the diversity of life

Tadpoles

THE BIOLOGY OF ANURAN LARVAE

Tadpoles

The Biology of Anuran Larvae

Edited by Roy W. McDiarmid

and Ronald Altig

“A brilliant compilation of information on

the world’s most common aquatic amphib-

ians."—Richard Wassersug, Copeia

“The present volume, a true magnum opus,

is a comprehensive account of larval anu-

ran biology and will no doubt become a

reference tome comparable with Duellman

and ‘Trueb’s classic on amphibian biology.

... Tadpoles is a substantial contribution to

the literature and a,;welcome addition to

my bookshelf. . . . [A] rich mine of infor-

mation to which many of us will be refer-

ring for years to come.”—Trevor Beebe,

Trends in Ecology and Evolution

458 pages 118 halftones, 181 line drawings,

23 tables

Paper $40.00

The Ecology of

Invasions by

Animals and Plants

Charles S. Elton

With a Foreword by Daniel Sinberloff

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Elton’s book is. The topic is even more

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most important cause of species extinction.

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of my students to read.” —Stuart Pimm,

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Paper $13.00

New from Chicago

In Search of the

Golden Frog

Marty Crump

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fascinating stories about birds, insects,

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them very readable. . . . The story of Marty

Crump’s life in the field is fascinating, and

told with respect for both the reader and

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320 pages 16 color plates, 60 halftones, 19 maps

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A Naturalist’s Guide

to the Tropics

Marco Lambertini

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an enlightening explanation of much to

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348 pages 11 color plates, 57 color photos,

21 maps, 76 line drawings

Paper $25.00

The University of Chicago Press

1427 East 60th Street, Chicago, IL 60637

www.press.uchicago.edu

FOUNDER AND EDITOR

Craig Hassapakis

Amphibian and Reptile Conservation

Modesto, California

Allison C. Alberts

Center for Reproduction of Endangered Species

Zoological Society of San Diego

Jonathan D. Ballou

National Zoological Park

Smithsonian Institution

Aaron M. Bauer

Department of Biology

Villanova University

Andrew R. Blaustein

Department of Zoology

Oregon State University

Harold G. Cogger

Australian Museum

Sydney, AUSTRALIA

C. Kenneth Dodd, Jr.

Biological Resources Division

U.S. Geological Survey

Lee A. Fitzgerald

Department of Wildlife and Fisheries Sciences

Texas A&M University

Julian C. Lee

Department of Biology

University of Miami

AMPHIBIAN ECOLOGY AND CONSERVATION

Jamie K. Reaser »)

U.S. Department of State, Bureau of Oceans and International

Environmental, and Scientific Affairs

ASSOCIATE EDITOR

Jack W. Sites, Jr.

Department of Zoology

Brigham Young University

ADVISORY BOARD

Joseph T. Collins

Natural History Museum

University of Kansas

Carl Gans

Adjunct Professor of Zoology

University of Texas at Austin

Roy W. McDiarmid

Herpetology, Biological Resources Division

U.S. Geological Survey

Russell A. Mittermeier

President, Conservation International

Washington, D.C.

EDITORIAL REVIEW BOARD

Harvey B. Lillywhite

Department of Zoology

University of Florida

Peter V. Lindeman

Department of Biology and Health Services

Edinboro University of Pennsylvania

Joseph C. Mitchell

Department of Biology

University of Richmond

Henry R. Mushinsky

Department of Biology

University of Florida

SECTION EDITORS

George B. Rabb

President, Chicago Zoological Society

Vice chair, Communications, Species Survival

Commission, The World Conservation Union

Hobart M. Smith

Department of Environmental, Population and

Organismic Biology

University of Colorado

Michael Soulé

Research Professor, University of California at

Santa Cruz

President, The Wildlands Project

Jaime E. Péfaur

Ecologia Animal, Facultad de Ciencias

Universidad de Los Andes, Mérida, VENEZUELA

Christopher J. Raxworthy

Department of Herpetology

American Museum of Natural History

Andrew T. Storfer

Wildlife Ecology and Conservation

University of Florida

Robert J. Wiese

Assistant Director of Animal Programs

Fort Worth Zoo

STATISTICS AND EXPERIMENTAL DESIGN

Anthony J. Krzysik

School of Arts and Sciences

Embry-Riddle Aeronautical University

Office of Ecology and Terrestrial Conservation, Washington, D.C.

COPY EDITOR

Mark L. Goodwin

COLUMNIST

Craig M. Hoover

Senior Program Officer, TRAFFIC North

America, World Wildlife Fund

Washington, D.C.

ARGENTINA

Maria E. Bridarolli

Instituto Argentino de Investigaciones de las Zonas

Aridas, Centro Regional de Investigaciones Cientificas

y Tecnoldogicas, Mendoza

GALAPAGOS ARCHIPELAGO (ECUADOR)

Marco Altamirano

Museum of Southwestern Biology, The University of New

Mexico and Charles Darwin Research Station

Galapagos, ECUADOR

INDIAAND MALAYSIA

Indraneil Das

Institute of Biodiversity and Environmental Conservation

Universiti Malaysia Sarawak, Sarawak, MALAYSIA

CONTRIBUTORS

GRAPHIC DESIGN

Kenneth W. Sholar

INDEXING AND ABSTRACTING

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Michael J. Tavares

WEBSITE DEVELOPMENT AND

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Jay G. Henry

COUNTRY LIAISONS

ITALY AND MADAGASCAR

Franco Andreone

Sezione di Zoologia, Museo Regionale di Scienze

Naturali, Torino, ITALY

INDONESIA

Darmawan Liswanto

Executive Director, Yayasan Titian

Jakarta

PERU

Antonio W. Salas

Museo de Historia Natural

Universidad Ricardo Palma

SOUTH AFRICA

Marius Burger

National Coordinator, South African Frog Atlas

Project, University of Cape Town

COMPUTER SUPPORT

Mark Warren

ZOO LIAISON

Chris Banks

Curator of Herpetofauna, Invertebrates & Education

Animals and Co-ordinator South East Asian

Conservation Programs, Melbourne Zoo, AUSTRALIA

TAIWAN

David McLeod

National Experimental High School at Science

Based Industrial Park, Huinchu

U.S. VIRGIN ISLANDS

Father Alejandro J. Sanchez Munoz

Pastor of Saint Anne's Parish

Saint Thomas

VENEZUELA

Jaime E. Péfaur

Ecologia Animal, Facultad de Ciencias

Universidad de Los Andes, Merida

photocopying for internal or personal use provided the appropriate fee is paid directly to the

(978) 750-8400; fax: (978) 750-4470; email: info(@copyright.com; website: www.copyright.com

- Editorial

WORDS FROM THE EDITOR—

With the completion of this issue,

we have successfully published our

second volume. Our mission is

further set, and I think it easy to

visualize our intended mission, based

on the observation of each new issue.

Amphibian and Reptile Con-

servation (ARC) now numbers over

700+ subscribers and has printed and

distributed 3,000 copies each of both

issues 2 and 3, thus making ARC the

largest distributed, English language,

herpetological scientific journal in

the world. I view my role with the

journal as “carrying the torch” as

editor and publisher, while many

interested in assisting the journal

have come “on board” as advisors or

editors, special section editors,

contributors, in-country liaisons, and

in other ways. I am grateful for all of

this assistance, and would like to

invite others with a special interest

or talent to contact me for more

information and discussion. I am

always open to feedback, new input, | x

and ideas regarding the journal and/

or the conservation of amphibians

and reptiles worldwide, as well as all

life. Again, I invite all to participate

in what I feel is one of the most important concerns for all

herpetologists, not just ones doing conservation related work. Now

I would like to briefly outline our progress thus far.

The idea for a journal devoted strictly to the conservation of

amphibians and reptiles evolved after I sadly left a brief stint at

dental school in 1991, much too early to ever practice. Though it

was a rather down time for me then (and not the last hurdle I would

have to overcome), I found that it would be several more years

before I would determine what my life’s work could be. After spending

about three years doing a lot of bookwork in the libraries of primarily

Utah after leaving dental school, I met a publisher, zoologist, and

a former classmate of mine [in my first graduate level class as a

student at Brigham Young University (BYU)], Jay Vilhena. It was

he that most directed my interest in herpetology into something

more concrete and credit should be kindly given. On a long road

trip to New Mexico, to deliver a computer system, I told Jay I

needed help with focusing my ideas of writing, research,

conservation, and herpetology. After a long and contemplated

discussion, as well as his background as a long-time friend and

associate, Jay announced, that I should publish a newsletter devoted

to my interests. This was 1994. After much thought, more time,

and finally, some strong persuasions of a spiritual nature, I agreed

to undertake the task. Eventually, I progressed the idea from a

newsletter, to a scientific journal, to lastly, what I now refer to as

amagazine styled, scientific journal. In October of 1996 we published

our first modest journal. Fifteen hundred copies of the first issue

z Oe it

Helping right a wrong. Our first day filming a documentary at

Hetch Hetchy reservoir (background) and holding one of our

extras, a bullsnake Pituophis melanoleucus, which wandered

across our trail. Reference: www.hetchhetchy.org

Authors

Amphibian and Reptile Conservation 2(2):40.

proved to be far too few as we quickly

ran out of stock in about February of

1997, as we continued to have a

strong interest in the journal. After

a move to the University of Arizona

in Tucson, Arizona from my alma

mater of BYU in Provo, Utah, in

the late fall of 1997 as the first snow

began to fall in Provo, the journal

and myself were exposed to a new

and exciting landscape. Work on the

journal was agonizingly slow, but I

continued to presevere and make

some progress. To help move the

journal forward, I again packed up

and moved to my hometown of

Modesto, California in the late

summer of 1999, to be closer to my

family and their influence. At each

step of the way, throughout the years,

I continued in good faith to steadily

work and make needed associations

to further the work on the journal,

though not all of these associations

were fruitful. I have left out many of

the gory details for lack of space

here, and/or left to some future

conference gathering (informally), I

can attest to the fact that it hasn’t

been an easy path to follow with many

uncertainties and setbacks. One

certainty is my unwavering commitment to continue forward

regardless of what must be overcome along the way. Further, as

conservation biologist Michael Soulé so accurately stated by telling

me in a telephone conversion, it “takes a lot of gumption” to start

a journal as I have done, and l-would add, even more to continue in

the face of adversity and uncertainty. However, with all struggles

and tribulations comes moments of great joy and satisfaction for

those who continue on. With the continued publication of this

journal I invite all to share in our success. It is my rock solid,

determined, and solemn devotion to continue on with the idea that

there should be a journal accessible by as many people as possible,

devoted to the conservation and preservation of amphibians and

reptiles, and their habitats worldwide. Words of my favorite musical

artist, whom I had the pleasure to meet just before I left Tucson,

comes to mind as he eloquently sang in the album, Late for the Sky,

and what I truly feel about my mission with the journal: “Keep a

fire burn’n in your eye... Dont let the uncertainty turn you around...

Go ahead and throw some seeds of your own and somewhere

between the time you arrive and the time you go, may lie a reason

you were alive.” (Jackson Browne 1974, select lines from the song,

For a Dancer). It is with the publication of this issue that we gain

momentum in contributing to the conservation of amphibians and

reptiles worldwide.

Craig Hassapakis

Founder, Editor, and Publisher

JAIME E. PEFAUR is Professor of Ecology at the Faculty of Science, University of Los Andes in Mérida, Venezuela. He

graduated as a Veterinarian from the University of Chile and received a Masters and Ph.D. (in 1979) degrees from the University

of Kansas. Dr. Péfaur has published over 90 papers dealing with taxonomy and ecology of South American vertebrates, as well as

edited several books. He was the Executive Secretary of the II Latin American Congress of Herpetology in 1990 and of the HI Latin

American Congress of Ecology in 1995. He was a member of the International Union for Conservation of Nature (IUCN)'s De-

clining Amphibian Populations Task Force Board of Directors.

JUAN A. RIVERO is Distinguished Professor at the University of Puerto Rico, Mayagiiez. He received his Ph.D. from Harvard

University in 1953. His dissertation was on the Salientia of Venezuela, which has become the classic text book for any herpetology

student of Latin America. He has published over 200 papers, mainly dealing with frogs of Venezuela, Colombia, and Ecuador. He

has also authored a book on the Herpetofauna of Puerto Rico.

CONTENTS Volume 2 Number 2 a

AMPHIBIAN & REPTILE

CONSERVATION

TNT ONAL AL DEVOTED 10. RLDWIDE PRESERVATION

ND.

i) RL

OB AMPHIBIAN AND REPTILIAN DIVERSITY

FOUNDER AND EDITOR

Craig Masapakis

ASSOCIATE EDITOR

Sack W. Sites. Jr

SPECIAL FEATURE ISSUE

VENEZUELA ty

reptiles

42 Distribution, species-richness, _.

endemism, and conservation of et

Venezuelan amphibians and 3

reptiles

Jaime E. Péfaur and Juan A. Rivero

Cover

The polkadot tree frog (Hyla punctata) is

DEPARTMENTS a common inhabitant of the largest

South American ecosystem, the

EDITORIAL Amazon, and it is also found within the

borders of Venezuela. It dwells in both

40 Words from the editor primary and secondary humid forests

. . and is common in marshy areas along

Craig Hassapakis river borders, lakes, Ana flooded

grasslands of lowland South America

from Venezuela and Trinidad south to

Authors the Paraguayan Chaco, east of the

Andes mountains. This treefrog

becomes almost entirely red at night.

Their diet consists of small inver-

tebrates. Photo kindly provided by Gail

Shumway.

Background screen photograph (see in color on page 69): A

view of the inside of San Eusebio cloud forest, in the Venezuelan

Andes. Tree ferns and bromelids are characteristics. Venezuela,

Mérida. Photo: Jaime E. Péfaur.

Scope: Amphibian and Reptile Conservation (ISSN: 1083-446X) [ARC] and the accompanying online edition (ISSN: 1525-9153) is a popularly accessible, peer-reviewed scientific journal of international

scope, which is devoted to the worldwide preservation and management of amphibian and reptilian diversity. Online edition: The full-text online edition is available to subscribers FREE-OF-CHARGE as

PDF (Portable Document Format) files through ingenta at: www. ingenta.com. The online edition may vary slightly from the print edition due to our reducing file sizes for efficient downloading over the Internet.

Some background screens (photographs) are removed which are deemed not essential to the content of the article(s). There is also some loss of clarity to photographs in reducing article file sizes to a minimum.

If clarity of photos is a problem, the print edition of the journal should be consulted. Audience: ARC is intended for a wide readership from nonprofessional to professional herpetologists, the general public,

and scientists. Frequency: ARC publishes two issues per year (semi-annually). Focus: ARC concentrates on publishing timely information in the form of feature articles, original papers and data, reviews,

reports, short communications, columns, commentaries, book reviews, editorials, and news and notes. Distribution: ARC is distributed worldwide by subscription as well as quality newsstands, bookstores,

and select vendors. Delivery is guaranteed. ARC is also available as pay-for-view full-text articles online with ingenta. Included and available online at ingenta are full abstracts and complete bibliographic

entries. Much of this same material plus more, such as FREE sample article(s), are available at the ARC website (www. herpetofauna.com). Publisher: Craig Hassapakis, Amphibian and Reptile Conservation

2525 Iowa Avenue. Modesto, California 95358-9467, USA. Fax: (509) 695-8747; email: publisher@herpetofauna.com; website: www. herpetofauna.com. Postmaster: Please send change of address to AR‘ }

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Distribution, species-richness, endemism, and

conservation of Venezuelan amphibians and reptiles

JAIME E. PEFAUR*2 AND JUAN A. RIVERO?

‘Ecologia Animal, Facultad de Ciencias, Universidad de Los Andes, Mérida, VENEZUELA *Departamento de Biologia, Universidad de

Puerto Rico, Mayagiiez, PUERTO RICO

Abstract.—This report assesses the macrodistribution of amphibian and reptile taxa, and compares the species-richness

of the various biogeographical zones in which Venezuela is herein divided. The macrodistribution of 252 amphibians and

299 reptiles species was established for the eight defined biogeographical regions. On the basis of the presence or absence

of species, these regions have been categorized into three units: a) those with fewer than 100 herpetological species

(Maracaibo Lake, Lara-Falcén, and the Islands), b) those with more than 100 but less than 200 species (Andes, Coastal

Range, Llanos, and Amazonas), and c) with more than 300 species (Guayana). An index of species-richness shows that some

regions of Venezuela are among the most distinguished herpetological places worldwide; the Andes region is the highest

with an index of 0.34. Endemics are given particular attention in the light of the current knowledge of the geographical

distribution of every taxon. There are 181 amphibian and 119 reptile species endemic to Venezuela. In terms of conserva-

tion, turtles and crocodiles are the most threatened species, but some frogs, particularly those of the highlands, are also

endangered. The existence of protected areas, together with management and scientific projects designed to protect

Venezuela’s amphibians and reptiles, and the increasing international concern for them, strengthen the hope of preserv-

ing its valuable herpetofauna populations, and other animal resources, for future generations.

Resumen.—La macrodistribucién de 252 especies de anfibios y 299 especies de reptiles fue establecida para las ocho

regiones biogeograficas en las que Venezuela se divide corrientemente. Sobre la base de la presencia o ausencia de

especies las regiones se han subdividido en tres unidades: a) las que tienen menos de 100 especies (Lago de Maracaibo,

Lara-Falcén y las Islas), b) las que tienen mas de 100 pero menos de 200 especies (Andes, Cordillera de la Costa, Llanos, y

Amazonas), y c) con mas de 300 especies (Guayana). El indice de riqueza de especies demuestra que algunas de las regiones

de Venezuela estan entre las regiones herpetologicas mas notorias en el Ambito mundial; particularmente Los Andes

sobresalen con un indice de 0.34. Se presta una atencion particular a las especies endémicas a la luz del actual conocimiento

de la distribucién geografica de cada taxén. Hay 181 especies de anfibios y 119 especies de reptiles consideradas como

endémicas en Venezuela. En términos de conservacion, las tortugas y los cocodrilos son los grupos mas amenazados, pero

algunos anfibios, particularmente los de alturas, también estan en peligro debido a la declinacién numérica de sus

poblaciones. La existencia de areas protegidas, unido a proyectos cientificos y de manejo disefados para proteger las

especies amenazadas, y el aumento de la preocupaci6n internacional, fortalecen la esperanza de que estos recursos

faunisticos puedan preservarse para futuras generaciones.

Key words. Checklist, distribution, species-richness, endemism, conservation, Venezuela, amphibians, reptiles, herpetofauna

Introduction sula, and along the coast, as well. The terrestrial boundaries of

The great variety of ecosystems in South America has induced Venezuela are continuous with Colombia on the west and south-

the development of a greatly diversified herpetofauna which is west, Guyana on the east, and Brazil on the south. The existence

about a fourth of the total number of amphibian and reptile of many large rivers and mountain ranges help make the country

species in the world. an environmental mosaic with a diverse fauna of amphibians and

Venezuela is one of the 13 countries comprising South reptiles (Fig. 1).

America. It lies in its northernmost portion and receives the This report assesses the macrodistribution of amphibian

climatic influences of the Caribbean Sea and Atlantic Ocean. In and reptile taxa, and compares the species-richness and the

the north and northwest it is encased by the Andean Cordillera faunistical similarity of the various biogeographical zone:

which in the Mérida Andes may attain a height of 5000 m. The which the country is divided. Endemics are given particular 2

extensive savannas in the central portion and the Amazonian tention in the light of the current knowledge of the geographical

forest of the south, contribute to a complex climate with an distribution of every taxon. Remarks on the conservation o1

abundance of rainfall throughout most of the country, although these faunas are discussed.

there are drier areas in the region of Falcon, the Goajira Penin- Efforts have been made by several researchers to unravel

the macrodistributional patterns of the diverse Venezuelan ver-

*Correspondence. Fax: (58) (74) 401286; email: pefaur@ tebrate groups. The distributional pattern of fishes were de-

ciens.ula.ve scribed by Mago-Leccia (1970); of amphibians by Rivero (1961,

VENEZUELA

twice the size of California). It lies in the.

ided into 24 states (political divisions), and’

water bodies exist. The climate is tropical, hot, and hursid in the

ng the coast. Venezuela is one of the main producers of oil in the

Is, s, hydropower and agricultural g goods, such as sugar cane, coffee,

ten countries in regards to biodiver: sity worldwide

fresh water fishes, about 250 species

th about 20,000,000 inhabitants and.

roblems, such as soil erosion and

concern are mining operations

station, urban and industrial pollution,

rship are not clarified in law, and continued

nomy thus, is of great environmental

conservation by the government as

vilderness intact and more than.

Figure 1. Relief map of Venezuela. The density of the stippled areas shows land elevations. Only the larger rivers are shown.

1963a, b, c, 1964a, b, c), Duellman (1988), Frost (1985), and

La Marca (1992); of reptiles by Roze (1966), Medem (1981,

1983), Pritchard and Trebbau (1984), and Lancini (1986);

and of mammals by Eisenberg and Redford (1979) and Bodini

and Pérez-Hernandez (1985). Some comprehensive studies

on the herpetofauna from several areas of Venezuela have

been provided by Staton and Dixon (1977), Duellman (1979),

Hoogmoed (1979), Hoogmoed and Gorzula (1979), Rivero-

Blanco and Dixon (1979), Péfaur and Diaz de Pascual (1982),

Durant and Diaz (1996), and Yustiz (1996). Many other

contributions about the distribution of orders (1.e., Brame

and Wake 1963), families (1.e., Dixon and Hendricks 1979),

genera (i.e., Dixon 1980; Di Bernardo 1992; Péfaur 1993;

Sefiaris et al. 1994), and/or species (1.e., Gallardo 1965, 1969;

Dixon and Michaud, 1992) have been consulted. Additional

references can be checked in Vanzolini (1978), Duellman and

Trueb (1986), La Marca (1992), Péfaur (1992), and Duellman

(1995), among others.

VENEZUELAN AMPHIBIANS AND REPTILES

Figure 2. Sketch map of Venezuela, showing the main biogeographical regions: |. Maracaibo Lake, II. Andes, Hl. Falcon-Lara, IV.

Coastal Range, V. Llanos, VI. Guayana, VII. Amazonas, and VIII. Islands.

Materials and methods

In order to assess the distributional records of each species, a

map of Venezuela with its herein defined biogeographical re-

gions was used. A biogeographic sketch of Venezuela is pre-

sented in Fig. 2. It contains the eight biogeographical zones

commonly accepted for the country. This physiographic en-

semble takes into account mainly the relief, climate, and vegeta-

tion (Marrero 1964; Ewel and Madriz 1968; Huber and Alarcon

1988). Although the boundaries of each region are not well de-

fined, especially in some ecological nodules, they serve our pur-

poses well. Different researchers have used this framework for

their studies (Rivero 1963, 1964; Eisenberg and Redford 1979;

Bisbal 1988; Péfaur and Rivero 1989).

The extent of each region is shown in Table 1. The land

bordering Maracaibo Lake corresponds to Region I, which is

covered by seasonally dry tropical forest in the north and by

tropical lowland rain forest in the south. Wetlands also cover an

extensive section of this region representing more than 3,500,000

ha. The southern extent of the Maracaibo Lake region is bor-

dered by Region II, corresponding to the elevated mountains of

the Cordillera de Mérida, and by the Cordillera de Perija, on the

northwest. Both ranges comprise the Andes region. For this

study, all lands above 500 m are considered within the Andean

unit and its biota as Andean; lands below 500 m are considered

as piedmont. The Andes are environmentally complex (Ewel

and Madriz 1968; Diaz et al. 1997), and may include premontane,

dry and humid forests, montane wet and cloud forests, xero-

phytic valleys, hot and dry landscapes, and the impressive

Paramos, highland tundras, occurring above 3000 m. This region

covers about 4,200,000 ha of the country’s surface.

The Falcon-Lara region or Region III also borders the

northeastern part of the Maracaibo Lake region. This land is

affected by the easterly drying winds of the Caribbean Sea and

exhibit mostly a xerophytic landscape, catalogued as

premontane dry shrub or dry forest. A large portion of the

north sealine of Venezuela is bordered by the Coastal Range

(Region IV), made up of forested mountains, with elevations

up to 2765 m, and covered by premontane tropical rain and

montane cloud forests. The Andes and the Coastal Range, slo]

down into the lowlands of the Llanos, Region V, which extends

to the Orinoco River in the south, to the border with Colombia

on the west, and extending to the Orinoco River delta in the

east. The approximately 27,000,000 ha of the Llanos are cov-

ered by savannas or extensive prairies intermingled with dry

Plate 3

Sinead . '

Plate 5 Plate 6

Plate captions: 2. Atelopus carbonerensis. Once very abundant, this bufonid frog is now extinct from the Andean cloud forests.

Venezuela, Mérida. 3. Bufo granulosus. An inhabitant of all Venezuelan biogeographical zones. Venezuela, Aragua. 4. Bufo marinus.

One of the largest toads from Venezuela and the one with the largest distribution; it remains abundant country wide. Venezuela,

Tachira. 5. Bufo typhonius. Venezuela, Guatopo. Photo courtesy of Laurie J. Vitt. 6. Dendrobates leucomelas. This attractive frog is

found in the southern part of the country, south of the Orinoco River. Venezuela, Bolivar.

VENEZUELAN AMPHIBIANS AND REPTILES

Table 1. Extent of land surface and number of life zones present in the biogeographical regions of Venezuela. (Source: Bisbal 1988;

Ewel and Madriz 1968.)

Estimated land surface

Biogeographical region km? % Number of Life Zones

I Maracaibo Lake 35,000 3.88 5

Il Andes 42,000 4.66 16

Hl Falcon-Lara 30,000 33.333) 6

IV Coastal Range 68,000 7.54 10

V Llanos 270,000 29.95 3

Vi Guayana 350,000 38.82 10

Vil Amazonas 105,000 11.65 Dy

Vill Islands 1,500 0.17

Total 901,500 100.00 22

Table 2. Surface of altitudinal belts in Venezuela. (Source: Ewel and Madriz 1968.)

Area

Altitudinal belt km? %

Tropical lowlands 640,283 71.30

Premontane 227,390 25.00

Low montane 27,987 3.05

Montane 4,570 0.52

Subalpine and alpine 1,270 0.13

Total 901,500 100.00

forests and riverine gallery forests.

The largest biogeographical region of Venezuela is the

Guayana, Region VI, which includes about 35,000,000 ha,

most of which are part of the Venezuelan Guayana shield.

The landscape is complex, including lowland, premontane

and montane wet and rain forests; savannas; and wet oases,

called morichales, rich in palms and herpetofauna. The land-

scape is defined by profound valleys bordered by fepuys,

table mountain remains of geological lands of the Precam-

brian-Paleozoic era. The area belonging to the Orinoco

River’s delta is also included in this region. Some 10,500,000

ha of tropical forests covering the lowlands of the upper

Orinoco River basin comprise Region VII, Amazonas, which

is continuous with South America’s large green core, the

Amazonia. An important part of this area is covered by wet

tropical forest.

A small portion of the country is made up of islands. All

of them.are included in Region VIII, which for the purpose of

the distribution of turtles also includes the Venezuelan Carib-

bean Sea. This region is included to call attention to the par-

ticular distribution of the island herpetofauna, especially that

of Isla Margarita, as well as maritime herpetofauna.

Each of these biogeographical regions is ecologically di-

verse and complex. In their ecological map, Ewel and Madriz

(1968) described 22 life zones for Venezuela (Table 1). The

most homogeneous biogeographical region, Amazonas, has

only two life zones, and the most complex of all, the Andes,

includes 16 life zones in its relatively small territory. Diver-

sity of ecosystems is proportional to the steepness of ver-

sants, the sides of the mountains, in the tropics, but most of

Venezuela is lowland. About three-fourths of the country is

comprised of lands below 500 m; lands above 1000 m do not

exceed 10% of the territory, yet these are the most ecologi-

cally diverse of all (Table 2), a situation with strong conse-

quences on the biological features of the country.

Several methods have been used to obtain the data pre-

sented in this paper: museum records, bibliographic informa-

tion, and personal observations. Reports made on the hold-

ings of relevant museums in Venezuela, Puerto Rico, Colom-

bia, Brazil, France, and the United States have been taken

into account, but in most instances they have been improved

with notes taken by the authors during their trips and mu-

seum visits. In order to update previous reports on the Ven-

ezuelan amphibians and reptiles (Péfaur 1992; Péfaur and

Rivero 1989), new lists had to be produced. To the best of

our knowledge the checklist database (Appendix |) contains

all known species up to December 31, 1996.

The distributional aspects of the list are given by the

presence or absence of data for every species in a biogeo-

graphical region. The eight regions were thus delimited by the

known information regarding the species’ geographical distri-

bution. The number of species present 1n a region is an indi-

cation of its species diversity and of the geological, climato-

logical, and biological evolution of the faunal elements con-

cerned. However, the index of species-richness, rather than

the simple number of species, is a better expression of evolu-

tionary trends. A Species-Richness Index (SRI) was calcu-

lated by considering the number of extant species divided b

the area multiplied by (x) 100. To obtain the faunistical sit

larity between regions, a four-fold contingency table wa

ated for every pair of compared regions and the Dice coeffi-

cient, as shown by Hayek (1994), was calculated.

Endemism, an ecological term, was determined by find-

ing whether there was a unique relationship between a spe-

cre-

JAIME E. PEFAUR AND JUAN A. RIVERO

Table 3. Taxonomic composition of the Venezuelan herpetofauna.

Class/Order Common name

Amphibia

Anura Frogs and toads

Caudata Salamanders

Gymnophiona Caecilians

Subtotal

Reptilia

Testudines Turtles and tortoises

Crocodylia Crocodiles and alligators

Amphisbaenia Amphisbaenians

Lacertilia Lizards

Serpentes Snakes

Subtotal

Total

Table 4. Allocation of amphibian and reptile species in the biogeographical regions of Venezuela.

Taxa I Il

Frogs 18 72

Salamanders 0 ]

Caecilians 2 2

Turtles 6 1

Crocodiles l 0

Amphisbaenians ] 2

Lizards 20 26

Snakes 44 38

Total 92 142

Families Genera Species

10 42 238

] 1 2

3 i] 12

14 50 252

fi 14 23

2 6

1 2 6

8 37 113

8 56 151

26 112 299

40 162 551

Biogeographical region nie

Hl IV Vv Vi Vil Vill Total

17 63 36 124) 53 5 238

0 ] 0 0 0 0 2

] 2 0 7 2 0 12

3 6 9 13 1] 6 23

] 2 2 3 5 ] 6

Ds 2 2 5 4 0 6

25 39 19 59 36 16 113

47 75 43 96 74 Oe 151

96 190 111 310 185 50 551

Table 5. Species-Richness Index (SRI) value calculations for herpetofaunal species in the biogeographical regions of Venezuela.

SRI = (Species number/area) x 100. The area to calculate the Index is provided in Table 1.

Biogeographic Amphibian species Reptile species Total Total

region n SRI SRI n SRI

I 20 0.05 72 0.20 92 0.26

ll 75 0.18 67 0.16 142 0.34

Hl 18 0.06 78 0.26 96 0.32

IV 66 0.10 124 0.18 190 0.28

V 36 0.01 75 0.03 111 0.04

Vi 134 0.04 176 0.05 310 0.09

Vil 55 0.05 130 0.12 185 0.18

Total 252 0.03 299 0.03 551 0.06

cies and a geographical region. As used in this study, when-

ever a species dwelled and apparently originated in a single

region, it was considered a biological endemic. However, if a

species was considered endemic because it occupied a single

region in Venezuela but also extended into another country, it

was considered as a political endemic with respect to the first

country. In the case of Venezuela, there are mainly biological

endemics, but there are also several political endemics, that

is, species extending beyond the boundaries to some neigh-

boring countries, such as Colombia, Brazil, and/or Guyana in

the mainland, or Trinidad-Tobago, and the Dutch Islands

(Bonaire, Aruba, Curag¢ao), in the Caribbean Sea.

To determine the conservation status of the herpe-

tofauna, only a few quantitative assessments are available.

Thus, a general impression rather than an accurate census

supports the cataloging for the species considered.

Plate 7 Plate 8

Plate 9 7 Plate 10

Plate 11 Plate 12

Plate 13 Plate 14

Plate captions: 7. Mannophryne collaris. The ventral side of the females of this species present a black collar and a yellow gular region.

Venezuela, Mérida. 8. Nephelobates alboguttatus. A very abundant species in the past, today it has disappeared from their geographical range.

Venezuela, Mérida. 9. Nephelobates haydeeae. This frog is found only in the western state of Tachira, where its populations are declining.

Venezuela, Tachira. 10. Nephelobates meridensis. This was the largest dendrobatid frog of western Venezuela. This species is probably

extinct. Venezuela, Mérida. 11. Nephelobates serranus. A very restricted frog, inhabiting only a part of the Sierra Nevada mountains. Venezuela,

Mérida. 12. Flectonotus pygmaeus. A marsupial frog that inhabits the northern mountains. Venezuela, Mérida. 13. Hyla crepitans. This species

dwells in most environments in the country. It remains very common. Probably conforms a taxonomical group of related species. Venezuela,

Tachira. 14. Hyla lanciformis. This large and slender brown frog has a wide distribution in the country. Venezuela, Tachira.

| =<

JAIME E. PEFAUR AND JUAN A. RIVERO

Table 6. Matrix of amphibian similarity among Venezuelan biogeographical regions. Note: This matrix shows in bold the amount of

species of every region. Above the diagonal line are the similarity values between a pair of regions, while under it are the numbers

of shared species. (Source of data: Appendix 1.)

REGION

I i il IV Vv vl vil

| 20 0.15 0.32 0.26 0.43 0.17 0.27

R i 7 5 0.15 0.18 0.18 0.11 0.15

E MT 6 7 18 0.33 0.33 0.14 0.22

4 IV i 13 14 66 0.45 0.25 0.28

o| Vv 12 i 9 24 36 0.36 0.46

N/} vi 13 12 in 25 31 134 0.40

Table 7. Matrix of reptile similarity among Venezuelan biogeographical regions. Note: See comments under Table 6. (Source of data:

Appendix 1.)

REGION

| Il ill IV Vv Vi Vil

I 72 0.47 0.64 0.56 0.52 0.30 0.32

R Il 33 67 0.51 0.44 0.37 0.42 0.24

= Hl 48 Si] 78 0.66 0.58 0.37 0.39

A IV 55 42 66 124 0.56 0.47 0.46

Oo V 38 26 44 56 75 0.56 0.62

N Vi 37 28 47 ial 65 176 0.72

vil 32 24 4] 59 59 110 130

Table 8. Number of endemic species and percentage of endemism in the herpetofauna of the biogeographic regions of Venezuela.

(Source Data: Appendix 1.)

Amphibians Reptiles me

Regions Species by region Endemics % Species by region Endemics %

| 20 4 DD: 2 12 10.1

Il 75 Sy7/ 31.5 67 20 16.8

Ml 18 0 0.0 78 5 4.2

IV A 66 82 17.7 124 19 16.0

Vv 36 l 0.6 75 2 tod.

Vi 134 75 41.4 176 48 40.3

Vil 55 12 6.6 130 13 10.9

Country’s total 252 181 60.3 299 119 39.7

Results toises have 23 species contained within seven families.

Crocodiles and alligators have six species belonging to two

Taxonomic composition families. The amphisbaenians are represented by only two

The herpetological component of the Venezuelan fauna con- genera and six species. The lizards have 113 species com-

sists of 551 species of which 252 are amphibians and 299 are prising eight families, with Gonatodes as its most specious

reptiles. A taxonomic list of species is provided by a country genus, with 13 species. Snakes make up the most diversi-

checklist (Appendix 1) and summarized in Table 3. The am- fied group with 151 species belonging to eight families. Its

phibians of Venezuela are included in three orders, of which largest family, the Colubridae, contains 104 species. All

the largest is Anura. The most specious family is Hylidae, other families of this class have less than 40 species each.

containing 77 species, followed by Leptodactylidae with 72 Its most specious genus is Afractus with 16 species; other

species. Other rather large families are Bufonidae, quite large genera are Liophis and Micrurus with 10 and 12

Centrolenidae, and Dendrobatidae. The rest of the frog families species, respectively.

have a very low number of species. The families of salamanders

and caecilians are also of minor number. Distribution

The reptiles are comprised within five orders of which The distribution of every species in the eight biogeographical

the largest are Serpentes and Lacertilia. Turtles and tor- regions, as considered in Appendix 1, is summarized in Table 4.

VENEZUELAN AMPHIBIANS AND REPTILES

By far, the largest assemblage of amphibians and reptile

species is found in Region VI, Guayana, followed by the Coastal

Range, Amazonas, and the Andes. The lowest number of spe-

cies in the country is found in the northern regions of Maracaibo

Lake and Falcon-Lara. In the maritime islands region there are 50

species of amphibians and reptiles.

Species-richness

As surface area differences among the biogeographical divisions

do not allow a direct comparison of the biodiversity by the

number of herpetofaunal species alone, an index (SRI) 1s re-

quired to more accurately express the results. The SRI index

values are shown in Table 5.

The country itself has a low value for the index, 0.06.

However, the four smallest regions achieve the highest SRI and

the opposite three largest regions have the lowest indexes.

Faunistic similarity

The herpetological similarity between the different biogeographi-

cal regions of Venezuela is presented in Table 6 and 7. Similarity

based on amphibian species 1s rather low (Table 6); the least pair

bound of regions is the Andes-Guayana, which share 12 species

and have a similarity value of 0.11; the strongest bound is the

Llanos-Amazonas pair, which have 21 species, with a similarity

value of 0.46, followed by the Llanos-Coastal Range and LI-

anos-Maracaibo Lake pairs. :

Regional similarity based on the reptile component of the

fauna produces different results (Table 7). In general, the simi-

larity region based on reptile species has higher values than

when amphibians are considered. Here, the least similar pair

region is the Andes-Amazonas, with a similarity value of 0.24,

while the strongest bounded pair is Guayana-Amazonas, with

110 shared species and a similarity value of 0.72.

Endemism

There are 181 species of amphibians and 119 of reptiles consid-

ered to be biological and/or political endemics (Table 8). Overall

there are more endemic species among the amphibians than among

reptiles. However, four regions have less endemic amphibians

than endemic reptiles. The Andes, the Coastal Range, and the

Guayana are the places with more endemics in both taxa.

Conservation

Actual data on the conservation status of the herpetofauna is

relatively scarce in Venezuela. The most relevant information on

population status is provided by Ramo (1982), Praderio (1985),

Silva et al. (1985), Péfaur and Diaz de Pascual (1987), Péfaur et

al. (1987), Silva and Valdéz (1989), La Marca and Reinthaler

(1991), Thorbjarnarson (1991), Péfaur and Pérez (1995), and

Durant and Diaz (1996), among others. The Red Data Book of

Venezuela (Rodriguez and Rojas-Suarez 1995) was helpful in

establishing herpetofauna endangerment status.

The report elaborated by the senior author to the Declining

Amphibian Populations Task Force (DAPTF) of The World

Conservation Union [IUCN] (in Vial and Saylor 1993), on the

declining status of amphibians was also used to generate a list of

endangered species (Appendix 2). Several species of amphib-

ians, among which there are seven species of Bufonidae (5

Atelopus, 2 Oreophrynella), four Centrolenidae (2 Centrolene, 2

Hyalinobatrachium), 15 Dendrobatidae (5 Colostethus, 2

Mannophryne, and 8 Nephelobates), five Hylidae (1 Gastrotheca,

4 Hyla), one Allophrynidae (1 Allophryne), seven

Leptodactylidae (2 Ceratophrys, 3 Eleutherodactylus, |

Leptodactylus, | Pseudopaludicola), and two Plethodontidae (2

Bolitoglossa) are reported as having a decline in their population

numbers and thus species of special concern. Among reptiles

there are several species on the verge of extinction, such as

Crocodylus intermedius, C. acutus, Caiman niger, and

Podocnemis expansa, as well as all the sea turtles.

Discussion

Taxonomic composition

The taxonomy of the Venezuelan herpetofauna changes con-

tinuously due to new research findings and systematic rear-

rangements. Substantial changes in the number of known spe-

cies for every region have taken place in Venezuela during

recent years. Descriptions of numerous new taxa have occurred

in the last decades mainly among amphibians (Péfaur 1985;

Rivero 1982a, b, 1985; Ayarzagiiena 1992, among others).

Important changes have recently been introduced to the tax-

onomy of the country’s herpetofauna thus, changing the sys-

tematic scenery. Among frogs, the Centrolenid family has been

divided into several genera (Centrolene, Cochranella, and

Hyalinobatrachium) by Ruiz-Carranza and Lynch (1991). The

Dendrobatidae was also generically rearranged with the intro-

duction of Epipedobates and Minyobates by Myers (1987),

Aromobates by Myers et al. (1991), and Mannophryne and

Nephelobates by La Marca (1995). The Hylidae has also suf-

fered some modification with the rearrangement of some

Ololygon into the resurrected genus Scinax (Duellman and Wiens

1992), and the elaboration of a new genus Tepuihyla by

Ayarzagtiena et al. (1992b). Among the caecilians, a general

taxonomic rearrangement of families and genera was proposed

by Nussbaum and Wilkinson (1989) and Wilkinson (1996).

Among the reptiles, there have also been some taxonomic

changes. Within the lizards, Iguanidae was divided into several

families (Corytophanidae, Iguanidae, Polychrotidae, and

Tropiduridae) by Frost and Etheridge (1989), while the genus

Anolis was divided into five genera, of which two are present in

Venezuela, Dactyloa and Norops (Savage and Guyer 1989). The

Teiidae was also divided into two units, Gymnophthalmidae

(small te1ids) and Teiidae [macroteiids] (Presch 1980). Within

the snake group, some colubrid genera have been revised modi-

fying the taxonomy for the Venezuelan members; for instance,

Dixon (1989) reviewed Liophis and other associated genera.

Among the Crotalidae there was also a strong change with the

partitioning of Bothrops into several genera: Bothriechis,

Bothriopsis, Bothrops, and Porthidium, with considerable effect

to the Venezuelan fauna (Campbell and Lamar 1989). Of course,

classification will continue to change as it is a dynamic science,

and new systems and names will be introduced changing subse-

quent lists of Venezuelan amphibian and reptile species. All

systematic allocations and changes should be taken as tempo-

rary arrangements that will be modified by the collection of new

data and insights by researchers.

Distribution

There is no single area in Venezuela where an amphibian an

areptile species is not present. From the most luxurious tropical

wet forest of Amazonas to the vegetationally depauperate

Paramos at the top of the Andean mountains, where numerous

Plate 16

eh ae eee Plate 18

Plate 19

Plate 21 Plate 2

Plate captions: 15. Hyla luteocellata. Venezuela, Guatopo. Photo courtesy of Janalee P. Caldwell. 16. Hyla microcephala. This small

frog has one of the largest biogeographical distribution in northern South America. Venezuela, Trujillo. 17. Hy/la vigilans. One of the

smallest frogs of the country; inhabits the Lago de Maracaibo Zone. Venezuela, Zulia. 18. Phyllomedusa trinitatis. Venezuela Guatopo.

Photo courtesy of Laurie J. Vitt. 19. Scinax rostratus. A medium sized frog with a distribution along the northern lowlands. Venezuela,

Mérida. 20. Eleutherodactylus lancinii. An inhabitant of the paramos cold streams. Venezuela, Mérida. 21. Eleutherodatylus lentiginosus.

A small frog from the Andes versants. Venezuela, Mérida. 22. Eleutherodactylus vanadise. A small frog occurring in the cloud forests

of the Venezuelan Andes. Venezuela, Mérida.

al ¢ : ~ 28h X

ue ee eee a

Plate 2

Plate 25 a 7 = Plate 26

Plate 27 Can oh le Plate 28

Plate 29 Plate 30

Plate captions: 23. Leptodactylus wagneri. A very elusive frog from most Andean environments, that can be detected by its ps r call.

Venezuela, Mérida. 24. Pleurodema brachyops. One of the most popular frogs in the country by its color and dots in the rear | the body.

Venezuela, Portuguesa. 25. Pipa pipa. This aquatic frog lives in the Llanos of Venezuela and Colombia. The picture shows a museum specimen

with eggs imbedded in its back. Venezuela, Barinas. 26. Rana palmipes. A typical frog from the lowlands of western Venezuela but that can

occasionally seen in the forested valleys of the Andes. Venezuela, Tachira. 27. Pseudis paradoxus. This medium sized frog is to be found in the

marshes of the country’s lowlands. Venezuela, Apure. 28. Bolitoglossa orestes. One of salamanders that dwells in the cloud forests of the

Andes. Venezuela, Mérida. 29. Caecilia subnigricans. A worm-like amphibian of northern Venezuela, that extends its range into Colombia.

Venezuela, Tachira. 30. Rhinoclemmys punctularia. This small turtle inhabits the Lake of Maracaibo region. Venezuela, Zulia.

JAIME E. PEFAUR AND JUAN A. RIVERO

frogs and lizards species dwell (Hoogmoed 1979; Rivero-Blanco

and Dixon 1979; Durant and Diaz 1996; Diaz et al. 1997),

species abound; however, snakes have not invaded the paramos

(Péfaur and Diaz de Pascual 1982; Diaz et al. 1997). Different

physiological and behavioral adjustments are found in the fauna

of each ecosystem, to cope with the contrasting climatological

factors impinging on every zone. Several of the more notewor-

thy adaptive strategies are the development of several unique

reproductive modes: developing eggs and tadpoles in tree cavi-

ties or bromeliads, or eggs on dorsum of females, or tadpoles

carried on dorsum of males, such as occur in anurans (Duellman

1985), or the use of collective nest deposits under rocks, as

occurs in the lizards of the Paramos.

Newly discovered and new locality records of species

have changed the known distributional patterns for many

amphibians and reptiles. For instance, in the Andean region,

species totals have changed from 16 frog, 4 lizard, and 10

snake species (as reported by Duellman 1979), to 56 frog and

15 lizard species (as reported by Péfaur and Diaz de Pascual

1982), to 72 frog, 26 lizard, and 38 snake species reported in

this study. The well-documented work by Lancini (1986)

reports 133 snake species for the country to 151 species

reported here. Our knowledge of the distribution of the Ven-

ezuelan amphibians and reptiles is quite acceptable at the

present time, but an increase could be expected as new data is

being collected all the time by researchers in the field. Both

the Andes and the Guayana regions have been actively ex-

plored by groups of researchers from the University of Los

Andes in Mérida and from Museo de Ciencias La Salle in

Caracas, respectively (Ayarzagtiena et al. 1992a, b; Péfaur

1993; Sefiaris et al. 1994; Durant and Diaz 1996), while ac-

tive research on the herpetofauna of the Coastal Range is

underway by researchers from Central University, Museum

of the Agrarian Zoology Institution in Maracay (Manzanilla

et al. 1995, 1996).

The distributional range of certain species is of concern.

There is a dramatic difference between the distribution of

some taxa extending over the whole country (e.g., Bufo

marinus, Hyla crepitans) compared to the punctual distribu-

tion of those tepui-associated taxa (e.g., Oreophrynella huberi,

O. vasquezi), or the narrow distribution of dendrobatids in

the Mérida Andes. Many examples of these are known in any

biogeographical region.

The fast change in the Venezuelan landscapes will have a

severe impact in the distribution of the herpetofauna. In less

than half a century most forests in Region I have been cut

down and replaced by prairies. Amphibian species prevail in

the south, while reptiles dominate the north. In general, most,

but mainly the xerophytic fauna, 1s shared with that of xeric

Region III, a relationship extending to the rich and large val-

ley of the Colombian Magdalena river. The valley of the

Catatumbo river might have played a role in acting as a pass-

way between the faunas of these regions.

The existence of many humid environments in the Andes

(Region II) permits the presence of a large number of frog

species (Duellman 1979; Péfaur and Diaz de Pascual 1982).

This region has few biogeographical contacts with other parts

of the country, although it has served as a passway for dis-

persing faunas (Péfaur and Pérez 1995; Rivero 1979; Rivero

and Solano 1977). Something similar occurs in the wet and

elevated lands of Region IV, the Coastal Range. The Andes do

not have within them any important large geographical bar-

rier, but there are rich microclimates that act as ecological

refuges. In the Coastal Range region, however, there are at

least two well differentiated kinds of lands: wet elevated and

xerophytic lowlands close to the Caribbean Sea (Rivero 1964a;

Manzanilla et al. 1995, 1996). Moreover, an important selec-

tive biogeographical barrier has acted in this region, as is the

case of the Unare river valley, where many herpetofauna spe-

cies distributions are discontinued past this point, such as

Colostethus mandelorum, L. insularum, and Eleutherodactylus

terraebolivaris, among others (Rivero 1964a; La Marca 1992).

In the Llanos (Region V) there exists a fairly large her-

petological fauna, which is common with the other regions

(Staton and Dixon 1977; Rivero-Blanco and Dixon 1979;

Péfaur and Diaz de Pascual 1987). The Llanos are climati-

cally and vegetationally homogeneous at the macrogeograph-

ical level, although they are intermingled with a web of ri-

parian forest communicating with the other surrounding re-

gions. Bordered by the Orinoco river in the south, this re-

gion does not separate faunistically from Regions VI and

ViI—the largest Venezuelan river seems not to be a selec-

tive biogeographical barrier for amphibians and reptiles

(Rivero 1961).

The southern regions of Venezuela, Guayana and

Amazonas (Regions VI and VII) are closer associated with

the Amazonian sector of South America than with the north-

ern areas of the country. The geological changes, the climate,

the topography, and the diverse vegetation make these two

areas the richest in herpetofauna species. This is especially

true of the Guayana where a multitude of habitats facilitate

the process of evolution. With the exception of salamanders

and crocodilians, the taxa are more numerous in the Guayana,

than in any other region. For the rest of the groups, the num-

ber is almost double in the Guayana, as compared to any of

the other biogeographical regions. The ecological complexity

of this region has played several roles in the evolution of the

biota. On the one hand, it has constituted a selective barrier

for the extension of some faunal elements from the Brazilian

Amazonian lowlands to the Llanos of Venezuela and vice

versa; on the other hand, it is the seat for many endemic

species (Hoogmoed 1979; this study). At the same time, it

has shared species with other regions of Venezuela and Brazil

(Avila-Pires 1995). One particular exception is Leptodactylus

labyrinthicus. This species lives north and south of this re-

gion, leaving a distributional hiatus in the Guayana shield

(Péfaur and Sierra 1995).

Depending on the total number of species present in

each region, three categories of regions can be proposed: a)

those with less than 100 species, comprised by the Maracaibo

Lake, the Lara-Falcon, and the Islands regions; b) those with

more than 100 but less than 200 species, which includes the

Andes, the Coastal Range, Llanos, and Amazonas; and c)

with more than 300 species, represented by the Guayana

region. Moreover, the limits of every biogeographical region,

are very imprecise. For instance, there is a sector where Re-

gions II, HI, and V converge, and thus there is an uncertainty

about the region to which some species belong. Another bio-

geographical nodule is the sector where Regions III and IV

merge. There is greater need for a more detailed analysis of

presence species in these conflicting sectors. On the other

hand, collecting records are very scarce in the Cordillera de

Perija in western Venezuela and the Orinoco delta region in

eastern Venezuela. It is possible that the Delta might be con-

VENEZUELAN AMPHIBIANS AND REPTILES

sidered a separate biogeographical region when more biologi-

cal information is gathered and analyzed.

Species-richness

The faunistic richness of an area would be better expressed by

an Index of Species-Richness (SRI) that takes into account

both the area and the number of species. In essence, this index

is a species-density index—that is, at the same number of

species, those biogeographical areas with small surfaces will

have higher values in SRI than areas with larger surfaces. SRI

numbers allow the comparison of any area or region of the

world with respect to species-richness. In the case of Venezu-

ela, its large size and the existence of extensive territories, such

as the Llanos, with a low number of species, account for a low

SRI value (0.06), similar to the one known for the Yucatan

Peninsula in Mexico (Lee 1980). This statistical artifact warns

about the use of such index for countries with a high diversity

of landscapes, but to use it for particular regions on small

countries. For instance, among published data, Oaxaca in Mexico

has an SRI value of 0.37 and is usually considered one of the

richest herpetological regions in the world (Casas-Andreu et

al. 1996), whereas Costa Rica has been reported as having the

largest SRI value (0.71) in the world (Johnson 1989).

If both amphibians and reptiles are analyzed together,

the highest SRI value is found in the Andes, followed by the

Falcon-Lara region. The SRI values of 0.34 and 0.32, respec-

tively (Table 8), are distinguished even when compared to the

SRI richest regions of the world. When the analysis is carried

out separately for amphibian species, the Andes is the only

region that stands out as the most diverse region. This rela-

tively small area, with several elevational belts and an abun-

dance of humidity and wet life zones, has a condensed den-

sity of amphibian species. In turn, when the reptile fauna is

analyzed separately, the largest value 1s found in Region III

(Falcon-Lara), which is also a small area covered with xero-

phytic vegetation—a fitting place for reptiles.

Faunistic similarity

Similarity is a concept that brings together biogeographical ele-

ments and evolutionary aspects of the fauna. In the comparison

of the seven regions of Venezuela, all obtained values for am-

phibians are under 0.50, indicating a low degree of species shared

between biogeographical regions. The most similar amphibian

faunas are those of Regions IV-V and V-VI]—the values of their

similarity indexes are the highest (0.45 and 0.46, respectively).

It is not a surprise to find these relationships, for there is a

geographical continuity between these regions, but what is

Table 9. Causes of amphibian population decline.

|. Environmental changes due to human activity

a) Destruction and/or fragmentation of habitats

b) Agriculture frontier expansion

c) Inadequate use of plaguicides

d) Water pollution

e) Ozone’s cover destruction or weakness

f) Acid rain

g) Introduction of predator and/or competitor species

h) Expansion and/or introduction of diseases

biogeographically relevant is the high similarity between the

Llanos and the Maracaibo Lake region, which are separated by

ranges and dry lands, but that share some species with disjunct

distributions such as Leptodactylus bolivianus, L. wagneri,

Physalaemus pustulosus, and Pseudis paradoxa, among others.

On the other hand, elements of Region II differ considerably

from the other areas, an indication of the greater amphibian

independence of the Andes with respect to the other regions in

Venezuela. A similar pattern is provided by the Guayana region,

which exhibits an independence as a biotic source.

Because reptile species usually have a rather ample distri-

bution, the number of shared species and the similarity values

are higher in this group than in the amphibians. The overall

similarity values are higher, especially between Regions VI and

VIL, and with the lowest level between the Andes and Amazonas.

Considering the reptiles, the Andes does not stands out as a

faunistically independent region because of this region’s rela-

tively strong relationships to the Maracaibo Lake and Falcon-

Lara regions.

Endemism

The formation of new animal species, as the result of genetic and

ecological processes, has been one of the major evolutionary

features in many areas of Venezuela. Wherever the distribution

of a species is reduced in space, an endemism process is at work.

Herpetological endemism is different in the several biogeo-

graphical zones of Venezuela. The Andes and the Guayana are

the regions with the highest percentages of amphibian endemics,

most of them biological endemics, reflecting an active speciation

process, something that has been acknowledged in other animal

groups (Brown et al. 1974). On one side, at the Andes there has

been an active process of contraction and expansion of forests as

consequence of the glaciation periods, and on the other side, at

the Guayana, there has been an appearance of different vegeta-

tional formations oriented by a long history of erosive changes.

In the Coastal Range almost half of the amphibian species are

biological endemics, too. The fact that most amphibian endemics

are found in elevated lands is evidence favoring the close rela-

tionship between abundance of life zones and diversity of am-

phibians. The Andes has 16 life zones, and both the Coastal

Range and the Guayana regions have 10 each.

The situation is different with the reptiles. Though there

are 119 species in the country considered as biological or politi-

cal endemics, only one region has a greatest number of endemics,

the Guayana, with 40% of their total species endemic. Reptiles

are less restricted to a geographical place and have a wider eco-

logical tolerance, thus extending their distributions into different

ll. Environmental changes due to astronomic factors

a) Global climatic changes

1) in temperature patterns

2) in precipitation and relative humidity patterns

b) Ultraviolet radiation increases

c) Not-yet-evaluated factors

1) Cosmic dust impact

2) Micrometeors

3) X-rays

4) Gamma-rays

Plate 31 a ) Plate 32

Plate 36

Plate 37 Plate 38

Plate captions: 31. Geochelone carbonaria. A common and popular turtle. Most peasants rise this species as food and pet.

Venezuela, Guarico. 32. Thecadactylus rapicaudus. A gekkonid lizard with an ample distribution in the country. Venezuela, Mérida. 33.

Norops nitens. Venezuela, Guatopo. Photo courtesy of Laurie J. Vitt. 34. Polychrus marmoratus. A chameleon-type lizard, common in

the wet forests. Venezuela, Aragua. 35. Anadia bitaeniata. A member of a taxonomically very complex group of lizards from the

Andean paramos and cloud forests. Venezuela, Mérida. 36. Gymnophthalmus speciosus. One of the smallest lizards; it lives in the litter

of most types of forests. Venezuela, Mérida. 37. Ameiva bifrontata. A medium sized lizard with an ample distribution in the lowlands.

Venezuela, Tachira. 38. Cnemidophorus lemniscatus. A ground dwelling lizard that probably has the largest distribution in all ecological

zones of the country. Venezuela, Zulia.

VENEZUELAN AMPHIBIANS AND REPTILES

regions and life zones within a region.

The Falcon-Lara and the Llanos have the smallest numbers

of endemics in the country. These areas are macroenvironmentally

homogenous, not favoring speciation as much as in heterogeneous

elevated lands with unstable environments. Both regions do not

have strong biogeographical barriers; this translates into a more

extensive sharing of herpetological species between bordering re-

gions. We hypothesize that any zoological taxa will have fewer

endemic species in the lands of Falcon-Lara or the Llanos of

Venezuela. The Maracaibo Lake Region is unusual, as the amphib-

ian endemic numbers are quite low compared to the reptile en-

demic species and can be explained by the extensive dry lands.

The finding of only a low number of amphibian endemics in

Amazonas is probably due to insufficient research in its territory.

Exemplified by the literature, animal groups exhibit more

endemics at elevated lands primarily due to isolation. For in-

stance, when the mammalian fauna, bats excluded, is studied, the

Andes region shows the largest percentage of endemic species of

terrestrial mammals (Eisenberg and Redford 1979). Supposedly,

any taxon closely associated with a geographical territory (as

opposed to a loose association, as in bats, birds, flying insects,

etc.) has experienced a high degree of speciation in the Andes,

mainly due to the glaciation processes during the Pleistocene

(Schubert and Vivas 1993) and the corresponding vegetational

responses of expansion and contractions (Duellman 1982). A

general review is needed, but the preliminary work by Diaz et al.

(1997) seems to be an indication that this actually occurs.

Conservation

Venezuela has a human population with a high degree of envi-

ronmental concern, but it is also a country with a rather strong

push for development. This contradictory situation has statisti-

cal expressions in the high and rapid destruction of natural envi-

ronments, with a rate of 600,000 ha of deforestation yearly and

the existence, at the present time, of 43 National Parks, two

Biosphere Reserves, and seven Wildlife Refuges (MARNR

1992). Such a great number of protected areas should help to

conserve the herpetofauna within their limits.

Unfortunately, actual study cases on the decline of animal

populations are few (Ojasti 1967; La Marca and Reinthaler 1991).

Few documented cases are known where herpetological species

have been known to be on the verge of extinction or have verifiable

population declines. Undoubtedly, the large reptiles have been the

most affected by an extractive commerce (e.g., collecting for the

skin trade, meat consumption) developed prior to the 1980s when

several regulations were established. Today strict regulations are

in action to protect crocodiles and turtles.

Amphibians are also imperiled, as can be derived from the

provided list of endangered species (Appendix 2). As in many

other places of the world (Blaustein and Wake 1990), a decima-

tion of amphibians population is occurring in the highlands of

Venezuela. The loss or reduction of amphibian species may

have some important ecological consequences, for they are

crucial to food chains. Thus, the energy linkage, as well the

predator and/or the prey populations would be altered. This

ecological biodiversity impoverishment would carry a loss of

genetic material of potential use as well as a waste of food and/

or pharmacological resources. The only pattern found among

most of the Venezuelan declining amphibian species is that a

high number are found at elevation. Why this occurs is not

totally understood but may be due to water contamination or

higher UV radiation levels at elevation, thus exhibiting vulner-

ability of amphibian species to this region particularly

(Blaustein et al. 1994). Many causes can be responsible for the

amphibian and other vertebrate’s decline, but most can be framed

into two kinds of environmental changes: human activities and

astronomic factors (Table 9; Péfaur 1993).

Many conservation problems are faced and intended to

be solved at the international level and at the national level by

different government offices. As a nation, Venezuela has par-

ticipated in several international agreements to protect the

environment and/or the fauna. Among the more important

ones are the Biodiversity Convention and The Amazonic Co-

operation Treaty, designed to protect large areas; and the

Interamerican Agreement for Protection and Conservation of

Sea Turtles, as well as the Freshwater Convention, which is

concerned with quality and quantity of the resource, its ba-

sins, the area drained by a river and its branches, and its

fauna. Projects to protect crocodiles (Gorzula 1985;

Thornbjarnarson 1991; Velasco and Ayarzagiiena 1995;

Baquero de Pedret and Quero de Pefia 1996; and many tech-

nical reports listed under Seijas 1993) or turtles (Licata et al.

1996) under the responsibility of the Venezuelan Ministry of

the Environment (MARNR) should help ensure proper pro-

tection and preservation of these animals. At several Venezu-

elan university laboratories, there are projects studying the

biological and ecological aspects of amphibian and reptile

species providing grounds to protect these valuable resources.

As a corollary, it is usually accepted wisdom that the better

we know a fauna and its associated ecosystems, the better

protection we can offer.

A list of imperiled species (this study), the existence of

protected areas, together with management and scientific projects

designed to protect Venezuela’s amphibians and reptiles, and

the increasing international concern for them, strengthen the

hope of preserving its valuable herpetofauna populations, and

other resources, for future generations.

Conclusions

The existence of elevated ranges, extensive forests, and a myriad of

rivers and other aquatic habitats, makes Venezuela an environmen-

tal mosaic where a diverse fauna of 252 amphibians species and 299

reptiles species live. Each of the eight current biogeographical zones

has a relatively numerous herpetofauna—the most diverse are the

Guayana and the Andes, with 310 and 142 species, respectively.

Due to the large area covered by certain territories, such as

the Llanos and Guayana, the obtained value for the country’s

Species-Richness Index is relatively low. However, the regions

situated in northern Venezuela appear high in the worldwide

species-richness ranking.

The similarity among the biogeographical regions ts rather

low when comparing shared amphibian species, but it is higher

when comparing reptiles. Selective biogeographical barriers work

mainly in the mountainous regions and are less effective in the

ecological continuous landscape of the rest of the country. The

higher amphibian similarity is found among the Llanos-Amazonas

regions, while for the reptiles is found among the Guayans-

Amazonas regions.

Endemism in amphibians develops more in elevate ds,

such as in the Guayana, the Andes, and the Coas‘a! Range.

There are less endemic reptiles than amphibians, due to their

greater ability to disperse. The area with more endemic reptiles

Plate 40

Plate 41 __ meat Plate 42

Plate 4300” ee Plate 44

Plate 45 eee an i feerG

Plate captions: 39. Tupinambis teguixin. The largest lizard of the country, is common in the Llanos and in the Lake Maracaibo zone.

Venezuela, Zulia. 40. Leptotyphlops affinis. This small ground dwelling snake, can be found in restricted parts of the Andean region.

Venezuela, Mérida. 41. Helminthophis flavoterminatus. This curious little snake distributes in many environments of the northwestern

states. Venezuela, Mérida. 42. Leptodeira annulata. This species is probably one of the most common snakes and the one with the

largest distribution in northern South America. Venezuela, Tachira. 43. Oxybelis fulgidus. This colored snake inhabits the southern

lowlands. Venezuela, Bolivar. 44. Phylodryas viridissimus. An attractive snake distributing in the southern states of the country.

Venezuela, Bolivar. 45. Micrurus mipartitus. Venezuela, Guatopo. Photo courtesy of Laurie J. Vitt. 46. Bothrops venezuelensis. A large

snake usually found in the forested environments of the northern mountains. Venezuela, Tachira.

JAIME E. PEFAUR AND JUAN A. RIVERO

is the Guayana region.

In terms of conservation, it is known that the more com-

mon environmental changes, destruction and fragmentation of

habitats and agricultural expansion, will have severe consequences

for the herpetofauna in Venezuela. As many species are endemics

and restricted to small areas, the destruction of a few kilometers

of the habitats could eliminate several species. The rapid de-

struction and/or contamination of natural environments are caus-

ing a decline in herpetofauna populations and numbers so as to

put some species at extinction risk. Most imperiled amphibians

are those living in the highlands of northern Venezuela, while the

most endangered reptiles are the marine turtles, crocodilians,

alligators, and turtles. It is hoped that the existence of several

National Parks and other protected areas, and the increasing

awareness of the citizens, would help to preserve the Venezu-

elan herpetofauna.

Acknowledgements.—We thank the people of the Animal

Ecology group at the Universidad de Los Andes and of the

Department of Biology at the Universidad de Puerto Rico,

Mayagiiez, for their permanent support to continue with our

herpetological studies in Venezuela. In particular we are in-

debted to William E. Duellman, James R. Dixon, Pedro Durant,

Nancy M. Sierra, Alberto Veloso, Jesus Manzanilla, Gustavo

Casas-Andreu, César Molina, and the late Adao J. Cardoso for

their help and continuous provision of data and references.

Thanks are extended to the curators of the visited museums,

especially to the Colecciédn de Vertebrados, Universidad de

Los Andes, Mérida, Venezuela; Estacidn Biolodgica Rancho

Grande del Ministerio del Ambiente y de los Recursos Natu-

rales, Maracay, Venezuela; Museo de Historia Natural La Salle,

Caracas, Venezuela; Museo de Biologia, Facultad de Ciencias,

Universidad Central de Venezuela, Caracas, Venezuela; Museo

de Zoologia Agricola, Universidad Central de Venezuela,

Maracay, Venezuela; Museo de Biologia, Universidad de Puerto

Rico, Mayagiiez, Puerto Rico; Instituto de Ciencias Naturales,

Universidad Nacional, Bogota, Colombia; Instituto de Biologia,

Universidade Estadual de Campinas, Sao Paulo, Brazil; Mu-

seum of Natural History, The University of Kansas, Lawrence,

Kansas, USA; Field Museum of Natural History, Chicago,

Illinois, USA; Department of Wildlife and Fisheries Sciences,

Texas A&M University, College Station, Texas, USA; Mu-

seum of Vertebrate Zoology, The University of California,

Berkeley, California, USA; Museum of Comparative Zoology,

Harvard University, Cambridge, USA; and Museum National

d’Histoire Naturelle, Paris, France. Comments and corrections

made by Jack Sites, William W. Lamar, Craig Hassapakis, and

two anonymous reviewers are deeply appreciated. Errors re-

main ours. Belkis Rivas and Yhilda Paredes were helpful in the

preparation of previous drafts. Marisela Angelino prepared

the final version of the manuscript. This study has been granted

by CDCHT-Universidad de Los Andes (C-805-96) and

CONICIT (PI-011).

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Manuscript received: 26-April-1997

Accepted: 15-January-1998

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Plate 48

Plate 51 mats Plate 52

Plate captions: 47. The Venezuelan Andes. The versants of the mountains are covered with lush forests, mainly of cloud type; at the

top, the paramo develops. The Bolivar Peak, covered by ice, is the highest point in the country. Venezuela, Mérida. 48. Coastal Range in

north-central Venezuela. The northern tradewinds dry up the sea-facing environments producing a semixeric or xeric ecosystem.

Venezuela, Aragua. 49. The Llanos of Venezuela is one of the largest and more homogeneous biomes of the county These flatlands

develop between the Andes and the Coastal Range to the north, and the Apure—Orinoco rivers to the south. Venezuela, Barinas. 50. In

the Venezuelan Guayana appears the tepuis, isolates tabletop mountains, that dominated the landscape. The lower lands are covered by

praliies, as the Gran Sabana, or by forests, as the largest part of the states of Bolivar and Amazonas. Venezuela, Bolivar. 51. A view of

he inside of San Eusebio cloud forest, in the Venezuelan Andes. Tree ferns and bromelids are characteristics. Venezuela, Mérida. 52.

Where dense clouds coming from the Caribbean Sea hit the mountains, a dense cloud forest appears at the upper parts of the Coastal

Range. One of the best known is Rancho Grande, on the road from Maracay to Ocumare. Venezuela, Aragua.

0} ejanzeue/ Wo ‘sep

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Asiatic Herpetological Research

Asiatic

Herpetological

Research

Volume 8 ® 1999

Editor

Ermi Zhao

Chengdu Institute of Biology, Academia Sinica. Chengdu, Sichuan, China

Associate Editors

Kellar Autumn

Lewis & Clark College. Portland, Oregon, USA

J. Robert Macey

Department of Biology, Washington University, St. Louis

Missouri, USA

Theodore J. Papenfuss

Museum of Vertebrate Zoology, University of California

Berkeley. California, USA

Editorial Board

Kraig Adler

Cornell University, Ithaca, New York, USA

Natalia B. Ananjeva

Zoological Institute, St. Petersburg. Russia

Steven C. Anderson

University of the Pacific. Stockton, California, USA

Aaron Bauer

Villanova University, Villanova, Pennsylvania, USA

Christopher Bell

University of Texas. Austin. Texas, USA

Leo Borkin

Zoological Institute, St. Petersburg, Russia

Bihui Chen

Anhui Normal University, Wubu. Anhui. China

I-jiunn Cheng

Institute of Marine Biology, National Taiwan Ocean University

Keelung, Taiwan. China

Ilya Darevsky

Zoological Institute, St. Petersburg, Russia

Indraneil Das

Madras Crocodile Bank, Vadanemmeli Perur. Madras, India

William E. Duellman

University of Kansas. Lawrence, Kansas. USA

Hajime Fukada

> He

Xiang Ji

Hangzhou Normal College, Hangzhou, Zhejiang, China

Pi-peng Li

Yantai Normal College. Yantai, Shandong. China

Ronald Marlow

University of Nevada, Las Vegas, Nevada, USA

Robert W. Murphy

Royal Ontario Museum, Toronto, Ontario, Canada

Goren Nilson

University of Gdteborg, Goteborg, Sweden

Nikolai Orlov

Zoological Institute. St. Petersburg, Russia

Hidetoshi Ota

Department of Biology, University of the Ryukyus. Nishihara

Okinawa. Japan

James F. Parham

University of California. Berkeley, Califormia, USA

Soheila Shafi

University of Shahid Bahonar, Kerman, Iran

i-tao Shi

Hainan Normal University, Haikou, Hainan, China

Xiu-ling Wang

Xinjiang Normal University. Urumqi. Xinjiang, China

Yue-zhao Wang

Sennyuji Sannaicho, Higashiyamaku, Kyoto, Japan Chengdu Institute of Biology, Academia Sinica, Chengdu,

Sichuan, China

Carl Gans Yehudah Werner

University of Michigan, Ann Arbor, Michigan, USA Hebrew University. Jerusalem. Israel!

Robert F. Inger Ken-tang Zhao

Field Museum, Chicago, Illinois, USA Suzhou Railway Teacher's College, Suzhou. Jiangsu China

Asiatic Herpetological Research is published by the Asiatic Herpetological Research Society (AHRS) and the Chinese So-

ciety for the Study of Amphibians and Repules (CSSAR) at the Museum of Vertebrate Zoology, University of California. The

editors encourage authors from all countries to submit articles concerning but not limited to Asian herpetology. All correspon-

dence outside of China and requests for subscription should be sent to AHR. Museum of Vertebrate Zoology, University of

California, Berkeley, California, USA 94720, or by email to asiaherp @uclink.berkeley.edu. All correspondence within Chi-

na should be sent to Ermi Zhao, Editor, Chengdu Institute of Biology, P-O. Box 416, Chengdu, Sichuan Province, China, Au-

thors should consult Guidelines for Manuscript Preparation and Submission at the end of this issue. Subscription and

membership are $25 per year ($45 for libraries). Postage outside of the USA and China, please add $5 per issue for surface

mail or $10 per issue for air mail. Make checks or money orders payable in US currency to AHRS. If you do not have access

to US currency. please notify us, and we will make other arrangements.

Asiatic Herpetological Research Volume 8 succeeds Volume 7 published in,1997. Vol. 6 published in 1995, Vol. 5 published

in 1993, Vol. 4 published in 1992, Vol. 3 published in 1990 and Chinese Herpetological Research Vol. 2, which was pub-

lished at the Museum of Vertebrate Zoology, 1988-1989 as the journal for the Chinese Society for the Study of Amphibians

and Reptiles. Volume 2 succeeded Chinese Herpetological Research 1987, published for the Chengdu Institute of Biology

by the Chongqing Branch Scientific and Technological Literature Press, Chongqing, Sichuan, China. Acta Herpetologica

Sinica ceased publication in June, 1988

Cover: Cuora galbinifrons. Diaoluoshan, 18 km N. of Nanxi, Hainan Province, China. Photo by James F. Parham

= “o a

These articles will appear in Volume 9 - Due out in SPRING 2001

Asiatic Herpetological Research

Museum of Vertebrate Zoology, University of California, Berkeley, California 94720

Fax: (510) 643-7706; email: asiaherp@uclink.berkeley.edu