny and usually or nocturnal, but some highland species have no cranial crests or have them poorly 2002). are diurnal. They are usually terrestrial or to partially developed (SAVAGE
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-
Fig. 225. LnciLius canaliferu, ,A inca Irlanda, Photo: G. Kahler Chiapas, Mexico, 1050 m).
fncilius aucoinae (O'NEILL & MENDELSON %004), J. Herpetol. 38: 487; type locality: "airport 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. Fig. 226. Incilius coccifer (near San Jose El Idolo, SuchitepBquez, Guatemala, 100 m). M u s bocourti (BROCCHI 1877c), Bull. Soc. Photo: G. Kiihler 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 k m W Puerto Santo %mBs, 104 m, Montaiias del Mico, Departam a t o de Izabal, Guatemala." Males to 62.6 mm, females to 86.2 mm. Northern and east81p 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.227. Incilius coniferus (Fortuna, Chiriqui, Photo: G. Kohler Panama, 1030 m).
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 Photo: M. Garcia Paris renas, Costa Rica).
Fig. 230. Incilius holdridgei (Cerro Chompipe, Heredia, Costa Rica, 2090 m). Photo: G. Kohler
Incilius fastidiosus (COPE1876), J. Acad. Nat. sei. Philadelphia Ser. 2, 8: 96; type locality: "2500 feet elevation on the slope of the Pico Blanco ICant6n de Talamanca. Provincia 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. 1952b1, Univ. Incilius holdridgei ( T A ~ O R Kansas Sci. Bull. 35: 607; type locality: "at a n 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. 19541, Roc. Biol. Sm. Incilius i b m a i (STUART 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 MENDELSONet al. 20051, 1360-2050 m elevation.
Fig. 231. Incilius ibarrai (San Miguel Ixtahuach, San Marcos, Guatemala, 2050 m). Photo: M. Acevedo
Fig. 232. Lncilius karenlipsae (holotype). Photo: J. R. Mendelson
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.
I
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), b a u e Nacional La MuralIa. 950 m elevation, ~ e p k a m e n t ode Olancho, ~ondurag."Males Disjunct Popto 64.4 mm,females to 90.7 ulations on the Atlantic versant of north-central Honduras, sea level to 1600 m elevation.
Fig. 234. InciLius luetkenii (P.N. El Imposible, Ahuachaph, El SaIvador, 720 m). Photo: G. Kohler
Incilius luetkenii (BOULENGER 18911, Ann. Mag. Nat. Hist. Ser. 6, 8: 455; type locality: "Cartago, Costa Rica"; type locality considered (1974). Males to 96 mm, erroneous by SAVAGE 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 &tatus (Raydn, Chiapas, Fig. 235. Inciliusmat %tween La Gloria and Cerro Azul (presumMexico, 1888 m). Photo: A. Ramirez Velizquez ably 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.
rg.haw. LnciLius macrocristatus (Rayon, Chiapas, Mexico, 1888 m). Photo: A. Ramirez Velizquez
Jndius melanochlom (COPE1877),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 V1967 ~ "1966"), Rev. Biol. Tropical 14: 153; type locality: "Costa
I Fig. 237. hcilius melanochloms (Rio San Photo: G. Kohler Juan, Nicaragua, 30 m).
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.
Fig. 238. Incilius periglenes (male, Monteverde, Photo: M. Franzen Costa Rica).
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.
WILLIAMS, SHEIL Incilius porteri (MENDELSON, & 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.
Fig. 239. bcilius porteri (Cerro Uyuca, FranPhoto: G. Kiihler cisco Morazh, Honduras).
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. 240. bcilius tacanensis (Top of Cerro Mozotal, Chiapas, Mexico, 3016). Photo: S. M. Rovito
Fig. 241. InciIius vallicep~\ l v l v - ~ h e r , Toledo, Belize, 7 m). Photo: G. Kiihler
Ag. 242. h d u s valIiceps (Cerro El Toro,Atlantico Norte, Nicaragua, 830 m). Photo: G. Kiihler
Bufonidae Incilius tutelarius (MENDELSON19911, Herpetologica 53: 15; type locality: &Colonia Rodulfo Figueroa, 19.0 km N W 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.
i
hcilius 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 Ret al. 2006), sea level to 2000 m elevation. Bufo valliceps wilsoni BAYLOR& STUART1961 is , regarded as a synonym of I. valliceps according to M E N D ~ Q(1998). N
lncilius valliceps
'I
coccifer 'lncilius 'ncilius ibarrai
PL - 'nciliusporteri r lncilius tutelariws
*--
lncilius canalifem
lncilius carnpbelli Incilius bocourti lncilius karenlipsa
lncilius rnacrocristatus r lncilius tacanensis
I Fig. 241. Lncilius tutelarius (Jiquipilas, Chiapas, Mexico, 1214 m). Photo: A. Ramfrez VelAzquez
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
b
4 a b 5 a
b
6 a
wide and its anterior border closely attached to the posterior border of eye; postocular crest well developed; tarsal fold pronounced ....................... hcilius bwourti 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 t a c a n d Males uniform orange in life (yellow in preservative), females olive-green to black with red spots .............. hcilius peziglenes Ground color grayish brown to black with parotoid glands and some warts reddish brown ........................................................ 5 Cranial crest low, obsolescent; parotoid gland less than 2/3 of area of upper eyelid; venter pale with some dark mottling on chest ............................... Inciliusholdridgai 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 Dorsum without enlarged warts; venter almost uniform pale tan; posterior thigh dark black, contrasting to dorsum; shank with moderate rugose warts .......................
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
..................................... ..hciliuspenpenpetetes 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 ...................................... Inciliusfastidiosus Fig. 246. Incilius conifems (Bartola, Rio San Photo: G. Kijhler 7 a Finger I shorter than Finger 11; upper sur- Juan, Nicaragua, 30 m). faces covered with widely spaced darktipped, 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 .................. Fig. 247. Incilius melanochlorus (Soltis Center, .....................................Inciliu8 ktiledp~at? 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 Drawings: M. Vesely d. I. valliceps; note variation in shape and size of parotoid glands.
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
Fig. 250. Foot of Lncilius aucoinae. Drawing: M. Vesely
b
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); 13a parietal crests present or absent .......... 10 b 10a ~arotoidgland significantly smaller than the size of area of upper eyelid (Fig. 248~); 14a 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 b a lateral row of tubercles ......................13 l l a Parotoid gland round to oval, area 112 or less area of upper eyelid (Fig. 248c); suborbital, and labial crests well-developed ... 1Sa
........................................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
b 16a
absent or poorly developed; throat and chest pale ........................ In&w aucainae Transverse folds between parietal crests present; pretympanic and preorbital crests well-developed; throat and chest black (Fig. 247) ........................hcik'ue melanochlaru8 Parietal crest absent . In&~8 marmomue Parietal crest present, usually prominent . .................................................................. 14 Subarticular tubercles on fingers and toes bifid or double; suborbital and labial crests usually well-developed; usually a pale interorbital bar ....................................... 15 Subarticular tubercles on fingers and toes single; suborbital and labial crests usually absent or weakly developed; interorbital bar, if present, dark ................................ 18 Supratympanic crests large and bulbous (Fig. 251a) ............................................... 16 Supratympanic crests relatively thin (Fig. 251b) ........................................................17 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
19a
b
20a
b
I 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
b
17a b 18a
b
southern Guatemala as well as southwestern Honduras .................... Incilius ibarrai 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 Venter with no, or few, relatively indistinct dark brown markings; cranial crests relam tively thick and high ........hcilius d Venter boldly marked with a distinct marbled pattern; cranial crests relatively thin and low ............................. h&us Bignifer 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 ~osteriorto confluence with supraorbital crest ......................................................... 19 Tympanum larger (ratio tympanum length / SVL usually >0.50);parietal and supraor-
21a
b
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 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 ............................. h c i l i u s valliceps 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 .... h c i l i u s~~mpbelIi Tympanum smaller (ratio tympanum hind leg shortlength / SVL usually ~0.38); er (ratio hind leg length / SVL about 0.41 in cilius tutdan'u~ males, 0.38 in females) h Tympanum larger (ratio tympanum length / SVL usually >0.38); hind leg longer (ratio hind leg length / SVL 0.42-0.49 in males, 0.42-0.46 in females) .............................21 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 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
Further Reading
PORTER1963, SAVAGE1966, 1972b, NOVAK& ROBINSON 1975, MCDL~RM~I & FOSTER1981, DUNDEE& ROSSMAN 1987, LEE & SALZBURG 1989, JACOBSON & VANDENBERG 1991, ROSSMAN & DONNELLY 1992, 1991, FREED1992, SAVAGE BLAIRet al. 1995,LIPS & KREMPELS 1995, FLEET & AUTREY1997, HAAS & KOHLER 1997, MENDELSON 1997, 1998, A'ITUM & EASON1999, FRANKLIN & FRANKLIN1999, MENDELSON et al. 1999, KOHLERet al. 2000, McC& KOHLER 2000, McCRANIE & WILSON2000, MULCAHY & MENDELSON 2000, DAYTON et al. 2001, MCGEHEE et al. 2001, MENDELSON 2001, JOHNSON 2002, KOHLER2003, O'NEILL& MENDELSON 2004, MENDELSON et al. 2005, JARAMILLO et al. 2009, L ~ P Eet Z 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 obvioas "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- Fig. 253. Rhaebo haematiticus (P.N. Saslaj Mt of the forest floor where it can be Atlantic0 Norte, Nicaragua, 720 m). h m e d foraging for small ants and other Photo: G. Kijhler e o p o d s in the leaf litter (LIEBERMAN 9.986). During the breeding season (March Rhaebo haematiticus (COPE 1862b), Proc. July), Rhaebo haematiticus typically Acad. Nat. Sci. Philadelphia 14: 157; type nrngregates in rocky pools along the borlocality: "Region of the Truando, New Granada ,& of forest streams and rivers (SCOTT [= Chocb, Colombia]." Males to 62 mm, females '*a). It is an explosive breeder that lays to 80 mm. From eastern Honduras (Caribbean slopes) and from southern Costa Rica (Pacific $ggs in paired strings. After metamorphoversant) south to northern Colombia and the froglets are about 19 mm SVL and northwestern Ecuador (and isolated populaU y identifiable by having a black venter tions on the eastern slopes of the Cordillera wrinkled with white spots in life; in Oriental of Colombia extending into adjacent Venezuela), sea level to 1300 m elevation. gmxsemative, the pale spots lose their i&&ensity (SAVAGE 2002).
'e
Rhaebo haematiticus
Y 3 a
Further Reading
CRUZDfAZ & WUSON 1983, SCOTT1983a, BARRIO
er
~ M O R 2001 ~ S
115
-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.
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 Rhinella data (THQMINOT 1884), Bull. Soc. content and hierachical level of this taxoPhilomath., Paris, Ser. 7, 8: 151; type locality: nomic group (e.g., CHAPARRO et al. 2007, Wsthme de Panama ... Obispo." Males to 43 SAVAGE & BOLAROS2009b, PAULYet al. mm, females to 55 mm. Possibly western Panama to northern Colombia, but the geo2009, VANBOCXLAER et al. 2010). Clearly graphic distribution of R. d a t a and its mormore research is needed in order to clarify phological differentiation &om R. acutirostris the phylogenetic relationships of the are poorly understood (HOOGMOED 1989, heterogeneous assemblage of species curBARRIO-AMOR~S 19998, ~oFuuL.4 & S ~ A R I S 1999, B A R R I O - ~ O B ~2004). S rently 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 for- Keytomella mer species of Rhamphophryne in 1 a Finger I shorter than Finger 11; tympanum absent or indistinct ..................................... Rhinella. GRAYBEAL & CANNATELLA (1995) ...................................... Rhinella auvlopha had already noted that there is no unarnb Finger I same length or longer than Finger biguous evidence in support of the mono11; tympanum distinct ................................ phyly of former Rhamphophzyne. .............................................. Rhinella alata The small toads Rhinella acrolopha and R. alata both can be found during daytime amidst leaf litter on the forest floor; gravid females of R. acrolopha were obtained during the months of February, March, and July (TRUEB1971, IBAE-JEZ et al. 1999b). Both species feed mainly on ants and other small arthropods (TOFT 1981). Rhinella data is an explosive breeder that uses areas in the forest ponds and reproduce, primarily near the end of the r a i season ~ and in the early dry .wason (BAREzet al. 1999b).
-
a
fi
pqF
&i.
k'
Fig. 254. &inella acrolopba (Cerro Mali, Da,.i&, Photo: W. E. Duellman (COAS~
of the Biodiversity Institute,University of Kansas)
r Rhinella acrolopha
Fig. 255. Rhinella d a t a (Nusagandi, Cuna, Photo: G. Klihler Panama, 350 m).
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 distribnted from southern Mexico across Central America and tropical South America (to Peru on the Pacific versant and to southcentral 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 centrolenids 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).
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 (VILLA1977, MCDIAR~WD 1983). Adult males of H. valerioi attend their egg clutches during night and day, being conspiciously similar in appearance & glassfrogs I are nocturnal and can be to their egg clutches (Fig. 273): the frog's &md in vegetation along small rivers and reticulate green pattern on a yellowish backb s . During the rainy season, the ground is thought to be an evolutionary &&-pitched call of the males can be response to visually hunting diurnal 1983). This resem&ad. .The females of most species deposit predators (MCDIARMID #bireggs on leaves overhanging fast-mov- blance between frogs and eggs is not &reams, on moss on branches, or on present in species that desert their clutch 1975, during the day, such as H. colymbiphyllum, above streams (MCDIARMID ARRANZA & LYNCH1991), although which is uniform green with small yellow ella granulosa is reported to breed dots. After night falls, the males of this nds occasionally (KUBICKI2007). species return to their egg sites. ding on the species, the eggs can be on the upper leaf surface, at the tip, Depending on local weather conditions, the neath the leaf (GUAYASAMINet al. larvae are ready to hatch after 2-3 weeks. . During development and with the Often clutches develop to a certain point of water, the gelatinous egg and then hatch during substantial rains ell and increase their volume. In when they are washed or fall into the species of glassfrogs,varying degrees stream. One benefit of hatching during care are present, such as in the rain is that tadpoles dropping into water Hyalinobatrachium. It is the 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 environment (SAVAGE 2002). The tadpoles take several months to a year before metamorphosis (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 creamy lavender to dark lavender in preservative; dentigerous process of the vomer present, with or without teeth; ventral parietal peritoneum white anteriorly and transparent posteriorly or completely transparent (Fig. 259cf) ..........................8 2 a Two to six conspicuous, clearly demarcated large, bright yellow dots present on the
H*Obawum b No conspicuous, clearly demarcated large, bright yellow dots present, although numerous small yellow flecks can be present ............................................................ 3 3 a Substantial webbing between fingers I1 and I11 (Fig. 257a) .......................................
......................Hyalinobatmchium chimi.. b Only vestigial webbing between fingers I1 and 111 (Fig. 257b) ..................................... 4
4 a Dorsum with a dark green vertebral stripe; pericardium colorless ...............................5 b Dorsum green with small yellow spots or reticulated with green and yellow in life, no dark green vertebral stripe; pericardium white or colorless ...................................... 6 5 a Dark green vertebral stripe bordered on either side by yellow stripes in life (the dark stripe indicated by purplish or brownish pigment in preservative) .......................
............. Hyalinobatrachium vhmittatum
b Dark green vertebral stripe not bordered on either side by yellow stripes in life ....... .................m o b a t r a c h i u m talamancae 6 a Dorsum with yellow blotches surrounded by a reticulum of green pigment and dark punctations in life (a reticulum of black punctations in preservative); pericardium
I
,.
F I ~ao . nana or a. H.chmpoi and b. H. Photos: B. Kubicki colymbiphylllum.
24
1
a.
Fig. 258. Profile of head in a. Sachatamia albomaculata and b. Cochranella euknemos. Drawings: G. Kohler
usually white (members of some populations lack a completely white-pigmented pericardium) . Hjdhobatrachium valerid b Do'sum with yellow spots but not Burrounded by a reticulum of green pigment and dark punctations in life (whitish or yellowish, often with a minute brown or purpelish punctations in preservative); pericardium white or colorless ................7 7 a Tympanum absent or very indistinct; pericardium white or colorless; nostrils in very slightly indicated fleshy swellings .............
...............qyalinobatrachiumfleischmanni
b Tympanum distinct; pericardium colorless; nostrils in strongly protuberant fleshy swellings ......................................................
.......... Hplinobatrachium c o l ~ p 4 v I I u m
8 a Prepollex with a free, spiny distal end; dorsum without distinct spots; SVL to 23 mm ......................................Teratohyla@OM b Prepollex without a free, spiny distal end; dorsum with or without distinct spots; SVL variable ..................................................... 9 9 a Skin texture strongly granular; dorsum uniform green or bluish-green without yellow spots (although evenly marked with tiny bluish to whitish flecks) but usually with scattered purple or black spots; a 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 present (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 dorsal 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
TAYLOR1949b, 1958a, SAVAGE& STARRETT1967, STARRETT & SAVAGE1973, M c D m & ADLER 1974, Villa 1974, DUELLMAN 1977, VILLA1977, HAYES & STARREIT1980, VILLA 1984, RuuCARRANZA & LYNCH1991, MCCRANIE & WILSON 1997c, BARREU-RODRIGUEZ 2000, SAVAGE2002, 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 sides of leaves overhanging the water, usually near the tip of the leaf. In both Central American species, the eggs are darkly pigrnented and the jelly masses can contain 40-70 eggs (KUBICKI 2007).
I 1 Fig-260. C ~ c h m e f l amdcnemos (Nusagan& Kuna Yda, Panama, 350 m). Photo: G. Kohler
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 (3k m SE Ayapal, Nicaragua, 200 m). Photo: G. Hiohler Zochranella euknernos
Further Reading
MCCRANIEet al. 1999,SAVAGE 2002,GUAYASAMIN & BONACCORSO 2004,KUBICIU2007, 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 vegalong fast-moving streams. Males soblepon call from the upper surleaves throughout the year with Fig. 262. Espadarana prosoblepon (male,P.N. ed vocal activity during the rainy Saslaya, Nicaragua, 920 m). Photo: G. Kohler
a prosoblepon (BOETTGER1892), Samml. Mus. Senckenb. Natur-
b
Fig.263. Espadarana prosoblepon (female,P.N. Saslaya, Nicaragua, 920 m). Photo: G. Kohler
C
alinobatrachium
>
!s b
Espadamna prosoblepon
Further Reading
JACOBSEN 1985, CRUZ-DIAZ & MCCRANIE1999, SAVAGE 2002, KUBICRI 2007, HOFFMANN 2010 I
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 vegetation overhanging streams. During daytime, these small frogs hide well camouflaged within streamside vegetation. The females deposit their eggs, which are green when laid, on the underside of leaves (RUIZ-CARRANZA & LYNCH1998, SAVAGE 2002). Males in this genus typically guard the eggs (KUBICKI2007). Hyalinobatrachium fleischmanni is the glassfrog with the widest geographical and elevational 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 <23.3mm, females to 23.9 mm.Eastern cordilleras af the Dari6n Province in Panama to northwestern Ecuador, 45-1570 m elevation.
:
'472: 66; type locality: "Costa Rica: Provincia de :Sari Jose: Canun Perez Zeledon: 0.5 k m 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 #.CostaRica and in the area of El Valle de Ant6n i n west-central Panama (KUBICKI 2007). 1 i630-1100 m elevation. I
.-
.Hyalinobatrachium chirripoi (TAYLOR1958a), 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 & WILSON1997 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& WILSON1997 is regarded as a synonym of H. colymbiphyllum according to CISNEROSHEREDIA & MCDIARMID(2007).
Hyalinobafrachium
A
Y. aureogutatum I. fleischmanni
t
J
4 H. vireovittatum r H. talamancae
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], Caribbean 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.
Hyalinobafrachium
f. colymbiphyllum - I. chirripoi
Centrolenidae
Fig. 265. Hyalinobatrachium aureoguttatunl (Ecuador). Photo: M. R. Bustamante 1 BiographicaO
I
Fig.264. Hyalinobatrachium colymbiphyllum (near Parrita, Puntarenas, Costa Rica, 250 m). Photo: B. Kubicki
Fig. 266. Hyalinobatrachium aureoguttatum (Ecuador). Photo: M. R. Bustamante 1BiographicaB
Fig. 267.Hyalinobatrachium chimipoi (Aguas Zarcas, Limbn, Costa Rica). Photo: G. Kohler
Fig. 270.Hyalinobatrachium chirripoi (P.N. Patuca, Olancho, Honduras). Photo: G. Kiihler
Fig. 268.Hyalinobatrachium fleischmanni (Salto lab^, Atlantic0 Norte, Nicaragua, 480 m). Photo: G.Kohler
Fig. 271.Hyalinobatrachium fleischmanni (Piedra de Candela, Chiriqui, Panama, 1470 m). Photo: G.Kohler
Fig. 269.Hyalinobatrachium talamancae (La Nevera, Ngobe Bugle, Panama, 1600 m). Photo: A. Hertz
Fig. 272.Hyalinobatrachium vireovittatum (nearParrita, Puntarenas, Costa Rica, 250 m). Photo: B. Kubicki
Fig. 273. Hyalinobatrachium valerioi (male with eggs, Guayacan, CRARC, Costa Rica, 550 m).
Photo: G. Kijhle
Fig. 274. Hyalinobatrachium valerioi (male, Guayacin, CRARC, Costa Rica, 550 m). Photo: G. Kohler
Fig. 275. Hyalinobatrachium valerioi (female, Photo: G. Kiihler Costa Rica).
Further Reading
GOIN 1964, GREER& WELLS1980, MCDIARMID 1983, VILLA1984, IBAREZ & JARAMILLO1997, JARAMILLO et al. 1997, MCCRANIE & WILSON1997, bAFmz et al. 1999a, BARRERA-RODRIGUEZ 2000, SERARIS 2001, SAVAGE 2002, KUBICRI 2004a, KUBICKI 2007, HOFFMANN 2010
125
Sachatamia The generic name Sachatamia was proet d. (2009)for three posed by GUAYASAMIN 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 but is absent in S. albomaculata.
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 (KUBICKI2007, HOFFMANN 2010).Occasionally, clutches are discovered on rocky surfaces alongside wateret al. 2004).Clutch size falls (PUSCHENDORF 2007). is 16-25 eggs in S. ilex (KUBICKI
&
I Fig. 276. Sachatamia albomaculata (Guayach, CRARC, Costa Rica, 550 m). Photo: G. Kiihler
Fig. 277. Sachatamia ilex (Soltis Center, Sachatamz'a albomaculata TAYLOR 1949b), Univ. Kansas Sci. Ed.33: 267; type locality: Alajuela, Costa Rica, 460 m). Photo: G. Kiihler '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 (SAVAGE1964), 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.
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.
Fig.278. Teratohyla spinosa (Guayach,
CRARC, Costa Rica, 550 m). Photo: G. Kohler
: The two Central American species of this 1
1 [ /
I i i
1 t
genus are found along streams in forests, where males call from the upper surfaces of leaves at night. In T. spinosa, the egg masses (clutch size 15-25 eggs) are usually attached to the undersides of leaves, along their margins, whereas in T. pulverata the eggs (clutch size 30-75 eggs) are laid on the upper surface and toward the tip of leaves (KUBICKI 2007). During early stages of development, the eggs of T. spinma are greenish white, whereas those of T. Fig. 279. Teratohyla pulverata (Cerro El Toro, pulverata are pale greenish blue (KUBICKI Atlantic0 Norte, Nicaragua, 480 m). Photo: G. Kohler 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 TAYLOR1949b, 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.
I A T.T.pulverata spinosa F'urther Reading KUBICKI2007, 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 NOBLE1926) has always been strongly supported by previous studies (VENCES et al. 2000, SANTOSet al. 2003). The taxonomic split of Dendrobatidae into two families (i.e., Allobatidae and Dendrobatidae), as proposed by GRANTet al. (2006), was rejected as unnecessary by SANTOS et al. (2009), who returned to the et more traditional taxonomy. Also, SANTOS 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 Dendrobates. 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 MYERSet 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. et al. 2004). 1980,1992,1994, DUMBACHER
Key to genera of Dendrobatidae
1 a Discs of outer fingers (Fingers III and TV)
b 2 a
b 3a b
4 a
b 5a
b
6a
b
and toes (Toes IV and V) usually at least two times width of digits; no maxillary teeth ....................................... Dendrobah Discs of outer fhgers (Fingers 111 and IV) and toes (Toes IV and V) only slightly wider than digits; maxillary teeth present ....... 2 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 Toes with basal or no webbing; dorsal color pattern variable; SVL of adults variable 3 Finger I as long as Finger 11; dorsum with pale spots, no stripes; SVL of single known specimen 18.9 mm ..................... AmFinger I longer than Finger 11; dorsum without pale spots, but usually dorsolateral or lateral stripes present; SVL of adults variable .....................................................4 Ventral surfaces of body and thighs black with pale mottling .................. Phyllobates Ventral surfaces of body and thighs white, although throat and chest can be black in adult males ............................................... 5 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 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 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 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 Fig. 280. Hand of Silverstonia flotatoq note swollen Finger 111 (arrow). Photo: G. Kijhler
SAVAGE 1968a, MYERS1987,- Z & ZIMMEFMANN 1988, DALYet al. 1992, BAUER 1994, CLOUGH& SUMMERS 2000, GRANTet al. 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 Fig. 281. Allobates talamancae (near Miramar, periods); the call consists of a series of Bocas del Toro, Panama, 20 m). Photo: G.Kohler 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 ~etzal. 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 develop2002). ment (IBAREzet al. 1999b, SAVAGE mobaces taIamancae ('OpE 1876)9 J' Nat. Sci. Philadelphia Ser. 2,s: 102;Type localitv: "near old H~~~~~~[= puerto vieio1, on the ~ * a s tcoast", [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.
Fig.282. Allobates talamancae (Lagarto Lodge, Alajuela, Costa Rica, 90 m). Photo: G. Kohler
-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).
-
Further Reading
SAVAGE 1968a, IBAREZet al. 1999b, SAVAGE 2002
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.
M e r Reading 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 nonriparian frog usually found on the floor of o s t e u s wet montane forest, including cloud forest, del Toro, Panama, 10 m). away from streams, whereas C. panamensis is riparian, occurring only in the immediate adjacency of forest streams, often along rocky sections (IBAREZ et al. 199913, GRANT2004). Males and females are territorial; the call of male C. panamensis consists of one to four whistled, moderately high-pitched notes in rapid succession (IBAREZet al. 1999b). In C. panamensis, eggs are laid in the moist leaf litter, and after hatching, the female carries the tad-
-he, Bocas 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 (COPE1863),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 GRANT2004). 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.
Dentrobatidae
.
-
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.
Dendrobates
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 (DALYet 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, MEBSet 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 mc -m predominantly lowland rainforests; most are terrestrial or semiarboreal,but at least g. 285. Foot of a. Colostethus panamensis Photos: G. Kohler D. arboreus and D. vincentei are strictly d b. C. pratti. arboreal, living in epiphyte-laden trees (MYERS et al. 1984, JUNGFER et al. 1996). &ey to Colost%t&us 1a Toes with lateral fringes and distinctly Dendrobates pumilio and D. granuliferus webbed at base (Fig. 285a); SVL of adult exhibit a considerable diversity in color males 18.8-27.1 mm, of adult females patterns (SUMMERSet al. 2003, BATISTA & 24.9-29.2 mm ..... Colostethuep a n a m d KOHLER 2008). There are examples of disb Toes without lateral fringes and webbing tinct populations in close proximity withabsent between all toes or barely detectable between toes I11 and TV (Fig. out any obvious physiographic barriers 285b); SVL of adults variable .................. 2 that show no evidence of hybridization; O a A dark stripe along posterior surface of these abrupt transitions have been interthigh; throat immaculate white in females, preted as evidence of lack of or minimal mottled with gray to dark brown in adult gene flow (BATISTA & KOHLER2008). DALY males; no toe webs; SVL of adult males to & MYERS(1967: 973) stated that for D. 24 mm, of adult females to 25 mm ............. ........................................ Colostetbuspratti pumiLio "a static subspecies concept does b No dark stripe along posterior surface of not seem well suited to the dynamics of the thigh; throat brown with white spots form- situation". However, the descriptions of the ing mottled pattern in females, darker in phenotypes by these authors agree well #. adult males; webbing absent between all with the observations by BATISTA& toes or barely detectable between toes 111 KOHLER(2008), indicating that over the I and TV; SVL of adult males to 24.5 mm, of adult females to 34 mm .............................. course of at least four decades the pheno9- .................................. Colostethus latinasus typic pattern had remained stable. The underlying mechanisms and barriers that prevent mixing of the phenotypes in the natural populations are poorly understood.
d
1
I
Dentrobatidae 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 (MYERSet 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. Dendrobates arboreus, D. granuliferus, D. pumilio, and D. speciosus a harsh, insect-like "chirpchirp-chirp" (MYERSet 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 Y -: water such as water-filled leaf or stem Dendrohafes 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, BRUST1993).
Dendrobates arboreus MYERS, DALY & MAR1984, 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 weste 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.
L ~ T ~ E&R S Dendrobates claudiae JUNGFER, J ~ R G E2000, N ~ Herpetofauna, Weinstadt, 22: 12; type locality: "Panama: Pronvincia Bocas
3. auratus
1
4
7. granuliferus
I
I. minutus I.
arboreus
pumilio fulguritus claudiae vicentei
Dentrobatidae
arboreus (Camino Palo Bocas del Toro, Panama, 1090 m). Photo: K.-H. Jungfer
Fig. 880. Uendrobates claumae (mainland opposite of Isla Loma Partida, Bocas del Toro, Panama, 20 m). Photo: K.-H. Jungfer
Rg. 287. Dendrobates auratus (Bartola, Rio Sari Juan, Nicaragua, 30 m). Photo: G.K6hler
Fig. 291. Dendrobates claudiae (mainland opposite of Isla Loma Partida, Bocas del Toro, Panama, 20 m).
Fig. 288. Dendrobates auracus wan Antonio ae Damas, Puntarenas, Costa Rica, 60 m). Photo: G. K6hler
r ~ gg. . 2 ~uendrobaces . mgmitus (huuagandi,
Kuna Yala, Panama, 330 m). Photo: K.-H. Jungfer
I Fig.289. Dendrobates auratus (Nusaganm, Kuna Yala, Panama, 350 m). Photo: G.Kohler
Fig. 293. Dendrobates minutus (Nusagandi, 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, Departamento 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 lower Rio Anchicaya, 160-800 m elevation. Dendrobates granuliferus TAYLOR1958b, 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. ~ Occas. Dendrobates minutus S m t ~ v1935, Pap. Boston Soc. Nat. Hist. 8: 212; type locality: "Barro Colorado Island, Panama Canal Zone", Panama. Males to 15.0 mm, females 15.5 mm. Central Panama (El Valle and Cerro Campana) to Colombia (south to Rio Saija), near sea level to 1100 m elevation.
Fig. 294. Dendrobates grandifems (Rincbn de Osa, Puntarenas, Costa Rica, 10 m). Photo: G. Kiohler
Fig. 295. Dendrobates granuliferus (San Antonio de Damas, Puntarenas, Costa Rica, Photo: G. Kohler 60 m).
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 SCHMIDT1857, 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.
r lg. LJO.
U~IIU~VU~LG ~U @U I I I I V\LUU C/ULLILLL L L ~ ,
Rio San Juan, Nicaragua, 30 m).Photo: G. Kijhler
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 1km 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. 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, Chiriqui, Panama, 1100 m). Photo: K.-H. Jungfm
Fig. 300. L,Jrobates pumilio (Isla Bastimen-
Fig. 304.Dendrobates vicentei (Rio BelBn, Photo: K-H. Jungfer Veraguas, Panama).
;,Bocas del Toro, Panama). Photo: G. Kohler
tos,Bocas del Toro, Panama). Photo: G. Kohler
Fig.301.Dendrobates pumilio (Cerro Brujo, Bocas del Toro, Panama).
Photo: G. Kiihler
Fig.305.Uendrobates vicentei (El Cop6, Coc16, Panama, 680 m). Photo: R-H. Jungfer
I
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
I
Fig. 306. Dendrobates pumilio (Guayachn, CRARC, Costa Rica, 550 m). Photo: G. Kohl1
Key to Dendzvbatee
1 a SVL of adults 2 5 4 2 mm .........................2 b SVL of adults 13-24 mm .........................3 2 a Dorsum and venter bluish green, green, or
b 3 a b 4 a
b 5a
b
6a
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 sexes 20-22 mm .... Dandrobates arbomus b Coloration and pattern of dorsal and ventral 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 adults of both sexes 19-22 mm ..................
........................... Dandrobatesgran* b Dorsum smooth;hind limbs coloration vari-
able; SVL of adults variable .................... 9 cream with large black or dark brown 9 a Throat coloration in males not different from that in females; Coloration and patblotches, bands, or stripes ........................... tern of dorsal and ventral surfaces of body ................................... Dandrobates auratus and limbs variable; SVL of adults Dorsum red in life, with or without black 14-24 mm . Dandrobata pumilio (inpart) flecks; venter uniform red ........................... b !mroat in males dark, in females pale (Fig. ................................ Dandrobates s p e C j m 307); dorsal and ventral surfaces of body Dorsum with longitudinal stripes ........... 4 and limbs olive green, yellowish green, or Dorsum patternless, or with speckles, bluish green in life, with or without brown spots, or blotches ...................................... 7 or black blotches or reticulations; SVL of adults 12.0-20.3 mm Dendrobates M'eentei SVL of adults >17 mm; dorsolateral pale stripe, if present, not continuous from eye to sacrum .. Dandrobatespurnilio (inpart) SVL of adults c17 mm; a continuous dorsolateral pale stripe from eye to sacrum .... 5 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 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 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, GRAVES1999, green, or yellow in life; SVL in adults of JUNGFER et al. 2000, SAVAGE 2002, SUMMERSet al. 2003, LOTTERSet al. 2007, HAUSWALDT et aL both sexes 13.5-16.5 mm ............................ ...............................hdrobates fidguritus 2010
indeed very similar in their external morphology and coloration, and have been found to be sister species in the molecular MYEW (1991), referred several specimens phylogeny published by WIDMERet al. i h m Colombia and Ecuador, and one indi- (2000). Furthermore, they readily vidual from Panama to "Colostethus hybridize under captive conditions (J. pers. com.). These frogs are diurnal chocoensis" [= Hyloxalus chocoensis VARGAS (BOULENGER 1912)l.MYERS & GRANT (2009) terrestrial denizens in the leaf litter of assigned populations from northwestern forested stream valleys where they move Ecuador to the new Anomaloglossus con- about predominantly by short hops inter2002). Males fusus and restricted the range of Hyloxalus spersed with walking (SAVAGE chocoensis to its type locality and one local- typically call from within holes found ity in Antioquia, both in Colombia, thereby between roots or rocks and are difficult to unpubl. obs.). The call of rendering the single Panamanian speci- locate (KOHLER both Central American species consists of a men (KU 172790 from El Llano-Cartf km 11.7, 150 m elevation, Province low rasping trill lasting 2-6 seconds and ' road, 2002). PanamA) formerly referred to as repeated after a pause (SAVAGE 'Colostethus chocoensis" without an iden- Oviposition is preceeded by a complex tification. Thus, I have refrained from including a species account under this genus.
Hyloxalus
Pnrther Reading MYERS1991, COLOMA1995, MYERS & GRANT 2009
As currently understood, the genus
PPlylobates contains five species, two of which are found in Lower Central America MYERSet al. 1978, 1995, GRANTet 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~ons (MYERS et al. 1978, DALYet al. 1980). Although the two Central American P. lugubris @es are not as dangerously toxic as is P. P. vittatus 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 (DALYet al. 1980). WAGE (1968a) considered P. vittatus as a synonym of P. lugubris, but SILVERSTONE (1976) recognized both as valid species. Phyllobates Phyllobates lugubris and P. vittatus are _i
Dentrobatidae
-
-
1
courtship; there is no amplexus, instead Key to PhyIobates the animals have been observed sitting 1 a Dorsolateral stripe yellow in life, usually narrow, bordering upper eyelid laterally; with their rear ends touching (SILVERSTONE upper surfaces of limbs marbled with gold, 1976, LEENDERS 2001). Eggs are deposited yellow, or yellowish green in life; SVL of adult males 19-21 mm, of adult females in the moist leaf litter and the clutches are 20-24 mm ................. Phyllobates lzzgrzblis not guarded, although males have been b Dorsolateral stripe orange-red in life, usuobserved to moisten them when they are ally broad, nearly as wide as upper eyelid; among dry leaves (SAVAGE2002). After upper surfaces of limbs marbled with hatching, the tadpoles wriggle onto the bluish green in life; SVL of adult males back of the male, who carries one or sever22-26 mm, of adult females 26-31 mm ..... ....................................Phyllobates vittatus al tadpoles to small puddles on the forest floor, water-filled depressions in logs, and backwater areas near streams (SILVER- Further Reading STONE 1976, LEENDERS 2001). Metamor- SAVAGE 1968a, SILVERSTONE 1976, MYERSet al. et al. 1990a, phosis occurs about two months later, and 1978, DALYet al. 1980, DONNELLY et al. 2000, RYAN2002 the froglets are 12-13 mm SVL (SAVAGE WLDMER 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 (COPE1893), 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, Photo: G. Kohler Puntarenas, Costa Rica).
Silverstoneia
GRANTet 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 sympatrically 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 "peetpeet-peet", resembling a trill that last about two seconds (IBMEZ & SMITH1995). 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 (DUNN1924b), 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 Silverstonia flofator
Fig. 311. Silverstoneia nubicola (Almirante, Bocas del Toro, Panama, 30 m). Photo: A. Hertz
-b 7 ; :
Silverstonia nubicola
Fig. 312. Silverstoneia nubicola (P.N. G. D. Omar Torrijos H., near El Cop6, Cocl6, Panama). Photo: R. Brenes
Further Reading
1
SAVAGE 1968a, 1-z 2002
& SMITH1995, SAVAGE
6
Terrarana New World direct developing ogs)
Key to the species of the genera Craugastor,Diaapom, Eleuthdactglus; and Aistimantis of northern Central America (southern Mexico to Nicaragua)
New World direct developing frogs com- 1 a Toe V longer than Toe I11 when each prise a diverse assemblage of species that adpressed to Toe IV;webbing between toes reduced (no webbing or only basal has a confusing taxonomic history. During webbing; Figs. 314a,b) ..............................2 the last decades of the lgth and the early b Toe V shorter than Toe I11 when each part of the 2oth centuries, these frogs were adpressed to Toe nT; webbing between toes assigned to the genus Hylodes, a name provariable ...................................................15 posed by FITZINGER (1826). However, 2 a A pale transverse line demarcating finger STEJNEGER(1904) determined that and toe discs (Fig. 313); most digits with Eleutherodactylus was the correct name palmate or spadate covers (Fig. 3150; SVL of adult males 16-21 mm, of adult females for the species that formerly had been 18-24 mm .................. Diaspom diadems referred to Hylodes because the latter b No pale transverse line demarcating finger name has to be applied to a group of frogs and toe discs; disc covers on Fingers 111-IV related to H. nasus (type species of the and on all toes round or truncate (Figs. genus Hylodes, today in the family 315c,d); SVL variable ............................... 3 Hylodidae of FROST et al. 2006a). This con- 3 a Disc covers on the outer two fingers (I11 clusion was supported by G. S. MYERS and IV) moderately expanded, rounded (Fig. 314b, 315c) ........................................ 4 (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-groupname 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, 313. Diasporus diastema; note pale line 24 genera, 11subgenera, 33 species series, Fig. demarcating discs (arrows). Photo: G. Kohler 56 species groups, and 11 species subgroups. Thus, Terrarana includes about 28 percent of anurans in the New World tropics and about 13 percent of frogs worldwide (DUELLMAN & LEHR2009). In Central America, there occur representatives of three families of Terrarana (Craugastoridae, Eleutherodactylidae, and Strabomantidae) with a total of 123 recognized species.
Further Reading CRAWFORD 2003, CRAWFORD & SMITH 2005, HEINICKE et al. 2007, HEDGES et al. 2008, DUELLMAN & LEHR2009
Fig. 314. Foot of a. Diasporus diastema; Drawings: M. Vesely b. Pristimantis ridens.
rig. 315. Finger disc covers of a. Craugastorpodiciferus (unexpanded, even); b. C. jota (slightly expanded); c. Pristin~antiscaryophyllaceus (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
Fig. 316. I'mger d~scpads 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 Lliasporus diastema (even, triangular).
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.c r u e n b , j. P. museosm.
Photos: a.-i. G. Kiihler;j. A. Hertz
141
I m
J
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 Wshaped 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 Hshaped 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 n f l d w 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 ................... ................................... Craugadorcdmpbelli 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 obtuse, pungent, or pointed; SVL of adult males 24.0-26.8 mm (females unknown) ...................................... .-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 ..........Cmugaetorglaucus 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 . 3 5in females); vocal slits present in males ........
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
-.-
...................................... ~ b o c o u r t i E ig. azu. Larerw neaa or a. ~raugastorglaucus Photos: G. Kiihler b Ear opening large (ratio tympanum length and b. C. xucanebi.
/ eye length >0.50 in males, >0.35 in
females); no vocal slits in males ........... 11 lla Overall dorsal coloration pale without dis-
b
12a
b
13a
tinct 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 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 Venter dark brown; ratio tympanum length / eye length >0.50 in females; SVL of single known female 40.2 mm ................ .................................... Craugador siIv1'cola Venter dirty white or pale cream color; ratio tympanum length / eye length variable; SVL of adults variable ................. 13 Occurs in the lowlands (sea level to 600 m elevation) &om central Veracruz, Mexico, to western El Pet&, Guatemala ................
Fig. 321 doration of thigh in a. Craugasl yucatanensis, b. C. stuarti, and c. C. xucaneor. Photos: G. Kohler
,. .................................... 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 (111and 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 lse 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 t .s :digital pads slightly expanded ............................... ........................ J5'leuthedac&lus pipilans 19a Finger and toes with supernumerary tubercles ................................................. 20
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 membranous web between toes ..................... 21 b No inner metatarsal tubercles; vocal slits in males present or absent; nuptial pad on thumb in males present or absent; basal webbing usually present between Toes 11-IV ........................................................ 22 21a All fingers and toes bearing ventral pads .
............................................
I .
I
Craugastorloki
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
Fig. 324. Craugastormimus; note dark eye mask along lateral head (Cerro El Toro, Atlantico Norte, Nicaragua. Photo: G. Kijhler
................................. 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 ............
...................................Craugastorlauraster
b A nuptial pad on thumb in males present (Fig. 342) ............... Craugastor b&ordii 26a Posterior dorsum with a definite laterosacral system of either V-shaped or nonconverging 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 3 0 4 3 mm, of adult females 50-70 mm ......................................
.......................... Craugastormegamphalus
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. 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 fingers (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 ....................................... Craugaetornoblei b Inner tarsal fold absent; sole of foot smooth (Fig. 326b,c); dorsum granulate; glandular supratympanic ridge not extending posteriorly to axilla ................... ..............................Craugastor talamancae
33a Heel pustules weak; row of outer tarsal tubercles very weak or absent; usually no la I11 3 - 4 IV (Fig. 326c); toes with wellpale vertebral stripe, although a weak, thin developed lateral fleshy fringes continuous ridge similarly colored as adjacent dorsum with webs (Fig. 326c) .. Crau.gahr m i m ~ present in most individuals; SVL of adult Toes with basal webbing, modal formula III males 30-36 mm, of adult females 40-58 3+-4 l/2 IV; toes with non-fleshy lateral mm .............................Craugastor rastralis keels ........................................................31 b Heel pustules well-developed; row of outer Heel smooth to rugose but without pustutarsal tubercles distinct; a thin pale vertelar tubercles or calcars; no distinct dark bral stripe usually present; SVL of adult stripe along outer thigh margin; most finmales 22-27 mm, of adult females 31-39 ger disc covers slightly to lanceolate, with mm ................................... chac swollen to cuspidate disc pads; no dark seat patch mark ................. Craugagtorlatioeps 34a Inner tarsal fold absent; tympanum usually hidden or indistinct, rarely distinct .... Heel with 1-4 more or less distinct small QX pustular tubercles or 1-2 large calcars; a b Inner tarsal fold present; tympanum distinct dark stripe along outer thigh marprominent ........................................ 48 gin; finger disc covers round, with even disc pads; a dark seat patch mark present .. 32 36a Body with a pair of prominent dorsolateral ridges .......................................................36 Heel with 1-2 large tubercles (calcars); without paired dark spots at the apex of b Body without a pair of dorsolateral ridges, the triangular dark seat patch mark ......... although irregular ridges and tubercles ....................................C r a m lineatus can be present ........................................ 38 Heel with 1-4 more or less distinct small 36a Dorsum smooth to rugose but never with pustular tubercles; paired dark spots prenumerous pustules or tubercles (Fig. 328a); sent at the apex of the triangular dark seat no nuptial pads in males ............................ patch mark ..............................................33 ........................................ Craugastor b Dorsum strongly granular with numerous large tubercles (Fig. 328b);males with nuptial 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; 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 1213 - 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 .........................
30a Toes with moderate webbing, modal formu-
b Sla
b
S2a
b
.............................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, moderatesized, or large tubercles; SVL of adult males 21-34 mm, of adult females 2 8 4 7
Fig. 326. Foot of a. Craugastor noblei; b. C. talamancae; c. C. mimus. Drawings: M. Vesely
40a Ratio disc width on Finger I11 / tympanum length 0.69-0.93 .....................................41
-,
.
4
b Ratio disc width on Finger I11 / tympanum length 0.38-0.58 .....................................42 41a Upper jaw flared beneath orbit (Fig. 327); snout truncate i n 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 (condition 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 Fig. 327. Craugastor trachydermus; note flarea toes basal to moderate, modal webbing for- upper jaw (arrow). Photo: G. Kohler mula 1113 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 ........................ h u g a s t o r 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 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 3 9 4 1 mm ............. ~m'ugamkrs a l ~ m b More toe webbinp; modal webbing formula I 2-2 415 11 2-3 3 4 1113-4 ~4 IV4 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 males; usuallyless webbgg between Toes Fig. 328. Dorsum of a. Craugastor g r e g - b. C. 111-V, modal formula 1113 4 1/4 IV 4 114 -2 d&; cc.C. adamastus. Photos: G. Kiihler
I 1
I
........... ....................................h u g a s t a r i&cundu~
3/4 V; occurs in northern Honduras
f
r w . 329. Craugastor fitzingen a. thighs; b. - -- . -
Photo: G. Kohler
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 ..... ......................................... CraugastormilmBB1 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~torwffellg 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 ........................ ............................. h u g a e t o r chingopetam b Inner tarsal fold flap-like; discs on Fingers 111-IV narrowly expanded ( ~ 2 . 5times 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 ....................................
................................Craugastarsandans0mIU
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) ....
Fig. 330. Foot of a. Craugastor fitzingeri, b. C. rupinius, and c. C. chingopetaca. Drawings: a,b M. Vesely, c. L. Czupalla
................................... h u g a s t o r-if b Discs on Fingers 111-IV narrowly expanded
( ~ 2 . 5times 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 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 ros-
tralis rounded to moderately angular; dorsum smooth, rugose, or tuberculate; 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 b Discs on Fingers 111-IV narrower expanded ( ~ 1 . 5times 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 reddish in life; SVL of adult males 26-36 mm, of adult females 53-65 mm .........................
....................... 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 37-51 mm, of adult females 66-83 mm .....
...................................... C r a w 57a Toes with a lateral narrow flange; dorsum rugose; ratio shank length / SVL 0.55-0.64; canthus rostralis moderately angular; occurs on the Pacific versant of south-central Honduras .............. Craugastar emleni b Toes with a lateral keel; dorsum granulate; ratio shank length 1 SVL 0.51-0.57; canthus rostralis rounded; occurs on the Atlantic versant in east-central Guatemala
.....................................Craugagtor inachurr
58a Inner tarsal fold flap-like (Fig. 332); discs on Fingers 111-IV ca. 1.5 times width of digit just proximal to disc; venter whitish or yellowish in life; vocal slits absent in males; nuptial pads present in males ........
................................... Chugastarpalanque b Inner tarsal fold ridge-like, or, if flap-like, then discs on Fingers 111-IV ca. 2.0 times width of digit just proximal to disc; coloration of venter variable; vocal slits and nuptial pads present or absent in males ...
..................................................................51 59a Discs on Fingers 111-N narrower expanded ( d . 5 times width of digit just proximal
a.
fig. 331. Hand of a. Craugastorfitzingeri, and b. C. 1aevissimus. Drawings: M. Vesely
3
P Fig. 332. Craugasfnrpalenque; note flap-like inner tarsal fold (arrow). Photo: G. Kohler
b
60a
b 61a
b
to disc); canthus rostralis sharply angular; venter whitish to cream in life; SVL of adult males 34-38 mm, of adult females 65-67 mm ...............hugestor d w l a Discs on Fingers 111-IV broader expanded (1.5 to >2.0 times width of digit just proximal to disc); canthus rostralis rounded to sharply angular; coloration of venter variable; adult SVL variable ........................60 Canthus rostralis rounded .....................61 Canthus rostralis moderately to sharply angular .................................................... 65 Males with single dextral vocal slit (however, possibly an anomaly in single known male); number of vomerine teeth 2-3 in males, 4 4 in females; discs on Fingers 111-IV ~ 1 . times 5 width of digit just proxima1 to disc; SVL of adult males 27-35 mm, of adult females 43-51 mm; only known from two sites in the Montaiia de Celaque, Departments of Lempira and Ocotepeque, western Honduras ..... C r a w anciano vocal in males, if present, bilateral, number of vomerine teeth 2-11 in males, 5-12 in females; relative width of discs on Fingers 111-IV variable; adult SVL variable ..........................................................62
$2a Discs on Fingers 111-IV broader expanded 66a Less toe webbing (modal formula I11 3 4 + (ca. 2.0 times width of digit just proximal to IV); usually with bold lip bars; number of disc); vocal slits and nuptial pads present vomerine teeth 5-9 in males, 7-10 in in males ...................................................63 females; SVL of adult males 29-38 mm, of b Discs on Fingers HI-IV narrower expandadult females 51-67 mm ............................ ed (ca. 1.5 times width of digit just proxi..................................Craugastorcharadra mal to disc); vocal slits and nuptial pads b More toe webbing (modal formula I11 3 - 4 absent in males ......................................64 IV);usually lacking bold lip bars; number 6&i Largest toe discs c1.5 times width of digit of vomerine teeth 3-6 in males, 6-7 in just proximal to disc; venter white, cream, females; SVL of adult males 25-28 mm, of or yellow in life; SVL of adult males 3144 adult females 44-57 mm ............................ mm, of adult females 69-81 mm ................ ..............................Craugastor hepisaimus ............................... Craugestor auzilegulw 67a Vocal slits absent in males; nuptial pads b Largest toe discs >1.5 times width of digit present in males; discs on Fingers 111-IV just proximal to disc; venter yellowish broader expanded (ca. 1.5 times width of green in life; SVL of adult males 33-43 digit just proximal to disc) .................... 68 mm, of adult females 59-62 mm ................ b Vocal slits present in males; nuptial pads ..................................Cra-pecaabsent in males; discs on Fingers 111-IV (Ida Ratio tympanum length / eye length narrower expanded (usually ~1.5times . 1.00-1.14 in males; SVL of adult males width of digit just proximal to disc) ..... 69 18-30 mm (females unknown) ................... 68a Canthus rostralis moderately angulate; ................................. Craugwbr olanchano SVL of adult males 25-36 mm, of adult b Ratio tympanum length / eye length females 58-64 mm ..... C r a e n d m 0.66-0.71 in males; SVL of adult males b Canthus rostralis sharply angulate; SVL 26-45 mm, of adult females 40-74 mm of adult males 41-57 mm, of adult females ....................................Crar;rgerrtorranoidee 76-110 mm ................ C r a e r P e l m @5a Discs on Fingers 111-IV narrower expand- 69a Dorsum rugose; SVL of adult males 28-40 ed (ca. 1.5 times width of digit just proximm,of adult females 58-72 mm ............... mal to disc); vocal slits and nuptial pads ...................................Crampinius absent in males ...................................... 66 b Dorsum smooth to granulate; SVL of adult b Discs on Fingers 111-IV broader expanded males 38-44 mm, of adult females 69-83 (> 1.5 times width of digit just proximal to mm ........................... C r a w t o r #*us disc); vocal slits and nuptial pads present 67 or absent in males ..................................
.....
Figs.333. Craugastor greggi (Finca Santa Julia, San Marcos, Guatemala, 1490 m). Photo: J. Sunyer
Fig. 334. Craugastor charadra (near Tegucigalpita, Honduras, 40 m). Photo: G. Kiihler
Key to the epeciea of the genera Craugastor, ~ a s p oE~l ,e u t h ~ l u sRia, timantis, and S e m a n t i s of southern Central America (Nicaragua to Panama)
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~u3qutqutddiW 6 a Fingers I1 and I11 with palmate disc covers and broadened, non-triangular disc pads; adults with vomerine teeth ........................
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 ....................................DiaspoRu3 diastema 2 a A pale transverse line demarcating finger b Fingers I1 and I11 with spadate disc covers and toe discs on dorsal surface (Fig. 313); and triangular disc pads; adults with or discs on outer fingers (111 and IV)subequal without vomerine teeth ........................... 6 or only slightly larger than those on inner fingers (I and 11) (Fig. 314a); some digits 6 a Venter in most individuals with distinct with palmate, spadate, lanceolate, or papilblack and white blotches; dorsum and dorlate disc covers (Figs. 315f,g); most disc sal surfaces of arms and legs in some indipads broadened or triangular (Figs. viduals bright red in life ............................ 316d,e); Finger I shorter than Finger 11; ........................Diaaponte ventrtpaculatus SVL of adult males 11-22 mm, of adult b Venter patternless or with a few small females 13-26 mm ................................... 3 black dots; dorsum and dorsal surfaces of b No pale transverse line demarcating finger arms and legs brown, cream, or yellow in and toe discs; discs on outer fingers (111 and life .............................................................7 IV) usually distinctly larger than those on inner fingers (I and 11) (Fig. 314b); digits 7 a Posterior surface of thigh unpigmented (yellow in life); adults with vomerine teeth usually with round or truncate disc covers .......................................Diaspom tigrill0 (Figs. 315c,d); disc pads usually ovoid or b Posterior surface of thigh pigmented 316a,b); relative lenghts broadened (Figs. (brownish, often suffused with red in life); of Fingers I and I1 variable; SVL variable -. adults without vomerine teeth .................. .................................................................... 8 ...............................D i a q o h~y~l a e f b d 3 a Disc cover lanceolate or papillate (Fig. 315g) .......................................................... 4 8 a Surfaces of groin and anterior thigh with large contrasting pale blotches on a dark b Disc cover palmate or spadate (Fig. 3150 L: gray to black background (Fig. 335) ....... 9 b Surfaces of groin and anterior thigh with4 a Dorsum shagreened; fingers without thick out large contrasting pale blotches on a lateral fkinges; Toe V not partially fused 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 . .thigh orange blotches of groin and anterior 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 tubercle on upper eyelid present or not ........ 15 Fig. 335. Plistimantis pardalis (Cerro Manl l a Snout long and pointed in dorsal view (Fig. posa, Veraguas, Panama); note white blotche 337a), ratio snout length / head width usuon anterior surface of thigh (arrow). ally >1.1, ratio snout width / SVL usually Photo: G. Kohler ~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 snout length 1head width ~ 1 . 1ratio manuwidth / SVL usuallv >0.37; dorsum 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 subarticu-
lar 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 q a y ) .................... ................................. 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 preseNative) ...............................P 18a Width of largest finger discs greater than 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 latera1 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 b Venter granulate or coarsely areolate; toes 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........................... a
--
Fig. 338. Lateral view of head in a. Pristimantis taeniatus and b. P. ridens. Photos: a. C. Hernandez J.; b. G. Kohler
Fig. 339. Pristimantis gaigei (Nusagandi, Kuna Yala, Panama, 350 m). Photo: G. Klihler
......................................
b Coloration not as above; width of larges finger discs >0.5 times tympanum length; vocal slits present in m a l ~ s........................ ...............................PhEltunantie achatinus 21a Larger £rogs, SVL of adult males to 28 mm, of adult females 47 mm; disc covers truncate ..................... Eleuth-1b Smaller frogs, SVL of adult males to 37 mm, of adult females 52 mm; disc covers 22 usually rounded ...................................... 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 males to 28 mm, of adult females 47 mm ..
b.
Fig. 340. Foot of a. Pristimantis cerasinus; b. Drawings: M. Vesely P. cruentus.
..................... 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 ...................................
..................... Eleuthemdactylusjohnstonel !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 ..........hqgastarjota b Heel smooth to rugose or with 1-3 heel tubercles, no conspicuous large, triangular calcar; no prominent supratympanicsuprascapular 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 .............................
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
................................craugaetorpodi*
b Heel smooth to rugose or with several very small subequal tubercles; subarticular in lateral view; nuptial pads present or tubercles on hands projecting; venter absent in males; SVL of adults variable 28 coarsely areolate; ventral disc V-shaped 28a Finger I1 longer than Finger I; SVL of with its posterior margin reaching beyond males to 18 mm, of females to 23 mm; nuplevel of insertion of hind limbs; nuptial tial pads absent in males .......................29 pads in males present or absent; SVL of b Fingers I and 11 subequal; SVL of adults males to 26.6 mm, of females to 30 mm 27 variable; nuptial pads present or absent in Toes I11 and IV usually without supernumales ....................................................... 30 merary tubercles; subarticular tubercles of 29a Toes with basal webbing; a weak inner hands and feet low and round, usually tarsal fold present ... C r a mIaurantar obtuse in lateral view; nuptial pads present b No thin membranous web between toes; in males; SVL of adult males 16-26 mm, of inner tarsal fold barely evident .................. adult females 18-30 mm ............................. ................................. craugaetorperaimilie ...............................Craugagtor undarwoodi Toes 111and IV with supernumerary tuber- 30a Thenar tubercle much smaller than palmar tubercle. about the same size as subarticucles; subarticular tubercles of hands and lar tubercles of Fingers II-IV (Fig. 343a); feet projecting and pointed, usually obtuse
b
31a
b 32a
b 33a
b
34a
b
36a
b
36a
nuptial pads absent in males; SVL of adult males 12-18 mm, of adult females 13-22 mm .................. Craugastar stejnegehnus 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 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 Drawings: M. Vessljr and b. C.bransfordii. 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 elongatDorsum essentially smooth, lacking ed supraocular tubercle, although several small superciliary tubercles usually presuprascapular ridges, but in juveniles charsent; eyes brown or grayish brown in life acteristically figured weak suprascapular ........................ Strabomantis b&ridges present, which disappear during 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 to contrastingly barred lips, then a distinct Dorsum of adults and juveniles with distympanic stripe present above and posteritinct paired suprascapular ridges; SVL of or to tympanum (Figs. 345 and 346); snout adult females to 70 mm .........................33 slightly protruding with rounded upper Posterior surface of thigh with pale and edge in prome; finger disc non-emarginate dark spotting or mottling; posterior dorsum .................................................................. 38 with a definite laterosacral system of b No dark eye mask as described above, or, if either V-shaped or non-converging parallel dark eye mask present, then finger disc rows of tubercles or ridges .......................... emarginate; snout usually rounded in pro.......................... C r a ~ m e g a o 8 p a a l u s file ............................................................41 Posterior surface of thigh with alternating black and red (white in juveniles and in 38a Discs on outer two fingers (111 and Tt3 markedly larger than those on inner finpreservative) or brown to black with or gers (I and 11) ......................................... 39 without may areas; posterior dorsum without a d e h b laterosacral system of either b Discs on fingers subequal ...................... 40 V-shaped or parallel rows of tubercles or ridges .......................................................34 39a Inner tarsal fold present; sole of foot with definite low plantar tubercles (Fig. 326a); Posterior surface of thigh brightly barred dorsum smooth; glandular supratympanic black and red in life (black and white in ridge extending at least one-third of body . juveniles and in preservative); dorsum ........................................ Craugtwtor noblai often with pale dorsolateral stripes; dorb Inner tarsal fold absent; sole of foot smooth sum strongly tuberculate with numerous (Fig. 326b,c); dorsum granulate; glandular large warts and ridges Craugastor rugasus supratympanic ridge not extending posteriPosterior surface of thigh not brightly orly to axilla ........ Craugastor talamancae barred black and red (or white), although barred with black and tan in some individ- 40a Heel with one or two well-developed calcars; toes with moderate webbing, modal uals; no pale dorsolateral stripes on dorformula I11 3 - 4 IV (Fig. 326c); toes with sum; dorsum smooth with scattered low well-developed lateral fleshy fringes conpustules .......................Craugaetor opitnus tinuous with webs (Fig. 32613; dark seat Finger discs not expanded; head very patch mark forming an inverted V or U .... broad, head width >45% SVL ................36 ....................................... Cmugastor mimus Finger discs expanded; head usually not b Heel without tubercles; toes with basal conspicuously broad, head width ~ 4 0 % webbing, modal formula 111 3 4 IV (Fig. SVL ..........................................................37 344b); toes without lateral fleshy fringes (Fig. 344b); dark seat patch mark usually Upper eyelid with prominently elongated reduced to a pair of dark spots above and supraocular tubercle; eyes blue or grayish lateral to vent, never forming an inverted V blue in life ........... Sh.abamantis laticcupus 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 ......
.......................................
h u g e s t o r 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
Fig. 344.Foot of a. Craugastor crassidigitus; b. C.gollmeri; c. C. melanostictus. Drawings: M. Vesely 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 width of digit just proximal to disc) ...... 43 b Posterior surface of thigh bra- or black 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 surface of thigh with pale mottling; nuptial pads absent in males .... CrarTgastar taupas
r a e evanemm
I I Fig. 345. Craugastormimus (Selva Negra, Matagalpa, Nicaragua, 1300 m). Photo: G. Kiihler
m Fig. a*o. b h u g ~ t o r ~ o l l m e(Nusagandi, ri 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 posterior surface of thigh grayish white without any distinct pattern .. C ~ ~ U @ & b Coloration not as above .........................52 52a Posterior surface of thigh with dark transverse 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, nuptial 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 ......... h u g a s t a r 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 bluishgray 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 C W ~ I1 longer than Finger I; a moderate57a Finger ly-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 -lf 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 59 mottling ................................................... 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 ...........................Cra60a 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
b
6%
i
b
63a
imal to disc); snout subelliptical in dorsal aspect in males, subovoid in females; canthus rostralis distinctly angular; vocal slits and nuptial ads present in males .......62 Discs on Fingers III-IV narrowly expanded (<2.5 times width of digit just proximal to disc); snout usually nearly rounded in dorsal aspect; canthus rostralis rounded or moderately angular; vocal slits and nuptial pads present or absent in males ...........63 Spots on posterior thigh usually discrete and sharply demarcated, at least in adults (Fig. 329a);in life, thigh spots normally yellowish green in individuals in central and eastern Panama, occasionally tan or gray in western Panama; median white gular stripe always present (Fig. 329b), although if throat is generally sparsely pigmented, the stripe may be only weakly indicated as an area devoid of melanophores; SVL of adult males 23-35 mm, of adult females 36-53 mm ................ Craugaetar f i u Posterior surface of thigh with many pale flecks or with larger, coalescing spots that are crowded and poorly defined, these markings normally pale brown or tan in life; melanophores normally distributed across entire with a median white line or very faint ill-defined stripe; SVL of adult males 27-43 mm, of adult females 52-74 mm (individuals of both sexes smaller on the Pearl Islands, Panama) ................. CraugaetorToe discs greatly enlarged, discs on Toes III-V almost 2.0 times the width of the digit; canthus rostralis rounded; posterior surface of thigh with numerous sharply
Fig. 348. Craugastor laevzssmus (&lambe, Nicaragua). Photo: J. Sunyer
contrasting small pale spots on dark brown background; vocal slits and nuptial pads absent in males ........ Craugaetor ranoidw b Toe discs moderately enlarged, discs on Toes III-V 1.5 or less times the width of the digit; canthus rostralis rounded or moderately angulate; posterior surface of thigh with pale spotting or mottling; vocal slits and nuptial pads present or absent in males .......................................................64 64a Toe discs greatly enlarged, discs on Toes 1114 almost 2.0 times the width of the digit; canthus rostralis rounded; posterior surface of thigh with distinct small yellow spots; vocal slits and nuptial pads present in males; SVL of adult males 3 0 4 5 mm, of adult females 45-75 mm .............................
................................... Craugaetor catalinae b Toe discs moderately enlarged, discs on
Toes III-V 1.5 or less times the width of the digit; canthus rostralis moderately angulate; posterior surface of thigh with a few rather small light spots or with pale spotting or mottling; vocal slits present or absent in males; nuptial pads absent in males; SVL of adult males 24-37 mm, of adult females 33-57 mm ........................66 M a Dorsum granulate; posterior surface of thigh with a few rather small light spots; discs on Toes III-V about 1.5 times the width of the digit; vocal slits present in d males ....................Crauga&~ m b Dorsum smooth; posterior surface of thigh with pale spotting or mottling; discs on Toes III-V g1.5 times the width of the digit; vocal slits absent in males ................
..............................C r a u g m h laevissimw
~'lg.349. Craugastor monnichonun (Aha Chiquero, Volch Bani, Chiriqui, Panama, 1820 m). Photo: G. Kohler
Until a few years ago, with over 700 Neotropicd species, the genus Eleu&emdac@lus sensu lato ranked as the most species-rich of all vertebrate genera ( C R A ~ O& RD 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, m d Strabomantidae RAWFOR FORD 80 S~MPPH 2005, ~ C S E et S d.2008). cmrentIy understood, Crauga8bridaecontains 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 (FaosT2010).
~ O S U specia S 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.Ytask this group of frogs. This is especially true with p r e m e d spechens that have lost theb life colaration, which in some w e s may be helpful in distingdehing among ~pecies.
Species of Cmugaator are ge11erdly emtic 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 & WILSON2002). Others are diurnal leaf litter inhabitants and can reach high papulation densities lo& (e.g., members of the br;aasfor& specie8 group; S~Cvrr1983b).Also diurnal leaf lib ter species, the members of tbe goheri species group may pass unnoticed by a person ,walking through the rainforest because of their cryptic domal pattern and coloration, but typically dart across the Recently, CRAWFORD 6t SMun (20051, w d d forest floor in a zigzag pattam sf long leaps on molecular genetic data, elevated the for- when approached too closely (SAVAGE mer subgenus Craugwtor to the rank of 1987). The species of Craugasbr are food genua, This genus represents a group .of generalists feeding on a wide variety of frogs Wt has its greatest diversiiy i-n arthropads and lager species may eat Middle America, and with 93 species, this 8.mall fkogs (SAVAGE 2002). Sexual dimorgenus is well reprersented in Central phism in tympanum she, with males havAmerica, Many epecies of Craugmtur are ing a larger tympanwn k h a n females, is notorious for the high degree of polymor- typical far khe majority of species in phism and polycbomatism, meaning that Craug~tstorQ Y N 2000; ~ Fig. 350).Also, in intraspecific variation 11 coloration, dorsal most species of this genus, females are skin te~ture,patterns of skin tubercles, larger than males (CAMPBELL& SAVAGE war& and ridges can be asto&hingly 2000, SAVAGE 2002). The males of the great in some species (SAVAGE & EMERSON majority of species in this genus have the 1970). Because of this phenomenon, ability to produce vocalizations to attract species diversity can easily be over- or females during the breeding season underestimated. In the case ofthe C.ru&rr- ( S C ~ F F E&R FSGEROA-S~mf 1998). losw specties group, early studies s e e d However,some species (e.g., those of the to grossly cwersimpliEy the situation, refer- goUmd species group) apparently do not ring most Mexican and Nuclear Central emit any kind of calls (%PAGE 19873.As f a American populations to a single species as is known?dl spe@esin this genus have (LYNCH1965d, SAVAGE 1975); currently, a direct development, i.e., they lay encapsutotd of 34 rrpeciaa is recognized in the C. lated eggs in moist terrestrial sites, and
~ i g3.550. ~raugastorpodiciferus;note sexual dimorphism in tympanum size. 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& CALDWELL2009). Nest construction and female attendance of the clutch has been reported for several species of Craugastor (HAYES 1985, MENDOZAQUIJANO 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,8g013Wand is drained by the headwaters of the Rio Las Caiias, which flows into Lago de Izabal", easte m Guatemala. Males to 31.0 mm, females to 40.8 mm. Known only from the type locality, 600-650 m elevation.
1898a),Roc. Craugastor a E e d i (BOULENGER 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. & SAVAGE Craugastor amniscola (CAMPBELL 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. sol. braugastor &edi Chiapas, Mexico).
(El Ocote, Photo: G. Kijhler
Fig. 352.Craugastor amniscola (El Zapotal Photo: W. E. D u e l L m ~ Chiapas, Mexico). (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 elevation.
O,~UU-1- - m
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; typelocality: "Costa Rica: Provincia de San Jose: Cant611 de Coronado: 0.8 km N juncture 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 (SAVAGE19751, 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 (CAMPBELL1994a), Herpetologica 50: 297; type locality: 92.0 km WSW Puerto Santo Tom&, 786 m, De~artamentode 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. Craugastor azueroensis (SAVAGE 1975), Copeia 1975: 277; type l o c a l i ~T : ~prOvincia ~ de 10s Santos: N slope Cerro Cambutal, 480 Males to 40 mm, females to 65 -. Endemic
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 m m (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 BROCCHI1882), 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 (SAVAGE1987), 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. charadra (CAMPBELL & SAVAGE ~Craugastor : 2000), Herpetol. Mono@. 14: 247; type locality: W P W reaches ~ of Webrada San Antonio, SW
-Fig. Costa 355. Rica). Craugastor angelicus Photo: (Monteverde, M.fianzen Fig. 359.Costa CRARC, Craugastor Rica, 550 bransfordii m). Photo: (Guayach, G. Kijhler I
Fig. 356. Craugastor aphanus (Guatemala). Photo: J. A. Campbell
Fig. 360. Craugastor bmchi (Guatemala). Photo: J. A. Campbell
Fig.35;. ~raugastcl~ aJrilegulus (QuetLa,, ~d Om,Atlhtida, Honduras). Photo: J. R. McCranie
Fig. 36;. v~,,,,,~r CL~, (M~-~..iias del Mico, Izabal, Guatemala). Photo: J. A. Campbel
Fig. 358. Craugastor bocourti (Guatemala). Photo: J. A. Campbell
Fig. 362.Craugastor chac (Cockscomb b s e r v e , 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 & WILSON19891, 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 cyanochthebius MCCRANIE& SMITH2006, 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 & SAVAGE19841, 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.
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 (TAYLOR1952b), 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 3 1 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 (SAVAGE1980), 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.
dig. 363. Craugastor daryi (3.8 k m SE PurulhB, Baja Verapaz, Guatemala). Photo: W. E. Duellman (Courtesy of the Biodiversity Institute,University of Kansas)
Fig. 364. Craugastor emcelae (north slope Cerro Pando, Bocas del Toro, Panama, 1350 m). Photo: W. E. Duellman (Courtesyof the Biodiversity Institute,University of Kansas)
Fig. 365. Craugastor chrysozetetes (Quebrada de Oro, Atlantida, Honduras). Photo: J. R. McCranie
Fig. 369. Craugastor charadra (near Teguci Photo: G. KO----pita, Honduras, 40 m).
Fig.366. Craugastor coffeus(holotype). Photo: G. Kohler
Fig. 370. Craugastor chingopetaca (Rio San Juan, Nicaragua, 30 m). Photo: J. Sunyer
I
Fig. 367. Craugastor crassidigitus (Guayach, CRARC, Costa Rica, 550 m). Photo: G.Kiihler
Fig. 371. Craugastor emleni (Los Golondrinos, Francisco Morazh, Honduras). Photo: J. R. McCranie
Fig. 368. Craugastor cyanochthebius (San Isidro, C o p h , Honduras). Photo: J. R. McCranie
,Fig. 372. Craugastor epochthidius (Quebrada 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.
I Fig. 373. Craugastor escoces (Bajo La Hondura, San Jos6, Costa Rica). photo: W. van Devender
Craugastor evaneseo RYAN,SAVAGE,LIPS & GIERMAKOWSKI 2010, Copeia 2010: 405; type locality: "Panama, Cocl6 Province, Parque ~acionalG. D. 0mar Tonijos H. (El CO&, headwaters of Rio Guabal. 8"39.59'N. -.. - . ..80°35.33W, 769 m.a.s.1. elevation." Males to 50.2 mm, females to 64.3 mm.Eastern portion of the Serrania de TabasarB, west-central 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.
Fig. 374. Crauga--- - ---:o
(holotype). Photo: M. Ryan
Craugastor emled ( D m 1932), Copeia 1932: 97; type locality: "San Juancito, in the cloud forest at 6600 feet and ... at 6000 feet 11829 m]," Departamento Francisco Morazh, Honduras. Males to 28.1 mm, females to 49.8 mm. On the Pacigc versant of south-central Honduras CRio Choluteca drainage system between Tegucigalpa and about El Zamorano, Departamento Francisco Morazh), 800-2000 m elevation. Craugastor epochthidius (MCCRANIE & WILSON 1997a),Alytes 14: 169; type locality: YRio Seco (14"55'N, 85'56'W), elevation 1050 m, Ixibutary of Rio Guayape N of Catacamas, Sierra de Agalta, Departamento de Olancho, Honduras." Males to 27.4 mm, females to 36.7 mm. Sierra de Agalta and Sierra Punta Piedra in the departments of Olancho and ColBn, east-central and northeastern Honduras, 150-1459 m elevation.
Craugastor fitzingen (SCHMIDT 18571, Sitzungsber. Akad. Wiss. Wien Phys. Math. Naturwiss. K1. 24: 12; type locality: "Neugranada"; SAVAGE'S (1974) designation of LACM 76859, from Barro Colorado Island, Panama Canal Zone, as neotype was consid& MYERS(1983). Males ered invalid by LYNCH to 35 mm, females to 53 mm. Northeastern Honduras to northwestern Colombia, near sea level to 1520 m elevation. Craugastor fleischmanni (BOETTGER 18921, &t. Batr, Samml. Mus. Senckenb. Naturforsch. Ges.: 27; type locality: "San Jod, [Cantbn de San Jos6, Provincia San Jose,] Costa Eta." Males to 47 mm, females to 75 mm. On the Pacific slopes of the volcanoes Po& and Barva, and on the Atlantic slopes of the volcanoes Irma and Turrialba, and on both slopes of the western portion of the Cordillera de Talamanca, Costa Rica, 1050-2286 m elevation. Craugastor glaueus (LYNCH1967a1, Trans. K m a s Acad. Sci. 70: 177; type locality: "1.6 km SW San Cristobal [de las Casas], Chiapas, M6xico,2100 m." Males to 31.7 mm (maximum sbe for females not available). Only known h m the type locality.
Fig.375. Craugastorfecundus (Quebrada de Oro,Atlhtida, Honduras). Photo: J. R. McCranie
Fig. 379.Craugastorglaurms (San Crld6bal de Photo: T. Bille las Casas, Chiapas, Mexico).
Fig. 376. Craugastor fitzingen (Lagarto Lodge, Alajuela, Costa Rica, 90m). Photo: G. Kijhler
r'ig.mu. craugastor golunen (nusagami, Kuna Yala, Panama, 350 m). Photo: G. Kohler
Fig.377. Craugastor fitzingen' (Lagarto Lodge, Alajuela, Costa Rica, 90m). Photo: G. Kohler
Fig. 381. Craugastorgreggi (Volch Tacand, Chiapas, Mexico). Photo: L. Gray
1 Kg.378.Craugastor fitzingeri (near Uvita, P ~ t a r e n a sCosta , Rica, 10 m). Photo: G. Kohler
Fig. 382.Craugastor gulosus (Cerro Frio, Photo: E. Boza Bocas del Toro, Panamd).
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 correctingthe 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. 19551, Copeia Craugastor greggi (BUMZAHEM 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 (COPE18761, 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, n e w 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.
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& ESPINAL1996), 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. 1879),Bull. Soc. Craugastor lineatus (BROCCHI 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 20001, sea level to 2100 m eleSalvador (LYNCE vation. 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
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 ~ O2002). N 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
Fig. 383. Craugastor inachus @om near type locality). Photo: J. k Campbell
Fig.384. Craugastorjota (Rio Changena, Bocas Photo: A. Hertz del Toro, Panama, 1765 m).
Fig. 388.Craugastor lineatus (Santa Barbara, S slope of Volcfin Atitlb, Suchitepbquez, Guatemala, Guatemala). Photo: G. Kohler
IIF!
Fig.385.Craugastor laevissimus (P.N Aff --eico Norte, Nicaragua, 780 m). Photo: G. Kohler
intla, Guatemala, 300 m).
Fig. 386. Craugastor laticeps (Cerro San Gil, Photo: G. Kohler Izabal, Guatemala).
Fig. 390.Craugastor longirostris (Cana, DariBn, Panama, 1597 m). Photo: A.J. Crawford
n g . 38.1. craugastor lauraster (P.N. Saslaya, Atlantic0 Norte, Nicaragua). Photo: G. Kohler
Fig. 391. Craugastor megacephalus (Cerro El Toro, Nicaragua, 830 m). Photo: G. Kohler
Photo: G. Kohler
part of Guayas Province in Ecuador, sea level to 1200 m elevation. Craugastor matudai (TAYLOR1941), 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.
the slope of Chiriqui volcano above Boquete, ChiriauIi Province". Panama. SVL to 65.7 mm. west& cordillera de Talamanca of Panama, 610-1830 m elevation. Craugastor montanus (TAYLOE1942), 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 HEDGESet al. (2008).
20001, Rev. Craugastor myllomyllon (SAVAGE 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, Herpetologica 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 Guatemala, 800-1000 m elevation. & DUNN1921), Craugastor noblei (BARBOUR 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 (BARBOUR19281, 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.
Craugastor mimus (TAYLOR19551, Univ. Kansas Sci. Bull. 37: 517; type locality: "5 km. NNE T i l a r h , [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 (DUNN19403, Proc. Acad. Nat. Sci. Philadelphia 92: 107; type locality: "Valley of the Velo, Finca Lerida, on
Fig. 392. Craugastor melanostictus (Pico Picacho, Chiriqui, Panama). Photo: G. Kohler
It R8.393. C r a u ~ a s t omilesi r (Y.N.I;usuco. Photo:
-
PIE.391. t i r a u ~ a s t onetrens r (rmca ~a
Kolby
Firmeza, ~ o r i e sIzabal, , Guatemala). Photo: C. R. Vhsquez Almazhn
Fig.394.Craugastor mimus (Lagarto Lodge, Alajuela, Costa Rica, 90 m). Photo: G.Kohler
Fig.398. Craugastor noblei (Selva Negra, Matagalpa, Nicaragua, 1300 m). Photo: G. Kohler
m , ,
I
395.Craugastormonnichom (AteCbique,Wqui, Panama, 1820 m). Photo: G.Kijhler
b &g. YYY. tiraugastor oDesus ( ~ o ~ q Photo: K-H. Jungfer Panama, 1050 m).
w
1 Fig.396.Craugastor montanus (Volchn Tacanh, Chiapas, Mexico, 900 m).Photo:k Ramirez Velslzquez
Fig.400.Craugastor olanchano (Quebrada El 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. & WILSON Craugastor omoaensis (MCCRANIE 1997a),Alytes 14: 155; type locality: "about 10 airline km WSW San Pedro Sula on road to Pen3 (15"28'N, 88"06W), elevation 1150 m, 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. & MYERS2002), Craugastor opimus (SAVAGE 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.
Craugmtor paIenque (CAMPBELL & SAVAGE 20001, Herpetol. Monogr. 14: 221; type locality: "10.7 km SE Cristbbal Col6n on road to Bonampak, ca. 350 m, Chiapas, Mexico. This site lies in Bemontane Wet Forest at about 17"08'N, 91°34W." Males to 43.1 mm, females to ~ 5 mm. . ~c&bbean slopes of northern Chiapas, xi^^, and adjacent ~ ~ where it is horn eom tributaries that drain into the lower mo Grijalva, 200400 elevation.
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Fig. 401. Craugastor opimus (Altos de Cerro Azul, PanamB, Panama, 820 m). Photo: A. J. Crawford
Fig. 402. Craugastor pechonun (Quebrada Machin, Col6n, Honduras).Photo: J. R. McCranie
drain into the lower Rio Grijalva and K O Usumacinta, 350-1700 m elevation. Craugastorpersimilis (BARBOUR 1926), Occas. Pap. Boston Soc. Nat. ~ ~ Hist. 5:~193; type locali~ ty: "Suretka, Costa Rica, near the boundary of Bocas del Toro Province, Panamd." Males to 17.8 mm, females to 23 mm. On the Atlantic versant in central to southern Costa Rica, 40-1200 m elevation.
& WILSON Craugastor pechorum (MCCRANIE 1999b1, Proc. Biol. Soc. Washington 112: 515; Craugastor phasma (LIPS & SAVAGE1996), dam alonga type locality: "from near a Roc. Bid. Sot. Washington 109: 7 e type small tributary of the Quebrada de Las Marfa8 locality: "Costa Rica: Puntarenas Province, (15017.64'N, 84021.29~),about 12 airline km Cmt6n Coto Brus, Zona Protectors Las L~ coloda, D~~~~~~~~~~ de 0lancho, Finca Jaguar, ~ ~ 680 rn elev." ~ ~ d a to 42.5 l ~ ~-, ~ ~ Tabla, ~ ~ 8"55N , 82"44W on the 20 km of La Lucha, females to 62.1 mm. On the Atlantic versant of elevation." Single known specimen 47.9 mm and eastern portions of the the Known from the type locality. Honduran departments of Olancho and Colbn, respectively, i50-680 m elevation. Craugastor podiciferus (COPE1876), J. Acad. Nat. Sci. Philadelphia Ser. 2,s: 107; type localcraugastor pelorus (C-BEL~ & sAVAGE ity: "from the level of 5000 to 7000 feet ... on 2000), Herpetol. 14: 218; type locality: "2.7 km N ~ ~ l ~ca. 400 ~ m, ~ chiapas, ~ h i the ~ Pico ~ Blanco", ~ , Costa Rica; corrected to "Cerro Utyum, 5000-7000 ft, Cant6n de ~ ~hi^ site ~ lies iin premontane ~ ~ wet , Provincia de Limbn: 1524-2134 F~~~~~ at about 17027'N, 93003v." ~ a to l Talamanca, ~ ~ m", Costa Rica by SAVAGE (1974). Males to 28 56.8 =,females to l l O mm. . ~ caribbean Both 'lopes of the to 40 slopes of northern Chiapas, Mexico, where it is cordilleras of Costa Rica and adjacent western upper tributaries that known from the Panama, 1089-2650 m elevation.
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Fig. 403. Craugastor persirnilis (Guayacan, CRARC, Costa Rica, 550 m). Photo: G. Kohler
Fig.407.Craugastor podiciferus (Guacuyo, Heredia, Costa Rica, 1850 m). Photo: G. Kohler
Fig.404.Craugastorpodiciferus (Alto Chiquero, Chiriqui, Panama, 1820 m). Photo: G. Kiihler
Fig.408. Craugastor podicifem P a s o Llano, Heredia, Costa Rica, 1700 m). Photo: G.Kohler
Fig.405.Craugastorpodicifem(Volch Barva, Heredia, Costa Rica, 2000 m). Photo: G.Kiihler
Fig. 409. Craugastor polyptychus (P.N. Manza nillo, Lim6n, Costa Rica, 30 m). Photo: G. Kijhler
Fig.404.Craugastorpodzciferus(Alto Chiquero, Chiriqui, Panama, 1820 m). Photo: G.Kiihler
Fig.410. Craugastor polyptychus (same as in 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. & SAVAGE 19951, J Craugastor pozo (JORNSON Herpetol. 29: 501; type locality: "12 km NW oBerriozAbal (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.
Fig. 411. Craugastorpozo (12.5 kilometers North of Berriozabal, Chipas, Mexico, 1100 m). Photo: T. J. Papenfuss
Craugastor psephosypharus (CAMPBELL, SAVAGE & MEYER19941, 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 Fig. 412. Craugastor psephosypharus (11.8 k m Belize, 150-1170 m elevation. WSW Puerto Santo Tom& Izabal, Guatemala). Photo: J. A. Campbell Craugastor punctariolus (PETERS1863b), (Courtesy of the Biodiversity Institute,University of Kansas) 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 elevaEscudo de Veraguas), exclusive of the Golfo tion. Dulce region of southwestern Costa Rica, sea level to 1300 m elevation. 1937a), Trans. Craugastor pygmaew (TAYLOR Kansas Acad. Sci. 39: 352; type locality: 1 ' mile Craugastor ray0 (SAVAGE & DEWEESE1979), north of Rodriguez Clara, Vera Cruz", Mexico. Bull. S. California Acad. Sci. 78: 107; type SVL to 18 mm. Extreme southern Michoach locality: "second sabana on the trail from Finca and M6xico (state), Mkco, south and east El Helechales to Sabanas Esperanza, 5 km, along the Pacific versant to western airline,east of Finca El Helechales, Cant611 de Guatemala; also in Veracruz and eastern Buenos Aires, Provincia Puntarenas, Costa Oaxaca, Mexico, on the Atlantic slope; sea level Rica, 1640 m elevation." Males to 45 mm, to 2000 m elevation. females to 71 mm. Pacitic slope of the Cordillera de Talamanca of southern Costa 18961, Craugastor raniformis (BOULENGER Rica, 1480-1820 m elevation. Ann.Mag. Nat. Hist. Ser. 6,17: 19; type locality: "Buenaventura and Cali", Departamento Craugastor rhodopis (COPE18671, Proc. Acad. Valle del Cauca,] Colombia. Males to 43.2 mm, Nat. Sei. Philadelphia 18: 323; type locality: females to 74.0 mm. East-central Panama to "Vera Cruz, at Orizava [= Orizabal and western Colombia, near sea level to 1500 m Cordova I= Cbrdobal", Veracruz, Mexico; elevation. Mexico, by lecrestricted to Orizaba, Vera-, totype designation (SMITH & TAYLOR1950). Craugastor ranoides (COPE1886a), Proc. Am. Males to 28.5 mm, females to 45.7 mm. Philos. Soc. 23: 275; type locality: "Nicaragua." Western V e r a m and adjacent Hidalgo and Males to 45 mm, females to 74 mm. Eastern Puebla in Mexico; also in central and southNicaragua on the Atlantic slope and northeastern Chiapas and adjacent Oaxaca, mostly western Costa Rica on the Pacific versant to 100-1000 m elevation. extreme western Panama (including Isla
!
FSg.413.Craugasiarpunotbuiolus(El Valle de Anttin, C d d , Panama, 800 m). Photo: M. Ryan
Fig. 416. Craugastorranaides (5 lun W A h h n t e , 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 ~
Fig.414. Craugastorpygmaats (30 km N San Gabriel Itfixtap, Oaxaca, Ikhdco).
Photo: W.E. Duehan
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~ g415. . Craugmtorradio&
Dmidn, Panama, l322;m).
(Chepigam Photo: M. Ponce
~ S B B )
fig. 417. Cmugwtor rh~dopis ( r m a Rancho Alegre*Ghiapaa, Msxioo, ilOO m). Phob: G. &@or
ff
Fig.418.Cmugastor dpulw. Photo: J. A. Campbell
I
m
Craugastor rhyacobatrachus (CAMPBELL & SAVAGE 2000), Herpetol. Monogr. 14: 274; type locality: "Quebrada Fortuna, 19 km N San Isidro de El General, 1700 m, Cant6n PerezZeledbn, 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 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 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 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.
Craugastor rostralis (WERNER1896), 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 elevation.
Craugastor silvicola (LYNCH 1967b),Proc. Biol. Soc. Washington 80: 211; type locality: "12 mi. NNE Zanatepec, Oaxaca, Mkxico, 4900 ft. elevation." SVL 40.2 in single known female. Only known from the type locality, 1450-1600 m elevation.
1
1
3
r
1
i
Craugastor rugosus (PETERS 1873),Monatsber. Preuss. Akad. Wiss. Berlin 1873: 610; type locality: "Chiriqui", Panama. Males to 44 mm, Paciiic versant of central females to 69 k. and southern Costa Rica and adjacent Panama, sea level to 1220 m elevation.
i
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 Fig. 419. Craugastor rugosus (Refugio El Chiapas, Mexico, southeastward through Photo: G. Kohler Rodeo, Costa Rica, 995 m). 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 Fig. 420. Craugastor rostralis (P.N. Cusuco, about 900 m elevation. Photo: J. Kolby Cartes, Honduras, 660 m).
L I
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m
Fig. 421. Craugastor rupinius (El Imposible, Ahuachaph, El Salvador, 720 m). Photo: G. Kiihler
Fig.425. C s a u g a ~ t stadRZmd o~ ( I airline b NNE of La Fofttma,Yoro, Honduras). Photo: J. R. McCrmie
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Fig.426.Craugastor+-w Fig. 422. CPauga&r 5abau8 (Gem San GI1, Mal,Guatemda, 160 m). Photo: G.Kahle~ Puntarenas, Costa Rica, 20 m).
I
I
Wends Bani,
Photo: G. Kijhler
Fig. 423. Craugastor saltuarius (south slope of Cerro Blifalo, Atlhtida, Honduras). Photo: J. R. McCranie
Fig. 427. Craugastor stuarti (Jaltenango E l Triunfo], Chiapas, Mexico, 1960 m). Photo: A. Ramirez Veldzquez
Fig. 424.Craugastor sandersoni (Sierra de Chinajd, Alta Verapaz, Guatemala, 615 m). Photo: J. Sunyer
Fig. 428. Craugastor tabasarae (La Nevera, Photo: G. Kohler Ngobe Bugl6, Panama).
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. Craugastor stejnegerianus (COPE18931, Proc. Am. Philos. Soc. 31: 338; type locality: "Palmar"* Provincia Puntarenas, Costa @& Males to 18.0 mm, females to 22.4 mm. Pacific versant of Costa Rica and western Panama, sea level to 1330 m elevation.
I Fig. 430. Craugastor t a m s (Las Mellizas, Punts Burica, Chiriqui, Panama, 120 m). Photo: M. Ponce
Craugastor stuarti (LYNCH1967a). 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,HOLLINGSWORTH, LIPS& JASMW 20041, Herpetologica 60: 521; type locality: "9.7 k m NNW El Cop6, on the continental divide at sawmill, La Pintada Fig. 431. Craugastor trachydermus (from type District, Cocl6 Province, Panama, 600400 m locality). Photo: J. A. Campbell (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 (TAYLOR1958b), 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 known from-the type locality. Craugastor trachydermus (CAMPBELL 1994b), 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
Fig. 432. Craugastor talamancae (Nusagandi, Kuna Yala, Panama, 350 m). Photo: G. Kohler
33.5 mm, females to 46.1 mm. Known only from the type locality 1896b1, Craugastor underwoodi (BOULENGER Ann. Mag. Nat. Hist. Ser. 6,18: 340; type local-
ity: "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, 1krn. 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.
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 BiodiversityInstitute,University of Kansas);
b. C. R. VBsquez Almazh
L
C. crassidigitus C. chingopetaca
Craugastor
I
C. fitzingen
C. talarnancae
Craugastor
'
Craugastor
5ransfordii lauraster
2I
f
A
2. xucanebi
..
:. loki :. rnyllomyllon
,.
rhodopis 2. chrysozetetes >.ornoaensis 8 C. coffeus
a
yucatanensis C. nefrens .L'. : glaucus taylori C. silvicola
r C. rostralis A C. epochthidius
--2. melanostlctus
C. saltuarius
.,
tabasarae S. monnichorum
r C. pygmaeus
Craugastor
r
8 C. stadelrnani
r C. rnatudai A
C. fecundus
S. erncelae S. longirostris C.cuaquero C. phasma
C. Iaeviss~mus
I C.rupintus
C. sabrinus C. pelorus C. pozo C. emleni C. montanus
C. inachus
Craugastor sandersoni
I c. rivulus
1C. amniscola
% C. olanchano
C. merendonensis C. pechorum
V C. rhyacobafrachus
C. chingopetaca
I
I
C. ranoides
E C. escoces IC.punctario/us -
V C. catalinae
I
I
Further Reading
SAVAGE 1975. SAVAGE & DEWEESE1979, LYNCH 1980b, SAVAGE 1980, SAVAGE & DEWEESE1981, MIYAMOTO 1983, Ford & SAVAGE 1984, MIYAMOTO & TENNANT 1984, LYNCH 1985a, M~PAMOTO 1986, et SAVAGE 1987, SAVAGE et al. 1988, MCCRANIE al. 1989, C A ~ B E L1994, L CAMPBELL et al. 1994, & SAVAGE 1995, LYNCH 1996, SAVAGE et JOHNSON al. 1996, MCCRANIE & WILSON1997a, KOHLER 1998, LYNCH1998, DONNELLY 1999, HOBEL 1999a, M c C m & KOHLER1999b, MCCRANIE & WILSON 199913, CAMPBELL & SAVAGE 2000, & WILSON LYNCH 2000, SAVAGE 2000, MCCRANIE & MYERS2002, F'USCHENDORF & 2002, SAVAGE CRAWFORD 2003, SAVAGE et al. 2004, CRAWFORD & S m 2005, SMITH2005, KOHLER& SUNYER 2006, HEDGES et al. 2008
Eleutherodactylidae
Eleutherodactylidae The recognition of the family Eleutherodactylidae containing two subfamilies (Eleutherodactylinae and Phyzelaphryninae), four genera, and a total of 201 species follows HEDGESet 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 ~ G e r i c aand 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 jumping (Scorn 1983~). Food consists of a variety of small arthropods, including ants (TOW1981).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, W n c m s m 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 petioles, and other cavities (TAYLOR1955, 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 (COPE18761,J. Acad. Nat. Sci. Philadelphia Ser. 2, 8: 155; type 1ocalit;pr "camp Mary C m t t a , Panama." Males to 2% mm, females to 24 mm. Southeastern Honduras to eastern Panama; sea level td 1620 m elevation.
Diasporu~ih 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 eleLim&n,Costa Rica; corrected to vation", ROV. "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 (LYNCH20011, 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,
n I
.Kn o localitie valley of the Rio Lmi, in Limb W v i n c e , 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 k m 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.I
Fig. 434.Diaspom ventrimaculatus (Valle del Silencio, Costa Rica). Photo: R. Vargas
~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; icentral Panama also on the Atlantic versanl EL&w@@~ to 1220 m elevation. ;Q p
.‘
7,
Figs. 435-436.Diaspom ventrimaculatus (Valle Photo: R. Vargas del Silencio, Costa Rica).
Fig. 43Y. Uiasporus dastema (tiuayacan, CRARC, Costa Rica, 550 m). Photo: G. Kohler Fig. 437.Diasporus diastema (Guayach, CRARC, Costa Rica, 550 m). Photo: G. Kohler
Fig. 4;-. 3 i a s p o m vocatc- .-Jusagandi, Kuna Yala, Panama, 350 m). Photo: G. Kijhler
Fig. 440.Diaspon Panama).
-1.
(Fortuna, Chiriqui, 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., STEJNECER1904, CRAWFORD & SMITH2005). After the splitting of the genus Eleutherodactylus into several families and genera, Eleutherodactylua, as currently understood, forms a group of 185 species that has its center of et al. diversity in the West Indies (HBDGES 2010). The genus is distrib2008, FROST uted throughout the West Indies, Florida, and southern Texas, USA, Mexico, Belize, and Guatemala (HEDGES et al. 2008); introduced to Honduras, Costa Rica, and & RAND1990, m z et al. Panama (1-z 1999b, SAVAGE 2002, MCCRANIEet 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 (Eleutheroda~l~~). The wide distribution and ability to successfully 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 subgenus Sphophus are nocturnal inhabi-Rsading SCOTT1983c,LYNCH2001, OVASKA& RaMD 2001, tants of open areas, including altered habibackyards; MCCRANIE et al. 2002, SAVAGE 2002, CHAVEBet tats such as gardens and they can be locally abundant (Vow et al. a1.2009 1997). ~ o u g h o u the t 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 Eleutherodactylus 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. 444. Eleutherodactylus coqui (Turrialba, Cartago, Costa Rica, 620 m). Photo: G. Kohler
Fig. 441. Eleu therodactylus an tillensis Photo: B. Barker (Panama City, Panama).
I d Fig. 442. Eleutherodactylus johnstonei (Mandeville, Jamaica, 650 m). Photo: K.-H. Jungfer -
& 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 THOMAS1966, 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. 1914, Eleutherodactylus johnstonei BARBOUR 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.
rclg. 44s.fi~eutherodactylusp~a.rurosms w a n Pedro Sula, Cortb, Honduras). Photo: J. R. McCranie
Eleutherodactylus leprus (COPE18791, 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
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 (TAYWR19401, Roc. Biol. SOC.Wa8hbgton 53: 95; type lucality: '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 ap ~l lhwh 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. Fig.447. Eleutherodactylus pipilans (Sierra Madre de Chiapas, Arriaga, Chiapas, Mexico, Eleutheroda&tlw rubrimactzlam (TAYLOR & Photo: A. Ramirez Veliizquez SMITa IS&), Proc. U.S. Natl. M w 96: 583; type 690 m). 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 1000m elevation.
-
~ e u t h e r o d a C ~ l mbrimacula u s tus (Finca Irlanda, Chiapas, Mexico, 1050 m). Photo: G. Kijhler
1
LYNCH1970a, SCKWARTZ & HENDERSON 1991, Fig. 446. Eleutherodactylus lepnrs (Xalapa, KAISER& HARDY 1994. KAISER1997, IBAI?EZet Photo: P. ~ e i m e s d . 1999b, KAISERet d.'2002, ~AvA~E.2002 ~eAcruz,Mexico).
Strabomantidae HEDGESet al. (2008)introduced the new family Strabomantidae to accommodate some 560 species of Neotropical directdeveloping frogs distributed from eastern Honduras m o s s 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) (FROST2010). 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.
-
On the basis of molecular evidence, SIEINICKE et al. (2007) resurrected the genus PTistimantis from synonymy with [email protected] 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 (FROST2010). The majority of species is distributed in tropical South America, whereas only 14 species are known to occur in Central America ( D ~ L L M A&NLEHR2009).
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. Readin€! HFINICKEet al. 2007, HEDGES et al. 2008, PADIAL cerasinus and P. ridens can be found in the et al. 2009 1986).Other species, leaf litter (LIEBERMAN 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 Fig.448.Pristimantis achatinus (Cerro ~ u i is a direct and froglets hatch out of the eggs Photo: W. E. Duellman without a free-swimming aquatic tadpole Daribn, Panama). (Courtesy of the Biodiversity Institute, Universiw of Kansas) stage (SAVAGE 2002).
rn I Fig. 449. histimantis adnus (holotype). Photo:A. J. Crawford
A noteworthy phenomenon is the suggested mimicry relationship between &tiinantis gaigei and poison-dart frogs of the genus Phylobatea (DUELUUN & TRUEB 1986,TOLEDO & HADDAD 2009).The palatable Pristimantis gaigei has a color pattern closely resembling the highly toxic potential models Phyllobates aurotaenia and Phyllobates lugubris. Pristimantis gaigsi occurs syntopically with the latter species in soubheastern Costa Rica and adjacent northwestern Panama, and with
P. aurotaenia in northwestern Colombia (LYNCH1985b). Unlike most other Pristimantis, 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 -, females to 46:l mm. Extreme eastern Panama (DariBn Province) to southern Ecuador, sea leveI to 900 m elevation. Pristimantis adnus CRAWFORD, RYAN & m 2010, Herpetologica 66 (2):194; type locality: "along a trail approximately 700 m in elevation at the foot of Cerro Piiia in the Serrania del Sapo, above the Rfo Piiia on the Pacific coast of the Dari6n Province, Republic of Panama." Males to 20.0 m, females unknown. Known only from the type locality.
J
i road k m NW Fig. 450. Pristimantis altat S& Ram6n, Alajuela, Costa Rica, 1233 m). Photo: A. J. Crawforfl
F'! -*,-
-
--.- - -
.
-. -
- .- -
-.
.
_-_
--
-.
,/
Fig. 451. Pristimantis caryophylaceus (La Nevera, Ngobe Bugl6, Panama). Photo: G. Kiihler
Pristimantis altae (DUNN1942b), 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 d e . 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 locality: "La Loma on the trail from Chiriquicito to Boquete, Bocas del Toro 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 v m a n t from extreme southern Costa Rica to western Panama, sea level to 1900 m elw?tion. Pristimantis cerasinus (COPE18761, J. Acad. Nat. Sci. Philadelphia Ser. 2,B: 112; type locality: *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. Northeastern Honduras to eastern Panama
Fig. 452. Plistimantis caryophyllaceus (female with eggs; Alto de Piedra, Veraguas, Panama, Photo: A. Hertz 910 m).
I
dm-l
Fig. 453. Pristimantis educatoilD&om type Photo: A. J. Crawford locality, 800 m).
.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 & WILSON1999 is regarded as a synonym of P. cerasinus according to MCCRANIE &WILSON2003b.
~ i454. ~ fiStUdtiS . bGx asinLu ,Lbancho~ ~ ralista, Costa Rica, 900 m). Photo: G. Kohler
I
Fig. 455. Pristimantis cerasinus (Nusagandi, Kuna Yala, Panama, 350 m). Photo: G. Kohler
Fig. 456. Pristimantis cruentus (Albergue Monterreal, Alajuela, Costa Rica, 1850 m). Photo: G. Kohler
Pristimantis cruentus . (PETERS 18731, Monatsber. Preuss. Akad. 'Wiss. Berlin 1873: 609; type locality: "Chiriqui", Panama (see comment under Ameerega maculata). Males to to 42 mm. On the Atlantic vert 28 mm, ~ females sant from northern Costa Rica to western-central Panama, 40-1800 elevation. 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 k m 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 (DUNN19311, 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 BRANDONJONES et al. (2007) in the usage of the species' name as originally spelled, and not the emended name ("gaigeae"), as proposed by T A ~ O R (1952b). Pristimantis moro (SAVAGE19651, 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; "Culebra (08"52'14"N, type locality: 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. 457. Pristiman tis gaigei (Changuinola, Bocas del Toro, Panama, 10 m). Photo: M. Ponce
Fig. 458.Pristimantis moro (ElCop6, Cocl6, Panama, 700 m). Photo: B. Kubicki
Fig. 462.Pristimantis ridens (near Rio Sereno, Chiriqui, Panama, 1210 m). Photo: G.Kohler
Fig. 459.Pristimantis museosus (Cerro Negro, Veraguas, Panama, 770 m). Photo: A. Hertz
Fig. 463.Pristimantis ridens (9k m NW Fortuna, Alajuela, Costa Rica). Photo: G. Kohler
Fig. 460.Pristimantis pardalis (Cerro Mariposa, Veraguas, Panama, 870 m). Photo: G. Kijhler
Fig. 464.Pristimantis ridens (P.N.Saslaya, Atlantic0 Norte, Nicaragua, 820 m). Photo: G. Kohler
Fig. 461.Pristimantis pirrensis (Cana, Darikn, Photo: A. J. Crawford Panama, 1300 m).
Fig. 465.Pristimantis taeniatus (Los Santos, 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&& northwet~ wSern P50-1qQ J&< elevation :. ,, L
a
J%stimantisg&nais (& CRAWOW,' 2004), J. Herpetol. 38: 241; type locality: ' ' a i o ~ , 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 1250-1550 m elevation.
m.
I
I
caryophY1lalaceus 9 pardalis 9 pirrensis
1 d to 18fB &I @?m&.Qian. I'rfamg8 @OULEpBieTp 1.SWh Ann. Mag, 'W.%. Bex+$10: 18% Qp ladity: "NoE. N-al, Rio Raa J m Chco, $.X COED*.^^ p.1mm* f d w tjg 8g.k 'wecentral P (--A& &fib Haita Tabmar4 I$&@ 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 Pristimantis
SAVAGE 1981b, MIYAMOTO1984, LYNCH1985b, k4Fmz et al. 1994, LYNCH& ARDILA-ROBAYO 1999, IBAREZ& CRAWFORD 2004, CRAWFORD et al. 2010, RYANet al. 2010
1 P. cruentus I? moro
Strabomantis
1896a),Ann.Mag Nat. Hist. Ser. 6,17:19;type locality: UBuenaventuran, Colombia. Males to 69 mm, females to 94mm.On the Atlantic versant from extreme southern Costa R i a to western Colombia, 16-350 m elevation.
The genus Strabomantis was removed from synonymy with Eleutherodactylus by HEINICKEet al. (2007). It includes 17 Strabomantis laticorpus (MYERS & LYNCH species of broad-headed frogs distributed 1997),Am. Mus. Nwit. 3196: ;4; type locality: in southern Costa Rica and Panama, as "montane forest on southwest sector of Cerro well as through the wet tropics of ColomTacarcuha mass% 1590 m elevation, Dari6n Province, Republic of Wnama. Approximatsly bia, northern Venezuela, Ecuador, eastern 8"10W, 77O4OW." Males to 30 mm, females to Peru, and western Brazil (FROST2010). 40 mm.Cerro Tacarcuna and Cerro MaIf in the? Just as with most of the species in this southeastern Daribn Province, Panama, genus, both Central American representa1460-1630 m elevation. , rb:.. ,, = . ' tives are toad-like terrestrial frogs. Strabomantis laticorpus is reported ti be active Rsading in the leaf litter by day, but also encoun- Further MYERS & LYNCH 1997, wz et al. 1999b, tered at the edge of a stream at night sAVAGE 2002, DUELLMAN & LEHR2009 (MYERS & LYNCH 1997).Strabomantis bufoniformis is a nocturnal terrestrial streamassociated frog that can be found on the ground along small forest streams at night (K~HLER unpubl. obs.). I B ~etZal. (1999b) reported males calling from the edges of small streams just after dusk. The call of S. bufoniformisconsists of paused short low bark-like notes followed by even shorter knock-like ones ( I B ~etzal. 1999b). -7
Fig. 466. Strabomantis bufoniformis(Nusagandi, Kuna Yala, Panama, 350 m). Photo: G. Kijhler
8Z.L
8..-
&a
3
Hemiphractidae- - - +
-
, -
W W -S S W --S P -
Fig. 467. He1111phractusfasciatus (Valle de Anton, Cocl6, Panama).
Photo: B. Wilson
Hemiphractidae Recently, GUAYA~AMIN et al. (2008) found that the species of the former families Amphignathodontidae, Cryptobatrachidae, and Hemiphractidae form a monophyletic group that they recognized as a single family, Hemiphractidae. This family is a predominantly South American group of frogs consisting of 93 species in five genera, with only three species in two genera occurring in Lower Central America (FROST2010). Most species in this family have direct development, with eggs brooded by the female in a dermal pouch (e.g., i n Gastrotheca) or attached to the dorsal skin (e.g., in Hemiphractus) (DUELLMAN 1970, MENDELSON et al. 2000). In all hemiphractids, eggs are carried on the female's dorsum for some or all of development; eggs hatch directly into froglets or into tadpoles that are deposited in ponds and other bod-
& ~MANEss 1980, ies of water (DUELLMAN WASSERSUG & DUELLMAN 1984,WIENSet al. 2007).
f i y to the species of Hemiphractidae 1 a A fleshy proboscis; head triangular in dorsal view; no brood pouch in females
.............................. Hemi-
fzleciatue
b NOfleshy proboscis; head not triangular in
dorsal view; a brood pouch present in females ...................................................... 2 flap on upper eyelid; a A triangular flank and posterior surfaces of thigh not darker than dorsum Gestrotheca m u t a b No t f i m d a r dermal flap on upper eyelid; flank and posterior surfaces of thigh darker than dorsum ......... GWmtheca nioefhn'
-er
Re*
TRUEB1974, DUELLMAN & ~ E S 1980, S WAS~ERSUG & DUELLMAN 1984, m ~ ~ et s al. 2000, GUAPASAMIN et al. 2008
o
c Gastrotheca
I
The 58 species of Gastrotheca are widespread throughout much of South America with only two species occurring in Lower Central America (FROST 2010). Gastrotheca cornuta is a nocturnal canopy dweller in tropical rainforest and low montane forest characterized by high humidity throughout the year (DUELLMAN 1970, SAVAGE 2002). The males call from high in I the trees, with the call having been 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). Photo: E-H. Jungfer dorsal skin; the embryos are partly or completely encased in large, thin bell-shaped
m
~
I
I
-
gills providing a surface for gas exchange
(PINO1980, 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, WIENSet al. 2007).
Fig. 468. Gastrotheca cornuta (female with eggs; Valle de Antdn, Cocl6, Panama). Photo: B. Wilson
'
I
Fig. 469. Gastrotheca nicefori (Ridge between Rio Jaque and Rfo Imamado, Daribn, Panama). Photo: C. W. Myers (Co-
of the Biodiversity Institute,University of Kansas)
Fig. 471. Gastrotheca cornuta (P.N. Chagres, Photo: M.Ponce Col6n, Panama, 790 m).
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1
Hemiphractidae 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 (1983). G. cornuta according to DTPLLMAN Gastrotheca nicefon' GAIGE1933, 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
DUELLMAN1980, DUELLMAN & MAN!XSS 1980, DUELLMAN 1983,WASSERSUG & DUELLMAN 1984, DUELLMAN 1989, WIENSet al. 2007
Fig. 472. Hemphractus fasciatus (Nusagandi, Kuna Yala, 320 m)Photo: K.-H. Jungfer
Hemiphractus Gastrotheca comuta Gastrotheca nicefori
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1
Herniphractus fasciatus
i
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 (MYERS1966). 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 (WIENSet 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 t o , 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, TRUEB1974, MENDELSONet d. 2000, SHEIL& MENDELSON 2001, SHEILet 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 & WILSON2002, SAVAGE 2002
Phyllomedusinae The subfamily Phyllomedusinae (learfrogs) 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 (DUELLMAN1970, 2001). The I Central American phyllomedusines are Fig. 474. Agalychnis callidryas (Soltis Center, placed into three genera: A g a 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 with vertical black bars; iris gray in life; occurs at moderate elevations, the Central SVL of adult males 50-81 mm, of adult American phyllomedusines are primarily a females 60-87 mm .. ~ o 4 y . cal& denizens of the lowland rainforest (SAVAGE b Lower eyelid reticulate (Fig. 474); heel without flap-like calcar; flanks blue, pur2002). During daytime, the Central ple, or brown in life with cream or white American species hide in trees, often high vertical bars; iris red in life; SVL of adult up in the forest's canopy (KUBICKI2004b). males 39-59 mm, of adult females 51-71 The breeding takes place primarily during mm ........................... the rainy season, and the females deposit 5 a Fingers with reduced webbing, webbing their eggs on leaves, vines, roots, bark, or between Fingers 111 and IV not reaching to penultimate subarticular tubercle of wood overhanging water; on hatching, tadFinger 111 (Fig. 477e); flanks blue or purpoles fall or are washed by rain into the plish blue; dorsum usually with wavy water body below (SAVAGE 2002, KUBICKI transverse lines; SVL of adult males 34-54 2004b). Phyllomedusine tadpoles, except mm, of adult females 52-66 mm ................ those of C. calcarifer, are often observed ..................................... saltator orienting themselves with the head up b Fingers with substantial webbing, webbing between Fingers 111 and IV reaching to near the water surface, moving just the tip penultimate subarticular tubercle of of the tail in a rapid motion (SAVAGE 2002, Finger I11 (Fig. 477a,c,d); flank coloration KUBICKI 2004b). variable blue; dorsum without wavy trans-
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Key to the species of phyllomedusine frogs (genera A g a l y c h i ~ , Cciuziohyla, and l?hyflomedwsa)
1 a Webbing between fingers and toes, if present, 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 flaplike calcar; flanks yellow or orange in life
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 j m h k 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 I
I
II
Fig. 475. Agalychnis callidwas (Volch Mombacho, Granada, Nicaragua, 1000 m). Photo: G. Kohler
Fig. 476. Agalychnis moreletii (El Refugio, Ahuachaph, El Salvador, 225 m). Photo: G. Kohler
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
SAVAGE 2002). Males have been reported to gather in breeding aggregations on vegetation above the breeding site (SCOTT& STARRETT 1974, ROBERTS1994, 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
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 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. calLidryas 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). Photo: A. Ramirez-Velazquez
Fig. 479. Agalychnis moreletii (El Occ-, Chiapas, Mexico). Photo: G. Kohler
Hylidae
Fig. 480.Agalychnts a m a e (Plan de Birri, Heredia, Costa Rica, 1630 m). Photo: G.Kohler
Fig. 482.Agalychnis annae (La Nevera, Ngobe Photo: G. Kohler BuglB, Panama).
Fig. 481.Agalychnis callidryas (KilambB, Jinotega, Nicaragua, 1015 m). Photo: G.Kohler
Fig. 483.Agalychnis callidryas (Peninsula 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 (PIBURN1963,1964, 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 (KUBICKI2004b). 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, KUBICKI2004b). Parachuting ability has been reported for A. sdtator and A. ~purrelli (SCOTT& Fig. 484. Agalychnis spurreLLi (Bani, PuntareSTARRETT 1974, ROBERTS 1994). Photo: G. Kohler nas, Costa Rica).
--=
Fig. 486. Agalychnis lemur (Guayach, CRARC, Costa Rica, 550 m). Photo: G. Kiihler
I Fig. 485. Agalychnis spurrelli (Guayach, 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. Agalychnis callidryas (COPE 1862a), Proc. Acad. Nat. Sci. Philadelphia 14: 359; type locality: "Panama." Males 56 mm, femalesto 71 mm. On the Atlantic versant from southern 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 COPE1885 is regarded as a synonym of A. cdidryas according to SAVAGE & HEYER (1967).
I ~ i487. ~ ~ .g a l ~moxletii ~ f i ~( ~~ ~i l ~~ photo: G. ~m~~ Chiapas, Mexico, 1050 m).
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: 'VeraPaz", 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
d~
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 T i l a r h [= 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).
Fig. 489. Agalychnis spurrelli (near Golfito, Puntarenas, Costa Rica, 50 m). Photo: B. Kubicki
.
A. moreletii
.
A. spumlli A. annae
Agalychnis
I
Further Reading
PWURN1963,1964, SAVAGE & HEYER1967, PYBURN 1970, SCOTT & STARRETT 1974, PROY 1992, PROY 1993a, JUNGFER & WEYGOLDT 1994, ROBERTS 1994, D'ORGEM& TURNER 1995, GRAY& RAND 1997, PROY2000, MCCRANIE et al. 2003, HOFFMANN 2005a
Fig. 490. Cruziohyla calcarifer (Guayacan, CRARC, Costa Rica, 550 m).
w
Photo: G.Kahler
utilize permanent or temporary ponds for breeding; rather, the females deposit the eggs on substrate (leaves, aerial roots, The genus Cruziohyla was introduced by wood, or bark) over water-filled cavities of FAIVOVICH et al. (2005) for two species of living trees, depressions in the tree trunk leaf-frogs, one of which, C. calcarifer, or its buttresses or similar habitats on falloccurs in Central America, whereas C. en trunks (MARQUIS et al. 1986, ROBERTS craspedopus is restricted to the Amazonian 1995, KUBICKI 2004b). The eggs hatch withlowlands in South America. Cruziohyla in five to ten days and on hatching, the tadcalcarifer is a large nocturnal treefrog that poles fall or are washed by rain into the 2002). The tadpoles inhabits the rainforest canopy and only water below (SAVAGE descends to find suitable breeding sites feed on organic material such as algae and during the rainy season (SAVAGE 2002). decaying vegetable matter in their puddle; Males call from trees surrounding or one to due to the low concentration of organic seven meters above the breeding sites food in this environment, it can take up to (KUBICKI 2004b). The call of C. calcarifer a year or more before metamorphosis takes has been described as a soft single note, place (KUBICKI 2004b). Under laboratory "rurm" or "whuunk"; several minutes to conditions, tadpoles metamorphosed after hours can pass between the calls (SAVAGE a minimum of 211 days (DONNELLY et al. 2002, KUBICK~ 2004b).This species does not 1987).
Cruziohyla
Hylidae Cruzioh~lacalcarifer (BOULENGER 19021, Ann. Mag. Nat. Hist. Ser. 7,9: 52; type locality: "Rio Duraneo. 350 feet". Provincia Esmeraldas. ~ c u a d & . . ~ a l etos 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.
1
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
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Phyllomedusa ---. -
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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 (RENJIFO& LUNDBERG 1999). The five type specimens of this species were found on vegetation in a swamp forest "where the species probably breeds" (DUELLMAN 1970:
~ ~
i494. ~ p.yllomedusa . venusta (Montes de bsucre,~ colombia). , Photo: A. Acosta
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 D a r i k Province, Panama, a s well a s 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 (Canamain camp, P.N.Darien, D d n , Panama, 500 m). Photo: A. J. Crawford
Further Reading
DUELLMAN & TRUEB1967, 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 (SAVAGE2002). Fig. 496. Snout profile: a. Trprion petasatus; Photos: G. Kohler Parental care is uncommon but occurs in b. Scinax elaeochroa. gladiator frogs (genus Hypsiboas) and two species of Smilisca (KLUGE1981, 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 1head 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 Fig. 497. Anotheca spinosa (Rancho Naturalists, blotches outlined with white ..... Anotheca b No bony occipital, canthal, or frontal near Platanillo, Costa Rica, 900 m). Photos: G. Kijhler 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. 498. Osteopilus septentrionalis (Florida, USA). Photos: T. Leenders
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Fig. 500. Trachycephalus venulosus (Scotland Photo: G. Kohler Halfboon, Belize, 10 m).
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. 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, CRARC, Costa Rica, 550 m); note mental gland brow). Photo: G. Kohler
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Fig. 503. Ventral surfaces of Isthmohyla insolita. Photo: J. R. McCranie
Fig. 504. Dendropsophus subocularis (Rionegro, Santander, Colombia, 831 m). Photo: C. Hernhdez-Jaimes
rig. sus. ~oloratlonor thigh in Isthmohyla lancasteri (Guayach, Vueltade Queque, Costa Rica). Photo: G. Kohler
transverse, not extending posterior to choanae; inner tarsal fold present or absent; toe webbing variable; dorsal coloration variable ...................................... 11 ........................................cllaraela Chin and throat boldly marked with a conlla b Head conspicuously large and deep (Fig. spicuous pale (yellow to white in life, dirty 501b); discs moderately large, width of disc white in preservative) central area, flanked on Finger I11 only slightly larger than the by solid, dark pigment (Fig. 503); dorsum length of tympanum; vocal sac absent or, if with numerous wart-like tubercles; SVL of present, single, median, subgular ............... adult males 30-36 mm, of adult females ...................................................Plecbhyla 34-38 mm bthmohyh (in park 1 insolit.) 10a Bones pale green in life; a circular whitish b Chin and throat not as described above; mental gland (Fig. 502) in adult males and dorsum with or without numerous wartsome adult females; vomerine tooth patchlike tubercles; SVL of adult variable .... 12 es acute, angulate, extending posterior to choanae; a low inner tarsal fold present; toe 12a Posterior surface of thigh contrastingly webbing extensive, modal formula I11 1- 2 barred black and yellow in life (black and IV or I11 1+ - 2 IV; dorsum usually pale or white or cream in preservative) (Fig. 505); olive green, sometimes pale brown (yellowone or more dark blotches in groin, SVL of ish white in preservative) ....... I.ipl08cirt~~ adult males 27-34 mm, of adult females 31-38 mm .................................................... b Bones white in life; no circular whitish .............bthmohyla (in park I. lancasteri? mental gland; vomerine tooth patches
9 a Head not conspicuously large and deep; discs very large, width of disc on Finger I11 nearly twice the length of tympanum ........
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 (inpart: 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.ldyla (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. Trachycephalus venulosus (Guaquira,Venezuela); c. Scinax staufEeri (San Juan de Dios, Nicaragua); 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
24a
posterior surface of thigh not as described above; SVL of adult variable ................ 24 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- 11/2 ....-la (in part. P.d e n t h u m ) Combination of characters not as above, particulary, if large frogs, then with less finger and toe webbing .......................... 25 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 ............................ 26 Adult males with a brown non-spinous pad or a patch of very tiny spinules on base of thumb (Fig. 507b,c); no prominent ventrolateral gland in adult males ................. 27 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 ....
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 b posterior margin of choanae ....................... ..................lethmohyla (in part: I. de-1 b A continuous pale lateral stripe from head %a to groin or from axilla to groin present; vomerine tooth patches positioned between choanae ...................................................22 2% No suborbital pale spot, labial stripe, ifpreb sent, not expanded below eye and terminating anterior to tympanum ..................... ................... la (in part. P. 1egk13 b A suborbital pale spot present or pale labi- 26a al 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 ............ lethm06gla (in park I. melacaena) genw except D. soralia) b Adult males with a patch of more than 30 23a An axillary membrane present, extending black spines on base of thumb; ventrolateral gland usually well-developed in adult at least one-fourth distance towards elbow; males; modal toe webbing formula 111 1+iris red in life; dorsal surfaces of head and 2 IV to I11 1 - 2- W,SVL of adult males body with a lichenous pattern of some shade of green in life (a pale lichenous pat30-39 mm, of adult females 35-46 mm ptscaohyla (in part: P. &pomy&my P. tern on a dark brown to dark gray backmacrotpmpanum, P.spinipoUe~) ground in preservative); posterior surface of thigh lemon yellow with dusting of 27a Small frogs, SVL of adult males 21-32 brown pigment in life (cream with brown mm, of adult females 24-35 mm; overall flecking in preservative); SVL of adult coloration tan, yellowish brown to yellowmales 28-32 mm, of adult females 35-38 ish green, often with accumulation of dark mm ....Duellmanohyla (in part: D. saralia) pigment in loreal region; finger webs vestib Axillary membrane absent, or if present, gial; toe webbing reduced (modal toe webthen iris not red in life and color pattern of bing formula I11 1 213 - 3 IV to I11 2 - 3 IV); axillary membrane absent; vomerine dorsal surfaces of head and body and of
nn
a.*.......................................
I
r 1
B2
b -
28a
b
29a
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:Izelkdd, I.picadrn3 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 webbing and tympanum distinct .................28 A well-developed a x i h r y 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 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 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 ~ mD. , phlebudes)
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 indicated 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'cta, 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
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
with pale and dark mottling, never with a series of discrete pale spots; venter without extensive dark pigment; finger webbing 33 variable ................................................... 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 axillary membrane (arrow). Photo: G. Kohler
Fig. 509. Smilisca baudinii (both from Mazatenango, Guatemala, 360 m). Photos: G. Kohler
............ (in park I.pmctip, I.xanthosticta) b Posterior surface of thigh patternless or
b No white or cream stripe from eye to groin; 1 l/4 -2 213 W,SVL of adult males 2 9 3 4 axillary membrane, if present, extending mm, of adult females 33-37 mm ................ no more than one-fourth of upper arm; ................ 3kthuo4pla (in part: I. r i v 7 l k b ) snout nearly rounded in dorsal aspect; canVenter immaculate or with Erie brown stipthus rostralis rounded to nearly angular; pling; snout shape variable; tympanum disSVL of adults variable ...........................39 tinct or hidden, length variable; modal toe webbing formula variable; SVL of adult 39a Tarsal fold distinct; subarticular tubercles under Fingers 111 and IV single, rounded; variable ................................................... 34 modal toe webbing formula 1111- 2' IV to Finger webbing vestigial or absent; white III 1 l/4 - 2 l/4IV; adult males with paired, stripes present on upper lip, forearm, and internal, subgular vocal sacs; groin and tarsus; dorsum with two elongate blotches posterior thigh reticulated with dark extending fmm head to sacrum, usually brown surrounding cream ar pale blue anastomosing at one or several points flecks; SVL of adult males 31-54 mm, of along midline; transverse supracloacal adult females 44-64 mrn ............................. white stripe present; posterior surface of .......... SiniIieca (inpark 5. rdla, s. EmIdida) thigh in lilfe and in preservative dark b Tarsal fold weak or absent; subarticular brown; SVL of adult males 32-38 mm,of tubercles under Fingers Ill or IV often douadult females 40-46 mm ............................. ble or b z d ; modal toe webbing formula ......................... S$lilieca (in part: S.puma) variable; adult males with single, median Combination of characters not as above, or bilobate, internal, subgular vocal sacs; particulary, if webbing vestigial, then dorcoloration of groin and posterior thigh not sal coloration not as above .....................36 as described above; SVL of adult males Small species, SVL of adult males 20-30 2 7 4 5 mm, of adult females 33-52 mm 40 mm, of adult females 18-36 mm, a mdi40a Dorsum lime green with scattered pale mentary or small (extending along upper green dots in life; flanks with a narrow arm one-fourth to one-half distance toward zone of reticulatian where dorsal and venelbow) axillary membrane usually present tral coloration meet; adult males with a ..................................................................36 patch of tiny black spines on base of Larger species, SVL of adult males >30 thumb; SVL of adult males 29-33 mm, of mm, of adult females >36 nm, or, if adult adult females 41-51 mm ............................. males <31 mm and adult females <37 mm, .......ptscho4rh (in part: P.sauc&mcmCie) then without a trace of an axillary memb Dorsum pale tan, dull brown, or reddish brane .......................................................38 brown with or without scattered dark Toe webbing reduced, modal formula 111 1+ brown blotches in life; flank coloration not - 3' IV; SVL of adult males 26-30 mm, of as described above; adult males with a adult females 27-36 mm .....Brnmelloil~h brown non-spinom pad on base of thumb; Toes extensively webbed, modal formula I11 SVL of adults variable ........................... 41 1 - 2 l V t o I I I 1 - 2 l/2IV;SVLofadults 41a Larger species, SVL of adult males 37-45 variable ....................................................37 mm, of adult females 41-52 mm ................ .......................................... Snout nearly rounded in dorsal aspect, can...........(inpart: I. Mizcata, I, peeudamma) thus rostralis rounded to nearly angular; SVL of adult males 27-35 mm, of adult b Smaller species, SVL of adult males 27-35 femdes 34-42 mm .................................... mm, of adult females 3 3 4 2 mm ...........42 ....... P@&*a (inpart: P.salved-) 42a Dorsum and upper surfaces of limbs granSnout acuminate or pointed in dorsal view ulate to tuberculate; modal toe webbing (Fig. 544-551); canthus rostralis usually formula 111 1-2 IV ....................................... angular; SVL of adult males 2 0 3 3 mm, of ........................ lidlrn*. (inpark I. ti=) adult females 18-36 mm ............................. b Dorsum and upper surfaces of limbs ................E.arodanta lin park E. bhcata, smooth; modal toe webbing formula I11 1 E. catracha, E. c h i m d q q E. mmidmdal l/2 -2 IV ....................................................... A discontinuous, irregular, white or cream ........&mhbyIa (inpark P. e a l v y i d d ) 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 (inpark E.p a r ! 3
..........am
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, strikingly 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 & FOSTER1996, CAMPBELL 1998, K O ~ R2001, K ~ E R et al. 2006). Anotheca spinos'a is a diflicult to find, nocturnal, arboreal frog (TAYLOR1954, DUELLMAN1970, SESSIONS19783. 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
1996). The bromeliad leaf axils (JUNGFER call was described as a loud "boop-boopboop* (DUELLMAN1970, JUNGFER 1996). This species uses bromeliads, wate~fflled tree cavities, and water-filled bamboo internodes for breeding (TAYLOR 1954, ROBINSON1961, DUEW 1970, J U N G ~ R 1996). Afker egg deposition, the male disappears, 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 & CALDWEZL2009). 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). Bromeliohyla bromeliacia is a small nocturnal cloud forest frog. that lives in arboreal tank I bromeliadi (MCCRANIE& WILSON2002). Males call from arboreal bromeliads or infrequently from vegetation such as on Fig. 513.Bromeliohyla bromeliacia (Quebrada 1300 m). C0ph7 top of leaves of branches outside of Photo: G. Kiihler bromeliads; the call consists of five or six soft notes repeated at intervals of 45-70 seconds (DUELLMAN -1970, MCCRANIE& WILSON2002). The female deposits the eggs into water-filled arboreal bromeliads and the tadpoles complete their development within the bromeliads (STUART 1943b, DUELLMAN 1970, MCCRANIE&. WILSON2002). In Honduras, eggs and tadpoles were found in bromeliads from May through August (MCCRANIE& WILSON 2002).
I
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 & WILSON2002
Fig. 515. Charadrahyla chaneque (extreme northwestern Chiapas on Hwy 190, Mexico, Photo: L. Gray ca. 700 m).
-
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-breedingfrogs 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: . 2*. -
I
F
Further Reading
DUELLMAN 1961c, DUELLMAN1965,1970, PEREZHIGAREDA 1981, MENDELSON & CAMPBELL1999, 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 attached to vegetation near the water surface (DUELLMAN 2001, SAVAGE 2002). The tadpoles of the Central American ~ e n d r o ~ s o ~have h u sbeaks but lack denticles (SAVAGE 2002). In D. ebraccatus, tadpole development
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 (FROST2010). The five Central American species of Dendropsophus are moderately to very small treefrogs that are widespread and common in the humid lowDendropsophus ebraccatus (COPE18741, Proc. lands. Throughout the rainy season, the Acad. Nat. Sci. Philadelphia 26: 69; type localmales can be heard calling from tall grass, ity: "region of Nicaragua." DUNN & STUART (1951:58) noted that the type specimen is reeds, or sedges at the edge of water, or labeled "Machuca", and that it surely came while perched on emergent vegetation; from "Nicaragua along the San Juan River." they produce insect-like calls, those of D. Males to 27.8 mm, females to 36.5 mm. ebraccatus sounding like a prolonged Atlantic versant of southern Mexico across northern Guatemala and Belize to Colombia "wreeeek" or "creeeek", those of D. microand northwestern Ecuador, sea level to 1600 m cephalus, D. phlebodes, D. robertmertensi elevation. like "creek-eek-eek-eek", and those of D. subocularis like a short trill (DUELLMAN Dendropsophus rnicrocephalus (COPE18861, Proc. Am. Philos. Soc. 23: 281; type locality: 1970, SAVAGE 2002). Calling usually begins "along a mountain stream in the department at dusk and continues until near dawn; of Chiriqui", Panama; at the time of the males usually sing in large choruses and description "Chiriquinincluded both Atlantic they will respond to conspecific males and and Pacific versants of extreme western Panama according to MYERS(1982). Males to males of other species within a vocalizing 27 mm,females to 32 mm.On the Atlantic vergroup (SCHWARTZ & WELLS1985, SAVAGE sant from southern Veracruz and northern 2002). Preferred breeding areas are tempoOaxaca, Mexico, to Panama; also widely disrary ponds, roadside ditches, flooded areas, tributed in South America (Colombia, Veneand marshes (DUELLMAN & FOUQUETTE
Fig. 516. Dendropsophus ebraccatus (male; La Selva, Heredia, Costa Rica, 70 m). Photo: G. Kohler
Fig. 517. Dendropsophus ebraccatus Itemale; Bartola, Rio San Juan, Nicaragua, 30 m). 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). 19061, Dendropsophus phlebodes (STEJNEGER 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. Fig. 520. Dendro~Oflus micmoeMus (Hopby 1934),Am. Dendropsophus subocularis (DUNN 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. 518. Dendropsophus robertmertensi (road from Hacienda San Cayetano to San Jose El Idolo, Suchitepbquez, Guatemala, 100 m). Photo: G. Ktihler
Fig. 521. Dendropsophus phlebodes (Turrialba, Cartago, Costa Rica, 600 m). Photo: G. Kohler
I Fig. 522. Dendropsophus subocularis (Rionegro, Santander, Colombia, 831 m). 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 hourmark; SVL of males Fig. 523. Eggs of Dendropsophus ebraccatus 23-27 rnm, of adult females 30-35 mm ..... ......................... Dendrupsophus ebraocatm CRARC*Cartago* Rita* 550 m). Photo: G. Kijhler b Head narrow; narrow transverse dashes or longitudinal dark line on dorsal surface of shank, dorsum patternless, with brown dashes, longitudinal dark lines, or reticulaD. phlebodes tions; SVL of adult males 18-25 mm, of D. subocularis adult females 23-31 m m .........................4 (Gua~acbp
A
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 ........ Dendrupsophurrpldebodes 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
n
-
-1 ID. microcephalus
D. robertmertensi
l?urther Reading Dendropsophus
-
DUELLMAN & F O U Q U1968, E ~ 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).
Fig. 524. Duellmanohyla chamulae (6.2 k m S
Ray6n Mescalapa, Chiapas, Mexico). Photo: W. E. Duellman (Courtesy of the Biodiversity Institute, University of Kansas)
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 1970, and terrestrial aroids (DUELLMAN 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 ("boop525. Duellmanohyla lythrodes (Rio Chanboop-boop-boop"), that of D. d o c u l i s as a Fig. guinola near Quebrada Guabo, Bocas del Toro, short rattle or "scraape" consisting of three Panama). Photo: C. W. Myers or four short notes, that of D. lythrodes as (Courtesyof the Biodiversity Institute, University of Kansas) a weak bell-like "pings' in bursts of five to seven notes, that bf 5.soralia as a single Duel2manohyla chamulae (DuEuMAN 1961b1, low-pitched "peep", repeated every 20-30 a t .13: 364; seconds, and that of D. chamulae and D. Univ, Kansas Publ. Mns. Nat. E type localiQ: 'a stream above (6.2 kilometers schmidtonun as a series of short, raucous by r o d 8 o d ) Ray6n Mescalapa, Chiapas,; low-pitched notes (DUELLMAN1970, Mgxico (1690 meters)." Males to 80.5 mm,, MCCRANIE& WILSON2002, SAVAGE 2002). females to 31.8 mm. North- sIopes of tha. central h i g h k h of Chiapas, Mexico, from' The tadpoles live in pools of the stream Jitotol to Soluschiapa,350-1700 m elevation. where they adhere to pebbles, stones, and Duellmanohyla fgthrodes (SAVAGB 1968b),! rocks in the relatively quiet water; when Bd. S. CaliforniaAcad. Sci. 67: 1;type locali~! disturbed, they seek refuge amidst debris ty:"Cash Rica: Pmvhcia de Lim6n; Cant6n dm \ on the bottom of th6 pools (DUELLMAN Lim6n: Alta Talamanca: 21 km SW Amubri a 6 1970). Although these species are apparconfluence of Rio Lari and Rio Dipari, 800 m"; elevation is actually 440 m according to ently active throughout the year, breeding SAVAGE (1974). Males to 32.6 mrn, females activity seems to peak in the early to midunknown. Atlantic versant in southern Costa dle rainy season (May to July); some Rica and adjacent northwestern Panama, species such as D. uranochroa are reported 170440 m elevation. to have a second reproductive peak in the Duehanohyla mfioctd.is (TAYLOR 1962b), late rainy season (DUELLMAN1970, Univ. Kansas Sci. Bull. 35: 827; type locality: MCCRANIE & WILSON2002, SAVAGE 2002).
"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 & WILSON19861, 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 de Nombre de Dios, Departamento de Atlhtida, Honduras." Males to 28.0 mm, females to 34.3 mm. Cordillera de Nombre de 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. ~ ~ ~ lsoda l (wlu0~ ~ ~ & ~M ~ ~ h 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.
Fig. 526. Duellmanohyla rufioculis ( Center,Alajuela, Costa Rica, 460 m). Photo: G. Kohler
Fig. 527. Duellmanohyla rufioculis (Rancho Naturalista, G.~K6hler ~c l ~~ Costa ~ Rica, 900 ~ m).Photo: ~
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,
b 2 a
b
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 & Dorsal color pattern not as above; a white lateral stripe usually present; adult SVL variable .....................................................2 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 White labial stripe expanded below eye; undersurfaces of foot heavily suffused with dark pigment; dorsal coloration variable; an inner tarsal fold present or absent; rela3 tive eye to nostril distance variable Distance from eye to nostril equals eye length; an inner tarsal fold present; dorsum leaf green in life (bluish to purplish in preservative) .... D u e l t n a n w f a salvavida 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 No ventrolateral glands in males; a brown nuptial pad on base of thumb in males; relative size of tympanum variable ............. 6 Ventrolateral glands present in males; no nuptial pad on base of thumb in males; tympanum length >50%of eye length .... 6 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 ......................
Fig. 529. Duellmanohyla salvadda (Liberacibn, Atlhtida, Honduras, 1050 m). Photo: J.H. Townsend
........
3a
b
4a b 6 a
............................ ouellman*k
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 )
doculiB
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 .... Fig. 531. Duellmanohyla uranochroa (Palo ........................... D u e l l m a n w l a chamulae Seco, Ngobe Bugl6, Panama, 1060 m). b Dorsum brown in life (and in preservative); Photo: A. Hertz venter white in life; no mental gland in males ...........-la ~chmidtmum
F'urther Reading
DUELLMAN1961b, SAVAGE1968b, DUELLMAN 1970, WILSON & MCCRANIE1985, 1986, MCCRANIE & WILSON1990, 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).It was hypothesized that the thick, rugose skin and the co-ossification 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 (SAVAGE2002). 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
WILSON2002, SAVAGE 2002). However, in captivity it was observed that the attendant 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 1970). FAIVOVICH et al. pools (DUELLMAN (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. m i o t p p a num on the other, certainly supports this view.
I
Fig. 533. Ecnomiohyla miotympanum (Hidalgo, Photo: G. Kohler Mexico).
Fig. 534. Ecnomiohyla milida
(Reserva
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 (COPE1886), 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, (2001: 898) suggested 630-1830 m. DUELLMAN 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 (SanJosB, 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. C o p b , Honduras." Males to 86.0 mm. females 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 & KUBICKI2010, 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 'lopes of Volch Tunialba), 400-1000 m elevation. 1966131, Ecnomiohyla thysanota (DUELLMAN 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.firmthe identity of this population, which probably belongs to another species.
Fig. 537. Ecnom'oh~lar a b b o m (Cemo Negro, Veraguas, Panama, 540 m). Photo: A. Hertz
Fig. 538. Ecnomiohyla s a l v e (P.N. Cusuco, Cortbs, Honduras, 1355 m). Photo: J. Kolby
E. miotympanum
rig. oad. Ecnomiohyla fimbrimembra (Bajo Mono, Chiriqui, Panama, 1820 m).
Photo: A. Hertz
prominent and scalloped; texture of dorsum variable; conditions of prepollex and nuptial excrecences variable; skin of dorsal surface of head in adults co-ossified with skull or not ................................................ 3 .......................E ~ o ~ & ' l a m o ~ P a n u m3 a Dorsum uniform green in life; dorsum smooth to minutely granular; skin of dorsal b Large, mostly brownish frogs, SVL in adult surface of head not co-ossified with skull; males 60-110 mm, in females 70-100 mm; heel without tubercles but with an acutely fingers at least two-thirds webbed; no white scalloped dermal fold .................................. stripe present on margin of upper lip .....2 ............................ Ecnomiohyla t h p m o t a 2 a Webbing between Fingers 11-IV not b Dorsum brown, reddish brown, or brown extending beyond penultimate subarticuwith green or darker brown markings or lar tubercle on Finger 111; supratympanic mottling; dorsum granular or tuberculate; fold continuing posteriorly behind tympaskin of dorsal surface of head in adults conum to terminate above axilla; dermal ossified with skull or not; condition of heel fringe along lateral edge of forearm and variable .....................................................4 tarsus narrow and not or only weakly scalloped; dorsum smooth to minutely granular 4 a Prepollical projection in adult males (tuberculate in juveniles); prepollex in recurved, terminating in a spine ..............6 terminus and scatmales with b Prepollex in adult males elongate, flat, tered black nuptial spines; skin of dorsal rounded, ending blunt, not terminating in surface of head in adults co-ossified with a spine ....................................................... 7 skull ..............Ecnomiohyla fhbrimembra 6 a Prepollex in adult males with scattered Webbing between Fingers II-IV extending small black spines; heel without tubercles; well beyond penultimate subarticular humera1 projection in adult males present; tubercle on Finger 111; supratympanic fold hands fully webbed (to base of disc on all not continuing posteriorly behind tympadigits) ........................Ecnomiohyla minera num towards axilla; dermal fringe along b Prepollex in adult males without scattered lateral edge of foreand tarsus usually
Key to E ~ n 0 ~ o h y l 8
1 a Small green frogs, SVL of adult males 27-38 mm, of adult females 37-51 mm; fingers about one-third webbed; distinct white stripe present on ~?argin of upper lip
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 coossified with skull; SVL of adult males 86-110 mm, of adult femalesto 86 mm
.................................. E
..a.
o o h
Fig. 541.Hand of a. Ecnom.Oh~lahbrimembra; 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 b Prepollex with scattered small black spines; heel without tubercles; humeral projection in adult males present ........... 8 8 a Dorsum smooth to granular; skin of d ~ r s a l Fig. 542. Hand of female Ecnomiohyla minera surface of head not co-assified with skull .. m~ Photo: Q Kohler A 24958). ...............................EuloziIiohyIa rabLnnum b Dorsum tuberculate, skin of dorsal surface of head co-ossified with skull ...................... ....................................Ecnozniohyla salvqje
I
I
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 Fig. 540. Ecnomiohyla sukia (from type locality). Photo: B. Kubicki
DUELLMAN 1970, SAVAGE 198lc, Wilson et d. 1985, HAYESet al. 1986, IBAREZ et al. 1991, DUELLMAN 2001, MCCRANIE& WILSON2002, SAVAGE 2002, MCCRANIE et al. 2003, MENDELSON et al. 2008
Exerodonta
+'-.
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.
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 Exerodonta catracha (PORRAS &WILSON 19871, forests and pine-oak forests where they Copeia 1987: 478; type locality: "Zacate Blanco breed in streams and ponds (DUELLMAN(14"20'N, 88"15W), a village located 14 k m WNW of La Esperanza, 2020 m, Sierra de 1970, MCCRANIE & WILSON 2002). During Opalaca, Departamento de IntibucB, the day, frogs of the genus Exerodonta Honduras." Males to 28.7 mm, females to 32.1 have been recorded hiding in arboreal tank mm. Sierra de Opalaca and Sierra de bromeliads and in axils of giant elephantMontecillos, Montafia de La Sierra, in southwestern Honduras, 1800-2160 m elevation. ear plants, some at considerable distance from the nearest streams of ponds, sugExerodonta chimalapa (MENDELSON& gesting these species might sometimes CAMPBELL 1994), Proc. Biol. Soc. Washington 107: 399; type locality: "Colonia Rodulfo migrate considerable distances to breed Figueroa, 19 k m NW Rizo de Oro (Chiapas), (CAMPBELL & BRODIE1992, MCCRANIE& 1542 m, Oaxaca, Mexico." Males to 24.9 mrn, WILSON 2002). During the breeding season females to 26.6 mm. Chimalapas region (June to August in E. catracha and E. (Sierra Atravesada in southeastern Oaxaca and adjacent Chiapas), Mexico, 850-1540 m sumichrasti?, males call at night from low elevation. vegetation and on the ground around slow moving streams and small ponds Exerodonta perkinsi (CAMPBELL & BRODIE 19921, J. Herpetol. 26: 187; type locality: "21.1 (STARRETT1960, DUELLMAN1970, km (by road) NNE Barillas, Finca Chiblac, MCCRANIE & WILSON 2002). Departamento de Huehuetenango, Guatemala, ... 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.
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 1879, Bull. notes ("haah-haah-haah") (DUELLMAN Exerodonta sumichrasti BROCCHI Soc. Philomath., Paris, Ser. 7,3: 20; type local1970, M c C m & WILSON 2002). Tadpoles ity: "Santa Efigenia pr&s de Tehuantepec", of E. catracha were collected from slow Oaxaca, Mexico. Males to 27.7 mm, females to moving streams and small ponds; after 33.0 mm. On the Pacific versant from central Guerrero and eastern central Oaxaca to completing metamorphosis, the froglets Chiapas, Mexico, 200-2000 m elevation. apparently move uphill from the streams and hide in bromeliads (MCCRANIE & WILSON 2002).
Fig. 544. Exerodonta bivocata (6.2 km S Ray6n Mescalapa, Chiapas, Mexico). Photo: W. E. Duellman (Courtesyof the Biodiversity Institute, University of Kansas)
I KeytoI5!zmdmte
Fig. 545.Exerodonta catracha (Zarciadero, Comayagua, Honduras, 1460 m). Photo: J. H. Townsend
Fig. 546.Exerodonta chimalapa (a. Guatemala; 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
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 onefourth 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 ......................E m d a n t a Bumichraeti b Inner tarsal fold present; dorsum usually with dark spots; snout nearly rounded in dorsal view; tympanum distinct or indistin& SVL of adult males 24.9-27.4 mm, of adult females 28.6-32.1 mm .......................
................................... Eremdanta c a m 3 a A broad continuous pale stripe along sides larger (ratio tympanum of body; tympm~~ll 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 parb No white or cream stripe &om eye to groin; axillary membrane extending about onefourth of upper arm; SVL of adult males 20-25 mm, of adult females 18-27 mm ................................. lbrodmta c?himsla-
Hyla
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 BiodiversityInstitute, University of Kansas)
Further Reading DUELLMAN & HOYT1961, PORRAS& WILSON1987, & BRODIE 1992,MCCRANIE et al. 1993a, CAMPBELL MENDELSON & CAMPBELL 1994
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 (FROST2010). 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 lowpitched 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, 13001536 m elevation. Hyla walkeri STUART1954, 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 a t 1725 meters." Males to 35.9 mm, females to 37.8 mm. Central Chiapas, Mexico, to the Sierra de 10s Cuchumatanes, Guatemala, 1450-2900 m elevation.
Hylidae
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 .....
Fig. 552. Hyla bocourti (3.8 km W PurulhB, Baja Verapaz, Guatemala). Photo: J. A. Campbell
...............................................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
(Coultesy of the B i o d i d t y Institute,University of I(ansas)
Fig. 555. Thigh pattern in a. Hyla bocourti; b. H. wdkeri. Photos: G. Kijhler
E'lg. 563. Hyla walkefi (18 Inn NW Gomitiin, Photo: W. E. Duellman Chiapas, Mexico). (Courtesyof the Biodiversity Institute,University of Kawad
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 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 is an old name proposed by PETERS (1882b) that was resurrected .from synonymy with Hyla by F ~ O V I Cet H al. (2005).As currently understood, the genus Hyloscirtus contains 30 species, all except two being distributed in South America Hyloscirtus palmen' (BOULENGER 19081, Ann. Mag. Nat. Hist. Ser. 8, 2: 515; type locality: (FROST 2010). The two species treated here "Jimenez", Department of Valle, Colombia. are unique among Central American Males to 45.0 mm, females to 50.0 mm.On the hylids by having a whitish or greenish-yel- Atlantic versant from central Costa Rica to low mental gland present in adult males central Panama, and on the Pacific versant and usually in adult females (SAVAGE from western Colombia to northwestern Ecuador, 100-920 m, in Colombia up to 1600 m 2002). Both H. colymba and H. paheri are elevation. nocturnal stream-breeding treefrogs that inhabit small to moderate-sized fast flowing watercourses with rocky beds in lowland and premontane rainforests (SAVAGE 2002, KOHLERunpubl. 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 1m Fig. 557. Hyloscirtus colymba (El Cop4 CoclB, Photo: B. Kubicki above the ground in the forest some hun- Panama, 750 m). dred meters away from the nearest stream (HERTZpers. comm. September 2010). Breeding is assumed to be restricted to the drier parts of the year (SAVAGE 2002).
Fig. 556. Hyloscirtus colymba (Alto de Piedra, Veraguas, Panama, 910 m). Photo: A. Hertz
Fig. 558. Hyloscirtus palmeri (Guayach, CRARC, Costa Rica, 550 m). Photo: G. Kohler
Hypsiboas
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 .............................
The genus Hypsiboas contains 82 species of moderately sized to very large arboreal nocturnal treefrogs (FROST 2010). Hypsiboas was removed from the synet al. (2005). onymy of Hyla by FAIVOVICH 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 (KLUGE1981).
Gladiator frogs have a special breeding system in which the males usually con.................................... Hylmcidus ml+ struct basin-like nests on the margins of b No yellow stripe extending along canthus ponds, roadside ditches, or slow-moving rostralis and margin of upper eyelid to level of axilla; no dermal calcar on heel; streams, the nests being fed with by seepSVL of adult males 36-45 mm, of adult age from these water bodies (KLUGE1981, females 36-50 mm ..... E&lmcidus palmme HOBEL2000). 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 footprints as substitutes for male-built nests (HOBEL1999b).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 (HOBEL2000). 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, M e r Reading DUELLMAN 1970, MYERS& DUELLMAN 1982, females seemed to inspect the nest of bArjEz et al. 199913, DUELLMAN 2001, SAVAGE males prior to amplexus, being rather 2002 selective, and at least in half the time they
I I !
I
1
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) (KLUGE1981). 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~iboasspecies 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, SAVAGE2002). 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 1970, areas in swamp forest (DUELLMAN SAVAGE 2002, K~HLER unpubl. obs.). The advertisement call consists of a series of clucks (DUE1970). The eggs are deposited as a surface film in shallow water, and the tadpoles develop in the shallow 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 absence of red on the digital webs (see Fig. 568).
Fig. 560. Hypsiboas boans (Laguna Purriche, Darien, Panama, 449 m). Photo: A. J. Crawford
Fig. 561. Hypsiboas crepitans (Ponuga, Photo: A. Hertz Veraguas, Panama, 35 m).
1i H e b o a s born (LINNAEUS17581, 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. (SCHMIDT 18571, Hypsiboas pugnax 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. 1898b1, Hypsiboas rosenbergi (BOULENGER Proc. Zool. Soc. London 1898: 123; type locality:"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. 19611, Hypsiboas rufitelus (FOUQUETTE 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 t o H. albomarginatus are provisionally 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
3a
b
4a
b
between Fingers I11 and IV reaching at least penultimate subarticular tubercle on Finger 111 (Fig. 563b); flank pattern variable; SVL of adults variable ....................3 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 Calcar absent; posterior surfaces of thighs with narrow vertical bars; flanks with distinct dark bars; palpebral membrane not reticulated; SVL of adults variable .........4 Webbing not extending beyond penultimate 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 Hqrpaibaeslpugnax 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 ........................... ~ b o t t s m e n b e l g i
Hylidae
Fig.568. &ad 4ql~enW.
d a. -?ha
arepiha%h. a. 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. Fig. 569. Isthmohyla angustilineata (Pianists angustilineata, I. lancasteri, and I. pseudo- Trail, Ngobe Bugl6, Panama, 1800 m). Photo: M. Ponce puma 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 (SAVAGE2002). 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 Fig. 570. Isthmohyla calypsa (north slope vegetation near or overhanging the Cerro Pando, Bocas del Toro, Panama, stream, as well as from rocks in or along 1920 m). Photo: W. E. Duellman watercourses during most of the year (Courtesy of the Biodiversity Institute, University of Kansas) (DUELLMAN 1970, LIPS 1996, SAVAGE 2002). The call of I. debilis and I. tica has been described as a series of cricket-like "chirpchirp-chirps", that of I. rivularis as a series I. pseudopuma) lay the eggs in the water in 2002). Females of high-pitched "cheep-cheep-cheeps", that single long strands (SAVAGE of I. pictipes as a single low-pitches, pulsed of I. pseudopuma appear to select breeding "eeck", and that of I. calypsa and I. lancas- sites lacking conspecific tadpoles ( C R ~ teri as a low-pitched, vibrating extended 1991). The tadpoles of Isthmohyla species note (SAVAGE1968b, DUELLMAN 1970). inhabit the quieter areas of the stream Amplexus is axillary (SAVAGE 2002). These such as slow-moving portions and backwa2002). In some frogs reproduce primarily in the drier ter pools (LIPS1996, SAVAGE months of the year when stream flow is species, such as I. rivularis, the tadpoles reduced or during the early part of the have very large oral discs to adhere to 2002). rainy season (SAVAGE 2002). Some Isthmo- large rocks in the stream (SAVAGE hyla species (e.g., I. calypsa) deposit the eggs in a mass on leaf surfaces of plants directly above the stream; on hatching, Isthmohyla angustilineata ( T A ~ O1952b), R tadpoles fall or are washed by rain into the Univ. Kansas Sci. Bull. 35: 850; type locality: stream below (WILSON et al. 1994, LIP^ 1996). "La Palma, [Provincia San JosB,] Costa Rica." Males to 34.2 mm, females to 36.8 mm. Isthmohyla insolita lays eggs in masses of Cordilleras de Tilaran, Central, and moss, which are brooded by the male Talamanca of Costa Rica and western (WILSONet al. 1994). Other Isthmohyla Panama, 1100-2300 m elevation. species (e.g., I. angustilineata, I. lancasteri,
isthmohyla caiypsa (LIPS1996), 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) eastern 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.
' Fig. 572. Isthmohyla debilis (Cerro Tugri, Ngobt Bugl6, Panama, 1570 m). Photo: A. Hert
1
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 detoTalamanca 38.7 mm, females and to 41.4 mm. de Fig. 573. Isthmohyla debilis (La Nevera, Ngobe Tabasan5 of Panama, 1120-1710 m elevation. Photo: A. Hertz Bugl6, Panama, 1643 m).
...
-
Fig. 571. Isthmohyla graceae (Fortuna, ChiriPhoto: IC-H. Jungfer quf, Panama, 950 m).
pig. 575. Isthmohyla graceae (Quebrada de Arena, Chiriqui, Panama). Photo: C. W. Myers (Courtesy of the 3iodiversit.v Institute,University of Kansas)
Fig. 574. Isthmohyla graceae (Cerro Sagui, Ngobe Bug14 Panama, 1710 m).Photo: A. Hertz
1 1
1
-
5
1
Fig. 576. Isthmohyla infucata (Bocas uGl L U L ~ , Panama). Photo: E. Boza
Fig. 579. Isthmohyla insoLita (2.5 airline km NNE of La Fortuna, Yoro, Honduras). Photo: J.R. McCranie
rig. u I I . r u u u u u u y x a ~ r u u w ~ a r\ rrw u vuuugrrru,
s.r e vuu.
Bocas del Toro, Panama, 1605 m).Photo: A. Hertz
CRARC,
~ca).
Photo: G. Kohler
Ishohyla infucata ( D m1968331,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
Fig.578.Lshnohyla iniizcata ( h m type l d t y ) . Photo:W. E. Duellman (Courtesyof the Biodivdty Wtuta, University of -8)
(15"26%, 87"18W), 1550 m elevation, Cordillera 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' (BARBOUR19281, ROC. New England Zool. Club 10: 31; type locality: "high forest a t Peralta, [Cant611 de Turrialba, Provincia Cartago,] Costa Rica, altitude, 500 meters." Males to 33.6 mm, females to 41.1
I
I
Fig. 581. Isthmohyla lancasteri (Fortuna, Chiriqui, Panama). Photo: G. Kohler
tago, Costa Rica, 1000 m).
rig. sax. lstnmonyla melacaena w.n.&I uusuco, Photo: J. R. McCranie CortBs, Honduras).
Fig. 584. Isthmohyla picadoi (Volcdn Barva, Heredia, Costa Rica, 2260 m) Photo: G. Kohler
mm. Cordillera de Talamanca of Costa Rica and western Panama, 90-1450 m elevation.
Photo: B. Kubicki
I
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." Males to 32.8 mm, females to 35.2 mm. Cordilleras Central and Talarnanca of Costa Rica and western Panama, 1920-2770 m elevation.
Fig. 585. Isthmohyla zeteki (La Nevera, Ngobe Photo: G. Kahler Bugl6,Panama)
Fig,586. Isthmohyla pictim (SE slope of Volch Bama, Heredia, Costa Rica, Photo: E-H. Jungfer ca. 2000 m).
jMhm~&la pl'c@xw (GQPE1816),J. A d .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 I c a , 1930-2800 m ekevation. lethmo3lpa pseudapuma (G1901). 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 ~ 1952b), R Univ. Ransas Sci. B d . 35: 847; type localiQ: "American Cinohona l?lantati~nEIsla Bonita), eastern slope of V O W Potis, [Cant611 de Alajyla, Provincia de AlajuelaJ Costa Rka, elevabon 5,500 ft." I1209 m elevation]. W e m 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 4 a c e n t western Fbama, 1210-2840 m elevation.
Fig. 587. Isthmohyla pictipes (1km W Poasito, Alajuela, Costa Rica). Photo: W. E. Duellman (Courtesyof the Biodiversity Xnatitute, Universityof Kansa~)
Ikbhmahyls tica (STARRETT1966), 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 (DUE1968b), Univ. Kansas Publ. Mw. Nat. a t . 17: 662; type localits "south Fork of the Rio Las Vueltas on the south s l q e 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& (GAIGE19291, 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. 588. Isthmohyla pseudopuma (female; road fiom Copey to Providencia, San Jos6, Costa Photo: G. Kohler Rica, 2540 m).
Fig. 589. Isthrnohyla pseudopuma (male; road &om Copey to Providencia, San JosB, Costa Photo: G. Kohler Rica, 2230 m).
Fig. 592. Isthmohyla tica (Tapanti, Cartago, Photo: K.-H. Jungfer Costa Rica, 1150 m).
Fig. 590. Isthmohyla rivularis (Jurutungo, Chiriqui, Panama, 2070 m; variation in ventral coloration). Photo: G. Kahler
Fig. 593. Isthmohyla tica (Finca Bambito, El Volch, Chiriqui, Panama). Photo: W. E. Duellman (Courtesy of the Biodiversity Institute,University of Kansas)
Fig. 591. Isthmohyla rivularis (Jurutungo, Photo: A. Hertz Chiriquf, Panama, 2070 m).
Fig. 594. Isthmohyla xanthosticta (Rama Sur Rio Las Vueltas, Heredia, Costa Rica). Photo: W. E. Duellman (Courtesy of the Biodiversity Institute,University of Kansas)
Hylidae
I
.
I. pseudopuma I. infucata
.
I. lancasteri I. tica I. calypsa
! 1
Key to I d l m e l a
1 a A broad (greatest width varying from onehalf tympanum length to greater than tym-
b
2 a
b
3 a
panum 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 m m ......... Isthm06pla graaeae 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 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 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 Males with 7-10 large black spines on thumb; skin on dorsum smooth; posterior part of dorsum transIucent in life; ventral
b
4 a
b 6 a
b
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 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 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 Chin and throat not as described above; dorsum with or without numerous wartlike tubercles; SVL of adults variable ..... 5 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 heasten' 31-38 mm ............... Lthmohy.. 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 continutan, or brown in preservative); snout ous white or golden yellow lateral stripe or rounded or truncate in lateral view; texan elongate pale blotch on glandular ridge ture of dorsum variable; SVL of adults between midlateral region and groin, tymvariable ................................................. 12 panum small, its length about one-third l2a Snout short and truncate in profile; doreye length; SVL of adult males 25-30 mm, sum and upper surfaces of limbs usually of adult females 27-32 mm ......................... strongly tuberculate; modal toe webbing ....................................... Isthmohyla d e w formula I11 1-2 TV; no discrete yellow spots b No pale suborbital spot; no continuous pale or a single yellow area in groin, although lateral stripe or elongate pale blotch preflank usually mottled brown and yellow, or sent between midlateral region and groin; with irregular pale spots; no dark brown tympanum size variable; SVL of adults lateral band between axilla and groin in variable ..................................................... 7 breeding males; SVL of adult males 27-34 mm, of adult females 3 3 4 2 mm ............... 7 a Posterior surface of thigh with discrete ........................................... lethmohyh tica pale spots (yellow in life, white in preservative); venter usually dark ...................... 8 b Snout moderately long and acurninate in lateral view; dorsum and upper surfaces of b Posterior surface of thigh immaculate or limbs smooth to granular; modal toe webwith mottling, but never with discrete pale bing formula III1 112 -2 l/2 IV; groin ususpots; venter usually pale ........................ 9 ally dark brown with indiscrete yellow 8a A bronze stripe on canthus rostralis; large spots or a single yellow area encroaching yellow spots on groin and posterior thigh; on lateral dark band from below; breeding tympanum length about 50% eye length; males usually with a distinct dark brown single known adult female 29 mm SVL lateral band between axilla and groin; SVL .............................. lethmohyla xanthosticta of adult males 37-45 mm, of adult females b No stripe on canthus rostralis; moderate41-52 mm ......... I8thmohyla peendopuma sized yellow spots on groin and posterior 13a Venter usually peppered with dark spots thigh; tympanum length about 30% eye (Fig. 590); dorsum gold orange, yellowish length; SVL of adult males 31-39 mm, of tan, to olive brown in life; modal toe webadult females 40-45 mm ............................. bing formula I11 1114-2 2/3 IV (Fig. 595b); ..................................... Istlunohyla pPct!*p SVL of adult males 29-34 mm, of adult 9a Dorsal surfaces of head, body, and limbs females 33-37 mm .. Isthmohyla rivduie with large, tubercular spines; adult males b Venter imaculate; dorsum grayish brown with small black spines on thumb; SVL of or tan in life; modal toe webbing formula adult males 26-35 mm, of adult females I11 1 2/3 -3 IV to III 2-3 IV (Fig. 595~); 31-40 mm ..................... Isthmohyla calgpaa SVL of adults variable .......................... 14 b Dorsal surfaces of head, body, and limbs 14a Flanks and ventral surfaces of arms smooth, granular, or slightly tuberculate, smooth and not glandularly thickened; but without large, tubercular spines; nupeyes directed forward at about a 45" angle; tial pads of adult males without small tympanum distinct and directed dorsolatblack spines; SVL of adults variable ..... 10 erally; a single odontoid present at the tip 10a Toes with extensive webbing, modal formuof lower jaw; prepollical pad greatly la I11 1-2 IV or I11 1 l/2 -2 112 N (Fig. enlarged (Fig. 507g); SVL of adult males 595a); moderate webbing between Fingers 21-24 mm, of adult females 24-27 mm .... 111and IV, modal formula 1112 l/3 -2 IV 11 lethm*la zeta b Toes with moderate webbing, modal formub Flanks and ventral surfaces of arms disla I11 1 l/4 -2 2/3 IV to I11 2 -3 IV (Fin. tinctly granulate to glandularly thickened; 595b);webbing between Fingers I11 and fi eyes directed forward at less than a 45" vestigial, modal formula I11 2 2/13 -2+ IV 13 angle; tympanum indistinct or hidden and directed vertically; a pair of odontoids prel l a Groin with discrete yellow spots edged by sent at the tip of lower jaw; prepollical pad reddish brown; hidden surfaces of groin, moderately enlarged (Fig. 5079; SVL of anterior and posterior thigh red or maroon adult males 27-32 mm, of adult females in life (pinkish tan in preservative); snout 34-35 mm ................... Isthmohyla picadoi 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, ante- Further Reading rior and posterior thigh orange yellow, tan, CRUMP1984,1989b, 1990, CRW & TOWNSEND brown, or orange brown in life (dirty white, 1990, AROSEMANA & I B ~1991, Z CRW 1991
.....................................
Osteopilus (West Indian treefiogs) is an Antillean genus of eight moderate-sized to very large species (FROST2010); 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 Fig. 596. Osteopilus septentrionalis (Florida, Photo: T. Leenders ponds, roadside ditches, drainage ditches, USA). 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 2002). described as rasping snarl (SAVAGE 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 M-I (1950).
Fig. 597. Osteopilus septentrionalis (pet trade). Photo: P. Janzen
Further Rsading
MIT~LEMAN 1950, DUELLMAN & CROMBIE 1970, TRUEB& TPLER 1974, SCHWARZ & HENDERSON 2002 1991, MESHAKA2001, SAVAGE
Fig.598. Plectrohyla exquisita (P.N. Cusuco, CortBs, Honduras, 1833 m).
Photo: J. Kolby
mats; individuals of P. exquisita were sleeping on vegetation three to six meters above a stream in early afternoons The genus Plectrohyla has been recognized (MCCRANIE & WILSON2002). At night, for a distinctive group of cloud forest hylids Plectrohyla species have been found along from southeastern Mexico and northern cascading streams where they were clingCentral America since the early 1940s ing to rocks and large boulders in the (e.g., HARTWEG1941, DUELLMAN & splash zone of cascades or behind waterCAMPBELL 1992). Currently, with the inclu- falls, or perching on branches of bushes or sion of the species formerly assigned to the on tree trunks at the edge or above the Hyla bistincta group by FAIVOVICH et al. stream, often at places receiving mist from (2005), 42 species of Plectrohyla are con- cascades (DUELLMAN & CAMPBELL 1992, sidered valid, 18 of which occur in the area MCCRANIE & WILSON2002). I encountered treated by this book (FROST2010). The juveniles of an undetermined Plectrohyla Central American Plectrohyla species at Cerro Saslaya, Nicaragua, about 20 inhabit cloud forests and pine-oak forests meters away from the nearest stream. in the highlands (615-3500 m elevation) Plectrohyla species apparently have an where they remain along streams through- extended breeding season, possibly more out most, if not all, of the year (DUELWIAN or less continuous throughout the year & CAMPBELL 1992). Daytime retreats of (DUELLMAN & CAMPBELL 1992). Males of these nocturnal frogs include arboreal some species are known to vocalize; the tank bromeliads, rock crevices, and moss call of P. guatemalensis has been
r~ectron yla
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 & CMBELL 1992).
-a
Fig. 599. Plectrohyla exquisita (P.N. Cusuco, Photo: J. Kolby Cortks, Honduras, 1833 m).
described as a "single low-pitched note 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 knifelike spine, whereas in those species having a blunt prepollex or bifid prepollex, males and females are about equal in size (DUELLMAN & CAMPBELL1992). Although no combat has been reported in any Plectrohyla species, the presence of scars on bodies and forelimbs has been interpreted as evidence for rnale-rnale aggressions (DUELLMAN & CAMPBELL1992). 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 "inoculate" females with secretions of the male's mental gland (BROCK& VERRELL lgg4)McCFUN1~ et al. (IgB7)found a clutch of P. g~atemalensiswith 240 eggs that were adherent to the roots of a shrub on the bottom of a shallow stream.
Fig. 600. Plectrohyla dasypus (P.N. CUSUCO, Codes, Honduras, 1385 m). Photo: J. Kolby
*
Fig.601. Ampleding pair of Plectrohyla matudai urith eggs submergedin a stream (volch Tacanl, Chiapas, Mexico, 1750 m). Photo: A. Ramirez-Vellzquez
Hylidae Plectrohyla acanthodes DUELLMAN & CAMPBELL 1992, Misc. Publ. Mus. Zool. Univ. Michigan 181: 2; type locality: "a stream 6.2 k m (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.
Fig. 602. Plectrohyla acanthodes (San Crist6bal de las Casas, Chiapas, Mexico). Photo: l? Heirnes
603. Plectrohyla avia (Volck TacanB, Chiapas, Mexico, 1805 m). Photo: A. Ramirez-Velhquez
, .g.
Fig. 604. Plectrohyla chrysopleura (La Liberacibn, Atlktida, Honduras, 1030 m). Photo: J. H. Townsend
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-DfAZ1994, F'roc. Biol. SOC.Washington 107: 68; type locality: "along the Quebrada de O m (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 Cusuco 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). Photo: J. H. Townsend
rig. 605. rrectrohyla
Fig. 606. Plectrohyla dasypus (P.N. Cusuco, CortBs, Honduras, 1348 m). Photo: J. Kolby
Fig. 608. Plectrohyla guatemalensis (P.N. Montaiia de Yoro, Francisco MorazBn, Photo: J. H. Townsend Honduras, 1850 m).
dasypus (P.N. Cusuco, CortBs, Honduras, 2040 m). Photo: J. H. Townsend
Plectrohyla guatemalemi~BROCCHI187Ta, 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 eastcentral 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 Francisco (about 25 km. northeast of Nebaj),
Fig. 609. Plectrohyla hartwe@ (east slope of Cerro Celaque, Lempira, Honduras). Photo: J. R. McCranie
Fig. 610. Pkckrohyla id (Tapalapa, Chispass, Mexico, 1757 m). Photo: A. Ramirez-VelgZquez
Fig. 611. Plectdyla i d (San Jose Maxbal, B d a s , Huehuetenango, Guatemala, 1390 m). Photo: C. R. VAsquez Almazh
[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~aB ~ & 5 Mm 1854, Herpetolagica 110: % type locdity: "Etegi6nde ~ 0 c o n u 8 ~ ~ FMexico? t 8 8 ~ Males to 47.8 mm, females fa 48.1 mm. R n m only from h'0 l&bl B ~ CkpSkE P (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 f o m t 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 . P I ~ c h h p l ap 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 hdwae*d~ElPiW@*r~Of El Sdvadar d Hcadr2ras'I, 2M0-2580 m elevation.
P1-rL! pmn&a B a a 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 cfill. Mus. Zool, Wpix nsichigein U:1;@pelocality: "SCTeam in BarrL 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 A l h Vmpm and the Siema de,lss Abwl, 615-160 m elevation.
Fig. 612. Plectrohyla 1acertos.a (Chiapas, Mexico), Photo: A. Ramhe-Velhquez
-
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Fig. 613. Plectrohyla psilodema (holotype). Photo: G. Kohler
Fig. 616. Plectrohyla matudai (r'lnca lrlanaa, Chiapas, Mexico, 1050 m). Photo: G. Kohler
Fig. 614. Plectrohyla matudai (Volch Tacana, Chiapas, Mexico, 1750 m). Photo: A. Ramirez-Veldzquez
Fig. 617. Plectrohyla quecchi (Finca Los Alpes, Senahu, Alta Verapaz, Guatemala, 1000 m). Photo: C. R. Vasquez Almazdn
Fig. 615. Plectrohyla pokomchi (Eslope C e m Quisis, Baja Verapaz, Guatemala). Photo: J. A. Campbell
Fig. 618. Plectrohyla quecchi (3.8 km SE Purulhti, Baja Verapaz, Guatemala). Photo: J. A. Campbell
(Courtesy of the Biodiversity Institute, University of Ransasf
(Courtesyof the Biodiversity Institute, University of Kansas)
Hylidae
*
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,'lectrohyla sagonun H A X ~ 1941, G 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, southcentral Chiapas, Mexico, southwest to Guatemala and also in northern El Salvador, 1000-2400 m elevatiofi.
Fig. 619. Plectrohyla sagorum (VolcBn Tacana, Chiapas, Mexico, 1810 m). Photo: A. Radez-VelBzquez
,.y..% 4 ,":+' -?
'-,;:
& !'
. $.
Plectrohyla tecunumani DUELLMAN& CAMPBELL 1984, Copeia 1984: 391; type locality: "a cave 1k m 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 Cuchumat a m s in western Guatemala, 3200-3395 m elevation. Plectrohyla teuchestes DUELLMAN & CAMPBELL 1992, Misc. Publ. Mus. Zool. Univ. Michigan 181: 15; type locality: "Finca Los Alpes, ca. 1000 m, Departamento Alta Verapaz, Guatemala (15"22'N, 90°01'W)." Males to 67.1 mm, females to 73.6 mm.Known onlv from the tvpe locality.
Fig. 620. Plectrohyla s a g o m (Rgserva de la Biosfera El Triunfo, Chiapas, Mexico, Phatn: S. M. Rovito ca. 2000 m).
Fig. 622. Plectrohyla sp. (subadult; Saalaya, Atlantic0 Norte, Nicaragua). Photo: G. Kahler
Fig. 621. Plectrohyla teuchestes (Finca Los Alpes, Alta Verapaz, Guatemala). Photo: W. E. Duellman
Fig. 623. Plectrohyla exquisita (P.N. Cusuco, CortBs, Honduras). Photo: J. R. McCranie
(Courtesy of the Biodiversity Institute,University of Kansas)
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 6 8 4 1 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 ................
................................-lP
he&&
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 projecting 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, Photos: G. Kiihler f P. guatemalensis.
-1
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 pycnd 14a Dorsal surfaces of head and body uniformly green in lie; dorsum smooth; vocal slits 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&dai d
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 & CAMPBELL 1984, MCCRANIE 1981, DUELLMAN et ale 1987, DUELL- & CAMPBELL 1992, Fig. 626. Flank coloration in a. Plectrohyla WILSON. MCCRANIE& ~ ~ J z - D ~ 1994, Az sagorum; b. P. quecchi; c. P. S,d. P. matudai. Photos: G. Kohler M C C R &~ WILSON1998,1999a, 2002 -
Fig. 627. Ptychohyla legleri (Alfombra,Puntarenas, Costa Rica, 880 m).
Photo: M. Ryan
spinipollex have been described as "a rapidly repeated series of five or six short, high-pitched notes almost like a stuttered The genus Ptychohyla contains 13 species whistle" (MCCRANIE & WILSON1989: 15). of stream-breeding treefrogs that inhabit DUELLMAN (1970) described the call of P. moderate elevation forests (broad-leafed euthysanota, P. macrotympanum and P. and mixed pine, 100-2200 m above sea hypomykter as a single soft note level) from south-central Guerrero and ("wraaack"). A clutch of about 135 eggs, northern Oaxaca, Mexico, across Central presumably from P. spinipollex, was found America to western Panama with a distri- attached to the undersurface of a large, butional gap in the Nicaraguan De- rather flat rock in a seepage area pression; nine species are known to occur (MCCRANIE & WILSON2002). In captivity, in Central America (FROST2010). Adults of females of P. legleri attached the eggs to this nocturnal treefrog are found on low the roof of a cavity, usually below the vegetation, logs, boulders, and on the waterline (PRoY 1993). The tadpoles of ground within, above, or near clear, swift Ptychohyla species develop in the slowerstreams in the forest (MCCRANIE & WILSON moving shallows or gravel-bottomed pools 2002). Males have been reported calling of the streams where they rest and feed on from bushes and thick grasses along the bottom (DUELLMAN 1970, SAVAGE 2002; streams and seepage areas (MCCRANIE& KOHLER unpubl. obs.). WILSON2002). The vocalizations of P.
Ptychohyla dendrophasma (CAMPBELL, SMITH & ACEVEW2000), 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 Chalatenango, 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.
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
Ptychohyla leg& (TAYLOR 1958b), Univ. Kansas Sci. Bull. 39: 33; type locality: "15 k m 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 (197411. Males to 36.7 mm, females to SAVAGE 37.0 mm. Pacific slopes of the Cordillera de Talamanca of Costa Rica and extreme western Panama, 880-1600 m elevation. 19571, Ptychohyla m a m o ~ p a n u m(TANNER 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 escarpment 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.
m
Fig. 630. Ptychohyla hgpomykter (Selva Negra, Matagalpa, Nicaragua, 1300 m). Photo: G. Kiihler
I Fig. 631. Ptychohyla leglen' (Las Cruzes, Puntarenas, Costa Rica, 1220 m). Photo: J. Vargas
257
Hvlidae
Fig. 632.Ptychohyla salvadorensis (Nahuaterique, M o r a h , El Salvador, 1390 m). Photo: T. Leenders
Fig.635.ptychohyla macrotympanum (Tapalapa, Chiapas, Mexico, 1741 m). Photo: A. Ramirez-Vekizquez
Fig.633.Ftychohyla macrotympanum (Berriozaba.,Chiapm, Mexico, 1241 m). Photo: A. Ramirez-Velbquez
Fig. 636.Ptychohyla panchoi (Los Amates, Aldea Vista Hermosa, Izabal, Guatemala). Photo: J.A. Campbell
Fig. 634. Ptychohyla macrotympanum (same locality as in Fig.635). Photo:A.Emhz-VeUzquez
as in Fig.639).
(Courtesy of the Biodiversity W t u t e , University of I(ansae)
:same localil Photo: J.H. Townsend
, 37. . ,:hohyla spin
I
Hylidae
d Fig. 638. Ptychohyla hypomykter (P.N. Saslaya, Atlantico Norte, Nicaragua).
'
Fig. 639. Ptychohyla spinipollex (La Liberacibn, Atlhtida, Honduras, 1030 m). Photo: J. H.
Photo: J. Sunyer
Ptychohyla salvadorensis (MERTENS 1952b1, Senckenb. Biol. 33: 169; type locality: "Hacienda San Jose, 800 m H., MetapanGebirge, 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 southcentral Honduras, 700-2050 m elevation. Ptychohyla sanctaecrucis CAMPBELL & SMITH 1992, Herpetologica 48: 155; type locality: "near Chinamococh, Finca Semuc, Departamento de Izabal, Guatemala; ...This locality is located at 640 m elevation in the Rio Sauce drainage of the Sierra de Santa Cruz at about 15"37'N, 89O29W." Males to 32.5 mm, females 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 Photo: J. H. Townsend as in Fig. 639).
259
SVL of single known female 84 mm ........... .......................... Pip&ohyla dendrophasma 1 a A distinct lateral white stripe from axilla to groin; width of disc on Finger I11 disb Fingers about one-fourth to one-half tinctly greater than tympanum length; iris webbed, modal formula I11 2+ -2 IV to I11 2 red or reddish bronze in life (usually redl/2 - 2 l/2 IV;toes with less webbing than dish brown in preservative) ..................... 2 described above; inner tarsal fold distinct, weak, or absent; SVL of adults <60 mm . 6 b No lateral white stripe from axilla to groin; width of disc on Finger I11 smaller than or 6 a Dorsum lime green with scattered pale only slightly greater than tympanum green dots in life; fingers about one-fourth length; iris not red in life (not reddish webbed, modal formula I11 2 l/2 - 2 l/2 IV; brown in preservative) ............................. 5 flanks with a narrow zone of reticulation where dorsal and ventral coloration meet; 2 a Venter with scattered brown flecks; adult adult males with a patch of tiny black males with a patch of 50-55 moderately spines on base of thumb; a distinct inner large black spines on base of thumb; dorsal tarsal fold present present; SVL of adult surfaces of head and body grayish tan with males 29-33 mm, of adult females 41-51 green flecks, especially prominent on eyemm .................... &ychohyla sanctaecrucis lids; a vertical rostral keel present; modal webbing formula of fingers I11 2+ - 1 IV; b Dorsum pale tan, dull brown, or reddish SVL of adult males 31-34 mm, of single brown with or without scattered dark known female 37 mm &ychohyla p c h o i brwon blotches in life; fingers about onehalf webbed, modal formula I11 2+ -2 IV; b Venter immaculate; adult males with a flank coloration not as described above; brown non-spinous pad on base of thumb or nuptial excrecences in males variable; an with a patch of 125-135 small to moderateinner tarsal fold distinct, weak, or absent; ly small black spines; dorsal surfaces of SVL of adults variable .............................7 head and body dark brown, uniform or mottled with small pale, bronze, or green 7 a Adult males with a non-spinous pad on flecks, or pale reddish brown with dark base of thumb (Fig. 641a); no ventrolateral brown reticulations; no vertical rostral gland in adult males; SVL of adult males keel; modal webbing formula of fingers I11 27-35 mm, of adult females 34-42 mm ..... 2+ - 2 IV;SVL of adult males 31-37 mm, of ............................&ychohyla salvado& adult females 36-39 mm ......................... 3 b Adult males with a patch of black spines on 3 a Width of disc on Finger I11 distinctly base of thumb (Fig. 641b); ventrolateral greater than tympanum length; adult gland weakly to prominently developed, or males with a brown non-spinous pad on only scattered small glands present base of thumb; ventrolateral gland absent between axilla and groin in adult males; in adult males; iris red in life; SVL of adult SVL of adults variable ............................. 8 males 31-37 mm, of adult females 36-39 8 a Flank without pale spots or mottling; adult mm ................................Pip&& yla leglmeR males with a patch of 90-100 small to modb Width of disc on Finger I11 about subequal erately small black spines on base of to tympanum length; adult males with a a thumb; inner tarsal fold weak, low and patch of 125-135 small to moderately rounded; SVL of adult males to 38 mm, of small black spines on base of thumb; venadult females to 45 mm .............................. trolateral gland present in adult males; iris macro@npanum ...................... -&la reddish bronze in life; SVL of adult males b Flank with pale spots or mottling; adult to 37 mm, of adult females to 53 mm ..... 4 males with a patch of 27-78 moderately 4 a Venter immaculate; adult males with a a large black spines on base of thumb; inner patch of 44-143 small to moderately small tarsal fold distinct; SVL of adults variable 9 black spines; iris reddish bronze in life; 9 a Belly and ventral surfaces of thighs SVL of adult males to 37 mm, of adult immaculate or with widely scattered females to 53 mm Rydiohyla e u ~ ~ o t a flecks; ventrolateral gland prominently b Venter with scattered brown or black developed in adult males; SVL of adult flecks; adult males with a a patch of males 30-35 mm, of adult females 3 5 4 0 90-100 small to moderately small black mm ....................... &&+la hypomyktar spines; iris coppery bronze in life SVL of b Belly and ventral surfaces of thighs usualadult males to 38 mm, of adult females to ly peppered throughout with brown flecks; 45 mm ......... &ychohyla macro@npanum ventrolateral gland not well-developed, 5 a Fingers about two-thirds webbed, modal only scattered small glands present formula I11 2 l/2 - 2 112 IV; toes nearly between axilla and groin in adult males; fully webbed, modal formula I11 1 -1 l/2 IV; SVL of adult males 33-39 mm, of adult a distinct inner tarsal fold present present; females 39-46 mm P&.hohyh spini'IIex
Key to ptschohyla
P: salvadorensis
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 & WILSON1989, CAMPBELL & 1982, MCCRANIE SMITH 1992, MCCRANIE & WILSON 1993b, CAMPBELL et al. 2000, DUELLMAN 2001
Hylidae
Fig. 643.Scinax boulengeri (La Selva, Heredia, Photo: G. Kiihler Costa Rica, 70 m).
Fig. 644.Scinax elaeochroa (Punta Uva, Limbn, Costa Rica, 10 m). Photo: G. Kohler
restricted to the rainy season, given the strong seasonality that prevails where this species occurs (CAMPBELL 1996). During About 100 species of small to moderate- the dry season, individuals of S. s t a d e r i sized anurans are currently assigned to have been found hiding under logs or rocks the genus Scinax, the vast majority of on the ground, beneath the bark of standthem occurring in South America (FROST ing trees, and within bromeliads near 2010). Central America is home to six water bodies (SMITH1941, CAMPBELL 1996, species of this genus. Some controversy LEE 1996). The call of S. elaeochroa has had existed as to the gender of the genus been described sounding like "waak", that name Scinax (DUELLMAN & WIENS 1992, of S. boulengeri like "wraak", and that of S. KOHLER & B O m 1996, SAVAGE 2002).I fol- boulengeri Like "ah-ah-ah-ah" (LEE 1996, low SAVAGE (2002) in regarding Scinax as SAVAGE 2002). Scinax species typically of masculine gender. All Scinax are noctur- breed in shallow temporary bodies of water nal and arboreal. Scinax staufferi is found throughout most of the wet season (LEE in a wide variety of habitats including dry, 1996, SAVAGE 2002). After the first heavy moist, and wet forests, usually in relative- rains hit the breeding site, males, especially open situations, avoiding deep forest ly of the common species, such as S. (LEE 1996). Scinax elaeochroa and S. elaeochroa and S. staufferi, suddenly conboulengeri, on the contrary, are more typi- gregate, and soon numerous males produce cal lowland rainforest species, inhabiting a raucous chorus (SAVAGE 2002). Amplexus forest habitats, often found near breeding is axillary in Scinax and eggs are laid as habitats (SAVAGE 2002, SCOTT1983d). small clumps in the shallow water where These two species are prolonged breeders, the tadpole hatch and complete their and males can be heard calling throughout development (DUELLMAN 1970, LEE 1996, most of the year (DUELLMAN 1970). The SAVAGE 2002). breeding activities of S. staufferi are
Hylidae
Scinax d t a e ~ 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 (COPE1887), 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.
Fig. 645. Scinax altae (Los Algarrobos, Photo: A. Hertz Chiriqui, Panama, 125 m).
Scinax elaeochroa (COPE18761, 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 a n 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 (LAURENTI17681, Spec. Med. Exhib. Synops. Rept.: 5; type locality: "America"; neotype from "Paramaribo, Surinam" (DUELLMAN & WIENS1993)). 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(COPE1865), 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. 648. Scinax staufferi(Santa Lucia, Intibuca, Honduras, 370 m). Photo: G. Kohler
Fig. 646. Scinax elaeochroa (Guayach, CRARC, Costa Rica, 550 m). Photo: G. Kohler
Fig. 647. Scinax staufferi (near San Jose E l Idalo, Suchitep6quez, Guatemala, 70 m). Photo: G. Kijhler
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
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 weakly, 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 ........................
S. stauffen' S. altae
........................................ 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 8tab 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,
I
'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 [
J
Fig. 650. Scinax ruber (Wacu~u,Panama, Photo: A. J. Crawford Panama, 130 m).
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. fodiens, traditionally were assigned to a separate genus, Pternohyla, before FANOVICHet 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-wonkwonk" (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 (KOHLERunpubl. 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, MALONE2004). 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 Fig. 651. Scinax rostratus. (Wacuco, PanamB, Photo: A. J. Crawford species (DUELLMAN Panama, 120 m). 1970). The pond-breeding species of Smilisca exhibit a generalized reproductive mode with eggs depositFurther Rsading RIVERO 1969, DUELLMAN 1970, FOUQUETTE & ed as a surface film in lentic water; the two PY~URN 1972, FOUQUETTE & DELAHOUSSAYE 1977, SCOTT1983d, DUELLMAN 1986, JUNGFER stream-breeding species on the other hand, 1987, DUELLMAN & WIENS1992, LANGONE1992, construct basins for egg deposition DUELLMAN & WIENS1993, K o m & ~ B ~ H M E (MALONE2004). During amplexus, females 1996, NIETO1999, SAVAGE 2002 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 (MALONE2004). 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).
Smilisca baudinii ( D U ~ I &L BIBRON18411, 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 embayment 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), Herpetologica 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 (COPE18851, 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.
Fig. 652. SmiLisca baudinii (Mazatenango, Photo: G. Kiihler Guatemala, 395 m).
;
,
I Fig. 653. Smilisca baudinii (Lagarto Lodge, Alajuela, Costa Rita, 90 m) photo: G. Kohler
I
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 central Panama to northern Colombia and the middle Magdalena Valley, sea level to 1350 m elevation. Smilisca sordida (PETERS 1863b), Monatsber. Preuss. Akad. Wiss. Berlin 1863: 460; type locality: veragua", Males to 45.0mm, females to 64.0 mm. On the Atlantic versant 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 SanPhoto: Juan, G. Kohler Nicaragua, 30 m).
Hylidae
Fin. 655. SmiIisca ~haeota(near Pueblo Wiso, Photo: G. ~ 6 h l k r Nicaragua). u
I 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
b Fig. 656. Smilisca phaeota (Cerro El Toro, Atlantic0 Norte, Nicaragua). Photo: G. Kohler
2 a
b
Fig. 657. fj'milisca slla JosB, Costa Rica, 990 m).
Rafael Norte, Sari Photo: G. Kohler
Fig. 658. Smilisca sordaa (ruco ae neara, Veraguas, Panama, 900 m). Photo: G. K'ohle
3 a
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 m m ............. 2 No dark brown or black postorbital mark encompassing tympanum present; SVL of adult males 31-54 mm, of adult females 40-62 mm ................................................. 4 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 .baudinii 56-90 mm ...................... S 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 Groin as well as anterior and posterior Surfaces of thighs with blue spots; SVL of adult males 50-56 mm, of adult females
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 4 0 4 6 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 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); dorsal surfaces smooth; snout longer, sloping. rounded: blotches on dorsum having th;;r'tendencytoform transverse bars; vocd sacs of breeding males white; SVL of adult males 32-54 mm, of adult females 56-64 mm .................................... SmiliecaBardida
a.
C. I
Fig. 660. Hand of a. Smilisca baudinii; b. S. Puma; c- S. sila; d. S. sordida. Drawings: M. Vesely -
I
I
Smilisca
:. baudinii
4 Smilisca
S. sila S. puma
. L
*
Smilisca
I
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 distribution in Mexico (DUELLMAN 2001). Tlalocohyla loquax is a widespread noctur. nal frog that is found in and around tem- Fig. 661. Tlalocohyla loquax (Guayach, porary and permanent ponds and small CRARC, Costa Rica, 550 m). Photo: G. Kohler lakes (DUELLMAN 1970, MCCRANIE& WILSON2002). 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 Fig. 662. Tlalocohyla picta (near Chetumal, (DUELLMAN 1970). The females deposit Quintana Roo, Mexico, 20 m). Photo: G. K6hler large gelatinous egg masses attached to submerged plants near the water surface; one such clutch contained about 250 eggs nent ponds and small lakes, forest pools, (DUELLMAN 1970, CAMPBELL 1998, SAVAGEgrassy depressions in pastures, and road2002). The tadpoles are exceedingly wary side ditches and call from emergent vege1970, LEE 1996). and inhabit the deeper portions of the tation (DUELLMAN water body where they prefer an abun- Amplexus is axillary in both T. loquax and T. picta (LEE1996).The call of T.picta condance of aquatic vegetation (DUELLMAN 1970). T2alocohyla picta is a common frog sists of series of high-pitched, insect-like that inhabits a variety of lowland habitats chirping notes, described as "cric-ic-ic-ic" 1970, LEE 1996, ROBLE1997). such as savannas, forest edges, and tall for- (DUELLMAN est (DUELLMAN 1970, LEE1996).During the Clutch size in T. picta was determined to day and in the dry season, individuals of T. vary between 160 and 956 eggs with an picta have been found in the axils of the average of 543 eggs per clutch (ROBLE 1997). The females deposit the eggs in leaves of elephant-ear plants (DUELLMAN 1970).After the onset of the rainy season, small packages at the water surface o r males congregate at temporary and perma- attached to aquatic plants (ROBLE1997).
Tlalocohyla loquax (GAIGE& STUART1934), 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 * l a
loquax
,.
loquax (near San Ignacio, , 663. , Photo: G. Kijhler Cayo, Belize, 240 m). x,,,uA,
Trachycephalus
b A distinct pale (white or yellowish) lateral or dorsolateral stripe between levels of The genus Trachycephalus consists of 12 axilla and groin; fingers moderately species of large, robust frogs with a center webbed, webbing between Toes 111 and ZV of diversity in South America; only a single reaching penultimate tubercle on Toe IV; webbing yellow or yellowish tan in life; pos- species, T. venulosus (milky treefrog), . terior surface of thigh uniform yellow in occurs in Central America (FROST 2010). life; SVL of adult males 19-21 mm, of adult The latter species was assigned to females 19-22 mm .......... 111aloa&la picta Phrynohyas for the past decades before FAIVOVICH et al. (2005) placed Pluynohyas i n the synonymy of Trachycephalus.The --. _ __ n..a..l.l-.__ milky treefrog is a widespread, nocturnal, T: loquax i arboreal inhabitant of primary and disr: picta t turbed habitats. It passes the dry season 1 sequestered in bromeliads, tree holes and crevices, under bark on standing trees, and in the axils of the leaves of heliconias or 2002).At night, bananas (LEE1996,SAVAGE 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 ( C o n 1926). Characteristic for T. venulosus are the thick dermal glands on the dorsum that can secrete a large Further Reading amount of sticky white, poisonous, waterDUELLMAN1970, ROBLF: 1985, LEE 1996, ROBLE insoluble mucus, that is highly irritating to 1997, CAMPBELL1998, M c C m & WILSON mucous membranes and can cause sneez2002, SAVAGE2002 I
1
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 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).
P
', Fig. 664. Trachycephalus venulosus (near Chetumal, Quintana Roo, Mexico, 20 m). Photo: G. Kohler
I Trachycephalus venulosus ( L A ~ N T 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 Tamaulipas 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 (Alamikamba, Atlantico Norte, Nicaragua, 20 m). Photo: G. Kohler
Further Reading DUELLMAN1956, HEMMING1958, ZWEIFEL 1964a, PYBURN 1967, MCDIARMID 1968, DUELLMAN 1970, 1971a, LEE 1996, CAMPBELL et al. 1998, MCCRANIE& 1998, MANZANILLA 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. SMITHet 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).
6
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, k'lg. 666. Triprion petasatus (near 'I'eabo, Photo: G. Kohler M c C m & WILSON2002). During the Yucath, Mexico, 20 m). 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 Trprion petasatus (COPE18651, Proc. Acad. Nat. Sci. Philadelphia 17: 193; type locality: 1935, LEE 1996, KOHLER1997). The frogs "vicinity of Mbrida, Yucatan ... f?om a hole in plug the openings with their casque-like the rocky wall of the Cenote Pamanche, on the heads, a behavior called "phragmosis", and new road to Progreso" 117 k m north of Mbrida, are almost impossible to extract; the mYucatsn, Mexico]. Males to 60.8 mm, females ossified dorsal head surface obviously to 74.2 mm. Yucath Peninsula, Mexico, southhelps to reduce cutaneous water loss (LEE ward to Guatemala and Belize; also at Lago de Yojoa in northwestern Honduras, sea level to 1996). During the first heavy rains of the 740 m elevation. wet season, males congregate at temporary ponds, limestone solution pits, and sink holes at night, and call from shrubs and Triprionpetasatus small trees as well as from the edge of the water bodies (LEE 1996, MCCRANIE & WILSON2002). The call was described as a series of low-pitched notes reminiscent of 1970, the quacking of a duck (DUELLMAN 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,LEE1996).Tadpoles F'urther Reading prefer shaded areas and seek refuge in DUELLMAN & KLAAS 1964, DUELLMAN1970, decaying vegetation on the bottom of the '~'RUEB 1970, LEE1996, KOHLER1997, CAMPBELL & WILSON2002 1998, MCCRANIE water body (DUELLMAN 1970).
I
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 GRANTet al. (2006). As currently understood, this family contains 79 species distributed from southern Mexico across Central America and South America (FROST2010). Two genera of Leiuperidae with one species in each occur in Central America, the &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 dedicated to its ecology, life history, and sexual selection kg., RYAN1985, RYANet al. 1996, RON et al. 2005, LG~'~PERT et al. 2007).
wars
-
Fig- 667. EngJ'stomo~s~ u s t d o s u (Guaquira, s Yaracuy, Venezuela). Photo: J. Sunyer
laemus where it had been placed by LXNCH (1970b). FUNR et al. (2008) suggested that 1 a A prominent deep purple to black colored the data provided by NASCIMENTO et al. (2005) did not just* the separation of s PhJ'salaemus. Pubfaces of groin, and posterior surface of E n ~ s t o m o ~from 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. pusb 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(brown same "lor as adjacent to grayish brown);ventral surface usually lations north of Panama (RYANet al. 1996, et al. 2005, with small to large dark spots .................... WYNN & HEYER2002, WEIGT .............................~ o p e p u s t u l o ~ BuO ~H L et al. 2006).
Key to Central American species of Leiuperidae
~ ~ ~ ~ U ~ l ~ & ~ ~ $ h ~ " , ~ f ~
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. DUE^ (1960) reported on a series of Eight species are currently recognized in 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-
Further Reading GRANT et al. 2006
-
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 (RYAN1985). 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 con- Fig. 668.Engystomops pustulosus (Nusagandi, struction of the foam nest by this species is Kuna Yala, Panama, 350 m). Photo: G.Kohler provided by HEYER& RAND (1977). The foam can prevent egg desiccation during short periods without rain (RYAN1985). 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 (RAND1983, RYAN 1985).
Aead. Nat. Sci. Philadelphia 16: 180; type locality: 'New Grenada, on the River Truandow,Colombia. Males to 34 rmn,females to 35 mm.On the Atlastic versaat from eouthern Vera~uz,Mexico, and on the Pa&c versant imm Oaxaca, Mexico, to e ~ t e r Panama; n also in northern South America, sea level to 1540 m elevation. Engystornops pustulosus !
Further Reading RAND 1983, CANNATELCA& DUE~LLMAN 1984, RYAN1985, RYAN et al. 1996, RON et al. 2005, LAMPERT et al. 2007
Fig. 669. Enggstomops pustulo~us(6:
Lorenzo, Suchitep6quez,Guatemala, 210 m, ~ ~ ~ t ~ ~ ~ Photo: l ~G. Kiihler ) .
Pleurodema As currently understood, the genus Pleurodema consists of 14 species with a predominantly South American distribution (FROST2010). 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 & VELOSO1977, HOOGMOED & Go= 1979, M. RYAN pers. com. October
1
I I
!
/
2010). When handled, this fiog inflates its body and elevates the hind parks of the body displaying the conspicuously colored lumbar glands and the orange coloration of the groin and posterior thighs; this supposedly 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 call can be heard on any rainy night for the rest of the rainv " season: toward the end of the season, the aggregations seem to consist almost exclusively of males (HOOGMOED & GORZULA 1979). For breeding, P. ~ i g670. . 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 ~ etZ al. 1999b). The eggs are laid in a foam nest floating freely on the surface of the water (HOOGMOED & GORZULA 1979).
I
P l e d e m a bra&pop% COPE 186% g868", h e . M.Nat, f i .PbhLIphia 20: 312;type bdty: "Wqddena Bveq New Cm& I= Colambia]." SVL to 44 ma. Penfmd8 d0 huwo and central Pamum -Do nozthernSouth Am& (northern Cohmbia, Venemela to Ou-' BFd" in Panamtt eea l d ta k?OU m elmathon.
Fig. 6'71. PIemdema brachyops (Cien6ga of Caimanera, Sucre,Colombia). Photo: A. Acosta
Fig. 672. Pleurodema brachyops (El Rodeadero DU~UMAN& VELOSO 1977, HOOGMOED & de Gaifa, Magdalena, Colombia). GORZULA 1979, MARTINS 1989, FERRARO & Photo: W. E. Duellman (Courtesy of the Biodiversity Institute,Univerai@ of Kansas) CASAGRAN~A 2009
Further Rsading
Leptodactylidae
or are placed hidden in vegetation, under debris, or in the mouth of a burrow in streambeds or other seasonal flooded sites (BREDER1946, SAVAGE2002). 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.
Prior to 2006, the Leptodactylidae was a very large unwieldy family of frogs. RWINSKY& MAXSON(1996) suggested that Leptodactylidae (sensu lato) was polyphyletic and FROSTet al. (2006a) partitioned the former Leptodactylidae into several reformulated families. Subsequently, GRANTet al. (2006) split the Leptodactylidae into the Leptodactylidae (as now recognized) and the Leiuperidae. As currently understood, the family Leptodactylidae contains four genera, one of Leptodactylus savagei is the largest which, Leptodactylus, occurs in Central Central American species in this genus. America (FROST 2010). Most species in this During the day, this nocturnal frog hides in family are moderate-sized frogs, but some burrows and under logs; juveniles can be species can reach a very large (up to 185 found active at daytime on leaf litter in mm SVL) adult size (VITT & CALDWELLshady forest situations (SAVAGE2002). They are important forest floor predators 2009). and eat just about any live prey they can fit into their mouths, including arthropods, M e r Reading GORHAM1966,L m c 1971, ~ RUVINSKY& MAXSON frogs, lizards, snakes, birds, and mammals 1996, FROST et al. 2006, GRANTet al. 2006 (SCOTT1983e). When caught, these frogs sometimes emit a loud high-pitched scream (SCOTT1983e). The skin secretions of adult individuals contain noxious compounds that are highly irritating to mucous membranes and little cuts in the The genus Leptodactylus consists of 88 rec- skin (VILLA1969); the mucus compounds ognized species, distributed from southern can be irritating by diffusion through the North America, across Central America air without direct contact, resulting in and South America, and into the West heavy sneezing and swelling of the eyes 2002). These secretions can disIndies (FROST2010). HEYER(1969) rede- (SAVAGE fined the genus, and in numerous subse- courage potential predators; R U N et al. quent papers contributed much to the cur- (2010) reported on a Bothrops asper that rent taxonomic knowledge of this group of they encountered while it was attempting frogs (e.g., HEYER1970a,b, 1972, 1974, to consume a specimen of L. savagei. The 1978, 1979, 1994, 2002, 2005). Lepto- snake regurgitated the frog and showed dactylus are terrestrial or semiaquatic and signs of poisoning including the loss of the nocturnal, occupying a variety of habitats righting reflex; it took more than an hour including tropical rainforest, marshes, for the snake to recover. Males call solitarlakes, ponds, and roadside ditches through- ily from margins of ponds and streams; the out Lowland and Premontane Dry, Moist, call is a loud "whorup" repeated at inter1946, and Wet Forest zones (sensu HOLDRIDGEvals of five to ten seconds (BREDER 1967) (SAVAGE 2002). All Leptodactylus VILLA 1969). An attracted female is build foam nests in which the eggs are laid; grasped by the male by axillary amplexus nest of L. savagei can contain 2-7 liters of and held firmly by the muscular forearms foam (SCOTT1983e). The foam nests are and the stout nuptial spines on thumbs 2002). either floating on the surface of the water and chest (SAVAGE
Leptodactylus bolivianw BOULENGER 1898~ 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 M i p e l and Saboga I~lands(Panama), sea level to 1400 m BARBOUR elevation. Lephikctylm ins1906 is regarded as a synonym of L. bolivianus, according to REYER (1970a, 1974) and (2002). SAVAGE Fig. 674. Leptodactylus boliviii~lus(male, Leptodadyllue h& (BROCCRI 1877~1,Bull. Ponuga, Veraguas, Panama, 35 m). Soc Philomath., Paris,Ser, 7,l: 182;type localPhoto: G. Kahler ity: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., HEYER1971),which is now a junior synonym of L. mystacjnus (see HEYElR 1978,2002). Leptodact-JImf u s ~ t(SCHNEIDER l~ 1799), Hist. Amph. Nat.: 130; type locality: implied to be from Surinam; neotype from "Surinam" (HJWIIR19681). 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. 673. Leptodactylus fragilis (road from FIacienda Sari Cayetano to Sari Jose El Iddo, Suchitep6quez, Guatemala, 70 m). Photo: G. Kohler
Fig. 675. Leptodadylus bolivianus (male, Ponuga, Veraguas, Panama, 35 m). Photo: G. K6hler
Fig. 676. Leptodadylus &agjlis (Cem Mariposa, Veraguas, Panama, 900 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.
Fig. 677. Leptodactylus savagei (juvenile; San Ram6n de La Virgen, Heredia, Costa Rica, Photo: G. Kohler 10 m).
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 sea level to 1150 Leptodactylus savagei HEYEa 2005, Arq. Zool. 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 versant south to ~olombia, sea level to 1200 m elevation.
WILSON Leptodactylus silvanimbus MCCRANIE, & 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. 678. Leptodactylus savagei (Guayacln, CRARC, Costa Rica, 550 m). Photo: G. Kohler
Fig. 679. Leptodactylus savagei (Nusagandi, Kuna Yala, Panama, 350 m). Photo: G. Kahler
Fig. 680. Leptodactylus silvanimbus (fiom type locality). Photo: J. R. McCranie
1
L. poecilochilus
I
Lepfodactylus
ck - 1
C -
L. savagei
Fig. 681. Leptodactylus melanonotus (Lagarto Lodge, Alajuela, Costa Rica, 90 m). Photo: G. Kohler
<._I.
I
I
melanonotus L. fuscus
I
Leptodactylus
-4
Leptodactylus
--,.
Key to Leptodactglus
1 a A pale longitudinal stripe present on poste-
Fig. 682. Posterior surfaces of thigh and shank in Leptodactylus poecilochilus; note pale Ion@Photo: G. Kohler tudinal stripe (arrow).
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); males with internal vocal sacs ............... 4 2 a Dorsum with six distinct longitudinal 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 3 a Ventral surfaces of foot and tarsus with prominent tubercles (Fig. 683a)...................
1
.................................. 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 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 3 5 4 8 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 sab 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 ..............
I
lb.
Fig. 683. Ventral surface of foot in a. Leptodactylus fi-agilis(foot and tarsus with prominent tubercles); b. L. poecilochilus (foot and tarsus smooth). Photos: G. Klihler
I Fig. 684. a. Hand of Leptodactglus melanonotus; b. Posterior thigh of L. b ~ l i ~ a n u s . Photos: C:Kiihler
............................ 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 3 5 4 3 mm, rarely to 55 mm .......... ........................ Leptodactglus melanonotus b Toes with lateral keels or only weakly developed lateral fleshy fringes; adult males with hypertrophied arms; SVL in adult males 40-53 mm, in adult females 4 3 4 8 mm ...... Leptodactglus ailvanimbus
--
Fig. 685. Leptodactyluspoecilochilus (Meseta de Chorcha, Chiriqui, Panama, 250 m). Photo: G. Kohler
Fig. 686. Leptodactylusfuscus (near Minacu, Goias, Brazil). Photo: W. Van Devender
Fig. 688. Leptodactylus melanonotus (Ponu,-, Veraguas, Panama, 35 m). Photo: G. Kohler
FurtherRaading
' Fig. 687. Leptodactylus melanonotus (9 km NW Fortuna, Alajuela, Costa Rica). Photo: G. Kohler
HEYER 1970a,b, 1971,1972,1974, STRAUGHAN & HEYER1976, HEYER1978, 1979, HEYER & MAX~ON1982, Scom 1983e, MCCRANIE et al. 1986, SOLANO 1987, MARTINS 1988, D w o ~ s& HEYER1992, HEYER1994, HEYER et al. 1996, 2002, HEYER2002, HEYER & REID2003, HEYER 2005, HEYER et al. 2005, PONSSA 2008, HEYER et 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 (SAVAGE2002). 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).
282
Photo: G. Kahler
Key to the genera of Central American Mimhylidae
1 a Inner and outer metatarsal tubercle pre2 sent (Fig. 690) ........................................... 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). Photo: G. Kiihler ly uniform glossy black .....Nelemoplqme b Toes with basal or no webbing; venter with or without markings; dorsal surfaces of head, body, and b b s essentially not uniform 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 6 without webbing ....................................... 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- NelBOnoPhryneate-a, Drawings: M. Vesely terior surface of thigh with flashy red markings in life; no stripe on po&erior surface of thigh ......................RaZictivrnner
FurtherRsading PARKER1934, Durn 1949, DE CARVALHO 1954, WALKER1973, ZWEIFEL 1986, WILD 1995, VAN DER MEIJDENet al. 2007, & CALDWELL 2009
-
Fig. 692. Relictivomerpearsei (ventral surfaces; Malagana, Bolivar, Colombia, 30 m). Photo: M. Garcia Paris
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 Fig. 693. Chiasmocleis panamensis (Gamboa, and rocks (RENJIFO & LUNDBERG 1999).The CoMn, Panama, 29 m). Photo: A. J. Crawford species is reported to be locally common in central Panama where individuals primarily occupy open areas with bushy vegetation; they are only encountered occasional-*->-*. ' . ->...,*..-.**, ..- 7 ly in the forest ( I B ~ et z al. 1999b). Chiasmocleispanamensis 1 E 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. ,~,,.,saA.~,~-~.~,..
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.
Further Reading
>
~
-,
DUNNet al. 1948, DUNN1949, 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, TOLEDO2010, TOLEDOet al. 2010). CARAMASCHI (2010) discussed the taxonomic position of E. ovaLis (SCHNEIDER 1799) and considered this taxon as a nomen dubithereby rendering the Panamanian populations formerly referred to as "Elachistocleis ovalid' without an identification. Thus, I have refrained from including a species account under this genus. Elachistocleis are small nocturnal, fossoria1 frogs. As is typical for most microhylids, 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, LAVILLAet al. 2003, 2010, TOLEDO et al. TOLEDO 2010, CARAMASCHI 2010
b*.n;
i
-/
Fig. 694. Elachistocleis sp. (P.N. G. D. Omar Torrijos H., near El Cop6, Cocl6, Panama). Photo: R. Brenes
Fig. 695. ~ ~ ~ ~elegans ~ ~( ~~ ~hde la ir ~y ~n f Cruz, Atlhntida, Honduras, 5 m). Photo: J. Sunyer
GastrODhrvne Five species are currently recognized in the predominantly North American genus Gastrophryne, distributed from the southern USA to Costa Rica (NELSON1973b). 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 frequently found in secondary growth (SAVAGE 2002). All species of Gastrophryne feed on ants, termites, small beetles, and other small insects. Typical for explosive breeders, they arrive at their breeding sites - shallow pools and flooded ditches after heavy rains, mostly during the months from May to August (STUART 1934, CAMPBELL 1998, SAVAGE 2002). The mating
E I
call of G. elegans has been described as a prolonged "baaaaaaa" (CAMPBELL1998) or a high-pitched, rather nasal "naaaaaaa" (LEE1996,2000). The males of G. pictiventris have been observed calling while floating in the water, with synchronous calling of several males (NELSON1973c7 SAVAGE 2002). The eggs are laid on the surface of the water and hatch within a few days (MEYER& FOSTER 1996). The tadpole of G. pictiventris has been described and illusFig. 696. Gastropluyne pictiventris (Guayachn, trated by DONNELLY et al. (1990b). 1
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 (COPE18861, Proc.
Am. Philos. Soc. 23: 272; type locality: "Nicaragua." Males to 31 mm, females to 38 mm. Atlantic slopes of southeastern Nicaragua and Costa Rica, sea level to 500 m elevation. Gastrophgme usta (COPE1866), 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 NELSON1963, LEGLER 1964, NELSON1972a,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 . variolosushas 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, KOHLERet 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 1m underground (SAVAGE 2002). They feed mostly on ants, termites and other small arthropods. Typical for an explosive breeder, these frogs emerge after heavy downpours, 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& FOSTER1996, 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). Photo: J. H. Townsend
Fig. 699. Hypopachus variolosus (Morgans Photo: J. Sunyer Rock, Rivas, Nicaragua).
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 (LEE1996, 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 (COPE1866), 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.@?&uBibanban' 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. 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 Fig. 700. Foot of a. Hypopachus variolosus; Drawings: M. Vesely only two specim, one of which, N.atenima, b. H. barben'. occurs in lower Central America. Nel~onophqyneatenima 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 KUBICMpers. comm. July 2010). Gravid females have Hypopachus 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 Further Reading NELSON 1973a, 1974, MCCRANIE& WILSON of N. aterrima from a small puddle at the edge of a logging road in early October, 2002, K~RLER et al. 2006
w
N e ~ m n o p ~a et 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
1
PARKER1934, DE CARVACHO 1954, BOGART& NELSON1976, 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 & LUNDBERG1999). 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). 1914), Proc. Relictivomer pearsei (RUTHVEN 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& CANNATELLA1986). Males in this genus lack vocal cords but are able to communicate 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 vascularized skin of the female's dorsum. The freeliving 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, TRUEB1984). 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 TRUEB1984, 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).
fig. 705. Lithobates vibican'us (subadult;near entrance to P.N. Juan Castro BIanco, AIajuela, Photo: G. Kiihler Costa Rica, 1900 m).
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 Guatemalan 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 HILLIS & DE SA (1988) published a revision the Meseta Central of Chiapas and the of the Lithobates palmipes species group. Central Plateau of Guatemala. Because However, our state of taxonomic knowledge there is no way to morphologically delimit regarding the Lithobates pipiens species L. macroglossa from L, forreri and L. group is less advanced. Much uncertainty b r o w n o m and no published evidence is is associated with the nominal species L, available to support the species status of macroglossa (BROCCHI1877~). Some this form, I prefer not to recognize L. authors considered it to be synonym of L. macroglossa as a valid species until a taxmaculata (e.g., STUART 1963, HOLMAN & onomic revision of the Central American BIRKENHOLZ 1963, h~1976). S m et al. members of the L, pipiens species complex (1966:171) designated MNHN 6321 as the has been published (see also comments in lectotype of Lithobates macroglossa and HILLIS& DE SA [1988], who considered L. stated that this specimen-"clearly repre- macmglossa to be a synonym of L. berlansents the species Rana pipiens." MCCRANIEdiem?. The Central American s~eciesof &WILSON (2002) analyzed the morphologi- Lithobates are semiaquatic denizens of cal variation in frogs of the L. pipiens ponds, slow-moving streams, river overspecies complex in Honduras and provided flows, swamps, and marshes active both at a lengthy discussion of the complex situa- day and at night. They are characteristic tion there. They concluded that at this inhabitants of the aquatic margin point the available data on Honduran (DUELLMAN 1963a); when disturbed, these material can be interpreted to support frogs jump into the water and swim to the
bottom t o conceal t h e m s e l v e s a m o n g debris. Males call w i t h their paired vocal sacs inflated while floating o n the water surface. The eggs are deposited in globular masses, often a t t a c h e d t o submerged vegetation o r bottom debris (HILLIS& DE SA
1988, SAVAGE 2002).
Lithobates brownorum (SANDERS19731, 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. 1883a1, AM. Lithobates f0rFer.i (BOULENGER 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.
Fig. 706. Lithobates brownorum (near Chetumal, Quintana Roo, Mexico, 10 m). Photo: G. Kohler
Lithobates maculatus ( B R O C C1877~1, ~ 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.
Fig. 707. Lithobates forreri (Zona Protectors El Rodeo, Alajuela, Costa Rica). Photo: G. Kohler
Lithobates taylon. (SMITH19591, 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 (BROCCHI18771, Bull. Soc. Philomath., Paris, Ser. 7, 1: 175; type locality: "sur les rives de la rivi&rede 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.
Fig. 708. Lithobates taylori (Cerro Mariposa, Veraguas, Panama, 900 m). Photo: G. Kohler
Lithobates vibicarius (COPE18941, 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 (SCHMIDT18571, 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. 711. Lithobates vibicarius (adult; near entrance to P.N. Juan Castro Blanco, Alajuela, Photo: G. Kohler Costa Rica, 1900 m).
Fig. 712. Lithobates maculatus (Laguna del Cem, Cop&, Honduras, 770 m). Photo: G.Kiihler
Fig. 709. Litho,,,.s miadis (Isla de Maiz ~equeiia,Nicaragua). Photo: J. Sunyer
I
I
Fig. 710 (above).Lithobates julim' (Columbia River Forest Reserve. Toledo. Belize). Photo: J. C. Lee (Courtesyof the Biodiversity Institute,University of Kansas) Figg. 714 (right). Lithobates vaillanti (Bartola, Rio San Juan, Nicaragua, 30 m).Photo: G. Kijhler
1
Fig. 713. Lithobates vaillanti (P.N. Ma~~llnillo, L i m b , Costa Rica, 10 m). Photo: G. Kohler
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 m a s k no distinct pale supralabial stripe .............................
......................................Lithobates vaillanti b Tympanum smaller (less than eye diame-
3 a Fig. 715. Lithobates warszewitschii (3 k m SE Ayapal, Atlantic0 Norte, Nicaragua, 200 m). Photo: G. Kohler
b
4 a
b
I
1 I Fig. - 716. Lithobates warszewitschii iRio San Photo: G. Kohler Juan, Nicaragua).
ter); distinct dark face mask present; a distinct pale supralabial stripe present ....... 3 Two metatarsal tubercles; toe pads present; webbing on hind feet reduced (Fig. 719b) .......................................................... 4 One metatarsal tubercle; toe pads absent; webbing on hind feet nearly complete (Fig. 719c) ......................................................... 5 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 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 .........................
................................... Lithobates vibimCBRus 5 a Tympanum length almost as large as eye diameter; flanks uniform dark brown; no vocal slits in males ....... Lithobatesjuliani b Tympanum length approximately half the diameter of the eve or less: flanks with dark blotches; vocd slits present in adult males ....................... Lithobates madatus 6 a Dorsal surfaces of lower hind limbs with irregular longitudinal dark bands; restricted to Isla de Maiz Pequeiia, Nicaragua .....
........................................ Tithobatesmiadis b Dorsal surfaces of lower hind limbs with dark crossbars; occurs on mainland Central America .......................................7
Fig. 717. Lithobates warszewitschii (Guayach, CRARC, Costa Rica, 550 m); note different coloration of male (right) and female (left). Photo: G. Kohler
Fig. 718. Coloration of posterior thigh in Lithobates warszewitschii (Rio Curinwas, BOSAWAS, Atlantico Norte, Nicaragua). Photo: G. Kohler
7 a Dorsolateral folds usually continuous, posterior portion not inset medially; dorsal blotches usually 15 or more; vocal sacs darkly pigmented in adult males; Miillerian ducts present in adult males; occurs on the Pacific versant ...................... ........................................ Lithobah h r i b Dorsolateral folds usually interrupted, with the posterior portion inset medially;
dorsal blotches usually 13 or fewer; vocal sacs not pigmented in adult males; Miillerian ducts present or absent in adult males; occurs on the Caribbean versant .8 5 a ~~~u~~~~ducts present in adult males ....
............................... Lithobates brownorum b Miillerian ducts absent in adult males ........................................ Lithobates taylon'
Fig. 719. Foot of Lithobates vaillanti;b. L. warszewitschii; c. L. maculatus;d. L. taylori. Drawings: M. Vegely .....
-L------
------.>
....a.
L. brownoturn i L. forreri
Y
L. rniadis
L. rnaculatu:
+ L. juliani
.
1
L. warszewitschii L. vibicarius t
i
6
I Lithobates
a d
$
1B
1 Lithobates *..a
.a
Further Reading ZWEIFEL 1964b, SMITHet al. 1966, ZWEIFEL 1967, LEE 1976, HILLISet al. 1983, HILLIS& FROST1985, H n m 1988. HILLIS& DE S A 1988. VILLA1988, 1990, ACOSTA-GALVIS 1999, RAM[REz et al. 1998,
Rhinophrynidae
a loud, low pitched "wh-o-o-o-a"with a rising inflection a t the end (FOUQUETTE 1969, FOSTER & MCDIARMID 1983). Large chorusThe Mexican burrowing toad (Rhino- es of R. dorsalis may be heard for several p h q u u s dorsalis) is the only living species kilometers (FOSTER& MCDIARMID1983). of the genus Rhinophrynus and the family Amplexus is inguinal. Both the female and Rhinophrynidae. The family was once more the male are submerged during egg laying; widespread, including species ranging as clutch size may range from 2000 to 8000 far north as Canada, but these became eggs (STUART1935, FOSTER& MCDIARMID extinct during the Oligocene (SAVAGE 1983).The eggs take several days to hatch, 2002). Rhinophgmus dorsalis is a peculiar and the tadpoles develop over one to three frog with a small, cone-shaped head and months, depending on food availability 1961).Adults reach sexual maturifour short, robust limbs projecting from a (STUART large, globular body. Its eyes are relatively ty at 60-65 m m (FOUQUETTE 1969). small, and the tympanum is not visible. It Rhinophrynus dorsalis D ~ R I & L BIBRON feeds underground, specialized on a diet of 1841, Erp. Gen. 8: 758; type locality: "Veraants and termites. The tongue that is not Cmz," Mexico. Males to 75 mm, females to 89 attached at the front can be protruded well mm. On the Atlantic versant from southern beyond the tips of the jaws - an adaptation Texas, USA to northeastern Nicaragua; and on for feeding on subterranean insects (TRUEB the Pacific versant from the Rio Balsas (Mexico) to Costa Rica, sea level to above 700 & GANS1983). m elevation.
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 '
->".--
.
*
--- -
Fig. 720.Rhinophrpus dorsalis (2k m W Tegucigalpita, Honduras, 40 m). Photo: J. R. McCranie
----#-~..----
Rhinophrynus dorsalis
Fig. 721.Rhinophrynus dorsalis (near Tegucigalpita, 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 STUART1961,FOUQUETTE & ROSSMAN 1963,G o m 1966,NELSON & NICKERSON 1966,F O U Q U E ~ 1969,DUELLMAN 1971c,FOSTER& MCDIARMID 1983,TRUEB& GANS1983,GARRETT & BARKER 1987
Tadpoles
The Tadpoles of Central American Anurana Most species of Central American anurans gives an overview on the tadpole body and have fiee-living tadpoles; exceptions are tail morphology. The spiracle is the exit for the species in the families Craugastoridae, respiratory water and its number, location, Eleutherodactylidae,Hemiphractidae, and and morphological configuration varies Strabomantidae that have direct develop- among species. The vent tube is the postement without aquatic free-swimming tad- rior opening of the digestive tract and its poles (MCDIARMID & ALTIG 1999).Available position (medial versus dextral) is useful data on the tadpole morphology of the for tadpole identification. The oral disc conmajority of Central American anurans tains the keratinous and soft tissue strucmake possible species identificationsbased tures adjacent to the mouth. The anterior on tadpoles. Tadpoles usually remain (called "upper" by some authorstposterior longer and at greater individual densities (called "lower" by some authors) direction at the breeding site than the adults and, of the oral disc is the same as the head-tail therefore, are easier to detect than the direction. Oral discs can have emarginaadults. Thus, including tadpoles in region- tions (Fig. 725 left side) or not (Fig. 725 al amphibian inventories can increase the right side).The jaw sheaths of the beak are detectability threshold and time period of usually ornamented by serrations. Tooth the species' presence at the study site, rows consist of keratinous spicules and are especially in the so-called explosive breed- found anteriorly and posteriorly to the ers. However, tadpoles of several Central mouth. The labial tooth row formula American anurans remain unknown describes the number and arrangement of (Table 2). The species with unknown tad- the labial tooth rows. The numbers before poles, mostly bufonids, dendrobatids, and the forward slash indicate how many tooth hylids, are either rare or difficult to find, rows are on the anterior labium and the are explosive breeders with short larval numbers after the forward slash indicate periods, or have not been differentiated the number of tooth rows on the posterior from congeners (ALTIG1987).The following labium. Tooth rows are always counted in key is useful for specimens between an anterior to posterior direction. Tooth GOSNER (1960)Stages 25 and 40; generally, rows in the anterior labium are representStage 40 tadpoles are two to two and a half ed by a letter A followed by the respective times as long as those at Stage 25 (SAVAGErow number, whereas rows in the posterior 2002). In some few cases, it is not possible labium are represented with a letter P folto distinguish between the tadpoles of two lowed by its respective number. Thus, as species, and in these cases they are placed examples, the first tooth row on the anteriin a single portion of a couplet and sepa- or labium is termed A-1, and the second rated by "or." More than 30 ecomorphologi- row on the posterior labium is called P-2. cal guilds have been distinguished among The oral disc usually is bordered by rows of tadpoles (ALTIG & JOHNSTON 1989, SAVAGEmarginal papillae. The marginal papillae 2002). These larval adaptations reflect can occur in one (uniserial), two (biserial), mostly ecological differences related to the or several (multiserial) rows. Submarginal type of habitat (lentic versus lotic water), papillae are situated on the face of the oral the niche in the water column, and the disc, particularly common near the lateral feeding mode and food items (SAVAGEends of the lower tooth rows besides the jaw sheaths and the emarginations. Other 2002). abbreviations used in the following tadpole Terminology of tadpole morphology follows key are BL (body length) and BW (body MCDIARMID& AZTIG (1999). Figure 724 width).
Tadpoles
\
b.'urnbelliform
\
Vent tube
\ ventral fin
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 Drawings: A Schulze dorsal fin high,extending onto body).
anterior tooth rows
anterior ("upper')
emarginate disc
posterior tooth rt marginal papillae (in two rows)
posterior ("lower")
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 Crepidophryneguanacaste 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 & LYNCH1969 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 & BRODIE1992 O'NEILL& MENDELSON 2004 ALTIG 1987 MCCRANIE & WILSON2002 KOHLER et al. 2006 MENDELSON & MULCAHY 2010 O'NEILL& MENDELSON 2004 SAVAGE 1972, SAVAGE & DONNELLY 1992 MENDELSON et al. 2005 ALTIG1987 DUELLMAN 1970, 2001 MCCRANIE & WILSON2002 MCCRANIE & CASTENEDA 2006 SAVAGE 2002 ZWEIFEL 1967 & CAMPBELL 1992 DUELLMAN DUELLMAN & CAMPBELL 1992 2001 CAMPBELL et al. 2000, DUELLMAN TRUEB1971 R. IBAREZpers. comm. Oct. 2010
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 .......... Fig. 726. Oral flaps of a. Hypopachus barberi; ................................R b i n o d~ ~ d b. Gastrophryne usta. Drawings: M. Vesely 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 Fig. 727. Tadpole of Dendropsophus ebraccatus tail muscle ...................- e d (GuayacBn,CRARC, Costa Rica, 550 m). Photos: G. Kiihler 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; 13a Vent tube medial, in preservative, dorsal lateral surface of body brown, mottled with and lateral surfaces of body dark brown, pi&vantriS white ................ ~~e mottled with t a n and a white stripe b Caudal fins uniform brown; lateral surface extending from mouth posteriorly below of body uniform black .................................. eye along body; caudal fins pale with gray ..................................G@@Pkme elor black blotches (colorful in life; Fig. 727). .........................DenmOp~ophusebraaxtus 10a Labial tooth rows absent .......................11 b Vent tube dextral, in preservative, dorsal b Labial tooth rows present ......................16 and lateral surfaces of body black with a lla Tail terminating in a thin flagellum .....12 dorsolateral tan stripe; caudal fins black b Tail not terminating in a thin flagellum 15 with irregular tan vertical marks .......................DandropBophae subocalans 1% Posterior labium with papillae-like protuberances ................................................ 13 14a Postorbital dark stripe absent or extending onto tail; no dark spots on tail b Posterior labium without papillae-like pro..................Lhdlvpeophus PLI-QD~~*UB tuberances ...............................................14
Tadb Postorbital dark stripe restricted to body;
dark spots ........ Dandropophw &bod& 16a Byes dorsal; moderate oral disc, protuber-
ant labia, velum covering anterior jaw Iethmela picadai (inpart) sheath b Eyes lateral; large umbelliform oral disc (Figs.724b, 728) ...... dlilvtmnku~aiaflotatnr l6a Enlarged ventral sudorial disc present 17 b Enlarged ve'ntral sucturial disc absent . 18 178 Tail musculature black with contrasting 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 ............................ .......................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 Vent tube medid no papillae on upper disc ................... 22 margin .......................... 20a Labial tooth rows l/0 or 210; umbeIliform oral disc ................... eia n&aola b Labial tmth rowa l / l z m a l loral disc
...........
b.'
L
.
I I
............ ..................................
. . .
Fig. 728. Tadpole of Silverstoneia flotator (Guayach, CRARC, Costa Rica, 550 m). Photos: G'
..................................................................21 21a Labial tooth rows l/1 .................................. .......................Ietbmo4yla picadai (inpart) b Labial tooth rows Y2 ...h % ~ ~ a h pd la
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 later8.1margins of upper jaw sheath ..............24 24a Tail tip rounded; beak huge, occupying D a n d m h &WI most of oral disc 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 29 b Labial tooth rows 2/3 or greater 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 27a Oral disc completelybordered by one row of tiny papillae; oral disc without emargina-
I
A Fig. 729. Tadpole of Dendrobatespumilio (Maria Luisa, Limbn, Costa Rica). Photos: G.Kiihler
................................
......
.....
............
...........
..................................................................
Fig. 730. Tadpole of Denohhates auratus (Guaya&, CRARC,Costa Rica). Photos: G. Kiihler
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 (inpart) b Papillae restricted to lateral and ventral portions of oral disc; oral disc with small emargination on medial portion of posterior 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 saddles 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, onehalf 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; on usually one row of labial papillae present
.............................................................. 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, serrated edge with a broad, medial shallow indentation; labial papillae in one row ...... ...................................Rhaebo haematitim 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 serrated edge; labial papillae at least partially in two rows ............... Chaunus marinus 39a Submarginal papillae present only on postenor disc area .............hcilius periglenes b Submarginal papillae present only on lateral 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
33
2
b Dorsal caudal fin with brown flecks (Fig. 732b) .......................... p4VIIobatea lugthis 49a Two complete rows of labial papillae on osterior margin of disc; caudd .fin with spots ............. Allobdm Erown
)RiarnnaFnn
b One complete row of labial papillae on pastenor margin of disc; caudal fin without brown spots 60 Mla Body dark-brown or blaclr; tail lightly marked with brown ..................................... Calcmtet&wparrurmaaaie b Dorsal caudal fin without brown blotches
.........................................
.a. . . . . . . . . . . . . . . . . . . . . . . . . . .
Fig. 732. Tadpole of P6gUobates lugubh (EN. h a d l o , Cwta Rim, 10 m). Photos: G. K6hler 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~ciliwC u W w b Tail fins transparent, with scattered, small pigment granules and a few large pigment granules; spiracle walls of nearly equal length, aperture ovoid ,..:....
............................
.................................................................. 51 5la Body tan, tail lightly marked wit21 brown; no scattered very large white guanophores on body and tail ...... Calo~te#iw&ib 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 65a Gap in papillae of posterior labium ...........
................................l?kmdbaXIlatcihgDpe b No gap in papillae of posterior labium . 5 4 ....................... e A-2 complete or with narrow gap ......... 66 ...............................~ c a k u / 3 ~ t a64a b Median gap in A-2 broad, longer than, or 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 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 syst e n ................................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 (inpart) 46a Spiracle ventrolateral ...........................47 b Spiracle lateral, located near midline of 49 side of body 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
...............................
...............
Dandzvbateaminu*
nearly as long as each lateral segment. 64 56a Labial papillae complete across anterior labium; oral disc greatly expanded; deep pictp folds lateral to mouth lkth~~&h b Gap in papillae of anterior labium; oral disc unexpanded; no folds lateral to mouth ~ p 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 col57 oration not reddish in life b Spiracle sinistral, but located more posteriorly, at 5341% of body length toward vent; body greatly depressed; eyes usually arescent-shaped; A-2 tooth mw segments very short, situated at lateral edges of jaw sheaths; overall coloration reddish in life (Fig. 734s) ............................................. 65 M a 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 ............. fndwh&d& 61a AU labial tooth rows complete ....................
Fig. 733. Oral disc of a. Leptodactylus melanonotns; b. L. &agilis. Drawings: M.Vesely
............................ 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 p a e d d w b A narrow median gap in P-1 present ........ ................................... lkphh&vIwibmw 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&lw8 & ~ 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 66a Spiracle located at 7281% of body length toward vent (at 71-79% in hatchlings) ..66 b Spiracle located a t 6345% of body length toward vent (at 51-71% in hatchlings) ...74 66a All parts of the sinus hyobranchialis conspicuously voluminous, its wings not separated medially, dominating the ventral buccopharyngeal body part ............................... ...................Taratolvlapulvarata (inpart) b Sinus hyobranchialis narrow and its wings broadly separated medially ................... 67 67a Dorsum with strong pigmentation, dark with broad (BL:BW = mostly c2.0) .......................... 68 b Dors- sparsely pigmented; body slender (BL:BW mostlp 2.0) .............................. 71 68a Interorbital distance narrow, ratio interorbital distance / oral disc width c0.45 (Fig. 736a) ................................. S a c h W i k b Interorbital distance broader, ratio interorbitd distance / oral disc width >0.45 69
....
Fig. 734. Tadpole of H-obatrachim fleischmanni (Costa Rica). Photos: G. K6hler
*
KC,S!@&-*Nq?
n
*
g&-&&
Fig.735. Tadpole of U,thobates taYjoi (near La Fortuna, Alajuela, Costa Rica); note emargination (arrow) of oral disc. Photos: G. Kiihler
Tadpoles I.
69a Sinus hyobranchialis with well developed
Teratohyia poiverata
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 .....
C
d Cachranella euknernos
I
I
Fig. 736. Centrolenid tadpoles (dorsal view). Photos: H. Hoffmann
........................... Sachatarnia albomacalata b Body not conspicuously flat (ratio BW/BH ~1.40);lateral part of the buccopharyngeal ventnun without numerous pigment spots ..............E s - m M e p a n (inpart) 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 . 1 7body ; slender (ratio BL:BW ~ 2 . 0 ..... ) ................... Taratohpapulverata (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 (ninpart) b Lower jaw sheath with small, uniform serrations throughout; sinus hyobranchialis narrow plain or convex; r?tio B F B H usu: ally >1.35 @ d n o b a h c b -ml f 74a Oral disc with more than 60 papillae in two rows; center of upper jaw narrowly keratinized and flexible ...................................... .........~ o b a ~ ladcwioi u m(in part) 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 .................Cockanellagranulosa b Sinus hyobranchialis large but the two wings of the RDO are mostly separated medially (Fig. 7368 t2ochmellaeuHnemos
:-Ru
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
78a b
79a b 80a
b.
Dorsum with few isolated spots; oral disc emarginated or notched, side-lobes of the oral disc smoothly separated from the upper lip ......4yaZinobafxachiumchimpoi .................................. 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 Oral disc with continuous labial papillae (Fig. 737a-c) ........................................... 79 Oral disc with gap in anterior labial papillae (Fig. 744,748a-c) ...........................104 Labial tooth rows not extending laterally beyond level of beak margins ................80 Labial tooth rows extending laterally well beyond level of beak margins ................ 81 Labial tooth row A-1 shorter than A-2
e
Fig. 738. Tadpole of Cruziohyla calcarifer (Guay a c h , CRARC, Costa Rica). Photos: G. Kijhler
Fig. 739. Tadpoie of Duellmanony~anrfiocuLis (Soltis Center, Alajuela, Costa Rica, 460 m). Photos: G. Kohler .
I Fig. 740. Tadpole of Duellmanohyla uranochroa (Rancho Naturalista, near Platanillo, Costa 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 (btpart) 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 i d 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 d&& a
b Four to eight rows of papillae below poste-
I
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 posterior 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
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 -IP ctulg9&m b Jaw sheaths bearing blunt serraiiom .. 94 94a Nostrils about midway between eyes and tip of snout ......PI-la guaW& b Nostrils about one-third distance &om eyes pokamclhi to tip of snout ......... J%&V&h 95a Caudal 5patternless; snout truncate
....
a. . . . . . . . . . . . . . . . . . . . . . . . . . .
..................................... 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~izedto those 'in labial rows; size
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~ h t 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 Fig.741. Tadpoles of Plectrohjla dasypus (P.N. b Oral disc with a single row of labial Cusuco, Corths, Honduras, 1370 m). Photo: 3. Kolby papillae ...................................................97
...........................
Tadpoles
b
98a b 99a
moderately small, total length <40 mm in Stage 36 ...........................Smilisca d d a 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 P-1, P-2, and P-3 tooth rows subequal in 99 length ..................................................... P-3 tooth row slightly shorter than P-1 and P-2 tooth rows ..............................101 Lateral processes of upper jaw sheath absent; iris coppery red in life ....................
a.
....................................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&claavia 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 ~ sagmum h weakly developed ... R b Lateral processes on upper jaw sheath absent ......................... ...................103 Fig. 742. Tadpoles of a. Agalyehnis annae;b. A. 103a Numerous small submarginal papillae spurrelli; c. A. lemur, d. A. cailidryas (a. from lateral to jaw sheaths ................................ Santo Domingo de Heredia, Costa Rica, ............................PIei9mhy.a tecmumani 1180 m; b . 4 . from Guayach, CRARC, Costa Photos: G.K i e r b No more than two to four large submar- Rica, 550 m). ginal papillae lateral to jaw sheaths ......... .............................PI-Ia g h d u l ~ ~ ~ 104a Gaps in anterior and posterior labial b Body with brown flecks or mottling (Fig. papillae; P-3 tooth row mounted on short, 742d) ..................... &&hz& vertically movable extension .................... .......................................S c i n a r b o u l ~ . lO9a Body a t least 15% deeper than wide; tail much deeper than body ............................ b No gaps in posterior labial papillae; P-3 .....................................Ag&ch& m a e tooth row not on extension ................ 106 b Body only slightly deeper than wide; tail 106a Spiracle ventrolateral ........................106 equal to or slightly deeper than body 110 b Spiracle lateral ...................................112 llOa Dorsal tail h not reaching to body (Fig. 106a Tail fins high, dorsal &d ventral fins 742b); well-developed lateral line system about subequal in height (Fig. 742a-c) 107 ................................. -epurralli b Tail fins lower, dorsal fin lower than venb Dorsal tail fin extending to body (Fig. tral fm (Fig. 742d) ..............................109 742c);lateral line system indistinct ... 111 107a P-3 tooth row with a median gap l l l a Tail fins transparent with scattered gray ...................................AgallPchnie saltator flecks; dorsal tail fin only slightly lower b P-3 tooth row complete .....................108 than ventral h ..... Agdjdmk m d e t i i 108a Body without brown flecks or mottling b Tail &SI transparent without scattered ..............................By..ameduaa van& gray flecks .................. Agdydds lemur
. .
, I
ll7a A narrow gap in A-1 tooth row; no gaps in posterior b o t h 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 posterior 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 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 l21a Ventral surface of body paler than dorsal Fig. 743. Tadpole of Tlalocohyla loquax (Guayach, CRARC, Costa Rica, 550 m). surface h 0 p z t O P ZmiOgmpanum t ~ . b Ventral and dorsal surfaces of body about Photos: G. Kijhler the same color .....................................122 122a Tail heavily flecked with brown; body uni1128 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. b Tail creamy tan with brown spots; body 743b) .......................... mmlalmottled bmwn and tan .............................. b No median gap in P-2 tooth row; P3 sube........................ 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 b P-3 tooth row subequal in length as or tail with high arch, considerably highonly shghtly shorter than P-1 and P-2 er than tail musculature a t midlength of tooth rows ........................................... 124 tail .......................................................114 124a Tip of tail pointed; gap in A-2 tooth row b Body as wide as, or, slightly wider than broad, over 70% length of either lateral high; dorsal tail fin without high arch, only segment ........................ hdbca baadinii slightly higher than, or same depth as, tail musculature at midlength of tail 117 114a Conspicuous dark dorsolateral stripe on body extending through eye ..................... 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
I
........
mw
a......................................
.........
..
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
Fig. 745. Tadpole of Isthmohyla pseudopuma (Albergue Montemeal, Alajuela, Costa Rica, 1820 m'l Photos: G. Kihler
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- Fig. 746. Tadpole of Isthmohyla graceae (Cemo shaped mark dorsolaterally on posterior part of body ................ wprianpeta~atue Sagui, Ngobe Bugl6, Panama, 1710 m). Photo: A. Hertz 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 posterior part of body (Fig. 749) L&~~%cBpheoth 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 venulosue u~
Vent tube dextral, gap in anterior labial papillae present or absent; number of tooth rows variable; dark brown lateral
Fig. 747. Oral disc of Trachycephalus venulosus. Drawing: M. Vesely
Fig. 748. Oral disc of a. Agalychnis calliwas, 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 ...........
of S&m Fig. 749. &, CRARC, Costa Rica).
phaeota (GuayaPhotos: G. EUihler
Fig. 75u. adpo pole of Bromeliohyla bromeliacia (P.N. Cusuco, Co&s, Honduras, 1616 m). Photo: J. Kolby
........................... Ekemdanta EIlmiM b Three to five posterior tooth rows; oral disc smaller, less wide than body ...... 134 134a E v e posterior tooth rows; tooth rows long, much longer than width of beak, two rows of labial papillae ........................................ .....F t y c Z ~ wsaZvadozmsisor P. legled 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;lasdlmiab P-1 tooth row with a narrow median gap; belly uniformly dark gray; tail with dense brown flecking, givin brown appearance
Fig. 751. Tadpole of Hypsiboas dtelus. Drawing: H. Hoffinann
......................... &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
..........................
........................ . .........................B m m a E ~ l al u v m h m.
without flecking (Fig. 750)
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 ventrolateral and ventral portions of oral disc
.....
Fig. 752. Oral disc of Hypsibom rufitelus. Drawing: H. Hoffmann
.................................
Tadpoles
--
-
b Labial papillae in a single row on ventrolateral and ventral portions of oral disc 142 142a Submarginal papillae present on lateral disc area
........................-b
b Submarginal papillae absent ............. 143 143a Tail shorter, ratio taiI length l body Iength
~ 1 . ........................ 7 -baasb Tail longer; ratio tail length / body length >1.7 ... @ p d h mpa'tane or EL pqgnar 144a Oral disc with continuous labial papillae
rn Fig. 753. TSidpole of Ptychohyla hypomykter (P.N. Cusuco, Cortks, Honduras, 1370 m). Photo: J. Kolby
.............................................................. 146 lae ........................................................152 146a Four anterior tooth rows (A-1 can be fi-agmented) ...............................................146 b Five to seven anterior tooth rows ..... 149 b Oral disc with anterior gap in labial papil-
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 posteriorby 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 Fig. 754. Hyloscirtus palmen' (Guayadn, b Papillae on anterior margin reduced to CRARC, Costa Rica, 550 m): a,b. Tadpole; Photos: G. G h l e r 151 c. metamorph. one row; labial tooth rows 5-717-10 150a Six anterior tooth rows, eight or nine posterior rows ........... b Five anterior tooth rows
..
...................................
..................................
........................
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 ................
......................................Lithobatasjuliani
..................
Four posterior tooth rows 155 166a ~ e d i a gap n in P-1 tooth row narrow, less than 10 teeth wide; labial teeth usually present in stages >30 ................................
b.
................................Lithobates maeuIatus
b Median gap in P-1 tooth row broader, more than 15 teeth wide; labial teeth absent (Fig. 7591 Lithobakw ~anizewitschii
A
1 Fig, 758. Tadpole of Lithobates vibicarius (neAlbergue Monterreal, Alajuela, Costa Rica, 1915 m). Photo: G. Kishler
Fig. 756. Tadpole of Litfiobates maculatus (P.N.
1
C&O,
C&S, Honduras, 520 m). Photo: J. Kolby
h M-I
&; -
Fig. 757. Metamorph of Lithobates vibicarius (near Albergue Monterreal, Alajuela, Costa Photo: G. Kijhler Rica, 1915 m).
Further Reading
STABRETT 1960, DUE& LYNCH1969, DUE1970, ~ T I G& BRANDON1971, ACTIG & CHRISTENSEN 1976, A ~ 1987, G DUELLMAN & CAMPBELL1992, MCDIARMID& ALTIG 1999, MENDELSONet al. 1999, DUELLMAN2001, M C C &~ WILSON 2002, SAVAGE 2002, Horn2005b, LYNCH 2006, HOFFMANN 2010, SANCHEZ 2010
rlg. .'(by.'mapole or urnonaces warszewztsmn (Soltis Center, Alajuela, Costa Rica, 460 m). 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 world- Furthermore, many of the reported wide. Hundreds of species are on the brink declines and disappearances have been of extinction or have already vanished labeled mysterious or enigmatic, because they took place in protected areas with no (POUNDS et al. 2006, WAKE & VREDENBURG 2008). Although declines of amphibians obvious changes in the environment such have been reported throughout at least the as habitat destruction (CRW et al. 1992, et al. 2006). last century (e.g., DICKERSON 1906, CORN& POUNDS FOGLEMAN 1984, KAGARISESHERMAN & MORTON1993), we are witnessing an Land use change obviously causes the alarming increase in regional declines and greatest anthropogenic threat to biodiveret al. 2006). Destruction, fragextinctions of frog and salamander species sity (GROOM since the late 1980s (RICHARDS et al. 1993, mentation, and severe modification of LIPS 1998, 1999, LIPS et al. 2004); popula- habitats through agricultural use, human tional data for the majority of caecilian settlements, logging, and burning are species are essentially unavailable and we, direct causes for species declines and therefore, have no information on the pop- extinctions. Fragmentation produces ulation trends in the species in this order smaller, more isolated populations that of amphibians (GOWER& WILKINSON 2005). have reduced options for dispersal and a
Fig. 760. Habitat degradation, destmction, contamination, and alteration are still the major cau-
ses for the loss of amphibian populations worldwide; Serrania de Tabasari, Ngobe Bugl6, Panama. Photo: G. Kiihler
rnoLo: m. r ranzen
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, medicines), 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 (LIPSet 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).
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 disap& pearance 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 species (40 percent of the anuran fauna) percent still being undescribed (PURm & had disappeared since 1987, including the HECTOR2000). This high percentage of golden toad (Incilius periglenes; Fig. 7611, undescribed species is highlighted by the Atelopus varius, eight hylids, and several 1980, results of a recent frog survey on Sri Lanka species of glass frogs (VANDEVENDER that resulted in the discovery of close to HAYES et al. 1989, POUNDSet al1997). 100 undescribed species, mostly of the et al. 2008); All Atelopus species have severely declined, genus PElautus (PETH~GODA a dramatic increase compared to the 50 and an estimated 67 percent of the known frog species recognized before these recent species of this bufonid genus have disappeared from their known localities (LA explorations (DUTTA& MANAMENDRAet al. 2005, POUNDS et al. 2006). ARACHCHI1996). However, many Sri WCA Lankan species known from 19th-century museum collections were not recorded in Although no long-term field study has ever & the recent surveys of natural habitats and been carried out on caecilians (Go2005), anectotal observations et WILKINSON are probably extinct (MEEOASWMBURA by M. W m in Costa Rica and Guatemala al. 2002). indicate that there are cases of caecilian For most areas of the world, baseline data population declines due to land use on the historical and current distribution changes (COLLINS& CRW 2009): a once or abundance of amphibian species are not high-density population of Dermophis available. Possible disappearances of popu- mexicanus that occurred at a coffee finca lations might have happened unnoticed in Guatemala that was been transformed and unreported because of the lack of prior from a shaded into an open growing planstudies. COLLINS& CRUMP (2009:45) tation resulted in a severe reduction in emphasize the importance to also publish caecilian density; in the time it would have results documenting stable or increasing taken M. Wake and colleagues "to find populations in order to not "paint a biased more than 100 individuals 30 years ago, & CRUMP 2009:20). they found 4." (COLLINS picture of worldwide changes." The evaluation of the numerous cases of Thanks to the efforts of the Global amphibian declines all over the world Amphibian Assessment, about 99 percent revealed several common traits: of the currently recognized amphibian species were evaluated; alarmingly, 1per- (1)Declines seem to affect mostly high elecent of those were considered extinct and vation species; this refers to elevations 32 percent threatened (IUCN categories above 500 m in Central America and above Critically Endangered, Endangered, and 1000 m in South America (YOUNG et al. Vulnerable); 23 percent of species were 2001, STUART et al. 2004, CRUMP 2005, LPS classified "Data Deficient" (STUART et al. et al. 2008). In many Neotropical species with a broader vertical distribution it was 2008). observed that these were declining at high In the late 1980s, awareness of the elevations, but populations of the same amphibian decline crisis xoalesced with species at lower elevations were not announcements of cases in Australia and (CRW 2005). 1994).To gain Central America (BLAUSTEIN a better understanding of what was going (2) Species that depended on water such as on, surveys of the amphibian species in streams or ponds for breeding were affectthese two regions were intensified ed most strongly by the declines and (RICHARDS et al. 1993). Fieldwork conduct- extinctions (LAURANCE et al. 1996a,b, LIPS ed in the early 1990s in the Monteverde 1998, STUARTet al. 2004). Also, streamregion of Costa Rica revealed that 20 frog 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, Eleutherodactylidae, and Strabomantidae) and plethodontid salamanders were not or were less affected (LIPSet 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& CRUMP2009). Therefore, widespread species tend to persist whereas species with very small ranges tend to become extinct when they are affected by declines (LIPSet al. 2003b, COOPER et al. 2008). In short, those s~eciesthat are streamassociated, 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 differentiate between normal population fluctuations and long-term trends? Frog populations are known to fluctuate widely in response to climatic factors. Over unfavorable 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 s H The difficulty of differfrogs ( ~ K ~2001). 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 credibility. BLAUSTEIN (1994:94) pointed out that "one must balance the risk of lost credibility, 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 unexplained cases of amphibian declines and extinctions occurring simultaneously on different continents could not be attributed to normal population fluctuations (BLAUSTEIN & WAKE1995, 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& CRUMP1994, 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 (POUNDSet 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 comparatively dry. POUNDSet 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 (2010) providconnection, ROHR& RAFFEL ed evidence for a link between climate change and Atelopus declines. Increased levels of ultraviolet B (UV-B) have been discussed as possible causes for et al. 1998, amphibian declines (BLAUSTEIN 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 (LINDERet 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 (LINDERet 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 (ROUSEet 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 circulating leukocytes in the frogs and, therefore, a pesticide-mediated immunosuppression was hypothesized (KIESECKER2002). 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 damage) have to be considered, too (LANNOO 2008).
Amphibian Declines
= Z T -+ -S S Z S
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
I
Fig. 763. Rediscovered: Isthmohyla debilis (CerroTugri, Ngobe BuglB, Panama, 1570 rn). Photo: A. Hertz
m
Fig. 764. Craugastormilesi was rediscovered at P.N. Cusuco, CortBs, Honduras, 1839 m. Photo: J. Kolby
Pesticides (insecticides, herbicides, fungicides) can cause a variety of effects in amphibian populations, ranging from feminization of males due to endocrine effects to mortality (HAYES et al. 2002). One of the most commonly used pesticides, atrazine, is a potent endocrine disruptor that can cause castration and feminization in male amphibians (HAYESet 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 (DAVIDSON & ~ A P 2007, P SPARLUVG et d. 2009). Wind-borne chemical deposition generally increases with elevation for
Amphibian Declines mountains a t middle and high latitudes (DALY& WANIA2005). Furthermore, an analysis carried out by HAGEMANet 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 & CRUMP2009).
Parasite infestations and infedious diseases occur in all animals. As seen above, trematodes can be a cause for morphological abnormalities in amphibians. And protozoa, 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).
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 et al. 2003, LIPS et al. Just a few years after the major amphibian declines (DASZAK die-offs in Australia, it was speculated that 2006): (1) the wave-like epidemiological a rapidly spreading epidemic disease phenology of the amphibian declines in might have caused the catastrophic Australia and Lower Central America declines (LAURANCE et al. 1996a,b).A major (LAURANCE et al. 1996a,b, LIPSet al. 2008); breakthrough in the research into the and (2)the low genetic variability observed et al. 2003). causes of the "mysterious" amphibian in Bd worldwide (MOREHOUSE declines was the isolation and description In Costa Rica and Panama, Bd has been of the putative causative pathogen, the isolated from dead or dying anurans in the chytrid fungus Batrachochytrium dendro- wild (LIPSet al. 2003a). batidis (Bd) (BERGER et al. 1998, LONGCORE et al. 1999, LIPSet al. 2003a). Bd is a basal LIPS et al. (2003b) analyzed the phenology fungal lineage in the Chytridiomycota of the cases of declining amphibians from which are distinct from other fungi in that four sites in Central America and identithey have a motile, flagellate zoospore fied ecological traits, with their "aquatic index" - which estimates the degree of (FISHER et al. 2009). associations with stream habitats over a At first, however, skepticism about the species' life span - being the most signifiactual role of this fungus was high cant factor associated with declines at all (BLAUSTEIN 1994):Is it a ubiquitous organ- four localities. Based on this analysis, they ism that has been present always in the predicted the likelihood of decline for amphibian environment worldwide? Is it species of the then unaffected site of El then only a secondary cause finally killing Cop6 in west-central Panama. The availthe amphibians after "something" has had able data also indicated a directional dis-
Amphibian Declines persal of the pathogen (Bd), similar to that et al. (1996a,b) hypothesized by LAURANCE in Australia. The epidemic wave was documented to move across Lower Central America at a rate of 15 to 43 kilometers per years (LIPSet 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).
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 physiological and ecological preferences of the fungus: Bd thrives best under typical cloud forest conditions: high humidity and temperature between 15" and 25" C (COLLINS & CRUMP2009). 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 An investigation of museum holdings world (LIPSet al. 2006, POUNDSet al. 20061, revealed that Bd has been present on frogs many unresolved issues and numerous collected in Puerto Rico at least since 1976 questions remain, and some authors argue (BURROWES et al. 20041, in Costa Rica at that the case for Bd as the primary agent least since the late 1970s or early 1980s has not yet been made conclusively (e.g., (PUSCHENDORF 2003, PUSCHENDORF et al. MCCALLUM 2005, POUNDSet al. 2006). 2006, LIPS et al. 20081, and on historical However, other investigators such as specimens of Xenopus laevis from Africa CYNTHIACAREYand KARENLIPS (among collected in the 1930s (WELDONet al. others), are convinced that no other factors 2004). Because Bd was prevalent as a sta- need to be invoked and "once the chytrid ble endemic infection in southern Africa infects a formerly chytrid-free population for 23 years before any positive specimen of toads, the toads die whether it's a warm was found outside Africa, it was proposed year, cold year, dry year, or wet year." that Africa is the origin of the amphibian (CAREYcited in COLLINS& CRUMP2009: chytrid fungus (WELDON et al. 2004). The 141). RYANet al. (2008) conducted a markavailable molecular data are all consistent recapture study of three subpopulations of with a single origin of Bd with subsequent the stream frog Craugastorpunctariolus a t regional and international spread (FISHER a mid-elevation site in central Panama between 1999 and 2005. Prior to the invaet al. 2009). sion of Bd, the subpopulations of C. puncThe regions with the highest suitability tariolus were large and showed a stable for Bd include habitats that contain the age-size distribution. Following detection world's most diverse amphibian faunas of Bd at this site in late September 2004, (RON2005). In Australia, Bd has infected all three subpopulations completely disap46 frog species of which 13 have declined peared from this site within two months. and 3 are considered to be extinct (COLLINS CRAWFORD et al. (2010) presented long& STORFER 2003). The wide range of condi- term field surveys and DNA barcode data tions under which Bd has been found and of an amphibian community in central its high detection rate in the Neotropics Panama; they documented a mass-mortalisuggests that Bd may already occupy a ty event caused by Bd resulting in the loss significant portion of the major natural of 41% of total amphibian lineage diversity, regions in Central and South America including five undescribed species. (RON2005).
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 progressive 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~irationor 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 (PESSIERet 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.
pathogen. The rediscovered individuals might be immune to Bd or at least able to & CRUMP tolerate its presence (COLLINS 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 outcompeting Bd (COLLINS &C R2009). ~ 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 witnessed 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 Reports on rediscovered frog species from major step toward this goal has been the both Central America (Figs. 762-764) and publication of the "Standard Methods for Australia have been published (BARRIO- Amphibians" (HEYERet al. 19941, a com~ ~ 0 2004, ~ 6 PUSCHENDORF s et a1 2005, pendium that recommends and defines LAMP0 et a1 2006, RODR~GUEZ-CONTRERAS et techniques for monitoring amphibian pop& CRUMP 2009, KOLBY & ulations including suggestions for field al. 2008, COLLINS MCCRANIE 2009, MCCRANIE et al. 2010). It design and statistical analyses. is not always clear whether these represent survivors of the disease outbreak Despite substantial achievements and (remnant populations), unaffected disjunct resolving some of the initial questions, populations in remote places, or new colo- much controversy remains and many nizations (LOTTERS et al. 2005). However, additional questions have been raised. We Bd is still present in these areas do not yet understand all the details of (RETALLICK et al. 2004, LOTTERS et al. 2005, this phenomenon, but a t least we now have explanations and hypotheses for the KOLBY & MCCRANIE 2009). "mysterious die-offs" of amphibians. In If Bd is still resent in the environment. addition, we have learned from the modhow can we explain the reappearances of ern amphibian extinctions that protected several frog species that have not been areas are no guarantee for the long-term seen for several (up to 25) years? The pep- survival of species. On the positive side tides in amphibian skin secretions are are reports of rediscoveries of "lost frog known to have antimicrobial properties species." Do these rediscoveries indicate (CLARKE 2007). Intraspecific variation in that the species involved are rebounding skin secretions can be high in frogs, and, in from the brink of extinction or that they the presence of a pathogen, a strong pres- are still in a period of decline (KOLBY & sure of natural selection will favor those MCCRANIE 2009)? Will evolutionary individuals that have peptide secretions processes such as mutations and natural that are more effective against the 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 multifactorial 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 distract us from research into other possible et causes. As was demonstrated by DASZAK 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.
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 declaration and management of nature reserves are still important components of any longterm 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 However, habitat degradation, destruction, alone are insufficient to address global contamination, and alteration are still the threats. Several possible solutions and conmajor causes for the loss of amphibian pop- servation tactics have been proposed in ulations worldwide. As BLAUSTEIN (1994) order to respond to the global decline of concluded, the amphibian decline problem amphibians (HOFFMAN et al. 2008). It must is part of the overall crisis in biodiversity. be emphasized that properly managed The world is steadily losing an unprece- biotic reserves still play a key role in longdented number of species in many differ- term conservation strategies. The emphaent groups of animals and plants (WILSON sis here lies on "properly managed" 1992). Ongoing deforestation, drainage because reserves that only exist on the and pollution of wetlands and other map are obviously useless. Currently, anthropogenic effects causes the alteration many protected areas in Central America and destruction of suitable habitat for lack adequate personnel and operational many organisms in all regions of the world, funding resulting in ongoing deforestation, including amphibians. For 90 percent of unregulated hunting, use of pesticides, and the amphibian species classified as threat- anthropogenic development (ACEVEDO et ened, habitat loss and degradation is a al. 2010, SUNYER & KOHLER 2010, WILSON major threat (HOFFMANet al. 2008). & TOWNSEND 2010). BECKERet al. (2007) have emphasized habitat split - the human-induced discon- For some species ex-situ programs might nection between habitats used by different be the only option to avoid extinction (e.g., life history stages of a species - as yet Atelopus zeteki and Nectophrynoides another important threat for amphibians. asperginis; ZIPPEL2002, KRAJICK 2006; Amphibians, including their larvae, play however, see criticism by POUNDSet al. an important role in energy flow and nutri- 2006). AmphibianArk, an alliance of conent recycling in all major terrestrial and servation organizations, has the goal to freshwater ecosystems worldwide. maintain in captivity breeding groups of Therefore, we have to address questions species that otherwise would become such as the ecological consequences of the extinct. The ultimate goal of these ex-situ disappearance of a substantial portion of 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 infrastructure 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 husbandry 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.
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 seemingly 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 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-
BLAUSTEIN 1994, HEYERet al. 1994, BLAUSTEIN & WAKE1995, LIPS1998,1999, SPARLING et al. 2000, YOUNG et al. 2001, SEMLITSCH 2003, LINDER et al. 2004, LIPSet al. 2003b, LIPS et al. 2004, STUARTet 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 amphib- (4) Normal population fluctuations should ians might be useful as bioindicators be relatively small so as not to mask because of their permeable skin and their declines resulting from environmental biphasic life style, leading to a kind of dou- problems on the one hand or to trigger ble jeopardy (BLAUSTEIN & W m 1995, false alarms on the other. However, frog KOHLER 2001). Disappearing frog popula- populations are known to fluctuate by tions might be interpreted as early warn- orders of magnitude in different years, and ing signals of environmental deterioration. the population dynamics of most species Although this concept was a great market- are unknown (COLLINS & CRUMP2009). ing tool and helped fund raising, it was Many amphibian populations have "boom eventually demonstrated that this hypoth- and bust" population cycles in which numesis was false (COLLINS & CRUMP 2009). To bers may vary by a factor of some 20 times be a good bioindicator, a group of species between successive years (SKELLYet al. should meet certain criteria, including the 1999). following ones (HILTY& MERENLENDER 2000, COLLINS & CRUMP 2009): (5) Changes in the group's abundance should occur quickly to the same environ(1)It should be readily accessible any time mental changes that might also influence when sampling might be done. However, the remaining biota and the ecosystem as most amphibians species are notoriously a whole. However, no evidence for this kind seasonal, and detectability is reduced over of relationship has been published. BOONE much of the annual cycle. & BRIDGES (2003) concluded that - depending on species studied - amphibians can be (2) Monitoring of the group should be either more or less sensitive to toxins than straightforward and cost-effective. other organisms. However, this is not the case for most amphibian species. Estimating an amphib- Consequently, there is no evidence to sugian species' population size is technically gest that amphibians are better bioindicators than other animals (BEEBEE& difficult and laborious. GRIFFITH2005, COLLINS& CRUMP2009). (3) The taxonomy and natural history of Any decline or disappearance of a n the species in the group should be fairly amphibian population is an indication that well known. However, for the majority of something is or was affecting that specific amphibian species life history data are population negatively; but this "somescarce andfor anecdotal, the taxonomy of thing" does not necessarily have an influmost major groups still is poorly under- ence on the remaining biota (PECHMANN & stood, and many species are known from WILBUR 1994). only a few individuals. For a large proportion of amphibians, the reproductive biolo- Further Reading & WILBUR 1994, HILTY & gy and activity periods are unknown. We PECHMANN 2000, BEEBEE& GRIFFITH lack basic data on habitat requirements for MERENLENDER & CRUMP 2009 numerous species that aggregate in aquat- 2005, COLLINS ic habitats for breeding but spend the rest of the year abroad in terrestrial environments.
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.
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 I appreciate the support I have received Lotzkat, James Randy McCranie, from forestry and wildlife officials i n Fernando Mendoza, Kirsten Nicholson, Central America. Collecting and exporta- Michael Obermeier, Marcos Ponce, Ardiel tion permits were provided by Z. Alain, Z. Quintana, Antonio Ramirez Vasquez, Edgar Arauz, Yariela Hidalgo, C. Medina, Fabian Schmidt, Robert Seipp, Jorge Soto, Abdiel Mendoza, Adalberto Montezuma Pablo Soto, Javier Sunyer, Joseph Vargas, M., Ricardo Rivera, and Aleida Salazar, and Jose Manuel Zolotoff. I am especially Autoridad Nacional de Ambiente (ANAM), thankful to these people for their valuable Panama City, Panama; Tito Perez, help. Many students and associates of the Congreso General Kuna, Panama City, department of herpetology a t Senckenberg Panama; Javier Guevara Sequeira, Rafael Museum, Frankfurt, Germany, have tested Gutierrez Rojas, and Marta Liliana the identification keys included in this Jimenez Fernandez, Ministerio de book. Their help is greatly appreciated. Recursos Naturales, Energia y Minas And I appreciate the support I have (MINAE), San Jose, Costa Rica; Milton G. received from my assistant a t SenckenCamacho, Mauricio Fonseca Cuevas, berg, Linda Acker. Carlos Peres-Roman, Carlos Rivas Leclair, and Sandra Tijerino Mejia, Ministerio del I am grateful to numerous colleagues for Ambiente y 10s Recursos Naturales photos of Central American reptile (MARENA), Managua, Nicaragua; Angel species-these photographers are acknowlBarahona, Karla Cantero, Ruben Castro edged in the captions of their respective Lobo, Tomas Garcia, Ana Patricia photos. All of the excellent pen and ink Martinez, Antonio Ortez Turcios, and drawings of anuran hands and feet are the Carlos Romero, Corporation Hondureiia de painstaking efforts of Milan Vesely, and Desarollo Forestal (COHDEFOR), Arne Schulze contributed illustrations of Tegucigalpa, Honduras; Patricia Quintana tadpoles. Carl Franklin, Andreas Hertz, Vda. De Lopez, and Andres Sanchez, Johannes Kohler, and Mason Ryan providMinisterio de Agricultura y Ganaderia, El ed unpublished data on maximum sizes of Salvador; J.I. Galvez Duiiionez, Oscar F. various amphibian species. Brian Kubicki,
I am particularly thankful to L a n y 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!
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.
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.
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 I owe a n immense debt of gratitude to Museum of Los Angeles County (LACMI, Andreas Hertz, Brian Kubicki, Sebastian Los Angeles; Douglas A. Rossman and Lotzkat, Joseph R. Mendelson 111, Mason Frank T. Burbrink, Museum of Natural Ryan, David B. Wake, Marvalee Wake, and Science, Louisiana State University Larry David Wilson, who patiently sup(LSUMZ), Baton Rouge; Jose Rosado and ported this project and read various porJames Hanken, Museum of Comparative tions of the manuscript; all individuals Zoology (MCZ), Harvard; Alain Dubois, provided valuable comments and advice Ivan Ineich and Annemarie Ohler, that significantly improved the text. Museum National d7Histoire Naturelle (MNHN), Paris; Eunice E. Echeverria, Special thanks are due to my father, Alfred Museo de Historia Natural de El Salvador A. Schmidt, for his constant support and (MUHNES), San Salvador; David B. Wake encouragement. and Carla Cicero, Museum of Vertebrate Zoology (MVZ), Berkeley; Franz Finally, I am especially grateful to my Tiedemann, Naturhistorisches Museum beloved wife Elke Kohler and daughter Wien (NMW), Wien; Janalee P. Caldwell, Isabelle for their emotional support, Amy D. Estep, and Laurie J. Vitt, Sam patience, and inspiring encouragement. Noble Oklahoma Museum of Natural Elke and Isabelle accompanied me i n the History (OMNH), Norman; James R. Dixon field and provided valuable assistance. and Katherine Vaughan, Texas Elke also merits special thanks for her Cooperative Wildlife Collection (TCWC), very careful editing of the manuscript. My College Station; Federico Bolafios and indebtedness to her cannot be exaggerated. 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,
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- Copeia 2005: 223-226.
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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 H a n a r d University Press. Cambridge, Massachusetts. WILSON,L.D. (1979): New departmental records for reptiles and amphibians from Honduras. - Herpetological Review 10 (1): 25. J.D. (2010): WILSON, L.D. & JOHNSON, 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. J.R. (1998): The bioWILSON,L.D. & MCCRANIE, geography 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 J.K., IBANEZ, R.. WILSON,L.D., MCCRANIE,J.R. & CRUZ-Dh,G.A. Yomc, B., LIPS, K.R., REASER, SALAS,A.W., CEDENO,J.R., COLOIMA, L.X. (1994):A new species of Plectrohyla (Anura: RON,S., LA W C A ,E., MEYER,J.R., Mu%-OZ. Hylidae) from a premontane rainforest in A., B O ~ O SI?.,, CHAVEZ,G. & ROMO,D. northern Honduras. - Proceedings of the (2001):Population declines and priorities for Biological Society of Washington 107 (1): amphibian conservation in Latin America. 67-78. Conservation Biology 15: 1213-1223. K.L. WILSON,L.D., MCCRANIE,J.R. & WILLIAMS, A., V. LE~N-REGAGNON & A. (1985): Two new species of fringe-limbed ZALD~VAR-RNER~N, NIETO-MONTES DE OCA.(2004): Phylogeny of hylid frogs from nuclear Middle America. the Mexican coastal leopard frogs of the Herpetologica 41 (2): 141-150. Rana berlandieri group based on mtDNA WILSON, L.D., MCCRANIE, J.R. & WILLIAMS, K.L. sequences. - Molecular Phylogenetics and (1994): Commentary on the reproductive Evolution 30: 3 8 4 9 . biology and systematic relationships of Hyla ZHANG,P. & WARE,M.H. (2009): A mitogenomic insolita MCCRANIE, WILSONand WILLIAMS. perspective on the phylogeny and biogeograCaribbean Journal of Science 30 (3-4): phy of living caecilians (Amphibia: 214-221. Gymnophiona). - Molecular Phylogenetics WILSON,L.D. & TOWNSEND, J.H. (2010):The herand Evolution 53: 4 7 9 4 9 1 petofauna of Mesoamerica: Biodiversity significance, conservations status, and future Z~MMERMANN,H. & ZIMMERMANN, E. (1988): Etho-Taxonomie und zoogeographische challenges. - Pp. 760-812. In WILSON, L.D., Artengruppenbildung bei Pfeilgiftfroschen TOWNSEND, J.H. & JOHNSON,J.D. (eds.): (Anura: Dendrobatidae). - Salamandra 24: Conservation of Mesoamerican Amphibians 125-160. and Reptiles. - Eagle Mountain Publishing, LC, Eagle Mountain, Utah. i-xviii + 1-812 ZIPPEL,K. (2002): Conserving the Panamanian PP. golden frog: Proyecto Rana Dorada. Herpetological Review 33 (1):11-12. WILSON, L.D., TOWNSEND, J.H. & JOHNSON, J.D. (eds.) (2010): Conservation of Mesoamerican ZUG, G. (1983): Bufo marinus (Sapo Grande, Amphibians and Reptiles. - Eagle Mountain Sapo, Giant Toad, Marine Toad. - In: JANZEN, Publishing, LC, Eagle Mountain, Utah. D.H. (ed.) Costa Rican Natural History i-xviii + 1-812 pp. Chicago & London (Univ. Chicago Press): 386-387. WYGODA, M. (1988): Adaptive control of water loss resistance in a n arboreal frog. - ZUG,G. & ZUG,P. (1979): The marine toad, Bufo Herpetologica 44 (2): 251-257. marinus: A natural history resum6 of native populations. - Smithsonian Contributions to WYGODA,M.L. (1989): Body temperature in arboreal and nonarboreal anuran amphibZoology 284: 1-58. ians: differential effects of a small change in ZWEIFEL, R. G. (1964a): Life History of water vapor density. - Journal of Thermal Phrynohyas venulosa (Salientia: Hylidae) in Biology. 14 (4): 239-242. Panama. - Copeia 1964 (1):201-208. WYNN,A. & HEYER,W.R. (2002 "2001"): Do geo- ZWEIFEL, R.G. (1964b): Distribution and life hisgraphically widespread species of tropical tory of the Central American frog Rana vibiamphibians exist? An estimate of genetic caria. - Copeia 1964 (2): 300-308. relatedness within the neotropical frog ZWEIFEL, R.G. (1986): A new genus and species Letpodactylus fuscus (Schneider 1799) of microhylid frog from the Cerro de la (Anura Leptodactylidae). - Tropical Zoology Neblina region of Venezuela and a discus14: 255-285. sion of relationships among New World YOUNG, B., SEDAGHATKISH, G., ROCA, E. & microhylid genera. - American Museum FUENMAYOR, Q. (1999): El Estatus de la Novitates 2863: 1-24. Conservacion de la Herpetofauna de R.G. (1967): Systematic status of the PanamB: Resumen del Primer Taller ZWEIFEL, Central American frog, Rana miadis. Internacional sobre l a Herpetofauna de Herpetologica 23: 54-56 Panama. - The Nature Conservancy. Arlington, Virginia.
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 helmet (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 surrounding 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 column 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-moving 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 recently 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 laping 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 different 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)anteromedian 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 texture 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 surface 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 external 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 resonating 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 animals
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,VASQUEZALMAZAN& 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). Bolitoglossa 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, 131 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 212f - spinosa 205,212f, 212,213 Ammobates 128 - nocturnus 128 A t e l o p ~ c u 97f s - 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 214 - 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 Cadta - iwokramae 27 Charadrahyla 215 - 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 Senckenberg 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-
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