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Amphibian and Reptile Conservation

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Amphibian and Reptile Conservation 2(1):4.

Editorial

WORDS FROM THE EDITOR—With the comple-

tion and publication of this issue, Amphibian and

Reptile Conservation (ARC) is now publishing semi-

annually, from our previously irregular publishing

schedule. We still have not given up our original

plans of being published as a quarterly serial, and

hope to be able to do so in the near future. We will

continue to pick up the pace, and add more pages to

each volume (when possible), as well as report on

some of the most exciting topics in the world, from

a balanced geographic perspective. ARC will con-

tinue its focus on publishing research in the field of

herpetology, and specifically herpetological conservation. We will

try our best to publish cutting edge herpetological conservation

research, but in a way that makes the subject matter accessible and

enjoyable to professional herpetologists, conservation managers,

naturalists, and nonprofessionals. This often requires an extra ef-

fort on part of reviewers, authors, and others involved. It is to

these unselfish individuals that deserve most of the credit for the

success of the journal.

This issue begins with an additional four pages (two in full-

color) being added from that of the previous issue

and volume (volume 1, number 1—premiere is-

sue). It also adds new standardized graphic design

throughout, important article contributions, full-

color illustrated maps, country sidebars, a new col-

umn titled “Herpetofauna and Humanity,” book

review(s), world news, glossary, and abbreviations

used. All these new improvements are imple-

mented to make this science journal as accessible

and interesting to as broad an audience as possible.

Many of these journal elements will continue to

be developed, as well as others that come to my

attention, with each successive volume. I am very pleased to

exhibit our latest effort and hope that you all will continue your

great enthusiasm for the journal, as we work hard to establish

what we think will become an important serial, specifically for

the conservation and preservation of amphibians, reptiles, and

their habitats worldwide.

Craig Hassapakis

Editor and Publisher

Authors

AARON M. BAUER received his Doctor of Philosophy

(Ph.D.) degree in Zoology from the University of California,

Berkeley. He is a Professor of Biology at Villanova University

and a research associate of the Smithsonian Institution, Cali-

fornia Academy of Sciences, and

Museum of Natural Science, Loui-

siana State University. He has pub-

lished several books and over 200

scientific articles. His research spe-

cialties are the systematics, bioge-

ography, and evolutionary mor-

phology of Southern Hemisphere

reptiles, especially geckos. Dr.

Bauer has had extensive field ex-

perience in New Caledonia and with Ross Sadlier has recently

written The Herpetofauna of New Caledonia, a new book on

the diverse lizard fauna of this French island territory.

CRAIG M. HOOVER, our columnist—Herpetofauna and Hu-

manity—is a Senior Program Officer with TRAFFIC North

America. Mr. Hoover has extensive experience in wildlife trade

issues and wildlife law enforcement. Prior to joining TRAF-

FIC in 1996, he worked for four years as a Wildlife Inspector

with the United States Fish and Wildlife Service, enforcing

state, federal, and foreign laws relating to the international

wildlife trade. Mr. Hoover has a Bachelor of Science (B.S.)

degree in Natural Resources from the University of Michigan

as well as a Juris Doctorate degree from Loyola Law School. In

his position with TRAFFIC, he has worked on a number of

reptile trade issues, including the role of the United States in

the live reptile trade, the trade in native turtles, and the trade in

turtles for food in East and Southeast Asia.

RONALD A. NUSSBAUM (RAN) is curator of herpetology,

professor of biology, and director of the Edwin S. George re-

serve at the University of Michigan in Ann Arbor. His research

interests include, broadly, ecology, evolution, and systematics

of amphibians and reptiles. His specific research projects in-

clude evolution of parental care in salamanders, evolution and

systematics of caecilians, and the systematics and biogeography

of the herpetofauna of the islands of the western Indian Ocean,

in particular the Seychelles archipelago and Madagascar.

BRIAN A. MAURER received a B.S. in Zoology from Brigham

Young University and a Ph.D. in Wildlife Ecology at Univer-

sity of Arizona. After 13 years in the Department of Zoology

at Brigham Young University, he recently accepted a position

as associate professor in the Department of Fisheries and Wild-

life and Department of Geography at Michigan State Univer-

sity. He has published two books and numerous articles on

population and community ecology of vertebrates, particu-

larly birds. His research interests include biogeography, quan-

titative ecology, and macroecology.

CHRISTOPHER J: RAXWORTHY is Associate Curator in

the Department of Herpetology at the American Museum of

Natural History in New York,

New York. His research interests

focus on the evolutionary biology

and ecology of reptiles, and in-

clude systematics, biogeography,

conservation, and behavior. Al-

though he has done much of his

most recent work in Madagascar,

his interest in reptiles is world-

wide, with ongoing fieldwork in ; ss

North America, Madagasacar, West Africa, and Southeast

Asia.

PAUL THOMPSON

ROSS A. SADLIER received his B.S. degree in Zoology from

Macquarie University in Sydney, Australia. He is a Senior

Scientific Officer in the Depart-

ment of Herpetology at the Aus-

tralian Museum. He has published

extensively on the herpetofaunas

of Australia and New Caledonia.

His research interests are centered

on the systematics of scincid liz-

ards of the Pacific region. Mr.

Sadlier has had extensive field ex-

perience in New Caledonia and is

co-author with Aaron Bauer of

The Herpetofauna of New Caledonia, anew book on the lizard

fauna of New Caledonia.

AARON BAUER

CONTENTS

Volume 2 Number 1

FEATURES

Commentary on conservation of “Sokatra,” the

radiated tortoise (Geochelone radiata) of Madagascar

Ronald A. Nussbaum and Christopher J. Raxworthy

Extinction and extinction vulnerability of amphibians

and reptiles in Madagascar

Christopher J. Raxworthy and Ronald A. Nussbaum

New data on the distribution, status, and biology of

the New Caledonian giant geckos (Squamata:

Diplodactylidae: Rhacodactylus spp.)

Aaron M. Bauer and Ross A. Sadlier

DEPARTMENTS

EDITORIAL

Words from the editor

Craig Hassapakis

Authors

COLUMN—HERPETOFAUNA AND HUMANITY

The United States role in the international live

~'*?

=e «

i tt

" . *

ladagascar, ** <%

‘adjated tortoises

Hefpefofauna extifiction

” “* ,

New Caled@hiia 7

Rhacodactylus getkos

Herpetofauna ang’Humahity (New Golumn)

The trade in live reptiles gneamphibighs R

Tha eral vppert the rec Perm taiass

Cover

Standing’s day gecko Phelsuma stand-

ingi, at Zombitsy Forest, Southwest

Madagascar. This species is restricted

to deciduous dry forest, a habitat that is

; i d declining due to clearing for cattle graz-

reptile trade ing and agriculture. Uncontrolled grass-

Craig M. Hoover land fires which, every year, burn the for-

BOOK REVIEWS est edge have scorched the tree stump.

These day geckos have also had dra-

32 Lizards of the Caribbean: ecology, evolution and matic population declines due to over-

plate tectonics

collecting for the pet trade in some ar-

Review by Brian A. Maurer

eas of its limited distribution. Photo: C. J.

NEWS AND NOTES

Raxworthy.

33 World news

Announcements.»

34__....Meetings

~~ Websites

New literature

Literature

Books and literature received

THE LAST PAGE

35 Glossary

Abbreviation usage

Errata

Acknowledgments “Wi

Dedication

Background screen photograph (see in color on

page 18): Some of the last known surviving Mantella

bemhardi forest habitat cleared for cultivation between

1993-4. Photo: C. J. Raxworthy.

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

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

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

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

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Commentary on conservation of “Sokatra,” the radiated

tortoise (Geochelone radiata) of Madagascar

RONALD A. NUSSBAUM" ? AND CHRISTOPHER J. RAXWORTHY?

Division of Amphibians and Reptiles, Museum of Zoology, University of Michigan, Ann Arbor, Michigan 48109-1079, USA

2Associate Curator, Department of Herpetology, American Museum of Natural History, Central Park West at 79th Street,

New York, New York 10024-5192, USA

Abstract.—The radiated tortoise Geochelone radiata of the desert regions of Southwestern Madagascar, known as “sokatra”’

among most Malagasy, has gained much attention recently as a result of increasing and highly publicized smuggling of

this commercially valuable species. Sokatra have been protected by Malagasy law since 1960 and have been classified as a

Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) Appendix I species since 1975.

Sokatra also are protected to some extent in the four reserves where they are known to occur. In the central part of their

range, on the Mahafaly and Karimbola Plateaux, sokatra are considered fady (taboo) by the Antandroy and Mahafaly people

who live in this area, and they generally are not killed for food in this region. They are, however, killed and eaten by the

Vezo and Antanosy people who largely occupy the northwestern and southeastern limits of the species’ range. It has been

suggested that this taboo is largely responsible for the survival of the species, and there is worry that the taboo may be

breaking down as a result of human famine and intermingling of tribal peoples, many of whom do not consider the flesh

of sokatra to be forbidden. In addition to the possible erosion of taboo barriers, there is strong evidence of increased illegal

trade in sokatra, increased deterioration of its habitat, and increased local consumption of them for food, all of which are

reasons for concern. Given the conspicuousness of this species in nature, its popularity among tortoise fanciers, and

concern for sokatra among conservationists, there is surprisingly little published information about them in their natural

environment that would allow for objective evaluation of their status. Especially needed are intensive studies of the life

history and ecology of sokatra, with special attention paid to determining their limiting environmental requirements. It

will also be important to examine the degree to which zebu (cattle) and goats compete with sokatra for food and to determine

the intensity of zebu and goat grazing that can be tolerated without causing local extinction of sokatra. We have no reason

to believe that the sokatra is threatened with extinction over the next 20 years, just as it obviously was not threatened over

the past 23 years, although classified as Appendix I during that period. We recommend that downgrading the sokatra to

CITES Appendix II might be beneficial to the survival of the species if certain conditions are met, such as careful controls

on the number of legally exported animals. We also strongly recommend the establishment of additional nature reserves

on the Mahafaly and Karimbola Plateaux in the central part of the range of the sokatra, both for survival insurance for this

species and for other rare and endemic species that occur in this area.

Key words. Radiated tortoises, Geochelone radiata, Madagascar, conservation, sokatra, protection, CITES, pet trade, education,

captive breeding, repatriation, monitored legal trade program

Introduction

The radiated tortoise, Geochelone radiata (Plate 1), is one of the

attention from conservationists. Recently, awareness of radi-

ated tortoises reached new heights as a result of a highly publi-

most spectacular of the larger tortoise species. It grows to a

maximum size of about 40 cm carapace length and may weigh up

to 14 kg. Radiated tortoises are readily identified by their color

pattern of bright yellow lines radiating from the center of each

dorsal scute and their yellow legs and throat. Because of their

beauty, and perhaps because they are members of the high pro-

file Madagascan fauna, radiated tortoises are highly coveted by

pet keepers. A large breeding pair may be valued up to $25,000

in the pet trade, and prices in the range of $5,000 for a single, not

necessarily mature, radiated tortoise are not uncommon. Be-

cause of their commercial value, and because they are killed for

food by local Malagasy and served as a delicacy in some Mala-

gasy restaurants, radiated tortoises have received considerable

3Correspondence. Tel: (734) 647-2201; fax: (734) 763-4080;

email: nuss@umich.edu

cized smuggling bust in Orlando, Florida that resulted in the

confiscation of about $250,000 worth of radiated tortoises, spi-

der tortoises, and Madagascan boas and in the conviction on 10

January 1997 of two smuggling partners, a German and a South

African. This headline news was followed by an article (Webster

1997) in a major news magazine that described the business of

smuggling rare animals and featured a color photograph of a

radiated tortoise on the cover.

Considering the great interest in radiated tortoises, it 1s

astounding that so little of scientific merit has been published

about them. Most of the sparse literature consists of anecdotal

and repeated observations. Not a single in-depth study of this

species in nature exists, and we are unaware of any ongoing or

planned research. The most recent field survey of the species

was sponsored by the World Wide Fund for Nature (WWF) and

was done by Richard Lewis, whose report (1995) to WWF-

RONALD A. NUSSBAUM AND CHRISTOPHER J. RAXWORTHY

Aires Protégées, Madagascar, is unpublished. Juvik (1975) re-

viewed the literature and presented some limited new informa-

tion on radiated tortoises. Durrell et al. (1989) commented on

captive breeding programs and the status of the species.

Goodman et al. (1994) reported road count data for a single trip

along one segment of road in the center of the range of the

species. Razafindrakoto (1987) provided the only information

available on food habits and other autecological‘ aspects of the

species in its natural environment at Beza-Mahafaly Réserve

Spéciale. Unfortunately, the population at Beza-Mahafaly has

subsequently been genetically polluted and otherwise influenced

by the release of numerous confiscated tortoises (Lewis 1995),

so that follow-up studies would be of limited value.

Distribution and habitat

“Sokatra,”! or radiated tortoises, are restricted to the xeric’ re-

gion of Southwestern Madagascar (Fig. 1), where they occur in

a variety of habitats ranging from brushy spiny desert domi-

nated by endemic Didieriaceae and euphorbs to gallery forests

dominated by deciduous species such as “kily,” or the tamarind

tree, Tamarindus indica. In this region, annual rainfall is low (<

400 mm) and highly unpredictable, and temperatures are very

high, especially during sunny summer (November-February)

days. It has been claimed that sokatra hibernate during the win-

ter, but, while there are undoubtedly periods of inactivity, we

have seen them active during every month of the year. Within

their range, sokatra are absent from open savannahs and from

forests with no understory vegetation, probably because of their

need for low vegetation for grazing and for frequent shady areas

to escape overheating from insolation. Historically, the eastern

limit of their range was probably determined by the dense, low

elevation rain forests near Télaiiaro. There are no records of the

species north of the Manombo River along the western sea-

board, although it is highly unlikely that the river itself poses a

barrier (they occur on both sides of much larger rivers). Their

restriction to a wide coastal band is somewhat mysterious, but it

may be that higher inland elevations limit them to the coastal

band. Before human occupation of Madagascar, the species prob-

ably occurred somewhat further inland.

Status and protection

Theoretically, sokatra have been protected since 1960 by na-

tional Malagasy law (Decree no. 60-126), which provides for

fines and/or imprisonment for unauthorized collecting. Since

1975, sokatra have been classified as Appendix I species ac-

cording to the Convention on International Trade in Endangered

Species of Wild Fauna and Flora (CITES), which regulates inter-

national commercial trade. The species is further protected in

limited parts of its range that are set aside as nature reserves.

These protected areas are Tsimanampetsotsa Réserve Naturelle

Intégrale (43,200 ha), Cap Sainte Marie Réserve Spéciale (1,750

ha), Beza-Mahafaly Réserve Spéciale (580 ha), and Parcel II of

Andohahela Réserve Naturelle Intégrale (12,420 ha). Two of

these reserves, Beza-Mahafaly and Parcel Il of Andohahela, are

at the limits of the species’ range in areas where sokatra are

uncommon (Andohahela) or consist of populations genetically

polluted by release of confiscated animals (Beza-Mahafaly).

Tsimanampetsotsa and Cap Sainte Marie are in the center of the

species’ range and are potentially important for conservation of

sokatra. Both of these reserves are overgrazed by zebu (cattle)

and goats and are subjected to limited woodcutting, but sokatra

seem to be abundant in both reserves (pers. obs.). There.are no

large, undisturbed reserves on the Mahafaly and Karimbola Pla-

teaux in the region of prime sokatra habitat. A large reserve in

this area is badly needed, not only for sokatra, but also for

protection of many other rare species of plants and animals that

occur in this unique and spectacular environment. These two

plateaux have not been adequately surveyed for biodiversity. A

brief survey done by us at the edge of the Mahafaly Plateau at

Lac Tsimanampetsotsa yielded several rare reptiles, including

two undescribed geckos of the genera Ebenavia (Malagasy leaf-

toed geckos) and Paroedura (Malagasy casque-headed geckos)

[Nussbaum and Raxworthy 1998].

The Antandroy and Mahafaly people of Southwestern

Madagascar regard sokatra as fady (taboo), and eating sokatra

flesh is forbidden. In some areas, especially where they are

associated with tombs, sokatra are believed to contain spirits of

ancestors and are, therefore, sacred as well as fady. It has fre-

quently been suggested (e.g., Juvik 1975) that their status as

forbidden and sacred animals 1s the best protection sokatra cur-

rently has, and without it they might already be extinct. It is

believed (Juvik 1975; Lewis 1995) that the range of sokatra 1s

shrinking and the populations diminishing at the northwestern

and southeastern limits of its range (Fig. 1). This may result

largely from exploitation for food by the Vezo and Antanosy

tribes to the northwest and southeast, respectively. Sokatra

have no taboo status among the members of these two tribes,

and these people eagerly seek sokatra for food. In addition to

exploitation for food, habitat destruction in these two areas is

advanced, especially east of Ambondro, and this undoubtedly

has a negative impact on local sokatra populations.

Sokatra are classified as “threatened/vulnerable” by the

World Conservation Organization (IUCN) [1996 Red List of

Threatened Animals}, which seems justified, given the conspicu-

ousness of sokatra in their habitat, their ease of capture, their

popularity for food and pets, and the ongoing degradation of

their habitat. However, their classification as a CITES Appen-

dix I species is highly questionable. Originally (1973), this clas-

sification was reserved for species that are threatened with ex-

tinction, or could be threatened with extinction within a five-

year period, or have a very limited range. This has been revised

so that now Appendix I species are those “threatened with

extinction which are or may be affected by trade.” Because

“threatened with extinction” can be interpreted very broadly

(there are no guidelines), the current criterion reduces the ques-

tion of status to a matter of opinion, so that almost any species

of commercial value arguably could be classified as Appendix I.

In our view, there is no evidence, published or otherwise, that

indicates sokatra currently are threatened with extinction, al-

though there are reasons for concern. There is not even any

strong evidence that sokatra are less abundant or more restricted

geographically now than they were in 1975 when placed on

Appendix I. Clearly, more objective criteria for CITES classifi-

cations are needed, and, most of all, in the case of sokatra and

many other threatened/vulnerable species, intensive research is

needed to determine the distribution and abundance of the spe-

cies and to identify environmental factors important for main-

taining viable populations.

Population densities

Most information on population densities of sokatra is anec-

dotal or involves estimates from road counts and other rapid

CONSERVATION OF THE RADIATED TORTOISE

survey methods. There are no data that would provide for

meaningful comparisons of past and present population densi-

ties. Juvik (1975) reported that after a rain storm in 1974,

sokatra were encountered at a rate of about one per km along

Route National 10 between T6laiiaro and Toliara, presumably

in the area around Beloha. Goodman et al. (1994) recorded road

counts of about 3 sokatra per km on 30 December 1992 after

heavy rain along 42.8 km of road between Ankororoka and

Beloha. These counts, however, included both living and many

dead sokatra, so that it is necessary to adjust for the accumu-

lation of dead animals. When that is done, the encounter rate is

closer to one sokatra per km. A count of sokatra in January/

February, 1995, along 15 km of road in the region of the

Menarandra River, yielded 30 individuals for a rate of two

sokatra per km (Lewis 1995).

We have traveled extensively, logging many thousands of

kilometers, over roads in this region every year and in every

season since 1989, and we can confirm observations of others

that sokatra activity is greatest during warm weather shortly

after rain. Not a single sokatra was observed during long periods

of drought along the Route National between Tsiombe and

Ampanihy, but, during warm and wet weather, numerous sokatra

can be observed, especially between 40 km south to 30 km north

of Beloha. Normally, sokatra will not be seen during the dry,

cool winters, but on 6 July 1995 we counted 0.36 sokatra/km

along this stretch of road. There had been rain the day before,

and the day of the count was partially cloudy with light sprinkle,

and the temperature was unseasonably warm.

Other than road counts, there are only two reports of den-

sities. Razafindrakoto (1987) indicated a density of 1.3 sokatra

per ha in Parcel I of Beza-Mahafaly Réserve Spéciale, based on

a mark-recapture study. Lewis (1995) estimated densities from

five transect counts in a variety of prime habitats from well

within the core of the species’ distributional area. Lewis’s den-

sity estimates ranged from 262 to 1,077 sokatra per km?. Using

these data, Lewis gave “conservative” total population esti-

mates of 1.6 to 4.0 million sokatra for the core area on the

Mahafaly and Karimbola Plateaux, an area of about 10,000 km’.

Juvik (1975) and Lewis (1995) believed the range of sokatra

was contacting and fragmenting at the northern and eastern ends

of the distributional area. While this is probably true, the his-

torical records that could confirm this unequivocally do not

exist. If humans are the main threat to sokatra, and if the range of

sokatra has been accurately identified, then sokatra have done

remarkably well over the past 2,000 years since humans colo-

nized Madagascar. Most of the negative impact on sokatra popu-

lations has probably occurred over that past 500 years, coinci-

dent with the arrival of Europeans, and more recently with the

explosion of the Malagasy population. There are reliable re-

ports of large numbers of sokatra being shipped to the

Mascarenes for food during the 18" and 19" centuries (Juvik

1975). Passing ships in the Western Indian Ocean regularly took

on large quantities of the larger tortoises from Southern Mada-

gascar and the Mascarene and Seychelles Islands for ship’s stores.

This activity is thought to be responsible for the extinction of

the giant tortoise (Geochelone gigantea) on the granitic Seychelles

Islands.

Prospect

Based on his limited survey taken in 1974, Juvik (1975:145)

believed that “the outlook for the radiated tortoise in Southern

Madagascar is not entirely gloomy, thanks to traditional taboos

on eating its flesh, improved government controls on exports,

and import restrictions in other countries. Its future depends on

the survival of some natural habitat; at the same time... modern

agricultural developments may be indirectly beneficial.” Now,

22 years after Juvik’s optimistic report, the outlook for sokatra

is still not “gloomy,” but there have been significant changes that

are worrisome. Increasing human population pressures in recent

years have resulted in a marked increase in habitat destruction,

harvesting of sokatra for food and pets is clearly on the rise, and

there is no evidence that the Malagasy government has been able

to do anything about it. We agree entirely with Richard Lewis

(1995) who writes, “One is left with the conclusion that there is

a lack of political will to enforce the law.”

Our evidence for increased habitat destruction and harvest-

ing of sokatra stems mainly from our field observations, which

began in the southwest in 1989. Habitat destruction is of four

kinds. First, land is being cleared to establish many new agricul-

tural plots, especially noticeable in the eastern part the range of

sokatra, near Ambovombe and Amboasary. These new fields

include small family-owned subsistence plots of corn, cassava,

sweet potato, and peanuts, but also larger plots for the commer-

cial growing of sisal®. Second, every year during the dry season,

much of Southern Madagascar is deliberately burned, as it is

widely believed that burning increases the growth of plants

needed for grazing of zebu and goats. Burning grasslands inevita-

bly leads to burning of spiny forest and brush, as the grass fires

are not controlled. A significant but unknown amount of sokatra

habitat is lost through uncontrolled burning every year, and it is

our impression that the intensity of burning has increased over

the past eight years. Furthermore, sokatra travel over terrain

slowly compared to most animals and can easily be overcome

by fire, and we suspect that large numbers are killed each year in

this manner. That some tortoises are killed in this way is certain,

as we have seen charred tortoise shells in several burned areas.

Third, the number of zebu and goats grazed is increasing with

human populations, and the intensity of grazing is exacerbated.

At the same time zebu and goat populations are increasing, the

amount of land available for grazing them is diminishing as a

result of conversion of land to agricultural use. It seems certain

that zebu, and especially goats, consume food necessary for the

survival of sokatra, but the intensity of competition is unstud-

ied. There is a critical need to study the effects of zebu and goats

on sokatra, because virtually every square meter of sokatra

range is subject to cattle and goat grazing, even in the reserves

and sacred forests of the Mahafaly. Fourth, and equally worri-

some, is the dramatic increase in woodcutting for firewood, char-

coal making, and construction of houses. Increased woodcutting

is proportional to population growth, and there is a direct effect

of increased agricultural clearing on woodcutting. For example,

now that the town of Amboasary is completely surrounded by

sisal plantations, woodcutters who supply the cooking fires for

that large village range out as far as 30 km to cut wood. Every

day, a steady stream of woodcutters with bundles of firewood

on their backs or in push carts can be seen along the Route

National east of Amboasary, and similar scenes can be seen

along the Route National east of the seaport of Toliara. Along

these same two stretches of national highways, entire villages

based on charcoal making and selling have sprung up. The char-

coal is made deep in the bush and transported to the villages

mainly on the backs of humans.

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found within villages. For example, Beloha is in the heart of

Antandroy country where sokatra are supposedly fady, yet iy

hundreds of discarded shells of eaten sokatra can be seen only

partially hidden in the sisal plants along the roads in the village.

It is not known whether these sokatra are being eaten by less

traditional Antandroy who no longer consider sokatra to be fady

or whether the significant harvest is done by the many non-

Antandroy who live in these regions. Increasingly, except in the

most remote areas of the southwest, there is a mixing of the

various Malagasy tribes, and this may be one of the greatest

threats to sokatra survival, as sokatra are neither fady nor sa-

cred to tribes other than the Antandroy and Mahafaly.

Sokatra are sold openly in restaurants in Southern Mada-

gascar, and neither federal law nor local taboo influences this

commerce. The Antandroy and the Mahafaly apparently place

no pressure on visitors to their lands to respect the fady status

of tortoises although they do insist that sacred tortoises be left

alone. Seated in a small hotely (Malagasy restaurant) on a

winter day, 1995, in Ejeda, a large village dominated by

Mahafaly tribespeople, one of us (RAN) overheard two sol-

diers and a policeman inquiring about sokatra on the menu. It

was available, but they declined it because the price 2,500

Francs Malagasy (FMG) was too high, choosing instead hena

kisoa (pork), which was only FMG 1,500 per plate. At that

time, FMG 2,500 was worth about US$0.55. The restaurant

owners made no attempt to hide their sokatra and the soldiers

200

Cape Ste. Marie

(Reserve)

Fig. 1. Distribution of sokatra in Southeastern Madagascar.

Evidence for increased harvesting of sokatra for local con-

sumption and illegal export is everywhere. Throughout coastal

Southwestern Madagascar, campsites very often are cluttered

with the remains of one to several sokatra that were killed and

eaten (Plate 2). Sometimes these campsites are outside the range

of sokatra, which means the animals were transported there for

the purpose of providing meals. Remains of sokatra are also

and policeman obviously weren’t interested in enforcing the

law. Nor, evidently, were the local Mahafaly opposed to sell-

ing sokatra for food.

On several occasions, we witnessed buses and other ve-

hicles stopping along Routes National within the range of sokatra,

so that occupants could debark to collect a hapless sokatra

observed near the road. RAN once followed a bus for about 15

CONSERVATION OF THE RADIATED TORTOISE

kilometers on the road south of Beloha, which stopped no less

than eleven times to collect sokatra... not one was passed up.

Some of these sokatra may have been destined to become pets,

but most were probably eaten. Some of the passengers of these

buses must have been Antandroy and Mahafaly, and yet there

was a festive atmosphere about collecting the tortoises and no

objections from any passengers.

Plate 2. Remains of a killed and eaten sokatra near a

campsite in Southeastern Madagascar.

In addition to local consumption, sokatra are harvested for

sale in the markets and restaurants in bigger cities, and at least

limited numbers of them are killed and exported for food from

the port of Toliara (Lewis 1995; pers. obs.). On 9 October 1995,

about 20 km east of Ampanihy, one of us (RAN) came across

five large oxcarts filled with.500-800 sokatra being transported

in disturbingly inhumane conditions. The sokatra were piled on

top of one another, fully exposed to the blazing sun, and were

bouncing up and down on the extremely rough road. The oxcart

drivers didn’t want to talk to us, but they didn’t seem worried

about exposure of their illegal cargo. We were informed in

Ampanihy that these tortoises were on their way to market in

Toliara. We have observed, and Lewis (1995) has reported,

dumpsites for sokatra shells numbering up to 300 shells per

dump near Toliara.

Droughts in Southern Madagascar between 1991 and the

present, which caused crop failures and human starvation and

death, resulted in increased killing of sokatra for food. During

the drought of 1992/93, numerous small sokatra appeared in

Tolafiaro where they were being sold to (mainly) the local

Antanosy for food. Inquiries led one of us (RAN) to the discov-

ery that nearly 3,000 sokatra were being sold openly in the

markets at Ambovombe, the Antandroy capital city. The 3,000

were present on a single market day; the rate of flow of sokatra

through the market is unknown. RAN was informed by many

local Malagasy of both Antandroy and Antanosy nationality

that during times of food shortages, nontraditional Antandroy

eat sokatra regardless of the fady (some Malagasy have ceremo-

nies for temporary lifting of various fadys), and of course the

Antanosy openly eat sokatra with great gusto any time they are

fortunate enough to have them. The danger now is that those

who were forced to eat sokatra during this unprecedented fam-

ine will continue to consume them after the famine.

Holidays in Madagascar are especially hard times for

sokatra. Poor families who can’t afford meat everyday will

usually find a way to provide their families with a special

holiday treat. This might be a fowl, but sokatra are often

available for nothing and may be more desirable than fowl for

special occasions.

Two other problems, perhaps of less concern, for sokatra

conservation are the killing of these animals as pests in agricul-

tural areas and for selling as stuffed curios to tourists, mainly in

Antananarivo and Toliara (Plate 3). As land is converted to

crops, sokatra are increasing obliged to forage in fields, and

many sokatra are now killed because of the damage they do in

fields. Many Antandroy who won’t eat them nevertheless do

not hesitate to kill them if their crops are threatened.

A surprising number of Malagasy citizens keep sokatra

for pets, in some cases for pleasure, and in others because they

believe their presence protects their poultry against diseases,

especially louse infestation. Keeping sokatra as pets is not re-

stricted to the south; families in the capital city (Antananarivo)

and other northern villages keep numerous sokatra. In one small

village near Antananarivo, about 30 sokatra are being kept by a

small group of Malagasy villagers (O. Pronk, pers. comm.).

The illegal harvesting and exportation of sokatra for pets is

obviously on the increase, as is indicated by the numerous re-

cent reports of arrests of smugglers and confiscation of these

animals (e.g., Webster 1997). It is easy for tourists to purchase

live sokatra in the larger cities, and customs officials confiscate

significant numbers. On two occasions, we witnessed sokatra

taken from the baggage of Japanese tourists at Ivato Interna-

tional Airport in Madagascar, and there are regular reports of

sokatra confiscated from European (mainly German) travelers

in the Malagasy newspapers distributed in Antananarivo. The

most recent report of which we are aware is in the 25 March

1997 issue of “Midi Madagasikara,” which told the story of 78

sokatra taken from the backpacks of two Japanese tourists at

Ivato International Airport.

Such is the current situation with the sokatra. Their habitat

is being degraded and destroyed at an increasing rate, they are

being harvested by the thousands every year, and local law

enforcement does little to mitigate the situation or stop such

activities. The laws protecting sokatra are well known to the

Malagasy, but they have learned that these laws can be com-

pletely ignored.

It is equally clear that laws to prevent international trade

in radiated tortoises are not working very well. Perversely, it

appears that laws restricting export of radiated tortoises may

do more harm than good. Banning exportation increases the

value of the tortoises, both in the legal and black market trades

(see Lamar 1997, for other species), which increases the deter-

mination of smugglers to find ways to get the tortoises out and

inevitably leads to more corruption of officials charged with

enforcing the laws. With higher values, smugglers can afford to

pay for illegal transport and lay out more money for bribes.

Evidence that embargoes on shipments of pet trade animals

from Madagascar don’t work can be seen in regard to the recent

CITES embargo on most species of day geckos (Phelsuma) and

chameleons from Madagascar. Since this embargo went into

effect 20 January 1995, the banned species have continued to

arrive in Europe and the United States in large numbers, and at

least in Europe the prices have declined suggesting they are

arriving in greater numbers than before (O. Pronk, pers. comm.).

Economically, this may be because the smaller day geckos and

RONALD A. NUSSBAUM AND CHRISTOPHER J. RAXWORTHY

chameleons can be smuggled in greater numbers than tortoises,

and because smugglers do not have to pay the export tax that is

levied by the Malagasy government against legal animal ex-

porters.

Bee yi

ate 3. C. J. Raxworthy examines a stuffed sokatra for sale to

tourists in the Analakely market in Antananarivo, Madagascar.

An additional negative effect of banning export of radiated

tortoises (and other species) is the inhumane treatment of the

exported animals. Smugglers do not have to comply with Inter-

national Air Transport Association (IATA) rules regarding hu-

mane packing and shipment of commercial animals and, indeed,

probably cannot comply with these conditions because of the

need to hide the animals and use circuitous and prolonged routes

for their movement. Undoubtedly, the percentage of dead, in-

jured, and unhealthy animals arriving at their destinations is

higher in illegal shipments than in legal shipments.

It seems certain that local traditions have had more impact

than federal and international laws in conserving the sokatra.

But it also seems certain that the forbidden status of sokatra

among the Mahafaly and Antandroy will erode as human popu-

lations and the need for protein and money increases and as

people from other tribes without fady constraints continue to

immigrate into the range of the tortoise. Much serious thought,

research, and considerable effort will be needed to insure the

survival of this increasingly vulnerable species.

THOMPSON

PAUL

Solutions

It has repeatedly been argued that education of the Malagasy is

the key to the survival of the sokatra. This seems unlikely.

Many well-educated Malagasy, who are well aware of conserva-

tion issues, keep sokatra as pets and regularly eat them because

they taste good. Poorly educated Malagasy, those who live in

the bush and survive off the land and eat tortoises because they

need food, are largely immune to education, and, in any case, all

the education in the world will not stop them from doing what is

necessary to survive. The dilemma these poor Malagasy face is

dramatically demonstrated in Webster’s (1997) article.

Education, especially in regard to resource management,

may be of some value if directed at land managers and guardians

of the Reserves. During a visit to Cap Sainte Marie, RAN was

proudly informed by the guardians of the Reserve that they

regularly collected tortoises observed on the limestone pla-

teaux and placed them in one of the steep-sided canyons where

the tortoise could not get out, find more food, and were pro-

tected from poachers. It apparently hadn’t occurred to these

guardians that the tortoises were numerous on the plateaux

because the conditions there are good for them and that taking

the tortoises from their familiar home ranges might be disas-

trous for them. Furthermore, the guardians had not considered

the negative effects of artificially concentrating tortoises in a

habitat which they should have realized was suboptimal for

the tortoise, otherwise there would have been more tortoises

there naturally. Finally, the guardians should have realized that

concentrating the tortoises in a canyon might actually make it

easier for poachers to collect them. Education might also help

to stop the genetic pollution caused by the irresponsible re-

lease of confiscated animals.

It has also been suggested that captive breeding programs

are a way to ensure survival of the species. Such a program is

the American Zoo and Aquarium Association’s Species Sur-

vival Plan (AZA/SSP) for the radiated tortoise, underway

through a consortium of zoos, in which tortoises would ulti-

mately be repatriated, to the natural environment under the

auspices of Malagasy guardianship. There are obvious prob-

lems with this approach, not the least of which is that it will do

no good to repatriate captive-bred tortoises to an environment

that will not sustain them. If the ecosystem no longer supports

wild-bred tortoises, then why should we expect captive-bred

tortoises to do any better? If the natural environment will not

support sokatra, then captive-bred animals are best kept where

they are... in zoos. If natural populations still exist, then they

should not be genetically polluted by release of captive-bred

animals. Captive-bred animals should be used to establish popu-

lations in nature only if natural populations no longer exist and

only if conditions that insure protection of the released ani-

mals are in place.

Aside from educating resource managers and guardians, what

are the solutions to conservation of sokatra? Something should

be done to (1) curtail habitat degradation, (2) reduce harvesting

of sokatra for local consumption, and (3) control the exporta-

tion of sokatra in the pet trade.

It seems highly unlikely, given the inexorable human popu-

lation growth and the Malagasy local traditions, that much can

be done to slow habitat degradation. The best that can be done is

to establish one or more large nature reserves on the Mahafaly

and Karimbola Plateaux in areas that are not currently being

converted to agricultural plots and have little value for agricul-

CONSERVATION OF THE RADIATED TORTOISE

= 4a ~ " 5

: oo) ee =; : soa

Hatchling ra

ture. This might be done in conjunction with the large and rela-

tively undisturbed sacred forests where Mahafaly kings are bur-

ied. These sacred forests are not immune to cattle grazing, and

neither are the reserves. Therefore, studies are needed to learn

the impact of grazing on tortoise populations and to determine

the maximum allowable grazing within reserves. Control of graz-

ing and poaching of tortoises on the reserves would be impos-

sible without the cooperation of local and federal authorities

and, more importantly, without some reward for local support

of law enforcement. This would have to include compensation

for lost income from reduced grazing of cattle‘and harvesting of

tortoises. Threats of punishment alone won’t work.

Reduction of local consumption of sokatra will be diffi-

cult, especially in areas outside of the Mahafaly and Antandroy

homelands. Antanosy and other Malagasy connoisseurs of

sokatra will always eat them, regardless of the law. Malagasy

law regarding the sokatra has been widely ignored for so long

that any attempt to enact strict enforcement locally would lead

to serious problems with which the Malagasy authorities are

not adapted to cope. However, we note that although lemurs

(family Lemuridae) are still consumed for food in Madagascar,

the law protecting lemurs is both more widely enforced and

respected than are the laws protecting sokatra, perhaps as a

result of more intense public scrutiny related to the economy of

tourism and because of greater international involvement.

If the international community, through agencies such as

United States Agency for International Development (US AID),

can be persuaded to act more responsibly by promoting mean-

ingful projects in Southern Madagascar that increase the food

supply (relative to human population density) and by reacting

0. = a . Po ae =

diated tortoise Geochelone radiata. Photo courtesy «

=, . <

of R. D. Bartlett.

more swiftly to drought and famine, then desperation consump-

tion of sokatra might be reduced. This would mean providing

food rich in protein, and not just surplus corn and bulger wheat,

during times of famine. In the area around Tolaiiaro, much of the

surplus food given to Malagasy by aid agencies during the recent

drought, especially the dried corn, was either fed to livestock or

sold for pittance so that the puzzled owners could buy real

food. The problem of consumption of sokatra is probably in-

tractable in peripheral areas, but creation of additional reserves

and strong rewards for respecting the boundaries of the reserves

would help to insure the survival of sokatra in the core area of

their distribution.

There are two options for reducing the impact of the illegal

commercial pet trade on natural populations of sokatra. The

first is to enact a monitored legal trade program that strictly

limits the number of wild-caught sokatra that could be exported

and would generate some income for the Malagasy government

through taxation on exports of sokatra. This would be espe-

cially desirable if the generated income were used to support

monitoring programs and research on the tortoise. Such a solu-

tion would require downgrading of the sokatra from Appendix I

to Appendix II, which theoretically shouldn’t be a problem,

because it is clear that the sokarra is currently misclassified by

CITES, as it is not currently “threatened with extinction.” If the

species was downgraded to Appendix II, then a limited number

of both wild-caught and first generation, captive-bred sokatra

could be legally exported as could confiscated animals (under the

monitored program), a better solution than genetic adulteration

of natural populations through random release into the environ-

ment. Limited legal export (with controls such as internal pas-

RONALD A. NUSSBAUM AND CHRISTOPHER J. RAXWORTHY

Plate 4. Confiscated sokatra at a Malagasy government station in lvoloina (September 1985).

sive integrated transponder [PIT] tag markers) would destroy or

greatly reduce the market for smuggled animals and prevent

inhumane shipping methods.

Realistically, downgrading to CITES IH might be politically

impossible, so a second solution would be to encourage captive

breeding programs in Madagascar that would eventually yield

second generation, registered, captive-bred tortoises that could

be legally exported under a monitored program. This could be

done under a partnership with the Malagasy government and

captive breeders in which the Malagasy government retains

ownership of the tortoises and in which some of the income

would be returned to the Malagasy to support tortoise conser-

vation. Perhaps third-party monitoring agencies would be re-

quired to insure against corruption. The nucleus for a monitored,

legal export program for sokatra already exists in Madagascar at

Ivoloina (Plate 4), where the Malagasy government (Eaux et

Foréts) keeps confiscated tortoises. Under either program, it

might be possible to supply tourists with legal tortoises, under

the same kind of regulation used for exporting semiprecious

stones and other valued objects from Madagascar. Such pro-

grams would not entirely eliminate illegal trade but might reduce

it considerably, as profit from illegal activity would be greatly

diminished.

Acknowledgments.—We thank Richard E. Lewis for giving

us access to his unpublished report on radiated tortoises and for

permission to cite it, and Olaf Pronk for useful insights into the

intricacies of the pet trade. Our research in Madagascar has been

funded by the National Science Foundation, National Geographic

Society, and Earthwatch. We have received logistic support from

Conservation International, WWF, and US AID.

References

Durrell, L., Groombridge, B., Tonge, S., and Bloxam, Q. 1989.

Geochelone radiata radiated tortoise, sokake, p. 96-98 in

Swingland, I. R. and Klemens, M. W. (editors). The Conserva-

tion Biology of Tortoises. Occasional Paper, IUCN No. 5, Gland,

Switzerland. Iv + 202 p.: illustrations, maps.

Goodman, S. M., Pidgeon, M., and O’Connor, S. A. 1994. Mass

mortality of Madagascar radiated tortoise caused by road con-

struction. Oryx 28:115-118.

IUCN. 1996. 1996 IUCN Red List of Threatened Animals. YUCN,

Gland, Switzerland. 448 p.

Juvik, J. O. 1975. The radiated tortoise of Madagascar. Oryx 13:145-

148.

Lamar, W. W. 1997. CITES: boon or boondoggle? Herpetological

Review 28:10.

Lewis, R. E. 1995. Status of the radiated tortoise (Geochelone ra-

diata). WWF-Madagascar, unpublished report.

Nussbaum, R. A. and Raxworthy, C. J. 1998. Revision of the genus

Ebenavia Boettger (Reptilia: Squamata: Gekkonidae).

Herpetologica 54(1):18-34.

Razafindrakoto, L. A. 1987. Contribution a l’etude bioecoetho-

logique de Geochelone radiata (Shaw 1802) (famille de

Testudinidae) dans la Réserve Spéciale de Beza-Mahafaly.

Mémoire de Fin d’Etude, EESSA, Universite de Madagascar,

Antananarivo, Madagascar.

Webster, D. 1997. The looting and smuggling and fencing and hoard-

ing of impossibly precious, feathered and scaly wild things: the

$10 billion dollar black market in endangered animals. New

York Times, February 16, 1997, Section 6, p. 26-33, 48-49, 53,

61.

Footnotes

The Malagasy name for the radiated tortoise varies regionally in Madagascar: “Sokatra”

(Merina), “sokaka” (Antandroy and Antanosy), and ‘kotroky” (Mahafaly) are commonly used.

These words are pronounced “sookot,” “sookock,”’ and “kootrook, ” respectively.

2For more information on protected areas (policy and legislation, international activities,

administration and management, system reviews, addresses, protected area information, defi-

nitions of protected area designations, as legislated, together with authorities responsible for

their administration, maps, and thé 1993 United Nations list of national parks and protected

areas), travel, health information, global biodiversity hotspots, and country information consult

the following sources which were used to write this country sunimary: Protected Areas: World

Conservation Monitoring Centre's Protected Areas website location: www.unep-wenic.org/

parks/index.hitml and the IUCN, 1992. Protected Areas of the World: a review of national

systems. IUCN, Gland, Switzerland.and Cambridge, United Kingdom. Xx + 352 p. Also, avail-

able via the Internet at: www.unep-weme.org/protected_areas/data/pa_world_text.html: Travel:

United States State Department, Bureau of Consular Affairs website: travel.state. gov; Health:

Shoreland’s Travel Health Online: www.tripprep.com: Global Biodiversity Hotspots:

‘wwnw.conservation. org/hotspots/default.htm: and Country information: United States Central

Intelligence eney’s (CIA) World Factbook, C/A, Washington, D.C. Also, available via the

Internet at: www.odei, gov/cia/publications/factbook

Manuscript received: 26-April-1997

Accepted: 15-January-1998

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

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

Extinction and extinction vulnerability of amphibians

and reptiles in Madagascar

CHRISTOPHER J. RAXWORTHY' AND RONALD A. NUSSBAUM?

1Associate Curator, Department of Herpetology, American Museum of Natural History, Central Park West at 79th Street,

New York, New York 10024-5192, USA *Division of Herpetology, Museum of Zoology, University of Michigan,

Ann Arbor, Michigan, 48109-1079, USA

Abstract.—In Madagascar, only two herpetofaunal extinction events are well documented. Both are extinctions of subfos-

sil giant tortoises, which coexisted with humans for more than 1,000 years. Modern extinctions of amphibians and reptiles

are also likely, but researchers and conservationists are probably overlooking these extinction events, because the most

vulnerable species, with small relict populations, are easily missed during regional surveys. To date, conservation pro-

grams in Madagascar have largely ignored many relict distribution species, restricted to transitional or rare habitat types.

We provide four examples of species with relict distributions that we consider vulnerable to extinction. Based on ongoing

surveys and systematic revisions, many new herpetofaunal species will be described in the future, some of which may

require rapid conservation efforts to prevent extinction.

Conservationists frequently measure extinction vulnerability using the World Conservation Union (IUCN) Red Lists,

and fifteen endemic Malagasy amphibians and reptiles are included in the IUCN Threatened List. However, this list

appears to reflect a historical bias towards conserving turtles and boas in Madagascar, listing eight species in these groups,

although they represent just 2 percent of the island’s actual endemic herpetofauna. Ironically, this taxonomic bias may

hinder attempts to prevent herpetofaunal extinctions, by promoting some species for conservation activities that are not

vulnerable (e.g., tolerant of human habitat modification, or widespread) and ignoring many species that are soon to be lost.

For Malagasy amphibians and reptiles, biogeographic data appear to provide more objective criteria with which to assess

extinction threats rather than suspected rates of population decline.

Key words. Extinction, conservation, Madagascar, herpetology, biogeography, reptiles, amphibians

Introduction Madagascar’s “megafauna” extinction event that occurred dur-

Despite the considerable interest in the amphibian and reptile ing the past two thousand years (Dewar 1984). The reasons for

faunas of Madagascar and the widely held view that the island their extinctions are unknown, but it is certain that they coex-

represents one of the world’s top conservation priorities (e.¢., isted with humans on the island between 2000 Before Present

Wright 1997), it is surprising that there has been so little discus- (B.P.), the date of earliest human occupation of the island

sion regarding the patterns of extinctions (past, present, or fu- (MacPhee and Burney 1991) and 750 B.P., the youngest carbon

ture) for these two highly diverse and largely endemic groups. date for giant tortoise subfossils in Madagascar (Burleigh and

This is of special concern because the objective of most conser- Arnold 1986). Raxworthy and Nussbaum (1996) have suggested

vation programs in Madagascar is to maintain biodiversity. There- that the modern day practice of regular (typically annual) burn-

fore these programs should prevent, or at least minimize, future ing of grassland and its peripheral forests, over much of the

extinctions. island, was responsible for destroying the original habitats of

The purpose of this paper is to provide a summary of grazing animals such as giant tortoises, while MacPhee and Marx

herpetofaunal extinctions and extinction vulnerability in Mada- (1997) consider their extinction may be due to the human intro-

gascar: first by reviewing the evidence of extinction; second by duction of exotic pathogens that caused hyperdisease.

providing examples of species we believe are at imminent threat The former distribution of both species of giant tortoises

of extinction; and third by evaluating and summarizing methods was vast, and their remains frequently represent the most com-

used to measure risks of extinctions by the conservation com- mon subfossil material recovered by paleontological excavations

munity using the World Conservation Union (IUCN) Red List (Dewar 1984). Because of their former large distribution and

criteria. fact that much of island of Madagascar has still been poorly

surveyed, there remains a chance that either species may survive

Subfossil extinctions as a small population in one of the remoter regions, although this

The only well documented cases of herpetofaunal extinctions in possibility must now be considered extremely unlikely.

Madagascar are for the two species of giant tortoises, No other extinct reptile or amphibian subfossil material is

Dispsochelys grandidieri and Dispsochelys abrupta, which once known from Madagascar, and there are no historically docu-

occupied a large area of the central and western region of the mented cases of species going to extinction.

island (Bour 1984). Both species went extinct during

Possible extinctions

‘Correspondence. Fax: (212) 769-5031; email: rax@amnh. One possible extinction claimed in the literature (IUCN/UNEP/

org WWE 1987) is the colubrid snake Liophidium apperti. This

Plate 2

Plate 3

Plate 7 Plate 8

Plate captions: 1. Antanosy day gecko Phelsuma antanosy, from a Petriky relict forest fragment that has now been lost. 2.

Chamaeleo belalandaensis, at Belalanda. One of Madagascar’s rarest chameleons. 3. Angonoka tortoise Geochelone yniphora,

photographed at Ampijoroa. 4. Adult big headed Madagascar side-necked turtle Erymnochelys madagascariensis, photographed at

the Andranomiditra River, Bevazaha village, Ankarafantsika Reserve. Photo: Gerardo Garcia Herrero. 5. Tomato frog Dyscophus

antongili, Antongil’s Bay. 6. Dumeril boa Boa dumerili. Photo courtesy of Kevin and Sue Hanley. 7. Madagascar boa Boa madagascariensis.

Photo: R. D. Bartlett. 8. Sanzinia tree boa Boa manditra. Photo by Peter Stafford, courtesy of The Natural History Museum, London. Photos

1 and 2: C. J. Raxworthy. Photos 3 and 5 courtesy of Franco Andreone, Museo Regionale di Scienze Naturali, Torino, Italy.

Plate 12

Plate 15 Plate 16

Plate captions: 9. Campan’s chameleon Furcifer campani. 10. Labord’s chameleon Furcifer labordi (female), Kirindy. 11. Minor

chameleon Furcifer minor (female), photographed at “Mandraka Breeding Centre.” 12. Standings day gecko Phelsuma standingi. 13.

Golden mantella Mantella aurantiaca, from Andasibe. Photo: C. J. Raxworthy. 14. Pyxis planicauda, photographed at Ampijoroa. 15.

Radiated tortoise Geochelone radiata. 16. Armored chameleon Brookesia peramata. Photos 9 and 12 courtesy of R. D. Bartlett. Photos

10, 11, and 14 courtesy of Franco Andreone, Museo Regionale di Scienze Naturali, Torino, Italy. Photos 15 and 16 courtesy of Kevin and

Sue Hanley.

CHRISTOPHER J. RAXWORTHY AND RONALD A. NUSSBAUM

species was described from a single specimen collected from a

forest 7 km north of Befandrina-sud in 1968 (Domergue 1983).

Domergue reported that the forest had been subsequently cleared,

which prompted IUCN/UNEP/WWE (1987) to consider that

“the survival of this snake must now be in question.” We think

it is premature to consider this species to be extinct, because this

region remains so poorly studied, and it appears that similar

forest types still survive in nearby sites in the Morombe region.

We have collected Liophidium c.f. apperti at several localities in

southern Madagascar, and although these specimens require fur-

ther taxonomic study, this also suggests this species is not con-

fined to Befandrina-sud. :

Possibly the best candidate we have for an extinction event

occurring during the past 100 years is for the colubrid snake

Pseudoxyrhopus ankafinaensis, which was not described until

1994. This very large species is represented by just a single

specimen collected in 1880 from montane forest of the High

Plateau. Forest of this type is now almost completely degraded

in this region (Raxworthy and Nussbaum 1994), and the absence

of observations of P. ankafinaensis from surviving forest of

lower elevation, or other montane forest sites suggests it was

endemic to this high elevational region of the High Plateau.

Dubious extinctions

Other species have not been found for more than one hundred

years, and, therefore, could also be considered extinct. However,

before this conclusion can be made with any certainty, it is

important to confirm that the species were collected in Mada-

gascar. During the 1800's, specimen localities were sometimes

confused or lost, frequently because museums were receiving

collections from throughout the world during this period. An

example is the colubrid snake Pseudoxyrphopus punctatus, which

was thought for a period of more than 50 years to have been

collected in Madagascar. Subsequently, this was identified as a

Brazilian snake in the genus Sordellina (see Raxworthy and

Nussbaum 1994).

Another possible example of a species that may not ever

have been collected in Madagascar is Ailuronyx trachygaster.

This giant gecko, known only from a single specimen MNHP

6679 (Muséum National d’ Histoire Naturelle, Paris) was col-

lected by an unknown person from a locality listed in the Paris

Museum as Madagascar. Interestingly, the two specimens cata-

logued before A. trachygaster: MNHP 6677-8, are both

Ailuronyx seychellensis collected in the Seychelles (6677 was

collected by Péron and Lesueur). Possibly A. trachygaster was

also collected on the same voyage. A. trachygaster 1s a valid

species, with very different characters compared to A.

seychellensis. Despite some uncertainty about the geographic

origin, we suspect A. trachygaster is extinct because no new

specimens (of what should be a very conspicuous gecko) have

been found in more than 140 years.

As aresult of recent surveying in Madagascar, most of the

rarer species in collections have been rediscovered, such as

Zonosaurus boettgeri, Phyllodactylus brevipes, Paragehyra petiti,

Uroplatus alluaudi, and Pseudoxyrhopus ambreensis, but a few

notable exceptions remain, especially among the most cryptic

groups. For example, among the skinks, the following species

have not been collected since their original description: Mabuya

betsileana (possibly African), Cryptoscincus minimus,

Pseudoacontias madagascariensis, and Paracontias rothschildi.

Their exact distributions remain unknown at present (in Mada-

gascar or elsewhere) but we consider it premature to. consider

any of them to be extinct.

Vulnerable to extinction

We consider species in the category “Vulnerable to Extinction”

to have populations that are sufficiently small that near-future

extinction in the wild can be considered highly likely (without

conservation action). These species are restricted to primary

habitat that has declined so dramatically, that they now survive

only in tiny isolated patches, which are continuing to decline.

As a result, the species dependent on this primary habitat are

now endemic to a very small region of the island. To illustrate

this type of extinction vulnerability, we have selected four spe-

cies to serve as examples. However, many other herpetofaunal

species exhibit similar extinction vulnerability in Madagascar.

Bernard’s mantella frog Mantella bernhardi (Plate 18A

and B). During a survey of Tolongoina made by Nussbaum in

1993, M. bernhardi was first discovered, in a single patch of

relict forest. Within months, commercial collectors visited the

locality and supplied André Peyrieras, a commercial exporter,

with animals to be sold into the pet trade. Vences et al. (1994)

described this species based on animals they obtained from

commercial collectors working for Peyrieras. They provided

no data on the exact locality, habitat requirements, or conser-

vation concerns of this species, because they never saw this

species in the wild.

A subsequent visit to the Tolongoina region by Raxworthy

in 1994 discovered that the only known M. bernhardi habitat

had been further cleared, so that no more than 20 ha of forest

survived (Plate 17). No other populations were discovered dur-

ing this visit, and almost all other primary rain forest had been

cleared from the area. Deforestation of primary forest has been

so extensive in this region that almost no fragments now survive

east of the Faraony River. Topographic maps indicate that this

forest was almost entire during aerial photography made of the

region between 1950 and 1965 (FTM 1974).

Although it is likely that M. bernhardi once had a more

widespread distribution, the deforestation pattern of low eleva-

tion rain forest (below 800 m elevation) in this region suggests

that little forest of this type now survives. The closest pro-

tected forest of this type occurs in the lowest elevational areas

of the Ranomafana National Park (the southern boundary limit

is 18 km to the northwest). It is not known if this species occurs

there, or even if the habitat is similar to that at Tolongoina. The

low elevation rain forest at Manombo Reserve (140 km to the

south) does not appear to have populations of M. bernhardi

based on a survey by Raxworthy in 1991.

Because no known populations of M. bernhardi occur

within a protected area and its habitat appears to have been

almost completely destroyed, we consider this species extremely

vulnerable to extinction.

Antanosy day gecko Phelsuma antanosy (Plate 1). This

day gecko is restricted to coastal fragments of forest in the

Tolagnaro region of southeastern Madagascar. Raxworthy and

Nussbaum (1993) described three sites (forest fragments) in the

description of this species at Petriky, Ste. Luce, and Tapera.

Since then, one new site has been discovered near Manambaro,

and one site (the Petriky fragment, area 81 ha in 1989) has been

completely destroyed. None of the three surviving forest frag-

HERPETOFAUNAL EXTINCTION IN MADAGASCAR

ments is greater than 191 ha, and the Manambaro site 1s decreas-

ing in area rapidly as a result of annual burning.

Unlike some Phelsuma species, P. antanosy does not sur-

vive in degraded or heavily modified habitats and appears to be

entirely dependent on the transitional dry-humid forests that

are restricted to a small region of the southeast. This habitat has

now been almost completely lost from the region, having been

degraded and cleared for charcoal production and agricultural

land. None of the forest fragments where P. antanosy occurs is

within a protected area.

The surviving populations are now so small that we con-

sider this species to be extremely vulnerable to extinction. Con-

servation efforts are required if the last fragments of habitat are

to be saved.

Belalanda chameleon Chamaeleo belalandaensis

(Plate 2). Very little information exists on either the distribution

or habitat requirements of this chameleon. However, all data to

date suggest it is endemic to a tiny region of Madagascar. The

only locality is Belalanda, Southwestern Madagascar, where

Raxworthy has recorded individuals as recently as 1995. How-

ever, we were unable to find this species at other sites, despite

intensive herpetofaunal surveys within 10-50 km of Belalanda.

The habitat where this chameleon was recorded is degraded gal-

lery forest, which has now been almost completely cleared.

Because C. belalandaensis was not found in the other surviving

primary forest habitats of the region, we suspect it is restricted

to gallery forest.

This chameleon is an example of a species for which bio-

geographic data are urgently needed. It would be valuable to

record the exact distribution limits for this chameleon, so that

conservation efforts could be directed at those populations, which

appear to be most viable. The species has already been subject

to some commercial collecting, but the impact on the population

is not known. The very localized distribution of C. belalan-

daensis, in part confirmed by our survey efforts in surrounding

areas, suggests that this species is vulnerable to extinction be-

cause of the apparently tiny surviving populations.

Angonoka tortoise Geochelone yniphora (Plate 3). This

tortoise is restricted to an area of less than 1,000 km’ in the Baly

Bay area of Western Madagascar. The distribution appears to be

relict, because the species is now confined to two isolated areas,

one to the east of the Baly Bay (Cap Sada), and the other to the

west (Belambo). Dispersal across the bay is likely to be very

limited or impossible. Therefore, the eastern and western popu-

lations appear to be genetically isolated. Only five sites are

known or suspected to have G. yniphora populations (Durrell

et al. 1994). The suspected sites are based on interviews with

local people and the occurrence of suitable habitat. The wild

populations are thought to have declined recently due to habitat

loss, predation of eggs, and juveniles by African bush pig

(Potamochoerus larvatus), and collecting by people (Durrell et

al. 1994). At Cap Sada the first detailed population study is

now underway.

The isolated eastern and western populations of G. yniphora

suggest this species was previously distributed to the south of

Baly Bay and that the distribution range has contracted since

this time. The habitat of this tortoise is a mosaic of deciduous

forest and bamboo scrub, which appears to be replaced by a

palm savanna as a result of frequent burning. Between 1949-

1973, Curl et al. (1985) reported only minor change in tortoise

habitat distribution, and suggested this habitat is no longer de-

clining. However, it should be noted that during this same pe-

riod, the setting of fires was both illegal and frequently enforced

by local communities.

The tiny isolated populations and the restricted area of

surviving habitat clearly indicate that G. yniphora is vulnerable

to extinction if further habitat decline continues. A major con-

servation program (Project Angonoka) coordinated by the Mala-

gasy Water and Forests Authority and Jersey Wildlife Preserva-

tion Trust is now underway in the Cap Sada region to protect

this population.

IUCN Threatened species

This section includes those species listed with a threatened cat-

egory (Critically Endangered, Endangered, or Vulnerable) in the

most recent IUCN Red List (IUCN 1996). Threatened species,

as recognized by IUCN, refers specifically to the level of risk of

extinction. Fifteen endemic Malagasy amphibians and reptiles

are currently listed as threatened in the Red List; three species

are classified as Endangered, and another 12 as Vulnerable. A

significant advance with the new IUCN categories 1s that threats

are presented in a quantified format. A summary of these en-

demic species, as well as their perceived risks of extinction, is

given in Table 1.

IUCN Endangered species

Big headed Madagascar side-necked turtle Erymno-

chelys madagascariensis (Plate 4). The criteria used for

considering this turtle as endangered is based on a 50 percent

population decline in 10 years or three generations. The genera-

tion time (average age of parents in the population, as used by

IUCN) for this turtle is unknown, but without doubt will be

greater than 10 years. For long-lived species IUCN suggests a

cap of 75 years IUCN 1996). Even working with this time

span, however, the problem is a lack of population data for this

species, both modern and historical.

There is no doubt that populations are being exploited by

fishing practices, and Kuchling and Mittermeier (1993) have

presented evidence that two populations have gone extinct out

of a sample of nine lakes. Nevertheless, these authors recog-

nized that the status of river populations have not been esti-

mated because of an almost complete lack of surveys within the

rivers of Western Madagascar. In addition, we still lack modern

data on either lake or river populations throughout much of the

species’ distribution range, especially the many remote regions

of the west.

Because our knowledge of the populations of E.

madagascariensis is so incomplete, it is questionable if we can

claim even a suspected population reduction of 50 percent. The

large historical distribution area of this turtle, with an extent of

occurrence of approximately 100,000 km’, does not suggest to

us that this species is yet at high risk to extinction, although

clearly more field work is required to determine the current

distribution of this species.

Angonoka tortoise Geochelone yniphora. Like E.

madagascariensis, the three-generation time for G. yniphora will

be greater than 10 years. We are unaware of data to support a 50

percent population decline over either 10 years or three genera-

tions. However, this tortoise does qualify for Endangered status based

CHRISTOPHER J. RAXWORTHY AND RONALD A. NUSSBAUM

fy a

Plate 17. Som

Raxworthy.

we pote as nee can Gi de : = Sn ea A MR LP ae s a 2

Plate 18A. Bernard’s mantella frog Mantella bernhardi. Probably Plate 18B. Mantella bernhardi (belly pattern). Photo courtesy of

the rarest Mantella species. Photo courtesy of Franco Andreone, Franco Andreone, Museo Regionale di Scienze Naturali, Torino,

Museo Regionale di Scienze Naturali, Torino, Italy. Ttaly.

HERPETOFAUNAL EXTINCTION IN MADAGASCAR

on the criteria of an extent of occurrence less than 5,000 km?, and

less than 5 isolated populations (see earlier).

Madagascar flat tailed tortoise Pyxis planicauda (Plate

14). Adult wild P. planicauda are reported to have 10-30 growth

rings (Kuchling and Bloxam 1988), which appear to reflect the

growth between each period of annual aestivation’. Using a mean

generation time of 20 years, three generations would represent

60 years. The Endangered category given to this tortoise is based

on a 50 percent decline in the population during this period.

Without question, populations of this tortoise are declin-

ing due to habitat destruction for cultivation (Kuchling and

Bloxam 1988; Bloxam et al. 1993; Raxworthy, pers. obs.). How-

ever, our knowledge of the species distribution is actually im-

proving. New localities are being discovered, including impor-

tant range extensions further to the north (Behler et al. 1993;

Bloxam et al. 1993). The extent of occurrence for P. planicauda

is currently about 500 km?(based on the localities given in Fig. 1,

Behler et al. 1993), with the species endemic to a small region of

coastal, western, deciduous forest between the Morondava and

Tsiribihina Rivers. This justifies P. planicauda being considered

Endangered (the extent of occurrence is significantly less than

5,000 km?) based on its small distribution, rather than the crite-

ria of rate of population decline. The localized distribution of

this species, and its dependence on native forest, suggests this

species is vulnerable to extinction. i

IUCN Vulnerable species

Of the 12 species classified as Vulnerable, 11 are so classifica-

tion based on a criteria of a 20 percent reduction in population

over 10 years or three generations. The mean wild generation

time is unknown for any of these species, although a study of

radiated tortoise Geochelone radiata at one site has yielded

some data on the population age structure (Razafindrakoto 1987).

The population size and rate of decline have never been mea-

sured for these species, although populations of some species

restricted to primary habitats are declining in areas subjected to

habitat loss. For those species restricted to primary habitats, it

may be reasonable to positively correlate rates of habitat loss

(e.g., based on satellite images) to rates of population decline.

Four of the Vulnerable species: the tomato frog Dyscophus

antongili (Plate 5), the Dumeril boa Boa dumerili (Plate 6), the

Madagascar boa Bea madagascariensis (Plate 7), and the

Sanzinia tree boa Boa manditra (Plate 8), are not restricted to

primary vegetation. D. antongili is found in secondary habi-

tats, such as around villages and in areas of cultivation

(Raxworthy 1991), and even in towns such as Maroantsetra

(Glaw and Vences 1994). All three of the Malagasy boas (B.

dumerili, B. madagascariensis, and B. manditra) are frequently

found in heavily degraded habitats and cultivated areas, even

including close proximity to villages where they are probably

feeding on commensal rats. For these species, the loss of pri-

mary forest does not lead to local extinction, although their

population changes (negative or positive), when primary for-

est is converted, are not known.

The most likely source of population decline for D.

antongili, and the three species of boa, is commercial collect-

ing, although now all four species are listed on the Convention

on International Trade in Endangered Species of Wild Fauna

and Flora (CITES) Appendix I, which prevents legal interna-

tional trade. Malagasy boas are collected in regions such as

Marovoay to supply a domestic leather trade. However, boas

continue to be among the most common snakes found in Mada-

gascar, and the scale of commercial collecting may be compa-

rable to the number of road kills that can be seen throughout

Madagascar during the rainy season. The approximate extent

of occurrence of each species is: D. antongili 10,000 km’, B.

dumerili, 120,000 km?, B. madagascariensis 40,000 km?, and

B. manditra, 100,000 km’. Based on the large distribution area,

and the broad tolerance to habitat degradation, none of these

species appear at risk to extinction.

Of the other eight vulnerable species which are restricted

to primary habitats, it is impossible for us to assess the criteria

used for the following species: Campan’s chameleon Furcifer

campani (Plate 9), Labord’s chameleon Furcifer labordi (Plate

10), the Minor chameleon Furcifer minor (Plate 11), Standing’s

day gecko Phelsuma standingi (Plate 12), and the golden

mantella Mantella aurantiaca (Plate 13), because these are based

on rates of population decline (observed or suspected) for

which we have been unable to find obvious supporting data.

Table 1. IUCN threatened species of endemic amphibians and reptiles of Madagascar (IUCN 1996).

SPECIES CATEGORY

Erymnochelys madagascariensis ENDANGERED

Pyxis planicauda ENDANGERED

Geochelone yniphora ENDANGERED

Dyscophus antongili VULNERABLE

Mantella aurantiaca VULNERABLE

Pyxis arachnoides VULNERABLE

Geochelone radiata VULNERABLE

Phlesuma standingi VULNERABLE

Brookesia peramata VULNERABLE

Furcifer campani VULNERABLE

Furcifer labordi VULNERABLE

Furcifer minor VULNERABLE

SUMMARY OF CRITERIA

50% population decline in 10 years or 3

< 5,000 km? extent of occurance and <

20% population decline in 10 years or 3 generations

< 20,000 km? extent of occurance, < 10 isolated populations

and decline in population

20% population decline in 10 years or 3 generations

< 20,000 km? extent of occurance, < 10 isolated populations

20% population decline in 10 years or 3 generations

and < 100 km? area of occupancy or < 5 locations

20% population decline in 10 years or 3

generations

generations,

5 isolated populations

generations

CHRISTOPHER J. RAXWORTHY AND RONALD A. NUSSBAUM

These species have been commercially traded, but the impact of

this trade on wild populations is unknown or only poorly known.

Two of these species have a very limited distribution: M.

aurantiaca is restricted to eastern rain forest with an approxi-

mate extent of occurrence of 3,000 km? and P. standingi re-

stricted to deciduous western forest in the Toliara region with

an approximate extent of occurrence of 2,500 km? (although an

unconfirmed report suggests this species may also occur fur-

ther north).

In the case of P. standingi we have found this gecko is

extremely rare (if not locally extinct) in forest within 5 km of

major roads at Ifaty (none were seen during at least 60 man-

days of searching). Commercial collecting has been occurring in

this area for some time, and in many cases collecting poles

could still can be found lying against baobab tree trunks, left

by the local collectors in the hope of catching any remaining

geckos. However, because the original. population size and

density in this region is unknown, the impact of commercial

collecting cannot be measured.

The impact of collecting on M. aurantiaca 1s also unknown.

Zimmermann and Zimmermann (1994) suggest that local popu-

lation extinctions occurred in a swamp system between 1966

and 1993 as a result of habitat loss. They therefore propose that

commercial collecting be prevented by placing M. aurantiaca on

Appendix I of CITES.

For M. aurantiaca and P. standingi, their limited known

distributions, primary habitat requirements, and commercial col-

lecting pressures, all suggest that both species may be vulner-

able to extinction.

Distribution data is used as criteria for Vulnerable status

for three species: Pyxis arachnoides, G. radiata (Plate 15), and

Brookesia perarmata (Plate 16). For the two tortoise species,

P. arachnoides and G. radiata, the extent of occurrence is actu-

ally greater than 20,000 km? (we estimate both around 30,000

km’), one of the criteria used to support the vulnerable status.

In addition, although the populations of these species are also

likely to becoming more fragmented, as a result of habitat loss,

there are without doubt many more than 10 isolated popula-

tions (another criteria used to assign these species vulnerable

status, see Table 1). Therefore, the only justification that can

be made for considering these tortoises, as Vulnerable, using

the IUCN criteria, is the rate of observed or suspected popula-

tion decline, for which we currently have very little quantita-

tive information. The large distributions of P. arachnoides and

G. radiata do not suggest that either species is yet threatened

by extinction.

B. perarmata is only known from one site: Bemaraha, in

Western Madagascar. Although the surface area of the Bemaraha

Reserve is large (152,000 ha), our survey of this protected area

at three well separated sites, yielded this chameleon at just one

locality indicating it is not continuously distributed throughout

the reserve. This clearly supports the “less than five site” crite-

ria for Vulnerable status.

Discussion

Despite the major environmental problems that have been de-

veloping for so long in Madagascar, it is perhaps surprising that

we have no evidence of any herpetofaunal extinction events

occurring on the island over the past 500 years. Although this

suggests the island’s species diversity has not declined during

this period, we believe herpetofaunal extinctions have been oc-

curring, and are occurring now, but that researchers and conser-

vationists have overlooked these events.

Species most vulnerable to extinction will have small popu-

lations, and therefore tiny relict distributions. This makes them

difficult to survey, and, therefore, easily missed. A modern ex-

ample of this is P. antanosy, distributed in a region of Madagas-

car that has been subject to herpetological collecting for over 100

years. We suspect many more herpetofaunal species are on the

verge of going extinct in transitional or rare habitat types. Re-

gions that experience highly localized climatic conditions fre-

quently have unusual transitional habitats in Madagascar. In

many cases these localized habitats have been further reduced to

small relict patches as a result of human exploitation and have

attracted little research attention because of their small size or

isolated locality. Examples of major sites we have surveyed,

which offer these conditions, include Analavelona, Isalo,

Bemaraha, Kelifely, Ambohijanahary, Namoroka, and Ankarana.

We are in the process of describing new species from all of these

sites, most of which are likely to be locally endemic, but active

conservation programs are now underway in just three sites

(Isalo, Bemaraha, and Ankarana). Many of these new species

are reduced to such small population sizes that they are obvi-

ously some of the most vulnerable herpetofaunal species in

Madagascar. Other potentially vulnerable species include

Phelsuma masohoala, Uroplatus malahelo, Pseudoxyrhopus

kely, and Alluaudina mocquardi.

Of the 15 endemic Malagasy amphibians and reptiles con-

sidered Threatened by the IUCN Red List, we consider (based

on distribution and habitat requirements) just three species to be

obviously vulnerable to extinction: P. planicauda, G. yniphora,

and B. perarmata. Another two species, M. aurantiaca and P.

standingi, appear to also have small distributions, but further

fieldwork is required to confirm this. The other 10 species do

not appear to be vulnerable to extinction because of their much

larger distributions, and in some cases, broad tolerance to human

modification of primary habitats.

In part, the IUCN Threatened List actually reflects a his-

torical bias towards conserving turtles and boas in Madagascar,

rather than actual risks to extinction. It is worth noting that 8 of

the 15 Red List species are boas or turtles (53 percent), although

this group represents just 2 percent of the island’s actual en-

demic herpetofauna. Ironically, this conservation bias is actually

hindering attempts to prevent herpetofaunal extinctions in Mada-

gascar, by promoting some species for conservation activities

that are not vulnerable and by ignoring many species that are

about to be lost.

Rates of past, present, or future population decline (ob-

served, estimated, inferred, or suspected) are used as IUCN

criteria for all but one of the 15 threatened species. This sur-

prises us, as we are unsure what types of evidence were used to

support or estimate these rates of population decline. For the

herpetofauna of Madagascar, we think that it would be far more

effective to base vulnerability to extinction on biogeographic

data, rather than estimated population declines. The advantage

is that biogeographic data is available for most species, and these

data can be compared between different taxa to determine con-

servation priorities. Of course, biogeographic data are never com-

plete for any species, but it can be obtained with far less effort

than undertaking population studies throughout a species distri-

bution range.

HERPETOFAUNAL EXTINCTION INMADAGASCAR

Table 2 includes four biogeographic criteria that we con-

sider to be the most important in determining vulnerability to

extinction for amphibians and reptiles in Madagascar. Other

population factors will also influence the risk of extinction (e.g.,

population density, generation time, reproductive output, pre-

dation, and human collecting), but since it is unlikely we will

ever have detailed information of this type over the entire distri-

butional range on any herpetofaunal species, we think the most

objective data set available to us is biogeographic.

Table 2. Measuring extinction vulnerability with biogeo-

graphic criteria.

EXTINCTION RISK

BIOGEOGRAPHIC CRITERIA

HIGH LOW

EXTENT OF OCCURANCE Small Large

NUMBER OF KNOWN SITES Few Many

DISTRIBUTION STRUCTURE Fragmented Continuous

HABITS Specialist of a

declining

habitat

Specialist of a

stable habitat

Provided that taxonomic biases are avoided, we believe that

the analysis of biogeographic data has the potential to reveal a

much more realistic picture of extinction threats to the amphib-

ians and reptiles in Madagascar, compared to the criteria cur-

rently being employed. This will identify those species of great-

est concern, which could then become the focus of conservation

programs aimed at maintaining existing levels of biodiversity in

Madagascar.

Acknowledgments.—This research was made possible with

the help of J.-B. Ramanamanyjato, A. Raselimanana, Angelin and

Angeluc Razafimanantsoa, and the cooperation of the Ministére

de |’Enseignement Supérieur, the Ministére de la Production

Animale et des Eaux et Foréts, and the Ministére de la Research

Scientifique et Technologie pour le Developpement. This re-

search was funded in part by grants from the National Science

Foundation (DEB 90 24505, 93 22600), the National Geographic

Society, and Earthwatch.

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Manuscript received: 05-November-1997

Accepted: 22-December-1997

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

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Amphibian and Reptile Conservation 2(1):24-29.

New data on the distribution, status, and biology of the

New Caledonian giant geckos (Squamata:

Diplodactylidae: Rhacodactylus spp.)

AARON M. BAUER’ AND ROSS A. SADLIER?

‘Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085-1669, USA ?Department of

Herpetology, Australian Museum, 6-8 College Street, Sydney, New South Wales 2000, AUSTRALIA

Abstract.—Recent collections and observations of the New Caledonian giant geckos (Rhacodactylus) result in range

extensions and new information regarding the biology of these lizards. Significant range extensions are reported for the

rough-snouted giant gecko (R. trachyrhynchus) and for the recently rediscovered Guichenot’s giant gecko (R. ciliatus). Field

observations confirm the association of the knob-headed giant gecko (R. auriculatus) with plants of the family Cunoniaceae

and that Leach’s giant gecko (R. leachianus) feeds on fruit. Range extension of some species, and data on local abundance

allows a reassessment of their conservation status. Despite implied increases in giant gecko density and range, significant

threats from habitat loss, introduced predators, and illegal trade leave all species at risk.

Key words. Rhacodactylus, geckos, New Caledonia, distribution, diet, conservation status

Introduction

Rhacodactylus is one of three carphodactyline* gecko genera

occurring in New Caledonia. The genus includes the largest liv-

ing species of gecko, R. leachianus (> 250 mm snout-vent length;

Russell and Bauer 1986) and has attracted scientific attention

because of unusual characteristics such as viviparity! (in R.

trachyrhynchus; Bartmann and Minuth 1979), the possession

of and specialized dentition (in R. auriculatus; Bauer and Russell

1990; Bauer and Sadlier 1994b), and prehensile tails (all species;

Bauer 1990; Bauer and Russell 1994). Species of this genus have

also attracted much popular attention, especially among ter-

rarium keepers, because most species thrive and reproduce well

in captivity (Henkel 1987, 1991, 1993; Henkel and Schmidt

1991; Tytle 1992).

The systematics and morphology of Rhacodactylus have

recently been the focus of significant investigation (Bauer 1990;

Bauer and Russell 1990; Bauer et al. 1993; Seipp and Klemmer

1994: Good et al. 1997). However, knowledge of the distribu-

tion and biology of these geckos remains largely incomplete

(Bauer and Sadlier 1993). Aside from brief reports of various

aspects of natural history (e.g., Mertens 1964; Meier 1979;

Sameit 1985; Bauer 1990; Bauer and Vindum 1990; Henkel 1991),

field-based data are limited to a few investigations of diet (Bauer

and DeVaney 1987; Bauer and Sadlier 1994b) and general ac-

counts of behavior and ecology in nontechnical works (de Vosjoli

1995; de Vosjoli and Fast 1995). However, increasing awareness

of the uniqueness of the flora and terrestrial fauna of New

Caledonia (Myers 1988; Mittermeier et al. 1996) has given new

impetus to the collection of basic distributional and biological

data for all members of the New Caledonian herpetofauna, and

previously understudied areas are being surveyed more system-

atically (e.g., Isle of Pines, Bauer and Sadlier 1994a). Observa-

tions made by the authors during several recent expeditions to

‘Correspondence. Tel: (6/0) 519-4857; fax: (610) 519-7863;

email: aaron.bauer@ villanova.edu

New Caledonia have yielded new distributional and/or dietary

information for all six of the recognized species of Rhacodactylus.

In addition, preliminary assessments of genetic variation within

certain species (reported more fully in Good et al. 1997) were

made on the basis of tissue samples accumulated over a series of

trips since the mid-1980’s. We present these data here as they

help to provide a more accurate picture of the geographic ranges

and biological requirements of the geckos and may be useful in

establishing the conservation status of Rhacodactylus species.

Materials and methods

Herpetological collections and observations were made on main-

land New Caledonia during trips in 1994 and 1995. Specimens

were collected under a series of permits issued by the conserva-

tion authorities of the Province Sud (Marcel Boulet) and Prov-

ince Nord (Christian Papineau) of New Caledonia to the au-

thors. Preliminary estimates of genetic divergence between popu-

lations were based on results derived from allozymicé data. De-

tails of the electrophoretic methodology employed are presented

in Good et al. 1997. Specimens cited are housed in the collec-

tions of the Australian Museum (Sydney)-AMS, the Natural

History Museum (London)-BMNH, and the California Acad-

emy of Sciences (San Francisco)-CAS.

Results and discussion

Knob-headed giant gecko (Rhacodactylus auriculatus)

[Plate 1]. B6hme and Henkel (1985) reported a striped color

phase of R. auriculatus, now known to be common. Although

polymorphisms were noted within a single population of this

species, there were no fixed differences among population samples

from four different localities and no suggestion of significant

intraspecific genetic variation. This tends to corroborate mor-

phological observations that this species is generally

polymorphic" throughout its range but that there are no geo-

graphically related trends in character variation (Bauer 1990).

Plate 1

Plate 5A

etal

Plate 5B Plate 6

Plate captions: 1. Knob-headed giant gecko, Rhacodactylus auriculatus, from Riviére Bleue. 2. Bavay’s giant gecko Rhacodactylus

chahoua. Photo courtesy of R. D. Bartlett. 3A. Guichenot’s giant gecko, Rhacodactylus ciliatus, from Riviere Bleue (adult with autoto-

mized tail). 3B. Rhacodactylus ciliatus, from Riviere Bleue (subadult with complete original tail and complex body patterning). 4.

Leach’s giant gecko, Rhacodactylus leachianus, from Mt. Koghis. 5A. Roux’s giant gecko, Rhacodactylus sarasinorum, from Kwa

Néie (adult retaining bold white dorsal markings). 5B. Rhacodactylus sarasinorum, from Riviére Bleue (adult with mottled dorsal

pattern). 6. Rough-snouted giant gecko Rhacodactylus trachyrhynchus, from Mt. Aoupinié. Photos 1, 3A and B, 4, 5A and B, and 6: Ross

A. Sadlier.

AARON M. BAUER AND ROSS A. SADLIER

Plate a Geissois sp. from Mt. Do, southern New Caledonia.

This plant is apparently utilized as a food source by Rhaco-

dactylus auriculatus.

Bavay (1869) first reported on the diet of this species,

indicating that it eats the flowers of Geissois (Cunoniaceae).

This was verified by the recovery of anthers, stamens, and (pos-

sibly) pollen belonging to either a member of this family (or the

Myrtaceae) from the stomach of one specimen (Bauer and Sadlier

1994b). On 9 January 1995 further evidence of the specific

association between R. auriculatus and Geissois was obtained

when geckos were found active on flowering specimens of

Geissois spp. (Plate 7) 1.3-2.6 km from the summit of Mount

(Mt.) Do (21°45'S, 166°00' E) in south central New Caledonia.

Bavay’s giant gecko (Rhacodactylus chahoua) [Plate

2]. The known distribution of this species in central and south-

ern mainland New Caledonia has been expanded by the capture

of a specimen from Sarraméa (AMS R144171) and by speci-

mens from unstated localities on the Isle of Pines (de Vosjoli

1995; de Vosjoli and Fast 1995) [Fig. 1]. This species is rather

polymorphic with respect to coloration (Bavay 1869; Béhme

and Henkel 1985; Bauer 1985), but the comparison of allozymes

from individuals separated by more than 100 km suggests rela-

tive genetic uniformity.

Guichenot’s giant gecko (Rhacodactylus ciliatus) [Plate

3A and 3B]. This species was numerous for the first 20 years

after its description (e.g., Bavay 1869) and then was not seen

again for over 100 years, despite extensive searches by several

researchers. It was regarded as extinct (see Bauer and Sadlier

1993). In 1994, the species was rediscovered and has since been

found at a variety of localities on the Isle of Pines (Seipp and

Klemmer 1994; Kullmann 1995) and several smaller offshore

islands (de Vosjoli 1995). De Vosjoli (1995) and de Vosjoli and

Fast (1995) recorded this species as common on the Isle of Pines

but stated that it was not present on the mainland of New

Caledonia. Despite their claims, Bavay (1869), whose data have

proved to be very accurate (see Bauer and Sadlier 1994b), re-

ported collecting seven specimens of this species at several (un-

specified) localities on the mainland, and the type locality given

by Guichenot (1866) was at Canala (21°35'S, 165°56' E), also

on the mainland. The persistence of this species on the mainland

was verified by two specimens (AMS 146594-5) collected by

R. A. Sadlier near Pont German, Riviere Bleue (22°06' S, 166°38'

E) in the extreme south of New Caledonia (Fig. 1). It has subse-

quently been taken at other localities on the mainland (Girard

and Heuclin 1998; Bauer and Sadlier 2000), suggesting that it

may be relatively widespread.

Leach’s giant gecko (Rhacodactylus leachianus) [Plate

4]. This species has a broad distribution in the wetter areas of

the New Caledonian mainland, especially along the east coast

(Bauer 1990). Boulenger (1885) first recorded the species from

the Isle of Pines (BMNH 53.8.16.13). Bauer and Sadlier (1994a)

confirmed the presence of this species on the island with a 194

mm female (CAS 182197). Subsequently, the species has been

recorded as fairly common on the Isle of Pines and nearby off-

shore islands (de Vosjoli 1995; de Vosjoli and Fast 1995). Al-

though most known mainland New Caledonian localities are in

low- to middle-elevation forests (Bauer 1990), specimens have

been recorded from up to 1100 m (Mertens 1964). Sight obser-

vations in January 1995 at 540 m on Mt. Mandjélia (20°24’ 15"

S, 164°31°18" E) extend the confirmed distribution of the spe-

cies to the northwest, almost to the limit of the humid forest on

the main island of New Caledonia.

The Isle of Pines population has recently been described

by Seipp and Obst (1994) as a distinctive subspecies,

Rhacodactylus leachianus henkeli. The validity of this form is

challenged on the basis of morphological and allozyme charac-

ters by Good et al. 1997. They found the henkeli color pattern

to occur among geckos in at least two regions of the New

Caledonian mainland and regarded behavioral differences as at-

tributable to reduced predation pressure on the insular form.

Because genetic distance data’ indicated no long separation of

Isle of Pines R. /eachianus from mainland populations Good et

al. 1997 regarded the split of the insular population to be very

recent. Indeed sea level minima of 100 m or more would have

connected New Caledonia to the Isle of Pines as recently as

16,000-20,000 years ago (Stevens 1973; Holloway 1979).

Although the diet in captivity of Rhacodactylus leachianus

has been well documented (Mertens 1964; Bauer and De Vaney

1987; Henkel and Schmidt 1991), and a few stomach contents

have been reported (Roux 1913), the natural diet remains poorly

documented. At Mt. Aoupinié, in January 1995, we observed

individuals feeding on fruit in humid forest trees. Examination of

feces of freshly captured individuals revealed only fig seeds and

partially digested fig fruit. It appears likely that this, and per-

haps other Rhacodactylus species, take advantage of seasonal

and local availability of figs and may play a role in seed dis-

persal.

Roux’s giant gecko (Rhacodactylus sarasinorum)

[Plate 5A and SB]. Bauer (1990) figured the type of R. sarasinorum

NEW CALEDONIA

| New Caledonia is a French overseas territory, consisting of the large island of New Caledonia and the Loyalty Islands. Its location is approximately 1,200

| km east of Australia (Geographic Coordinates 21°30’ S, 165°30’ E) in the South Pacific Ocean. These islands have an extraordinary diversity of fauna and —

| flora with an extreme level of endemism in many taxa including birds and reptiles. Naturally occurring plant species number 3,380 (vascular plants), birds

116, mammals 9, and reptiles 87 (71 terrestrial and 16 marine). No naturally occurring amphibian species exist on New Caledonia though a nonnative

species has been introduced from Australia (green and golden bell frog Litoria aurea). Total area is 19,060 square (sq) km (land 18,575 sq km and water

485 sq km) comparatively, slightly smaller than New Jersey. The terrain is west coastal plains with interior mountains (highest point Mont Panie 1,628

m) making up two-thirds of the island. The climate is subtropical (warm and humid) modified by southeast trade winds. There is little temperature change

throughout the year, averaging between 71°F and 75°F (22°C and 24°C). The natural vegetation comprises tropical evergreen rain forest up to 1,000 m and

' tropical montane rain forest above 1,000 m. Mangroves occur along western coasts. The major vegetation types are dense evergreen forest (22.8% of total

land area), Niaouli savanna woodland (13.8%), maquis vegetation in mining areas (25.1%), savanna grassland (21.7%), and scrub (8.3%). New Caledonia’s

human population numbers 191,003 (July 1997 estimate) with a 1.68% (1997 estimate) annual growth rate. New Caledonia’s moderately developed

economy is based on mining and has more than 20% of the world’s known nickel resources as well as other natural resources as chrome, iron, cobalt,

manganese, silver, gold, lead, and copper (thus mining is an important environmental issue). Only a negligible amount of the land is suitable for cultivation

and food accounts for about 25% of imports. In addition to nickel, financial support from France and tourism are key to the health of the economy. The

principal threats to the natural flora and fauna are mining, logging, and bushfires, reducing the forest cover from an estimated 90% cover to just 20%.'

and noted variation in color pattern and body pro-

portions in this species but did not elaborate. At

least two color morphs have been illustrated and

described by Henkel (B6hme and Henkel 1985;

Henkel 1987, 1988), but there has been no sug-

gestion of subspecific or specific distinction be-

tween these forms. Bauer’s (1990) and Bauer and

Vindum’s (1990) concept of R. sarasinorum was

based in part on typical specimens and in part on

an individual from Touaourou that is larger, darker,

and differs from other specimens in a number of

scale counts. Allozyme analysis (Good et al. 1997)

revealed that this specimen differed from a typi-_

cal R. sarasinorum from Riviere Bleue (AMS R

146596) by four fixed differences. This is a greater

genetic difference than that between R. ciliatus

and R. chahoua. Both allozyme and morphologi-

cal data thus suggest significant variation and are

Fig. 1. Distribution of Rhacodactylus ciliatus ie

being analyzed separately, which may result in (triangles). R. chahoua (squares), and R

h “4: f Rh il i trachyrhynchus (circles) in the New Caledonia

the recognition of a new Khacodactylus sara- region. Open symbols represent older records

i -li i as summarized by Bauer and Henle (1994), new

sinorum-like SPStles. records are indicated by solid symbols. Note

especially the range extension of R. trachyrhny-

chus to the west coast at Pindai and the docu-

Rough-snouted giant gecko (Rhacodacty- mentation of the occurence of the other species

lus trachyrhynchus) [Plate 6]. Bauer (1990) on the Isle of Pines.

recorded five mainland New Caledonian localities

for R. trachyrhynchus. Several of these, Coula-

Borearé, Ciu, and Mt. Gouemba are in the eastern humid forest scincus festivus), New Caledonian skink (Caledoniscincus

region of the island. The other two localities, La Foa and near austrocaledonicus), and a new species of elf skink (Nannoscincus

Nouméa are imprecise but probably are also humid forest locali- sp.). All previous records of Rhacodactylus spp. from the New

ties. All localities are at middle to low elevation. As briefly Caledonian mainland have originated from the wetter eastern

noted by Bauer (1995), recent censuses have expanded the portions of the island, or from rainforest or maquis vegetation‘

known range of the species, both elevationally and geographi- in the south (Bauer 1990; Henkel 1991, 1993; Bauer and Henle

cally (Fig. 1). Five specimens (AMS R146417-9, CAS 200266- 1994). The occurrence of R. trachyrhynchus at this site is thus

8) were found during rainstorms in humid forest (Fig. 2A) at intriguing and suggests a much broader habitat tolerance range

approximately 520 m on Mt. Aoupinié in Central New Caledonia than previously suspected for this species. Although normally

(21°09719" S, 165°19712" E), 27 km north and west of the associated with large, mature rainforest trees (Meier 1979), at

previously documented range of the species. A single specimen, Pindai, this gecko was collected less than five meters from the

CAS 200269, was obtained in sclerophyll forest’ at Pindai ground in the branches of a small tree. A very low genetic dis-

(21°20’02" S, 164°58’21" E) at approximately 20 m elevation tance from this specimen to specimens from Mt. Gouemba and

(Fig. 2B). This locality is also somewhat further north than Mt. Aoupinié (Good et al. 1997) suggests no significant differ-

earlier records, but it is unique in that it is a west coastal locality entiation in the dry forest population, and the specimen is typi-

in an area of low rainfall. The local vegetation is dominated by cal for the species in regard to morphology. Henkel (1991) sug-

largely endemic dry forest plants and is regarded as the most gested that there are two morphs in the species, one with a

threatened terrestrial habitat in New Caledonia (Jafrré et al. 1993; short, wide, robust snout, the other less so, but because these

Bouchet et al. 1995). The only other reptiles collected sympat- features were noted in captive born specimens of uncertain lo-

rically with R. trachyrhynchus at Pindai were Vieillard’s prehen- cality he did not imply any subspecific distinction. Our data,

sile-tailed gecko (Eurydactylodes vieillardi), Giinther’s New from three widely scattered localities, do not support the recog-

Caledonian gecko (Bavayia cyclura), sclerophyll forest gecko nition of any specific or subspecific subdivisions within R.

(Bavayia exsuccida), festive New Caledonian skink (Caledoni- trachyrhynchus.

AARON M. BAUER AND ROSS A. SADLIER

Fig. 2. Habitat of Rhacodactylus trachyrhynchus.

Caledonia.

The presence of this species on the Isle of Pines remains

problematic. Boulenger (1878) recorded the species (as

Chamaeleonurus trachycephalus) from this locality. Subsequent

collecting activity on this island, however (Bauer and Sadlier

1994a; de Vosjoli 1995) has not verified this occurrence. The

unexpected findings of this and other Rhacodactylus after many

years of extensive and intensive searching, however, argue against

dismissal of this early record. Bauer and Sadlier (1994a) identi-

fied appropriate habitat for the species on the island.

Conclusions

The new dietary observations confirm earlier reports of her-

bivory by Rhacodactylus species. It is especially noteworthy

that R. auriculatus was found active on flowering heads of plants

of the same genus on which Bavay observed them over 130

years ago. The significance of reptiles as seed dispersers and as

possible pollinators was recognized by Borzi (1911) but re-

mains largely unexplored. Its recent documentation for the closely

related carphodactyline geckos of New Zealand (Whitaker 1987)

suggests that at least some species of giant geckos are important

in this regard. The importance of plant material in the diets of

Rhacodactylus sp. and the significance of geckos as dispersers of

pollen or seeds, however, can only be adequately addressed by

a seasonal dietary study at a single site.

Bauer and Sadlier (1993) reviewed the conservation status

of all New Caledonian lizards on the basis of data then available.

They summarized both the extent of the geographic range of the

species and their apparent abundance within the appropriate

B. Sclerophyll forest habitat at Pindai on the west central coast

A. Overview of humid forest habitat at middle elevation on Mt. Aoupinié.

of New

habitat types. The data reported on here necessi-

tate a re-evaluation of that status report.

Rhacodactylus ciliatus, previously considered

possibly extinct, is now known to be common on

islands of the south coast of New Caledonia and

present, if somewhat less common, on the main-

land. Using the terminology of Bauer and Sadlier

(1993), its distribution is now regarded as re-

stricted and its status as locally common. With

the extension of its known range to the north and

to the west coast sclerophyll forest, the distribu-

tion of Rhacodactylus trachyrhynchus can now

be upgraded from restricted to moderately wide-

spread and its status to locally common. The pres-

ence of this species, as well as a regionally en-

demic Bavayia (B. exsuccida) and a new, appar-

ently endemic, Nannoscincus in the sclerophyll

forest in west coastal New Caldeonia (Bauer et al.

1998) adds impetus to efforts to protect the small

remaining tracts of this habitat. Rhacodactylus

leachianus was previously regarded as widespread

and uncommon. The extension of the east coast

range to the limit of humid forest underscores the

fact that this is the most widely distributed of all

Rhacodactylus species and observations of indi-

viduals under ideal weather conditions (warm and

wet) suggest that the species may best be catego-

rized as common in appropriate habitats. The sta-

tus of the remaining species is unchanged by the

new records.

Although at least several species of

Rhacodactylus do appear to be locally abundant,

even the most widely ranging species is endemic

to the New Caledonian mainland and adjacent sat-

ellite islands, an area about the size of Connecticut. Further, no

species is known to be present in all native habitat types and all

are excluded from agricultural or urban environments (although

they may be present at the periphery of human-modified areas).

Habitat destruction and the impact of introduced predators were

cited as the primary threats to the herpetofauna of New Caledonia

by Bauer and Sadlier in 1993. These factors remain the most

significant conservation concerns, but the illegal pet trade in

Rhacodactylus has increased significantly in the 1990’s and has

become a potential threat to wild populations.

AARON M. BAUER

Acknowledgments.—We thank the permit-issuing authori-

ties in New Caledonia for their continued support of our re-

search. We also thank Jean Chazeau (IRD) and Alain Renevier

and his family for their encouragement of our activities. Finan-

cial support for this work came from Villanova University, the

California Academy of Sciences (AMB), and the Australian

Museum (RAS).

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zucht. Eugen Ulmer, Stuttgart, Germany. 224 p.

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Jaffré, T., Morat, P., and Veillon, J. -M. 1993. Etude floristique et

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Der Zoologischer Garten, N. F. 29:49-47.

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a conservation imperative for an ancient land. Oryx 30:104-

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Russell, A. P. and Bauer, A. M. 1986. Le gecko géant Hoplodactylus

delcourti et ses relations avec le gigantisme et 1’endémisme

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Zeitschrift 38:279-281.

Seipp, R. and Klemmer, K. 1994. Wiederentdeckung von

Rhacodactylus ciliatus Guichenot 1866 im Stiden Neukale-

doniens (Reptilia: Sauria: Gekkonidae). Senckenbergiana

Biologica 24:199-204.

Seipp, R. and Obst, F. J. 1994. Beschreibung einer neuen Unterart des

neukaledonischen Rhacodactylus leachianus Cuvier 1829.

Senckenbergiana Biologica 74:205-211.

Stevens, G. R. 1973. The palaeogeographic history of New Zealand.

New Zealand Entomologist 5:230-239.

Tytle, T. 1992. The natural history and captive husbandry of the

New Caledonian lizard genus Rhacodactylus. The Vivarium

3(6):32-35.

Whitaker, A. H. 1987. The roles of lizards in New Zealand plant

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328.

Footnote

"The research sources used in writing the country sidebar.on page 10 (Madagascar), of this

volume, and footnoted on page 14 (foomote *), were also used in writing the New Caledonia

summary.

Manuscript received: January-1997

Accepted: April-1998

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

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

Amphibian and Reptile Conservation 2(1):30-31.

Column

The United States role in the international live reptile trade

CRAIG M. HOOVER"

TRAFFIC North America, World Wildlife Fund (WWF), 1250 24th Street, NW, Washington, D.C. 20037-1132, USA

Abstract.—In the 1990’s, the trade in live reptiles has grown substantially, and the United States (U.S.) is the world’s

most significant player in the international trade in live reptiles, both as an importer of exotic species, and as an exporter

of native and exotic species. In 1995, more than 2.5 million reptiles were imported into the U.S., primarily to supply the pet

trade. In 1996, over 9.5 million reptiles were exported or reexported from the U.S., primarily to Europe and Asia, to supply

the demand for reptiles as pets and food. Despite the large and apparertly growing number of reptiles and amphibians in

trade, we have yet to quantify the impacts of this trade on the conservation of these species in the wild.

Key words. Herpetofauna, import, export, reexport, live reptile, international trade

By way of introduction, I would like to give you some

background information about myself, TRAFFIC, and

this new column in Amphibian and Reptile Conserva-

tion titled, Herpetofauna and Humanity. Presently, ama

Senior Program Officer for TRAFFIC North America. As

a Senior Program Officer, I am responsible for the devel-

opment and oversight of wildlife trade studies and the

implementation of their findings and recommendations.

TRAFFIC North America is a part of the worldwide TRAF-

FIC Network, a program of World Wildlife Fund (WWF)

and The World Conservation Union (IUCN). TRAFFIC

is the world’s largest

wildlife trade monitor-

ing program with 21

offices covering most

regions of the globe.

TRAFFIC produces

reports and papers

documenting the find-

ings of its studies and

recommending mea-

sures necessary to

help ensure that the

trade in wildlife and

wildlife products are

conducted in a sus-

tainable and legal

manner. This work is

done by collecting

trade data via govern-

ment agencies such as the United States Fish and Wild-

life Service (USFWS), customs agencies, and interna-

tional bodies, carrying out market surveys, conducting

literature reviews and website searches, and other means.

In the three years that I have been with TRAFFIC, much

of my time has been devoted to investigating various

aspects of reptile and amphibian trade.

Prior to coming to TRAFFIC North America, I was a

wildlife inspector for the USFWS in Los Angeles, where

for over four years I was able to see firsthand the scope of

the trade in reptiles and amphibians. Equally important, I

gained valuable knowledge of the laws that govern the

trade, and the means by which these laws are implemented

and enforced in the United States (U.S.).

Ball python (Python regius). Photo

‘Correspondence. Tel: (202) 822-3452; fax: (202) 775-

8287; email: craig.hoover@wwfus.org

. World Wildlife Fund.

This background, along with an education in natural

resources and law, will influence the areas to be covered

in this column. Among the subjects I intend to tackle in

this space are reptile and amphibian trade and its implica-

tions for conservation; the use of reptiles and amphibians

as clothing, food and medicine; the enactment, implemen-

tation and effectiveness of wildlife trade laws; illegal trade

and the threat posed to reptiles and amphibians in the

wild; captive breeding and the private breeder’s role in

conservation; and the current events that shape the rela-

tionship between herpetofauna and humankind. As an

introduction to this new col-

umn, Herpetofauna and Hu-

manity, it would be useful to

provide an overview of the

U.S. role in the international

live reptile trade. The follow-

ing information summarizes

a report released by TRAF-

FIC in August, 1998, entitled

The U.S. Role in the Inter-

national Live Reptile Trade:

Amazon tree boas to Zulu-

land dwarf chameleons

(Hoover 1998).

The international trade

in live reptiles has grown

dramatically in the last de-

cade. The import, export, and

reexport of live reptiles sup-

plies a number of markets, including zoos and aquariums,

breeding facilities, research centers, private breeders and

keepers, and even food markets in some segments of soci-

ety. By far the most significant market for the live reptile

trade is the pet market (private breeders and keepers of

amphibians and reptiles).

The causes of the substantial rise in the international

trade in live reptiles are difficult to quantify but may include

an increase in the availability and variety of species; im-

proved reptile husbandry practices due to advances in tech-

nology and scientific knowledge; increased restrictions on

other wildlife trade; changing lifestyles that make reptiles

more suitable pets than other fauna; or simply an increased

popularity that has made reptiles today’s fashionable pets.

Whatever the reason, and there may be some truth to all of

these explanations, there can be no denying that the live

reptile industry has expanded dramatically.

CRAIG M. HOOVER

In 1970, prior to the passage of laws such as the U.S.

Endangered Species Act and adoption of the Convention

on International Trade in Endangered Species of Wild

Fauna and Flora (CITES), U.S. imports of live reptiles ap-

proached 2 million animals. Nearly 80 percent of this im-

port volume consisted of turtles, primarily red-eared slider

turtles (Jrachemys scripta elegans), with 12 percent liz-

ards, 6.5 percent crocodilians, and less than 2 percent

snakes (Busack 1974).

Based on analysis of USFWS trade data for a number

of reptile species, it is clear that the trade subsequently

declined significantly and remained relatively low through-

out the 1980s. However, imports increased again in the

early 1990s, and in 1995 more than 2.5 million live reptiles

were imported to the U.S. Yet the content of this trade

differed significantly from the reptiles imported in 1970.

By far the most commonly imported species was the green

iguana ([guana iguana), which made up more than 45

percent of the total trade in 1995, and only 8 percent of

imports in 1970. Snakes and lizards played a far more sig-

nificant role in current import levels than they did in 1970,

with 1995 turtle volumes greatly reduced and crocodilian

imports virtually nonexistent. In fact, the dramatic increase

in the total number of live reptiles imported is primarily

due to fluctuations in the number of iguanas imported.

However, the U.S. is not only a consumer in the

international live reptile trade, but also a significant sup-

plier. In fact, the U.S. presently exports or reexports more

live reptiles than it imports, due largely to the export of

farm-raised hatchling red-eared slider turtles, a species

even more influential on overall trade volumes than the

iguana. For example, in 1996, the U.S. exported or reex-

ported over 9.5 million reptiles, primarily to Europe, East

and Southeast Asia, yet over 88 percent of this trade

consisted of the red-eared slider, at a volume of nearly

8.4 million animals.

Setting aside the voluminous trade in red-eared slid-

ers, there are more than one million reptiles of other spe-

cies that are exported or reexported from the U.S. The

North American taxa nearest to the red-eared slider in ex-

port volume are the map turtles (Graptemys spp.). Ac-

cording to USFWS data, map turtle exports have risen

from less than 10,000 in 1990 to over 80,000 in both 1995

and 1996. Based on map turtle export data, just two of the

twelve map turtle species make up 90 to 95 percent of the

export trade: the common map turtle (G. geographica)

and the false map turtle (G. pseudogeographica) [Ventura

1997; data provided by Weissgold 1997].

The U.S. also plays a substantial and apparently ex-

panding role as an exporter of previously imported rep-

tiles. For example, the U.S. reexported less than 60,000

iguanas in 1993 and more than 270,000 imported iguanas

in 1996. This role as a supplier of previously imported

reptiles is influenced by a number of factors. Perhaps fore-

most is geography; for instance, U.S. dealers are well po-

sitioned to supply Asia and Europe with Latin American

reptiles. Another significant factor may be that U.S. deal-

ers have long-established connections with overseas sup-

pliers that may provide a competitive advantage as new

markets arise. For example, most of the live reptile trade in

Canada, where the trade in reptiles is less established,

appears to be supplied by U.S. traders (Chatel 1998).

US. trade in live reptiles also appears to make up a

substantial portion of the world trade in live reptiles. A

comparison of trade data for certain CITES-listed species

indicates that U.S. trade constituted approximately 28 per-

cent of the total world trade in 1983, but constituted more

than 82 percent of the world trade by 1992. However, these

numbers may be artificially high given the failure of many

countries to accurately report such trade.

There appears to have been an increase in illegal as

well as legal trade. Based on a review of press releases,

wildlife trade journals, and other sources, from 1970 to

1990 there were only 11 reported investigations of inter-

national live reptile smuggling, while from 1991 to 1997

there were at least 23 such cases reported. However, there

are a number of alternative explanations to these results,

including increased enforcement effort and better report-

ing of prosecutions.

Of course, all of this information on the growth in the

reptile trade raises the all-important “so what” question.

The overview study that TRAFFIC conducted was not

meant to answer that question, but to identify areas that

needed further examination so that, in some respects, the

“so what” question could be addressed. That, in large

part, will be the role of this column as well. In future is-

sues, we will look at several “so what” questions, such as:

What impact does the pet trade have on wild popula-

tions of reptiles and amphibians?

What is being done to monitor and protect native

species found in trade?

What other forms of reptile and amphibian trade may

be threatening species around the world, such as the skin,

food, and medicine trades?

What is the impact of commercial captive breeding,

farming, and ranching operations for iguanas, turtles, ball

pythons, and other species on wild populations of rep-

tiles and amphibians?

What is the threat posed by exotic species introduc-

tions that can occur with international trade, such as the

red-eared slider turtle in Europe?

What is being done to more effectively enforce exist-

ing laws and regulations to ensure that trade is not detri-

mentally affecting wild reptiles and amphibians?

The U.S. is clearly the world’s largest consumer of

live reptiles for the pet trade, as well as a significant sup-

plier and intermediary. Yet, this is only one piece of the

puzzle. The growth in popularity of reptiles and amphib-

ians as pets, along with the continued enormous demand

for these species for skins, meat, and medicine, present us

with a broad array of subjects for discussion and debate.

In coming issues, we will explore these fascinating topics

in an effort to gain a better understanding of the dynamic

and complex relationship between herpetofauna and hu-

manity.

References

Chatel, K. W. 1998 (June). Personal communication to Craig

Hoover, TRAFFIC North America, Washington, D.C.

Busack, S. D. 1974. Amphibians and reptiles imported into the

United States. U.S. Department of Interior, Washington,

D.C. 36 p.

Hoover, C. M. 1998. The U.S. Role in the International Live

Reptile Trade: Amazon tree boas to Zululand dwarf chame-

leons. TRAFFIC North America, Washington, D.C. 59 p.

Ventura, J. 1997 (May). Personal communication to Craig

Hoover, TRAFFIC USA, Washington, D.C.

Weissgold, B. 1997 (May). Personal communication to Craig

Hoover, TRAFFIC USA, Washington, D.C.

photocopying for mntemal or personal use provided the appropriate fee is paic d

Copvrizht Clearance Center, Tae., 222 Rosewood Dr. Danvers. MA 01923-4

(978) 750-8400; fax: (978) 7302 4470: ne info ruiteht co; website: wit)

BRIAN A. MAU RER*

Michigan 48824-1222, US.

versity Pres

Key words. Lizards, Au

islands, history, foraging, coexistence,

for ecologists,

surmountable.

s of study by

venus Anolis,

th its ability to make ae ree ane its choice

of prey items, can maximize its rate of

energy intake when constrained by food

abundances, oe and other 1m-

portant aspects of lizard biology. The abil-

ity of lizards to forage profitably is re=

lated tortheir fitness, which in tum, can

cies of lizards that use similar resources.

Using empirically based estimates of for-

aging rates Roughgarden shows that it is

not unreasonable to assume that much of

lizard diversity, especially on small is-

lands, is a consequence of constraints on

foraging of different-sized species as they

mteract together. Roughgarden sketches

out a reasonable picture of how these in-

teractions vary spatially from one island

to the next, and how one might begin to make predictions of

which species should be found together on which islands. Coy-

ering the literature on pairs of lizard species found together in

different combinations on different islands, Roughgarden builds

a convincing case that resource-based competition between dif-

ferent species is likely to explain many aspects of Anolis ecol-

ogy in the Caribbean.

But there is much more to the story, as Roughgarden goes

on to show in later chapters. Lizards evolve at relatively slow

rates, and Roughgarden shows how certain aspects of the his-

tory of the Caribbean basin can be traced by examining the

relationships among different species of lizards within the Car-

ibbean dwelling species. Many aspects of the distribution of

Anolis species among islands cannot be explained by rare coloni-

zation events, but must bea consequence of the geological his-

qeyeee tropidonotus

male), Las Cuevas, Cayo

istrict, Belize (April

4998). Photo hy Peter

Stafford, courtesy of The

Natural History Museum,

London.

‘Correspondence. Tel: (517) 353- Sethe Jax: (S17) 432-

1699; email: maurerb@msu.edu ~~

_ Depariment of isheries and Wildlife, Michigan State University, East Lansing,

Jonathan. 1995. Anolis Lizards of the Carib-

evolution, and plate tectonics. Oxford Uni-

ew York. 200 p. + xvi. (ISBN: 0-19

be related to the presence of other spe- cable. Melding together the two, how-

ever, yields deep insights in to the nature

Oxford Series in Ecology and Evolution

Anolis Lizards

of the

Caribbean

Ecology, Evolution, and

Plate Tectonics

Jonathan Roughgarden

-506731-2).

field experiments that a elucidate cer-

tai aspects of Anolis ecology, the real

meat of the story lies in weaving together

evidence from a wide variety of studies.

So ecology without history is incomplete,

while history without ecology ts inexpli-

of species assemblages that cannot be had

from investigations of limited scope. Her- ~

petologists and ecologists alike have much

to gain from examining Roughgarden’s

broad, sweeping approach.

Norops uniformis (male),

eux Bani, upper Ras-

ulo, Cayo District,

lize (A et, 1997). Photo

by Peter Stafford, courtesy

of The Natural History Mu-

seum, London.

Further suggested readings:

Crother, Brian I. (editor). 1999. Carib-

bean Amphibians and Reptiles. Academic Press, San Diego,

California. 495 p. (ISBN: 0-12-197955-5).

Powell, R. and Henderson, Robert W. (editors). 1996. Contribu-

tions to West Indian Herpetology: a tribute to Albert Schwartz.

contributions to herpetology, volume 12. Society for the

Study of Amphibians and Reptiles, Ithaca, New York. 457 p.

(ISBN: 0-916984-37-0).

Rivero, Juan A. 1976. Los Anfibios y Reptiles de Puerto Rico.

University of Puetto Rico Press. (ISBN: 0-8477-2317-8).

Schettino, Lourdes Rodriguez. (editor). 1999. The Iguanid Lizards

of Cuba. University Press of Florida, Gainesville, Florida. 384

p: (ISBN: 0-8130-1647-9). Availability: see inside cover ad.

Schwartz, Albert and Henderson, Robert W. 1991. Amphibians and

Reptiles of the West Indies: descriptions, distributions, and natu-

ral history. University Press of Florida, Gainesville, Florida. 736

p. (ISBN: 0-8130-1049-7), Availability: see inside cover ad.

Background screen photograph: Anolis erermanni (male), Ciales, Puerto Rico”

(24-August-1999). Photo courtesy of Father Alejandro J. Sanchez Munoz, Pastor

of Saint Anne’ Parish, Saint Thomas, U.S, Virgin Islands.

SUBMITTING BOOKS FOR REVIEW. Please send one deskcopy to: Book Keres Daphibion

and Reptile Conservation, 2525 Iowa Avenue, Modesto, California ee 467, ee

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

: News and Notes

WORLD NEWS

Amphibian declines: unraveling the

mystery. The apparent mysterious

declines of amphibian populations in

protected or relatively undisturbed areas

was discussed recently by university

scientists, government biologists, federal

administrators and representatives from

non-government organizations at a

workshop organized by Dr. James

Collins, Dr. Elizabeth Davidson, and Dr.

Andrew Storfer from Arizona State

University and sponsored by the

National Science Foundation in

Arlington, Virginia on May 28-29, 1998.

On Thursday morning, May 28, strong

evidence for declines of amphibian

populations in different geographic

locations around the world was

presented. The group consensus was

that there is a global amphibian decline

problem, but there is no single cause.

Rather, multiple factors are implicated,

including: habitat loss and alteration,

global change, pathogens, parasites,

toxic chemicals, ultraviolet radiation,

and invasive species. The potential

effects of four of these factors; UV,

toxic chemicals, pathogens, and global

change, were discussed in detail on

Thursday afternoon. On Friday,

participants outlined a research and

management plan and passed a

resolution (below) that summarizes the

plan. Follow-up workshops on specific

topics were outlined, and the National

Science Foundation has already funded a

workshop on disease in amphibians in

late July 1998.

Resolution: declining amphibian

populations. Whereas, there is compel-

ling evidence that, over the last 15

years, there have been unusual and

substantial declines in abundance and

numbers of populations of various

species of amphibians in globally

distributed geographic regions, and

Whereas, many of the declines are in

protected areas or other places not

affected by obvious degradation of

habitats, and Whereas, these factors are

symptomatic of a general decline in

environmental quality, and Whereas,

even where amphibian populations

persist, there are factors that may place

them at risk, and Whereas, some

patterns of amphibian population

decline appear to be linked by causative

factors, and Whereas, declines can occur

on multiple scales, in different phases

of amphibian life cycles, and can impact

species with differing ecology and

behavior, and Whereas, there 1s no

obvious single common cause of these

declines, and Whereas, amphibian

declines, including species extinctions

can be caused by multiple environmen-

tal factors, including habitat loss and

alteration, global change, pathogens,

parasites, various chemicals, ultraviolet

radiation, invasive species, and

stochastic events, and Whereas, these

factors may act alone, sequentially, or

synergistically to impact amphibian

populations, and Whereas, to under-

stand, mitigate and preempt the

impacts of these factors, a comprehen-

sive, interdisciplinary research program

must be undertaken, and Whereas, this

research program must be conducted in

several regions around the globe, both in

areas of known declines, and in areas

where declines have not been docu-

mented, and Whereas, this research

must examine issues ranging from

environmental quality of landscapes to

the condition of individual animals.

Now therefore be it resolved, the

signatories hereto call for the establish-

ment of an interdisciplinary and

collaborative research program, which

will specify and quantify the direct and

indirect factors affecting amphibian

population dynamics, and Be it further

resolved, that this program will include

basic research and monitoring that will

test hypotheses of causative factors

and examine patterns of change through

historical records, field-based correla-

tive data, and controlled, multi-factorial

experiments, and Be it further resolved,

that interdisciplinary, incident response

teams should be assembled in “hot

spots” of amphibian decline to identify

causative factors to facilitate the

mitigation of these sudden declines, and

Be it further resolved, that the

signatories hereto call upon both public

and private agencies and institutions, to

promote and support research, policies

and conservation measures that will

ameliorate losses and declines of

amphibian populations, and Be it

further resolved, that this broad-based

approach to the study of amphibian

population dynamics will serve as a

model for study of the global

biodiversity crisis. Submitted by

Amphibian and Reptile Conservation 2(1):33-34.

Andrew Storfer, Department of Biology,

Arizona State University, Tempe,

Arizona 85287-1501, USA. Tel: (602)

965-5857; fax: (602) 965-2519; email:

astorfer@asu.edu

ANNOUNCEMENTS

NARCAM. Since 1995, reports of

malformed amphibians have increased in

number, and public concern for the

health of our environment has grown.

The North American Reporting Center

for Amphibian Malformations

(NARCAM) was established as a

central repository for information on

this phenomenon. With the help of the

public and scientists, NARCAM strives

to convey an accurate account of the

frequency and distribution of malformed

amphibians. NARCAM is maintained

by the Northern Prairie Wildlife

Research Center in Jamestown, North

Dakota, a facility of the United States

Geological Survey, Biological Resources

Division. The United States Environ-

mental Protection Agency provides

additional support. NARCAM’s

worldwide website contains maps on

the geographic distribution of reports in

North America, along with a description

of the type of malformation present at

each site. The site also has photos of

the different types of malformations

that may be encountered. For help in

identification of species, NARCAM has

a growing online guide to the amphib-

ians of the United States and Canada

that most frequently are reported with

malformations. The public and

researchers can submit information

directly through the Web site (http://

Www.npwic.usgs.gov/narcam) by using

an online data reporting form. Individu-

als who do not have Web access can

phone in reports toll-free at 1 (800)

238-9801. The public is urged to report

sightings of malformed amphibians. If

appropriate, a local biologist will visit

the site to confirm the species identity

and collect additional information.

Submitted by Jeff A. Jundt, Coordinator,

North American Reporting Center for

Amphibian Malformations, Northern

Prairie Wildlife Research Center, United

States Geological Service (USGS)/

Biological Resources Division, 8711

37th Street Southeast, Jamestown, North

Dakota 58401, USA. Tel: (701) 253-

5580; fax: (701) 253-5553; email:

Jeffrey_Jundt @ usgs.gov

MEETINGS

Society for the Study of Amphibians and

Reptiles and the Herpetologists’ League

Annual Joint Meeting, 27-31 July 2000,

Indiana University Purdue University at

Indianapolis, Indianapolis, Indiana.

Two symposiums: amphibian popula-

tion declines (organized by David M.

Green and Karen Lips) and a one-day

symposium on herpetological research

in zoos: the academic connection. For

further information contact Henry R.

Mushinsky, Department of Biology,

University of South Florida, Tampa,

Florida 33620. Tel: (813) 974-5218;

email: MUSHINSK@ CHUMAI.CAS.

USF EDU

21st Annual Symposium on Sea Turtle

Biology and Conservation Philadelphia,

USA February 24-28, 2001. For more

information reference: http://www.

seaturtle.org/symposium/2001

WEBSITES

AmphibiaWeb

elib.cs.berkeley.edu/aw

Center for North American

Amphibians and Reptiles

Eagle.cc.ukans.edu/~cnaar/

CNAARHomePage.html

Conservation Breeding

Specialist Group

www.cbsg.org

International Zoo News

www.quantum-conservation.org/IZN/294/

IZN-294.html

Kansas Herpetological Society

eagle.cc.ukans.edu/~cnaar/khs/khsmain.html

Online Herpetologists Directory

www.smoky-hills.com/directory/search.asp

Species Survival Commission

www.iucn.org/themes/SSC

World Conservation Monitoring Centre

www.wemce.org.uk

World Zoo Organization

WWW.WZO.Org

NEW LITERATURE

Journal—A dvances in Amphibian

Research in the Former Soviet Union

(AARFSU) [ISSN: 1310-8840]. Volume

1, Kuzmin, S. L. and Dodd, Jr., C. K.

(editors). [ISBN: 954-642-017-4], vi +

233 p. (9 color photos). Volume 2

(ISBN: 954-642-019-0), vi + 189 p.,

11 color photos. Format 165 x 235

mm, paperback, color photos,

numerous black and white graphs,

photos, drawings, figures, maps, and

tables. Text in English. US$34.

Kuzmin, S. L. and Wilkinson, J. 1998.

Volume 3 (ISBN: 954-642-046-8), 165

x 245 mm, graphs, maps, black and

white drawings, and ~ 60 color

photographs. Text in English, 245 p.

US$34. Tarkhnishvili, D. N. and

Gokhelashvili, R. K. 1999. Volume 4,

The Amphibians of the Caucasus

(ISBN: 954-642-047-6). Format 165 x

235, graphs, color and black and white

drawings, photos, tables, and bibliog-

raphy. Text in English. 240 p. US$34.

Discounts are possible for ordering all

three volumes and subscribing to

future issues. Inquiries and ordering

information can be obtained from

PENSOPT Publishers (Sofia and

Moscow-based scientific publishers

and booksellers), Dr. Lyubomir D.

Penev, Akad. G. Bonchev Street, Block

6, 1113 Sofia, Bulgaria. Tel: +359-2-

716451; fax: +359-2-704508; email:

pensoft @ mbox.infotel.bg; website:

www. pensoft.net

Electronic Journal—Contemporary

Herpetology (CH). CH 1s a peer-

reviewed electronic journal devoted to

herpetology on-line at URL: http://

vmsweb.selu.edu/~pcsd4805. CH

plans to publish articles covering all

aspects of herpetology, including

ecology, ethnology, systematics,

conservation biology, and physiology.

CH also plans to publish monographs,

points-of-view, and faunistic surveys

of poorly known areas but will not

publish herpetocultural or anecdotal

papers. For more information contact

the editor, Joe Slowinski, at tel:(415)

750-7041 (or) by email: jslowins @ cas.

calacademy.org

LITERATURE

Burbrink, F. T. et al. 1998. A riparian

zone in southern I]linois as a poten-

tial dispersal corridor for reptiles

and amphibians. Biological Conser-

vation 86 (2):107-115.

News and Notes

de Silva, Anslem. 1997. Report on the In-

ternational Conference of the Biology

and Conservation of the South Asian

Amphibians and Reptiles. Lyrioce-

phalus. 3(1):40-43. [Faculty of Medi-

cine, University of Peradeniya, Sri

Lanka]

Gibbons, J. W. 1997. Measuring declines

and natural variation in turtle popu-

lations: spatial lessons from long-

term studies, p. 243-246 in

Abbema, J. V. (editor). Interna-

tional Conference on Conservation,

Restoration, and Management of

Tortoises and Turtles. New York

Turtle and Tortoise Society, Pur-

chase, New York.

Hager, H. A. 1998. Area-sensitivity of

reptiles and amphibians: are there in-

dicator species for habitat fragmen-

tation? Ecoscience 5:139-147.

Halliday, T. 1998. Ecology: a declining am-

phibian conundrum. Nature 394:418-

419.

Lips, K. R. 1998. Decline of a tropical

montane amphibian fauna. Conser-

vation Biology 12 (1):106-117.

Lovich, J. E. and Gibbons, J. W. 1997.

Conservation of covert species: pro-

tecting species we don’t even know,

p. 426-429 in Abbema, J. V. (editor).

International Conference on Conser-

vation, Restoration, and Management

of Tortoises and Turtles. New York

Turtle and Tortoise Society, Pur-

chase, New York.

Seigel, R. A., Sheil, C. A., and Doody, J.

S. 1998. Changes in a population of

an endangered rattlesnake Sistrurus

catenatus following a severe flood.

Biological Conservation 83:127-131.

Webb J. K., and Shine R.1998. Using ther-

mal ecology to predict retreat-site se-

lection by an endangered snake spe-

cies. Biological Conservation

86(2):233-242.

BOOKS AND LITERATURE

RECEIVED

Points of view on contemporary education

in herpetology. Herpetologica 54 (2)

{Supplement]. S82 p.

Laurance, W. F. and Bierregaard, Jr., R. O.

(editors). 1997. Tropical Forest Rem-

nants: ecology, management, and con-

servation of fragmented communities.

The University of Chicago Press, Chi-

cago, Illinois. 616 p. (ISBN: 0-226-

46899-2).

P. 6-14

P. 15-23

P. 24-29

Amphibian and Reptile Conservation 2(1):35.

| : The Last Page

GLOSSARY

Definitions of words footnoted in journal articles.

“qutoecological.—ts the study of the relationship of individual

organisms to their environment.

’xeric.—“Dry,” of or pertaining to a habitat having a low or

inadequate supply of moisture.

‘sisal.—A species of agave grown for commercial production of

fiber.

4qestivation.—A state of inactivity occurring in some animals

during prolonged periods of drought or heat.

Diplodactylidae (In title). —Family of southwest Pacific geckos

and pygopods, distributed in Australia, New Zealand, New

Caledonia, and in the case of pygopods, New Guinea.

‘carphodactyline—Adjectival form of Carphodactylini, a tribe

of the Diplodactylidae including Rhacodactylus.

/viviparity.—Reproductive mode in which development takes

place within the mother and young are born live, as opposed to

oviparity, in which eggs are laid.

£allozymic.—In reference to allozymes, the different forms of

proteins that can be separated by electrophoresis to yield evi-

dence about relationships or similarity of individuals within or

between populations and species.

"polymorphic.—Having many forms, usually refers to the pres-

ence of more than one manifestation of particular characters

within a single taxon.

‘genetic distance data.—Measurements of the relative similarity

between taxa or individuals being compared, based on direct or

indirect genetic information. In this instance, genetic distance

data were derived from the allozyme data obtained by electro-

phoresis.

Jsclerophyll forest —Critically endangered dry forest found only

in isolated pockets on the west coast of New Caledonia.

‘maquis vegetation.—The characteristic and highly endemic scrub

vegetation of the mineral-rich soils of parts of New Caledonia,

especially the south.

New abbreviations appear herein. Past issues should be consulted for previous abbreviation usage.

W = West N = North

E = East S = South

et al. = and others F = Fahrenheit

e.g.= forexample C = Celsius

cm =centimeter tel = telephone

mm = millimeter fax = fascimile

km = kilometer

kg = kilogram

email = electronic mail

ha = hectares

U.S. and US = United States

USA = United States of America

pers. = personal

m=meter

UK = United Kingdom

Pp. = pages, page

~ = approximately obs. = observation(s)

fig. = figure

ERRATA

comm. = communication(s)

The following journal corrections should be noted in Volume 1,

Number | (Premiere issue): Page 3. CONTENTS PAGE. The

article under Next Issue: Conservation of South African’s En-

demic Dwarf Chameleons has been postponed indefinitely.

MASTHEAD. The premiere issue (Volume 1, Number 1) is out-

of-print and no longer available, although nicely reproduced pho-

tocopies are available for $10 plus shipping and handling. All

back issues are available for $12 each while regular supplies last.

PAGE 10. In the “Key words” section the word Viridi_ should

have been spelled Viridis. PAGE 16. Under the photo caption

the word Plethedon should have been spelled Plethodon. PAGE

20. The third sentence of the body of the article should read

“Inbreeding depression may adversely affect small populations

by unmasking recessive deleterious alleles and reducing heterozy-

gosity.” Pages 24-26. COLUMN. 1). The personal communica-

tion on page 25 (Groombridge 1994) was not attributed to Dr.

Brian Groombridge of the World Conservation Monitoring Cen-

tre (WCMC), Cambridge, England. 2). The WCMC database,

referred to in the column, is in fact the CITES database of trade

records. 3). By comparing export records and import records,

and checking against known countries of origin, it is in fact

possible to get some idea of the extent of misreporting (The

updated commentary for this particular article was provided by

Brian Groombridge on November 8, 1996). PAGE 27. Meet-

ings. The meeting date for the conference Conservation and

Biodiversity of Amphibians and Reptiles of Tropical Rain For-

ests was re-scheduled for June 1999 and was then subsequently

cancelled.

NOTE.—The most up-to-date Writer’s Guidelines and

Manuscript Preparation Instructions can be located at the Am-

phibian and Reptile Conservation website at: www.

herpetofauna.com

ACKNOWLEDGEMENTS

As the journal grows and climbs towards international as well as

domestic prominence, a growing list of individuals and organiza-

tions must be credited with their help, assistance, and support.

Individuals who have kept their enthusiasm and support for

the project, as well as me individually, are Jack Sites, Harvey

Lillywhite, Mark Goodwin, Jamie Reaser, Craig Hoover, Carl

Gans, Peter Lindeman, Maria Bridarolli, John Megahan, Dave

Stone, Chris Raxworthy, Ron Nussbaum, Aaron Bauer, Kraig

Adler, Allison Alberts, Michael Soulé, George Rabb, Tony

Krzysik, Jude McNally, Trevor Hare, Dave Hardy, and Dennis

Caldwell. A special thanks goes to Dave Piper, Jeff Middleton,

and Jose Solorzano and to all the authors, contributors, and asso-

ciates of the journal. Without your generous support, help, and

patience this journal would not exist.

Organizations which should be acknowledged for their support

and use of facilities are the Information Technology Center staff and

facility (especially, Dave Piper, Jeff Middleton, and Jose Solorzano),

Arizona Health Sciences Library; Science Reference, Science-Engi-

neering Library; and Multimedia & Visualization Lab, Center for

Computing & Information Technology; all at the University of An-

zona, Tucson, Arizona; University Computing Services, Brigham

Young University, Provo, Utah; F. Wayne King and the Museum of

Natural History, University of Florida, Gainesville, Florida.

A special tribute is extended to GRM of Tucson, Arizona,

and specifically Arnell McSwain, Dennis Barreras, Albert D.

Jones, Darryl A- Lacey, John Roden, Everett E. Sellers, Donald

Batson, Chipper Ward, David Morrow, Tim Church, and Leland

McKirryher for support, help, and friendship.

A very special thanks goes to Kenneth Sholar, our graphic

designer, who when I had all but given up on finding a talented

graphics person to work with on the journal, took on the task regard-

less of the fact he was working three jobs and attending school full-

time. ARC and I extend a heart felt thanks for his commitment and

fine work in helping get the journal “just nght’ in every detail.

DEDICATION

This issue is kindly dedicated to my family: my mother

Frances, my dad Anthony, and both my brothers Steve,

and Greg Hasspakis.