AMPHIBIAN & REPTILE

CONSERVATION

THE INTERNATIONAL JOURNAL DEVOTED TO THE WORLDWIDE PRESERVATION

AND MANAGEMENT OF AMPHIBIAN AND REPTILIAN DIVERSITY

FOUNDER AND EDITOR

Craig Hassapakis

ASSOCIATE EDITOR

Jack W. Sites, Jr.

Brigham Young University

CHIEF ASSISTANT EDITOR

Csilla M. Csaplar

National Fish and Wildlife Foundation

ASSISTANT EDITORS

Deahn M. DonnerWright

North Central Research Station

Michael J. Dreslik

Illinois Natural History Survey

Stephen R. Johnson

William Penn University

Malcolm L. McCallum

Louisiana State University at Shreveport

CHAIRMAN OF THE BOARD

George C. Gorman

John D. McVay

Texas Tech University

Bethany J. Meisinger-Reiff

Marshall Erdman & Associates

Gad Perry

Texas Tech University

Craig D. Snyder

U.S. Geological Survey

ADVISORY BOARD

Allison C. Alberts

Zoological Society of San Diego

Aaron M. Bauer

Villanova University

Andrew R. Blaustein

Oregon State University

Joseph T. Collins

University of Kansas

Michael B. Eisen

Public Library of Science

Carl Gans

University of Texas at Austin

EDITORIAL REVIEW BOARD

Harold G. Cogger

Australian Museum

C. Kenneth Dodd, Jr.

U.S. Geological Survey

Lee A. Fitzgerald

Texas A&M University

Julian C. Lee

University of Miami

Harvey B. Lillywhite

University of Florida

Peter V. Lindeman

Edinboro University of Pennsylvania

James Hanken

Harvard University

Roy W. McDiarmid

U.S. Geological Survey

Russell A. Mittermeier

Conservation International

George B. Rabb

Chicago Zoological Society

Hobart M. Smith

University of Colorado

Michael Soulé

The Wildlands Project

Joseph C. Mitchell

University of Richmond

Henry R. Mushinshy

University of South Florida

Jaime E. Péfaur

Universidad de Los Andes

Christopher J. Raxworthy

American Museum of Natural History

Andrew T. Storfer

University of Florida

Larry David Wilson

Miami-Dade Community College

Continued on inside back cover

WORDS FROM THE EDITOR—The journal Amphibian and

Reptile Conservation (ARC) has made many advances since

our last published issue. A quick glance at this issue verifies

again our continued commitment and resolve to publish a jour-

nal of the highest standards devoted exclusively to the

conservation of amphibians and reptiles worldwide. To review

let me elaborate further.

The journal is now open access. This means anyone with an

Internet connection and web browser can access the contents of

the journal free-of-charge with absolutely no restrictions and/or

registration. Go to the PubMed Central website at http://www.

pubmedcentral.nih.gov/, review the list of journals, and click on

Amphibian and Reptile Conservation. The journal can be

retrieved as either exact reproduction of the actual journal pages

(PDF) and/or as HTML files. All the orginal photographs,

tables, graphs, etc. are available online in high resolution for all

the world to read, use, and exchange. The journal is also perma-

nently archived (online) by PubMed Central at the National

Library of Medicine (the world’s largest medical library) [NLM]

under the auspices of the US National Institutes of Health (NIH)

and National Center for Biotechnology Information (NCBI).

ARC does not charge any fees whatsoever for an author(s)

to publish in the journal. Authors publishing in the journal also

retain full copyright to all their material. Furthermore, the jour-

nal will always be available in hard copy (printed edition) for

those who prefer to read it in this format.

I feel it very important to elaborate on the importance of

open access publishing. In a perfect world, all journals (and lit-

erature) would be open access. To advance scientific study and

further the benefit to human health and especially environmen-

tal conservation (Fonseca and Benson 2003). It is imperative for

organizations, societies, and publishers to advance the open

access model for scientific publication. In a recent 194-page

report of the STM (Science, Technical, and Medical) journal

industry by Sami Kassab financial analysts at BNP Paribas

(http://www.bnpparibas.com/en/home/default.asp) it warned

about the impact of changes in the scientific publishing industry

(BioMedCentral 2003a). It was estimated in the report that the

global scientific research community could save more than 40%

in costs by switching entirely to an open-access model.

Societies should continue to increase organizational func-

tions and offer an ever-increasing number of member benefits

separate and therefore outside of the publication(s) aspect. In this

way, being a society member is a valued benefit separate from

receiving the publication(s). At the very least, members should

be able to access the societies publications full-text online from

home. I am a member of several scientific societies but cannot

access their content online from home; academic libraries are

where I need to go now for online access.

Editorial

Another added benefit for publications online is their inher-

ent ability to link to the full-text of other publications and

websites. This is a must in the electronic age we are heading into

full force (CrossRef [www.crossref.org] and PubMed Central

[www.pubmedcentral.nih.org]). For example, reading the most

recent articles from ARC one directly links to Proceedings of

the National Academy of Sciences of the United States of

America, AmphibiaWeb, and other articles, abstracts, and

important resources indexed and linked to the PubMed database.

Other important areas in an open access environment also

become a reality such as data mining and the use of the Digital

Object Identifier (DOI) [DeRisi, et al. 2003).

I support science and scientific societies, which advance

our knowledge of life on earth and all of scientific study in gen-

eral. I am especially partial to the Herpetologists’ League and

Herpetologica for it was the first science journal in the field of

herpetology I was exposed to. I remember riding my modest

Stingray bicycle 10 or 12 miles one way just to see the only

library in my area that had a subscription to Herpetologica! In

the electronic age, which is upon us and growing greater every-

day, this type of effort is very impractical. It is especially

important for people in developing countries to have good infor-

mation available online for this is where the most urgent

conservation work is being conducted (BioMed Central. 2003b).

The future is ready to explode further into the Information

Age caused primarily by the Internet and is thus literally chang-

ing the landscape of scientific publishing (Doyle, et al. 2003). It

is in the best interest of science and the public to make the nec-

essary steps toward the noble goal of open access of knowledge

(online) surely to advance society and surpass any of our wildest

dreams. The solutions to global warming, species extinctions,

developmental malformations as seen for example in some

amphibian species, will offer humankind unimagined advance-

ments in every field due to the uninhibited exchange of

information online. The brave, noble, and solution solvers now

and in the future will remain at the forefront of this exciting

human endeavor called open access.—Craig Hassapakis

References

BioMedCentral. 2003a. Financial analysts warn about impact of changes in the scientific

publishing industry. Open Access Now (15 December 2003).Reference online:http://

www.biomedcentral.com/openaccess/news/?issue=1 1

BioMedCentral. 2003b. Open Access in the developing world (15 December 2003).

Reference online:http://www.biomedcentral.com/openaccess/archive/?page=fea-

tures&issue=1 |

DeRisi, S., Rebecca, K., and Twyman, N. 2003. The what and whys of DOIs. PLoS

Biology 1(2):133.

Doyle, H. and Gass, A., and Lappin, D. 2003. A changing landscape. PLoS Biology

1(3):301.

Fonseca, G. and Benson, P.J. 2003. Biodiversity conservation demands open access.

PLoS Biology 1(2):163-165.

Authors

LARRY DAVID WILSON is a recognized authority on the reptiles and amphibians of Honduras, based on three and one-half decades

of field experience. He is the author of about 230 publications in his field, including the recently published The Amphibians of

Honduras, coauthored with James R. McCranie. He is also a professor of biology at Miami-Dade College in Miami, Florida, where he

has worked since 1972. He has traveled extensively in Mexico and Central America, especially Honduras. He is currently working on

books on Honduran reptiles (with James R. McCranie), the herpetofauna of the Honduran Mosquitia (with McCranie and Josiah H.

Townsend), and the herpetofauna of the Bay Islands and Cayos Cochinos of Honduras (with McCranie and Gunther KGhler).

JAMES R. (RANDY) MCCRANIE is also a recognized authority on the herpetofauna of Honduras, the result of more than a quarter

century of fieldwork. He is the author of about 160 papers in the field of herpetology and is the senior author of The Amphibians of

Honduras. He recently retired from 30 years of service with the U.S. Postal Service. His field experience is as extensive as is Wilson’s;

although he and Wilson usually travel to Honduras together, McCranie has made a number of trips by himself. He is involved with the

same three books as mentioned above, being senior author on two of them.

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 1

IPEN AGGESS JOURNAL

ISSN: 1083-446X

SUBSCRIBE AND/OR MAKE A DONATION

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Yes, I don’t want to be left out! Please sign me up for a subscription to ARC (four

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NOTE: Please add $15 per subscription for all orders outside the US.

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DONATIONS FOR THE JOURNAL ARE ACCEPTED

These resources are used to further the conservation of

amphibians and reptiles with goals of the journal.

Categories Friends of ARC $40

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Contributing $100

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Sustaining $500

Founder $1,000

Amount enclosed

Founders receive a lifetime subscription to ARC. All other

Unrestricted access to jour contributors receive four consecutively distributed issues.

pubmedcentral.nih.gov/) [PMC =

and managed by the United Staliyyiy . : ay RAAT mirranee ae , bank (check

journal orders must be paid in US currency drawn on a US bank (chee

(NLM) [the world's largest med or international money order). Journals are mailed to subscribers as they are

published. If available, previously printed journals are mailed with all new

subscriptions.

” <a

Amphibian Decline: An Integrated Analysis of

posi Multiple Stressor Effects

An Integrated Analysis of Multiple Stressor Effects

EEE Capturing the attention and imagination of the public and the scientific

community alike, the mysterious decline in amphibian populations drew

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amphibian decline and the role of stressors in these losses.

Greg Linder, Sherry Krest, Donald Sparling

ISBN 1-880611-55-4 $60/60€ members, $98/98€ nonmembers

Ecotoxicology of Amphibians and Reptiles

For contaminant ecologists interested in these seldom-studied but extremely

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physiological, and distributional considerations; responses and body burdens

of major contaminant groups; contaminant-related deformities, diseases, and

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Donald Sparling, Greg Linder, Christine Bishop

ISBN 1-880611-28-7 $72/72€ members, $118/118€ nonmembers

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Much more than simply a field guide, this

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“—

Plate 1. Plectrohyla dasypus. A Honduran endemic with all known populations believed to be declining.

DOI: 10.151 4/journal.arc.0000012.g001

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 6

the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduc- z

tion in any medium, provided the original work is properly cited. DOI: 10.151 4/journal.arc.000001 2 ( 004KB)

http://creativecommons.org/licenses/by/1.0/

The conservation status of the herpetofauna

of Honduras

LARRY DAVID WILSON’ AND JAMES R. MCCRANIE?

'Department of Biology, Miami-Dade Community College, Kendall Campus, Miami, Florida 33176-3393, USA

210770 SW 164th Street, Miami, Florida 33157-2933, USA

Abstract. —The conservation status of the members of the Honduran herpetofauna is discussed. Based on

current and projected future human population growth, it is posited that the entire herpetofauna is endan-

gered. The known herpetofauna of Honduras currently consists of 334 species, including 117 amphibians and

217 reptiles (including six marine reptiles, which are not discussed in this paper). The greatest number of

species occur at low and moderate elevations in lowland and/or mesic forest formations, in the Northern and

Southern Cordilleras of the Serrania, and the ecophysiographic areas of the Caribbean coastal plain and

foothills. Slightly more than one-third of the herpetofauna consists of endemic species or those otherwise

restricted to Nuclear Middle America. Honduras is an area severely affected by amphibian population decline,

with close to one-half of the amphibian fauna threatened, endangered, or extinct. The principal threats to the

survival of members of the herpetofauna are uncontrolled human population growth and its corollaries, habi-

tat alteration and destruction, pollution, pest and predator control, overhunting, and overexploitation. No

Honduran amphibians or reptiles are entirely free of human impact. A gauge is used to estimate environ-

mental vulnerability of amphibian species, using measures of extent of geographic range, extent of ecologi-

cal distribution, and degree of specialization of reproductive mode. A similar gauge is developed for reptiles,

using the first two measures for amphibian vulnerability, and a third scale for the degree of human persecu-

tion. Based on these gauges, amphibians and reptiles show an actual range of Environmental Vulnerability

Scores (EVS) almost as broad as the theoretical range. Based on the actual EVS, both amphibian and reptil-

ian species are divided into three categories of low, medium, and high vulnerability. There are 24 low vulner-

ability amphibians and 47 reptiles, 43 medium vulnerability amphibians and 111 reptiles, and 50 high vulnera-

bility amphibians and 53 reptiles. Theoretical EVS values are assessed against available information on cur-

rent population status of endemic and Nuclear Middle American taxa. Almost half (48.8%) of the endemic

species of Honduran amphibians are already extinct or have populations that are in decline. Populations of

40.0% of the Nuclear Middle American amphibian species are extirpated or in decline. A little less than a third

(27.0%) of the endemic reptiles are thought to have declining populations. Almost six of every ten (54.5%) of

the Nuclear Middle American reptilian species are thought to have declining populations. EVS values provide

a useful indicator of potential for endangerment, illustrating that the species whose populations are current-

ly in decline or are extinct or extirpated have relatively high EVS. All high EVS species need to be monitored

closely for changes in population status. A set of recommendations are offered, assuming that biotic reserves

in Honduras can be safeguarded, that it is hoped will lead to a system of robust, healthy, and economically

self-sustaining protected areas for the country’s herpetofauna. These recommendations will have to be

enacted swiftly, however, due to unremitting pressure from human population growth and the resulting defor-

estation.

Resumen.—Se discute el estatus de conservacion de los miembros de la herpetofauna de Honduras.

Basados en el crecimiento presente y proyectado de la poblacion del ser humano, se propone que toda la

fauna herpetologica de Honduras esta en peligro de extincion. Lo que se conoce de la fauna herpetologica

hondurena en el presente consiste de 334 especies, incluyendo 117 anfibios y 217 reptiles (incluyendo seis

reptiles marinos, que no se discuten en este articulo). La mayoria de las especies se presentan en bajas y

moderadas elevaciones en formaciones forestales de tierras bajas y/o humedas, en las Cordilleras

Septentrional y Meridional de la Serrania, y las areas ecofisiograficas de la costa y las faldas de la montana

del Caribe. Un poco mas de un tercero de la fauna herpetolégica consiste de especies endémicas o sino de

esas especies restringidas al Mesoamerica Nuclear. Honduras es una area severemente afectada por la dis-

minucion de las poblaciones de anfibios, con cerca de la mitad de la fauna anfibia amenazada, en peligro, o

extinta. Las principales amenazas a la sobreviviencia de los miembros de la fauna herpetoldgica son el crec-

Correspondence. ' Fax: (305) 237-0891, email: lwilson@mdcc.edu > acai Jmccrani@ bellsouth.net

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 7

L. D. Wilson and J. R. McCranie

imiento sin control de la poblacion humana y sus vastagos, la alteracion y destruction de habitacion, polu-

cion, el control de pestes y predadores, el exceso de caza y explotacion. Ningun anfibio o reptil hondureno

esta totalmente libre de el impacto humano. Se ha desarrollado una regla de medir para estimar la vulnera-

bilidad ambiental de las especies de anfibios, usando medidas de extension del rango geografico, amplitud

de distribucion ecoldégica, y estado de especializacion del modo de reproduccion. Se ha desarrollado una

medida similar para los reptiles, usando las dos primeras medidas de vulnerabilidad usados con los anfibios,

y una tercera medida para el grado de persecusion humana. Basados en estas medidas, los anfibios y rep-

tiles muestran un rango actual de una marca de vulnerabilidad medioambiental (EVS) casi tan amplia como

el rango teorético. Basados en la EVS, ambas especies de anfibios y reptiles estan divididas en tres cate-

gorias, de baja, media, y alta vulnerabilidad. Hay 24 especies de anfibios y 47 de reptiles de baja vulnerabili-

dad, 43 especies de anfibios y 111 de reptiles de media vulnerabilidad, y 50 especies de anfibios y 53 de rep-

tiles de alta vulnerabilidad. Teoréticamente, los valores de EVS son determinados de acuerdo de informacion

disponible del estado presente de las taxas endémicas de Mesoamérica Nuclear. Casi la mitad (48.8%) de las

especies endémicas de anfibios hondurenos estan ya extintos o tienen poblaciones en disminucion.

Poblaciones de 40.0% de las especies de anfibios de Mesoamerica Nuclear estan extintas o en disminucion.

Un poco menos de un tercio (27.0%) de los reptiles endémicos se piensa que tienen poblaciones en dismin-

ucion. Casi seis de cada diez (54.5%) de las especies de reptiles de Mesoamerica Nuclear se piensa que

tienen poblaciones en disminucion. Los valores de EVS proporcionan un indicador util del riesgo potencial,

el cual muestra que las especies cuyas poblaciones actuales estan disminuyendo, o son extintos o extirpa-

dos tienen EVS relativamente altos. Todas las especies con un EVS alto necesitan ser observadas de cerca

para anotar los cambios en el estado de las poblaciones. Ofrecemos un grupo de recomendaciones, asum-

iendo que las reservas bidticas de Honduras pueden ser preservadas, se espera que esto resulte en un sis-

tema de areas protegidas que es robusta, saludable, y sostenible economicamente para la fauna her-

petologica del pais. Estas recomendaciones tienen que ser observados rapidamente, debido a la presion con-

tinua causada por el crecimiento de la poblacion humana y la resultante destruccion de los bosques.

Key words. Conservation status, amphibians, reptiles, herpetofauna, Honduras, distribution

“To the extent that we depend on prosthetic devices to

keep ourselves and the biosphere alive, we will render

everything fragile. To the extent that we banish the rest of

life, we will impoverish our own species for all time. And if

we should surrender our genetic nature to machine-aided

ratiocination, and our ethics and art and our very mean-

ing to a habit of careless discursion in the name of

progress, imagining ourselves godlike and absolved from

our ancient heritage, we will become nothing.”

E. O. Wilson

Consilience: the unity of knowledge, 1998

Introduction

The portion of the closing paragraph of E. O. Wilson’s (1998)

powerful book quoted above provides an extremely serious

warning to our species, a warning that in continuing with our

plan to place all the natural world in service to ourselves, we

risk erasing any meaning for our continued existence. This

concept is antipodal to the usual thinking that we encounter

our raison d’étre as we continue to subjugate Nature to our

own designs. One of the central goals of conservation biolo-

gy, then, is to attempt to bridge the gap between these anti-

thetical worldviews in an effort to salvage and restore as much

of the remaining global biodiversity as possible in the shortest

time possible.

It is common knowledge among biologists that the great-

‘est amount of biodiversity resides in the area between the

Tropics of Cancer and Capricorn—the tropics. It is frequently

stated that 40-80% of the diversity of life occurs in this region

Amphib. Reptile Conserv. | http://www.herpetofauna.org

(Miller 2001; Raven and Berg 2001). Unfortunately, this

region also is subject to the highest rates of human population

growth. For example, in the Western Hemisphere, there are

thirty-one countries that lie wholly within the tropics. The

average natural increase for these thirty-one countries is

1.71% (data obtained from the 2000 World Population Data

Sheet of the Population Reference Bureau, an insert in Raven

and Berg 2001). This translates to an average doubling time of

40.9 years (using the formula DT = 70/natural increase).

The countries of Central America, however, are the

fastest growing ones in the American tropics (data obtained

from the 2000 World Population Data Sheet of the Population

Reference Bureau, an insert in Raven and Berg 2001). Natural

increase ranges from a low of 1.7 in Panama to a high of 3.0

in Nicaragua, with doubling times ranging from 23 years for

Nicaragua to 41 years for Panama.

Growth rates, however, are significantly higher for the

nations of northern Central America than are those for lower

Central America. Costa Rica and Panama have growth rates of

1.8 and 1.7, respectively, whereas those for Belize,

Guatemala, El Salvador, Honduras, and Nicaragua range from

2.4 to 3.0. For the latter five countries, these figures translate

to doubling times ranging from 23 (Nicaragua) to 29 years (El

Salvador). The natural increase of Honduras, at 2.8%, is the

third highest in Central America, being exceeded only by

those of Guatemala (2.9%) and Nicaragua (3.0%). Thus, its

doubling time is the third fastest in the region, at 25 years.

The senior author has been working on the herpetofauna

of Honduras since 1967. In the 35 years since then, the human

population of the country has grown from about 2.4 million to

a figure somewhat in excess of 6.7 million (the former figure

Volume 3 | Number 1 | Page 8

The conservation status of the herpetofauna of Honduras

is from Golenpaul, 1968, and the latter one is from data

obtained from the 2001 World Population Data Sheet of the

Population Reference Bureau, an insert in later copies of

Raven and Berg 2001). In other words, in that 35-year period

of time, the population of Honduras has doubled and increased

by almost half again as much.

Habitat degradation and destruction are recognized as the

major threats to biodiversity today (Raven and Berg 2001).

Such degradation and destruction in Honduras is primarily

fueled by deforestation (E. Wilson and Perlman 2000), occa-

sioned by shifting agricultural practices, ranching, logging,

and fuel gathering. The deforestation models in E. Wilson and

Perlman (2000) indicate that the amount of forest remaining in

1995 amounted to 4.1 million hectares. Honduras, however,

contains 43,277 sq. mi. or 11,208,935 hectares. Thus, in 1995

only about 37% of the original forested area of the country

(i.e., once the entire country) remained. The E. Wilson and

Perlman (2000) deforestation model for Honduras also indi-

cates that the time to halve the remaining forest is 30.1 years.

Thus, the 1995 figure of 4.1 million hectares will be down to

2.05 million hectares by about 2025. The deforestation rate

indicated by E. Wilson and Perlman (2000) is -2.3% and will

reduce the remaining forest in the country to 0.5 million

hectares by the year 2085. It can be expected that, if these

rates continue, no forest will remain in Honduras by the end

of the present century.

Measured against this backdrop, it is abundantly clear

that the Honduran herpetofauna, and indeed the entire biota, is

endangered, in the best sense of the term. Equally clear, thus,

is the rationale for an examination of the conservation status

of the herpetofauna of the country. If we do not examine it

now, we can only look to further deforestation, fueled by the

uncontrolled growth of the human population, and increasing

threats to the survival of the herpetofauna. We have no idea

what the herpetofauna of Honduras looked like at the time of

Columbus’ arrival at Cabo de Honduras, opposite Trujillo, in

1502, but at least we do know that the known herpetofauna

that existed when the senior author began to work in the coun-

try in 1967 is not the herpetofauna known today (see below).

It is the purpose of this paper to assess the conservation

status of the known members of the Honduran herpetofauna

and to construct a set of conservation and research priorities

for the foreseeable future. It is hoped that the brutal honesty

with which we have approached this work will act to spur the

necessary steps to enable these priorities before this segment

of the Honduran patrimony is lost for all time.

Status of our knowledge of the Honduran

herpetofauna

The modern history of the study of the amphibians and rep-

tiles of Honduras began with the first trip to the country made

by John R. Meyer in 1963. Meyer was “in country” for three

months with a field crew from Texas A&M University led by

the mammalogist Gerald V. Mankins. It was during this trip

that Meyer began to formulate an idea for a dissertation topic

dealing with a survey of the herpetofauna of Honduras. With

his transfer to the University of Southern California under the

mentorship of Jay M. Savage, the idea became a reality.

At about the same time, Larry D. Wilson was also work-

Amphib. Reptile Conserv. | http://www.herpetofauna.org

ing on his dissertation at Louisiana State University in Baton

Rouge. Unaware of Meyer’s dissertation work, Wilson began

to survey various collections around the country to see what

material from Honduras existed there. The word got around to

Meyer, who then began to correspond with Wilson. In time,

Meyer suggested that Wilson join him on a three-month field

trip to the country during the summer of 1967. A second three-

month journey ensued in the summer of 1968.

At this point, Meyer began to write his dissertation,

which was completed in 1969 (Meyer 1969). The known her-

petofauna as of that publication consisted of 196 species. Two

years later, Meyer and Wilson (1971) provided a checklist of

the amphibian fauna containing 52 species and in 1973 a

checklist of the turtle, crocodilian, and lizard fauna listing 59

species (not 58, as stated in their abstract and introduction).

Wilson and Meyer (1985) treated 95 species of snakes then

known to occur in Honduras (Wilson and Meyer 1982, had

treated 91 species of snakes in Honduras).

In 1976, Wilson began to work with James R. McCranie,

and their first paper together (joined by Louis Porras) on

Honduras appeared in 1978 (Wilson et al. 1978). These same

three authors described in 1980 the first new species to result

from the fieldwork up to that point (McCranie et al. 1980). In

1983, Wilson produced the first list of amphibians and reptiles

for the country since the work of Meyer and Wilson (1971,

1973) and Wilson and Meyer (1982). That list consisted of

208 species (56 amphibians and 152 reptiles). Wilson and

McCranie (1994) produced a second update of the Honduran

herpetofauna, listing a total of 277 species (89 amphibians and

188 reptiles).

The latest accounting of the species of amphibians is in

McCranie and Wilson (2002). This book lists 117 species for

Honduras, including two species of caecilians, 25 species of

salamanders, and 90 species of anurans (one of which is

reported in an addendum). The most recent list of the reptiles

is in Wilson and McCranie (2002), in which are included 217

species (14 turtles, two crocodilians, 88 lizards, and 113

snakes). The total known herpetofauna, thus, as of these two

publications, consists of 334 species (including six marine

reptiles).

McCranie and Wilson (2002) hypothesized that seven

additional species of amphibians probably reside in Honduras.

A similar work in progress on the reptiles of Honduras

(McCranie and Wilson, in preparation) lists 13 species of

probable occurrence. At the present time, then, we know the

herpetofauna consists of 334 species, and we think it may con-

tain as many as 20 more species, apart from any new taxa that

may be discovered. The above summarizes our current under-

standing of the composition of the Honduran herpetofauna.

Our understanding of the geographic and ecological dis-

tribution of the members of the herpetofauna of Honduras is

summarized in McCranie and Wilson (2002) for the amphib-

ians and, to a lesser extent, in Wilson et al. (2001). The latter

situation is the case because Wilson et al. (2001) spent over

five years in press and could not be consistently updated to the

point it appeared in print. For example, Wilson et al. (2001)

considered 276 species of amphibians and reptiles, but did not

include five species of marine turtles, one species of marine

snake, and six reptile species restricted in Honduras to the

Swan Islands and the Miskito Keys. Inclusion of these 12

Volume 3 | Number 1 | Page 9

dante wt

Plate 2 DOI: 10.1514/journal.arc.0000012.g002

Plate captions: 2. Primary forest in Parque Nacional El Cusuco,

Cortés. Photograph taken from 1820 m elevation on 13 April

1979. 3. Primary forest along Rio de Cusuco, Parque Nacional El

Cusuco, Cortés. Photograph taken at 1670 m elevation on 13

April 1979. 4. Primary forest in Parque Nacional de Celaque,

Lempira. Photograph taken from 2440 m elevation on 28 April

1982. 5. Primary forest along Rio Seco, Parque Nacional Sierra

de Agalta, Olancho. Photograph taken at 990 m elevation on 8

August 1986. Primary forest like that shown in Plates 1-4 exists

today only within the boundaries of some of the biological

reserves of Honduras. 6. Primary forest along Quebrada de Oro,

Parque Nacional Pico Bonito, Atlantida. Photograph taken at 950

m elevation on 4 June 1980. 7. Quebrada de Oro, Parque

Nacional Pico Bonito, Atlantida, showing destruction caused by a

large landslide in November 1988. Photograph taken at 940 m

elevation on 7 August 1989.

Amphib. Reptile Conserv. | http://www.herpetofauna.org

Plate 4 DOI: 10.1514/journal.arc.0000012.g004

¢ : > on” pl f, y- : With a2 nan , mf ~ ES

Plate 3 DOI: 10.1514/journal.arc.0000012.g003

Plate 5 DOI: 10.1514/journal.arc.0000012.g005

2 Mae Ne Ee

DOI: 10.1514/journal.arc.0000012.g007

Plate 7

Volume 3 | Number 1 | Page 10

Plate 8 DOI: 10.1514/journal.arc.0000012.g008 Plate 9 DOI: 10.151 4/journal.arc.0000012.g009

Plate 11 DOI: 10.151 4/journal.arc.000001 2.9011

Plate 10 DOI: 10.1514/journal.arc.0000012.g010

Plate captions: 8. Collapsed ridge on slope N of Quebrada de Oro, Parque Nacional Pico Bonito, Atlantida. This ridge is part of the two

large landslides that severely damaged a large portion of the Quebrada de Oro in November 1988 and November 1995. Photograph

taken at ca. 1000 m elevation on 28 May 1996. 9. Quebrada de Oro, Parque Nacional Pico Bonito, Atlantida. Portion of stream through

primary forest that disappeared underground between 28 May and 2 June 1996. Colonies of army ants had invaded the dry stream bed

to feed on the perished tadpoles (mostly Atelophryniscus chrysophorus and Ptychohyla spinipollex) and invertebrate carcasses.

Photograph taken at 960 m elevation on 3 June 1996. 10. El Portillo de Ocotepeque, Ocotepeque. This area was “protected” as part of

the Reserva Bioldgica GUisayote in 1987, even though the vast majority of this reserve was already deforested at that time. Photograph

taken at 1900 m elevation on 14 April 1978. 11. Quebrada Grande, Parque Nacional Cerro Azul, Copan. The haze in the photograph is

smoke from slash and burn agriculture. The only forest remaining today in this national park is on some of the steep slopes above this

village. The national park was created in 1987 and the photograph was taken on 6 May 1988 (from 1500 m elevation).

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 11

L. D. Wilson and J. R. McCranie

species would have raised their tally to 288 species, which is

46 species fewer than the number now known to occur in the

country. Thus, the information presented below is somewhat

more accurate for the amphibians than it is for the reptiles,

although the major distributional patterns discussed are not

affected much by the relative lack of currency of the informa-

tion for the reptiles, nor will it have much affect on the con-

clusions reached in the remainder of this paper.

Both Wilson et al. (2001) and McCranie and Wilson

(2002) discussed ecological distribution of Honduran amphib-

ians and reptiles with respect to ecological formations, phys-

iographic regions, elevation, and ecophysiographic areas.

They also discussed the broad patterns of geographic distribu-

tion of these animals.

With regard to distribution in ecological formations

(modified from those of Holdridge 1967), Wilson et al. (2001)

indicated that the greatest number of species occur in lowland

formations (Lowland Moist Forest, Lowland Dry Forest, and

Lowland Arid Forest formations) and mesic formations

(Lowland Moist Forest, Premontane Wet Forest, Lower

Montane Wet Forest, and Lower Montane Moist Forest for-

mations). For the amphibians alone, however, the greatest

numbers of species are found in only three of the four mesic

formations (Premontane Wet Forest, Lowland Moist Forest,

and Lower Montane Wet Forest formations).

With reference to distribution in physiographic regions,

Wilson et al. (2001) noted that the greatest numbers of species

are found in the Northern Cordillera and the Southern

Cordillera, these two areas comprising the Serrania of

Honduras. The same pattern was discovered for the amphib-

ians when considered alone (McCranie and Wilson 2002).

Analysis of distribution with respect to elevation indi-

cates that the greatest number of amphibians and reptiles

occur at low elevations (0-600 m), although moderate eleva-

tions (601-1500 m) harbor almost as many (Wilson et al.

2001). When amphibians are considered alone, however, there

is a significantly greater number of species known from mod-

erate elevations (88 species) than from low elevations (65

species). In addition, a sizable number of species (56) also

occurs at intermediate elevations (1501-2700 m).

Combining ecological formations and physiographic

regions gives rise to ecophysiographic areas (see Wilson et al.

2001 for a discussion). Thirty-eight such areas were recog-

nized by Wilson et al. (2001), of which 28 were subjected to

analysis. McCranie and Wilson (2002), however, presented

data on amphibian distribution in 32 of the 38 areas (see

McCranie and Wilson 2002 for a map showing the distribu-

tion of these areas). Wilson et al. (2001) showed that the

highest numbers of species occurred (in decreasing order) in

the Eastern Caribbean Lowlands, the West-central Caribbean

Lowlands, the Sula Valley, and the Central Caribbean Slope,

all of which are Caribbean lowland regions or the foothills

above such areas. When the amphibians are considered alone,

however, a slightly different pattern emerges. The highest

numbers of species of amphibians are found in the Eastern

Caribbean Lowlands, the Eastern Caribbean Slope, the

Central Caribbean Slope, and the Western Caribbean Slope.

The prevalence of foothill regions in this list is reflective of

the sizable presence of amphibians at moderate elevations in

the country (see above).

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Analysis of the broad patterns of geographic distribution

by Wilson et al. (2001) showed that the largest numbers of

species are endemic to the country or otherwise restricted to

Nuclear Middle America (about a third of the herpetofauna

therein considered). Slightly more than 90 percent of the her-

petofauna were distributed in the area from Mexico to South

America. The amphibians, when considered alone (McCranie

and Wilson 2002), show the same pattern, with 56.9% either

endemic to Honduras or to Nuclear Middle America and

94.0% distributed in the area from Mexico to South America.

The overall outcome of the research on the Honduran

herpetofauna that has taken place since 1967 is the description

of a large number of new taxa, the discovery of a sizable num-

ber of species new to the herpetofauna, and a few

resurrections of formerly synonymized taxa. More recently,

however, we have entered a new era in our studies in

Honduras, as detailed by McCranie and Wilson (in press) for

the amphibians. As noted above, McCranie and Wilson (2002)

treated 116 species of amphibians (and another one in an

addendum). The majority of these 116 amphibian species are

either endemic to Honduras (41 species) or otherwise endem-

ic to Nuclear Middle America (25 species). Thus, 56.9% of

the amphibian fauna falls into these two distributional cate-

gories, as noted above. The analysis presented by McCranie

and Wilson (in press) indicates that of the 41 endemics, six

apparently have already disappeared. The populations of an

additional 14 are in apparent decline (field work in 2001 indi-

cated that one of the 14 species thought to be in decline by

McCranie and Wilson, in press, has also disappeared) and

there are four species for which we do not currently know the

population status. Thus, only 17 of 41 species (41.5%) appear

to have stable populations at the present time. Of the 25

species otherwise restricted to Nuclear Middle America, the

populations of nine species appear to be in decline and those

of one species appears to have been extirpated in Honduras.

We have no data on the populations of an additional four

species. Thus, only 11 of 25 species (44.0%) appear to have

populations that are stable at this time. Of the 50 remaining

amphibian species not discussed above, McCranie and Wilson

(in press) determined that 25 (50.0%) of them require rela-

tively undisturbed forest regions to survive, and, thus, have

lost much of their habitat in recent years. In summary, the

populations of only 53 of 116 species of Honduran amphib-

ians (45.7%) appear to be stable or nearly so. Thus, close to

half the known amphibian fauna of Honduras is threatened,

endangered, or now extinct. This sad picture is being repeated

throughout much of Latin America (Young et al. 2001).

In a following section, we attempt to establish a set of

conservation priorities for all the members of the Honduran

herpetofauna, using revised environmental vulnerability

scores, first developed and used by Wilson and McCranie

(1992).

Threats to the survival of amphibians and reptiles of

Honduras

Wilson et al. (2001:109) opined that, “The most serious of the

plethora of environmental problems impacting the planet cur-

rently, perhaps, is biodiversity decline, for this is the only one

that is irreversible. As species of organisms are pushed to

Volume 3 | Number 1 | Page 12

The conservation status of the herpetofauna of Honduras

extinction, the information stored in their genomes is irre-

trievably lost. What importance such creatures have in

maintaining the planet’s life support systems and what more

immediate or direct value that information content may have

for humanity is most often extremely imperfectly known to

completely unknown. Upon the extinction of the organisms,

such enlightenment becomes permanently unattainable.” This

opinion is based on a cascade of modern research concerning

the nature and extent of environmental problems, most specif-

ically about the above-discussed problem of biodiversity

decline (see, for example: Ehrlich and Ehrlich 1981, 1996; E.

Wilson 1984, 1988 [ed.], 1992; E. Wilson and Perlman 2000;

Miller 2001; Raven and Berg 2001).

The anthropogenic threats to the Earth’s biota are fairly

clearly identified. E. Wilson and Perlman (2000), for example,

identify the following threats as most important:

¢ Habitat loss and fragmentation

e Exotic species

¢ Overhunting

¢ Degradation of air, water, and soil

e Synergistic pressures

Raven and Berg (2001) listed the following factors as

most important for U.S. plants and animals:

¢ Habitat loss and degradation

e Exotic species

¢ Pollution

¢ Overexploitation

McCranie and Wilson (2002) identified habitat alteration

and destruction, pollution, and pest and predator control as the

threats of greatest importance to Honduran amphibians. When

one considers the reptile segment of the herpetofauna, then

overhunting and overexploitation must be added to the list.

However, it may be shown that the synergistic interactions of

these various threats will represent the ultimate threat (E.

Wilson and Perlman 2000), pushing the existing natural sys-

tems in Honduras beyond any hope of recovery. Given the rate

at which habitat alteration and destruction is proceeding, as

especially measured by the rate of deforestation (see the

Introduction), it may be hypothesized that the collapse of most

to all of the populations of the country’s amphibians and rep-

tiles will be complete at or before the end of the present

century. In the same period of time, based on Honduras’s

human population doubling time of 25 years (data obtained

from the 2000 World Population Data Sheet of the Population

Reference Bureau, an insert in Raven and Berg 2001), its pop-

ulation will increase theoretically by a factor of 16 times! One

of the most basic questions facing the populace of Honduras

is what the country will be doing with its 107.2 million people

it is scheduled to have by the year 2101.

In recent years, additional threats have been manifested.

One such threat comes in the form of a chytrid fungus that

has been implicated as a proximate cause of mortality for

anurans in Australia, Costa Rica, and Panama (see Berger et

al. 1998, Lips 1999). This effect is especially startling, inas-

much as it has been occurring “... in pristine areas at

moderate to intermediate elevations” (McCranie and Wilson

2002, p. 539). Many tadpoles of several Honduran species of

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montane hylids of the genus Plectrohyla, as well as a species

of Ptychohyla, have been found to have deformed keratinized

mouthparts, likely a symptom of infection by a chytrid fun-

gus (McCranie and Wilson 2002; also see Fellers et al. 2001).

Another threat may be connected to “documented climatic

changes associated with recent warming” (McCranie and

Wilson 2002, p. 527-528), strongly implicated by Pounds et

al. (1999) to be responsible for amphibian population crashes

in a Costa Rican montane habitat. We suspect “these same

climatic changes are also likely taking place in montane habi-

tats within Honduras” (McCranie and Wilson 2002, p. 528)

and may be implicated in what is looking like a general trend

in the decline or disappearance of several anuran species in

pristine regions at moderate to intermediate elevations

(essentially above 900 m; see McCranie and Wilson 2002 for

a more extended discussion).

What is especially frightening about these recent devel-

opments involving pathogens and climatic change is that they

produce unanticipated changes that make it difficult to impos-

sible to predict their effects. As such, it becomes difficult to

impossible to plan for these effects. They appear to have the

potential to become an environmental “super-problem,” in the

sense of Bright (2000). Bright (2000) uses this term to

describe environmental synergisms resulting from the interac-

tion of two or more environmental problems, so that their

combined effect is greater than the sum of their individual

effects. These problems represent an environmental worst-

case scenario—the point when environmental problems

become so serious that they produce unanticipated results, the

successful resolution of which threaten to slip forever from

the grasp of humanity. It is against this terrifying backdrop

that we proceed with the effort to assign conservation priori-

ties for the members of the herpetofauna of Honduras. It may

be stated without fear of contradiction that there are no popu-

lations of Honduran amphibians and reptiles that are entirely

free of anthropogenic impact (Wilson et al. 2001, McCranie

and Wilson 2002, McCranie and Wilson, in press).

Establishment of conservation priorities for the

Honduran herpetofauna

Prior attempts have been made by us to assess the effective-

ness of the current system of biotic reserves in Honduras in

protecting the country’s herpetofauna (Wilson et al. 2001), to

determine the status of amphibian populations (McCranie and

Wilson, in press), and to anticipate the future of the amphib-

ian faunal component (McCranie and Wilson 2002). Each of

these efforts has pointed to significant threats to the integrity

of herpetofaunal populations. In a very real sense, this is all

we have been able to do—to point to these threats. Addressing

these threats in any meaningful way is the responsibility of the

people of Honduras—through their government, information

media, educational systems, and environmental organizations.

We have written this paper in the hope that looking at these

problems in a different way than has been done heretofore

may act to focus sufficient attention before it is too late—if it

is not too late already. An overriding problem is that there is

little consensus in the literature concerning the number and

individual sizes of the protected areas in the country (see

Table 15 in Wilson et al. 2001; Anonymous 2001).

Volume 3 | Number 1 | Page 13

DOI: 10.1514/journal.arc.0000012.g012 Plate 13

SE *

Plate 14 DOI: 10.151 4/journal.arc.0000012.g015

; e S y- “se ne a 2 4 oS . eo | : ee oa . ‘ tai i : “ A .

Plate 16 000012.g016 Plate 17 : 10.1514/journal.arc.0000012.g017

Plate 18 DOI: 10.1514/journal.arc.0000012.g018 Plate 19 DOI: 10.1514/journal.arc.0000012.g019

Plate captions: 12. Reserva Bioldgica El Pital, Ocotepeque. Almost no original forest remains in this reserve. Photograph taken from 1480 m ele-

vation showing secondary gallery forest in foreground and denuded hillsides in background. 12 August 1997. 13. Reserva de la Bidsfera Rio

Platano, near Quebrada de Las Marias, Olancho. The forested hillsides in the background lie about 1 km from the southern edge of the “nuclear

zone” of this Biosphere Reserve. Photograph taken at 660 m elevation on 1 August 1997. We rode on horseback through this same locality in August

1998, and found the human population to have substantially increased from the previous year, as had the deforestation. 14. 3.7 km NW of

Zambrano, Francisco Morazan. Photograph taken at 1450 m elevation in June 1976. These pine forests are burned annually, thus the trees in this

area are now considerably more fire scarred. In addition, tree stumps and logs lying on the ground are now largely bumt remains, offering little

refuge for ground dwelling snakes. 15. Eleutherodactylus anciano. 16. Eleutherodactylus chrysozetetes. 17. Eleutherodactylus milesi. 18.

Eleutherodactylus stadelmani. Plates 15 through 18. Honduran endemics now feared extinct. 19. Bolitoglossa car. A Honduran endemic with all

known populations believed to be declining.

The conservation status of the herpetofauna of Honduras

Many others share these concerns, of course. In fact,

Honduras is one of the countries in the Western Hemisphere

that figures into the Mesoamerican Biological Corridor Project

(“Paseo Pantera”), as described by Illueca (1997). While

expansive and desirable in concept, there are serious problems

in its design and prospects in Honduras. The map of the com-

ponents of this project in Mesoamerica includes a number of

“protected areas” (incidentally, one of these “protected areas,”

the Mayan ruins of Copan, Honduras, is mismapped; what is

shown apparently is the Parque Nacional Montecristo-Trifinio)

and “desired green connections.” We have previously dis-

cussed the pressures existing in the “protected areas” (here and

in Wilson et al. 2001). Even more significantly, however, are

the problems associated with attempting to turn the “desired

green connections” into anything actually “green” (i.e., eco-

logically restored). For example, one of these connections

traverses the area between the Maya Mountains Biosphere

Reserve in Belize, the Copan Maya Ruins in the department of

Copan in extreme western Honduras, and the Rio Platano

Biosphere Reserve in northeastern Honduras. The intervening

area encompasses about the western two-thirds of Honduras, in

which area lives the large majority of the human population of

the country. This is also the area that has suffered greatly at the

hands of agriculturists for centuries, to the point that

Hondurans, especially the landless poor, are moving in signif-

icant numbers to the less heavily exploited Mosquitia in

eastern Honduras. Creating a “green connection” through this

area of the country appears to us to be an impossibly large task.

Several years ago (Wilson and McCranie 1992), we

developed an environmental vulnerability gauge for use with

amphibian populations. We then (McCranie and Wilson 2002)

updated it for use with the 116 species of amphibians treated

in The Amphibians of Honduras. For this paper, we have

developed a similar gauge for the reptiles. The gauge for

amphibians and that for reptiles resemble one another in using

scales for extent of geographic range and ecological distribu-

tion. The two gauges differ from one another in that

susceptibility of reproductive mode to anthropogenic pressure

is used for amphibians and extent of human persecution is

used for reptiles (see below).

We use these gauges to establish a set of conservation

priorities for the remaining species of the Honduran her-

petofauna. This is an approach different from the one we

adopted in Wilson et al. (2001), which attempted to evalu-

ate the effectiveness of the existing system of biotic

reserves to protect al] members of the herpetofauna known

at the time, and to make suggestions about where addition-

al reserves needed to be established. In essence, we have

been forced to adopt a different approach, given the mute

testimony provided in recent years by disappearing

Honduran amphibians.

As noted above, this environmental vulnerability gauge

for both amphibians and reptiles has three components, which

are described below. The first component of the gauge, appli-

cable to both groups, deals with the extent of the geographic

range using the following scale:

1 = widespread in and outside of Honduras

2 = distribution peripheral to Honduras, but widespread

elsewhere

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3 = distribution restricted to Nuclear Middle America

(exclusive of Honduran endemics)

4 = distribution restricted to Honduras

5 = known only from the vicinity of the type locality

As is evident, in a rough sense, the degree of restriction

of geographic range increases as the scale number increases.

The second gauge component, also applicable to both

groups, indicates the extent of ecological distribution, based

on a modified version of the forest formations of Holdridge

(1967), using the following scale (omitting consideration of

the Montane Rainforest formation, the herpetofauna of which

is almost completely unknown):

1 = occurs in eight formations

2 = occurs in seven formations

3 = occurs in six formations

4 = occurs in five formations

5 = occurs in four formations

6 = occurs in three formations

7 = occurs in two formations

8 = occurs in one formation

The degree of restriction of ecological range increases as

the scale number increases, similar to that of geographic range

in the previous component.

In gauging the degree of specialization of reproductive

mode in amphibians, as it relates to the effect of environmen-

tal modification, especially deforestation, we use the

following scale:

1 = both eggs and tadpoles in large or small bodies of

lentic or lotic water

2 = eggs in foam nests, tadpoles in small bodies of lentic

or lotic water

3 = tadpoles occur in small bodies of lentic or lotic water,

eggs elsewhere

4 = eggs laid in moist situations on land or moist arbore-

al situations, direct development

5=eggs and tadpoles in water-retaining arboreal

bromeliads or water-filled tree cavities

Again, increase in number signifies probable increase

in reproductive vulnerability to the effects of habitat

degradation.

In light of the fact that reptiles are amniote vertebrates

and, thus, do not possess the biphasic life cycle or the range

of reproductive modes typical of amphibians, it is necessary

to develop another gauge of human pressure on the popula-

tions of these animals. In addition, reptiles, being vertebrates

fully adapted to life on land, are often more noticeable to

humans and more frequently encountered than are amphib-

ians, especially larval amphibians. Moreover, many, if not

most, reptiles are the subjects of superstition, ignorance,

fear, and, as a consequence, outright killing upon sight.

Finally, given that all Honduran reptiles are scaled verte-

brates and some are large enough to be of commercial

interest for their hides, meat, and/or eggs, these species are

hunted (i.e., actively sought) for these products. Taking

these biological and sociological features into consideration,

we developed the following scale to indicate the degree of

human persecution:

Volume 3 | Number 1 | Page 15

L. D. Wilson and J. R. McCranie

1 = fossorial, usually escape human notice

2 = semifossorial, or nocturnal arboreal or aquatic, non-

venomous and usually nonmimicking, sometimes

escape human notice

3 =terrestrial and/or arboreal or aquatic, generally

ignored by humans

4 = terrestrial and/or arboreal or aquatic, thought to be

harmful, may be killed on sight

5 = venomous species or mimics thereof, killed on sight

6=commercially or noncommercially exploited for

hides and/or meat and/or eggs

As with the previously discussed components, the degree

of threat from human beings roughly increases as the scale

number increases.

In order to obtain this rough idea of environmental vul-

nerability, thus, each of the three applicable scores has been

determined for each Honduran amphibian and reptilian

species. Then the numbers associated with the three scales

have been added to obtain a composite score. These compos-

ite scores can range theoretically from a low of three to a high

of 18 for amphibians and from a low of three to a high of 19

for reptiles.

The composite environmental vulnerability scores (EVS;

used either in singular or plural form, as determined by con-

text) for amphibians (Table 1) actually range from a low of

three to a high of 17, almost the entire gamut. The numbers of

species attaining the various EVS are as follows:

EVS 3-1 species EVS 11-12 species

EVS 4-1 species EVS 12-13 species

EVS 5-5 species EVS 13-13 species

EVS 6-7 species EVS 14-15 species

EVS 7-2 species EVS 15-17 species

EVS 8-2 species EVS 16-10 species

EVS 9-6 species EVS 17-8 species

EVS 10-5 species

Using this measure, the least vulnerable amphibian

species are Bufo marinus, B. valliceps, Hyla microcephala,

Phrynohyas venulosa, Scinax staufferi, Smilisca baudinii,

and Rana berlandieri. They are all 1-1-1, 1-2-1, or 1-3-1

species (species widespread geographically in and outside of

Honduras, of broad ecological occurrence, and having the

least derived reproductive mode). The most vulnerable

species are Bolitoglossa carri, B. decora, B. longissima,

Nototriton lignicola, Eleutherodactylus chrysozetetes, E.

coffeus, E. cruzi, and E. merendonensis. They are all 5-8-4

species (species known only from the vicinity of the type

locality, in one forest formation, with eggs laid in moist sit-

uations on land or moist arboreal situations). In addition,

three of the four species of Eleutherodactylus (save for E.

coffeus for which there are no data available) appear to have

already disappeared or are in decline (McCranie and Wilson,

in press).

We have used the same method in this paper as

McCranie and Wilson (2002). Thus, we have divided the

species of Honduran amphibians into three categories of

environmental vulnerability, i.e., low vulnerability, of medi-

um vulnerability, and high vulnerability. This categorization

provides an initial rough means of gauging the degree of

Amphib. Reptile Conserv. | http://www.herpetofauna.org

attention that ought to be focused on the various taxa. Thus,

the species that can be expected to have the best chance to

survive in the face of continued environmental degradation

are those in the first category. These 24 species make up

only 20.5% of the Honduran amphibian fauna. A larger

group of 43 species, making up 36.8% belongs to the medi-

um category; nonetheless, this is a heterogeneous grouping,

created due to a lack of weighting of the three categories

used to compute the EVS, in which relatively widespread

species, such as Agalychnis callidryas, are grouped with

highly restricted ones, such as Plectrohyla chrysopleura. A

larger group of 50 high vulnerability species, making up

42.7%, can be expected to have the poorest chance for sur-

vival. Almost all of these species are endemic to Honduras

or are otherwise restricted to Nuclear Middle America.

Additionally, recent declines or disappearances in amphibian

populations from moderate to intermediate elevation, pris-

tine habitats were not considered in this analysis. The

importance of these declines and disappearances, however,

is discussed in the following section.

The composite environmental vulnerability scores (EVS)

for reptiles (Table 2) actually range from a low of four to a

high of 19, only one number less than the entire theoretical

range (marine species not included). The numbers of species

attaining the various EVS are as follows:

EVS 4 - | species EVS 12 - 42 species

EVS 5 -1 species EVS 13 - 28 species

EVS 6 -2 species EVS 14 - 15 species

EVS 7 - 9 species EVS 15 - 23 species

EVS 8-11 species EVS 16 - 11 species

EVS 9 - 23 species VS 17 - 2 species

EVS 10 - 19 species EVS 18 - | species

EVS 11 - 22 species EVS 19 - | species

The least vulnerable reptilian species, by this measure, are

Norops tropidonotus, Enulius flavitorques, Imantodes cenchoa,

and Ninia sebae. They are 1-1-2, 1-1-3, or 1-3-2 species (wide-

spread geographically, occurring in six or eight forest

formations, and semifossorial or terrestrial/arboreal, sometimes

escaping human notice). The most vulnerable reptile is

Ctenosaura bakeri, 5-8-6 species (known only from the vicini-

ty of the type locality, in one forest formation, and used for its

meat and eggs locally). The next most vulnerable is Crenosaura

oedirhina, a 4-8-6 species (a Honduran endemic, occurring in

one forest formation, and used for its meat and eggs locally).

As for the amphibians, we have divided the species of

Honduran reptiles into three categories of environmental vul-

nerability, as indicated in Table 2. As above, this

categorization is intended as a coarse gauge as to the degree of

attention that should be brought to bear on the various species.

There are 47 low vulnerability species, making up only 22.3%

of the Honduran reptilian fauna. A slightly larger group of 53

species, making up 25.1% of the taxa, comprises the high vul-

nerability category. Many of these species (35) are endemic to

Honduras. The largest group of 111 species, as with the

amphibians, is composed of taxa of intermediate vulnerability

(52.6% of total). Most of these species (93) are geographical-

ly widespread, although in many cases occurring peripherally

to Honduras, and many (66) are known from only one or two

forest formations.

Volume 3 | Number 1 | Page 16

The conservation status of the herpetofauna of Honduras

Table 1. Environmental vulnerability scores (EVS) for the 117 species of amphibians of Honduras. Numbers for each gauge explained in text.

The table is broken into three parts: low vulnerability species (EVS of 3-9; 24 species; 20.5%); medium vulnerability species (EVS of 10-13;

43 species; 36.8%); and high vulnerability species (EVS of 14-17; 50 species; 42.7%). Updated from Table 33 in McCranie and Wilson (2002).

Amphibian Species

Geographic

Distribution

Ecological

Distribution

Low

Bolitoglossa mexicana

Bufo coccifer

Bufo luetkenii

Bufo marinus

Bufo valliceps

HAyalinobatrachium fleischmanni

Hyla loquax

Hyla microcephala

Ayla picta

Phrynohyas venulosa

Plectrohyla guatemalensis

Ptychohyla hypomykter

Scinax staufferi

Smilisca baudinii

Eleutherodactylus laevissimus

Leptodactylus labialis

Leptodactylus melanonotus

Physalaemus pustulosus

Hypopachus variolosus

Rana berlandieri

Rana forreri

Rana maculata

Rana vaillanti

Rhinophrynus dorsalis

Medium

Dermophis mexicanus

Gymnopis multiplicata

Bolitoglossa rufescens complex

Oedipina cyclocauda

Atelophryniscus chrysophorus

Bufo campbelli

Bufo haematiticus

Bufo leucomyos

Centrolene prosoblepon

Cochranella albomaculata

Cochranella granulosa

Cochranella spinosa

Hyalinobatrachium pulveratum

Agalychnis calcarifer

Agalychnis callidryas

Agalychnis moreletii

Agalychnis saltator

Duellmanohyla salvavida

Duellmanohyla soralia

Hyla catracha

Hyla ebraccata

Plectrohyla chrysopleura

Plectrohyla dasypus

Plectrohyla exquisita

Plectrohyla hartwegi

Plectrohyla matudai

Plectrohyla psiloderma

a cc eee ce ce SS ce ce ce ee ee ee ce ed

WwWwWRRHYE BPWENNKFNNNYNNNFSNN PR WHe

NTYnF DR fPWWWWNWYANHAWAIW fFOWWN ff

ADA WAWAiAIANA RAIMA AAA AIYIMAAYAYNYAYNAADAAAYNA N~ ~4

Reproductive Total

Mode Score

SS SS SS SW) BS SS SSS is ee

OANDAWAAAAABHMAOUONOADWOSA AN DA Oo

pe es fe a ys US) Sy Sh ey US) US) (OS) WS WS) WY) Wey WO yy Sy SN IS IS IN

Amphib. Reptile Conserv. | http://www.herpetofauna.org

Continued on page 18.

Volume 3 | Number 1 | Page 17

L. D. Wilson and J. R. McCranie

Table 1. Continued.

Geographic Ecological Reproductive Total

Amphibian Species Distribution Distribution Mode Score

Ptychohyla salvadorensis 3 7 1 11

Ptychohyla spinipollex 4 6 1 11

Scinax boulengeri 2 8 1 11

Smilisca phaeota 2 7 1 10

Smilisca sordida 2 8 1 11

Triprion petasatus 3 8 1 12

Eleutherodactylus charadra 3 6 4 13

Eleutherodactylus fitzingeri 2 7 4 13

Eleutherodactylus mimus D 7 4 13

Eleutherodactylus noblei 2 7 4 13

Eleutherodactylus ridens 1 7 4 12

Leptodactylus pentadactylus 2 7 2 11

Leptodactylus silvanimbus 4 7 2 13

Gastrophryne elegans 2 8 1 11

Hypopachus barberi 3 7 1 11

Rana warszewitschii 2 8 1 11

High

Bolitoglossa carri 5 8 4 17

Bolitoglossa celaque 4 8 4 16

Bolitoglossa conanti 3 7 4 14

Bolitoglossa decora S) 8 4 17 |

Bolitoglossa diaphora 4 8 4 16 )

Bolitoglossa dofleini 3 7 4 14

Bolitoglossa dunni 3 7 4 14

Bolitoglossa longissima 5 8 4 17

Bolitoglossa occidentalis 2 8 4 14

Bolitoglossa porrasorum 4 7 4 15

Bolitoglossa striatula 2 8 4 14

Bolitoglossa synoria 3 8 4 15

Cryptotriton nasalis 4 7 4 15 |

Dendrotriton sanctibarbarus 4 8 4 16 |

Nototriton barbouri 4 7 4 15 |

Nototriton lignicola 5) 8 4 17

Nototriton limnospectator 4 8 4 16

Oedipina elongata 3 8 4 15

Oedipina gephyra 4 8 4 16

Oedipina ignea 3 7 4 14

Oedipina stuarti 4 7 4 15

Oedipina taylori 3 8 4 15

Hyalinobatrachium cardiacalyptum 4 7 3 14

Hyalinobatrachium crybetes 5) 7 3 15 |

Anotheca spinosa D 8 5 15

Hyla bromeliacia 3 i] 5 IS

Hyla insolita 5 8 3 16

Hyla salvaje 3 8 S) 16

Eleutherodactylus anciano 4 7 4 15

Eleutherodactylus aurilegulus 4 6 4 14

Eleutherodactylus chac 3 7 4 14 |

Eleutherodactylus chrysozetetes 5 8 4 17 |

Eleutherodactylus coffeus 5 8 4 17

Eleutherodactylus cruzi 5 8 4 17 |

Eleutherodactylus emleni 4 6 4 14 |

Eleutherodactylus epochthidius 4 7 4 15

Eleutherodactylus fecundus 4 wf 4 15

: Continued on page 20. |

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 18

Se OU

rig ee Me te

Plate 21 DOI: 10.1514/journal.a arc.0000012.g021

Rages as RTTET a me

Plate 22 DOI: 10. 1514/journal arc. 0000012. 9022 Plate 23 DOI: 10.1514/journal.arc.0000012.g023

Plate a . DOI: 10.1514/journal.arc.0000012.g025

Plate 27 DOI: 10. Tenanolinnal! arc. 0000012. 9027

Plate captions: 20. Oedipina gephyra. 21. Atelophryniscus chrysophorus. 22. Duellmanohyla salvavida. 23. Hyla catracha. 24. Plectrohyla

chrysopleura. 25. Eleutherodactylus epochthidius. 26. Eleutherodactylus fecundus. 27. Eleutherodactylus pechorum. Plates 20 through 27.

Honduran endemics with all known populations believed to be declining.

DOI: 10.1514/journal.arc.0000012.9026

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 19

L. D. Wilson and J. R. McCranie

Table 1. Continued.

Geographic Ecological Reproductive Total

Amphibian Species Distribution Distribution Mode Score

Eleutherodactylus laticeps 2 8 4 14

Eleutherodactylus lauraster 3 7 4 14

Eleutherodactylus loki 2 8 4 14

Eleutherodactylus megacephalus 2 8 4 14

Eleutherodactylus merendonensis >) 8 4 17

Eleutherodactylus milesi 4 7 4 15

Eleutherodactylus olanchano 4 8 4 16

Eleutherodactylus omoaensis 4 8 4 16

Eleutherodactylus operosus 4 7 4 15

Eleutherodactylus pechorum 4 7 4 15

Eleutherodactylus rostralis 3 I 4 14

Eleutherodactylus saltuarius + 8 4 16

Eleutherodactylus stadelmani 4 7 4 15

DOI: 10.1514/journal.arc.0000012.t001

Categorization of EVS provides a means to assign con-

servation priorities, with high vulnerability species given

highest priority, medium vulnerability species intermediate

priority, and low vulnerability species lowest priority. The

highest priority taxa include 50 amphibians and 53 reptiles

(total of 103 species or 31.4% of 328 total species); the inter-

mediate priority taxa consist of 43 amphibians and 111

reptiles (total of 154 species or 47.0%); and the low priority

taxa comprise 24 amphibians and 47 reptiles (total of 71

species or 21.6%).

Current population status of members of the

Honduran herpetofauna

The above discussion attempts to assign conservation priori-

ties to the members of the Honduran herpetofauna on a largely

theoretical basis, with the assumption that there are features of

distribution (geographic and ecological), life history (repro-

ductive mode), and human persecution that can act as a rough

gauge of vulnerability to anthropogenic environmental pres-

sures, in a similar manner as has been done for threatened and

endangered species in general (see Raven and Berg 2001 for a

discussion of such features).

As noted in a previous section, however, there are factors

at work in Honduras, as elsewhere in the world, the effect of

which were not predicted by the typical models of species

endangerment. The unanticipated factors apparently of great-

est importance are chytridiomycosis (Berger et al. 1998) and

climatic warming (Pounds et al. 1999), although neither has

been conclusively demonstrated to be in effect in Honduras.

Whatever the causative factors that may be involved, it is

apparent that populations of many members of the Honduran

herpetofauna are in decline or have disappeared since the

early years of the 1990s (Wilson and McCranie 1998,

McCranie and Wilson 2002, in press). The declines have been

substantiated best among amphibian populations.

Unfortunately, these declines have involved the two most

important groups of amphibians, those endemic to Honduras

and those otherwise restricted to Nuclear Middle America

Amphib. Reptile Conserv. | http://www.herpetofauna.org

(Table 3). As noted by McCranie and Wilson (in press), of the

41 species of endemic amphibians, six are feared extinct and

14 appear to have declining populations (field work in 2001

indicated that one of the 14 species, Eleutherodactylus stadel-

mani, thought to be in decline by McCranie and Wilson, in

press, has also disappeared). In addition, we have no data for

four species. Only 17 species appear to have stable popula-

tions. Thus, 20 of the 41 endemic species of Honduran

amphibians (48.8%), or almost half, are already gone or are in

decline.

The seven endemic amphibian species feared extinct

have EVS ranging between 14 and 17 (mean 15.6). The 13

species whose populations are in decline have EVS from 12 to

17 (mean 14.4). Of considerable interest is the fact that the

EVS for the 17 endemics thought to have stable populations

range from 11 to 17, with a mean value of 15.0. The implica-

tion of these data are that there is an urgent need to monitor

populations of these supposed “stable” species, because 14 of

the 17 have scores indicative of high vulnerability to environ-

mental pressures.

McCranie and Wilson (in press) also discussed the pop-

ulation status of 25 amphibian species not endemic to

Honduras, but restricted in distribution to Nuclear Middle

America. They considered nine species to be in decline and

one to probably have been extirpated. The EVS of the nine in

decline range from nine to 16 (mean 12.1). The one species

thought extirpated (Bolitoglossa occidentalis) has an EVS of

14. These data indicate that EVS of 13 and above are indica-

tive of species that need to be monitored, but that scores below

that level do not insulate a species from anthropogenic pres-

sure. As we have noted above, there is no species of Honduran

amphibian safe from human depredation, although there are

clearly some species capable of persisting as commensals of

human beings.

The picture for Honduran reptiles is somewhat less clear.

This is due to the fully terrestrial life cycle of most reptiles,

which allows for habitation of niches removed from water, in

turn increasing the potential breadth of occurrence.

Nonetheless, it is possible to comment on the current popula-

Volume 3 | Number 1 | Page 20

a ee eee: ie

The conservation status of the herpetofauna of Honduras

Table 2. Environmental vulnerability scores (EVS) for the 211 species of reptiles of Honduras (marine species are not included). Numbers

for each gauge explained in text. The table is broken into three parts: low vulnerability species (EVS of 4-9; 47 species; 22.3%); medi-

um vulnerability species (EVS of 10-13; 111 species; 52.6%); and high vulnerability species (EVS of 14-19; 53 species; 25.1%).

Geographic Ecological Human Total

Reptilian Species Distribution Distribution Persecution Score

Low

Rhinoclemmys pulcherrima

Kinosternon leucostomum

Kinosternon scorpioides

Sphaerodactylus millepunctatus

Basiliscus vittatus

Laemanctus longipes

Sceloporus malachiticus

Sceloporus variabilis

Norops cupreus

Norops laeviventris

Norops lemurinus

Norops sericeus

Norops tropidonotus

Mabuya unimarginata

Sphenomorphus cherriei

Gymnophthalmus speciosus

Ameiva undulata

Cnemidophorus deppii

Cnemidophorus motaguae

Leptotyphlops goudotii

Boa constrictor

Adelphicos quadrivirgatus

Coniophanes fissidens

Conophis lineatus

Dryadophis melanolomus

Drymarchon melanurus

Drymobius margaritiferus

Enulius flavitorques

Hydromorphus concolor

Imantodes cenchoa

Lampropeltis triangulum

Leptodeira annulata

Leptodeira septentrionalis

Leptophis ahaetulla

Leptophis mexicanus

Ninia diademata

Ninia sebae

Oxybelis aeneus

Rhadinaea godmani

Sibon nebulatus

Spilotes pullatus

Storeria dekayi

Tantilla melanocephala

Thamnophis proximus

Tretanorhinus nigroluteus

Micrurus nigrocinctus

Porthidium ophryomegas

SoS oS eS SS aS eS SSeS Se SS SS SS SP Se Se Se Se Se Se SSS Se Se Se SS eS See Se Se oS See Se eS

WWNMEDHDEUNDHKHKH NHAWWEWWHWWAWNHKK HK KRNHWUHAUNEWRWWHEWUHNNWEUNWWUUAN

AmnstnnFsFtvnrnrFvnwnFFFPUNWNnwnnskPHHPHYW FH WW WW WW WWW WW WW WW WY W W W

OOWOOOUOWOOFL WA WDWOMWAODAOAIOOCOOWMOHOHNIOAY MWY IYMATYO WOON WOMONANWO WO WO

Medium

Crocodylus acutus 1 6 6 13

Chelydra serpentina 1 6 6 13

Rhinoclemmys annulata Dy, 8 3 13

Rhinoclemmys areolata Dy) 7 3 1

Continued on page 24.

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 21

Plate 28 DOI: 10.151 Afisuinnelh arc. 0000012 2. 028 Plate 29 DOI: 10.1514/journal.arc.0000012.g029

Plate 30 DOI: 10. 151 4fjournal.a arc. c.000001 2. 9030 Plate 31 DOI: 10.1514/journal.arc.0000012.9031

Plate 32

Plate 34 DOI: 10.151 4/journal.arc.0000012.g034 Plate 35 DOI: 10. ‘eyeucal arc.0000012. 9035

Plate captions: 28. Eleutherodactylus saltuarius. 29. Leptodactylus silvanimbus. 30. Abronia salvadorensis. 31. Norops kreutzi. 32. Norops

muralla. 33. Norops ocelloscapularis. 34. Norops wampuensis. 35. Typhlops stadelmani. Plates 28 through 35. Honduran endemics with all

known populations believed to be declining. a

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 22

ey ee BE ie

Plate 36 DOI: 10.1514/journal.arc.0000012.g036 _—~Plate 37 DOK: 10.1514/journal.arc.0000012.g037

~*~

X %®

Plate 38 DOI: 10.1514/journal.arc.0000012.g038 Plate 39 DOI: 10.151 4/journal.arc.000001 2.9039

Plate40 =~ DOI: 10.1514/journal.arc.0000012.g040 _—S~PPlatte 44 DOI: 10.1514/journal.arc.0000012.9041

Er al 7 ree - ai a tt 54 Ty

Plate 42

~ ae -

DOI: 10.1514/journal.arc.0000012.9042 Plate 43 DOI: 10.1514/journal.arc.0000012.g043

Plate captions: 36. Enulius bifoveatus. 37. Tantilla tritaeniata. 38. Bothriechis marchi. 39. Bolitoglossa dofleini. 40. Bolitoglossa synoria. 41.

Duellmanohyla soralia. 42. Plectrohyla guatemalensis. 43. Plectrohyla matudai. Plates 36 through 38. Honduran endemics with all known pop-

ulations believed to be declining. Plates 39 through 43. Nuclear Middle American Restricted Species with all known Honduran populations

believed to be declining.

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 23

Table 2. Continued.

L. D. Wilson and J. R. McCranie

Ecological Human Total

Distribution Persecution Score

Geographic

Reptilian Species Distribution

Trachemys scripta 1

Celestus bivittatus

Mesaspis moreletii

Coleonyx mitratus

Aristelliger georgeensis

Aristelliger praesignis

Gonatodes albogularis

Hemidactylus brookii

Hemidactylus frenatus

Hemidactylus mabouia

Phyllodactylus tuberculosus

Sphaerodactylus glaucus

Sphaerodactylus notatus

Thecadactylus rapicauda

Basiliscus plumifrons

Corytophanes cristatus

Corytophanes hernandesti

Laemanctus serratus

Ctenosaura flavidorsalis

Ctenosaura similis

Iguana iguana

Leiocephalus carinatus

Sceloporus squamosus

Anolis allisoni

Norops biporcatus

Norops capito

Norops crassulus

Norops humilis

Norops limifrons

Norops lionotus

Norops pentaprion

Norops petersit

Norops rodriguezii

Norops sagrei

Norops uniformis

Polychrus gutturosus

Eumeces sumichrasti

Mesoscincus managuae

Sphenomorphus assatus

Sphenomorphus incertus

Ameiva ameiva

Ameiva festiva

Cnemidophorus lemniscatus

Lepidophyma flavimaculatum

Typhlops costaricensis

Typhlops stadelmani

Loxocemus bicolor

Corallus annulatus

Ungaliophis continentalis

Alsophis cantherigerus

Amastridium veliferum

Chironius grandisquamis

Clelia clelia

Coniophanes bipunctatus

Coniophanes imperialis

Re eRe ENN WRK BNR RF KE NNNNF RF NNNNYK NNN WRN YF NRF RP WNNK NRF NNYE NNN YK NN W WW

Amphib. Reptile Conserv. | http://www.herpetofauna.org

NNDANAIWAAIWHAHiAIAWAAAAWAIMWAIANAAMANMA AIA AYAANANIACAOMFAIAAYATNAANAWAAAAAAAAAAA WSN ~~)

10 |

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Continued on page 25. |

BHP HPNWNYHY FH KH FW WW WW WWW WW WW WWW WW WW WWW DAD WWW WW BPW WWW WW WWW BWWAD

Volume 3 | Number 1 | Page 24

ee DE ee:

Table 2. Continued.

Reptilian Species

Geographic

Distribution

Ecological

Distribution

The conservation status of the herpetofauna of Honduras

Human Total

Persecution Score

Coniophanes piceivittis

Dendrophidion nuchale

Dendrophidion percarinatum

Dipsas bicolor

Dryadophis dorsalis

Drymobius chloroticus

Elaphe flavirufa

Erythrolamprus mimus

Ficimia publia

Geophis fulvoguttatus

Geophis hoffmanni

Imantodes gemmistratus

Imantodes inornatus

Leptodeira nigrofasciata

Leptodrymus pulcherrimus

Masticophis mentovarius

Ninia espinali

Ninia maculata

Nothopsis rugosus

Oxybelis brevirostris

Oxybelis fulgidus

Oxyrhopus petola

Pliocercus elapoides

Pseustes poecilonotus

Rhadinaea kinkelini

Rhadinaea lachrymans

Rhadinaea montecristi

Scaphiodontophis annulatus

Senticolis triaspis

Sibon carri

Sibon dimidiatus

Sibon longifrenis

Stenorrhina degenhardtii

Stenorrhina freminvillei

Tantilla impensa

Tantilla lempira

Tantilla schistosa

Tantilla taeniata

Tantillita lintoni

Thamnophis marcianus

Trimorphodon biscutatus

Tropidodipsas fischeri

Tropidodipsas sartorii

Urotheca guentheri

Xenodon rabdocephalus

Micrurus diastema

Atropoides nummifer

Bothriechis schlegelii

Bothrops asper

Cerrophidion godmani

Crotalus durissus

Porthidium nasutum

High

Caiman crocodilus

Amphib. Reptile Conserv. | http://www.herpetofauna.org

1 ee

SSS Ss Se ies oS G9) eS td CSS ES COS SH) iS DS] WH) WD DOS SSS OW PD Wis Ss Ss aS WH Wis Ss SHS we SS

oO’

NDNDADADAAMNAAAWAAAYTMA AMAA AYAYANYNAMA YADA AYMAAPDAIAAYANFAIYMNAATNWAOAADADMAAYAAAIAYNAAANA N~> xn ~

MANN nnrnrarnurnst FPnwonwnnnnrnst F$nwmnrFnwmFwrnNMNnwnnwnmFse urns snvwvnwnnsks$FHnwnnnnvnwnws HPnwvosrs

Continued on page 27.

Volume 3 | Number 1 | Page 25

Plate 44

c.0000012.g045

< ex

ies

DOI: 10.151 4/journal.arc.0000012.g044 Plate 45 DOI: 10.1514/journal.ar

a - a

Plate 46 DOI: 10.1514/journal.arc.0000012.g046

Plate 47 DOI: 10.1514/journal.arc.0000012.g047

a . ae. ee

" Lise Phi) pe ee o's iad af ‘a

i . 2 : : : g = rn fea om : — si : : Se 4° Ne 2 Sina > : — -~

Plate 48 DO}: 10.151 4/journal.arc.0000012.g048 Plate 49 DOI: 10.1514/journal.arc.0000012.g049

, a

Plate 50 DOI: 10.151 4/journal.arc.000001 2.9050

Plate captions: 44. Plectrohyla psiloderma. 45. Ptychohyla hypomykter. 46. Abronia montecristoi. 47. Celestus bivittatus. 48. Corytophanes

percarinatus. 49. Tropidodipsas fischeri. 50. Bothriechis thalassinus. Plates 44 through 50. Nuclear Middle American Restricted Species with

all known Honduran populations believed to be declining.

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 26

The conservation status of the herpetofauna of Honduras

Table 2. Continued.

Geographic Ecological Human Total

Reptilian Species Distribution Distribution Persecution Score

Rhinoclemmys funerea 2 8 6 16

Staurotypus triporcatus 2 7 6 IS

Abronia montecristoi 3 8 4 15

Abronia salvadorensis 4 8 + 16

Celestus montanus 4 7] 3 14

Celestus scansorius 4 7 3 14

Phyllodactylus palmeus 4 8 3 15

Sphaerodactylus dunni 4 7 3) 14

Sphaerodactylus rosaurae 4 8 3 15

Corytophanes percarinatus 3 8 3 14

Ctenosaura bakeri 5 8 6 19

Ctenosaura melanosterna 4 7 6 17

Ctenosaura oedirhina 4 8 6 18

Norops amplisquamosus 5) 8 3 16

Norops bicaorum 5) 8 3 16

Norops cusuco 5 8 3 16

Norops heteropholidotus 3 8 3 14

Norops johnmeyeri 4 8 3 15

Norops kreutzi 5) 8 3 16

Norops loveridgei 4 7 3 14

Norops muralla 4 8 3 15

Norops ocelloscapularis 5 7 3 15

Norops pijolensis 4 7 3 14

Norops purpurgularis 4 8 3 15

Norops roatanensis 4 8 3 15)

Norops rubribarbaris 5 8 3 16

Norops sminthus 4 8 3 15

Norops utilensis 5 8 3 16

Norops wampuensis 5) 8 3 16

Norops yoroensis 4 7 3 14

Norops zeus 4 7 3 14

Crisantophis nevermanni 2 8 4 14

Drymobius melanotropis 2 8 4 14

Enulius bifoveatus 5 8 2 15

Enulius roatanensis 5 8 2 IS)

Geophis damiani 5) 8 2 15

Leptophis modestus 3 8 4 15

Leptophis nebulosus 2 8 4 14

Omoadiphas aurula >) 8 2 15

Oxybelis wilsoni 4 8 3 15

Rhadinaea tolpanorum 5 8 2) 15

Rhinobothryum bovallii 2 8 5 15

Scolecophis atrocinctus D, 7 5) 14

Sibon anthracops 1 8 5) 14

Tantilla tritaeniata 5 8 2 15

Thamnophis fulvus 3 7 4 14

Micrurus alleni 2 8 5 15

Micrurus browni 2 8 5 15

Micrurus ruatanus 4 8 5) 17/

Agkistrodon bilineatus 2 8 5) 15

Bothriechis marchi 4 7 > 16

Bothriechis thalassinus 3 7 5) 15

’ Based on specimens without precise locality data and one sight record in the Middle Choluteca Valley.

? However, this species is extirpated on the Swan Islands, the only place where this species is known in Honduras.

DOI: 10.1514/journal.arc.0000012.t002

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 27

L. D. Wilson and J. R. McCranie

Table 3. Current status of populations of Honduran amphibian endemics and species otherwise restricted to Nuclear Middle

America. Stable = at least some populations stable; Declining = all populations believed to be declining. Extinct category applies to

Honduran endemics; extirpated category applies to Nuclear Middle American endemics (excluding those endemic to Honduras).

Extinct or

Species Stable Declining Extirpated No Data

Honduran endemics

Bolitoglossa carri x

Bolitoglossa celaque

Bolitoglossa decora

Bolitoglossa diaphora

Bolitoglossa longissima

Bolitoglossa porrasorum

Cryptotriton nasalis

Dendrotriton sanctibarbarus

Nototriton barbouri

Nototriton lignicola

Nototriton limnospectator

Oedipina gephyra x

Oedipina stuarti x

Atelophryniscus chrysophorus x

Bufo leucomyos

Hyalinobatrachium cardiacalyptum

Hyalinobatrachium crybetes x

Duellmanohyla salvavida x

Hyla catracha x

Hyla insolita x

Plectrohyla chrysopleura x

Plectrohyla dasypus x

Plectrohyla exquisita x

Ptychohyla spinipollex x

Eleutherodactylus anciano x

Eleutherodactylus aurilegulus x

Eleutherodactylus chrysozetetes x |

Eleutherodactylus coffeus x |

Eleutherodactylus cruzi x

Eleutherodactylus emleni X

Eleutherodactylus epochthidius

Eleutherodactylus fecundus

Eleutherodactylus merendonensis

Eleutherodactylus milesi x |

Eleutherodactylus olanchano x |

Eleutherodactylus omoaensis x

Eleutherodactylus operosus x

Eleutherodactylus pechorum x

Eleutherodactylus saltuarius x

Eleutherodactylus stadelmani x :

Leptodactylus silvanimbus x

KX KM KX KO OK

x xX

KK XM

Honduran species otherwise restricted to Nuclear Middle America

Bolitoglossa conanti x

Bolitoglossa dofleini xX

Bolitoglossa dunni x

Bolitoglossa occidentalis x

Bolitoglossa rufescens complex x

Bolitoglossa synoria x

Oedipina elongata x

Oedipina ignea x

Oedipina taylori xX

Duellmanohyla soralia x |

Continued on page 29.

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 28 |

a ee ee eS aa eee

The conservation status of the herpetofauna of Honduras

Table 3. Continued.

Extinct or

Species’ . Stable Declining Extirpated No Data

Hyla bromeliacia x

Ayla salvaje x

Plectrohyla guatemalensis x

Plectrohyla hartwegi x

Plectrohyla matudai

Plectrohyla psiloderma

Ptychohyla hypomykter

Ptychohyla salvadorensis

Triprion petasatus x

Eleutherodactylus chac

Eleutherodactylus charadra

Eleutherodactylus lauraster

Eleutherodactylus rostralis

Hypopachus barberi

Rana maculata

x x

Kx mK KM MX

DOI: 10.1514/journal.arc.0000012.t003

Table 4. Current status of populations of Honduran reptile endemics and species otherwise restricted to Nuclear Middle America.

Stable = at least some populations stable; Declining = all populations believed to be declining. Extinct category applies to Honduran

endemics; extirpated category applies to Nuclear Middle American endemics (excluding those endemic to Honduras).

Extinct or

Species Stable Declining Extirpated No Data

Honduran endemics

Abronia salvadorensis x

Celestus montanus x

Celestus scansorius x

Phyllodactylus palmeus

Sphaerodactylus dunni

Sphaerodactylus rosaurae

Ctenosaura bakeri

Ctenosaura melanosterna

Ctenosaura oedirhina

Norops amplisquamosus

Norops bicaorum

Norops cusuco

Norops johnmeyeri

| Norops kreutzi x

Norops loveridgei

Norops muralla x

Norops ocelloscapularis x

Norops pijolensis

Norops purpurgularis

Norops roatanensis

Norops rubribarbaris x

Norops sminthus x

Norops utilensis x

Norops wampuensis x

Norops yoroensis x

Norops zeus x

Typhlops stadelmani x

Enulius bifoveatus x

Enulius roatanensis x

Geophis damiani x

Omoadiphas aurula x

Continued on page 30.

mM KM PM OM OO OK OS OP

mm PX

Amphib. Reptile Conserv. | http://www.herpetofauna.org

Volume 3 | Number 1 | Page 29

L. D. Wilson and J. R. McCranie

Table 4. Continued.

Stable

Oxybelis wilsoni x

Rhadinaea tolpanorum

Tantilla lempira

Tantilla tritaeniata

Micrurus ruatanus

Bothriechis marchi

Species

Honduran species otherwise restricted to Nuclear Middle America

Abronia montecristoi

Celestus bivittatus

Mesaspis moreletii

Corytophanes percarinatus

Ctenosaura flavidorsalis

Norops crassulus

Norops heteropholidotus

Sphenomorphus incertus

Ungaliophis continentalis

Dryadophis dorsalis

Geophis fulvoguttatus

Leptophis modestus

Ninia espinali

Rhadinaea kinkelini

Rhadinaea lachrymans

Rhadinaea montecristi x

Sibon carri

Tantilla impensa

Tantilla taeniata

Thamnophis fulvus x

Tropidodipsas fischeri

xxx KM XK

Extinct or

Declining Extirpated No Data

BotiriecWis SNGUASSUMUS Te

DOI: 10.1514/journal.arc.0000012.t004

tion status of reptiles endemic to Honduras or otherwise

restricted to Nuclear Central America. Thirty-seven species of

reptiles are endemic to Honduras (Table 4). Of these 37

species, only 19 species (51.4%) are thought to have stable

populations. Ten (27.0%) are considered to have declining

populations, primarily on the basis of destruction of habitat

within their ranges. Finally, eight species (21.6%) are poorly

known enough so that we are uncertain of their status.

The ten endemic reptile species considered to have

declining populations have EVS ranging between 12 and 16

(mean 14.9). The EVS for the 19 endemics thought to have

stable populations range from 14 to 19 (mean 15.4), which is

higher than the mean for those species thought to have declin-

ing populations. It is interesting that the reptilian endemics

thought to have stable populations also have a higher mean

EVS than those thought to have declining populations. The

implication of these data is same as that for the analogous data

for amphibians. The populations of these endemics need to be

monitored carefully, inasmuch as all have scores indicating

high vulnerability to environmental pressures.

We also determined the population status for those rep-

tile species not endemic to Honduras but restricted in

Amphib. Reptile Conserv. | http://www.herpetofauna.org

Pad: - F ee ee | ay

distribution to Nuclear Middle America. Of these 22 species,

only eight (36.4%) are considered to have stable populations,

at least somewhere in their known ranges in Honduras.

Twelve species (54.5%) are thought to have declining popula-

tions. Finally, two species (9.1%) are too poorly known to

judge their current population status.

The 12 Nuclear Middle American reptile species that

appear to have declining populations have EVS ranging

between ten and 15 (mean 12.8). Following the same pattern

as indicated above, the EVS for the eight species appearing to

have stable populations range from 12 to 14 (mean 12.9),

which is slightly higher than the mean for the declining popu-

lation Nuclear Middle American species. The populations of

these species also need to be closely monitored.

In general, it should be understood that the population sta-

tus of amphibian and reptile species in Honduras potentially

can change relatively rapidly. As habitats are degraded, the

fabric of community structure unravels. The community inhab-

itants depend on the integrity of this structure in order to obtain

the materials and energy necessary to support their life

processes. Thus, they are links in biogeochemical cycles and

food webs, through which these materials and energy move,

Volume 3 | Number 1 | Page 30

The conservation status of the herpetofauna of Honduras

respectively. Thus, for example, given that amphibian popula-

tions are undergoing apparent increasing decline, this can be

expected to adversely affect the populations of amphibian-eat-

ing snakes. In turn, decline of these snake populations should

affect the populations of ophiophagous snakes, and so on. Thus

does the straight edge of much human thinking cut deeply.

Plates 2-14 show some of the primary forest left in

Honduras, plus some of the extensive deforestation taking

place in the country. Plates 15-18 show some Honduran

endemic species now feared extinct. Plates 19-38 show some

of the Honduran endemic species in which all known popula-

tions are believed to be declining. Finally, plates 39-50 show

some of the Nuclear Middle America-restricted species

(exclusive of the Honduran endemics) in which all known

populations are believed to be declining.

Recommendations

Biodiversity decline is one of the most serious environmental

problems, if not the most serious (Wilson et al. 2001). Since it

is a problem, it cries out for solutions. Unfortunately, one of

the tenets of the problem solving critical thinking strategy (see

Chaffee 1994 for a description of the strategy) is that a prob-

lem cannot be solved by simply treating its symptoms.

Biodiversity decline is a symptom of habitat loss and degra-

dation, in turn a symptom of runaway human population

growth. Uncontrolled population growth is, in turn, a symp-

tom of the mismanaged human mind, to use a phrase coined

by E. O. Wilson (1988). The “cascade of deeper problems

arising within the human psyche” (Wilson et al. 2001, p. 109)

referred to by E. O. Wilson (1988) has been explored at length

by L. D. Wilson in a series of papers (1997 a, b, 1998, 1999,

2000, 2001). L. D. Wilson (2001) concluded, after a lengthy

argument presented in this series, that the sustainable society

described by the better environmental science texts (see for

example Miller 2001, and Raven and Berg 2001) will only

come about (if it ever does) by a fundamental reform of the

educational process, so as to enable us to use education as a

kind of species-wide psychotherapy. This view, then, treats

the “mismanagement of the human mind” (E. O. Wilson

1988) as a pervasive psychological illness in need of broad-

based therapy.

Until and unless the “mismanaged human mind” is treat-

ed successfully, then we argue that none of the problems that

cascade from it, which are, after all, the persistent problems of

humankind, will ever encounter workable and lasting solu-

tions. Having said this, then it must be understood that the

recommendations we outline below will only work if the geo-

metrically advancing problems of uncontrolled human

population growth and its corollaries, habitat loss and degra-

dation, are solved. If not, then the exercise below is merely a

monument to futility.

Given the above, we have to assume that it is possible to

guard the integrity of established biotic reserves in Honduras.

Based on our decades-long field experience, this is only hap-

pening in a limited way. It is still the case that most biotic

reserves in the country exist only on paper, without the

appropriate resources dedicated to establish boundaries, hire

personnel to police them, build facilities for housing admin-

istrative, scientific, and security personnel, and fund the

Amphib. Reptile Conserv. | http://www.herpetofauna.org

scientific studies necessary to make such reserves sustain-

able. This situation will have to change and change rapidly,

for the pressure of a 25-year doubling time will brook no

idleness.

It is also evident that we have been idle too long, and that

the study of the Honduran herpetofauna has turned a corner

into a torturous maze from which there is no easy exit. It is

already clear, as is discussed above, that a new era has been

breached—one in which advances in our cataloguing of the

herpetodiversity of Honduras is being offset by documented

losses of that same diversity over the last decade or so. We

are, thus, fighting an uphill battle on very slippery slopes.

In full light of the provisos identified in this section

above, the following recommendations concerning the protec-

tion of the members of the Honduran herpetofauna are made:

e The system of biotic reserves should be expanded to

include areas for protection of species not currently

known to reside in any legally established reserve. The

locations of such areas are discussed by Wilson et al.

(2001) and McCranie and Wilson (2002). Of the

Honduran endemics, there are 14 such species. For the

Nuclear Middle American species, seven species are

involved.

The entire system should be evaluated to ascertain the

health of the populations of amphibians and reptiles

resident within the various reserves. At least an initial

effort can be accomplished by use of Rapid Ecological

Assessment Program methodology (see Parker and

Bailey 1991).

Following this evaluation, the system of reserves

should be adjusted to the extent possible to provide

maximal protection of the remaining populations of

resident amphibians and reptiles. Undoubtedly, this

step also would involve establishment of additional

reserves. Wilson et al. (2001) and McCranie and

Wilson (2002) provide some guidance for such deci-

sions.

Steps then should be taken to clearly identify the lim-

its of the reserves, build facilities to house personnel,

involve local people in planning and decision making,

make employment available to local people, and put

the resulting revenues into local communities for

future improvements. Meyer and Meerman (2001) dis-

cussed this type of “participatory” management strate-

gy, which they advocate to replace the traditional

“exclusionary” management strategy maintained by

them to be ineffective over the long term. These steps,

which need to occur as rapidly as possible, will obvi-

ously require appropriate allocation of governmental

funds. The administration of the new Honduran presi-

dent, Ricardo Maduro Joest, is just beginning. It

remains to be seen what priority is established by the

new government to address these issues.

Once facilities are available for housing personnel,

then the longer-term scientific survey work and other

sorts of scientific studies can begin, with the goal of

establishing the biological worth of the various

reserves. Opportunities for cooperation in such studies

between resident and foreign scientists should be

Volume 3 | Number 1 | Page 31

L. D. Wilson and J. R. McCranie

explored. We continue to explore such collaborations

with various Honduran biologists.

¢ With completion of facilities and scientific studies can

come educational and ecotourist programs, with the

goal of making the reserves economically self support-

ing. Again, cooperative undertakings should be

encouraged. Such steps would involve reaching out to

various Honduran and foreign governmental and non-

governmental organizations.

¢ Our strongest recommendation is that the steps out-

lined above be taken with all dispatch possible. We

have demonstrated that populations of a highly signif-

icant number of species of Honduran amphibians and

reptiles are already in decline or have disappeared,

especially of the most important segment containing

the endemic species and those whose distribution is

otherwise restricted to Nuclear Middle America. In

addition, deforestation has been demonstrated to be

increasing at an exponential rate, commensurate with

the increase in human population. Deforestation is the

principal type of habitat destruction in Honduras,

which is, in turn, the major threat to the highly distinc-

tive and important Honduran herpetofauna. There is, in

the final analysis, no time to dawdle.

“We must learn to use our intelligence to live more

lightly on the land, so that we do not degrade the only

home we have—and the only one we can leave to our chil-

dren.”

E. O. Wilson and D. L. Perlman

Conserving Earth’s Biodiversity, 2000

Acknowledgments.—Over the decades that we have

worked on the herpetofauna of Honduras, we have incurred

indebtedness to uncounted individuals and organizations. First

and foremost, this work would not have been possible without

the support of the personnel of Recursos Naturales

Renovables and Corporaci6n Hondurefia de Desarrollo

Forestal (COHDEFOR), who made the necessary collecting

and export permits available to us over the years. In addition,

our good friend Mario R. Espinal has acted as our agent in

Honduras to assure that the gaining of these permits went as

smoothly as possible, that vehicles were ready when we need-

ed them, and that other myriad details upon which the success

of our field work depended were appropriately managed.

Finally, it has been our great and continuing pleasure to work

with uncounted Hondurans who have made we gringos feel

welcome in what is, in a very real sense, our second home.

Without the unstinting efforts of these Honduran friends, we

would never have unraveled the secrets of the animals to

which we have devoted our scientific lives.

We also wish to thank Nidia Romer, who kindly translat-

ed the Abstract into Spanish for use as the Resumen. Finally,

we Owe a sincere debt of gratitude to Louis Porras, Jay M.

Savage, and Jack W. Sites, the reviewers of this paper, whose

suggestions materially improved our presentation.

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Manuscript received: 21 December 2001; Accepted: 4 April 2002

Published online: October 2003; Printed: January 2004

Volume 3 | Number 1 | Page 33

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The herpetofauna of the cloud forests of Honduras

LARRY DAVID WILSON' AND JAMES R. MCCRANIE?

'Department of Biology, Miami-Dade Community College, Kendall Campus, Miami, Florida 33176-3393, USA

210770 SW 164th Street, Miami, Florida 33157-2933, USA

Abstract.—The cloud forest amphibians and reptiles constitute the most important herpetofaunal segment in

Honduras, due to the prevalence of endemic and Nuclear Middle American-restricted species. This segment,

however, is subject to severe environmental threats due to the actions of humans. Of the 334 species of amphib-

ians and reptiles currently known from Honduras, 122 are known to be distributed in cloud forest habitats. Cloud

forest habitats are found throughout the mountainous interior of Honduras. They are subject to a Highland Wet

climate, which features annual precipitation of >1500 mm and a mean annual temperature of <18°C. Cloud for-

est vegetation falls into two Holdridge formations, the Lower Montane Wet Forest and Lower Montane Moist

Forest. The Lower Montane Wet Forest formation generally occurs at elevations in excess of 1500 m, although

it may occur as low as 1300+ m at some localities. The Lower Montane Moist Forest formation generally occurs

at 1700+ m elevation. Of the 122 cloud forest species, 18 are salamanders, 38 are anurans, 27 are lizards, and

39 are snakes. Ninety-eight of these 122 species are distributed in the Lower Montane Wet Forest formation and

45 in the Lower Montane Moist Forest formation. Twenty species are distributed in both formations. The cloud

forest species are distributed among restricted, widespread, and peripheral distributional categories. The

restricted species range as a group in elevation from 1340 to 2700 m, the species that are widespread in at least

one of the two cloud forest formations range as a group from sea level to 2744 m, and the peripheral species

range as a group from sea level to 1980 m. The 122 cloud forest species exemplify ten broad distributional pat-

terns ranging from species whose northern and southern range termini are in the United States (or Canada) and

South America, respectively, to those species that are endemic to Honduras. The largest segment of the her-

petofauna falls into the endemic category, with the next largest segment being restricted in distribution to

Nuclear Middle America, but not endemic to Honduras. Cloud forest species are distributed among eight eco-

physiographic areas, with the largest number being found in the Northwestern Highlands, followed by the

North-Central Highlands and the Southwestern Highlands. The greatest significance of the Honduran herpeto-

fauna lies in its 125 species that are either Honduran endemics or otherwise Nuclear Middle American-restricted

species, of which 83 are distributed in the country’s cloud forests. This segment of the herpetofauna is seriously

endangered as a consequence of exponentially increasing habitat destruction resulting from deforestation,

even given the existence of several biotic reserves established in cloud forest. Other, less clearly evident envi-

ronmental factors also appear to be implicated. As a consequence, slightly over half of these 83 species (50.6%)

have populations that are in decline or that have disappeared from Honduran cloud forests. These species pos-

sess biological, conservational, and economic significance, all of which appear in danger of being lost.

Resumen. —Los anfibios y reptiles de los bosques nublados constituyen el segmento mas importante de la her-

petofauna de Honduras, debido a la prevalencia de especies endémicas y restringidas a la Mesoamerica Nuclear.

Este segmento, sin embargo, esta sometido a fuertes amenazas medioambientales debido a acciones humanas.

De las 334 especies de anfibios y reptiles que se conocen en Honduras en el presente, 122 se conocen que estan

distribuidas en las habitaciones de los bosques nublados. Las habitaciones del bosques nublados se encuentran

a través de las montanas del interior de Honduras. Ellos estan sujetos a un clima Iluvioso de tierras altas, el cual

tiene una precipitacidn anual de mas de 1500 mm y una temperatura anual promedia de menos de 18 grados

centigrados. La vegetacion de los bosques nublados cae entre dos formaciones de Holdridge, la de Bosque

Lluvioso Montano Bajo y la de Bosque Humedo Montano Bajo. La formacion de Bosque Lluvioso Montano Bajo

generalmente occure a elevaciones en exceso de 1500 m, aunque puede ocurrir tan bajo como 1300 m en algu-

nas localidades. La formacion Bosque Humedo Montano Bajo generalmente ocurre a 1700 m o mas de elevacion.

De las 122 especies de los bosques nublados, 18 son salamandras, 38 son anuros, 27 son lagartijas y 39 son cule-

bras. Noventa y ocho de estas 122 especies estan distribuidas en la formacion Bosque Lluvioso Montano Bajo y

45 en la formacion Bosque Humedo Montano Bajo. Viente especies estan distribuidas en ambas formaciones.

Las especies de los bosques nublados estan distribuidas entre categorias distribucionales restringidas, amplias,

y periféricas. Las especies restringidas se encuentra como grupo en un rango de elevaciones de los 1340 a los

Correspondence. !Fax: (305) 237-0891, email: lwilson@mdcc.edu email: jmccrani@betlsouth.net

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 34

The herpetofauna of the cloud forests of Honduras

2700 m, las especies que tienen una distribucion amplia en al menos entre una de las dos formaciones de los

bosques nublados como grupo tiene un rango desde el nivel del mar hasta 2744 m, y las especies periféricas

como grupo tiene un rango desde el nivel del mar hasta 1980 m. Las 122 especies de los bosques nublados ejem-

plifican 10 patrones distribucionales amplios con rangos de especies para las cuales los rangos terminales

norteno y sureno estan en los Estados Unidos (o Canada) y América del Sur, respectivamente, hasta esas

especies que son endémicas de Honduras. El segmento mas grande de la herpetofauna cae en la categoria

endémica, con el proximo segmento mas grande siendo restringido en distribucion a la Mesoamerica Nuclear,

pero no endémico de Honduras. Las especies de los bosques nublados estan distribuidas entre ocho areas

ecofisiograficas, con el grupo mas grande encontrandose en las tierras altas hacia el noroeste y seguido por las

tierras altas norte-central y las tierras altas del suroeste. La importancia mas grande de la herpetofauna hon-

durena cae en sus 125 especies que son endémicas de Honduras o de otra manera restringidas a la

Mesoamerica Nuclear, de las cuales 83 estan distribuidos en los bosques nublados del pais. Este segmento de

la herpetofauna esta seriamente amenazado a consequencia de la destruccion exponencial de sus habitaciones,

el cual es el resultado de la destruccion de los bosques, aunque existen varias reservas bidticas establecidas en

los bosques nublados. Otros factores medioambientales menos claramente evidentes parecen estar implicados.

Como consequencia, un poco mas de la mitad de estas 83 especies (50.6%) tiene poblaciones que estan dis-

minuyendo o que han desaparecidos de los bosques nublados hondurenos. Estas especies poseen significancia

bioldgica, de conservacion, y economica, todas las cuales parecen estar en peligro de ser perdidas.

Key words. Cloud forests, Honduras, amphibians, reptiles, herpetofauna

Introduction

After decades of warnings by environmental scientists, popu-

lation biologists, and demographers (see especially Osborn

1948: Carson 1962; Ehrlich 1968; Meadows et al. 1972), it is

becoming increasingly apparent to an enlarging group of peo-

ple that the Earth is entering a sixth spasm of mass extinction

of life, at least comparable to and, perhaps, exceeding in scope

the five episodes that have preceded it (Ehrlich and Ehrlich

1981, 1996; E. Wilson 1988, 1992; E. Wilson and Perlman

2000). What has come to be known as biodiversity decline is

best documented in areas where the flora and fauna are most

completely understood, e.g., the United States, and corre-

spondingly less well understood in the areas of the world

supporting the greatest amount of biodiversity—the tropics.

To use as an example the country that has been the focus

of our research for more than three decades—Honduras—and

the group upon which we have specialized—the herpetofauna,

it is evident that the modern study of the Honduran herpeto-

fauna began with the research of John R. Meyer that led to his

dissertation, which appeared in 1969. Meyer’s (1969) study

documented a known herpetofauna of 196 species, including

53 amphibians and 143 reptiles. The current tally is 334

species, including 117 amphibians and 217 reptiles (McCranie

and Wilson 2002; Wilson and McCranie 2002). With respect

to the total count, there has been an increase of 138 species or

41.3% in the 33 years since 1969 to the present (although

Meyer did not include five marine turtles species then known

to occur in Honduran waters, nor five species of reptiles

known in Honduran territory only from the Swan Islands,

which are included in the total count of 334). Meyer (1969)

included 35 species in the cloud forest herpetofauna of

Honduras, although one species included by him (Ungaliophis

continentalis) is not so included by us. Presently, we can doc-

ument the presence of 122 species in one or more cloud forest

regions of Honduras. This increase of 88 species (or 72.1% of

the total now known) is largely a result of our field work in the

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country. Forty-two of these 88 species (47.7%) have been

described as new species since 1979. In addition, populations

of two species reported from cloud forest by Meyer (1969)

have been described as new species (Ptychohyla spinipollex

and Ninia lansbergi cloud forest populations of Meyer equal

P. hypomykter and N. espinali, respectively).

There is still significant mountainous terrain in Honduras

supporting cloud forest that has been incompletely sampled

herpetofaunally. Such is the case with the Yoro Highlands, the

Agalta Highlands, and the Santa Barbara Highlands. Given

the frequency with which new taxa have been added to the

Honduran cloud forest herpetofauna (2.3 taxa per year since

1972), it can be expected that additional forms await discov-

ery in these yet poorly known ranges.

Acting in contraposition, however, is a more recent trend

toward decline of herpetofaunal populations, which has been

documented in Honduras by Wilson and McCranie (1998,

2003 a and b) and McCranie and Wilson (2002). This trend

has been most evident in regions of the country in excess of

900 m in elevation and has most obviously affected the

species composing the most distinctive group, i.e., those that

are endemic to Honduras or otherwise restricted in distribu-

tion to Nuclear Middle America. Of the 125 species belonging

to this group, 52 or 41.6% are considered to have declining

populations, to be extinct, or to be extirpated in Honduras.

This trend is extremely alarming, given the fact that the 125

species involved do not occur outside of Nuclear Middle

America.

In light of the importance of the cloud forest environ-

ments of Honduras as centers of herpetodiversity and the

accumulating evidence of the decline and disappearance of a

significant amount of this diversity, it is the purpose of this

paper to update our current understanding of the composition

and distribution (both geographic and ecological) of this her-

petofauna, to discuss its biodiversity significance, to examine

its current conservation status, and to speculate on the future

for this segment of the Honduran herpetofauna.

Volume 3 | Number 1 | Page 35

L. D. Wilson and J. R. McCranie

Materials and methods

Fieldwork upon which this paper is based has been conducted

by one or both of us since 1968. The material collected has

been reported in a number of publications written by one or

both of us since 1971 and summarized in Meyer and Wilson

(1971, 1973), Wilson and Meyer (1985), and McCranie and

Wilson (2002, in preparation).

The Coefficient of Biogeographic Resemblance algo-

rithm (Duellman 1990) was used to demonstrate herpetofaunal

relationships among the cloud forest ecophysiographic areas

examined in this study. The formula is CBR = 2C/(Nj + N2),

where C is the number of species in common to both forma-

tions, Nj is the number of species in the first formation, and N2

is the number of species in the second formation.

Physiography

Honduras contains within its borders a major segment of the

mountains of Nuclear Middle America (West 1964). Many of

the ranges found within the country have portions high

enough to support cloud forest (Fig. 1). Descriptions of the

physiography of Honduras have appeared in Wilson and

Meyer (1985) and McCranie and Wilson (2002), so this

description will be limited to only those mountain ranges upon

which cloud forest vegetation occurs.

Elevations high enough to support cloud forest are dis-

tributed throughout the Serrania, the mountainous interior of

Honduras, which is a portion of the Nuclear Middle American

highlands (Fig. 1). The Serrania is traditionally divided into

the Northern Cordillera and the Southern Cordillera, the latter

distinguishable from the former by an overlay of Pliocene

volcanic ejecta deposits (Wilson and Meyer 1985). Both of

these cordilleras are interrupted by an irregular graben, called

the Honduran depression, traceable from north to south

through the Ulua-Chamelecon Plain, the Valley of Humuya,

the Comayagua Plain, and the Valley of Goascoran (Wilson

and Meyer 1985). In effect, these physiographic features

divide the mountainous interior of Honduras into four sectors,

three of which are recognized as ecophysiographic areas on

the basis of this division. They are the Northwestern

Highlands, the Southwestern Highlands, and the Southeastern

Highlands. The fourth sector is significantly larger than any of

the other three and is broken into four ecophysiographic areas

(see below).

Climate

Savage (2002), in his opus on the amphibians and reptiles of

Costa Rica, noted “the term cloud forest is often applied to

forests that develop at an altitude where the temperature (6 to

10°C) causes water condensation that produces clouds, fog,

and rain. This zone may be at any elevation, and its degree of

development is related to the amount of water vapor in the air.

Cloud forests usually occur where there are prevailing

onshore winds that have their air masses uplifted along ocean-

facing mountains. In Central America, cloud forests develop

principally on the windward slopes affected by the northeast

trade winds. In the Holdridge (1967) system, cloud forests are

regarded as atmospheric association within bioclimates that,

in Central America, usually develop in the lower portion of

the lower montane life zone under the influence of strong pre-

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vailing winds. During much of the year these forests receive

precipitation in the form of light mists. In the drier seasons,

much of the time they are enveloped in dense, dripping fog.”

Areas supporting cloud forest in Honduras are generally

subject to a Highland Wet climatic regime (Wilson and Meyer

1985). This climatic type is broadly characterized by annual

rainfall of >1500 mm and a mean annual temperature of

<18°C. The cloud forest regions occurring in the Southern

Cordillera generally receive less rainfall than do those in the

Northern Cordillera, part of the general effect of the dissipa-

tion of moisture in clouds carried by the prevailing winds

arising over the Caribbean Sea as they sweep inland.

Climatic data are available for the nuclear zone and the

buffer zone of Parque Nacional El Cusuco, a cloud forest

reserve in the Sierra de Omoa in northwestern Honduras

(Fundacion Ecologista “Hector Rodrigo Pastor Fasquelle”

1994). Annual precipitation in the nuclear zone is 2995 mm

and in the buffer zone 2580 mm. The rainiest months, in both

cases, are October, November, and December, accounting for

45.1% of total rainfall in both zones. The least rainiest months

are March, April, and May, when only 12.1% of rainfall

occurs in both zones. Monthly temperatures range from

12.9°C in December to 20.2°C in April, with a mean of

16.7°C, in the nuclear zone and from 17.5°C in December to

23.1°C in April, with a mean of 20.6°C, in the buffer zone.

Vegetation

The vegetation of the Honduran cloud forests is referable to

two forest formations, as slightly modified from the work of

Holdridge (1967), which differ from one another on the basis

of the amount of annual precipitation (Wilson and Meyer

1985). The formation characteristic of the cloud forests of the

Northern Cordillera is the Lower Montane Wet Forest forma-

tion. It is characterized by annual precipitation of >2000 mm.

The formation typical of the cloud forests of the Southern

Cordillera is the Lower Montane Moist Forest formation. It

features an annual precipitation of <2000 mm.

Wilson and McCranie (in preparation a) presented infor-

mation on the vegetation of Parque Nacional El Cusuco

(Lower Montane Wet Forest formation), as follows:

“Fundacion Ecologista “Hector Rodrigo Pastor Fasquelle’

(1994) indicated that this forest formation, called “Zona de

Vida Bosque Muy Humedo Montano Bajo Sub-Tropical,’ is

characterized by the presence of three strata. The uppermost

stratum consists of a closed canopy of trees attaining heights

of 35 to 40 m of the following species: Quercus spp.;

Podocarpus oleifolius; Clusia massoniana; and Liquidambar

styraciflua. The middle stratum is composed of the forgoing

species lying in the shade of the taller conspecifics mixed with

Persea vesticula and Myrica cerifera. The lowermost stratum

is comprised of seedlings of the species in the middle and

uppermost strata intermixed with palms such as Chamaedorea

costaricana and C. oblongata, as well as Geonoma congesta

and a great variety of ferns. Many epiphytic orchids, bromeli-

ads, and mosses are present, as well as lianas and vines.”

Espinal et al. (2001) presented similarly limited data on

floristic composition at two sites (at 1570 and 1650 m) in

Parque Nacional La Muralla (both in Lower Montane Wet

Forest formation), located in the Ocote Highlands of the

northwestern portion of the department of Olancho. They stat-

Volume 3 | Number 1 | Page 36

—

The herpetofauna of the cloud forests of Honduras

Table 1. Geographic and ecological distribution, relative abundance, and conservation status of the cloud forest herpetofauna (122

species) of Honduras. Abbreviations include: Formations—LMWF = Lower Montane Wet Forest formation, LMMF = Lower Montane

Moist Forest formation; Forest Formation Distribution—W = widespread in that formation, R = restricted to that formation, P = periph-

erally distributed in that formation; Primary Microhabitat—A = arboreal, T = terrestrial, F = forest inhabitant, P = pondside inhabitant,

S = streamside inhabitant; Relative Abundance—C = common, | = infrequent, R = rare; Conservation Status—S = stable populations

at least at one cloud forest locality, D = all known cloud forest populations declining, E = extinct or extirpated from all known cloud

forest localities, N = no data on population status. See text for explanation of Broad Distribution Pattern abbreviations.

Elevational Broad

Range Distribution Primary Relative Conservation

Species LMWF LMMF (m) Pattern Microhabitat Abundance’ Status

Salamanders (18 species)

Bolitoglossa carri — R 1840-2070 J A,F,S C D

Bolitoglossa celaque — R 1900-2620 J A, T, F, S C S

Bolitoglossa conanti W WwW 1370-2000. I A, F C S

Bolitoglossa decora R — 1430-1550 J A, F C S

Bolitoglossa diaphora R — 1470-2200 J A, F I S

Bolitoglossa dofleini iB — 650-1370 I 18 I D

Bolitoglossa dunni W — 1200-1600 I A, F I S

Bolitoglossa longissima R — 1840-2240 J A, F C S

Bolitoglossa porrasorum W — 980-1920 J A,F,S C S

Bolitoglossa rufescens complex P — 30-1400 I A, F Cc D

Bolitoglossa synoria — R 2150 I A,S R D

Cryptotriton nasalis W — 1220-2200 J AGE R S

Dendrotriton sanctibarbarus W — 1829-2744 J A, T, F C S

Nototriton barbouri W — 860-1990 J A, F C S

Nototriton lignicola R — 1760-1780 J T,F I S

Nototriton limnospectator R — 1640-1980 J ASE C S

Oedipina cyclocauda P — 0-1780 H T,F I S)

Oedipina gephyra R — 1580-1810 J T, F C D

Anurans (38 species)

Atelophryniscus chrysophorus W — 750-1760 J T, F,S C D

Bufo coccifer — W 0-2070 E ‘W, 1 C S

Bufo leucomyos W — 0-1600 J iliegls C S

Bufo valliceps P — 0-1610 E Dee C S

Hyalinobatrachium fleischmanni —_—P — 0-1550 D A,S C D

Duellmanohyla soralia P — 40-1570 I A,S C D

Ayla bromeliacia W — 1250-1790 I A, F C S

Hyla catracha — R 1800-2160 J A,S C D

Ayla insolita R —_ 1550 J A,S C S

Ayla salvaje R — 1370 I A, F R D

Phrynohyas venulosa P — 0-1610 D A, T, P C S

Plectrohyla chrysopleura WwW — 930-1550 J A, T,S I D

Plectrohyla dasypus R — 1410-1990 J A,S C D

Plectrohyla exquisita R — 1490-1680 J A,S C S

Plectrohyla guatemalensis W WwW 950-2600 I A,S C D

Plectrohyla hartwegi — R 1920-2700 I A, F,S I N

Plectrohyla matudai P W 7710-1850 I T,S C D

Plectrohyla psiloderma — R 2450-2530 I A, T,S Cc D

Ptychohyla hypomykter WwW W 620-2070 I A,S C D

Ptychohyla salvadorensis — W 1440-2050 if A, T,S C S

Ptychohyla spinipollex P — 160-1580 J A,S C S

Smilisca baudinii P —_ 0-1610 B A,P C S

Eleutherodactylus anciano —_ W 1400-1840 J T,S I E

Eleutherodactylus aurilegulus P —_ 50-1550 J Hes C E

Eleutherodactylus charadra P — 30-1370 I r,S C E

Eleutherodactylus cruzi R — 1520 J T,S R E

Eleutherodactylus emleni — W 800-2000 J TS R E

Eleutherodactylus laevissimus — P 100-1640 H T,S I E

Eleutherodactylus loki R = 1370 F 1, F R N

Continued on page 38.

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L. D. Wilson and J. R. McCranie

Table 1. Continued.

Elevational Broad

Range Distribution Primary Relative Conservation

Species LMWF LMMF (m) Pattern Microhabitat Abundance’ Status

Eleutherodactylus milesi WwW — 1050-1720 J T,S C E

Eleutherodactylus rostralis W — 1050-1800 I TF I D

Eleutherodactylus saltuarius R — 1550-1800 J 10, 8 I D

Eleutherodactylus stadelmani W — 1125-1900 J T,S C E

Leptodactylus silvanimbus _— W 1470-2000 J ee. C D

Hypopachus barberi — WwW 1470-2070 I 1,2. C S

Hypopachus variolosus P — 0-1610 B 1, C S

Rana berlandieri? P W 0-2200 Cc T,P C S

Rana maculata WwW WwW 40-1980 I Us C D

Lizards (27 species)

Abronia montecristoi R — 1370 I A,F R D

Abronia salvadorensis — R 2020-2125 J A, T, F R D

Celestus bivitattus — P 1510-1980 I TF C D

Celestus montanus P — 915-1372 J A, F R N

Celestus scansorius R — 1550-1590 J A, F R N

Mesaspis moreletii WwW W 1450-2530. I T,F C S

Sceloporus malachiticus W W 540-2530 H A, F C S

Norops amplisquamosus R — 1530-1720 J A, F C S

Norops crassulus — W 1200-2020 I A, F (C S

Norops cusuco R — 1550-1935 J A, F C S

Norops heteropholidotus —_ R 1860-2200 I A, F C S

Norops johnmeyeri R — 1340-1825 J A, F C S

Norops kreutzi R — 1670-1690 J A, F I D

Norops laeviventris WwW W 1150-1900. E A, F I S

Norops loveridgei Pp — ca. 550-1600 J A, F I S

Norops muralla R — 1440-1740 J A, F C D

Norops ocelloscapularis P — 1150-1370 J A, F I D

Norops petersii R — 1340-1370 F A, F R N

Norops pijolensis WwW — 1180-2050 J A, F C S

Norops purpurgularis R — 1550-2040 J A, F C S

Norops rubribarbaris R — 1700 J T,S R N

Norops sminthus — W ca. 1450-2200 J A, F C Ny

Norops tropidonotus P P 0-1900 F ASE: C S

Norops uniformis P — 30-1370 F A, T, F C D |

Norops yoroensis P — 1180-1600 J A, F I S )

Sphenomorphus cherriei P P 0-1860 E T,F C S

Sphenomorphus incertus P — 1350-1670 I Tr, Je R S |

Snakes (39 species) |

Typhlops stadelmani P — 850-1370 J T, F I D |

Boa constrictor P = 0-1370 D T,F I N

Adelphicos quadrivirgatus P — 0-1740 F T,F C S

Coniophanes bipunctatus P — 0-1370 E ml, P I N

Dryadophis dorsalis WwW W 635-1900 I T,F I S

Drymarchon corais P — 0-1555 A T,F I N

Drymobius chloroticus W W 780-1900 F TFS I D

Drymobius margaritiferus P — 0-1450 A TN deh 12 C S

Geophis damiani R — 1750 J ‘Th Jel R N

Gevnhis fulvoguttatus W W 1680-1900 I T, F R D

Imaniodes cenchoa Pp — 0-1620 D A, F C S

Lampropeltis triangulum P — 0-1370 A T,F I N

Leptodeira septentrionalis W WwW 0-1940 A A,P,S I S

Leptophis ahaetulla P — 0-1680 D A, T, P, S C N

Leptophis modestus — R 1890-2020 I T,F R D

Ninia diademata P — 0-1370 Dak oe T,F I D

Continued on page 40.

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 38

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Volume 3 | Number 1 | Page 39

Amphib. Reptile Conserv. | http://www.herpetofauna.org

——————————

L. D. Wilson and J. R. McCranie

Table 1. Continued.

Elevational Broad

Range Distribution Primary Relative Conservation

Species LMWF LMMF (m) Pattern Microhabitat Abundance’ Status

Ninia espinali W W 1590-2242 I T,F Cc D

Ninia sebae P — 0-1650 E T,F C S

Pliocercus elapoides P — 0-1670 F 1, 1! I S

Rhadinaea godmani W W 1450-2160 H T,F I S

Rhadinaea kinkelini W W 1370-2085 I T,F I D

Rhadinaea lachrymans R — 2050 I i sl R N

Rhadinaea montecristi W W 1370-2620 I T,F I S

Rhadinaea tolpanorum R — 1900 J T, F R N

Sibon dimidiatus P — 950-1600 E A,F I D

Sibon nebulatus P — 0-1690 D A,F,S Cc S

Stenorrhina degenhardtii P — 100-1630 D TF I N)

Storeria dekayi — P 635-1900 Cc T,F R N

Tantilla impensa W — 635-ca. 1600 J T,F R D

Tantilla lempira — P 1450-1730 J 1, F I D

Tantilla schistosa P — 950-1680 E T, F I S

Thamnophis fulvus — WwW 1680-2020 I TT PSS C S

Tropidodipsas fischeri — W 1340-2150 I 1, F I D

Micrurus browni — R 1900 F T,F R N

Micrurus diastema P — 100-1680 F T, F I S

Micrurus nigrocinctus P — 0-1600 G T, F C S

Bothriechis marchi W — ca. 500-1840 J A,S I D

Bothriechis thalassinus W W 1370-1750 I A,S R D

Cerrophidion godmani W W ca. 1300-2620 H 1, J I S

Total 122 species

‘Historical. For example, species that were common at one time during our field experience, but may now be declining or extinct.

?LMMEF specimens represent Rana berlandieri x Rana forreri hybrids (see McCranie and Wilson 2002).

DOI: 10.1514/journal.arc.0000013.t001

ed the following (p. 102): “At the 1570 m site, of 38 species

with chest-high diameters of 5 cm or more, seven species were

considered most important [based on numerical prevalence].

These species, in order of importance, are: Persea sp. (agua-

cate); Calatola mollis (nogal); Quercus sapotaefolia

(encinillo); Calophyllum brasiliense (aceite de maria);

Elaeagia auriculata (oreja de macho); Quercus skinneri (bel-

lota); Chamaedorea neurochlamys (palma pacaya).

At the 1650 m site, a group of 6 species (out of 30) were

judged most important, based on frequency of occurrence.

These species, in order of importance, are: Calophyllum

brasiliense (aceite de maria); Quercus sapotaefolia (encinil-

lo); Persea sp. (aguacate); Quercus skinneri (bellota);

Elaeagia auriculata (o[reja] de macho); Alchornea latifolia

(amargoso). Five of these six species are of greatest impor-

tance as well at the 1570 m site.”

Wilson and McCranie (in preparation b) included data

on the floristic makeup of the vegetation of the Lower

Montane Moist formation in Parques Nacionales de Celaque

and La Tigra, as follows: “The undisturbed forest is composed

of three strata. The upper stratum is composed of trees 25 to

30 m in height, principally of the species Quercus skinneri

(bellota), Liquidambar styraciflua (liquidambar), Pinus pseu-

dostrobus (pinabete), P. oocarpa (ocote), and Persea sp.

(aguacate sucte). These trees carry a moderate amount of epi-

Amphib. Reptile Conserv. | http://www.herpetofauna.org

phytic mosses, orchids, bromeliads, and aroids. The middle

stratum consists of Quercus sp. (curtidor), Q. oleoides (enci-

no), Clethra macrophylla (alamo blanco), Cedrela oaxacensis

(cedro), Inga sp. (guama), and various species of laurals. The

lower stratum consists of shrubs belonging to the families

Compositae, Myrsinaceae, Rubiaceae, Saurauiaceae, and

Verbenaceae and the genera Cleyera, Miconia, Piper,

Psidium, and Vismia.”

Composition of the cloud forest herpetofauna

The herpetofauna of the cloud forests of Honduras is known

to consist of 122 species (Table 1), including 18 salamanders

(14.8% of total), 38 anurans (31.1%), 27 lizards (22.1%), and

39 snakes (32.0%). The salamanders are all members of the

family Plethodontidae. The anurans belong to six families,

including the Bufonidae (4 species), Centrolenidae (1

species), Hylidae (17 species), Leptodactylidae (12 species),

Microhylidae (2 species), and Ranidae (2 species). The lizards

are members of four families, the Anguidae (6 species),

Phrynosomatidae (1 species), Polychrotidae (18 species), and

Scincidae (2 species). The snakes belong to five families,

including the Typhlopidae (1 species), Boidae (1 species),

Colubridae (31 species), Elapidae (3 species), and Viperidae

(3 species).

Volume 3 | Number 1 | Page 40

2 EOE

The herpetofauna of the cloud forests of Honduras

Distribution and distributional relationships of the

cloud forest herpetofauna

Distribution within forest formations

More than twice as many of the 122 cloud forest species are

distributed in the Lower Montane Wet Forest formation (98 or

80.3% of total) than in the Lower Montane Moist Forest for-

mation (45 or 36.9%). Twenty-one species (17.2%) are found

in both formations (Table 1). The Coefficient of

Biogeographic Resemblance (CBR) for these two forest for-

mations is 0.29.

The species distributed in cloud forests fall into three dis-

tributional categories, viz., restricted, widespread, and

peripheral (Table 1). Restricted species are those whose dis-

tribution is limited to a particular cloud forest formation.

Widespread species are those that are widespread in distribu-

tion in a particular cloud forest formation or both cloud forest

formations, as well as, perhaps, outside those forest forma-

tions. Finally, peripheral species are those whose distribution

is largely peripheral to a particular cloud forest formation.

The Lower Montane Wet Forest formation is inhabited

by 26 restricted species (26.5% of the total of 98 in this for-

mation), including six salamanders, seven anurans, ten lizards,

and three snakes. Thirty-two species (32.7%) are widespread

in this formation, including six salamanders, ten anurans, four

lizards, and 12 snakes. Finally, 40 species (40.8%) are periph-

erally distributed in this formation, including three

salamanders, 11 anurans, eight lizards, and 18 snakes.

The Lower Montane Moist Forest formation is home to

ten restricted species (22.2% of the total of 45 in this forma-

tion), including three salamanders, three anurans, two lizards,

and two snakes. Twenty-nine species (64.4%) are widespread

in this formation, including one salamander, 11 anurans, five

lizards, and 12 snakes. Finally, there are six species (13.3%)

peripherally distributed in this formation, including one anu-

ran, three lizards, and two snakes. Notably, there are

proportionately more peripheral and widespread species than

restricted species in the Lower Montane Wet Forest forma-

tion. In the Lower Montane Moist Forest formation, most

species are widespread ones, followed by relatively few

restricted and peripheral species. The relative prevalence of

peripheral species in the Lower Montane Wet Forest forma-

tion apparently is due to the grading of this type of cloud

forest into highland rain forest (Premontane Wet Forest for-

mation) at elevations usually around 1500 m, whereas the

Lower Montane Moist Forest formation grades into upland

pine forest (Premontane Moist Forest) typically.

As noted above, 21 species are distributed in both cloud

forest formations (Table 1). The largest number of these

species (17) are widespread in both formations. Two species

are peripheral in distribution in both formations, and, finally,

two species are widespread in one formation and peripheral in

the other.

Distribution with respect to elevation

The Lower Montane Wet Forest formation is generally found

at elevations in excess of 1500 m, although in some locales it

occurs at elevations down to 1300+ m. The Lower Montane

Moist Forest formation usually occurs at 1700+ m elevation.

Thus, it is expected that patterns of elevational occurrence

would be related to the patterns of occurrence in the two forest

formations elucidated above. That is to say, the widespread and

peripheral species would be expected to have broader overall

elevational ranges than those whose distribution is restricted to

cloud forest vegetation, with the peripheral species more

broadly distributed overall than the widespread ones.

The restricted species, as a group, range from 1340 to

2700 m. The mean elevational range for this group of 36

species is 209.6 m. The species that are widespread in at

least one of the two cloud forest formations, as a group,

range from sea level to 2744 m. The mean elevational range

for this group of 44 species is 1000.4 m. The species that

occur peripherally in at least one of the two cloud forest for-

mations, as a group, range from sea level to 2200 m. The

mean elevational range for this group of 44 species is 1260.3

m (two species are peripheral in one formation and wide-

spread in the other).

Broad distribution patterns

As did Wilson and Meyer (1985), Wilson et al. (2001), and

McCranie and Wilson (2002), we placed the cloud forest species

into a set of distributional categories based on the entire extent of

their geographic range. Two of the categories used by Wilson et

al. (2001) do not apply to this paper (marine species and insular

and/or coastal species). The applicable categories are as follows:

A. Northern terminus of the range in the United States

(or Canada) and southern terminus in South America.

B. Northern terminus of the range in the United States

and southern terminus in Central America south of the

Nicaraguan Depression.

C. Northern terminus of the range in the United States

and southern terminus in Nuclear Middle America.

Table 2. Summary of numbers of taxa exhibiting various Broad Patterns of Geographic Distibution (See text for explanation of categories).

Groups Broad Patterns of Distribution

A B Cc D E F G H I J

Salamanders (18 species) — 1 5 12

Anurans (38) — yD) ] 2) 2 1 — 1 13 16

Lizards (27) — 2 3 — 1 6 15

Snakes (39) 4 — 1 5) 4 6 1 2 11 5

Totals 122 4 2 2 7 8 10 1 5 35 48

DOI: 10.1514/journal.arc.0000013.t002

Amphib. Reptile Conserv. | http://www.herpetofauna.org

Volume 3 | Number 1 | Page 41

L. D. Wilson and J. R. McCranie

Table 3. Distribution of the Honduran cloud forest herpetofauna within eight ecophysiographic areas. Abbreviations are: W = wide-

spread in that area; R = restricted to that area; P = peripherally distributed in that area; HL = Highlands.

Santa

SE SW N-Central Yoro Ocote Adgalta NW Barbara

Species HL HL HL HL HL HL HL HL Total

Bolitoglossa carri R

Bolitoglossa celaque R

Bolitoglossa conanti P

Bolitoglossa decora R

Bolitoglossa diaphora

Bolitoglossa dofleini P

Bolitoglossa dunni

Bolitoglossa longissima R

Bolitoglossa porrasorum W

Bolitoglossa rufescens complex P

Bolitoglossa synoria R

Cryptotriton nasalis W

Dendrotriton sanctibarbarus R

Nototriton barbouri W

Nototriton lignicola R

Nototriton limnospectator R

Oedipina cyclocauda W

Oedipina gephyra

Atelophryniscus chrysophorus

Bufo coccifer W W

Bufo leucomyos P P W

Bufo valliceps P P

Hyalinobatrachium fleischmanni'! P P

Duellmanohyla soralia

Hyla bromeliacia W

Hyla catracha R

Ayla insolita R

Hyla salvaje R

Phrynohyas venulosa P

Plectrohyla chrysopleura P

Plectrohyla dasypus

Plectrohyla exquisita

Plectrohyla guatemalensis W

Plectrohyla hartwegi

Plectrohyla matudai

Plectrohyla psiloderma

Ptychohyla hypomykter

Ptychohyla salvadorensis

Ptychohyla spinipollex P

Smilisca baudinii P P P P P

Eleutherodactylus anciano P

Eleutherodactylus aurilegulus P W

Eleutherodactylus charadra P

Eleutherodactylus cruzi R

Eleutherodactylus emleni W

Eleutherodactylus laevissimus

Eleutherodactylus loki

Eleutherodactylus milesi

Eleutherodactylus rostralis

Eleutherodactylus saltuarius R

Eleutherodactylus stadelmani W W

Leptodactylus silvanimbus

Hypopachus barberi

Hypopachus variolosus P

ae]

ZEA ERS

~ UTADA

ape

ae)

2 =R

meee NRF NR RR RP RP HE NK HK KY BD BN BR BB Be Be ee BP PN WN FB FB FB i i i ii i i i i i

Continued on page 43.

Amphib. Reptile Conserv. | http://www.herpetofauna.org Volume 3 | Number 1 | Page 42

See eee ee ~'”— m0. OOoOoees

Table 3. Continued.

The herpetofauna of the cloud forests of Honduras

Species

Rana berlandieri*

Rana maculata

Abronia montecristoi

Abronia salvadorensis

Celestus bivitattus

Celestus montanus

Celestus scansorius

Mesaspis moreletii

Sceloporus malachiticus

Norops amplisquamosus

Norops crassulus

Norops cusuco

Norops heteropholidotus

Norops johnmeyeri

Norops kreutzi

Norops laeviventris

Norops loveridgei

Norops muralla

Norops ocelloscapularis

Norops petersii

Norops pijolensis

Norops purpurgularis

Norops rubribarbaris

Norops sminthus

Norops tropidonotus

Norops uniformis

Norops yoroensis

Sphenomorphus cherriei

Sphenomorphus incertus

Typhlops stadelmani

Boa constrictor

Adelphicos quadrivirgatus

Coniophanes bipunctatus

Dryadophis dorsalis

Drymarchon corais

Drymobius chloroticus

Drymobius margaritiferus

Geophis damiani

Geophis fulvoguttatus

Imantodes cenchoa

Lampropeltis triangulum

Leptodeira septentrionalis

Leptophis ahaetulla

Leptophis modestus

Ninia diademata

Ninia espinali

Ninia sebae

Pliocercus elapoides

Rhadinaea godmani

Rhadinaea kinkelini

Rhadinaea lachrymans

Rhadinaea montecristi

Rhadinaea tolpanorum

Sibon dimidiatus

Sibon nebulatus

Santa

SE SW N-Central Yoro Ocote Agalta NW Barbara

HL HL HL HL HL HL HL HL Total

W W P P 4

WwW W P P WwW 5

WwW 1

P 1

R i

WwW WwW WwW W 4

WwW Ww WwW P P WwW WwW 7

R I

WwW 1

R 1

WwW 1

R 1

P P P WwW 4

W 1

x x 2

P 1

R ]

WwW 1

R 1

WwW 1

P P WwW 3

P 1

P P 2

P P P 3

P 1

WwW P P P 5

P 1

P P WwW 4

P it

R 1

WwW W 2

P P P 3

P 1

WwW W P 3

WwW WwW 2

R 1

P 1

WwW WwW 2D

P 1

iP WwW 1p 3

WwW WwW W 3

Ww W P 3

R 1

W 2

R 1

P ]

W 1

Amphib. Reptile Conserv. | http://www.herpetofauna.org

Continued on page 44.

Volume 3 | Number 1 | Page 43

L. D. Wilson and J. R. McCranie

Table 3. Continued.

Santa

SE SW N-Central Yoro Ocote Agalta NW Barbara

Species HL HL HL HL HL HL HL HL Total

Stenorrhina degenhardtii P P D;

Storeria dekayi P 1

Tantilla impensa W |

Tantilla lempira P 1

Tantilla schistosa W 1

Thamnophis fulvus P W 2

Tropidodipsas fischeri W 1

Micrurus browni R 1

Micrurus diastema W 1

Micrurus nigrocinctus Pp 1

Bothriechis marchi Pp W 2

Bothriechis thalassinus Pp W P 3

Cerrophidion godmani W W W W 4

Totals 19 39 39 5 21 4 60 11 -

'North-Central Highlands records based on calling males that could not be located.

Rana berlandien from the Southeastem and Southwestem Highlands equal R. berlandieri x R. forreri (see McCranie and Wilson 2002).

DOI: 10.1514/journal.arc.0000013.t003

D. Northern terminus of the range in Mexico north of the

Isthmus of Tehuantepec and southern terminus in

South America.

E. Northern terminus of the range in Mexico north of the

Isthmus of Tehuantepec and southern terminus in

Central America south of the Nicaraguan Depression.

F. Northern terminus of the range in Mexico north of the

Isthmus of Tehuantepec and southern terminus in

Nuclear Middle America.

G. Northern terminus of the range in Nuclear Middle

America and southern terminus in South America.

H. Northern terminus of the range in Nuclear Middle

America and southern terminus in Central America

south of the Nicaraguan Depression.

I. Restricted to Nuclear Middle America (exclusive of

Honduran endemics).

J. Endemic to Honduras.

The data on broad distributional patterns in Table | are

summarized in Table 2. These data indicate that the largest

number of species (48 or 39.3% of the total of 122 species)

fall into the J category, i.e., that containing the species endem-

ic to Honduras. The next largest category is I, with 35 species

(28.7%), containing those species not endemic to Honduras

but restricted in distribution to Nuclear Middle America.

Together, these two categories contain 68.0% of the cloud for-

est species. The other eight categories contain from one to ten

species and harbor, as a group, 32.0% of the total number.

These data again point to the biogeographic and conservation

importance of the Honduran cloud forest herpetofauna.

Primary microhabitat distribution

We used the same microhabitat categorization as did Espinal

et al. (2001). In terms of vertical positioning, we scored

species as either terrestrial or arboreal. With respect to occur-

rence in the three major microhabitats found in cloud forest,

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species were scored as being found in the forest proper, along

streams, or around ponds (Table 1).

In terms of vertical positioning within the primary micro-

habitats, 49 species (40.2%) were usually found only in

arboreal situations, 62 species (50.8%) only in terrestrial situ-

ations, and 11 (9.0%) in both. With respect to occurrence in

the three major microhabitats (forest proper, streamside,

pondside), 76 species (62.3%) were found exclusively in the

forest proper, 26 (21.3%) only along streams, eight (6.6%)

only around ponds, seven (5.7%) in the forest and along

streams, three (2.5%) around ponds and along streams, and

two (1.6%) in the forest and around ponds (Table 1).

If the two sets of categories, vertical positioning in pri-

mary habitat and microhabitats, are combined, it can be

demonstrated that 94 species (77.0%) fall into four groups, as

follows (Table 1): 40 terrestrial forest inhabitants (32.8%); 31

arboreal forest inhabitants (25.4%); 12 arboreal streamside

inhabitants (9.8%); and 11 terrestrial streamside inhabitants

(9.0%). The terrestrial forest inhabitants include four sala-

manders, four anurans, four lizards, and 28 snakes. The

arboreal forest inhabitants are eight salamanders, two anurans,

19 lizards, and two snakes. The arboreal streamside inhabi-

tants are one salamander, nine anurans, and two snakes. The

terrestrial streamside inhabitants are ten anurans and one

lizard.

Relative abundance

In discussing relative abundance, we used the following catego-

rization: common (C: found on a regular basis, many individuals

can be found); infrequent (I: unpredictable, few individuals

seen); rare (R: rarely seen). These classifications are historical

(i.e., based largely on earlier trips to cloud forest localities) and

do not take into consideration the population declines taking

place for many species (see Biodiversity significance and con-

servation status of the cloud forest herpetofauna). Sixty-three

species (51.6%) are classified as being common (11 salaman-

Volume 3 | Number 1 | Page 44

)

The herpetofauna of the cloud forests of Honduras

NCH

13 2

NWH fs -

SWH

SEH

1g)

Figure 2. Greatest shared species diagram of eight cloud forest areas in Honduras. See text for explanation of abbreviations.

Numbers in boxes are the number of species in each area; numbers on arrows indicate the number of species shared between

areas connected and represent the greatest shared value for each area. Position of the boxes in the diagram is roughly reflective

of their geographic relationships in Honduras.

DOI: 10.1514/journal.arc.0000013.g002

ders, 28 anurans, 15 lizards, and nine snakes), 37 (30.3%) as

being infrequent (five salamanders, six anurans, five lizards, and

21 snakes), and 22 (18.0%) as being rare (two salamanders, four

anurans, seven lizards, and nine snakes).

Patterns of distribution among ecophysiographic

areas

Wilson et al. (2001) recognized eight ecophysiographic areas

that contain cloud forest vegetation. Two of these areas, the

Southeastern Highlands and Southwestern Highlands, are

located in the Southern Cordillera. The remaining six areas are

situated in the Northern Cordillera. The distribution of the

members of the Honduran cloud forest herpetofauna among

these eight cloud forest ecophysiographic areas is indicated in

Table 3. Perusal of the data in this table allows for several

conclusions, as follows:

1. The numbers of species in these eight areas range from

four (Agalta Highlands) to 60 (Northwestern Highlands).

2. Significantly more species are known from the

Northern Cordillera areas (98 or 80.3% of total) than

those in the Southern Cordillera (45 or 36.9%) areas.

Only 20 species (16.4%) are distributed in both

cordilleras (the Rana berlandieri listed in Table 3 from

the Southern Cordillera are considered R. berlandieri

x R. forreri hybrids—see McCranie and Wilson 2002).

. The above pattern is seen in each of the major her-

petofaunal groupings. Only four salamanders are

found in the Southern Cordillera cloud forests, com-

pared to 15 in the Northern Cordillera cloud forests.

Only a single species (Bolitoglossa conanti) is dis-

tributed in both cordilleras (although the population

in the Southern Cordillera likely represents an unde-

scribed species). Fifteen species of anurans occur in

the Southern Cordillera cloud forests, as opposed to

28 in the Northern Cordillera forests. Only four

species (Plectrohyla guatemalensis, P. matudai,

Ptychohyla hypomykter, and Rana maculata; the

Table 4. CBR matrix of herpetofaunal relationships for the eight ecophysiographic areas supporting cloud forest. N = species in

each region; N = species in common between two regions; N = Coefficients of Biogeographic Resemblance. See text for explana-

tion of the abbreviations. No distinction is made between Rana berlandieri and R. berlandieri x R. forreri for this analysis.

SEH SWH NCH YH OH AH NWH SBH

SEH 19 13 7 1 7 10 2

SWH 0.45 39 9 7) 6 1 18 2

NCH 0.24 0.23 39 5 12 1 13 1

YH 0.08 0.09. 0.23 5 it 3 |

OH 0.35 0.20 0.40 0.31 21 2 12 |

AH 0.09 0.05 0.05 0.22 0.16 4 I 0

NWH 0.25 0.36 0.26 0.09 0.30 0.03 60 5

SBH 0.13 0.08 0.04 0.13 0.06 0.00 0.14 11

DOI: 10.1514/journal.arc.0000013.t004

Amphib. Reptile Conserv. | http://www.herpetofauna.org

Volume 3 | Number 1 | Page 45

L. D. Wilson and J. R. McCranie

NCH

ay)

Figure 3. Coefficient of Biogeographic Resemblance diagram for the eight cloud forest areas in Honduras. See text for explana-

tions of abbreviations. Numbers in boxes are the number of species in each area; decimal numbers on arrows indicate the CBR

value shared between areas connected and represent the highest value for each area; absolute numbers in parentheses indicate

the number of species shared between the areas connected. Position of the boxes in the diagram is roughly reflective of their geo-

graphic relationships in Honduras.

DOI: 10.1514/journal.arc.0000013.g003

Rana berlandieri listed in Table 3 from the Southern

Cordillera are considered R. berlandieri x R. forreri

hybrids—see McCranie and Wilson 2002) occur in

both regions. Ten species of lizards are distributed in

the Southern Cordillera forests, whereas 22 are in the

Northern Cordillera forests. Only five species

(Mesaspis moreletii, Sceloporus malachiticus,

Norops laeviventris, N. tropidonotus, and Spheno-

morphus cherriei) are found in both areas. Finally, 16

species of snakes occupy the Southern Cordillera

cloud forests and 33 the Northern Cordillera forests.

Ten species (Dryadophis dorsalis, Drymobius

chloroticus, Geophis fulvoguttatus, Leptodeira

septentrionalis, Ninia espinali, Rhadinaea godmani,

R. kinkelini, R. montecristi, Bothriechis thalassinus,

and Cerrophidion godmani) are distributed in both

areas.

4. Most of the 122 species (102 or 83.6%) occur in only

one or two cloud forest ecophysiographic areas. The

most broadly-distributed species occur in seven eco-

physiographic areas (Sceloporus malachiticus) or in

five ecophysiographic areas (Plectrohyla guatemalen-

sis, Smilisca baudinii, Rana maculata, and Dryadophis

dorsalis). The average area occurrence is 1.6.

A greatest shared species diagram of the eight cloud for-

est physiographic areas is presented in Figure 2. The areas are

abbreviated as follows: Southeastern Highlands - SEH;

Southwestern Highlands - SWH; North-Central Highlands -

NCH; Yoro Highlands - YH; Ocote Highlands - OH; Agalta

Highlands - AH; Northwestern Highlands - NWH; Santa

Barbara Highlands - SBH. The number of species shared

between areas ranges from two to 18. In general, the greater

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the total herpetofaunas of any two compared areas, the greater

is the number of species shared.

Generation of Coefficient of Biogeographic

Resemblance (CBR) values allows for a more robust analy-

sis of herpetofaunal resemblances. Thus, a matrix of CBR

values for the eight ecophysiographic areas is summarized in

Table 4, and these values are used to produce a CBR diagram

(Fig. 3) indicating highest values for each ecophysiographic

area. These values indicate that the herpetofauna of a given

ecophysiographic area most closely resembles that of anoth-

er area occupied by the same forest formation and/or lying in

close geographic proximity. For example, the Southeastern

Highlands and the Southwestern Highlands are both occu-

pied by the Lower Montane Moist Forest formation and they

share 13 species. Also, as an example, the Northwestern

Highlands and the Southwestern Highlands are in close geo-

graphic proximity and share 18 species. Geographic

proximity, however, appears to be the more important deter-

minant of the degree of herpetofaunal resemblance,

inasmuch as Figure 3 illustrates a western and southern

grouping of areas (NWH, SBH, SWH, and SEH) and a

northern and eastern grouping of areas (NCH, OH, YH, and

AH). These two groups are connected by a relatively high

CBR value between SEH and OH.

Averaging all CBR values provides a gauge of herpeto-

faunal distinctiveness, as follows: SEH (0.23); SWH (0.21);

NCH (0.21); YH (0.16); OH (0.25); AH (0.09); NWH (0.20);

SBH (0.08). The most distinctive herpetofauna is that of the

SBH (average CBR value of 0.08), the least that of the OH

(average CBR value of 0.25). The distinctiveness of the SBH

herpetofauna is an artifact of being poorly known. The fewer

the species known from a given area, the fewer there are to be

shared with other areas.

Volume 3 | Number 1 | Page 46

The herpetofauna of the cloud forests of Honduras

Biodiversity significance and conservation status of

the cloud forest herpetofauna

As noted in the Introduction, the herpetofauna of Honduras is

being subjected to the same anthropogenic pressures as have

been demonstrated to be in effect elsewhere in the tropics. The

most substantial pressure is created by habitat loss as a result

of deforestation (Wilson et al. 2001; Wilson and McCranie

2003 a and b). Also significant is a threat of unsubstantiated

origin (but see Duellman 2001, for a discussion of events

elsewhere in the tropics) that is decimating amphibian popu-

lations in the country occurring at elevations in excess of 900

m (Wilson and McCranie 1998; McCranie and Wilson 2002),

thus conceivably impacting all cloud forest areas.

That these threats are impinging on herpetofaunal popu-

lations at 900 m and above is especially poignant, inasmuch as

the herpetodiversity of greatest significance is distributed in

these regions, especially those supporting cloud forest. This

most significant herpetodiversity consists of those species

endemic to Honduras and those otherwise restricted to

Nuclear Middle America. Of the 334 species now known to

constitute the Honduran herpetofauna (including six marine

reptiles), 78 are country endemics (23.4% of total) and 47 are

Nuclear Middle American-restricted species (14.1%). A

greater percentage of the amphibian species fall into these two

categories than do the reptilian species. There are 41 amphib-

ian Honduran endemics (35.0% of total of 117 species) and 25

Nuclear Middle American-restricted amphibian species

(21.4%), compared to 37 (17.1% of total of 217 species) and

22 (10.1%) such reptilian species, respectively. Thus, a total

of 125 species of amphibians and reptiles (37.4%) are either

endemic to Honduras or otherwise restricted to Nuclear

Middle America.

Of these 125 species, 83 or 66.4% are distributed in

cloud forests in Honduras (Table 2). Of the remaining 209

Honduran species not found in cloud forests, only 42 species

or 20.1% are Honduran endemics or Nuclear Middle

American-restricted. It is obvious that the large majority of the

species of greatest biodiversity significance is found in cloud

forests.

As indicated above, deforestation is eroding forest

resources throughout the country. Wilson and McCranie

(2003 a) presented estimates, based on a computer model in E.

Wilson and Perlman (2000), suggesting that the current defor-

estation rate is -2.3%, giving rise to a halving rate of 30.1

years. At this rate, only a half a million hectares of forest will

remain in Honduras by the year 2085 and none will remain by

the end of the current century.

This trend has been affecting cloud forests in Honduras,

just as it has everywhere else in the country, and continues to

the present day. It has been abated somewhat by the establish-

ment of biotic reserves in several of the ranges supporting cloud

forest (Wilson et al. 2001). This establishment largely has been

the result of an effort to secure water supplies for populated

areas. As noted by Wilson et al. (2001), however, most of these

reserves are incompletely developed, such that deforestation

still proceeds in many, if not all of them (e.g., Espinal et al.

2001), as a result of illegal logging and subsistence farming.

It has been demonstrated in recent years that populations

of many Honduran amphibians and reptiles are in decline or

Amphib. Reptile Conserv. | http://www.herpetofauna.org

have disappeared altogether, as part of a global pattern

(Duellman 2001). Wilson and McCranie (2003 a) have pro-

vided the most recent assessment of this trend for the

Honduran herpetofauna. However, their assessment differs

somewhat from the one undertaken here. Wilson and

McCranie (2003 a) considered the range as a whole for each

species when classifying whether a given species had stable

populations somewhere in their range. However, a few species

may have stable populations at some low elevation localities,

but may be extirpated from their known cloud forest localities

(e.g., Eleutherodactylus charadra). Thus, the conservation

status categories in this paper refer only to cloud forest popu-

lations. Table 1 lists the conservation status for each of the

122 species at their known cloud forest localities. These data

indicate that 40 species (32.8%) have populations that are in

decline, eight species (6.6%) have disappeared altogether

from cloud forests, and 16 species (13.1%) are too poorly

known to determine their status in Honduran cloud forests.

Fifty-eight species (47.5%) appear to have stable populations

in at least one cloud forest locality.

When one considers only the two most important com-

ponents of the Honduran cloud forests (the Honduran

endemics and the Nuclear Middle American-restricted

species), then 15 of the 48 Honduran endemics (31.3%) have

declining populations, six endemics (12.5%) have disap-

peared, five endemics (10.4%) are too poorly known to

determine their status, and 22 endemics (45.8%) appear to

have stable populations in at least one cloud forest locality. Of

the 35 Nuclear Middle American-restricted species, 20

(57.1%) have declining populations, one (2.9%) has disap-

peared, two (5.7%) are too poorly known, and 12 (34.3%)

appear to have stable populations in at least one cloud forest

locality. Thus, about one half of the 83 Honduran endemics or

Nuclear Middle American-restricted species have declining

populations (35 species or 42.2%) or have disappeared from

Honduran cloud forests (seven species or 8.4%). Of the six

Honduran endemic species that have disappeared from cloud

forests, five are feared extinct. These are shocking statistics,

considering the importance of these species not only biologi-

cally, but also from conservation and ecotourist standpoints.

From simply a biological standpoint, the systematics of

the majority of the 83 cloud forest notables (Honduran

endemics and Nuclear Middle American-restricted species)

are insufficiently understood to be subjected to cladistic

analysis, a requirement for reconstructing their phylogenies,

and, beyond this, their biogeographic histories. These species

are particularly important in our effort to understand the gen-

eral patterns of evolution of the herpetofauna and to take that

understanding beyond the work done on this subject to date.

From the perspective of conservation biology, we have

demonstrated here and elsewhere (Wilson and McCranie

1998, 2003 a and b; Wilson et al. 2001; McCranie and Wilson

2002, in press) that the herpetofauna is anything but the

pedestrian compendium alluded to in Lynch and Fugler’s

(1965, p.15) conclusions when they wrote that, “The anuran

fauna seems to be derived from largely widespread species

and species with northern affinities.” Quite to the contrary, the

work that has been accomplished since Lynch and Fugler pub-

lished their paper 38 years ago has shown that slightly more

than a third (37.4%) of the Honduran herpetofauna is com-

Volume 3 | Number 1 | Page 47

L. D. Wilson and J. R. McCranie

posed of endemics or otherwise Nuclear Middle American-

restricted species. Our work in cloud forests has provided the

major support for that conclusion.

The economic value of the Honduran cloud forests for

ecotourism is only beginning to be calculated. It is stunningly

evident to us, however, based on the several decades of our

field work in the country, that efforts to develop an ecotourist-

generated component to the Honduran economy is likely to be

doomed by the uncontrolled human population growth that

continues to stymie efforts to conserve the considerable biodi-

versity of the country.

Acknowledgments.—Over the decades that we have

worked on the cloud forest herpetofauna in Honduras, we

have become indebted to innumerable individuals and organ-

izations. First and foremost, this work could not have been

accomplished without the support of the personnel of the

governmental agencies Recursos Naturales Renovables and

Corporacién Hondurefia de Desarrollo Forestal (COHDE-

FOR), who made the necessary collecting and export permits

available to us over the years. In addition, our inestimable

friend Mario R. Espinal has acted as our agent in Honduras in

assuring that the garnering of these permits went as smooth-

ly as possible, that vehicles were ready when we needed

them, and that other myriad details upon which the success of

our field work depended were appropriately handled. Finally,

it has been our great and continuing pleasure to work with

uncounted Hondurans who have made us gringos feel at

home in Honduras. Without the unflagging efforts of these

friends, we would never have unearthed the secrets of the

amphibians and reptiles to which we have devoted our scien-

tific lives.

We also extend our gratitude to Nidia Romer, who gra-

ciously translated the Abstract into Spanish for use as the

Resumen. We are also grateful to Louis Porras and Jay M.

Savage, the reviewers of this paper, for their improvements.

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Published online: October 2003; Printed: January 2004

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