wie UN

‘Significant Trade in Wildlife:

A Review of Selected Species
in Cites Appendix II

Volume 2: Reptiles and Invertebrates

Ss

Compiléd by IUCN’ Conservation Monitoring Centre

SIGNIFICANT TRADE IN WILDLIFE:

A REVIEW OF SELECTED SPECIES IN CITES APPENDIX Il

COMMERCE IMPORTANT DES ESPECES SAUVAGES: ENQUETE AU SUJET

DE CERTAINES ESPECES INSCRITES A L’ANNEXE Il DE LA CITES

COMMERCIO SIGNIFICATIVO DE VIDA SILVESTRE: ESTUDIO DE

DETERMINADAS ESPECIES INCLUIDAS EN EL APPENDICE Il DE CITES

VOLUME 2: REPTILES AND INVERTEBRATES
VOLUME 2: REPTILES ET INVERTEBRES

VOLUMEN 2: REPTILES E INVERTEBRES

Edited by Publié par Publicado por
Richard Luxmoore, Brian Groombridge and Steven Broad.

IUCN Conservation Monitoring Centre, 219c Huntingdon Road, Cambridge, UK.

1988
International Union for Conservation of Nature and Natural Resources

Secretariat of the Convention on International Trade in Endangered Species
of Wild Fauna and Flora

A joint publication of the International Union for Conservation of Nature and
Natural Resources (IUCN), Gland, Switzerland and Cambridge, UK and the
Secretariat of the Convention on International Trade in Endangered Species of
Wild Fauna and Flora, Lausanne, Switzerland.

1988.

The publishers acknowledge the financial support of the governments of Canada,
Denmark, Federal Republic of Germany, Luxembourg, Netherlands, Portugal and
the United States of America and of WWF-USA, WWF-Switzerland, Pet Industries
Joint Advisory Council.

This report was prepared under contract to the Secretariat of the Convention
on International Trade in Endangered Species of Wild Fauna and Flora by IUCN
Conservation Monitoring Centre, which is supported by the United Nations
Environment Programme and the World Wide Fund for Nature (WWF).

1988. International Union for Conservation of Nature and Natural
Resources and the Secretariat of the Convention on International Trade in
Endangered Species of Wild Fauna and Flora.

ISBN 2-88032-954-K

Printed in Canada by the Canadian CITES Management Authority (Canadian
Wildlife Service), Ottawa.

Cover photo: Python molurus WWF/Romulus Whitaker

The designations of the geographical entities in this book and_ the
presentation of the material do not imply the expression of any opinion
whatsoever on the part of IUCN or the CITES Secretariat concerning the legal
status of any country, territory, or area, or of its authorities, or
concerning the delimitation of its frontiers or boundaries.

The views of the authors expressed in this publication do not necessarily
reflect those of IUCN or the CITES Secretariat.

ACKNOWLEDGEMENTS

This report was prepared with the assistance of many of the staff at IUCN

Conservation Monitoring Centre. The first drafts of the species accounts were

written by the following authors:

Brian Groombridge Geochelone chilensis, Testudo spp., Crocodylus_ spp.,
Varanus spp., Python spp.,

Diana Evans Geochelone pardalis, Malacochersus tornieri, Phelsuma
spp., Chamaeleo spp.

Richard Luxmoore Podocnemis expansa, Caiman crocodilus, Iguana iguana,

Dracaena guianensis, Tupinambis Spp., Boa
constrictor, Eunectes spp.

Mark Collins Ornithoptera spp.

Sue Wells Papustyla pulcherrima, Cirrhipathes anguina.

Tim Inskipp was responsible for the over-all co-ordination of the contract and
assisted with the literature research. John Caldwell produced the basic trade
tabulations from the Wildlife Trade Monitoring Unit computerised database.
Wendy Coombes and Andrew Desforges carried out much of the analysis of the
CITES trade data. The accounts for Geochelone chilensis, Malacochersus
tornieri, Testudo graeca, Testudo hermanni, Podocnemis expansa, Caiman
crocodilus, Crocodylus n. novaeguineae and Crocodylus porosus were based
on the accounts written for the IUCN Amphibia-Reptilia Red Data Book, Part l,
Testudines, Crocodylia, Rhynchocephalia.

We should like to thank all those who responded to requests for information
sent out by ourselves and the CITES Secretariat,, and to those who reviewed
the earlier drafts of this report. They include: M. Abu Jafar, R.T. Adams,
W. Aguilar, J.B. Alvarez, S. Ambu, P.A. Anadu, R.D. Auerbach, W. Auffenberg,
T. Belokapova, J.M. de Benito, E. Bennett, M.A. Bereteh, S.S. Bist, Q. Bloxam,
Bodiopelli, E. Boomker, L. Bortolotti, D.M. Botello, D.G. Broadley,
W.Y. Brockelman, S.D. Budd, A.A. Burbidge, P. Butler, G. Ceballos,
M.K. Cheung, Chew Hong, P.B. Clark, J.A. Cranwell, J.A. Crespo, G. Davison,
P.G. Diaz, R. Dipouma, F. Djedjo, C. Dudley, Dr Fischer, H.S. Fitch,
A.S. Gardner, A. Gaski, E. Ghamba, E.O. Gonzalez Ruiz, P. Gopalakrishnakone,
S. Gorzula, A. Haynes, J. Hebrard, G. Hemley, M.S. Hoogmoed, K. Howell,
. d'Huart, R.F. Inger, T. Jalel, C.J. Kalden, B.H. Kiew, W. King,
. Mackay, W.E. Magnusson, M. Mamane, E.J. Martinez, G.T. Masina,
. Mitchell, D. Momo, J.V. Morales Molina, J.A. Mortimer, J.R. Navas,
.P. Ntsekhe, F.H.O. Opolot, J.A. Ottenwalder, J.D. Ovington, A.W. Owadally,
J.T. Palazzo, A.L. Peal, D. Perry, P.C.H. Pritchard, M. Romero Pastor, Rubini
Atmawidjaja, N.M. Sarker, fF. Serracino-Inglott, K. Shirazi, M.P. Simbotwe,
P. Singsouriya, A. Stimson, I. Swingland, Syafii Manan, J. Thomsen, R. Thorpe,
E. Vallester, M. Vinas, H. Watson, D. Werner, Yang Peng Hwa,

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Digitized by the Internet Archive
in 2010 with funding from
UNEP-WCMC, Cambridge

htto://www.archive.org/details/significanttradeO2luxm

INTRODUCTION

Background

The Convention on International Trade in Endangered Species of Wild Fauna and
Flora (CITES) was drawn up in 1973 to control trade in wildlife. It does so
by affording to species either of primarily two levels of protection. Those
species (or smaller geographical populations) which are threatened with
extinction are listed in Appendix I, and are thereby banned from international
commerce under most circumstances. Species which are not currently threatened
with extinction, but which may become so unless their trade is regulated, are
listed in Appendix II. Such species may be traded internationally, but
nations must ensure that the levels of trade do not endanger the remaining
wild populations. This requirement is expressed formally in the text of the
Convention in Article IV, paragraph 2a, which demands that the authorities in
exporting countries must have advised that the export of specimens of such a
species “will not be detrimental to the survival of that species". Article
IV, paragraph 3 indicates that the trade in a species "should be limited in
order to maintain the species throughout its range and at a level consistent
with its role in the ecosystem in which it occurs and well above the level at
which that species might become eligible for inclusion in Appendix I". The
authorities in the exporting country must monitor the exports and take steps
to limit them whenever they determine it to be necessary.

At the fourth meeting of the Conference of the Parties to CITES, held in 1983
in Gaborone, Botswana, it was recognised that many countries exporting
Appendix II wildlife were unable, on their own, to determine whether the
levels of trade were having a detrimental effect on the wild populations.
Therefore it was recommended (CITES Resolution Conf. 4.7) that the CITES
Technical Committee should provide assistance by identifying those Appendix II
species which were currently being traded internationally in significant
quantities, but for which there was insufficient scientific information on the
capacity of the species to withstand such levels of trade to satisfy “the
requirements of Article IV, paragraph 3, of the Convention as determined by
the range states". It was recommended that once the species of particular
concern had been identified, the Technical Committee, together with the range
states involved, importing states and organisations experienced in the
management of wildlife, “develop and negotiate measures required to ensure
that continued trade in these species is within the terms of Article IV,
paragraph 3". ;

Initial discussions of the means by which the Technical Committee could
identify those species of particular concern (as recommended by Resolution
Conf. 4.7) were based on the premise that a high volume of trade was
sufficient evidence alone to justify concern. However, an unpublished report
produced in 1984 by WIMU for the CITES Secretariat, on the perception of the
issue of high trade-volume, came to the following conclusions:

- The concept of high trade-volume may be approached in two ways: high
volume may be considered in absolute terms (i.e. large numbers), or in
relative terms (i.e. large numbers in relation to the population and
biology of the species).

- Absolute high trade-volume does not alone have any bearing on whether a
species is threatened by trade. However, species traded in high absolute
numbers are likely to be of considerable ecological significance.

- Relative high trade-volume is of direct relevance to the survival of the
species involved, but there is no evidence that this is correlated with
absolute high trade-volume. By virtue of their designation on the

iii

Appendices, trade in all CITES-listed species is of concern, and should be
monitored.

- Consideration of absolute high trade-volume as a major criterion for
selecting species for special attention is thus not only irrelevant in
terms of species conservation, but may divert attention from more
important cases.

The Technical Committee Working Group on Significant Trade in Appendix II
Species produced a paper, based on its meeting in Switzerland in December
1984, which aimed to formulate a procedure or course of action to enable the
Technical Committee to fulfil the recommendations of Resolution Conf. 4.7. It
was decided that the Group should restrict its attention to fauna, as a Plant
Working Group was already in existence. The conclusions of the WIMU report on
high trade-volume were endorsed, in that the Working Group agreed that it was
not possible to identify those Appendix II taxa of greatest concern on the
basis of trade data alone. Information on biological status, population
trends and a whole range of other factors was needed in order to assess
properly the impact of the trade in those taxa.

A five-part procedure was established as the most appropriate mechanism for
implementing Resolution Conf. 4.7. This plan was presented to the fifth
meeting of the Conference of the Parties which was held at Buenos Aires,
Argentina in 1985 (Doc. 5.26). Steps 1-3 have already been carried out.

Step 1: Production of list "A"

It was acknowledged that, with a very few exceptions, all taxa listed in
Appendix II should be able to withstand some degree of exploitation for
international trade. The Working Group chose an arbitrary “safe” level of
trade for any such taxon of an average of 100 individuals taken from the
wild (globally) and entering trade per year. By eliminating all taxa
traded at a level within that considered “safe", a list of “potential
candidate" taxa could be produced (List "A"). These taxa were defined as
those that might be the subject of significant international trade.

List A was prepared by WIMU on the basis of average trade volume over the
pericd 1980-1982. Figures relating to live specimens (excluding those
recorded as captive-bred), whole or substantially whole skins, skin
flanks/sides, furskin plates, shells, trophies and other worked material
were included in the analysis. Species never recorded in trade, with the
exception of those included in Appendix II as part of a higher taxon or
for look-alike reasons, were listed separately in order that consideration
could be given to their deletion from the Appendices.

Step 2: Production of list "B"

The Working Group agreed that some taxa might be eliminated from
consideration as “significant trade" species on the basis of knowledge
readily available to the Group regarding their status. After this
process, the remaining taxa constituted list "B", which contained those
taxa which could be classified as a “possible problem". In addition, two
species (Tupinambis rufescens and Papustyla Pulcherrima) were added to
this list under special circumstances where there was evidence of a
problem despite only a low volume of trade being recorded.

Step 3: Production of list "c™

The next phase in the procedure was to assess the information available
for each of the species in list "B", and to eliminate those species which

iv

were, on the basis of expert knowledge, known not to be a problem. This
part of the operation entailed the collection of information on as many
aspects of each species as possible and the assessment of the impact of
the known trade on the known population. The Working Group agreed that
for each species the global situation should be of paramount importance,
but that if a species were apparently being affected by trade on a
national or regional scale, this fact should be noted in an addendum to
the list. List "C" was to be divided into two groups: those species for
which current information or knowledge of their biology and/or management
indicated that the population was being detrimentally affected by
international trade (List 1), and those species for which there is
insufficient information available on which to base such a judgement (List
2

Step 4: Development of remedial measures

The Technical Committee, or a working group of the Technical Committee,
was to examine the lists "1" and "2" and establish priorities within each
list. For species of high priority in list "1", workshops were to be
convened to formulate recommendations for remedial measures. Such
measures would include, but not necessarily be limited to: preparing
proposals for transfer to Appendix I; establishment of additional
Management procedures both for wild populations (hunting quotas, seasons,
size limits, etc.) and for trade controls (such as export quotas); and
listing of taxa for look-alike reasons.

For species of high priority in list "2", projects were to be established
to collect information on the biology and management of the species.
Where such information indicated the need, the species were to be
transferred to list "1".

Step 5: Implementation of remedial measures

The remedial measures identified were to be carried out by the range
states involved on the basis of the recommendations arising from the
workshops.

This five-step procedure was approved at the Buenos Aires meeting in 1985 and
steps 1-3 were implemented by the IUCN Conservation Monitoring Centre. List
"C" was prepared in time for the second meeting of the Technical Committee,
held in June 1986 in Lausanne, Switzerland. For each species in list "C", a
draft report was prepared presenting a summary of all available information,
including a detailed analysis of available trade data and information on the
population status and other factors thought to be of relevance. On the basis
of this information, each species was assigned to the two recommended lists
(list 1, problem species; list 2, possible problems). At this stage it was
also discovered that some species, originally included in list "C", were
probably not being significantly affected by the current levels of trade.
These were assigned to a third group (list 3, no problem). The Significant
Trade Working Group reviewed the information provided by CMC and the suggested
listings, and made a number of recommendations for further action which are
outlined below. The Technical Committee also decided that, after further
review, the report prepared by CMC should be published.

Further action

The Significant Trade Working Group presented a paper at the second Technical
Committee meeting outlining proposals for further action (WGR.TEC. 2.2). The
recommendations of this report, some of which were amended at the Technical

Committee meeting, are detailed below for the reptile species involved.

Vv

List 1 (4 taxa)

Podocnemis expansa The international trade data are probably greatly
exaggerated as a result of false identification. The Secretariat should
notify the Parties of current trade bans, and Venezuela will consider
developing a proposal to transfer the species to Appendix I bearing in mind
the possible insignificance of international trade.

Caiman crocodilus The problem is being addressed by the Secretariat's
project in Bolivia, Brazil and Paraguay and the results of this study should
be useful in the establishment of appropriate quotas. Funding for the second
project (Brazil, Colombia and Venezuela) is needed. Studies of the population
status in Central America should be considered. Historical trade data
including numbers and sizes of skins should be collected and analysed. Where
the species is legally exploited, national harvest quotas should be
established.

Geochelone chilensis Management action already taken by Argentina is
considered adequate.

Tupinambis spp. Steps are being taken by Argentina to review their harvest
quotas. TJIllegal trade may continue to be a problem and the Secretariat should
assess the 1984/85 trade statistics, identify any importing countries
permitting trade from exporting countries with export bans and request that
the necessary action be taken to halt this problem. The Nomenclature
Committee should clarify the taxonomy.

List 2 (17 taxa)
The Working Group recommended that the following taxa should receive attention
as priority species or groups of species for the collection of information (in

order of importance):

1. Asian Varanidae (three species, i.e. Varanus salvator and the Appendix I
species V. bengalensis and V. flavescens).

2. Asian pythons (three species, i.e. Python curtus, P. molurus
bivittatus and P. reticulatus).

3. African monitor lizards (two species, i.e. Varanus exanthematicus and

V. niloticus) - noting that the project should concentrate on aspects
necessary to assess the levels of exploitation that the species can
sustain.

4. South American Boidae (three species, i.e. Boa constrictor, SEunectes
murinus and E. notaeus).

5. Papustyla pulcherrima.

In addition the Group made the following special recommendations:

Dracaena guianensis The Parties having populations of Dracaena. spp.
should clarify the situation with respect to distribution and consider listing
Dracaena spp. in Appendix II.

Crocodylus porosus The IUCN/SSC Crocodile Specialist Group should be asked

urgently to review the management programme for this species in the light of
the recent Indonesian report.

vi

List 3 (15 taxa)

It was agreed that available information indicated that these taxa were
essentially unaffected by international trade.

METHODS

This report comprises the review of the biological and trade status of species
included in list "C". It was carried out by the IUCN Conservation Monitoring
Centre under contract to the CITES Secretariat over the period September 1985
to April 1986. As a first step, the CITES Secretariat circulated a request
for information to all of the countries in which the species occurred,
contacting the CITES Management Authorities in the countries party to CITES
and designated wildlife management or equivalent authorities in others. The
responses to this request were passed to CMC and are referenced in the
following format: Name of country CITES MA, 1987. Comments received from
wildlife management authorities in non-Party states are referenced by the name
of the government department involved. Information was also solicited from
relevant specialists (individuals or agencies), and amongst the major sources
were the specialist groups of the IUCN Species Survival Commission. Trade
organisations and other interested parties were also approached. A draft
report was presented to the 2nd meeting of the CITES Technical Committee in
June 1986. This report was discussed and amended by the Committee and review
copies were again circulated by the CITES Secretariat to all range states and
interested parties, including the Pet Industry Joint Advisory Council. Final
modifications to the text and recent trade data were added by CMC during 1987.

In a small number of cases the designation of category of a species at the
time of the second Technical Committee meeting has subsequently been amended
in the light of new information, in particular the 1985 trade data which have
been added to the reports.

Information was collected and collated under the following headings:
distribution; population; habitat and_ ecology; threats to survival;
international trade; conservation measures; and captive breeding.

CITES trade data were analysed for the years 1980 to 1985 using the Annual
Reports of Parties to the Convention for which the statistics are held on
computer at CMC. These data contain records of imports and exports of species
listed in the CITES Appendices and of their products. They contain
information on the species involved, a description of the type and quantity of
product and, in the case of imports, the exporter or re-exporter and primary
source country, and, for exports, the destination and original source. For
trade between two CITES Parties, each transaction should therefore be reported
twice, once by the importer and once by the exporter. As suggested by the
Significant Trade Working Group, the analysis was largely restricted to trade
in live animals and unworked products, however, in a small number of
exceptional cases worked products were included.

Various problems impair the value of CITES trade data in the assessment of
levels of world trade. For example: not all trading nations are CITES
Parties; not all CITES Parties produce annual reports; and the reports of
those that do, vary in quality and regularity of submission. Some countries
may report the number of specimens covered by the permits issued, while others
report the actual number for which the permit was used. Furthermore exports
from a country at the end of one year may arrive in the importing country
early in the next and in such cases it is possible that the same transaction
may be recorded in the trade tables for both years. These factors and others
have to be taken into account when analysing CITES data, but for most species

vii

these statistics are the only detailed source of information on their
international trade and generally CITES reports are of great value in
assessing approximate levels of legal trade, the geographical patterns in
such trade and the trends in volume and commodity preference over time.

In most cases the trade data are presented, in the following accounts, in two
tables. The first (usually Table 1) details the net imports of importing
countries, the total of which gives an estimate of the minimum volume of world
trade for each year. The second (usually Table 2) shows the origin, or where
no origin was given, the exporter, of specimens in trade. When specimens have
been exported to an intermediate country and subsequently re-exported, the
minimum net trade was calculated, ensuring that the numbers were only recorded
once. The table therefore shows, for each year, the minimum number of items
in trade from each country of origin. However, because some items may be
re-exported without the country of origin being specified, they may be
recorded twice in Table 2. The totals are therefore usually higher than those
in Table 1.

viii

INTRODUCTION
Informations générales

La Convention sur le commerce international des espéces de faune et de flore
Sauvages menacées d'extinction (CITES) a été élaborée, en 1973, pour contréler
le commerce des espéces de faune et de flore sauvages. Elle agit en offrant a
ces espéces deux niveaux principaux de protection. Les espéces (ou de plus
petites populations géographiquement isolées) qui sont menacées d'extinction
sont inscritent a l'Annexe I, ce qui signifie que leur commerce international
est interdit dans la plupart des cas. Les espéces qui ne sont pas actuellement
menacées d'extinction, mais qui pourraient le devenir si leur commerce n'‘était
pas réglementé, sont inscritent a l'Annexe II. Le commerce international de
ces derniéres espéces est autorisé, a condition que les pays s'assurent que le
volume du commerce ne mette pas en danger la survie des populations sauvages
restantes. Cette exigence est formellement énoncée a =1'Article IV,
paragraphe 2 a, du texte de la Convention, qui prévoit que les autorités des
pays d'exportation émettent l'’avis que l'’exportation de spécimens de ces
espéces "ne nuit pas a la survie de l'espéce intéressée". Le paragraphe 3 de
l'article IV indique que le commerce d'une espéce “devrait é6étre limité pour la
conserver dans toute son aire de distribution, a un niveau qui soit a la fois
conforme a son réle dans les écosystémes ot elle est présente, et nettement
supérieur a celui qui entrainerait l'inscription de cette espace a
l'Annexe I". Les autorités des pays d'exportation doivent surveiller les
exportations de facon continue et prendre les mesures qui s'imposent pour les
limiter lorsqu'elles le jugent nécessaire.

Lors de la quatriéme session de la Conférence des Parties 4 la CITES, tenue en
1983 a Gaborone, Botswana, il fut reconnu que maints pays exportateurs
d'espéces de faune et de flore sauvages figurant a l'Annexe II étaient dans
l'incapacité de déterminer par eux-mémes si les niveaux de commerce avaient un
effet nuisible sur les populations sauvages. C’est pourquoi, il fut recommandé
(résolution CITES Conf. 4.7) que le Comité technique de la CITES assiste ces
pays en identifiant les espéces de l'Annexe II faisant actuellement l'objet
d'un commerce international important, mais pour lesquelles, selon l'‘avis des
Etats de l'aire de répartition, les données scientifiques portant sur leur
capacité & supporter le commerce a un tel niveau sont insuffisantes au regard
des exigences de l'Article IV, paragraphe 3, de la Convention. Il fut
recommandé que, une fois les espéces présentant un intérét particulier
identifiées, le Comité technique, en collaboration avec les Etats de l'aire de
répartition intéressés, les Etats importateurs et les organisations ayant une
expérience en gestion de la faune et de la flore sauvages, "mette au point et
négocie les mesures nécessaires pour assurer le maintien du commerce continu
de ces espéces dans les limites prévues a l'Article IV, paragraphe 3, de la
Convention".

Les discussions initiales sur les moyens a utiliser par le Comité technique
pour identifier les espéces représentant un intérét particulier (selon la
recommandation de la résolution Conf. 4.7) ont été fondées sur le principe
qu'un volume de commerce important est, & lui seul, une indication suffisante
pour justifier un intérét. Toutefois, un rapport non publié, produit en 1984
par le WIMU pour le Secrétariat CITES et traitant de la maniére dont il
percevait la question du volume important du commerce, parvenait aux
conclusions suivantes:

- Le concept du volume important du commerce peut é6tre abordé de deux
maniéres: un important volume peut étre considéré en terme absolu (soit de
grandes quantités) ou en terme relatif (soit de grandes quantités par
rapport a la population et a la biologie de l'espéce).

ix

- Un important volume de commerce, au sens absolu du terme, n'a pas en soi
de rapport avec le fait qu'une espéce soit menacée ou non par le commerce.
Toutefois, il est probable que les espéces dont de grandes quantités de
spécimens, en terme absolu, sont commercialisés aient une importance
écologique considérable.

- Un important volume de commerce, au sens relatif du terme, a un rapport
direct avec la survie de l'espéce en question, mais rien ne prouve qu'il y
ait corrélation avec un important volume de commerce au sens absolu du
terme. Le seul fait que ces espéces soient inscrites aux annexes A la
CITES signifie que leur commerce est motif a préoccupation et qu'il
devrait faire l'objet d'une surveillance continue.

- Considérer un important volume de commerce au sens absolu du terme comme
critére majeur de sélection des espéces nécessitant une attention
particuliére est donc non seulement hors de propos en ce qui concerne la
conservation des espéces mais, qui plus est, risquerait de distraire
l'attention de cas plus importants.

Le Groupe de travail du Comité technique sur le commerce important d'espéces
de l’Annexe II a élaboré un document, fondé sur sa session tenue en Suisse en
décembre 1984, session dont l'objectif était de formuler une procédure ou une
ligne de conduite permettant au Comité technique de remplir ses obligations au
titre des recommandations de la résolution Conf. 4.7. Il fut décidé que le
groupe devait limiter ses discussions a la faune en raison de l'existence d'un
Groupe de travail sur les plantes. Les conclusions du rapport du WTMU sur le
volume important du commerce furent endossées, en ce sens que le groupe de
travail convint qu'il n'était pas possible d'identifier les taxons les plus
préoccupants de l'Annexe II sur la base des seules données commerciales. Des
informations sur l'état biologique des taxons, sur les tendances de leurs
populations et sur toute une série d'autres facteurs sont nécessaires pour
évaluer correctement l'effet du commerce sur ces taxons.

Une procédure en cing étapes, constituant le mécanisme le plus favorable pour
l'application de la résolution Conf. 4.7, fut établie. Ce plan d'action fut
présenté a la cinquiéme session de la Conférence des Parties qui eut lieu a
Buenos Aires, Argentine, en 1985 (document Doc. 5.26). Les étapes 1 a 3 ont
déja été réalisées.

lére étape: Production de la liste "A"

Il fut reconnu que, A trés peu d'exceptions prés, on peut raisonnablement
assumer que tous les taxons inscrits a l'Annexe II peuvent supporter un
certain niveau d'exploitation pour le commerce international. Le groupe de
travail choisit un niveau de commerce arbitraire et "sfr" pour tout taxon,
soit en moyenne 100 individus prélevés dans la nature (globalement) et
entrant dans le commerce chaque année. En éliminant tous les taxons dont
le commerce était considéré d'un niveau "sfir", une liste de taxons
“candidats potentiels" (liste A") put alors étre établie. Ces taxons sont

définis comme étant ceux qui peuvent faire l'objet d'un commerce
international important.

La liste A a été établie par le WIMU sur la base d'un volume de commerce
moyen couvrant la période 1980-1982. Les chiffres ayant trait aux
spécimens vivants (sauf les spécimens enregistrés en tant qu'élevés en
captivité), aux peaux entiéres ou substantiellement entiéres, aux flancs,
aux nappes de peaux, aux carapaces, aux trophées et a d'autres articles
travaillés ont été inclus dans cette analyse. Les espéces qui n'ont jamais
été enregistrées dans le commerce, a l'exception de celles inscrites a
l'Annexe II en tant que partie d'un taxon supérieur ou pour des raisons de

x

ressemblance, ont été énumérées séparément en vue de leur éventuelle
élimination des annexes.

2e étape: Production de la liste "B"
Le groupe de travail convint que, sur la base des connaissances dont le

groupe pouvait disposer aisément au sujet de leur état, certains taxons ne
devaient plus étre considérés comme des espéces faisant l'objet d'un

“commerce important". Aprés cette opération, les taxons restants ont
constitués la liste "B", laquelle contient les taxons qui pourraient étre
classés en tant que "“probléme possible". En outre, deux espéces

(Tupinambis rufescens et Papustyla pulcherrima) ont 4té ajoutées a la
liste dans des circonstances particuliéres: la mise en é4vidence d'un
probléme en dépit de l'enregistrement d'un faible volume de commerce.

3e 6tape: Production de la liste "C"

L'étape suivante de la procédure revenait a évaluer les informations
disponibles pour chacune des espéces de la liste "B" et a éliminer les
espéces qui, sur la base des connaissances des experts, ne posent pas de
probléme. Cette partie de l'opération fut réalisée en rassemblant des
informations sur autant d'aspects que possible relatifs A chaque espéce et
en évaluant l'effet du commerce connu sur la population connue. Le groupe
de travail convint que, pour chaque espéce, la situation globale devait
avoir une importance’ primordiale, mais que, si une espéce était
apparemment affectée par le commerce a l'échelle nationale ou régionale,
ce fait devait figurer dans un supplément a la liste. Les espéces de la
liste "C" devaient 6tre réparties en deux groupes: d'une part les espéces
pour lesquelles les informations courantes ou la connaissance de leur
biologie et/ou de leur gestion montrent que la population est affectée par
le commerce international (liste 1) et d‘autre part les espéces pour
lesquelles les informations disponibles sont insuffisantes pour servir de
base a un tel jugement (liste 2).

4e étape: Mise au point de mesures correctives

Le Comité technique, ou un groupe de travail du Comité technique, devait
examiner les listes "1" et "2" annotées et établir des priorités au sein
de chaque liste. Pour les espéces de la liste "1" ayant un ordre de
priorité élevé, des sessions de travail devaient é6tre convoquées dans le
but de recommander des mesures correctives. De telles mesures devaient
comprendre, sans nécessairement s'y limiter: la préparation. de
propositions de transferts de taxons a l'Annexe I; la mise en place de
procédures de gestion supplémentaires, aussi bien en faveur des
populations sauvages (telles que quotas de chasse, saisons de chasse,
tailles limites des spécimens, etc.) qu'en ce qui concerne les contréles
du commerce (telles que quotas a l'exportation), et l'*inscription de
taxons pour des raisons de ressemblance.

Pour les espéces de la liste "2" ayant un ordre de priorité élevé, des
projets devaient étre élaborés afin de collecter des informations sur leur
biologie et leur gestion. Lorsque ces informations en montraient la
nécessité, l'espéce devait étre transférée 4 la liste "1".

Se étape: Mise en vigueur des mesures correctives
Les mesures de correction identifiées devaient é6étre prise par les Etats de

l'aire de répartition intéressés, sur la base des recommandations
formulées lors des sessions de travail.

xi

Cette procédure en cing étapes a été approuvée 4 la session de Buenos Aires,
en 1985, et les étapes 1 & 3 ont été réalisées par le Centre UICN de
surveillance continue de la conservation de la nature (CMC). La Liste "C" a
été élaborée pour la deuxiéme session du Comité technique, qui s'est tenue a
Lausanne, Suisse, en juin 1986. Pour chaque espéce de la liste "C", un projet
de rapport a été rédigé, lequel présentait un résumé de toutes les
informations disponibles, dont une analyse détaillée des données sur le
commerce et des informations sur l'état des populations et d'autres facteurs
jugés pertinents. Sur la base de ces informations, chaque espéce a été
assignée 4 l'une deux listes recommandées (liste 1, espéces a probleémes;
liste 2, problémes possibles). A ce stade, on a également découvert que
certaines des espéces figurant a l'origine sur la liste "C" n'étaient
probablement pas affectées de maniére significative par les niveaux actuels de
commerce. Celles-ci furent assignées & un troisiéme groupe (liste 3, sans
probléme). Le Groupe de travail sur le commerce important d'espéces de
l'Annexe II a étudié les informations fournies par le CMC, ainsi que les
listes proposées, et a fait un certain nombre de recommandations quant aux
activités futures qui sont décrites ci-aprés. Le Comité technique a également
décidé que, aprés un nouvel examen, le rapport élaboré par le CMC devait étre
publié.

Activités futures

Le Groupe de travail sur le commerce important d'espéces de l'Annexe II a
présenté un document a la deuxiéme session du Comité technique, document qui
ébauchait des projets d'activités futures (WGR. TEC. 2.2). Les recommandations
de ce rapport, dont certaines ont été modifiées lors de la session du Comité
technique, sont présentées ci-dessous de facon détaillée en ce qui concerne
les reptiles.

Liste 1 (4 taxons)

Podocnemis expansa - Les données sur son commerce international sont
certainement grandement exagérées, en raison d'une identification erronée. Le
Secrétariat devrait informer les Parties des interdictions actuellement
imposées au commerce de cette espéce, et le Venezuela envisagera 1'élaboration
d'une proposition de transfert de l'espéce a l’Annexe I, en ne perdant pas de
vue le fait que son commerce international pourrait étre insignifiant.

Caiman crocodilus - Le probléme est adressé par le projet du Secrétariat
réalisé en Bolivie, au Brésil et au Paraguay et les résultats de cette étude
devraient étre utiles a 1'établissement de quotas adéquats. Le financement du
deuxieme projet (Brésil, Colombie et Venezuela) doit encore étre trouvé. Des
études sur l'état des populations en Amérique centrale devraient étre
envisagées. Des données historiques sur le commerce de cette espéce, notamment
sur les quantités de peaux et leur taille, devraient 6tre réunies et

analysées. Lorsque l'espéce fait l'objet d'une exploitation légale, des quotas
nationaux devraient &tre établis.

Geochelone chilensis - La gestion de cette espéce déja entreprise par
l'Argentine parait adéquate.

Tupinambis spp. - L'Argentine entreprend les démarches nécessaires a la
révision des quotas de prélévement de cette espéce. Il se pourrait que le
commerce illégal de ces espéces soit toujours un probléme et le Secrétariat
devrait évaluer les statistiques commerciales de 1984-85, identifier tous les
pays d'importation qui permettent le commerce avec des pays d'exportation
ayant des interdictions d'exporter et demander que les mesures nécessaires

soient prises pour mettre un terme a ce probléme. La taxonomie devrait étre
clarifiée par le Comité de la nomenclature.

xii

Liste 2 (17 taxons)

Le groupe de travail a recommandé que les taxons suivants recoivent une
attention particuliére en tant qu'espéces ou groupes d'espéces hautement

prioritaires en ce qui concerne la collecte d'informations a leur sujet (par
ordre d'importance):

1. Varanidae asiatiques (trois espéces, soit Varanus salvator et les
espéces V. bengalensis et V. flavescens inscrites A l'Annexe I).

2. Pythons asiatiques (trois espéces, soit Python curtus, P. molurus
bivittatus et P. reticulatus).

3. Varans d'Afrique (deux espéces, soit Varanus exanthematicus’ et
V. niloticus) - a noter que le projet devrait se concentrer sur les
aspects nécessaires 4 1'évaluation des niveaux d'exploitation que l'espéce
peut supporter.

4. Boidae sud-américains (trois espéces, soit Boa constrictor, SEunectes
murinus et E. notaeus).

5. Papustyla pulcherrima.
En outre, le groupe a fait les recommandations spéciales suivantes:

Dracaena guianensis - Les Parties ayant des populations de Dracaena spp.
devraient clarifier la situation en ce qui concerne leur répartition et
envisager l‘inscription de Dracaena spp. a 1'Annexe II.

Crocodylus porosus - I1 serait urgent de demander au Groupe de spécialistes
des crocodiles de la Commission de sauvegarde des espéces de 1'UICN de réviser
le programme de gestion de ces espéces & la lumiére du récent rapport
indonésien.

Liste 3 (15 taxons)

Il a été convenu que les informations disponibles indiquent que ces taxons ne
sont pas au premier chef affectés par le commerce international.

METHODES

Ce rapport comprend l'examen de l'état biologique des espéces contenues dans
la liste "C" et des données commerciales les concernant. Il a été élaboré par
le Centre UICN de surveillance continue de la conservation de la nature, sur
la base d'un contrat avec le Secrétariat CITES, au cours de la période
septembre 1985 - avril 1986. Dans un premier temps, le Secrétariat CITES a
adressé, par l'intermédiaire des organes de gestion CITES des pays Parties a
la Convention ou des autorités responsables de la gestion de la faune ou
équivalentes des pays non-Parties, une demande d'informations 4 tous les pays
dans lesquels se rencontrent les espéces de la liste "C". Les réponses recues
ont été envoyées au CMC et il y est fait référence en indiquant le nom de
l'organe de gestion de la Partie CITES en 1987. Il est fait référence aux
commentaires recus des autorités responsables de la gestion de la faune dans
les pays non-Parties en indiquant le nom du département gouvernemental
intéressé. Des informations ont également été demandées aux spécialistes
(personnes ou organisations), et les groupes de spécialistes de la Commission
de sauvegarde des espéces de 1*UICN en furent parmi les principales sources.
Il a également été fait appel aux organisations du commerce et autres parties
intéressées. Un projet de rapport a été présenté a la deuxiéme session du

xiii

Comité technique CITES en juin 1986. Ce rapport a été examiné et amendé par le
comité et des versions révisées ont été transmises par le Secrétariat CITES
aux Etats de l'aire de répartition et aux personnes intéressées, dont le Pet
Industry Joint Advisory Council. Les modifications finales et des données
commerciales récentes ont été ajoutées au texte, par le CMC, au cours de 1987.

Dans un petit nombre de cas, la catégorie a laquelle une espéce avait été
attribuée lors de la deuxiéme session du Comité technique a, par la suite, été
modifiée sur la base des nouvelles informations recues, en particulier les
données commerciales de 1985 qui ont été ajoutées aux rapports.

Les informations ont été recueillies et rassemblées sous les titres suivants:
répartition; population; habitat et écologie; menaces pour la survie; commerce
international; mesures de conservation; et élevage en captivité.

Les données commerciales CITES ont été analysées pour les années 1980 a 1985,
sur la base des rapports annuels des Parties a& la Convention dont les
statistiques sont conservées sur ordinateur par le CMC. Ces _ données
comprennent les importations et exportations des espéces figurant aux annexes
a la CITES et de leurs produits. Elles contiennent des informations sur les
espéces en question, une description du type de produits et leur quantité et,
dans le cas des importations, mentionnent l'exportateur ou le ré-exportateur
et le premier pays producteur, et, pour les exportations, la destination et la
source d'origine. En ce qui concerne le commerce entre deux pays Parties a la
CITES, chaque transaction devrait donc étre enregistrée deux fois, une fois
par l'importateur et une fois par l'exportateur. Ainsi que le Groupe de
travail sur le commerce important d'espéces de l'Annexe II l'avait suggéré,
l'analyse a été, pour l'essentiel, limitée au commerce des animaux vivants et
aux produits non-travaillés; cependant, dans un petit nombre de cas
exceptionnels, des produits travaillés y ont été inclus.

Divers problémes réduisent la valeur des données commerciales CITES pour
l'évaluation des niveaux du commerce mondial. Par exemple: toutes les nations
faisant du commerce ne sont pas Parties a la CITES; les Parties a la CITES ne
présentent pas toutes des rapports annuels; et les rapports présentés sont de
qualité variable et le sont de maniére irréguliére. Certains pays font état du
nombre de spécimens couverts par les permis émis, tandis que d'autres
indiquent le nombre réel de spécimens pour lesquels le permis a été utilisé.
En outre, il se peut que des exportations ayant lieu en fin d'année arrivent
dans le pays d'importation au début de l'année suivante et, dans de tels cas,
il est possible que la méme transaction soit enregistrée dans les tableaux
relatifs aux données commerciales des deux années. Il s'agit de tenir compte
de ces facteurs, et d'autres encore, dans l'analyse des données de la CITES;
toutefois, pour la plupart des espéces, ces statistiques constituent l'unique
source d'informations détaillées sur leur commerce international, et les
rapports CITES sont en général précieux pour évaluer les niveaux approximatifs
du commerce légal, la répartition géographique des voies empruntées par le
commerce international et les tendances, au cours des ans, en ce qui concerne
le volume du commerce et l'évolution des préférences a l'égard des produits.

Dans la plupart des cas, les données commerciales sont présentées en deux
tableaux dans les exposés qui suivent. Le premier (le tableau 1 en régle
générale) énumére, dans le détail, les importations nettes des pays
d*importation dont le total donne une estimation du volume minimal du commerce
mondial pour chaque année. Le second (le tableau 2 en régle générale) indique
l'origine ou, dans les cas ot l'origine n'a pas été indiquée, 1l'exportateur
des spécimens commercialisés. Lorsque des spécimens ont été exportés vers un
pays intermédiaire et réexportés par la suite, le commerce net minimal est
alors calculé, en s'assurant que les quantités n'ont été enregistrées qu'une
fois. Ainsi, le tableau indique, pour chaque année, la quantité minimale

xiv

d’articles commercialisés a& partir de chaque pays d'origine. Cependant,
certains articles pouvant étre réexportés sans que le pays d'origine ne soit
spécifié, il est possible qu'ils soient enregistrés deux fois dans le
tableau 2. C'est la raison pour laquelle les totaux du tableau 2. sont
généralement plus élevés que ceux du tableau 1.

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INTRODUCCION

Antecedentes

La Convencién Sobre el Comercio Internacional de Especies Amenazadas de Fauna
y Flora Silvestres (CITES) fue elaborada en 1973 con el objeto de controlar el
comercio de vida silvestre. Ese control se efectua asignando a las especies
dos niveles de proteccién. Aquellas especies (0 pequenas poblaciones
geograéaficas) que se encuentran amenazadas de extincién estdn incluidas en el
Apéndice I de la Convencién, y su comercio internacional estdé prohibido,
excepto bajo circunstancias excepcionales. Aquellas especies que no corren
peligro de extincién, pero que podrian estar amenazadas si su comercio no
estuviera reglamentado, se incluyen en el Apéndice II de la Convencién. Dichas
especies pueden comercializarse a nivel internacional, pero las naciones
concernidas deben asegurarse de que los niveles de comercio no representan una
amenaza para las poblaciones silvestres remanentes. Este requisito se explica
formalmente en el texto de la Convencién, Articulo IV, pdrrafo 2 a), que exige
que las autoridades de los paises exportadores informen que la exportacién de
especimenes de esas especies "no perjudicard la supervivencia de esa especie".
En el articulo IV, parrafo 3 se indica que el comercio de esas especies “debe
limitarse a fin de conservarlas, a través de su hdébitat, en un nivel
consistente con su papel en los ecosistemas donde se hallan y en un nivel
suficientemente superior a aquel en el cual esa especie seria susceptible de
inclusién en el Apéndice I". Las autoridades del pais exportador deberan
controlar las exportaciones y tomar medidas para limitarlas cuando asi se lo
estime conveniente.

Durante la cuarta reunién de la Conferencia de las Partes en CITES, realizada
en 1983 en Gaborone, Botswana, se reconociéd que varios paises exportadores de
especimenes de especies del Apéndice II no podian determinar por si solos si
los niveles de comercio perjudicaban a las poblaciones silvestres. Por lo
tanto, se recomendéd (por medio de la Resolucién Conf. 4.7) “que el Comité
Técnico de CITES identifique las especies del Apéndice II que son objeto de un
comercio internacional considerable, para las cuales la informacién cientifica
disponible sobre su capacidad de resistir a tales niveles de comercio resulta
insuficiente como para satisfacer los requisitos estipulados en el Articulo
IV, parrafo 3 de la Convencién, segtin la opinién de los Estados involucrados
en el area de distribucién". Se recomendéd que, una vez que determinadas
especies se hayan identificado, el Comité Técnico, junto con los Estados
involucrados en el 4drea de distribucién, los Estados importadores y las
organizaciones que poseen una experiencia en el manejo de la fauna y de la
flora, "“elaboren y negocien las medidas necesarias para assgurar el
mantenimiento del comercio continuo de esas especies dentro de los limites
previstos en el Articulo IV, parrafo 3, de la Convencién”.

Las discuciones iniciales respecto a la manera como el Comité Técnico
identificaria las especies en cuestiédn (tal como se recomienda en la
Resolucién Conf. 4.7) se basaron en la premisa de que un importante volumen de
comercio era evidencia suficiente como para justificar la preocupacién. Sin
embargo, un informe no publicado, que fue realizado en 1984 por el WTMU para
la Secretaria CITES, llegé a las siguientes conclusiones en lo que se refiere
a la percepcién del problema relativo al volumen significativo de comercio:

- El concepto de volumen significativo de comercio puede definirse de dos
maneras: el volumen significativo puede considerarse en términos absolutos
(i.e. grandes cantidades), o en términos relativos (i.e. grandes
cantidades en relacién con la poblacién y la biologia de la especie).

xvi

- El volumen de comercio significativo absoluto no implica por si solo que
la especie esté amenazada por el comercio. Sin embargo, la
comercializacién de especies en ntmeros significativos absolutos puede
tener un significado ecoldédgico importante.

- El volumen de comercio significativo relativo estdé directamente ligado a
la supervivencia de las especies concernidas, pero no se tienen pruebas de
que esto este correlacionado con el volumen de comercio significativo
absoluto. Debido a su designacién en los Apéndices, todo comercio de
especies incluidas en CITES es de interés y debe ser vigilado.

- Considerar el volumen de comercio significativo absoluto como un criterio
para la seleccién de especies para un cuidado especial es por lo tanto no
solamente irrelevante en términos de conservacién de especies, sino que
puede también distraer la atencidén de casos mds importantes.

El Grupo de Trabajo del Comité Técnico sobre el comercio significativo de
especies del Apéndice II produjo un documento, basado en su reunién en Suiza
en Diciembre de 1984, cuyo fin consistia en formular un procedimiento o una
linea de conducta que permitiera al TEC cumplir con sus obligaciones en virtud
de la Resolucién Conf. 4.7. Se decidid que el Grupo debia limitar sus
discusiones a la fauna, pues ya existia un Grupo de Trabajo para las plantas.
Las conclusiones del informe del WIMU sobre gran volumen de comercio fueron
endosados, y el Grupo convino en que no era posible identificar los taxa del
Apéndice II mas preocupantes basdndose solamente en los datos comerciales.
Para evaluar correctamente el efecto del comercio sobre esos taxa era
necesario poseer informacién sobre la situacién bioldgica, sobre la tendencia
de las poblaciones y sobre toda una serie de otros factores.

Se convino en un procedimiento de cinco etapas como siendo el mecanismo mas
favorable para la aplicacién de la Resolucién Conf. 4.7. Dicho procedimiento
se presenté durante la quinta reunién de la Conferencia de las Partes que se
realiz6 en Buenos Aires, Argentina, en 1985 (Doc. 5.26). Los pasos 1-3 ya han
sido realizados.

lra. etapa: Elaboracién de una lista "A"

Se reconociéd que, salvo algunas pocas excepciones, se puede razonablemente
asumir que un taxén incluido en el Apéndice II puede soportar un cierto
grado de explotacién con fines de comercio internacional. El Grupo decidié
fijar una cantidad a un nivel “prudente” de comercio para todos los taxa
del Apéndice II, en término medio, de menos de 100 ejemplares por ano de
un tax6n incluido en el Apéndice II, que son obtenidos de la naturaleza
(en forma global) y que entran anualmente en el comercio.

De esta forma, eliminando todos los taxa que no estén concernidos por el
comercio internacional o que estdn concernidos solamente a un nivel
minimo, se obtiene una lista de taxa "candidatos potenciales" (lista "A").

Esos taxa se definen como aquellos que podrian ser objeto de un comercio
internacional significativo.

La lista "A" fue preparada por la WIMU, utilizando el promedio de las
estadisticas comerciales CITES ofrecidas por las Partes en el periodo
1980-1982. Se incluyeron en el andlisis los datos relativos a los
especimenes vivos (excluyendo los especimenes criados en cautividad), las
pieles enteras fo) substancialmente enteras, las pieles de los
flancos/lados, las napas de pieles, los caparazones, los trofeos y otros
articulos no trabajados, etc. Las especies que nunca fueron registradas en
el comercio, con excepcién de aquellas incluidas en el Apéndice II como
parte de un taxo6n superior o por razones de semejanza, fueron listadas

XVii

separadamente para que se tomara en consideracién su retiro de los
Apéndices.

2da. etapa: Elaboracién de una lista "B"

El Grupo convino que algunos taxa pueden ser eliminados de las especies de
"comercio significativo" basdndose en los conocimientos disponibles
relativos a su situacién. Luego de este proceso, los taxa remanentes
constituyen la lista "B", formada por aquellos taxa con “posibles
problemas". Ademdés, agregaron a esa lista dos especies (Tupinambis
rufescens y Papustyla pulcherrima) bajo circunstancias especiales,
donde se pone en evidencia un problema, a pesar del bajo volumen de
comercio registrado.

3ra. etapa: Elaboracién de una lista "C"

El paso siguiente del procedimiento consistia en evaluar las informaciones
disponibles para cada una de las especies de la lista "B" y en eliminar
las especies que, sobre la base de la opinidédn de expertos, no presentan
problemas. Esta parte de la operacién significaba tener que reunir el
maximo de informacién posible con respecto a cada especie y evaluar el
efecto del comercio conocido sobre la poblacién conocida. El Grupo convino
que, para cada especie, se debia acordar una importancia primordial a la
situacién global, pero que, si una especie estaba aparentemente afectada
por un comercio a nivel nacional o regional, se lo debia mencionar en un
suplemento anexado a la lista. Las especies de la lista "C" deberian
distribuirse en dos grupos: en primer lugar las especies para las cuales
las informaciones corrientes o el conocimiento de su biologia y/o de su
manejo demuestran que la poblacién se halla afectada por la explotacién
debido al comercio internacional (Lista 1); y, en segundo lugar, las
especies para las cuales las informaciones disponibles o los conocimientos
son insuficientes como para servir de base a un juicio de ese tipo (Lista
7

4ta. etapa: Elaboracién de medidas correctivas

El TEC, o un grupo de trabajo del TEC constituido a ese efecto, debia
examinar las listas "1" y "2", y establecer prioridades dentro de cada
lista. Para las especies o grupos de especies de la lista “1" de gran
prioridad, se debian convocar sesiones de trabajo con el objeto de
recomendar medidas correctivas. Las medidas correctivas examinadas debian
comprender, sin necesariamente limitarse a esto: la preparacién de
propuestas para transferir las especies en cuestion al Apéndice I, la
elaboracioén de procedimientos de manejo suplementarios ya sea en favor de
las poblaciones silvestres (tales como cupos de caza, temporadas de caza,
tamanos limites de los especimenes, etc.) o bien en lo que se refiere a
los controles del comercio, y la inclusién de taxa por razones de
semejanza.

Para las especies de la lista "2", de gran prioridad, se deberian
establecer proyectos con el objeto de recabar informacién sobre su
biologia y manejo. Cuando esas informaciones demuestren la necesidad, la
especie deberia transferirse a la lista "1".

Sta. etapa: Aplicacién de las medidas correctivas
Las medidas correctivas deberian ser desarrolladas por los Estados del

&rea de distribucién concernida, sobre la base de las recomendaciones
formuladas en las sesiones de trabajo.

xviii

Este procedimiento de cinco etapas fue aprobado en la reunién de Buenos Aires
en 1985 y las etapas 1-3 ya fueron desarrolladas por el Centro UICN de
Vigilancia Continua de la Conservacién. La lista "Cc" fue preparada a tiempo
para la segunda reunién del Comité Técnico realizada en Junio 1986 en
Lausanne, Suiza. Para cada especie incluida en la lista "C", se preparé un
borrador presentando un resumen de toda la informacién disponible, incluyendo
un andlisis detallado de referencias e informacién disponible sobre el
comercio y sobre el estado de la poblacién y otros factores que se
consideraron importantes. Basado en esta informacién, cada especie fue
asignada a las dos listas sugeridas (lista 1, especies con problemas; lista 2,
problemas posibles). En esta etapa se descubrié también que era posible que
algunas especies, originalmente incluidas en la lista "C", no se vieran
afectadas en forma significativa debido a los .presentes niveles de comercio.
Dichas especies fueron incluidas en un tercer grupo (lista 3, sin problemas).
El Grupo de Trabajo del Comité Técnico sobre el comercio significativo de
especies revisé la informacién proporcionada por el CMC, asi como los listados
presentados, y preparé recomendaciones para una accién ulterior, las cuales se
ennumeran a continuacién. El Comité Técnico decidiéd asimismo que, después de
revisién ulterior, el informe preparado por el CMC debia ser publicado.

Accién ulterior

El Grupo de Trabajo sobre el Comercio Significativo de Especies presenté un
documento durante la segunda reunién del Comité Técnico en el que se
delineaban propuestas para acciones ulteriores (WGR.TEC. 2.2). A continuacidén
se describen las recomendaciones de dicho informe para las especies de
reptiles concernidas, las cuales fueron modificadas durante la reunién del
Comité Técnico.

Lista 1 (4 taxa)

Podocnemis expansa - Cabe suponer que los datos relativos al comercio son
sumamente exagerados como consecuencia de una identificacién errdédnea. La
Secretaria deber& informar a las Partes acerca de las prohibiciones vigentes
sobre el comercio y Venezuela se encargardé de preparar una propuesta para la
transferencia de la especie al Apéndice I teniendo en cuenta de que el
comercio internacional es’ probablemente insignificante.

Caiman crocodilus - Para tratar este problema, la Secretaria estdé realizando
proyectos en Bolivia, Brasil y Paraguay. Se necesita financiacién para un
segundo proyecto (Brasil, Colombia y Venezuela). Se deberian realizar estudios
sobre la poblacién en América Central. Deberia recolectarse y analizarse
informacién sobre antecedentes en el comercio, incluyendo cantidades y tamano
de las pieles,. En los lugares en los que las especies sean legalmente

explotadas, deberdén establecerse cupos nacionales para la extraccién del medio
silvestre.

Geochelone chilensis - Se considera apropiado el plan de manejo que ha
asumido Argentine.

Tupinambis spp. - Argentina ha iniciado una accién para revisar sus cupos de
extracciédn del medio silvestre. Es posible que el comercio ilegal siga siendo
un problema y la Secretaria deberd& evaluar las estadisticas relativas al
comercio en 1984/1985, identificar cualquier pais importador que autorice el
comercio a partir de paises exportadores donde existe una prohibicién sobre
las exportaciones, y pedir que se tomen las medidas del caso a fin de poner

término al problema. El Comité de Nomenclatura deberé aclarar el aspecto
taxondémico.

xix

Lista 2 (17 taxa)

El Grupo de Trabajo recomendé que los siguientes taxa (enumerados por orden de
importancia) se consideraran como especies o grupos de especies prioritarios:

Ae Varanidos asidticos (tres especies, es decir, Varanus salvator y las
especies del Apéndice I V. bengalensis y V. flavescens).

7s Pitones asidticos (tres especies, es decir, Python curtus, P. molorus
bivittatus y P. reticulatus).

I Lagartos Africanos (dos especies, es decir, Varanus exanthematicus y
Varanus niloticus), haciendo notar que el proyecto deberd concentrarse
en los aspectos necesarios para evaluar los niveles de explotacidén
sostenidos para las especies.

4. Boas sudamericanas (tres especies, es decir, Boa constrictor, Eunectes
murinus y EBunectes notaeus).

Sie Papustyla pulcherrima.
Ademés, el Grupo formulé las recomendaciones especiales siguientes:

Dracaena guianensis: Las Partes que poseen poblaciones de Dracaena spp.
deberén aclarar su situacién en lo que se refiere a la distribucién y
considerar la inclusién de Dracaena spp. en el Apéndice II.

Crocodylos porosus: Deber& recomendarse con urgencia al Grupo UICN/CSE una
revision del programa de manejo para esta especie a la luz de los resultados
del reciente informe de Indonesia.

Lista 3 (15 taxa)

Se acordéd en que la informacién disponible indicaba que esos taxa no estaban
esencialmente afectados por el comercio internacional.

METODOS

Este informe incluye la revisién del estado biolédgico y comercial de especies
que aparecen en la lista "C". Este informe ha sido realizado por el Centro
UICN de Vigilancia Continua de la Conservacién, bajo contrato con la
Secretaria CITES, cubriendo el periodo Septiembre de 1985 a Abril de 1986.
Como paso inicial, la Secretaria CITES circuléd, a traves de las Autoridades
Administrativas CITES de los Estados miembros en la Convencién, o a traves de
las Autoridades Administrativas responsables de fauna u otras autoridades
equivalentes en los estados no Partes en la Convencién, una solicitud de
informacién a todos los paises en los que se encuentran las especies de la
lista "C". Los comentarios recibidos fueron enviados a la CMC y se
clasificaron de la siguiente manera: Nombre del pais de la Autoridad
Administrativa CITES, 1987. Los comentarios recibidos de las autoridades
responsables de los Estados no Partes fueron clasificados por nombre de la
autoridad gubernamental concernida. También se solicitéd informacién de
especialistas concernidos (personas o agencias), y entre las fuentes
principales se encontraban los grupos de especialistas de la Comisién de
Supervivencia de Especies de la UICN. También fueron consultadas algunas
organizaciones comerciales y otras Partes interesadas. Un informe borrador se
presenté en la segunda reunién del Comité Técnico CITES en Junio de 1986. Este
informe fue discutido y corregido por el Comité y las copias, una vez
revisadas, fueron nuevamente enviadas por la Secretaria CITES a todos los
paises concernidos y a las partes interesadas, incluyendo el Pet Industry
Joint Advisory Council. Las modificaciones finales al texto, asi como la

xx

informacién sobre el comercio reciente, fueron incluidas por el CMC durante
1987.

Por lo tanto, en la minoria de los casos, la designacién de la categoria de
una especie al realizarse la segunda reuniédn del Comité Técnico ha sido
modificada a la luz de nueva informacién, en particular la informacién
comercial de 1985 que ha sido agregada a los informes.

Se recolecté6 e incluyé6 la informacién bajo los siguientes titulos:
distribucién; poblacién; habitat y ecologia; amenazas a la supervivencia;
comercio internacional; medidas de conservacién; y cria en cautividad.

Los datos sobre el comercio CITES fueron analizados para los afos 1980 a 1985,
utilizando los Informes Anuales de las Partes de la Convencién, cuyas
estadisticas han sido procesadas en el computador del CMC. Esta informacién
incluye el registro de importaciones y exportaciones de especies de los
Apéndices de CITES, asi como sus productos, y'contienen informacién sobre las
especies concernidas, una descripcién del tipo y la cantidad del producto, y,
en el caso de importaciones, el exportador o re-exportador y los principales
paises de origen, y, para las exportaciones, el destino y la fuente de origen.
En lo que concierne al comercio entre dos Partes en CITES, cada transaccién
deberia por lo tanto registrarse dos veces: una vez por el importador y otra
por el exportador. Tal como sugirié el Grupo de Trabajo sobre el Comercio
Significativo, el andlisis se restringid al comercio de animales vivos o de
productos no trabajados, sin embargo, también se incluyeron productos
terminados en un numero pequenho de casos excepcionales.

Varios problemas reducen el valor de la informacién comercial de CITES en la
evaluacién de los niveles del comercio mundial. Por ejemplo: no todas las
Maciones que realizan comercio son Partes en CITES; no todas las Partes en
CITES elaboran informes anuales, y la presentacién de los informes varian en
calidad y regularidad. Algunos paises pueden proporcionar informacién sobre
lacantidad de especimenes que cubren los permisos expedidos, mientras que
otros proporcionan informacién sobre la cantidad real por la cual se utilizé
el permiso. Mas atn, las exportaciones de un pais al finalizar un ano pueden
arrivar al pais importador al comienzo del afio siguiente, y en tales casos es
posible que, por la misma transaccién, se registren en los cuadros comerciales
para ambos anos. Estos factores y otros deben tomarse en cuenta cuando se
analizan los datos de CITES, pero para la mayoria de las especies, estas
estadisticas representan la tnica fuente detallada de informacién respecto a
su comercio internacional y generalmente los informes CITES son de gran
utilidad al evaluar los niveles aproximados de comercio legal, asi como los
patrones geograéficos en tal comercio y las tendencias relativas a los
volumenes de productos preferenciales, en un determinado lapso de tiempo.

En la mayoria de los casos, los datos comerciales son presentados en los dos
cuadros siguientes. En el primero (normalmente Cuadro 1), se detallan las
importaciones netas de paises importadores, cuyo total nos proporciona una
cifra estimada del volumen minimo de comercio mundial anual. El segundo
(normalmente Cuadro 2) muestra el origen, o en los casos en los que el origen
no se menciona, el exportador de los especimenes en cuestién. Cuando los
especimenes han sido exportados a un pais intermediario y posteriormente
reexportados, el comercio minimo neto ha sido calculado, asegurdndose de que
los nuimeros sélo fueron registrados una sola vez. Por lo tanto, el cuadro
muestra, anualmente, la cantidad minima de articulos de comercio de cada pais
de origen. Sin embargo, ya que algunos articulos pueden ser reexportados sin
que necesariamente aparezca especificado el pais de origen, éstos pueden ser
registrados dos veces en el Cuadro 2. Por lo tanto, los totales son
usualmente mas altos que los que aparecen en el Cuadro 1.

xxi

CHACO TORTOISE Recommended list: 1
[Problem]
Geochelone chilensis (Gray, 1870)

Order TESTUDINES Family TESTUDINIDAE

SUMMARY AND CONCLUSIONS A medium size terrestrial tortoise, restricted to
rather arid lowlands, mainly in the Chaco region, in southern South America.
Ranges from Paraguay and possibly adjacent Bolivia through much of Argentina.
Is classified as Insufficiently Known in the IUCN Amphibia-Reptilia Red Data
Book. Mainly herbivorous. Clutch comprises up to six eggs. Two new species
have recently been described from within the range of G. chilensis sensu
lato; these are not widely recognised and may not be distinct species,
although new information suggests that the large, montane form
(donosobarrosi) may be sympatric with G. chilensis.

Central and northern populations in Argentina are reportedly in marked decline
due to over-exploitation for the international live animal trade. Large
numbers have been exported through Bolivia mainly to U.S.A. From 1980 to
1984, the number in reported trade increased very sharply from 5 to 8111, but
declined again in 1985.

Field study on population levels and the sustainability of the reported and
alleged unreported trade is required, as is clarification of the claimed
ranching or farming operations in Argentina

DISTRIBUTION Rather widespread in the dry lowlands of central South
America, centred on the Chaco zone; extends from Paraguay south through much
of north and central Argentina, into the northern fringes of Patagonia.

One authority (Freiberg, 1973, 1981) regards tortoises in the western part of
the Argentinian Chaco, around Santiago del Estero and La Rioja, as a full
species G. petersi, and those from the Patagonian portion of the range,
between La Pampa (37.3°S) and the Rio Negro (41°S), as G. donosobarrosi.
Most other authorities have preferred to retain the conventional treatment
pending further study. It has recently been proposed (Bour, 1980) that
several taxa usually recognised as subgenera of Geochelone should be
elevated to generic rank, as Chelonoides in the case of the chilensis
complex. This usage is not widespread.

Argentina Rather widely distributed, from the Chaco zone in the north,
south to around 41°S; recorded from the provinces of Buenos Aires, Catamarca,
Cordoba, Chaco, Chubut, Formosa, La Pampa, La Rioja, Mendoza, Neuquen, Rio
Negro, Salta, San Juan, San Luis and Santiago del Estero (Auffenberg, 1969,
Freiberg, 1981; N. Scott, in litt. 23 December 1981). Waller (1986) listed
known localities and estimated that the distribution covered 29-30% of the
continental land area of the country.

Bolivia Although no published records exist, the species “undoubtedly”
(Auffenberg, 1969) occurs in the Gran Chaco of southern Bolivia.

Paraguay Widely distributed in the Chaco zone of north-west Paraguay;
recorded from the departments of Boqueron, Chaco, Nuevo Asuncion and
Presidente Hayes (N. Scott, in litt. 23 December 1981). Waller (1986)
listed two records from Boqueron.

Uruguay The species has been reported to occur in Uruguay (Honegger, 1981),
but there appear to be no published records from this country.

Geochelone chilensis

POPULATION Reportedly in marked decline in the main part of the range,
comprising the Argentinian Chaco, apparently relatively secure in Paraguay.

Argentina Northern and central populations reported to be declining
severely, but southern populations ('G. donosobarrosi' ) appear’ secure,
although existing in low density (Freiberg, 1974, 1981). In the late 1960s a
maximum density of 15-20 G. chilensis per acre were recorded in optimum
habitat in Cordoba (Auffenberg, 1969). In general, reported to be not so
widespread and abundant as often said, and to be patchy in distribution (J.M.
Cei, in litt. to F.W.King, 14 June 1978). Gruss and Waller (1986) cited no
evidence of population decline but claimed that habitat destruction was a
major threat and had caused a severe reduction in range.

Bolivia No information.

Paraguay Reasonably abundant in much of the range, the majority of which is
difficult of access; no evidence for decline (N. Scott, in litt. 23 December
1981).

HABITAT AND ECOLOGY A medium-size terrestrial tortoise, inhabiting arid
lowlands, most typically in thorny chaco habitats (Auffenberg, 1969;
Pritchard, 1979). Population density at one site, near La Posta, Cordoba
(Argentina) was between 15-20 tortoises per acre. Although the species ranges
into semi-arid intermontane basins in western Argentina, densities are highest
in the chaco region. Mainly herbivorous, consuming the fruit of various trees
and shrubs, pads and fruit of cacti, and grasses (Auffenberg, 1969). Shallow
pallets are excavated, deep enough to cover the anterior third or half of the
shell, in which nights and much of the day are spent; a somewhat deeper pallet
is formed for shelter during cold and dry periods. Each tortoise has an
activity range of about 30 m from the pallet being used. Courtship occurs in
November-December and a clutch of 1 to 4 round white eggs, c 4.5 cm diameter,
is laid in January; there is some evidence that two clutches of 1-3 eggs may
be laid in one year (Auffenberg, 1969).

THREATS TO SURVIVAL Threatened by heavy exploitation for the live animal
trade, by increasing habitat destruction owing to timber extraction and
conversion of forest and scrub to agricultural land (Salas, 1985; Gruss and
Waller, 1986); local utilisation as a food source may also have an adverse
effect in conjunction with these factors. These activities seem to be most
prevalent in the province of Santiago del Estero, said to show more severe
habitat destruction than any other province, and a centre for the wildlife
trade (Salas, 1985). Tortoises collected that are not sold for the wildlife
trade may be eaten by local inhabitants, although the meat is not especially
favoured (Salas, 1985). Said to be exported to Santiago (Chile) for food use
(Auffenberg, 1969). In the 1970s very large numbers of young and adults were
sold or exported, about 4000 tortoises monthly in Mendoza Province. Most of
these animals were sent to Chile and other countries, with an estimated
mortality in transit of around 70% (J.M. Cei, in litt. to F.W.King, 14 June
1978). Gruss (1986) conducted a questionnaire survey in Buenos Aires which
concluded that 63% of the population had bought or owned G. chilensis as
pets. He estimated that this could account for an offtake of 75 000 tortoises
from the wild each year to supply the demand in the city and its surroundings.

INTERNATIONAL TRADE Reportedly “tens of thousands" of G. chilensis are
collected annually in Argentina for the national and international pet trade
(Salas, 1985). In the north and centre of Argentina G. chilensis (including
G. petersi) are collected, taken to Buenos Aires and then exported in
thousands (with other wildlife) via Bolivia to USA (notably Los Angeles)

Geochelone chilensis

(Freiberg, 1981). A number of ‘hatcheries' or farms of some description exist
which claim to supply captive-bred tortoises to the trade; it is alleged that
most such operations exist simply as a cover for massive collection from the
wild (Salas, 1985). European trade in live G. chilensis may increase in the
future as EEC countries prohibited the import of European Testudo species in
1984 (Honegger, 1984), although no such increase was apparent in 1985. The
numbers of chilensis in trade reported to CITES is not as large as the
overall numbers said to be entering the pet trade (Salas, 1985; M.A. Freiberg,
in litt. 16 March 1981); this implies that there is an extensive unreported
trade. The number of live animals in reported trade increased very steeply
between 1980 and 1985, but then fell again in 1985.

Table 1. Apparent minimum net imports of live G. chilensis reported to
CITES, 1980-85 (including one shell imported to USA).

1980 1981 1982 1983 1984 1985
a EE EE EE EEE ee
Bermuda - - ~ 6 - -
Canada - - - - 15 -
Chile - - - ~ 50 -
Denmark - - - 6 10 6
Germany, F.R. - 195 - 2 570 ~
Japan - 42 - 361 72 ~
Netherlands - ~ ~ - 162 ~
Switzerland - 32 - - - -
UK = 24 = = = =
USA 5 540 1430 2653 7225 647
USSR - - - - 7 6
TOTAL 5 833 430 3028 8111 659
a a Ns ee a eS
Table 2. Reported countries of origin (or exporting country if no original

source reported) and quantities of transactions in live G. chilensis
reported to CITES.

1980 1981 1982 1983 1984 1985

a

a. Countries having or possibly having wild populations of G. chilensis

Argentina 4 215 1400 2928 8111* 655*
Bolivia 1 7 - 100 - -
Chile - - - - - 4
Paraguay - 611 30 - - -

*2951 of these said to be captive bred in 1984 and 530 in 1985.
b. Countries without wild populations

U.S.A. - - - - 6 -

ee

Geochelone chilensis

CONSERVATION MEASURES Existing laws should be adequately enforced and
international trade restricted. The taxonomic status of '‘G. petersi' and
'G. donosobarrosi' requires investigation.

Argentina All tortoises are protected from hunting and trade in Argentina
(Ley No. 22.421), but this legislation is said not to be enforced (Freiberg,
1974).

Bolivia A ban on the export of all wildlife was imposed in 1984 (Resolucion
Ministerial No 85/84). This was extended in 1985 to cover all wildlife
products, with very few exceptions (Resolucion Ministerial No 2262/85).

Paraguay Export of wildlife from Paraguay is prohibited except under
permit; reportedly (N. Scott, in litt. 23 December 1981) permits are now
issued only for scientific specimens.

Uruguay All hunting and trade of wild animals, with very few exceptions, is
prohibited in Uruguay under Ley No. 9.481.

CAPTIVE BREEDING A number of operations said to be ‘commercial breeders’ of
G. chilensis exist in Argentina (Salas, 1985). A recent inventory (Slavens,
1985) reports 23 chilensis in 8 collections responding to a questionnaire.
Captive breeding prospects are reportedly poor; survival rate in captivity is
very low, due to distinct environmental requirements (Honegger, 1979).

REFERENCES

Auffenberg, W. (1969). Land of the Chaco Tortoise Geochelone chilensis.
International Turtle and Tortoise Society Journal 3(3): 16-19, 36-37.

Bour, R. (1980). Essai sur la taxinomie des Testudinidae actuels (Reptilia,
Chelonii). Bulletin du Muséum National d'Histoire Naturelle, Paris. 4
sér., 2, section A, No 2: 541-546.

Freiberg, M.A. (1973). Dos nuevas tortugas terrestres de Argentina. Boletin
de la Sociedad de Biologia, Concepcion, Chile, 46: 81-93.

Freiberg, M.A. (1974). The Argentine Land Tortoise, Geochelone chilensis,
an endangered species. Bulletin of the Maryland Herpetological Society
10(2): 39-41.

Freiberg, M.A. (1981). Turtles of South America. T.F.H. Publications, Inc.
Ltd., Hong Kong.

Gruss, J. (1986). Geochelone chilensis en cautividad. Descripcion. Amphibia
& Reptilia (Conservacion) 1(2): 29-35.

Gruss, J. and Waller, T. (1986). Un jugueta amenazado. Vida Silvestre 5(19):
28-32.

Honegger, R. (1979). Red Data Book, Vol. 3, Amphibia-Reptilia. IUCN, Gland
(third edition, partly revised).

Honegger, R. (1981). Family Testudinidae. In, CITBS Identification Manual,
Vol. 3: amphibia, reptilia, pisces. CITES Secretariat, Geneva.

Honegger, R. (1984). In litt., 4 June, to D. Jelden. Remarks on the
situation of Geochelone chilensis and G. carbonaria in Argentina.

Pritchard, P.C.H. (1979). Encyclopedia of turtles. T.F.H. Publications,
New Jersey and Hong Kong, 895 pp.

Salas, S. (1985). Project proposal: determination of the status and
exploitation of the Chaco Tortoise, Geochelone chilensis (unpublished).
Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity,

current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

Waller, T. (1986). Distribucion, habitat y registro de localidades para

Geochelone chilensis (Gray, 1870) (Syn. donosobarrosi, petersi)
(Testudines, Testudinidae). Amphibia & Reptilia (Conservacion 1(2):
36-48.

LEOPARD TORTOISE Recommended list: 2*
[Potential problem]

Geochelone pardalis (Bell, 1828)
Order TESTUDINES Family TESTUDINIDAE

*The category has been changed from 3 to 2 since approval by the CITES TEC
Meeting in 1986 owing to the increased trade reported in 1985.

SUMMARY AND CONCLUSIONS A widely distributed species occurring over much of
southern and eastern Africa from Sudan, Ethiopia and Somalia south through
Kenya, Uganda, Tanzania, Mozambique, Malawi, Zambia, southern Angola,
Zimbabwe, Botswana, Namibia to South Africa. Two subspecies have been
recognised but their validity is disputed. Likely to be relatively rare in
easily accessible areas, although no data are available on population size.
An inhabitant of coastal plains and upland savanna, this species is primarily
herbivorous. Breeding takes place in spring in South Africa and over a longer
period elsewhere. Up to thirty eggs are laid. Threats to this species appear
to be hunting for food (eggs and meat) and possibly capture for the pet
trade. It breeds readily in captivity.

The apparent minimum trade volume reported to CITES for the period 1980-1985
was 3392 specimens, virtually all live animals. Tanzania was the major
exporter in 1985, taking the place of Kenya which predominated in 1980 and
1981. F.R. Germany and the USA were the principal importers. Trade increased
sharply in 1985, mostly because of exports from Tanzania to F.R. Germany and
the USA. The species is theoretically protected in Tanzania.

It seems unlikely that the overall trade levels reported to CITES pose a
significant threat to the species, but the apparent sudden rise in illegal
exports from Tanzania, and to a lesser extent Kenya, is of concern. They may
be in response to the prohibition on imports of European tortoises to the EEC
imposed in 1984.

DISTRIBUTION Widely distributed over much of southern and western Africa,
having been reported from Sudan, Ethiopia and Somalia south through Kenya,
Uganda, Tanzania, Mozambique, Malawi, Zambia, southern Angola, Zimbabwe,
Botswana, Namibia to South Africa, possibly including Lesotho and Swaziland
(Loveridge and Williams, 1957; Greig and Burdett, 1976). Two subspecies are
recognised, although Greig and Burdett (1976) doubt their validity. However,
Pritchard (1979) was of the opinion that further study would lead to naming of
more such taxa.

G. pardalis pardalis Now only known from Namibia, though it may previously
have occurred in South Africa. Although there are reports to the contrary,
it does not occur in Angola (Loveridge and Williams, 1957). This subspcies is
not thought to be distinct from G. p. babcocki (South African CITES MA,
1987).

Namibia Probably only south of the line from near Luderitz Bay to
Keetmanshoop (Pritchard, 1979; Loveridge and Williams, 1957). It may,
however, occur as far north as Rehoboth (Loveridge and Williams, 1957).

South Africa In the past it probably also occurred within South Africa,
possibly as far south as the Cape of Good Hope, and around the Oliphants River
(Loveridge and Williams, 1957).

Geochelone pardalis

G. p. babcocki More widely distributed and occurs from Sudan, Ethiopia and
Somalia, southward through Kenya, Uganda, Tanzania, Mozambique, Malawi,
Zambia, Zimbabwe, Botswana, Angola, to Namibia and South Africa. Loveridge
and Williams (1957) refuted reports from Madagascar and the Senegambia region.

Angola Recorded inland from Benguela, Capangombe, Cubal, Dombodola, Forte
Rocadas, Mopa and the interior of Mossamedes. Absent from the north of the
country (Loveridge and Williams, 1957).

Botswana Recorded from Gemsbok, Kalahari, Magalapsi, Makarikari, Maun,
Serowe, Shaleshonto, Shorobe, Tsotsoroga Pan and Mababe Flats (Loveridge and
Williams, 1957; Greig and Burdett, 1976).

Ethiopia Recorded from the south of the country at: Ado-Shebeli Valley,
Araro, Arenda, Artu, Arussi Gallaland, Boorgha, Caschei, Dabas, En, Erre,
Javello, Lake Abaya, Lake Zwai, Neghelli, Shoa, Sibi, Harrar, Soddu,
(Loveridge and Williams, 1957).

Kenya Recorded from numerous localities from around the country (Loveridge
and Williams, 1957). Widespread in drier savannah, though not deserts (A.
Mackay, in litt., 26 March 1986).

Lesotho The species is said to occur in Lesotho (Ministry of Agriculture
and Marketing, Lesotho, in Jlitt., 1985), although Greig and Burdett (1976)
located no records, and Branch and Brack (1987) indicate that it may be absent.

Malawi Occurs south of the Mwanza River and west of the Shire in both
Chikwawa and Port Herald Districts (Loveridge and Williams, 1957).

Mozambique Recorded from Cape Delgado, Kasumbadedza, Sena and _ Tete
(Loveridge and Williams, 1957). The species is said to be widespread
throughout the country (Mozambique CITES MA, 1986).

Namibia Recorded from several localities, it meets with G. p. pardalis in
southern Namibia, but is dominant north of 25°S (Loveridge, 1957).

Somalia Recorded from Hargeisa, Nogal Valley, Toyo Plain, between Bardale
and Condut, Pozzi-Ircudt, Jet, "Sahaaieroi” (Loveridge and Williams, 1957).

South Africa Recorded from Natal, Transvaal, and numerous localities in the
Cape Province (Loveridge and Williams, 1957). There are no records from the
Western Cape, the central Cape Karroid areas, the highveld regions of the
Orange Free State, southern Natal or Transkei (Greig and Burdett, 1976; Branch
and Braak, 1987). Transport and release of pets around the country regularly
occurs and results in their occurrence in semi-urban areas (Greig and Burdett,

1976). This may mask any local genetic variation (South Africa CITES MA,
1987).

Sudan Recorded from the south of the country at: Bahr el Ghazal, Bahr el

Jebel, Bor, Gondokoro, Lado Enclave and Mongalla (Loveridge and Williams,
1957).

Swaziland Recorded as being included in the distribution (Groombridge,

1983), but no records have been located although it occurs very close to the
border (Greig and Burdett, 1976).

Tanzania Recorded from numerous localities from around the country

(Loveridge and Williams, 1957). Said to be widespread (K. Howell, in litt.,
15 March 1986).

Geochelone pardalis

Uganda Recorded from Mount Debasien (Loveridge and Williams, 1957). Said

to occur widely throughout the country (Uganda Game Department, in Jlitt.,
1987).

Zambia Recorded from Luangwa River and Pitauke (Loveridge and Williams,
1957). Said to be widespread throughout the country but absent above 1 830 m
on the Nyika Plateau (Wilson, 1968).

Zimbabwe Recorded from Birchenough Bridge, Sabi River, Gwamayaya River and
Mount Darwin (Loveridge and Williams, 1957; Greig and Burdett, 1976).

POPULATION Little information is available regarding the current status of
the populations of this species. Loveridge and Williams (1957) did not
provide evidence to suggest that G. p. babcocki was rare, indeed they quote
Peel's (1900) observation of “an army of monsters migrating across the
desert". Pritchard (1979) reported that, in easily accessible areas, Leopard
Tortoises may be relatively rare.

Angola No information.
Botswana No information.

Ethiopia No information.

Kenya The species is widespread, but never occurs in large numbers
(A. Mackay, in litt., 26 March 1986).

Malawi No information.

Mozambique The species is said to be abundant in Gorongosa National Park
and the Gilé Game Reserve, common in Banhine and Zinave National Parks and
Niassa Game Reserve, and rare in Maputo and Marromeu Game Reserves. It has
declined and is now rare in cultivated areas. As the species is actively
hunted for food (Mozambique CITES MA, 1986), it is likely to have come under
severe pressure in the recent famine.

Namibia Said to be abundant in Etosha and fairly abundant in other
conservation areas in the north-east (Bushmanland, Kavango, etc.). It is
locally extinct in all communal areas (Department of Agriculture and Nature
Conservation, Namibia, in litt., 1987).

Somalia No information.

South Africa Said to be very common in the eastern Cape Province, and the
lowveld regions of Transvaal, but scarce elsewhere (Branch and Brack, 1987).

Sudan No information.

Tanzania Not considered threatened in Tanzania (K. Howell, in Jlitt., 15
March 1986).

Uganda No information.
Zambia Said to be common throughout the country (Wilson, 1968).
Zimbabwe Broadley (in Jlitt., 18 March 1986) considered this tortoise to

be scarce near densely populated communal lands but common in sparsely
inhabited areas.

Geochelone pardalis

HABITAT AND ECOLOGY Leopard Tortoises inhabit coastal plains and upland
savanna and may be found in rather sandy, thornbush steppes and on kopjes and
stone strewn hills with scattered scrub. Not found in of primary forest.
G. p. babcocki is often found on mountain masses, and has been recorded at a
height of 9 000-10 000 feet on Mt. Elgon, Uganda (Loveridge and Williams,
1957). The diet includes grasses, a variety of succulents, also fallen fruit
and crop plants such as beans and watermelons (sources in Pritchard, 1979).
The species can apparently exist without water for long periods.

Males compete for females in the breeding season by pushing and butting.
Nesting takes place in spring in South Africa (September-October) but the
season is longer in tropical Africa. Nest holes vary in size to a maximum
recorded depth of 1 foot and may contain up to 30 eggs. These hatch in 8-18
months depending on the weather conditions (Loveridge and Williams, 1957).
Leopard Tortoises appear to have a well developed homing ability and have been
observed moving en masse (Peel, 1900 in Loveridge and Williams, 1957;
Pritchard, 1979).

THREATS TO SURVIVAL Flooding and grass fire have both been identified as
the cause of deaths of large numbers of leopard tortoises They are preyed on
by a variety of animals - crows, ground hornbills, ratels, hyenas and soldier
ants, and eggs may be dug up and devoured by jackals, dogs, and viverrids
(Loveridge and Williams, 1957). The food value of G. p. pardalis may have
led to its extermination in parts of the Cape (Loveridge and Williams, 1957).
Bushmen also prize G. p. babcocki for its shell which may be used for
ladling or holding water and also to make snuffboxes (Fitzsimmons, 1935 in
Loveridge and Williams, 1957).

Botswana Auerbach (1985) reports that this species is prized by man for its
meat, eggs and shell.

Kenya In Kenya generally the species seems to little persecuted, if at all,
and is often accorded a respectful title, such as ‘tortoise elder' (A. Mackay,
in litt., 26 March 1986).

Lesotho There is said to be no trade in any wild fauna in Lesotho (Ministry
of Agriculture and Marketing, Lesotho, in litt., 1985).

Mozambique The species is not traded but is said to be actively hunted for
food and to be particularly depleted by fires which are common in cultivated
areas for bush clearance (Mozambique CITES MA, 1986).

Namibia The species has been hunted for food to the point of extinction in
all communal areas. Hunting also occurs to a lesser extent in other areas
(Department of Agriculture and Nature Conservation, Namibia, in litt., 1987).

South Africa Greig and Burdett (1976) reported that the meat and eggs of
G. pardalis were highly regarded in some areas; and attributed its absence
from the western Cape to past persecution. Branch and Braak (1987) suggested
that the absence from southern Natal and Transkei might be similarly due to
human predation. They reported that one tortoise had died after becoming
trapped in an Aardvark hole.

Tanzania GC. pardalis is said not to be persecuted in Tanzania, though

increasing numbers are exported for the live pet trade (K. Howell, in litt.,
15 March 1986).

Uganda Some tribes consider this species a delicacy, mainly in the north of
the country and Karamoja, but there is little other persecution. No trade has
been reported (Game Department, Uganda, in ditt., 1987).

8

Geochelone pardalis

INTERNATIONAL TRADE Listed on CITES Appendix II. The principal data
available relating to international trade are those contained in the Annual
Reports of the Parties to CITES. These are summarized in Tables 1 and 2.

Table 1: Minimum net imports of live Geochelone pardalis reported to CITES.

1980 1981 1982 1983 1984 1985
Australia - - 6 * = = =
Austria - 4 - - - 4
Canada - - - - ~ 4
Denmark 4 2 - - - -
Dominica 4 = = = = =
Dominican Rep. - 1 - ~ - -
German D.R. 10 - ~ - - 2
Germany, F.R. 345 9 2 - - 1009
Israel - - - - - 1
Japan - - ~ 10 12 71
Namibia - - ~ - 1 -
Switzerland 82 260 - - - 16
UK 1 34 - 4 - 3
USA 376 413 4 81 35 558
Total 822 723 12 95 48 1675
Table 2: Reported countries of origin (or exporting country if no original

source reported) and quantities of export of live Geochelone pardalis
reported to CITES.

1980 1981 1982 1983 1984 1985

Countries having or possibly having wild populations of G. pardalis

Kenya 736 617 - - 41 100
Namibia 66 10 - - - -
Somalia - - - - - 2
South Africa - 2 Ox 34 * 6 * -
Tanzania - ~ - 150 - 1497
Uganda - 2 - - - -
Zimbabwe - - - - - 18

Countries without wild populations of G. pardalis

Australia 4 = = = = =
Germany, F.R. - - - = = 1*
Country unknown - - 3 = 1 =

* Captive-bred

Over 99% of all imports were live specimens, presumably mostly destined for
the pet trade. The principal importers of G. pardalis were the USA and F.R.
Germany. Imports dropped substantially in the years 1982, 1983 and 1984, but
rose to a peak in 1985.

Geochelone pardalis

Kenya was the principal exporter until 1981 when an export ban was imposed,
after which few exports were indicated. In 1983 and 1985, Tanzania was the
source of the great majority of G. pardalis. South Africa principally
exported captive-bred specimens. F.R. Germany re-exported a large number of
animals notably to the GDR and Switzerland in 1980 and 1981.

The precise effects of the trade in this species cannot be fully assessed due
to lack of population data. It seems unlikely that the total reported trade
could cause significant long-term depletion of the species, or of many local
populations. However, the recent rise in exports reported from Tanzania is of
potential concern, partially because such exports are theoretically prohibited
and partially because they may be a response to the EEC prohibition on import
of European tortoises (q.v.) imposed in 1984. To a lesser extent, exports
from Kenya also appear to have risen in 1984 and 1985.

CONSERVATION MEASURES Unless otherwise stated, all information on legal
protection status is from African Wildlife Laws by IUCN Environmental Law
Centre. There is no information on protection in other countries within the
range, namely Angola, Botswana, Malawi, Sudan, Uganda and Zambia.

Ethiopia G. pardalis is totally protected under the Wildlife Conservation
Regulations, 19 January 1972. It may only be hunted for scientific purposes.

Kenya In 1981, Kenya prohibited the export of G. pardalis without the
written permission of the Minister for the Environment and Natural Resources
(Legal Notice 152; 25 September, 1981). Occurs in Tsavo East, Tsavo West,
Meru and Nairobi National Parks (A. Mackay, in litt., 26 March 1986).

Lesotho All tortoises are protected under Proclamation of Monuments,
Relics, Fauna and Flora, L.N. 36 of 1969. There is said to be no trade in any
wild fauna in Lesotho (Ministry of Agriculture and Marketing, Lesotho, in
Jitt., 1985).

Mozambique The species is not protected in Mozambique (Mozambique CITES MA,
1986).

Namibia Protected by the Nature Conservation Ordinance. Occurs in all
Nature Conservation areas (Department of Agriculture and Nature Conservation,
Namibia, in litt., 1987).

Somalia The hunting of all wildlife is prohibited in Somalia under a ban
enacted on 13 October 1977.

South Africa Fully protected in Natal under Nature Conservation Ordinance,
No. 15 of 1974, Proc. 164/1974. Also protected in Transvaal, Cape Province

and the Orange Free State. Occurs in several protected areas (South Africa
CITES MA, 1987).

Tanzania G. pardalis is totally protected under Wildlife Conservation
(National Game) Order, 1974.

Zimbabwe G. pardalis is not specially protected.

CAPTIVE BREEDING Honegger (1980) reported that this species breeds readily
in captivity. Loveridge and Williams (1957) described the breeding behaviour

of several captive specimens of G. p. babcocki. Leakey (in Loveridge and
Williams, 1957) reported that mating was a common occurrence among captive
tortoises of this subspecies. There has been extensive captive breeding in

10

Geochelone pardalis

South Africa, one person having bred over 200. Up to 1981, there were 1985 in
captivity in Transvaal alone; permit records showed that there were at least
1500 in captivity in the Cape Province in 1987 (South Africa CITES MA, 1987).
Commonly breeds in semi-captivity in gardens in Windhoek, Namibia (Department
of Agriculture and Nature Conservation, Namibia, in litt., 1987).

A recent inventory by Slavens (1985) reported that G. p. babcocki is
maintained in 9 localities and G. pardalis_ ssp. in a further 19
collections. A total of 80 specimens was maintained in these localities but
total numbers in captivity may be higher. Several members of the California
Turtle Club have bred this species in large numbers (P.H.C. Pritchard, in
litt., 29 December 1987).

REFERENCES |

Auerbach, R.D. (1985). The Reptiles of Gaborone. Botswana Book Centre,
Gaborone, 47 pp.

Branch, W.R and Brack, H.H. (1987). Reptiles and amphibians of the Addo
Elephant National Park. Koedoe 30: 61-111.

Greig, J.C. and Burdett, P.D. (1976). Patterns in the distribution of

southern African’ terrestrial tortoises (Cryptodira: Testudinidae).
Zoologica Africana 11: 249-273.
Groombridge, B. (1983). World checklist of threatened amphibians and

reptiles. 3rd edition. Nature Conservancy Council, UK, 65 pp.
Honegger, R.E. (1980). Geochelone pardalis, CITES Identification Manual.
Loveridge A. (1957). Checklist of the reptiles and amphibians of East Africa

(Uganda; Kenya; Tanganyika; Zanzibar). Bulletin of the Museum of
Comparative Zoology 117(2): 153-362.
Loveridge, A. and Williams, E.E. (1957). Revision of the African tortoises

and turtles of the suborder Cryptodira. Bulletin of the Museum of
Comparative Zoology 115(6): 163-557.

Peatehard, P.C.H. (1979). Encyclopaedia of Turtles. TFH Publications.
895 pp.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341pp.

Wilson, V.J. (1968). The Leopard Tortoise, Testudo pardalis babcocki, in
eastern Zambia. Arnoldia (Rhodesia) 3(40): 1-11.

11

PANCAKE TORTOISE Recommended list: 3*
[No problem]
Malacochersus tornieri (Siebenrock, 1903)

Order TESTUDINES Family TESTUDINAE

*See last sentence of Summary and Conclusions

SUMMARY AND CONCLUSIONS A small, terrestrial tortoise, inhabiting
scattered rocky areas in southern Kenya and north and east Tanzania. Reported
to be declining in number, no precise estimates are available. Present
populations are in areas with fairly dense human populations. Occurs in the
Ruaha National Park, Tanzania. This species is maintained in captivity and
apparently breeds readily. Listed as ‘Insufficiently Known' in the IUCN
Amphibia-Reptilia Red Data Book.

Until 1981, Kenya exported virtually all animals in trade, F.R. Germany and
the USA taking most of these. From 1982 to 1984, only 1 captive specimen was
reported in trade, but the USA reported importing 65 specimens from Tanzania
in 1985. There is further evidence of continuing and possibly increasing
exports from Tanzania in 1987. Kenyan legislation appears to have effectively
controlled the trade. Although the species is also protected in Tanzania, and
so the recent exports appear to have been illegal.

The reported trade cannot be said to pose a problem to the species; however
there may be increasing trade from Tanzania. The suggested 'C3' listing rests
upon the continuation of Kenya's export ban, and the absence of a substantial
illegal trade from Tanzania.

DISTRIBUTION Confined to Kenya and Tanzania.

Kenya Present in isolated localities in southern Kenya, south of a line
joining Njoro in the west and Malindi on the coast (Loveridge and Williams,
1957) although R. Wood (pers. comm. to P.C.H. Pritchard) reported a specimen
from 29 km west of Isiolo in the Samburu district, and pointed out that the
distribution was much wider than had previously been realised (see Pritchard,
1987). Recent available reports are from the Kitui District (Kenya). There
is some doubt over the veracity of Loveridge's reports of the species from
Njoro and Mida Creek (A.D. Mackay, in litt., 26 March 1986).

Tanzania Found in north and east Tanzania (from Lake Victoria to Lindi near
the Mozambique border (Loveridge and Williams, 1957). Recent available
reports are from the Dodoma District and the Ruaha National Park (Tanzania).
There is some doubt over the veracity of Loveridge's reports of the species
from Njoro and Mida Creek (A.D. Mackay, in litt., 26 March 1986).

POPULATION No precise estimates available, but reported to be declining

(C.E. Norris, in litt., 31 January 1981) and threatened in some degree
(Honneger, 1979).

Kenya Reported by local informants to be still fairly abundant in Kitui
District (A.D. Mackay, in Jitt., 26 February 1981), where a healthy
population exists in the gneissic inselbergs around Mwingi (A.D. Mackay, in
litt., 26 March 1986). R. Wood (see Pritchard, 1987) reported that they were

“not really rare" and that a helathy population existed in an area where
tortoises had previously been heavily collected.

Tanzania Said to be quite frequently encountered in Ruaha National Park,
Tanzania (K. Howell, in Jitt., 1 January 1984). Eleven individuals were
once found under one rock; so density may be locally high (Loveridge, 1923).

12

Malacochersus tornieri

HABITAT AND ECOLOGY A small (around 160-180 mm carapace length) terrestrial
tortoise, inhabiting rocky hills (kopjes) and outcrops in arid thornbush or
savanna, from 30 m to 1800 m altitude. The species may aestivate in hot
weather and apparently remains in shelter during the day (Loveridge and
Williams, 1957). In the wild, dry grass has been recorded as a food item and
probably a variety of vegetation is consumed. Mating occurs in January and
February and nesting in July or August. The clutch comprises a single egg (c
47 x 28 mm), one or two clutches may be laid per year (Pritchard, 1979).

THREATS TO SURVIVAL Exploitation for the pet trade has been the chief
threat, but Wood reported (see Pritchard, 1987) that this had virtually ceased
in Kenya, and considered that the clearance of thorn scrub for agricultural
purposes was now the most serious threat. The presently known populations are
in areas with fairly dense human populations and are particularly vulnerable
(A.D. Mackay, in litt., 26 February 1981).

INTERNATIONAL TRADE The Pancake Tortoise has been moderately exploited for
the pet trade (Groombridge, 1982; Honegger, 1979; Norris, 1980). The only
available data on international trade in this species are those contained in
the Annual Reports of the Parties to CITES which are summarized below.

Table 1 Minimum net imports of live Malacochersus tornieri reported to
CITES. All figures represent live wild specimens except when marked with an
asterisk when captive bred.

1980 1981 1982 1983 1984 1985
German D.R. 10 2 - = = =
Germany, F.R. 123 212 - = iss z
Japan - 6 - = ~ ws
Switzerland 9 19 - s = =
UK - 30 - = = =
USA 59 318 - 1* - 65
Total 201 587 0 il 0 65

Table 2 Reported countries of origin (or exporting country if no original
source reported) and quantities of transactions of live Malacochersus
tornieri reported to CITES. * captive-bred.

1980 1981 1982 1983 1984 1985

Countries having or possibly having populations of HM. tornieri

Kenya 196 567 = = a =
Tanzania - = = = = 65

Countries without populations of M. tornieri

Germany, F.R. - = 1* = 2
Country unknown 5 20 - = = =

Malacochersus tornieri

International trade in this species principally occurred in 1980-1981. The
only other transaction in wild animals occurred in 1985.

Kenya exported 97% of the known exports; the USA and F.R. Germany were the
principal importers. Since Kenya prohibited export in 1981, there have been
no reports of tortoises originating in Kenya. Pritchard (in Jlitt., 29
December 1987) reported substantial exports from Tanzania in 1987, one
exporter alone having 300 ready for export. He noted many in stock at pet
wholesalers in Florida, USA, but considered that the market might be
saturated, as the price per animal had fallen from US$300 to US$40 (Pritchard,
1987). There were indications in 1987 that the trade in Europe might be
increasing D. Morgan, pers. comm.) and this may be in response to the ban on
trade in European species.

CONSERVATION MEASURES Listed as ‘Insufficiently Known’ in the IUCN Red Data
Book (Groombridge, 1982).

Kenya In 1981, Kenya prohibited the export of M. tornieri without the
written permission of the Minister for the Environment and Natural Resources
(Legal Notice 152; 25 September, 1981).

Tanzania The Pancake Tortoise is totally protected under’ Wildlife
Conservation (National Game) Order, 1974. It occurs within the Ruaha National
Park, Tanzania (K. Howell, in litt., 1 January 1984).

CAPTIVE BREEDING Numerous individuals occur in zoos and private collections
(Honegger, 1979). In a recent inventory, Slavens (1985) listed 9 collections
containing 26 specimens. Total numbers in captivity are likely to be higher.
The species has been bred in captivity (Shaw, 1970), and breeding potential is
good under suitable conditions (Honegger, 1979). Breeding may take place at

any time of the year in captivity. Captive animals accept a wide variety of
fruits and vegetables.

REFERENCES

Groombridge, B. (1982). The Reptilia-Amphibia Red Data Book. IUCN Gland and
Cambridge, 426 pp.

Honegger, R. (1979). Red Data Book, Vol. 3: Amphibia and Reptilia. IUCN,
Gland (third edition, revised).

Loveridge, A. (1923). Notes on East African tortoises collected 1921-1923,
with the description of a new species of soft-shelled tortoise.
Proceedings of the Zoological Society of London 1923: 923-933.

Loveridge, A., and Williams, E.E. (1957). Revision of the African tortoises

and turtles of the Suborder Cryptodira. Bulletin of the Museum of
Comparative Zoology 115(6): 163-577.

Norris, C.E. (1980). Excerpts from the report on trade in fauna and flora in
Kenya for 1979. Traffic (International) Bulletin 11(7): 61-68.

Pritchard, P.C.H. (1979). Encyclopaedia of Turtles. T.F.H. Publications,
Hong Kong, 895 pp.

Pritchard, P.C.H. (1987). Pancake Tortoise survey. IUCN Tortoise and
Freshwater Turtle Specialists Group Newsletter 2: 4-5.

Shaw, C.E. (1970). The hardy (and prolific) soft-shelled tortoises.
International Turtle and Tortoise Society Journal 4(1): 6-9, 30-31.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

14

SPUR-THIGHED TORTOISE Recommended list: 2
(Possible problem]
Testudo graeca (Linnaeus, 1758)

Order TESTUDINES Family TESTUDINIDAE

a

SUMMARY AND CONCLUSIONS A medium size terrestrial tortoise. Present in a
variety of habitats including open woodland, scrub, heath, dunes and
semi-desert. Widespread in countries bordering the Mediterranean and in the
Middle East. Recorded from: Algeria, Bulgaria, Egypt, Greece, Iran, Iraq,
Israel, Jordan, Lebanon, Libya, Morocco, Romania, Spain, Syria, Tunisia,

Turkey, USSR, Yugoslavia. Introduced to Cyprus, France and Italy.
Populations in North Africa depleted in areas of heavy commercial collecting,
also threatened by habital destruction. Depleted in Spain. The nominate

subspecies is classified as Vulnerable in the IUCN Amphibia-Reptilia Red Data
Book.

Legally protected in several countries. Import into the EEC region prohibited
since 1984. This appears to have resulted in the virtual cessation of
T. graeca trade reported to CITES, with numbers falling from an annual mean
of 63 033 in 1980-1983 to 321 in 1985.

There seems little doubt that populations could not sustain the level of
exploitation existing previously. If trade has indeed fallen to levels
recorded by CITES, it can no longer be considered a problem; however, the
apparent decline should be substantiated, and trade with non-EEC countries
should be monitored to ensure that it does not show a compensatory increase.

DISTRIBUTION Widespread in countries bordering the Mediterranean and in the
Middle East. Recorded from: Algeria, Bulgaria, Egypt, Greece, Iran, Iraq,
Israel, Jordan, Lebanon, Libya, Morocco, Romania, Spain, Syria, Tunisia,
Turkey, USSR, Yugoslavia. Introduced to Cyprus, France and Italy. Not
present on Malta (Ministry of Agriculture and Fisheries, in litt., 1985).
Testudo graeca is so named because the appearance of the carapace scales
reminded Linnaeus of a Greek mosaic, it is not the typical and widespread
tortoise in Greece; the subspecies T. g. ibera is named after Iberia in
Transcaucasia and not the Iberian Peninsula (Pritchard, 1979).

Four sub-species are generally recognised (Wermuth and Mertens, 1977),
although the race from the Western Transcaucasus (USSR) has recently been
described as a fifth (Khikvadze and Tuniev, 1986).

Testudo graeca graeca: Native to North Africa and Spain; introduced to Italy
and France.

Algeria Confined to the north of the country (Loveridge and Williams, 1957;
Lambert, 1983).

Egypt A single specimen is recorded from El Daba, north-west of El Alamein
(Lambert, 1983).

France Introduced along the south coast (G.H. Parent, in Jitt., 1980,
J.P. Rische, pers. comm., 1980).

Italy Introduced into peninsular Italy, Sicily and probably Sardinia (Bruno
and Maugeri, 1977).

Libya Extending eastwards to Cyrenaica (Lambert, 1983) including Kouf
National Park, where it intergrades with T. g. terrestris (Schleich, 1984).

15

Testudo graeca

Morocco Widespread below 1900 m, but absent from the southern part of the
Moroccan Atlantic plateau and from the arid areas in the south and south-east
(Lambert, 1983).

Spain Occurs in isolated colonies in southern Spain (Murcia-Almeria, Coto
de Donana Reserve and possibly the la Mancia-New Castille Region), south-west
Mallorca and possibly the Pityusen Islands (Ibiza and Formentera) (Lopez
Jurado et al., 1979).

Tunisia Found in the north (Loveridge and Williams, 1957; Lambert, 1983).

T. g. ibera: From northern Greece, thgrough Asia Minor to western Iran.
Albania Listed as present (CITES, 1980), but no records have been located.

Bulgaria Widespread south-east of a line from Zemen to Novgrad (Honegger,
1981; Beshkov, 1984).

Cyprus Isolated specimens have been recorded, but it is doubtful that they
are indigenous (Demetropoulos and Hadjichristophorou, 1981).

Greece Found in parts of north-eastern Greece, Samothrace, Samos, and
certain other islands (with a questionable record from Euboea) (Gruber, 1982),

Iran Confined to the west of the country (Anderson, 1979; Lambert, 1980).
Iraq Found in the north (Mahdi and Georg, 1969; Anderson, 1979).

Romania Occurs in the coastal range to the east of the Danube (Fuhn and
Vaneca, 1961).

Syria Anderson (1979) maps the distribution as passing just outside the

north-eastern border, but Lambert (1980) claimed that this subspecies had been
recorded from Syria.

Turkey Widespread, but absent from the Black Sea Coast to the north of the
Pontic Alps (Basgolu and _ Baran, 1977; Lambert, 1980). Meets with
T. g. terrestris in the south-east (Anderson, 1979).

USSR Occurs in the eastern Caucasus on the Caspian coast (Bannikov et
al., 1977; Lambert, 1980). The race from the Western Transcaucasus has
recently been described as T.g. nikolskii (Khikvadze and Tuniev, 1986).

Yugoslavia Apparently confined to the mountains in southern Macedonia,
although there may be an introduced population in the north (Iverson, 1986).
T. g. terrestris: Around the eastern Mediterranean, from southern Turkey to

Egypt.

Egypt Confined to the north of the country from the Libyan border to Sinai
(Wermuth and Mertens, 1977, Schleich, 1984).

Israel Found almost throughout the northern half of the country in

cultivated as well as natural areas (Israel CITES MA, 1987; Mendelssohn,
1983).

Jordan Restricted to the Upper Jordan valley and Mediterranean habitats
(Jordan CITES MA, 1986).

16

Testudo graeca

Lebanon Wermuth and Mertens (1977) indicate that the range of this
subspecies would include Lebanon, but no records have been located. Lambert
(1980) claimed that T. g. ibera had been recorded from Lebanon, but it seems
that there may have been confusion over the subspecies.

Libya Schleich (1984) confirmed that T. g. terrestris occurs in the Kouf
National Park, in the north-east of Libya, together with T. g. graeca.

Syria Anderson (1979) indicates that this subspecies occurs in the north
and east of Syria.

Turkey Confined to the south-east of Anatolia (Wermuth and Mertens, 1977;
Anderson, 1979).

T. g. zarudnyi: found only in eastern-central Iran, on the Central Plateau.
Although it occurs close to the border, there are no records from Pakistan
(Anderson, 1979).

POPULATION Said to be severely depleted in some western parts of the range,
most notably in Morocco, also parts of Algeria. No details are available for
the greater part of the species's range.

Algeria In the last century, tortoises were reported to be extremely common
in northern Algeria. Tortoises have been collected in the Oran region for
export to France as pets since the end of the last century (Lambert, 1980,
J.P. Rische, pers. comm., 1980). Ina 10-hour search in the Oran region one
recent worker reported finding only one individual (Lambert, 1980). Elsewhere
the species would appear to be relatively abundant. In some regions,
tortoises are reported to be found at sighting frequencies of 4 to 10 per hour
(Anon., 1981). Still found in reasonable numbers in the Algiers region,
despite well developed agriculture (Lambert, 1980).

Bulgaria Formerly widespread, but now depleted in places, particularly the
lowlands (Beshkov, 1984). Tortoises had virtually been eliminated from the
south-west of the country in the 1960s but populations have since recovered
slightly (T. Belokapova, in litt., 20 April 1987).

Cyprus Only isolated specimens have been recorded (Demetropoulos and
Hadjichristophorou, 1981).

France Introduced. Population status uncertain, not clear if regular
breeding occurs (G.H. Parent, in litt., 1980; J.P. Rische, pers. comm.,
1980).

Greece Population status in general appears satisfactory, although the
species is potentially threatened by collecting (Gruber, 1982).

Israel No surveys have been made but T. graeca populations are said to be
healthy and substantial and there is no indication that they are declining
(Israel CITES MA, 1987).

Italy Introduced to peninsular Italy, Sicily, probably also into Sardinia
(Bruno and Maugeri, 1977). Uncertain if regular breeding occurs in all
localities. Precise status unknown, considered rare (M. Capula, in litt.,
1980) and threatened (S. Bruno, in litt., 1980; M.G. Di Palma, in litt.,
1980). A study conducted in April 1986 classified them as vulnerable (Italy
CITES MA, 1987).

Jordan The total population is estimated at 10 000-15 000 (Jordan CITES MA,
1986).

17

Testudo graeca

Morocco Traditionally supplied most specimens for the pet trade in Europe
until 1978 when Morocco ratified CITES and banned the export of all wild
animals including tortoises. At its peak the trade exported several hundred
thousand tortoises annually. Populations now severely depleted. In heavily
collected areas tortoises are now scarce: one worker recorded an average
Sighting frequency of 0.41 per man-hour in June 1978 (Lambert, 1980).
Comparisons between west Turkey and Morocco suggest that collection over the
last 80 years could have reduced abundance by as much as 90% in some areas
(Lambert, 1980).

Spain Populations exist at low densities and have disappeared from many
areas of the peninsula (Andrada, 1980). The western population in the Donana
National Park is said to be secure, but populations to the east, in the
provinces of Almeria and Murcia, are much sparser (Spain CITES MA, 1987).
Despite reports of the species's presence on the Pityusen Islands (Ibiza and
Formentera), populations do not appear to exist at present, or at least are
represented by very few individuals, possibly escaped animals (Lopez Jurado
et al., 1979).

Tunisia In the nineteenth century tortoises were reported to be common on
the Tunisian plains. In the past small numbers have been exported to Italy
(Lambert, 1980). It is said to be common throughout the coastal zone and in
the central plains (Tunisia CITES MA, 1987).

No information is available for remaining countries within the range: Egypt,
Iran, Iraq, Israel, Lebanon, Libya, Romania, Syria, Turkey, Yugoslavia.

HABITAT AND ECOLOGY A medium sized terrestrial species, large individuals
reaching more than 25 cm in length. Found in a variety of habitats, including
sparsely vegetated coastal heathlands, sand dunes, semi-desert areas, and
Mediterranean woodlands, often with Cork Oak Quercus suber and_ Ilex
Q. ilex. Ranges up to 1900 m in Morocco. Mainly herbivorous. The species
hibernates during the winter but may emerge temporarily during warm days.
Emerges finally as early as February in hot coastal areas. Mating occurs in
the spring and eggs are usually laid in May and June (reported to be June and
July in Israel). Clutch size ranges from 2-5, egg size approximately 3 x 4
cm. Although survival is possible in the higher latitudes of north-west
Europe, the success of reproduction is limited by insolation generally being

insufficient for incubation and hatchling survival (Lambert, 1983; Pritchard,
1979).

THREATS TO SURVIVAL The main threats to this species are habitat
destruction and bulk trade collection. Habitat destruction is particularly
Significant in the Mediterranean coastal belt where large scale tourist
development is occurring. Agricultural habitat modification also influences
reproductive success (Lambert, 1981). Large-scale collection for the pet
trade has resulted in populations in northwest Africa being severely depleted;
numbers may have been reduced by as much as 90% in some areas. Selective size
collection both to meet voluntary agreements in the U.K. and to maximize
profits has had an effect on the size and weight of tortoises remaining in the
wild. In general, tortoises in areas of heavy collection tend to be smaller
and lighter. Females tend to be larger than males and selective collection of
the middle size range may have resulted in small males being left to mate with
especially large females. This may affect reproductive success (Lambert,
1980, 1981). Following Morocco's ban on tortoise exports the bulk of this
trade has switched to Turkey (Anon., 1981). Studies in Germany suggest that
after arrival in northern Europe more than 80% of imported tortoises die in
the first year. This is mainly due to stress caused by transportation, poor
conditions in pet shops, and disease and parasite attacks encouraged by
unsuitable climate (Anon., 1981; Lambert, 1980). A further threat may be

18

Testudo graeca

posed by the desertification of the range; the northward extention of the
Sahara has resulted in retraction of T. graeca distribution in Libya
(Lambert, 1981). Not collected or persecuted in Jordan, where an encounter
with a tortoise is believed to bring good luck (Jordan CITES MA, 1986). There
is not thought to be any trade in tortoises in Israel, although they are
occasionally kept as pets when they have been found in gardens or fields.
This is not thought to pose a significant threat (Israel CITES MA, 1987).
Illegal commercial collection of tortoises is thought to be a problem in
Bulgaria, as well as locals and tourists who collect them for pets (T.
Belokapova, in litt., 20 April 1987).

INTERNATIONAL TRADE There is good evidence that large-scale collecting for
the pet trade, for long centred in Morocco and Algeria, has resulted in severe
depletion of tortoise populations. Although most animals have remained in
Europe, significant numbers have been re-exported to Japan and the USA, mainly
by F.R. Germany and the UK. No data are available on the effects of
collecting in Turkey, which assumed greater importance after Morocco's 1978
export ban, but in view of the very large numbers involved, local depletions
are likely to have occurred there also. Although reporting is not
comprehensive, available CITES figures for 1984 indicate an almost complete
cessation of trade in Testudo graeca into the EEC, although there were
substantial imports to Austria. By 1985, the only significant trade was to
the USA, the Austrian trade having ceased. The import ban imposed in the EEC
in 1984 appears to have been highly effective in halting trade, one result of
which was a very dramatic increase in the retail price of tortoises in the UK
(Luxmoore and Joseph, 1986). There can be little doubt that populations of
this species cannot sustain trade on the scale occurring in the 1970s and
early 1980s.

Table 1. Apparent minimum net imports of live Testudo graeca reported to
CITES. Virtually all reported imports are commercial not private.

1980 1981 1982 1983 1984 1985
___.. eee es
Argentina - - - - 4 -
Austria - - 2985 10800 11000 -
Belgium - 7000 - - - =
Brazil - - - - - 2
Denmark - - - - 8 -
France 100 - - - - -
German D.R. - - - - - 10
Germany, F.R. 41985 48302 - 3 - -
Italy = 8x = = = =
Japan 43 12 - - - -
Netherlands - - - 1 - 3
Spain - - - - - 7
Sweden 50 - - - - -
Switzerland 1658 3530 2021 50 1 -
UAE ~ - - 500 - -
UK 39946 38625 22240 27935 - -
USA 352 1449 275 2265 1 299

*xinc. 4 captive bred

TOTAL 84134 98926 27523 41550 11014 321

se SSS eS ea

19

Testudo graeca

Table 2. Reported countries of origin (or exporting country if no original
source reported) and quantities of transactions in live T. graeca reported
to CITES.

1980 1981 1982 1983 1984 1985

a. Countries with wild populations.

Greece - 61 2 - - 3
Italy 50 - - - = -
Morocco 20 - - - - -
Spain ! 45 - - : - - -
Turkey 55003 89115 24520 35000 - 301
Yugoslavia 28990 9742 3000 6550 11000 -

b. Countries without wild populations.

Hungary - - - = 1 =
Switzerland - 4 - = = n

United Kingdom - - = = 8x 3

USSR - = = = 4x 3x
*captive bred

Country Unknown 18 - = 12 1 13

CONSERVATION MEASURES With effect from 1 January 1984 the import of
Testudo graeca and T. hermanni into the EEC region is prohibited.

Listed on Appendix II of the Convention of European Wildlife and Natural
Habitats, Bern 1979 (Honegger, 1981). This obliges contracting parties to
take appropriate legislative and administrative measures to ensure strict
protection of the species.

Algeria T. graeca is listed as a protected species under Decret No.
83.509, 20 August 1983 (Algeria CITES MA, 1987).

Bulgaria Collection and killing of tortoises is prohibited by Order No. 128
of 1981 (Beshkov, 1984). A further Order, No. 729, was issued in 1986 which
made all forms of collection illegal. A poster on tortoise protection was
distributed in 1986, and the police conducted road checks to look for illegal
tortoise dealers. Further distribution of posters and television coverage of
the problem were planned for 1987 (T. Belokapova, in litt., 20 April 1987).

Israel 7. graeca is banned from all trade in Israel. It has also been

designated a “protected asset of nature" and, as such, may not be harmed
(Israel CITES MA, 1987).

Italy 1. graeca is not legally protected in Italy, although it occurs
within some protected areas (Italy CITES MA, 1987).

Morocco After ratifying CITES in 1978 Morocco banned the export of all wild
animals, including tortoises (Anon., 1981).

Romania Legally protected, and declared a national monument. Occurs in
several reserves (Honegger, 1981).

20

Testudo graeca

Spain Legally protected under Decreto 2573 of 5 October, 1973; occurs in
the Coto de Donana reserve in Spain; listed as Vulnerable in the Spanish Red
Data Book (Spain CITES MA, 1987).

Tunisia 7. graeca has been protected since 1974, and its export has been
forbidden (Tunisia CITES MA, 1987).

CAPTIVE BREEDING Large numbers of this species are kept as pets. Regular
breeding in captivity is possible, given appropriate care and conditions
(Kirsche, 1980). It has been suggested that captive-bred tortoises should
replace wild-caught specimens in the live animal trade (Kirsche, 1980).

REFERENCES
Anderson, S.C. (1979). Synopsis of the turtles, crocodiles, and amphisbaenians
of Iran. Proceedings of the California Academy of Sciences 4th ser.,

41(22): 501-528.
Andrada, J. (1980). Guia de campo de los Anfibios y Reptiles de la Peninsula
Ibérica. Omega, Barcelona.

Anon. (1981). The Tortoise Trade. A report on an investigation carried out
by the Royal Society for the Prevention of Cruelty to Animals, UK.
Bannikov, A.G., Darevskii, Ess. 5 Iszczenko, W.G., Rustamov, A.K. and

Shcherbak, N.N. (1977). Opredelitelj zemnovodnye i presmykajuscichsja
fauny SSSR, Moscow.
Basgolu, M. and Baran, I. (1977). Tirkiye Siiringenlen. Kisim 1. Kaplumbaga

ve Kertenkeleler. (The reptiles of Turkey. Part 1. Turtles and
lizards). Fen. Fak. Kitaplar Ser. Eye University. No. 76. Bornova,
Izmir.

Beshkov, V.A. (1984). On the distribution, relative abundance and protection
of tortoises in Bulgaria. Bulgarian Academy of Sciences, Ecology 14:
14-33 (in Bulgarian, English summary).

Bruno, Sis and Maugeri, Si: (1977). Retilli d'Italia, Vol. Tg
Tartarughe-Sauri, Martello-Giunti, Firenze.

Demetropoulos, A. and Hadjichristophorou, M. (1981). The Chelonians of
Cyprus. Biological Society of Cyprus Bulletin 1: 13-17.

Fuhn, I.E. and Vancea, S. (1961). Fauna Republicii Populare Romine.

Reptilia (Testoase, Sopirle, Serpi). Ed. Acad. Rep. Pop. Romine., Vol.
XIV F.2.: 158-170.

Gruber, U. (1982). In litt. (text of paper read at scientific meeting of
the Hellenic Society for the Protection of Nature, Karlsruhe, 27 March
1982).

Honneger, R.E. (1981). Threatened Amphibians and Reptiles in Europe.
Supplementary Vol., of Handbuch der Reptilien und Amphibien BEuropas,
Wiesbaden, Akademische Verlagsgesellschaft.

Iverson, J.B. (1986). A checklist with distribution maps of the turtles of
the world. Earlham College, Richmond, Indiana.

Khikvadze and Tuniev (1986). On the taxonomic status of modern land tortoises
of the Western Transcaucasus. Bulletin of the Academy of Sciences of the
Georgian SSR 124(3): 617-620.

Kirsche, W. (1980). Conservation of Tortoises by breeding. Proceedings of
the European Herpetological Symposium, Oxford C.W.L.P. pp. 125 (Abstr.),
also in ASRA Journal 1(3).

Lambert, M.R.K. (1980). The Mediterranean Spur-Thighed Tortoise, Testudo

graeca, in the wild and in_ trade. Proceedings of the Suropean
Herpetological Symposium, Oxford C.W.L.P., pp. 17-23.

Lambert, M.R.K. (1981). The conservation of Mediterranean (W-Palaearctic)
tortoises: the problem posed. (Abstract). First Ordinary General
Meeting Societas Europaea Herpetologica. 13-16, September, Vienna;
Abstracts.

21

Testudo graeca

Lambert, M.R.K. (1983). Some factors influencing the Moroccan distribution of
the western Mediterranean spur-thighed tortoise, Testudo graeca graeca
L., and those precluding its survival in NW Europe. Zoological Journal
of the Linnaean Society, London 79: 149-179.

Lopez Jurado, L.F., Talavera Torralba, P.A., Ibanez Gonzalez, J.M., MacIvor,

J.A., and Garcia Alcazar, A. (1979). Las tortugas terrestres Testudo
graeca y Testudo hermanni en Espana. Naturalia Hispanica 17, ICONA,
Madrid.

Loveridge, A. and Williams, E.E. (1957). Revision of the African tortoises

and turtles of the suborder Cryptodira. Bulletin of the Museum of
Comparative Zoology 115(6): 163-557.

Luxmoore, R. and Joseph, J. (1986). UK trade in tortoises. Traffic
Bulletin 8(3): 46-48.

Mahdi, N. and Georg, P.V. (1969). A systematic list of the vertebrates of
Iraq. Iraq Natural History Museum, University of Baghdad Publication No.
26.

Mendelssohn, H. (1983). Herpetological nature protection. Israel - Land and
Nature, 9(1): 21-27.

Pritchard, P.C.H. (1979). Encyclopedia of Turtles. T.F.H. Publications,
Hong Kong and New Jersey, 895 pp.

Schleich, H.-H. (1984). Studies on the herpetology of Kouf National Park and
adjacent areas of Cyrenaika with a checklist of the amphibians and
reptiles of Libya. Final Report, IUCN/WWF Project 9044 - Libya -Kouf N.P.

Wermuth, H. and Mertens, R. (1977). Testudines, Crocodylia, Rhynchocephalia.
Das Tierreich, Walter de Gruyter, Berlin, 100: 1-174.

22

HERMANN'S TORTOISE Recommended list: 2

[Possible problem]
Testudo hermanni (Gmelin, 1789)

Order TESTUDINES Family TESTUDINIDAE

SUMMARY AND CONCLUSIONS A small terrestrial tortoise, restricted to
southern Europe. The range extends around the Mediterranean, often in coastal
areas, from north-east Spain eastward to European Turkey; the species also
occurs in parts of Romania and Bulgaria and is present on several islands.
Inhabits a variety of rather dry habitats (scrub woodland, heath or maquis,

for example), in areas with hot summers, often at coastal sites. Mainly
herbivorous. Clutch comprises 1-12 eggs, often around three, there may be two
clutches per. season. Western populations from Spain to Italy (T. h.

robertmertensi) are in general extremely localized and depleted; eastern
populations (T. h. hermanni) are more widespread and sometimes with adequate
numbers, but heavily exploited for the pet trade in many areas and declining
in parts. Listed as Vulnerable in the IUCN Amphibia-Reptilia Red Data Book,
Part 1.

Import into EEC countries has been prohibited since 1984. Numbers in reported
trade fell significantly, from an annual mean of 16 787 in 1980-1983, to 7371
in 1984, but then rose again to 13 349 in 1985. Most of the imports in 1984
and 1985 were to Austria, Switzerland, Hungary and the USA (not EEC members).
Turkey was formerly the major source, but recently Yugoslavia has emerged as
the main supplier. In 1985, 4000 were reported as originating in the USSR,

possibly indicating misidentification for fT. horsfieldii. Listed on
Appendix II of the Convention on the Conservation of European Wildlife and
Natural Habitats. Nominally protected by legislation in much of the range,

present in a few protected areas.

There seems little doubt that populations were adversely affected by the level
of exploitation existing prior to 1984. If imports to the EEC since the
import ban have in fact fallen to the level reported to CITES, it is unlikely
to pose a serious problem; but the growing imports to non-EEC countries,
particularly Austria, is of concern. Imports from the USSR_ should be
investigated.

DISTRIBUTION Restricted to southern Europe. Present in Albania, Bulgaria,
France, Greece, Italy, Romania, Spain, Turkey and Yugoslavia. Although
reported to occur on Malta (see Iverson, 1986), any individuals of
T. hermanni found may be escapes, it is occasionally imported by pet shops
but does not live or breed in the wild (Ministry of Agriculture and Fisheries,
in litt., 1985).

This species is usually (Wermuth and Mertens, 1977) treated as comprising two
subspecies; T. h. robertmertensi from central Italy, Sardinia and Corsica
westwards, and T. h. hermanni from southern Italy and the Balkans. However,
some doubt has been raised about the proper assignment of animals from
peninsular Italy (Bruno and Maugeri, 1977). Bour (1987) has proposed that
what Wermuth and Mertens (1977) treated as T. h. robertmertensi should be
called T. h. hermanni and that what they treated as T. h. hermanni should
be assigned to T. h. boettgeri.

Albania Few details available (see Iverson, 1986), probably widespread.

Bulgaria Formerly widespread, although now absent from several parts of its
former range (Beshkov, 1984).

23

Testudo hermanni

France Localized, occuring in the Albéres hills along the French-Spanish
border in the east Pyrenees, and the Maures and Estérel mountains in Var
province (south-east France) (Cheylan, 1981; Devaux et al., 1986). Also
present (though probably introduced) on Corsica, mainly along the east coast
(Honegger, 1981).

Greece The most widespread tortoise in Greece where it appears to occur
over much of the mainland including the Peloponnese (Arnold and Burton,
1978). It may be absent from parts of the south-east, around Corinth, and
becomes progressively rarer east of Thessaloniki (Stubbs et al., 1981).
Present on the Ionian Islands, including Corfu (Honegger, 1981).

Italy Hermann's Tortoise is present in peninsular Italy, mainly on the
Ligurian and Tyrrhenian coasts and also occurs on Sicily, Sardinia, and
certain smaller islands including Elba, Lampedusa and Pantelleria (Bruno and
Maugeri, 1977).

Romania Recorded from about 40 localities in the counties Mechedinti,
Caras-Severin and Gorj; all near the Yugoslavian border in the extreme
south-west (Fuhn, 1981; Fuhn and Vancea, 1961).

Spain Present in small numbers in the south of Menorca and two populations
in north-east and south-east Mallorca (Balearic Islands), also on the
mainland, in a small area in the north of Gerona province (Lopez Jurado et
al., 1979; Cheylan, 1981). Also reported from the area of Valencia (Andrada,
1980), with scattered reports from elsewhere. There is a possibility that
most T. hermanni colonies on the mainland are not indigenous (Lopez Jurado
et al., 1979).

Turkey Restricted to European Turkey (Basoglu and Baran, 1977; Eiselt and
Spitzenberger, 1967; Pritchard, 1979).

Yugoslavia The species ranges along the Adriatic coast from Dalmatia
southwards, then south-east through parts of Macedonia (Meek and Inskeep,
1982; Meek, 1985).

POPULATION
Albania No data.

Bulgaria Formerly widespread (Beskov and Beron, 1964), but numbers of both
Testudo in Bulgaria have declined sharply over the last decades; both are
now absent from several parts of the country (Beskov, 1984).

France During Neolithic times T. h. robertmertensi ranged across’ the
entire Mediterranean region of mainland France, and also much further to the
north (Cheylan, 1978). The species has since undergone a severe regression,
due apparently to climatic and human influences, and is now restricted to
hills in the province of Var in the extreme south-east (the Massifs des Maures
and Estérel), and the Albéres in the east Pyrenees. The tortoise is now
regarded as severely threatened in France (Cheylan, 1978, 1981). Extirpated
in the early 19th century in Iles d'Hyéres owing to over-exploitation (Rische,
1979). Although the general area of distribution may not have diminished
greatly for several decades, the density of individuals is progressively
decreasing (Dumont, 1974). Small populations remain in parts of the Massif
des Maures, but at densities as low as 3-4 per ha, where they are regarded as
under severe threat (Devaux et al., 1986). Populations may be locally
adequate in Corsica (J.-P. Rische, pers. comm., 1980). However, available
evidence indicates that the Massif des Maures population is slowly declining
due to egg predation and changes in land use (Stubbs and Swingland, 1985).

24

Testudo hermanni

Greece Populations are widespread and still generally in satisfactory
condition, although an overall decline has been observed (Gruber, 1982). In
the north-east, the species becomes progressively rarer east of Thessaloniki
(Stubbs et al, 1981). At a major 75-ha study site on coastal heath at Alyki
(Macedonia), average T. hermanni density was 55 per ha, with a_ total
population of around 5,000. In optimum dry sandy heath habitat within this
site, a maximum density of over 150 tortoises per hectare was observed (Stubbs
et al., 1981). After a severe fire swept the area in 1980 the population
was estimated to have fallen to 25 per ha in 1982 (Stubbs et al., 1985).

Italy In general, rare and localized (Bruno and Maugeri, 1977); has
disappeared from much of the coast, but probably retains natural densities in
some National Parks (eg. Maremma) or on private land (Bruno, 1971, 1973).

Romania Reportedly in marked decline (Honegger, 1981).

Spain Both Testudo species in mainland Spain are reported to exist in low

density and to have disappeared from many areas (Andrada, 1980). In the
Balearics, populations are extremely localized in the south of Menorca and
local in Mallorca (Lopez Jurado et al., 1979). Fire is said to be an

important factor causing a population decline in Gerona (Spain CITES MA, 1987).

Turkey Apparently not common within its restricted range, but no details
available.

Yugoslavia Declining locally due to collecting for the pet trade (Honegger,
1981). Healthy populations are known in parts of Dalmatia and Montenegro
(Meek and Inskeep, 1982; Windolf, 1980). Around 125 individuals were noted in
a 2 km* area in Montenegro (Meek and Inskeep, 1982). A further study, in
1983, estimated a mean density of 39.2 per ha (Meek, 1985).

HABITAT AND ECOLOGY A small terrestrial tortoise, to around 20 cm length,
individuals in western populations may be a little smaller. Inhabits a variety
of often rather dry habitats, in areas with hot summers. Generally in
lowlands and low hills, and in open deciduous woodland, wood edges, scrub
fields and hillsides, maquis and garrigue vegetation, etc. (Arnold and
Burton, 1978; Meek, 1985; Windolf, 1980; Bruno and Maugeri, 1977). At one
study site in northern Greece, maximal tortoise density was found in areas of
dry sandy heath with a ground cover of lichens and herbs, Artemesia clumps,
Hawthorn and Ruscus aculeatus (Stubbs et al., 1981).

Mainly herbivorous, although invertebrates, carrion and faeces may be eaten on
occasion. At one scrubland site in southern Yugoslavia (Meek and Inskeep,
1982) leguminous plants of the subfamily Papilionoidea provided the bulk of
the diet.

Sexual maturity may be attained (in females) at around seven years. There is
typically a period of winter hibernation, courtship may occur sporadically
throughout the summer. Eggs are generally laid in May-June, with the
hatchlings emerging in August-September. A flask-shaped nest 7-8 cm deep is
excavated by the hind feet. The clutch comprises from one to 12 eggs, usually
three. Eggs are hard-shelled, slightly elongate. There may be two clutches
laid per season (Pritchard, 1979; Street, 1979).

THREATS TO SURVIVAL Threatened mainly by large-scale commercial collecting
and habitat destruction. Western populations assigned to T.. h.
robertmertensi, particularly those in mainland France, are at risk due to the
extremely localized distribution. About 103 450 ha of maquis and pine woods
have been destroyed by fire in Var (France) in the last ten years, and 20% of
the French hermanni population may have been destroyed in the fire of July

25

Testudo hermanni

1979 (Dumont, 1974). In northern Greece, Stubbs et al. (1985) estimated
that a heath fire reduced the population by about 40%, but they found that the
growth rate of juveniles rose sharply subsequently, and they concluded that
tortoise populations were relatively resiliant to this type of catastrophe.
In France, tortoises are occasionally collected for commercial purposes, but
the collection of individual animals as pets is also a serious problem owing
to the low population numbers (Devaux et al., 1986). Urbanization is a
further threat (Cheylan, 1981). Habitat alteration is cited as the primary
cause of decline in Bulgaria, including intensification of agriculture,
expansion of pasture-land, and coniferous afforestation, but with
industrialisation, urbanisation and collecting for food as secondary causes.
Collecting now occurs mainly around Plovdiv and vicinity, but was concentrated
in the east where T. hermanni used to be abundant, and shifted to the west
in the 1970s (Beshkov, 1984). On the Tyrrhenian coast of Italy,
transformation of the coastal environment by building speculation, collecting
for the pet trade, and use as food by local inhabitants, are cited as major
threats (Chelazzi, in Jitt., 1980). Populations in Yugoslavia have been
subjected to extremely heavy collection pressure; as many as 40 000 animals
were exported for the pet trade in the early 1970s (Honegger, 1981). The
tortoises are eaten by man in some parts of the range, eg. Bulgaria (Sura,
1981) and are widely offered as tourist souvenirs, in Yugoslavia, for example.

Table 1. Apparent minimum net imports of live Testudo hermanni reported to
CITES (eight preserved specimens were also reported but do not appear below).

1980 1981 1982 1983 1984 1985
Australia = - - - - 2
Austria - - - 5350 3355 85
Bermuda - - - 8 - -
Canada - - - 7 - 6
Chile - - - - 1 -
Denmark - - - - 8 -
France 650 499 - 400 300 -
German D.R. 1400 - 6 - a -
Germany, F.R. 14217 2 - 2 7 Ce sed
Hungary - - - - 400 -
Italy - - - 15 - -
Japan 20 800 - 210 5 43
Korea (Rep. ) - - 2x - - -
New Zealand - 3 - - - -
Saudi Arabia - = = = = bs
Spain - - - 2 2 1
Switzerland 3855 4788 5277 4751 2549 2019
Tanzania - = = = = 2
UAE = = = 2 = 30
UK 9993 10000 3 2503 - -
USA 347 71 841 1126 734 1354
Country Unknown - 4 = E: wz =
* captive bred
Total 30478 16171 6129 14372 7371 13349

26

Testudo hermanni

INTERNATIONAL TRADE Large numbers of live T. hermanni have been involved
in the pet trade in recent decades. Austria, F.R. Germany, Switzerland and
the UK have all been major importers in the 1980s, the last two countries most
consistently. A majority of these specimens have remained in Europe but some
have been re-exported to the USA. Turkey was formerly the major source, but
recently Yugoslavia has emerged as the main supplier. In 1985, 4000 were
reported as originating in the USSR, possibly indicating misidentification for
T. horsfieldii. Since the ban on imports into the EEC in 1984, numbers in
reported trade have fallen significantly, from an annual mean of 16 787 in
1980-1983 to 7371 in 1984, but then rose again to 13 349 in 1985. Most of the
imports in 1984 and 1985 were to Austria, Switzerland, Hungary and the USA
(not EEC members).

Table 2. Reported countries of origin (or exporting country if no original
source reported) and quantities of transactions in live T. hermanni reported
to CITES.

a. Countries having wild populations of T. hermanni

Bulgaria - - 1 - - -
Greece 13 at 3% 3 4 -
Italy - 73 ~ - - -
Turkey 24000 10350 255 2500 - 286
Yugoslavia 6131 5720 5784 11866 7358 9054

b. Countries without wild populations of T. hermanni

Austria 29 68 - = = e
Belgium = = 83 = cS =
Czechoslovakia 2 - - = = =
German D.R. - - = = 4x =
Germany, F.R. 303 - = = 2 s
Iceland - - = - 5 =
Switzerland - - = = x 42*
UK - - - - 10* 2
USSR = - - - 2 4000
Country Unknown - - 5 1 4 3

*captive bred

a

CONSERVATION MEASURES Listed on Appendix II of the 1979 Convention on the
Conservation of European Wildlife and Natural Habitats (the Berne Convention)

(Anon., 1979). This obliges contracting parties to _ take appropriate
legislative and administrative measures to ensure strict protection of the
species. Import of Mediterranean Testudo species into the EEC has been

prohibited since 1 January 1984.

Bulgaria Collection and killing of tortoises is prohibited by Order No. 128
of 1981 (Beshkov, 1984). A further Order, No. 729, was issued in 1986 which
made all forms of collection illegal. A poster on tortoise protection was
distributed in 1986, and the police conducted road checks to look for illegal
tortoise dealers. Further distribution of posters and television coverage of
the problem were planned for 1987 (T. Belokapova, in litt., 20 April 1987).

France Protected by law (No. 76269, 10 July 1976, and Décret No. 77.1295,

27

Testudo hermanni

25 November 1977). A three-year research programme has recently been
completed in the Massif des Maures (Var) and a cooperative tortoise
conservation project was initiated. This involves habit restoration,

protection of nest sites, and captive propagation (Devaux et al., 1986).

Italy 7. hermanni is not legally protected in Italy, although it occurs
within some protected areas (Italy CITES MA, 1987), including the Parco
Naturale Regionale della Maremma, in coastal Tuscany (E. Balleto, pers. comm.,
1981).

Romania Legally protected, and declared a national monument. Occurs in
several reserves (Honegger, 1981).

Spain Legally protected under Decreto 2573 of 5 October, 1973; listed as
Vulnerable in the Spanish Red Data Book (Spain CITES MA, 1987). A major
population of T. hermanni occurs on private land in Mallorca and is the main
subject of the Son Cifre Conservation Project (Kramer, 1981).

CAPTIVE BREEDING Regular breeding in captivity is possible, given

appropriate care and conditions (Kirsche, 1980). It has been suggested that
captive-bred tortoises should replace wild-caught specimens in the live animal
trade (Kirsche, i980). As part of a conservation programme in southern

France, eggs have been collected from the wild and artificially incubated to
reduce predation (Devaux et al., 1986).

REFERENCES

Andrada, J. (1980). Guia de campo de los Anfibios y Reptiles de la Peninsula
Ibérica. Omega, Barcelona.

Anon. (1979). Explanatory report concerning the Convention on_ the
Conservation of European Wildlife and Natural Habitats. Council of
Europe, Strasbourg.

Arnold, E.N., and Burton, J.A. (1978). A Field Guide to the Reptiles and
Amphibians of Britain and Europe. London: Collins.

Basgolu, M. and Baran, I. (1977). Tiirkiye Siiringenlen. Kisim 1. Kaplumbaga

ve Kertenkeleler. (The reptiles of Turkey. Part 1. Turtles and
lizards). Fen. Fak. Kitaplar Ser. Eye University. No. 76. Bornova,
Izmir.

Beshkov, V.A. (1984). On the distribution, relative abundance and protection
of tortoises in Bulgaria. Bulgarian Academy of Sciences, Ecology 14:
14-33 (in Bulgarian, English summary).

Beskov, V., and Beron, P. (1964). Catalogue et Bibliographie des Amphibiens
et des Retiles en Bugarie. Ed. Acad. Bulg. Sci., Sofia, pp. 1-39.

Bour, R. (1987). L'identité des tortues terrestres européennes. Revue
Francaise d'Aquariologie 13(4): 111-122.

Bruno, S. (1971). Red Book: Testudo hermanni_ Gmelin. Notiz. Unione
Erpetol. Ital. 1(2): 30.

Bruno, S. (1973). Problemi di conservazione nel campo dell’ erpetologia.
Atti 3rd Simp. naz. Conserv. Nat. Bari 2: 117-226.

Bruno, So and Maugeri, S. (1977). Retilli d'Italia, Vol. J ie

Tartarughe-Sauri, Martello-Giunti, Firenze.
Cheylan, M. (1978). Species account for Testudo hermanni, p. 76, In: Anon.,

(Societe Herpetologique de France). Atlas preliminaire des Reptiles et
Amphibiens de France. Montpellier.
Cheylan, M. (1981). Actual status and future of Hermann's Tortoise in

western Europe. Paper presented at the 2nd European Chelonian Symposium,
3 October, Oxford.

Devaux, B., Pouvreau, J.-P. and Stubbs, D. (1986). Programme de sauvegarde

des tortues d'Hermann dans le Massif des Maures (France). SOPTOM, Les
Mayores, France.

28

Testudo hermanni

Dumont, M. (1974). Les chéloniens de France. Leur avenir, leur protection.
Le courrier de la nature 33: 224-227. Also published (1972) in Natur.
Orléan 3(5): 10-12.

Eiselt, dicts and Spitzenberger, F. (1967). Ergebnisse zoologischer
Sammelreisen in der _ Turkei: Testudines. Annalen Naturhistorischen
Museums in Wien 70: 357-378.

Fuhn, I.E. (1981). Rare and endangered amphibians and reptiles in Romania,
proposal for conservation (text of paper read at Societas Europaea
Herpetologica meeting, 3-16 September, Vienna).

Fuhn, I.E. and Vancea, S. (1961). Fauna Republicii Populare Romine.
Reptilia (Testoase, Sopirle, Serpi). Ed. Acad. Rep. Pop. Romine., Vol.
XIV F.2.: 158-170.

Gruber, U. (1982). Herpetofauna Griechenlands (text of lecture given at
meeting of the Scientific Commission of the Hellenic Society for the
Protection of Nature, 27 March, Karlsruhe).

Honneger, R.E. (1981). Threatened Amphibians and Reptiles in _ Surope.
Supplementary Vol., of Handbuch der Reptilien und Amphibien Suropas,
Wiesbaden, Akademische Verlagsgesellschaft.

Iverson, J.B. (1986). A checklist with distribution maps of the turtles of
the world. Earlham College, Richmond, Indiana.

Kirsche, W. (1980). Conservation of Tortoises by breeding. Proceedings of
the European Herpetological Symposium, Oxford C.W.L.P. p. 125 (Abstr.),
also in ASRA Journal 1(3).

Kramer, T. (1981). Statement at first meeting of IUCN/SSC Tortoise Specialist
Group, 1-2 October, Oxford.

Lopez Jurado, L.F., Talavera Torralba, P.A., Ibanez Gonzalez, J.M., MacIvor,

J.A., and Garcia Alcazar, A. (1979). Las tortugas terrestres Testudo
graeca y Testudo hermanni en Espana. Naturalia Hispanica 17, ICONA,
Madrid.

Meek, R. (1985). Aspects of the ecology of Testudo hermanni in southern
Yugoslavia. British Journal of Herpetology 6: 437-445.

Meek, R., and Inskeep, R. (1982). Aspects of the field biology of a
population of Hermann's Tortoise (Testudo hermanni) in southern
Yugoslavia. British Journal of Herpetology 6: 159-164.

Pritchard, P.C.H. (1979). Encyclopedia of Turtles. T.F.H. Publications,
Hong Kong and New Jersey, 895 pp.

Risch, J.-P. (1979). Les tortues terrestres paléarctiques (Testudo spp.) en
France: présence a l’*état sauvage, maintien et reproduction en captivité,
protection (Reptilia, Testudines, Testudinidae). Bulletin Societes
Zoologique de France 103(4): 524-527.

Street, D. (1979). The Reptiles of Northern and Central Europe. London,
Batsford.

Stubbs, D., Hailey, A., Tyler, W., and Pulford, L. (1981). University of
London Natural History Society, Expedition to Greece 1980. Report, pp.

1-136.
Stubbs, D., and Swingland, I.R. (1985). The ecology of a Mediterranean
tortoise (Testudo hermanni): a declining population. Canadian Journal

of Zoology 63: 169-180.

Stubbs, D., and Swingland, I.R., Hailey, A. and Pulford, E. (1985). The
ecology of the Mediterranean tortoise (Testudo hermanni) in Northern
Greece (The effects of a catastrophe on population structure and density.
Biological Conservation 31: 125-152.

Sura, P. (1981). Notes on the reptiles of Bulgaria. British Herpetological
Society Bulletin 3: 25-28.

Wermuth, H. and Mertens, R. (1977). Testudines, Crocodylia, Rhynchocephalia.
Das Tierreich, Walter de Gruyter, Berlin, 100: 1-174.

Windolf, R. (1980). Zur biologie, Gkologie und zum artenschutz der
Griechischen Landschildkréte (Testudo hermanni h.) in Jugoslavien.
Okol. (Z. £. Okologie, Natur- und Umweltschutz) 2(4): 14-20.

29

AFGHAN or HORSFIELD's TORTOISE Recommended list: 2
[Possible problem]

Testudo horsfieldii Gray, 1844

Order TESTUDINES Family TESTUDINIDAE

SUMMARY AND CONCLUSIONS A small Central and South-west Asian tortoise,
ranging from Kazakhstan (USSR) and the western fringe of the Dzungarian Basin
(China) south through Afghanistan and east Iran to the Baluchistan region of
Pakistan. Occurs in a variety of semi-arid habitats, including semi-desert
and steppelands. Reproductive potential is low; rather late maturing, with
first breeding at 11-14 years, and with two clutches per year, of only 5-6
eggs in total.

The species, is affected by habitat loss due to agricultural expansion in parts
of the range. Around 150 000 specimens annually have been collected in
Kazakhstan alone in recent years, about half of which are sold within the
USSR. Between 24 000 and 100 000 annually have been recorded in international
trade from 1980 to 1985.

Despite substantial populations remaining in the USSR, this level of harvest
may well be non-sustainable; management and further field research have been
recommended.

DISTRIBUTION A Central and Southwest Asian species, extending from the
eastern shores of the Caspian Sea eastward through the Turkestan region just
into the Dzungarian Basin of China, and south-east through eastern Iran and
much of Afghanistan to north-west Pakistan.

Following Khozatsky and Mlynarski (1966), some authors assign this species to
the genus Agrionemys rather than Testudo; it differs most noticeably from
members of the latter genus in possessing four rather than five digits on the
forelimbs, also in other features.

Afghanistan Apparently occurs widely, with the exception of the Afghan
portion of the Seistan basin and the complex knot of mountain ranges in the
centre and north-east (Anderson, 1979, Fig. 12).

China To the east of Lake Balkhash (USSR) the species extends into the
Dzungarian Basin in the Xinjiang Uygur (Sinkiang Uigur) Autonomous Region of

China, where it occurs in a 120 sq. km hill area north of Huocheng County
(Anon., 1984).

Iran The range extends south from Turkestan into eastern parts of the
Iranian plateau, where the species is recorded from the provinces of

Mazanderan (the north-east only) and Khorasan, but not south of the Seistan
basin (Anderson, 1979).

Pakistan Despite its apparent absence from Iranian Baluchistan (Zarudny,
cited in Anderson, 1979), the Afghan Tortoise is generally distributed in the
north and west of the Baluchistan region of Pakistan and into Wazirstan in the
tribal frontier areas (Minton, 1966).

USSR Widely distributed through the Turkestan region, in Turkmenistan,
Uzbekistan and southern and central parts of Kazakhstan. Apparently largely
absent from the Pamir and Tien Shan highlands of Tadzhikistan and Kirgizia, in
the southwest of this region (Bannikov et al., 1977).

30

Testudo horsfieldii

POPULATION Little information is available on populations of
T. horsfieldii in Afghanistan, China, Iran and Pakistan, but useful data are
available from the Russian portion of the range.

Afghanistan Noted as numerous on the plains near Gulran in the extreme
north-west of the country in 1884 (Aitchinson, 1889). Reportedly quite
abundant in the Dasht-i-Leili between Maimana and Shibarghan in northern
Afghanistan (Toynbee, 1961); no recent information available for most of the
country.

China One source (Anon., 1984) terms the species “rare” and states that the
population has been “decimated”.

Iran Reports from the turn of the century (Zarudny, cited in Anderson,
1979) suggest that the species was common at (at least) one locality in
eastern Khorasan, but rare in the southern part of its range; no recent
information available.

Pakistan Cited as “rare” (Ghalib et al., 1980), but general information
presented by Minton (1966) suggests the species is not uncommon locally; for
example, seven individuals were seen at one time in a meadow near a small
watercourse.

USSR Quite extensive surveys have been carried out in many parts of
Kazakhstan S.S.R. (Alma-Ata, Taldy-Kurgan, Dzhambul, Chimkent oblasts)
(Brushko and Kubykin, 1982), and the southern parts (Maryj region) of Turkmen
Stick. (Frolov et al., 1985). Tortoise population density varied
considerably over the 2130 ha surveyed in Kazakhstan, between 0.5 and
(exceptionally) 26 specimens per hectare, but about seven per hectare was
rated as “comparatively high" and most sites appear to have yielded between
two and three per hectare (Brushko and Kubykin, 1982). A specific instance of
regional decline is provided by the Kerbulak massif in Kazakhstan where at
present between 11 and 15 individuals may be found per hectare, yet the same
sites reportedly held about 40 per hectare in the 1950s (Brushko and Kubykin,
1982). The higher of the densities found in Kazakhstan are reportedly rare in
other parts of the species' range in USSR (see Russian language sources cited
in Brushko and Kubykin, 1982). This seems to be confirmed by recent data from
southern Turkmenia, where the highest local mean density in the Maryj region
was about 4.5 specimens per hectare (in argillaceous and sandy desert
foothills), and the lowest 0.44 per hectare (in the valley of the Murghab
river system) (Frolov et al., 1985). Total numbers in the 45% of the
86 000 sq. km area Maryj region that was surveyed, were estimated at 26.8
million. The species is not included in the USSR national Red Data Book.

HABITAT AND ECOLOGY A rather small tortoise, males attain a maximum length
of about 16.5 cm (1100 g), and females about 19.5 cm (1750 g) (Brushko and
Kubykin, 1982). Various habitat types are occupied, including sand or stone
deserts and mountain slopes, but tortoise density varies appreciably between
them. In Kazakhstan the largest populations occur in desert areas with
wormwood Artemisia spp. and a variety of ephemerals, but in general good
populations may be found in areas of broken relief, steppe areas with good
grass cover, along seasonal river beds and ravines with abundant shelter.
Numbers are lowest in saline zones, and in sandy or stony deserts (Brushko and
Kubykin, 1982). In Pakistan, Afghan Tortoises frequent rocky hill country,
especially between 1500 and 2100 m (5000-7000 ft), and can be seen most
readily at grassy sites near springs or streams. They do range into
moderately arid rocky desert but appear to avoid areas with sand or clay
substrates (Minton, 1966).

31

Testudo horsfieldii

In USSR the species is only active for 2.5-3 months, from the end of March
until mid-June; the rest of the year is spent in shelter (although there may
be sporadic activity in summer and autumn). Mature males both enter
aestivation earlier and become active earlier than females and juveniles.
Activity over a 24 hr period is similarly restricted, typically to between
10 a.m.-12 noon and 4-6 p.m., but this simple two-peak pattern is obscured in
bad weather when activity is more widely distributed, and also varies
according to the nutritional state of the individual. Shelters occupied are
simple scrapes in the ground, to carapace depth, or burrows under rocks -
apparently deeper during the winter aestivation period (Brushko and Kubykin,
1982; Minton, 1966).

The diet is largely vegetarian; adults feed on leaves, fruit and flowers
(rarely on grass) while young appear especially attracted to fruit and flowers
red in colour (Minton, 1966). Considerable fat reserves are accumulated
during spring and feeding (Brushko and Kubykin, 1982).

Sexual maturity is attained relatively late: at 11-12 years of age, carapace
length 11.2-11.7 cm, weight 380-480 g, in males, and 13-14 years, 13.4-14 cm,
560-620 g, in females (Brushko and Kubykin, 1982). Copulation (males always
smaller and younger than females) occurs in March-April, egg-laying typically
at the end of April and the beginning of June (although varying in females of
different ages) (Brushko and Kubykin, 1982). The reproductive rate is low: in
south-east Kazakhstan two clutches are laid per year, five or six eggs in
total. There appears to be appreciable, although unquantified, mortality of
eggs and young due to predation and climatic factors. For example, young
tortoises form a considerable portion of the diet of rooks Corvus frugilegus
in Kazakhstan (Brushko and Kubykin, 1982).

Individual movements do not appear to be extensive; for example, 32% of 132
specimens marked in 1975 were recaptured in 1976-1978 between 40 and 1220 m
from the original site (Brushko and Kubykin, 1982).

THREATS TO SURVIVAL Threatened by exploitation for the live animal trade,
for food, and in the USSR portion of the range at least, by habitat changes
associated with agricultural development.

In China, hunting for food is cited as the major threat (Anon., 1984).
Similarly, in parts of the USSR, tortoise soup is much appreciated and the
liver and eggs are highly regarded. Tortoises are exported for food use and

were even used as food for fur-bearers during the 1950s (Brushko and Kubykin,
1982).

Large numbers of tortoises enter the live animal trade. The number collected
for this purpose in Kazakhstan has increased steeply: 43 000 in 1967, 110 000
in 1973, 126 000 in 1975 and about 150 000 annually in recent years (Brushko
and Kubykin, 1982). Brushko and Kubykin (in a paper prepared in 1981) stated
that large-scale exploitation for the world market has been permitted “during
the last five years", ie. since about 1976. However, almost half of the
animals collected are still sold within the USSR Collecting in Kazakhstan
tends to take place soon after the start of aestivation, so a greater
Proportion of males than females are caught (they cease activity before
females); they are also preferred by collectors because of their smaller
size. However, it seems that collecting can lead to complete removal of
tortoises from large areas; because of the restricted nature of tortoise
movement the re-colonisation of such areas may take a long time or may not
take place at all (Brushko and Kubykin, 1982).

Decline in the USSR is also attributed to the extension of mechanised
agriculture into hitherto untouched terrain, leading to direct injury during
ploughing and other operations, and to the loss of suitable habitat. In

32

Testudo horsfieldii

much-modified areas, tortoises are confined to field edges and to remnants of
suitable habitat. Significant juvenile mortality is caused by road traffic
and fire, and (juveniles) by trampling by livestock. Tortoises are said to
avoid areas with heavy pasturage of cattle and sheep (Brushko and Kubykin,
1982); on the other hand, in Pakistan tortoises are said to be oblivious to
the presence of goats (Minton, 1966).

Table 1. Minimum net imports of T. horsfieldii reported to CITES

1980 1981 1982 1983 1984 1985
Austria : - - 70 = = =
Argentina - - - - 4 6
Belgium 5000 ~ 7000 10000 - -
Bermuda - - - 6 - -
Canada - - - 4 - -
Denmark 1000 3250 5910 - - -
German D.R. - - 15000 15000 - 20000
Germany, F.R. 41146 4599 62940 29835 4 3
Hungary - - - - - 4000
Italy = - = = 40314 =
Japan - - 12 50 - -
Kuwait - - 200 30 - -
Luxembourg - - 50 100 - -
Netherlands - - ~ - 1 -
Saudi Arabia - - 30 270 ~ -
Spain - 5060 5300 400 - 2
Sweden - - - - 2 -
Switzerland 850 - 120 185 - -
UK - i - 6 10 -
USA 4 750 873 320 - -
Country unknown - 65000 65000 - - -
TOTAL 48000 78660 97512 56206 40335 24011

a

Table 2. Reported countries of origin and quantities of transactions reported
to CITES.

1980 1981 1982 1983 1984 1985

eee ODD

a. Countries having wild populations of T. horsfieldii

Afghanistan - = = 6 = ce
USSR 48000 78660 97500 56206 14 24008

b. Countries without wild populations of T. horsfieldii

German D.R. - - - = 4* es
Germany, F.R. - - 12 60 = =
Turkey - - - - 40314 -
Country unknown - - 5 - 3 3

*captive bred

pa

33

Testudo horsfieldii

INTERNATIONAL TRADE See preceding section for an outline of collecting and
trade in the USSR; virtually all horsfieldii in trade reportedly originate
from this country. Data from this major part of the species's range suggests
that populations can withstand at least a moderate level of harvest, but, in
conjunction with evidence for habitat modification and the large domestic
trade in the USSR, it may be suspected that levels of international trade
prevailing in the early 1980s are excessive. See Conservation Measures
section, below, for proposals to mitigate the effects of exploitation.

The 1984 transaction appearing in the above tables, involving 40 314 tortoises
supposedly shipped from Turkey to Italy is anomalous; these animals may in
fact have been of one or both the Testudo species occurring in Turkey
(horsfieldii does not), or may be horsfieldii in transit from USSR, which
otherwise had an unusually small export figure for these years.

CONSERVATION MEASURES

China Nominally protected by legislation in China (hunting prohibited)
(Anon., 1984).

Pakistan Occurs in Hazarganji-Chiltan National Park near Quetta (Shafique,
1984).

USSR Present in the Badkhyz and Repetek protected areas in southern

Turkmenistan, USSR The following measures (paraphrased) have been proposed as

a basis for rational utilisation of T. horsfieldii in Kazakhstan (Brushko

and Kubykin, 1982).

1. Continue and extend surveys of tortoise population, status and ecology.

2. Ban the sale of tortoises as household pets, thus halving the total
harvest (N.B. from the context, it seems that this should be interpreted
as meaning a ban in the USSR, not a ban on international trade for the pet
market); limit the collection of males and immatures; prohibit total
harvest collecting thus leaving a portion of all populations undisturbed.

3. Prehibit collecting before May in order to allow breeding and weight
recovery.

4. Prohibit continued collecting at single sites, but collect in rotation,
with intervals of several years.

CAPTIVE BREEDING A recent inventory (Slavens, 1985) lists 133 individuals
in 26 collections in 1984. Successful breeding in captivity seems to be rare,
but is recorded at Tel Aviv in 1980 (Olney, 1983).

REFERENCES

Aitchinson, J.E.T. (1889). The zoology of the Afghan Delimitation Commission.
Transactions of the Linnaean Society, London, Series 2, Vol V, Zoology
Part III, pp. 53-142.

Anderson, S.C. (1979). Synopsis of the turtles, crocodiles, and amphisbaenians
of Iran. Proceedings California Academy Science 4th Series 41: 501-528.

Anon. (1984). The four-toed tortoise of Xinjiang. China Pictorial, No. 3.

Bannikov, A.G., Darevskii, I.S., and Rustamov, A.K. (1977). Zemnovodnye i
Presmykayshchie SSR, Izdatel’stvo "Mysl", Moscow (2nd edition).

Brushko, Z.K., and Kubykin, R.A. (1982). Horsfield's Tortoise (Agrionemys
horsfieldi Gray, 1844) and the way of its rational utilization in
Kazakhstan. Vertebrata Hungarica 21: 55-61.

Frolov, V.E., Makejev, V., and Bozhanskii, A. (1985). Distribution of the
Testudo horsfieldi in the south of the Turkmen SSR. Paper presented at
Third Ordinary General Meeting of Societas Europaea Herpetologica, Prague,
August 1985. Abstract published in programme booklet, full text in press.

34

Testudo horsfieldii

Ghalib, S.A., Rahman, H., Iffat, F., and Hasnain, S.A. (19807). A checklist

of the reptiles of Pakistan. Records of the Zoological Survey of
Pakistan 8: 37-59.
Khozatsky, L.I., and Mlynarski, M. (1966). Agrionemys - nouveau genre de

tortues terrestres (Testudinidae). Bulletin of the Academy of Political
Science, Series Science Biology 2: 123-125.

Minton, S.A. (1966). A contribution to the herpetology of West Pakistan.
Bulletin of the American Museum of Natural History 134(2): 27-184.

Olney, P.J.S. (Ed) (1983). Reptiles bred in captivity and multiple generation
births, 1981. In, International Zoo Yearbook, 22.

Shafique, M. (1984). Hazarganji-Chiltan National Park. WWF-Pakistan 3(4):

8-11.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in
captivity, current January 1, 1985. F.L. Slavens, Seattle, Washington.
Toynbee, A.J. (1961). Between Oxus and Jumna. Oxford University Press,

London.

35

SOUTH AMERICAN RIVER TURTLE Recommended list: 1
[Problem]

Podocnemis expansa (Schweigger 1812)

Order TESTUDINES Family PELOMEDUSIDAE

a

SUMMARY AND CONCLUSIONS One of the largest species of freshwater turtle,
formerly very abundant in the Amazon and Orinoco drainages of Brazil, Bolivia,
Colombia, Peru, Venezuela, and possibly Ecuador and the Essequibo in Guyana.
Now extremely depleted throughout the range and locally extinct. Inhabits
lakes and major rivers, moving into flooded forest to feed on vegetation and
fruits. Nests in large aggregations on sand banks exposed in the dry season.
The primary cause of its decline is over-exploitation for meat and eggs.
Although protected in all countries within its confirmed range, it is still
hunted illegally and sold at exorbitant prices ($150 in Manaus).

CITES reports reveal few international transactions, with the exception of one
shipment of 1292 skins in 1981. This may well represent misidentification of
sea turtle skins. International trade is probably of little significance
compared with the levels of domestic exploitation. Some cross-border smuggling
of turtles for meat occurs around the Amazon basin.

There is little evidence of international trade in this species. The severely
depleted state of all remaining wild populations means that any commercial
trade is potentially damaging. Therefore Appendix I listing may well be
appropriate to help control the current low level of illegal trade and to
ensure that no new trade routes emerge.

DISTRIBUTION Widespread in the Amazon, Orinoco and Essequibo drainages.

Bolivia Found in the Mamoré and Guaporé Rivers (Pritchard and Trebbau,
1984).

Brazil Formerly found thoughout the Amazon drainage system, but now
virtually extinct in the upper Amazon (Pritchard, 1979), the chief breeding
areas are now on the rivers Trombetas, Branco, Purus and Kingu (Alho, 1985).

Colombia Found in the Solimoes, Putumayo and Caqueta rivers in the Amazon
drainage; and in the Rio Meta and its tributaries in the Orinoco system
(Pritchard and Trebbau, 1984). The Cahuinari watershed, in the Caqueta system,
is now the main refuge in Colombia (Hildebrand, 1985).

Ecuador Said by Pritchard and Trebbau (1984) to be possibly present,
although no specimens exist; the occurrence was confirmed by Hoogmoed (in

Jitt., 26 August 1986). Iverson (1986) mapped one occurrence on the Peruvian
border.

French Guiana Recorded by Groombridge (1982) as possibly present, but
Pritchard and Trebbau (1984) concluded that there was no good evidence that
the species had ever occurred in the country. Fretey (in press) mentions a
specimen captured at Ilet Bache in the Orapak Estuary in the Paris Museum
(MNHNP 1980-1462) but indicated that confirmatory observations were needed.

Guyane Said to be present in several rivers (Mittermeier, 1978), including
the Essequibo system (Pritchard, 1979). However there are no authenticated
records from the country, although shells in the possession of indians have

been observed at Lethem, on the Takutu River, a tributary of the Rio Negro
(Pritchard and Trebbau, 1984).

36

Podocnemis expansa

Peru Found in the Amazon River system, including the Rio Maranon and Rio
Ucayali, although it is said to have been extirpated from the region of

Iquitos (Dixon and Soini, 1986). The main surviving nesting area is now on the
Rio Pacaya (Pritchard and Trebbau, 1984).

Suriname Pritchard and Trebbau (1984) concluded that there was no good
evidence that the species had ever occurred in the country. This was
corroborated by Hoogmoed (in litt., 26 August 1986).

Trinidad and Tobago Individuals are sometimes washed up on the south coast
of Trinidad by the Orinoco in flood, but no breeding has ever been recorded
(Pritchard and Trebbau, 1984).

Venezuela Found thoughout the Orinoco drainage system from the delta to
Boca Mavaca in the upper Orinoco; mostly in the main Orinoco, but some ascend
the Rio Capanaparo, Rio Cinaruco, Rio Meta, and occasionally the Rio Apuré;
the main surviving nesting beach is on the Orinoco, downstream from Puerto
Paez, near the Colombian border. There is no evidence of exchange with the
population in the Amazon via the Casquiare Canal (Pritchard and Trebbau,
1984). Introduced to Lake Valencia in the Andean Highlands (Pritchard, 1979).

POPULATION Very abundant in previous centuries (Bates, 1863; Goeldi, 1906;
Mittermeier, 1975; Smith, 1979), Podocnemis expansa was already rapidly
declining in the latter half of the nineteenth century, and is now severely
depleted throughout most of its range.

Bolivia No infromation.

Brazil In Brazil, the core of the range, many former nesting beaches are
deserted (Vanzolini, 1967), it is rare today to find a single P. expansa in
the Upper Amazon (Mittermeier, 1975). The species was still plentiful enough
in the mid-19th century on the Rio Madeira for example, for the gathering of
nesting females to impede river traffic, ana on one occasion (also on the
Madeira) rows of turtles eight to ten deep stretched along the waterside for
six to seven miles (Smith, 1974). Similarly, in the late eighteenth century
P. expansa were reported to be exceptionally large and abundant around
Itacoatiara (Amazonas), they were still an important dietary item in the
mid-nineteenth century (Smith, 1979), but today they are virtually eliminated
and the few that appear on the market come mainly from the Rio Uatuma, 80 km

away (Smith, 1979). The Instituto Brasileiro de Desenvolvimento Florestal
(IBDF) has embarked on a programme to prevent hunting and to protect
P. expansa nesting beaches in the Lower Amazon. Between 1976 and 1978 at

Leonardo Beach on the Rio Trombetas, there is reported to have been an
increase of around 50% in the number of hatchlings produced (Groombridge,
1982). However, Ayres and Best (1979) suggested that such apparent increases
might reflect nesting females moving into the relatvely undisturbed nesting
beach from other areas, as there was not sufficient time for the reproductive
segment of the population to have increased. More recently, this view and the
data on which it was based have been questioned as there is still considerable
human disturbance. In 1983, 1984 and 1985, there was virtually no successful
nesting on the Rio Trombetas. In 1985, out of a previous pcpulation of
6000-8000 females on the Rio Trombetas, only 200-300 females nested and only
one nest is known to have hatched successfully, the remaining nests having
been plundered by the local people (J.A. Mortimer, in litt., 18 October
1986). The most recent estimates of the main breeding populations are given
in Table 1. Groombridge (1982) reported that population numbers appeared to
be stable on the Guaporé, Branco, Purus and Jurua, and increasing on the
Trombetas, Tapajos and Xingu. In view of the fact that the population on the
Trombetas still appears to be under severe pressure, the security of
populations in other rivers deserves more critical scrutiny.

37

Podocnemis expansa

Table 1. Estimates of the numbers of nesting females and hatchlings in
Brazil. Figures in 1979 from Padua (in litt.) cited by Groombridge (1982),
for 1978-1982 from Alho (1985), and for 1985 from Mortimer (in litt., 18
October 1986).

River Hatchlings Hatchlings Nesting Nesting

females females

1979 1978-1982 1978-1982 1985
Purus 49000 104300 1117
Jurua 80000 21400 291
Guapore . 12000 11470. 226
Branco 153000 180756 1935
Tapajos 18000 18566 353
Xingu 146000 98450 1859

Trombetas 480786 * 393345 5184 300

* data from Leonardo Beach in 1978 (Alho et al., 1979).

Colombia Populations of P. expansa have declined drastically in Colombia,
having disappeared from most of the rivers in which they used to occur. In the
Orinoco system they used to be common 20 years ago on the Rios Manacacias,
Ete, Casanare and Meta, reaching to within 25 km of Villavicencio. There is
still one nesting beach on the Rio Meta, below Orocué (Pritchard and Trebbau,
1984). In the Amazon drainage the Caqueta River is practically the only one
where they still exist, nesting at beaches between Pedrera and Araracuara. The
decrease in numbers is particularly noticable each year in the nesting season
(Hildebrand et al., 1983). An intensive study was initiated in 1983 in
conjunction with INDERENA (Instituto para el Desarrollo de los Recursos
Naturales Renovables) and Fundacion Estacion de Biologia Puerto Rastrojo. 34
breeding beaches were located, and four main beaches were selected for
intensive study. On these four beaches, 87 nests were located in 1983/84, and
43 in 1984/85 (Hildebrand, 1985).

Ecuador No information.

Peru The species is rare in most areas of Amazonian Peru, having been
extirpated from the region of Iquitos, where it was once common (Pritchard and
Trebbau, 1984; Dixon and Soini, 1986). Viable populations still exist in a
few isolated areas, for example in the National Reserve of Pacaya-Samiria
(Soini, 1980).

Venezuela The overall decline of the species is exemplified by data from
the upper Orinoco in Venezuela. Whereas Humboldt estimated 330 000 animals in
1799 and 1800, numbers were down to an estimated 123 600 in 1945, 36 100 in
1950, and 13 800 in 1965 (Mittermeier, 1978; Ojasti, 1971). Surveys in 1981
indicated that the population was only one-third of that estimated in 1965.
The site of this investigation, Playa del Medio on the Rio Orinoco, together
with Pararuma beach are the major nesting areas for P. expansa in
Venezuela. The 1981 season was atypical climatically, with unexpected summer
flooding, and it is possible the results were affected by this; however, the
results are thought to adequately represent the general situation in
Venezuela. Pararuma beach, formerly one of the most important sites in
Venezuela, had only 30 nesting animals in 1981 (Groombridge, 1982). Another
population, on the Rio Meta along part of the Colombia-Venezuela border, is
now also greatly reduced although it was still possible to see aggregations of
up to 6000 turtles basking on the beaches even recently (Pritchard and

38

Podocnemis expansa

Trebbau, 1984). The species is considered to be threatened or in danger of
extinction (Venezuela CITES MA, 1987).

HABITAT AND ECOLOGY A large freshwater turtle, by far the largest of the
living Podocnemis species, a maximum length of 89 cm has been recorded,
(Pritchard and Trebbau, 1984) although the mean length of a more typical

sample of 38 was around 70 cm (Vanzolini, 1967). Males are smaller than
females. Individuals may reach 50 kg (Mittermeier, 1978). Occurs in large
rivers, oxbow lakes, and during the flood season, in lakes formed by rivers
and in flooded forest areas (igapos) (Groombridge, 1982). Turtles re-enter

rivers at the start of the dry season as nesting beaches become exposed.

Podocnemis species are omnivorous but predominantly vegetarian, feeding on
aquatic plants and fruits that fall into the water, but also consuming animal
material such as dead fish (Mittermeier, 1978). Best (1984) provides a table
of over 31 species of plant consumed by P. expansa.

The growth rate in the wild has been estimated at 0.5 cm a year for mature
females or 1.5 cm a year for smaller females (Ojasti, 1971). In captivity in
Manaus, hatchlings have grown from a carapace length at emergence of 55 mm
(22 g) to 85 mm (85 g) after one year and 112 mm (187 g) after two (Alho and
Padua, 1982b). Alho (1985) reports extremely fast growth in captivity,
hatchlings attaining 400 g after one year. The age at first breeding is not
known, but it appears that females do not mature until they attain a carapace
length of 50 cm (Alho and Padua, 1982a). Pritchard and Trebbau (1984) suggest
that they may normally not mature until about 15 years old. In captivity they
have been known to breed at 8 years old (Alho, 1985).

The nesting season varies from June - July in the upper Amazon in Brazil, to
October - November in the lower Amazon. In the Iquitos region, and the basins
of the Rio Tapiche and Rio Pacaya, nesting is in August - September. In

Venezuela, the species nests from mid-February to early March, when water
levels in the Orinoco and its tributaries are at their lowest (hatching here
is in early May) (Groombridge, 1982). On the Rio Trombetas in Brazil the
nesting season is in the middle of October, turtles begin to gather off the
principal nesting beaches in September. On the Rio Caqueta in Colombia there
is an extended nesting period from October to about March. Eggs laid in
October and November are particularly at risk from flooding (Hildebrand, 1985).

Sand beaches are required; nesting usually occurs in large aggregations on a
few selected beaches, although individuals may occasionally nest in outlying
areas (Pritchard and Trebbau, 1984, Magnusson, pers. comm.). Basking
behaviour by the females, which can be observed until the end of the laying
season, begins 15 days before the season begins. This behaviour occurs during
the hottest hours of the day, generally from 10.00 until 16.00 hrs. At times
500 animals can be seen basking, with others swimming in the shallow water
with their heads above water, breathing or watching the beach (Alho and Padua,
1982a; Alho, 1985).

The turtles emerge silently from the water, led initially by a few
individuals, and head toward the higher surfaces of the beach. The actual
nesting process is complex, and completion of the nest hole itself may take
around 100 minutes (Vanzolini, 1967). An irregular shallow body pit is first
scooped out by forceful sweeps of the body and scooping with all four limbs.
A deeper pit is then formed, using first one hindlimb then the other, and
swinging the body back and forth through 90° as each hindlimb is used in
turn. The finished pit is 70-100 cm deep and 100-150 cm in diameter at the
mouth. At this stage the turtle's head is about level with the sand surface,
and the egg chamber is then formed at the bottom of the pit, partly using the
hind margin of the carapace. Egg-laying takes around 15-35 minutes, during

39

Podocnemis expansa

this time the body is at an extreme angle, more than 45° and sometimes almost
vertical. Afterwards the nest is filled and the sand is compacted by the
raising and rapid lowering of the plastron onto the surface, before the turtle
returns to the water (Alho and Padua, 1982a). The clutch size normally varies
from 63 to 134 eggs with a mean of 91.5 (Alho, 1985). The largest recorded was
156, laid by a 75-cm (56-kg) female (Pritchard and Trebbau, 1984); in Peru the
range in examined nests was 80-133 (Groombridge, 1982). Research at the Rio
Trombetas in Brazil showed that clutch size was positively correlated with
carapace length. An average of 86 hatchlings emerged (94%) after a mean
incubation period of 48 days (Alho and Padua, 1982a). In Colombia, eggs from
15 natural nests hatched with 69% success, with incubation periods between 50
and 60 days; the temperature in the nest chambers varied from 28°C to 35°C,
with a mean of 31°C (Hildebrand, 1985). The nesting season coincides with the
hottest time of year, and temperatures in the nest chambers at the Rio
Trombetas site were found to fluctuate between 30°C and 39°C (Alho and Padua,
1982a). As in other reptiles, the sex of the hatchlings is influenced by the
incubation temperature, males predominating when the temperature is low (Alho,
1984; Alho et al., in press).

Tagging of turtles in Venezuela indicates that females breed once a year, and
may continue nesting for at least 15 years (Pritchard and Trebbau, 1984).

After emerging the hatchlings move straight down the beach, running the
gauntlet of assembled predatory birds, and into the water, where they become
prey to a variety of predators, ranging from fish and caimans to herons,
vultures and otters (Best, 1984). There is very little natural predation of
eggs in the nests, but predation of hatchlings is heavy. Alho (1985) asserted
that fewer than 20% of the hatchlings return to the beaches to breed.

Podocnemis expansa and P. sextuberculata are notable in that nesting is
restricted to relatively low beaches or bars of pure sand; they are thus
particularly vulnerable to fluctuations in water level at the nest site
(Vanzolini, 1977). For example, in 1973 all P. expansa nests on the Rio
Trombetas (the largest known present-day nesting area in Brazil) were flooded
and destroyed before hatching (Mittermeier, 1978), and in 1980, 99% of the
eggs were destroyed (Alho and Padua, 1982a); at Playa del Medio on the
Venezuelan Orinoco 25-80% of the annual egg production may be lost in this
Manner (Mittermeier, 1978). In Venezuela this factor certainly appears to be
the principal source of egg and _ hatchling mortality; natural non-human
predation seems to be much less significant (Groombridge, 1982). Whereas most
Podocnemis nest singly or in small groups, P. expansa is exceptional in
nesting principally in large, synchronized aggregations, although some
solitary nesting also takes place. The social nesting of P. expansa recalls
the synchonized mass nesting (arribada) of most populations of marine Olive
Ridley Turtles (Lepidochelys olivacea). However, it differs in several
respects. For example, in P. expansa, nests are concentrated in a
restricted area of beach, and the nesting females are highly tolerant of
disturbance by other females (Vanzolini, 1967). The reverse is the case in
gregarious sea turtles. P. expansa digs deeper nests than other species of
Podocnemis, and so it is possible that the choice of nest site is more
restricted owing to the greater danger of flooding.

THREATS TO SURVIVAL The major current threat to the species, apart from the
natural mortality caused by flooding of nests, is the very heavy exploitation
of adults (including nesting females) and eggs for food. An adult
P. expansa can provide up to 13-14 kg of meat (Smith, 1974), and this turtle
has always been a valuable food resource for the human population, both the

local inhabitants and through more distant market outlets (Mittermeier, 1978;
Smith, 1974; Smith, 1979).

40

Podocnemis expansa

Although P. expansa is one of several turtles that provided a valuable food
resource (also a medium of exchange and a source of raw materials) for the
indigenous peoples of Amazonia, and is often embodied in tribal mythology, the
intensity of exploitation only rose to critical levels following European
colonisation (Smith, 1974; Smith, 1979). European traders and missionaries
were perhaps mainly interested in P. expansa as a source of oil for cooking,
lighting and other purposes. The finest oil was prepared by boiling fresh
turtle fat, the rest of the turtle was typically thrown away (Smith, 1974). A
coarser oil was prepared by crushing eggs. The extent of this exploitation in
the eighteenth century was enormous; it was estimated that 5000 jars of oil
were harvested annually from three major nest beaches on the Orinoco, each jar
contained 25 bottles, each bottle the contents of about 200 eggs. Depending
on the precise clutch size, this would represent the wasted reproductive
effort of about 400 000 females (Mittermeier, 1975; Mittermeier, 1978). A
Similar intensity of exploitation was maintained in the Brazilian Amazon
(Smith, 1974; Smith, 1979). The females, after laying, were gathered into
artificial enclosures along the rivers to provide a food store, for use
notably when fish were harder to catch during the flood periods; it is
estimated (Smith, 1979) that two million Podocnemis were killed annually in

the mid-eighteenth century in the state of Amazonas alone. Intensive
predation on adults and eggs, sustained over three centuries, has brought
P. expansa to its present severely depleted condition. The survival of the

species may be largely due to the introduction of kerosene and vegetable oils
in the latter part of the nineteenth century, replacing turtle products
(Smith, 1974).

Habitat modification, notably clearance of floodplain forests that provide a
food source for P. expansa and other Podocnemis, and changes in river
regime following construction of hydroelectric dams, constitutes an as yet
unquantified threat (Smith, 1979).

Brazil Trade in P. expansa for human consumption still occurs. In Manaus
important occasions are celebrated with a banquet of turtle, for which very
high prices may be paid (Alho, 1985). The increasing rarity of the species
has raised the market price until this food source is out of reach of those
people who would most need it. Fifty years ago in Manaus, a large
P. expansa cost the equivalent of US$0.01, but the present-day equivalent
may be up to US$80 or $100 (Mittermeier, 1978; Smith, 1979). An animal of
30 kg or more is worth $150 (Groombridge, 1982). In Itacoatiara, much closer
to remaining P. expansa localities, the price is still $60, and probably
fewer than 50 are consumed each year (Smith, 1979).

In 1983 a nesting beach at Monte Cristo, near Fordlandia in Para, was
completely devastated when hunters caught all the nesting females (Alho,
1984). Human disturbance of nest sites is an important factor on the Rio
Trombetas, the problems being compounded by surveying activity for the
hydro-electric dam which is to be built upstream. In spite of protection
which is supposed to be given by the staff of IBDF, of a total of an estimated
200-300 nests laid along the Rio Trombetas in 1985, only one is known to have
hatched successfully, the remaining nests having been plundered by the local
people (J.A. Mortimer, in litt., 18 October 1986).

During regular inspections at Manaus and in control operations on the Rio
Purus, 192 turltes were confiscated by IBDF, of which 19% were P. expansa.
68% of the turltes were female, 31% male, and the rest juvenile. A total of 75
boats were boarded, and 10 of these were transporting illegally acquired
wildlife products (Rebélo, 1984). Turtles caught in Colombia are also
smuggled into Brazil for sale in Tefé (see below).

41

Podocnemis expansa

Bolivia Mallinson (1966, fide Pritchard and Trebbau, 1984) reported that
there was heavy exploitation of P. expansa at Gujara-Mirim, on the Rio
Mamoré, the border with Brazil. During a 6-month season, 3000-4000 turtles
were shipped out of this region to Porto Velho.

Colombia Although there has long been subsistence use of turtles by
indians, the commercial traffic in P. expansa developed mainly in the late
1950s. It became particularly intense in the 1960s, with respites in 1965-1967
and 1971-1976, when the traffic in other wildlife skins (cats, caiman, otters)
caused a reduction in the numbers of turtles taken. Trade in turtle skin is
virtually non-existant, the major commerce being in adult animals, which are
smuggled into Brazil (Hildebrand, 1985).

The local indians along the Caqueta exploit’ all stages in the life cycle of
P. expansa: eggs are removed from the nests, hatchlings are caught as they
emerge, adult females are taken on the nesting beaches, and adults of both
sexes may be caught in the lakes and rivers. A survey of 45 of the 80 families
living along the Caqueta River showed that they destroyed over 250 nests of
eggs and 60 nests of hatchlings in 1983/84, and 200 nests of eggs and 40 of
hatchlings in 1984/85. 250 adult females were caught on the nesting beaches in
the first year and 200 the following year. A further 50 or so animals were
caught along the rivers in each year. It is estimated that this resulted in a
total loss of about 59 000 and 35 000 eggs destroyed in the two years
respectively (Hildebrand, 1985).

In 1984 there were approximately ten boats involved in taking turtles caught
on the Caqueta down the river to Brazil, where they were mostly offloaded at
Tefé. In 1983/84, they took a total of 400 P. expansa, 150 of which were
consumed on the boats before they reached their destination. The following
year only about 70 turtles were taken, owing to improved control measures. It
was thought that the presence of the research team had caused a decrease in
the volume of trade in 1983/84 to half its previous levels. The price of each
turtle rises from CPs700-1000 near the nesting beach, to CPs1500-2500 at La
Pedrera, to CPs5000-10000 in Tefé (US$1 = CPs90). There is no commercial
traffic in eggs or hatchlings, and all of the adult turtles are shipped out
down the river towards Tefé (Hildebrand, 1985).

Venezuela Collection of nesting females was legally prohibited in 1962, and
patrolling of the beaches by the Guardia Nacional has reduced the numbers
taken to low levels. Some turtles are still captured in the feeding grounds by
local fishermen. The creoles tend to use baited hooks and lines, while the
indians prefer harpoons and arrows (Pritchard and Trebbau, 1984). A similar
price rise to that seen in Brazil has occurred in Venezuela (Smith, 1974). On
the illegal market in Venezuela, one animal can fetch Bs400 in Puerto Ayacucho
(capital of the Territoria Federal Amazonas); higher prices can be reached, in
San Fernando de Apure and Caicara del Orinoco for example (Groombridge, 1982).
For a brief period in the early 1960s, hatchlings were exported as pets, but
they seldom survived the low temperatures to which they were exposed, and the
trade was soon stopped (Pritchard and Trebbau, 1984).

The population of P. expansa in Venezuela is thought to be seriously
affected by the number of motorized cargo-boats travelling daily between
Puerto Ayacucho and the lower Orinoco; this traffic causes great disturbance
to the nesting beaches during the breeding season, when turtles are
aggregating, basking and nesting (Groombridge, 1982).

INTERNATIONAL TRADE Apart from the illegal cross-border trade in live
P. expansa for human consumption, noted in the previous section, there are
very few reports of international trade in this species. Between 1980 and 1983
the total trade reported to CITES amounted to 1437 skins, five live animals

42

Podocnemis expansa

and three shells, with a few manufactured products such as handbags. No trade
in these products was reported in 1984 or 1985 (Table 2). The great majority
of this trade comprised a single shipment of 1292 skins, country of origin
Colombia, exported from F.R. Germany to Italy in 1981. The skin of
Podocnemis is normally considered to be of little value and it is possible
that these were actually the skins of marine turtles which were declared as
P. expansa to evade CITES controls (P. Dollinger, pers. comm.). Were it not
for this shipment, P. expansa would not have featured in the survey of
significant trade in Appendix II species. In view of the extensive
exploitation for human consumption, it would appear that international
commercial trade in skins, if it occurs at all, is insignificant.

Table 2. Minimum net imports of products of Podocnemis expansa reported to

CITES.

_ EE ae eee ee Ss ee
1980 1981 1982 1983 1984 1985

a ee eee eee ee ee

Canada ~ 8 h'bags - ~ - -

Italy 35 skins 1292 skins - - - -

Japan - - 3 live 2 live - -

UK - 16 h'bags 1 shell - -

USA 2 shells 1 shell ~ 1 shell - -

CONSERVATION MEASURES The protection status of P. expansa is summarised
in Table 3, where it can be seen that it is nominally protected throughout all
its confirmed range. This legislation is difficult or impossible to enforce,
and effective protection is largely limited to the major nesting beaches, with
occasional attempts to control river traffic. Exploitation is unchecked in
other parts of Amazonia (Mittermeier, 1978); in Peru, for example, where
legislation simply means that trading continues covertly (Groombridge, 1982).

Table 3. Legal prohibition on the hunting, internal trade and commercial
export of P. expansa. Dates are those on which the legislation came into
force. A - All live animals & parts; P - Allowed under permit (source, mostly
from Fuller and Swift, 1985).

CITES Hunting Trade Export
___ Ee
Bolivia 1979 A 1979 A 1979 A 1979
Brazil 1975 A 1967 A 1967 A 1967
Colombia 1981 A A A
Ecuador 1975 - - A 1981
Guyana 1977 - - P
Peru 1975 A 1973 A 1973 A 1973
Venezuela 1978 A 1962 A 1962 A 1962

Ee OOOO

The primary requirements in future are to limit continuing over-exploitation,
continue and extend protection of nesting beaches, and continue ecological
studies with a view to rational management. It has _ been suggested that
controlled exploitation is a preferable option to total protection in Brazil,
and outline plans for a ranching scheme have been put forward (Mittermeier,

43

Podocnemis expansa

1978; Alho, 1984; Alho, 1985). Action in Brazil, Peru and Venezuela provides
an essential foundation. Specific data should be sought on the movements and
fate of hatchlings translocated in the IBDF programme, to aid in design of
future conservation action (Groombridge, 1982).

The IUCN/SSC Freshwater Chelonian Specialist Group plans highest priority
projects on the ecology and conservation of this and related species.

Brazil Nesting beaches are supposedly protected by the Instituto Brasileiro
de Desenvolvimento Florestal (IBDF) on the Rios Trombetas, Xingu, Tapajos,
Guaporé, Branco, Purus and Jurua. But the efficacy of the protection, on the
Trobetas at least, has been cast into doubt (J.A. Mortimer, in 1litt., 18
October 1986). The protection planned . extends from the pre-nesting
aggregation of females until hatching of the eggs. Hatchlings are protected
from predation in enclosed waters until full absorbtion of the yolk sac
(assumed to be the greatest attractant to predators); they are then released
in shallow water near lakes, far from predators congregated at the nest
beach. This work is financed by Programas de Polos Agropecuarios e Minerais
de Amazonia (POLAMAZONIA). The IBDF, assisted by the Universidade de
Brasilia, is collecting data on population size and trends, migration,
hatchling and adult behaviour, etc., and attempting to develop an economic
model relating to the feasibility of rearing turtles (Groombridge, 1982;
Rebélo, 1984; Alho, 1985). Plans for a comprehensive ranching and release
programme were put forward by Alho (1985). At present there are no data on
the effects of the IBDF programme of translocating hatchlings to lakes away
from the nest beach. Doubts have been expressed as to the likelihood of such
hatchlings finding the traditional nesting areas or establishing a new site,
and the success of the programme cannot be confirmed until translocated
hatchlings have been observed to breed and nest successfully (Groombridge,
1982). Sporadic checks are made on boats passing up and down the Amazon in
Brazil (Rebélo, 1984), but turtles are still sold openly in the market at
Manaus.

Colombia A research project was established on the Caqueta river in 1983.
Major nesting beaches were located and monitored. Eggs were collected from
nests about to be flooded, and were moved to new sites to complete their
incubation. Hatchlings from natural nests were also kept in pens for four to
six weeks to reduce neonatal predation; they were subsequently released in
nearby lakes or islands in the river. The presence of the research staff
reduced the numbers of nesting females captured by commercial traders, but it
was thought that this effect would cease when the researchers withdrew, unless
more effective policing could be established. The local indians and other
inhabitants were encouraged to control the exploitation of eggs and
hatchlings, and to ensure that any nests opened were fully utilised to prevent
wastage. They were encouraged not to take females on the nesting beaches
before laying had been completed. They were also persuaded not to work with
Brazilian turtle dealers; and river patrols intercepted several boats bound
for Brazil. This was reflected in a five-fold decrease in the number of
turtles smuggled downriver in 1984/85 (Hildebrand, 1985).

For the future it was thought essential that the research programme on the Rio
Caqueta should continue, in order to encourage the participation of the local
inhabitants in the conservation measures, and to deter the commercial turtle
traders from returning. More effective inspection and control of Brazilian
river traffic was advocated. It was suggested that the Cahuinari watershed be
turned into a national park to ensure continued protection (Hildebrand, 1985).

Venezuela The effect of rearing and releasing hatchlings in Venezuela in
the 1950s and 1960s was uncertain and the schemes were abandoned (Smith,

44

Podocnemis expansa

1974). At present in Venezuela, hatchlings are sometimes rescued from nests
in danger of flooding, but the effectiveness of this is unknown. The major

nesting beaches in Venezuela are protected by the National Guard (Groombridge,
1982).

CAPTIVE BREEDING Captive breeding has been achieved at the Museu Goeldi in
Belem (Alho, 1985). The majority of the other schemes to rear P. expansa in
captivity involve the use of eggs or hatchlings taken from the wild (Alfinito,
1980). Experience of the incubation of eggs in artificial nests has been
documented by Hildebrand (1985). Eggs, threatened by flooding, were moved to
new sites to complete their incubation. Of 15 clutches moved in this way,
hatchlings were produced from 11, the hatching success varying from 14% to
92%. The low success was thought to be associated with problems of maintaining
temperature and humidity.

Apart from the head-starting programmes in Brazil and Colombia, artificial
rearing of hatchlings has also been attempted in Brazil at Rio Trombetas,
Manaus and Brasilia. High mortality was experienced in Brasilia owing to the
low temperatures, and the hatchlings had to be moved to indoor tanks, after
which they grew at rates similar to those experienced at the two warmer sites
(Alho and Padua, 1982b).

The Centro Experimental para Criagao em Cativiero de Animais Nativos de
Interesse Cientifico e Economico (CECAN), established in 1977 by IBDF and the
Ministerio da Agricultura outside Manaus, originally had plans to keep
P. expansa, but these were subsequently abandoned.

Young P. expansa are commonly kept as pets by people living in Amazonia
(Alho and Padua, 1982b). A survey of 260 zoological collections in 1985
revealed a total of 18 P. expansa in six different collections (Slavens,
1985).

REFERENCES

Atfanieco, J. (1980). A tartaruga verdadeira do Amazonas - sua criacao.
Belém, FCAP. Servico de Documentacao e Informacao. FCAP Informe Tecnico
No.5.

Alho, C.J.R. (1984). A ciéncia do manejo da fauna silvestre. Revista
Brasiliera de Tecnologia 15(6): 24-33.

Alho, C.J.R. (1985). Conservation and management strategies for commonly
exploited Amazonian turtles. Biological Conservation 32: 291-298.

Alho, C.J.R., Carvalho, A.G., and Padua L.F.M. (1979). Ecologia da tartaruga
da Amazonia e avaliacao de seu manejo na Reserva Biologica de Trobetas.
Brazil Florestal 9(38): 29-47. IBDF, Brazil.

Alho, C.J.R. and Padua L.F.M. (1982a). Reproductive parameters and nesting
behaviour of the Amazon turtle Podocnemis expansa (Testudinata:
Pelomedusidae) in Brazil. Canadian Journal of Zoology 60: 97-103.

Alho, C.J.R. and Padua L.F.M. (1982b). Early growth of pen-reared Amazon
Turtles Podocnemis expansa (Testudinata, Pelomedusidae). Revista
Brasiliera de Biologia 42(4): 641-646.

Alho, C.J.R., Padua L.F.M. and Danni, T.M.S. (in press). Temperature-
dependent sex determination in Podocnemis expansa (Testudinata:
Pelomedusidae). Biotropica.

Ayres, J.M., and Best, R.C. (1979). Strategies for the conservation of the
fauna of Brazilian Amazonia. Acta Amazonica 9(4), Suppl: 81-101.

Bates, H.W. (1863). The naturalist on the River Amazon. John Murray,
London.

45

Podocnemis expansa

Best, R.C. (1984). The aquatic mamamals and reptiles of the Amazon. In: Sioli,
H. (ed), The Amazon. Limnology and landscape ecology of a mighty tropical
river and its basin. Dr W. Junk, Netherlands, pp. 371-412.

Dixon, J.R. and Soini, P. (1986). The reptiles of the Upper Amazon Basin,
Iquitos Region, Peru. Part 1, Lizards anbd Amphisbaenians; Part 2,
Crocodilians, Turtles and Snakes. Milwaukee Public Museum.

Fretey, J. (in press). Les tortues de Guyane Francaise.

Fuller, K.S. and Swift, B. (1985). Latin American wildlife trade laws,
Second Edition. World Wildlife Fund-US, 392 pp.

Goeldi, E.A. (1906). Chelonios de Brasil. Boletim do Museu Paraense Emilio
Goeldi 4: 699-756.

Groombridge, B. (1982). The IUCN Amphibia - Reptilia Red Data Book, Part 1,
Testudines, Crocodylia, Rhynchocephala. IUCN, Galnd Switzerland, 426 pp.
Hildebrand,-P. von (1985). Biology and conservation of Podocnemis expansa in
the Rio Caqueta of eastern Colombia. Final Report. Fundacion Estacion de

Biologia Puerto Rastrojo.

Hildebrand, P. von, Andrade, G.I. and Defler, T. (1983). Project proposal -
Requests for funds from WWF and IUCN.

Iverson, J.B. (1986). A checklist with distribution maps of the turtles of
the world. Earlham College, Richmond, Indiana.

Mallinson, J.J.C. (1966). The river turtles of the Amazon. International
Turtle and Tortoise Society Journal 1: 34-35.

Mittermeier, R.A. (1975). A turtle in every pot. Animal Kingdom 78(2):
9-14.

Mittermeier, R.A. (1978). South America's river turtles: saving them by use.
Oryx 14(3): 222-230.

Ojasti, J. (1971). La tortuga arrau del Orinoco. Defensa de la Naturaleza
1(2): 3-9.

Pritchard, P.C.H. (1979). Encyclopedia of Turtles. T.F.H. Publications,
New Jersey and Hong Kong, 895 pp.

Pritchard, P.C.H. and Trebbau, P. (1984). The turtles of Venezuela. Society
for the Study of Amphibians and Reptiles, Oxford, Ohio, 403 pp. + plates.

Rebélo, G.H. (1984). Projeto Quelénios. Relatorio No. 01/84. IBDF Unpublished
Report, 7 pp.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Washington, 341 pp.

Smith, N.J.H. (1974). Destructive exploitation of the South American River
Turtle. Yearbook of the Association of Pacific Coast Geographers 36(c):
85-102. Oregon State University. (Reprinted 1975 in Chelonia 2(5):
3-9,30).

Smith, N.J.H. (1979). Aquatic turtles of Amazonia: an endangered resource.
Biological Conservation 16(3): 165-176.

Soini, P. (1980). Informe de Pacaya No.2: reproduccion, manejo y conservacion
de los quelonios del genero Podocnemis (Charapa, Cupiso y Taricaya).

Vanzolini, P.E. (1967). Notes on the nesting behaviour of Podocnemis
expansa in the Amazon valley (Testudines, Pelomedusidae). Papéis
Avulsos do Departamento de Zoologia, Secretaria da Agricultura, Sao Paulo
20(17): 191-215.

Vanzolini, P.E. (1977). A brief biometrical note on the reproductive biology
of some South American Podocnemis (Testudines, Pelomedusidae). Papéis

Avulsos do Departamento de Zoologia, Secretaria da Agricultura, Sao Paulo
31(5): 79-102.

46

SPECTACLED CAIMAN Recommended list: 1

[Problem]
Caiman crocodilus (Linnaeus 1758)

Order CROCODYLIA Family ALLIGATORIDAE

SUMMARY AND CONCLUSIONS Widely distributed in Latin America from Mexico to
Paraguay, this small caiman still occurs in appreciable numbers in many swamps
and rivers, occasionally in estuaries. Populations are reported to have
recovered slightly as a result of protection in Ecuador, Brazil and Venezuela,

but are still declining in many countries, including Paraguay, Bolivia and
Colombia.

The species is hunted intensely for skins, over a million of which are in
trade each year, the great majority being exported illegally from their
countries of origin. Recent protection measures mean that skin exports are now
only permitted from El Salvador, Guyana, Venezuela, Bolivia and Suriname.

A major project to assess the distribution and population status of C.
crocodilus is currently under way in South America, and it would be premature
to predict its findings. However it seems pointless to set further controls,
as most trade is currently illegal and the existing legislation is obviously
being flouted by importers and exporters alike.

Taxonomic Note The taxonomy of what may be termed the ‘Caiman crocodilus
complex’ is in a highly confused state. In this analysis four subspecies have
been recognised. C. crocodilus apaporiensis is in Appendix I, and so has not
been covered, but the remaining three, C. c. crocodilus, C. c. fuscus, and
C. c. yacare, have been treated separately as regards distribution, status,
ecology and exploitation. The international trade has been considered together
as there is evidence that the subspecies in trade are regularly
mis-identified. Some of the major taxonomic controversies may be summarised as
follows (references in following accounts):

(1) While a major field authority on South American crocodilians proposed
several years ago that the southernmost populations (C. c. yacare)- are
distinct enough to require full species status (C. yacare), this, opinion
has been ignored by most subsequent workers.

(2) The form ranging from Mexico south through Central America to Colombia and
north-west Venezuela has usually been referred to as C. c. fuscus; it
has been proposed that this name should only be applied to the central and
eastern populations in Colombia and those in north-west Venezuela, while
populations from Pacific Ecuador and Colombia north through Central
America are distinct at subspecies level and named C. c. chiapasius.
While this treatment may be reasonable it is not based-= on
readily-available published work.

(3) Two new subspecies C. c. mattogrossiensis and C. c. paraguayensis have
recently been described, and accepted in a major checklist of the group,
while these taxa are regarded as totally without foundation by others more
familiar with the field situation.

Overall it appears that variation within the C. crocodilus complex is far
from adequately represented by the present taxonomy. These might be

47

Caiman crocodilus

considered fairly academic questions, but these uncertainties do appear to
hinder effective conservation. It is clearly difficult to design or justify
legislation when there is not a consensus position on the taxonomic treatment
of animals from a particular area of concern, nor on which populations are
most distinct and might thus deserve priority protection (if other factors are
equal).

The taxonomic and population status of the whole genus is currently being
assessed in a major project involving many South American countries, but
principally Brazil, Bolivia and Paraguay. When the results of this are
available it should be possible to assess the levels of sustainable harvest
with greater certainty.

DISTRIBUTION

Caiman crocodilus crocodilus (Linnaeus 1758)

Northern parts of South America, east of the Andes.

Brazil Throughout the Amazon basin of Brazil with the exception of a few
southern tributaries (Brazaitis, 1973).

Colombia Occurs to the east of the Andes, in the Amazon and Orinoco
drainages.

Ecuador Found in the Amazonian region (Asanza, 1984).

French Guiana Present (Medem, 1973).

Guyana Present (Medem, 1973).

Peru Found in the Amazonian region (Medem, 1973).

Suriname Found throughout the northern plains and swamps (Glastra, 1983).
Trinidad and Tobago In Tobago the range is now limited to a few rivers
along the south coast and to the lake behind the Hillsborough Dam. Also
present in Trinidad. An adult animal was caught swimming in the sea, 5 km

north-east of Tobago in 1979 (Hardy, 1982).

USA Introduced, probably as a result of escapes of pet animals, there is a
small breeding population in South Florida (Behler and King, 1979).

Venezuela Found throughout the Llanos and the Guyanan region as far as
Amazonia (Maness, 1982; Gorzula and Paolillo, 1986). Also recorded from
numerous localities along the north coast. The north-west of the country is
populated by C. crocodilus fuscus and the exact range of the two subspecies
is not known. However it is thought that the boundary is the Yaracuy River,
with only C. crocodilus crocodilus eastwards from there (Seijas, 1986).

Caiman crocodilus fuscus (Cope 1868) (Brown Caiman)

Extends from southern Mexico and Nicaragua, south through Central America to
the Pacific slopes of Ecuador, Colombia and to north-west Venezuela.

F. Medem (the major authority on the genus Caiman) recommends (Smith and
Smith, 1979) that the populations of the Magdalena and Sinu river systems in

48

Caiman crocodilus

Colombia (extending into north-west Venezuela) be considered distinct from
those of the Colombian Pacific coast, Central America and Mexico (Medem,
1973). By this interpretation, which does not appear to be widely followed,
the coastal form is referred to as Caiman crocodilus chiapasius (Bocourt)
and the name C. crocodilus fuscus is restricted to the Magdalena-Sinu and
Venezuela population.

Colombia West of the Andes, including the Magdalena and Sinu rivers, and
along the north coast to the Venezuelan border; also found on Isla Gorgona
(Medem, 1979).

Costa Rica Occurs along both the Pacific and the Caribbean coasts (smith
and Smith, 1979; Mena Moya, 1978; Wermuth and Fuchs, 1978).

Cuba Introduced into Lanier Swamp, in the south-west of Isla de Pinos
(Juventud) in about 1959 (Varona, 1980; Garrido and Jaume, 1984).

Ecuador Occurs west of the Andes in the extreme north of the country.
El Salvador Present (Serrano, 1978).

Guatemala Occurs along the Pacific coast (Smith and Smith, 1979; Wermuth
and Fuchs, 1978).

Honduras Occurs along both the Pacific and the Caribbean coasts (Wermuth
and Fuchs, 1978; Smith and Smith, 1979).

Mexico Recorded from the Chiapas and the extreme south of Oaxaca, on the
Pacific coast (Smith and Smith, 1979).

Nicaragua Occurs along both the Pacific and the Caribbean coasts (Smith and
Smith, 1979; Wermuth and Fuchs, 1978).

Panama Occurs throughout the length and breadth of the country (Wermuth and
Fuchs, 1978; Panama CITES MA, 1985).

Puerto Rico Introduced to Puerto Rico; there are reports of specimens from
Rio Manati and from areas east of San Juan (Schwartz et al., 1978; Schwartz
and Henderson, 1985).

Venezuela Occurs around Lake Maracaibo, and along the north coast as far as
the Yaracuy River, although the exact eastern limit is not certain (Maness,
1982; Seijas, 1986).

Caiman crocodilus yacare (Daudin 1802) (Yacare Caiman)

Central-southern South America; in the Paraguay River drainage from the
Pantanal and Mato Grosso to north Argentina; also the southern tributaries of
the Amazon.

Although generally treated as one of the several subspecies of Caiman
crocodilus, F. Medem, the authority on the genus, prefers to regard this form
as a full species, Caiman yacare (Medem, 1960). Two recently described taxa
accepted by some authorities (Wermuth and Mertens, 1977), C. crocodilus
matogrossiensis and C. crocodilus paraguayensis, are regarded as
C. crocodilus yacare by Medem (Groombridge, 1982).

49

Caiman crocodilus

Argentina Found from the Chaco to Corrientes as far south as 30°N (Fitch
and Nadeau, 1980). Recorded in Buenos Aires, Misiones, Corrientes, Entre
Rios, Santa Fe, Santiago del Estero, Chaco, Formosa and Salta (Waller, 1987).

Bolivia The taxonomic status of subspecies of C. crocodilus in Bolivia is
under dispute. C. crocodilus yacare undoubtedly occurs in the south of the
country, but some of the populations in the north may be C. crocodilus
crocodilus. The species's distribution includes the southern tributaries of
the Amazon (the Mamoré, Itenez and Beni); also the Guaporé on the
Bolivia-Brazil border. In the south it has been recorded from the drainages of
the Rio Paraguay and the Rio Pilcomayo (Groombridge, 1982).

Brazil Pantanal and Mato Grosso regions of south-west Brazil also the
southern tributaries of the Amazon (the Araguaia above its confluence with the
Tapirape), also the Guaporé on the Bolivia-Brazil border (Groombridge, 1982).

Paraguay Widespread.

POPULATION Populations of C. crocodilus crocodilus generally appear to be
at least stable throughout much of South America, with the exception of
Colombia, Peru and some parts of Guyana. This taxon might well be more
appropriately considered non-threatened, were it not for the reported recent
increase in hunting pressure, and the lack of data - due to trans-—border
smuggling - concerning the extent of exploitation in particular areas.

C. crocodilus fuscus has been reported as depleted in Colombia and Ecuador,
relatively abundant in parts of southern Mexico. Few recent data on
populations in Central America. Further information is given below for those
countries for which it is available.

The overall status of C. crocodilus yacare is uncertain; each of the four
countries in the range is reported to hold some apparently adequate
populations, while the taxon is depleted or extirpated elsewhere. It is under
considerable hunting pressure.

Argentina Reportedly approaching extinction (Groombridge, 1982). However,
a population estimated at 200 000 individuals in 1979 (not confirmed by a full
census) remained in the Esteros de Ibera, an immense swamp in Corrientes,
200 km x 100 km at its widest point (Fitch and Nadeau, 1980).

Bolivia A 1978 census in the Mamoré, Beni and Itenez regions, covering an
area of 693 082 km2, resulted in an approximate total of 3 500 000
individuals. Reportedly relatively common in 1973 in the Rio Madre de Dios.
Recent studies at Lugo Tumi-Chucua (Beni) indicated over-hunting had occured;
immatures were common, an average of ten caimans were seen each night.
Reported to be relatively common in the eastern lowlands in 1976-77, including
the Isiboro-Sécure National Park, also reported extirpated from parts of the
eastern plains (Groombridge, 1982).

Brazil C. crocodilus crocodilus populations are recovering rapidly since
professional hunting was banned (Vanzolini and Gomes, 1979). CC. crocodilus
is now caught fairly frequently in gill nets in floodplain lakes, even near
Manaus, and can be seen without difficulty on tributaries of the Amazon such
as the Rio Negro and Rio Tocantins (Smith, 1980). Magnusson (1982) reported
that it occurred in high numbers in most areas, and that populations appeared
to be limited by habitat preference rather than by disturbance by man. It has
even colonised the artificial lakes formed behind the Transamazonian Highway,

50

Caiman crocodilus

and has increased in areas where populations of Melanosuchus niger have been
reduced by hunting.

C. crocodilus yacare was reportedly relatively common in parts of the Mato
Grosso area, although severely depleted locally. A significant population
occurs in the Caracara Reserve and at the Paraguai Pantanal (Crawshaw and
Schaller, 1980; Groombridge, 1982). In 1977, 30-40 adults were observed from
May to June along a 20-km stretch of road from Cuiaba to Poconé, 20-30 large
individuals were also observed basking at a cattle ranch within 10 km
(Groombridge, 1982). In the Pantanal there has been a drastic decline in
populations of Yacare over the last five years. Among other reasons, this is
causing concern as it is reported to be accompanied by a corresponding
increase in numbers of Piranhas (Hyman, 1985). Brazaitis (1985) reported that
the populations in the Pantanal were barely stable if not declining; most of
the caimans seen were less than 2 m long, smaller than in previous years.

Colombia C. crocodilus crocodilus was reported in late 1970s to be rarely
hunted on the Vaupes and Guayabero-Guaviare rivers because the price of
petrol, salt and air transport to Villavicencio and Bogota were too high.
Juveniles are now seen often, with some adults reappearing. There is a
healthy population in the '‘'Tomo-Tuparro’ Faunistic Territory, a 600 000 ha.
reserve maintained by INDERENA. In 1975, 350 caimans, mostly juveniles were
counted in the lagoons and backwaters of the Capanaparo River (Medem,
undated). However, some hunting continues in the Llanos (Orinoco plains), the
fact that most hides are between 30-60 cm may suggest that adults have been
virtually wiped out from this area (Medem, 1980).

C. crocodilus fuscus was reportedly seriously depleted (Medem, 1971; Medem,
1973), mainly by hunting, and particularly along the Caribbean coast between
Ciénaga Grande and the Magdalena River, also on the lower and middle Magdalena
(Medem, 1980). There is a healthy and abundant breeding population in the
Laguna de Cabrera, Isla Gorgona, comprising at least 28 adults and sub-adults
(Medem, 1979).

Costa Rica Included in a list of animals threatened with extinction,
although C. crocodilus fuscus is said to be abundant in the Rio Tempisque
and in Tortuguero National Park, where it is "almost impossible not to see it”
(Mena Moya, 1978).

Cuba The introduced population has thrived, to the detriment of the endemic
Crocodylus rhombifer because of predation of the young (Varona, 1980;
Garrido and Jaume, 1984).

Ecuador C. crocodilus crocodilus was reported to be not_ seriously
endangered in 1973 (Medem, 1973). In 1983, populations were said to be
thriving in some areas, and indians reported that they had increased in recent
years (Asanza, 1984). However, populations of C. crocodilus fuscus have
been described as seriously depleted; must be considered endangered (Medem,
1971; Medem, 1973).

El Salvador Said to be in danger of extinction, and to be in need of a
10-year ban on hunting to allow the population to recover (Serrano, 1978).

French Guiana Reported in 1973 to be in no serious danger of extinction
(Medem, 1973).

Guatemala The population of C. crocodilus fuscus in the country is
thought to be in the region of 10 000-12 0000 (0. Menghi, vers. comm.).

Si

Caiman crocodilus

Guyana In 1973 Guyana populations were still abundant in most areas but
were declining in areas where habitat had been lost to agriculture,
particularly rice fields (Medem, 1973).

Honduras C. crocodilus is relatively more abundant than Crocodylus
acutus, and its population is said to be “in balance with the habitat” on the
north coast (Honduras CITES MA, 1985).

Mexico Reported to be relatively abundant (Alvarez del Toro, 1974; Medem,
1973). At Chiapas still common in some areas but declining due _ to
over-exploitation (King and Brazaitis, 1971). In recent years exploitation
has diminished somewhat as the species has become more scarce (Alvarez del
Toro, 1974), and it is now considered to be in danger of extinction (Flores
Villela, 1980).

Nicaragua No information.
Panama Said to be moderately common and widespread (Panama CITES MA, 1985).

Paraguay Years of excessive hunting have drastically reduced populations.
However, some individuals remain even in traditional hunting areas. Much of
the range is relatively inaccessible and some populations are likely to
persist (Groombridge, 1982). C. crocodilus was said to be “scarce” in the
Parque Nacional Defensores del Chaco in 1978 (Torres Santibanez, 1978).

Peru. Reported seriously depleted in 1973, judging by a rapid decline in
numbers of hides exported (Medem, 1973). In the Parque Nacional del Manu
C. crocodilus was said to be “frequently observed" in 1979 (Pereya, 1979).

Puerto Rico No information.

Suriname Reported in 1973 to be abundant in suitable habitat in Suriname,
with populations as close to an untouched state as could be found (Medem,
1973). By 1983, some large-scale hunting had depleted populations in the
Coesewijne river and swamp area, although the species was still locally common

along the north coast and its status was not thought to be critical (Glastra,
1983).

Trinidad and Tobago Abundant in parts of Trinidad including the vicinity of
the capital (Medem, 1973). In Tobago it was formerly widespread throughout
most of the rivers and marshes but its range is now limited to a few rivers
along the South coast. It was reported to be quite abundant in the lake behind
the Hillsborough Dam (Hardy, 1982).

Venezuela C. crocodilus crocodilus was described as still relatively
abundant (Medem, 1973); since hunting was made illegal in 1973, the
populations of C. crocodilus have increased in most regions, particularly in
the Llanos, where farmers have complained about the high densities. The
construction of dams and reservoirs on cattle ranches has greatly increased
available habitat. The Ministerio del Ambiente y de los Recursos Naturales
Renovables (MARNR) censused 17 private ranches in the states of Apure, Guarico
and Bolivar in 1982/83 and extrapolated the results to a further 39. They
estimated the total population on these ranches to be 700 000 to 1 000 000
from which a cull of 70 000 was authorised (Quero de Pena, 1984). In Estado
Bolivar it is difficult to find a body of water where this species does not
occur. In the Guyana region Gorzula and Paolillo (1986) estimated the density
of C. crocodilus in lakes and ponds to be 6.64 ha-! or 23.42 km-1 of
shore. Rivers in the same region supported densities of 2.52 km]. It was
not thought that the populations of caiman in this region were under threat,

52

Caiman crocodilus

at the prevailing level of human development (Gorzula and Paolillo, 1986). The
density of C. crocodilus in the state of Anzoategui' (assumed to be
C. erocodilus crocodilus) was estimated in 1984 to be 26.3 km-! in the Rio
Unare, and 1.2 km! in the Rio Neveri. It was thought that the populations
were not only abundant, but may even have increased. Reasons for this included
the creation of new habitat, in the form of reservoirs, and the elimination of
Crocodylus acutus, a potential competitor. In areas where the two species
coexisted the populations of C. crocodilus were lower than elsewhere and
only increased where the numbers of Crocodylus acutus had declined. The only
thing which appeared to be limiting the spread of C. crocodilus populations
was adverse ecological factors, such as saline conditions (Seijas, 1986).

Populations of C. crocodilus fuscus were censused in 1984. A total of 830
was counted in five reservoirs, the mean density being 5.8 indiviuals per km
of perimeter. The density in rivers in the states of Zulia and Falcon varied
from 0.7 km! to 196.6 km-1, the latter in a short stretch of the Rio
Sanare. Large populations of C. crocodilus are also known to occur in the
South of the Lago de Maracaibo, which was not surveyed on this occasion. It
was thought that the populations were abundant throughout the north-coastal
region, and were increasing in some parts (Seijas, 1986).

HABITAT AND ECOLOGY Caiman crocodilus crocodilus seems to prefer. the
quiet waters of lakes, ponds, swamps and marshes, sometimes in brackish waters
(Groombridge, 1982), or along the bends and meandering tributaries of large
rivers where the currents are slight. However it may also occur in fast
flowing water and near rapids (Vanzolini and Gomes, 1979). It is rarely found
in small forest streams, although it occasionally ventures into them

(Magnusson, 1982). C. crocodilus fuscus has similar habitat preference,
occurring in swamps, lagoons, small streams, and tributaries, rather than in
the larger rivers. It does, however, enter brackish and salt water, which

accounts for it occurrence in forest ponds on _ off-shore’ islands.
C. crocodilus yacare prefers open waters, marshy savannah, lakes, lagoons
and rivers; avoids brackish or salt water (Groombridge, 1982).

The maximum size of C. crocodilus crocodilus is about 2.5 m, males growing
larger than females, and also maturing at a larger size. Growth of juveniles
is thought to be in the region of 30 cm a year up to lm, slowing down after
maturity is reached (Rebelo and Magnusson, 1983). C. crocodilus fuscus is a
small form, reaching 1-2 m in length (Groombridge, 1982). C. crocodilus
yacare is reported to attain lengths of 2.5-3m (Fitch and Nadeau, 1979;
Medem, 1973), maximum size recently encountered in Bolivia (Groombridge, 1982)
was 0.88 m snout-vent length in females, 1.2 m snout-vent length in males.

Studies in the Venezuelan Llanos suggest that behavioural adjustment to the
daily pattern of solar radiation plays a major role in determining daily
activity. Individuals tend to submerge during the hottest part of the day,
particularly at the end of the dry season. At this time of year they may also
be found buried in the mud of shallow pools and in leaf litter in shaded parts
of the forest. When water temperatures are seasonally high (29°-30°C)
daylight hours are spent in the water, in the evenings many individuals move
on shore and remain there for much of the night. When water temperatures
fall, individuals spend several hours in the morning on land on sunny days.
Body temperatures are maintained relatively constant (30-33°C during the day,
and 26-30°C at night) during the transition from dry to wet seasons (Lang,
1977).

53

Caiman crocodilus

Juvenile C. crocodilus crocodilus of up to nearly a meter in length feed
entirely on aquatic invertebrates, such as crustaceans and insects. Adults
are opportunistic feeders taking whatever they can kill, ranging from snails
to small deer and pigs (Magnusson, 1982; Maness, 1982; Gorzula, 1978). A
majority of all food items found in the stomach contents of caimans caught
during the wet season in Bolivar State, Venezuela was frogs, especially Bufo
granulosus and Pleurodema brachyops (Gorzula, 1978). There seemed to be
little feeding during the dry season (Gorzula, 1978). Anurans have been
reported in the diet from Venezuela, armoured catfish as the main food item in
Guarico state, whilst the aquatic snail Pomacea ursus and crabs are most
frequently taken in Apure state (Gorzula, 1978). Adult C. crocodilus fuscus
feed on snails, crustaceans and fish (Alvarez del Toro, 1974; Brazaitis, 1973;
Medem, 1973). C. crocodilus yacare: snails, crabs (Brazaitis, P., cited in
Groombridge, 1982) and fish (Fitch and Nadeau, 1980) are major food items;
rodents, snakes and turtles are also consumed (Fitch and Nadeau, 1979;
Crawshaw and Schaller, 1980). In Corrientes a principal prey is Serrasalmus
spilopleura, a species of pirana. This species has increased rapidly in the
last decade in Corrientes, possibly as a result of the decrease in Yacare, and
is reportedly preying heavily on the offspring of other species including game
fish (Fitch and Nadeau, 1980). Where Yacare have been eliminated, the
incidence of schistosomiasis among cattle has risen sharply, possibly in
proportion to an increase in abundance of snails (an intermediate host for the
parasite causing schistosomiasis) upon which Yacare feed (Medem, 1973).

In the case of C. crocodilus crocodilus copulation usually takes place near
the end of the dry season, with nesting from mid-August to early November. In
the Venezuelan Llanos courtship and breeding was observed from May-August
during the early wet season, with nesting beginning in August, and reaching a
peak in September. Nesting strategies are related to rainfall patterns and
vary depending on local conditions. Mound nests, constructed of vegetable
materials, average 117 cm length, 104.5 cm width and 44.5 cm height (Stanton
and Dixon, 1977). 14-40 eggs are laid and hatch in 70 to 90 days, at nest
temperatures of 28° to 32°C. In areas where hunting activity has not altered
behaviour the female defends the nest very actively (Groombridge, 1982). Eggs
vary from spherical to elliptical, average size 63.8 mm x 40.7 mm average
weight 59.9 g. The major egg predator in the Venezuelan Llanos is the lizard
Tupinambis sp. (Stanton and Dixon, 1977). In Venezuelan Guayana young
caimans remained together for around 18 months (Gorzula, 1978).

Research involving C. crocodilus fuscus in Colombia showed that breeding
takes place all year, but usually from January to March. In Chiapas, nesting
is reported to occur between April and September (Flores Villela, 1980).
Fifteen to 30 eggs are laid in a mound nest, constructed by the female from
organic debris. Nests are usually within 30 m of water, but they have been
recorded up to 200 m away. A well developed pattern of maternal care has been
reported from observations made in Mexico. Hatching takes place in 75-80 days
(Alvarez del Toro, 1974).

In Brazil nesting of C. crocodilus yacare takes place during the rainy
season between December and April. Eggs are laid in a mound nest of organic
material, constructed by the female. Average nest size in Brazil was 134 x
117 cm, and 40.5 cm high. Eges are elliptical, white, hard-shelled and
rugose, around 68 x 43 mm, weighing around 73 g. Average egg size varies
considerably between clutches (Crawshaw and Schaller, 1980). Average clutch
size in Brazil is 31 (range 21-38), in Bolivia 33.6 (23-41) (Groombridge,
1982). The peak of hatching is in March. During incubation, the female
visits the nest at intervals, usually at night. Nests are often abandoned
when disturbed by man. At hatching time the female opens the nest, allowing

54

Caiman crocodilus

the hatchlings to escape, there is some evidence that she may crack open the
eggs to ensure a simultaneous hatch. It is not known how long the female
guards hatchlings after they enter the water (Crawshaw and Schaller, 1980).
Major egg predators at Paconé, Brazil include the Coati Nasua nasua and the
Crab-eating Fox Cerdocyon thous, with local reports of predation by tegu

lizard Tupinambis sp. and Capuchin Monkey Cebus apella (Crawshaw and
Schaller, 1980).

The production of excrement by large populations of caimans may form an
important input of nutrients in certain otherwise nutrient-poor aquatic
ecosystems. The decline in caiman populations in certain mouth lakes of the
central Amazon region has been followed by diminished fish populations,
attributed to a decline in the invertebrate prey of young fishes (Fittkau,
1970; Glastra, 1983).

Caiman crocodilus crocodilus is highly adaptable, and is quick to
re-establish itself in habitat vacated by Melanosuchus niger and Crocodylus
intermedius. This may account for the temporary increase in some populations
as Melanosuchus becomes extinct. The species has in fact established itself
in man-made ditches and canals in parts of Florida, USA, and in newly-formed
artificial lakes behind the Transamazon Highway in Brazil (Magnusson, 1982).
In studies carried out at Lago Amana (an effluent of the Rio Japura) and the
Parque Nacional da Amazonia, numbers appeared to be limited by habitat
preference rather than disturbance by man, despite heavy hunting pressure.

The distribution of C. crocodilus fuscus overlaps that of Crocodylus
acutus and Crocodylus moreletii, but C. crocodilus fuscus is_ only
sympatric with C. acutus. C. crocodilus yacare occurs in sympatry with
Melanosuchus niger, when it retreats to smaller creeks and streams to avoid
the larger species, and with Caiman latirostris in the south. It appears to
be capable of out-competing C. latirostris for available habitat (Crawshaw
and Schaller, 1980). However, this may be the result of selective hunting for
C. latirostris, with the consequent expansion of C. crocodilus yacare into
the vacant niche (Fitch and Nadeau, 1980). Yacare are migratory and may
travel overland for considerable distances while moving from stream to stream.

THREATS TO SURVIVAL Legal and illegal hide hunting and the lack of adequate
enforcement to control smuggling, are the major factors contributing to the
pressures placed upon the species (Fitch and Nadeau, 1980; Medem, 1971; Medem,
1973).Although the belly skin of C. crocodilus contains well-developed
osteoderms (Hunt, 1969), renewed hunting pressures are being applied to the
species as the more desirable hides of Melanosuchus niger and true
crocodiles become unavailable. It is reported that commercial interests, with
the renewed legal trade in hides of some populations of Alligator
mississippiensis, have rekindled the demand for crocodilian products in world
fashion. The hide of C. crocodilus is an economically attractive product
with which to meet this demand (Groombridge, 1982).

C. crocodilus yacare produces the most desirable and largest hide of all the
Caiman crocodilus group. Although Yacare does bear ventral osteoderms,
these are not as extensive as in other Caiman crocodilius. Flank hides are
comparable in size and near in quality to the Black Caiman Melanosuchus
niger. Thus, shoes made from Yacare may retail at prices nearly equal to 4H.
niger, are made of more readily available hides, and are cheaper to purchase
raw. Profits for those items become greater, and commercial demand is
therefore great. It is difficult for law enforcement agents to distinguish

55

Caiman crocodilus

small products made from Yacare hide from those made from of other members of
the genus Caiman (Brazaitis, P., cited in Groombridge, 1982).

It is possible that C. crocodilus crocodilus may not be greatly affected by
low levels of hunting because of its small size at sexual maturity (130 cm).
Hunters usually select animals longer than 110 cm, and as the hunting is
seasonal, many females can reach breeding size before they are subject to
hunting pressure. C. crocodilus is therefore thought to be able to withstand
low levels of exploitation better than larger species, such as M. niger,
which is taken at a size much: smaller than that at which it is likely to
breed. However one other aspect of the trade in C. crocodilus skins is that
it is usually accompanied by a small trade in the skins of M. niger. This
this enables populations of M. niger to be exploited at levels below those
which would be commercially viable in isolation, and could cause the
extinction of M. niger (Magnusson, 1982; Rebélo and Magnusson, 1983).

Conversion of habitat for agricultural purposes has caused a decline in some
regions, such as the rice fields in Guyana. Consumption of the meat on a
non-commercial basis by local people is not an important threat at present
(Medem, 1971; 1973; 1980).

Argentina An estimated 20 000 caimans, largely Yacare, are taken annually
from Corrientes (Fitch and Nadeau, 1979). Hunting is usually carried out at
night, either with baited hooks, or with rifles and harpoons. Many of the
larger ranchers are conservationists, and will not allow hunters on their
land, but the surrounding smaller farms often provide a means of access (Fitch
and Nadeau, 1980). In Corrientes much former habitat has been lost to
agriculture or ranching. In areas where C. crocodilus co-exists with
C. latirostris the severe depletion of the latter species by selective
hunting seems to have benefited C. crocodilus (Fitch and Nadeau, 1980).

Bolivia Bolivia is the main tanning country in South America and skins are
imported from surrounding countries for processing before being re-exported
(Groombridge, 1982). From January lst to August 3lst 1976, 124 114 Yacare
hides were exported (Medem, undated). Although Bolivian protective laws are
now more strongly enforced the hide industry remains powerful. Large numbers
of hides are taken from Brazil and Paraguay and are then tanned and exported
from Bolivia (Medem, 1973). Most skins of a shipment of 300 000 caiman hides
from Bolivia intercepted in Florida are thought to have originated in the
Pantanal region of south-west Brazil (Groombridge, 1982).

Brazil Hunting has been illegal since 1967, and although some still occurs,
there is evidence that populations of C. crocodilus have recovered. There is
no direct trade in crocodilian skins through major Brazilian ports. However
because of the size, and remoteness of much of the Amazon basin within Brazil
it is very difficult to enforce the law and hundreds of thousands of
crocodilian skins are transported annually across remote parts of the Colombia
border (Magnusson, 1980). An analysis of reptile skins confiscated in Amazonia
by the Instituto Brasiliero de Desenvolvimento Florestal (IBDF) showed that
the great majority were of C. crocodilus. The size distribution of skins
showed that hunters had apparently selected animals longer than 110 cm
(Magnusson, 1982; Rebelo and Magnusson, 1983).

There is still considerable illicit hunting of C. crocodilus yacare in the
Pantanal, organised on a large scale. The skins are mostly smuggled over the
border in light aircraft to Bolivia and Paraguay, where they are often
bartered for cocaine. In August 1983 a major law enforcement operation was
carried out by the wildlife authorities supported by military personnel and

56

Caiman crocodilus

equipment. Many poachers were arrested, and 1800 Yacare skins were
confiscated, although this is a tiny percentage of the annual illegal harvest.
Each skin is worth US$1-4 to the hunter, who may collect up to 100 in a night,
providing an income considerably in excess of the minimum monthly wage of
US$70. The penalties for poaching are low; many of the poachers arrested were
released, and the excercise has had little effect in controlling the trade
(Hyman, 1985). The poachers are often better equipped than the law enforcement
officers, and new road construction will make access possible all year round.
There was reported to be pressure to allow a legal harvest in 1986, but no
management programme had been formulated, and it was hoped that the plan would
not receive approval. The poaching in the Pantanal was considered to be
largely driven by overseas demand for caiman skin, particularly from France,
F.R. Germany, Itlay and the USA (Brazaitis, 1985).

Colombia Although the export of caiman skins under 1.5 m has _ been
prohibited in Colombia since 1973, a trade in this species continues. In
1974, 556 422 C. crocodilus were exported, 84% under the minimum legal
length (Smith, 1980). Leticia, in the Colombia Amazon, is a major outlet
(Medem, 1973; Smith, 1980). Many skins come from individuals killed in Peru
and Brazil and smuggled over the border. The minimum legal size restrictions
are not enforced (Smith, 1980). Along the Caribbean coast between Ciénaga
Grande and the Magdalena River illegal hunting is common, also on the lower
and middle Magdalena. Almost all hides are of hatchlings or juveniles (Medem,
1980). Some refuge is found in lagoons along the lower and middle Magdalena
Valley, where aquatic vegetation has become more extensive with the decline of
manatees (Medem, 1980). Additionally, hatchlings are killed in large numbers
and are preserved as curios for sale to tourists. For many years,
C. crocodilus hatchlings from Colombia and Ecuador had provided crocodilian
pets for the world pet trade. It is estimated that legal export figures
reflect about one half of the true number of hides exported from Colombia and
Peru, a large number of those animals killed spoil before reaching the
tanneries. In many instances, the animal is killed for its flanks only, the
remainder of the body left to waste (Groombridge, 1982). Marked habitat
destruction has occurred in Colombia including the officially protected
mangroves of the Isla de Salamanca National Park (Medem, 1980).

Costa Rica There is said to be illegal exploitation for skins in Costa Rica
(Mena Moya, 1978).

Ecuador For many years, hunters and traders have been taking skins
illegally over the border into Peru and Bolivia; however, recently, the border
controls have been improved and the trade has declined. There is some
traditional use of caiman products for medicinal purposes (Asanza, 1984). Oil
developments have opened up new towns in Amazonia, and new roads have been
built, with accompanying deforestation. There are reports of increasing
pollution of the waterways by the uncontrolled dumping of oil/water mixtures
(Asanza, 1984).

Honduras Hunting of C. crocodilus is illegal in Honduras unless’ the
hunter operates a farm, and has achieved growth and breeding. No farms were
functioning in 1985 (Honduras CITES MA, 1985).

Mexico Caimans are much persecuted in Chiapas, primarily for the skin
trade. They are protected under legislation which sets closed seasons and a
minimum size of 1.5 m, but in general the laws are not well enforced and they
are considered as a free resource. There are several skin companies which
exploit C. crocodilus fuscus: one company was reported to process 1000 skins
a month. Dried juvenile specimens are also often sold as curios. In 1970,

57

Caiman crocodilus

small ones (60 cm) sold for Ps50, and larger ones (120 cm) for Ps150. In 1980,
corresponding prices were Ps300 and PsS0OO in San Cristobal las Casas, or Ps150
and Ps200 in Tuxtla Gutierrez (Flores Villela, 1980).

Panama Trade in wildlife in Panama has been giving cause for concern since
at least 1978, when "caiman skins" were amongst the species being traded. At
that time there were ten companies involved in the import, export and
re-export of wildlife products. Legislation did not control the shipment of
goods in transit through Panama, and companies were said to import animals
from overseas without adequate documentation, and then to re-export them.
Occasionally additional animals, caught within Panama, would be included in
the re-export consignment without being declared (Vallester, 1978).
Currently, the major threat is more from persecution by people who think that
caimans are dangerous. Hunting for skins is very much less important (Panama
CITES MA, 1985).

Paraguay Many hides have been exported by Paraguay to France, F.R. Germany
and the United States despite national protective legislation. Paraguayan
authorities claim that the export documents are forgeries (Groombridge, 1982).
Paraguay provides an important staging post for skins brought illegally out of
Brazil. Figures collected by the Ministerio de Agricultura y Ganaderia showed
that around 1000 skins of C. crocodilus were exported in 1984 (Acevedo
Gomez, 1987).

Peru A total of 101 641 C. crocodilus hides were exported from Peru from
1962 to 1967 (Smith, 1980).

Suriname Large-scale hide-hunting was reported in two consecutive dry
seasons around November 1978 and 1979 in the Coesewijne Swamp area. This
resulted in the loss of 1500-2000 caiman. Although the species is not legally
protected in Suriname, conservation organisations launched a press campaign to
stop the hunting, and, in 1980, none was reported (Glastra, 1983).

Trinidad and Tobago Abundant in parts of Trinidad, but illegally hunted on
a limited scale. The population in Tobago is not subject to hide hunting
(Medem, 1973).

Venezuela Crocodilians have traditionally been exploited by indigenous
peoples in Venezuela for food, medicinal and religious purposes, but these
uses probably had a negligible effect on the populations (Maness, 1982).
Commercial hide-hunting caused the systematic degradation of caiman
populations to such an extent that the species was declared totally protected
in 1973. Over the next 20 years the populations built up until renewed
exploitation was not only thought feasible, but was demanded by farmers to
allow the control of the growing numbers of caiman. A proposed harvesting
programme was discussed at length at a meeting of the IUCN/SSC Crocodile
Specialist Group which was held in Caracas in 1983, and received provisional
approval for a trial period. In 1982/83 MARNR conducted a survey of 56
ranches, and recommended a harvest of 70 000 caiman, being 7-10% of the
estimated population. Hunting was only allowed on private ranches in three
states, between 1 January and 30 April, and size limits of 1.5 m to 1.7m
snout-tail length were imposed. Export of raw hides was prohibited, and 5-6
tanneries were operating in the country to provide tanned skins for export
(Quero de Pena, 1986). After the harvest had been permitted for two seasons,
it was suspended by a Ministerial Resolution in October 1985. This was
reportedly due to abuse and mismanagement in issuing hunting tags. Tags were
issued in excess of the quotas set, and hunters were reported to have taken
additional skins to make up for those spoiled by improper preservation

58

Caiman crocodilus

(Gorzula, 1985). There is known to be a small amount of illegal hunting taking
place, particularly in the Orinoco Delta, whence skins have been smuggled to
Trinidad and Guyana. Indigenous peoples also hunt alligatorids for
subsistence: in the Amazon Territory they accounted for 30% by weight of the
wild animal meat consumed by the Ye'kwana Indians in one 7-month study. Over
the same period the comparable proportion for the Yanomamé Indians was only
2%. A figure of 2.5% was quoted for the 350 Pemon Indians in Estado Bolivar.
There is very little commercial use of caiman meat, and none was seen being
sold in markets or restaurants, although the Criollo people in the Guyanan
region are known to eat them occasionally (Gorzula and Paolillo, 1986). Caiman
hunting in the Llanos is so easy that populations can be quickly eliminated
from many areas.

The populations of caimans in the north-west of Venezuela probably escaped the
worst of the effects of the over-hunting which afflicted the remainder of the
country in the 1950s and 1960s; one exception was that in the Maracaibo basin,
which probably supported a large industry (Seijas, 1986). There has been no
legal hunting of C. crocodilus fuscus in Venezuela since the ban was imposed
in 1973. When hunting of the other subspecies was temporarily re-started in
1984 in the Llanos, no licences were granted for the north-western states in
which C. crocodilus fuscus occurs (Quero de Pena, 1984). Some illegal
hunting is known to occur, and official figures show that 616 caiman skins
were confiscated in the Maracaibo region in 1982 (Seijas, 1986).

In very dry years populations in savannas may be reduced by as much as 80%
because of mortality in small individuals. However, populations seem able to
recuperate (Gorzula, cited by Groombridge, 1982). Many of man's impacts on
the environment, in particular, the creation of reservoirs, and the digging of
drainage ditches, seem to be mainly beneficial to C. crocodilus (Gorzula and
Paolillo, 1986; Seijas, 1986). The region in which C. crocodilus fuscus
occurs in Venezuela is the most heavily affected by urban and industrial
development, the Maracaibo basin being the main oil-producing area. Some of
the effects may be deleterious to caimans, but other impacts, in particular,
the creation of reservoirs, and the digging of drainage ditches, seem to be
mainly beneficial to C. crocodilus. The species also appears to have
benefited from the removal of competition from Crocodylus acutus, whose
populations have decreased (Seijas, 1986).

INTERNATIONAL TRADE Caiman crocodilus is the second most commonly
exported species of reptile in South America, after Tupinambis. Caiman skin
is traditionally considered inferior to classic crocodile skin, and levels of
exploitation only rose after the supplies of the more valuable crocodilians
became depleted. However its popularity is increasing, partially owing to
improved tanning procedures, and it was reported to be replacing crocodile in
the high-quality handbag and shoe market in the USA in 1985. Caiman products
commanded the same prices as crocodile ones, wallets selling for US$60-180,
shoes for US$475-600, and handbags from US$375 to over US$1000. Most of the
caiman skin is C. crocodilus yacare. Artificial embossed crocodile leather
was reported to be improving in quality, and fetching similar prices to the
real item. This was thought to be associated with a rise in price of real
skin, in turn dependent partially on its decreasing availability (Brazaitis,
1985). Caiman sides were reported to be worth about US$60 each in the USA
(Hyman, 1985). French and Italian reptile leather dealers report falling
demand for reptile skin products. Most of the C. crocodilus fuscus products
available in Italy range from 17 cm to 35 cm belly width (Bodiopelli, in
lJitt., 4 March 1986).

59

Caiman crocodilus

Small numbers of live animals are traded, mainly as pets, but the quantities
are insignificant in comparison with the skin trade.

There have been several reports published recently detailing the international
trade in caiman skins (Anon., 1984; Hemley and Caldwell, 1986), using CITES
and Customs statistics as sources of data. It is impossible to arrive at an
accurate estimate of even the legal trade in caiman skin from CITES reports,
because of the widely varying methods of reporting transactions. Most caiman
hide is shipped as sides or flanks, as this is usually the only part to be
used, but there are also some whole skins in trade. CITES reports contain
reference to both skins and sides, but there is good evidence that the terms
are used interchangably; sometimes it is possible to see that the importer has
reported sides and the exporter skins for what is obviously the same
transaction. Confusion is increased, as the quantity may be reported by
Number, weight, length or area; and finally the taxon may be described to
species or subspecies level. There are thus at least 16 ways in which any
shipment can be reported without giving any false information. Added to which,
it often suits the unscrupulous trader to falsify the subspecies, either to
conceal the true country of origin of the skins, or else to circumvent
protective legislation, such as the USA Endangered Species Act. These factors
all compound to make the analysis of CITES statistics difficult and
potentially inaccurate. This should be borne in mind when considering the
analysis presented in the following tables, but it is thought that they do
give an indication of the order of magnitude of the trade. The CITES reports
are summarised in Tables 1 and 2. As explained in the introduction, the
analysis involves many potential systematic errors, but with caimans there is
an additional error involving double reporting, where imports and exports were
reported in different units, or where the terms "skins" and “sides” were
confused. The actual trade may therefore be lower than the totals reflected
in the tables.

Table la contains an analysis of the minimum net trade in skins and sides of
all subspecies of C. crocodilus, including those reported as "Caiman spp."
The total volume has fluctuated around a value of about three-quarters of a
million. It peaked in 1985, at just over a million skins. The totals for 1980
to 1982 are all higher than those reported by Hemley and Caldwell (1986)
partially owing to different analysis techniques and partially because more
data have subsequently been added. However they are not unrealistically high,
as it has been estimated that the number of caiman skins taken annually in
southern Brazil, Bolivia and Paraguay exceeds one million (Hemley and
Caldwell, 1986). The major net importing countries are the USA, Italy, F.R.
Germany, France, Austria, Switzerland, Hong Kong and Japan. Japan had high
gross imports in all years but also re-exported large quantities, with the
result that it often appeared as a net exporter.

The composition of reported trade in different subspecies is shown in Table
1b. C. crocodilus crocodilus was traded in the largest numbers in all years
except 1982 and 1983, when C. crocodilus yacare took precedence.
C. crocodilus fuscus was mostly traded in low volumes except in 1980, 1984
and 1985, when figures of over 200 000 were reported, exceeding half a million
in 1985. The relative numbers of the subspecies takes on greater importance
when the countries of origin are considered.

The declared sources of the skins of C. crocodilus crocodilus are shown in
Table 2a. Bolivia has been included under countries without wild populations
as, although it may have a few of this subspecies in the north, it is not
thought to have an exploitable population. Up to 1984, the main declared
sources were Bolivia, Paraguay, Colombia and Panama, only one of which,

60

Caiman crocodilus

Colombia, can be correct. Only about 10-20% of the declared trade has been
from countries having this subspecies in the wild. The apparent exports from
Bolivia are comparable with the figures quoted by Hemley and Caldwell (1986)
obtained from the Bolivian wildlife authority, which give exports of 85 551,
62 155 and 29 823 for 1982, 1983 and the first six months of 1984,
respectively. It is thought that a large number of skins of C. crocodilus
yacare are reported as C. crocodilus crocodilus. This subterfuge has been
used to import skins to the USA, where the Yacare is protected by the
Endangered Species Act, and some successful prosecutions have been brought
against skin dealers (Brazaitis, in press). In 1984 and 1985, Guyana and
Venezuela emerged as the major sources, the skins from the latter country
coming from a newly introduced management programme.

The declared sources of the skins of C. crocodilus fuscus are shown in Table
2b. The majority apparently came from Colombia and Panama, although Honduras
and El Salvador emerged as significant sources in 1984 and Guatemala appeared
to export around 347 700 in 1985. All these countries do have wild populations
of this subspecies but these levels of offtake seem excessive. In the case of
Guatemala the total is over twenty times the estimated wild population, and
the skins probably originated in other countries and were re-exported. Panama
is known to be a major entrepot for wildlife trade originating in South
America (Vallester, 1978).

The declared sources of the skins of C. crocodilus yacare are shown in Table
2c. The major sources were Paraguay and Argentina, although Panama and
Colombia had significant exports in 1982 and 1984, and these, if correctly
identified, must represent re-exports. The Yacare thus presents a different
picture to the other subspecies: most of the skins are reported to come from
possible source countries, and the reported levels of exploitation are not
grossly inflated compared with probable wild population sizes. This does not
imply that the Yacare is not being over-exploited, but merely that there is no
incentive to falsify the reported countries of origin for skins declared as
being C. crocodilus yacare. The total volume of trade in skins of this
subspecies is probably far higher, as many of the skins are thought to have
been incorrectly recorded as C. crocodilus crocodilus.

Two outstanding features of the trade in C. crocodilus are its size and the
fact that the great majority of it is illegal. Since 1980, the only legal
sources of skins in South America have been Venezuela (from 1983 to 1985),
Colombia (only stocks held by two companies, prior to 1984), Bolivia (only
C. crocodilus yacare, prior to 1985) and the Guianas. Small amounts of trade
in C. crocodilus fuscus from Central America and Colombia may also be legal.
The only mainland countries currently permitting commercial exports are El
Salvador and the Guianas. Ignoring the illegal and undeclared trade, less than
20% of the declared trade in C. crocodilus up to 1984 was even potentially
legal. Questions of whether the limits on trade set by the producing countries
are within sustainable levels are therefore irrelevant, as the existing
legislation was obviously being flouted by importers and exporters alike. In
1985, there have been some improvements, with a greater percentage of the
trade apparently coming from the legal harvest in Venezuela, but some of the
newly emerging trade routes, in particular Guatemala, Guyana and El Salvador,
together with the continuing exports from Bolivia, Paraguay and Panama, give
cause for concern.

61

Caiman crocodilus

Table la. Minimum net commercial imports of skins and sides of all subspecies
of Caiman crocodilus reported to CITES (2 sides = 1 skin).

1980 1981 1982 1983 1984 1985
Albania - 200 - - - -
Argentina - - - 4103 - -
Australia = - 3 972 - -
= 2 4 628 m2 = =
Austria 37029 47089 36567 20927 37534 35240
= 167 m? 72 m2 355 m2 562 m2 557 m2
- - 6 kg - - 3m
- - 32 m - - -
Belgium 1550 1488 441 156 - 2955
= = 5 m2 = = =
Canada 5004 1316 685 651 4067 8329
75 m2 = S 172 m@ 346 m2 342 m2
China - - - - - 9 m2
Costa Rica - - 4000 - - 244 m
Demark 141 58 - 22 - -
Egypt - 681 1228 2101 - -
Finland 232 - 200 115 - 117
France 94290 248900 1111 - 328336 137569
= 769 m2 110 m2 1042 m2 2604 m2 41 m2
- 20460 kg 1934 m - - -
Germany DR ~ - - 2427 - -
= = = 146 m2 = =
Germany FR 211451 46074 128299 185024 - 14005
197 m2 112 m2 = 788 m2 417 m2 19 m2
- 196 kg - - - -
- 326 m 93 m - - -
Greece - - - 20 785 385
- - - - 23 m -
Hong Kong 4158 6292 15665 13242 34356 10547
356 m@ 664 m2 630 m2 3011 m2 538 m2 1429 m2
134 kg 1354 kg 353 kg 70 m - -
Ireland - - 4 - - -
Italy 358041 173868 322325 305087 151315 513803
é 2269 m2 987 m2 = = 2713 m?
- 16723 kg 8217 kg 6360 kg 423 kg 646 kg
Japan 36325 - - 61884 153 m 94268
- - 127 kg 131518 kg 123846 kg 65797 kg
Kuwait - - = = 18 +
Lebanon 142 - 30 961 27 -
Luxembourg 12 - 10 2 - -
Malaysia - - - - 122 -
Mexico 96 - - 1 kg - -
Monaco 16 - - = = S
Morocco = = = = uf 86
Netherlands 18 9 - 367 15000 -
N. Antilles - - - - 2965 -
New Zealand - 5 27 - 12 -
Portugal 210 446 60 - - -
= = 3 m2 = = =

62

Table la continued.

Qatar
Singapore

S. Arabia
S. Africa

S. Korea
Spain

Sweden
Switzerland

Syria
Taiwan
Thailand
Turkey
UK

USA

Uruguay

Yugoslavia
Unknown

TOTAL

1980

842214
818 m2
5283 m
134 kg

1981

9383
76
123112
43

764871
4034
3840

38786

kg

ke

1982

54
962

674803
9799
3285
8791

1983

kg

734849
m2 8410
m 133
kg 137879

Caiman crocodilus

1984

4
3690
1788

87
7658

m2

104173
40 m2
lm

32

10071

460 m
50
5 m2
20
238

702512
4622 m2
636 m

124269 kg

1985

27
1039
276
1243
241
1302

103870

13
3491
511
129643
19

20

46

1057978
6270
267
66719

nee SEEEEEEIEEES SESE EERE

kg

kg

63

Caiman crocodilus

nn ———_—$—

Table 1b Minimum net trade in skins and sides of three subspecies of
C. crocodilus (2 sides = 1 skin).

1980 1981 1982 1983 1984 1985
ee ee ————————EEEeee
C. c. crocodilus

555492 546189 209889 297372 436988 649730
630 m2 3224 m2 2066 m? 6976 m2 2428 m2 6267 m2
5283 m 3514 m - 363 m 613 m 229 m
134 kg 25429 kg 4793 kg 13168 kg 120981 kg 56065 kg
Cc. c. fuscus
277016 181738 98444 69609 206182 552202
a“ “s 18 m2 125 m 423 kg 10654 kg
- 326 m 93 m - 23 m 264 m
Cc. c. yacare
19320 138358 385307 392826 89032 32703
= 108 m2 401 m 601 m2 2086 m2 3m
= 11085 kg 3785 kg 6052 kg 1432 m

Table 2a
skins
skin).

Reported countries

of origin or export for commercial exports of

and sides of C. crocodilus crocodilus reported to CITES (2 sides = 1

Countries with wild or farmed populations of C. crocodilus crocodilus

Brazil
Colombia

F. Guiana

Guyana
Peru
Suriname

Taiwan
Venezuela

64

1980 1981

104495 71414

= 155
= 1363
6307 8804
276 m2 83
103 kg a

= 12

as 108

93 m2 3000
44554 s

1982

35254

1983

132809
1488 m2
3793

85 m2

1984

835

144127
1466 m2
1550 kg
945

1985

78166
1832
922
1379

kg

121340

270424
3981

Caiman crocodilus

Table 2a continued

1980 1981 1982 1983 1984 1985

Countries without wild populations of C. crocodilus crocodilus

Argentina 4659 10516 = = 1568 4
= 75 m2 = = s if
Bolivia 164666 106530 102067 62574 31326 162733
139 m2 2313 m¢ 1223 m2 772 m@ 2 367 m2
31 kg 21634 kg 533 kg 560 kg = 6626 kg
- - - 48 m - -
Canada 4 3 4 1 < =
El Salvador - - - - 5253 15042
France 26206 46645 4413 28 - -
75 m? = = = 2 =
Germany, F.R. 6958 7305 31 - - -
2 m2 - = = = &.
Haiti 4747 - - 55 175 -
Indonesia 625 298 = 52 m2 = s
Italy 13022 19905 - 300 - -
Japan - - ~ - - 1940
Madagascar 8 ~ - - - -
Netherlands 498 - - - - -
N. Antilles 1 719 kg 450 4 - -
Nicaragua - - - - 1 ~
Panama 59161 7246 7063 34582 88 -
z = 2 4750 kg 2085 kg =
Paraguay 239498 379619 56319 49808 169609 7424
E 962 m2 130 m2 108 m? 95 m2 91 m2
- 14m - 230 m lm -
- 2910 kg - 123444 kg 114374 kg 48389 kg
Singapore - 12 - - - -
S. Africa - - - - 28 -
Switzerland - 89 - - 1076 -
UK 88 1262 412 5151 - 154
2 m2 - - - - -
21m - - - - -
USA 8948 - 1 3200 - -
= - 47 m2 - - -
Unknown 66270 39796 16456 59966 6307 288366
46 m2 106 m2 411 m2 4298 m2 873 m2 231 m
5263 m 3500 m - 85 m -

= = - 3195 kg 2972 kg -

eee ee

65

Caiman crocodilus

Table 2b.

skins and sides of C. crocodilus fuscus reported to CITES (2 sides

1980

1981 1982

1983 1984

Countries with wild or farmed populations of C. crocodilus fuscus

Colombia 161170
Costa Rica -
El Salvador -

Guatemala -
Honduras -
Mexico -
Nicaragua

Panama 176150

Countries without wild

Argentina 100
Bolivia 1488
France -

Fr. Guiana -
Germany, F.R. -
Haiti =
Indonesia —
Italy 3723
Paraguay
Spain =
S. Africa -
Thailand -
USA -
Unknown 1584

Table 2c.

45968 19788

130905 88985

or farmed populations of C. crocodilus fuscus

- 4

- 19
3501 -
500 -
1255 -
760 100
402 -
100 3086
326 m 93

38253 8958

12 -

2479 79389
- 423 kg

- 55484

= 1

59646 10062
- 23 m

435 -

195 m2 £

6 m2 -

962 1090

= 1

3 =

4665 66747

Reported countries of origin or export for commercial exports of

1 skin).

1985

8746

121835
10654 kg
347700
36524

264 m
23845

Reported countries of origin or export for commercial exports of

skins and sides of C. crocodilus yacare reported to CITES (2 sides 1 skin).
1980 1981 1982 1983 1984 1985
Countries with wild or farmed populations of C. crocodilus yacare
Argentina 13073 36874 15585 7827 10861 1200
Bolivia - - 602 3919 - -
Brazil 4 - - - - -
Paraguay 4833 131094 333933 416427 70926 27747
= 112 m2 379 m2 18 m2 2014 m2 3m
- 11085 kg 3779 kg 6052 kg 2865 kg
Countries without wild or farmed populations of C. crocodilus yacare
Colombia - - - 1234 11208 3756
Ecuador - 1 - - - -
France 1678 651 - - - -
Fr. Guiana - - - 1328 - -
Italy 660 8574 - - - -
Panama - - 46316 3626 342 32
Unknown 721 4338 1492 23086 3 233
= = 6 kg 620 m2 139 m =

Caiman crocodilus

CONSERVATION MEASURES The legal protection status of C. crocodilus is
summarised in Table 3. A discussion of the legislation is given below as it is

so complicated in many countries. Unless otherwise stated all information was
from Fuller et al. (1987).

It has been suggested (Brazaitis, cited in Groombridge, 1982) that Cc.
crocodilus fuscus subspecies should be upgraded from Appendix II to Appendix
I of CITES. However, this may be premature until studies have be made to
determine the status of Venezuelan populations, as well as those in the
various Central American countries.

C. crocodilus yacare is specifically listed on the USA Endangered Species
Act, which prevents the import of this taxon to the USA.

A major requirement is to limit smuggling by proper enforcement of existing
laws. It has been proposed by some that C. crocodilus yacare should be
upgraded from Appendix II to Appendix I of CITES, at least until adequate
studies have been completed, evaluating the status of populations and the

effect of hunting on their reproductive potential. This proposal is not
endorsed by all authorities. There appear to be good populations still in
parts of Bolivia, Brazil and Paraguay. Present uncertainty about’ the

taxonomic status and distribution of Yacare, in particular in relation to the
remainder of the ‘Caiman crocodilus compiex', should be resolved when the
current research project has been completed.

Argentina C. crocodilus yacare is designated as a protected species under
Ley 22.421, Art. 7, and is protected by a total ban on commercial hunting,
import and export in Argentina. Although the implementing legislation was
enacted in 1981, the ban did not become effective until 1983. The export of
raw hides has been banned since 1976 (Resolucion No. 134).

Bolivia Decreto Supremo No. 16605 (June 1979) prohibits the hunting and
trade in species of native wildlife listed as protected, except that trade in
captive-bred specimens is permitted. C. crocodilus is included in _ the
protected list, however there is official disagreement over the validity of
this decree, and it has largely been disregarded. A temporary ban on the
export of live animals was imposed from May 1984 until August 1985, when it
was extended to July 1986 and expanded to cover wildlife products as well.
Decreto Supremo No. 21312 of June 27 1986 extended the ban on the export of
all wildlife products for three years with the sole exception of an annual
quota of 50 000 skins of C. crocodilus, all of which must be tanned before
export.

Brazil Complete protection is afforded the species in Brazil (Decreto
Presidential No. 58.054, 23 Mar 1966; Decreto-Lei No. 289, Feb 1967).

Colombia From 1982 until 1984, only inventoried stocks of hides were
permitted (from 1983 onwards this included only two companies), but all
further exports were banned from November 1984. Under Resolucion No. 847
(August, 1973), hunting and trade of Caiman crocodilus is prohibited in the
Orinoco basin, except for subsistence purposes, and is banned along the
Atlantic coast from April to July. A minimum size limit of 1.5 m is in force.
Only tanned skins were allewed to be exported (Resolucion No. 16, October
1969), but even this has now been prohibited.

Costa Rica Commercial trade and exports of non-marine wildlife has been

prohibited since 1970. All hunting of endangered species (including
C. crocodilus) was prohibited in 1985 (Decreto No. 15895-MAG).

67

Caiman crocodilus
Ecuador No commercial exports of indigenous wildlife are permitted.

El Salvador There are no protective laws for the species in El Salvador
other than those resulting from membership of CITES.

French Guiana C. crocodilus is listed under Article 3 of Arrété of 15 May
1986, which prohibits their use, taxidermy, purchase or sale. Their transport
is permitted only if they have been legally acquired outside the territory.

Guatemala In response to queries from the CITES Secretariat about the large
numbers of caiman skins exported early in 1985, a total prohibition on
hunting, capture, local trade, export and re-export was imposed by Resolucion
No. 410-86, 23 June, 1986.

Guyana All exports were prohibited on 28 February 1987. The ban was lifted
on 1 October 1987 and an export quota system was introduced. An annual export
quota of 20 000 live C. c. crocodilus and 40 000 skins was proposed for 1987
and 1988.

Mexico Annual closed seasons are imposed for C. crocodilus, and there is
a minimum size limit of 1.5 m (Flores Villela, 1980). Exports of live animals
and parts and derivatives have been prohibited since 1982.

Nicaragua Commercial hunting and export of wildlife has been prohibited
since 1977 (Decreto No. 625), but non-commercial tourist exports of up to two
objects made from C. crocodilus are permitted. The species is not listed as
endangered, and hunting seasons may be restricted.

Panama The capture, hunting, sale or export of endangered’ species
(including C. crocodilus) has been prohibited since 1980 (Resolucion No.
002-80). There are few regulations covering the import and re-export of
wildlife and their products.

Paraguay All hunting, trade and exports of indigenous wildlife has been
prohibited since 1975 (Decreto No. 18.796). There was confusion over whether
the law still applied after the ratification of CITES in 1977 but it has now
been confirmed, and no export permits have been issued since 1982.

Peru) C. crocodilus only occurs in the Amazonian lowlands (Selva region),
and under Decreto Supremo No. 934-73-AG (October 1973), no trade in species
from this area is permitted, except for animals hunted for subsistence
purposes. Caiman are not on the list of huntable species.

Suriname There are no protective laws for the species in Suriname.

Trinidad and Tobago The species is protected in Trinidad (Ordinance No. 26,
Page 13, 1958).

Venezuela C. crocodilus is listed as a game species in Venezuela, but
exports were prohibited from 1974 until 1983. Experimental harvest quotas were
set in 1983, but these were suspended in 1985 Exports were prohibited for one
year by Resolucion No. 61, 23 October 1985. When this expired, a new quota
system was introduced, the quota for 1987 being set at 150 000 animals.
Hunting is only authorised in the states of Apure, Barinas, Cojedes and
Portuguesa Region (Venezuela CITES MA, 1987).

68

Caiman crocodilus

Table 3. Legal prohibition on the commercial hunting, internal trade and
commercial export of C. crocodilus. Dates are those on which the legislation
came into force. A - All live animals & parts; L - Live animals only;
S - Semi-finished or finished skins allowed; P - Allowed under permit under
special circumstances; C - Closed seasons or quotas may be imposed; B -
Animals and parts from captive-breeding facilities allowed; * - these
territories are Overseas Départements of France with which the EEC may trade
without the imposition of CITES controls; ? - no information. Note
C. crocodilus apaporiensis, which is on CITES Appendix I, occurs within
Colombia, where it is accorded full protection (Fuller et al., 1987 modified
by G. Hemeley in litt. 1987).

CITES Hunting Trade Export
Argentina 1981 A 1983 A 1983 A 1983
Bolivia 1979 A 1986 A 1986 B 1979/ S/P/C 1986
Brazil 1975 A 1967 A/B 1967 A/B 1967
Colombia 1981 A 1983 B/P 1978 B/S/P/C 1974
Costa Rica 1975 A 1983 A/B 1983 A/B 1984
Cuba - ? ? ?
Ecuador 1975 Cc P A 1981
El Salvador 1987 - = P 1985
Fr. Guiana 1978 * A 1986 A 1986 A 1986
Guatemala 1980 A 1986 A 1986 A 1986
Guyana 1977 - - A 1986/C 1987
Honduras 1985 - - -
Mexico - Cc 1951 - A/B 1982
Nicaragua 1977 A 1977 P 1979 A 1977
Panama 1978 A 1980 A 1980 A 1980
Paraguay 1977 A 1975 A 1975 A 1975
Peru 1975 Cc 1973 A 1973 A 1973
Puerto Rico 1975 ? ? P
Suriname 1981 - ~ P
Trinidad & Tobago 1984 A 1958 A 1958 A 1958
Venezuela 1978 C 1982 P 1982 S/P/C 1982

CAPTIVE BREEDING C. crocodilus is kept in a large number of zoological
collections and breeds regularly. There are aslo several farming operations
orientated towards the commercial production of caimans for skins; these are
summarised below. Many of these farms keep crocodilians primarily as a tourist
attraction, and others are orientated more towards research. There are
currently none which derive most of their income from skin sales, and
Magnusson (1984) has suggested that this may not be possible, owing to the low
value of C. crocodilus hides. The largest commercial operations for
C. crocodilus exist in Taiwan, and these depend mainly on the sale meat and
medicinal products.

Bolivia ASICUSA (Asociacion de Curtidones de Cueros de Saurios), based in
Cochabamba and comprising four companies, Tomy, Alligator, Dorado and Moxos,
all involved in the tanning and processing of caiman skins in Bolivia, was
reported in 1982 to be establishing a crocodilian farm in Bolivia. In 1983 it
was claimed that the stock included 2000 crocodilians comprising
C. crocodilus crocodilus, C. crocodilus yacare, Melanosuchus niger and
“another species of Caiman". However Quaino, Director of “Moxos” Alligator

69

Caiman crocodilus

Ranch, stated that his ranch was only in a very preliminary phase and
contained only C. crocodilus in 1983. It is extremely unlikely that any
breeding has taken place on any farm in Bolivia. A report in 1982 stated

categorically that no breeding had occurred at that stage (Luxmoore, et al.,
1985).

Brazil No commercial crocodilian farming has taken place in Brazil but IBDF
(Instituto Brasileiro de Desenvolvimento Florestal) has initiated some
experimental farming of C. crocodilus. One breeding centre operated by IBDF
outside Manaus, the Centro Experimental de Criacao em Cativeiro de Animais
Nativos de Interésse Cientifico e Econdémico (CECAN), formerly held small
numbers of C. crocodilus and Paleosuchus trigonatus but these have since
been disposed of. ;

An experimental farm for C. crocodilus yacare was set up in 1981 in the
Pantanal about 150 km from Pocone, Mato Grosso. The aim of the project,
managed by IBDF, is to evaluate the possibility of caiman ranching in the
Pantanal. Eggs are collected from the wild, incubated and hatchlings are to be
released later. Few results are available as yet (Luxmoore et al., 1985).
Several cattle farmers in the Pantanal have expressed an interest in caiman
farming, and two are reported to have started farms (Brazaitis, 1985).

Colombia No commercial crocodilian breeding operations exist in Colombia,
but there are two (formerly three) breeding centres for conservation or
scientific purposes. One of these, the Estacion de Biologia Tropical at
Villavicencio, kept 17 C. crocodilus in 1983. Another, at Cienaga Grande,
formerly kept C. crocodilus fuscus, but this is believed to have closed
(Luxmoore et al., 1985).

Costa Rica A commercial farm was set up in the country in 1984 by a
commercial company acting in conjunction with the Ministerio de Agricultura y
Ganadaria. They had a stock of 96 C. crocodilus, mostly yearlings in 1985
(Luxmoore et al., 1985; Anon., 1985).

Honduras It was reported that a commercial farm for C. crocodilus was
under consideration in 1985 (Anon., 1985).

Italy Only one crocodilian farm is reported to have operated in Italy, but
it is now thought to have closed down. In 1981 it held a stock of 406 caimans,
probably C. crocodilus, originating in Colombia (Luxmoore et al., 1985).

Suriname There was reported to be a caiman farm in the Saramanca district
in 1979 (Luxmoore et al., 1985), but the farm never materialised, being
merely used as a cover for a commercial hunting operation (M.S. Hoogmoed, in
litt., 26 August 1986).

Taiwan There are 35 crocodilian farms in Taiwan, the first of which was
established in 1976. The bulk of the stock is C. crocodilus, of which about
8000 were kept in 1984. The annual production from all farms in 1984 was
12.5 t of skins, 30 t of meat and 7.5 t of other products, worth a total of
around NT$30 million (US$1 = NT$39). Most of this is sold within Taiwan but
some is exported to Japan and Korea. The meat is sold for food rather than
medicinal use, except those parts of the skeleton, blood, and male genitalia
which have medicinal value. The skin is processed for leather manufacture.
About 2000 hatchlings are produced each year on the farms. Each mature female
lays about 25-45 eggs with a hatching rate of 45% (Luxmoore et al., 1985).

70

Caiman crocodilus

Thailand The Samutprakan Crocodile Farm, which stocks mainly Crocodylus
siamensis and Crocodylus porosus, also had about 225 C. crocodilus in
1984 (Luxmoore et al., 1985).

Venezuela The Ministerio del Ambiente y los Recursos Naturales Renovables
(MARNR) runs an experimental centre to investigate techniques of captive
rearing of crocodilians, mainly C. crocodilus. Two private ranches in the
Llanos region, Hato Masaguaral and Hato El Frio, have been keeping Crocodylus
intermedius and have begun experiments on collecting and incubating eggs of
C. crocodilus, to evaluate the effect of releasing juveniles on the
potential harvest of the wild population (Luxmoore et al., 1985).

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Pereya, G.R. (1979). Fundamentos y programa de manejo para uso publico del
Parque Nacional del Manu. Ministerio de Agricultura y Alimentacion, Peru,
183 pp.

72

Caiman crocodilus

Quero de Pena, M. (in press). Politica de administracion del recurso fauna
silvestre en relacion al aprovechamiento comercial experimental de la
especie baba (Caiman crocodilus) en los Llanos Venezolanos. Paper
presented at the 7th Working Meeting of the Crocodile Specialist Group of
the Species Survival Commission of IUCN, Caracas, 21-28 October 1984.

Rebélo, G.H. and Magnusson, W.E. (1983). An analysis of the effect of hunting
on Caiman crocodilus and Melanosuchus niger based on the sizes of
confiscated skins. Biological Conservation 26: 95-104.

Schwartz, A. and Henderson, R.W. (1985). A guide to the identification of the
amphibians and reptiles of the West Indies, exclusive of Hispaniola.
Milwaukee Public Museum, 165 pp.

Schwartz, A., Thomas, R. and Ober, L.D. (1978). First supplement to a
check-list of West Indian amphibians and reptiles. Carnegie Museum of
Natural History Special Publication 5: 1-35.

Seijas, A.E. (1986). Situacion actual de las poblaciones de babas y babillas
(Caiman crocodilus) en la region norte costera de Venezuela. In:
Crocodiles. Proceedings of the 7th Working Meeting of the Crocodile
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21-28 October 1984. IUCN Publication New Series, pp. 28-36.

Serrano, F. (1978). Informe de actividades de la Unidad de Parques Nacionales
y Vida Silvestre de El Salvador. In: Morales, R., MacFarland, C., Incer,
J. and Hobbs, A. (eds), Primera Reunion Regional Centroamericana Sobre
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Turrialba, Costa Rica, pp. 117-124.

Smith, H.M., and Smith, R.B. (1979). Synopsis of the herpetofauna of Mexico.
Vol. V., Guide to Mexican Amphisbaenians and Crocodilians. John Johnson,
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Smith, N.J.H. (1980). Caimans, capybaras, otters, manatees and man in
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Staton, M.A. and Dixon, J.R. (1977). Breeding Biology of the Spectacled
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Torres Santibanez, H. (1978). Conservacion y manejo de la fauna silvestre del
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6/PAR/02/T.

Vallester, E. (1978). Informe de Panama sobre la situacion de la fauna
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Zoologia 32(17): 205-216.

Varona, L.S. (1980). Protection in Cuba. Oryx 15(3): 282-284.

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especies del genero Caiman (Crocodylia, Alligatoridae) en Argentina.
Amphibia y Reptilia (conservacion) 1(3): 68-75.

Wermuth, H. and Fuchs, kK. (1978). Bestimmen von krocodilien und ihrer
Haute. Verlag, Stuttgart, 100 pp.

Wermuth, H. and Mertens, R. (1977). Testudines, Crocodylia, Rhychocephalia.
Das Tierreich 100: 1-174.

73

NBW GUINEA CROCODILE Recommended list: 3
[No problem*]
Crocodylus novaeguineae Schmidt, 1928

Order CROCODYLIA Family CROCODYLIDAE

*Note last paragraph Summary and Conclusions

SUMMARY AND CONCLUSIONS A small to moderate size crocodilian, occurring
mainly in freshwater habitats, almost restricted to the island of New Guinea
(Papua New Guinea and Irian Jaya). Intense exploitation for hides, at a peak
in the 1960s, has led to widespread depletion of populations, but adequate
populations are known or thought to persist in both Irian Jaya and Papua New
Guinea, mainly in remote and poorly accessible lowland grass _ swamps.
Difficult terrain and secretive habits' inhibit reliable estimation of
population size or location but may also hinder exploitation. Remains largely
submerged in water during daylight, rarely basks, emerges at night. Feeds
mainly on fish and waterfowl, also other small vertebrates. Sexual maturity
in females generally attained at 1.8-2 m, age 6-8 years (possibly up to 10
years in the wild), and 2.5 m, age uncertain, in males. A 1.7 m wild female,
belly width 34 cm, has been recorded with 30 eggs. A mound-nesting species,
females guard nest. Mean clutch size in northern PNG 35, in southern PNG 22;
eggs of southern females tend to be larger, to 77 cm and 97 g. Females and
males may excavate nest, assist hatchlings to water, and females associate
with own hatchlings.

Hide hunting and collection of young to stock rearing farms are present
threats to survival. Few protected areas exist. In Papua New Guinea a
project developed with FAO/UNDP assistance exports ranched hides. The initial
emphasis on village ‘farms' has now given way to ranching in a small number of
larger commercial establishments, using stock collected by villagers from the
wild. The effect on wild populations is unknown in full, but early results of
the PNG monitoring scheme indicate a small but steady rise in nesting numbers
in the survey area (Sepik River). In this area at least, a continued harvest
of young crocodiles has been compatible with survival of a healthy crocodile
population, given that breeding adults and nests are adequately protected.
Present legal size range of skins is 7-20 inch belly width; the upper limit
should be decreased to provide more complete protection of the smaller
breeders. Rearing projects, similar to the PNG model, are to be promoted in
Irian Jaya, where numbers of novaeguineae appear to be adequate. Large
numbers of skins of this species have appeared in trade reported to CITES in
recent years: extremes of 5812 and 25 304 between 1980 and 1985, with a mean
of 16 746, from PNG and 1455-17 319, mean 7079, from Indonesia. The PNG
figures, derived from reported imports, do not correspond well with export
numbers reported by PNG itself, which are rather higher. Similarly, with the
exception of 1985, the Indonesian figures are not thought to reflect the true
volume of exports.

There seems no doubt that this species is able to sustain a substantial trade,
based on ranching of wild caught young and controlled harvest of sub-adults,
provided that protection of breeding adults and of nests is rigorously
enforced. There are indications that the control of trade in Indonesia is not
adequate and there have been substantial illegal exports. However a new FAO
programme has been initiated which aims to manage the harvest and regulate the
trade. It is too early to say if this is working.

74

Crocodylus novaeguineae

DISTRIBUTION Virtually restricted to the island of New Guinea (Papua New
Guinea and Irian Jaya) (Neill, 1971; Hollands, 1987; Whitaker et al., 1985).

The freshwater crocodile in New Guinea was originally described as a full
species (Schmidt, 1928), but several authors have treated this form and the
freshwater crocodile in the Philippines as two subspecies of one species,
C. n. novaeguineae and C. n. mindorensis respectively. A recent tendency
is to regard the Philippines crocodile as a full species C. mindorensis, in
which case the New Guinea form also reverts to a monotypic. species

C. novaeguineae (Wermuth and Mertens, 1977). A significant number of
differences exist - in cranial osteology, scalation, and aspects of
reproductive biology - between populations north and south of the central
highlands in Papua New Guinea (Hollands, 1987, and references therein). A

case could be made for regarding these population groups as separate species
(Cox, 1984; Hollands, 1987 and sources cited therein).

Indonesia Seemingly widespread in Irian Jaya, extending from the border with
Papua New Guinea westward to the Kepala Burung (Vogelkop) region (Whitaker et
al., 1985). This species retains significant populations in large tracts of
swamp country, especially when deep inland or otherwise remote; some 40 000 sq
km of such habitat occurs in Irian Jaya, concentrated in the southern lowlands
(from Merauke to Timuka), the Mamberamo and its tributaries in the Meervlakte
region (north of the central highlands), and the southern section of the
Kepala Burung (Whitaker et al., 1985).

Papua New Guinea Widespread in areas of suitable freshwater habitat both
north and south of the central highlands, extending from the border with Irian
Jaya eastward to the vicinity of Robinson River near Abau (Central Province),
almost at the eastern tip of the island. Absent from the island provinces of
Papua New Guinea (Whitaker, 1980).

POPULATION Overall, populations of C. movaeguineae appear to be widely
depleted to some extent, severely depleted in places, but adequate densities
are retained in parts of both Irian Jaya and Papua New Guinea. The difficult
terrain and secretive habits inhibit accurate estimation of population status,
but regular aerial surveys in selected areas of the Sepik system from 1981 to
date - a component of the crocodile management project in PNG - have now
provided the foundation for essential long-term monitoring, using nest numbers
as an indicator of population trends (Hollands, 1984, 1985).

Indonesia Although populations of both crocodilians in MIrian Jaya
(C. novaeguineae and C. porosus) are reported significantly reduced from
primordial levels due to excessive hide-hunting, “stable populations” or
“considerable numbers" of novaeguineae remain in large tracts of remote or
poorly accessible swamp country (Whitaker et al., 1985). Given the great
extent of such potential habitat available, Irian Jaya has been estimated to
possess the carrying capacity for several hundred thousand crocodiles
(Whitaker et al., 1985). A recent survey recorded only 0.27 crocodiles per
kilometre of the 847 km of river covered; about 4% of the total of 226
crocodiles were C. porosus, the rest, C. novaeguineae. (Whitaker et
al., 1985). These same consultants give a conservative estimate, based on
their survey results and on habitat availability, that 7 000-10 000 small
novaeguineae could be collected annually in Irian Jaya to form the basis for
the establishment of a sustainable crocodile industry along the lines of the
PNG model. Another study, carried out by the Forestry District Office, Irian
Jaya, and the Faculty of Forestry, Bogor, (Anon., 1986) concluded that the
population of crocodiles (both C. porosus and C. novaeguineae) in Irian
Jaya was 2 596 808, based on sample surveys in a few localities. These
estimates were thrown into doubt by Cox (1987), who extrapolated the

75

Crocodylus novaeguineae

population density estimates from Papua New Guinea to the amount of available
habitat in Irian Jaya and concluded that a better estimate for the latter
country would be 350 000 C. novaeguineae and 63 000 Cc. porosus.

Papua New Guinea Aerial surveys of selected nesting areas along the Sepik,
yielding data from which a nesting index is calculated, suggest a 38% increase
in novaeguineae numbers between 1981 and 1983 (Hollands, 1984). The Sepik
region produces nearly 40% of the country's skin crop, and, since the
situation here is considered likely to be representative of the overall
national picture, it was concluded in 1984 that the current crocodile
Management and exploitation regime is not only not causing further depletion
of wild stocks, but is allowing significant population recovery (Hollands,
1984). The last survey incorporated in the above summary was carried out
during the high-water phase in March 1984 but novaeguineae nesting is at a
peak during low-water, in October/November; a subsequent survey at this time
in the same year suggested a substantial reduction from the 1983 level to a
little below that recorded for 1981 (Hollands, 1984, Annex 1). The latest
survey for which results are available was carried out in October 1985. This
indicates a substantial rise in nesting activity, to a level above that in
1983, and 43% above that recorded at the start of the monitoring programme in
1981 (Hollands, 1984, Supplement dated November 1985). It was concluded at
the end of 1985 that rainfall and water level patterns were not conducive to
nesting during 1984, and that the novaeguineae population in the Sepik area
is continuing to rise steadily (Hollands, 1984, Supplement dated November
1985).

The report cited above highlights the case of Kamiemu as demonstrating the
best aspects of the PNG cropping programme; here, some 3.5 sq. km of floating
mat vegetation enclosed within an overgrown oxbow loop supports markedly
increased nesting by novaeguineae, up to 27 nests in 1985 (Table 1 in
Supplement) . At this site both nests and breeding adult crocodiles are
protected but a good number of young are harvested by local inhabitants to be
sold to commercial crocodile farms. In contrast, at another site, nesting
numbers continue to fall, mainly due to hunters killing adults for meat and
taking skins (Hollands, 1984, Supplement dated 1985).

Population data of comparable quality are not available for other parts of
PNG. In general, good numbers are said to remain in large tracts of inland
Brassy swamp, especially in more remote areas (Downes, 1971; Whitaker, 1980).
The upper and middle Strickland River, with up to 2.23 crocodiles/km of river,
primarily adults and subadults, may remain at historic population levels
(Montague, 1981). Populations have been locally overhunted (Downes, 1971;
Whitaker, 1980), with severe decline noted, for example, in Fly and Sepik
River populations, and Lake Murray (Neill, 1971). Recent surveys in the Sepik
region (see above) now suggest significant population recovery (Hollands,
1984). There is some evidence for slight recovery of novaeguineae in Lake
Murray and the Fly River, due in the former to Government releases (10 adults
in 1980) and escapes from the Baboa crocodile station, and in the latter to
reduced hunting pressure (Montague, 1981, 1982a). Crocodile density is still
(1978-1980) only 0.18 crocodiles/km on Lake Murray, and 1.8, 0.35, on the
lower and middle Fly (Montague, 1981). In Papua New Guinea a density of 2 or
more crocodiles/km appears indicative of healthy crocodile populations
(Montague, 1981). It has been suggested that crocodile hunting should be
banned on rivers with a density index of under 0.5 (Montague, 1981). A survey
of navigable portions of the Purari River revealed a low density of
crocodiles, subject to substantial exploitation; hunters interviewed suggested
that the population had declined in the last decade (Pernetta and Burgin,
1980). However, significant numbers of novaeguineae may exist undetected in
large areas of grass swamp connected by creeks to the main river. It was
suggested by local hunters that crocodiles move into remote Swamps during the

76

Crocodylus novaeguineae

breeding season and only appear in more accessible river channels as water
levels in the swamps decline (Pernetta and Burgin, 1980). It has been
stressed that although hunting pressure may be less in remote grass swamps, it
is probably never negligible, due mainly to the versatility of native canoes
(Pernetta, 1982). In the Lake Murray area, low water surveys of 1302 km
resulted in a count of 1112 crocodiles, probably indicating about 1765
actually present. Yet in 1980 alone, 2002 young live crocodiles (including
1.1% C. porosus) and 1100 skins were taken out of the area; it is considered
that the permanent swamps adjacent to main navigable areas provided the
additional crocodiles (Montague, 1981).

HABITAT AND ECOLOGY A relatively small crocodilian, average adult size
1.8-2 m (Brazaitis, 1973), maximum male length possibly around 3.5 m
(Whitaker, 1980), maximum actually recorded 3.35 m (Montague, 1982a). Mainly
occurs in freshwater habitats, but sometimes in brackish areas (Brazaitis,
1973), for example near the mouth of the Fly River (source in Whitaker,
1980). Primarily a lowland species, maximum known elevation is August River
(West Sepik) c 600 m above sea level (source in Whitaker, 1980).

Most remaining good populations appear to occur in remote areas of extensive
grass-swamp (Downes, 1971; Whitaker, 1980; Whitaker et al., 1985). The New
Guinea crocodile was formerly abundant, for example, in Lake Murray, a vast
shallow lake basin, with highly indented shoreline and forested islands, the
surroundings with rain forest, Melaleuca swamp and sago (Neill, 1971). In
Waigani Swamp, the species was reported to avoid deep open water of the larger
lakes, and prefer deep slow-flowing channels and small lakes overgrown with
herbaceous aquatics (Neill, 1971).

Generally very secretive in habits. Juveniles, in particular, disperse into
flooded swamplands during high water, possibly in response to harassment by
larger individuals (Montague, 1981). Smaller crocodiles rarely enter the
midstream of rivers but prefer near-shore cover such as emergent vegetation,
overhanging branches and fallen trees (Montague, 1981). Captive individuals
(at Moitaka Crocodile Farm, PNG) would emerge to bask in direct sunlight only
occasionally, usually during periods of cool rainy weather. Most of the
daylight hours were spent submerged or with only the head at the water
surface. The crocodiles would emerge in late afternoon and remain out of the
water all night and into the early morning. Body temperature was typically
close to 30°C during daytime. The major behavioural pattern involved heat
avoidance rather than heat gain (Lang, 1980a). Similar behaviour is recorded
in wild populations (Neill, 1971). In Papua New Guinea about 12 times as many
C. novaeguineae could be found on land at night than in day time (Montague,
1981, 1982a).

An opportunistic feeder, the known diet includes waterfowl such as crakes,
rails and grebes (Neill, 1971; Whittaker, 1980) but fish are thought to be the
main food (Montague, 1982a). Stomach contents of a 135 cm novaeguineae
included a 60 cm watersnake Amphiesma mairii, a rallid bird, grasshoppers,
leaves, and 40 gm of pebbles (Whitaker, 1980). Presumably small mammals are
also taken. Hatchlings feed on invertebrates such as mosquitos, grasshoppers
and water bugs (source in Burgin, 1980b).

Sexual maturity may generally be attained in females at 1.8-2 m, age 6 to 8
years, and 2.0-2.5 m, age uncertain, in males (Lang, 1981). Females estimated
at 1.5 m total length have been observed guarding nests in Waigani Swamp
(Neill, 1946), and a female of 34 cm belly width, 1.7 m length, was found to
contain 30 eggs (Jelden, 1981). First breeding in the wild may typically
occur at around 10 years of age in both sexes (Montague, 1982a). It should be
noted that the PNG legislation prohibiting trade in skins over 20" (51 cm)
belly width exposes young breeding females to legal hunting mortality for 1-2

77

Crocodylus novaeguineae

years until this size is attained (Montague, 1982b). A near 1:1 sex ratio was

found in a sample of 2031 wild novaeguineae from the Fly River (Montague,
1982b).

At Moitaka (PNG) courtship begins with the first rains, egg deposition follows
after 4-8 weeks. The northern populations in Papua New Guinea nest during a
rather short period in the dry season between August and November, hatching
occurs as water levels increase; in dry zones of the south, nesting occurs at
the start of the wet season, between Auguat and May, but with a peak in the
middle of the wet season (Hollands, 1987; Cox, 1984). This timing may reflect
hatchling requirements for increased vegetative cover and food _ supply
(Whitaker, 1980). In a captive group (Moitaka Crocodile Farm, PNG) it appears
likely that territorial behaviour and operation of a well-defined social
hiearchy occur all year round, not only during the breeding season when most
observations have been made (Lang, 1980a). However, individuals of the

species appear more tolerant of conspecifics than individuals of C. porosus
(Lang, 1980a).

In the Sepik River region, nearly 90% of nesting takes place amid floating
vegetation mats involving various plant associations. Most of these nests are
placed either in the flooded parts of ‘scroll’ zones (areas of parallel curved
ridges and gulleys formed by sequential erosion and deposition at river bends)
or in floating vegetation fringing open lakes and lagoons, and in vegetated
oxbow lakes (Cox, 1984). Land nests are more frequent in southern Papua New
Guinea where floating mat vegetation is less widespread (Cox, 1984). Both the
general site and the nest itself are often reused from year to year. The nest
is constructed of a mound of vegetable debris derived from plants at the nest
site; mean length, width and height for floating nests in the Sepik region
were 153, 131 and 50 cm, respectively (Cox, 1984). Mean clutch size in
northern populations is 35; 22 in southern populations. Mean egg length and
weight is 70 cm and 77 g, and 77 cm, 97 g, respectively. Northern animals
thus lay more, but smaller eggs.

THREATS TO SURVIVAL The primary factor leading to depletion of
C. novaeguineae populations over the past few decades has been over-hunting
for skins. Collection of eggs for food is also widespread (Downes, 1971;
Jelden, 1981). Present evidence from PNG indicates that harvesting of young
can be compatible with survival and even recovery of novaeguineae
populations providing that killing of breeding adults and nest predation are
curtailed.

In Papua New Guinea the two indigenous crocodilians, C. porosus and
C. novaeguineae, are utilized in a management scheme developed with the
assistance of FAO/UNDP. It is argued that since there is a very high
mortality of hatchling crocodiles in the wild, a harvestable surplus exists
that could be taken and reared to commercial size in captivity, without
affecting the status of wild populations. Rearing stations were mainly
village-based and owned by local people and it was intended that rational
Management instead of indiscriminate hunting would be of long-term economic
benefit. By the end of 1979 over 200 village crocodile-rearing stations were
established (Kwapena and Bolton, 1980). However, only about 15% of the
‘farms’ were rearing crocodiles satisfactorily, due to lack of expertise in
animal husbandry, seasonal fluctuations in water, difficulty in obtaining
crocodile food, and other factors (Kwapena and Bolton, 1980). The emphasis
now is on collecting of young crocodiles for sale to the three large-scale
commercial farms. It has been suggested that this buying scheme has increased
hunting pressure on wild populations (Burgin, 1980a). It has also been argued
(Burgin, 1980a) that the basic assumption, that there is a very high
density-dependent hatchling mortality, may only apply to populations at the
carrying capacity of the environment. Density-dependant mortality would be

78

Crocodylus novaeguineae

expected to be lower in populations already depleted by exploitation. The
effect of the rearing programme on wild populations is uncertain although
results of the monitoring programme (see ‘Population’, above) are very
promising (Hollands, 1984, 1987). The PNG programme can not be considered a
confirmed success until it is demonstrated that exploitation is being limited
to a level consistent with long-term survival of the resource, but, despite
valid reservations, the PNG project appears to constitute the closest approach
yet achieved toward rational utlization of any crocodilian resource. It is
claimed that the scheme not only maintains rural economies and counters urban
drift, but also conserves wild crocodile populations since the hatchlings
collected are considered a harvestable surplus and recruitment into the
breeding segment, itself untouched, is able to continue. Full scale FAO/UNDP
involvement ended in 1981 (except for persons assisting with monitoring) .

A similar pattern of trade in the 1950s and 1960s is reported in Irian Jaya
(Lever, 1980b). Hunting for the skin trade is the main factor affecting
crocodile populations in Indonesia and collection of young for rearing appears
to be on a smaller scale at present. Hunting in Irian Jaya is often organised
by Chinese traders using transmigrants from Java or Sulawesi to liase with
villagers who perform the actual hunting (Lever, 1980a). In 1972-1973 on the
Eilanden River, Indonesian soldiers based at Agats were hide hunting and
Organising local people into hunting parties (Webb, 1981). Reportedly
thousands of juvenile crocodiles (proportion of novaeguineae unknown) were
being purchased ready for shipment to rearing farms in Singapore, a collection
station was controlled by merchants in Jayapura and another located on Biak;
these crocodiles and hides were traded through Jakarta or direct with
Singapore (Webb, 1981). The scale of illegal skin exports is discussed in the
next section. Dixon et al. (in press) showed that the average belly width
of C. novaeguineae skins imported to Japan from Indonesia varied between
26.9 cm and 39.6 cm, while those from PNG were between 20.3 cm and 28.2 cm,
being both smaller and less variable in size.

INTERNATIONAL TRADE The number of C. novaeguineae skins in trade in the
1960s increased as hunters turned to this species after shooting-out most
C. porosus in accessible waterways, but this was followed by a marked
decline after 1966 as C. novaeguineae populations in turn became depleted
(Downes, 1971). The volume of C. novaeguineae skins produced in the PNG
trade fell from 62 948 in 1974 to 17 690 in 1975 after the introduction of new
control measures, but then gradually increased again to 35 374 in 1979
(Hollands, 1987). More recent export figures are given in Table 3. Minimum
net imports of skins of C. novaeguineae recorded in CITES Annual Reports are
shown in Tables 1 and 2. The principal importers have been Japan, the USA,
Italy and France. The number of skins reported to have originated in PNG
(Table 2) shows fairly poor correlation with those elsewhere reported to have
been exported (Table 3), due partially to the fact that PNG did not submit
annual reports to CITES in 1981 or 1982. The number of skins said to have
Originated in Indonesia in Table 2 similarly gives a very poor indication of
the true volume of the trade, and this subject was examined in detail by
Luxmoore (1986). Lever (1980a) considered that about 10000 skins were
exported annually from Irian Jaya, around 90% of these being C. novaeguineae
(Lever, 1980b). Dixon et al. (in press) examined the records of skin
dealers in Japan and estimated the total numbers of skins of C. novaeguineae
imported into Japan directly from Indonesia and via Singapore to be 518, 669,
976, 513, 1680, 745, 832 and 2856 in the eight years from 1977 to 1984
respectively. Similar imports from PNG over the same peiod were estimated at
5108, 9785, 10 624, 10 728, 7920, 7956, 8363 and 9706. However they pointed
out that independant evidence from Japanese Customs import records indicated
that the figures supplied by the dealers only accounted for between 5 and 48%
of the total amount of crocodile skins imported from Indonesia over this
period. Correlation between the dealers’ and Customs data was much better for

79

Crocodylus novaeguineae

imports from Papua New Guinea (over 70%). If the imports from Indonesia are
increased proportionately to take account of this discrepancy, they would vary
from 10 861 in 1977 to 6012 in 1984. These figures are closer to Lever's
(1980a) estimates of the trade.

Table 1. Apparent minimum net imports of skins and live animals (L) of
C. novaeguineae reported to CITES. Some transactions were reported in terms
of length or weight but the quantities involved were insignificant and these
data have been excluded from the table. A few live imports, although noted in
the table, are ignored in the figure for total volume for each year.

1980 1981 1982 1983 1984 1985

Australia 2L 2 2 21 - -
2L
Austria 383 - 66 210 107 107
Bahamas - ~ - 2 - -
Belgium 37 - - 24 - -
Canada 1368 - il jt - - -
Denmark 63 - - - - -
France - - - - 8863 12660
Greece 74 - - 3 - -
German D.R. ~ - - 29 - -
1L
Germany, F.R. 1522 286 85 940 - -
14L

Hong Kong 279 262 34 2010 169 180
Iceland - - - 110 - -
Israel 11 - - - - -
Italy 5838 21865 5102 5426 - -
Japan 5098 493 1680 15915 16428 24904
Korea 10 - - - - -
Kuwait 11 - - - - -
Liberia 367 - - 12 - -
Mexico - 1 - - - 56
Morocco - - - - - 6
New Zealand 84 - - - - -
Saudi Arabia - - - 82 - 4
Singapore - - 874 - - -
South Africa 12 - - - - 8
Spain 88 - - - 3 3
Sweden - - - 10 - -
Switzerland 4693 4306 4088 2117 2454 2393
Taiwan - - - - 4 -
UAE - - - 12 - -
UK ~ - 3 - -
USA 13029 21282 2396 1125 1128 1160
Total 32967 48496 14330 28048 29156 41481

80

Crocodylus novaeguineae
Table 2. Reported countries of origin (or exporting country if no origin
reported) and quantities of transactions in skins of C. novaeguineae

reported to CITES. Transactions reported in terms of length or weight have
been excluded from the table.

1980 1981 1982 1983 1984 1985

a. Countries having wild populations of C. novaeguineae

Indonesia 3851 1455 3503 8957 7391 17319
PNG : 11060 15097 5812 25304 22525 20678

b. Countries without wild populations of C. novaeguinae.

Argentina - - - - - 1232
Australia 200 = = = = &
Austria ~ = = 12 i 2
Denmark ~ = 157 = pa =
France 842 154 - = = =
Guinea - - - 24 165 x
India - - - - - 396
Malaysia 100 - = = # 115
Philippines - 60 - - - -
Singapore 10781 20882 3051 170 686 2201
South Africa - = = = 54 2
UK 36 - = = = =
Unknown 8487 11204 1886 1010 12 -
Table 3. Exports of C. novaeguineae skins from Papua New Guinea (Hollands,
1987)

1980 1981 1982 1983 1984 1985

27709 15021 24733 15111 21268 n/a

CONSERVATION MEASURES

Indonesia Nominally protected to some extent by legislation in Irian Jaya.
The species is listed as protected in Irian Jaya (Law No. 327/Kpts/Um/5/1978),
but it has been proposed that limited hunting should be allowed by permit
under control of the Ministry of Agriculture (360/Kpts/Um/8/1975) (Lever,
1980b).

In Irian Jaya the species appears to occur in the Gunung Lorentz Nature
Reserve, although severely depleted, and in Pulau Dolok and Wasur Game
Reserves (Lever, 1980b), also in proposed Mamberamo-Foja, Jayawijaya, Bintoni
Bay and Danau Bian Nature Reserves (Petocz, 1982).

Despite legal protection, illegal trade in crocodile skins from Irian Jaya is
proceeding at an alarming rate; and illegal exports exceed by a large margin
those officially authorised (Lever, 1980b; Whitaker et al., 1985; Luxmoore,
1986; Cox, 1987; Anon., 1987; Dixon et al., in press). The size of the
skins in trade, 9-38 inch (c 23-96.5 cm) belly width (Lever, 1980b), are

81

Crocodylus novaeguineae

generally larger than those exported from PNG (Dixon et al., in press),
indicating that breeding adults are being killed.

Commercial utilization has been identified as the only route to effective
conservation (Petocz, 1982). The real difficulties of preventing
over-exploitation are recognized, and a management project, funded by FAO, has
been initiated which hopes to establish a controlled harvest and ranching
programme similar to that operating in neighbouring PNG (Cox, 1987). At the
6th meeting of the Conference to the Parties to CITES in 1987, an agreement
was reached with the Japanese CITES Management Authority that all imports of
crocodile skins from Indonesia should be verified with the Indonesian
Management Authority before they were allowed into Japan. Singapore, the
other main importer, did not give such an undertaking.

Papua New Guinea Crocodile management is controlled by the Crocodile Trade
Ordinance, 1966 and the Crocodile Trade (Protection) Act. Hunting may only be
carried out by land-owners or with their permission, but beyond that hunting
is unregulated. Control is achieved by licensing those who wish to trade in
skins. The selling of skins having a belly width greater than 20" (51 cm) or
of less than 7" (18 cm) is prohibited. Smaller animals may be caught and
traded live for sale to ranches, but they may not be killed for skin trade.
All exports must be _ licensed (Hollands, 1987). It appears that this
programme has been relatively successful in managing the crocodile harvest.
Populations of both C. novaeguineae and C. porosus have increased in thr
Sepik River area. The numbers of skins in international trade are in
relatively close aggreement with the numbers authorised for export, and the
Japanese skin importers report that the great majority of skins from PNG fall
within the legal size limits (Dixon et al., in press), all of which indicate
that there is little unregulated trade. It has been suggested that the
maximum legal skin size in Papua New Guinea trade should be reduced from 51 cm
to 41 cm (16 inches) to ensure protection of females in the first year of
breeding (Montague, 1982b).

CAPTIVE BREEDING Most of the farms currently rearing C. novaeguineae in
PNG and Indonesia depend for their stock on hatchlings captured in the wild
and there is little if any captive breeding. There are in the region of 11
farms operating in PNG, but the great majority of the stock is held in only
two large commercial farms. In 1983 the stock of C. novaeguineae was
estimated to be 11 250 (Luxmoore, et al., 1985). The percentage of skins
exported by the farms was small until 1983 but increased to around 50% of the
wild harvest in 1984 (Hollands, 1987). There are currently 21 ranches in
Indonesia which have a capacity of 20-30 000 head. It is estimated that annual
production from these could rise to 15-20 000 in the next three years, of
which 75% would be C. novaeguineae (Anon., 1987). Some farms in Singapore
are believed to be keeping C. novaeguineae (Luxmoore et al., 1985).

REFERENCES

Anon. (1986). Feasibility study of the crocodile skin industry in Irian Jaya,
prepared by the Forestry District Office, Irian Jaya and the Faculty of
Forestry, Bogor. Summary translation by A. Mitchell and A. Pugu.

Anon. (1987). Maintenance of the Indonesian population of Crocodylus
porosus in Appendix II, with an increase in export quota to 4000 hides
per annum. Proposal submitted by the Republic of Indonesia to the 6th
meeting of the Conference of the Parties to CITES.

Brazaitis, P. (1973). The identification of living crocodilians. Zoologica
(NY), 58(4): 59-101.

82

Crocodylus novaeguineae

Burgin, S. (1980a). Crocodiles and crocodile conservation in Papua New
Guinea. Background paper given 30 October at Conference on Traditional
Conservation in Papua New Guinea: Implication for Today. Port Moresby,
27-31 October.

Burgin, S. (1980b). The status of the biology and ecology of Papua New
Guinea's Crocodile, Crocodylus mnovaeguinea (Schmidt) Science in New
Guinea 7(3): 163-170.

Cox, J. (1984). Crocodile nesting ecology in Papua New Guinea. FO:
DP/PNG/74/029. Field Document No 5. FAO and PNG Dept. of Primary Industry.

Cox, J. (1987). A strategy for sustained development of the crocodile
resource in Irian Jaya. Working Paper Number One, GCP/INS/060/JPN.

Dixon, A.M., Milliken, T. and Tokunaga, H. (in press) Japanese imports of
crocodile and alligator skins, 1970-July 1986. Report prepared for the
International Alligator and Crocodile Trade Study.

Downes, M.C. (1971). Regional situation report - Papua and New Guinea. In:
Crocodiles, IUCN Publications New Series, Suppl. Paper 32: 41-43.
(Proceedings of the First Working Meeting of Crocodile Specialists, Bronx
Zoo NY, March 1971).

Hollands, M. (1984). A preliminary examination of crocodile population trends
in Papua New Guinea from 1981-1984. Paper presented at 7th Working
Meeting of IUCN Crocodile Specialists Group, October 1984, Caracas,
Venezuela. (With two Supplements covering October 1984 and March 1985 nest
surveys).

Hollands, M. (1987). The management of crocodiles in Papua New Guinea. In:
Webb, G.J., Manolis, S.C. and Whitehead, P.J. (eds) Wildlife Management:
crocodiles and alligators. Surrey Beaty, Chipping Norton, N.S.W.,
Australia, pp. 73-89.

Jelden, D.C. (1981). Preliminary studies on the breeding biology of
Crocodylus porosus and Crocodylus novaeguineae on the Middle Sepik
(Papua New Guinea). Amphibia-Reptilia 3/4: 353-358.

Kwapena, N., and Bolton, M. (1980). The National Crocodile Project in Papua
New Guinea. Unpublished report pp 1-7.

Lang, J.W. (1980a). Behaviors of the Saltwater Crocodile (Crocodylus
porosus) and the Freshwater Crocodile (Cc. novaguineae) and
recommendations on captive breeding. Unpublished preliminary report to
the Wildlife Division, Dept. of Lands and Environment, Papua New Guinea.
Pp 1-7.

Lang, J.W. (1980b). Reproductive behaviours of New Guinea and Saltwater
Crocodiles. Text of paper presented at SSAR symposium on reproductive
biology and conservation of crocodiles, 7-9 August 1980, Milwaukee.

Lang, J.W. (1981). Thermal preferences of hatchling New Guinea Crocodiles;
effects of feeding and ontogeny J. them. Biol. 6: 73-78.

Lang, J. (1982). In litt., 7 February (comments on draft RDB account).

Lever, J. (1980a). Crocodile status in Irian Jaya. 2-page report to IUCN/SSC
Crocodile Specialist Group, unpublished.

Lever, J. (1980b). Crocodile conservation and industry development in Irian
Jaya. Report on consultancy sponsored by FAO, prepared for the
Directorate of Nature Conservation, Directorate General of Forestry.
Bogor, Indonesia, pp. 1-53 plus Appendices 1-9.

Lever, J. (1982). In litt., 16 February (comments on draft RDB account).

Luxmoore, R. (1986). Exploitation of the saltwater Crocodile in Indonesia.
Traffic Bulletin 7(5): 78-80.

Luxmoore, R.A., Barzdo, J.G., Broad, S.R. and Jones, D.A. (1985). A directory
of crocodilian farming operations. IUCN, Gland, 204 pp.

Montague, J.J. (1981). Characteristics of a crocodile population in Papua New
Guinea. Unpublished M.Sc. thesis, submitted to Michigan State University,
Dept. of Fisheries and Wildlife.

Montague, J.J. (1982a). In Jlitt., 19 February (comments on draft RDB
account).

83

Crocodylus novaeguineae

Montague, J.J. (1982b). Morphometric, injury and growth analysis of
Crocodylus novaeguineae from the Fly River drainage. Unpublished Ph.D.
Thesis, submitted to Michigan State University, Dept. of Fisheries and
Wildlife.

Neill, W.T. (1946). Notes on C. novaeguineae. Copeia 1946 (1): 17-20.

Neill, W.T. (1971). The last of the ruling reptiles. Colombia Univ. Press,
New York. 485 pp.

Pernetta, J.C. (1982). In litt., March (comments on draft RDB account).

Pernetta, J.C. and Burgin, S. (1980). Census of crocodile populations and
their exploitation in the Purari area. Purari River (Wabo) Hydroelectric
Scheme. Environmental Studies Vol. 14. Office of Environment and
Conservation, and Dept. of Minerals and Energy, PNG.

Petocz, R. (1982). In litt., 14 March (comments on draft RDB account).

Schmidt, K.P. (1928). A new crocodile from New Guinea. Field Mus. Nat. Hist.
Zool. Ser., Publ. 247, 12(14): 177-181.

Webb, G.J.W. (in preparation 1981) A guide to the status, conservation and
management of world crocodilians. Report to Australian National Parks and
Wildlife Service.

Whitaker, R. (1980). Interim report on the status and biology of crocodiles
in Papua New Guinea. Field Document No. 1, FAO Project PNG/74/029,
Assistance to the Crocodile Skin Industry. Wildlife Division (Dept. of
Lands and Environment) and FAO, Port Moresby.

Whitaker, R., Sukran, P., and Hartono, C. (1985). The crocodile resource in
Irian Jaya. WWF/IUCN Report 12. P. 1528-Consultancy Report. WWF/IUCN
Conservation for Development Programme in Indonesia.

Wermuth, H. and Mertens, R. (1977). Testudines, Crocodylia, Rhynchocephalia.
Das Tierreich 100: 1-174. Berlin, Walter de Gruyter.

B4

ESTUARINE or SALTWATER CROCODILE Recommended list: 2

[Possible problem]
Crocodylus porosus Schneider 1801

Order CROCODYLIA Family CROCODYLIDAE

SS ee ee ee ee

SUMMARY AND CONCLUSIONS A widespread species, occurring from Sri Lanka,
eastern India and Bangladesh, through coastal southeast Asia to the
Philippines, Western Carolines, and south through Indonesia to Papua New
Guinea and northern Australia, east to the Solomon Islands and Vanuatu.
Individuals have been recorded at sea and on land far beyond the breeding
range. The largest extant crocodilian, reported to have attained around 9m
length. Typically asssociated with brackish waters, such as coastal mangrove
swamp-forest, but also extends to freshwater rivers and grass-swamp. Males
are sexually mature at around 3.2 m, 16 years, and females at 2.2 m, 10
years. Clutch size 25-90 eggs, usually around 50. Very severely depleted,
rare and declining through most of the range, adequate population levels are
Maintained in parts of north Australia and New Guinea. Listed as Endangered
in the IUCN Red Data Book. Listed in CITES Appendix I, except the populations
in Australia, PNG and Indonesia which are listed on Appendix II. Depleted,
and still threatened, by extensive hide-hunting and habitat destruction.
Nominally protected by legislation through much of the range. Active research
and management programmes are operating in Australia, India and Papua New
Guinea. Breeds on a few commercial farms. The PNG rearing scheme is designed
to improve rural economies by trade in skins from hatchlings reared in
captivity from wild-collected young, and conserve wild populations by
decreasing indiscriminate hunting. A monitoring programme has been developed
to determine the effect of the scheme on wild populations.

The Australian porosus population was very severely depleted by hunting
between 1945-1972. Expert opinion is divided over the status of this
population; some workers believe a substantial recovery has taken place since
protection of the species, with an estimated total population of 30 000-40 000
representing 30-50% of pre-war levels though there is uncertainty of what
these were, others believe that recovery is slight and assert that the species
has no future in the country outside protected areas. Published data indicate
a small rise in numbers in many waterways and an increase in the proportion of
large individuals. It seems likely, although this is disputed by some, that
the Australian population can sustain the limited harvest neccessary for
commercial ranching; ca 4350 porosus are currently held, with an projected
1986 export of 470 skins. Monitoring in the Sepik River area of Papua New
Guinea has indicated a progressive increase in nesting numbers, with 1985
levels being 160% of 1982 levels; it is suspected that this is an adequate
reflection of the national situation. Present management and trade in ranched
crocodiles, with a mean annual export of 3163 hides in 1980-1984, seems to be
compatible with survival and recovery of the resource, providing that adults
are strictly protected.

The species appears to be severely depleted throughout its range in Indonesia,
with the possible exception of parts of Irian Jaya, but even here a recent
consultancy established that the species requires effective protection and
rehabilitation before commercial ranching can be considered. There has been a
small legal export from Indonesia in recent years but there is said to be a
substantial illegal trade. Transfer of the Indonesian population from
Appendix I to II was premature since there is no evidence that it can sustain
the heavy illegal trade. A recent programme, set up with assistance from FAO,
has attempted to bring the trade under control, but it is too early to say
whether it has been effective.

85

Crocodylus porosus

N.B. Appendix I and Appendix II populations are treated separately in the
Distribution and Population sections of this account.

DISTRIBUTION Crocodylus porosus has an extensive distribution in south
and South East Asia, extending eastward into Melanesia. Basic distribution
data are given below; see ‘Population’ section for further details. Breeding
populations are highly localized within the overall range due to extreme
depletion of the species in most areas and to habitat preferences (large
rivers, estuaries, coastal and inland swamps).

Although a specimen was obtained from southern China in 1912, in the vicinity
of Hong Kong (Smith, 1931), there are no recent records; it is uncertain
whether there was a breeding population at that time. Formerly present in
Singapore, but there have been no resident porosus for over 30 years (King
et al., 1979).

Wandering individuals have been seen at sea and recorded from localities far
outside the known breeding range, such as the Fiji Islands and Cocos-—Keeling
Islands. One 3.8-m male was found at Ponape in the eastern Caroline Islands,
some 1360 km from the nearest population (Allen, 1974), and a specimen has
been sighted at sea 480 km north of New Zealand's North Cape (Robb, 1980).

Appendix I populations

Bangladesh Formerly occurred in most rivers south of 23°15'N, throughout
the Sunderbans and much of the Chittagong area in the east. By the early
1950s all populations in Chittagong, the Meghna and Balleshwar estuaries and
northern parts of the Sunderbans had disappeared (Khan, 1980) and the species
is now restricted to southern parts of the Sunderbans. Preferred rivers
(based on interviews with locals, not actual sightings) appear to be the
Balleshwar, Bhola, Sela, Katka, Ambaria Ghat, other tributaries of the Pusur,
Bhadra and part of the Sibsa between 22°N and 22°40'N (Khan, 1980).

Brunei Present in coastal wetlands; no details available.

Burma Mainly confined to the Irrawaddy Delta, but also present in Arakan,
between Ramree Island and the mainland (Salter, 1983).

Federated States of Micronesia Occasional strays have been reported from
Pohnpei and Truk Islands which are considered to originate from New Guinea
(USA CITES MA, 1987).

India Restricted to two sites on the mainland; the Sunderbans of West
Bengal (in the north-east) and the Bhitarkanika Wildlife Sanctuary in the
Brahmani-Baiterani delta of Cuttack District, Orissa (northern sector of the

east coast). Formerly more widespread and present in suitable mangrove
habitat at many parts of the east coast, extending to Kerala on the west
coast. Also present in the Andaman and Nicobar Islands. (Bustard and

Choudhury, 1980; Whitaker and Daniel, 1978)

Kampuchea Probably present formerly; no recent information.

Malaysia Recorded from Peninsular Malaysia but now virtually extinct
although formerly widespread and common (King et al., 1979). Present in

Sabah (Whitaker, 1984) and Sarawak (Proud, 1981 cited in Groombridge, 1982).

Palau Islands [Belau] (U.S. Trust Territory of the Pacific Islands)
Restricted to the Palau group in the western Caroline Islands (Allen, 1974).

86

Crocodylus porosus

Philippines Present in the Cagayan River (draining north-east Luzon),
Mindoro Oriental, Catanduanes, Palawan, and in seven provinces of Mindanao;
probably more widespread formerly (Ross, 1984).

Solomon Islands Present on most larger islands and many smaller islands in
the group (McCoy, 1980).

Sri Lanka The main breeding population is in the south-west wet zone,
centred on the rivers draining into the formerly more extensive swampland
stretching from Puttalam south to the tip of the island (Whitaker and
Whitaker, 1979).

Thailand Probably extinct, the last confirmed record was from Ko Tarutao in
Changwat Satun in 1971 (Bain and Humphrey, 1980).

Vanuatu The main population was centred on the Sulphur River-Alligator
River area, north of Port Patteson, on Vanua Lava in the Banks Islands group
(Luders, 1983); this population now appears to be virtually extinct. Single
individuals have been reported form Espiritu Santo and Malo.

Viet Nam Reported to persist in the lower Mekong and Rung Sat Swamp (Nowak,
1976).

Appendix II populations

Australia Present in coastal regions of the far north in western Australia,
Northern Territory and Queensland (Cogger, 1979).

Indonesia Populations are widespread although generally small and
scattered. Recorded from the following areas, in approximate west to east
order: Sumatra, Siberut, Java, Kalimantan, Lesser Sundas, Sulawesi, Moluccas,
Irian Jaya, Timor.

Papua New Guinea The species has been reported from all of Papua New
Guinea's nine lowland provinces, also the island provinces of Manus, New
Ireland, West and East New Britain, North Solomons, and Milne Bay ‘includes
portion of mainland) (Whitaker, 1980). Formerly widely distributed and common
in the mangrove areas at the mouths of the Sepik and Fly, and around the Gulf
of Papua, now largely absent from easily accessible areas and found more
frequently in inland swamp habitat (Whitaker, 1980). Main centres are the
Swamps along the Sepik and Ramu rivers in the north, and swamps of the Fly
River and other rivers draining into the Gulf of Papua in the south (Bolton,
1978). On the Sepik River, the species occurs inland to the Irian Jaya
border, although numbers decrease in relation to C. novaeguineae, and on the
Fly porosus has been recorded 500 km from the sea (Whitaker, 1980).

POPULATION Severely depleted and at risk almost throughout its range (King
et al., 1979). Adequate population levels are maintained in only a few
localities, notably parts of northern Australia and parts of New Guinea (Irian
Jaya and Papua New Guinea).

Appendix I populations
Bangladesh The current population, restricted to southern parts of the
Sunderbans (but formerly extending from 23° 15' N throughout the Sunderbans

and much of the Chittagong area), is estimated at fewer than 200 individuals
in an area of c 780 sq. km out of the Sunderban area of 6000 sq. km (Khan,

87

Crocodylus porosus

1980). This population is of considerable importance, it stands a good chance
of surviving since it is within a protected reserve for Tiger Panthera
tigris and the Sunderbans area remains relatively hazardous and difficult of
access for humans (Whitaker, 1981 cited in Groombridge, 1982).

Brunei No detailed information available, scattered specimens are sometimes
seen in the coastal mangrove and Nipa palm swamps. A 1978 report indicated a
trade in young crocodiles, bought mostly by operators of commercial rearing
stations (King et al., 1979).

Burma Only small isolated populations remain (Whitaker and Daniel, 1978).
The main concentration, comprising c. 4000 individuals of all size classes
spread over the entire Irrawaddy Delta, but mainly in the east, is depleted
and decreasing (Caughley, 1981 cited in Groombridge, 1982). The Delta
population was reportedly (King et al., 1979) subject to heavy exploitation
in the late 1970s following control of Karen insurgents in the area. The
Arakan coast population also appears to have greatly declined since the early
1960s, due to conversion of mangrove habitat to agriculture and to hunting
(Salter, 1983).

India Formerly extended from the Cochin area of Kerala on the west coast,
southward around the tip of the peninsula, and northward along the east coast
to the Sunderbans in West Bengal. Also present in the Union Territory of the
Andaman and Nicobar Islands. Previously abundant wherever suitable habitat,
preferably estuarine mangrove forest, occurred; now severely depleted and rare
or extinct in most of its former range in India (Bustard and Choudhury,
1980). Extinct in Kerala, Tamil Nadu and Andhra Pradesh (a 3.2-m specimen
captured January 1979 in the Krishna estuary in A.P. is thought to have been a
wandering individual) (Bustard and Choudhury, 1980). The species persists at
two localities on mainland India; in West Bengal, where a small number occur
in the Sunderbans (Bustard and Choudhury, 1980; Kar, 1981 cited in
Groombridge, 1982), and in Orissa, where the major mainland Indian population
occurs in Bhitarkanika Wildlife Sanctuary (Bustard and Choudhury, 1980;
Daniel, 1980). The Sanctuary was founded on Kanika Island in _ the
Brahmani-Baiterani delta area. The entire Bhitarkanika population comprises
only 35 adult C. porosus, and at the end of 1976 there were in addition 61
crocodiles of 1-1.4 m length, the hatchlings of the 1974 season. The young of
1973 had virtually all disappeared by late 1975, and increased survival of the
1974 cohort is attributed to the protection afforded by declaration of the
area as a Sanctuary in 1975 (Bustard and Choudhury, 1980). The Bhitarkanika
area is probably unique in that ten per cent of the adults in this protected
population exceed 6 m in length (Bustard and Choudhury, 1980), and the largest
male exceeds 7 m (23'-24') (Kar, 1981 cited in Groombridge, 1982). Such a
high proportion of extremely large animals is unknown elsewhere. The species
was said to be abundant in the first half of the present century in several
islands of the Andaman and Nicobar groups (Choudhury and Bustard, 1979;
Whitaker and Whitaker, 1978), but the populations here are now depleted
(Bustard and Choudhury, 1980; Choudhury and Bustard, 1979; Whitaker and
Whitaker, 1978). On North Andaman Island, probably the best remaining area
for C. porosus in the Andaman group, a June-July 1978 survey suggested
presence of 36 nesting females (Choudhury and Bustard, 1979). Other workers
(Whitaker and Whitaker, 1978) provide the following approximate figures for
breeding females, based on field surveys and interviews; North Andaman - 50,
Middle Andaman - 20, South Andaman - 10. The total porosus population may
be 170-330. Populations are reported to be healthy in the Nicobars (Whitaker
and Whitaker, 1978), although no precise information is yet available. Human
predation on eggs, killing of adults, and loss of habitat exert a continuing
pressure on the Andaman-Nicobar populations (Choudhury and Bustard, 1979;
Whitaker and Whitaker, 1978).

88

Crocodylus porosus

Malaysia West Malaysia, uncommon and declining, considered essentially
extinct (Bullock in King, 1973; King et al., 1979; Scriven, 1972; Wycherley,
1971), a few porosus may remain in Tasek Bera Lake and the Pahang River
(King et al., 1979). Sarawak Cox and Gombek (1985) surveyed several river
systems and concluded that populations of C. porosus were as low as 0.054
per km. Comparing this with similar habitats in PNG and northern Australia,
they concluded that the species could be considered endangered, being
seriously depleted to less than 1% of the estimated population levels before
the advent of hunting. Sabah, becoming rare, reported extinct in much of the
former range (source in King et al., 1979). During 32 days of fieldwork in
April-June 1983, 56 porosus were seen along the 1146 km of river surveyed.
This suggests a very low density of 0.46 per km, in contrast to 4-12 per km in
healthy populations. By extrapolation Sabah's total population may be around
2600 (Whitaker, 1984).

Palau Islands [Belau] Reported to be still relatively common in the Palau
group in the late 1970s (sources cited in Webb, 1978). A total of 300
crocodiles were killed in a 1975 control programme, present status unknown. A
recent report stated that some illegal hunting occurs; skins from _ such
activities are exported to Japan. Overall, the population was thought to have
not recovered since the control programme eradicated the larger animals and
the population size was considered to be declining owing to habitat loss (USA
CITES MA, 1987).

Philippines Considered threatened, although isolated and depleted

populations persist. Present in the Cagayan River (draining north-east
Luzon), Mindoro Oriental, Catanduanes, Palawan, and in seven provinces of
Mindanao; probably more widespread formerly (Ross, 1984). Local populations

were reported healthy in 1978 in Lake Danau, Camotes Island, along the north
shore of Moro Gulf in Minadanao, perhaps in the Sulu Archipelago and Leyte
(King et al., 1979). However, more recent information (1981) indicates a
marked decline; no crocodiles are now known to survive on Camotes Island or in
Lake Danau (Ross, 1982).

Solomon Islands Declined in numbers since World War II, largely due to
hide-hunting. Only an occasional individual is now encountered on the larger
islands. A relatively large population, perhaps up to 300 individuals, occurs
on one small island where nesting occurs regularly. The persistance of this
population is attributed to the fact that crocodiles are sacred totemic
animals to the local people (McCoy, 1981 cited in Groombridge, 1981).

Sri Lanka The species appears to have been abundant around much of the
coast until the early part of the 20th century, but is now severely depleted
(Whitaker and Whitaker, 1978), and rapidly dwindling outside the two main
National Parks (Whitaker, 1986). A recent estimate suggests around 250
individuals (excluding first year hatchlings), with 25 breeding females, along
the southwest coast, and 125 individuals, with 15 breeding females, in the
rest of the island (Whitaker and Whitaker, 1978). The main breeding
population is in the heavily populated south-west wet zone and is concentrated
in remnants of formerly more extensive swampland, extending from Puttalam on
the west coast, southward to the southern tip of the island (Whitaker and
Whitaker, 1978). The best areas are centred around the main rivers draining
into this coast, the Maha Oya, Kelani Ganga, Bentota Ganga, Gin Ganga and
Nilwali Ganga (Whitaker and Whitaker, 1978). Breeding unconfirmed elsewhere
(King et al., 1979). Still present at some east coast sites where formerly
more common e.g. Batticaloa lagoon (Whitaker and Whitaker, 1978). Also
reported in the Mahaweli Ganga up to Mahiyangana (about 125 km inland) in the
east, and within the Yala National Reserve (Hoffman, 1981 cited in
Groombridge, 1982). Population in Sri Lanka continues to decline, chiefly due
to habitat loss. The Mutharajavela swampland between Negombo and Colombo, a

89

Crocodylus porosus

former main breeding site, is being cleared as it lies within a ‘Free-Trade
Zone' (Farook, 1980; King et al., 1979; Whitaker and Whitaker, 1978). A
suitable reserve area will be necessary to ensure survival of C. porosus
(Farook, 1980; Whitaker and Whitaker, 1978).

Thailand Probably extinct (Bain and Humphrey, 1980), no more than ten
adults may remain (King et al., 1979), the last confirmed specimen came from
the area of Ko Tarutao in Changwat Satun in 1971 with no confirmed sighting
since (Bain and Humphrey, 1980).

Vanuatu Single individuals have been reported during 1980-1981 from south
Espiritu Santo and west Malo, but the main population appears to have been
located in the Sulphur River-Alligator River area on Vanua Lava in the Banks
Islands group (Dickinson, 1981). Local tradition is that this population is
descended from a single colonizing female. Breeding appears to have occurred
in the recent past since individuals of all sizes could be seen; the total
population may have numbered around 200 (Luders, 1983). Hunting does occur
sporadically; in 1973 a group of Solomon Islanders shot seven crocodiles for
hides and a 4.8 m individual (maximum recorded length for Vanuatu) was shot by
an Australian in 1980 (Dickinson, 1980, 1981 cited in Groombridge, 1982).
This population appears to have become virtually extinct by 1983; this is
attributed largely to the effects of the 1972 cyclone which, with associated
flooding, caused massive destruction in the area (Luders, 1983).

Viet Nam Status and distribution little-known. Reported to persist in the
lower Mekong River and Rung Sat Swamp (Nowak, 1976). Not recorded north of Ho
Chi-Minh (Saigon) (Smith, 1931).

Appendix II populations

Australia Populations declined greatly during the 25 years following World
War II due to extensive hunting for skins (Jenkins, 1979; Messel, 1977). The
species may be responding to protection, although slowly (Messel and Vorlicek,
1986), and may not be at risk while this is maintained (Webb et al., 1984).

The published body of data concerning the habitat, biology, numbers, and size
distribution of C. porosus in various water bodies in northern Australia is
vastly greater than for any other crocodilian, with the possible exception of
the American Alligator. However, the interpretation of these data, in terms
of population status and trends, remains controversial.

In an overview of the major ten-year research programme undertaken by the
University of Sydney, the scientists involved state that the C. porosus
population “appears to be recovering at a very slow rate...it may take many
decades to recover - if ever". Further, “we can realistically and
unfortunately conclude only this about the saltwater crocodile's future: it
has none" (Messel, et al., Monograph 18 [Part 4, Abstract] in 1979-1984
series; Messel and Vorlicek, 1986).

On the other hand, in the Australian proposal to transfer its porosus
population from Appendix I to Appendix II of CITES (documentation prepared by
the Conservation Commission of the Northern Territory), it is stated that
available data (including those from the University of Sydney) “provide
unequivocal evidence of a population which has been increasing since
protection: not one which is decreasing or threatened” (Webb et al., 1984).

This dichotomy of views, or at least of emphasis, persists in the estimate for
the total C. porosus numbers in Australia. Messel et al. (1984) and
Messel and Vorlicek (1986) state that their 1979 maximum figure of 15 500
crocodiles over 3' in length was likely to apply also to the 1983 population.

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Crocodylus porosus

This estimate comprises 10 000 in Northern Territory; 2,500 in the Kimberley
region of Western Australia; 3000 in northern Queensland. These figures are
extrapolations from estimates based on actual surveys of a major proportion of
the total length of tidal waterways in the Northern Territory but only a small
proportion of the tidal waterways in Western Australia and Queensland.
Freshwater swamps and other freshwater habitats remain inadequately surveyed.
According to the Sydney team (Messel, 1981 cited in Groombridge, 1982; Messel
et al., 1979-1984), populations in the Northern Territory are slowly
recovering in tidal waterways of northern Arnhem Land and the Alligator Rivers
Region; populations from the Adelaide River westward to the Western Australian
border are steady or still falling; populations in the Gulf of Carpentaria are
at extinction levels with the exception of the Roper and Towns Rivers. In
Western Australia, populations probably recovering very slowly in major
sections of the Kimberley tidal waterways. Burbidge and Messel (1979)
estimated that the total non-hatchling population in Western Australia was
about 2000. Taplin (1987) reviewed the size of crocodile populations in
Queensland, and concluded that some recovery of crocodile populations had
occurred since protection was implemented, but that it was very difficult to
quantify. The major populations of C. porosus occur in the Weipa area of
north-western Cape York Peninsula. Messel et al. (in prep.) resurveyed this
area in 1987 and concluded that there was good evidence of a slow population
recovery since 1979. However they cautioned that human population pressures
on crocodiles were very much greater on the east side of the peninsula.

Messel et al (1979-1984) and Messel and Vorlicek (1985, 1986) provide
abundant evidence of the highly dynamic nature of porosus populations in
northern Australia; although a population model involving density-dependent
mortality is consistent with this evidence, these authors play down the
possible role of this factor in the present situation and stress instead their
belief that major recovery in numbers and age-structure of porosus
populations is a very long term process.

Webb et al. (1984) estimate that at least 30 000 individuals, and probably
closer to 40 000, remain in Northern Territory alone. This figure is in part
an extrapolation based on the extent of suitable habitat, and appears to
include hatchlings, or at least crocodiles up to 3' in length. These authors
use data from the Blyth-Cadell Rivers system to exemplify the general Northern
Territory trend. In this system there has been a 35% population increase
since protection in 1972, and an increase in the proportion of large animals.
Continuing recruitment, decreasing numbers of 3-5' crocodiles, and an
increasing number of crocodiles of 6' and over, are consistent with the
effects of density-dependent mortality. It may well be that this system is
approaching its carrying capacity. Populations along much of the Northern
Territory coast region are said to show similar trends, though usually with a
greater proportional increase in total population size (Webb et al., 1984).

One likely cause of these differing interpretations is the uncertainty over
the magnitude of decline suffered by porosus populations during the 27 years
of concentrated hunting between 1945 and 1972 (when federal prohibition on
import-export was implemented). While the present population has_ been
estimated as a small fraction of the pre-hunting population, other authors
argue that populations in the Northern Territory are 30-50% of pre-war levels
(Webb et al., 1984). This divergence of views, apparently caused in part by
differing concepts of primordial population levels, is superimposed on that
caused by differing approaches to the interpretation of raw survey data in
terms of population levels and structures.

However difficult it may be to fully reconcile the opposing views that have
been expressed on the Australian porosus population, the evidence appears to
indicate that some degree of population recovery has taken place in the the

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Crocodylus porosus

later 1970s and early 1980s, with a small observed increase in the number of
non-hatchling crocodiles in some waterways, and, perhaps more importantly, a
more general increase in the proportion of large animals (over 6" in length).
Furthermore, the Sydney team, while generally taking a more pessimistic view
of the recovery prospects of C. porosus than other workers, now appears to
agree (Messel and Vorlicek, 1986: 110) that crocodile farming should be
encouraged, with eggs being removed from nests known to be flooded during
January-March. Webb et al. (1987) provide a discussion of crocodile
populations in Australia in relation to proposed exploitation and management.

Indonesia Regional information available is as follows, in approximate west
to east order. Sumatra, becoming rare everywhere but still present in most
large rivers of the east coast (MacKinnon; 1981). Present in the Kluet area
of Leuser National Park in northern Sumatra, in general survival prospects are
poor due to disturbance and diminishing habitat (Wind, 1981 cited in
Groombridge, 1982). Siberut, formerly common but now depleted, due to hunting
in the south at least. Presence of porosus is confirmed in part of the Bay
of Tiop in the south, presence reported but as yet unconfirmed in estuaries
and swamps in the uninhabited north-west (within Reserve area), including
rivers Simarausau, Bolot, Tobakle, Torokatkat, Taio, Togilitte, Sinlingkle,
Tamaerak, also lakes at Gobjib and Bolot (Mitchell, cited by MacKinnon,
1981). Java, almost extinct, a few left in Ujung Kulon (MacKinnon, 1981), no
confirmed reports in the last three years (up to 1981) from East Java and very
few unconfirmed reports (Blouch, cited by MacKinnon, 1981). Kalimantan,
becoming very rare (MacKinnon, 1981), a 1972 survey of 200 miles of the
Mahakan River revealed only two adult porosus (King et al., 1979). Lesser
Sunda islands, rare (MacKinnon, 1981). Sulawesi, a few small surviving
populations, e.g. Randangan estuary on North Peninsula (MacKinnon, 1981).
Formerly common at the mouths of the Sausu and Tambarana Rivers (Teluk Tomini)
but no longer, following the increase in human population due to two large
transmigration schemes; overall, rare in Sulawesi with poor survival prospects
(Watling, cited by MacKinnon, 1981). Maluku (Moluccas), reported common in
Aru (MacKinnon, 1981) also reported severely depleted here (King et al.,
1979), still occurs in Wae Apu estuary on Buru, reported from Wahai near
Pasahari on the north coast of Ceram, otherwise rare (MacKinnon, 1981).
Timor, status unknown, breeding reported a decade ago on south coast (source
cited in King et al., 1979). Irian Jaya, scarce throughout its range in the
province, although apparently more secure here than in other parts of
Indonesia. Pulau Dolok, half of which is a Reserve, is a major stronghold
(Lever, 1980). There is no evidence of any recovery after the 1980 ban on
hunting and trade, and illegal skin exports continue (Whitaker et al.,
1985). Of the current populations in Indonesia, that in Irian Jaya appears to

have the’ greatest potential for recovery. Whitaker et al. (1985)
recommended that conservation of this species should be given high priority in
any management programme; indicating that adequate restoration of the

population was necessary before it could be included in any ranching scheme.
A subsequent study (Anon., 1986) concluded that the combined population of
C. porosus and C. novaeguineae was around 2 596 800, but this result was
questioned by Cox (1987), who estimated that there were around 63 000
C. porosus in Irian Jaya.

Papua New Guinea The species remains widespread, but is depleted throughout
its accessible range due to heavy exploitation for hides in the 1950s and
early 1960s (Bolton, 1978; King et al., 1979). Although exploitation is
less in remote areas, it is never negligible, and there is at present almost
no locality that is not subject to some kind of hunting (Pernetta, 1982 cited

in Groombridge, 1982). While formerly common in the mangrove areas at the
mouths of the Sepik and Fly, and around the Gulf of Papua, it is now scarce
and occurs more frequently in inland swamp habitat (Whitaker, 1980). Main

centres are the swamps along the Sepik and Ramu rivers in the north, and

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Crocodylus porosus

swamps of the Fly River and other rivers draining into the Gulf of Papua in
the south (Bolton, 1978). On the Sepik River, the species occurs inland to
the Irian Jaya border, although numbers decrease in relation to
C. novaeguineae, and on the Fly porosus has been recorded 500 km from the
sea (Whitaker, 1980). In the islands, C. porosus is extremely depleted on
Manus, New Ireland, somewhat depleted in North Solomons, sparse in East New
Britain but more widespread and in greater numbers in West New Britain
(Whitaker, 1979).

An active rearing programme, aided until 1982 by FAO/UNDP, is in operation in
Papua New Guinea. Wild-caught young are reared for their hides. A major aim
is to expand rural economies while conserving wild crocodile populations.
Early results from the monitoring scheme in PNG, a component of the crocodile
Management ‘programme, indicate that both crocodile species in PNG are now
increasing in numbers. While there has been a small decrease in the number of
porosus nesting in lake fringe habitats, probably as a result of greater
vulnerability to hunting and disturbance, there has been a_ substantial
increase in numbers nesting in overgrown oxbows and channels, and other
river-margin habitats. The overall trend has been upward, with the March 1985
nesting index being 160% of the baseline 1982 figure (Hollands, 1984, 1987).
The Sepik area is suspected to be adequately representative of the national
Situation. One urgent requirement is to continue and extend the protection of
breeding adults by decreasing the upper size limit of skins permitted in trade
(20" belly width) and further curtailing the taking of adults from the wild.

HABITAT AND ECOLOGY The largest extant crocodilian, C. porosus is
dubiously reported to have attained around 9 m (30 feet) in length (Taylor,
1979) (based on a probably erroneous extrapolation from length of a preserved
skull (Lang, 1981 cited in Groombridge, 1982)). The greatest authenticated
length in recent years is 6.2 m (20'4") recorded on a male drowned in a
barramundi net in the Fly River near Obo, Papua New Guinea (Whitaker, 1980);
however, a very large skull preserved by the Raja of Kanika, Orissa, belonged
to an individual estimated to have been around 7 m (23') in length (Daniel,
1980; Daniel and Hussain, 1974), and it is reported that the largest male in
the Bhitarkanika area at present exceeds 7 m in length (Kar, 1981 cited in
Groombridge, 1982).

The species typically occurs in brackish waters, inhabiting large river
estuaries and deltas with associated coastal mangrove swamp-forest, but also
extends into deep rivers far above tidal influence, also freshwater pools and
swamps (Bustard and Choudhury, 1981; Smith, 1931; Webb, 1977). Through much
of its range C. porosus is now mainly restricted to the mangrove system, in
India for example, the remaining porosus populations occur only in this
habitat (Bustard and Choudhury, 1981; Daniel, 1980). The same applies to

populations in southeast Asia and Indonesia. In the Andamans C. porosus
occurs in coastal streams, with mangroves, notably Rhizophora mucronata and
R. apiculata, and Brugiera, giving way inland to cane _ brakes with

Calamus and creeping cane, and semi-evergreen or evergreen riverine forest,
with bamboo and scattered trees of Dipterocarpus, Planchoria and Pinsonia
(Choudhury and Bustard, 1979).

However, while in Papua New Guinea the species has been typically associated
with coastal mangrove, nipa and sago swamp, in some areas it is now more
common in inland grass-swamp, and in the island provinces is most often found
in inland lakes (even upland crater lakes and hill streams) (Whitaker, 1980).
In Australia many C. porosus occur in tidal reaches with mangrove
vegetation, and also occur in adjacent floodplain billabongs and spring-fed
swamps which tend to be dominated by Paperbark Melaleuca and Pandanus palm,
with a variety of sedges, grasses and vines. They may also extend 150 km
upstream into freshwater non-tidal areas, and are well established in coastal

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Crocodylus porosus

freshwater swamps with floating mats of vegetation (including Phragmites,
Typha and Cyclasaurus fern) (Webb, 1977; Webb, 1981 cited in Groombridge,
1982). As in Papua New Guinea, the present distribution of porosus and
nesting activity through these habitats partly reflects differential hunting
pressure before protective legislation (Magnusson et al., 1978), as well as
location of optimum nest sites. Hunting is easiest when crocodiles are
exposed on mudflats of coastal mangrove.

C. porosus appears to be a largely opportunistic feeder, and the nature and
size of the prey taken varies with age and habitat (Taylor, 1979; Webb,
1977). Feeding occurs during the day and night, when food is available, and
on both ebb and flood tides in tidal regions. They are mainly shallow water
or water-edge feeders. Crustaceans and insects comprise most of the diet of
hatchlings and juveniles, mainly grapsid crabs of the subfamily Sesarminae and
Palaemonid shrimps of the genus Macrobrachium (Taylor, 1979). Crustacea and
insects are also the main diet of subadults in the India-Burma region (Smith,
1931), the stomach of a 2.5 m individual from the Irrawaddy was packed with
crabs (Smith, 1931). Juveniles feed mainly along the water's edge and among
Mangroves at high tides. Larger juveniles, over 1.2 m length, take an
increasing proportion of vertebrate prey (Taylor, 1979; Webb, 1977). Food
items recorded include sharks, archer fish, barramundi, pop-eye mullet,
mullet, mudskippers, white-bellied mangrove snakes, cormorants and magpie
geese (Messel, 1981 cited in Groombridge, 1982; Messel et al., 1979-1984).
As size increases an individual becomes able to deal with progressively larger
prey, including sharks, turtles, cattle, horses and humans. Large crocodiles
may be cannibalistic and will take porosus hatchlings and small juveniles
(Messel, 1981 cited in Groombridge, 1982; Messel et al., 1979-1984).

Sexual maturity appears to be attained at around 3.2 m, 16 years, in males and
2.2 m, 10 years in females (Webb et al., 1978). The grouping behaviour
shown by hatchlings is lost at about 8 months and territorial behaviour begins
at about 2.5 years, several years before first breeding. The time of nesting
varies between localities through the extensive range of the species, but
often coincides to some extent with the annual wet season (Lang, 1980; Webb,
1977; Webb et al., 1977). Nesting may be spread over a 3-5 month period and

females at a given locality nest asynchronously (Lang, 1980). In Papua New
Guinea (Whitaker, 1979), nesting occurs in September-January (or into March),
and begins with or before the rains. This timing may indicate hatchling

requirements for shelter, food, and fresh water (in tidal areas), all of which
are enhanced by the first rains (Lang, 1980; Whitaker, 1979). Nesting is in
April-September in northeast India and the Andamans (here coincident with the
onset of the southwest monsoon (Choudhury and Bustard, 1979). In Australia
nesting occurs mainly in the wet season; the first nests are made in
October/November and the majority of nests are made in January/February.
However, sporadic nesting occurs at least until August, and may occur
throughout the year in suitable habitat (Jenkins, 1981 cited in Groombridge,
1982). Minor peaks in nesting may correspond with falling flood levels in
tidal areas, but in freshwater swamps such late nests are often next to (and
in one case on top of) an earlier nest strongly suggesting multiple nesting
from the one female (Webb, 1981 cited in Groombridge, 1982). On the west
coast of Sri Lanka nesting takes place in July-August (Deraniyagala, 1939),
the height of the monsoon season (Whitaker, 1981 cited in Groombridge, 1982).

A mound nesting species, the nest may be constructed from a variety of
vegetable debris. Along the tidal rivers of northern Australia nests are
sited in adjacent freshwater swamps in downstream areas, and at the riverside
or the edge of flood plain billabongs further upstream (Webb, 1977). In some
coastal swamps almost all nests are constructed on floating mats of vegetation
(Webb, 1981 cited in Groombridge, 1982).

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Crocodylus porosus

The clutch comprises 25-90 eggs, but samples of nests in north Australia (Webb
et al., 1977) and in the Andamans (Choudhury and Bustard, 1979) showed
remarkably similar mean clutch sizes of 50 and 51 respectively, range 40-62
(18 nests) and 42-67 (6 nests). Incubation period is 80-90 days (56,65)
although this can be greatly extended in low temperatures (Webb, 1981 cited in
Groombridge, 1982).

The female apparently remains near the nest for much of the incubation period,
and on bank nests is often in one of the one to four wallows typically found
immediately adjacent to the nest. Males appear to take no part in the nesting
processs (Webb et al., 1977). The female in captivity, and apparently in
some areas in the wild (Shelford, 1916; Bustard and Kar, 1980), becomes
aggressive during this phase and actively deters intruders, including other
female C. porosus (Lang, 1980). This behaviour means that nest-guarding
females are highly vulnerable to human hunters, and it has been suggested that
the trait is now less frequently shown in wild populations (Bustard and
Choudhury, 1980). Males in captivity also defend territories throughout the
year (Lang, 1980). The female does not attend the nest continually for the
entire incubation period, but frequently leaves it for short intervals
(Bustard and Choudhury, 1980). She assists the hatchlings by opening the
nest, probing with the snout and digging with fore and hind limbs, in response
to their calling within (Lang, 1980; Webb et al., 1977). Females are also
reliably recorded (Bustard and Choudhury, 1980) to gather hatchlings in the
throat pouch in times of danger, and probably to carry them in this way from
the nest to water. At least some adult C. porosus remain in the water with
grouped hatchlings for around two months (Webb et al., 1977), in Arnhem Land
rivers (Australia) for example, but creche formation appears not to be
universal in Australia and may vary with river type and parental age (Messel
et al., 1979-1984, Monograph 1).

A high proportion of nests are lost due to various factors. In north
Australia egg predation, mainly by Monitor Lizards Varanus spp. and humans,
is minimal but losses due to flooding are extremely high. It is estimated

that in the four year period up to 1977 over 90% of nests in areas studied
were destroyed by flooding (Webb, 1977; Webb et al., 1977). By contrast in
the Andamans of 30 nests found, 28 were destroyed by predators (human
predation - 22 nests, Monitor lizard - 2, wild pigs - 2) and only 1 was
flooded (Choudhury and Bustard, 1979). There is a possibility that some
females nest more than once in a year (Messel, 1981 cited in Groombridge,
1982). In a captive breeding colony in Papua New Guinea (Lang, 1980) only one
clutch was laid per season.

Adult crocodiles appear to tolerate hatchlings, and small crocodiles up to
3-4' in length in their vicinity, but not larger crocodiles in the same size
class as themselves or the next smaller size class. It is stated (Messel et
al., 1979-1984) that this single factor is of critical significance, and
alone largely determines the dynamics of C. porosus populations. Once a
crocodile reaches the 3-4' (.93-1.24m) and 4-5' (1.24-1.55m) size classes, it
is likely to be challenged increasingly by crocodiles near its own size and in
the larger size classes and be excluded from the area it was able to occupy
when it was smaller. A substantial fraction (80%) of the 3-6' (.95-1.86m)
sized crocodiles may thus be excluded from the river or be predated upon by
larger crocodiles. The overall numbers missing, presumed dead, remain high
and appear to be some 60-70%.

THREATS TO SURVIVAL The drastic depletion of C. porosus throughout its
range is attributed to commercial hide hunting (see below), a second factor is
widespread loss of habitat; general animosity to the species and local use for
food are factors of minor or local importance.

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Crocodylus porosus

Habitat loss appears to reduce the recovery prospects of populations already
depleted by hunting, and has been more severe in India than in most other
areas of the range (Bustard and Choudhury, 1981). The coastal mangrove
habitat, with which C. porosus is particularly associated, is a highly
vulnerable system: such areas have very high soil fertility when reclaimed,
and are often, as in India, accorded low management priority as their
intrinsic economic and ecological value is not appreciated (Bustard and
Choudhury, 1981). Similarly, coastal swamps, including mangrove, in Sri Lanka
have been widely cleared and drained for use as agricultural or building land;
this habitat loss is continuing (Farook, 1980; Whitaker and Whitaker, 1978).
Habitat loss is the primary current threat in the Philippines, caused by
expanding agriculture and aquaculture schemes (Ross, 1981).

In Australia the overall (Messel, 1981 cited in Groombridge, 1982; Messel et
al., 1979-1984) recovery of C. porosus could be halted or reversed by two
main factors. Firstly, C. porosus nesting habitat on Northern Territory
waterways is very vulnerable to destruction by trampling by feral water
buffaloes Bubalis bubalus. The destruction of riverine and swamp habitat by
these animals can be extreme and on rivers such as the South Alligator was
reportedly almost complete by the late 1970s. The anchorage of floating mats
of vegetation is broken down so that the whole mat drifts away in the wet
season (Webb, 1981 cited in Groombridge, 1982). On Adelaide and Daly Rivers,
Murgenella Creek, the East Alligator, West Alligator and Wildman Rivers the
habitat has also been extensively destroyed; however, the last three of these
(and the South Alligator River) are now within Kakadu National Park, which is
subject to an intensive campaign to control Water Buffaloes. Elsewhere, the
continued presence of these animals poses a severe threat to the long-term
recovery of C. porosus populations. The second important factor is the
toleration of net fishing for barramundi well upstream of tidal river mouths
and often right up into the breeding areas (Messel et al., 1979-1984;
Jenkins, 1980). This is done legally and often illegally; it has been
suggested that this is leading to marked depletion of stocks of barramundi and
of C. porosus (Messel et al., 1979-1984). In one river 23 large
C. porosus were found drowned in barramundi fishermen's nets within a two
week period and a considerable number of large specimens are probably drowned
annually (Messel et al., 1981). It is alleged (Messel et al., 1981) that
no effective action is being taken to counter this. It is often observed that
rivers with large C. porosus populations also produce good commercial
catches of barramundi, and it may be that the crocodile is beneficial- to
barramundi populations, perhaps by eating large numbers of predatory fishes
(such as catfish) that would otherwise consume small barramundi (Jenkins,
1980).

Another factor is that general animosity toward crocodiles is often directed
against C. porosus due to its large size, often aggressive behaviour if
encountered near a nest, and tendancy to eat humans on occasion. This factor
is likely to assume increasing importance where recovery of C. porosus
populations is occuring (Webb, 1981 cited in Groombridge, 1982; Jenkins,
1980). Proud (1981 cited in Groombridge, 1982) reported that the few large
individuals in the Rejang river, Sarawak, had been trapped due to man-eating
habits, the local authority at Sibu offering a bounty for each specimen killed.

The species is sometimes used for food, for example eggs are heavily-collected
in the Andamans (Choudhury and Bustard, 1979), or its parts used medicinally
(Bustard and Choudhury, 1981). Eggs and meat are consumed in Papua New Guinea
(Lever, 1982 cited in Groombridge, 1982).

C. porosus has been very extensively hunted for the leather trade; this has
undoubtedly been the major factor in its decline (King et al., 1979).

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Crocodylus porosus

The hide of porosus yields the finest quality leather due to the small scute
size and lack of bony osteoderms in the belly skin, resulting in a relatively
large area of attractive, flexible, and commercially useable hide from any
given skin (King and Brazaitis, 1971; Whitaker, 1979; Lever, 1982 cited in
Groombridge, 1982).

Hide-hunting, carried out both by local people and by expatriates armed with
high-velocity rifles and often with motor boats, was on a particularly large
scale in the 1950s and 1960s. In this period there was a rapid worldwide
increase in the price of reptile leather and it is estimated that hundreds of
thousands of C. porosus were killed annually (Neill, 1971). In Papua New
Guinea the estuarine and delta zones were mainly exhausted by the early 1950s
(Whitaker, 1979). In Irian Jaya, and elsewhere, C. porosus occurred mainly
in these readily-accessible areas and was thus the first crocodilian species
to be over-exploited (Lever, 1980). Selective hunting for large crocodiles
has resulted in a severe reduction in the breeding population and continued
hunting has resulted in minimal recruitment into the population, although, in
PNG at least, the larger crocodiles are+ not at present the most valuable
commercially (Lever, 1980; Lever, 1982 cited in Groombridge, 1982; Whitaker,
1981 cited in Groombridge, 1982). Legislation in PNG (see below) forbids the
sale of skins having a belly width of more than 20" (51 cm). Dixon et al.
(in press) examined the sizes of skins imported by various skin dealers to
Japan, and found that those from PNG varied in mean width from 17.8 cm in 1977
to 27.4 cm in 1984. In no year was the maximum skin size greater than 51 cm,
indicating that the regulations in PNG are largely being adhered to. However
the skins of C. porosus from Indonesia were much larger, varying in mean
width from 46.4 cm in 1977 to 48.8 cm in 1984. The maximum skin size was
119.4 cm for a skin imported in 1984. This demonstrates that the hide hunting
in Indonesia relies to a greater extent on the mature breeding adults, and is
therefore far more damaging to the population recovery.

Collection of eggs and young for ‘farm' reering to marketable size poses a
further threat to the species, when this is not accompanied by a suitable

Management programme. Several rearing farms operate in Indonesia (in Irian
Jaya, Kalimantan and Java), dependant on eggs and young taken from wild
populations (King et al., 1979). In Kalimantan, for example, as_ the

subadult and adult population became depleted to commercial exhaustion by hide
hunting, pressure switched to eggs and young to be sold to rearing farms; most
farms were closed by the mid-1970s as eggs and young could no longer be
readily obtained (King et al., 1979).

INTERNATIONAL TRADE There has been a steep decline in total world trade
volume of C. porosus skins during the 1970s, from 100 000 per year to 20 000
per year, with prices rising at the same time (King et al., 1979). However,
demand for porosus hide continues and several commercial operations attempt
to meet this, either through large-scale captive breeding (as at Samut Prakan)
or through captive rearing, whereby hatchlings are taken from the wild and
reared in captivity to a suitable size (Luxmoore et al., 1985).

Minimum net trade in skins of C. porosus reported to CITES is shown in Table
1. Of the three regional populations now listed on Appendix II of CITES, that
of Papua New Guinea has been so listed since 1979. Australia and Indonesia
have been on Appendix I until 1985 when both were transferred to Appendix II.
There has been no legal trade from Australia apart from a few skins exported
in 1980, 1984 and 1985: the first exports from Australian farms. The number
of skins reported to have originated in PNG (Table 2) shows fairly poor

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Crocodylus porosus

Table 1 Minimum net imports of skins of C. porosus reported to CITES.

1980 1981 1982 1983 1984 1985
Austria 17 181 155 287 240 -
Australia - ~ - 33 - -
Canada - - - 1 - -
France - - - - 1392 2331
- - - - 14 m 351 m
Germany, F.R. 248 11327 50155 490 29 3
Hong Kong — 35 738 - - 1 1
10 m - - - - -
Italy 890 2341 423 389 - -
- 66 kg - - - -
Japan 73 - 109 2050 2538 2414
Korea 9 - - 25 - -
Malaysia - 60 - - - -
Mexico - - - 4 kg - -
PNG - 1 - - - -
Singapore 7 - 100 371 300 231
Spain 20 - - - - -
Switzerland 2003 551 772 1717 1044 216
Turkey - - - 1 - -
UK 89 130 57, 34 - 362
USA - - - - 10 -
Total 3391 5134 2117 5398 5553 5758
10 m 66 kg - 4 kg 16m 351 m

correlation with those elsewhere reported to have been exported (Table 3), due
partially to the fact that PNG did not submit annual reports to CITES in 1981
or 1982. Indonesia has recorded an annual mean of 487 skins in 1980-1985, but
there is apparently a substantial illegal trade (Luxmoore, 1986; Whitaker et
al., 1985; Anon., 1987); it is very likely that the large numbers of skins in
reported trade with country of origin listed as Singapore or ‘Unknown' also in
fact originated in Indonesia. The number of skins said to have originated in
Indonesia in Table 2 gives a very poor indication of the true volume of the
trade, and this subject was examined in detail by Luxmoore (1986). Lever
(1980) considered that about 10 000 skins were exported annually from Irian
Jaya, around 10% of these being C. porosus. Dixon et al. (in press)
examined the records of skin dealers in Japan and estimated the total numbers
of skins of C. porosus imported into Japan directly from Indonesia and via
Singapore to be 78, 1121, 1820, 1188, 1070, 207, 310 and 1402 in the eight
years from 1977 to 1984 respectively. Similar imports from PNG over the same
peiod were estimated at 1278, 3094, 3603, 3217, 2481, 2062, 2263 and 3436.
However they pointed out that independant evidence from Japanese Customs
import records indicated that the figures supplied by the dealers only
accounted for between 5 and 48% of the total amount of crocodile skins
imported from Indonesia over this period. Correlation between the dealers’
and Customs data was much better for imports from Papua New Guinea (over
70%). If the imports from Indonesia are increased proportionately to take
account of this discrepancy, they would vary from 1635 in 1977 to 2951 in 1984.

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Crocodylus porosus
Table 2 Reported countries of origin (or exporting country if no Origin

reported) and quantities of transactions in skins of C. porosus reported to
CITES.

Countries having wild populations of C. porosus.

Australia 35 - - - 100 98
Indonesia - 1160 126 345 200 1094
Malaysia ~ 186 - 56 - -
PNG 1803 3147 651 6315 6538 4266
10 m 66 kg - 4 kg 15 m 351 m

Singapore 1233 2504 606 3 74 74
Thailand - - - - - 300
Countries without wild populations of C. porosus.
Canada - ~ - - 3 -
France 1938 1084 - - - -
Unknown 1205 2841 1470 537 46 -
Table 3: PNG exports according to Department of Primary Industry figures
(Hollands, 1987).

1980 1981 1982 1983 1984 1985

5797 4281 4853 3456 5145

There seems little doubt that the Australian population can withstand the
moderate exploitation necessary for the planned ranching programme in Northern
Territory (Webb et al., 1985). The large numbers exported from PNG perhaps
Bive some cause for concern, but harvesting does seem compatible with
continued population recovery, given the clear evidence for the latter in the
Sepik region (Hollands, 1984; 1987). On the contrary, the evidence for
widespread and continued depletion of C. porosus in Indonesia, combined with
reports of substantial illegal trade, suggest that harvesting here is not
likely to allow any population recovery until appropriate management has been
implemented (Whitaker et al., 1985). Since 1985, a new management programme
has been initiated with the assistance of FAO, but it is too early to say
whether it has been effective.

CONSERVATION MEASURES Listed on CITES Appendix I with the exception of
populations in Australia, Indonesia and Papua New Guinea, which are listed on
Appendix II. Reservations with regard to Appendix I listing are held by Japan
and Austria (reservations held by EEC countries, Italy and France, were
withdrawn on 1 January 1984, and by Thailand on 17 August 1987). The
protective legislation covering C. porosus through much of the range appears
to be poorly enforced in most countries and with a continued demand for
porosus hides, long-term survival of the species may depend on provision of
strictly protected reserves, coupled with evolution of adequate management
strategies where utilization is to continue.

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Crocodylus porosus

The species is not protected in Burma, Palau Islands [Belau], Philippines,
Thailand, Vanuatu. No information countries not listed below.

Australia Nominally protected by legislation in Australia (Western
Australia, 1969; Northern Territory, 1971; Queensland, 1974) (Webb et al.,
1985). The Australian population of C. porosus was transferred from

Appendix I to II in 1985 under the provisions of CITES Resolution Conf. 3.15
on ranching.

The species occurs in reserve areas in Australia, such as the Ord River Nature
Reserve in Western Australia (A.A. Burbidge, in litt., 5 February 1981) and
a sizeable population exists in the 20 000 sq km Kakadu National Park in
Northern Territory (Jenkins, 1979); this park now includes much of the East,
South and West Alligator River systems and the Wildman River system (including
the downstream reaches of all four) and is subject to an intensive campaign to
control Water Buffalo.

Bangladesh Listed on Schedule 3 of the Wildlife (Preservation) (Amendment)
Act, 1973, crocodiles may not be hunted, killed or captured. C. porosus
occurs within the Sunderbans Tiger reserve.

India All crocodilians are listed in Schedule 1 of the Wildlife Protection
Act, 1972, which confers the highest order of protection, but all states had
to ratify the Act individually, and Kashmir and Assam had still not ratified
the Act ten years later (Whitaker, 1987). The Government of India/FAO/UNDP
Crocodile Breeding and Management Project has resulted in important
conservation action. This scheme was initiated in 1975 with the dual aim of
rehabilitating, depleted wild populations of India's three crocodilian
species, and allowing the possibility of sustained utilization of wild or
farmed crocodiles (Jayal, 1980; Saharia, 1981). The State Government of
Orissa declared the entire remaining mangrove forest of the Brahmini-Baitarani
delta (Bhitarkanika), comprising 176 sq. km, a sanctuary in May 1975. Fishing
(a cover for crocodile poaching) was banned in the area in the same month, and
felling of mangroves ceased in 1976. Eggs are collected from natural nests,
incubated in captivity and the hatchlings reared to a size of 1 metre (Bustard
and Choudhury, 1980). Other centres were set up at Port Blair, Andaman
Islands, and the Sunderbans Tiger Reserve. A total of 408 C. porosus had
been released up to 1984 (Whitaker, 1987). The species was extinct in Andhra
Pradesh but the sanctuary was declared in July 1978 with the aim of
re-establishing C. porosus, using hatchlings captive reared at Hyderabad
from eggs from the Andaman Islands (Bustard and Choudhury, 1980).

Indonesia Nominally protected to some extent by legislation in Irian Jaya.
The species is listed as protected in Irian Jaya (Law No 716/Kpts/Um/10/1980),
in which hunting, except for capture for ranching purposes, is prohibited.
Since November 1986, only processed skins may be exported. The Indonesian
population of C. porosus was transferred from Appendix I to II in 1985,
subject to an annual export quota of 2000 skins under the provisions of CITES
Resolution Conf. 5.21. The quota was increased to 4000 in 1987.

Despite legal protection, illegal trade in crocodile skins from Irian Jaya is
proceeding at an alarming rate; and illegal exports exceed by a large margin
those officially authorised (Lever, 1980; Whitaker et al., 1985; Luxmoore,
1986; Cox 1987; Anon., 1987; Dixon et al., in press).

A management project, funded by FAO, has been initiated which hopes to
establish a controlled harvest and ranching programme similar to that
Operating in neighbouring PNG (Cox, 1987). At the 6th meeting of the
Conference to the Parties to CITES in 1987, an agreement was reached with the
Japanese CITES Management Authority that all imports of crocodile skins from

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Crocodylus porosus

Indonesia should be verified with the Indonesian Management Authority before
they were allowed into Japan. Singapore, the other main importer, did not
give such an undertaking.

C. porosus occurs in reserves on Siberut, Sumatra (Leuser Nat. Park) and
Irian Jaya. Pulau Dolok Game Reserve is probably the most important crocodile
area in Irian Jaya in that it provides good habitat for C. porosus, the
rarer of the two Irian Jaya crocodiles (Lever, 1980).

Malaysia Sabah Crocodiles are on Schedule 1 of the Fauna Conservation
Ordinance of 1963 (Amendment of 1 November 1982). They may only be killed in
self-defence or under special permit for scientific purposes (Whitaker,
1984). Small populations may occur in Klias N.P. and in Crocker Range N.P. in
Sabah (King et al., 1979). Not protected in Sarawak.

Papua New Guinea The two indigenous crocodilians, C. porosus’ and
C. novaeguineae, are utilized in a FAO/UNDP-aided scheme (FAO involvement
ended 1982) whereby hatchlings are taken from the wild, reared in ranches.
Hunting of wild animals is also allowed. It is claimed that the scheme not
only maintains rural economies and counters urban drift, but also conserves
wild crocodile populations since the hatchlings collected are considered a
harvestable surplus and recruitment into the breeding segment, itself
untouched, is able to continue. It appears that this programme has been
relatively successful in managing the crocodile harvest. Populations of both
C. novaeguineae and C. porosus have increased in thr Sepik River area.
Crocodile management is controlled by the Crocodile Trade Ordinance, 1966 and

the Crocodile Trade (Protection) Act. Hunting may only be carried out by
land-owners or with their permission, but beyond that hunting is unregulated.
Control is achieved by licensing those who wish to trade in skins. The

selling of skins having a belly width greater than 20" (51 cm) or of less than
7" (18 cm) is prohibited. Smaller animals may be caught and traded live for
sale to ranches, but they may not be killed for skin trade. All exports must
be licensed (Hollands, 1987). The Japanese skin importers report that the
great majority of skins from PNG fall within the legal size limits (Dixon et
al., in press), which indicates that there is little unregulated trade.

Sri Lanka Listed in Schedule IV of the Fauna and Flora Protection
Ordinance, Ammendment. No. 44, November 1964, crocodiles may only be killed
under special permit issued by the warden. Occurs in Yala and Wilpatu
National Parks (Whitaker, 1986).

Thailand Not protected. Much former habitat near Ko Turutao in Thailand is
within Turutao Marine National Park (Bain and Humphrey, 1980).

CAPTIVE BREEDING Breeds in ceptivity in suitable conditions, but few zoos
keep adult pairs (Honegger, 1979). The species is currently being bred at the
Samut Prakan commercial crocodile farm near Bangkok, Thailand, where about
1755 C. porosus were held in 1984 and a number of hybrids’ with
C. siamensis (Luxmoore et al., 1985). Animals are killed for their skin
at around three years of age, and the meat is also sold. In 1979 it appeared
(Lang, 1981 cited in Groombridge, 1982) that most Crocodylus breeding at
Samut Prakan involved C. siamensis, and there was little or no C. porosus
breeding. There is an experimental crocodile farm at Edwards River,
Queensland, Australia, which was started in 1973, but which did not export its
first commercial quantity of skins until 1985. There are three ranches in the
Northern Territory, which take primarily eggs collected from the wild, and had
a total stock of 2849 C. porosus in 1985. Most of the farms currently
rearing C. porosus in PNG and Indonesia depend for their stock on hatchlings
captured in the wild and there is little if any captive breeding. There are
in the region of 11 farms operating in PNG, but the great majority of the

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Crocodylus porosus

stock is held in only two large commercial farms. In 1983 the stock of
C. porosus was estimated to be 9 000 (Luxmoore, et al., 1985). The
percentage of skins exported by the farms was small until 1983 but increased
to around 50% of the wild harvest in 1984 (Hollands, 1987). There are
currently 21 ranches in Indonesia which have a capacity of 20-30 000 head.
The total stock of C. porosus was 5703 in 1987. It is estimated that annual
production from these farms could rise to 15-20 000 in the next three years,
of which 25% would be C. porosus. There is currently no captive-breeding,
though this is planned (Anon., 1987). A ranching operation was established in
Burma in 1978, taking eggs from the wild (Luxmoore et al., 1985). It was
reported to have a stock of around 1000 C. porosus in 1987, with an annual
turnover of 300-500 crocodiles (S. Agarwal, in litt. to B. Bunting, 29 July
1987). Some farms in Singapore are keeping C. porosus; breeding programmes
have been initiated (Luxmoore et al., 1985), and have met with some success
(Cox and Gombek, 1985). A farm at Kuching, Sarawak is reported to have been
breeding C. porosus regularly since 1980, 38 clutches being produced up to
October 1985 (Cox and Gombek, 1985). A large facility has recently been set
up in the Philippines with the intention of breeding C. porosus (J.L. Diaz,
in litt., 8 December, 1986).

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105

ABBOTTS DAY GECKO Recommended list: 2
[Possible problem)
Phelsuma abbotti Stejneger, 1893

Order SQUAMATA Family GEKKONIDAE

SUMMARY AND CONCLUSIONS Found in the Seychelles (Aldabra and Assumption),
and Madagascar, two subspecies have been characterised in the Seychelles and
others are considered to exist in Madagascar. The estimated population size
of P. a. abbotti is over 1 million specimens; there are no estimates for P.
a. sumptio. An arboreal lizard, P. abbotti may be found in plantations and
native forest. Threatened by loss of habitat on Assumption. Aldabra is a
Strict Nature Reserve and the species may also occur in a reserve in
Madagascar. Apparently breeds readily in captivity.

The minimum trade volume for the years 1980 - 1985 was 484 specimens, of which
the majority, 247, were reported as dead specimens for scientific purposes.
The remainder were live animals, presumably used for the pet trade. The
principal exporters were the Seychelles and Madagascar. The UK, Italy, and
the USA were the principal importers.

Exports in identified specimens of this species would probably not harm the
population although additional data are needed to verify this. However, the
Seychelles reported a large export of unidentified Phelsuma which, if they
were P. abbotti, could adversely affect the status of certain populations.
The restricted distribution of P.a. sumptio renders it particularly
vulnerable to exploitation. Expert opinion favours maintaining trade controls.

DISTRIBUTION Known from the Seychelles and Madagascar.

Taxonomic note Three former subspecies of P. abbotti are now generally
considered to be subspecies of P. longinsulae; i.e. P. 1. longinsulae,
P. 1. pulchra, P. 1. menaiensis (Cheke, 1982; 1984). All three are known
from the Seychelles and Cheke (1982; 1984) summarized their distribution.
P. v-nigra is sometimes treated as a subspecies of P. abbotti - here it is
treated as a full species following Cheke (1984) and Mertens (1962). It is
probable that Phelsuma befotakensis and P. chekei Borner and Minuth, 1984
from Madagascar should be referred to as subspecies of P. abbotti (Gardner,
1985). Borner and Minuth (1984) also considered that P. a. sumptio should
be upgraded to P. sumptio; as they are apparently the only authorities to
use this nomenclature, it is here maintained as a subspecies of P. abbotti.

Madagascar Mertens (1966) considered that the distribution of
P. a. abbotti included Nossi Bé, and the north-west coast of Madagascar. It
is likely however that specimens found here are uncharacterised subspecies
other than P. a. abbotti (Cheke, 1982; 1984)

Seychelles P. a. abbotti is known from Aldabra Atoll (Cheke, 1984; Blanc,
1972). Found on all the major islands and many of the smaller ones (Gardner,
1985). P. a. sumptio is known from Assumption Island (Cheke, 1982).

POPULATION
Madagascar No information is available.

Seychelles P. abbotti has been found at densities of 100-200
individuals/ha in coconut groves, and at lower densities in native forest
(Niedzwiedzi pers. comm. in Cheke, 1984).

106

Phelsuma abbotti

P. a. abbotti: The entire population numbers over a million specimens at a
conservative estimate. Highest densities are in mixed scrub vegetation and
coconut trees (A.S. Gardner, in litt., 23 January 1986).

P. a. sumptio: Abundant in coconut and Casuarina plantations, the
population of this subspecies is now restricted following clear felling over
much of the island. No detailed density estimates are available (A.S.

Gardner, in litt., 23 January 1986).

HABITAT AND ECOLOGY An arboreal lizard, P. abbotti abbotti is known from
various forest types. A solitary species, breeding takes place throughout the
year with eggs being deposited under bark in trees. The clutch size is
typically two. (Niedzwiedzi in Cheke, 1984; Cheke, 1984).

The lizard is diurnal and feeds on insects, nectar from flowers and liquid
from fruits (Niedzwiedzi in Cheke, 1984). A feeding association has been
observed with giant tortoises (Geochelone gigantea) and the geckos forage on
mosquitoes attracted to their soft parts and also insects disturbed by the
tortoises' movement through the vegetation (Honneger, 1966; Stoddart and
Wright, . 1967). Predators have _ been identified as drongo (Dicrurus
aldabranus), doves (Streptopelia picturata), herons (Ardeidae spp) and
coucal (Centropus toulou) (Benson and Penny, 1971; Frith, 1979).

THREATS TO SURVIVAL Clear felling is taking place on Assumption Island for
Buano extraction and this is restricting the population of P. a. sumptio
(A.S. Gardner, in litt., 23 January 1986). No information has been found
relating to exploitation of this species within the countries of origin.

INTERNATIONAL TRADE All Phelsuma_ species were originally included in
CITES Appendix II as they were or could be subject to heavy pet trade and the
species are difficult to tell apart. The only data on international trade are
those contained in the Annual Reports of the Parties to CITES which are
summarized in Tables 1 and 2.

Most of the reported trade was 247 dead specimens imported to the UK for
scientific purposes. The remaining trade was in live animals, probably to be
used as pets. Over the six years 1980-1985 inclusive, the Seychelles was the
largest exporter of this species, followed by Madagascar. Most of the live
animals were imported by Italy, followed by the USA and F.R. Germany.

Table 1: Minimum net imports of Phelsuma abbotti reported to CITES. All
specimens were live except for 247 scientific specimens, indicated by #.

1980 1981 1982 1983 1984 1985
Canada - - 2 - - -
Denmark 10 - - - - -
Germany, F.R. - 15 - 30 - -
Italy - - - - 100 -
UK - 247 # - - - 1*
USA - é1 18 - - -
Total 10 323 20 30 100 1

* Captive-bred

nn

107

Phelsuma abbotti

Table 2: Reported countries of origin (or exporting country if no original
source reported) and quantities of export of Phelsuma abbotti reported to
CITES. All exports were live except for 247 scientific specimens, indicated
by #.

1980 1981 1982 1983 1984 1985

Countries having or possibly having wild populations of P. abbotti

Madagascar - - - - 100 -

Seychelles - 247 tt 20 30 - -
- 25

Countries without wild populations of P. abbotti

Comores 10 51 - = = =
Germany, F.R. - 10 = = =

Indonesia - - : - = = 1*
USA - = 2 a Pe =

* Captive-bred

61 specimens were declared as having originated in the Comores, where
P.abbotti does not occur; so it is possible that this is case of mistaken
identity, in which case they could be P. comorensis, P. dubia, P. laticauda,
P. v-nigra, (all species treated under this contract) or P. robertmertensi.

In addition to the trade in this species, the Seychelles reported exports of

unidentified Phelsuma species. These are likely to be one of the following
species: Phelsuma astriata, P. abbotti, P. laticauda, P. longinsulae or
P. sundbergi. These exports were substantial and were as follows: 1980, 60

specimens; 1981, 1015 kg; 1982, 72 kg; 1983, 28 kg; 1984, 41 kg of bodies.
The largest single importer of these was the United Kingdom. F.R. Germany and
the USA also took large numbers. If these figures can be relied on and if
all, or a substantial proportion, of these imports, were P. abbotti, this
would be a cause for concern since they could make a great impact on wild
populations. Using the maximum weight of P. a. sumptio (11.7 g), the total
number of Phelsuma exported by the Seychelles in 1981 would be in excess of
85 000 animals, although only around 3% of this figure was exported in 1983,
(2400 animals). It should be noted that no such shipments, nor even a
substantial part of them, were reported as imports by other countries and it
is possible that errors were made in the Annual Report of the Seychelles.

Due to lack of population and life history data it is not possible to assess
the precise effects of international trade on this species. The reported
trade in P. abbotti is not large and, if the specimens were P. a. abbotti,
there would be little cause for concern, since the minimum population is
estimated as one million. If the subspecies exported from the Seychelles was
P. a. sumptio, which has a restricted distribution and is subject to
pressure from logging, it will probably be less able to withstand the
pressure. Gardner (in Jitt., 1986) reports that P. a. abbotti is fully
protected and not exploited and that P. a. sumptio is also not exploited.
However, he considers that their handsome appearance combined with their
rarity could make them attractive to collectors. Several experts have
expressed reservations about the advisability of reducing the trade controls
for Phelsuma (Q. Bloxam, in Jitt., 1986; A.S. Gardner, in Jlitt., 23
January 1986; R. Thorpe, in litt., 21 January 1986).

108

Phelsuma abbotti

CONSERVATION MEASURES It is possible that this species occurs within the

protected area of Lokobé on Nossi Bé (Madagascar). Aldabra is run by the
Seychelles Island Foundation as a strict Nature Reserve which affords
protection to P. a. abbotti. Gardner Cine eel Ger. 1986) reports that

P. a. abbotti and P. a. sumptio are not exploited.

CAPTIVE BREEDING. Phelsuma abbotti apparently breeds relatively easily in
captivity. Jacobi (1982, unpublished data) reported that, in the Netherlands,
5 specimens were bred in 1980 and 11 in 1981. A recent inventory by Slavens
(1985) did not report any specimens in captivity. There are however likely to
be many.

REFERENCES

Borner, A.R. and Minuth, W. (1984). On the taxonomy of the Indian Ocean
lizards of the Phelsuma madagascariensis species group (Reptilia,
Geckonidae) . Journal of the Bombay Natural History Society
81(2): 243-281.

Benson, C.W. and Penny, M. (1971). The landbirds of Aldabra. Philosphical
Transactions of the Royal Society of London 260: 417-527.

Blanc, C.P. (1972). Les reptiles de Madagascar et des iles voisines. In:
Battistini; R. and Richard-Vindard, G. (eds.). Biogeography and Ecology
in Madagascar. Dr W. Junk, The Hague, pp. 501-614.

Cheke, A.S. (1982). Phelsuma Gray 1825 in the Seychelles and neighbouring
islands: a re-appraisal of their taxonomy and description of two new
forms. Senckenbergiana biologica 62: 181-198.

Cheke, A.S. (1984). Lizards of the Seychelles. In: Stoddart, D.R. (ed).
Biogeography and ecology of the Seychelles Island. Dr W. Junk, The
Hague, pp. 332-360.

Frith, C.B. (1979). Feeding ecology of land birds on West Island, Aldabra
Atoll, Indian Ocean: a preliminary survey. Philosophical Transactions of
the Royal Society of London 286: 195-210.

Gardner, A.S. (1985). An identification key to the geckos of the Seychelles,
with brief notes on their distribution and habits. Herpetological
Journal 1: 17-19.

Honnegger, R.E. (1966). Beobachtungen an der Herpetofauna der Seychellen.
Salamandra 1966: 21-36.

Mertens, R. (1962). Die Arten und Unterarten der Geckonengattung Phelsuma.
Senckenbergiana biologica 43: 87-127.
Mertens, R. (1966). Die Nichtmadagassischen Arten und Unterarten der

Geckonengattung Phelsuma. Senckenbergiana biologica 47: 85-110.
Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.
Stoddart, D.E. and Wright, C.A. (1967). Geography and ecology of Aldabra
Atoll. Atoll Research Bulletin 118: 11-52.

109

Recommended list: 3*
[No problem]
Phelsuma astriata Tornier, 1901

Order SAURIA Family GEKKONIDAE

*See last paragraph of Summary and Conclusions

SUMMARY AND CONCLUSIONS This species is endemic to and widespread in the
Seychelles. Two widespread forms, Pr astriata astriata and P. a.
semicarinata, both appear to be abundant (numbering millions) and capable of
living in a wide variety of natural and manipulated habitats. Two further
subspecies, each on a single island, may be recogniseable. Present in a
Number of reserves in the Seychelles, this species is apparently relatively
easy to breed in captivity.

The minimum trade volume reported to CITES during the period 1980-85 was 891
specimens, the majority of which comprised a single shipment of 537 dead
specimens, imported to the UK for scientific purposes. Most of the remainder
were reported as live exports from the Seychelles and the Comores. As the
species does not occur in the Comores, there may have been a mistake over the
identity. A large quantity of unidentified Phelsuma spp. were reported as
exports from the Seychelles.

The volume of trade in lizards identified as P. astriata reported to CITES
seems certain not to have significant adverse effects on population levels.
However, the high number of unidentified Phelsuma reported as exports from
the Seychelles could, if all or the great majority were P. astriata, have
caused severe pressure on local populations; the inclusion in list 3 rests on
the assumption that this is not the case. Further information on the
identification and provenance of these lizards, and the veracity of the
reported transactions, is required. Expert opinion favours maintaining trade
controls.

DISTRIBUTION Endemic to the Seychelles. Cheke (1984) summarized the
distribution as follows:

P.a.astriata: Mahé, Long, Saint Anne, Cerf, Conception, Thérése, Cachee,
Anonyme, Islette and Silhouette Islands (Cheke, 1984). Meier (1982) however
additionally reported this subspecies from Praslin, Curieuse and La Digue.

P. a. semicarinata: St Joseph, D'Arros, Denis, Praslin, Curieuse, Round, La
Digue, Felicité, Petite Soeur, Grand Soeur, Marianne, Cousin, Cousine, Aride
(Cheke, 1984), Chauve Souris, Cocos (Thorpe, in Cheke, 1984).

Cheke (1984) recognises two further subspecies, P. a. astovei restricted to
Astove Island (Blanc, 1972; Cheke, 1982; 1984) and 'P. a. intermediate form'
on Frégate; both these forms appear to be treated as semicarinata by
others, such as Gardner (1985).

POPULATION

P. a. astriata: apparently abundant in most lowland habitat including
coconut plantations and gardens, has been recorded at densities of up to 643
per 100 trees on Silhouette (the actual density probably being at least double
this figure). The entire population must be numbered in millions (A.S.
Gardner, in litt., 23 January 1986).

P. a. semicarinata: Thorpe and Crawford (1979) and Evans and Evans (1980)

found that the highest densities on Praslin were in native palm forest, with
225-250 individuals/ha. Gardner (in Jitt., 23 January 1986), however,

110

Phelsuma astriata

reported densities of 400 per ha on Praslin. Cheke (1984) provided data for

Cousin where there was a maximum density of 175 individuals/ha. Diamond
(1976) reported that it was quite common there. The species may be excluded
from coconut plantations to some extent by the larger P. sundbergi. The

entire population is numbered in millions (A.S. Gardner, in Jlitt., 23
January 1986).

No information is available on the population size of P. a. astovei and the
putative Frégate form; these are not likely to be large since both have a
restricted distribution.

HABITAT AND ECOLOGY PP. a. semicarinata: On both Praslin and Cousin, this
arboreal subspecies is found in forests. Thorpe and Crawford (1979) found it
in coconut groves, gardens, natural lowland forests, eroded hillsides (low
densities) and hillside forest (highest density). It lives under bark in the
trees and at the leaf bases (Evans and Evans, 1980). A diurnal subspecies,
P. a. semicarinata feeds on insects (ants, bugs and beetles), spiders and
nectar and pollen (Cheke, 1984). On Praslin, breeding takes place all year
round and two eggs are laid under bark, in hollow rotting wood or in crevices
of buildings. Approximately 10-12 eggs are laid per year (Gardner, 1984). No
other information is available on the ecology of the other subspecies.

THREATS TO SURVIVAL None known. No information has been found relating to
exploitation of this species within the Seychelles.

INTERNATIONAL TRADE. The whole genus Phelsuma was listed on _ CITES
Appendix II as the species are difficult to separate. The only data on
international trade are those contained in the Annual Reports of the Parties
to CITES which are summarized in Tables 1 and 2. The majority of reported
trade comprised a single shipment of 537 dead specimens, imported to the UK
for scientific purposes. The remainder was in live animals, presumably for
the pet market. Over the six years 1980-1985 inclusive, the Seychelles was
the largest exporter of P. astriata, followed by the Comores, where the

species does not occur. It is likely therefore that this is a case of
Mistaken identity, in which case they could be P. comorensis, P. dubia,
P. laticauda, P. v-nigra or P. robertmertensi. The major importers of live

animals were F.R. Germany and the USA. Of the 13 reported shipments, five
involved the Netherlands as a re-exporting country and one as an exporting
country.

Table 1: Minimum net imports of Phelsuma astriata reported to CITES. All
figures represent live animals except for 537 dead specimens, indicated by #.

i —$—$

1980 1981 1982 1983 1984 1985
eee ater eee ee ee eee
Austria = = = 10 2 pos
Denmark 12 - = = = 2
Germany, F.R. 10 58 99 90 - -
Switzerland - = 2 oS = =
UK - 537 # - ~ - 7 *

- 29
USA = 46 = a = -
Total 22 661 101 100 (0) 7

* Captive-bred

ee ee

111

Phelsuma astriata
Table 2 Reported countries of origin (or exporting country if no origin

reported) and quantities of export of Phelsuma astriata reported to CITES.
All represent live animals except for 537 dead specimens, indicated by #.

1980 1981 1982 1983 1984 1985

Countries having or possibly having wild populations of P. astriata

Seychelles - 69 1 100 - -
- 537*

Countries without wild populations of P. astriata

Comores 12 55 100 - - -
Germany, F.R. - 7 - - -

Indonesia - - - - - 7 *
Madagascar 10 - - - - -
Netherlands - - 1 - - -

* Captive-bred

In addition to the trade in this species, the Seychelles reported exports of

unidentified Phelsuma species. These are likely to be one of the following
species: Phelsuma astriata, P. abbotti, P. laticauda (all species treated
under this contract), P. longinsulae or P. sundbergi. These exports were

substantial, reportedly mainly to the UK, F.R. Germany and USA and were as
follows: 1980, 60 specimens; 1981, 1 015 kg; 1982, 72 kg; 1983, 28 kg; 1984,
41 kg bodies. However no country reports importing such shipments (or even a
substantial part of them) and it is possible that errors exist in the Annual
Report of the Seychelles to CITES. If these shipments took place, in 1981
tens of thousands of specimens would have been transported. If these were
all, or a substantial part of them, P. astriata, populations could seriously
have been depleted.

The levels of identified specimens of P. astriata in trade do not appear to
be large enough to jeopardize the survival of this species. P. a. astovei
and the Frégate form are most likely to be vulnerable to collections, since
they have a restricted distribution. Gardner (in litt., 23 January 1986)
considered that this species could sustain a large export trade if managed
sensibly. However, Thorpe (in jJlitt., 21 January 1986) observed that
sustained and organised collection could threaten the species, particularly on
small islands. Both considered that commercial trade should not be encouraged.

CONSERVATION MEASURES The following reserves on the Seychelles contain P.
astriata: Cousin Island Special Reserve (managed by ICBP International),
Aride Island (managed by the Royal Society for Nature Conservation), La Digue
Veuve Reserve (on La Digue, managed by the Ministry of National Development),
Morne Seychelles National Park (managed by the Ministry of National
Development) and Vallee de Mai (on Praslin, a World Heritage Site).

CAPTIVE BREEDING Phelsuma spp. apparently breed relatively easily in
captivity. Jacobi (1982, unpublished data) reported that in the Netherlands,
17 specimens were bred in 1980 and 26 in 1981. In a recent inventory, Slavens
(1985) listed holdings of 3 specimens in two localities. There are however
likely to be considerably greater numbers in captivity.

112

Phelsuma astriata

REFERENCES

Blanc, C.P. (1972). Les reptiles de Madagascar et des iles voisines. In:
Battistini, R. and Richard-Vindard, G. (eds), Biogeography and Ecology in
Madagascar. Dr W. Junk, The Hague, pp. 501-614.

Cheke, A. (1982). Phelsuma Gray 1825 in the Seychelles and neighbouring
islands: a re-appraisal of their taxonomy and description of two new
forms. Senckenbergiana biologica 62: 181-198.

Cheke, A. (1984). Lizards of the Seychelles. In: Stoddart, D.R. (ed.),
Biogeography and ecology of the Seychelles Islands. Dr W. Junk, The
Hague, pp. 332-360.

Diamond, A.W. (1976). Cousin Island Nature Reserve Management Plan
1975-1979. International Council for Bird Preservation, London, 61 pp.
Evans, P.G.H. and Evans, J.B. (1980). The ecology of lizards on Praslin

Island, Seychelles. Journal of Zoology 191: 171-192.

Gardner, A.S. (1984). The evolutionary ecology and population systematics of
day geckos (Phelsuma) in the Seychelles. Ph.D. Thesis, Aberdeen
University..

Gardner, A.S. (1985). An identification key to the geckos of the Seychelles,
with brief notes on their distribution and habits. Herpetological
Journal 1: 17-19.

Meier, H. (1982). Zur Taxonomie und Okologie der Gattung Phelsuma auf den
Seychellen mit Nachtragen zu dieser Gattung auf den Komoren. Salamandra
18(1/2): 49-55.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

Thorpe, R.S. and Crawford, C.M. (1979). The comparative abundance and
resource partitioning of two green gecko species (Phelsuma) on Praslin,
Seychelles. British Journal of Herpetology 6: 19-24.

113

GREATER DAY GECKO Recommended list: 3
[No problem]
Phelsuma cepediana (Merrem, 1820)

Order SAURIA Family GEKKONIDAE

SUMMARY AND CONCLUSIONS Found in Mauritius and introduced on Réunion,
reports indicate that P. cepediana is abundant on Mauritius, numbering tens
of thousands. Destruction of native forest habitat and cultivation may affect
this species. Protected against capture, killing and export in Mauritius,
this gecko may occur in nature reserves. Apparently readily bred in captivity.

CITES Annual Reports indicate a minimum net trade of 1608 P. cepediana in
the period 1980-85; however only 180 specimens originated from Mauritius, the

only country with a wild population. Other reported exports, from the
Comores, Madagascar and the Seychelles, are likely to have been
misidentified. The Netherlands, F.R. Germany and the USA were the major

importers of P. cepediana.

The low level of reported trade before 1980, and the small number of specimens
originating in Mauritius, suggests that trade in recent years has not been a
significant threat to the species. Expert opinion does not favour relaxing
trade controls on Phelsuma.

DISTRIBUTION This species is native to Mauritius and its islets (Blanc,
1972; Cheke, 1982; 1984; A.S. Gardner, in litt., 23 January 1986; Rendahl,
1939) and introduced into Réunion (Cheke, 1975). Mertens (1963b) reported
that 50 had been released at Ivoloina on the east coast of Madagascar.

POPULATION Although reported as localized and declining by Temple (1974),
Owadally (Mauritius CITES MA, 1985) considered that it is the most abundant of
the Phelsuma in Mauritius with a population of hundreds of thousands.
P. cepediana was reported as the commonest and most widely distributed gecko
on the island (Mertens, 1963a); probably relatively secure (Q. Bloxam, in
Jitt., 22 January 1986).

HABITAT AND ECOLOGY Commonest in upland vegetation including banana clumps
and gardens, but also found at sea level (A.S. Gardner, in litt., 23 January
1986). The species usually hides in trunk crevices, beneath bark or under
sheltered branches. Eggs are usually laid in coupled pairs concealed in wood
crevices or at the base of the leaf sheaths of monocotyledons (Vinson and
Vinson, 1969)

THREATS TO SURVIVAL None confirmed; possibly affected by the large scale
modification of native vegetation.

INTERNATIONAL TRADE Phelsuma spp. were listed on CITES Appendix II
because it was likely that some species were or could be in international
trade and the different species are difficult to identify. Temple (1974)
reported that foreign dealers had expressed interest in the export of
Phelsuma from Mauritius. The only data available on this trade are those
contained in the Annual Reports of Parties to CITES which are summarized in
Tables 1 and 2. All reported trade was in live animals, probably mostly for
the pet market, although some were identified as being for zoological
purposes. The Netherlands and F.R. Germany were the main importers, followed
by the USA and Switzerland.

114

Phelsuma cepediana

Table 1 Minimum net imports of live Phelsuma cepediana reported to CITES.

a ae eee

1980 1981 1982 1983 1984 1985

Austria = = = 12 = es
Canada - - = = = 3x
Denmark 14 = = = = =
German D.R. 10 - = = = =
Germany, F.R. 90 237 181 26 - 12
Japan - = = 8 4 =
Netherlands ~ = = = = 524
South Africa 10 = = = = =
Switzerland 20 - 34 36 8 15
UK - 20 2 - = =
USA 100 55 - - 38 -

- - - - 54 * 98 *
Total 244 312 217 82 104 649

* Captive-bred

Table 2 Reported countries of origin (or exporting country if no origin
reported) and numbers of live in Phelsuma cepediana reported to CITES.

1980 1981 1982 1983 1984 1985

Countries having or possibly having wild populations of P. cepediana
Mauritius 110 2 2 - - 42

Countries without wild populations of P. cepediana

Comores 14 110 215 86 3 500
Germany, F.R. - 3 = = 4 x 23 *
Indonesia - - = = ee 2*
Madagascar 100 - - = = =
Netherlands 20 - - - 43 =

- - - - 50 * 76 *
New Zealand - - - - 4 =
Seychelles - 50 - - - -
Togo - - - 4 = =
Country unknown - 150 - = = =

* Captive-bred

The great majority of trade in P. cepediana was reported to have originated
in the Comores, where the species does not occur. It is likely that the

species was misidentified, possibly being P. comorensis, P. dubia,
P. laticauda, P. v-nigra or P. robertmertensi. Similarly, the 100
specimens reported as originating from Madagascar in 1980 are likely to be
species other than P. cepediana. Mauritius, where P. cepediana does occur

supplied a total of 156 specimens over the period 1980-1985 (mostly in 1980

115

Phelsuma cepediana

and 1985) to the USA, South Africa and the UK. 24 of the specimens in 1985
were identified on re-export from Switzerland as being for zoological
purposes. Owadally (Mauritius CITES MA, 1985) considered that no export of
P. cepediana from Mauritius now takes place. 21 of the 38 shipments during
this period either originated from or were re-exported from the Netherlands.

The number of animals originating from Mauritius is unlikely to harm the
population of P. cepediana since it is generally considered abundant.
Systematic collecting could, however, pose problems and Bloxam (in litt.,
1986) and Gardner (in litt., 1986) both consider that trade in Phelsuma
should not be encouraged.

CONSERVATION MEASURES A law was enacted in 1973 to protect endemic reptiles
from captute or killing and specifically forbids export from the island
(Temple, 1974). P. cepediana was protected under the Wildlife Act No. 33 of
1933 (Mauritius CITES MA, 1987). Macchabee/Bel Ombre and Ile Plate are
protected as nature reserves (Anon., 1985) and it is not inconceivable that
P. cepediana occurs within these. Data are required on the ecology and life
history of this species.

CAPTIVE BREEDING Phelsuma spp. apparently breed relatively easily in
captivity. Jacobi (1982, unpublished data) reported that, in the Netherlands,
1 specimen was captive bred in 1972, 9 in 1973, 1 in 1976, 4 in 1977, 6 in
1978, 14 in 1979, 6 in 1980, 24 in 1981. Trade in captive-bred specimens
reported to CITES in 1984 and 1985 indicates that breeding has occurred in the
Netherlands and F.R. Germany. A recent inventory (Slavens, 1985) listed 21
specimens in 6 collections; the total numbers are likely to be much higher.

REFERENCES

Blanc, C.P. (1972) Les reptiles de Madagascar et des iles voisines.
In: Battistini, R. and Richard Vindard, G. (eds). Biogeography and
Ecology in Madagascar. Dr W. Junk, The Hague, pp. 501-614.

Cheke, A.S. (1975). Un lézard malgache introduit a la Réunion. Info-Nature,
Ile Réunion 12: 94-96.
Cheke, A.S. (1982). A note on Phelsuma Gray 1825 of the Agalega Islands
Indian Ocean (Sauria: Gekkonidae). Senckenbergiana biologica 62: 1-3.
Cheke, A.S. (1984). Lizards of the Seychelles. In: Stoddart, D.R. (ed.).
Biogeography and Ecology of the Seychelles Islands. Dr W. Junk, The
Hague, pp. 332-360

Anon. (1985) 1985 United Nations List of National Parks and Protected Areas,
IUCN Gland and Cambridge, 171 pp.

Mertens, R. (1963a). The geckos of the genus Phelsuma on Mauritius and
introduced islands. Mauritius Institute Bulletin 5: 299-305.
Mertens, R. (1963b). Zwei neue Arten der Geckonengattung. Phelsuma.

Senckenbergiana biologica 44: 349-356.
Rendahl, H. (1939). Zur Herpetologie der Seychellen I. Reptilien. Zoologisches
Jahrbuch 72: 157-328.
Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.
Temple, S.A. (1974). Wildlife in Mauritius today. Oryx 12(5): 584-590.
Vinson, J. and Vinson, J.M. (1969). The saurian fauna of the Mascarene
Islands. Mauritius Institute Bulletin 6: 203-320.

116

Recommended list: 2
[Possible problem]
Phelsuma comorensis Mertens, 1966

Order SAURIA Family GEKKONIDAE

SUMMARY AND CONCLUSIONS Known only from Grande Comore, Comores Islands. No
information on population size or ecology is available, this is urgently
required. The restricted range is itself a cause for concern. Reputedly
easily bred.

CITES Annual Reports indicate a minimum net trade of 1236 live specimens of
P. comorensis and a further 222 unidentified Phelsuma in the_ period
1980-85, mostly from the Comores. F.R. Germany was the principal importer of
this species (followed by the USA and Switzerland). Specimens were probably
destined for the pet trade.

Although no assessment can be made of the precise effects of the exports on
the wild population, trade in such a restricted species must give some cause
for concern. Expert opinion does not favour relaxing trade controls on
Phelsuma.

DISTRIBUTION Known from the island of Grand Comore, Comores Islands (Blanc,
1972; Mertens, 1966). Meier (1982) noted that the distribution extends from
the type locality (La Grille at an altitude of 1000 m) to the north coast of
the island.

POPULATION Said to be less common on the north coast than elsewhere (Meier,
1982), otherwise nothing known.

HABITAT AND ECOLOGY No information available.

THREATS TO SURVIVAL At risk due to the restricted range, and perhaps the
effects of trade.

INTERNATIONAL TRADE All Phelsuma spp. were listed on Appendix II as it
was likely that they were, or might be, in trade and the individual species
are difficult to identify. The only data that are available are those
contained in the Annual Reports of the Parties to CITES and these are
summarized in Tables 1 and 2.

All reported trade was in live animals, presumably for the pet trade.
Virtually all the exports were from the Comoro Islands, accounting for 97% of
animals in trade over the period 1980-1985. The major importer was F.R.
Germany, taking 77% of the animals, the USA imported 12% and Switzerland 8%;
virtually all transactions were for commercial purposes.

The exports from the F.R. Germany probably originated from the Comores. It is
likely that the exports from the Seychelles were not P. comorensis but one
of the following: P. astriata, P. abbotti, P. laticauda, P. longinsulae or
P. sundbergi.

In addition to the trade in this species, there were several reported imports
to F.R. Germany and Austria from the Comores of unidentified Phelsuma
species. These are likely to be one of the following species: Phelsuma
comorensis, P. dubia, P. laticauda, P. v-nigra or P. robertmertensi. These
were as follows: 1980, 150 specimens; 1981, 20; 1983, 20; 1984, 30, 1985, 2.

117

Phelsuma comorensis

Table 1 Minimum net imports of live Phelsuma comorensis reported to CITES.

1980 1981 1982 1983 1984 1985
Argentina - - - - 2 -
Austria - - - 20 - -
Canada - 3 - - - -
Denmark 10 - 15 - - -
Germany, F.R. - 473 149 170 63 100
Japan - - 6 - - -
Switzerland - - 30 40 25 -
USA J 2 82 =e 60 10 -
Total 12 558 200 290 100 100

Table 2 Reported countries of origin (or exporting country if no origin
reported) and numbers of live Phelsuma comorensis reported to CITES.

1980 1981 1982 1983 1984 1985

Countries having or possibly having wild populations of P. comorensis
Comores 10 536 200 290 100 100
Countries without wild populations of P. comorensis

F.R. Germany 2 5 - = = =
Seychelles - 25 - = = =

Since no information is available on the population status of this species, it
is not possible to assess the effect of trade. However, the number of animals
exported from the Comores should be viewed with concern given the very
restricted range of this species.

CONSERVATION MEASURES Information on distribution, population status and
ecology is required.

CAPTIVE BREEDING Phelsuma spp. are reputedly easily bred. However no
information is available to suggest that this species has been bred in
captivity. A recent inventory (Slavens, 1985) listed 5 specimens in 3
collections; there are however likely to be more.

REFERENCES

Blanc, C.P. (1972). Les reptiles de Madagascar et des iles voisines.
In: Battistini, R. and Richard-Vindard, G. (eds). Biogeography and
Ecology in Madagascar. Dr W. Junk, The Hague, pp 501-514.

Meier, H. (1982). Zur Taxonomie und Okologie der Gattung Phelsuma auf den
Seychellen mit Nachtragen zu dieser Gattung auf den Komoren. Salamandra
18: 49-55.

Mertens, R. (1966). Die nichtmadagassischen Arten und Unterarten der

Geckonengattung Phelsuma. Senckenbergiana biologica 47: 85-110.
Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

118

Recommended list: 3
[No problem]
Phelsuma dubia (Boetteger, 1881)

Order SAURIA Family GEKKONIDAE

SUMMARY AND CONCLUSIONS A widely distributed species occurring in the
Comores, Mayotte, Madagascar, Mozambique and Tanzania. Little is known of the
population status or life history of this species. Habitat destruction may

threaten it in parts of its range. It may occur within the Lokobe Reserve on
Nosy Bé in Madagascar.

Over the six years 1980-1985 inclusive, minimum net trade in P. dubia
reported to CITES amounted to 1460 live animals. The Comores were the largest
exporter and F.R. Germany was the principal importer.

It is not possible to indicate with certainty the effect that this trade has
on the species since the population size is unknown. The level of recent
trade does not appear excessive for such a wide-ranging gecko. Experts have
warned against encouraging trade in Phelsuma.

DISTRIBUTION Known from Madagascar, the Comores, Mayotte, Tanzania
(including Zanzibar) and Mocambique Island.

Comores Inhabits the islands of Grande Comore, Anjouan and Mohéli (Blanc,
1972; Mertens, 1966). Angel (1942) and Mertens (1966) listed localities.

Madagascar P. dubia may be found in the north-west, south-west, west and
south-central regions as well as on the island of Nossi Bé (Angel, 1942;
Blanc, 1972; Jenkins, 1987).

Mayotte Recorded from Mayotte (Mertens, 1966), a dependency of France.

Mozambique Recorded only from Mocambique Island (D.G. Broadley, in litt.,
18 March 1986).

Tanzania Recorded from Zanzibar and the mainland of Tanzania (Loveridge,
1957; Mertens, 1966).

POPULATION Apparently numerous around Majunga, Madagascar (Q. Bloxam, in
litt., 22 January 1986). No information is available for other parts of the
range.

HABITAT AND ECOLOGY Observed on coconut palms in Majunga (Q. Bloxam, in
litt., 22 January 1986) and on tree trunks and fences on Mocambique Island
(D.G. Broadley, in Jlitt., 18 March 1986). No further information is
available.

THREATS TO SURVIVAL None known, but habitat destruction could presumably be
a problem in certain parts of its range. No information has been found
relating to exploitation of this species within the countries of origin.

INTERNATIONAL TRADE Listed on Appendix II of CITES as Phelsuma spp. were
or were likely to be in trade and the species are difficult to differentiate.
The only data available on trade are those contained in the Annual Reports of
the Parties to CITES which are summarized in Tables 1 and 2.

119

Phelsuma dubia

Table 1 Minimum net imports of live Phelsuma dubia reported to CITES.

1980 1981 1982 1983 1984 1985
Austria - - 30 12 - -
Canada 2 - - - - 12
Denmark 4 - 10 - - -
German D.R 20 - - - - -
Germany, F.R. 110 150 213 203 260 190
Japan - - 4 20 - -
Spain - - - - 10 -
Surinam - - Sty - - -
Switzerland 10 10 30 - - -
UK - 20 - - - -
USA - 60 2* - 30 10

- - - - - 25 *

Total 146 240 292 245 300 237

* Captive—bred

Table 2 Reported countries of origin (or exporting country if no origin
reported) and numbers of live Phelsuma dubia reported to CITES.

1980 1981 1982 1983 1984 1985

Countries having or possibly having wild populations of P. dubia
Comores 134 240 290 235 300 212
Countries without wild populations of P. dubia

Czechoslovakia - - 2+2* - = -
Germany, F.R. - - 3 - =
Indonesia - = = = me 21 *
Mauritius 2 - - = = =
Netherlands 30 - = = Z 4 x
Sri Lanka - = = 10 2 »

* Captive-bred

Over the six years 1980-1985 inclusive, minimum net trade in P. dubia
amounted to 1460 live animals. The Comores were the largest exporter,
accounting for 97% of total specimens exported. Exports from Czechoslovakia,
F.R. Germany and the Netherlands were likely to have been re-exports or
captive bred specimens. The principal importer was F.R. Germany.

In addition to the trade reported in this species, F.R. Germany and Austria

reported imports from the Comores of unidentified Phelsuma species. These
are likely to be one of the following species: Phelsuma comorensis, P. dubia,
P. laticauda, P. v-nigra or P. robertmertensi. These were as_ follows:

1980, 150 specimens; 1981, 20; 1983, 20; 1984, 30; 1985, 2. In 1983,

120

Phelsuma dubia

Madagascar exported 32 live unidentified Phelsuma which could have included
P. dubia.

It is not possible to assess the precise effect of trade on the populations of
P. dubia in the absence of population data. The number of identified
specimens together with the possible addition of unidentified Phelsuma does
not appear to be great enough to jeopardise the security of P. dubia, given
its extensive distribution. Bloxam (in litt., 22 January 1986), however,
warned against encouraging trade in Phelsuma as did Gardner (in litt., 23
January 1986) and Thorpe (in litt., 21 January 1986).

CONSERVATION MEASURES
Comores No information. Not a Party to CITES.

Madagascar All wildlife (except vermin) are protected under the Ordinance
on hunting, fishing and the protection of Wildlife (3 October 1960). This
provides for the requirement to obtain permits for the commercial hunting,
possession, sale and of wildlife. It is possible that P. dubia occurs
within the protected area of Lokobe on Nosy Bé (Jenkins, 1987).

Mayotte Mayotte is a dependency of France, intermediate in status between
an Overseas Department and an Overseas Territory. It is not in the EEC, nor
is it thought to be covered by the French ratification of CITES.

Tanzania Under the Wildlife Conservation Act, No. 12, 30 June 1974 (amended
Act No. 21, 1978), all vertebrates are protected and may not be killed,
captured, traded, imported or exported without a permit.

CAPTIVE BREEDING Phelsuma spp. are reputedly easily bred. Jacobi (1982,
unpublished data) reported that, in the Netherlands, 6 were bred in 1977, 4 in
1978, 23 in 1979, 92 in 1980 and 89 in 1981. CITES Annual Reports indicate
the export of captive-bred animals from Indonesia, the Netherlands and
Czechoslovakia. A recent inventory (Slavens, 1985) reported only 1 specimen
in captivity; this is likely to be an underestimate.

REFERENCES

Angel, F. (1942). Les lézards de Madagascar. Memoires de 1'Academie
Malgache 36: 1-193.

Blanc, C.P. 1972. Les reptiles de Madagascar et des iles voisines. In:

Battistini, R. and Richard-Vindard, G. (eds). Biogeography and Ecology
in Madagascar. Dr W. Junk, The Hague, pp 501-614.

Jenkins, M.D. (ed.) (1987). An environmental profile of Madagascar. IUCN
Conservation Monitoring Centre, Cambridge, UK.

Loveridge, A. (1957). Checklist of the reptiles and amphibians of East Africa

(Uganda; Kenya; Tanganyika; Zanzibar). Bulletin of the Museum of
Comparative Zoology 117(2): 153-362.
Mertens, R. (1966). Die nichtmadagassischen Arten und Unterarten der

Geckonengattung Phelsuma. Senckenbergiana biologica 47: 85-110.
Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

121

GOLDDUST DAY GECKO Recommended list: 3*
[No problem]

Phelsuma laticauda (Boettger, 1880)

Order SAURIA Family GEKKONIDAE

*See last paragraph of Summary and Conclusions

SUMMARY AND CONCLUSIONS Widely distributed, occurring in the Comores,

Mayotte, Madagascar and the Seychelles; observations indicate that
P. laticauda is usually abundant. Habitat alterations could affect this
gecko. Possibly occurring in a reserve on Madagascar, the species is

reputedly easily bred.

Over the six years 1980-1985, minimum net trade in live P. laticauda
reported to CITES amounted to 5356 specimens, most of which were in 1985. The
largest exporter overall was the Comores, but in 1985, Madagascar exported 64%
of all specimens. F.R. Germany, the USA, Switzerland, and latterly the
Netherlands were the main importers.

The trade in P. laticauda does not appear to threaten populations of this
species, particularly in view of its wide distribution. However, the high
Number of unidentified Phelsuma reported as exports from the Seychelles
could, if it was correct and if all or the great majority were P. laticauda,
have caused significant pressure on local populations. The increasing exports
form Madagascar are probably not excessive at present, but might become so if
they were to continue. Expert opinion favours maintaining controls on trade.

DISTRIBUTION A widely distributed species occurring in the Comores,
Madagascar, Mayotte and the Seychelles (Blanc, 1972; Cheke, 1984).

Comores P. 1. laticauda has been found on Anjouan (Blanc, 1972; Mertens,
1962). Rehndahl (1939) noted a subspecies, P. 1. comorensis, not identified
in later work.

Madagascar Both P. 1. laticauda and P. 1. angularis have been’ reported
from Madagascar; the species appears to be widespread on the main island, also
on Nosy Bé (Blanc, 1972; Mertens, 1964; Angel, 1942)

Mayotte P. 1. laticauda has been found on Mayotte (Mertens, 1962).

Seychelles P. 1. laticauda has been reported from Farquhar (Blanc, 1972;
Cheke, 1984), Cerf (near Farquhar), Providence, St Anne and Cerf (Mahé group)
(Cheke, 1984).

POPULATION Abundant in the coastal regions of north Madagascar and the
Comores, reaching a high density (Gardner, 1984). Apparently numerous on Nosy
Bé and Nosy Kamba (Q. Bloxam, in litt., 22 January 1986; A.S. Gardner, in
lJitt., 23 January 1986). Density has reached 672 per 100 trees on Farquar;
the species is abundant here, and on Providence (A.S. Gardner, in litt., 23
January 1986).

HABITAT AND ECOLOGY Observed on buildings on the edge of degraded forest in
Nosy Bé and Nosy Kamba (Q. Bloxam, in litt., 22 January 1986).

THREATS TO SURVIVAL Habitat alterations are occurring within the range of
this species and could conceivably affect it. No information is available to
indicate that this species is subject to exploitation within the countries of
origin.

122

Phelsuma laticauda

INTERNATIONAL TRADE Phelsuma spp. were originally included on CITES
Appendix II as they were or could be subject to heavy pet trade and the
species are difficult to tell apart. The only data on international trade are
those contained in the Annual Reports of the Parties to CITES.

Table 1 Minimum net imports of Phelsuma laticauda reported to CITES. All
specimens are live except 29 bodies for scientific purposes, indicated by #.

1980 1981 1982 1983 1984 1985
Argentina = - - - 2 2
Austria - - 30 32 10 40
Canada 2 - - - - 17 +8*
Denmark 20 - Z2 30 - -
German D.R. 20 ~ - - - -
Germany, F.R. 70 670 382 770 469 1332
Japan ~ - - - Yas) 26
Netherlands - - - - 100 325
Spain - - ~ - 10 -
Suriname ~ - 6 - - -
Switzerland 50 10 40 60 53 25
UK - 20 +29# - - - 50
USA 102 170 - 10 131 178 +8*
Total 264 899 480 902 800 2011

* Captive-bred

Table 2 Reported countries of origin (or exporting country if no origin
reported) and numbers of Phelsuma laticauda reported to CITES.

1980 1981 1982 1983 1984 1985

Countries having or possibly having wild populations of P. laticauda

Comores 20 870 480 892 650 700
Madagascar 90 25 - - 150 1295
Seychelles - 29 # - 10 - -

Countries without wild populations of P. laticauda

Czechoslovakia 4 = - - - -
Germany, F.R. - - 6 - - 15
Mauritius 2 - - - - -
Netherlands 70 - - - 25 16 *
Sri Lanka - - - - 10 -

Country unknown 100 - - = = =

* Captive-bred

nc

123

Phelsuma laticauda

The great majority of reported trade was in live animals, presumably for the
pet market. Over the six years 1980-1985, minimum net trade reported to CITES
amounted to 5356 specimens, most of which were recorded in 1985. The largest
exporter of P. laticauda overall was the Comores, but in 1985, Madagascar
exported 64% of all specimens. The F.R. Germany was the largest importer of
this gecko, the numbers imported remaining high over the whole period
covered. The USA, Switzerland, and latterly the Netherlands were also
significant importers.

In addition to the trade reported in this species, the Seychelles reported

exports of large quantities of unidentified Phelsuma species. These are
likely to be one of the following species: Phelsuma abbotti, P. astriata, P.
laticauda, P. longinsulae or P. sundbergi. These exports, mostly to The

UK, F.R. Germany and the USA, were as follows: 1980, 60 specimens; 1981, 1015
kg; 1982, 72 kg; 1983, 28 kg; 1984, 41 kg of bodies. The Comores also
reported exports of unidentified Phelsuma species which are likely to be one
of the following species: Phelsuma comorensis, P. dubia, P. laticauda, P.
v-nigra or P. robertmertensi. These exports, mostly to F.R. Germany and
Austria, were as follows: 1980, 150 specimens; 1981, 20; 1983, 22; 1984, 30;
OBS yee)

The export of unidentified Phelsuma from the Seychelles could cause concern
if the figures can be relied on and if all, or a substantial number, were to
be P. laticauda. The trade could amount to several tens of thousands of
specimens in 1981 which could make an impact on the wild populations. It
should, however, be noted that no such shipments were reported as imports by
other countries and it is possible that errors were made in the Annual Reports
of the Seychelles to CITES. The export of unidentified Phelsuma from the
Comores would not cause such concern even if they were all P. laticauda
since the numbers involved were relatively small.

The Golddust Day Gecko is apparently common over much of its extensive range,
and therefore is unlikely to be deleteriously affected by the trade in this
species which amounted to around 5356 specimens over six years. However, the
recent substantial trade from Madagascar should be monitored and should the
Seychelles trade in unidentified Phelsuma be predominantly P. laticauda,
this could affect the wild populations there. Several experts have expressed
reservations about the advisability of reducing the trade controls (Q. Bloxam,
in 1litt., 22 January 1986; A.S. Gardner, in litt., 23 January 1986; R.
Thorpe, in litt., 21 January 1986).

CONSERVATION MEASURES
Comores No information. Not a Party to CITES.

Madagascar All wildlife (except vermin) are protected under the Ordinance
on hunting, fishing and the protection of Wildlife (3 October 1960). This
provides for the requirement to obtain permits for the commercial hunting,
possession, sale and of wildlife. It is possible that P. laticauda occurs
within the protected area of Lokobe on Nosy Bé (Jenkins, 1987).

Mayotte Mayotte is a dependency of France, intermediate in status between
an Overseas Department and an Overseas Territory. It is not in the EEC, nor
is it thought to be covered by the French ratification of CITES.

CAPTIVE BREEDING Phelsuma spp. are reputedly easily bred. Jacobi (1982,
unpublished records) reported the following captive breeding success in the
Netherlands: 1972, 1 specimen; 1973, 2 specimens; 1974 10 specimens; 1976, 4
specimens; 1977, 45 specimens; 1980, 79 specimens; 1981, 85 specimens. CITES

124

Phelsuma laticauda

Annual Reports also indicate that captive-breeding is occurring in the
Netherlands. A recent inventory (Slavens, 1985) reported 25 specimens of P.
laticauda in 9 collections; the numbers are likely to be higher.

REFERENCES

Angel, F. (1942). Les lézards de Madagascar. Mémoires de 1'Académie de
Malgache 36: 1-193.

Blanc, ¢€.P. (1972). Les reptiles de Madagascar et des iles voisines. In:

Battistini R. and Richard Vindard, G. (eds) Biogeography and Ecology in
Madagascar. Dr W Junk, The Hague, pp. 501-614.

Cheke, A.S. (1984). Lizards of the Seychelles. In: Stoddart, D.R. (ed.)
Biogeography and ecology of the Seychelles Islands. Dr W Junk, The
Hague, pp. 332-360.

Gardner, A.S. (1984). The evolutionary ecology and population systematics of
Day Geckoes (Phelsuma) in the _ Seychelles. Ph.D. Thesis, Aberdeen
University.

Jenkins, M.D. (ed.) (1987). An environmental profile of Madagascar. IUCN
Conservation Monitoring Centre, Cambridge, UK, 374 pp.

Mertens, R. (1962). Die Arten und Unterarten der Geckonengattung
Phelsuma. Senckenbergiana biologica 43: 81-127

Mertens, R. (1964). Funf neue Rassen die Geckonengattung Phelsuma.
Senckenbergiana biologica 45: 99-112

Mertens, R. (1966). Die Nichtmadagassischen Arten und Unterarten der
Geckonengattun Phelsuma. Senckenbergiana biologica 47: 85-110.

Rendahl, H. C4939). Zur Herpetologie der Seychellen, I. Reptilien.

Zoologisches Jahrbuch 72: 157-328.
Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

125

Recommended list: 3
[No problem]
Phelsuma madagascariensis Gray 1831

Order SAURIA Family GEKKONIDAE

SUMMARY AND CONCLUSIONS Endemic to Madagascar, populations of this species
occur over much of the island and the species seems to be abundant, at least
in places. Known from forests, the loss of habitat by fire and for slash and
burn cultivation could affect the status of this gecko. It may occur in the
Réserve Naturelle Integrale de Lokobé on Nosy Bé.

The minimum net trade in P. madagascariensis reported to CITES from 1980 to
1985 amounted to 2189 live animals and 33 scientific specimens. Levels of
trade remained low until 1984, when Madagascar, the only country which has
wild populations of this species, reported exporting substantial quantities.
The main importers were F.R. Germany, the USA and the Netherlands.

It is difficult to assess the significance of the trade, since nearly half of
the specimens in trade appear to have been misidentified. The volume reported
to CITES was not large, considering that the species is widespread in
Madagascar, and it is unlikely to have any adverse effects on the
populations. However the trade from Madagascar seems to be increasing and it
should continue to be monitored. Expert opinion does not favour encouraging
trade in Phelsuma.

DISTRIBUTION P. madagascariensis is endemic to Madagascar (Blanc, 1972)
and appears to occur in most parts of the island (Angel, 1942).

POPULATION Little information is available on the population size of this
species. In 1942, Angel reported that it was very common around Diego Suarez
and frequent in the forests of the north-west. Bloxam (in litt., 23 January
1986) considered that P.m. kochi and P.m. grandis could be_ fairly
numerous.

HABITAT AND ECOLOGY South of Tamatave, P. madagascariensis is found in
coastal regions . In the north-west it is known from forest (Angel, 1942).
Bloxam (in Jlitt., 23 January 1986) observed P. m. kochi in the deciduous
forest of Morandava and P. m. grandis in degraded forest on Nosy Bé.

THREATS TO SURVIVAL Vegetation destruction is likely to affect this
species. Forest is disappearing for slash and burn cultivation and fire is
also depleting the forest cover. Rice and manioc are being grown on Nosy Bé
(Jenkins, 1987). No information has been found relating to exploitation of
this species within Madagascar.

INTERNATIONAL TRADE Originally included on CITES Appendix II as Phelsuma
spp. were or could be subject to heavy pet trade and the species are difficult
to differentiate. The only data on international trade are those contained in
the Annual Reports of the Parties to CITES (Tables 1 and 2).

The minimum net trade in P. madagascariensis reported to CITES from 1980 to
1985 amounted to 2189 live animals and 33 scientific specimens. Levels of
trade remained relatively low until 1984, when Madagascar, the only country
which has wild populations of this species, reported exporting substantial
quantities. The specimens reported as being exported from the Seychelles may
be P. sundbergi (formerly P. madagascariensis sundbergi); this Seychelles
endemic is widespread and abundant (A.S. Gardner, in litt., 23 January
1986). Specimens exported from the Comores are likely to have _ been

126

Phelsuma madagascariensis

misidentified and could be one of the following: P. comorensis, P. dubia,
P. laticauda, P.v-nigra or P. robertmertensi. The main importers. of
P. madagascariensis were F.R. Germany, the USA and the Netherlands.

Table 1 Minimum net imports of Phelsuma madagascariensis reported to CITES.
All specimens were live except 33 scientific specimens, indicated by #.

oo —————eeeeeeeeeeeeeeeeoeowNUN____

1980 1981 1982 1983 1984 1985
re eee eee eee eee eee
Austria - = = = & 32 43%
Canada d - 2 8 - - -
Denmark 16 - 2 - 17 * -
Germany, F.R. - 75: 61 87 17 682
France - - - - 60 -

Japan - - 6 14 4 26
Netherlands - - - 2 200 56
Sweden - - - 4 - -
Switzerland 2 - 16 34 14 15 +12*
UK - 33)# - - - 38

USA 44 54 72 8 30 +134* 162 +180*
Total 62 164 165 149 476 1206

* Captive-bred

Table 2 Reported countries of export (or exporting country if no origin
reported) and numbers of Phelsuma madagascariensis reported to CITES. All
specimens were live except 33 scientific specimens, indicated by #.

1980 1981 1982 1983 1984 1985

Countries with wild populations of P. madagascariensis
Madagascar - - - - 460 1169

Countries without wild populations of P. madagascariensis

Austria = - = 4 = =
Comores 16 30 85 34 - -
Czechoslovakia 7 - - - - -
Germany, F.R. 5 15 - - 44 * -
German D.R. - - - 14 6 * 12 *
Guyana - - - 4 - -
Indonesia - - - - - 12 x
Mauritius 20 - - - - -
Netherlands - - - 4 107 * 168 *
Seychelles - 103 +33* 80 103 - -
USA - - 8 - - -
Country unknown 14 - - 10 - -

* Captive-bred

127

Phelsuma madagascariensis

It is difficult to fully assess the trade in this species, an endemic of
Madagascar, since nearly half of the specimens in trade appear to have been
misidentified. The volume reported to CITES not large, considering that the
species is widespread in Madagascar, and it is unlikely to have any adverse
effects on the populations. However the trade from Madagascar seems to be
increasing and it should continue to be monitored. Bloxam (in Jlitt., 22
January 1986) considers that trade in Phelsuma should not be encouraged.

CONSERVATION MEASURES All wildlife (except vermin) are protected under the
Ordinance on hunting, fishing and the protection of wildlife (3 October
1960). This provides for the requirement to obtain permits for the commercial
hunting, possession, sale and export of wildlife. It is likely that this
species occurs in the Réserve Naturelle Intégrale de Lokobe on Nosy Bé.
(Jenkins, 1987).

CAPTIVE BREEDING Phelsuma spp. are reputedly easily bred. Jacobi (1982,
unpublished data), reported that, in the Netherlands, 21 specimens were bred
in 1972, 14 in 1978, 7 in 1979, 65 in 1980 and 97 in 1981. A recent inventory
(Slavens, 1985) reported 111 specimens of P. m. grandis in 24 localities, 19
specimens of P. m. kochi in 4 locations, 1 specimen of P. m.
Madagascariensis in 1 location and 48 specimens of P. madagascariensis
(subspecies unidentified) in 5 locations. This is likely to be an
underestimate. CITES trade reports indicate that significant numbers may be
being bred in the Netherlands and possibly also F.R. Germany.

REFERENCES

Angel, F. (1942) Les lézards de Madagascar. Mémoires de _ 1'Académie
malgache 36: 1-190

Blanc, C.P. (1972). Les reptiles de Madagascar et des iles voisines. In:

Battistini, R. and Richard-Vindard, G. (eds). Biogeography and Ecology
in Madagascar. Dr. W. Junk, The Hague, pp 501-614.

Borner, A.R. (1972). Revision der Geckonengattung Phelsuma Gray 1825.
Saurologia 1: 1-145.
Jenkins, M.D. (1987). An environmental profile of Madagascar. IUCN

Conservation Monitoring Centre, Cambridge, UK.
Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Salvens, Seattle, Washington, 341 pp.

128

Recommended list: 3

[No problem]
Phelsuma v-nigra (Boettger, 1913)

Order SAURIA Family GEKKONIDAE

—————— ES ES EE EE ee ee ee ee eee

SUMMARY AND CONCLUSIONS Found in coastal regions of the Comores Islands and
Mayotte; no data are available regarding status and life history.

P. v-nigra may be affected by habitat destruction. Reputedly easily bred in
captivity.

Minimum net trade in P. v-nigra reported to CITES amounted to 641
specimens. All trade was in live animals, presumably for the pet trade. The
principal exporter was the Comores where 90% of the reported trade
originated. Those specimens reported from Mauritius and the Seychelles are
likely to be wrongly identified specimens since this gecko does not occur in
either locality. F.R. Germany and the USA were the main importers.

Declared volume of trade is low and is unlikely to have a detrimental effect
on the species as a whole, it is conceivable, however, that populations are
locally affected. Expert opinion does not favour encouraging trade in
Phelsuma spp.

DISTRIBUTION Known only from the Comores and Mayotte. P. v-nigra is
sometimes considered a subspecies of P. abbotti.

Comores Known only from islands of Grande Comore, Anjouan and Mohéli
(Blanc, 1972; Mertens 1966).

Mayotte Recorded from Mayotte (Mertens 1966).
POPULATION No information.

HABITAT AND ECOLOGY This species occurs in coastal regions up to altitudes
of 300 m (Angel, 1942).

THREATS TO SURVIVAL Vegetation destruction is occurring and is particularly
serious on Anjouan where extensive areas of the coastal zone have been cleared
(Tattersall, 1977). No information has been found relating to exploitation
within the Comores.

INTERNATIONAL TRADE The only data available are those contained in the
Annual Reports of the Parties to CITES and are summarized in Tables 1 and 2.

Minimum net trade in P. v-nigra reported to CITES amounted to 641
specimens. All trade was in live animals, presumably for the pet trade. The
principal exporter was the Comores where 90% of the reported trade
Originated. Those specimens reported from Mauritius and the Seychelles are
likely to be wrongly identified specimens since this gecko does not occur in
either locality. F.R.Germany and the USA were the main importers.

In addition to the trade in this species, imports of unidentified Phelsuma
species were recorded from the Comores. These are likely to be one of the
following species: Phelsuma comorensis, P. dubia, P. laticauda, P. v-nigra
(all species treated under this contract) or P. robertmertensi. These
imports, mostly to F.R. Germany and Austria, were as follows: 1980, 150
specimens; 1981, 20; 1983, 20; 1984, 30; 1985, 2. Should all the unidentified

129

Phelsuma v-nigra

Phelsuma spp. have been P. v-nigra, a possible addition of 222 specimens
were in trade. The effect of this trade cannot be ascertained precisely since
there are no data relating to population size or life history requirements.
In view of the fact that this species is found on four islands, it is likely
that there is no great impact on the species, although populations in
accessible areas could be depleted. Bloxam (in litt., 22 January 1986),
Gardner (in litt., 23 January 1986) and Thorpe (in litt., 21 January 1986)
all consider that trade in Phelsuma should not be encouraged.

Table 1 Minimum net imports of live Phelsuma v-nigra reported to CITES.

1980 1981 1982... 1983 1984 1985
Austria - - - 2 10 -
Denmark 20 = 10 5 - -
Germany, F.R. - 48 110 123 - 80
Switzerland 30 - 20 29 - -
UK - 20 - - - -
USA 12 112 - - 10 -
Total 62 180 140 159 20 80

Table 2 Reported countries of origin (or exporting country if no origin
reported) and numbers of live Phelsuma v-nigra reported to CITES.

eee

1980 1981 1982 1983 1984 1985

ees

Countries having or possibly having wild populations of P. v-nigra
Comores 20 160 140 159 20 80

Countries without wild populations of P. v-nigra

Germany, F.R. 2 2 — = = =
Mauritus 10 = = = = =
Netherlands 30 = = = = =
Seychelles - 20 = = = =

CONSERVATION MEASURES Information is required on the population status and
ecology of this species.

Comores No information. Not a Party to CITES.

Mayotte Mayotte is a dependency of France, intermediate in status between
an Overseas Department and an Overseas Territory. It is not in the EEC, nor
is it thought to be covered by the French ratification of CITES.

CAPTIVE BREEDING Phelsuma spp. are reputedly easily bred. Jacobi (1982,
unpublished data) reported that, in the Netherlands, 10 P. v-nigra were bred
in 1978, 17 in 1979, 49 in 1980 and 35 in 1981. A recent inventory (Slavens,

130

Phelsuma v-nigra

1985), only reported one specimen in captivity; this is likely to be an
underestimate.

REFERENCES

Angel, F. (1942). Les lézards de Madagascar. Mémoires de _ 1'Académie
malgache 36: 1-190.

Blanc, C.P. (1972). Les reptiles de Madagascar et des iles voisines. In:

Battistini, R. and Richard Vindard, G. (eds). Biogeography and Ecology
in Madagascar, Dr W. Junk, The Hague, pp 501-614.

Mertens, R. (1966). Die michtmadagassischen Arten und Unterarten der
Geckonengattung Phelsuma. Senckenbergiana biologica 47: 85-110

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

Tattersall, ibe (1977). The lemurs of the Comore’ Islands. Oryx
8(5): 445-448.

131

Recommended list: 3*
[No problem]

Chamaeleo bitaeniatus Fischer, 1884

Order SAURIA Family CHAMAELEONIDAE

*See last sentence of Summary and Conclusions

SUMMARY AND CONCLUSIONS Widespread in eastern Africa from Ethiopia and
Sudan south through Kenya and Uganda to Tanzania, possibly including Somalia
and Zaire; no information is available on the population status of
C. bitaeniatus. Little is known of the life history in the wild, some
studies have been made on captive specimens. Apparently difficult to maintain
in captivity and requiring special conditions. Possibly found within
protected areas. ;

Trade in this species for the period 1980-83 was only reported in 1980 and
1981, Kenya being responsible for all exports (around 3500 in total) and the
F.R. Germany and USA importing most specimens. Export from Kenya (and thus
all reported trade) appears to have ceased since 1981, despite
C. bitaeniatus not being listed on 1981 legislation which specifically
banned export of three other Kenyan Chameleon species.

There is no evidence of the species, or local populations thereof, having been
adversely affected by trade volumes as reported to CITES. Although unreported
trade may exist, the known volume of international trade in the early 1980s is
almost certain not to have been a significant problem. The inclusion in list
3 is dependent on Kenya maintaining her apparent ban on export of the species.

DISTRIBUTION Widespread; reported from Ethiopia and Sudan, south through
Kenya, Uganda to north-west Tanzania (Loveridge, 1957; Mertens, 1966; Rand,
1963), possibly including Somalia and north-eastern Zaire.

Ethiopia Recorded from Addis Ababa, between Sancurrar and Amarr, and
between Badditu and Oime (Rand, 1963).

Kenya Most Kenyan records are from the Rift Valley and adjacent areas as
far as Mt Elgon in the West (Rand, 1963; J. Hebrard, in litt., 1 April
1986).

Somalia Although Loveridge (1957) reported that the distribution of C. b.
bitaeniatus included Somalia, Rand (1963) indicated that the only record from
Somalia (Coronna) was doubtful. Simonetta and Magnoni (1986) did not list the
species from Somalia.

Sudan Recorded from the Podocarpus forest on Mt Kinyeti in the Imantong
Mountains (BOhme and Klaver, 1980).

Tanzania Restricted to Longido West (Rand, 1963; K.M. Howell, in litt.,
15 March 1986).

Uganda Loveridge (1957) reported that the distribution of C. b.
bitaeniatus extended from Ethiopa and Kenya “south through Uganda". Rand
(1963) recorded no specimens from Uganda, but included some from the Kenyan
side of Mt Elgon. The locality in the Imantong Mountains of Sudan (BoOhme and
Klaver, 1980) is close to Uganda's northern border.

Zaire Howell (in Jitt., 15 March 1986) noted that this species occurred

in north-eastern Zaire, but no other records have been located in the
literature.

132

Chamaeleo bitaeniatus

POPULATION No information available. This species is said to be small and
extremely cryptic (J. Hebrard, in Jlitt., 1 April 1986), and so may be
overlooked.

HABITAT AND ECOLOGY A small and extremely cryptic chameleon. Occurs in
grasslands, with or without scattered bushes, and attains high population
density in stands of Acacia drepanolobium (where ants form the main prey)
(J. Hebrard, in litt., 1 April 1986).

A viviparous species; 3-4 broods may be produced per year, each with around 17
young (Bustard, 1966; Schmidt and Inger, 1957; de Vosjoli, 1979). The
gestation period is not accurately known, but is at least three months. Both
males and females tend to have a regular perch for basking and spending the
night.

THREATS TO SURVIVAL None known other than capture for the live animal trade.
INTERNATIONAL TRADE The only data available on international trade are
those contained in Annual Reports of the Parties to CITES which are summarized

in Table 1.

Table 1 Minimum net imports of live Chamaeleo bitaeniatus reported to CITES.

1980 1981 1982 1983 1984 1985
Germany, F.R. 1460 474 - = = <
Switzerland 90 35 — - = =
UK - 57 - = = =
USA 500 1010 - = = =
Total 2050 1576 0 0 0 0

All specimens reported in trade were live, suggesting that they were intended
for the pet trade. Trade only occurred in 1980 and 1981 and all reported
exports originated in Kenya. The F.R. Germany and USA were the principal
importers of this chameleon, taking 53% and 42% of the total imports,
respectively

Although no trade in this species took place after 1981, prior to that date,
the reported trade did not seem unduly high in view of the extensive
distribution of this species. Specimens were however taken from only a
limited part of the total range. Since population and life history data are
lacking, it is not possible to assess the true impact on wild populations.

CONSERVATION MEASURES Unless otherwise stated, all of the information on
protection is from IUCN Environmental Policy and Law Occasional Paper No. 3,
African Wildlife Laws.

Ethiopia Under the Wildlife Conservation (Amendment) Regulations, 12
February 1974, Chameleons are classified as game, and may only be captured by
licensed trappers and dealers under a Game Capture Permit, the value of which
was set as Eth$0.10 per animal.

133

Chamaeleo bitaeniatus

Kenya Kenya has prohibited the export of three other species of chameleon,
but these do not include C. bitaeniatus. There are several protected areas
where this species may occur including the National Parks at Lake Nakuru, and
Mt Kenya.

Somalia A ban on all hunting was instituted on 13 October 1977.

Sudan Chameleons are not listed in the Ordinance for the Preservation of
Wild Animals, 1935 (Amended 1974), as species for which hunting is permitted.
Licences are only required if firearms are used for hunting. The Hides and
Skins (Export) Regulations, 1969, specify a grading system for the export of
reptile skins.

Tanzania Under the Wildlife Conservation Act, 30 June 1974, the capture of
all live animals requires a valid capture permit, and the hunting of all
animals requires a valid hunting permit.

Uganda No reptiles, other than crocodiles, are protected.

Zaire The commercial capture of unprotected animals requires a permit under
the Hunting Act, 28 May 1982. Export can only be authorised if the exporter
holds a certificate of lawful possession.

CAPTIVE BREEDING Chameleons are reputedly difficult to keep in captivity
over long periods. De Vosjoli (1979) provided details of suitable breeding
conditions, particularly regarding temperature, humidity, lighting, cage size
and layout and feeding; see also Bustard (1966). Formerly maintained at the
Chameleon Research Center (de Vosjoli, 1979); this centre no longer exists. A
recent inventory (Slavens, 1985) does not record any individuals in live
collections, although there are likely to be some.

REFERENCES

Bohme, W and Klaver, C.J.J. (1980). The systematic status of Chamaeleo
kinetensis Schmidt, 1943, (Sauria: Chamaeleonidae) from the Imantong
Mountains, Sudan, with comments on lung and hemipenial within the
C. bitaeniatus-group. Amphibia-Reptilia 1(1): 3-17.

Bustard, H.R. (1966). Observations on the life history and behaviour of
C. bitaeniatus. Herpetologia 22: 13-23.

Loveridge, A. (1957). Checklist of the reptiles and amphibians of East Africa

(Uganda, Kenya, Tanganyika, Zanzibar). Bulletin of the Museum of
Comparative Zoology 117: 153-362.

Mertens, R. (1966). Liste der rezenten Amphibien und Reptilien. Das
Tierreich 83: 1-37.

Rand, A.S. (1963). Notes on the Chamaeleo bitaeniatus complex.

Bulletin of the Museum of Comparative Zoology 130: 1-29.

Schmidt, K.P. and Inger, R.I. (1957). Living reptiles of the world.
Hamish Hamilton, 287 pp.

Simonetta, A.M. and Magnoni, M.L. (1986). Status and conservation problems of
Somali lower’ vertebrates. Revista di Agricoltura Subtropicale e
Tropicale 53(3): 405-432.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

de Vosjoli, P. (1979). The care, maintenance and breeding of the African
Chamaeleo. Journal of the Chameleon Research Center 2: 7-38.

134

Recommended list: 3*

[No problem]
Chamaeleo gracilis Hallowell, 1842

Order SAURIA Family CHAMAELEONIDAE

*See last sentence of Summary and Conclusions

—_—.eO LLL

SUMMARY AND CONCLUSIONS A wide-ranging species, extending across tropical
Africa from Senegal east to Somalia, and south to Zaire and Tanzania. Present
in Angola, Equatorial Guinea (Bioko), Cameroon, Central African Republic,
Congo, Ethiopia, Gabon, Gambia, Ghana, Guinea, Ivory Coast, Kenya, Liberia,
Nigeria, Senegal, Sierra Leone, Somalia, Sudan, Tanzania, Uganda, Zaire, and
possibly Benin, Togo, Chad and Mali. Little information is available on the
life history and population status of C. gracilis. An insectivore which
prefers open country, international trade is the only known possible threat.
Breeding can occur in captivity, but chameleons are reputedly difficult to
maintain.

Declared trade in the period 1980-85 was only reported in 1980, 1981 and 1985
and amounted to a total of around 2200 specimens. The principal importers
were the USA and F.R. Germany. Kenya was responsible for the export of 77% of
all specimens exported in 1981 and 1982, but then the Kenyan export ceased
following legislation to prohibit trade. Tanzania and Togo were the main
suppliers in 1985.

It is unlikely that the relatively low volume of trade declared up to 1981 had
a deleterious affect on the species, although in the absence of population and
life history data this cannot be stated with certainty, and it is possible
that populations were affected locally. International trade cannot be
regarded as a Significant threat to the species, provided that there is no
appreciable unreported trade and that the Kenyan export ban remains effective.

DISTRIBUTION A very wide-ranging species, C. gracilis extends across
tropical Africa from Senegal east to Somalia, and south to Zaire and
Tanzania. Present in Angola, Equatorial Guinea (Bioko), Cameroon, Central
African Republic, Congo, Ethiopia, Gabon, Gambia, Ghana, Guinea, Ivory Coast,
Kenya, Liberia, Nigeria, Senegal, Sierra Leone, Somalia, Sudan, Tanzania,
Uganda, Zaire (Loveridge, 1957; Mertens, 1966; Schmidt 1919), and possibly
Benin, Togo, Chad and Mali (Groombridge, 1983). The nominate’ form
C. g. gracilis occupies most of the range, giving way to C. g. etiennei in
Gabon, Congo, Zaire and Angola (Mertens, 1966).

Angola C. g. etiennei has been recorded from Angola (Mertens, 1966;
Laurent, 1964).

Benin Not recorded by Mertens (1966), although as it occurs in nearby
countries, Nigeria and Ghana, it might be expected to occur there and was
listed by Groombridge (1983). Loveridge (1957) pointed out that it had not
been recorded from Dahomey.

Cameroon C. g. gracilis was recorded by Mertens (1966).

Central African Republic C. g. gracilis was recorded by Mertens (1966).

Chad Listed as present by Groombridge (1983), but no record has been
located.

135

Chamaeleo gracilis

Congo Both C. g. gracilis and C. g. etiennei were recorded from Congo
(Brazzaville) (Mertens, 1966). Loveridge (1957) stated that C. g. gracilis
had not been recorded from the French Congo.

Djibouti Groombridge (1983) listed the species as probably occurring in
Djibouti, but this seems unlikely as in neighbouring Somalia it appears to be
confined to the south (Simonetta and Magnoni, 1986).

Equatorial Giunea C. burchelli, listed as a synonym of C. g. gracilis
by Mertens (1966), was recorded from Fernando Poo (Bioko), although Mertens
did not include either the island or the mainland of Equatorial Guinea in his
distribution.

Ethiopia C. g. gracilis was recorded by Mertens (1966).

Gabon Both C. g. gracilis and C. g. etiennei were recorded from Gabon
(Mertens, 1966).

Gambia Recorded by Hakansson (1981).

Ghana C. g. gracilis was recorded by Mertens (1966).

Guinea C. g. gracilis was recorded by Mertens (1966).

Ivory Coast C. g. gracilis was recorded by Mertens (1966).

Kenya Occurs in two different habitats in Kenya: arid thorn scrub and
semi-desert; and well watered agricultural land in western Kenya (J.L Hebrard,
in litt., 1 April 1986).

Liberia C. g. gracilis was recorded by Mertens (1966).

Mali Listed as present by Groombridge (1983), but no record has _ been
located.

Nigeria C. g. gracilis was recorded by Mertens (1966). Butler and Shitu
(1985) described traditional uses of the species in Nigeria.

Senegal C. g. gracilis was recorded by Mertens (1966).
Sierra Leone C. g. gracilis was recorded by Mertens (1966).

Somalia Recorded from Middle Shebelle, Mogadishu, Lower Shebelle, Bay, Gedo
and Lower Juba, all in southern Somalia (Simonetta and Magnoni, 1986).

Sudan C. g. gracilis was recorded by Mertens (1966).

Tanzania Found only at Longido and Mt Meru in northern Tanzania (K.M.
Howell, in litt., 15 March 1986).

Togo Not recorded by Mertens (1966), although as it occurs in nearby
countries, Nigeria and Ghana, it might be expected to occur there, and was
listed by Groombridge (1983).

Uganda C. g. gracilis was recorded by Mertens (1966).

Zaire Both C. g. gracilis and C. g. etiennei were recorded from Zaire
(Mertens, 1966) Lanza and Vanni (1976) recorded C. g. gracilis from the
north.

136

Chamaeleo gracilis

POPULATION Virtually no information is available on the population size of
this animal. Marked population fluctuations have been noted in dry parts of
the range (J. Hebrard, in litt., 1 April 1986).

Congo C. gracilis is said to be very abundant in the country (Congo CITES
MA, 1986).

Kenya In Tsavo National Park, for example, the habitat may become uniformly
saturated with young animals when they hatch during the short rains (around
November); most of these subsequently die and only a few isolated adults may
remain by the time of the long rains (April). The species was said to be
extremely rare in Kora National Reserve in 1984 (J. Hebrard, in litt., 1
April 1986).

Sudan Schmidt (1919) reported that C. gracilis was abundant in the east.

HABITAT AND ECOLOGY Apparently an inhabitant of open country, although
occasionally recorded from forest (in Cameroon and Liberia) (Schmidt, 1919).
The species occurs in two distinct habitats in Kenya; in well-watered
agricultural land in western Kenya, and in arid thorn scrub and semi-desert in
other parts of the country (J. Hebrard, in litt., 1 April 1986). Lanza and
Vanni (1976) recently collected specimens in a savannah habitat in Zaire.

This species deposits its eggs in a hole dug in the ground. The eggs are laid
when the rainy season is ending and hatching occurs before the next rains
(Menzies, 1958). Schmidt (1919) reported that C. g. gracilis lives chiefly
on Orthoptera and that C. g. etiennei feeds on a variety of insects
especially grasshoppers, crickets, cockroaches and flies.

THREATS TO SURVIVAL Butler and Shitu (1985) reported that the Yorubas in
Nigeria use various parts of C. gracilis in tribal medicine. They also
believe that stepping in the excreta causes elephantiasis. In Congo, the only
use is said to be in traditional medecine (Congo CITES MA, 1986). No exports
are said to have originated in Uganda, and the chief threat is thought to be
from fires in the dry season (Uganda Game Department, in Jitt., 28 March
1987).

INTERNATIONAL TRADE The only available data are those provided in the
Annual Reports of the Parties to CITES which are summarized in Tables 1 and
2. This species was only reported in trade in 1980, 1981 and 1985. The
principal importers were the USA and F.R. Germany. The specimens imported by
the G.D.R. in 1980 were re-exports from the F.R. Germany. Kenya was
responsible for the export of 77% of all specimens exported in 1981 and 1982,
but then the Kenyan export ceased following legislation to prohibit trade.
Tanzania and Togo were the main suppliers in 1985.

Table 1 Minimum net imports of live Chameleo gracilis reported to CITES.

1980 1981 1982 1983 1984 1985
Germany, F.R. 350 230 - - - 370
German D.R. 30 ~ = = = =
UK 300 34 - - - -
USA 375 503 - - - 5
Total 1055 767 0 0 0 375

Chamaeleo gracilis
Table 2 Reported countries of origin (or exporting country if no origin
reported) and numbers of live Chamaeleo gracilis reported to CITES.

1980 1981 1982 1983 1984 1985

Countries having wild populations of C. gracilis

Ghana 300 - - - - 5
Kenya 755 692 - = = ix
Tanzania - = = = = 250
Togo = = = - - 120
Country unknown - 75 - ‘ = = =

Exports until 1985 were not so large as to cause concern for the overall
status of this species, however, populations in accessible regions could be
affected. The precise effects of trade in this species cannot however be
ascertained since population and life history data are lacking.

CONSERVATION MEASURES Unless otherwise stated, all of the information on
protection is from IUCN Environmental Policy and Law Occasional Paper No. 3,
African Wildlife Laws.

Angola Apparently not protected.
Benin Chameleons are not protected.

Cameroon Under the Forests, Wildlife and Fisheries Act, 27 November 1981,
bag limits are set for the hunting of all species.

Central African Republic The Ordinance concerning the protection of
wildlife and regulating hunting, 27 July 1984, does not list chameleons, but
all wildlife species may only be taken by customary hunters or by the holders
of hunting licences.

Chad No information.

Congo The Act concerning the conservation and exploitation of wild fauna,
21 April 1983, vests in the State ownership of all wild animals of economic
value, and requires the issuing of licences for commercial capturing.

Ethiopia Under the Wildlife Conservation (Amendment) Regulations, 12
February 1974, Chameleons are classified as game, and may only be captured by
licensed trappers and dealers under a Game Capture Permit, the value of which
was set as Eth$0.10 per animal.

Gabon The Wildlife and Forests Act, 22 July 1982 requires the issuing of

licences for the commercial capture of all wildlife. Traditional hunting for
subsistence pruposes is permitted.

Gambia Under the Wildlife Conservation Act, 14 February 1977, all wildlife
except game and vermin are protected.

138

Chamaeleo gracilis

Ghana The Wildlife Conservation Regulations, 4 March 1971, define all
wildlife, other than protected species, as game animals and establish hunting
restrictions.

Guinea No information.

Guinea-Bissau The Hunting Regulations, 12 May 1980, require the issuance of
a licence for the capturing of live wild animals.

Ivory Coast Chameleons are not specifically listed in the Wildlife and
Hunting Act, 4 August 1965, but Arrété No. 15, 26 December 1972, establishes
licence fees for the capturing of all live reptiles.

Kenya Kenya has prohibited the export of C. gracilis unless the written
permission of the Minister for Environment and Natural Resources is obtained
(Legal Notice 152, 25 September, 1981).

Liberia Apparently not protected.

Mali A general hunting ban (Decree 325/PG-RM) throughout the country was
imposed on 6 November 1978.

Nigeria Apparently not protected.

Senegal The Game and Wildlife Protection Regulations, 30 May 1967, lay down
regulations governing the issuance of licences for hunting and commercial
capturing of wildlife.

Sierra Leone Apparently not protected.
Somalia A ban on all hunting was instituted on 13 October 1977.

Sudan Chameleons are not listed in the Ordinance for the Preservation of
Wild Animals, 1935 (Amended 1974), as species for which hunting is permitted.
Licences are only required if firearms are used for hunting. The Hides and
Skins (Export) Regulations, 1969, specify a grading system for the export of
reptile skins.

Tanzania Under the Wildlife Conservation Act, 30 June 1974, the capture of
all live animals requires a valid capture permit, and the hunting of all
animals requires a valid hunting permit.

Togo Apparently not protected.
Uganda No reptiles, other than crocodiles, are protected.

Zaire The commercial capture of unprotected animals requires a permit under
the Hunting Act, 28 May 1982. Export can only be authorised if the exporter
holds a certificate of lawful possession.

CAPTIVE BREEDING Menzies (1958) described the breeding behaviour of this
chameleon in captivity and reported that a captive specimen laid 45 soft
shelled eggs, which hatched after 219 days. De Vosjoli (1979) provided
guidance on the care, breeding and maintenance of the African chameleons,
which are difficult to maintain over a long period of time. Formerly kept at
the Chameleon Research Center (Riva, 1979); this centre no longer exists. A
recent inventory (Slavens, 1985) reported that three specimens were maintained
in two collections. This is likely to be an underestimate.

139

Chamaeleo gracilis

REFERENCES

Butler, J.A. and Shitu, E. (1985). Uses of some reptiles by the Yoruba people
of Nigeria. Herpetological Review 16(1): 1516.

Groombridge, B. (1983). World checklist of threatened amphibians and
reptiles. 3rd Bdition. Nature Conservancy Council, UK, 65 pp.

Hakansson, N.T. (1981). An annotated checklist of reptiles known to occur in
the Gambia. Journal of Herpetology 15: 155-161.

Lanza, B. and Vanni, S. (1976). On a small collection of reptiles from north
Zaire. Monitore Zoologica Italiano 8(3): 129-160.

Laurent, R.F. (1964). Reptiles et amphibiens de 1l'Angola (Troisiéme
contribution). Mus. Dundo Publ. Cult. 67: 1-165.

Loveridge, A. (1957). Checklist of the reptiles and amphibians of East Africa
(Uganda, Kenya, Tanganyika, Zanzibar). Bulletin of the Museum of
Comparative Zoology 117:153-362.

Menzies, J.I1. (1958). Breeding behaviour of the Chameleon (Chamaeleo
gracilis) in Sierra Leone. British Journal of Herpetology 2: 130-132.
Mertens, R. (1966). Liste der rezenten Amphibien und Reptilien.

Chamaeleonidae. Das Tierreich 8: 31-37

Riva, D. (1979). Present CRC Breeding Projects. Journal of the Chameleon
Research Center 2: 4.

Schmidt, K.P. (1919). Contributions to the herpetology of the Belgian Congo
based on the collection of the American Museum Congo Expedition
1909-1915. Part 1. Turtles, Crocodiles, Lizards and Chamaeleons.
Bulletin of the American Museum of Natural History 39: 385-624.

Simonetta, A.M. and Magnoni, M.L. (1986). Status and conservation problems of
Somali lower vertebrates. Revista di <Agricoltura Subtropicale e
Tropicale 53(3): 405-432.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

de Vosjoli, P. (1977). The care, maintenance and breeding of the African
Chamaeleo. Journal of the Chameleon Research Center 2: 7-42.

140

Recommended list: 3%
[No problem]
Chamaeleo hoehnelii Steindachner 1891

Order SAURIA Family CHAMAELEONIDAE

*See last sentence of Summary and Conclusions
ea ee ee eee err ree!

SUMMARY AND CONCLUSIONS Found in upland areas of Kenya and Uganda,
C. hoehnelii is known to be abundant in some areas and thrives in areas of
dense human habitation. An inhabitant of forest edges and low shrubby
vegetation, this viviparous species may produce around 14 young twice a year.
Probably occurring within some protected areas, no specific threats to this
species are known. It will breed regularly under correct conditions but is
reputedly difficult to maintain in captivity for any length of time.

The only reported trade in the period 1980-85 was in 1980, 1981 and 1985 and
totalled some 12 700 specimens; 90% of these were recorded as origin Kenya.
Export from Kenya appears to have ceased since 1981, despite C. hoehnelii
not being listed on 1981 legislation which specifically banned export of three
other Kenyan Chameleon species.

The legal status of this species in Kenya should be determined. it) is
possible that the relatively high volume of trade recorded in 1980 and 1981
could have led to local depletions, but if, in addition to the apparent
cessation of reported trade since 1981, there has been no appreciable
unreported trade, international trade is most unlikely to pose a significant
threat to the species.

DISTRIBUTION Known from Kenya and Uganda.

Kenya Loveridge (1957) reported it from the highlands of Kenya. Rand
(1963) listed localities in south-western Kenya between 1660 m and 3050 n.

Uganda Loveridge (1957) reported it from the highlands of eastern Uganda
(Mt Elgon). Rand (1963) listed localities between 1220 m and 3350 m.

POPULATION The species may be extremely abundant in some suitable habitats
(J. Hebrard, in litt., 1 April 1986). Abundant at Karatina (Kenya) (Lin and
Nelson, 1981). Apparently increasing in the Aberdare range in Kenya, where it
may be displacing C. jacksonii (Riva, 1979). No data are available for
other parts of its range.

HABITAT AND ECOLOGY Restricted to well watered upland habitats above about
1600 m; may be extremely abundant in forest edges, hedgerows and in ornamental
vegetation - appears to thrive in areas of dense human habitation (J. Hebrard,
in litt., 1 April 1986). Known at altitudes up to 11 000 feet (3350 m) on
the west slope of the Mt Elgon and above 3000 m in the Aberdare range (Rand,
1963; Hebrard et al., 1982). Found in the tree heath zone immediately below
the alpine zone in the Aberdare mountains (Loveridge, 1935); at Karatina
(Kenya), found principally in small localized areas of open secondary scrub
and rarely on trees over 2 m high (Lin and Nelson, 1981). Hebrard et al.
(1982) found this species inhabiting shrubs in bushland.

Females mature at approximately 9.5 months old and males about one month later
(Lin and Nelson, 1981). This viviparous species breeds throughout the year,
females producing 2 litters of young a year in the wild. The average litter
size is 13.7 and newborn young measure less than 30 mm SVL. Mean longevity

141

Chamaeleo hoehnelii

for females is 2.3 years. (Lin and Nelson, 1981). Young take insects
(Bustard, 1965). Frequently basks on trees (Lin and Nelson, 1981). Shows
particular adaptations to the low temperatures typical of high altitude
(Bustard, 1965; Hebrard et al., 1982).

THREATS TO SURVIVAL None known other than trade (see below).
INTERNATIONAL TRADE The only data on international trade are _ those
contained in the Annual Reports of the Parties to CITES which are summarized

in Tables 1 and 2. All reported trade was in live animals.

Table 1 Minimum net imports of live Chamaeleo hoehnelii reported to CITES.

1980 1981 1982 1983 1984 1985
Denmark 10 - = = = =
Germany, F.R. 2146 1455 - - - =
German D.R. 50 - = = = =
Switzerland 154 140 = = = fa
UK 25 107 - = = =
USA 3975 4640 - - - 25
Total 6360 6342 0 0 0 25

Table 2 Reported countries of origin (or exporting country if no origin
reported) and numbers of live of Chamaeleo hoehnelii reported to CITES.

_— hc —— oa ———————— eee

1980 1981 1982 1983 1984 1985

A Kw Ceeererrrr

Countries having or possibly having wild populations of C. hoehnelii
Kenya 6060 5417 = = = =

Countries without wild populations of C. hoehnelii

Germany, F.R. 2 - = = = =
Mozambique - = = = = 25
Country unknown 300 925 — = = =

eee

Trade in this species was only reported for the years 1980, 1981 and 1985.
During this period, Kenya was the largest exporter of C. hoehnelii,
accounting for 90% of all specimens reported in export, although no Kenyan
specimens were reported after 1981. The USA and F.R. Germany were the largest
importers taking 68% and 28% respectively of the total imports.

The trade volume reported to CITES in 1980 and 1981 is relatively large,
however, it seems most unlikely that this could have any significant effect on
the species in view of its frequent abundance (including around human
habitation). Trade could perhaps reduce populations in easily accessible
areas. There has been no recent trade in the species other than a single
shipment from Mozambique, where the species does not occur.

142

Chamaeleo hoehnelii

CONSERVATION MEASURES There are several protected areas where this species
May occur. These include the National Parks at Lake Nakuru, Mt Elgon and Mt
Kenya. Additional data are required on the population status of this species.

Kenya Kenya has prohibited the export of three other species of chameleon,
but these do not include C. hoehnelii.

Uganda No reptiles, other than crocodiles, are protected.

CAPTIVE BREEDING This species apparently breeds regularly in the correct
conditions and Angel (1933) and Bustard (1965) reported live births of captive

specimens. Chameleons are however difficult to maintain in captivity for a
long time and suitable environmental conditions have to be produced. De
Vosjoli (1979) described the care and conditions required for breeding. A

recent inventory (Slavens, 1985) did not record any collections of this
species, but did note that some had existed.

REFERENCES

Angel, M.F. (1933). Sur la naissance de jeunes caméléons au vivarium du
museum. Bulletin du Museé national d'Histoire naturelle 2éme_ series
5: 360-362.

Bustard, H.R. (1965). Observations on the life history and behaviour of
Chamaeleo hoehneli (Steindachner). Copeia 1965(4): 401-410.

Hebrard, J.J., Reilly, S.M., Guppy, M. (1982). Thermal ecology of Chamaeleo
hoehnelit and Mabuya varia in the Aberdare Mountains: Constraints of
heterothermy in an alpine habitat. Journal of the Bast African Natural
History Society and National Museum 176: 1-7.

Lin, J.¥. and Nelson, C.E. (1981). Comparative reproductive biology of two
sympatric tropical lizards Chamaeleo jacksonii Boulenger and Chamaeleo

hoehnelii Steindachner (Sauria: Chamaeleonidae). Amphibia-Reptilia 1
(3/4): 287-311.

Loveridge, A. (1935). Scientific results of an expedition to rainforest
regions in eastern Africa. Bulletin of the Museum of Comparative

Zoology 79(1): 1-19.

Loveridge, A. (1957). Checklist of the reptiles and amphibians of East Africa
(Uganda, Kenya, Tanganyika and Zanzibar). Bulletin of the Museum of
Comparative Zoology 117: 142-362.

Rand, A.S. (1963). Notes on the Chamaeleo bitaeniatus complex. Bulletin
of the Museum of Comparative Zoology 130: 1-29.

Riva, D. (1979). The chameleons dismal future and the need for conservation.
Journal of the Chameleon Research Center 2: 3.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

de Vosjoli, P. (1977). The care, maintenance and breeding of the African
Chamaeleo. Journal of the Chameleon Research Center 2: 7-42.

143

THREE-HORNED CHAMELEON Recommended list: 2*
[Possible problem]
Chamaeleo jacksonii Boulenger, 1896.

Order SAURIA Family CHAMAELEONIDAE

* The category has been changed from 3 to 2 since approval by the CITES TEC
Committee meeting in 1986.

SUMMARY AND CONCLUSIONS MInhabits highland regions in Kenya and Tanzania;
one of the two subspecies, C. j. merumontanus, is found only at Mt Meru in
Tanzania. Little information is available on population’. status. An
OvOViviparous species, producing around 22 young once ae_e year. The
Three-horned Chameleon probably occurs within several protected areas and is
export from Kenya is. prohibited. Apparently difficult to maintain in
captivity for any length of time, captive breeding has taken place.

Recorded trade in this species in the period 1980-85 only took place in 1980,
1981 and 1985, and amounted to c. 26 600 specimens. It appears to have been
the most heavily traded Chamaeleo species in 1980/81. At that time, 91% of
the number in trade were recorded as originating in Kenya. The only trade
since then has been from Tanzania, and may therefore be of C. j.
merumontanus . Most specimens were imported by the USA and F.R. Germany.
Kenyan legislation (enacted in 1981) appears to have effectively stopped its
exports.

It is not possible to assess the impact of previous trade on the overall
population status since detailed population data are lacking. Local
populations in accessible areas could have been affected but the species as a
whole is most unlikely to be significantly threatened by trade, providing,
firstly, that Kenya's protective legislation continues to be implemented, and

secondly, that there is no appreciable unreported trade. The subspecies
C. j. merumontanus is particularly vulnerable due _ to its restricted
distribution. As its only known range (Mt Meru) is a game reserve, it is

difficult to see how any could have been legally acquired. The recent trade
from Tanzania is therefore of greater concern.

DISTRIBUTION Inhabits highlands in Kenya and Tanzania. Two subspecies are
recognised, C. j. jacksonii being endemic to Kenya and C. j. merumontanus
to Tanzania.

Kenya C. j. jacksonii occurs in the uplands from Mt Kenya south to
Nairobi (Loveridge, 1957; Mertens, 1966; Rand, 1958). It also occurs in the
Aberdare range and around Karatina (Lin and Nelson, 1981; Riva, 1979b).

Tanzania C. j. merumontanus is only known from Mt Meru, Tanzania, where
it has been found at altitudes of 7500 - 9000 feet (Rand, 1958). Loveridge
(1957) stated that C. j. jacksonii also occurred in Tanzania, but Rand
(1958) pointed out that this was incorrect, the only specimen from Tanzania
being correctly attributable to C. j. merumontanus.

POPULATION No information is available on the overall status of this
species. Density can be very high locally, in various secondary habitats as
well as in native forest (J. Hebrard, in litt., 1 April 1986).

Kenya Abundant around Karatina, Kenya, in 1981 (Lin and Nelson, 1981).

Said to be disappearing from the Aberdare mountain range due _ to
over-collecting and a population explosion of C. hoehnelii (Riva, 1979b).

144

Chamaeleo jacksonii

Tanzania The population of C. j. merumontanus is likely to be very
limited due to its restricted range.

HABITAT AND ECOLOGY A comparatively large species of chameleon, inhabiting
highland areas up to 2000 m (de Vosjoli, 1979a). Occurs primarily in forest
and former forest, but also in coffee plantations where some original trees
have been left standing, and in stands of ornamental or exotic trees, and in
hedges in Nairobi (J. Hebrard, in litt., 1 April 1986). Around Karatina C.
jJacksonii occurred predominantly in woodlands mixed with herbs and shrubs
particularly above 2 m, and rarely in thick undergrowth. Bushes were the
principal perch site (Lin and Nelson, 1981). On Mt Meru, C. j. merumontanus
was found in bushes and in low trees (Rand, 1958).

This species is ovoviviparous (Lawrence, 1985). Females mature at around 13.5
months and males 3 months later. Copulation has been observed from March to
May, parturition occurring from mid January to March. The average litter size
is around 22 animals, the new born ranging from 25-32 mm SVL. Birth takes
place during the period of maximum insect abundance (Lin and Nelson, 1981).

THREATS TO SURVIVAL An increase in numbers of C. hoehnelii is said to be
reducing numbers of C. jacksonii in the Aberdares. Apparently heavily
exploited in the Aberdares by collectors for the pet trade (Riva, 1979b).

INTERNATIONAL TRADE The only information available on the international
trade are data contained in the Annual Reports of the Parties to CITES which
are summarized in Tables 1 and 2.

All the figures represent live specimens primarily in trade for commercial
purposes, suggesting that they were destined for the pet trade. Kenya was the
chief exporting country over the years 1980-1981, exporting 24 345 specimens,
91% of the total in trade. The majority of these chameleons were exported in
1980. The principal importing countries were the USA and F.R. Germany,
accounting for 73% and 24% respectively of all imports reported in 1980 and
1981. The Netherlands re-exported a total of 190 specimens which originated
largely from Kenya. De Vosjoli (1979b) reported that this is the most
frequently imported chameleon (to the USA).

Table 1 Minimum net imports of live Chamaeleo jacksonii reported to CITES.

1980 1981 1982 1983 1984 1985
Argentina 6 7 - - - -
Austria - 10 - - - -
Canada 174 35 - - - -
Denmark 10 - - - - -
Germany, F.R. 4681 1595 - - - 250
German D.R. 60 - - - - -
Italy - 12 - - - -
Japan - 13 - - = -
Switzerland 224 135 - - = =
UK 50 600 - - - -
USA 12491 6580 - - - =
Total 17696 8997 0 0 0 250

145

Chamaeleo jacksonii
Table 2 Reported countries of origin (or exporting country if no origin
reported) and numbers of live Chamaeleo jacksonii reported to CITES.

1980 1981 1982 1983 1984 1985

Countries having or possibly having wild populations of C. jacksonii

Kenya 17130 7215 - = = =
Tanzania - - - - - 250

Countries without wild populations of C. jacksonii

Germany, F.R. 7 - = = = =
Thailand 4 = = = = =
Country unknown 510 1735 - = = =

Relatively large numbers have been reported in trade, however, the overall
effect on the species is most unlikely to be _ significant. It is not
inconceivable that numbers of the species could be reduced in accessible
areas. No trade from Kenya has been reported since 1981. The only trade
since then has been from Tanzania, and may therefore be of C. j.
merumontanus. Since this sub-species has such a restricted distribution, it
could be particularly vulnerable to any trade. As its only known range (Mt
Meru) is a game reserve, it is difficult to see how any could have been
legally acquired.

CONSERVATION MEASURES Data are required on the population status of this
species.

Kenya The Aberdare Mountain range is a National Park. Kenya has prohibited
the export of this species without written permission from the Minister for
Environment and Natural Resources (under Legal Notice 152 enacted on 25
September 1981).

Tanzania Under the Wildlife Conservation Act, No. 12, 30 June 1974 (amended
Act No. 21, 1978), all vertebrates are protected and may not be killed,
captured, traded, imported or exported without a permit. Mount Meru is
protected as a Game Reserve.

CAPTIVE BREEDING De Vosjoli (1979b) produced guidelines on the care,
Maintenance and breeding of the African chameleons, which are apparently
difficult to maintain in captivity. This included information on feeding,
lighting, temperature, humidity and environment. C. jacksonii requires
large enclosures. Formerly maintained at the Chameleon Research Center (Riva,
1979a); this centre no longer exists. A recent inventory (Slavens, 1985)
reported six specimens in three collections. This may be an underestimate.
It is the only chameleon to have been bred over three generations (Lawrence,
1985).

REFERENCES

Lawrence, K. (1985). Lizards. In: Cooper, J.E., Hutchinson, M.F., Jackson,
O.F. and Maurice, R.J. (eds). Manual of exotic pets. British Small
Animal Veterinary Association, Cheltenham, pp. 165-172.

146

Chamaeleo jacksonii

Lin, J.¥. and Nelson, C.E. (1981). Comparative reproductive biology of two
sympatric tropical lizards Chamaeleo jacksonii Boulenger and Chamaeleo
hoehnelii Steindachner (Sauria: Chamaeleonidae). Amphibia-Reptilia
1(3/4): 287-311.

Loveridge, A. (1957). Checklist of the reptiles and amphibians of East Africa
(Uganda, Kenya, Tanganika, Zanzibar). Bulletin of the Museum of
Comparative Zoology 117: 153-362.

Mertens, R. (1966). Liste der rezenten Amphibien und Reptilen. Chamaeleonidae.
Das Tierreich 88: 1-37.

Rand, A.S. (1958). A new subspecies of Chamaeleo jacksoni Boulenger and a
key to the species of three horned chameleons. Breviora 99: 1-8.

Riva, D. (1979a). Present CRC Breeding Projects. Journal of the Chameleon
Research Center 2: 4.

Riva, D. (1979b). The chameleon's dismal future and the need for conservation.
Journal of the Chameleon Research Center 2: 3.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

de Vosjoli, P. (1979a). Preliminary observations on intraspecies communication
in the Jackson's chameleon. Journal of the Chameleon Research Center
2: 70-87.

de Vosjoli, P. (1979b). The care, maintenance and breeding of the African
Chamaeleo. Journal of the Chameleon Research Center 2: 7-38.

Welch, K.R.G. (1982). Herpetology of Africa. R.E. Krieger Publishing
Company, Malabar, 293 pp.

147

GREEN IGUANA Recommended list: 2*
[Possible problem)

Iguana iguana (Linnaeus, 1758)
Order SAURIA Family IGUANIDAE

* The category has been changed from 3 to 2 since approval by the CITES TEC
Committee meeting in 1986 owing to the increasing levels of trade in 1985.

SS

SUMMARY AND CONCLUSIONS Extremely widespread in Latin America, I. iguana
is adaptable and found in a great variety of habitats from Mexico and the West
Indies to Paraguay. It is large, up to 45 cm snout-vent length, almost
exclusively herbivorous and principally arboreal, being found particularly
along the margins of rivers. Sexual maturity is reached at an age of 2-3
years, and: broods, averaging 35-40 eggs, hatch at the beginning of the wet
season. Most population data are available for Central American and Caribbean
countries, where the species is said to be depleted in many areas, chiefly by
intense hunting for meat and eggs, and also habitat destruction. In mainland
South America hunting pressure seems to be less severe, except in Colombia,
and populations may be less affected.

Although formerly traded in small quantities for skins, the main international
trade is now primarily for live animals as pets, 109 000 being reported in
1985. The main scurce countries were El Salvador and Guatemala. There is
substantial cross-border trading in Central America of iguanas for human
consumption, but this is mostly illegal, and is unlikely to be affected by
CITES. There is a low level of international trade in skins.

The main countries currently recorded as supplying live animals to
international trade, El Salvador and Guatemala, both have depleted populations
of Iguanas. Few data are available for South America, but it seems that
populations are not under threat, except possibly in Colombia. In Central
America the major threat is from internal trade for food, which is outside the
control of CITES, but the current level of international trade in live animals
from this region seems likely to be exacerbating the problem.

DISTRIBUTION Widespread in the Neotropics from Mexico to Paraguay,
including many Caribbean Islands. On mainland South America I. iguana is
widely distributed in most lowland areas north of the Tropic of Capricorn. The
highest altitude record is from 1000 m in Colombia (Etheridge, 1982). Lazell
(1973) has been followed in not recognising any subspecies, although some
authorities consider that the form on the South American mainland is different
from that occurring in the Antilles.

Aruba Recorded from the island (Lazell, 1973).

Belize Highly localized, as much of the habitat is unsuitable (Fitch et
al., 1982).

Bolivia Widespread, including the Amazonian region (Hoogmoed, 1979)

Brazil Widespread in Amazonia (Hoogmoed, 1979) and the Pantanal, and
throughout north-east and central Brazil as far as Bahia (Cunha, 1961)

British Virgin Islands Including Tortola (Etheridge, 1982), although this

population may now be extinct, Peter Island and Virgin Gorda (UK CITES MA,
1986).

148

Iguana iguana

Colombia Widespread on the mainland, including the Amazonian’ region
(Hoogmoed, 1979); and including the islands of Gorgona, Providencia, San
Andrés and Santa Catalina (Lazell, 1973; Gallego 1978).

Costa Rica Found mostly in the lowlands in the dry, north-western part; it
is limited to gallery forests along streams (Fitch et al., 1982).

Dominica The early record from Dominica (Boulenger, 1885) is probably
erroneous (Lazell, 1973).

Ecuador Widespread from the coast (Lazell, 1973) to the Amazonian region
(Hoogmoed, 1979)

El Salvador’ Chiefly in the coastal mangrove forests (Fitch et al., 1982).
French Guiana Occurs throughout the country (Hoogmoed and Lescure, 1975).

French West Indies Including the arid, leeward coast of La Guadeloupe,
especially abundant on the Iles du Pigeon ou Goyave, and the Iles des Saintes
on La Coche, Grande Ilet, central and eastern Terre de Haut and Ilet-a-Cabrit,
but absent from Marie Galante, La Désirade and surrounding small islands
(Lazell, 1973).

Grenada Including the islands of Kick-"'em-Jenny, Mabouya Cay,
Isle-a-Caille, Isle-a-Ronde, Carriacou, but probably extinct on Glover Island
(Lazell, 1973).

Guatemala Chiefly in the Caribbean lowlands (Fitch et al., 1982).
Guyana Widespread (Hoogmoed, 1979).

Honduras Occurs on both coasts, but is most abundant in the warm, humid
Caribbean lowlands (Fitch et al., 1982); including Las Islas de la Bahia
(Utila, Roatan and Guanaja) (Etheridge, 1982), Half Moon Cay and the Swan
Islands (Lazell, 1973).

Mexico Occurring southwards from about 24° 30' on the Pacific coast and 21°
30° on the Atlantic coast, in the states of Campeche, Chiapas, Colima,
Durango, Guerrero, Jalisco, Mexico, Michoacan, Morelos, Nayarit, Oaxaca,
Puebla, Quintana Roo (including Cozumel Island), Sinaloa, Tabasco, Tamaulipas,
Veracruz and Yucatan (Smith and Smith, 1976), although it is rare in Yucatan
(Etheridge, 1982).

Montserrat Locally abundant in the South (Lazell, 1973).

Netherlands Antilles Recorded from Bonaire, Klein Bonaire, Curacao and, in
the Leeward Islands, Saba (Lazell, 1973).

Nicaragua Widely distributed, but occurs mainly on the Caribbean coast and
along major rivers (Fitch et al., 1982); also recorded from the Corn Islands
(Lazell, 1973).

Panama Including the Archipelago de Las Perlas (Etheridge, 1982).

Paraguay Confined to the Chaco (Etheridge, 1982).

Peru Widespread, including the Amazonian region (Hoogmoed, 1979). Recorded
from Iquitos, the Rio Napo or upper Maranon (Dixon and Soini, 1986).

149

Iguana iguana

Puerto Rico Mostly in the San Juan area, but there may also be a population
near Trujillo Alto; also recorded from Isla Magueyes, Cayo Icacos and
Whistling Cay (Schwartz et al., 1978).

St Lucia Found chiefly on the north-eastern coast of the main island; but
probably extinct on Pigeon Island (Lazell, 1973), and absent from the Maria
Islands (Corke, 1987).

St Vincent Throughout the lowlands and coastal cays, and including the
islands of Bequia, Quatre, Battowia, Petite Mustique, Mustique, Petit St
Vincent, Union Island, Frigate Cay, Tobago Cays, Cannouan and Savan (Lazell,
1973).

Suriname Occurs throughout the country in a wide variety of habitats, from
sea level to at least 500 m (Hoogmoed, 1973).

Trinidad and Tobago Occurs on both islands (Lazell, 1973).

US Virgin Islands Including St Croix, but absent from the small cays of the
St Croix bank (Lazell, 1973), occurring on St Thomas, St John, Water Island,
Patricia Cay and Hassel Island (Etheridge, 1982).

Venezuela Found in most mainland regions (Rivero-Blanco and Dixon, 1979);
and including the islands of Margarita, Los Testigos, Los Frailes, Los
Hermanos, La Blanquilla, La Tortuga, Orchilla, Los Roques, Isla Aves (Lazell,
1973). Widespread in lowland habitats, from primary forest to arid areas, and
up to the splash zone along rocky coasts, even adapting to live in parks and
gardens (S. Gorzula, in litt., 11 April 1986).

POPULATION Most population data are available for Central American and
Caribbean countries, where numbers are said to be declining in many areas.
There are few data for mainland South America, but there are no suggestions
that the populations are declining, except in Colombia.

Aruba No information.
Bolivia No information.

Brazil Said to be extremely common around all major waterways (W.E.
Magnusson, pers. comm.).

British Virgin Islands The population on Tortola is said to be greatly
reduced or extinct; Peter Island has a healthy but small population; and
Virgin Gorda has a small population in the North Sound area at Bird Creek (UK
CITES MA, 1986).

Colombia Populations of I. iguana are thought to be declining in Colombia
as a result of over-hunting (Harris, 1982). The populations in the islands of
San Andrés and Providencia are said to “much persecuted" (Gallego, 1978). On
Isla Gorgona, iguanas are scarce, but can still be seen without much
difficulty, although they are very abundant on Isla Gorgonilla (Medem, 1979).

Costa Rica As in Nicaragua, populations are thought to be scarce or
declining rapidly except in areas too remote from settlements to be hunted
regularly (Fitch et al., 1982).

Ecuador No information.

El Salvador The populations of JI. iguana which once thrived in the
coastal mangrove forests are reported to have almost disappeared as the

150

Iguana iguana

forests have been destroyed. Although iguanas were still relatively common in
1974, the forests have now been reduced to 1% of their original area, and,
within remaining stands, iguana populations may be as low as 1% of their
Original density. Iguanas are reported to be still surviving in the area
around Volcan San Miguel, although they are probably mostly Ctenosaura
similis (Fitch et al., 1982).

French Guiana No information.

French West Indies On the Iles des Pigeons, Guadeloupe, it was especially
abundant; on Saba it was everywhere common, and on St Croix it was locally
abundant in the East End District, while the Iles des Saintes were said to be
infested with iguanas (Lazell, 1973).

Grenada I. iguana is said to be rare (Ministry of Agriculture and
Tourism, in litt., 15 November 1985).

Guatemala Both hunting and habitat destruction are reported to have
contributed to a “drastic reduction" of iguana populations (Fitch et al.,
1982). Populations in Alta Verapaz are reported to have declined as a result
of over-hunting (Flores Villela, 1980), but Fitch et al. (1982) assert that
they have been little affected in Alta Verapaz and Peten, on the Caribbean
coast. They report that the worst affected areas are on the Pacific coast
where the coastal mangrove forests have been reduced to 7% of their original
extent by agricultural development. The increasing use of pesticides has also
been implicated.

Guyana No information.

Honduras Most of the country, particularly the North, is sparsely populated
and iguana populations are not in danger of extermination or even of drastic
reduction, however around Choluteca in the Pacific lowlands the population may
have been reduced by as much as 90% (Fitch et al., 1982).

Mexico I. iguana is reported to have declined in numbers as a result of
hunting, and is generally considered to be endangered in Mexico (G. Ceballos,
in litt., 2 December 1986). In the coastal mangrove forests of Chiapas,
most of the animals captured are adolescents and juveniles, and the population
is estimated to be as low as 5% of its original size (Fitch et al., 1982).
Northwards from Chiapas, up the Pacific coast to Sinaloa, I. iguana is still
relatively common and substantial populations have been reported in Jalisco
and Colima (G. Ceballos, in litt., 2 December 1986). On the gulf coast and
in parts of the Yucatan Peninsula there were reported to be substantial
populations (Fitch et al., 1982). Huerta and Mondragon (1975) reported that
iguanas had declined enormously in Tabasco, particularly around Villahermosa
and Teapa.

Montserrat No information.

Netherlands Antilles I. iguana is said to have declined in recent years
on Curacao (Bakhuis, 1983).

Nicaragua Drastic reductions in numbers were reported in many areas,
particularly towards the South of the Pacific coastal strip, although there it
was probably Ctenosaura similis which was worst affected. In many of the
country's sparsely populated areas, iguanas were thought to be somewhere near
their original abundance (Fitch et al., 1982).

Panama Evidence from the volumes of iguanas sold in markets suggests that
their populations declined markedly between 1950 and 1969 (Fitch et al.,

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Iguana iguana

1982). Werner (1986) reported that they had disappeared from several areas
Owing primarily to over-hunting.

Paraguay No information.

Peru Said to have become extremely rare around Iquitos, supposedly because
of over-hunting (Dixon and Soini, 1986).

Puerto Rico No information.

St Lucia Said to be “extremely rare in St Lucia” although “little is known
about their status in the wild” (Corke, 1983).

St Vincent ‘No information.

Suriname I. iguana is reported to common and is not considered to be
endangered (Kalden, C.J., in litt., 16 January 1986).

Trinidad and Tobago At least locally common in 1973 (Lazell, 1973).
US Virgin Islands No infromation.

Venezuela Said to be common in spite of the hunting pressure which may be
severe in the north of the country (S. Gorzula, in litt., 11 April 1986).
In April, after the young have hatched, the Llanos are reported to be
inundated with iguanas, which even spread into areas of human habitation (Ramo
and Ayarzaguena, 1983).

HABITAT AND ECOLOGY A large, herbivorous lizard, Iguana iguana is
sexually dimorphic, the maximum size in males being about 3.8 kg, 45 cm
snout-vent length (SVL) and 1.9 kg, 34 cm in females (Dugan, 1982). It occurs
in a very wide variety of habitats, usually favouring river banks and the
Margins of ponds and lakes, but also found in arid or rocky areas with
xeromcorphic vegetation. Ceballos (in litt., 2 December 1986) noted that
sparse populations occurred in sugar cane crops and along water ditches in
other cultivated areas. It is described as "primarily arboreal and strongly
heliothermic". Adults spend most of their time perched in trees, but younger
animals often bask on the ground during the day, returning to the vegetation
only at night. It is reported to be relatively sedentary, some animals never
moving far from the site of hatching (Dugan, 1982; Van Devender, 1982).

Juveniles hatch at a length of 72-79 mm SVL (Harris, 1982). Van Devender's
(1982) studies of growth rate indicate a linear increase in SVL with age for
the first two years of life, reaching 180 mm after one year and 290 mm after
two. Sexual maturity in females is achieved after two or three years when they
attain a length of over 200 mm (Wiewandt, 1982). Eggs are laid in burrows,
usually in sandy soil, the clutch size being reported to vary from 14 to 76,
with the mean between 35 and 40. Egg weights vary from 9 g to 14 g (Wiewandt,
1982). Incubation has been found to last for 74 to 81 days at 31°C (Harris,
1982). Mating is relatively synchronous throughout the population, and occurs
in the dry season, hatching taking place at the beginning of the wet season.
This pattern has been reported to be common to populations throughout most of
the geographic range, and is thought to ensure not only that a flush of new
vegetation is available for the hatchlings but also that the soil temperatures
at the end of the dry season are high enough for incubation (Rand and Greene,
1982). Some nests are communal, up to eight females sharing the same deep,
branching burrow. This allows for the deeper burial of the eggs, which are
often extremely difficult to locate as a result (Rand and Dugan, 1984).
Repeated nesting at the same site may help to maintain loose soil structure
and keep the vegetation clear. Females have been reported to guard nests after

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Iguana iguana

laying has been completed (Wiewandt, 1982). Males defend mating territories
and females tend to select the larger males (Dugan, 1982). They may copulate
with more than one male (Dugan and Wiewandt, 1982). After reaching maturity,
females may breed annually to an age of ten years (Rand and Greene, 1982). The
oldest recorded Iguana iguana reached 12 years, 5 months in captivity in
Florida (Bowler, 1977).

Survival of hatchlings is very low, 75% dying in the first three months and
50% dying in each of the next three 3-month periods, with not less than 2.6%
of the original hatchlings surviving at the end of one year (Harris, 1982).
Van Devender (1982) provides a similar mortality model.

Adult I. iguana are exclusively herbivorous, feeding primarily on leaves of
trees, but-fruits and other vegetation are also taken. It has been reported
that there is an ontogenetic shift in diet, the juveniles being carnivorous,
but this is now thought to be incorrect, even the juveniles being herbivorous
(Hoogmoed, 1973; Van Devender, 1982). Instead, young animals have been shown
to satisfy their higher requirements for protein and energy compared to adults
by selecting better quality vegetation (Troyer, 1984a). Lazell (1973) observed
I. iguana eating birds eggs. Food undergoes microbial fermentation in the
specially enlarged hind-gut, 30-40% of the total metabolisable energy being
obtained from this source (McBee and McBee, 1982). The overall digestive
efficiency (54% for fibre and 86% for protein) is comparable with values found
in ruminants (Troyer, 1984b).

The chief predator of JI. iguana is undoubtedly man, but Basiliscus
basiliscus is also locally important. Other predators include the reptiles
Boa constrictor, Crocodylus acutus, Ctenosaura similis and Trimorphodon
biscutatus, the birds Sarcorhamphus papa, Elanoides forficatus, Quiscalus
spp., and Philander opossum, Felis pardalis and other large cats and
domestic dogs amongst the mammals (Hoogmoed, 1973; Van Devender, 1982; Dugan
et al., 1981; G. Ceballos, in litt., 2 December 1986). When disturbed,
iguanas may dive from overhanging branches into the water, where they may swim
submerged for considerable distances. They can defend themselves with biting
and lashing of the tail when cornered (Hoogmoed, 1973).

THREATS TO SURVIVAL The main use of I. iguana since prehistoric times has
been for food, both its flesh and its eggs being widely esteemed. Iguanas are
also reputed to have medicinal values. In Mexico they are thought to be
effective against renal disorders (Flores Villela, 1980), and aphrodisiac
properties have been attributed to the eggs (Jacobs, 1984) and flesh (Fitch
et al., 1982). The eggs are usually preferred to the flesh, and indeed in
some areas, such as northern Colombia, the flesh is not consumed at all. Where
they occur sympatrically, the flesh of Ctenosaura similis (Fitch et al.,
1982) or Iguana delicatissima (Lazell, 1973) is usually preferred to that of
I. iguana. In Mexico, I. iguana is usually preferred to Ctenosaura spp.
(G. Ceballos, in litt., 2 December 1986). As well as subsistence hunting,
iguanas are often available in markets and restaurants throughout Central and
South America. They are usually sold live, the legs being tied across the top
of the back; the mouth may also be sewn shut, in which condition they can be
kept alive for many days (Fitch et al., 1982). In recent years a lucrative
pet trade in live animals has grown up, and they may also be dried or stuffed
and sold as curios. The skins are tanned for the speciality leather trade, and
there is a small demand for iguanas as laboratory animals, for both anatomical
and physiological studies (Lazell, 1973, Flores Villela, 1980).

The usual methods of hunting are by shooting with a small-calibre rifle or a
sling-shot. Snaring is also used, and occasionally hunters have a noose on the
end of a long pole to capture iguanas in trees. Dogs are often used to run

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Iguana iguana

down iguanas flushed in the open, to locate iguanas in burrows or to retrieve
wounded animals (Fitch et al., 1982; Flores Villela, 1980). Eggs are
sometimes dug from the nests but, in a common and misguided attempt at
sustainable utilization, the eggs are often excised from living females which
are later stuffed with ash or dead leaves, sewn up and released, in the belief
that they will survive to breed again. In other cases the cosmetic surgery may
be dispensed with (Harris, 1982; Flores Villela, 1980).

Belize Trade in I. iguana appears to have declined in Belize. In four
weeks in 1971, 300 were sold in the market in Belize City; none were sold
during a week of observation in 1976; and only about 24 were sold over a
3-week period in 1977. Only gravid females were observed on the last occasion,
the eggs being more highly esteemed than the flesh. The majority of the
I. iguana sold in Belize City in 1971 had been obtained in the Cayo
District, near the Guatemalan Border, and' further animals were noted in
markets in Cayo and Stan Creek (Fitch et al., 1982).

Brazil I. iguana is one of the species hunted frequently by the Coboclos
living along the banks of the Amazon and its tributaries. It is much prized
for its flesh and also hunted for its skin (Cunha, 1961). Several tribes of
Indians, including the Caraja and the Waiwai, are known to have hunted iguanas
regularly (Fitch et al., 1982).

British Virgin Islands As far as is known, deliberate killing of iguanas is
insignificant, the main threats coming from human disturbance of the habitat
and exotic predators, including cats, dogs and mongooses (Budd, S.D., In
litt., 7 January 1986).

Colombia Over-exploitation of JI. iguana in Colombia is giving concern.
Eggs are the main product, and are usually sold boiled, when they fetch double
the price of hens eggs. The meat is considered a low class food but Indians
and other poor Colombians may depend heavily on it for protein (Harris, 1982).
In the islands of Providencia and San Andrés the species is much persecuted
for its meat and eggs. Live animals, captured in Providencia, are sold in the
market in San Andrés (Gallego, 1978).

Costa Rica There is thought to be no commercial hunting in Costa Rica, but
local hunting is heavy in some areas. In 1963, 40 gravid females were konwn to
have been caught by locals along four miles of beach at Tortugero, on the
north-east coast, and several nests were known to have been destroyed by dogs
(Fitch et al., 1982).

El Salvador Iguanas are absent from most of their former range in El
Salvador but are still hunted in appreciable numbers in the area around San
Miguel, although there they are thought to be mostly Ctenosaura similis
rather than I. iguana. Even where they are scarce and commercial hunting is
unprofitable, subsistence hunting-pressure is still intense owing to the
scarcity of protein. Retail prices of I. iguana in markets in Santa Ana, San
Miguel and Le Union in 1979 ranged from US$1.40 for animals without eggs to
US$2.40 for females bearing eggs. Prices in the markets in San Salvador were
up to US$5.20 each, representing a 3- to 8-fold increase since 1976. El
Salvador is a major importer of iguanas, most carried illegally on foot across
the borders from Honduras and Nicaragua. The small town of Santa Rosa de Lima,
near the Honduran border is of central importance, hundreds of iguanas passing
through the market for redistribution to other towns. In 1976, large numbers
were being imported from Nicaragua by boat across the Gulf of Fonseca, but by

1979 there were few or none, as numbers had declined and export had been made
illegal (Fitch et al., 1982).

Iguana iguana

French West Indies On the Iles des Saintes the people are reported to
Slaughter iguanas in large numbers. Some are stuffed and sold as curios
(Lazell, 1973).

Grenada Iguanas are considered a delicacy by the Grenadians, but the
numbers hunted are not known (Ministry of Agriculture and Tourism, Grenada,
in litt., 15 November 1985).

Guatemala Over-hunting of iguanas has been reported in Guatemala (Flores
Villela, 1980), and the numbers sold are reported to have declined with the
falling population of wild animals. The reduction in the supply and the rising
prices have apparently caused some consumers to abandon iguanas as a food
source. Adult I. iguana were being sold in 1979 in the market at Mazatenango
at US$3 each, and juveniles at US$1.50. Some are traded across the border to
El Salvador (Fitch et al., 1982).

Honduras Hunting and habitat destruction are thought to be a threat only in
the Choluteca region where iguana populations may be only 10% of their former
levels. In other more sparsely populated regions hunting pressure for home
consumption is slight, and restricted transport prevents the ready export of
iguanas, although illegal exports undoubtedly occur along the border to El
Salvador. The price of I. iguana in the market in Tegucigalpa in 1979 was
reported to be US$1.25-3. Unrestricted use of pesticides is suspected to have
had a part in the reductions in some areas (Fitch et al., 1982). I. iguana
is considered a game species in Honduras (Fuller and Swift, 1984).

Mexico JI. iguana is very popular as food in Mexico and is much preferred
to Ctenosaura spp., particularly in Jalisco and Colima (G. Ceballos, in
dJitt., 2 December 1986). The main areas where iguanas are hunted are
Chiapas, Veracruz, Michoacan, Oaxaca, Guerrero, Tehuantepec (Flores Villela,
1980), the Yucatan Peninsula (Fitch et al., 1982) and Tabasco (Huerta and
Mondragon, 1975). They are frequently found in markets in Guerrero (Flores
Villela, 1980) and Tabasco, and are extensively hunted in Chiapas, where they
are now much reduced in numbers. In some markets in southern Mexico vendors
were reported to be selling small helpings of iguana stew at US$0.45 to
US$1.35, the inflated price reflecting its supposed medicinal properties.
There is substantial evidence that the aboriginal peoples of Mexico hunted

iguanas (Fitch et al., 1982). The main hunting season is in March-April
when gravid females are selectively caught because their flavour is
preferred. Occasionally the female is discarded after the eggs have been

removed. Some iguanas are also killed for their skins; males are selected for
this purpose because the skin needs to be larger than 30 x 40 cm (Huerta and
Mondragon, 1975). The skin trade has flourished recently for the manufacture
of belts and boots, the latter fetching up to US$100 a pair (G. Ceballos, in
Jitt., 2 December 1986).

Netherlands Antilles Iguanas are a popular food item in the Netherlands
Antilles and a survey conducted on Curacao revealed that 60% of the population
eat iguanas and 32% of the male population hunt iguanas, mainly for their own
consumption. This is related to the fact that most people like their iguanas
either live or recently dead, as the flesh deteriorates fast (Bakhuis, 1983).
The price of iguanas was approximately AG10 (AG1 = US$0.56) in 1983 (Bakhuis,
pers. comm.).

Nicaragua Large-scale exploitation has been reported to be threatening both
species of iguana with extinction. In 1968 it was estimated that 150 000 were
eaten annually within the country and there was reported to be a mass export
to El Salvador, where they had already been depleted. Hunters in some areas
were reported to have stopped hunting by 1976 as the animals were so scarce.
Only in the sparsely populated, inaccessible areas was the hunting pressure

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Iguana iguana

low. There was a well organised distribution system in 1976, with a group of
professional hunters near the Honduran border supplying iguanas to the rest of
the country. Other sources were in San Francisco de Carniceria, on Lago de
Managua, and San Carlos, on Lago de Nicaragua. Dealers collect iguanas from
the hunters in weekly rounds, and take them to the city markets. Prices were
highest, and iguanas scarcest, in the South-east (Fitch et al., 1982).

Panama In 1950 iguanas were reported to have been sold in large numbers in
the market in Panama City, but very few were available by 1969. Sale of
iguanas was made illegal in 1967, and the ban was apparently well enforced,
but it was lifted again in 1979 because of a beef shortage (Fitch et al.,
1982).

Peru Over-hunting for skins and the pet trade is thought to have reduced
iguana populations around Iquitos. A number’ of skins and a few live animals
are exported from Iquitos each year. Iguanas are not though to be eaten in
the region (Dixon and Soini, 1986).

Suriname I. iguana is not thought to be hunted in Suriname, but skins
reported to have originated there have probably been smuggled over the border
from Brazil (M.S. Hoogmoed, in litt. 26 August 1986).

Venezuela Hunting of iguanas is now illegal apart for sport hunting
(Venezuela CITES MA, 1987), although they were formerly much prized by some
indigenous peoples (Fitch et al., 1982). Illegal hunting still occurs in
the north of the country and the lizards are often sold along the road from
Puerto La Cruz to Caracas and in Estado Sucre. Such sales are illegal and the
meat is mever seen in markets nor served in restaurants. There is less
hunting in the south of the country, although females are often caught to
extract their eggs, which are locally popular (S. Gorzula, in Jlitt., 11
April 1984). There are plans to develop a commercial management plan for this
species (Venezuela CITES MA, 1987).

INTERNATIONAL TRADE Apart from the illegal trade in iguanas for food in
Central America, mentioned in the previous section, the only other data on
international trade are those contained in the Annual Reports of Parties to
CITES. Virtually all the transactions recorded were in skins or live animals,
and only these have been included in the following analysis. The CITES reports
are summarised in Tables 1 and 2.

The tables show that the majority of the reported trade is in live animals,
the chief importers being the USA and F.R. Germany (Table 1). This implies
that they are used primarily as pets rather than food items. The trade
reached a peak of over 109 000 in 1985. The major suppliers of live animals
were Colombia, El Salvador, Suriname, Guatemala, Guyana and Mexico. Of these
Colombia has exported few recently, while El Salvador, Guatemala and Guyana
have increased their exports (Table 2). A pricelist, published by an animal
supplier in the USA (South American Unlimited, New York) in 1985, gave the
price of live Green Iguanas from Suriname as US$12 for 1 ft (30 cm) animals to
US$35 for 3-4 ft ones.

Significant quantities of skins were traded in 1980 and 1981, but the volume
has since declined sharply, and none was reported in 1984. The skins
originated mostly in Colombia, and this may be associated with the fact that
Colombia implemented a new plan to control the export of wildlife in 1978,
under which only two companies were allowed to export their remaining stocks
of hides. The skins appeared to follow a circuitous route, passing first
through the USA then to the UK, from where they were re-exported mainly to
F.R. Germany, Canada, Spain, Egypt and even back to the USA. The 16 633 skins

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Iguana iguana

reported from unknown countries of origin in 1980 in Table 2 were mostly
re-exported by the UK, and it is reasonable to assume that the bulk of these
were skins from Colombia, of which the UK imported 20 000 in 1980. This
suggests that the international trade routes are not well established and that
there may be little regular trade in iguana skins. The only other major source
of skins was Paraguay, which produced 5000 in 1982.

Of the countries with no wild populations of I. iguana from which exports
were reported (Table 2), it seems likely that the majority of the exports were
in reality re-exports, the country of origin not having been specified.
Exceptions are probably Indonesia and Singapore, from which significant
exports of skins were reported in 1980 and 1981; it is possible that the skins
were wrongly identified, and that they may have been Varanus spp. Similarly
the skins ‘exported from Argentina may have been of Tupinambis spp. The
export from Ghana may result from a confusion with Guyana, or refer to
Varanus sp.

Table 1. Minimum net commercial imports of live I. iguana (L) and their
skins (S) reported to CITES.

1980 1981 1982 1983 1984 1985
Argentina 6L 10 L - - - 228 L
Austria - 114 L 80 L 16 L 122 L 40s.
Belgium - - - - - 101 L
Canada 4955 $ 7185 $ - 1125 Ss - 15s
404 L 442 L LI3 0 929 L 1639 L 2624 L
China 4 L - - - - -
Costa Rica - - 5 kg S - - -
Denmark 24 L 16 L TS 86 L 527 L 292 L
Dominica - = - - - 40 L
Egypt 3105 $s - - - - -
Finland - - 3b - - -
France 4L 3216S 13 Ss 20 2L 2L
German D.R. 180 L - - - 4L -
Germany, F.R. 9896 L 2489 L 1792 L SS 20 6403 L 10900 L
Guyana - - - 50 L - -
Israel 60 §S 10 L 355 - Sls 1L
Italy 19 L - - 166 L - 620 L
Japan 108 L 42 L 75 G 496 L 736 L 1022 L
Malaysia 2L - - - - -
Netherlands 2L 102 L 589 L 362 L 481 L 559 L
New Zealand - - - - - 6L
Singapore - - 10 L - - 10 L
S. Africa - 6s - - - 6L
S. Korea - - - - 5 L -
Spain - 599 Ss 2L 27 L 10 L 20 L
Sweden - - - - 6L 4L
Switzerland - 23 Ss - - - 750 §
180 L 304 L 210 L 288 L 326 L -
UK 3367 S - - - - -
498 L 922 L 337 L 988 L 1104 L 733 L
USA 10016 $ 26485 S 5243 Ss - - -
51387 L 51562 L 37120 L 37750 L 34303 L 92450 L
TOTAL 21503 §S 34614 §$ 5256 S + 1125 s - 791 S
62714 L 56013 L 40349 L 42712 L 45723 L 109786 L

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Iguana iguana

Table 2 Reported countries of origin or export for commercial exports of live
I. iguana (L) and their skins (S) reported to CITES.

1980 1981 1982 1983 1984 1985

oe ee ——————————e

Countries with wild populations of I. iguana

Colombia 20500 S$ 8978 S 2s 1125 $s - 750 S
3301 L 34108 L 320 L - - -
Costa Rica - - 121 Ss - - -
Ecuador - - - - - 222 L
El Salvador: 54 L 8140 L 31146 L 23668 L 13737 L 24152 L
Fr. Guiana - 60L 630 L 50 L - -
Guatemala 61584 L 3781 L 25 L 4400 L 19850 L 72181 L
Guyana 56 L 41 L 839 L 3952 L 7591 L 7180 L
Honduras 75 L 85 L 120 Ss - - 222 L
Mexico 38 - - - - -
76 L 10 L 3487 L 891 L 225 L -
Neth. Antilles - - - 43 L - -
Nicaragua - - 5 kg S - - -
Panama 1200 - - 150 L 50 L 2L
Paraguay - - 5000 S - - -
Peru ~ 20 L 100 L 330 L 285 L 820 L
Suriname 300 L 10212 L 5847 L 9047 L 3960 L 4605 L
Venezuela - - - - 10 L -
Countries without wild populations of I. iguana
Argentina 1000 Ss 5294 S - - - -
- 14838 L - - - -
Australia - - - - - 52 L
Chile - - - - 80 L -
Germany, F.R. - - 11 L - - =
Ghana 14 L - - - - -
Guinea - - - - 20 L -
Haiti - 23 L - - - =
Indonesia 1959 S$ 1000 S$ - - - -
Netherlands 10 L - - - = =
Singapore 1950 S 1875 Ss - - - -
Tanzania - - - - - 350 L
USA * Ih AG 1L 12 L 283 L 74 L 274 L
Uruguay - - = 9 ay = =
Unknown 16633 S 9056 S$ 13 Ss - - 41 Ss
38 L 6988 L 45 L 31 L 70 L 2 L

* It is not clear whether the US report includes Puerto Rico or the US Virgin
Islands, in which I. iguana does occur.

CONSERVATION MEASURES The legislation controlling the hunting, trade and
export of I. iguana is summarised in Table 3. Owing to the wide distribution
of this species it can be found in the majority of National Parks within its

range. In these it will generally receive greater protection than in the rest
of the country.

There has been one attempt at the re-introduction of I. iguana to a National
Park in Panama (see Captive Breeding Section).

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Iguana iguana

Table 3. Legal prohibition on the hunting, internal trade and commercial
export of I. iguana. Dates are those on which the legislation came into

force. A - All live animals & parts; L - Live animals only; S - Skins;
P - Allowed under permit; C - Closed seasons may be imposed; * - these
territories are Overseas Départements of France with which the EEC may trade
without the imposition of CITES controls; ? - no information (mostly from

Fuller et al., 1987).

CITES Hunting Trade Export
Aruba - - ? 4
Belize : 1981 - A 1981 A 1981
Bolivia 1979 - s 1979 A 1985
Brazil 1975 - A 1967 A 1967
Br. Virgin Is 1976 ? ? P
Colombia 1981 P - L 1978
Costa Rica 1975 A 1985 A 1970 A 1970
Dominica 7 ? ? ?
Ecuador 1975 - - A 1983
El Salvador 1987 - ~ -
Fr. Guiana 1978 * - - P
Fr. W. Indies 1978 * P
Grenada - - - -
Guatemala 1980 - - P
Guyana 1977 - - A 1986 +
Honduras 1985 Cc P P
Mexico - Cc - A 1982
Neth. Antilles ~ - ? 2
Nicaragua 1977 (o; P P
Panama 1978 A 1980 A 1980 A 1980
Paraguay 1977 A 1975 A 1975 A 1975
Peru 1975 Cc A 1973 A 1973
St Lucia 1983 A 1980 A 1980 A 1980
St Vincent - ? ? ?
Suriname 1981 c - P
Trinidad & Tobago 1984 ? fe P
US Virgin Is 1975 2 ? P
Venezuela 1978 Cc A 1970 A 1970

+ Export quotas of 8400 live I. iguana were suggested for 1987 and 1988.

CAPTIVE BREEDING Because of their widespread use as a food item, their
relatively high fecundity and their herbivorous habit, I. iguana has been
Suggested as a suitable candidate for commercial captive breeding.
Experimental projects have been initiated in at least two countries.

In Curacao at the Government research institute of Carmabi, Pescadera Bay,
iguanas were initially kept in small cages measuring 2 x 3 x 5 m. Later an
open enclosure of 10 x 10m was constructed into which over 50 iguanas were
introduced. Shelters comprising tiers of wooden drawers, each having an
entrance hole small enough to exclude cats, were used by the animals at night.
They were fed on leaves and fruits and a breeding success of 100% was
reported. Further trials were under way to see whether they could be reared
economically (Bakhuis, 1983).

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Iguana iguana

The Smithsonian Tropical Research Institute in the Republic of Panama started
an Iguana Management Project in 1983 under Dr Dagmar Werner, in collaboration
with the Directorate of Renewable Natural Resources, the Institute for
Agricultural Research and the University of Panama. The project aims to
repopulate areas which have been depleted of iguanas, using captive-raised
stock. Artificial incubation is claimed to raise survival to one year from 5%
to 95% (Werner, 1986). Initially 400 wild-caught hatchlings were divided into
groups of 10 to 20 and introduced into small, fenced enclosures, measuring
about 12 square yards (10 m?) containing bamboo shelters, and were fed on
fruit and natural vegetation. Later it was found that up to 60 iguanas could
be kept in each enclosure. In breeding trials adult females were captured from
the wild and induced to lay eggs in containers buried in artificial clearings
(Werner and Miller, 1984). Eggs were collected and incubated artificially to
produce over 700 hatchlings. Some of the hatchlings were used to carry out a

small repopulation experiment in central Panama in 1984 (Jacobs, 1984). A
further 1200 were released in December 1985 around farms, and the initial
survival appeared to be high. The possibility of reforesting areas with

fodder trees in an attempt to regenerate natural habitat has been suggested
(Werner, 1986).

Iguanas are regularly kept as pets in several countries, and may breed in
captivity. CITES Annual Reports contain records of 6 captive-bred specimens in
1982 and 2 in 1983. It is apparent that the pet trade in I. iguana depends
almost entirely on wild-caught animals.

REFERENCES

Bakhuis, W.L. (1983). Exploitation of the Green Iguana in the Netherlands
Antilles. Trade Wind 5(5): 66-68.

Boulenger, G.A. (1885). Catalogue of the lizards in the British Museum. Vol.
2, ed. 2, 492 pp.

Bowler, J.A. (1977). Longevity of reptiles and amphibians in North American
collections. SSAR and Phil. Herp. Soc. Misc. Pub. Herp. Cir. (6): 1-32.
Corke, D. (1983). The reptiles of the Maria Islands, St Lucia. Unpublished

report to the Eastern Caribbean Natural Area Management Programme.

Corke, D. (1987). Reptile conservation on the Maria Islands (St Lucia, West
Indies). Biological Conservation 40: 263-279.

Cunha, O.R. da (1961). II Lacertilios da Amazonia. Os Lagartos da Amazonia
Brasiliera com especial referencia aos representados na colecao do Museu
Goeldi. Boletim do Museu Paraense Emilio Goeldi, Nova Serie, Zoologia
(39): 1-189.

Dixon, J.R. and Soini, P. (1986). The reptiles of the Upper Amazon Basin,
Iquitos Region, Peru. Part 1 Lizards and Amphisbaenians. Part 2
Crocodilians, Turtles and Snakes. Milwaukee Public Museum, 154 pp.

Dugan, B. (1982). The mating behaviour of the Green Iguana Iguana iguana.
In: Burghardt, G.M. and Rand, A.S. (eds) Iguanas of the world, their
behaviour, ecology and conservation. Noyes Publications, New Jersey, pp.

320-341.

Dugan, B.A., Rand, A.S., Burghardt, G.M. and Bock, B.C. (1981). Interactions
between nesting crocodiles and iguanas. Journal of Herpetology 15:
409-415.

Dugan, B. and Wiewandt, T.A. (1982). Socio-ecological determinants of mating
strategies in iguanine lizards. In: Burghardt, G.M. and Rand, A.S. (eds)
Iguanas of the world, their behaviour, ecology and conservation. Noyes
Publications, New Jersey, pp. 303-319.

Etheridge, R.E. (1982). Checklist of iguanine and malagasy lizards. In:
Burghardt, G.M. and Rand, A.S. (eds) Iguanas of the world, their

behaviour, ecology and conservation. Noyes Publications, New Jersey, pp.
7-37.

160

Iguana iguana

Fitch, H.S., Henderson, R.W. and Hillis, D.M. (1982). Exploitation of iguanas
in Central America. In: Burghardt, G.M. and Rand, A.S. (eds) Iguanas of
the world, their behaviour, ecology and conservation. Noyes Publications,
New Jersey, pp. 397-417.

Flores Villela, 0O.A. (1980). Reptiles de importancia economica en Mexico.
Thesis, Universidad Nacional Autonoma de Mexico, 278 pp.

Fuller, K.S., Swift, B., Jorgenson, A. and Brautigam, A. (1987). Latin
American wildlife trade laws. Second Edition (Revised). World Wildlife
Fund-US, 418 pp.

Gallego, H.C. (1978). Informe de la Comision Efectuada a San Andrés y
Providencia: fauna silvestre. Unpublished report, 25 pp.

Harris, D.M. (1982). The phenology, growth and survival of the Green Iguana
Iguana iguana in northern Colombia. In: Burghardt, G.M. and Rand, A.S.
(eds) Iguanas of the world, their behaviour, ecology and conservation.
Noyes Publications, New Jersey, pp. 150-161.

Hoogmoed, M.S. (1973). Notes on the herpetofauna of Surinam IV. The lizards
and amphisbaenians of Surinam. Biogeographica 4: 1-419.

Hoogmoed, M.S. (1979). The herpetofauna of the Guianan region. In: Duellman,
W.E. (ed.) The South American herpetofauna: its origin, evolution, and
dispersal. Museum of Natural History, University of Kansas, Monograph
(7): 241-280.

Hoogmoed, M.S. and Lescure, J. (1975). An annotated checklist of the lizards
of French Guiana, mainly based on two recent collections. Zoologische
Mededelingen 49: 141-172.

Huerta Mercado, P. and Mondragon, J. (1975). Generalidades sobre la Iguana en
el Estado de Tabasco. Bosques y Fauna 12(3): 16-21.

Jacobs, M. (1984). Scientists make breakthrough in effort to cultivate iguanas
as food source. Smithsonian Institution Research reports (43): 1-8.

Lazell, J.D. (1973). The lizard genus Iguana in the Lesser Antilles.
Bulletin of the Museum of Comparative Zoology 145(1): 1-28.

McBee, R.H. and McBee, V.H. (1982). The hindgut fermentation in the Green
Iguana Iguana iguana. In: Burghardt, G.M. and Rand, A.S. (eds) Iguanas
of the world, their behaviour, ecology and conservation. Noyes
Publications, New Jersey, pp. 77-83.

Medem, F. (1979). Los anfibios y reptiles de las Islas Gorgona y Gorgonilla.
In: Prahl, H. von, Guhl, F. and Grogl, M. (eds). Gorgona. Universidad de
los Andes, Colombia, pp. 189-217.

Ramo, C. and Ayarzaguena, J. (1983). Fauna Jllanera, apuntes sobre su
morfologia y ecologia. Cuadernos Lagoven, Venezuela, 84 pp.

Rand, A.S. and Dugan, B. (1983). Structure of complex iguana [Iguana iguana]
nests. Copeia 1983(3): 705-711.

Rand, A.S. and Greene, H.W. (1982). Latitude and climate in the phenology of
reproduction in the Green Iguana Iguana iguana. In: Burghardt, G.M. and
Rand, A.S. (eds) Iguanas of the world, their behaviour, ecology and
conservation. Noyes Publications, New Jersey, pp. 142-149.

Rivero-Blanco, C. and Dixon, J.R. (1979). Origin and distribution of the
herpetofauna of the dry lowland regions of northern South America. In:
Duellman, W.E. (ed.) The South American herpetofauna: its origin,
evolution, and dispersal. Museum of Natural History, University of
Kansas, Monograph (7): 281-298.

Schwartz, A., Thomas, R and Ober, L.D. (1978). First supplement to a
check-list of West Indian amphibians and reptiles. Carnegie Museum of
Natural History Special Publication 5: 1-35.

Smith, H.M. and Smith, R.B. (1976). Synopsis of the herpetofauna of Mexico
Vol. III. Source analysis and index for Mexican reptiles. John Johnson,
Vermont, 23 pp.

Troyer, K. (1984a). Diet selection and digestion in Iguana iguana: the
importance of age and nutritional requirements. Oecologia 61(2): 201-207.

Troyer, K. (1984b). Structure and function of the digestive tract of
a herbivorous lizard, Iguana iguana. Physiological Zoology 57(1): 1-8.

161

Iguana iguana

Van Devender, R.W. (1982). Growth and ecology of Spiny-tailed and Green
Iguanas in Costa Rica, with comments on the evolution of herbivory and
large body size. In: Burghardt, G.M. and Rand, A.S. (eds) Iguanas of the
world, their behaviour, ecology and conservation. Noyes Publications, New
Jersey, pp. 162-183.

Werner, D. (1986). Iguana management: conservation of Green Iguanas through
rational use? Notes from Noah 14(1): 11-13.

Werner, D. and Miller, T. (1984). Artificial nests for female Green Iguanas.
Herpetological Review 15(2): 57-58.

Wiewandt, T.A. (1982). Evolution of nesting patterns in iguanine lizards. In:
Burghardt, G.M. and Rand, A.S. (eds) Iguanas of the world, their
behaviour, ecology and conservation. Noyes Publications, New Jersey, pp.
119-141.

162

CAIMAN LIZARD Recommended list: 2
{Possible problem]
Dracaena guianensis Daudin, 1802

Order SAURIA Family TEIIDAE

SUMMARY AND CONCLUSIONS A large and little-known semi-aquatic lizard, found
throughout much of the Amazon basin. There are conflicting reports of its
distribution, but the only reliable records are from Brazil, Ecuador and Peru;
it does not occur in the Guianas. A closely related species, Dracaena
Paraguayensis, occurs in the Matto Grosso of Brazil, Paraguay and Bolivia.
D. guianensis grows to about 1.2 m, and has large, blunt teeth, adapted to
crushing snail shells, which form its diet almost exclusively.

It is hunted chiefly for its skin, up to 90 000 of which appear to be traded
annually. The main importing countries are the USA and Japan, and the main
reported countries of origin are Paraguay, and the Guianas.

Nothing is known about the population size, and so it is impossible to say
whether this level of exploitation is excessive. However almost all of the
reported trade in D. guianensis is from countries which have no wild
populations of the species, and a large proportion is from Paraguay, which has
a ban on wildlife exports. The trade should therefore virtually cease were
CITES controls to be implemented correctly by the importing countries. Many of
the skins traded may be from the Pantanal and therefore of D. paraguayensis.
Consideration should therefore be given to listing the two taxa as "Dracaena
spp." on Appendix II.

DISTRIBUTION Found chiefly within the Amazonian drainage basin, where it is
widespread.

Brazil Occurs throughout the Amazonian region from Belem (Duellman, 1987)
at least as far as Tefé (W.E. Magnusson, pers. comm., 1986), but is absent
from the Atlantic forests (Dixon, 1979). Amaral (1950) indicates that it
occurs up the Tocatins River to the Sao Francisco Basin, and that it is
particularly abundant in the tidal, marshy sections of the Amazon.

French Guiana the type locality of D. guianensis is Cayenne, French
Guiana, and most subsequent discussions of the distribution of the species
have included the country on the basis of this information. However Hoogmoed
and Lescure (1975) showed that there were no other records from the country,
and consider that the type locality is erroneous, representing the port of
export rather than the true origin of the type specimen. They conclude that
D. guianensis does not occur in French Guiana.

Guyana Hoogmoed (1973) indicates that this species is absent from the
western part of the Guianan region (Suriname and Guyana), although Crawford
(1931) lists it in his key to the herpetofauna of the country, which includes
species for which "there seems to be an authentic record of collection within
the colony". No record has been located.

Ecuador Duellman (1987) recorded the species from Santa Cecilia and it was
included, without comment, in a checklist of the reptiles of Ecuador (Miyata,
1982).

Paraguay There are records of three museum specimens of D. guianensis
from the Matto Grosso region (Peraccia, 1902; Burt and Burt, 1931), but
Vanzolini and Valencia (1965) identified these as the subsequently described

163

Dracaena guianensis

and closely related species Dracaena paraguayensis Amaral 1950. Therefore
D. guianensis has not been recorded from Paraguay, and it is unlikely that
it occurs there, as it is outside the Amazon drainage basin.

Peru Recorded from Cashiboya in the Amazonian region (Boulenger, 1885;
Hoogmoed, 1979) and Cuzco Amazonica Reserve on the Madre de Dios River
(Duellman, 1987). In the Iquitos Region, Dixon and Soini (1986) obtained
specimens from Centro Union, Moropon and Iquitos.

Suriname There have been suggestions that this species may occur in
Suriname, but Hoogmoed (1973) refutes them, indicating that it is absent from
the western part of the Guianan region (Suriname and Guyana).

Groombridge’ (1983) states that the species possibly occurs in other countries
bordering the Amazon, but Hoogmoed (1979) indicates that there are as yet no
records from the Amazonian regions of Colombia and Bolivia, nor are there any
from Venezuela (S. Gorzula, in litt., 11 April 1986).

POPULATION There is no information available on the size of the population
of D. guianensis. The paucity of knowledge of the species may indicate that
it is scarce, but it could equally reflect its inconspicuous nature and
tendency to inhabit swamps and the smaller waterways, which are seldom
visited. Duellman (1987) conducted a study of lizards at Cuzco Amozonico
Reserve, Peru, and captured only one D. guianensis, but because of its size
it accounted for 15.5% of the lizard biomass. Dixon and Soini (1986) reported
that during the flood season these lizards were occasionally seen swimming in
the flooded streets of towns and villages in the Iquitos region.

HABITAT AND ECOLOGY A large, semi-aquatic lizard inhabiting river margins
and areas of flooded forest, D. guianensis has seldom been studied in the
wild, and little is known of its ecology. It is particularly characteristic of
the tidal, marshy areas around the mouth of the Amazon. Fully grown it reaches
a length of 4 feet (1.2 m) and is sometimes mistaken for a caiman, having
powerful jaws (Breen, 1974), although the teeth are blunt and rounded, adapted
for crushing snail shells (Darymple, 1979; Preghill, 1984). One of its local
names is Jacuaruxi, and it is regarded as a species of caiman by the fishermen
around Tefe (W.E. Magnusson, pers. comm.). The lizard preys almost exclusively
on snails, which it catches by foraging in the water. Rand (1964) reports that
it hunts underwater, walking along the bottom, and searching under the leaves,
while Vanzolini (1961) observed it feeding walking on its hind legs, with its
head above the water. The snails are usually brought to the surface and
crushed; the fragments of shell are not found in the faeces and are expelled
from the mouth before the snail is eaten (Dixon and Soini, 1986; Vanzolini,
1961).

When not feeding, it usually lies along the branches of trees and bushes,
relying on cryptic colouration to avoid detection (Vanzolini, 1961). The back
is olive brown and the belly is yellowish with black marbling (Cunha, 1961).
Young animals are reported to have green bodies and reddish brown heads (Dixon
and Soini, 1986). It swims well, with strong strokes of the flattened tail,
and may take refuge in holes underwater when disturbed. The habit of resting
in trees may enable it to avoid aquatic predators, chief amongst which are
probably caimans (Vanzolini, 1961).

Very little is known of the breeding biology of this species. There are two

records of females both having 17 yolked ovarian follicles, and a single
report of a nest lodged in a termite mound (Best, 1984; Dixon and Soini, 1986).

164

Dracaena guianensis

THREATS TO SURVIVAL There is very little information regarding exploitation
of this species, although Cunha (1961) reports that it is much persecuted in
Amazonia for its flesh and principally its skin. Dixon and Soini (1986)
reported that a number of skins and a few live individuals are shipped out of
Iquitos each year. Hoogmoed (in litt., 26 August 1986) considered that the
skins of this species reportedly originating in Suriname had almost certainly
been smuggled in from Brazil. In common with other Amazonian species, its
habitat is threatened by deforestation and the extensive hydro-electric dam
projects now under consideration.

INTERNATIONAL TRADE The only reports of international trade in
D. guianensis are those contained in the Annual Reports of Parties to CITES.
Since 1980; there have been no reports of trade in live animals, the bulk
being of trade in skins; the great majority of these were reported as numbers
of skins, with insignificant quantities being reported by weight or length.
These last two categories were therefore excluded from the following
analysis. The CITES reports are summarised in Tables 1 and 2.

Table 1 shows that the minimum world trade volume of D. guianensis skins
varied from 8925 to 85 335 skins between 1980 and 1985. The chief importers
were Japan and the USA. In 1981 and 1984 Japan was a net exporter, although
its gross imports were over 10000 skins. Most of the skins eventually
destined for the USA in 1984 went via Japan, suggesting that they were
probably tanned there. Other major importers are Belgium, Italy and Spain. To
judge by reported countries of export or origin (Table 2), France must also
have imported appreciable quantities of skins, although until 1984 it did not
report imports of Appendix II species.

Table 1. Minimum net imports of skins of D. guianensis reported to CITES,
excluding transactions reported by weight or length.

1980 1981 1982 1983 1984 1985
Argentina - 160 - - - -
Belgium - - - 9117 - -
Canada - - 2 - - -
France - - - - 4636 -
Italy - 200 6163 886 - -
Hong Kong - - - - 188 -
Japan 5000 - 40300 8655 - 1695
Korea - - - 120 116 -
Mexico - 1000 - - - -
Spain - - - 2307 - -
Switzerland 180 - ~ - 8 -
USA 3745 71062 38870 16631 43499 23506
Total 8925 72422 85335 37716 48447 25201

The distribution of D. guianensis is incompletely known, however Table 2
shows that the only country known to have a wild population which is reported
as a country of origin is Brazil. Venezuela may also have some D. guianensis
in the extreme south, and it was given as the country of origin of 100 skins
in 1981. Both Brazil and Venezuela have bans on the hunting of this species
(see Table 3).

165

Dracaena guianensis

Of the source countries not having wild populations of D. guianensis, the
exports from France, Japan and Spain probably indicate re-exports where the
country of origin was not specified. The skins from Indonesia could fall into
this category or they could represent incorrect identification. But the
remaining countries, Argentina, Paraguay, French Guiana, Guyana and Suriname,
all border on Brazil. It is known that large quantities of Caiman skins are
smuggled into Paraguay out of Brazil, where their hunting is prohibited, and
are then exported to consumer countries. It appears that the same is also
happening with D. guianensis, both into Paraguay and also Guyana and
Suriname. The possibility of confusion of the skins with the similar species
Dracaena paraguayensis is discussed below. The whole of the world trade in
this species therefore appears to involve illegally acquired skins, the vast
majority having been smuggled to Paraguay, French Guiana, Suriname and Guyana.

Table 2. Reported countries of origin and quantities of transactions in whole
skins of D. guianensis reported to CITES.

1980 1981 1982 1983 1984 1985

Countries known to have wild populations of D. guianensis
Brazil - 67 - - - -
Countries without wild populations of D. guianensis

Argentina 1092 5000 3000 - = =

France 90 4966 5000 - - -
Fr. Guiana 5180 8417 8909 1104 - -
Guyana - 10141 22680 6715 786 7935
Indonesia - - - 1000 1520 -
Japan - - - - 2000 -
Mexico - - - - 2 -
Paraguay - 28205 47096 34665 31386 18428
Spain 962 - - - - -
Suriname 6490 26339 6653 2913 2538 -
Venezuela - 100 - - - -
Unknown 1253 2893 3992 877 16274 1181

CONSERVATION MEASURES The protection status of D. guianensis in _ the
countries surrounding its range is given in Table 3. With a few marginal
exceptions, the great bulk of the population of D. guianensis lives within
Brazil, a country in which it is totally protected. In spite of this,
large-scale poaching is clearly taking place. The Brazilian Government is
fully aware of this problem and has initiated measures using troops to combat
the poaching and smuggling in the Pantanal region in the South of the country,
where many wildlife products are known to be smuggled out to Bolivia and
Paraguay. The scale of the problem is such that these measures have not yet
had any detectable effect in significantly curtailing the trade. Table 2
indicates that the northern border of Brazil is equally permeable to illegal
wildlife trade, but, so far as is known, no concerted effort has been made to
prevent this from the Brazilian side. Earlier confusion over the distribution
of D. guianensis led to the belief that it occurred in French Guiana, Guyana
and Suriname. It is possible that this mis-apprehension has been exploited by
traders, who have discovered that these countries are acceptable countries of
origin when applying for import permits. This loophole could be closed if the

166

Dracaena guianensis

distribution of the species were specified unequivocally to the CITES
Management Authorities. Measures have been initiated by CITES to ensure that
Suriname is not accepted by importing countries as a possible country of
origin, after the Secretariat was informed by the Suriname Management
Authority that the species did not occur in the country. The same procedure
could be followed with the other two countries.

D. guianensis does not occur in Paraguay, and, in any event, all wildlife
exports from the country are prohibited. Nevertheless all the major importing
countries, notably the USA and Japan, have accepted imports of Paraguayan
skins, apparently without question. The closely related species, Dracaena
Paraguayensis, lives in the Pantanal region of Paraguay and southern Brazil,
and looks very similar to D. guianensis. Vanzolini and Valencia (1965)
remark that "while there is no actual evidence indicating that the two forms
are races of a same species, the possibility cannot at present be ruled out."
It is thought that a large proportion of the illegal trade in Dracaena may
originate in the Pantanal, and therefore be in D. paraguayensis. There is no
information on the status of this species, but it is at least more accessible
than D. guianensis (W.E. Magnusson, pers. comm.). The scarcity of records
for D. paraguayensis (Vanzolini and Valencia, 1965) indicates that its
population density may be similar to that of D. guianensis. Consideration
should therefore be given to including the whole genus in CITES Appendix II, a
step which is, in any event, justified on the grounds that the two species
look very similar (Amaral, 1950).

Table 3. Legal prohibition on the hunting, internal trade and commercial
export of D. guianensis. Dates are those on which the legislation came into
force. A - All live animals & parts; L - Live animals only; S - Skins;
P — Allowed under permit; C - Closed seasons may be imposed; * - these
territories are Overseas Départements of France with which the EEC may trade
without the imposition of CITES controls (Fuller et al., 1985).

CITES Hunting Trade Export
Bolivia 1979 - S 1979 A 1985
Brazil 1975 A 1986 A 1967 A 1967
Colombia 1981 P - L 1978
Ecuador 1975 - - A 1981
Fr. Guiana 1978 * - - P
Guyana 1977 - - A 1986
Peru 1975 A 1973 A 1973 A 1973
Suriname 1981 - - P
Venezuela 1978 Cc A 1970 A 1970

CAPTIVE BREEDING No records have been found of D. guianensis having bred
in captivity, and only one animal was listed in an international survey of 260
zoological collections in 1985 (Slavens, 1985).

REFERENCES

Amaral, A. (1950). Two new South American lizards. Copeia 1950: 281-284.

Best, R.C. (1984). The aquatic mamamals and reptiles of the Amazon. In: Sioli,
H. (ed), The Amazon. Limnology and landscape ecology of a mighty tropical
river and its basin. Dr W. Junk, Netherlands, pp. 371-412.

Boulenger, G.A. (1885). Catalogue of the lizards in the British Museum. Vol.
2, ed. 2, 492 pp.

167

Dracaena guianensis

Breen, J.F. (1974). Bncyclopedia of septiles and amphibians. T.H.F.
Publications, Neptune City, New Jersey, 576 pp.

Burt, C.E. and Burt, M.D. (1931). South American lizards in the collection of
the American Museum of Natural History. Bulletin of the American Museum
of Natural History 61: 227-395.

Crawford, S.C. (1931). Field key to the lizards and amphibians of British
Guiana. Annals of the Carnegie Museum 21: 11-42.

Cunha, O.R. da (1961). II Lacertilios da Amazonia. Os Lagartos da Amazonia
Brasiliera com especial referencia aos representados na colecao do Museu
Goeldi. Boletim do Museu Paraense Emilio Goeldi, Nova Serie, Zoologia

(39): 1-189.
Darymple, G.H. (1979). On the jaw mechanism of the snail-crushing lizards,
Dracaena guianensis (Reptilia, Lacertilia, Teiidae). Journal of

Herpetoiogy 13(3): 302-312.

Dixon, J.R. (1979). Origin and distribution of reptiles in lowland tropical
rainforests of South America. In: Duellman, W.E. (ed.) The South American
herpetofauna: its origin, evolution, and dispersal. Museum of Natural
History, University of Kansas, Monograph (7): 217-240.

Dixon, J.R. and Soini, P. (1986). The reptiles of the Upper Amazon Basin,
Iquitos Region, Peru. Part 1 Lizards and Amphisbaenians. Part 2
Crocodilians, Turtles and Snakes. Milwaukee Public Museum, 154 pp.

Duellman, W.E. (1987). Lizards in an Amazonian rain forest community:
resource utilization and abundance. National Geographic Research 3(4):
489-500.

Fuller, K.S., Swift, B., Jorgenson, A. and Brautigam, A. (1987). Latin
American wildlife trade laws, Second Edition (Revised). World Wildlife
Fund-US, 418 pp.

Groombridge, B. (1982). World checklist of threatened amphibians’ and
reptiles. N.C.C., London, 65 pp.

Hoogmoed, M.S. (1973). Notes on the herpetofauna of Surinam IV. The lizards
and amphisbaenians of Surinam. Biogeographica 4: 1-419.

Hoogmoed, M.S. (1979). The herpetofauna of the Guianan region. In: Duellman,
W.E. (ed.) The South American herpetofauna: its origin, evolution, and
dispersal. Museum of Natural History, University of Kansas, Monograph
(7): 241-280.

Hoogmoed, M.S. and Lescure, J. (1975). An annotated checklist of the lizards
of French Guiana, mainly based on two recent collections. Zoologische
Mededelingen 49(13): 141-172.

Miyata, K. (1982). A check list of the amphibians and reptiles of Ecuador with
a bibliography of Ecuadorean herpetology. Smithsonian Institution
Herpetological Information Service (54): 1-70.

Peraccia, M.G. (1904). Viaggio del Dr A. Borelli nel Matto Grosso brasiliano e
nel Paraguay, 1899. IX Reptili ed aAmfibi. Bollettino dei Musei di
Zoologia e di Anatomia Comparata della R. Universita Torino.

Preghill, G. (1984). Durophagous adaptations in an amphisbaenid. Journal of
Herpetology 18(2): 186-191.

Rand, A.S. (1964). An observation on Dracaena guianensis foraging
underwater. Herpetologica 20: 207.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

Vanzolini, P.E. (1961). Notas bionomicas sobre Dracaena guianensis no Para
(Sauria, Teiidae). Papéis Avulsos do Departamento de Zoologia, Secretaria
da Agricultura, Sao Paulo 14(25): 237-242.

Vanzolini, P.E. and Valencia, J. (1965). The genus Dracaena, with a brief

consideration of macroteiid relationships (Sauria, Teiidae). Arquivos de
Zoologia 13: 7-35.

168

TEGU LIZARDS Recommended list: 1
[Problem]
Tupinambis spp.

Order SAURIA Family TEIIDAE

SUMMARY AND CONCLUSIONS A genus of large lizards, growing to about 1.3 m,
very widely distributed in South America, from Darien to northern Patagonia.
The taxonomy is confused, but there are probably two or three species, the
cured skins of which are virtually indistinguishable. They inhabit many
different habitats, from tropical forests to arid scrub, and are generally
hardy and opportunistic. They are diurnal, retiring to burrows at night, and
are omnivorous, eating mainly invertebrates, small vertebrates, fruits and,
when adult, carrion. In the South of the range, hibernation occurs from about
March to September in burrows or rock crevices. Broods of about 20-35 eggs are
laid in burrows, and usually hatch in late December to January.

They are very extensively hunted for skins, but also locally for meat.
Argentina and Paraguay are the main exporters, the former exporting about a
Million skins annually, mostly to the USA and Western Europe. There is an
insignificant trade in live animals. Argentinian law sets closed seasons and
quotas for hunting on a provincial basis, but these are not based on a
knowledge of population size or biology, and the numbers of Tupinambis are
slowly declining.

The genus is described as common or abundant throughout some of its wide
range, and can undoubtedly withstand a substantial level of trade. There is
currently a large illegal trade in skins from Paraguay which could be halted
if CITES controls were implemented by the importing countries. The bulk of the
skins are legally exported from Argentina under a management scheme, but this
appears to be inadequate at present. A project currently in progress aims to
assess the population status and revise hunting quotas and seasons in the
country. The taxonomy of the genus is in urgent need of revision.

DISTRIBUTION There is considerable confusion over the taxonomy and
distribution of the genus Tupinambis. Peters and Donoso-Barros (1970)
recognise four species: Tupinambis duseni, from a single record in Parana;
Tupinambis nigropunctatus, from the Amazon basin; Tupinambis rufescens,
from western Argentina; and Tupinambis teguixin, from northern Argentina,
Uruguay, Brazil and the Guianas. However the species occurring in the Guianas
is normally considered to be fT. nigropunctatus by most authors (e.g.
Hoogmoed and Lescure, 1975). Presch (1973) attempted a review of the genus in
which he suggested that TT. nigropunctatus was included in T. teguixin,
T. duseni in T. rufescens, and that the species distributed throughout the
arid eastern and south-eastern regions of Brazil and Uruguay, was
T. rufescens and not T. teguixin, as had been previously held. Hoogmoed
and Lescure (1975) questioned his treatment of Tf. nigropunctatus, asserting
that it was distinct, and should at least be considered a subspecies of
T. teguixin. Presch's treatment of the species occurring in southern and
eastern Brazil and Uruguay has received strong opposition from Vanzolini
(1976), Gudynas (1981), and Donadio and Gallardo (1984), who concur with
Achaval and Langguth (1972) in regarding the species as T. teguixin, though
there is some suggestion that T. rufescens may also occur in Brazil
(Vanzolini, 1976). As regards the implementation of CITES controls, there
seems little point in differentiating beyond the level of the genus, as the
skins are not readily distinguishable when cured.

The taxonomic status of fT. ‘nigropunctatus'’ poses few problems, as its
distribution does not overlap with T. ‘teguixin', and it can either be

169

Tupinambis spp.

regarded as a full species (e.g. Hoogmoed, 1973), or as a subspecies of
T. teguixin (e.g. Mertens, 1972). In Argentina the status of T. rufescens
is not in dispute, but in southern Brazil, Uruguay, Paraguay and perhaps
Bolivia, more work is needed to resolve the distinction between this species
and T. teguixin. In the outline of the distribution, the following three
taxa have been used:

Tupinambis ‘nigropunctatus' (Spix): Widespread in northern South America,
north of about 15°S, mainly to the east of the Andes, but reaching the Isthmus
of Darien in Colombia.

Tupinambis ‘teguixin' (sensu Boulenger): Widespread from eastern Argentina
through Uruguay, Paraguay and Bolivia to Brazil, south of about 15°S.

Tupinambis ‘rufescens' (Linnaeus): Mainly ‘in the drier areas of western
Argentina, but possibly extending through Paraguay into Brazil and perhaps
Bolivia.

Argentina Two species occur in Argentina. T. ‘teguixin' is found in the
damper, eastern regions from Missiones, Corrientes, Entre Rios, Formosa,
Chaco, Santa Fe, Santiago del Estero, Cordoba (especially the east), Buenos
Aires and the east of La Pampa (Donadio and Gallardo, 1984). T. ‘rufescens'
is widespread in the drier areas in the west of the country, especially
Sanitago del Estero, south of Jujuy, east of Salta, north-east and south-east
of Tucuman, La Rioja, San Juan, Mendoza, with unconfirmed reports from the
north-east of Neuquen, central La Pampa, south of San Luis, and possibly the
north-west of Chaco (Donadio and Gallardo, 1984) and western Formosa (G.
Hemley, in litt., 25 March 1986). The most southerly record of this species
is from Gran Bajo de Gualicho, south of the Rio Negro (Cei and Scolaro, 1982).

Bolivia Presch (1973) recorded one species, "T%. teguixin", from the Rio
Iténez and the Rio Mamoré, but Hoogmoed and Lescure (1975) suggested that the
species occurring in South America, north of 15°S was T. ‘nigropunctatus’.
This would include the northern part of Bolivia, although in the southern part
T. ‘teguixin' would certainly occur, and the range of T. ‘rufescens' may
also extend into the country. Clearly, detailed studies are necessary.

Brazil The genus occurs thoroughout the country, from Amazonia to the arid
regions in the east. Presch (1973) considered the species in the latter region
to be "“T. rufescens", although its earlier description as "T. teguixin" is
probably correct (Vanzolini, 1976). Within the Amazonian drainage and the
Pantanal, the species occurring to the north of about 15°S is considered to be
T. ‘nigropunctatus', and to the south, T. ‘teguixin' (Hoogmoed and
Lescure, 1975). Presch (1973) considers both of these to be "TP. teguixin".
Vanzolini (1976) indicates that two species of Tupinambis, T. "teguixin' and
T. ‘rufescens', occur sympatrically in the cerrado vegetation region of
Brazil. This implies that three species occur in the country. The description
of Tupinambis duseni Lénnberg and Anderson, 1910 is not generally accepted.

Colombia T. 'nigropunctatus' is widespread, from the Amazonian region
(Hoogmoed, 1979), through Meta, Magdalena, Bolivar and Antioquia to the
Isthmus of Darien (Presch, 1973).

Ecuador Duellman (1987) recorded "T. teguixin" from Santa Cecilia in the
Amazonian region of Ecuador, and Miyata (1982) included "?. teguixin"” in a
checklist of the herpetofauna of the country, but it is not clear what
taxonomy they were following. It is likely to be fT. ‘nigropunctatus' and
not T. ‘teguixin' that occurs there.

170

Tupinambis spp.

French Guiana T. ‘nigropunctatus' occurs in French Guiana, and is
probably present throughout the country (Hoogmoed and Lescure, 1975).

Guyana Both "T. teguixin" and "T. nigropunctatus" are included in
Crawford's (1931) checklist for British Guiana, although it is likely that
only one form occurs there, and that this should be T. ‘nigropunctatus'
(Hoogmoed, 1973; Hoogmoed and Lescure, 1975). Presch (1973) lists several
records from the Demerara district.

Paraguay Two species of Tupinambis have been recorded from Paraguay
(Hellmich, 1960, fide Presch, 1973), probably corresponding to
T. ‘teguixin' and T. ‘rufescens'. The genus is widespread, recorded from
the Departments of Chaco, Central, Concepcion, Caaguazu and Alto-Parana
(Presch, 1973).

Peru Present in the Amazonian region (Hoogmoed, 1979), and recorded from an
altitude of 550 m at Rio Comberciato, near the great head of Urubamba (Presch,
1973). Dixon and Soini (1986) reported that T. teguixin occurred in all
habitats in the Iquitos Region except in direct contact with human
habitation. Duellman (1987) recorded T. teguixin from Cuzco Amazonico
Ecological Reserve on the Madre de Dios River. If Hoogmoed and Lescure's
(1975) suggestion that the species occurring in South America, north of 15°S
is T. ‘nigropunctatus’ is correct, then this is the only form to occur in
Peru. Tupinambis spp. are said to have been observed at Iquitos, Iparia,
Rio Pachitea, Pucallpa, Sarayucu, and Satipo, though the claim that both
T. teguixin and T. rufescens occur (Anon., 1985) is clearly erroneous.

Suriname T. ‘nigropunctatus' occurs throught the country (Hoogmoed, 1973;
in litt., 26 August 1987).

Trinidad and Tobago T. ‘nigropunctatus' is present on both islands
(Hardy, 1982). Presch (1973) has records from Milford Bay, Tobago, and
Princetown, Trinidad.

Uruguay Widespread, from the sandy coastal environments, inland at least to
Tacuarembo (Achaval and Langguth, 1972; Gudynas, 1981). These authors consider
the species to be "7. teguixin", while Presch (1973) refers it to
"T. rufescens".

Venezuela T. ‘nigropunctatus' is widespread throughout the country,
including the llanos region (Rivero-Blanco and Dixon, 1979) and also Amazonia
(Hoogmoed, 1979).

POPULATION

Argentina Said to be common in the east of Buenos Aires (Bella Vista, Campo
de Mayo, San Miguel, Lujan, Jauregui, Lobos, Loma Verde, Otamendi, Zelaya,
Rojas, and in the delta of the Parana), frequent in Cordoba (Laguna del Monte,
Sampacho, Emblase Rio III, Dique la Vina, La Paz and Bialet-Massé), San Luis,
Santa Fe, Entre Rios, Misiones, but not frequent in D'Orbigny (Gallardo,
1977). The largest populations of Tupinambis occur in Formosa and the
western “impenetrable” zone of the Chaco province. Populations are slowly
declining in Salta and Santiago del Estero. Local extinctions near towns have
been noted in these two provinces as well as Formosa and Chaco (Hemley, G. in
dJitt., 7 March 1986).

Bolivia No information.

Brazil Said to be very common in Amazonia (Cunha, 1961), and one of the
most abundant lizards in southern Brazil (Milstead, 1961). T. teguixin was

171

Tupinambis spp.

reported to be seen frequently in the Parque Nacional da Tijuca, Rio de
Janeiro (Anon., 198la).

Colombia No information.
Ecuador No information.
French Guiana No information.
Guyana No information.

Paraguay Populations of Tupinambis were said to have declined in most
inhabited areas. Hunters reported having to walk further to find lizards and
the scarcity can be attributed to overhunting. In some regions, where forests
have recently been cleared, tegu populations are reported to have increased,
but this may be due to the increased ease of observation (Norman, 1987)..
T. rufescens was abundant in the Parque Nacional Defensores del Chaco
(Torres Santibanez, 1978). TT. teguixin is “seen frequently" in Parque
Nacional Ybycui (Anon., 1982).

Peru Dixon and Soini (1986) reported that T. teguixin was observed on
numerous occasions in the Iquitos Region and was more numerous than their
small sample (5 animals) indicated. Duellman (1987) captured two
T. teguixin in a survey of Cuzco Amazonico Ecological Reserve on the Madre
de Dios River and concluded that it accounted for 22% of the lizard biomass.

Suriname Said to be common throughout but nowhere abundant (M.S. Hoogmoed,
in litt., 26 August 1987).

Trinidad and Tobago No information.

Uruguay Although still locally abundant, populations are said to have
decreased as a result of human exploitation (Gudynas, 1981).

Venezuela No information.

HABITAT AND ECOLOGY The large teiid lizards of the genus Tupinambis are
found in a wide variety of habitat types from dense tropical forest and
marshes to arid scrubland. They are very opportunistic and hardy, and may do
well in cattle farming areas. T. rufescens is particularly associated with
arid areas, but even this species preferentially selects the margins of rivers
and ponds (Donadio and Gallardo, 1984).

In the south of the range, at least, Tupinambis hibernates in burrows over
the winter. In Argentina, the hibernation period is usually from March to May
until August to October, but this varies depending on climatic conditions.
Further North the hibernation period shortens, as temperatures increase; and
T. nigropunctatus is active all year round in Venezuela. Burrows may be
excavated to a depth of 0.5-1.5 m, or old vizcacha burrows may be taken over.
Other favoured sites are crevices in rocky areas, hollow tree trunks and gaps
under roots. During the summer they are diurnal, returning to burrows at night
an emerging in the morning and basking before starting foraging (Millstead,
1961; Gudynas, 1981; Donadio and Gallardo, 1984).

Tupinambis is omnivorous, eating a variety of snails, slugs, myriapods,
insects, and occasionally small reptiles, mammals and birds. Fruits and honey
are also consumed, the lizards climbing trees to raid wasp and bee nests.
Predation on the eggs of caimans, turtles, wild and domestic fowl is also
reported (Palermo, 1983). Juveniles eat mostly insects, and as adulthood
approaches there is an increasing trend towards carrion eating (Gudynas, 1981;

172

Tupinambis spp.

Donadio and Gallardo, 1984). This ontogenetic shift in diet is reflected in a
corresponding change in the dentition (Presch, 1974). Juvenile Tupinambis
are preyed upon by snakes and birds of prey. The adults have few natural
predators except man and some larger cats. When cornered they may become
aggressive, with displays of mouth-opening, hissing and tail-lashing. A more
usual response to disturbance is to flee down the burrow (Donadio and
Gallardo, 1984).

Courtship occurs in Argentina in October and November. The males emerge from
their hibernation burrows first, and pursue the females when they appear.
Mating usually takes place in mid-November, after which the female retires to
a breeding burrow. This is usually 0.5 m deep and 1.5 m long, and the nest
chamber is furnished with a layer of dry vegetation into which the eggs are
laid. Some-reports suggest that the female stays with the eggs until they
hatch, usually from late December to early January. The clutch size varies
from 20 to 54. In Pernambuco smaller clutch sizes of around 13-29 are
reported. Egg dimensions are about 41-48 mm by 30-34 mm. Nests. of
T. nigropunctatus are reported to be often associated with arboreal termite
nests (Donadio and Gallardo, 1984). There is some evidence that a second brood
may be laid towards they end of the summer (Gallardo, 1977). The lizards in
Argentina hatch at a length of 19-20 cm and grow to about 35 cm in their first
five months. Adult T. teguixin are normally 1-1.35 m (3.5-4.0 kg), and
slightly smaller for T. rufescens. The tail comprises about half of the
total length (Donadio and Gallardo, 1984). Cunha (1961) reports a maximum size
of 75 cm for his collection of T. nigropunctatus. Males are generally larger
and more robust than females (Palermo, 1983).

THREATS TO SURVIVAL Tupinambis species are probably the most extensively
hunted reptiles in South America. Some are hunted for food and for the pet
trade, the fat is thought to have medicinal properties, but it is exploitation
for skins that provides the biggest commercial incentive. The levels of trade
are so great that they inevitably pose a potential threat to population
levels, although there are few data to suggest that populations are declining
at alarming rates. An Italian reptile leather dealer reported that the market
was very depressed in 1985. The usual size of Tupinambis skins in trade is
17-30 cm belly width (Bodiopelli, in litt., 4 March 1986). There is a minor
trade in live animals for pets. A supplier in the USA (South American
Unlimited, New York) listed "Golden Tegues" (T. nigropunctatus) on its price
list in 1985 for US$15-20 each, depending on size. Habitat modification is
also encroaching on the range of the species, but they are opportunistic
animals, and may be able to establish themselves in cattle grazing areas,
where they can capitalise on the abundant insect life associated with this
type of farming, although population declines are still reported (G. Hemley,
in litt., 7 March 1986). Intensive arable farming and the use of pesticides
would obviously pose a far greater threat.

Argentina Both species are hunted heavily for skins and meat. The
consumption of meat varies from region to region; in some areas it is much
sought-after, while in others it is totally rejected. The fat is supposed to
cure a variety of ailments, including piles, sprains and infections, and is
used to make poultices and as a laxative. It is frequently sold in markets in
Santiago del Estero. Several hunting methods are used, the most common being
with dogs, snares and digging out burrows. Trained dogs, particularly near
Santiago del Estero, are used to locate the lizards or chase them into
burrows. The skin is removed after making an incision along the back, and is
then stretched out to dry. The importance of hunting varies from being the
sole source of income in a few cases, to serving merely as a useful supplement
(Donadio and Gallardo, 1984). The average value of skins exported between 1976
and 1979 was US$4.64 (Mares and Ojeda, 1984). In 1986 the value of skins in
crust was reported to be about US$5-6 each. Previously most of the skins were

173

Tupinambis spp.

transported to tanneries in Buenos Aires or Santiago del Estero but recently a
number of tanneries have opened in the North of the country. This has made the
exploitation still easier. Habitat alteration is also contributing to the
population decline, particularly the removal of the forest and thicket
vegetation to make way for cattle grazing. The western Chaco region (Salta and
Santiago del Estero) is badly affected, and some 80-90% of the habitat has
been altered for grazing and farming (G. Hemley, in litt., 7 March 1986).

Bolivia Large numbers of skins, possibly up to 300 000 a year, are believed
to be illegally exported to Argentina (G. Hemley, in litt., 25 March 1986).

Brazil Tupinambis is frequently hunted for its flesh and skin in Amazonia
(Cunha, 1961), and also in Bahia, where it is thought to have a good flavour
(Anon., 1981b).

Colombia No information.
Ecuador No information.
French Guiana No information.
Guyana No information.

Paraguay Tegus are widely hunted by farmers for skin and meat and for
protection of their poultry. The meat is usually eaten by the hunter and has
no cash value. Sixty-seven farmers interviewed killed an average of 12.7 tegus
each during a summer hunting season, which they sold for a mean total of
US$35, equivalent to 20 days wages for casual labour. The skins are sold to
travelling merchants who usually pass them on to specialist exporters in
Ascuncion (Norman, 1987). Illegal skin exports to Argentina are still
continuing (G. Hemley, in litt., 25 March 1986). Figures compiled by the
Ministerio de Agricultura y Ganaderia showed exports in 1984 of about 91 000
skins of T. teguixin and 36 000 of Tupinambis spp. (Acevedo Gomez, 1987).

Peru Dixon and Soini (1986) reported that a large number of skins and
several live T. teguixin were shipped from Iquitos each year.

Peru No information.

Suriname There is said to be no skin hunting in the country but skins are

though to be smuggled over the border from Brazil (M.S. Hoogmoed, in 1litt.,
26 August 1987).

Trinidad and Tobago No information.

Uruguay Tupinambis is greatly affected by human activity in Uruguay. It
is hunted for skins, meat or sport, often using dogs. The meat of the tail is
particularly favoured, although it only provides an “occasional human meal".
In some localities it is considered a pest and is _ blamed, probably
incorrectly, for raiding eggs from chicken houses (Gudynas, 1981). The
species is also affected by habitat destruction, particularly from
hydro-electric dams and new forest plantations (Gudynas, 1981).

Venezuela Tupinambis is persecuted by rural people in Venezuela, as it is
suspected of raiding chicken houses (Ramo and Ayarzaguena, 1983). There is
not thought to be any systematic exploitation (Venezuela CITES MA, 1987).

Tupinambis spp.

Table la Minimum net imports of Tupinambis skins reported to CITES.

1980 1981 1982 1983 1984 1985
Albania - 1201 - - - -
Australia 4 - 1075 50 - -
Austria 3972 13756 25276 24000 14671 17957
Belgium 6 - 22 25232 - 12913
Canada 25712 20393 1464 10864 33013 40481
= 892 m = 837 m2 = e
Chile - 3 - = = =
China C - = = = = 5000
Colombia 14000 33977 14515 - - -
Cyprus 79 - - - - -
Egypt - 2600 - - - -
France - - 124 - 1122 115179
German D.R. “= — - 8 m2 - -
Germany, F.R. 22747 3075 m 22347 96729 - -
Greece 5454 590 409 180 1474 2793
Hong Kong 12028 35809 32603 14406 88242 10042
3409 kg - 68 kg 3900 kg 3644 kg 10 kg
Hungary 557 - - - - -
Ireland 216 - - - - -
Italy 202402 321853 452506 282436 302660 113439
- 301 kg 16 kg 1000 m - -
Japan - 1504 kg - 12452 kg 14783 kg 1873 kg
Lebanon 2775 1507 - 2300 131 -
Kuwait - - - - - 1
Malaysia - - - - - 124
Mexico 12515 46800 34144 106414 100966 47428
5 kg 114 kg 1481 kg 261 kg 202 kg 770 kg
21m 266 m - - 52 m -
Netherlands 32 - - - - -
New Zealand - - 2 - - -
Panama 1 - - 75203 - -
Paraguay - - - - - 10651
Portugal 80 - - - - -
S. Arabia - - - 4 39 3
S. Africa 1515 5683 100 4 - -
- - - 786 m - -
S. Korea - - 6 - 4621 21950
Spain - - - 80854 70003 96515
Sweden - - 2 - - -
Switzerland 27856 65796 49971 20354 26561 37154
- - 46 m2 - -
Taiwan = - - - 24202 12270
Turkey - - - 50 33 1000
UK 76160 6996 5 m 25099 - 16074
USA 1454299 3044137 1128277 772640 1465389 918468
20297 m 9806 m 106 m - 2m 318 m
Uruguay - - - 2025 260 -
Venezuela 1895 1615 1333 1400 2230 1800
Unknown 4100 1372 22834 - - -
Total 1868405 3604088 1787010 1540244 2135617 1481242
3414 kg 1919 kg 1565 kg 16613 kg 18629 kg 2653 kg
20297 m 13773 m 111m 1786 m 2m 318 m
21 m2 266 m2 = 891 m2 52 m? 2

175

Tupinambis spp.

Table 1b. Minimum net commercial imports of live Tupinambis reported to
CITES.

1980 1981 1982 1983 1984 1985
Austria 1 10 - - 4 4
Belgium - - 20 - - 1
Canada 5764 29 46 23 59 135
Denmark 15 6 - 5 39 6
German D.R. - = - - - 2
Germany, F.R. 223 231 247 162 225 309
Israel ° - - 3 1 - 2
Italy - 4 14 i 43 - 20
Japan 12 98 69 66 69 21
Mexico 10 - - - - -
Netherlands 4 12 - - 2 55
New Zealand 2 - - - - -
Poland - ~ - - 20 -
S. Korea - - - - 5 -
Spain - 6 2 18 - -
Sri Lanka - - - 2 - -
Switzerland 68 70 60 27 - 12
UAE - = 3 = = =
UK - 179 202 209 229 201
USA 7a 1867 1811 2539 2569 2731
Total 6670 2512 2477 3095 3221 3499

INTERNATIONAL TRADE There have been several analyses published in recent
years which include details of international trade in Tupinambis_ skins
(Anon., 1983; Hemley, 1984; Mares and Ojeda, 1984). Most have been based
either on the Annual Reports of Parties to CITES or on the reported exports
from Argentina. The CITES reports are summarised in Tables 1 and 2; only
trade in live animals and skins was considered and all reports of Tupinambis
spp., T. rufescens, T. teguixin and T. teguixin nigropunctatus were lumped
together. °

Minimum net trade in skins of Tupinambis (Table la) varied from over
1 481 000 to over 3 600 000 between 1980 and 1985. Trade reached a peak in
1981, and subsequently declined (see also Hemley, 1984), although there was an
increase in 1984. The main importing countries have been the USA, Canada,
Mexico, Hong Kong, and several European countries, especially Italy, France,
F.R. Germany, Austria, UK and Spain. It should be noted that France did not
report imports of Appendix II species before 1984, and, as it was a major
importer in 1985, the totals shown in the earlier years in Table la may be
artificially low. Japan has usually featured as a net exporter of skins,
indicating that large numbers of skins. are imported and subsequently
re-exported, presumably after tanning. The minimum net imports of live
Tupinambis (Table 1b) have varied between 2477 and 6670, and are therefore
insignificant in comparison with the skin trade. The major net importing
countries have been the USA, Canada, F.R. Germany and the UK.

Tupinambis spp.

Table 2a. Reported countries of origin or export for commercial exports of
skins of Tupinambis reported to CITES.

1980 1981 1982 1983 1984 1985

Countries with wild populations of Tupinambis

Argentina 1263395 1890834 894490 1070300 1448929 1312881
3414 kg 1714 kg 1050 kg 2561 kg 2013 kg 780 kg
= 3936 m 111m - 3m 318 m
29 m2 263 m2 a = 52 m2 1 m2
Bolivia 2 = = 99999 12500 130301
Brazil 30 137000 551 : 8 238
Colombia 535 16817 2062 10132 318 3500
Paraguay 604926 1893658 731854 465464 329529 59203
= 301 kg 515 kg 12739 kg 16013 kg 1873 kg
20297 m 2 E 1786 m z =
Peru 5371 ~ - - - -
Uruguay = e 59355 = 6000 3413

Countries without wild populations of Tupinambis

Canada 4 3 4 1 = =
France 13973 9619 - ~ - -
Germany, F.R. 489 12818 - - - -

~ 590 m - - - -
Indonesia 820 2250 - - - -
Italy 18224 37876 - - - -
Japan - 64740 ~ 4849 1946 26500

- 6172 m - - - -
Panama 2 ~ 213189 71392 10830 -
Netherlands 1980 - - - - -
Nigeria 2884 - - - - -
Singapore - 3612 - -. - 8000
S. Korea - 5280 m - - - =
Spain 12876 13303 = = = -
Sweden 1000 - - - - -
Switzerland - - 27 - - -
Thailand 7 - - - - -
UK 3438 17013 532 1010 1357 -
USA 9249 508 751 16629 5189 8020

s ss = 1764 kg = =
Unknown 40125 73865 64579 55846 150826 1825

= 3801 m 2 865 m2 603 m es

The apparent countries of origin of the skins in trade are shown in Table 2a.
These show that Argentina was the major exporter in all years except 1981,
when Paraguay took over this position. Exports from Argentina have varied
between 0.8 million and 1.9 million. This is remarkably consistent with the
trade revealed by the Argentinian Customs export statistics quoted by Mares
and Ojeda (1984), which showed annual exports between 0.7 million and 1.6
million from 1976 to 1979. Other major source countries in South America have
been Paraguay, Brazil, Colombia, Uruguay and Bolivia. The 99 999 skins in
1983, country of origin Bolivia, were reported as an import to the USA from
Haiti, but is possible that the quantity was an error. Some 19 000 Paraguayan
skins were imported from Haiti in the previous year, and there is known to be
a reptile leather tannery in Haiti (G. Hemley, in litt., 25 March 1986).
Brazil, Paraguay and Uruguay all have bans on the export of wildlife products.

177

Tupinambis spp.
Table 2b Reported countries of origin or export for commercial exports of
live Tupinambis reported to CITES.

1980 1981 1982 1983 1984 1985

Countries with wild populations of Tupinambis

Argentina 4031 - 1 2 48 241
Bolivia 2 - - 10 408 -
Brazil - - - - - 6
Colombia 108 763 1 - - -
Fr. Guiana - 1 - - - -
Guyana : 2570 1720 1916 2849 2637 2903
Paraguay 7 14 2 , - - -
Suriname - 10 20 190 95 43
Venezuela - - = = 5 =

Countries without wild populations of Tupinambis

El Salvador - - 520 - 30 261
Germany, F.R. 10 - - - - -
Guatemala 22 - - - 3 -
Honduras 2 4 - - - -
Italy - - - - 10 -
Panama 36 - - - - -
Thailand 4 = - - - 48
Togo - 10 - - - -
UK 5 - 2 - - -
USA 7 ~ 31 16 3 -
Unknown 10 168 29 50 20 2

Most of the apparent exports from countries without wild populations of
Tupinambis probably represent re-exports where the country of origin was not
specified. Panama is notable as the source of considerable numbers of skins
between 1982 and 1984, as it is known to feature as an entrepot for wildlife
trade leaving South America (Vallester, 1978). A large tannery in Panama,
which closed in 1985, was owned by an Argentinian skin trader (G. Hemley, in
Jitt., 25 March 1986). The majority of these were systematic errors, as
neither Canada nor Switzerland reported countries of origin. The skins from
Indonesia, Nigeria and Thailand may represent incorrectly identified Varanus
skins. The source countries for the live Tupinambis are shown in Table 2b.
Since 1980, the major supplier has been Guyana, which also supplies numerous
other birds and reptiles to the European and North American pet trade.

Unlike many other South American wildlife products in trade, the majority of
Tupinambis skins appear to have been legally acquired and exported from
Argentina. However there are still very large quantities which have apparently
originated in Paraguay, which has a total ban on wildlife exports. It is
possible that some of the skins declared as originating in Bolivia also came
from Paraguay. Argentina already has a management programme for Tupinambis,
which sets quotas for skin hunting and export, although these may not be
adequately enforced. Furthermore they cannot be related to knowledge of
sustainable offtakes at present as the size of the wild population is not
known. A research programme is under way to provide such information, and if
this can be used to revise the quotas, then there is a good potential for
establishing a rational and sustainable exploitation plan. The fact that
Tupinambis populations are reported to be falling slowly in Argentina

Suggests that the current levels of exploitation are excessive, but not
grossly so.

178

Tupinambis spp.

CONSERVATION MEASURES The legal protection status of Tupinambis spp. is
summarised in Table 3.

Table 3 Legal prohibition on the commercial hunting, internal trade and
commercial export of T. teguixin. Dates are those on which the legislation

came into force. A - All live animals & parts; L - Live animals only;
S - Skins; P - Allowed under permit; C - Closed seasons may be imposed; * -
these territories are Overseas Départements of France with which the EEC may
trade without the imposition of CITES controls; ? - no information (Fuller et
al., 1987).

CITES Hunting Trade Export
Argentina 1981 Cc P P
Bolivia 1979 - s 1979 L 1984 +
Brazil 1975 A 1967 A 1967 A 1967
Colombia 1981 P - L 1978
Ecuador 1975 = = A 1981
Fr. Guiana 1978 * ~ A 1986 A 1986
Guyana 1977 - - A 1986 ++
Paraguay 1977 - A 1975 A 1975
Peru 1975 P A 1973 A 1973
Suriname 1981 - - P
Trinidad & Tobago 1984 ? ? P
Uruguay 1975 A 1978 A 1978 A 1978
Venezuela 1978 Cc A 1970 A 1970

+ Bolivia banned the export of all wildlife products from July 1985 for one
year.
++ Export quotas of 7200 live T. nigropunctatus were suggested for 1987 and
1988.

Argentina Exports of raw skins are prohibited, and skins must be tanned or
semi-processed (in crust). The harvest is regulated by the setting of quotas
and closed seasons, which vary from province to province; neither mechanism
appears to be effective (G. Hemley, in litt., 7 March 1986). Attempts are
being made to regulate the harvest on a Federal level; it is hoped to set a
maximum total export quota of 1 million skins, but agreement from the
provincial authorities has not yet been achieved (E.0. Gonzalez Ruiz, in
litt. to G. Hemley, 12 March 1986). In Santiago del Estero, hunting is
permitted all year round. The closed seasons rarely have any basis in a
knowledge of the biology of the animal, and are set more for the convenience
of the hunters. It was suggested that hunting should be prohibited during the
breeding season (November to January), and that a minimum size limit of 20 cm
be set (Donadio and Gallardo, 1984).

Uruguay Tupinambis is not specifically protected in Uruguay, although
there is a blanket ban on wildlife trade. It was thought that population
surveys and further conservation measures were needed to prevent further
population decline (Gudynas, 1981).

CAPTIVE BREEDING Tupinambis is regularly kept in captivity, and has been
successfully bred. A total of 48 animals in 21 different collections were
listed in an international survey of 260 zoological collections in 1985
(Slavens, 1985).

179

Tupinambis spp.

A project is under way to investigate captive-breeding of Tupinambis in
Formosa, Argentina, jointly co-ordinated by the provincial Government and a
local tannery. Two other traders have started trying to breed Tupinambis in
captivity in Argentina since 1985, but no results are available yet.
T. rufescens adapts readily to captivity and tames easily (G. Hemley, in
litt., 7 March 1986). All of the breeding operations are still experimental,
and do not have authorization to operate commercially. To achieve this they
will need to comply with Resolution No. 144 (1983) on the rearing of species
included in Appendix II (E.0. Gonzalez Ruiz, in litt. to G. Hemley, 12 March
1986).

REFERENCES

Acevedo Gomez, C. (1987). Especies de fauna amenazadas por comercio en el
Paraguay. Ministerio de Agricultura y Ganaderia, Unpublished Report.

Achaval, F. and Langguth, A. (1972). Nota sobre habitos anfibios de
Tupinambis teguixin (L) (Sauria, Teiidae). Boletin de la Sociedad
Biologica Uruguaya 2:107.

Anon. (1981a). Plan de manejo, Parque Nacional da Tijuca. IBDF, Brasilia.

Anon. (1981b). Plan de manejo, Reserva Biologico de Poco das Antas. IBDF,
Brasilia. :

Anon. (1982). Plan de manejo, Parque Nacional Ybycui. Ministerio de
Agricultura y Ganaderia, Asuncion.

Anon. (1983). International trade in skins of monitor and tegu lizards,
1975-1979. Traffic Bulletin 4(6): 71-79.

Anon. (1985). Iguanas, In: Manejo de fauna silvestre y desarrollo rural.
FAO, Santiago, Documento Technical No. 2, pp.46-73.

Boulenger, G.A. (1885). Catalogue of the lizards in the British Museum. Vol.
2, ed. 2, 492 pp.

Cei, J.M. and Scolaro, J.A. (1982). A population of Tupinambis from northern
Patagonia, South of the Rio Negro, Argentina. Herpetological Review
L3G) ZGr

Cunha, O.R. da (1961). II Lacertilios da Amazonia. Os Lagartos da Amazonia
Brasiliera com especial referencia aos representados na colecao do Museu
Goeldi. Boletim do Museu Paraense Emilio Goeldi, Nova Serie, Zoologia
(39): 1-189.

Crawford, S.C. (1931). Field key to the lizards and amphibians of British
Guiana. Annals of the Carnegie Museum 21(1): 11-42.

Dixon, J.R. and Soini, P. (1986). The reptiles of the Upper Amazon Basin,
Iquitos Region, Peru. Part 1 Lizards and Amphisbaenians. Part 2
Crocodilians, Turtles and Snakes. Milwaukee Public Museum, 154.

Donadio, O.E and Gallardo, J.M. (1984). Biologia y conservacion de las
especies del genero Tupinambis (Squamata, Sauria, Teiidae) en el
Republica Argentina. Revista del Museo Argentino de Ciencias Naturales
“Bernardino Rivadavia", Zoologia 13: 117-127.

Duellman, W.E. (1987). Lizards in an Amazonian rain forest community:
resource utilization and abundance. National Geographic Research 3(4):
489-500.

Fuller, K.S., Swift, B., Jorgensen, A. and Brautigam, A. (1987). Latin
American wildlife trade laws, Second Edition (Revised). World Wildlife
Fund-US, 418 pp.

Gallardo, J.M. (1977). Reptiles de los alrededores de Buenos Aires.
Editorial Universitaria de Buenos Aires, 213 pp.

Gudynas, E. (1981). Some notes from Uruguay on the behaviour, ecology, and
conservation of the macroteiid lizard, Tupinambis teguixin. Bulletin of
the Chicago Herpetological Society 16(2): 29-39.

Hardy, J.D. (1982). Biogeography of Tobago, West Indies, with special

reference to amphibians and reptiles: a review. Bulletin of the Maryland
Herpetological Society 18(2): 37-142.

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Tupinambis spp.

Hellmich, W. (1960). Die Sauria des Gran Chaco und seiner Randgebeite.
Bayerische Akademie der Wissenschaften Math-Natur. Klasse. Abhandlungen
(Neue Folge) (101): 1-131.

Hemley, G. (1984). World trade in tegu skins. Traffic Bulletin 5(5/6): 60-62.

Hoogmoed, M.S. (1973). Notes on the herpetofauna of Surinam IV. The lizards
and amphisbaenians of Surinam. Biogeographica 4: 1-419.

Hoogmoed, M.S. (1979). The herpetofauna of the Guianan region. In: Duellman,
W.E. (ed.) The South American herpetofauna: its origin, evolution, and
dispersal. Museum of Natural History, University of Kansas, Monograph
(7): 241-280.

Hoogmoed, M.S. and Lescure, J. (1975). An annotated checklist of the lizards
of French Guiana, mainly based on two recent collections. Zoologische
Mededelingen 49(13): 141-172.

Mares, M.A. and Ojeda, R.A. (1984). Faunal commercialization and conservation
in South America. Bioscience 34(9): 580-584.

Mertens, R. (1972). Herpetofauna Tobagana. Stuttgarter Beitrdge zur
Naturkunde 252: 1-22.

Milstead, W.W. (1961). Notes on the teiid lizards in southern Brazil. Copeia
1961(4): 493-495.

Miyata, K. (1982). A checklist of the reptiles of Ecuador with a bibliography
of Ecuadorean herpetology. Smithsonian Herpetological Information Service
No. 54: 1-70.

Norman, D.R. (1987). Man and tegu lizards in Eastern Paraguay. Biological
Conservation 41: 39-56.

Palermo, M.A. (ed.) (1983). La Iguana Overa. Fauna Argentina 22: 1-32.

Peters, J.A. and Donoso-Barros, R. (1970). Catalogue of the Neotropical
Squamata: Part II. Lizards and Amphisbaenians. Bulletin of the US
National Museum 297: 1-293.

Presch, W. (1973). A review of the Tegus, Lizard genus Tupinambis (Sauria:
Teiidae) from South America. Copeia 1973(4): 740-746.

Presch, W. (1974). A survey of the dentition of the macroteiid lizards
(Teiidae: Lacertilia). Herpetologica 30(4): 344-349.

Ramo, C. and Ayarzaguena, J. (1983). Fauna Llanera. Cuadernos Lagoven,
Caracas, 84 pp.

Rivero-Blanco, C. and Dixon, J.R. (1979). Origin and distribution of the
herpetofauna of the dry lowland regions of northern South America. In:
Duellman, W.E. (ed.) The South American herpetofauna: its origin,
evolution, and dispersal. Museum of Natural History, University of
Kansas, Monograph (7): 281-298.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

Torres Santibanez, H. (1978). Conservacion y manejo de la fauna silvestre del
Parque Nacional Defensores del Chaco. FAO Documento de Trabajo FAO/SFN:
6/PAR/O2/T, 61 pp.

Vallester, E. (1978). Informe de Panama sobre la situacion de la fauna
silvestre. In: Morales, R., MacFarland, C., Incer, J. and Hobbs, A. (eds),
Primera Reunion Regional Centroamericana Sobre Vida Silvestre. Centro
Agronomico Tropical de Investigacion y Ensenanza, Turrialba, Costa Rica,
pp. 43-64.

Vanzolini, P.E. (1976). On the lizards of a cerrado-caatinga contact:
evolutionary and zoogeographical implications (Sauria). Papéis Avulsos,
Zoologica, Sao Paulo 29(16): 111-119.

181

SAVANNA MONITOR Recommended list: 2
[Possible problem]
Varanus exanthematicus (Bosc, 1792)

Order SAURIA Family VARANIDAE

SUMMARY AND CONCLUSIONS A large diurnal lizard, widespread in sub-Saharan
Africa, it has been recorded in: Angola, Botswana, Cameroon, Central African
Republic, Chad, Ethiopia, Gambia, Guinea, Guinea-Bissau, Ivory Coast, Kenya,
Liberia, Malawi, Mali, Mauritania, Mozambique, Namibia, Nigeria, Senegal,
Sierra Leone, Somalia, South Africa, Sudan, Tanzania, Togo, Uganda, Yemen A.R,
Yemen P.D.R., Zaire, Zambia and Zimbabwe. It is likely to occur in Benin,
Burkina, Djibouti, Niger and Swaziland, and may be found in Burundi and
Rwanda. Inhabits typically savanna and open woodland habitats. The varied
diet includes small vertebrates, eggs and invertebrates. No detailed
information on population sizes and trends is available, but the species is
said to be common in protected areas and other sparsely inhabited regions in
southern Africa, and scarce in settled regions due to human predation. The
same is likely to hold for northern parts of the range.

Widely used locally for food and for leather’ goods. Internationally,
relatively small numbers are in the live animal trade, with an annual mean of
some 1708 specimens in recent years, but very large numbers of skins are
traded, an annual mean of 88 138 from 1980 to 1985. Nigeria has been the
reported origin of most of these skins, with Mali, Sudan and South Africa also
contributing large numbers in different years. Most of the live animals
originated in Ghana, Kenya and Togo.

While there is no evidence that the species as a whole is threatened, this
scale of exploitation, coupled with local utilisation and persecution, is
likely to lead to local depletions. Field data on population status in
selected countries are required.

DISTRIBUTION Widespread in Africa south of the Sahara, from Senegal east to
Ethiopia and Somalia, and south through the continent to South Africa, but
apparently absent from areas of tropical rain forest in west and central
Africa (Loveridge, 1957; Mertens, 1942; Mertens, 1963, Schmidt, 1919).
Specific records are sparse but the species has been recorded in: Angola,
Botswana, Cameroon, Central African Republic, Chad, Ethiopia, Gambia, Guinea,
Guinea-Bissau, Ivory Coast, Kenya, Liberia, Malawi, Mali, Mauritania,
Mozambique, Namibia, Nigeria, Senegal, Sierra Leone, Somalia, South Africa,
Sudan, Tanzania, Togo, Uganda, Yemen A.R., Yemen P.D.R., Zaire, Zambia,
Zimbabwe. It is likely to occur in Benin, Burkina, Djibouti, Niger and
Swaziland, and may be found in Burundi and Rwanda.

Mertens (1942) recognised four subspecies: V. e. exanthematicus from Senegal
to Egypt; V. e. microstictus from Ethiopia to Tanzania; V. e. albigularis

from southern Africa; and V. e. angolensis which is confined to Angola and
southern Zaire.

Angola Mertens (1942) noted the occurrence of V. e. microstictus and
cited numerous records of V. e. angolensis.

Benin No records have been located, but the species is most likely to occur
as it found in neighbouring countries, Togo and Nigeria.

Botswana Mertens (1942) cited several records of V. e. microstictus. It
is particularly associated with the drier areas (Auerbach, 1985).

182

Varanus exanthematicus

Burkina No records have been located, but the species is most likely to
occur as it found in neighbouring countries, Togo, Ghana, Ivory Coast and
Mali. V. exanthematicus is mentioned by name in the national legislation.

Burundi Mertens (1942) noted no localities in Burundi, but recorded
V. e. microstictus from the Tabora District of Tanzania and from
south-eastern Zaire.

Cameroon Two species of Varanus occur in Cameroon, V. niloticus’ and
what is referred to as V. griseus, the "Varan de terre" (Tsague, 1986).
V. griseus is also referred to in the national legislation. However, it is
unlikely that V. griseus occurs in Cameroon as it is confined in Africa to
the extreme north (Mertens, 1942), and it is most probable that "V. griseus"

refers to V. exanthematicus. Mertens gave no localities for the country,
but he cited records for V. e. exanthematicus as "Lake Chad", without
specifying which shore. "V. griseus" is said to occur in Kalamaloué, Waza,

Boubandjida, Benoué and Faro protected areas (Tsague, 1986), all of which are
located in the north of the country.

Central African Republic V. e. exanthematicus was recorded from Batafango
(Mertens, 1942).

Chad V. e. exanthematicus was recorded from around Lake Chad and Bahr el
Ghazal (Mertens, 1942).

Djibouti No records have been located, but the species is most likely to
occur as it found in neighbouring areas of Somalia (Simonetta and Magnoni,
1986) and Ethiopa.

Egypt Mertens (1942) noted one 1826 record from “Aegypten", but it seems
unlikely that this refers to Egypt.

Ethiopia Mertens (1942) cited several records of V. e. exanthematicus
from Eritrea and of V. e. microstictus from Abyssinia.

Gambia Records of V. e. exanthematicus noted by Mertens (1942), including
MacCarthy Island in the Gambia River.

Ghana Mertens (1942) noted one record of V. e. exanthematicus from the
Gold Coast and one from Jendi (=Yende) in what was then Togo but is now Ghana.

Guinea Klaptocz (1913) recorded V. exanthematicus from "Am Niger", French
Guinea, presumably meaning the Niger River. Mertens (1942) cited another
record of V. e. exanthematicus from Guinea.

Guinea-Bissau Recorded from Bissalanca, Brene and Tor, on Bissau Island;
Canchungo, Pecixe and Pitche (Manacgas, 1955). Records of V. e.

exanthematicus were noted by Mertens (1942)

Ivory Coast V. exanthematicus occurs in Comoe National Park (Anon.,
1979), and is likely to occur in other dry areas.

Kenya Numerous records of V. e. albigularis were noted by Mertens (1942),
from Ludwar to Malindi.

Lesotho V. exanthematicus is said to be extinct in Lesotho (Lesotho,
Ministry of Agriculture and Marketing, in litt., 1986).

Liberia One record of V. e. exanthematicus was noted by Mertens (1942).

183

Varanus exanthematicus
Malawi Mertens (1942) noted records of V. e. albigularis from Nyassaland.

Mali V. e. exanthematicus was recorded from Tombouctou (Chabanaud, 1917)
and Goundam (Angel, 1933), Kidal and north of Bamako (Mertens, 1942).

Mauritania Mertens (1942) cited Flower (1929) indicating that the
distribution of V. e. exanthematicus extended from Mauritania to the Niles,
though no records from Mauritania were given.

Mozambique Mertens (1942) cited records of UV. e. microstictus from
Chifumbazi and V. e. albigularis from Quitangonha, Sena, Tette and Pungwe
River.

Namibia Numerous records of V. e. albigularis noted by Mertens (1942).
Niger V. exanthematicus is said to be "unknown" in Niger (Niger CITES MA,
1986). Mertens gave no localities for the country, but he cited records for

V. e. exanthematicus as “Lake Chad", without specifying which shore.

Nigeria Said to occur throughout the savanna zone (Nigeria CITES MA,
1987). Dunger (1967) reported it to be common in grassland in the north.

Rwanda Mertens (1942) noted no localities in Rwanda, but recorded
V. e. microstictus from the Tabora District of Tanzania and from
south-eastern Zaire. V. exanthematicus is not mentioned in national

legislation, although V. niloticus is.
Senegal Numerous records of V. e. exanthematicus noted by Mertens (1942).

Sierra Leone Reported to be present in the Outamba-Kilimi area, but no
specimens were collected (Teleki, 1980).

Somalia V. e. microstictus has been recorded from Wewst Galbeed,
Togdheer, Bari, Nugal, Mogadishu, Lower Shebelle, Bay, Gedo and Lower Jubba
(Simonetta and Magnoni, 1986). Mertens (1942) cited numerous records.

South Africa Found throughout the country except in the western and
southern Cape Province (McLachlan, 1978). Numerous records of
V. e. albigularis noted by Mertens (1942).

Sudan Numerous records of V. e. exanthematicus were cited by Mertens
(1942), including Dongola, Kordofan, Sennaar, Roseires, Blue and White Niles.

Swaziland No records have been located, but the species is most likely to
occur as it found in neighbouring countries, South Africa and Mozambique.

Tanzania Numerous records of V. e. microstictus were noted by Mertens
(1942), from the Tabora District to Lindi. Pakenham (1983) refutes the
earlier records from the island of Zanzibar, but noted a sight record from the
island which could have been of this species. Laurent (1964) described an
endemic subspecies, V. e. ionidesi, from south-east Tanzania.

Togo Records of V. e. exanthematicus were noted by Mertens (1942),
including Bismarckburg, Mangu, Basari and Sokodé.

Uganda One record of V. e. microstictus was cited by Mertens (1942).
Yemen Arab Republic Arnold (1986) noted the occurrence of
V. exanthematicus from North Yemen, but said that no specimens had been

collected from there. There is some dispute that the form occurring in Yemen

184

Varanus exanthematicus

may represent a new species.

Yemen Peoples' Democratic Republic Two specimens were collected from El
Khobar (Arnold, 1986).

Zaire V. e. exanthematicus was recorded from Mauda, in the north-east;
V. e. microstictus from Kikondja, Kansenia, Lukafu, Kiambi and Lukulu in the
south-east; and V. e. angolensis from Kabengere (Mertens, 1942).

Zambia V. e. albigularis was recorded from the “bank of Loangwa" and
north of the Zambezi (Mertens, 1942). Said to be widespread, but most
abundant in the Zambezi and Luanga valleys (Broadley, 1971).

Zimbabwe Numerous records of V. e. albigularis were noted (Mertens, 1942).

POPULATION Said to be still common in sparsely inhabited areas of southern
Africa generally, including the Kalahari region, National Parks and cattle
ranches, but scarce in densely populated communal land due to human predation
(D.G. Broadley, in litt., 18 March 1986). The same is likely to be true for
other parts of the range. Little information is available for individual
countries.

Botswana V. exanthematicus is said to be more common than V. niloticus
around Gaberone (Auerbach, 1985).

Guinea-Bissau Said to have been common in 1896 (Manacas, 1955).

Kenya Common along the coast and in Tsavo East and Tsavo West National
Parks (A.D. Mackay, in litt., 26 March 1986).

Malawi Said to be uncommon and seldom seen (C. Dudley, in litt., 14 April
1986).

Mali Said to have been very rare around Goundam (Angel, 1933).

Mozambique Uncommon in most areas, though there are good populations in
conservation areas (Mozambique CITES MA, 1986).

Namibia Said to be abundant in some Nature Conservation Areas, such as
Etosha, but marginal in others. Otherwise, scarce in heavily populated
communal lands, but relatively common elsewhere (Namibia Department of
Agriculture and Nature Conservation, in litt., 1987).

Nigeria Said to be common, but threatened (Nigeria CITES MA, 1987).
Formerly reported common in grassland in the north (Dunger, 1967).

South Africa Considered to be vulnerable outside game reserves, though
common in reserves such as Ndumu and the southern Kruger Park (McLachlan,
1978). The species is said to be less common than V. niloticus, but often
encountered in the larger Zululand reserves (South Africa CITES MA, 1987).

Uganda Said to be common in the north-east of the country, where cattle
keep the grass short (Uganda Game Department, in litt., 1987).

Zambia Said to be most abundant in the Zambezi and Luanga valleys
(Broadley, 1971). Common in wooded savannas of Zambia and other parts of
southern Africa, but threatened by utilisation and habitat changes (M.P.
Simbotwe, in litt., October 1983).

185

Varanus exanthematicus

Zimbabwe Still common in sparsely populated areas, though scarce in the
densely populated communal lands due to predation (D.G. Broadley, in litt.,
18 March 1986).

HABITAT AND ECOLOGY A large diurnal lizard, up to nearly 1.5 m in length.
Typically associated with dry areas, especially the variety of savanna-type
habitats and Brachystegia woodland. Occupies animal burrows, holes in the
ground or rock crevices, and may aestivate therein during acute drought. The
diet mainly comprises small mammals and birds, but vertebrate eggs and
invertebrates are also taken. In South Africa, the diet consists mainly of
millipedes and tenebrionid beetles. An able swimmer; often climbs into trees
when disturbed or while resting. Females lay a clutch of 30-40 eggs,
ca 60 x 35 mm, in a hole which is covered over for the incubation period
(Auerbach, 1985; Broadley, 1971; Dunger, 1967). Smaller clutches of 7-15 eggs
are reported for V. e. albigularis (South Africa CITES MA, 1987).

THREATS TO SURVIVAL Apparently adversely affected by human predation for
both food and skins (D.G. Broadley, in litt., 18 March 1986; Dunger, 1967;
M.P. Simbotwe, in litt., October 1983). No details are available on the
effect of these factors on lizard populations, or on their relative intensity
in different parts of the continent.

Botswana There seems to be no large-scale commercial collecting although
local food use is important (R.D. Auerbach, pers. comm., 1986).

Mozambique There has been very little commercial exploitation of the
species over the last ten years, but it is hunted for food in rural areas, and
human predation is said to be significant. Bush fires may also pose a threat
(Mozambique CITES MA, 1986).

Namibia Although many V. exanthematicus are killed on roads, the species
is not particularly sought after as food (Namibia Department of Agriculture
and Nature Conservation, in litt., 1987).

South Africa Said to be exploited for skins and pets. It is also often
killed out of fear and is sometimes used by witchdoctors for medicinal
purposes (McLachlan, 1978).

Uganda The species is not used for food in the north-east of the country,
but is regarded as a delicacy in the West-Nile region. It is also persecuted
as a notorious predator of chickens’ eggs (Uganda Game Department, in litt.,
1987).

INTERNATIONAL TRADE CITES reports, summarised in Tables 1 and 2, indicate
that large numbers of V. exanthematicus are involved in the skin trade.
Reported quantities have ranged between 14 010 and 215 952 between 1980 and
1985, with an annual mean of 88 138. Nigeria has been the reported origin of
most of these skins, with Mali, Sudan and South Africa also contributing large
numbers in different years. Many of the skins were re-exported from F.R.
Germany in 1985. It is not known whether skins exported from Nigeria are
collected in that country or in other parts of West Africa. In the absence of
good population information for any part of the range, the sustainability of
trade cannot be assessed. Although local declines have been noted, there is
no evidence that the species as a whole is significantly threatened.

Relatively small numbers of live animals (mostly from Ghana, Kenya and Togo)
are in reported international trade, between 434 and 5448 from 1980 to 1985,
with an annual mean of 1708; most of these were imported by the USA, with
lesser numbers to F.R. Germany and other parts of Europe.

186

Varanus exanthematicus

Table 1 Minimum net imports of skins of Varanus exanthematicus reported to
CITES. Only transactions recorded by number of skins were included

— eRe —OSOSo.SOSOSS ee eee — — — — —————

1980 1981 1982 1983 1984 1985
ao eee ee ee el Be ee |
Albania - 287 - - - -
Australia - - 8 - - -
Austria 2835 4431 1044 6360 - -
Belgium 329 - - - - -
Canada 1128 80 60 420 360 507
France 4517 5030 - - - -
German D.R. - - - 1 - -
Germany, F.R. 37456 8363 = = = =
Hong Kong 60 25709 486 - 450 ~
Italy 29925 - - - 13113 302
Lebanon 24 - - - - -
Netherlands - - 1350 - - -
New Zealand - - 1 - - -
South Africa 248 72 100 - - -
Spain - - - 7301 - -
Switzerland - 12007 3 - 8 -
Uruguay - - 22 - - -
UK 5018 - - 9315 - -
USA 30032 159973 15812 8692 - 143450
Yugoslavia - - - - 79 94
Country Unknown - - 300 - ~ -
TOTAL TAS 72 215952 19183 23760 14010 144351

Table 2 Reported countries of origin (or exporting country if no origin
reported) and quantities of transactions in skins of V. exanthematicus
reported to CITES.

1980 1981 1982 1983 1984 1985

a. Countries with wild populations of V. exanthematicus.

Mali - - 1 1460 - -
Nigeria 118916 168810 19495 14340 153 143692
Sudan - 1809 514 10366 - 60
South Africa - 4418 - - 13927 -
Senegal - 849 - - - -

b. Countries without wild populations of V. exanthematicus.

Afghanistan - - = = 450 ie
German D.R. - 100 - = = =
Germany, F.R. 2 - - = = Z
Indonesia - 14905 - = = =
Philippines - 2500 = = = =
Singapore - 5454 - = = =
Thailand - 2000 - 47 = x
Togo - - = 3 = es
UK 6280 6806 - 420 10 -
Country Unknown 35960 28328 128 361 - -

Varanus exanthematicus

CONSERVATION MEASURES Unless otherwise stated, all of the information on
protection is from IUCN Environmental Policy and Law Occasional Paper No. 3,
African Wildlife Laws.

Angola Not protected under the Hunting Regulations, 11 December 1955.

Benin Under the Decree concerning hunting and capturing licences, bag
limits and professional hunters, 11 February 1980, Varanidae are designated as
small game species. They may only be hunted by the holders of traditional
hunting rights or under hunting permits.

Botswana The hunting of Varanus spp. is controlled under the Fauna
Conservation (Unified Hunting) Regulations 12 March 1979, which set out the
conditions of issue of hunting licences and the number of animals which may be
taken by each hunter.

Burkina V. exanthematicus and V. niloticus are partially protected
under the Wildlife Conservation and Hunting Act, 31 December 1968, which
establishes bag limits and close seasons, protects nests and eggs, and
regulates the sale of meat. Commercial capture and export are controlled
under the same Act by Decree No 68-314. Ordonnance No 68-58, December 1985,
fixed the open season from 15 December 1985 to 28 February 1986 and stipulated
that partially protected species could only be hunted under special permit and
only in the province of Gourma and Tapoa.

Burundi No information.

Cameroon Varanus niloticus and Varanus griseus (sic.) are partially
protected under the Forests, Wildlife and Fisheries Act, 27 November 1981.
They may only be taken under a valid hunting licence. No more than two

animals may be taken in one day.

Central African Republic The Ordinance concerning the protection of
wildlife and regulating hunting, 27 July 1984, designates Varanus as totally
protected.

Chad No information.

Congo The Act concerning the conservation and exploitation of wild fauna,
21 April 1983, vests in the State ownership of all wild animals of economic
value, and requires the issuing of licences for commercial capturing. A
minimum size is set for the taking of Varanus, and fees are set for the
issuing of certificates of lawful possession or export of monitor lizards.
Under a later Order (18 May 1984), Varanus flavescens (sic.) is designated
partially protected, and may therefore only be taken under a hunting licence.

Djibouti No information.

Ethiopia Under the Wildlife Conservation (Amendment) Regulations, 12
February 1974, Varanus niloticus and Varanus exanthematicus are
protected. They may only be hunted for scientific purposes and require a
special permit, the value of which was set at Eth$1.00 for each Varanus.

Gambia Under the Wildlife Conservation Act, 14 February 1977, all wildlife
except game and vermin are protected.

Ghana The Wildlife Conservation Regulations, 4 March 1971, define Varanus

niloticus as totally protected and V. exanthematicus as partially
protected, for which hunting is permitted under licence during the open season.

188

Varanus exanthematicus

Guinea No information.

Guinea-Bissau The Hunting Regulations, 12 May 1980, require the issuance of
a licence for the capturing of live wild animals.

Ivory Coast Varanus niloticus’ and V. exanthematicus are listed as
predatory animals in the Wildlife and Hunting Act, 4 August 1965, and may be
hunted without licence anywhere except in protected areas. Arrété No. 15, 26
December 1972, establishes licence fees for the capturing of live reptiles.
Under the Order regulating the commercial hunting of crocodiles and monitor
lizards, 29 September 1967, a special permit is required to capture monitor
lizards for commercial purposes. The possession of specimens having a skin
width of less than 25 cm is prohibited.

Kenya Protected by legislation in Kenya, export being banned (Legal Notice
No. 152, 25 September 1981).

Lesotho Varanus spp. are totally protected under the Proclamation of
Monuments, Relics, Fauna and Flora, 1969.

Liberia The Wildlife Conservation Regulation (in draft, 1985) lists
V. niloticus as partially protected, thereby providing for closed seasons.
Trade in and export of fully and partially protected species is prohibited
except for educational or scientific purposes.

Malawi Hunting, possession, trade and export of game is controlled under
the Game Act, 1953. No reptiles, other than crocodiles, are protected.

Mali All Varanus species are classified as game species under the Hunting
Act, 11 November 1969. There is a general close season from 1 June to 31
October. A general hunting ban (Decree 325/PG-RM) throughout the country was
imposed on 6 November 1978.

Mauritania Under the Hunting and Wildlife Protection Act, 15 January 1975,
the capture, possession, sale and export of live wild animals requires a
commercial capture permit. The sale of game meat is prohibited.

Mozambique V. exanthematicus is fully protected by national legislation.
It occurs in Gorongosa National Park (Mozambique CITES MA, 1986).

Namibia Protected by the Nature Conservation Ordinance (Namibia Department
of Agriculture and Nature Conservation, in litt., 1987).

Niger All hunting for utilitarian purposes has been prohibited in Niger
since 1972 (Niger CITES MA, 1986).

Nigeria V. niloticus and V. exanthematicus are listed as fully
protected in the Endangered Species (control of International Trade and
Traffic) Decree, 20 April 1985. Their hunting, capture, trade and export is
prohibited. On 28 May 1982, the CITES Secretariat sent a notification (No.
218) to all Parties requesting them not to accept imports from Nigeria as the
Nigerian Management Authority did not issue any export permits.

Rwanda Varanus niloticus is partially protected under the Hunting
Regulations, 31 December 1974. A licence is required for hunting and close
seasons may be set.

Senegal The Game and Wildlife Protection Regulations, 30 May 1967, lay down
regulations governing the issuance of licences for hunting and commercial
capturing of wildlife.

189

Varanus exanthematicus

Sierra Leone All species of Varanus are listed as prohibited animals in
the Wildlife Conservation Act, No. 27, 1972. It is illegal to hunt or possess
them.

Somalia Varanus are listed as game animals under the Law on Fauna
(Hunting) and Forest, 25 January 1969. They may only be hunted or exported
under licence. A ban on all hunting was instituted on 13 October 1977.

South Africa V. exanthematicus is legally protected in the Cape Province,
Natal and partially in the Transvaal (McLachlan, 1978). Occurs in 14 out of
60 reserves and resorts in Natal (South Africa CITES MA, 1987).

Sudan V. exanthematicus and V. niloticus are listed in the Ordinance
for the Preservation of Wild Animals, 1935 (Amended 1974), as species for
which hunting is permitted. Licences are only required if firearms are used
for hunting. The Hides and Skins (Export) Regulations, 1969, specify a
grading system for the export of reptile skins.

Swaziland Apparently no reptiles are protected in Swaziland.

Tanzania Under the Wildlife Conservation Act, 30 June 1974, the capture of
all live animals requires a valid capture permit, and the hunting of all
animals requires a valid hunting permit.

Togo V. exanthematicus and V. niloticus are classified as pest species
under the Ordinance on Wildlife Protection and Hunting, 16 January 1968. An
enabling decree, dated 4 June 1980, sets out licence fees for hunting and
commercial capture, but there is no bag limit for these species.

Uganda No reptiles, other than crocodiles, are protected.

Zaire The commercial capture of unprotected animals requires a permit under
the Hunting Act, 28 May 1982. Export can only be authorised if the exporter
holds a certificate of lawful possession.

Zambia V. exanthematicus and V. niloticus are listed as game animals in
the National Parks and Wildlife Act, 3 December 1968, under which their
hunting, possession, trade and export are controlled and fees are set for the
export of trophies and live animals.

Zimbabwe The collection of live reptiles and their eggs and their breeding
in captivity are controlled under licence by the Parks and Wildlife (General)
Regulations, 1981.

CAPTIVE BREEDING A recent inventory lists 26 specimens in 16 collections in
1984 (Slavens, 1985); breeding is recorded in Rotterdam Zoo in 1979 (Olney,
1982).

REFERENCES

Angel, M.F. (1933). Sur quelques reptiles et batraciens du nord du Soudan
Francais. Bulletin du Muséum d'Histoire Naturelle, Paris, (ser. 2),
5: 68-69.

Anon. (1979). Etat actuel des parcs nationaux de la Comoe et de Tai ainsi
que de la Réserve d'Azagny et propositions visant a leur conservation et a
Jeur développement aux fins de promotion du _ tourisme. Tome II: Parc
National de la Comoe. Partie 1: Inventaire des conditions écologiques et
biologiques. GTZ, Eschborn.

Arnold, E.N. (1986). A key and annotated check list to the lizards and
amphisbaenians of Arabia. Fauna of Saudi Arabia 8:385-435.

Auerbach, R.D. (1985). The Reptiles of Gaborone, Botswana Book Centre, 48 pp.

190

Varanus exanthematicus

Broadley, D.G. (1971). The Reptiles and Amphibians of Zambia. The Puku
6: 1-143.

Chabanaud, M.P. (1917). Enumération des reptiles non encore étudiés de
l'Afrique occidentale, appartenant aux collections du Muséum, avec la
description des espéces nouvelles. Bulletin du Muséum d'Histoire
Naturelle, Paris 23: 83-105.

Dunger, G.T. (1967). The Lizards and Snakes of Nigeria (Part 3: the Monitors
and a Plated Lizard). Nigerian Field 32: 170-178.

Flower, S.S. (1929). List of vertebrated animals exhibited in the gardens of
the Zoological Society of London, 1828-1927, 3 Reptiles. London.

Klaptocz, A. (1913). Reptilien, Amphibien und Fische aus FranzdOsisch Guinea.
Zool. Jb. Syst. 34: 279-290.

Laurent, R.F. (1964). A new subspecies of Varanus exanthemeticus (Sauria,
Varanidae). Breviora 199: 1-5.

Loveridge, A. (1957). Checklist of the reptiles and amphibians of East Africa

(Uganda; Kenya: Tanganyika; Zanzibar). Bulletin of the Museum of
Comparative Zoology, Harvard 117(2): 153-362 and i-xx + vi.
McLachlan, G.R. (1976). South African Red Data Book - Reptiles and

Amphibians. South African National Scientific Programmes Report No. 23.
Manacas (1955). Saurios e ofidos de Guiné Portuguesa. Anais de Junta de
Investigacoes do Ultramar (Estudios de Zoologia) 10(4): 190-193.

Mertens, R. (1942). Die Familie der Warane (Varanidae). Abhandlungen der
Senckenbergischen Naturforschenden Gesellschaft 466: 235-391.

Mertens, R. (1963). Liste der rezenten Amphibien und _ Reptilien.
Helodermatidae, Varanidae, Lanthanotidae. Das Tierreich 79. Walter de

Gruyter, Berlin.

Pakenham, R.H.W. (1983). The reptiles and amphibians of Zanzibar and Pemba
Islands. Journal of the Bast Africa Natural History Society and National
Museum 177: 1-40.

Olney, P.J.S. (ed.) (1982). Reptiles bred in captivity and multiple
generation births, 1979. International Zoo Yearbook 21. Zoological
Society of London.

Schmidt, K.P. (1919). Contributions to the herpetology of the Belgian Congo
based on the collection of the American Congo Expedition 1909-1915.
Bulletin of the American Museum of Natural History 39(2): 385-624.

Simonetta, A.M. and Magnoni, M.L. (1986). Status and conservation problems of
Somali lower vertebrates. Rivista di Agricoltura Subtropicale e
Tropicale 80(3): 405-432.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity,
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

Teleki, G. (1980). Status report on the Wildlife Survey Project, Sierra
Leone, conducted November 1979 through May 1980. WWF-US Project Report
(#1728).

Tsague, L. (1986). Contribution a l'étude des parcs nationaux et réserves
analogues du Cameroun. MSc dissertation, Dépt. de Biologie et Physiologie
Vegétales, Université de Yaoundé.

191

PACIFIC or MANGROVE MONITOR Recommended list: 3*
[No problem]
Varanus indicus (Daudin, 1802)

Order SAURIA Family VARANIDAE

* The category has been changed from list 2 to 3 since approval by the CITES
Technical Committee meeting, 1986, owing to the absence of recent trade.

SUMMARY AND _ CONCLUSIONS A moderately large monitor’ lizard, rather
widespread in the western Pacific. Ranges from eastern Indonesia, east
through New Guinea to the Solomon Islands, south to northern Australia and
north to several island groups in Micronesia (much of the present distribution
here is due to introduction by man, often as late as World War II). Said to
be less common than previously in some places, but also reportedly common in
Papua New Guinea and parts of the Solomons and Guam. Prefers semi-open areas
to dense forest, often along the littoral and in mangroves, often in coconut
plantations. Has been shown to feed on various pest species, such as Giant
African Snails, and rats in coconut plantations (and has been examined as a
possible rat control agent), but widely regarded as a pest itself due its
habit of feeding on chickens and other human food items.

Apparently not heavily exploited and probably not in excessive numbers in
international trade. CITES Annual Reports show that the only substantial
trade was 13 080 skins from Indonesia in 1980, since when trade has been
negligible.

Although it is unlikely that international trade at the reported levels poses
a significant threat to the species, it would be advisable to obtain data on
population levels and exploitation rates, particularly for Indonesia.

DISTRIBUTION A western Pacific species, ranging from eastern Indonesia
north-east to the Marianas and Marshall Islands, and east through New Guinea
and northern-most Australia to the Solomon Islands. The natural range in
Micronesia is uncertain; the species is known to have been very widely
introduced in the region, during the German administration prior to WwWl and by
Japanese and U.S. forces around WW2, and quite possibly at various times
previously by the indigenous population of the region (for use as food).
Present in the following countries, with details of distribution where
available. Three subspecies have been recognised: V. i. spinulosus, which
is confined to “Georges Island" in the Solomon Islands; V. i. kalabeck,
which is confined to Waigeo Island, Indonesia; and V. 4. indicus, which
occupies the remainder of the range (Mertens, 1942).

Australia Rainforest and coastal mangroves of eastern Cape York Peninsula
(Queensland), coastal mangroves of Arnhem Land (Northern Territory), and the
islands of Torres Straits (Cogger, 1975; Cogger et al., 1983).

Federated States of Micronesia Present on Kosrae, Mortlock, Woleai, Ifalik,
Yap (Dryden, 1965; Uchida, 1966); also found on Ulithi and several islands in
the Truk Atoll (USA CITES MA, 1987).

Guam Present (Dryden, 1965; Dryden and Taylor, 1969; USA CITES MA, 1987).

Indonesia Apparently widespread in the eastern sector of the archipelago,
including Irian Jaya and adjacent islands, north to the Talaud group and west
to Halmahera and the Moluccas (Ambon, Buru, Seram etc.) and Timor in the
Lesser Sundas. De Rooij (1915) includes Sulawesi within the range of this
species but Mertens (1963) does not.

192

Varanus indicus

Mariana Islands Present on Agiguan, Rota, Tinian, Sarigan, Saipan, (Dryden,
1965, Uchida, 1966) and on some of the islands north of Sarigan (USA CITES MA,
1987); no information on other islands in the group.

Marshall Islands Introduced into Japten before WwW2 (Dryden, 1965); no
information on other islands in the group.

Palau Present on Ngeaur (Angaur), Oreor (Koror), Babeldaob (Babelthaup),
Ncheangel (Kayangel) (Dryden, 1965, Uchida, 1966).

Papua New Guinea Occurs widely throughout the main island and the island
provinces (Whitaker et al., 1982).

Solomon Islands Widespread, including the main islands of Guadalcanal,
Isabel, Malaita, San Cristobal, (McCoy, 1980) but apparently not recorded from
Choiseul or New Georgia.

POPULATION A popular source (Schmidt and Inger, 1957) states that this
species is not as abundant as it once was on Pacific islands. Published
comments on status are given below, but no adequate population estimates
appear to be available for any part of the species's range.

Australia The species is said to be uncommon although there is no
information on population size. It is encountered infrequently in the wild
but populations are considered to be stable (Australia CITES MA, 1986).

Federated States of Micronesia Fairly common on islands where they occur
(USA CITES MA, 1987).

Guam Relatively abundant in the northern sector (Dryden and Taylor, 1969).
Indonesia No information.

Mariana Islands Relatively abundant on Agiguan; relatively common in
uninhabited forested areas on Rota and Saipan; uncommon on Tinian (USA CITES
MA, 1987).

Marshall Islands No information.

Papua New Guinea Reportedly the most common of the six Varanus species in
PNG (Whitaker et al., 1982)

Solomon Islands Uncommon on most of the larger islands but can occur as
large populations on many smaller islands, such as the Olu Malau group (McCoy,
1980).

HABITAT AND ECOLOGY An active diurnal lizard, relatively large in size,
growing to around 1.5 m total length (50 cm body length). Olive-brown to near
black dorsally, patterned with small evenly arranged yellow spots. Mainly
terrestrial, but an able climber and fond of water. In northern Australia at
least (Bustard, 1970) said to rest on trees overhanging water into which it
dives when alarmed. Often found near the sea or in mangrove swamps; prefers
semi-open areas to dense forest and in many places frequents coconut
plantations.

In the Solomons, grapsoid crabs in the littoral zone form a major part of the
diet (McCoy, 1980); this may well be true elsewhere in the range. Also feeds
on small lizards and snakes, small mammals, sea turtle eggs, sometimes birds
and their eggs. A study on Guam (Dryden, 1965) showed that V. indicus there
fed mainly on animals often regarded as contrary to human interests, such as

193

Varanus indicus

the introduced Giant African Snail Achatina sp., rats, shrews and hermit
crabs. However, in Micronesia generally, indicus is persecuted for its
habit of feeding on chickens, land crabs and coconut crabs, all used as human
food (Uchida, 1966).

The eggs (five in a captive female) are laid under decaying ground vegetation,
or in rotting timber; hatchlings feed on insects and small scincid lizards.

THREATS TO SURVIVAL Available information does not suggest any confirmed
threats to the species, although data are sparse for much of the range.
Probably affected locally by persecution, in particular for its real or
alleged stock-raiding activities. Some Aboriginal or Torres Strait Island
communities may use the species for food (Australia CITES MA, 1986).

Table 1 Minimum net imports of live animals (L) and skins (S) of V. indicus
reported to CITES.

1980 1981 1982 1983 1984 1985
Australia - = 2L = = 2
Austria 150 Ss ~ = = = és
Belgium - 51S - = = =
Denmark Ze - - = = a
France 12295 S$ - = = = =
Japan - - - - = 4L
Switzerland 645 S 3014 Ss = = = =
UK = a = = 2 =
USA 2L 120 s 100 Ss - 3 =
TOTAL 4 L - 2L = = 4 L
13080 S 3185 Ss 100 Ss = = eS

Table 2 Reported countries of origin (or exporting country if no original
source reported) and quantities of transactions in live animals (L) and skins
(S) of V. indicus reported to CITES.

1980 1981 1982 1983 1984 1985

a. Countries having wild populations of V. indicus

Indonesia 2L - = = bs =
12590 s 120 S$ - = = &

Papua New Guinea - = = s ss 4L

b. Countries not having wild populations of V. indicus

Chad 645 Ss 3185 S$ 100 $s - - =
India 500 S$ 17 $s - - - -
Singapore - - 2 L - - -
Thailand 2 ib - - - - -

Varanus indicus

INTERNATIONAL TRADE CITES Annual Reports (Tables 1 and 2) show that there
has been very little international trade in V. indicus since 1980. In that
year there was an export from Italy to France of 12 440 skins of Indonesian
origin and another of 645 skins from France to Switzerland. Since then, most
of the trade reported to have been of this species has been declared to have
originated in Chad (Table 2), and was therefore probably misidentified. There
has been a negligible trade in live animals.

It seems unlikely that the species could be seriously affected by present
levels of exploitation, although local populations could perhaps be _ so
affected, and no population data are available for Indonesia, the source of
the majority of specimens reported to CITES. However, the population
information at hand does not provide an adequate basis for recommendations for
Management or for expanding trade volumes.

CONSERVATION MEASURES

Australia Export for commercial purposes is prohibited under the Wildlife
Protection (Regulation of Exports and Imports) Act (Australia CITES MA, 1986).

Indonesia Protected since 1978 (Indonesia CITES MA, 1986).

CAPTIVE BREEDING A recent inventory (Slavens, 1985) reports 10 individuals
in four collections in 1984, with breeding in one group.

REFERENCES

Bustard, R. (1970). Australian Lizards. Collins, Sydney and London, 162 pp.

Cogger, H.G. (1975). Reptiles and Amphibians of Australia. A.H. and A.W.
Reed Pty. Ltd., Sydney.

Cogger, H.G., Cameron, E.E. and Cogger, H.M. (1983). Amphibia and Reptila,
Vol. 1, in: Zoological Catalogue of Australia. Bureau of Flora and Fauna,
Australian Government Publishing Service, Canberra, 313 pp.

De Rooij, N. (1915). The Reptiles of the Indo-Australian Archipelago,
Vol.1, Lacertilia, Chelonia, Emydosauria. E.J. Brill Ltd., Leiden.
Reprinted 1970, A. Asher and Co. N.V. Vaals.

Dryden, G.L. (1965). The food and feeding habits of Varanus indicus on
Guam. Micronesia 2: 72-76.

Dryden, G.L. and Taylor, E.H. (1969). Reptiles from the Mariana and Caroline
Islands. Kansas University Scientific Bulletin 58(8): 269-279.

McCoy, M. (1980). Reptiles of the Solomon Islands. Wau Ecology Institute,
Handbook No. 7, 80 pp.

Mertens, R. (1942). Die Familie der Warane (Varanidae). Abhandlungen der
Senckenbergischen Naturforschenden Gesellschaft 466: 235-391.
Mertens, R. (1963). Helodermatidae, Varanidae, lLanthanotidae. In: Das

Tierreich 79, Walter de Gruyter and Co., Berlin.

Schmidt, K.P., and Inger, R.F. (1957). Living Reptiles of the World.
Doubleday and Co., Inc., N.Y., 287 pp.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity,
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

Uchida, T.A. (1966). Observations on the Monitor Lizard, Varanus indicus,
(Dardin) as a rat control agent on Ifaluak, Western Caroline Islands. WHO
Document WHO/EBL/66.64.

Whitaker, R., Whitaker, Z. and Mills, D. (1982). Reptiles of Papua New
Guinea. Wildlife in Papua New Guinea, No. 82/2. Department of Lands and
Environment, Konedobu, 53 pp.

195

NILE MONITOR Recommended list: 2
[Possible problem]

Varanus niloticus (Linnaeus, 1766)

Order SAURIA Family VARANIDAE

SUMMARY AND CONCLUSIONS A large diurnal lizard, occurring virtually
throughout sub-Saharan Africa (except the arid south-west). Has been recorded
in: Angola, Benin, Botswana, Cameroon, Central African Republic, Congo, Egypt,
Ethiopia, Equatorial Guinea, Gabon, Gambia, Ghana, Guinea, Guinea Bissau,
Ivory Coast, Kenya, Lesotho, Liberia, Malawi, Mali, Mauritania, Mozambique,
Namibia, Niger, Nigeria, Sao Tome and Principe, Senegal, Sierra Leone,
Somalia, South Africa, Sudan, Tanzania, Togo, Uganda, Zaire, Zambia, Zimbabwe.
Probably present in: Burkina, Burundi, Chad, Djibouti, Rwanda, Swaziland, and
possibly in’ the extreme south of Algeria. Typically found in association with
rivers, streams, lakes and other permanent water sources. Often common,
especially in protected areas and areas with sparse human activity, but
utilisation for food and skins is widespread and the latter in particular
poses a threat to local populations.

A substantial skin trade was reported to CITES, with an annual mean of 408 292
between 1980 and 1985. The skins were mostly imported to Europe, especially
France, Italy, Switzerland and F.R. Germany. Most originated in Mali,
Nigeria, Sudan and Cameroon. Relatively small numbers of live animals were
reported international trade, with an annual mean of 816 from 1980 to 1985.

While the species as a whole is not threatened, exploitation appears to be a
cause of local depletion. Field information on population status and trends
is required to help assess the long-term sustainability of the present high
trade levels.

DISTRIBUTION Very widespread in sub-Saharan Africa from Mauritania in the
west, eastward through Mali, Niger and Sudan, and south almost throughout the
continent to South Africa's Cape Province, but generally absent from desert
areas. Penetrates north of the Sahara into Egypt down the Nile valley. Has
been recorded in: Angola, Benin, Botswana, Cameroon, Central African Republic,
Congo, Egypt, Ethiopia, Equatorial Guinea, Gabon, Gambia, Ghana, Guinea,
Guinea Bissau, Ivory Coast, Kenya, Lesotho, Liberia, Malawi, Mali, Mauritania,
Mozambique, Namibia, Niger, Nigeria, Sao Tome and Principe, Senegal, Sierra
Leone, Somalia, South Africa, Sudan, Tanzania, Togo, Uganda, Zaire, Zambia,
Zimbabwe. Probably present in: Burkina, Burundi, Chad, Djibouti, Rwanda,
Swaziland, and possibly in the extreme south of Algeria.

Mertens (1942) recognised two subspecies, V.n. niloticus in the larger,
drier, part of the range, and V. n. ornatus in forest areas of West Africa,
its distribution partly interrupted on the Guinea coast by the nominate form.

Algeria Mertens (1942) gave a record from Adrar Mountain, which is in
southern Algeria; however as this is an extremely arid area, it is likely that
the term may refer to the Adrar region which extends into northern Mali along
the edge of the Niger drainage.

Angola Manacas (1955) listed localities from Angola, including Rio Quilo,
Cabinda, Luanda, Rio Cuando, Mayumba, Duque de Braganca, Dondo, Rio Cuanza,
Hanha, N'dala Tando, Golungo Alto, Kuvangu and Rio Mbalé. V.n. ornatus is
said to occur on the Loango coast in what is now Cabinda, while the nominate
subspecies occurs elsewhere in the country (Mertens, 1942).

196

Varanus niloticus

Benin Recorded from Agouagon and Haut-Dahomey (Chabanaud, 1917). Mertens
(1942) cited records of V.n. niloticus from Ajuda, Agouagon, and Haut
Dahomey.

Botswana Present along rivers and near permanent water, but absent from the
true Kalahari (Auerbach, 1985). V. mn. niloticus has been recorded at Maun
and along the Chobe River (Mertens, 1942).

Burkina No records have been located, but the species almost certainly
occurs in the country as it is listed in the national legislation and is found
in the surrounding countries, Mali, Ivory Coast, Ghana, Togo, Benin and Niger.

Burundi No records from Burundi have been located, but the species almost
certainly ~- occurs there, because Mertens (1942) gives records of
V. n. niloticus from Luvungi and Uvira in Zaire and Ujiji in Tanzania all
less than 20 km from the Burundi border.

Cameroon V. niloticus is said to occur in Kalamaloué, Boubandjida,
Benoué, Faro and Douala-Edéa protected areas (Tsague, 1986), all of which are
located in the north of the country, except for the last, which is on the
coast. Mertens (1942) listed many localities for V. n. ornatus.

Central African Republic Mertens (1942) quoted a record from Avakubi in
“Central Africa" which may represent the CAR and another from Fort Sibul in
"French Equatorial Africa.

Chad V. niloticus occurs in Kalamaloué in northern Cameroon (Tsague,
1986) and therefore almost certainly occurs in Lake Chad.

Congo V.n. ornatus is widespread (Congo CITES MA, 1986), occurring at
numerous localities, including Brazzaville and Niari (Mertens, 1942).

Djibouti No records have been located from Djibouti but the species occurs
in the Awash River of Ethiopia (Neumann, 1905), and may therefore extend down
to Lake Abbe on the Djibouti border.

Egypt V.n. niloticus occurs along the Nile as far north as Giza, just
upstream of Cairo (Flower, 1933).

Equatorial Giunea V.n. ornatus occurs on the island of Bioko and has
been recorded on the mainland at Cap Saint-Jean (Mertens, 1942).

Ethiopia Neumann (1905) noted the fisrt record of V. niloticus from the
river-less “Hauasch" region, which probably refers to the Awash. Other
records from Ethiopa are from the west, from the Blue Nile and the Akobo
Rivers.

Gabon Said to be distributed more or less uniformly throughout the country
(Gabon, Ministere des Eaux et Forets, in Jlitt., 29 November 1985).
V. mn. ornmatus has been recorded at numerous localities in Gabon, including
Setto Kama, and Lambaréné (Mertens, 1942).

Gambia V.n. niloticus occurs in Gambia, including on MacCarhty Island
(Mertens, 1942).

Ghana Mertens (1942) cited records of V. n. niloticus from the Goldcoast,
including Abruri, Akropong, Elima and Accra.

197

Varanus niloticus

Guinea V.m. ornatus is said to occur on the west coast of Guinea
(Mertens, 1942). Chabanaud (1921) said the species was widespread.

Guinea-Bissau Recorded from Brene and Bijimita, on Bissau Island; Formosa
and Bambadinca (Manacas, 1955).

Ivory Coast Recorded from Ivory Coast (Chabanaud, 1917). Occurs in Comoe
National Park (Anon., 1979)

Kenya Said to be present in all rivers and freshwater lakes in Kenya below
about 5000 feet (1500 m) (A.D. Mackay, in litt., 26 March 1986). Mertens
(1942) listed many localities for V. n. niloticus.

Lesotho The species is said to occur in the country (Lesotho Ministry of
Agriculture and Marketing, in litt., 23 January 1986).

Liberia Said to occur throughout the country (Liberia CITES MA, 1986).
Mertens (1942) cited records of V.n. niloticus from Monrovia, Lenga Town,
Farmington River, Gbanga and Ganta. V.n. ornatus has been recorded at Cape
Mount, Monrovia, Cavally River, Tala, Javillo, Mahfah River and Fisherman's
Lake (Mertens, 1942)

Malawi V.n. niloticus occurs at many places along the west shore of Lake
Malawi (Mertens, 1942), and along the Shire River (C. Dudley, in litt., 14
April 1986).

Mali vV.n. niloticus occurs in the Adrar Mountain Region in the north and
at Bourem (Mertens, 1942).

Mauritania Mertens (1942) cited Flower (1929) indicating that the
distribution of V. n. niloticus extended from Mauritania eastwards, though
no records from Mauritania were given. The species certainly occurs on the
south bank of the Senegal River at Bakel.

Mozambique V. niloticus said to be found throughout the country, having a
wider distribution than V. exanthematicus (Mozambique CITES MA, 1986).
Manacas (1955) listed localities from Mozambique, including Cabeceira, Boror,
Quelimane, Tete, Beira, Lumbo, Caia and Charre.

Namibia The species only occurs along the rivers bordering the north and
south of the country, and in the Caprivi Strip. The distribution extends to
the mouths of the rivers (Namibia Department of Agriculture and Nature
Conservation, in litt., 9 March 1987).

Niger V. niloticus occurs along the Niger River and around other
permanent water courses and lakes (Niger CITES MA, 1986).

Nigeria Associated particularly with the forests of southern Nigeria and
the riverain forest of the savanna zone; recorded from Lagos, Umuahia,
Claabar, Egbe, Mokwa, Makurdi, Jos, Bauchi and Malam Fatori (Dunger, 1967).
Mertens (1942) provided records of V.n. niloticus from Benoué and Lagos,
and of V. n. ornatus at Warri, Sapelli, Calabar and the Niger Delta.

Rwanda No records from Rwanda have been located, but the species almost
certainly occurs there, because Mertens (1942) gives records of
V. n. niloticus from Rutshuru in Zaire, less than 40 km from the Rwanda

border. He also cites a probable record from Lake Kiwu, half of which is in
Rwanda.

198

Varanus niloticus

Sao Tomé and Principe V.m. ornatus occurs on the island of Principe
(Mertens, 1942).

Senegal Numerous records of V.n. niloticus from Senegal were quoted by
Mertens (1942), extending up to Bakel on the Senegal River.

Sierra Leone Present in the Outamba-Kilimi and Mamunta-Mayoso areas
(Teleki, 1980). Mertens (1942) cited records of V.n. niloticus from
Freetown and the Grand-Galbar River.

Somalia Recorded from the Shebelle and Juba Rivers and the region of Ola
Uager (Simonetta and Magnoni, 1986).

South Africa In the Cape Province, the species extends along the south

coast no further west than the Gamtoos River valley. It also extends along
the Orange River to the Atlantic coast in the north (Branch and Braack,
1987). Widespread in Natal; also present in the Transvaal, Transkei and the

Orange Free State (Mertens, 1942).

Sudan V.n. niloticus was said to occur along the Nile in the provinces
of Halfa, Dongola, Berber and Khartoum, and up the Blue Nile as far south as
Roseires. It was not seen on the main White Nile, but was found on the Bahr
el Gebel and the Bahr el Zeraf as far south as Lado (Flower, 1933).

Swaziland No records have been located, but the species almost certainly
occurs as it is found in neighbouring parts of South Africa and Mozambique.

Tanzania Occurs on Zanzibar Island but not on Pemba (Pakenham, 1983), also
on Mafia Island and at numerous localities on the Mainland from the coast to
the western border (Mertens, 1942).

Togo V.n. ornatus occurs at several localities, including Bismarckburg
and Sebbe. V. n. niloticus was indicated at Tabligbo, Ancho-Bezirk, Sokodé,
Mangu, Bogu and Moba (Mertens, 1942).

Uganda The species is widely distributed along all river’ systems,
particularly those connected with the Nile, and all lakes (Uganda Game
Department, in Jlitt., 28 March 1987). Mertens (1942) cited records of
V.n. niloticus from Uganda, including the Sesse Islands, Entebbe, and
Murchison Falls.

Zaire Mertens (1942) cited records of V.n. niloticus from Gamangui,
Niangara, Niapu, Ngayu, Poko, Uvira, Luvungi, Eala, Koteli, Mahagi, Faradje,
Medje, Panga, Buta, Stanleyville, Leopoldville, Kikondja, Nyonga, Kansenia,
Kakyelo, Lukafa, Kasenga, Kando, Kiato, Kiambi, Leopoldville, Katobwe and
Ekibondo. V.n. ornatus appears to be confined to the more eastern regions,
including Banana, Loango, Kituri, Katanga, Mayumbé, Upoto, Maringa (Mertens,
1942).

Zambia Found wherever there are permanent rivers and streams (Broadley,
1971). Mertens (1942) cited records from the Zambezi and Luangwa rivers.

Zimbabwe Common in and around large rivers and lakes (D.G. Broadley, in

litt., 18 March 1986). Mertens (1942) gave records from the Zambezi and
Matabele Land.

199

Varanus niloticus

POPULATION Little information is available on population trends or current
status. In southern Africa generally, said to be very common in and around
large rivers and lakes, and not threatened except where exploitation for the
skin trade is allowed (D.G. Broadley, in litt., 18 March 1986).

Cameroon No surveys have been carried out, but the species is said not to
be in danger of extinction (Cameroon CITES MA, 1987).

Egypt V. niloticus was said to be fairly numerous in the Nile in the El
Derr district and around Abu Simbel, becoming rarer further downstream
(Flower, 1933).

Gabon Said to be more numerous along the coast where human predation is
less severe:(Gabon Ministere des Eaux et Forets, in litt., 29 November 1985).

Guinea Said to be very common throughout Guinea (Chabanaud, 1921).

Kenya Apparently still abundant at least around Lake Victoria (A.D. Mackay,
in litt., 26 March 1986).

Malawi The species is said to be very common around Lake Malawi and in the
Shire River (C. Dudley, in litt., 14 April 1986).

Mozambique V. niloticus said to be more abundant than V. exanthematicus
and to be especially common in the National Parks and Game Reserves
(Mozambique CITES MA, 1986).

Niger V. niloticus is said to be fairly common in suitable habitat (Niger
CITES MA, 1986).

Nigeria Common, formerly at least, in forests of southern Nigeria and in
the riverain forest of the savanna zone (Dunger, 1967).

Somalia Relatively abundant in Somalia (Fagotto, 1985).

South Africa Said to be common and widespread in Natal (South Africa CITES
MA, 1987).

Sudan The species was described as “not uncommon" on the main Nile in the
north, and “really numerous" on the Blue Nile between Wad Medani and Roseires
(Flower, 1933).

Tanzania Common in many parts of Tanzania; apparently not widely exploited
for food or skins (K. Howell, in litt., 15 March 1986). Not uncommon on
Zanzibar Island (Pakenham, 1983).

Uganda The species is said to be widespread, but particularly abundant in

the Bugosa and Buganda areas (Uganda Game Department, in litt., 28 March
1987).

Zambia Good populations occur in Zambia (Broadley, 1971).

Zimbabwe Very common in and around large rivers and lakes (D.G. Broadley,
in litt., 18 March 1986).

HABITAT AND ECOLOGY A large diurnal lizard, reaching up to 2 m in length.
Found only in association with permanent rivers, streams and lakes, and thus
absent from many arid regions. Occupies a variety of holes and crevices as a
burrow, usually on abandoned site of some other animal. Excellent swimmers;
will dive into water if alarmed. Diet includes a variety of small vertebrates

200

Varanus niloticus

and eggs (a noted consumer of crocodile eggs), also crabs and mussels and
readily scavenges food remains, such as fish guts and chicken bones, from the
vicinity of human habitation (Edroma and Ssali, 1983). The female often lays
the clutch of 40-60 eggs, ca 58 x 35 mm, in a live termite nest where the
termites maintain a thermally stable environment (Cowles, 1930; Branch and
Erasmus, 1982)

THREATS TO SURVIVAL Widely used, apparently throughout the continent, for
food and skins. In Nigeria, for example, numerous dried lizards can be seen
in Ibadan market, and shoes and handbags made from monitor lizard skin can be
obtained anywhere in the country (Butler and Shitu, 1985). The skin trade
poses a threat to some local populations (D.G. Broadley, in litt., 18 March
1986).

Cameroon Skins are supplied to the commercial trade from the north and
extreme north of the country (Cameroon CITES MA, 1987).

Congo V. niloticus is expoited for its skin and meat (Congo CITES MA,
1986).

Gabon The people of the coast apparently do not eat monitors, but those of
the interior are particularly fond of it, and may have depleted the
populations. Internal trade in skins is said to be negligible, and there is
not thought to have been any export in 1984 and 1985 (Gabon Ministere des Eaux
et Forets, in litt., 29 November 1985).

Kenya Often trapped as vermin around Lake Victoria due to poultry and egg
eating, though this seems not to have an adverse effect on the local
population (A.D. Mackay, in litt., 26 March 1986).

Mozambique There is said to be some human predation (Mozambique CITES MA,
1986), but its effects have not been assessed.

Namibia The species is not particularly threatened, and there is said to be
no commercial exploitation (Namibia Department of Agriculture and Nature
Conservation, in litt., 9 March 1987).

Tanzania Not threatened. There is not thought to be much skin trade and
few people eat monitors (K. Howell, in litt., 15 March 1986).

Uganda The skins are used by the Bantu tribes to cover their long,
cylindrical drums and substantial quantities are thought to be used each
year. There are no known authorised exports of live animals and few skins.
50 were exported in 1981, and a shipment of 176 skins bound for Lebanon was
intercepted at Entebbe airport in 1986 (Uganda Game Department, in litt., 28
March 1987). :

INTERNATIONAL TRADE While relatively small numbers of live animals (mostly
from Kenya, Nigeria and Togo) are in reported international trade, between 169
and 1392 from 1980 to 1985, with an annual mean of 816, very large numbers of
skins are traded, between 183 830 and 634 639 from 1980 to 1985, with an
annual mean of 408 292. Most live animals were destined for the USA, with
lesser numbers to F.R. Germany and other parts of Europe.

The skin trade reported to CITES is summarised in Tables 1 and 2. The skins
were mostly imported to Europe, especially France, Italy, Switzerland and F.R.
Germany. It should be noted that France did not report imports of Appendix II
material before 1984, and so the totals in Table 1 from 1980 to 1983 are
artificially low. Most of the skin originated in Mali, Nigeria, Sudan and
Cameroon.

201

Varanus niloticus

Table 1 Apparent minimum net imports of skins of Varanus niloticus reported
to CITES.

1980 1981 1982 1983 1984 1985
Australia 126 - 340 4917 - -
Austria 7478 6008 29918 14272 3749 1071
Belgium 735 54 415 5745 30000 -
Canada 79 1609 - 4 2 10
Cyprus 1474 - - - - -
Denmark : 39 60 - - - -
Egypt z . mtg = = 4
Finland - - 379 - - 3
France - - - - 82653 274829
Germany, F.R. 1592 8478 22595 13440 906 546
Greece - 132 - 50 - -
Hong Kong 18 - 1220 103 8 -
Italy 412046 464528 123355 129216 130341 52342
Japan 381 773 - - - 30
Kenya - - - - - 2
Korea - - - - 100 -
Kuwait - - - - 1 -
Lebanon - - 20 179 -
Macao - - - - - 1
Malaysia - - - - 396 -
Mexico 6000 1379 - - - -
Netherlands - 10 - - - -
New Zealand - - - 90 - -
Portugal - 1033 - - - -
Saudi Arabia - - - 78 8 -
South Africa - - 109 120 - 37
Singapore - - - - - 8
Spain 2135 - - - = =
Sweden - - - = = 8
Switzerland 88736 - 52630 61359 75144 60870
Togo - - = = = 3
Turkey - - - - 100 72
UAE 9 - - - 1
UK - - - 2420 - 53
USA 113791 58522 33553 20117 3678 5124
USSR = = Z = = 2
Uruguay - - 930 200 - -
Yugoslavia - - = = = 204
Country Unknown - 2 71305 - - 30000
Total 634639 542588 336390 183830 327086 425216

————— en ee _ eee eee

202

~_—

Varanus niloticus

Table 2 Reported countries of origin (or exporting country if no original
source reported) and quantities of transactions in skins of V. niloticus
reported to CITES.

1980 1981 1982 1983 1984 1985

a. Countries with populations of V. niloticus.

Africa 12 - 440 = = z
Benin - - - - - 6950
Cameroon T7525 21175 41505 2400 25508 67276
Chad : 7500 111 4574 12214 6008 -
Egypt - 2182 - - - -
Kenya - TAS) - - - -
Mali 89005 53771 80334 43556 63013 192213
Nigeria 367522 174773 82913 51979 52640 38779
South Africa 5578 3938 1 - 2 1
Sudan 44479 311302 126704 153502 190751 131461
Zimbabwe ~ 1 9 170 529 304
b. Countries without populations of V. niloticus.

China - - 2185 - - -
France 1390 1350 - - - -
Indonesia 1200 - - 1871 344 3
Italy 967 - - - - -
Madagascar 2829 - - - - -
Netherlands 40 - - - - ~
Papua New Guinea - - - 4 - -
Paraguay - - - - 900 -
Philippines - 13270 - 249 995 -
Spain - 11539 - - - -
Switzerland 1 - - 200 - 2000
Thailand - - - 4 - 140
UK - 119 - - - -
Unknown 205500 94970 85885 3565 2131 5390

CONSERVATION MEASURES Unless otherwise stated, all of the information on
protection is from IUCN Environmental Policy and Law Occasional Paper No. 3,
African Wildlife Laws.

Angola Not protected under the Hunting Regulations, 11 December 1955.

Benin Under the Decree concerning hunting and capturing licences, bag
limits and professional hunters, 11 February 1980, Varanidae are designated as
small game species. They may only be hunted by the holders of traditional
hunting rights or under hunting permits.

Botswana The hunting of Varanus spp. is controlled under the Fauna
Conservation (Unified Hunting) Regulations 12 March 1979, which set out the
conditions of issue of hunting licences and the number of animals which may be
taken by each hunter.

203

Varanus niloticus

Burkina V. exanthematicus and V. niloticus are partially protected
under the Wildlife Conservation and Hunting Act, 31 December 1968, which
establishes bag limits and close seasons, protects nests and eggs, and
regulates the sale of meat. Commercial capture and export are controlled
under the same Act by Decree No 68-314. Ordonnance No 68-58, December 1985,
fixed the open season from 15 December 1985 to 28 February 1986 and stipulated
that partially protected species could only be hunted under special permit and
only in the province of Gourma and Tapoa.

Burundi No information.

Cameroon Varanus niloticus and Varanus griseus (sic.) are partially
protected under the Forests, Wildlife and Fisheries Act, 27 November 1981.
They may only be taken under a valid hunting licence. No more than two

animals may be taken in one day.

Central African Republic The Ordinance concerning the protection of
wildlife and regulating hunting, 27 July 1984, designates Varanus as totally
protected.

Chad No information.

Congo The Act concerning the conservation and exploitation of wild fauna,
21 April 1983, vests in the State ownership of all wild animals of economic
value, and requires the issuing of licences for commercial capturing. A
minimum size is set for the taking of Varanus, and fees are set for the
issuing of certificates of lawful possession or export of monitor lizards.
Under a later Order (18 May 1984), Varanus flavescens (sic.) is designated
partially protected, and may therefore only be taken under a hunting licence.

Djibouti No information.
Egypt All hunting is prohibited in certain regions.
Equatorial Guinea The Hunting Regulations, 29 April 1953, establish the

need to obtain hunting licences except for subsistence hunters. Reptiles are
not protected.

Ethiopia Under the Wildlife Conservation (Amendment) Regulations, 12
February 1974, Varanus niloticus is protected. They may only be hunted for
scientific purposes and require a special permit, the value of which was set
at Eth$1.00 for each Varanus.

Gabon The Wildlife and Forests Act, 22 July 1982 requires the issuing of
licences for the commercial capture of all wildlife. Traditional hunting for
subsistence pruposes is permitted. :

Gambia Under the Wildlife Conservation Act, 14 February 1977, all wildlife
except game and vermin are protected.

Ghana The Wildlife Conservation Regulations, 4 March 1971, define Varanus
niloticus as totally protected.

Guinea No information.

Guinea-Bissau The Hunting Regulations, 12 May 1980, require the issuance of
a licence for the capturing of live wild animals.

204

Varanus niloticus

Ivory Coast Varanus niloticus, V. exanthematicus is listed as predatory
animals in the Wildlife and Hunting Act, 4 August 1965, and may be hunted
without licence anywhere except in protected areas. Arrété No. 15, 26
December 1972, establishes licence fees for the capturing of live reptiles.
Under the Order regulating the commercial hunting of crocodiles and monitor
lizards, 29 September 1967, a special permit is required to capture monitor
lizards for commercial purposes. The possession of specimens having a skin
width of less than 25 cm is prohibited.

Kenya Protected by legislation in Kenya, export being banned (Legal Notice
No. 152, 25 September 1981).

Lesotho Varanus spp. are totally protected under the Proclamation of
Monuments, Relics, Fauna and Flora, 1969.

Liberia The Wildlife Conservation Regulation (in draft, 1985) lists
V. niloticus as partially protected, thereby providing for closed seasons.
Trade in and export of fully and partially protected species is prohibited
except for educational or scientific purposes.

Malawi Hunting, possession, trade and export of game is controlled under
the Game Act, 1953. No reptiles, other than crocodiles, are protected.

Mali All Varanus species are classified as game species under the Hunting
Act, 11 November 1969. There is a general close season from 1 June to 31
October. A general hunting ban (Decree 325/PG-RM) throughout the country was
imposed on 6 November 1978.

Mauritania Under the Hunting and Wildlife Protection Act, 15 January 1975,
the capture, possession, sale and export of live wild animals requires a
commercial capture permit. The sale of game meat is prohibited.

Mozambique V. exanthematicus is fully protected by national legislation.
It occurs in Gorongosa National Park (Mozambique CITES MA, 1986).

Namibia Protected by the Nature Conservation Ordinance (Namibia Department
of Agriculture and Nature Conservation, in litt., 3 March 1987).

Niger All hunting for utilitarian purposes has been prohibited in Niger
since 1972 (Niger CITES MA, 1986).

Nigeria V. niloticus, and V. exanthematicus are listed as fully
protected in the Endangered Species (control of International Trade and
Traffic) Decree, 20 April 1985. Their hunting, capture, trade and export is
prohibited. On 28 May 1982, the CITES Secretariat sent a notification (No.
218) to all Parties requesting them not to accept imports from Nigeria as the
Nigerian Management Authority did not issue any export permits.

Rwanda Varanus niloticus is partially protected under the Hunting
Regulations, 31 December 1974. A licence is required for hunting and close
seasons may be set.

Sao Tomé and Principe No information.

Senegal The Game and Wildlife Protection Regulations, 30 May 1967, lay down
regulations governing the issuance of licences for hunting and commercial

capturing of wildlife.

Sierra Leone All Varanus are listed as prohibited animals in the Wildlife
Conservation Act, No. 27, 1972. It is illegal to hunt or possess them.

205

Varanus niloticus

Somalia Varanus are listed as game animals under the Law on Fauna
(Hunting) and Forest, 25 January 1969. They may only be hunted or exported
under licence. A ban on all hunting was instituted on 13 October 1977.

South Africa V. niloticus is partially protected in the Transvaal and may
not be captured or kept except under permit. They are protected by ordinance
in the Cape Province and occur in at least 58 of the 60 protected areas in
Natal (South Africa CITES MA, 1987).

Sudan V. exanthematicus and V. niloticus are listed in the Ordinance
for the Preservation of Wild Animals, 1935 (Amended 1974), as species for
which hunting is permitted. Licences are only required if firearms are used
for hunting. The Hides and Skins (Export) Regulations, 1969, specify a
grading system for the export of reptile skins.

Swaziland Apparently no reptiles are protected in Swaziland.

Tanzania Under the Wildlife Conservation Act, 30 June 1974, the capture of
all live animals requires a valid capture permit, and the hunting of all
animals requires a valid hunting permit.

Togo V. exanthematicus and V. niloticus are classified as pest species
under the Ordinance on Wildlife Protection and Hunting, 16 January 1968. An
enabling decree, dated 4 June 1980, sets out licence fees for hunting and
commercial capture, but there is no bag limit for these species.

Uganda No reptiles, other than crocodiles, are protected.

Zaire The commercial capture of unprotected animals requires a permit under
the Hunting Act, 28 May 1982. Export can only be authorised if the exporter
holds a certificate of lawful possession.

Zambia V. exanthematicus and V. niloticus are listed as game animals in
the National Parks and Wildlife Act, 3 December 1968, under which their
hunting, possession, trade and export are controlled and fees are set for the
export of trophies and live animals.

Zimbabwe The collection of live reptiles and their eggs and their breeding
in captivity are controlled under licence by the Parks and Wildlife (General)
Regulations, 1981.

CAPTIVE BREEDING A recent inventory lists 15 specimens in 9 collections,
but lists no captive breeding (Slavens, 1985). Breeding took place in the
Transvaal Snake Park in 1986, 33 being produced from a single clutch (South
Africa CITES MA, 1987).

REFERENCES

Anon. (1979). Btat actuel des parcs nationaux de la Comoe et de Tai ainsi
que de la Réserve d'Azagny et propositions visant a leur conservation et a
leur développement aux fins de promotion du tourisme. Tome II: Parc
National de la Comoe. Partie 1: Inventaire des conditions écologiques et
biologiques. GTZ, Eschborn.

Auerbach, R.D. (1985). The Reptiles of Gaborone. Botswana Book Centre.

Branch, W.R. and Braack, H.H. (1987). Reptiles and amphibians of the Addo
Elephant National Park. Koedoe 30: 61-111.

Branch, W.R., and Erasmus, H. (1982). Notes on reproduction in South African
Water Monitors Varanus niloticus niloticus (Sauria: Varanidae).
Journal of the Herpetological Association of Africa 28: 4-9.

Broadley, D.G. (1971). The Reptiles and Amphibians of Zambia. The Puku
6: 1-143.

206

Varanus niloticus

Butler, L.A., and Shitu, E. (1985). Uses of some reptiles by the Yoruba
people of Nigeria. Herpetological Review 16(1): 15-16.

Chabanaud, M.P. (1917). Enumération des reptiles non encore étudiés de
l'Afrique occidentale, appartenant aux collections du Muséum, avec la
description des espéces nouvelles. Bulletin du Muséum d'Histoire
Naturelle, Paris, 23: 83-105.

Chabanaud, P. (1921). Contribution a l'étude de la faune herpetologique de
l'Afrique Occidentale Bull. Com. Etudes Hist. Sci. Occ. franc. 445-472.

Cowles, R.B. (1930). The life history of Varanus niloticus Linnaeus as
observed in Natal, South Africa. Journal of Entomological Zoology
22: 1-31.

Dunger, G.T. (1967). The Lizards and Snakes of Nigeria (Part 3: the Monitors
and a Plated Lizard). WMNigerian Field 32: 170-178.

Edroma, E.L., and Ssali, W. (1983). Observations on the Nile Monitor lizard
(Varanus niloticus, L.) in Queen Elisabeth National Park, Uganda.
African Journal of Ecology 21: 197-201.

Fagotto, F. (1985). Larger animals of Somalia in 1984, Enivironmental
Conservation 12: 260-264.

Flower, S.S. (1929). List of vertebrated animals exhibited in the gardens of
the Zoological Society of London, 1828-1927, 3 Reptiles. London.

Flower, S.S. (1933). The recent reptiles and amphibians of Egypt, with a list
of the species recorded from that Kingdom. Proceedings of the Zoological
Society of London 1933: 735-851.

Manacas (1955). Saurios e ofidos de Guiné Portuguesa. Anais de Junta de
Investigacoes do Ultramar (Estudios de Zoologia) 10(4): 190-193.

Mertens, R. (1942). Die Familie der Warane (Varanidae). Abhandlungen der
Senckenbergischen Naturforschenden Gesellschaft 466: 235-391.

Mertens, R. (1963). Liste der rezenten Amphibien und Reptilien.
Helodermatidae, Varanidsae, Lanthanotidae. Das Tierreich 79: 1-29.
Walter de Gruyter, Berlin.

Neumann, O. (1905). Uber nordost-afrikanische und arabische Kriechtiere.
Zool. Jahrb. Syst. 22: 389-404.

Pakenham, R.H.W. (1983). The reptiles and amphibians of Zanzibar and Pemba
Islands. Journal of the Bast Africa Natural History Society and National
Museum 177: 1-40.

Simonetta, A.M. and Magnoni, M.L. (1986). Status and conservation problems of

Somali lower vertebrates. Rivista di Agricoltura Subtropicale e
Tropicale 80(3): 405-432.
Slavens, F.L. (1985). Inventory of live reptiles and amphibians in

captivity, current January 1, 1985. F.L. Slavens, Seattle, Washington.
Teleki, G. (1980). Status report on the Wildlife Survey Project, Sierra
Leone, conducted November 1979 through May 1980. WWF-US Project Report
(#1728).
Tsague, L. (1986). Contribution a l'étude des parcs nationaux et réserves
analogues du Cameroun. MSc dissertation, Dépt. de Biologie et Physiologie
Vegétales, Université de Yaoundé. .

207

WATER MONITOR Recommended list: 2
[Possible problem]
Varanus salvator (Laurenti, 1768)

Order SAURIA Family VARANIDAE

SUMMARY AND CONCLUSIONS A potentially very large monitor lizard, widespread
in South and Southeast Asia, from Sri Lanka east to southern China and the
Philippines, and south through mainland Southeast Asia to Indonesia.
Typically associated with water and thus frequently found alongside forest
waterways but also occurs in rice fields and coastal mangroves. Eggs, 6 to 30
in number, are often laid in live termite mounds. In the early twentieth
century generally said to be common or very common; although now reportedly
depleted or extirpated locally, the species appears to remain relatively
common over much of its range. Populations are affected by habitat loss,
although utilisation appears to be a more important factor.

Lizards and eggs are widely, but not universally, utilised for food and
medicinal purposes. Moderate numbers of live animals (around 1000-4500
annually from 1980 to 1985) are in international trade, but very large numbers
of skins are traded, with over a million in 1985.

Field information on population status and trends is required to help assess
the long-term sustainability of the present high trade levels.

DISTRIBUTION A widespread south and South East Asian species, ranging from
Sri Lanka and parts of India, eastward to southern China and the Philippines,
and south-east through the Malay Peninsula to Indonesia. Present in the
following countries, with details of distribution in each, if available.

Bangladesh Occurs in the Sunderbans and extends east and south to the
border with Burma, near St. Martin's Island (Khan, 1982).

Brunei Presumed present, no records located.

Burma Present, at least formerly, in suitable sites throughout the country
(Smith, 1930).

China Present in the south only, in Yunnan, Kwangtung, Kwangshi and Hainan
(Anon., 1977).

Hong Kong Despite being present in southern China, the Water Monitor was
apparently unknown in Hong Kong until four specimens were reported at
different times in 1961 (Romer, 1963); no further sightings have been reported
since, and the species probably no longer occurs there (UK (Hong Kong) CITES
MA, 1987).

India Restricted in distribution. Absent from the peninsula except for the
200 sq. km Bhitarkanika Wildlife Sanctuary (an island in the
Mahanadi-Baitarani delta) in Orissa, and coastal parts of west Bengal
(Sunderbans), also recorded (Smith, 1935) in extreme north-east India up to
6000 ft (1800 m). Present in the Andaman and Nicobar Islands (Biswas & Kar,
1981; Whitaker and Whitaker, 1980).

Indonesia Widespread, including Sumatra, Java, Kalimantan, with adjacent
smaller islands; seemingly more sporadic in the east, but extends to Sulawesi,
Halmahera and Obi, and also Bali, Lombok, Sumbawa and Flores (Mertens, 1963).

Kampuchea No data.

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Varanus salvator

Laos Occurs along the Mekong River valley, which extends the length of the
country in the west (Laos Wildlife Conservation and Fishery Office, in
Jitt., 31 January 1986).

Malaysia Few details available; in Sabah reportedly widespread at lower
altitudes wherever water is available (Malaysia (Sabah) CITES MA, 1985), also
widespread in Peninsular Malaysia (G. Davison, in litt., 22 February 1986)
and in Sarawak (H. Watson, in litt., 17 March 1986).

Philippines Widespread, including the larger islands of Luzon and Mindanao,
also Leyte, Cebu, possibly Samar, and probably numerous smaller islands
(Mertens, 1963).

Singapore Has been recorded (Ridley, 1899), no details available.

Sri Lanka Widespread, although seemingly absent from the east’ and
south-east coastal areas (Whitaker and Whitaker, 1980).

Thailand Occurs throughout the country, typically along rivers and streams
(Taylor, 1963; Lekagul, 1969).

Viet Nam No data.

POPULATION Despite, or perhaps because of, the wide distribution and
sometime abundance of this species there is virtually no detailed population
information presently available (W. Auffenberg has results of fieldwork in
preparation). The sole exception relates to Flores (Lesser Sundas, Indonesia)
where Auffenberg (1981; 1982) has recorded an estimated 15 individuals per km
of river bank (the Nanga Look), and 8 per km of coastal mangrove (see below
for note on habitat tolerance). Population density can thus be rather high
locally. Older literature sources typically remark on the wide distribution
and abundance of the Water Monitor, eg. “very numerous in suitable localities
throughout the Malay Peninsula and Siam" (Flower, 1896). However, although
the present situation seems to be unknown in detail, the species is now
reportedly reduced in density, or extirpated, in many places where it was
common in the early 1970s (Auffenberg, 1982; in litt., 1 March 1984).
Comments on general status, often anecdotal in nature, are given below where
available.

Bangladesh Common (Khan, 1982); common in coastal districts (Bangladesh
CITES MA, 1986).

Burma A 1930 source (Smith) states that the species is "quite plentiful” in
Burma in suitable localities. Often encountered on islands of the Mergui
Archipelago, also reported particularly plentiful around Mandalay where the
lizards live in canals and irrigation channels. Lizards of 1.8 m (6 ft) or
more in length were “frequently seen dashing across the Mandalay-Maymyo main
road in front of motors" (Smith, 1930). No recent data on status are
available (Salter, 1983).

China Reported rare in e 1922 source (Mell, cited by Smith, 1935), and said
to be apparently rare (Romer, 1963); no recent data available.

Hong Kong Probably no longer occurs in Hong Kong (UK (Hong Kong) CITES MA,
1987).

India Uncommon within Bhitarkanika, where the population is restricted to
the Wildlife Sanctury. Said to be reduced to remnants of former abundance in
the Sunderbans (Whitaker and Whitaker, 1980). Fairly common before WW II in
the Garo Hills of Assam (Parry, 1935). Population levels in the Andamans and

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Varanus salvator

Nicobars seem to be good away from human settlements, and in the Jarawa Tribal
Reserve, parts of Little Andaman and the southern islands of the Nicobar group
are “still intact and close to original in composition" (Whitaker and
Whitaker, 1980). Officially considered severely depleted and given Endangered
status (Tikader, 1983).

Indonesia Cited as very common by a 1915 source (De Rooij); no recent data
other than Auffenberg's, cited above.

Kampuchea Said to be common in the early twentieth century (Smith, 1935),
no recent data.

Laos Believed to be declining due to habitat loss and exploitation for food
(Laos Wildlife Conservation and Fishery Office, in litt., 31 January 1986).

Malaysia Said to be still quite common and readily seen (B.H. Kiew, in
litt., 25 February 1986) in Peninsular Malaysia, or still very common but
declining rapidly (S. Ambu, in litt., 17 February 1986). However, although
the species is not unusually seen along larger rivers here, assessing its
abundance is difficult, one factor being to distinguish salvator from other
Malaysian Varanus (G. Davison, in Jlitt., 22 February 1986). The call
(Khan, 1969) for improved management and legislation implies that exploitation
has adversely affected population levels.

Said to be abundant in suitable habitat (which is very widespread) in Sabah
(Malaysia (Sabah) CITES MA, 1985) and widespread and common in Sarawak, even
in areas with high human populations, such as the suburbs of Kuching (H.
Watson, in 1litt., 17 March 1986). However, also reportedly seen
infrequently in primary rainforest in Sarawak, where during 654 days of
herpetological sampling at three sites, a total of only six salvator were
recorded (R.F. Inger, in litt., 5 March 1986).

Philippines No information.
Singapore No information.

Sri Lanka Apparently still a stronghold for the species in South Asia;
Water Monitors are abundant and commonly seen scavenging in village rubbish or
swimming in streams despite the presence of people washing clothes or bathing
(Whitaker and Whitaker, 1980). This is attributed to the fact that the
species is little persecuted or exploited in the country.

Thailand Reportedly common along rivers and streams, also often encountered
in deep forest far from water (Lekagul, 1969). Seen fairly often in Khao Yai
National Park; perhaps not markedly threatened in Thailand (W.Y. Brockelman,
in litt., 14 February 1986). ;

Viet Nam Said at one time to be common in extreme south Viet Nam (‘Cochin
China’) (Smith, 1935). No recent data.

HABITAT AND ECOLOGY A large diurnal, carnivorous, terrestrial—aquatic
monitor lizard, capable of attaining about 2.5 m total length. In Burma, most
frequently seen near streams in remote evergreen forests, including on islands
(Smith, 1930). Here, and apparently throughout the range, the presence of
permanent surface water is the primary factor determining the occurrence of
V. salvator (Auffenberg, 1981; 1982). For example, the banks of the river
Nanga Look (Flores, Lesser Sundas) bear coastal mangrove swamp, dense gallery
forest, savanna, and agricultural land, at various points along its course,
but the species ranges through all riverine habitats regardless of vegetation
structure, and also occurs in areas of rice cultivation away from the river.

210

Varanus salvator

Individuals rarely move away from water and are most frequently observed
swimming in the river, basking on its banks, or foraging along a narrow linear
strip parallel to the river, its width dependent on the adjacent vegetation
(up to 50 m in dense forest but only a few metres in Open areas). However,
the lizards may move 0.5 km or more inland in the monsoons (Auffenberg, 1981)
and Smith (1930) records a large female being found in a new rain pool ina
remote and very dry forest area of Burma. The core areas of each adult's
active range are typically 500 m apart (Auffenberg, 1981).

Feeding is largely opportunistic. The diet is varied, larger individuals tend
to take larger prey. Food items include crabs, fishes, frogs, birds, small
mammals, eggs and all manner of carrion, also domestic chickens.

In monsoon ‘areas, nesting coincides with the start of the rains, in June in
India (Whitaker and Khan, 1982) and Thailand (Smith, 1935) for example. In
India at least, nest sites are located above the high watermark and often in
live termite mounds (Whitaker and Khan; Biswas and Kar, 1982); elsewhere
(Burma; Smith, 1930) nest sites are reported in holes in trees near or
overlying water. Eggs are about 40 x 70 mm; around 15-30 (Smith, 1935) or
6-20 (Biswas and Kar, 1981) in a clutch. Hatchlings from eggs laid in June
emerge in March-April, after about 260 days incubation, and are around
30-32 cm in length. In one captive breeding group, both sexes were 1.4-1.5 m
in length.

THREATS TO SURVIVAL Widely exploited for skins, meat and fat (for medicinal
purposes); semi-developed eggs in gravid females are considered a delicacy and
body fat is either consumed, to improve general vigour, or applied, for muscle
and joint pains (Whitaker and Whitaker, 1980).

India In India, also affected by loss of mangrove habitat to farming, and
by timber and firewood extraction; these are likely to be significant in other
parts of the range although no details are available. Hunting with dogs is a
particular threat in the Andamans and Nicobars; 30 monitors were seen to be
collected this way in one month (Whitaker and Whitaker, 1980).

Laos Adversely affected by habitat loss and disturbance, also exploited for
food, in Laos (Laos Wildlife Conservation and Fishery Office, in litt., 31
January 1986).

Malaysia Said to be little exploited in Sabah, and not exported (Malaysia
(Sabah) CITES MA, 1985). In Sarawak, hunting of this species is generally for
personal food use, not for the commercial skin trade (H. Watson, in litt.,
17 March 1986). Said to be only moderately exploited, for food and leather,
in West Malaysia (B.H. Kiew, in litt., 25 February 1986).

Philippines A permit for the hunting of 19 796 head of V. salvator was
granted retrospectively in 1981 to allow the sale of stockpiled skins
(Philippines CITES MA, 1981).

Sri Lanka Water Monitor flesh is believed poisonous in Sri Lanka and this
seems to be one reason why it is little persecuted.

INTERNATIONAL TRADE CITES Annual Reports show that there is a substantial
international trade in the skins of V. salvator, varying from 256 683 in
1980 to 1 152 532 in 1985 (Table 1). The main importing countries have been
the USA, France, Italy, Austria and Japan. Until 1984, France did not report
the imports of Appendix II species, and so it is possible that the totals for
earlier years are artificially low. The reported source countries (Table 2)
were Indonesia, Thailand, Malaysia, the Philippines and Singapore. As the
latter country is not thought to have a substantial wild population of

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Varanus salvator

Table 1 Minimum net imports of skins of V. salvator reported to CITES. The
few transactions reported in terms of weight, length or area are not listed.

1980 1981 1982 1983 1984 1985
Argentina - 250 - - - -
Austria 1589 - 9849 18832 53322 72533
Australia 3159 - 16 2124 - -
Belgium - - 1181 - - -
Brazil - - - - - 32921
Canada 13651 1783 1004 , 14604 37617 108979
Denmark : 654 17931 92 - 171 -
Egypt - - 1176 - - -
Finland - - 151 - - -
France - - - - 188472 285685
Gambia - - - - 8125 -
Germany, F.R. 3864 - 19480 14857 17649 16606
Greece - - 40 - - 213
Hong Kong 17279 - 22133 31733 33675 26211
Hungary - - - 200 - -
Ireland - - 2 - - -
Italy 9203 37958 75344 71863 133349 121659
Japan 85438 - - - - 340411
Korea, S. - - - 9607 29172 21797
Lebanon 70 - - - - -
Mexico - - - - - 177
Netherlands - 5058 - - 362 1224
New Zealand - - 82 43 83 -
Norway 20 - - - - -
Panama 1 - - - - -
Singapore - - - 203934 - -
South Africa 108 10 - 112 - -
Spain - - - 726 10056 -
Sweden 141 - 160 - 55 -
Switzerland 6750 — 6372 10222 8604 27329
Turkey - - 20 83 300 1036
Taiwan - - - - - 2696
UK 22820 - 8782 - - 23395
USA 107436 689668 71077 118954 167069 67467
Yugoslavia - - - - - 193
Unknown - 17796 33 1 1 2000
Total 256683 770454 216994 497895 610118 1152532

V. salvator, it is likely that the skins reported to have originated there
were re-exports, probably deriving in Indonesia or other neighbouring
countries. Up to half of the skins in trade in some years were reported to
have come from unknown sources.

There was also a smaller reported trade in live V. salvator. Minimum net
trade amounted to 2283 in 1980, 1238 in 1981, 321 in 1982, 1694 in 1983, 4577
in 1983 and 4686 in 1985. These numbers, although large, are probably
insignificant in terms of the species's survival. However, it seems that at
least local populations are likely to be adversely affected by the extensive
international trade in skins. Instances of local decline and extirpation are
reported by Auffenberg (1982; in litt., 1 March 1984), although no details

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Varanus salvator
are available. This suggests that present trade volumes may be excessive, and

reinforces the need for quantitative information on population status and
trends, currently lacking.

Table 2 Reported countries of origin (or exporting country if no original
source reported) and quantities of transactions in Varanus salvator reported
to CITES.

1980 1981 1982 1983 1984 1985

a. Countries having or possibly having wild populations of V. salvator

Bangladesh - - 20 2502 8225 525
Brunei - 1000 - =~ = -
China - 40000 600 - - -
India 148 6200 - 1001 847 17 kg
Indonesia 81051 148895 147228 360841 404490 626036
Malaysia 1071 17816 - 30348 25863 1391
Philippines 330 33653 22904 4478 3634 82870
Singapore 73707 247453 69825 43728 14420 307644
Thailand 54552 115988 20629 34550 210582 214074
Viet Nam - - - aS - -
‘Asia' 7548 3400 - 10 - -

b. Countries without wild populations of V. salvator

Australia - 2454 = = a =
Canada 4 3 4 = = x
Colombia - = = 1000 = ee
France 54 - = = = wS
Germany, F.R. 3401 - = = = =
Japan 1298 43910 600 8736 48 51758
Netherlands 4000 - - = a =
Nigeria 3 - - - 100 111
Paraguay - 250 - - = x
Spain = = - = = 1
Sudan 974 - - 3600 24 -
Switzerland - = = 24 33 =
Taiwan - 17885 570 - - =
UK 794 9415 - 30 - -
USA 677 - 1 421 178 431
Unknown 166349 151097 92471 110985 113078 920

De

CONSERVATION MEASURES
Bangladesh Nominally protected by legislation (Bangladesh CITES MA, 1986).

Hong Kong Protected under Hong Kong legislation (UK (Hong Kong) CITES MA,
1987).

India Nominally protected by legislation in India (Schedule I of the
Wildlife Protection Act).

Indonesia Hunting quotas for V. salvator have been set for 1987 totalling
350 000 animals, divided between the different regions as follows: Aceh

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Varanus salvator

(65 000), Riau (30 000), Sumut (16 000), Sumbar (17 000), Bengkulu (8500),
Jambi (8000), Sumsel (20 000), Lampung (5000), Jabar (9000), Kalbar (17 500),
DKI (18 000), Jateng (500), Jatim (600), Kalsel (36 400), Kalteng (20 000),
Kaltim (40 000), Sulsel (10 000), Sulteng (1000), Sultra (2500), Sulut (-),
NTT (1000), NTB (12 000), Timtim (1000), Maluku (1000), Irja (10 000)
(Indonesia CITES MA, 1987).

Malaysia In Peninsular Malaysia a law prohibits trapping of lizards up to
and including 16 inches length (S. Ambu, in Jitt., 17 February 1986); also
listed on the 1972 Wildlife Protection Act, however, trading licenses are
readily obtainable (B.H. Kiew, in litt., 25 February 1986).

Philippines Permits for the export of V. salvator were granted
retrospectively to allow the sale of stockpiled skins. These totalled 1400 in
1983, 4020 in 1984 and 248 in 1985 (Philippines CITES MA, 1981). They are
very substantially less than the number of skins which appear from CITES
Annual Reports to have originated in the Philippines (Table 2).

Singapore All wild fauna in Singapore are protected by legislation
(Singapore, Primary Production Department, in litt., 11 January, 1986).

CAPTIVE BREEDING A recent inventory (Slavens, 1985) reports 47 individuals
in 26 collections in 1984, with breeding in two.

REFERENCES

Anon. (1977). Sichuan Biology Research Institute (ed.). A systematic key to
the reptiles of China. Science Press, Peking.

Auffenberg, WwW. (1981). The Behaviour Ecology of the Komodo Monitor
University Presses of Florida, Gainesville.

Auffenberg, W. (1982). Catch a lizard, use a lizard. International
Wildlife 12(6): 16-19.

Biswas, S., and Kar, S. (1981). Some observations on nesting habitats and
biology of Varanus salvator (Laurenti) of Bhitarkanika Sanctuary,
Orissa. Journal of the Bombay Natural History Society 78(2): 303-308.

De Rooij, N. (1915). The Reptiles of the Indo-Australian Archipelago.
Vol. 1. Lacertilia, Chelonia, Emydosauria. E.J. Brill Ltd., Leiden.
Reprinted 1970, A. Asher and Co. N.V., Vaals.

Flower, S.S. (1899). Notes on a second collection of reptiles made in the
Malay Peninsula and Siam...with a list of the species recorded from those

countries. Proceedings of the Zoological Society of London 1899:
600-697.

Khan, M.A.R. (1982). Wildlife of Bangladesh. University of Dhaka, Dhaka,
174 pp.

Khan, Md.K.b.M. (1969). A preliminary study of the Water Monitor, Varanus
salvator. Malayan Nature Journal 22(2): 64-68.

Lekagul, B. (1969). Monitors (Varanus) of Thailand. Conservation News,
S.BE. Asia. No. 8: 31-32.
Mertens, R. (1963). Helodermatidae, Varanidae, Lanthanotidae. Das

Tierreich (Liste der rezenten Amphibien und Reptilien) 79: 1-26. Walter
de Gruyter and Co., Berlin.

Parry, N.E. (1935). Some notes on Water Monitors in the Garo Hills, Assam.
Journal of the Bombay Natural History Society 35(4): 903-905.

Ridley, H.N. (1899). The habits of Malay reptiles. Journal of the Straits
Branch of the Royal Asiatic Society 32: 185-210.

Romer, J.D. (1963). Occurrence of the Common Water Monitor Varanus salvator
(Laurenti), in Hong Kong. Memoirs of the Hong Kong Natural History
Society 6: 12.

Salter, R.E. (1983). Summary of currently available information on
internationally threatened wildlife species in Burma. FO: BUR/80/006.
Field Document 7/83. FAO, Rangoon.

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Varanus salvator

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in
captivity, current January 1, 1985. F.L. Slavens, Seattle, Washington,
341 pp.

Smith, H.C. (1930). The Monitor Lizards of Burma. Journal of the Bambay
Natural History Society 34: 365-373.

Smith, M.A. (1935). Reptilia and Amphibia, Vol. II, Sauria. In, The Fauna
of British India. Taylor and Francis Ltd., London. Reprinted 1973,
Ralph Curtis Books, Fla.

Taylor, E.H. (1963). The lizards of Thailand. Kansas University Science
Bulletin 44: 687-1077.

Tikader, B.K. (1983). Threatened animals of India. Zoological Survey of
India, Calcutta, 307 pp.

Whitaker, R. and Whitaker, Z. (1980). Distribution status of Varanus
salvator in India and Sri Lanka. Herpetological Review 11(3): 81-82.

Whitaker, R. and Khan, M.A.R. (1982). Monitor Lizards: identity and sexing
problems. Hamadryad 7(2): 8-10.

215

BOA CONSTRICTOR Recommended list: 2
[Possible problem]
Boa constrictor (Linnaeus, 1758)

Order SERPENTES Family BOIDAE

SUMMARY AND CONCLUSIONS Widely distributed from Central America to
Argentina. Largely arboreal, but found in a variety of habitats, from moist
forests to arid scrubland. The largest subspecies, B. constrictor
constrictor, grows to up to 5.6m long. It is chiefly nocturnal, and feeds
mainly on small vertebrates. Brood sizes are usually between 30 and 50. There
is little information on population levels, but what there is may be
conflicting. In some areas it is said to be seriously depleted, and in others,
abundant. It is locally encouraged and protected to control rodent pests.

CITES Annual Reports show that from 1980 to 1983 most of the trade in the
species was in skins, possibly up to 100 000 a year, but that this dropped
considerably in 1984 and 1985. There is also a demand for pets, and trade in
live animals declined from 20 000 in 1980 to 1830 in 1983 but increased to
18 418 in 1985, thereby accounting for more snakes than the skin trade. The
major suppliers of live animals are in Central America, particularly El
Salvador; Paraguay, Argentina and Panama supplied most of the skins.

Insufficient data are available to set sustainable harvest levels; but it
seems likely that the current levels are not excessive on a global scale,
although they may be locally. The major threat is from the skin trade,
partially because of the higher numbers which were formerly involved, and
partially because it is liable to use older animals than the pet trade, but
the apparently increasing trade in live animals should be closely monitored.
The species is protected in all of the major countries that supply skins, and
imports from these countries should not be permitted. This requirement is
clearly being ignored. The major importing countries are the USA, France and
Italy. In 1987, B. c. occidentalis was transferred to Appendix I of CITES.

DISTRIBUTION Widespread in Central and South America from Mexico to
northern Argentina, including selected Caribbean islands. The species has been
split into several subspecies, which are morphologically distinct.

Boa constrictor imperator Daudin 1803:
Belize Present (Groombridge, 1983).

Colombia West of the Andes on the mainland (Peters and Orejas-Miranda,
1970), and including the islands of Providencia, San Andrés, Santa Catalina
(Schwartz and Thomas, 1975), Uraba and Gorgona. It is thought to have been
introduced to the islands of Providencia and San Andrés to control rodent
pests in the plantations around 1930 (Gallego, 1978). The few records from
Isla Gorgona are thought to indicate sporadic passive migration, and that
there is no established breeding population (Medem, 1979).

Costa Rica The highest altitude record for the country was said to be 914 m
(Pope, 1962).

Ecuador West of the Andes (Peters and Orejas-Miranda, 1970; Miyata, 1982).
A new subspecies, B. constrictor melanogaster, has recently been described

from Ecuador (Langhammer, 1983).

El Salvador The highest altitude record for the country was said to be
650 m (Pope, 1962).

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Boa constrictor

Guatemala Present (Groombridge, 1983).

Honduras Recorded from scattered locations throughout the mainland and
including Las Islas de la Bahia (Wilson and Meyer, 1982).

Mexico Recorded from the states of Baja California, Campeche, Chiapas,
Colima, Durango, Guerrero, Jalisco, Michoacan, Morelos, Nayarit, Oaxaca,
Puebla, Queretaro, Quintana Roo, San Luis Potosi, Sinaloa, Sonora, Tabasco,
Tamaulipas, Veracruz, Yucatan and Zacatecas (Smith and Smith, 1976). The most
northerly record is in Sonora, only 120 km from the Arizona border. There are
occasional, unverified reports from Arizona, but these are thought to
represent escapes (Baltosser, 1982). The highest altitude record for Mexico
was said to be 792 m (Pope, 1962).

Nicaragua Present (Groombridge, 1983).

Panama B. constrictor imperator occurs on the mainland in mountainous
areas, chiefly in the provinces of Chiriqui, Bocas del Toro, Panama and Darien
(Panama CITES MA, 1985). A seperate subspecies, B. constrictor sabogae, is
said to be confined to Taboga Island (Peters and Orejas-Miranda, 1970).

Peru West of the Andes (Peters and Orejas-Miranda, 1970); a seperate
subspecies, B. constrictor ortoni, is said to be confined to a region in the
North-West (Peters and Orejas-Miranda, 1970).

Boa constrictor constrictor Linnaeus, 1758:

Argentina Moist forests in the North (Freiberg, 1982).

Bolivia Widespread in the Amazonian region (Hoogmoed, 1979).

Brazil Widespread in the Amazonian region (Hoogmoed, 1979), and northern
forests, but absent from the Atlantic forests (Dixon, 1979).

Colombia East of the Andes (Peters and Orejas-Miranda, 1970), widespread in
the Amazonian region (Hoogmoed, 1979).

Ecuador East of the Andes (Peters and Orejas-Miranda, 1970; Miyata, 1982),
widespread in the Amazonian region (Hoogmoed, 1979).

French Guiana Widespread (Gasc and Rodrigues, 1980).

Guyana Present (Hoogmoed, 1982b).

Paraguay Found in moist forests (Freiberg, 1982).

Peru East of the Andes (Peters and Orejas-Miranda, 1970), widespread in the
Amazonian region (Hoogmoed, 1979). Recorded from Iquitos, Moropon, Mishana
and Pebas (Dixon and Soini, 1986).

Suriname Present (Hoogmoed, 1982b).

Trinidad and Tobago Recorded from Trinidad, where it is widely distributed
(Emsley, 1977), and Tobago (Emsley, 1977; Hardy, 1982).

Venezuela Widespread throughout most of the country below 1000 m (Roze,
1966).

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Boa constrictor
Boa constrictor amarali Stull, 1932:
Brazil South and south-west regions (Freiberg, 1982).

Bolivia Said to occur in south-east regions (Freiberg, 1982), although this
is given as the possible range of B. c. occidentalis (Anon., 1987).

Boa constrictor occidentalis Philippi, 1873:

Argentina Arid sub-tropical forests in the north and west (Freiberg, 1982;
Peters and Orejas-Miranda, 1970), in Salta, Jujuy, Chaco, Formosa, Catamarca,
La Rioja, Tucuman, Santiago del Estero, Cordoba, Santa Fe, San Luis, San Juan,
Mendoza and: La Pampa, possibly occuring in the west of Buenos Aires (Gallardo,
1977; Anon., 1987). The area of distribution is said to be declining
(Argentina CITES MA, 1987).

Bolivia This subspecies may occur in the extreme south-east adjacent to the
Argentinian and Paraguayan borders, but there is no accurate information
(Anon., 1987).

Paraguay Arid sub-tropical forests (Freiberg, 1982; Peters and
Orejas-Miranda, 1970) in the western part of the country (Anon., 1987).

Uruguay A map given by Hoogmoed (1982b) includes Uruguay in the range of
B. constrictor, but this must be considered to be erroneous.
Boa constrictor ortoni Cope, 1878:

Peru Confined to a region in the north-west (Peters and Orejas-Miranda,
1970).

Boa constrictor orophias Linaeus, 1758:

St Lucia Widespread on the main island, but absent from the Maria Islands
(Lazell, 1964; Schwartz and Thomas, 1975; Corke, 1983).

Boa constrictor nebulosa (Lazell, 1964):

Dominica Present (Schwartz and Thomas, 1975); widely distributed, observed
at La Chaudiére and River Canari (reported as 8B. constrictor orophias by
Swank and Julien, 1975).

Boa constrictor sabogae:

Panama Confined to Taboga Island (Peters and Orejas-Miranda, 1970).

POPULATION

Argentina The species is said to have become restricted in distribution
(Gallardo, 1977) and to have declined in numbers alarmingly in the country
(Tarak, A., in litt., 1 June 1981). The status in the provinces has been
summarised as follows: Cordoba, scarce; Tucuman, potentially endangered;
Mendoza, virtually extinct although formerly abundant; San Juan, major
population reduction; La Pampa, extremely rare; San Luis, threatened and
declining; Salta, declining and in need of protection; Catamarca, considerably

218

Boa constrictor
reduced from former abundance owing to hunting; La Rioja, vulnerable; Chaco,
Formosa and Santiago del Estero, relatively common in the extensive forests
(Anon., 1987). 8B. constrictor occidentalis was said to be already extinct
in the Reserva Ecologica Nacunan in Mendoza (Abraham de Vazquez and Wuilloud,
undated).

Belize No information.

Bolivia No information.

Brazil Said to be extremly common, especially around towns (W.E. Magnusson,
pers. comm., 1986).

Colombia No information.
Costa Rica No information.

Dominica Said to be common and widely distributed in 1975 (Swank and
Julien, 1975)

Ecuador No information.

El Salvador Included in the list of species "threatened with extinction" in
1978, and said to be in need of a 10-year ban on hunting to aid recovery
(Serrano, 1978).

French Guiana Said to be widespread (Gasc and Rodrigues, 1980).

Guatemala No information.

Guyana No information.

Honduras The population size is unknown, but it is thought to be very rare
(Aguilar, W., in litt. 29 November 1985). Distribution records indicate that
it is widespread within the country (Wilson and Meyer, 1982).

Panama Moderately common in mountainous areas (Panama CITES MA, 1985).
Paraguay There is no information for the whole country, but
B. constrictor was said to be abundant in 1978 in the Parque Nacional
Defensores del Chaco (Torres Santibanez, 1978).

Peru Described as common in the Iquitos Rgion (Dixon and Soini, 1986).
B. constrictor is listed as a “vulnerable species" in Peruvian legislation

(Fuller and Swift, 1985).

St Lucia Probably widespread, but present status unknown, thought to be
vulnerable (Corke, 1983).

Suriname Said to occur in reasonable numbers (Hoogmoed, 1982a).

Trinidad and Tobago Widely distributed (Emsley, 1977).

Venezuela Populations are said to have decreased (Venezuela CITES MA,
1987), although Gorzula (in litt., 11 April 1986) considered that they could
still be categorised as frequent.

HABITAT AND ECOLOGY A medium or large boid snake, B. constrictor commonly
grows to a length of 2-3 m (Freiberg, 1982), although may exceptionally reach
5.6 m (Stidworthy, 1969). Females generally grow larger than males (Stafford,

219

Boa constrictor

1979). B. constrictor constrictor is the largest subspecies, while
B. constrictor amarali is usually little over 1m in length (Freiberg,
1982). B. constrictor occidentalis seldom exceeds 2.2 m (Gallardo, 1977).

It is largely arboreal, commonly found in moist forests, along water-courses
and in coastal regions, although it is rarely seen in water. It may also live
in arid areas. It is chiefly nocturnal, taking refuge during the day in hollow
trees, crevices in rocks or under roots, or the burrows of mammals, especially
those of vizcachas in Argentina (Pope, 1962; Emsley, 1977; Flores Villela,
1980; Freiberg, 1982).

The colouring is very variable: B. constrictor imperator is largely dark,
with obscure cross-bars, while B. constrictor constrictor, the Red-tailed
Boa, has handsome reddish markings, particularly on the tail (Breen, 1974).

The growth rate is not known, although the greatest longevity recorded in
captivity was 40 years, 3 months (Engelmann and Obst, 1984). Pope (1962)
presents data on the growth in captivity of several individuals. The fastest
incremant in length was a female which grew from 51 cm to 269 cm in two years.
Another animal, 157 cm long, only grew to 165 cm in a little over a year. The
relationship between weight and length is exhibited by a snake which weighed
348 g at a length of 91 cm, 747 g at 119 cm, and 1992 g at 180 cm (Pope, 1962).

B. constrictor kills its prey by constriction. It feeds chiefly on small
mammals, especially rodents, agoutis, pacas, squirrels, bats and monkeys, but
it may also take birds and other reptiles, such as iguanas, tegus and other
snakes. Occasionally larger prey, such as ocelot, or even porcupine may be
taken (Emsley, 1977; Flores Villela, 1980). It is particularly useful in
agricultural regions in keeping rodent pests under control (Pope, 1962; Mena
Moya, 1978). Pope (1962) gives some data on the amount of food eaten by boas
in captivity: one young animal ate 104 mice, 22 hamsters, 5 guinea pigs and 5
sparrows over a 26-month period. The passage time can be as little as 7-9
days, but digestion can take longer, depending on temperature (Engelmann and
Obst, 1984).

Like all New World boids, it is ovoviviparous, commonly having broods of
30-50, the young emerging at a length of around 50 cm (Freiberg, 1982). The
largest brood recorded was 63 (Emsley, 1977). A brood of 59 B. constrictor
constrictor, in a private collection, averaged 14 inches (36 cm) in length
(Slavens, 1985).

The age at maturity in the wild is not known, but a captive animal started
sexual activity at an age of three years (Pope, 1962).

The gestation period is said to be 119-295 days (Engelmann and Obst, 1984). A
pair was seen copulating during the month of April in Mexico (Flores Villela,
1980), and in captivity it is usual for broods to emerge in May (Breen, 1974).
In Trinidad it is usual for copulation to occur in the dry season (January to
May), the young being released from May to September (Emsley, 1977). A pair in
a private collection were observed copulating six times between 15 February
and June 1985, the young being born on 7 August 1985 (Slavens, 1985).

THREATS TO SURVIVAL B. constrictor is hunted chiefly for its. skin,
although its flesh is apparently good, and there are reports of its having
been used as food by pre-Columbian peoples (Flores Villela, 1980), although
Pope (1962) considers that the consumption of snakes was not widespread in
South America. More recently, a Belgian restaurant is reported to have
featured Boa Constrictor on its menu (Chapman, 1982). Live animals are
extensively traded as pets. The pet trade relies primarily on young animals,

220

Boa constrictor

and differs in this respect from the skin trade, for which larger animals are
preferred. It is therefore likely to have less impact on exploited
populations. A pet supplier in the USA (South American Unlimited, New York)
listed "Guyana Redtail Boas" at US$100 each on its price list in 1985. There
is very little detailed information regarding exploitation.

Argentina Hunting for skins is thought to have caused a severe reduction in
the population of 8B. constrictor in the country (Tarak, A., in Jlitt., 1

June 1981). Habitat destruction is also implicated in some areas,
particularly Tucuman, East Chaco and Formosa, where the preferred habitat of
chaquena forest has been destroyed. Vizcacha are sometimes eradicated in

their burrows by burning or toxic gas, and this has a catastrophic effect on
the Boas, which regularly use the burrows. There is some hunting for meat and
for the fat which is supposed to have medicinal properties, but both are of
minor significance compared with the hunting for skins. The main domestic
market was in Buenos Aires, but internal trade was made illegal in 1986
(Anon., 1987).

Brazil In July 1984, IBDF (Instituto Brasileiro de Desenvolvimento
Florestal) seized a shipment of 140 tanned snake skins (B. constrictor and
Bunectes murinus) in Porto Alegre. The skins were en route from Belem to Sao
Leopoldo and were thought to be destined for export once they had been
manufactured into shoes and other products (3.T. Palazzo, in litt., 1984).

Wettenberg et al. (1976) conducted a survey of restaurants in Manaus to find
out which species of wildlife would be preferred as food by the public. At the
time of the survey none of the restaurants had any wildlife on their menus, as
sales of wildlife products were illegal. “Giboia" (Boa constrictor) was
mentioned by only one restaurant.

Panama Trade in wildlife in Panama has been giving cause for concern since
at least 1978, when “small boas" were amongst the species being traded. At
that time there were ten companies involved in the import, export and
re-export of wildlife products. Legislation did not control the shipment of
goods in transit through Panama, and companies were said to import animals
from overseas without adequate documentation, and then to re-export them.
Occasionally additional animals, caught within Panama, would be included in
the re-export consignment without being declared (Vallester, 1978).

Paraguay Figures compiled by the Ministerio de Agricultura y Ganaderia
(Acevedo Gomez, 1987) showed that some 2000 skins of 8B. constrictor were
exported from Paraguay in 1984, and that in 1986, 3114 pairs of shoes made
from the skin of this species were exported to the USA. It was estimated that
this accounted for the skins of 623 snakes, on the basis that each pair of
shoes needed 35-40 cm of skin and that there were an average of 2 m of useful
skin on each snake. Domestic trade was thought to exert a lesser effect on
reptile populations.

St Lucia Since B. constrictor was declared protected, in 1980, the only
recorded illegal trade was when an English tourist attempted to smuggle out
five live boas. He was subsequently deported (St Lucia CITES MA, 1985).

Suriname Hoogmoed (1982a) writes that "there is no hunting for hides, nor
is the natural habitat being destroyed. So, at least in Suriname, this species
seems to be safe".

Venezuela There is said to be some persecution by farmers but no organised
exploitation for the skin trade (Venezuela CITES MA, 1987). Gorzula (in
litt., 11 April 1986) considered that there might be some local trade as pets
or for skins as curiosities.

221

Boa constrictor

INTERNATIONAL TRADE The only reports of international trade in
B. constrictor are those contained in the Annual Reports of Parties to
CITES. Some transactions were identified to the level of subspecies, the
majority of these being 8. constrictor constrictor, with only small
quantities of B. constrictor imperator, but for the purposes of the
following analysis all records for the species were lumped together. Only
trade in live animals and skins was considered. The CITES reports are
summarised in Tables 1 and 2. Trade in skins was sometimes recorded by
length. It is notoriously difficult to convert this into numbers of skins, as
it depends on the size of the snakes killed and on the method of preparation
of the skin, however an average skin length of 1.4m has been suggested
(Anon., 1984). Acevedo Gomez (1987) considered that the average length of
skins from Paraguay was 2.5 m, of which 2m was usable skin. This may be
excessive if the subspecies in trade is B. constrictor amaralis, which is
usually nearer 1m in length.

Table la shows that the annual trade in skins of B. constrictor may have
reached nearly 125 000 in 1983, depending on the length conversion factor, and
that, if anything, the volume of trade increased from 1980 to 1983, although
the figures for 1984 and 1985 are markedly lower again. The chief net
importing countries were the USA, Italy, Mexico, F.R. Germany, Spain, France
and the UK.

The numbers of live animals in trade (Table 1b) were much lower, and have
declined from 21 735 in 1980 to 1830 in 1983, but then increased sharply again
to 18 418 in 1985. Between 70% and 92% of these were imported to the USA. It
is not known whether the temporary decline in the level of this trade was due
to a decline in demand for pet boas, a shortage of supply or stricter
controls. In 1985 the trade in live animals exceeded the trade in skins.

Table 2a shows that the great majority of the skins were declared as having
originated in Paraguay, although Argentina was the major source in 1984, with
large quantities also originating in Colombia, Guyana, Peru and Suriname. In
1983, Panama emerged as the source of about 25 000 m of skins.

The majority of live snakes originated in Central America, Colombia and
Suriname (Table 2b), which probably reflects the ease of supply to the main
market in the USA. The dramatic increase in the volume of trade in 1985 was
almost entirely attributable to snakes originating in El Salvador. The
countries of origin indicate that the main subspecies used in the pet trade is
B. constrictor imperator which is supposedly less suitable in temperament as
a pet than 8B. constrictor constrictor (Breen, 1974). Most of the _ skins,
where subspecies was declared, were of B. constrictor constrictor. Of the
countries with no wild populations of B. constrictor the majority probably
represent re-exports where the country of origin was not specified, or
possibly captive—bred snakes not specified as such.

222

Boa constrictor

Table la. Minimum net commercial imports of B. constrictor skins reported to
CITES.

1980 1981 1982 1983 1984 1985
Andorra - - = 50 = =
Austria - - 20 7 = 8
Belgium 3444 - 4 = 7m 2
Canada 1912 1972 150 167 31 1
211m 130 m 18 m = = eZ
Chile - 20 2 154 40 -
Denmark 46 - 95 = = =
Finland = = = s & 16m
France 108 616 393 - 3887 1050
- 10 m - = = =
German D.R. - - - 13 m - -
Germany, F.R. 7964 - 174 4185 95 m -
Hong Kong = - 44 = = =
Israel 43 47 - ~ 110 124 m
Italy 5550 18652 18676 17324 3740 6012
24064 m 547 m 8789 m - 7763 m -
= 750 m@ = 4 2 eZ
Japan ~ = = 76 2 Ss
Kuwait - = = = 2 1
Lebanon - = - = 67 m =
Mexico 26 5192 9576 5838 400 3
Netherlands 332 - - = =
New Zealand 14 - - - 105 -
Peru 6908 - = = = 2
Saudi Arabia - - 40 - - 14
Singapore - - - - 21 -
South Africa ~ 12 - - - -
Spain 3315 - 2216 7233 3924 -
— = - 2455 m - -
Sweden ~ - - - 2 -
Switzerland 484 617 102 121 10 11
- 5m - 4m = =
Turkey - - - 20 - 2500
UK - 5000 m 5000 m 17 - -
Uruguay - 3246 - 758 359 -
USA 19156 53643 135 1695 10302 4711
- 10971 m 20464 m 84865 m 698 m 800 m
= - 3290 kg - - 105 kg
Venezuela 18 319 119 - - 5
Unknown - - 167 - - -
Total 49320 84336 31913 37645 22931 14316
24275 m 16663 m 34271 m 87337 m 8630 m 940 m
= 750 m2 3290 kg 2 = 105 kg

Dee eS

223

Boa constrictor

Table 1b Minimum net commercial imports of live B. constrictor reported to
CITES.

1980 1981 1982 1983 1984 1985
Austria 203 8 3 - 63 41
Belgium - - 2 - 6 -
Canada 272 92 9 24 149 267
Colombia - - - - - 1
Czechoslovakia 9 1 1 - - -
Denmark 4 4 3 - 54 12
Finland : - - 4 - - -
France - 5 - 12 8 10
German D.R. 165 - - - - 1
Germany, F.R. 1764 356 373 117 1877 892
Greece - - - 2 1 a
Honduras - - 1 - - -
Indonesia - - - 3 - 5
Ireland - 1 - - - -
Israel - - - - 7 2
Italy 3 46 41 1 176 -
Jamaica - - - - 1 -
Japan 4 22 24 16 24 29
Korea, Rep of - - 1 - - -
Liberia - 3 - - - -
Malaysia 2 - - - - -
Martinique - - - 1 -
Mexico - 1 - - - -
Netherlands 26 3 - 9 13 192
Poland - - 16 - - -
Romania 2 - - - - -
Saudi Arabia - = 4 2 - -
Singapore - - - 2 - 2
South Africa - - 2 - 2 -
Spain - - 8 13 1 15
Sri Lanka 2 - - - - -
Sweden - 4 - - - -
Switzerland 381 78 87 - 15 -
Tunisia - - - - 1 -
UAE - - 4 - - -
UK 327 370 141 39 239 173
USA 18206 12794 3605 1589 6285 16774
USSR 2 2 9 - - -1
Unknown 3 64 = 1 = =
Uruguay - - - - 1
Total 21375 13854 4338 1830 8923 18418

Table 2a.

B. constrictor skins reported to CITES.
——————_— SS ee eee eee eee

1980

1981

Countries with wild populations of B. constrictor

Argentina

Bolivia
Brazil

Colombia

Costa Rica
Ecuador
Guyana
Honduras
Mexico
Nicaragua
Panama

Paraguay

Peru

Suriname

5174
3154 m

23543
21121 m

Trinidad & Tobago -

Countries without wild

Guadeloupe
India
Indonesia
Italy
Nigeria
Singapore

South Africa

Spain
Thailand
UK

USA
Uruguay
Unknown

11613
1674 m

1

104

5175

populations

250 m

2592 259
141 kg 1628
58 -
4126 270
- 5281

5 a

7 2570

at 1

- 1
9885 1585
6764 m 24825
16207 28313
14491 m 89057
9 kg -
3140 kg -
1986 m -
434 =
1915 m 200
of B. constrictor
2510 -
24 -

36 -
10991 m 6
2526 5461
110 m 5206

1 kg -

m

m

Boa constrictor

125
4521 m

Reported countries of origin or export for commercial exports of

225

Boa constrictor
Table 2b Reported countries of origin or export for commercial exports of
live B. constrictor reported to CITES.

1980 1981 1982 1983 1984 1985

Countries with wild populations of B. constrictor

Argentina ~ - - 2 7 52
Belize - 2 1 - - -
Bolivia 86 108 - 7 - -
Brazil - ak - - - 6
Colombia ; 1215 11299 443 - 110 -
Costa Rica - 2 1 - 2 -
Ecuador - 1 6 - - -
El Salvador 124 11 280 1247 1759 14389
Fr. Guiana ~ 2 3 - - -
Guatemala 15330 61 14 75 6327 1521
Guyana 130 354 154 227 247 429
Honduras 131 le, 1 - 290 348
Mexico - 4 94 50 25 -
Nicaragua = ~ - - - 1
Panama 1374 379 3174 - - 500
Paraguay 1802 34 - - 4 1064
St Lucia - - - 1 12 -
Suriname 3 1585 155 123 114 77
Trinidad & Tobago —- 1 - - - 1
Venezuela - 2 1 - 4 -

Countries without wild populations of B. constrictor

Australia - - - - 30 -
Austria - - - 6 2 3
Canada 6 4 8 2 20 4
Czechoslovakia - 1 - - - -
Denmark - - 1 - 1 -
France - 2 1 - - -
German D.R. - - 18 35 112 1
Germany, F.R. 145 - 1 2 16 1
Haiti = 3 - - - 9
Martinique - - - - 1 -
Netherlands - - 9 - - 1
Nigeria - - - 1 - -
Switzerland 2 11 - 89 6 48
Togo - - - 1 - -
UK 2 - - - 1 -
USA 304 109 11 19 65 127
Unknown 1312 532 108 20 4 3

226

Boa constrictor

CONSERVATION MEASURES The legal protection status of 8B. constrictor in
various countries throughout its range is summarised in Table 3. On the basis
of this information it appears that the species is protected in most of the
major supplying countries: Paraguay, Argentina, Colombia, Peru and Panama.
Furthermore the quantities of skins reported as originating in Paraguay are so
large that it seems likely that they did not originate in the country but were
smuggled over the border from Brazil.

Table 3. Legal prohibition on the commercial hunting, internal trade and
commercial export of 2B. constrictor. Dates are those on which’ the
legislation came into force. A - All live animals & parts; L - Live animals
only; S - Skins; P - Allowed under permit; C - Closed seasons or quotas may be
imposed; *-- these territories are Overseas Départements of France with which
the EEC may trade without the imposition of CITES controls; ? - no information
(Fuller et al., 1987).

a

CITES Hunting Trade Export

ee _

Argentina 1981 A 1983 A 1983 A 1986
Belize 1981 - A 1981 A 1981
Bolivia 1979 A 1979 A 1979 A 1979
Brazil 1975 A 1967 A 1967 A 1967
Colombia 1981 A 1973 A 1973 A 1973
Costa Rica 1975 A 1985 A 1970 A 1970
Dominica - ? ? ?
Ecuador 1975 - ~ A 1981
El Salvador 1987 - - P 1985
Fr. Guiana 1978 * - A 1986 A 1986
Fr. W. Indies 1978 * - - P
Guatemala 1980 - - C 1987
Guyana 1977 - - A 1986 +
Honduras 1985 - - Cc 1978
Mexico - c 1951 - A 1982
Nicaragua 1977 A 1977 A 1977 A 1977
Panama 1978 A 1980 A 1980 A 1980
Paraguay 1977 A 1975 +4 A 1975 A 1975 ++
Peru 1975 A 1973 A 1973 A 1973
St Lucia 1982 A 1980 A 1980 A 1980
Suriname 1981 - P
Trinidad & Tobago 1984 A 1958 A 1958 A 1958
Venezuela 1978 A 1970 A 1970 A 1970

+ Export quotas of 600 live B. constrictor and 500 skins were suggested

for 1987 and 1988.

++ On 17 February 1986, Decree 13806 allowed the export of 50 000 pairs of
shoes made from 8B. constrictor and BEBunectes spp. over the period of one
year. The ban on hunting was also relaxed for this purpose. A further
decree, No. 19815 of 6 February 1987, extended this dispensation for a further
year without specifying the number of shoes.

CAPTIVE BREEDING B. constrictor is regularly kept both in zoological
collections and as a pet. 8B. constrictor imperator is reported to be
difficult to tame, whereas B. constrictor constrictor adapts much better to
captivity (Breen, 1974).

227

Boa constrictor

An international survey of 260 zoological collections in 1985 (Slavens, 1985)
revealed the following information on the keeping and breeding of various
subspecies.

B. constrictor amarali: a total of 6 animals kept in 2 different
collections. No breeding recorded.

B. constrictor constrictor: a total of 223 animals kept in 38 different
collections. Breeding recorded in 8 collections, a total of 183 young
surviving.

B. constrictor imperator: a total of 22 animals kept in 10 different
collections. No breeding recorded.

B. constrictor nebulosa: a total of 8 animals kept in 3 different
collections. No breeding recorded.

B. constrictor occidentalis: a total of 34 animals kept in 12 different
collections. No breeding recorded.

B. constrictor orophias: a total of 7 animals kept in 3 £different
collections. No breeding recorded.

B. constrictor ortoni: a total of 8 animals kept in 4 different
collections. No breeding recorded.

B. constrictor sigma (=B. constrictor imperator ?): a total of 2 animals
kept in 1 collection. No breeding recorded.

B. constrictor’ subspp.: a total of 213 animals kept in 48 different
collections. Breeding recorded in 7 collections, a total of 221 young
surviving.

There is one record of commercial captive breeding of 8B. constrictor in
Latin America. A company in Panama, called Panama Sanchez, established a
captive-breeding operation for boa constrictors (Boa constrictor) in April
1981 near Panama City. The operation was allowed to collect adult snakes and
built up a breeding stock of 100 animals. Live snakes were exported to Miami,
USA, under five permits issued over the period 1981/1982.

375 snakes were exported in 1981 and 3355 in 1982. In July 1983 the operation
was closed after bad management practices caused the Government to suspend
permission to operate (D.M. Botello, in litt., 9 November 1983).

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Suchs. S.A., Costa Rica, 255 pp.

Miyata, K. (1982). A checklist of the reptiles of Ecuador with a bibliography
of Ecuadorean herpetology. Smithsonian Herpetological ' Information
Service No. 54: 1-70.

Peters, J.A. and Orejas-Miranda, B. (1970). Catalogue of the Neotropical
Squamata; Part 1, Snakes. Bulletin of the U.S. National Museum 297(1):
1-347.

Pope, C.H. (1962). The giant snakes. Routledge and Keegan Paul, London,
297 pp.

229

Boa constrictor

Roze, J. (1966). La taxonomia y zoogeografia de los ofidos en Venezuela. Ed.
Bibl. 28 Col. Cienc. Biol. 3: 1-362.

Schwartz, A. and Thomas, R (1975). A check-list of West Indian amphibians and
reptiles. Carnegie Museum of Natural History Special Publication 1:
1-216.

Serrano, F. (1978). Informe de actividades de la Unidad de Parques Nacionales
y Vida Silvestre de El Salvador. In: Morales, R., MacFarland, C., Incer,
J. and Hobbs, A. (eds), Primera Reunion Regional Centroamericana Sobre
Vida Silvestre. Centro Agronomico Tropical de Investigacion y Ensenanza,
Turrialba, Costa Rica, pp. 117-124.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens,, Seattle, Washington, 341 pp.

Smith, H.M. and Smith, R.B. (1976). Synopsis of the herpetofauna of Mexico
Vol. III. Source analysis and index for Mexican reptiles. John Johnson,
Vermont, 23 pp.

Stafford, P. (1979). Spotlight on a species: the Boa Constrictor. The
Herptile 4(3): 13-16.

Stidworthy, J. (1969). Snakes of the World. Hamlyn, London, 160 pp.

Swank, W.G. and Julien, C.R. (1975). Wildlife management and protection,
Dominica. The distribution and status of wildlife in Dominica FAO Project
Working Document FO:DP/DMI/74/001.

Torres Santibanez, H. (1978). Conservacion y manejo de la fauna silvestre del
Parque Nacional Defensores del Chaco. FAO Working Document FAO/SFN:
6/PAR/02/T.

Vallester, E. (1978). Informe de Panama sobre la situacion de la fauna
silvestre. In: Morales, R., MacFarland, C., Incer, J. and Hobbs, A. (eds),
Primera Reunion Regional Centroamericana Sobre Vida Silvestre. Centro
Agronomico Tropical de Investigacion y Ensenanza, Turrialba, Costa Rica,
pp. 43-64.

Wettenberg, G.B., Ferreira, M., Santos Brito, W.L. dos and Araujo, V.C. de
(1976). Amazonian fauna preferred as food. F.A.O. Technical Report, No.
6, FO:DP/BRA/71/545, 17 pp.

Wilson, L.D. and Meyer, J.R. (1982). The snakes of Honduras. Milwaukee
Public Museum, Wisconsin, 159 pp.

230

ANACONDA Recommended list: 2
[Possible problem]
Bunectes murinus (Linnaeus, 1758)

Order SERPENTES Family BOIDAE

—_e—e—eeeeeeeee COO nn ee

SUMMARY AND CONCLUSIONS The largest snake in South America, found in the
drainage basins of the Orinoco and the Amazon. Primarily aquatic, inhabiting
large rivers and swamps, and possibly growing to lengths in excess of 10m.
Prey includes a variety of mammals, up to the size of peccaries, and reptiles.
Brood sizes range from 14 to 82. Population levels are unknown.

The species is not extensively used for food, but may be persecuted for
cultural reasons. There is a substantial skin trade, probably over 20 000
animals a year, mostly declared as originating in Paraguay, Bolivia and
Guyana, but in reality probably deriving from Brazil. There is also a low
level of trade in live animals, probably for the pet trade.

Sustainable hunting rates cannot be estimated, owing to the almost complete
lack of population data, but the majority of the trade is currently illegal
and originates in countries where the species is protected. Importing
countries should not permit imports from these countries, but this requirement
is obviously being ignored. The chief importing countries are the USA, Italy
and France.

DISTRIBUTION Found in moist forests and swamps in the drainage basins of
the Orinoco and Amazon. The species has been divided into two subspecies:
Bunectes murinus scytale (formerly Bunectes murinus murinus), found in the
Amazonian region; and Bunectes murinus murinus (formerly Eunectes murinus
gigas), found chiefly in Venezuela and Guiana. However as there is not
agreement on the subspecific names, and as E. murinus murinus could apply to
either subspecies, giving considerable potential for confusion, the subspecies
are not considered separately in this account.

Bolivia Found in the Amazonian region (Hoogmoed, 1979).

Brazil Widespread in the Amazonian region, but absent from the Atlantic
forests (Dixon, 1979; Hoogmoed, 1979; Hoogmoed, 1982b).

Colombia Found in the Amazonian region (Hoogmoed, 1979).
Ecuador Found in the Amazonian region (Hoogmoed, 1979; Miyata, 1982).
French Guiana Present (Gasc and Rodrigues, 1980; Hoogmoed, 1982b).

Guyana Found in the Amazonian region (Hoogmoed, 1982b).

Peru Recorded from Quistococha, Iquitos, Rio Napo region (Dixon and Soini,
1986).

Paraguay There is a single record of &E. murinus from a swamp in the
Parque Nacional Cerro Cora (Anon., 1982). Scott (in Jitt., 2 April 1982)
reports that it is apparently found near all of the large rivers in the East
of the country. There is dispute as to whether these records may have been
confused with EB. notaeus.

Suriname Present (Hoogmoed, 1982a; 1982b).

231

Bunectes murinus

Trinidad and Tobago Recorded from Trinidad, but not from Tobago (Emsley,
1977; Hardy, 1982).

Venezuela The distribution covers the Orinoco and Maracaibo basins,
including all the tributaries. The species has not been found in the north of
the country except in the Orinoco Delta and in Caripito (Roze, 1966). Found
in the Llanos and Yunari Savanna of central Venezuela (Rivero-Blanco and
Dixon), and also in the Amazonian region (Hoogmoed, 1979; Hoogmoed, 1982b).
Probably does not occur above 500 m (S. Gorzula, in litt., 11 April 1986).

POPULATION

Bolivia No information.
Brazil No information.
Colombia No information.
Ecuador No information.

French Guiana Said to be relatively abundant, though rarely reported (Gasc
and Rodrigues, 1980).

Guyana No information.

Paraguay Before the single published record of FE. murinus, reported in
1982, the species was not considered to occur in the country (Peters and
Orejas-Miranda 1970). It must therefore be considered to be rare, if present
at all.

Peru Said to be common in and around flooded forests and large and small
streams where aquatic vegetation is dense (Dixon and Soini, 1986).

Suriname Said to occur in reasonable numbers (Hoogmoed, 1982a).

Trinidad and Tobago Locally common in the Narira Swamp and the larger
rivers of Trinidad, but does not occur in Tobago (Emsley, 1977; Hardy, 1982).

Venezuela Gorzula (in litt., 11 April 1986) considered that the species
was frequent in Venezuela, saying aestivating animals could readily be found
in receding lagoons in the Llanos in the dry season. In the right habitat in
southern Venezuela and Sucre, specimens are found more rarely - about one
every 40-50 man days.

HABITAT AND ECOLOGY Claimed by some to be the world's largest snake,
Eunectes murinus is almost certainly the heaviest, weighing up to a tonne,
although the maximum authenticated length of 7.63 m is exceeded by Python
reticulatus (Emsley, 1977; Freiberg, 1982). Unauthenticated reports put the
maximum length at 12-14 m (Pope, 1962), and it is not unlikely that snakes of
this size occur (Best, 1984).

E. murinus is- primarily nocturnal and aquatic, inhabiting swamps’ and
slow-flowing rivers. It occasionally emerges to rest on branches or sand banks
but is never found far from water, and always returns to water when
threatened. The young tend to be more arboreal than the adults (Pope, 1962;
Emsley, 1977). Usually solitary, there are isolated records of aggregations of
up to 11 snakes (Pope, 1962), although these may be associated with mating
behaviour. Owing to its large bulk, it is relatively sluggish on land but can
move with great rapidity in the water (Belloumini et al., 1976/77).

232

Bunectes murinus

The growth of E. murinus has been studied in captivity. Young emerge at a
length of about 70 cm, although hatchlings as small as 36-52 cm have been
reported (Belloumini et al., 1976/77). Holstrom (1980) reports three broods
of hatchlings averaging 74.8-87.7 cm in length and weighing 206-265 g. At 11
months Deschanel (1978) reported that young’ had grown to 140-157 cm
(1.65-1.91 kg), and at five years lengths of 3.1-3.2 m (17-27 kg) have been
recorded. After the age of three years, the increase in length slows but the
snake continues to increase in weight. Snakes of 4-5 m commonly weigh
50-100 kg (Belloumini et al., 1976/77). The record age in captivity is 29
years (Emsley, 1977).

Ecdysis occurs about six times a year in captivity. A young captive male
moulted 30 times during its first five years of life (Belloumini et al.,
1976/77), while six- to seven-year-old females moulted five to seven times a
year. The moulting frequency increases during gestation (Holstrom, 1980).

E. murinus is viviparous; the gestation period has been reported to vary
from six to nine months (Belloumini et al., 1976/77; Deschanel, 1978;
Holstrom, 1980; Holmstrom, 1982). Brood sizes have been reported to vary from
14 (Deschanel, 1978) to 82, larger mothers having more young, although the
size of the young is inversely correlated with the size of the mother
(Belloumini et al., 1976/77). Mating always takes place in the water and, in
Trinidad, usually occurs in December and January, the young emerging in July
and August (Emsley, 1977). In captivity, heterologous mating between
BE. murinus and Bunectes notaeus has been recorded, though no young were
produced (Veinert and Belloumini, 1980/81).

The prey is killed by constriction, a great variety of species being taken.
Emsley (1977) lists agoutis, Paca, cavies, peccaries, deer, monkeys, birds,
caiman, turtles, and some occasional domestic animals, but considers it
doubtful that they eat fish, although Pope (1962) reports considerable
quantities of fish from the stomachs of specimens taken in Guyana. The size of
the prey depends on the size of the snake. A 25-foot (7.6-m) Anaconda was
found in French Guiana with a 100-1b (45-kg) pig (possibly a peccary) in its
stomach (Pope, 1962). The prey is always taken near water, and is usually
drawn underwater for constriction. One snake was observed to remain underwater
for 17 minutes with its prey (Emsley, 1977). Young Anacondas tend to ingest
their prey underwater, while older animals often do so on the surface
(Belloumini et al., 1976/77).

A female in captivity ate a total of 88.9 kg of food in her first five years
of life, during which time she grew to a weight of 22 kg. A male sibling ate a
total of 79.9 kg and grew to 17 kg during the same period (Belloumini et
al., 1976/77).

THREATS TO SURVIVAL &. murinus is hunted chiefly for its skin, although
its flesh is occasionally eaten. Gasc and Rodrigues (1980) reported that it
was found "throughout" French Guiana on the menus of restaurants specialising
in exotic foods. Pope (1962) considered that the consumption of snakes was not
widespread in South America. The native peoples of French Guiana have a strong
aversion to Anacondas for cultural reasons, and are unwilling to kill, or even
look at them (Gasc and Rodrigues, 1980). Live animals are extensively traded
as pets. A pet supply company in the USA (South American Unlimited, New York)
included Anacondas on its price list in 1985 at US$60 each. Most of the
animals traded as pets are thought to be small, and therefore this trade is
likely to be less damaging to the population than the skin trade, which would
tend to use larger animals. There is very little detailed information
regarding exploitation. The species adapts readily to irrigation dams and
other man-made water bodies (S. Gorzula, in litt., 11 Aprii 1986).

233

Bunectes murinus
Brazil In July 1984, IBDF (Instituto Brasileiro de Desenvolvimento
Florestal) seized a shipment of 140 tanned snake skins (B. constrictor and
EBunectes murinus) in Porto Alegre. The skins were en route from Belem to Sao
Leppoldo and were thought to be destined for eventual export once they had
been manufactured into shoes and other products (Palazzo, J.T., in litt.
1984).

Paraguay Scott (in 1l4itt., 2 April 1982) reports that all 8. murinus
that are found are usually killed. The habitat where it is found, along the
large rivers in eastern Paraguay, is rapidly being settled. Figures compiled
by the Ministerio de Agricultura y Ganaderia (Acevedo Gomez, 1987) showed that
some 12 000 skins of E. murinus were exported from Paraguay in 1984.

Suriname Hoogmoed (1982a) writes that “there is no hunting for hides, nor
is the natural habitat being destroyed. So, at least in Suriname, this species
seems to be safe".

Venezuela Anacondas are killed very occasionally in Venezuela by ranchers
when they have taken cattle. There is thought to be no exploitation for meat
and little for the skin or pet trade. There are locally held beliefs that
anacondas, known as “madre de agua", prevent lagoons from drying out (S.
Gorzula, in litt., 11 April 1986).

INTERNATIONAL TRADE The only reports of international trade in &. murinus
are those contained in the Annual Reports of Parties to CITES. Only trade in
live animals and skins was considered.

Trade in skins was sometimes recorded by length. It is notoriously difficult
to convert this into numbers of skins, as it depends on the size of the snakes
killed and on the method of preparation of the skin, however an average skin
length of 2.1m has been suggested (Anon., 1984). It must be stressed that
with a snake which shows such a potential range of lengths, it seems unsafe to
put too much reliance on this figure.

Table la shows that the annual trade in skins of E. murinus declined from a
the high levels in 1980 and 1981 to about a third of the peak volume in 1985.
The reason for the temporary decline in 1983 is not known. It could be
associated with a decline in the demand for reptile skins reported by some
dealers (A. Sarkissian, in Jlitt., 27 January 1986); or with protection
measures in the countries of origin.

The numbers of E. murinus traded live (Table 1b) are insignificant compared
with the numbers of skins in trade. The small quantities suggest either that
the pet trade is very limited or that most of them are destined for zoological
collections.

The reported countries of origin of the skins are shown in Table 2a. The great
majority of skins are reported to have originated in Paraguay, a country in
which there are very few records of E. murinus. It is possible that these
skins, and those reported as originating in Argentina, were mistakenly
identified Eunectes notaeus skins, but independent evidence (Acevedo Gomez,
1987) confirms that substantial quantities of E. murinus are exported from
Paraguay, and so it seems likely that they are yet another example of the vast
illegal trade in wildlife skins which pours across Brazil's southern border.
There has been a much smaller number of skins reported as originating in
Brazil itself, which has a ban on all trade in wildlife. Skins originating in
Colombia made a significant contribution to the world volume of trade in 1983
in spite of the fact that exports are illegal. Bolivia and Guyana were major
suppliers in 1984 and 1985. Skins reported as originating in Indonesia,
Thailand and Nigeria are probably species of Python. Those from the other

234

Bunectes murinus

unacceptable countries of origin probably represent re-exports not specified
as such. Panama, the source of about a quarter of the world's supply of
E. murinus skins in 1983, is noteworthy as it is known to be a major
entrepot for wildlife products from South America (Vallester, 1978). The
major consignment from Singapore in 1985 may also have been a re-export.

Table la. Minimum net commercial imports of SE. murinus skins reported to
CITES.

1980 1981 1982 1983 1984 1985
Austria ‘ 6984 = = = - =
Brasil - ~ 82 = “3 =
Canada - 2 = at x a
- 29 m 36 m = = =
France - - 54 2 2645 =
- 2m - 12 m - -
Germany, F.R. - - = 3 = =
= - - 250 m - -
Hong Kong - - - = 2 =
Israel 215 m 89 m = - 1 =
Italy 675 500 942 5621 - -
- 1300 m 6778 m 4850 m 11053 m -
110 kg 1739 m2 S ef ay 4
Japan - - - - - 4000
Mexico - - 158 - - -
- - 22 m - - -
- - 120 kg - - 1 kg
New Zealand - - 1 - - -
South Africa 818 - - - - -
Spain - - - 39 m - -
Sweden - - - - 1 -
Turkey - - - 20 - -
UK 1443 m - - - - -
USA 11717 17730 4188 36 12938 4559
14041 m 23765 m 8029 m 632 m 2353 m 1500 m
- - 1010 kg - - 119 kg
Total 20194 18232 5425 5683 15587 8559
15699 m 25185 m 14865 m 5783 m 13406 m 1500 m
110 kg 1739 m2 1130 kg = a 120 kg

er een

235

Bunectes murinus

Table 1b. Minimum net commercial imports of live E. murinus reported to CITES

1980 1981 1982 1983 1984 1985

Australia
Austria
Belgium -
Canada 7 - -
Czechoslovakia
Denmark 2 - -
Dominican Rep 2 - = = = =
Germany, F.R. 39 35 26 | 24 3 70
Hungary - 3 - = = yy
Indonesia - = = 3 = <
Israel - = 1
Italy 7 - 2 17 - 4
2 5 4
1

a |
Ne I
i}

i]

i}

i}
(ew

Japan
Korea, Rep of - - - - 1
Mexico 2 - -
Netherlands - - -
Poland - -
Singapore - 2
South Africa - 2 - -
1
3

aon i
a
Nn
uw
Ls)

Spain =
Sweden =
Switzerland 2
Trinidad & Tobago - - = =

Table 2a Reported countries of origin or export for commercial exports of
skins of E. murinus reported to CITES.

1980 1981 1982 1983 1984 1985

Countries with wild populations of E. murinus

Bolivia 1 ~ - - 1101 20
- - - - 6300 m -
Brazil 1114 459 99 kg 2 kg - 166
872 m 1650 m 607 m 585 m - -
2 500 m2 = = = 1 kg
Colombia - 1 1 2500 - -
Guyana - - - 1888 4365 2100
- - - - 6000 m -
Paraguay 6730 11394 4438 1266 11014 2011
8091 m 5289 m 8264 m 2228 m 1407 m 1500 m
z 250 m2 2 = = 120 kg
Peru 6710 - 158 - - -
- 4749 m 21 kg 3 kg - -
Suriname 110 kg - - - - -
Trinidad & Tobago - 1 - - - -

236

Bunectes murinus

Table 2a continued

1980 1981 1982 1983 1984 1985

Countries without wild populations of &. murinus

Argentina 5639 5678 739 - ~ 825
499 m 8069 m 1579 m - - -
= 989 m2 = cf 2 A
Belgium - - - - 200 Es
Canada - - iT = = =
Indonesia ~ 636 - - 2 -
Netherlands 168 - - - - -
6215 m - = = = =
Nigeria 1 - 246 - - -
Panama - - - 3794 m - -
Singapore - - ~ - - 4000
Spain 136 1052 m - - - -
Thailand 228 - - - - -
22 m - - - - -
USA - - 2294 - - =
- - 4450 m - - -
Unknown 8992 2357 1130 kg 28 1627 -
1056 m 6026 m 550 m 12 m 946 m -

Table 2b. Reported countries of origin or export for commercial exports of

live E. murinus reported to CITES.

1980 1981 1982 1983 1984 1985

Countries with wild populations of EB. murinus

Bolivia 72 1
French Guiana - 3 1 = = =
Guyana 346 202
Paraguay 6 =
Peru - a = - = =
Suriname - 2
Venezuela - - - - 4 =

Countries without wild populations of E. murinus

Canada - 2 4 - - -
Finland - - - - 1 =
France - 3 - = >= =
Guatemala 3 - = = = =
Haiti 2 - = = =
Honduras - 2 - = = =
USA 8 11 9 5 8 2
Unknown 35 25 2 4 - =

237

Bunectes murinus

CONSERVATION MEASURES The legal protection status of 5. murinus_ in
various countries throughout its range is summarised in Table 3. On the basis
of this information it appears that the species is protected in most of the
major supplying countries: Paraguay, Argentina, Colombia, Bolivia, Brazil,
Peru and Panama. Furthermore the species does not occur in Argentina or
Panama, and is extremely rare in Paraguay, indicating that the skins were
acquired elsewhere. None of these countries should therefore have been
acceptable to the CITES Management Authorities who granted import licences for
the skins. The emergence of Guyana as a major source of skins in 1983 and 1984
is disturbing as E. murinus is not protected there. While it is possible
that the skins were acquired within the country it is also possible that they
derived from illegal imports from Brazil, as is the case with other wildlife
products. The Brazilian authorities have already begun action to curb the
poaching and illegal traffic in the South of the country, but the problem is
so vast that little improvement is yet apparent (Hyman, 1985).

Table 3. Legal prohibition on the hunting, internal trade and commercial
export of E. murinus. Dates are those on which the legislation came into
force. A - All live animals & parts; P - Allowed under permit; * - these
territories are Overseas Départements of France with which the EEC may trade
without the imposition of CITES controls (Fuller et al., 1987).

CITES Hunting Trade Export
Bolivia 1979 A 1979 A 1979 A 1979
Brazil 1975 A 1967 A 1967 A 1967
Colombia 1981 A 1973 A 1973 A 1973
Ecuador 1975 - ~ A 1981
Fr. Guiana 1978 * - A 1986 A 1986
Guyana 1977 - - A 1986 +
Paraguay 1977 - A 1975 A 1975 ++
Peru 1975 A 1973 A 1973 A 1973
Suriname 1981 - - P
Venezuela 1978 A 1970 A 1970 A 1970

+ Export quotas of 600 live B. constrictor and 500 skins were suggested
for 1987 and 1988.

++ On 17 February 1986, Decree 13806 allowed the export of 50 000 pairs of
shoes made from B. constrictor and Eunectes spp. over the period of one
year. The ban on hunting was also relaxed for this purpose. A further
decree, No. 19815 of 6 February 1987, extended this dispensation for a further
year without specifying the number of shoes.

CAPTIVE BREEDING E. murinus is regularly kept both in zoological
collections and as a pet. It is reported to be relatively aggressive in
temperament and to strike readily (Emsley, 1977).

An international survey of 260 zoological collections in 1985 (Slavens, 1985)
revealed a total of 48 animals kept in 15 different collections. Breeding was
not reported in 1985, although it had occurred in previous years.

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238

Bunectes murinus

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Veinert, T. and Belloumini, H.E. (1980/81). Observacoes do comportamento e da
copula heterologa da sucuris em cativiero - Bunectes murinus murinus

(Linnaeus) e Bunectes notaeus_ Cope, 1862. Memorias do Instituto
Butantan 44/45: 391-402.

240

YELLOW ANACONDA Recommended list: 2

[Possible problem]
Bunectes notaeus Cope, 1862

Order SERPENTES Family BOIDAE

SUMMARY AND CONCLUSIONS Distributed in large rivers and swamps in
sub-tropical South America, from Bolivia to Argentina. Primarily aquatic,
growing to a maximum length of 4 m. Very little is known about its ecology or
population size.

Apart from limited trade in live specimens, probably for zoological
collections, the main trade is in skins, mostly declared as originating in
Paraguay, Bolivia and Argentina. The volume of this trade appears to have
increased dramatically since 1980 to some 37 000 skins in 1984.

The data available are insufficient to set sustainable hunting rates, but the
species is protected in its main countries of origin. With the exception of a
few skins from Bolivia, all exports should not have been accepted by the
importing countries, but this requirement appears to have been ignored. The
main importing countries are the USA, France and Italy. From 1985 onwards the
species has been protected in all the source countries.

DISTRIBUTION Found in sub-tropical South America, from Bolivia to northern
Argentina.

Argentina Confined to the north-east, in the Chaco, Corrientes and Santiago
del Estero (Gallardo, 1977; Freiberg, 1982).

Bolivia Present (Peters and Orejas-Miranda, 1970).
Brazil Confined to the south-west of the country (Freiberg, 1982).
Paraguay Present (Peters and Orejas-Miranda, 1970).

Uruguay Present (Peters and Orejas-Miranda, 1970).

POPULATION There is no information on the status of populations in any of
the source countries.

HABITAT AND ECOLOGY fFunectes notaeus is similar in habits to Sunectes
murinus, being largely aquatic, and inhabiting large rivers and swamps. It is
smaller, usually between 2m and 3m in length, with a maximum of 4m
(Freiberg, 1982).

There is little information on the growth rate, but 37 young were born at New
York Zoological Park at lengths of 535-780 mm (mean 639 mm), weighing 95-180 g
(mean 137 g). At two years of age they had grown to an average length of
1.78 m, weighing 2.6 kg (Holmstrom, 1981). A male E&. notaeus at Sao Paulo
Zoo grew from 2.00 m (5.0 kg) to 2.25 m (9.0 kg) in four years (Veinart and
Belloumini, 1980/81).

Like all New World boids, EF. notaeus is ovoviviparous. Broods of 6 to 13
have been reported (Belloumini et al., 1976/77; Holmstrom, 1981). Gestation
appears to take from four to six months, and at New York Zoological Park,
births occurred from April to October (Holmstrom, 1981; Holmstrom, 1982). One

241

Bunectes notaeus

of the males in this collection began courtship at an age of 21 months
(Holmstrom, 1981), and a female gave birth at four years, two months of age
(Holmstrom, 1982). The oldest female gave birth to three broods, totalling 31
offspring, in successive years with 13-month intervals between each birth
(Holmstrom, 1981). In captivity, heterologous mating between E. notaeus and
Eunectes murinus has been recorded, though no young were produced (Veinert
and Belloumini, 1980/81).

Prey is killed by constriction, and is thought to consist of mammals and
reptiles. Young snakes born in a Zoo did not start feeding until one to four
months after birth (Holmstrom, 1981).

THREATS TO SURVIVAL 5. notaeus is hunted chiefly for its skin, and also
occasionally for meat. It is reputed to have the best flavour of all boids
(Gallardo, 1977). ;

Paraguay Scott (in litt., 2 April 1982) asserted that it was not hunted
much in Paraguay. The habitat where it is found, along the large rivers in
eastern Paraguay, is rapidly being settled. Figures compiled by the
Ministerio de Agricultura y Ganaderia (Acevedo Gomez, 1987) showed that some
10 000 skins of E. notaeus were exported from Paraguay in 1984.

Table la Minimum net commercial imports of E. notaeus skins reported to
CITES.

1980 1981 1982 1983 1984 1985
Australia - - 7 - - -
Canada - 4 289 52 37 1
France - - - 2m 12177 -
Germany, F.R. 1 2768 236 122 - -
- 3000 m - ~ - -
Hong Kong - - - 3 - -
Italy 58 123 - - - 2466
950 m2 = 42 m 8260 m 6376 m =
Japan - - - - - 253
Korea, Rep of - - - - 24 -
Mexico = = 539 603 1 m@ E
New Zealand - - 20 - 1 -
Panama - - 1317 - - -
Peru - - - 4375 - -
Singapore - - 1 - - -
Spain - 1380 - - - -
Switzerland 526 - 2 34 - 4
Turkey - - - 13 - -
UK - 5000 m - - 75 m -
USA 891 1721 13973 - 18113 14712
- 6226 m 34851 m 1721 m 2702 m 2389 m
- - - - 4 kg 13 m
Venezuela - 13 - - - -
Unknown - - - 3 - -
Total 1476 6009 16384 5205 30352 17435
- 14226 m 34893 m 9983 m 9153 m 2389 m
950 m2 a = 1 m2 13 m? 964 m2
= = = = 4 kg =

242

Bunectes notaeus

Table 1b. Minimum net commercial imports of live E. notaeus reported to
CITES.

1980 1981 1982 1983 1984 1985
Belgium - = = = = 7
Canada 1 2 - = a =
Czechoslovakia - 1 = = = as
Denmark - 2 = = = =
France - 3 = = =
German D.R. 2 - 8 2 = =
Germany, F.R. 40 3 10 3 - -
Hungary - 2 = = = A
Italy 2 - = 3 = 9
Japan - 2 = = + S
Netherlands - - = = = 8
Poland - - = 4 = =
Switzerland 7 = = = = re
UK 7 6 10 - - -
USA 9 47 2 13 23 -
USSR - - - - 2 =
Total 61 68 30 25 25 24

INTERNATIONAL TRADE The only reports of international trade in E. notaeus
are those contained in the Annual Reports of Parties to CITES. Only trade in
live animals and skins was considered. The CITES reports are summarised in
Tables 1 and 2. Trade in skins was sometimes recorded by length. It is
notoriously difficult to convert this into numbers of skins, as it depends on
the size of the snakes killed and on the method of preparation of the skin,
however an average skin length of 1.4 m has been suggested (Anon., 1984). It
must be stressed that with a snake which shows such a potential range of
lengths, it seems unsafe to put too much reliance on this figure.

The minimum net trade in skins of E. notaeus (Table la) has increased from
comparatively low levels in 1980 to about 37 000 in 1984 but declined again in
1985. This pattern differs significantly from the trade in skins of
E. murinus, which has been declining slightly since 1980. It is possible
that, as the supply of E. murinus skins has been curtailed, the trade has
switched to EB. notaeus. The chief importing countries appear to be the USA,
F.R. Germany, Italy and the UK.

The trade in live animals (Table 1b) has been minimal, and is probably mostly
connected with zoological collections.

The great majority of skins were declared as originating in Paraguay,
Argentina and Bolivia, and, for once, there is no real reason to doubt this,
as the species occurs in all of these countries. However it seems likely that
a certain number of skins will have originated in neighbouring Pantanal of
Brazil and have entered Paraguay along with the other illegal traffic in
wildlife products. Small numbers of skins were declared as originating in
Brazil and Uruguay. Among the countries of origin not having wild populations
of EF. notaeus, the majority probably represent re-exports without the
country of origin having been specified, or of captive-bred specimens not
reported as such. Skins originating in Indonesia and Thailand were possibly of
another species of snake, Python sp., for instance. In 1982 and 1983, Peru

243

Bunectes notaeus

Table 2a. Reported countries of origin or export for commercial exports of
skins of BE. notaeus reported to CITES.

1980 1981 1982 1983 1984 1985

Countries with wild populations of EB. notaeus

Argentina 4 - ~ 363 8844 2596
= 200 m2 = 7466 m 6376 m 1486 m
Bolivia - 1 - - 13489 2950
- - - - - 886 m
Brazil - 28 45 396 - -
Paraguay 420 2424 14526 - 9888 11368 11157
- 9482 m 33871 m 2551 m 2590 m -
= 750 m2 a 7 kg 4 kg =
Uruguay - - = 61 = ~
Countries without wild populations of E. notaeus
Colombia - - - - 185 m -
France - - 1022 m - - -
Guyana - 2 - - 645 -
Indonesia - - - 60 - -
Netherlands 1472 35 - 33 kg - -
Panama - - - - 190 2244
Peru - - 7802 m 1075 746 544
South Africa - - - - 60 1
Thailand - - - 319 165 -
Unknown - 3866 2211 228 - 50
’ - 6244 m 42 m 27 m - 17 m
- - - - - 13 m2

Table 2b. Reported countries of origin or export for commercial exports of.
live EB. notaeus reported to CITES.

1980 1981 1982 1983 1984 1985

Countries with wild populations of BE. notaeus
Paraguay 61 59 2 - - -
Countries without wild populations of E. notaeus

Austria - = = = = 2
Canada - - 8 14 19 =
Czechoslovaia - = 10 = = 3
Germany, F.R. -
Guyana -
Suriname
Switzerland
USA

Unknown

“ow!
nmin
a
fo)
Peni wi
Int nI
r
co

244

Bunectes notaeus

was declared as the origin of large quantities of skins, although FE. notaeus
does not occur in the country. These skins must therefore either have been
imported from elsewhere or have been misidentified skins of 5. murinus.
There is no evidence to suggest which of these two explanations is the more
likely.

CONSERVATION MEASURES The legal protection status of &. notaeus. in
various countries throughout its range is summarised in Table 3. The species
receives nominal protection in all its potential countries of origin, and all
exports of skins, except for skins from Bolivia (Table 2a), have therefore
been illegal. It is difficult to understand why such trade in the skins of
BE. notaeus has been sanctioned by importing CITES Management Authorities.
Since August 1985, when Bolivia banned the export of wildlife products, there
have been no legal sources of skins of this species.

Table 3 Legal prohibition on the hunting, internal trade and commercial
export of E. notaeus. Dates are those on which the legislation came into
force. A - All live animals & parts; L - Live animals only; S - Skins;
P - Allowed under permit; C - Closed seasons may be imposed; ? - no
information (Fuller et al., 1987).

CITES Hunting Trade Export
Argentina 1981 A 1983 A 1983 A 1986
Bolivia 1979 - S 1979 L 1984 +
Brazil 1975 - A 1967 A 1967
Paraguay 1977 A 1975 A 1975 A 1975 ++
Uruguay 1975 A 1978 A 1978 A 1978

+ all wildlife exports have been banned since August 1985.

++ On 17 February 1986, Decree 13806 allowed the export of 50 000 pairs of
shoes made from B. constrictor and Eunectes spp. over the period of one
year. The ban on hunting was also relaxed for this purpose. A further
decree, No. 19815 of 6 February 1987, extended this dispensation for a further
year without specifying the number of shoes.

CAPTIVE BREEDING &£. notaeus is regularly kept in zoological collections.
A survey of 260 zoological collections in 1985 (Slavens, 1985) revealed a
total of 61 animals kept in 18 different collections. Breeding was reported in
two collections, a total of 20 live being born.

REFERENCES

Acevedo Gomez, C. (1987). Especies de fauna amenezadas por comercio en el
Paraguay. Ministerio de Agricultura y Ganaderia, Paraguay, Unpublished.

Anon. (1984). A perception of the issue of high trade-volume. Unpublished
report prepared for CITES by the Wildlife Trade Monitoring Unit, IUCN.

Belloumini, H.E., Veinert, T., Dissman, F., Hoge, A.R. and Penha, A.M.
(1976/77). Notas biologicas a respeito do genero Bunectes Wagler, 1830
*sucuris' (Serpentes: Boinae). Memorias do Instituto Butantan 40/41:
79-115.

Freiberg, M. (1982). Snakes of South America. T.H.F. Publications, Neptune,
New Jersey, 198 pp.

Fuller, K.S., Swift, B., Jorgensen, A. and Brautigam, A. (1987). Latin
American wildlife trade laws, Second Edition (Revised). World Wildlife
Fund-US, 418 pp.

Gallardo, J.M. (1977). Reptiles de los alrededores de Buenos Aires.
Editorial Universitaria de Buenos Aires, 213 pp.

245

Bunectes notaeus

Holmstrom, W.F. (1981). Observations on the reproduction of the Yellow
Anaconda Bunectes notaeus. International Zoo Yearbook 21: 92-94.

Holmstrom, W. (1982). Life history notes, Serpentes. Herpetological Review
13(4): 126.

Peters, J.A. and Orejas-Miranda, B. (1970). Catalogue of the Neotropical
Squamata; Part 1, Snakes. Bulletin of the U.S. National Museum 297(1):
1-347.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

Veinert, T. and Belloumini, H.E. (1980/81). Observacoes do comportamento e da
copula heterologa da sucuris em cativiero - Bunectes murinus murinus
(Linnaeus) e Bunectes notaeus’ Cope, 1862. Memorias do Instituto
Butantan 44/45: 391-402.

246

BLOOD PYTHON, SHORT PYTHON Recommended list: 2

{Possible problem]
Python curtus Schlegel, 1872

Order SERPENTES Family BOIDAE

SUMMARY AND CONCLUSIONS A small South East Asian python, restricted to the
southern half of the Malay Peninsula, Bangka, Borneo and Sumatra; seemingly
local in distribution. Variously regarded as rare or moderately common,
although no population surveys are available. Consistently reported much
rarer than Python reticulatus. Said to prefer swamp forest or heavy jungle
along watercourses, also occurs in secondary growth. Nocturnal, often enters
water, feeds on small vertebrates. Clutch of 10-15 eggs is much smaller than
in congeneric species. Possibly threatened by habitat changes, although no
substantiating data are available; more likely to be threatened’ by
over-exploitation.

The volume of the skin trade as shown by CITES reports increased sharply from
935 in 1980 to some 58 500 in 1985. The USA, Italy, Japan and Canada were the
main importers. Most of the skins originated in Indonesia. The volume of
trade in live animals declined from 359 in 1980 to only 43 in 1985. This rise
in international skin trade may perhaps in part be attributed to legislation
affecting other exploited species.

It seems possible that this size of harvest may not be sustainable in the long
term, particularly as it appears to be increasing rapidly; a prime requirement
is to obtain relevant data on population levels, distribution, and the effects
of trade, with a view to appropriate management. Such data are required most
urgently for Indonesia, the reported origin of the great majority of animals
in the skin trade.

DISTRIBUTION A South East Asian species, with a somewhat more restricted
distribution than other Python in the region. Present in the southern half
of the Malay Peninsula and certain islands of the Indo-Australian
archipelago. As noted below, the species appears to be largely confined to
swamp forest and heavy jungle along watercourses, and is thus likely to be
sporadically distributed within its general range.

Brunei Almost certainly present, but no specific records are at hand.

Indonesia Restricted to Bangka, Sumatra and Kalimantan (De Haas, 1950, De
Rooij, 1917).

Malaysia Recorded from Peninsular Malaysia, Sabah, and Sarawak; seemingly
widely distributed but local (Tweedie, 1983; Malaysia, Sabah CITES MA, 1985).

Singapore First recorded from Singapore by Blandford, reporting in 1881 on
a collection made by Dennys. This record (of a single specimen) seems to be
the basis for reports of curtus from Singapore made by later authors (such
as Ridley, 1899; Flower, 1899; Boulenger, 1912), none of whom appear to have
had first hand experience of the species in that country. Sworder (1922), in
an annotated list of Singapore snakes, calls into question the accuracy of
locality data for many specimens in the Dennys collection; thus there may be
an element of doubt over the occurrence of the species in Singapore. However,
although no later authentic records can be traced, and recent information is
that curtus is not in fact present (P. Gopalakrishnakone, in Jitt., 13
March 1986), the species may have occurred in Singapore in the past.

Thailand Restricted to Changwat Pattani in the extreme southeast (Taylor,
1965; Soderberg, 1965).

247

Python curtus

Viet Nem Two specimens were reported from near Saigon, southern Viet Nam,
by Tirant in 1885. His identification seems justified by the description
provided (Campden-Main, 1970), but the distinct possibility has been raised
that these were introduced by man - there is an active trade in pythons and
other large snakes in the region (Saint Girons, 1972). There have been no
subsequent records from Viet Nam, but if the natural range does extend this
far to the northeast the species might be expected to occur in Kampuchea
also although it does not appear to have been recorded (Saint Girons, 1972).

POPULATION No detailed information is available on populations’ of
P. curtus anywhere in its range, nor on population trends or the effects of
exploitation. The species is generally reported to be less common than
Python reticulatus in the region.

Brunei No information.
Indonesia No information.

Malaysia Reported not rare in the peninsula at the turn of the century
(Ridley, 1899), and not uncommon in peninsular Malaysia in the 1950s (Tweedie,
1983). At Asahan in Malacca, only one P. curtus was seen during a period of
three and a half years, while P. reticulatus was said to be very common
(Batchelor, 1958). Among the snakes regularly brought to the University at
Bangi, Selangor, there is perhaps one curtus to every eight to ten
reticulatus (G. Davison, in Jlitt., 22 February 1986). In general, not
commonly encountered in peninsular Malaysia, but widespread and not heavily
exploited (S. Ambu, in litt., 17 February 1986). Here, also said to be not
frequently seen, but not markedly uncommon either; the relative frequency of
curtus to reticulatus is about 1:100 (B. Kiew, in Jlitt., 25 February
1986).

Said to be much rarer than Python reticulatus (common) in Borneo in the
early years of this century; this report apparently refers to Sarawak in
particular (Shelford, 1916). No curtus were found during long-term
herpetological sampling at three primary rainforest sites in Sarawak in
1962-64 and 1984 (R.F. Inger, in litt., 5 March 1985), and none were seen by
another fieldworker (working on rainforest lizards) (H. Watson, in litt., 17
March 1986). Similarly, reported less common and less widespread than P.
reticulatus in Sabah, but suitable habitat is found through most of the
country (Malaysia, Sabah CITES MA, 1985).

Singapore No information (see Distribution section above).
Thailand Cited as rare within its very restricted range (Soderberg. 1965).

HABITAT AND ECOLOGY A relatively small and very stout-bodied python,
reaching about 2.75 m total length. Preferred habitat variously cited as
swampy country (De Rooij, 1917), swamp forest (Bain and Humphrey, 1982), and
heavy jungle along watercourses (Reitinger, 1978). In West Malaysia
P. curtus has been found in the same kinds of habitat as P. reticulatus,
such as along streams, on the forest floor, and in secondary growth.
Nocturnal, spends much time in water, feeds on small vertebrates, reportedly
fond of rats (Ridley, 1899; Reitinger, 1978). This last seems confirmed by
the fact that 10 out of 11 specimens collected near Kuala Lumpur between 1948
and 1954 were found in rat traps at an experimental trapping area at Sungei
Buloh (Lim, 1955). An oviparous species, laying 10-15 eggs which are brooded
by the mother until hatching (Reitinger, 1978)

THREATS TO SURVIVAL None adequately documented, although collection for the
skin trade may be a serious threat (see below). Reportedly “most of" the

248

Python curtus

preferred habitat of the species has been destroyed (Bain and Humphrey, 1982);
it seems likely that this is intended to refer to Thailand rather than South
East Asia in general, although this is not entirely clear. It is further
uncertain whether mangrove forest should be regarded as P. curtus habitat as
is implied by these authors’ text, and in any case ‘preferred’ habitat is not
known. The species is certainly much consumed locally, although it is not as
popular as the much larger P. reticulatus (Irvine, 1954).

Malaysia In Sabah, the Chinese community uses python meat particularly for
soup, and occasionally the skins are used for decorative purposes. The levels
of exploitation are thought to be low, and capture for the pet trade is
thought to be insignificant (Sabah CITES MA, 1985).

INTERNATIONAL TRADE Although P. curtus has long figured in the live
animal trade, with many specimens being exported through Singapore (Irvine,
1954), CITES reports indicate that a much larger number of animals are now
used by the skin trade.

Table la Minimum net imports of skins of P. curtus reported to CITES.

1980 1981 1982 1983 1984 1985

Austria - - 89 - = =

Canada - - - 54 m - 11750

Finland - - - 217 m 481 m -

France - 76m 1427 2170 m - -

German D.R. - - - 8822 m - -

Germany, F.R. - - - 2391 655 -

= = - = 118 m rs

Greece - - 13 Jim - -

Italy 2. 2500 m2 13910 4571 2544 =

- 1440 m 7685 m 15732 m 1455 m -

Japan - - 22 - 971 kg 7527

Mexico - - - - 803 m -

Netherlands 72 - - - - -

Spain - 1189 “167 - 58 -

Switzerland - 4001 m - 1436 1449 -

Turkey - - - - 200 -

UK = - 222 - - -

USA 863 14891 3619 4083 9984 11902
- - 3762 m 700 m 15500 m 27393 m

= = = = 2 kg =

Total 935 16080 19469 12481 14890 31179
- 5517 m 11447 m 27766 m 18357 m 27393 m

a 2500 m? = e 973 kg =

International trade in P. curtus was examined by means of the Annual Reports
of Parties to CITES; only trade in live animals and skins being considered.
The volume of the skin trade is shown in Table la. Some transactions were
reported in terms of length, weight or area; it is difficult reliably to
convert these to numbers of skins, but as P. curtus rarely exceeds 3m in
total length, the mean length of skins in trade is likely to be in the region
of 1m. It must be recognised that this can only lead to an approximation of

249

Python curtus

Table 1b Minimum net imports of live P. curtus reported to CITES.

1980 1981 1982 1983 1984 1985
Argentina 1 - 2 - - -
Austria - - 5 2 - 1
Canada 1 - 1 3 6 6
Denmark 4 - - - - -
Germany, F.R. 127 40 46 16 20 10
Hong Kong - - - - - 6
Italy 13 1 8 30 - -
Japan - - - 16 - -
Mexico ’ = = 4 - = =
Neth. Antilles - - = 1 - -
South Africa - - 2 - 3 -
Spain - 2 - - - -
Switzerland 4 - 10 - - -
UAE - - 4 - - -
UK 52 13 17 5 - 2
USA 157 128 170 69 67 18
Total 359 184 269 142 96 43

Table 2a Reported countries of origin (or exporting country if no origin
reported) and quantities of transactions in live animals (L) and skins (S) of
P. curtus reported to CITES.

1980 1981 1982 1983 1984 1985

Countries having or possibly having populations of P. curtus

9256 9256 11568 16708 28090

Indonesia -
- 10517 m 3012 m 23948 m 11480 m~ 27393 m
= 2500 m2 = 285 kg 975 kg =
Malaysia - 715 - - - -
Thailand - - - - 1642 -

Countries without wild populations of P. curtus

India - 8 - - - -
Japan - 3479 - - 59 -
Singapore 935 7579 8435 m 1289 272 11359
- - - 8m 1514 m -
Unknown - 2899 24688 906 1642 6
- - - 8822 m 22559 m -
- - - - 975 kg -

250

Python curtus

the number of individuals involved, but on this basis the volume of trade
increased sharply from 935 in 1980 to 58 572 in 1985, ignoring transactions
reported by weight or area. The USA, Italy, Japan and Canada were the main
importers. The declared sources of the skins are given in Table 2a, from
which it appears that Indonesia was the major supplier; it is likely that the
large numbers recorded from Singapore also in fact originate from Indonesia.

Table 1b shows the volume of trade in live animals, which declined from 359 in
1980 to only 43 in 1985. The great majority of live snakes originated in
Thailand (see Table 2b), until 1984, when Malaysia emerged as the main source.

The sudden and dramatic increase in 1981 in the number of P. curtus skins in
international trade suggests that trade may be shifting to this species now
that its larger congeners are nominally protected in much of their range;
reduced availability of other Python used by the skin trade may also be a
factor, but this cannot be established at present. No data whatsoever are
available on population status of this species in Indonesia; this information
is required in order to assess the likely impact of the present harvest.

Table 2b Reported countries of origin (or exporting country if no origin
reported) and numbers of live P. curtus reported to CITES.

1980 1981 1982 1983 1984 1985

Countries having or possibly having populations of P. curtus

Indonesia 1 = = = en a
Malaysia 7 2 2 5 96 43
Thailand 354 185 279 135 - 6

Countries without wild populations of P. curtus

Canada 1 - - - = -
Germany, F.R. - - 1 - -
Ghana 1 - - = = =
USA 1 - 2 3

Unknown - - 3 2 - -

CONSERVATION MEASURES The species is largely unprotected by legislation; it
does not appear on available lists of taxa covered by conservation legislation
in Brunei.

Indonesia Hunting quotas for 1987 were set to total 25 000 skins, divided
between the regions as follows: Acoh, 8000; Sumut, 13 000; Sumbar, 2000; Riau,
2000 (Indonesia CITES MA, 1987).

Malaysia Not protected in Peninsular Malaysia, Sabah or Sarawak (E.
Bennett, in litt., 5 February 1986).

Singapore All wild fauna in Singapore is fully protected (Singapore Primary
Production Department, in litt., 11 January 1986).

Thailand Recently protected in Thailand under the Wild Animals Reservation
and Protection Act B.E.2503, effective 11 November 1985.

251

Python curtus

CAPTIVE BREEDING This species has reportedly proved very difficult to
Maintain successfully in captivity (Stafford, 1982). There were 38 captive
specimens in 18 collections in 1984; no captive breeding recorded (Slavens,
1985), nor in 1979-1981 (Olney, 1984).

REFERENCES

Bain, J.R. and Humphrey, S.R. (1982). A profile of the endangered species of
Thailand. Report No. 4, Office of Ecological Services, Florida State
Museum.

Batchelor, D.M. (1958). Some notes on the snakes of Asahan, Malacca.
Malayan Nature Journal 12(3): 103-111.

Boulenger, G.A. (1912). A vertebrate fauna of the Malay Peninsula from the
Isthmus of Kra to Singapore, including the adjacent islands. Reptilia
and Batrachia. London.

Campden-Main, S.M. (1970). A field guide to the snakes of South Viet Nam.
Smithsonian Institution, Washington.

Flower, S.S. (1899). Notes on a second collection of reptiles made in the
Malay Peninsula and Siam. With a list of the species recorded from those
countries. Proceedings of the Zoological Society of London 1899:
600-697.

Haas, C.P.J. de (1950). Checklist of the snakes of the Indo-Australian
Archipelago. Treubia 20(3): 511-625.

Irvine, FOR: (1954). Snakes as food for man. British Journal of
Herpetology, 1(10): 183-189.
Lim, B.L. (1955). Snakes collected near Kuala Lumpur. Malayan Nature

Journal 9(4): 122-125.

Olney, P.J.S. (Ed) (1984). Reptiles bred in captivity and multiple generation
births, 1981. In, International Zoo Yearbook 23. (See also vols. 21,
(274 ))ic

Reitinger, F.F. (1978). Common snakes of South Bast Asia and Hong Kong.
Heinemann Educational Books (Asia) Ltd., Hong Kong, 114 pp.

Ridley, H.N. (1899). The habits of Malay reptiles. Journal of the Straits
Branch of the Royal Asiatic Society 32: 185-210.

Rooij, N. de (1917). The reptiles of the Indo-Australian Archipelago, II
Ophidia. E.J. Brill, Leiden. Reprinted 1970, A. Asher & Co. N.V., Vaals.

Saint Girons, H. (1972). Les Serpents du Cambodge. Mémoires museum National
d'histoire Naturelle, N.S., Ser. A (Zool.), 74.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity,
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

Shelford, R.W.C. (1916). A naturalist in Borneo. London, T. Fisher Unwin.

Soderberg, P. (1965). The pythons of Thailand. Conservation News S.E. Asia
5: 11-12

Stafford, P.J. (1982). Further observations on the Blood Python in captivity
including an effective method of inducing feeding. The Herptile 7(1):
21-22.

Sworder, G.H. (1922). A list of the snakes of Singapore Island. Singapore
Naturalist 1(2): 55-73.

Taylor, E.H. (1965). Serpents of Thailand and adjacent waters. University
of Kansas Science Bulletin 45(9): 609-1096.

Tweedie, M.W.F. (1983). The snakes of Malaya (3rd edn). Singapore National
Printers (Pte) Ltd.

252

INDIAN PYTHON, ROCK PYTHON Recommended list: 2

[Possible problem]
Python molurus (Linnaeus, 1758)

Order SERPENTES Family BOIDAE
a

SUMMARY AND CONCLUSIONS A large and widespread South and South East Asian
python, ranging from Pakistan, Indian, Sri Lanka, Bangladesh and Burma
eastward to Hong Kong and southern China (including Hainan), and south through
Viet Nam, Kampuchea, and Laos to Thailand, although absent from the Malay
Peninsula. Present in Indonesia (Java, Sulawesi, Sumbawa) but reports from
Borneo may be in error. Occurs in a variety of habitats, often rather open
deciduous woodland, often with rock outcrops, and typically in the vicinity of
permanent water. Populations in the Indian sub-continent are assigned to P.
m. molurus, those from Burma eastward to P. m. bivittatus; the former taxon
is listed on CITES Appendix I, the latter on Appendix II. Population
information is sparse and anecdotal; said to be common in southern Viet Nam,
Sri Lanka and possibly Burma, and to be depleted or locally extinct in most of
the Indian sub-continent. No data available for most of the range of P. m.
bivittatus.

Widely used for food and medicinal purposes, particularly by tribal and
Chinese-speaking communities. The volume of the international skin trade
appears from the CITES reports to have increased from 52 572 in 1980 to
Z2o92o in 1985. Italy, France, the USA and F.R. Germany were the main
importers and Thailand, Indonesia and Viet Nam were the major suppliers. The
trade in live animals also showed a dramatic increase, from 4873 in 1981 to
25 255 in 1985. The USA was the major importer and the great majority of live
snakes originated in Thailand.

The sustainability of the recent trade cannot be objectively evaluated in the
absence of population data; field data on population sizes and trends are
required. It must be suspected that present trade is excessive and it appears
to be increasing steadily.

DISTRIBUTION Widespread in the Indo-Malayan region. Extends from the Sind
region of Pakistan east almost throughout India and across the lowlands of
Nepal to Bangladesh, also to Sri Lanka. From the Indian sub-continent the
species extends through much of mainland southeast Asia, north into
subtropical China and Hong Kong; although apparently absent from the Malay
Peninsula, it occurs on Sulawesi, Java, and adjacent smaller islands, and has
been reported to occur on Borneo (this may be questionable). As with Python
curtus, P. molurus virtually always lives in the vicinity of water, and its
actual distribution must thus be rather patchy within its overall range.

Taxonomic note Populations centred on the Indian subcontinent are assigned
to Python molurus molurus, Indian Python (Appendix I); populations from
Burma (and possibly Nepal) eastwards are assigned to P. m. bivittatus,
Burmese Python (Appendix II). Smith (1943) states that molurus ranges east
to Bengal (i.e. including present-day Bangladesh) and that bivittatus occurs
throughout the ‘Indo-Chinese subregion’ (in which he includes Nepal and
north-east India); however, animals from throughout the sub-continent,
including Nepal, are often treated as the nominate form (Stimson, 1969). The
south-west Bangladesh population has been referred to P. m. bivittatus (Kock
and Schroder, 1981). Although many authorities support the use of trinomials,
very few specimens from the supposed contact area of molurus’ and
bivittatus, in north-eastern parts of the Indian subcontinent, have been
examined. The distinguishing features of the two taxa are perhaps more
clearly expressed in this region than in more distant parts of the range (A.
Stimson, pers. comm., 1986).

253

Python molurus

For comparative purposes, brief information on P. m. molurus is given in the
Distribution and Population sections, and much of the Habitat and Ecology
section is perforce based on the sub-continent, but this taxon is not treated
elsewhere in this account.

Appendix I populations: P. m. molurus

Bangladesh Formerly widespread through all 21 districts, but now depleted
and very restricted in distribution and largely confined to the Sunderbans and
evergreen forests of the south-east (Khan, 1982), although the latter may have
been P. m. bivittatus (Kock and Schroder, 1981).

India Ranges virtually throughout, although reportedly extirpated from many
former localities (Whitaker, 1978).

Nepal No records are available from west or east Nepal, the species appears
to be restricted to central areas (Swan and Leviton, 1962).

Pakistan Records are restricted to Sind in the south-east, in the Indus
delta and lower valley (Minton, 1966) mostly east of the river, northward at
least to the Nawabshah district (Pakistan CITES MA, 1986); also in the
Tharparkar desert area (Ghalib et al., undated); the species may extend to
the Punjab (cited from the Punjab area of pre-partition ‘India’ by Smith,
1943).

Sri Lanka Occurs widely in the low country, occasionally ascending into the
hills (De Silva, 1980).

Appendix II populations: P. m. bivittatus
Bangladesh Reported from the CHittagong region (Kock and Schroder, 1981).

Brunei No specific records available, probably present if present at all on
Borneo (see under Indonesia).

Burma Records extend south to the Tavoy district (Smith, 1943).

China Occurs in mountain forests in Fujian, Guangdong (including Hainan
island), Guangxi, Guizhou and Yunnan (Anon., 1980).

Hong Kong Reported from many parts of the country, most frequently from
widely distributed sites on Hong Kong island (Romer, 1979)

Indonesia Present on Java (and adjacent islands east to Sumbawa) and
Sulawesi; also reported from Borneo (de Haas, 1950) and may thus occur in
Kalimantan, although the species's occurrence in Borneo is doubted by one
authority (R.F. Inger, in litt., 5 March 1986).

Kampuchea Widely distributed over most of the country (Saint Girons, 1972).

Laos Reportedly found in all provinces, although more rare _ than
P. reticulatus (Deuve, 1970), and more common in the south (Lao P.D.R.
Forest Department, in litt., 31 January 1986).

Malaysia There appears to be no confirmed record of the species from any
part of Malaysia. The occasional old report exists for west Malaysia, but any
such individuals are likely to have escaped from snake charmers (Flower, 1899;
Tweedie, 1983). There appear to be no specific records for Sabah or Sarawak;
although the species has been reported to occur in Borneo (Smith, 1943; de
Haas, 1950), and might thus be expected in Sabah or Sarawak, its occurrence in

254

Python molurus

Borneo is doubted by one authority (R.F. Inger, in litt., 5 March 1986).

Thailand Recorded from the provinces of Raheng, Lopburi and Chonburi

(Smith, 1943); also said to be found throughout the country, except for the
southern provinces (Soderberg, 1965).

Viet Nam Widespread in southern Viet Nam although absent from southern
parts of the delta (Campden-Main, 1970); no information for the northern half
of the country but quite probably similarly widespread.

POPULATION Virtually all information available on population levels or

trends is anecdotal in nature; appropriate fieldwork is required to generate
some quantitative data.

Appendix I populations: P. m. molurus

Bangladesh Uncommon generally but common in the Sunderbans (Khan, 1982;
Bangladesh CITES MA, 1986). Widely distributed and present in all 21
districts about three decades ago, but due to habitat changes there have been
only two records of sightings outside government controlled forests in the
past decade; small populations survive in evergreen forest in the east and
good populations remain in the Sunderbans (Khan, 1982).

India Widely distributed but heavily exploited and locally extirpated in
many areas (Whitaker, 1978), possibly in most of its former range, remains
common in certain locations in Rajasthan, Andhra Pradesh and Uttar Pradesh
(and perhaps elsewhere) (Whitaker and Whitaker, 1983); also cited as in severe
decline, extremely rare outside protected areas, considered a threatened
species (Tikader, 1983).

Nepal Regarded as an endangered species in the Kingdom, but common in the

grasslands of Chitwan National Park, in particular at Kans and Dhadi (Dhungel,
1985).

Pakistan Cited as threatened (Ghalib et al., undated). Populations have
not yet been surveyed, but they are thought to have been very much depleted
(Pakistan CITES MA, 1986). i

Sri Lanka Said to be one of the more common snakes of the country (Taylor,
1950), or common in the low country (De Silva, 1980).

Appendix II populations: P. m. bivittatus
Burma Pythons (sources do not distinguish between molurus and
reticulatus) appear to be widespread and reasonably common, although were
becoming rare in some areas even by the early 1900s (Salter, 1983).

China Said to be rare, except, perhaps, on Hainan (Pope, 1961).

Hong Kong Not common anywhere in the country (Romer, 1979). One source
(Hong Kong CITES MA, 1985) estimates a population of between 50 and 200
individuals.

Indonesia No information.

Kampuchea Although not rare, not really abundant, perhaps because of
hunting for food or commerce (Saint Girons, 1972).

Laos Reported more rare than P. reticulatus (apparently not uncommon)
(Deuve, 1970). Both this species and P. reticulatus are said to have

255

Python molurus

declined significantly due to local utilisation and the export trade (Lao
P.D.R. Forest Department, in litt., 31 January 1986).

Malaysia No data.

Thailand No details available. Literature records are from lowland areas,
which in some cases may not have good snake populations; animal dealers are
thought still to obtain specimens (W.Y. Brockelman, in litt., 14 February
1986)

Viet Nam Common in the southern parts (Campden-Main, 1970); no information
for the north.

HABITAT AND ECOLOGY A large and lethargic, heavy-bodied, diurnal and
nocturnal snake, capable of attaining a total length of around six metres
(18-19 £t) (although few, if any, such individuals are likely to exist at
present). May be found in a variety of habitats, but appears to prefer wooded
areas -— ranging from evergreen forest to more open deciduous woodland; known
localities often include rock outcrops or hollow trees used for shelter and
nesting, and typically will include still or flowing permanent water. The
species may also be found in the vicinity of rivers, lakes or marshy areas,
often in open semi-arid country, and in reed beds and mangrove stands. An
able swimmer, capable of remaining submerged for many minutes, and an able
climber, often ascending trees to seek prey or to ambush prey while concealed
among branches. Prey includes a wide variety of mammals, birds and reptiles.
Although mammals as large as deer, gazelle and leopard have been taken, small
mammals - rats in particular - appear to comprise the bulk of the diet (there
is growing awareness of their importance as rodent control agents in
agricultural areas).

In India, mating occurs during December-February (the colder season); the
clutch of 8-100 eggs, about 6 x 12 cm in size, is laid three to four months
later, in the hot weather months of March-June. The female incubates the eggs
which hatch around 58 days later. Hatchlings may measure near 75 cm and grow
rapidly in their first years. Maturity is attained at around five years of
age and a little over three metres length. Captive specimens have lived for
22 years.

THREATS TO SURVIVAL This species is used for food by a variety of
indigenous peoples throughout its range (Wall, 1912), and has been exported,
from India to China for example, for food and medicinal purposes (Irvine,
1954). In the 1950s python meat on sale in Hong Kong was more expensive than
beef (Irvine, 1954). Python fat and various organs are often attributed
medicinal properties, apparently by Chinese and tribals in particular; the
gall bladder is especially prized in parts of Burma for example (Wall, 1912).
Local utilisation for food is unlikely to pose a serious threat, but it has
been suggested that the extent of demand for medicinal and other purposes in
Chinese-speaking areas should be investigated. However, the primary threat is
exploitation for the skin trade, which appears to exist, often at an
unsustainable level, almost throughout the range of the species.

Bangladesh There is said to be no internal or external trade in this
species (Bangladesh CITES MA, 1986).

Pakistan The species was formerly ruthlessly hunted for its skin but this
is now illegal (Pakistan CITES MA, 1986).

INTERNATIONAL TRADE This species has long been prominent in the live animal
trade, being large, attractive and readily kept in captivity; captive breeding
is also relatively straightforward.

256

Python molurus

Table la Minimum net imports of skins of P. molurus reported to CITES.

ee

1980 1981 1982 1983 1984 1985
ae es ES et ey |
Australia - 30 - 33 - -
Austria - 160 47 140 336 150

- ~ - 14 m - -
Belgium - - - = 766 706
= = = - 248 m -
Canada 14 - 102 78 16 148
= = - - 58 m -
China 839 - = = = =
Cyprus - - - 38 m - -
Denmark 386 - = ~ = =
Egypt ~ - 81 - - -
France - - - ~ 23839 54658
- 2890 m - - 28000 m 7500 m
German D.R. - 5945 3000 m - - -
Germany, F.R. 11851 21200 671 5701 6247 6807
- - - - 37945 m 7603 m
- - - - - 80 kg
Greece - - 1 96 - 545
Hong Kong 10500 m - 1623 5499 - 3140
Hungary - - ~ 50 - 7
Israel - 20 5 34 16 18
Italy 79 5210 13598 22255 7566 62650
12000 m 7060 m 45477 m 180525 m 80226 m 28414 m
= 52869 m? Z Z = =
Japan 1847 — 409 5391 4060 18148
- - 134 kg 1736 m 1783 kg -
Korea, Rep. of - - - - 189 -
Lebanon - - - 10 30 -
Mexico ~ 64 - - 3333 893
- - - - 396 m -
Netherlands - - ~ 1399 726 1394
= = = - 7000 m 12150 m
New Zealand - ~ i 15 115 -
Saudi Arabia - - - 30 3 -
Singapore - - - - 5355 -
South Africa - 639 1 25 - 24
Spain - 348 83 15 6855 13858
- 2200 m - - 257 m -
Sweden 638 - - = = =
Switzerland 1 2m™ 1523 693 m 723 -
Taiwan - = 7 - 1080 450
= = = - 2000 m -
Turkey = - - - 22 100
= = = - - 583 m
UK 10000 m 16839 m 2136 3737 m 5000 m -
USA 6411 22320 16850 6204 20769 25372
19667 m 32341 m 5846 m 4401 m 17453 m 6404 m
- = 241 kg - 96 kg =
Unknown - 2260 m 14 - = =
Total 22066 55936 37262 46975 82046 189068
52167 m 63592 m 54323 m 191144 m 178583 m 62654 m
2 52869 m2 375 kg = 1879 kg 80 kg

257

Python molurus

Table 1b Minimum net imports of live P. molurus reported to CITES.

1980 1981 1982 1983 1984 1985
Argentina - - 1 5 - 5
Austria ~ - 73 12 - 14
Belgium - = - - 40 10
Canada 119 44 21 73 65 79
Czechoslovakia - - - - - 15
Denmark 40 - - - . 388 525
Finland i - 5 - - -
France - 13 - 17 9 82
German D.R. 15 - - - 6 1
Germany, F.R. 770 598 672 794 700 2001
Greece - - - 1 4 4
Hungary - - - 4 - 1 -
India - - = 2 - -
Israel - 10 - - - -
Italy 28 167 119 160 42 194
Japan - - - 246 326 1182
Korea, Rep. of - - - - - 4
Kuwait - - - - 12 10
Liberia - 4 - - - -
Mexico 8 - 25 - - -
Netherlands ~ ~ - - 601 1762
Poland - - - 2 - -
Portugal - - - 2 16 112
Qatar - - - 4 - -
Saudi Arabia - - - - 3 55
Singapore - - - - - 6
South Africa 1 - - - ~ -
Spain - - - 15 11 48
Sri Lanka - - - 4 - -
St Helena - - - - - 3
Sweden - - - 6 - 4
Switzerland 109 51 37 - - 1030
Tunisia - - - - 1 -
UAE - - 2 - 9 -
UK 97 521 798 631 1150 1421
USA 5360 3465 5486 5363 12626 16663
USSR - - - - 3 25
Unknown 5 - - - - -
Total 6553 4873 7239 7337 16013 25255

The only reports of international trade in P. molurus are those contained in
the Annual Reports of Parties to CITES. Most were reported = as
P. m. bivittatus, but a few were simply described as P. molurus. Only
trade in live animals and skins was considered. The volume of the skin trade
is shown in Table la. Some transactions were reported in terms of length,
weight or area; it is difficult reliably to convert these to numbers of skins,
but Fuchs (1975) suggested 1.7 m as the mean length of P. molurus skins in
trade. It must be recognised that this can only lead to an approximation of
the number of individuals involved, but on this basis the volume of trade
increased steadily from 52 752 in 1980 to 225 923 in 1985, ignoring
transactions reported by weight or area. Italy, France, the USA and F.R.

258

Python molurus

Germany were the main importers. Some of the apparent increase is artificial,
because France, a major importer in 1984 and 1985, did not report imports of
Appendix II material before 1984. The declared sources of the skins are given

in Table 2a. from which it appears that Thailand, Indonesia and Viet Nam were
the major suppliers.

Table 1b shows the volume of trade in live animals, which also showed a
dramatic increase, from 4873 in 1981 to 25 255 in 1985. The USA was the major
importer, but the UK, F.R. Germany, Japan, Switzerland and the Netherlands
also imported significant quantities. Almost all of the live _ snakes
Originated in Thailand (see Table 2b).

The effects of the trade cannot be satisfactorily evaluated in the absence of
any adequate population data, but the levels must be viewed with concern,
particularly as they seem to be increasing steadily.

Table 2a Reported countries of origin (or exporting country if no origin
reported) and numbers of skins of P. molurus bivittatus reported to CITES.
Transactions reported by length were converted to numbers using an average
length of 1.7 m.

1980 1981 1982 1983 1984 1985

Countries having or possibly having populations of P. m. bivittatus

China - 280 - - 233 -
Hong Kong - - - - - 471
Indonesia 1244 11665 4724 19494 10887 3184
Malaysia 104 770 - - - -
Thailand 19772 228772 53500 128321 159451 224315
Viet Nam 28654 25682 7062 27784 17075 1192
Burma - - - 26 - -
‘Asia’ - - - - il -
Countries without wild populations

Argentina 311 92 - - - -
Austria - - - = =
Cameroon - - - - - -
Canada 3 - - - - =
Denmark - - - = = =
Ghana — - - - 4 -
India - 4 - - = =
Italy - 57 - = = =
Japan - 160 - = = =
Netherlands 1176 - = - = =
Nigeria - - - - 27 =
Singapore 5 15334 1731 654 1026 1363
Spain 1 - = = = =
South Africa - 3 254 5 - -
UK - - 102 1 - ~
USA - = = 34 = =
Taiwan = = - 1 1690 =
Venezuela =— - 6 35 - -
Unknown 10598 6195 9176 6370 4099 94

259

Python molurus
Table 2b Reported countries of origin (or exporting country if no origin
reported) and numbers of live P. molurus bivittatus reported to CITES.

1980 1981 1982 1983 1984 1985

Countries having or possibly having populations of P. m. bivittatus

Burma - 3 - - - 2
Indonesia - 2 - - - -
Malaysia - - - 2 113 94
Thailand 6508 4847 1464 7334 15935 24903
Viet Nam - - - - - 308
‘Asia’ - 4 - - - -

Countries without wild populations

Austria - 2 1
Cameroon - - -
Canada - 13 11
Denmark - - -
German D.R. - - -
Germany, F.R. 24 13 3
Guatemala 2 - - - - -
Poland - - - - 4 4
South Africa - 4 - - -

Switzerland - - - - 15 10
Togo - 20 175 - - -
UK - - - - 2 -
USA 1 - 29 - 39 2
Unknown 2 3 3 - 7 3

fFuw!l YW
!
!

CONSERVATION MEASURES

Bangladesh The skins of pythons are mentioned in Schedule II of the
Wildlife (Preservation) Order, 1973. Their possession, transfer or import
requires a certificate of lawful posession.

Hong Kong Listed as Protected under the Wildlife Protection Ordinance No.

5, 1976, and as such hunting, killing and possession is prohibited (Hong Kong
CITES MA, 1985).

India Pythons are listed on Schedule I of the Widlife Protection Act, 1972,
and are thus totally protected.

Indonesia Protected since 1978 in Indonesia (Indonesia CITES MA, 1986).

Pakistan The species is protected, and may not be killed, trapped or traded
(Pakistan CITES MA, 1986).

Thailand Listed on Schedule 2 (protected wild animals of the second
category) of the Wild Animals Reservation and Protection Act B.E.2503 in
Thailand, effective 11 November 1985.

CAPTIVE BREEDING This species is very commonly held by zoos and pet
keepers, and is frequently bred in captivity. A recent inventory (Slavens,

260

Python molurus

1985) records 77 P. m. molurus in 21 collections and 198 P. m. bivittatus
in 56 collections in 1984; total numbers in captivity will certainly be much
higher than these figures. Second generation captive breeding has been
recorded at Berlin, Milwaukee, Pilsen and Little Rock zoos (Olney, 1984).

Reportedly reared (though not bred) until of commercially viable size for the
skin trade at several crocodile farms in South East Asia, including Singapore
(Platt, 1985).

REFERENCES

Anon. (1980). Atlas of Chinese Snakes. Technical Publishing House of
Shanghai University.

Campden-Main, S.M. (1970). A Field Guide to the Snakes of South Vietnam.
Smithsonian Institution, Washington.

De Silva, P.H.D. (1980). Smake fauna of Sri Lanka with special reference to
skull, dentition and venom of snakes. Spolia Zeylanica 34(1,II): 1-142.

Deuve, J. (1970). Serpents du Laos. Memoire O.R.S.T.O.M. No. 39, Paris.

Dhungel, S.K. (1985). Ordeal with a Rock Python. Tigerpaper 10(3): 7-8.

Flower, S.S. (1899). Notes on a second collection of reptiles made in the
Malay Peninsula and Siam. With a list of the species recorded from those
countries. Proceedings of the Zoological Society of London 1899:
600-697.

Fuchs, K. (1975). Chemistry and technology of novelty leather. FAO, Rome,
201 pp.

Ghalib, S.A., Rahman, H., Iffat, F., and Hasnain, S.A. (undated, post-1980?).
A checklist of the reptiles of Pakistan. Records of the Zoological
Survey of Pakistan 8(1-2): 37-59.

Haas, C.P.J. de, (1950). Checklist of the snakes of the Indo-Australian
Archipelago. Treubia 20(3): 511-625.

Irvine, F.R. (1954). Snakes as food for man. British Journal of Herpetology
1(10): 183-189.

Khan, M.A.R. (1982). On the endangered snakes of Bangladesh. In, Proceedings
of the IUCN/SSC Snake Group First Meeting, 8-12 November, 1982. Madras
Crocodile Bank.

Kock, D. and Schrodder, H. (1981). Die Gattung Python in Bangladesh.
Salamandra 17(3/4): 112-118.

Minton, S.A. (1966). - A contribution to the herpetology of West Pakistan.
Bulletin of the American Museum of Natural History 134: 27-184.

Olney, P.J.S. (Ed) (1984). Reptiles bred in captivity and multiple generation
births, 1981. In, International Zoo Yearbook 23. (See also vols. 21,
22).

Platt, CG. (11985). Regional Director's field trip to Asia 1985 (summary
report). Animals International (WSPA) 5 (17): 4-5.

Pope, C.H. (1961). The Giant Snakes. Routledge & Kegan Paul, London, 297 pp.

Romer, J.D. (1979) Second revised annotated checklist with keys to the snakes
of Hong Kong. Memoirs of the Hong Kong Natural History Society No. 14:
1-23.

Saint Girons, H. (1972). Les Serpents du Cambodge. Memoires museum National
d'histoire Naturelle, Paris, N.S., Ser. A (Zool.), 74.

Salter, R.E. (1983). Summary of currently available information § on
internationally threatened wildlife species in Burma. F.0O.: BUR/80/006.
Field Document 7/83. FAO, Rangoon.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity,
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

Smith, M.A. (1943). Reptilia and Amphibia, Vol. III, Serpentes. In, The
Fauna of British India. Taylor and Francis, Ltd., London. Reprinted
1973, Ralph Curtis Books, Florida.

Soderberg, P. (1965). The pythons of Thailand. Conservation News S.E. Asia
5: 11-12.

261

Python molurus

Stimson, A. (1969). Boidae. Das Tierreich, Walter de Gruyter, Berlin 89:
1-49.

Swan, L.W. and Leviton, A.E. (1962). The herpetology of Nepal: a history,
check list, and zoogeographic analysis of the herpetofauna. Proceedings
of the California Academy of Sciences 32(6): 103-147.

Taylor, E.H. (1950). The snakes of Ceylon. University of Kansas Science
Bulletin 33(2): 519-603.

Tikader, B.K. (1983). Threatened Animals of India. Zoological Survey of
India, Calcutta, 307 pp.

Tweedie, M.W.F. (1983). The Snakes of Malaya (3rd edn.). Singapore National
Printers (Pte) Ltd.

Wall, F. (1912). <A popular treatise on the common Indian snakes. Number 17,
Python molurus. Journal of the Bombay Natural History Society 21:

447-475.
Whitaker, R. (1978). Common snakes of India. Macmillan and Co., Madras,
154 pp.

Whitaker, R. and Whitaker, Z. (1983). Herpetological conservation in India.
Paper presented at Bombay Natural History Society Centenary Seminar.
December 1983. (Proceedings in press, 1986).

262

RETICULATED PYTHON Recommended list: 2

[Possible problem]
Python reticulatus (Schneider, 1801)

Order SERPENTES Family BOIDAE

SUMMARY AND CONCLUSIONS A potentially very large species. Widespread in
South East Asia from south-east Bangladesh east through Burma, Thailand,
Kampuchea and Laos to Viet Nam and south through the Malay Peninsula to
Singapore, thence east through much of the Indo-Australian Archipelago and the
Philippines. Present in the Nicobar Islands (India). Generally regarded as
common, or even very common, around the turn of the century. Little recent
information is available but the species is still reportedly common in parts
of the range (Malaysia, Thailand, Viet Nam); no information is at hand for
most of the range, and no sound data on population status or trends for any
part of it. Often a forest species and typically occurs in the vicinity of
water. Also present in secondary growth and quite frequently near human
settlements, presumably attracted by the availability of chickens and other
domestic stock. Females of four metres length typically lay a clutch of 30-40
eggs.

Widely used for food and medicinal purposes, particularly by tribal and
Chinese-speaking communities. Heavily exploited by the live animal trade, and
very heavily by the skin trade. The volume of the international skin trade
appears from the CITES reports to have increased from 166 000 in 1980 to
677 000 in 1985. Italy, the USA and F.R. Germany were the main importers and
Indonesia, Thailand and Malaysia were the major suppliers. The trade in live
animals also showed a dramatic increase, from 4909 in 1981 to 16 613 in 1985.
The USA was by far the major importer and the great majority of live snakes
originated in Thailand.

The sustainability of the recent trade cannot be evaluated in the absence of
Significant population data; such data should be gathered as a matter of
priority. It must be suspected that present trade is excessive and it is
particularly worrying that it seems to be increasing.

DISTRIBUTION Widespread in the Indo-Malayan region. Extends from south-
east Bangladesh eastward almost throughout mainland South East Asia, and south
through Peninsular Malaysia to Singapore (Smith, 1943). The range extends
across most islands of South East Asia, including most of the Philippines, and
most of Indonesia. Also present in the Nicobar Islands (a Union Territory of
India).

Bangladesh Restricted to the Chittagong and Sylhet areas (Khan, 1982).
Brunei Recorded as present (Ussher, 1979); no details available.

Burma One source suggests that the species is restricted to southern parts
of Burma, but others suggest that pythons (without differentiating between
reticulatus and molurus) are widespread in the country (Salter, 1983).

India Confirmed records appear to be restricted to the Nicobar islands in
the Union Territory of the Andamans and Nicobars; recorded from Car Nicobar,
Teressa, Trinkat, Nancowry, Great Nicobar and Little Nicobar (Whitaker and
Whitaker, 1983). Other sources (Daniel, 1984; Tikader, 1983) state that the
species also occurs in mainland India, in the extreme north-east, but these
reports seem to be unconfirmed.

263

Python reticulatus

Indonesia Widespread, west to Sumatra and the adjacent Mentawai group, and
east to Halmahera, Seram and Tanimbar (but not to New Guinea) (De Rooij, 1915;
de Haas, 1950). Present in Kalimantan, Sulawesi, Java, Flores, Lombok,
Natuna, Tanimbar, Sumba and Sumbawa (Indonesia CITES MA, 1986).

Kampuchea There appear to no recent records (Saint Girons, 1972), although
the species would be expected to occur throughout (Smith, 1943).

Laos Found in all provinces, although more common in the centre and south
(Deuve, 1970).

Malaysia No details available for Malaysia, but apparently occurs in
suitable habitat throughout the peninsula (B.H. Kiew, in litt., 25 February
1986); also present in Sabah, where recorded in forested areas throughout
(Malaysia, Sabah CITES MA, 1985), and in Sarawak.

Philippines Widespread, including all the larger islands (Leviton, 1963).
Singapore Present (Ridley, 1899), no details available.

Thailand Reportedly restricted to below 18°N (Smith, 1943), although others
(Taylor, 1965) regard the species' absence from the north as "possible" or,
(Soderberg, 1965), state that it occurs commonly in all provinces.

Viet Nam Occurs throughout southern Viet Nam (Campden-Main, 1970) and
extends north through most of the country at least to Yen Bai (near Hanoi)
(Smith, 1943). Generally more frequent in the south than the north in
Indochina as a whole (Bourret, 1936).

POPULATION The little information available on wild populations, often
dating from around the turn of the century, is very generalised or anecdotal
in nature. No sound data are available on population levels or trends in any
part of the species’s range; appropriate fieldwork is required.

Bangladesh While always confined to rain forests in the east and southeast,
habitat destruction has reduced and fragmented the range and only two small
populations remain, in Sylhet and Chittagong (Khan, 1982). Reportedly common
within its restricted range (Bangladesh CITES MA, 1986).

Brunei No data.

Burma No specific data; pythons (both P. molurus and P. reticulatus)
reportedly widespread and reasonable common, although with local declines
already noted after the turn of the century (Salter, 1983). Wall (1916)
reported that the species was regarded as common in Pegu, not uncommon in
Tenasserim (more common than P. molurus), but rare in lower Burma generally.

India Restricted distribution, numbers unknown (Whitaker and Whitaker,
1983); officially considered threatened (Tikader, 1983).

Indonesia One early twentieth century source cites this species as very
common (De Rooij, 1915), no recent information available.

Kampuchea No data
Laos Both this species and P. molurus are said to have declined markedly

in Laos due to local utilisation and the export trade (Lao P.D.R. Forest
Department, in litt., 14 February 1986).

264

Python reticulatus

Malaysia Cited at the turn of the century as one of the commonest snakes,
pythons of 6 m (20 ft) then being “by no means uncommon" (Ridley, 1899).
Reported very common in the 1950s at Asahan, Malacca (Batchelor, 1958). A
recent popular source (Lim, 1981) states that the species is “still quite
common", but questions how long this situation will persist. Similarly, said
to be still quite common despite exploitation, and can be readily seen (B.H.
Kiew, in litt., 25 February 1986). Still abundant in Perlis, within the
security area of northern peninsular Malaysia (S. aAmbu, in Jlitt., 17
February 1986), but disturbance, habitat loss, persecution and exploitation
for food reportedly causes appreciable mortality in other parts of the
peninsula, where, by implication, the species may often be less than abundant.

Reported common in Sarawak at one time (Shelford, 1916), and said to be still
widespread and common (H. Watson, in Jitt., 17 March 1986). During
intensive herpetological fieldwork at three primary rainforest sites in
Sarawak - Nanga Tekalit (366 days), Labang (128 days), Sengai Pesu (160 days)
- the reticulatus seen numbered only 8, 10 and 4, respectively (R.F. Inger,
in litt., 5S March 1986). Population levels unknown in Sabah, although
P. reticulatus is more common then P. curtus (Malaysia, Sabah CITES MA,
1985).

Philippines No data

Singapore Said to be “still far from rare” on Singapore in 1922, when
several specimens were captured within Municipal limits (Sworder, 1922), and
five individuals were included in a collection of snakes made over seven
months, chiefly around the Nee Soon Forest Reserve (Harman, 1961). Apparently
still relatively common (P. Gopalakrishnakone, in litt., 13 March 1986),
although no details of distribution or abundance are available.

Thailand Said to occur “commonly" in all provinces (Soderberg, 1965). On
Phuket Island (Frith, 1977) the species is apparently still common although
larger individuals are now very rarely seen because of human predation.
Numerous specimens can be seen in the possession of local residents who
collect them to sell for skins (Frith, 1977). Seen occasionally in Khao Yai
National Park, and more often in Khao Soi Dao in southeast Thailand (in
slightly wetter forest); probably present in all evergreen forests and
apparently relatively secure (W.Y. Brockelman, in litt., 14 February 1986).
At the end of the nineteenth century, said to be very common in the city and
suburbs of Bangkok (Flower, 1899); this is presumably no longer the case.

Viet Nam Said to be common throughout the south (Campden-Main, 1970). Half
a century ago reportedly not rare in the Indochinese region in general
(Bourret, 1936).

HABITAT AND ECOLOGY A potentially very large boid snake, the Reticulated
Python is perhaps the only snake which regulerly exceeded 6 m in length (Wood,
1982) - enormous old individuals of 8-9 m have been reported on occasion -
but the largest specimens living at present are unlikely to approach such
sizes, and an ‘average’ specimen would be perhaps 3-4 m long.

Said to be fond of water, and rarely to be found far from it (Smith, 1943;
Tweedie, 1983), the Reticulated Python often occurs in the vicinity of forest
rivers and streams, but may also be found around rice fields, and sometimes in
water-less rocky hills (Bourret, 1936). In southern Viet Nam, for example,
the species is reportedly "always" found near water, and in particular, often
found under bridges over rapidly flowing streams (Campden Main, 1970). Whilst
the species is said to favour dense forest (Lim, 1981), jungle growth
(Soderberg, 1965), or (in Burma) the most dense and least frequented jungle
(Wall, 1926), it also occurs in plantations or secondary growth (B.H. Kiew,

265

Python reticulatus

in litt., 25 February 1986), and Flower (1899) even stressed that it was
once common in Bangkok city and appeared to prefer the busiest parts of the
riverside.

The species is said to spend much of the day in hiding, often climbing trees
for the purpose, and to seek prey at night, often small or medium sized
mammals such as monkeys, civet cats, pangolins, small deer and pigs, but also
birds (Lim, 1981; Wall, 1912). Rats, mice, and domestic stock, especially
chickens, are taken; for this reason pythons are often found close to human
habitation and persecuted as a result.

In captive animals, mating occurs around the turn of the year, with egg
laying, a single clutch, usually between April and October (Wall, 1926), about
two months after the last mating. Clutch size tends to be greater with
increasing size and age of the female; extremes of 8 and 103 have been
recorded (Lim, 1981; Reitinger, 1978). Females 4m in length typically lay
30-40 eggs, about 64 x 118 mm. Pope (1961) cites a report of two nests in
Sumatra, one in a hollow log, the other in a hole under bamboo roots. The
female coils around the eggs and broods them for the entire incubation period
of 2.5-3 months, although she may leave them to drink and slough. Newborn
snakes are about 60 cm long. The youngest female known to lay viable eggs was
five and a half years old. Captive specimens have a maximum recorded age of
21 years.

THREATS TO SURVIVAL Widely used in South East Asia for food and medicinal
purposes, especially by Chinese communities, also by indigenous populations
(Ridley, 1899; Irvine, 1954).

Indonesia Python skins are used for leather crafts. Authorised exports of
this species were 37 590 in 1981, 67 377 in 1982, 75 775 in 1983 and 170 470
in 1984 (Indonesia CITES MA, 1986). These are considerably lower than imports
reported from Indonesia (see Table 2a, below).

Laos Both this species and P. molurus are said to have suffered from
habitat destruction and hunting for skins and food. The number of pythons
used locally as pets may equal those supplying the export trade (Lao P.D.R.
Forest Department, in litt., 14 February 1986).

Malaysia There is a low level of local exploitation in Sarawak: each rural
village may consume half a dozen pythons (up to 7 m long) each year (J.
Caldecott, in Jlitt., 30 January 1986), and there appears to be only one
restaurant in Kuching that regularly serves reticulatus (E. Bennett, in
litt., 5 February 1986). Although the species is caught and eaten or sold
whenever encountered in Sarawak, it remains widespread and common (H. Watson,
in litt., 17 March 1986). However, large numbers of Reticulated Pythons
appear in markets in West Malaysia, and python meat is readily available
throughout. The species is also suffering from habitat loss and general
persecution as rapid development proceeds. However, there are indications
that the demand for python meat has decreased recently, in parallel with an
increase in demand for meat, blood and other products of believed medicinal
value from venomous snakes (S. Ambu, in litt., 17 February 1986). Other
sources (B.H. Kiew, in Jitt., 25 February 1986) do not regard local
exploitation for food in West Malaysia as excessive. In Sabah, the Chinese
community uses python meat particularly for soup, and occasionally the skins
are used for decorative purposes. The levels of exploitation are thought to
be low, and capture for the pet trade is thought to be insignificant (Sabah
CITES MA, 1985).

Philippines Permits were granted retrospectively for the capture of 4842 m
of P. reticualtus skin in 1981, most of which was subsequently exported. A

266

Python reticulatus

total of 5 animals were authorised to be captured live from 1981 to 1985
(Philippines CITES MA, 1986).

INTERNATIONAL TRADE The only reports of international trade in
P. reticulatus are those contained in the Annual Reports of Parties to
CITES. Only trade in live animals and skins was considered. The volume of the
skin trade is shown in Table la. Some transactions were reported in terms of
length, weight or area; it is difficult reliably to convert these to numbers
of skins, but Fuchs (1975) suggested 1.8m as the mean length of
P. reticulatus skins in trade. It must be recognised that this can only
lead to an approximation of the number of individuals involved, but on this
basis the volume of trade fluctuated between 166 000 and 677 000 from 1980 to
1985, ignoring transactions reported by weight or area. Italy, the USA and
F.R. Germany were the main importers. The volume of trade appeared to
increase substantially from 1980 to 1984, although the level decreased
slightly in 1985. The declared sources of the skins are given in Table 2b.
from which it appears that Indonesia, Thailand and Malaysia were the major
suppliers.

Table 1b shows the volume of trade in live animals, which also showed a
dramatic increase, from 4909 in 1981 to 16 613 in 1985. The USA was by far
the major importer, but the UK, Italy and F.R. Germany also imported
significant quantities. The great majority of live snakes originated in
Thailand, with lesser numbers from Indonesia and, formerly, India.

The sustainability of this volume of exploitation cannot be assessed in the
absence of good population data. Exploitation would seem to be excessive and
increasing; although the species is said to retain healthy populations in many
areas, little is known of populations where exploitation is highest.

Table la Minimum net imports of skins of P. reticulatus reported to CITES.

1980 1981 1982 1983 1984 1985

Andorra - - - 19 = a

Argentina - - - 1852 - -

Australia 875 9 26 4002 18 -

- - - 82 m - -

Austria - - 22 85 1051 173

- 3m 112 m 184 m 77 m -

Belgium - - 27 28 646 -

- - 10885 m - 875 m -

Brazil - - - - 4 55

Canada 1568 4938 474 315 452 202

561 m 248 m - 188 m 50 m -

Cyprus - - 45 270 32 -

Denmark 235 233 342 31 45 -

Egypt - 4 - = - -

Finland - - - 19 3 -
- - - - 98 m 150 m

France - - - - 8898 22932
- 6138 m - - 8000 m 2172 m

German D.R. - 329 7142 - - -

Germany, F.R. - 8009 - 636 - 6976
55614 m - - 50913 m 38494 m 176 m

Greece - 442 64 113 1654 456

= = - 179 m 3118 m -

A. x = s 84 m2 =

267

Python reticulatus
Table la Continued.
1980 1981 1982 1983 1984 1985
Hong Kong 1576 - 228 250 4766 5911
- - - 3 3829 m -
Hungary 497 - - 50 - -
Indonesia - - - - - 4m
Israel 34 71 126 10 39 -
- 276 m - - 22 m -
Italy 13185 28251 61939 76866 54397 17399
600 m 4112 m 65570 m 182254 61066 m 41855 m
2 23832 m? 12 kg zs = =
Japan - - - - 7992 70578
= = - - 9271 kg 4 kg
Jordan - - - - 38 -
Korea, Rep. of - - - - 665 8
= - - 483 1851 m 1831 m
Lebanon 105 - - - 61 -
Malta - 550 m - - 100 40
Mexico 727 708 629 318 - 6730
- 7m 54 m - 3508 m 2206 m
221 m@ 44 m2 24 m2 bs 92 m@ a
2 kg 29 kg 329 kg - 340 kg 743 kg
Morocco - - - - - 3
Namibia - - - - - 11
Netherlands - 445 - - 883 308
- - - - 2927 m 11760 m
Neth. Antilles - - - 13 - -
New Zealand 553 700 257 713 1666 -
Norway 20 20 - - 241 1
Panama - - - - - 716
Saudi Arabia - - - 60 - 17
Singapore - - 7758 150779 131046 -
South Africa - 402 m¢ - - - -
Spain 5632 2942 - - 28307 m -
Sweden - - - - 8 10
Switzerland - 13517 m 1083 501 17752 m -
Taiwan - - - 2757 232 3366
Tunisia - - - - 71 71
Turkey - - - 55 148 368
- - - 50 617 m 104m
UAE - - - - 77 -
UK 834 5311 1201 - - -
- 15184 m - 25167 21785 m 56915 m
Unknown 1956 2456 665 4 - -
Uruguay - - - - 300 m -
USA 90038 110577 37623 65696 169789 184179
30429 m 104284 m 39212 m 59087 219624 m 300167 m
1873 kg 97 kg 81 m2 A 2341 m@ 89 m2
Venezuela - - - 13 - -
Total 117835 165445 119651 304954 385022 320510
87204 m 144319 m 115833 m 322864 412300 m 417340 m
221 m2 =. 24278 m2 105 m2 = 2517 m2 89 m2
1875 kg 126 kg 341 kg - 9611 kg 747 kg

Python reticulatus

Table 1b Minimum net imports of live Python reticulatus reported to CITES.
a a a Oe Eee eee

1980 1981 1982 1983 1984 1985
en ee a ee ee ee eee
Argentina 2 = 1 = 3 =
Austria - 4 42 14 18 14
Belgium ~ ~ - - - 10
Canada 126 48 19 29 58 40
Czechoslovakia - - - - - 10
Denmark 59 = = = 70 110
Egypt = - - 4 = -
Finland = = = = 1 =
France - 6 4 12 19 82
German D.R. 20 - = 14 4 2
Germany, F.R. 1022 26 405 660 671 1609
Greece - - = a 432 2
Hong Kong - - = = 15 2
Israel - 10 - 2 - 2
Italy 41 52 118 151 74 36
Japan - - - 156 144 1077
Korea, Rep. of - - - 4 2 -
Kuwait - - - - 14 8
Liberia - 2 - - - -
Mexico 2 - 11 - - -
Netherlands - - - - 2550 872
Portugal - - 1 - 15 126
Romania 2 - - - - -
Saudi Arabia - - 2 1 - 10
Singapore = - 1 - 1
South Africa ~ - - - - 1
Spain ~ 5 6 10 - -
Switzerland 95 93 22 8 - 496
Taiwan - - - 1 - ~
UAE - - - - 1 -
UK 116 284 355 466 1090 1151
USA 5841 4377 4342 4738 6713 10849
USSR 13 - - 4 2 6
Unknown - - - 1 - 99
Uruguay - 2 - = - =
Venezuela ~ - 2 - - =
Total 7339 4909 5330 6277 11896 16613

Table 2a Reported countries of origin (or exporting country if no origin
reported) and numbers of skins of Python reticulatus reported to CITES.

1980 1981 1982 1983 1984 1985

Countries having or possibly having wild populations of P. reticulatus.

India 21 510 a 900 900 3971
- 1334 m 1000 m - 254 kg -
= 1000 m2 = = 40 m ne!
Indonesia 13811 72646 68404 242922 265639 176849
5329 m 55248 m 56975 m z 203441 m 273355 m
254 m2 15333 m2 2 a 2433 m2 =
2 kg 125 kg = = 8447 ke 370 kg

269

Python reticulatus

Table 2a Continued.

1980 1981 1982 1983 1984 1985
Malaysia 19069 2771 14 2751 4642 3775
- 150 m - - - -
- 29 kg 213 kg - 221 kg 271 kg
Philippines 497 1435 188 2144 480 2198
286 m 3069 m 600 m - 1362 m 5349 m
= 500 m? = = z= ZI
Singapore 37702 62104 26535 36452 8754 90746
37535 m 55083 m 20745 m - 712 m 918 m
2 800 m2 125 kg = 84 m2 as
Thailand 36717 88025 31146 116110 119690 134959
37954 m 49115 m 32172 m - 86368 m 135431 m
= 6796 m@ 24 m2 © a 52 m? 89 m2
= = 116 kg = 878 ke 75 kg
Viet Nam - - 207 690 441 4 kg
Countries without wild populations of P. reticulatus.
Argentina - - - 15 4 56 kg
Canada - - - - - 520 m
Cameroon - 2 - - - -
Chad - - - - - 500 m
China - - - - 160 -
Denmark - - - - 1 8
France - - - 630 100 -
German D.R. 170 - - - - -
Germany, F.R. 9447 3972 - - - -
- 12001 m - - - -
Italy 4 - - - - -
Japan 873 1066 - 1212 8341 1
5000 m 251 m 329 m - 215 m -
- - - - 77 kg 387 kg
Nepal - - - 1265 - -
Netherlands 1179 - - - - -
Nigeria 1740 66 3 - - 5
- 61 m - - - -
Panama - - - - 77 707
Paraguay - - - 7 - -
South Africa 1718 1055 23 471 - -
- 1623 m - - - -
= 31 m2 = 2 * =
Spain 2 - - - - -
Sudan - - - 1 - 104
Switzerland - 259 23 - 8 -
- 466 m - - - -
Taiwan 100 18 18 1654 807 202
- 4074 m - - - -
Tanzania 23 - - - - -
Togo - - - 466 - -
UK 16 334 - 226 66 18
1720 m 891 m - - - -
USA - 350 - 13 49 8
Unknown 28509 21607 32461 57121 16853 3122
4380 m 6257 m 13832 m - 133034 m 2077 m
37 m2 > 104 m2 x a s
1873 kg - 15 kg - - -

270

Python reticulatus

Table 2b Reported countries of origin (or exporting country if no origin

reported) and quantities of transactions in live Python reticulatus reported
to CITES.

1980 1981 1982 1983 1984 1985

ee ee ee
Countries having or possibly having wild populations of P. reticulatus.

Burma - - - -
Indonesia -
India 1878 - a
Malaysia 513 161 37 101 449 114
Philippines ~ - - 1 1 3
Singapore - -
Thailand 6346 4518 5255 5771 9619 15751

Countries without wild populations of P. reticulatus.

‘Asia' =
Australia 4 =
Canada 1 1

German D.R. - -

Germany, F.R. 10 4 = 4 4 s
Ghana - 6

Guyana 1 - - = = =
Honduras 2

Japan - - - 2 = =
Netherlands 12 - 1 = = =
Poland - = = = 10 =
Sweden - = = = 1 x
Switzerland 1 - - - 15 7
Togo - 75 25 10 - 294
UK - 2 - = = a
USA 2 - 22 - 27 1
USSR = =
Unknown 475 5 US 19 5 1

CONSERVATION MEASURES

Bangladesh The skins of pythons are mentioned in Schedule II of the
Wildlife (Preservation) Order, 1973. Their possession, transfer or import
requires a certificate of lawful posession.

India Pythons are listed on Schedule I of the Widlife Protection Act, 1972,
and are thus totally protected.

Indonesia Harvest is "controlled" in Indonesia (Indonesia CITES MA, 1986).
Quotas for 1987 were set to total 140 000 skins, divided between the regions
as follows: Acoh, 6900; Sumut, 11 500; Sumbar, 4100; Riau, 8000; Jambi, 6000;
Bengkulu, 7000; Sumsel, 12 000; Lampung, 4500; DKI, 2400; Jabar, 1000; Jateng,
100; Jatim, 1000; Kalteng, 17 000; Kalsel, 7000; Kaltim, 30 000; Sulut, 2500;
Sulteng, 1000; Sulsel, 10 000; Sultera, 1500; NTT, 500; Timtim, 500; Maluku,
2500; Irja, 3000 (Indonesia CITES MA, 1987).

Malaysia Protected in Peninsular Malaysia (1972 Protection of Wild Life
Act) but not in Sabah or Sarawak (E. Bennett, in litt., 5 February 1986).

271

Python reticulatus

Singapore All wild fauna in Singapore is fully protected (Singapore Primary
Production Department, in litt., 11 January 1986).

Thailand Recently protected in Thailand under the Wild Animals Reservation
and Protection Act B.E.2503, effective 11 November 1985.

CAPTIVE BREEDING Widely held in public and private zoological collections;
a recent inventory (Slavens, 1985) reports 105 individuals in 40 collections.
Has bred quite frequently in captivity, in four zoos in 1981, with second
generation breeding at Pilsen, Czechoslovakia (Olney, 1985).

REFERENCES

Batchelor, D.M. (1958). Some notes on the snakes of Asahan, Malacca.
Malayan Nature Journal 12(3): 103-111.

Bourret, R. (1936). Les Serpents de 1'Indochine, II, Catalogue Systématique
Descriptif. Toulouse, Imp. H. Basuydu et Cie.

Campden-Main, S.M. (1970). A field guide to the snakes of South Vietnam.
Smithsonian Institution, Washington.

Daniel, J.C. (1984). The book of Indian reptiles. Bombay Natural History
Society, Bombay, 141 pp.

De Rooij, N. (1915). The reptiles of the Indo-Australian Archipelago, Vol.
III, Ophidia. E.J. Brill Ltd., Leiden. Reprinted 1970, A. Asher and Co.
N.V. Vaals.

Deuve, J. (1970). Serpents de Laos. Mémoire O.R.S.T.O.M. No. 39, Paris.

Flower, S.S. (1899). Notes on a second collection of reptiles made in the
Malay Peninsula and Siam. With a list of the species recorded from those
countries. Proceedings of the Zoological Society of London 1899:
600-697.

Frith, C.B. (1977). A survey of the snakes of Phuket Island and the adjacent
mainland areas of Peninsular Thailand. Natural History Bulletin of the
Siam Society 26: 263-316.

Fuchs, K. (1975). Chemistry and technology of novelty leather. FAO, Rome,

201 pp.

Haas, C.P. de (1950). Checklist of the snakes of the Indo-Australian
Archipelago. Treubia, 20(3): 511-625.

Harman, A.J.E. (1961). A collection of snakes from Singapore. Malayan
Nature Journal 15: 181-183.

Irvine, F.R. (1954). Snakes as food for man. British Journal of

Herpetology 1(10): 183-189.

Khan, M.A.R. (1982). On the endangered snakes of Bangladesh. In, Proceedings
of the IUCN/SSC Snake Group First Meeting, 8-12 November, 1982. Madras
Crocodile Bank.

Leviton, A.E. (1963). Remarks on the zoogeography of Philippine terrestrial
snakes. Proceedings of the California Academy of Sciences 4th ser.,
31(15): 369-416.

Lim, F.L.K. (1981). The Reticulated Python. Nature malaysiana 6(4): 14-17.

Olney, P.J.S. (Ed) (1984). Reptiles bred in captivity and multiple generation
births, 1981. In: International Zoo Yearbook, 23.

Reitinger, F.F. (1978). Common Snakes of South East Asia and Hong Kong.
Heinemann Educational Books (Asia) Ltd., Hong Kong.

Ridley, H.N. (1899). The habits of Malay reptiles. Journal of the Straits
Branch of the Royal Asiatic Society, 32: 185-210.

Saint Girons, H. (1972). Les serpents du Cambodge. Mémoires museum National
d'histoire Naturelle, Paris, N.S., Ser. A (Zool.), 74.

Salter, R.E. (1983). Summary of currently available information on
internationally threatened wildlife species in Burma. F.O.: BUR/80/006.
Field Document 7/83. FAO, Rangoon.

Shelford, R.W.C. (1916). A naturalist in Borneo. London, T. Fisher Unwin.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity,
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

272

Python reticulatus

Smith, M.A. (1943). Reptilia and Amphibia, Vol. III, Serpentes. In: The
Fauna of British India. Taylor and Frances, Ltd., London. Reprinted
1973, Ralph Curtis Books, Florida, 583 pp.

Soderberg, P. (1965). The pythons of Thailand. Conservation News S.E Asia
5: 11-12.

Sworder, G.H. (1922). A list of the snakes of Singapore Island. Singapore
Naturalist 1(2): 55-73.

Taylor, E..H. (1965). Serpents of Thailand and Adjacent Waters. University
of Kansas Science Bulletin 45(9): 609-1096.

Tikader, B.K. (1983). Threatened Animals of India. Zoological Survey of
India, Calcutta, 307 pp.

Tweedie, M.W.F. (1983). The Snakes of Malaya (3rd edn.). Singapore National
Printers (Pte) Ltd.

Ussher, C. (1979). Brunei's largest snake. Brunei Museum Journal 4(3): 180
(not seen, cited in Zoological Record).

Wall, F. (1916). The Reticulate Python Python reticulatus (Schneider).
Journal of the Bombay Natural History Society 31: 84-90.

Whitaker, R. and Whitaker, Z. (1983). Herpetological conservation in India.
Paper presented at Bombay Natural History Society Centenary Seminar.
December 1983. (Proceedings in press).

Wood, G.L. (1982). The Guiness Book of Animal Facts and Feats. Guiness
Superlatives, London, 252 pp.

273

AFRICAN PYTHON Recommended list: 2

[Possible problem]
Python sebae (Gmelin, 1789)

Order SERPENTES Family BOIDAE

a ee

SUMMARY AND CONCLUSIONS A potentially very large species, very widespread
in Africa south of the Sahara. Records exist for the following countries:
Angola, Benin, Botswana, Burkina, Burundi, Cameroon, Central African Republic,
Chad, Congo, Equatorial Guinea, Ethiopia, Gabon, Gambia, Ghana, Guinea, Guinea
Bissau, Ivory Coast, Kenya, Liberia, Malawi, Mali, Mozambique, Namibia, Niger,
Nigeria, Rwanda, Senegal, Sierra Leone, Somalia, South Africa, Sudan,
Swaziland, Tanzania, Togo, Uganda, Zimbabwe, Zaire, Zambia. The species is
also likely to occur in Mauritania. Typically found in the vicinity of water
and in savanna habitats but also ranges into forest regions. Seemingly still
common in mational parks and similar protected areas in southern parts,
possibly in the north also, but also reported to be locally depleted, although
little information is available on population sizes or trends.

Widely exploited for food, medicine and skins; no detailed information is
available on numbers utilized nationally. Relatively small numbers of live
P. sebae are in reported international trade, an annual mean of 465 animals
between 1980 and 1985, but large numbers of skins are traded, ranging between
641 and 15 260 in the same period, with an annual mean of 4403. In the skin
trade, the main importers were Italy and F.R. Germany, and the major sources,
Sudan, Nigeria, Ghana and Togo. The chief importers of live animals were the
USA and F.R. Germany, who obtained stock mostly from Ghana, Togo and Senegal.

This volume of trade seems unlikely to pose a threat to the species as a
whole, but, although it may well be adversely affecting local populations,
adequate population data are not available to evaluate this possibility; such
data should be gathered as a matter of priority.

DISTRIBUTION Widespread in the continent of Africa, south of the Sahara.
Present on a few offshore islands, such as Bioko, (Fernando Poo) and Zanzibar,
but absent from Madagascar. The northern limit of the range extends from
Senegal in the west, across the fringes of the Sahel to Ethiopia and Somalia.
The species is absent from the Kalahari and other arid parts of southern
Africa, where the southern. limit extends from the northern sector of Namibia,
across northern Botswana, to north-east South Africa.

Records exist for the following countries: Angola, Benin, Botswana, Burkina,
Burundi, Cameroon, Central African Republic, Chad, Congo, Equatorial Guinea,
Ethiopia, Gabon, Gambia, Ghana, Guinea, Guinea Bissau, Ivory Coast, Kenya,
Liberia, Malawi, Mali, Mozambique, Namibia, Niger, Nigeria, Rwanda, Senegal,
Sierra Leone, Somalia, South Africa, Sudan, Swaziland, Tanzania, Togo, Uganda,
Zimbabwe, Zaire, Zambia. The species is also likely to occur in Mauritania.

Taxonomic note A recent review (Broadley, 1984) recognises two subspecies,
P. s. sebae in northern parts of the range, and P. s. natalensis in
southern parts, from northern Angola, southern Zaire and Kenya south to
Natal. <A recently described species P. saxuloides Miller & Smith, 1979 is
reduced to the synonymy of P. s. natalensis.

Angola P.s. sebae has been recorded from Ambriz and Dundo in the north
of Angola. P. s. natalensis occurs in the south of the country (Broadley,
1984).

Benin P. s. sebae is recorded only from Zizonkame (Broadley, 1984).

274

Python sebae

Botswana Recorded localities for P. s. natalensis are Kasane, Khwai, Lake
Ngami, Maun, Metsimaklaba (Broadley, 1984).

Burkina Recorded localities for P.s. sebae are Natiaboani, Ougarou
(Broadley, 1984).

Burundi P. s. natalensis is recorded only from Bujumbura (Broadley, 1984).

Cameroon Recorded localities for P.s. sebae are Bafout, Banjo Bamendo,
Bipindi, Edea, Koum, Libamba-Makak, Metet, Mieri, Nachtigal, Tcholliré,
Victoria (Broadley, 1984).

Chad P. s. sebae is recorded only from Sar (Broadley, 1984).

Congo Broadly (1984) recorded P. s. sebae only from Nganchou, but it is
said to be widely distributed (Congo CITES MA, 1986).

Equatorial Guinea Recorded localities for P. s. sebae are Rio Benito on
the mainland and Bioko (Broadley, 1984).

Ethiopia Recorded localities for P.s. sebae are Amba Aradam, Godare,
Kaalam, Sodu (Broadley, 1984).

Gabon Distributed more or less uniformly throughout the country (Gabon
Ministére des Eaux et Foréts, in Jlitt., 29 November 1985). Recorded
localities for P.s. sebae are Belinga, Fernan Vas, Franceville, Omboué,
Talagouga (Broadley, 1984).

Gambia Listed as present (Hakansson, 1981).

Ghana Recorded localities for P. s. sebae are Abonu, Accra, Achimota,
Akosombo, Amisano, Asempanaye, Bolgatanga, Dabacrom, Gbefi, Kete Krakye,
Kpong, Kumasi, Legon, Mole National Park, Secondi, Suhum, Tamale, Volta River,
Wa (Broadley, 1984).

Guinea Recorded localities for P. s. sebae are Conarky, Mont Nimba
(Broadley, 1984).

Guinea Bissau Recorded localities for P. s. sebae are Bijagos
Archipelago, Bissao, Mansoa (Broadley, 1984).

Ivory Coast Recorded localities for P. s. sebae are Adio, Podoumé, Bouna,
Elima, Flampleu, Lampto (Broadley, 1984).

Kenya The species occurs in a variety of habitats from semi-desert to swamp
and forest, occasionally appearing near the centre of Nairobi (A.D. Mackay,
in litt., 26 March 1986). P. s. sebae has been recorded from several
localities in. southern Kenya, where it appears to intergrade with
P. s. natalensis (Broadley, 1984).

Liberia Said to be restricted to swamps and river courses (Liberia CITES
MA, 1986). Recorded localities for P.s. sebae are Du River, Harbel,
Monrovia, Mount Coffee, Peahtah, Robertsport, Suococo (Broadley, 1984).

Malawi Recorded localities for P. s. natalensis are Chibotela, Chipoka,
Kondowe to Karonga, Lujeri Estate, Misuku Hills, Nyungwe (Broadley, 1984).
Morgan-Davis et al. (1984) remarked that large individuals could be seen on
the Shire River south of Mvuu.

Mali P. s. sebae is recorded only from Bamako (Broadley, 1984).

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Python sebae

Mauritania There are no records from the country, but P. s. sebae has
been recorded from three localities on the southern bank of the Senegal River
(Broadley, 1984).

Mozambique Widely distributed near permanent water (Mozambique CITES MA,
1986). Recorded localities for P.s. natalensis are Alves de _ Lima,
Mocambique Island, Monzo, Setache Po, Vamizi Island and Zinave (Broadley,
1984).

Namibia Recorded localities for P.s. natalensis are Kaokoveld to Etosha
Pan, Rundu (Broadley, 1984). Said to be widespread in the north of the
country, often associated with artificial water points (Namibia Department of
Agriculture and Nature Conservation, in litt., 1987).

Niger P. s. sebae is recorded only from 11 km north-west of Niamey
(Broadley, 1984).

Nigeria Found throughout the country (Nigeria CITES MA, 1987). Recorded
localities for P.s. sebae are Bamendo, Lagos, Mundame, Samaru (Broadley,
1984).

Rwanda Recorded localities for P.s. sebae are Lac Ihema, Ngarama, Parc
National Kagera (Broadley, 1984).

Senegal Recorded localities for P. s. sebae are Bakel, Boughari, Dagana,
Darou, Diattacounda, Fatick, Rossombo, Rufisque (Broadley, 1984).

Sierra Leone Recorded localities for P.s. sebae are Bo, Sandaru
(Broadley, 1984).

Somalia Recorded in Middle Shebelle, Mogadishu, Bay, Lower Shebelle, Gedo,
Middle Jubba and Lower Jubba (Simonetti and Magnoni, 1986).

South Africa P.s. natalensis is widely distributed in Natal, Transvaal
and Transkei, and reaches the extreme north of the Cape Province. The last
specimen was captured in the Eastern Cape Province in 1927 (Fitzsimmons,
1962; Broadley, 1984), but 31 were reintroduced to a nature reserve in the
region between 1980 and 1986 (South Africa CITES MA, 1987).

Sudan Recorded localities for P. s. sebae are south of Khartoum, Torit
and the White Nile (Broadley, 1984).

Swaziland Recorded localities for P.s. natalensis are Groenpan Farm,
Hlane, Mliba (Broadley, 1984).

Tanzania Said to be widespread (K. Howell, in Jitt., 15 March 1986).
P. s. sebae has been recorded from several localities in northern Tanzania,
where it appears to intergrade with P. s. natalensis (Broadley, 1984).
Present on Zanzibar Island but probably not on Pemba (Pakenham, 1983).

Togo Recorded localities for P. s. sebae are Binaparba, Nagarou
(Broadley, 1984).

Uganda Said to be ubiquitous except at higher altitudes and in arid eastern

regions (Pitman, 1974). P. s. sebae was recorded only from the Sese Islands
by Broadley (1984).

Zaire P.s. sebae has been recorded from numerous localities in north,
west and central Zaire, including the northern end of the Rift Valley.
P. s. natalensis occurs in the south of the country (Broadley, 1984).

276

Python sebae

Zambia Recorded localities for P.s. natalensis are Chipangali, Chipata,
Chipopera, Kabompo, Kabwe, Kafue River, Kaputa, Kasama, Kasauta, Livingstone,
Lusaka West, Mbala (Broadley, 1984).

Zimbabwe P. s. natalensis is found throughout Zimbabwe, but is usually

most plentiful along major rivers and reed-fringed lakes (Broadley and Cock,
1975). Broadley (1984) listed numerous localities.

POPULATION The southern subspecies is said to be “still common in thinly
populated areas where there is good cover and an adequate food supply”
(Broadley, 1983); this general situation is also likely to apply to the
northern subspecies. In many cases such areas are likely to be in National
Parks, Game Reserves, and similar protected areas. In general the species is
under pressure due to habitat loss and killing for the skin trade. Although
python populations are said to be rapidly declining in many parts of Africa
(Madsen et al., 1985), the species is also said to be very common in many
regions, and to adapt readily to disturbed environments such as sugar cane
fields (and to be found frequently around human habitation) (Branch and Hacke,
1980). Most sources do agree that very large specimens are now rarely seen.
Given the widespread exploitation of the species and the concern in some
quarters for its population status, appropriate field data are required with
some urgency. Available status information is summarised below; there is no
information for the other countries.

Cameroon No surveys have been carried out, but the species is said to be
not in danger (Cameroon CITES MA, 1987).

Gabon Said to be more common in the coastal regions than inland, where it
has been depleted by exploitation (Gabon Ministére des Eaux et Foréts, in
Jitt., 29 November 1985).

Kenya The species is often quite common, although seldom seen (A.D. Mackay,
in litt., 26 March 1986). Fairly common along the Kora River in the Kora
Game Reserve (Madsen et al., 1985).

Malawi Morgan-Davis et al. (1984) remarked that the species must be more
common in Lilondwe National Park than the single record would suggest.

Namibia Described as widespread in the north and fairly abundant in Etosha
(Namibia Department of Agriculture and Nature Conservation, in litt., 28
February 1987).

Nigeria Described as rare and vulnerable (Nigeria CITES MA, 1987).

Somalia Populations are said to have been sadly depleted owing to hunting
and habitat destruction. In Juba and Shebelle they are rather rare an
represented almost entirely by young specimens (Simonetti and Magnoni, 1986).
Fagotto (1985) described the species as “not rare".

South Africa Listed in the South African Red Data Book as vulnerable
outside game reserves, but said to be common in Kruger N.P. (McLachlan, 1978)
and in several of the larger Zululand reserves (South Africa CITES MA, 1987).

Tanzania Perhaps not threatened in Tanzania, despite trade in skins (both
legal and illegal) and persecution (K. Howell, in litt., 15 March 1986).

Uganda Said to be locally common in suitable habitat, particularly on
Victoria and Nyanza Islands (Pitman, 1974). Described as quite common in the
extensive swamps of Lake Kyoga and Bunyoro-Luwero areas, although population
size is unknown (Uganda Game Department, in litt., 28 March 1987).

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Python sebae

Zaire Seemingly under heavy pressure in Zaire due to exploitation for food
and skins: pythons were difficult to find within about 30 km of Kinshasa and
specimens over 4 m long were not found at all (larger skins on sale may have
been stretched or have originated from the interior of the country) (Van
Wallach, in litt., 22 October 1983).

Zimbabwe Reportedly common in most National Parks and Safari Areas in
Zimbabwe (Broadley and Blake, 1979; cited by Broadley, 1983).

HABITAT AND ECOLOGY A large and heavy-bodied species; at present a snake of
4.5 m would be classed as large but occasional old specimens of 7-8 m have
been recorded in the past.

Mainly nocturnal or crepuscular, although not exclusively. Highly aquatic, a
strong swimmer, and a good climber. May occur in various vegetation types.
Whilst deserts and truly arid regions are avoided, the species is most
commonly associated with savanna habitats (or ‘bush country’) of various
kinds, and when present in forest regions it appears to prefer clearings
within them, and is often found around human habitation and cultivations
(Pope, 1961). Most frequently found near swamps, rivers or other permanent
water sources, but can also be common elsewhere. Often found in large
horizontal rock crevices near streams, among large boulders in rock outcrops,
and in ant-bear burrows or deserted termitaria. Extends up to 2250 m in
Uganda (Pitman, 1974).

The diet is varied; any mammal of appropriate size (ranging from rats to
antelopes of several species) or bird, may be taken, or occasionally reptiles,
frogs or fishes (Broadley and Cock, 1975). Prey is typically taken from
ambush; for example, in cover next to a game trail, on a branch overhanging
it, or partly submerged at the edge of a water hole (Broadley and Cock, 1975).

Breeding may occur at any time of year in equable humid conditions, but
apparently does not occur during dry seasons elsewhere. Clutch size is often
between 30 and 50, sometimes more, exceptionally to 100; about 40 is usual for
a female of about 4.5 m (14-15 ft) length (Cansdale, 1961). Eggs are about
6 x 9 cm (often many in a clutch being infertile) and are frequently laid in
an antbear hole, or similar situation (Broadley and Cock, 1975). Incubation
period is about 90 days; the female broods the eggs (assisting maintenance of
suitable temperature and moisture conditions), but leaves them frequently to
visit water. Hatchlings are 20-22 in (51-56 cm) long and 3 oz (85 g) in
weight (Cansdale, 1961). First mating in a captive group was at nearly six
years age, 2.4 cm length.

THREATS TO SURVIVAL This species is used for food, medicinal purposes and
skins, probably throughout its range. The fat is applied to ease muscle pain
and chest complaints, and various internal organs are used for other medicinal
purposes. In many parts of Africa, Pythons are revered for spiritual reasons,
and are therefore not interfered with (Fitzsimmons, 1962). In West Africa
generally, P. sebae is most commonly taken for its skin. Also widely
persecuted for its alleged threat to domestic stock such as chickens, goats

and calves, and thus greatly depleted in settled and farming areas (Broadley,
1983).

Botswana A few pythons are killed for the protection of livestock. There
are no legal exports (Botswana CITES MA, 1986), though a few manufactured
products are exported.

Cameroon There is little trade in raw skins, but a fairly intense trade in

manufactured goods, such as bags, wallets, etc. (Cameroon CITES MA, 1987).

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Python sebae

Congo Some tribes in the Congo region are said to relish the meat of

pythons (Fitzsimons, 1962). They are also hunted for their skins (Congo CITES
MA, 1986).

Gabon The people of the inland regions are fond of python meat, but it is
not eaten in coastal districts. There is said to be little internal trade and

no recorded exports in 1984 and 1985 (Gabon Ministere des Eaux et Foréts, in
litt., 29 November 1985).

Liberia Although the species is used for food, commercial exploitation for
skins is the primary form of exploitation (Liberia CITES MA, 1986).

Mozambique There is thought to be little trade, although some animals are
killed out of fear and their skins may be traded (Mozambique CITES MA, 1986).

Namibia There is said to be no commercial exploitation and little illicit
exploitation by reptile fanciers. Occasionally persecuted by farmers (Namibia
Department of Agriculture and Nature Conservation, in Jitt., 1987).

Nigeria Pythons are commonly killed for their skins. A 5- to 6-m skin can
fetch 100 naira ($150) or more; skins are openly sold to tourists on beaches
and in hotels in all major cities (Butler and Shitu, 1985).

Sierra Leone Pythons are rarely caught for food but more often for skins,
particularly for export. Skins are sold to trophy dealers who retail them to
tourists. They are also used by local citizens for house decorations (Sierra
Leone Ministry of Agriculture and Forestry, in litt., 23 March 1987).

Somalia Hunting and habitat destruction are both implicated in population
decline. Most riverine forest, one of the former main habitats, has now
disappeared, and smuggling of python skins is said to be still thriving
(Simonetti and Magnoni, 1986). Fagotto (1985) described the species as "not
rare".

Uganda Greatly persecuted for skins in Lake Kyoga and Bunyoro-Luwero
areas. Used for food in East Madi and possibly West Nile as a whole. From
1981 to 1986 legal exports of skins amounted to 63 in 1981, 16 in 1982, 15 in
1983 and 5 in 1985. Single live animals were exported in 1984 and 1986.
There is also thought to be a substantial illegal trade; 142 skins bound for
Lebanon were seized in 1986 (Uganda Game Department, in Jlitt., 28 March
1987).

INTERNATIONAL TRADE CITES Annual Reports record trade in skins in numbers
of pieces and by length. Transactions reported in terms of length have been
converted to numbers of skins using the figure of 1.7 m given as the mean
length of skins of P. molurus in trade (a similar sized species to
P. sebae) by Fuchs (1975). These data are summarised in Tables 1 and 2.
Relatively small numbers of live P. sebae are in reported international
trade, an annual mean of 465 animals between 1980 and 1985, but large numbers
of skins are traded, ranging between 641 and 15 260 in the same period, with
an annual mean of 4403. In the skin trade, the main importers were Italy and

F.R. Germany, and the major sources, Sudan, Nigeria, Ghana and Togo. The
chief importers of live animals were the USA and F.R. Germany, who obtained
stock mostly from Ghana, Togo and Senegal. This volume of trade seems

unlikely to pose a threat to the species as a whole, but, although it may well
be adversely affecting local populations, adequate population data are not
available to evaluate this possibility.

279

Python sebae

Table 1 Minimum net imports of live animals (L) and skins (S) of P. sebae
reported to CITES.

1980 1981 1982 1983 1984 1985
Australia 2S - - - 1s -
Austria - - eS - - 281 L
Belgium - - 4.L 1s - 1s
Canada 19 L 1L at St, - - 8 L
25 Ss - - 96S 30 Ss 13 $s
Chile - - - - 1s -
Cyprus - - - - - 100 S
Czechoslovakia - - - 1L - -
Denmark - - - 5 L 4L -
- - - 9 1s 1s
German D.R. 14 L - - 8 L -
Germany, F.R. 97 L 29 L 187 L 81 L - 44 L
1s 137 S 32 Ss 335 Ss 2s 548 S
France 4S IL 2L 14 L Je 2 L
- - - - 55s 7s
Greece - - - 1L 6 L 887 S
Israel 1068 S 21S - 131s - 88 S
- - - - - 1L
Italy 1L 50 L 40 L 35 L 24 L -
- 11553 Ss 5754 S$ - 162 S 148 S$
Ivory Coast - - - 15 Ss - -
Japan - 2L 2L 158 2L 21 L
Korea, Rep. - - - - 4 L -
Lebanon - - - 10 $s - -
Liberia - - - 3S - -
Lybia - - - - - 358
Malaysia - - - - 2 L -
Netherlands - - 1L 26 L 18 L Sy)
- - - - 100 Ss -
Philippines - - - 2L - =
Poland © - - 1s - +o aR— -
Romania 1L - - - - -
Saudi Arabia - - - 7) jt 35E 19 §
Singapore - - - 38 S 6s -
Spain - ibs its - 1L 3 L -
= a e s a os
Sweden - - 1s “= = 1s
Switzerland 3m 6 L 9 L 9 L 1L 4L
48 - 3 58 34 Ss 3 Ss 17 s
Thailand - - = = = 59s
UAE = = - = = 69 S
UK 84 L - 45 L 6L 7L 44 L
- - 50 s - 1s 738
USA 315 L 261 L 197 L 81 L 108 L 547 L
46 S 3549 S 401 S 128 Ss 279 S 258 S
USSR - - - 1L - 2s
Yemen - - - - - 1S
Yugoslavia - - - - - 15s
Zaire 1L - - - - -
Country unknown 1L - 51S - - -
TOTAL 536 L 351 L 488 L 265 L 193 L 958 L
1127 s 15260 S 6294 S$ 727 S 641 S 2369 S

280

Python sebae
Table 2 Reported countries of origin (or exporting country if no origin

reported) and numbers of live animals (L) and skins (S) of P. sebae reported
to CITES.

1980 1981 1982 1983 1984 1985

Countries having or possibly having wild populations of P. sebae

Benin - - - - 50 Ss 62 Ss
Cameroon - 35S 2s DS 1s 11 °S
Chad - 430 Ss - = = =
Congo - NES - - - -
Ghana 327, L 145 L 190 L TA) Jb 30 L 702 L
20S 35 Ss 185 s 38 Ss 4s -
Guinea - - - - 2s 10 Ss
Ivory Coast - aS - - - 5 Ss
Kenya 167 L 106 L 1s 38 - 1S
Liberia - 24 L 7} ht 42 L 2201 40 L
- 420 Ss - - 2aS -
Malawi 72 - - - - iss
Mali ~ - 335 S - 140 § AL
Mozambique - - - - is -
Nigeria 2107S 391 Ss 84S 656 S 1538S 413 s
Senegal 25 150 L - 100 L

WwW
7)
ry
w
wt
“n

|
w
Ww
“
ray
[o)
ol
“
|

Sierra Leone - - = = 7s 3s
South Africa ZS - 32 Ss - 2s -
Sudan ~ 11308 § 5695 $ 7s - 1660 S
Tanzania 2s SS - - - 2s
Togo 45 L UM 149 L 95 L 136 L 116 L
- - Z2ES 43 Ss 87 S 147 S$
Uganda - - 20 $s - - =
Zaire eS as SaaS S70 - 41 Ss
Zambia 2s - - - iS -
Countries without wild populations of P. sebae.
Austria - is - - - -
Germany, F.R. - - - - 8 L -
India es - - - - -
Indonesia 2s - - - - -
Jamaica - - - eS - -
Spain 2S - - - - -
Switzerland = - - 39 L - 1L
- - - Lis - 18
Thailand 6L 2718 Ss - - - -
UK oe - - - = =
Unknown 6L - - 4L 1L -
904 S 39 S$ 21S 21S 161 S 20 Ss

281

Python sebae

CONSERVATION MEASURES Unless otherwise stated, all of the information on
protection is from IUCN Environmental Policy and Law Occasional Paper No. 3,
African Wildlife Laws.

Angola All Pythons are protected under the Hunting Regulations, 11 December
1955.

Benin Under the Decree concerning hunting and capturing licences, bag
limits and professional hunters, 11 February 1980, Pythons are designated as
small game species. They may only be hunted by the holders of traditional
hunting rights or under hunting permits.

Botswana Pythons are totally protected under the Fauna Conservation Act,
1961.

Burkina Python sebae is partially protected under the Wildlife
Conservation and Hunting Act, 31 December 1968, which establishes bag limits
and close seasons, protects nests and eggs, and regulates the sale of meat.
Commercial capture and export are controlled under the same Act by Decree No
68-314. Ordonnance’ No 68-58, December 1985, fixed the open season from 15
December 1985 to 28 February 1986 and stipulated that partially protected
species could only be hunted under special permit and only in the province of
Gourma and Tapoa.

Burundi No information.

Cameroon Python sebae is partially protected (Class B) under the Forests,
Wildlife and Fisheries Act, 27 November 1981. They may only be taken under a
valid hunting licence. No more than two animals may be taken in one day.

Central African Republic The Ordinance concerning the protection of
wildlife and regulating hunting, 27 July 1984, designates Pythons as partially
protected; they may only be taken under a special hunting permit and sold or
exported if they are accompanied by documentary proof that they were acquired
lawfully.

Chad No information.

Congo The Act concerning the conservation and exploitation of wild fauna,
21 April 1983, vests in the State ownership of all wild animals of economic
value, and requires the issuing of licences for commercial capturing.

Equatorial Guinea The Hunting Regulations, 29 April 1953, establish the
need to obtain hunting licences except for subsistence hunters. Reptiles are
not protected.

Ethiopia Under the Wildlife Conservation (Amendment) Regulations, 12
February 1974, Python sebae are protected. They may only be hunted for
scientific purposes and require a special permit, the value of which was set
at Eth$12.50 for each python.

Gabon The Wildlife and Forests Act, 22 July 1982 requires the issuing of
licences for the commercial capture of all wildlife. Traditional hunting for
subsistence pruposes is permitted.

Gambia Under the Wildlife Conservation Act, 14 February 1977, all wildlife
except game and vermin are protected.

Ghana The Wildlife Conservation Regulations, 4 March 1971, define Python
sebae as totally protected.

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Python sebae

Guinea No information.

Guinea-Bissau The Hunting Regulations, 12 May 1980, require the issuance of
a licence for the capturing of live wild animals. Pythons are partially
protected, and may not be taken by sport hunters.

Ivory Coast Python sebae is listed as a predatory animal in the Wildlife
and Hunting Act, 4 August 1965, and may be hunted without licence anywhere
except in protected areas. Arrété No. 15, 26 December 1972, establishes
licence fees for the capturing of live reptiles.

Kenya Protected under Legal Notice No. 152 (25 September 1981), export
being prohibited (Broadley, 1983). Probably present in all Kenya's National
Parks, except Mt. Kenya and Sibilot, too high and too dry, respectively (A.D.
MacKay, in litt., 26 March 1986).

Liberia The Wildlife Conservation Regulation (in draft, 1985) lists Python
sebae as fully protected. Trade in and export of fully and partially
protected species is prohibited except for educational or scientific purposes.

Malawi Hunting, possession, trade and export of game is controlled under
the Game Act, 1953. No reptiles, other than crocodiles, are protected.

Mali All snakes are classified as game species under the Hunting Act, 11
November 1969. There is a general close season from 1 June to 31 October. A
general hunting ban (Decree 325/PG-RM) throughout the country was imposed on 6
November 1978.

Mauritania Under the Hunting and Wildlife Protection Act, 15 January 1975,
the capture, possession, sale and export of live wild animals requires a
commercial capture permit. The sale of game meat is prohibited.

Mozambique

Namibia Protected by the Nature Conservation Ordinance (Namibia Department
of Agriculture and Nature Conservation, in litt., 1987).

Niger All hunting for utilitarian purposes has been prohibited in Niger
since 1972 (Niger CITES MA, 1986).

Nigeria Python sebae is listed as fully protected in the Endangered
Species (control of International Trade and Traffic) Decree, 20 April 1985.
Their hunting, capture, trade and export is prohibited.

Rwanda Pythons are apparently not protected.

Senegal The Game and Wildlife Protection Regulations, 30 May 1967, lay down
regulations governing the issuance of licences for hunting and commercial
capturing of wildlife.

Sierra Leone Under the Wildlife Conservation Act, No. 27, 1972, pythons are
classified as game animals with an unlimited quota, they may only be hunted by
those in possession of a valid game licence.

Somalia Pythons are listed as game animals under the Law on Fauna (Hunting)
and Forest, 25 January 1969. They may only be hunted or exported under
licence. A ban on all hunting was instituted on 13 October 1977.

South Africa P. sebae occurs in 14 nature reserves in Natal and several
in Zululand (South Africa CITES MA, 1987).

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Python sebae

Sudan Pythons are listed in the Ordinance for the Preservation of Wild
Animals, 1935 (Amended 1974), as species for which hunting is permitted.
Licences are only required if firearms are used for hunting. The Hides and
Skins (Export) Regulations, 1969, specify a grading system for the export of
reptile skins.

Swaziland Apparently no reptiles are protected in Swaziland.

s

Tanzania P. sebae is classified as big game under the Wildlife
Conservation Act, 30 June 1974. It may not be hunted or killed without a
valid hunting licence. The capture of all live animals requires a valid

capture permit, and the hunting of all animals requires a valid hunting permit.

Togo PP. sebae is classified as a pest species under the Ordinance on
Wildlife Protection and Hunting, 16 January 1968. An enabling decree, dated 4
June 1980, sets out licence fees for hunting and commercial capture, but there
is no bag limit for the species.

Uganda Protected under the Game Act by Statutory instrument No. 20 of 31
October 1980 (Uganda Game Department, in litt., 28 March 1987).

Zaire The commercial capture of unprotected animals requires a permit under
the Hunting Act, 28 May 1982. Export can only be authorised if the exporter
holds a certificate of lawful possession.

Zambia Pythons are listed as game animals in the National Parks and
Wildlife Act, 3 December 1968, under which their hunting, possession, trade
and export are controlled and fees are set for the export of trophies and live
animals.

Zimbabwe Python sebae is declared a specially protected animal under the
Parks and Wildlife Act, 1975. The collection of live reptiles and their eggs
and their breeding in captivity are controlled under licence by the Parks and
Wildlife (General) Regulations, 1981.

CAPTIVE BREEDING A recent inventory (Slavens, 1985) lists 37 specimens in 9
collections, with breeding in two groups; the total number in captivity will
be much higher than this.

REFERENCES

Branch, W.R. and Hacke, D. (1980). A fatal attack on a young boy by an African
Rock Python Python sebae. Journal of Herpetology 14(3): 305-307.

Broadley, D.G. (1983). In litt., 10 August (draft RDB account’ for
P. s. natalensis).

Broadley, D.G. (1984). A review of geographical variation in the African
Python Python sebae (Gmelin). British Journal of Herpetology 6(10):
359-367.

Broadley, D.G. and Blake, D.K. (1979). A check list of the reptiles of the
national parks and other conservation areas of Zimbabwe (Rhodesia).
Arnoldia, Rhodesia 8(35): 1-15.

Broadley D.G., and Cock, E.V. (1975). Snakes of Rhodesia. Longman Rhodesia
(Pvt) Ltd., Salisbury, 152 pp.

Butler, I.A., and Shitu, E. (1985). Uses of some reptiles by the Yoruba
people of Nigeria. Herpetological Review 16(1): 15-16.

Cansdale G.S. (1961). West African Snakes. Longmans, London, 74 pp.

Fagotto, F. (1985). Larger animals of Somalia in 1984. Environmental
Conservation 12: 260-264.

Fitzsimmons, V.F.M. (1962). Snakes of Southern Africa. Purnell, Cape Town,
423 pp.

284

Python sebae

Fuchs, K. (1975). Chemistry and technology of novelty leather. FAO, Rome,
201 pp.

Hakansson, N.T. (1981). An annotated checklist of reptiles known to occur in
the Gambia. Journal of Herpetology 15: 155-161.

Madsen, T., Loman, J., Hebrard, J. and Duff-Mackay, A. (1985). A survey of
the reptile fauna in Kora Game Reserve. Unpublished report.

McLachlan, G.R. (1978). South African Red Data Book - Reptiles and
Amphibians. South African National Scientific Programme Report No. 23.
Miller, T., and Smith, H.M. (1979). The Lesser African Rock Python.

Bulletin of the Maryland Herpetological Society 15: 70-84.

Morgan-Davis, A.M., Dudley, C.O. and Meredith, H.M. (1984). Additions to the
vertebrate fauna of Lilondwe National Park, Malawi. Nyala 10(2): 91-98.
Pakenham, R.H.W. (1983). The reptiles and amphibians of Zanzibar and Pemba
Islands. Journal of the Sast African Natural History Society and

National Museum. 177: 1-40.

Pitman, C.R.S. (1974). A guide to the snakes of Uganda. Revised edition.
Wheldon and Wesley, Codicote, 290 pp.

Pope, C..H. (1961). The giant snakes. Routledge and Keagan Paul, London,

297 pp.

Simonetta, A.M. and Magnoni, M.L. (1986). Status and conservation problems of
Somali lower vertebrates. Rivista di Agricoltura Subtropicale e
Tropicale 80(3): 405-432.

Slavens, F.L. (1985). Inventory of live reptiles and amphibians in captivity,
current January 1, 1985. F.L. Slavens, Seattle, Washington, 341 pp.

285

Recommended list: 3
‘ [No problem]
Ornithoptera caelestis (Rothschild, 1898)

Phylum ARTHROPODA Order INSECTA
Class LEPIDOPTERA Family PAPILIONIDAE

SUMMARY AND CONCLUSIONS Ornithoptera caelestis, generally considered as a
subspecies of O. priamus, is endemic to the Louisiade Archipelago, Papua
New Guinea. Although not common, there are believed to be strong populations
of O. caelestis on three islands. It normally inhabits hills and mountains
and its feeding habits are similar to those of O. priamus.

Ornithoptera caelestis is not threatened, although it has a rather limited
distribution. International trading is virtually entirely in specimens reared
ex-pupae and sold via the PNG Government Insect Farming and Trading Agency
(IFTA). Present maximum sales in the order of 1000-1500 specimens per year
are well within acceptable limits and pose no threat to caelestis. Indeed,
the IFTA, through the PNG Government, could be encouraged to expand its
operations in deadstock and, more particularly, in livestock (for butterfly
houses). An increase in commercial contact between the IFTA and entrepreneurs
in retailing centres might be advisable. CITES statistics show very low
levels of trade in 1984 and 1985 but the numbers may have increased after the
the lifting of the EEC ban on birdwing imports in 1987.

Conservation and utilization of birdwings are mutually reinforcing in PNG The
IFTA and government agencies responsible for protected areas could work
together to ensure that sufficient wild habitat remains to ensure a
sustainable yield of caelestis and other species. The trading operations
could go hand in hand with conservation efforts on behalf of threatened
subspecies and species of birdwings. Threatened taxa are described in the
Swallowtail Red Data Book (Collins and Morris, 1985).

Taxonomic note: Only D'Abrera (1975) of the recent authors considers

caelestis to be a separate species. Haugum and Low (1978-79), after
detailed examination of genital and other characteristics, found this to be ‘a
highly unwarranted separation for which we fail to find any support’. Hancock

(1983) follows this decision, as do Collins and Morris (1985) in the
Swallowtail Red Data Book. The CITES listings follow D'Abrera and hence this
separate report becomes necessary, but on scientific grounds it is not
justifiable.

DISTRIBUTION Ornithoptera caelestis is confined to the Louisiade
Archipelago, off the south-eastern trip of the Papua New Guinea mainland
(Haugum and Low, 1978-79).

POPULATION Described by Haugum and Low (1978-79) as localised and generally
uncommon to fairly rare. However, Pyle and Hughes (1978) suggest that Misima,
Nemoa and Hemenahei islands all support strong populations.

Interestingly, there is evidence from caelestis that overcollecting has the
capacity to decrease populations or even cause local extinction. A number of

286

Ornithoptera caelestis

observers on the Louisiade island of Nemoa concurred that the population of
caelestis was badly depressed for some years after the removal of several
hundred specimens by expatriate collectors. The species was reintroduced to
the site from another part of the island and subsequently flourished (Pyle and
Hughes, 1978). In contrast, Pyle and Hughes (1978) present evidence that
fairly heavy, but nevertheless carefully controlled, levels of cropping were
sustained by caelestis on Misima Island. In this instance several dozen
larvae and pupae were removed from a colony every season.

HABITAT AND ECOLOGY 0. caelestis is a denizen of hills and mountains,
with small populations at around 100 m, but rarely at sea-level. The feeding
habits are similar to O. priamus with larvae on at least two species of
Aristolochia.

THREATS TO SURVIVAL The level of threat to caelestis is difficult to
assess. Haugum and Low (1978-79) indicate that the taxon is rather rare,
while Fenner (in Jlitt., 1983) and Hutton (in Jitt., 1983) concur and
express fears for its future. Conversely, Pyle and Hughes (1978) describe how
caelestis quickly developed a strong colony on Aristolochia vines growing
prolifically on the young tree invaders of an abandoned garden. This level of
adaptability is confirmed by the success of butterfly ranchers in the
Louisiades, who attract caelestis to their gardens. Aristolochia vines
are trained up trellises and the butterflies breed within the gardens. Some
farmers, able to recognise the wild Aristolochia, take care to protect the
plants when they create new gardens (Pyle and Hughes, 1978). On balance,
caelestis seems adaptable and resilient, unlikely to be_ seriously
threatened. However, the Louisiades are certainly in need of representative
protected areas, to ensure the survival of strong, wild colonies in natural
conditions.

INTERNATIONAL TRADE Table 1 shows the levels of imports of caelestis,
1980-85. The figures for 1980-1982 are of little value since Papua New Guinea
did not submit a report to CITES in those years. The report for 1983 is
probably an accurate representation of imports, while the data for 1984 are
difficult to assess. Recent data from the IFTA indicated 1985 sales of 932
specimens, slightly up on previous years (Clark, P.B., in litt., 1986) but
the Papua New Guinea report for that year included the export of only one
specimen. Commercial exports during 1986 reportedly totalled 851 specimens
(Papua New Guinea CITES MA, 1987).

Theoretically, none of the EEC countries should have licensed the import of
caelestis during the period 1983 to 1986 because of the birdwing inclusion
on Annex C(1) of EEC Regulation 3626/82. In this regulation to implement
CITES, all CITES Appendix II butterflies were effectively treated as Appendix
I, thus prohibiting import. All birdwings except Ornithoptera alexandrae
were removed to Annex C(2) of the regulation during 1987.

Papua New Guinea is the only country of origin caelestis, the taxon being

endemic to the Louisiades Archipelago. Table 2 reflects this situation;
figures for Germany and Japan being the result of incorrect reporting.

287

Ornithoptera caelestis

Table 1. Minimum net imports of Ornithoptera caelestis
dead specimens reported to CITES, 1980-1984.

1980 1981 1982 1983 1984 1985
Australia - - - 22 5 -
Canada - - - - 2 -
France - - - 229 - -
Germany, F.R. - - 275 45 65 -
Ireland - - - 4 - -
Italy - - - 4 - -
Japan - - - 64 - -
Norway - - - 2 - -
Singapore - - - 4 - -
Switzerland 2 96 42 , 14 8 1
UK - - - 29 - -
USA - - - 280 - -
Unknown . - - - 48 - -
Total 2 96 317 745 80 1

The caelestis material exported by the Insect Farming and Trading Agency, a
government controlled body, is all farmed from pupae (Clark, P.B., in litt.,
1983, Fenner, 1976, National Research Council, 1983). In other words, the
specimens are collected as pupae from Aristolochia vines tended specially
for the purpose of attracting birdwings. The pupae are hatched in protected
conditions and the adults killed and delivered to the IFTA for export. With
such perfect specimens in good supply, there would be no demand for
wild-caught material, which is invariably damaged. Although PNG did not
report to CITES 1980-1982, information in CMC files indicates that sales of
caelestis between 1978 and 1982 totalled 4400. All of these were via the
IFTA and the majority, if not all, were ex-pupa. According to recent
information, the IFTA had to slow down the ranching operation in the
Louisiades because of a glut of stock. No doubt this was partly the result of
the unwarranted EEC ban on imports. The international trade in all farmed or
ranched birdwings from Papua New Guinea is greatly to be encouraged.

Table 2: Reported countries of origin (or exporting country if no original
source reported) and quantities of transactions in. dead specimens of
Ornithoptera caelestis

1980 1981 1982 1983 1984 1985

Countries having wild populations

Papua New Guinea - 80 317 745 78 1
Countries without wild populations

Australia - 10 - = = ae
Germany, F.R. 2 6 - 4 - -
Japan - - - - 2 =

288

Ornithoptera caelestis

CONSERVATION MEASURES 0. caelestis is not protected in Papua New Guinea,
partly in recognition of its status, but largely because it is considered as a
subspecies of priamus, the most abundant ornithopteran. Seven less common
species are fully protected. The best way to conserve caelestis,
paradoxically, is to maintain and extend its importance in international
trading. Given sufficient natural habitat and encouragement from farmers who
plant foodplants, the caelestis population on the Louisiades could sustain a
far greater trade than is currently being achieved.

However, it is essential that the PNG government, in recognition of the
growing potential of the insect trade, should put aside managment areas and
protected areas in key sites for caelestis and other birdwing butterflies.
Detailed recommendations for the implementation of such a plan have already
been prepared by Parsons (1985).

CAPTIVE BREEDING It has not proved possible to breed Ornithoptera species
in captivity. Furthermore, it is not particularly desirable to to do so since
this would threaten the PNG ranching programme. As stated above, ranched
specimens are reared ex-pupae from wild parents.

REFERENCES

Anon. (1983). Butterfly Farming in Papua New Guinea. Managing Tropical
Animal Resources Series. National Research Council, National Academy
Press, Washington, D.C.

Collins, N.M. and Morris, M.G. (1985). Threatened Swallowtail Butterflies of
the World. The IUCN Red Data Book. IUCN, Cambridge and Gland;
vii + 401 pp. + 8 col. pls.

D*Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press,
Melbourne. 260 pp.

Fenner, T.L. (1976). Aspects of butterfly conservation in New Guinea. In:
Lamb, K.P. and Gressitt, J.L. (Eds), Ecology and Conservation in Papua
New Guinea. Wau Ecology Institute Pamphlet No. 2, 153 pp.

Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a
phylogenetic approach. Smithersia 2: 1-48.

Haugum, J. and Low, A.M. (1978-1979). A Monograph of the Birdwing
Butterflies. Vol. 1. The genus Ornithoptera. Scandinavian Science
Press, Klampenborg. 308 pp.

Parsons, M. (1985). A conservation study of the birdwing butterflies,
Ornithoptera and Troides (Lepidoptera: Papilionidae) in Papua New
Guinea. Unpublished report to the Department of Primary Industry, Bulolo,
PNG

Pyle, R.M. and Hughes, S.A. (1978). Conservation and utilization of the insect
resources of Papua New Guinea. Consultants' report to the PNG Wildlife
Branch. 157 pp.

289

NEW GUINEA, COMMON, COMMON GREEN OR Recommended list: 3
PRIAM'S BIRDWING {No problem]

Ornithoptera priamus (L., 1758)

Phylum ARTHROPODA Order INSECTA
Class LEPIDOPTERA Family PAPILIONIDAE

SUMMARY AND CONCLUSIONS Ornithoptera priamus, the most widespread species
in the genus, occurs from the Aru Islands in the Moluccas, down to the Torres
Strait Islands and Cape York's Iron Range in Australia. For the purpose of
this paper, priamus is deemed to include the subspecies priamus, euphorion,
hecuba, arruana, poseidon, gebeensis, boisduvali, admiralitatis,
bornemanni, and ~miokensis. Under CITES the taxa richmondia, caelestis
and urvillianus are treated separately. O. priamus maintains high natural
populations because its foodplant, Aristolochia tagala, is a vigorous vine
that occurs wherever forest vegetation is regrowing after disturbance. Many
other Aristolochia spp., and their associated birdwings, only survive in
climax forest. O. priamus is reportedly not threatened, but’ the
restricted subspecies miokensis (Duke of York Is.) is Endangered,
euphorion (Queensland) is Indeterminate, boisduvali (Woodlark Is.) is
Rare, the form demophanes (Trobriand Is.) of the subspecies poseidon is
Rare, the subspecies hecuba (Kai Is.) and arruana (Aru Is.) are cause for
concern and gebeensis is Insufficiently Known. The overwhelming source of
threat to subspecies of concern is destruction of their natural habitat.

The trade in O. priamus (sensu D'Abrera, 1975) apparently involved about
1500 specimens a year between 1980 and 1983, a maximum of 2500 specimens in
1984 and about 5000 in 1985. The bulk of the trade involved the subspecies
poseidon and admiralitatis, neither of which is threatened. Furthermore,
the trade is almost entirely via the Insect Farming and Trading Agency (IFTA)
in Papua New Guinea, a government-controlled organization which buys birdwings
that are reared from pupae by butterfly ranchers. Trading does not represent
a threat to O. priamus. Indeed, the IFTA, through the P.N.G. government,
could be encouraged to expand its operations in deadstock and, more
particularly, to begin trading in livestock (for butterfly houses) An
expansion of the industry to include more species might also be recommended.
An increase in contact between the IFTA and entrepreneurs in retailing centres
might be advisable, but since CITES statistics show that a large proportion of
the trade is with Europe, it is important that the EEC ban on birdwing imports
(Regulation 3626/82) be lifted as quickly as possible (except for the
Endangered O. alexandrae).

Conservation and utilisation of birdwings in Papua New Guinea are mutually
reinforcing. The P.N.G. Government could set up more protected areas as a
source of wild birdwing stock and of course to conserve birdwings and other
forms of wildlife. Trading operations could go hand in hand with conservation
efforts on behalf of threatened subspecies and species of birdwings.
Threatened taxa are described in the Swallowtail Red Data Book (Collins and
Morris, 1985).

Taxonomic note: conservation considerations are confused by nomenclatural
inconsistencies. While there is general agreement as to the taxa or forms to
which the ancestral priamus has given rise in its spread through the Papuan
region, there remains much disagreement as to the precise status of these
forms, i.e. whether they are still only subspecies, or whether their
appearance, behaviour and distribution merit full specific status.

290

Ornithoptera priamus

DISTRIBUTION The genus Ornithoptera (sensu Haugum and Low, 1978-79) is
confined to New Guinea and its offshore islands (part of Indonesia and all of
Papua New Guinea), the Solomon Islands, the Indonesian Maluku Islands
(Moluccas) and the Australian states of Queensland and New South Wales.

Ornithoptera priamus is the most widespread species in the genus, extending
throughout this range, from Ceram and Ambon through New Guinea and the islands
of the Papuan group to the Solomon Islands in the east and N.E. Australia in
the south (Haugum and Low, 1978-79). The taxonomic situation and distribution
of each taxon is summarized in Table 1 below.

POPULATION No precise information is available on densities and total
populations of any subspecies of O. priamus. Indeed, such data are rare for
any butterfly. However, there is general agreement that O. priamus is the
most widespread and successful of the Ornithoptera. It is classified as not
threatened in the IUCN Swallowtail Red Data Book (Collins and Morris, 1985).
The breadth of distribution of the numerous subspecies is so variable that
each needs to be considered separately.

Australia

O. p. euphorion, commonly known as the Cairns Birdwing, appears to have some
populations of a high density. It is recorded as common at various localities
around Cairns itself, but for other historically important locations there are
no recent statements (Haugum and Low, 1978-79). It is generally considered
that euphorion is less numerous than it was previously, although there are
few substantiated records. Haugum and Low, however, state that ‘there is no

doubt ... that at certain localities where the imagines have been collected in
quantity for commercial purposes, the numbers have been drastically reduced in
recent years.' Other populations have also suffered. D'Abrera (1975)

believed that a reduction in the number of adults in the Etty Bay area south
of Cairns is due to over-collecting, although this statement requires
substantiation.

Indonesia

O. p. priamus from Ceram and Ambon in the Moluccas is supposedly common in
mature, but there are few published observations and no recent ones (Haugum
and Low, 1978-79). For the present it is regarded as not threatened.

O. p. gebeensis, only recently described from Gebe island, between Waigeo
and Halmahera (Parrott, 1985), is lacking in all but taxonomic information.
It is classified as Insufficiently Known until such time as its conservation
status on this small, 500 sq. km., island may be properly assessed.

O. p. hecuba from the Kai Islands suffers from a lack of recent data and
little is known of its status, but it appears not to be rare in the coastal
areas of the main islands. It is known to have been abundant early this
century, when various aberrations were described (Haugum and Low, 1978-79).
This subspecies should perhaps be classified as Rare in view of its restricted
distribution. Data on the extent of suitable habitat are needed.

O. p. poseidon see notes under Papua New Guinea.

291

Ornithoptera priamus

Table 1: The systematics, distribution and IUCN category of the
Ornithoptera priamus subspecies listed by Haugum and _ Low
(1978-79). D'Abrera (1975), the authority used by CITES,
recognised three of these as separate species while Hancock (1983)
recognised two as separate species. Collins and Morris (1985), in
the Swallowtail Red Data Book, followed Hancock's classification.
Authors Distribution IUCN
Threat
Haugum D'Abrera Hancock Category
& Low (1975) (1983)
(1978)
1. richmondia x + + Australia (Queensland, I
New South Wales)
2. euphorion x x + Australia (N. Queensland) I
3. priamus x x x Indonesia (Ceram, Ambon) nt
4. gebeensis 0 0 0 Indonesia (Gebe Island) K
5. hecuba x x * Indonesia (Kai, Walim, R?
Laut)
6. arruana x x * Indonesia (Aru, off R?
Halmahera)
7. poseidon x x x Papua New Guinea and nt
Irian Jaya (Waigeo main-
land and adjacent islands),
Australia (Cape York)
8. boisduvali 2s te x Papua New Guinea R
(Woodlark Is.)
9. admiralitatis x x x Papua New Guinea nt
(Admiralty Is.)
10. bornemanni x x x Papua New Guinea nt
(New Britain)
11. miokensis x x x Papua New Guinea E
(Duke of York Is.)
12. caelestis x + x Papua New Guinea nt
(Louisiades)
13. urvillianus x + * Papua New Guinea (New nt

x

+
0

Hanover, New Ireland),
Solomon Is.

recognized by author(s) as a subspecies of O. priamus
recognized by author as a full species
unknown to these authors; described by Parrott (1985)

Various forms of O. p. poseidon are occasionally cited or sold as separate
subspecies.

292

These include:

teucrus from Biak and Schouten

archideus from Waigeo

Ppronomus or macalpinei from Cape York (Iron Range)
demophanes from Ferguson, Trobriands

cronius from SW coast of New Guinea

Ornithoptera priamus

O. p. arruana, from the Aru Islands, is unknown, but it is probably not too
uncommon. Most populations are apparently near the coast (Haugum and Low,
1978-79). The subspecies should perhaps be classified as Rare in view of its

limited distribution; data on habitat are needed to assess the question of
threat.

Papua New Guinea Parsons (1985) reports that the species O. priamus is
locally common to abundant, generally common and very widespread. This is
encouraging since P.N.G. is currently the main exporting country of origin.

O. p. poseidon probably ranges over the whole of mainland New Guinea and the
offshore islands. It certainly occurs from Waigeo in the west to the
Trobriands and D'Entrecasteaux in the east, and south into the extreme
northern Cape York Peninsula of Australia and in the Iron Range (Haugum and
Low, 1978-79). Locally abundant, it is usually found at low or medium
altitudes up to 1500 m, particularly in coastal areas. Generally little is
known of its status, but its range in P.N.G. is now known to span the whole
mainland (Parsons, 1985) and it is certainly the most abundant ornithopteran.
There is a marked lack of information on the populations of poseidon in
Irian Jaya, including all southern districts, most western, and the Indonesian
islands (Haugum and Low, 1978-79), although Parsons (1985) implies that it is
at least as abundant there as it is in the Papuan part of New Guinea. Morris
(in litt., 1986) recently travelled in Irian Jaya and found poseidon to be
very widespread. The subspecies as a whole is therefore listed as not
threatened. However, the form demophanes (sometimes listed as a separate
subspecies) is restricted to the Trobriand Islands, which are generally
heavily populated and deforested. With severe destruction of its habitat and
intermittent heavy collecting pressure, the status of demophanes is a matter
for concern (Fenner, in litt., 1983; Hutton, in litt., 1983). In 1978 it
was reported to be numerous in a small area of Kiriwina, but elsewhere in the
Trobriands to have very little suitable habitat remaining (Pyle and Hughes,
1978).

O. p. boisduvali is endemic to the small island of Woodlark, east of the
P.N.G. mainland and isolated between the South Solomon Sea and the Louisiade
Archipelago. It is very poorly represented in collections, and its status is
unknown, probably fairly rare (Haugum and Low, 1978-79). The P.N.G. Insect
Farming and Trading Agency has organized the collecting of some material since
the 1970s. It is classified as Rare on account of its very restricted
distribution and possible threats to its habitat.

O. p. admiralitatis, from Manus, the main island of the Admiralty group, is
believed to be widespread within the confines of its 80 km-long island, but
rather localised. Ebner (1971) found it to be common, flying throughout the
year. It is not regarded as threatened.

O. p. bornemanni from New Britain and the adjacent islands is reportedly
generally widespread and locally abundant, especially in suitable localities
along the seashores (Haugum and Low 1978-79). Females are said to be not
infrequent in certain coastal coconut plantations. Its present status is
unknown, but it is unlikely to be in serious decline and is classified as not
threatened.

O. p. miokensis is known only from the small group called the Duke of York

Islands. It was once referred to as not rare but is now known to have been
declining in recent years as a result of the high human populations and

293

Ornithoptera priamus

intense agricultural activity in its main habitat areas. Pyle and Hughes
(1978) visited the Duke of Yorks in the course of a consultancy on P.N.G.
butterflies. They concluded that ‘the Duke of Yorks ... have been nearly
obliterated as diverse butterfly habitat.' They considered miokensis to be
threatened with extinction, not from collecting but from sheer displacement.
Believed to be extirpated from its type locality of Mioko and with no suitable
habitat on Milmila, it may be confined to a 50 ha site on one end of Ulu
Island. It is therefore classified as Endangered. There is a possibility
that Miokensis is a hybrid between urvillanus and bornemanni, but so far
no substantiating evidence has been published.

HABITAT AND ECOLOGY Ornithoptera priamus, probably the most successful
species in the genus, has adapted to a range of diverse habitats and is
Oligophagous, feeding on a number of Aristolochia species in the larval
stage but usually on Aristolochia tagala. Aristolochia, commonly called
the Dutchman's Pipe Vine, has a distinctive flower with a single, prolonged
lip-like petal and typically a tubular throat and swollen, rounded, basal
chamber that encloses the stamens etc. The fléwer can be 90 mm in length and
the leaves are also large, being heartshaped and up to 200 mm or so long.
Vines reach up to 10 m but are more commonly 3-5 m high.

O. priamus occurs exclusively in marginal and open secondary forest, where
A. tagala, unlike other species, grows well. <A. tagala grows fast and can
host large numbers of caterpillars (Clark, P.B., in litt., 1986). Habitat
includes forest margins, large regrowth areas within mature secondary or
climax forest and strand vegetation above high tides. OO. priamus flies and
breeds up to 1000 m altitude and adults occur up to 1500 m.

Adults take just over two months to develop from the egg, and may survive for
over 3 months. Females could lay an estimated 630 eggs, at a rate of seven
eggs per day for three months, but lifespans will vary.

Parasitism by encyrtid wasps and predation by Oecophylla ants can be high at
times, but losses of larvae and pupae are relatively low as a result of
sequestration of unpalatable by-products from the foodplant (Clark, P.B., in
litt., 1986). As would be expected, a variety of opportunistic predators
(including man) have been observed, but attacks by chalcid wasps and other
parasitoids are rare (Parsons, 1985).

THREATS TO SURVIVAL The species Ornithoptera priamus is not under threat,
being widespread and abundant in many localities. However, some of the
subspecies give cause for concern (see Table 1 and comments under Populations).

Collecting and trade would not normally represent a threat to healthy and
widespread butterfly populations, and this is certainly the case for
subspecies priamus, poseidon, admiralitatis and bornemanni. Some other
subspecies, notably miokensis and possibly boisduvali, have suffered
setbacks through habitat destruction, often within the already narrow confines
of a restricted vegetation type or small island. In these cases trading needs
to be particularly carefully controlled, but it is at present extremely low
and represents no threat. The subspecies gebeensis, only described in 1985,
could be in demand as a novelty. The situation of this taxon needs to be
investigated and monitored.

INTERNATIONAL TRADE Table 2 shows the minimum net imports of Ornithoptera
priamus reported to CITES, 1980-85. Table 3 is a breakdown of these imports
between the various subspecies. The bulk of the trade was in subspecies
poseidon and admiralitatis, neither of which are under threat. Trade in

294

Ornithoptera priamus

threatened subspecies is minimal. The five major importers were Switzerland,
Federal Republic of Germany, Spain, France and the U.S.A. Theoretically, none
of the EEC countries should have licensed the import of priamus or any other
birdwing during 1983, 1984 or 1985 because of the inclusion of birdwings in
Annex Cl of EEC Regulation 3626/82. In this regulation to implement CITES,
all CITES Appendix II butterflies were effectively treated as Appendix I, thus
prohibiting import. All birdwings except Ornithoptera alexandrae were
removed to Annex C(2) of the regulation during 1987.

Table 2: Minimum net imports of Priam's birdwing (Ornithoptera priamus)
butterfly bodies (all subspecies) reported to CITES, 1980-1985.

Australia - = 4 48 36 5
Canada - - - 1 16 1
Colombia - = = = = 25
France - - - 243 - -
Germany, F.R. ~ - 371 75 - -
Ireland - - - 20 - -
Italy - - - 6 = =
Japan ~ - - 34 29 106
Norway - - - 2 - -
Singapore - - - 9 - -
Spain - 250 - 10 - -
Sri Lanka = = - - 1 -
Switzerland 59 94 38 21 23 6
UK - - - 82 11 3
USA - - - 451 8 28
Vanuatu - - - 2 - -
Unknown - - - 332 - -
Total 59 344 413 1336 124 174

Table 3: Breakdown of imports of Ornithoptera priamus into subspecies:

Subspecies 1980 1981 1982 1983 1984 1985
euphorion - 4 - - 2 34
hecuba - - - - 2 -
arruana - - - - 2 -
poseidon 1 - 228 1080 111 114
admiralitatis - 8 186 256 5 -
bornemanni - - 1(?) - = =
miokensis - - 1(?) - = =
Unknown 58 332 - - 2 26
Total 59 344 413 1336 124 174

295

Ornithoptera priamus

Table 4 gives the reported countries of origin, 1980-85. Reported export of
euphorion from Australia was quite low, probably because the taxon is
protected in Queensland and only available for export when accompanied by an
expensive licence.

No exports are reported from Indonesia, where O. priamus is protected (see

below). However, ‘protection’ implies that collecting is permitted under
licence. Only ‘strictly protected’ species may not be taken under any
circumstances. Indonesia must have been the origin of at least the two

specimens each of hecuba and arruana.

Papua New Guinea is the only significant exporter of Ornithoptera priamus
during the 1980s. Unfortunately, P.N.G. did not produce proper reports to
CITES in 1980, 1981 or 1982, so that CITES statistics are unreliable for those
years. However, from information on file at CMC, between 1400 and 1600
specimens were believed to have been exported by the Insect Farming and
Trading Agency (IFTA) during that year. Data for the year 1985 indicate that
sales of priamus through IFTA amounted to 2422 specimens (Clark, P.B., in
litt., 1986). This figure does not include a further 2383 specimens of
urvillianus and 932 specimens of caelestis and it is far higher than the
number reported to have been exported in the P.N.G. report to CITES for 1985.
Commercial exports for 1986 were reported to have included: 6201 poseidon;
695 admiralitatus; and 57 bornemanni (P.N.G. CITES MA, 1987). Fenner (in
litt., 1976) considered the world market could absorb 5000 pairs per year of
poseidon alone.

Table 4: Reported countries of origin (or exporting country if no original
source reported) and quantities of transactions in dead specimens of
Ornithoptera priamus.

1980 1981 1982 1983 1984 1985

Countries having wild populations:

Australia - 46 - 2 - 79
P.N.G. * - 250 413 1337 110 94
(1500) (1400) (1600)

Countries without wild populations:

Germany, F.R. 59 48 10 7 10 -
France - - = = ry a=
Philippines - - = = = 1
x Papua New Guinea did not report its exports to CITES in 1980-1982.

Figures in brackets are estimated from information held in CMC files.

The implication is that these figures are well within the limits that wild
populations can sustain and that trade in priamus_ subspecies during
1980-1985 posed no threat whatsoever to wild populations. Indeed, the
controlled marketing of birdwings by the IFTA is likely to encourage
conservation of natural habitat as a source of butterflies. This is essential
to sustain the birdwing butterfly ranching programme run by the IFTA, as well
as its other trading activities. About 30 per cent of all butterflies

296

Ornithoptera priamus

reaching the IFTA are ranched, the rest are collected as adults in the wild.
However, over 50 per cent of revenue is from the better quality ranched
specimens (National Research Council, 1983). Collecting of wild adult
birdwings is currently discouraged by IFTA because the specimens are
invariably damaged and are thus of little value (Clark, P.B., in litt.,
1986).

In recent years there has been a growing demand for birdwing pupae that can be
reared and released in ‘butterfly houses’ in temperate countries. Essentially
these places consist of glass-houses containing tropical plants, in which
butterflies fly freely whilst visitors walk around and watch, photograph, or
even touch them. The benefits in educational terms are very high, while
reductions in wild populations are minimal. For these reasons the trade in
livestock is to be encouraged.

CONSERVATION MEASURES

Australia The species Ornithoptera priamus is_- protected under the
Queensland Fauna Conservation Act of 1974. The intention of the law was
undoubtedly to protect all three Australian taxa, poseidon, euphorion and
richmondia, notwithstanding changes of taxonomic opinion since the law was
passed. Specimens may be collected only under licence. Although
over-collecting has undoubtedly been a problem in some areas, particularly for
euphorion and richmondia, habitat destruction probably constitutes the
gravest danger in the long term. Parsons (1985) is of the opinion that of all
the priamus subspecies only the Australian euphorion and richmondia
(which Parsons includes es a subspecies of priamus) warrant conservation
concern. There is, however, sufficient evidence to warrant concern for
miokensis too (see above).

Indonesia Indonesia has included O. priamus in its list of protected
butterflies, published in a decree of 1980. As stated above, this means that
butterflies may only be taken under licence. In effect this restricts world
trade in the subspecies gebeensis, hecuba and arruana, endemic to Gebe
Island, the Kai and Aru Islands respectively. It also restricts trade in
poseidon from Waigeo, Irian Jaya and the offshore islands, leaving the
market almost entirely to Papua New Guinea. The effect of this decree could,
in the long term, be counter-productive. Whilst hindering the development of
farming and ranching of this and other birdwing species, it does nothing to
protect habitat. Indonesia is currently developing a system of protected
areas for Irian Jaya; a marine reserve is proposed for the Aru Islands, and
the proposed Kai Besar Reserve will cover 37 000 ha. It remains to be seen
how effectively these protected areas will benefit birdwing butterflies.

Papua New Guinea P.N.G. protected seven species of Ornithoptera in 1968,
but these did not include priamus or any of its subspecies. The government
realised that 0. priamus and fTroides oblongomaculatus were common and
therefore did not warrant protection (Parsons, 1985). These two species are
the cornerstones of the P.N.G. butterfly ranching programme. More recently,
it has been proposed that O. victoriae, and possibly O. goliath, should be
removed from the protected list and brought into trade. So far, however, no
changes have been made.

In a recent confidential report to the P.N.G. Government, M.J. Parsons has
proposed a complex of protected sites throughout P.N.G., with a view to
safeguarding the -rarer_ species such as O. alexandrae, 0. chimaera,
O. paradisea and others. No response to the report has been made eveilable
to CMC.

297

Ornithoptera priamus

CAPTIVE BREEDING It has not proved possible to breed Ornithoptera species
in captivity. Indeed, it is not particularly desirable to to do so since this
would threaten the existing P.N.G. ranching programme as well as incipient
programmes in Irian Jaya and the Solomon Islands. As stated above, ranched
specimens are reared ex-pupae from wild parents.

REFERENCES

Anon. (1983). Butterfly Farming in Papua New Guinea. Managing Tropical
Animal Resources Series. National Research Council, National Academy
Press, Washington, D.C.

Collins, N.M. and Morris, M.G. (1985). Threatened Swallowtail Butterflies of
the World. The IUCN Red Data _ Book. IUCN, Cambridge and Gland;
vii + 401 pp. + 8 col. pls.

D'Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press,
Melbourne. 260 pp.

Ebner, A.J. (1971). Some notes on the Papilionidae of Manus Island, New
Guinea. Journal of the Lepidopterists' Society 25(1): 73-80.

Fenner, T.L. (1976). Aspects of butterfly conservation in New Guinea. In:
Lamb, K.P. and Gressitt, J.L. (Eds), Bcology and Conservation in Papua
New Guinea. Wau Ecology Institute Pamphlet No. 2, 153 pp.

Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a
phylogenetic approach. Smithersia 2: 1-48.

Haugum, J. and Low, A.M. (1978-1979). A Monograph of the _ Birdwing
Butterflies. Vol. 1. The genus Ornithoptera. Scandinavian Science
Press, Klampenborg. 308 pp.

Parrott, R.E. (1985). A new subspecies of Ornithoptera priamus, Linn. from
Gebe Island, Indonesia. Papilio International 2(3-4): 131-142.

Parsons, M. (1985). A conservation study of the birdwing butterflies,
Ornithoptera and fTroides (Lepidoptera: Papilionidae) in Papua New
Guinea. Unpublished report to the Department of Primary Industry, Bulolo,
P.N.G.

Pyle, R.M. and Hughes, S.A. (1978). Conservation and utilization of the insect
resources of Papua New Guinea. Consultants’ report to the PNG Wildlife
Branch. 157 pp.

298

MANUS GREEN TREE SNAIL Recommended list: 2

[Possible problem]
Papustyla pulcherrima Rensch, 1931

Phylum MOLLUSCA Order STYLOMMATOPHORA
Class GASTROPODA Family CAMAENIDAE

SUMMARY AND CONCLUSIONS This well known brilliant green tree snail is
endemic to the rain forest of Manus Island, northern Papua New Guinea. Its
restricted range makes it particularly vulnerable to large-scale deforestation
which is not a problem at present but could easily become one as there is no
protection of the Manus Island forests. Papua New Guinea is considering
passing protective legislation for this species. Listed as Rare in the IUCN
Invertebrate Red Data Book and as Endangered under the U.S. Endangered Species
Act, it is much sought after by shell collectors. Although there is little
information available, this species, like other Pacific tree snails, is
probably vulnerable to over-collection.

It is known that shells are exported to Japan, Australia and probably Europe,
both as commercial consignments and in personal effects but, despite CITES
listing, the trade has been very poorly monitored. A shipment of 300 shells
was exported to the UK in 1981 via Australia and 1072 were reportedly exported
to Thailand in 1985; there have been no other records of commercial shipments.

A study of the distribution, ecology and population of this species is
urgently required to provide the data for constructive recommendations for
management of the trade. This must be one of the easiest CITES-listed species
for a customs officer to identify because of the unique colouration of the
shell and accurate reporting of the trade should therefore be possible.
Monitoring of the trade should be improved and the number of specimens leaving
the country in personal as well as commercial consignments should be checked.

DISTRIBUTION Manus Island, Admiralty Archipelago, northern Papua New Guinea
(Clench and Turner, 1962). Few professional malacologists have collected
specimens and those in museums have been obtained mainly by traders, explorers
or incidentally by collectors of other animal material. Precise locality data
are therefore poor (Clench and Turner, 1962). The snail is known to occur
several kilometres inland from Lorengau, the provincial capital, and can be
collected along the main highway (Harrison Gagne, 1981; Wells, 1982) but may
well occur in other less accessible areas of the forest. Frequently referred
to under the old name of Papuina pulcherrima.

POPULATION Unknown.

HABITAT AND ECOLOGY No scientific studies have been carried out on this
species, and the only information available is from anecdotal accounts. The
snail is restricted to rain forest. The main host trees are Dillenia
(Dilleniaceae) and Astronia (Melastomataceae), although it may also be found
on other species and on large climbing aroids. Only a few individuals are
found in each tree. The snails are inactive during the day and are found
attached to the undersides of leaves, five or more metres above the ground
(Harrison Gagne, 1981).

299

Papustyla pulcherrima

THREATS TO SURVIVAL The Manus Green Tree Snail appears to be relatively
host specific to trees with timber potential, and logging is almost certainly
the main threat. In 1977 the snail was said to have retreated into the wilder
central forests of the island and populations were thought to have been much
reduced or eliminated by collecting and agriculture within a several mile
radius of Lorengau (Pyle, in litt., 1980). However, specimens were found
quite easily near Lorengau in 1981 and there was little evidence of heavy
collection. New roads were being opened up through the forest which could
pose a potential threat through loss of habitat and increased accessibility

for collectors. It has been reported that branches are cut down to obtain
single snails (Harrison Gagne, 1981) but this seems unlikely as collectors
generally climb the trees (Wells, 1982; Pitman, 1977). Nevertheless,

collecting could pose a threat, given the restricted range of this species,
and the possibility that the population is small and slow to replace itself.

INTERNATIONAL TRADE The shells are still used by Manus islanders for
decorative purposes (Harrison Gagne, 1981). Its intense green coloration
makes Papustyla pulcherrima unique among land molluscs and as a result it is
highly prized by shell collectors. The shell is often used in jewellery. In
the 1970s collection was said to be heavy with large numbers being bought by
tourists and exported by dealers (Pitman, 1977). However, the commercial
export trade seems to have declined. Between January and June 1981, 154
shells were exported by the New Guinea Shell Agency. The decline was partly
due to problems in the running of the shell agency; collectors were not
receiving their payment and the agency staff showed little interest in
promoting the business. 834 specimens had been collected and deposited at the
Lorengau Department of Business Development that year but were not being
forwarded to the agency until payment for past consignments was received
(Wells, 1982). The current status of the agency is not known. The following
figures were reported to CITES between 1980 and 1984:

Table 1 Minimum net imports of Papustyla pulcherrima shells reported by
CITES Parties. All of this trade reportedly originated in PNG.

1980 1981 1982 1983 1984* 1985
Australia - = = 1 = ee
Thailand - - - - - 1072
UK - 300 - 1 = a
USA - al - 8 = =

PNG did not supply annual reports to CITES for the years 1980-1982.

* 17 640 carvings of this species were reportedly imported by USA from Taiwan
in 1984 - this is almost certainly an error in the CITES report and this
figure probably refers to black coral, Antipatharia; it is most unlikely that
such large numbers of shells are being exported from Papua New Guinea and the
import would presumably have been illegal.

Despite the decline in the trade, the CITES reports clearly underestimate the
total international trade. For example, considerable numbers of shells go to
Japan (Wells, 1982; Kwapena in litt., 1985) and large numbers were on sale
in Australia in 1979 (Bruggen in litt., 1981).

300

Papustyla pulcherrima

CONSERVATION MEASURES It is listed as endangered under the U.S. Endangered
Species Act and imports into the USA are prohibited.

Logging poses the greatest threat to this species at present and reserves
should be created in appropriate sites to provide protection for snail
populations should large-scale logging commence. Papua New Guinea is
considering passing protective legislation for this species (Kwapena, in
Itt... , 1985). If the current trade is no larger than it appears, it is
unlikely to be having any major impact on populations, but it is possible that
there is a significant amount of unrecorded trade. Monitoring of the trade
should be improved, and the number of specimens leaving the country in
personal as opposed to commercial consignments should be checked. This must
be one of the easiest CITES-listed species for a customs officer to identify
because of the unique coloration of the shell and accurate reporting of the
trade should therefore be possible. A thorough study of the distribution,
ecology and population of this species is urgently required to provide the
data for constructive recommendations for management of the trade. In 1981,
local people and expatriates were largely ignorant of the CITES listing of
this species and the need for export permits (Wells, 1982), and there was a
clear requirement for a public awareness campaign. The Manus Green Snail is
only one of a number of land and tree snails being collected for export in
Papua New Guinea (Wells, 1982). Since most of these are barely known to
science the opportunity should be taken of obtaining live specimens for
anatomical studies, and surveys and ecological studies should be carried out
to determine more precisely their distribution patterns and population biology.

CAPTIVE BREEDING No attempts at captive breeding known but at one time
Manus villagers attempted to ‘farm' snails by collecting them from the wild
and raising them in large cages; this was unsuccessful.

REFERENCES

Clench, W.J., and Turner, R.D. (1962). Monographs of the genera Papustyla,
Forcartia, and Meliobba (Papuininae: Camaenidae). The Malacological
Society of Australia.

Harrison Gagné, B. (1981). Up a tree with the Manus green snail. Hawaiian
Shell News 24(5) N.S. 257:1, 8-9.

Pitman, R.W. (1977). Manus Island's green tree snails at home. Hawaiian Shell
News 25(4), N.S. 208: 9-10.

Wells, S.M. (1982). Marine Conservation in the Philippines and Papua New
Guinea with special emphasis on the ornamental coral and shell trade.
Report to Winston Churchill Memorial Trust, London.

301

BLACK (WHIP) CORAL Recommended list: 2
{Possible problem]
Cirrhipathes anguina Dana

Phylum CNIDARIA Order ANTIPATHARIA
Class ANTHOZOA Family ANTIPATHIDAE

SUMMARY AND CONCLUSIONS Cirrhipathes anguina is a whip coral with a wide
Indo-Pacific distribution and is found attached to rocks and coral reefs,
often at shallow depths unlike many black corals. Data on the ecology,
abundance and distribution of all black and whip corals, found in tropical and
subtropical waters in all major oceans, are scanty. The order Antipatharia is
listed in the IUCN Invertebrate Red Data Book as Commercially Threatened and
on Appendix II of CITES. Black corals are protected in a number of countries
in the Caribbean but as yet few Indo-Pacific species receive protection.

C. anguina accounts for most international trade as it is used exclusively
for the making of bracelets which are probably the most popular and cheapest
items in the precious coral market. It is exported to the U.S.A. in large
quantities, mainly from Taiwan and the Philippines, but there is no
information on collecting localities. There are no specific reports of
depletion of C. anguina, mainly because no studies have been carried out,
but there have been many general reports of over-collection of local
populations of black corals in shallow waters.

As a result of CITES reporting, the importance of C. anguina has _ been
revealed and efforts should now be made to study the ecology, distribution and
exploitation of this species in order to make recommendations for its
sustainable use.

Taxonomic note: Classification and identification of the 150 known species of
black coral is complicated. The genus Antipatharia includes the true black
corals, which form branched plant-like colonies, and the whip or wire corals
such as C. anguina which form unbranched stick-like structures which are
sometimes coiled. C. anguina is generally unbranched and may reach a length
of one metre or more. Colonies are strengthened by a brown or black skeleton
of horny material; the polyps which are situated in the living tissue around
this skeleton are short and cylindrical with six tentacles which cannot be
retracted. The lower end of the colony is attached to a firm substrate by a
flattened base or may simply extend into the sediment (George and George,
1979; Grigg, 1976; Warner, 1981).

DISTRIBUTION Black coral is found in tropical and subtropical waters in all
major oceans. The distributions of individual species are poorly known.
C. anguina occurs from 30-500 m throughout the Indo-West Pacific (George and
George, 1979).

POPULATION Unknown.

HABITAT AND ECOLOGY Black corals are usually found in deep water, between
30-110 m although some have been recorded from depths of 2000-3000 fathoms
(4000-6000 m). A number of species however are found in shallower water. For
example, off Curacao, black corals are found mainly between 15 and 50 m (Noome
and Kristensen, 1975). C. anguina also occurs at shallow depths (10-20 m)
which accounts its widespread use in trade. It is found attached to hard

302

Cirrhipathes anguina

substrates such as rocks or corals and is frequently present on coral reefs.
Work on branching black corals has shown that colonies are frequently
associated with terraces and undercut notches. Adult colonies can withstand
light intensities of up to 60 per cent of the surface incident light, but it
appears that larvae cannot settle and survive under a light intensity greater
than 25 per cent of the surface light. This would account for the greatest
densities being found below 35 m, colonies being found at shallower depths
only in turbid water or in shaded areas (Grigg, 1965; Grigg, 1976). The
normal method of feeding is probably trapping of animal plankton by the
nematocysts (Warner, 1981). Polyps are either male or female but colonies may
be hermaphroditic (George and George, 1979). Reproductive maturity in the
black coral Antipathes dichotoma is probably reached between 10 and 12.3
years and the reproductive cycle may be annual. Asexual reproduction occurs
naturally by fragmentation of branch ends. Colonies may have a life span of
70 years (Grigg, 1976). Further research is required to determine to what
extent the whip corals such as C. anguina have a similar life history.

THREATS TO SURVIVAL Black corals have been locally depleted in many areas
in shallow water, but deeper populations are unlikely to be exploited to
extinction due to their inaccessibility. Caribbean species have _ been
particularly heavily exploited (Castorena and Metaca, 1979; Goldberg, 1981;
Noome and Kristensen, 1975; De la Torre, 1978; Wells et al., 1983).
Extrapolating from the problems that have arisen in the Caribbean, it is
highly likely that local populations of C. anguina in south-east Asia are
being depleted by the trade. Black Corals may also be affected by habitat
disturbance, particularly species such as C. anguina which are found on
coral reefs.

INTERNATIONAL TRADE In the Middle and Far East black coral has long been
accredited with medicinal and magical properties (Hickson, 1924) but such
traditional use probably has little impact on populations. More recently,
black coral has become a very popular material for jewellery which is either
made for export or sold to tourists. Identification of the raw material to
species level is difficult and is almost impossible with worked or
Manufactured products; black coral, both raw and carved, is probably regularly
shipped under the wrong name. Furthermore, synthetic ‘Black Coral’ is often
sold to unsuspecting tourists (Grigg and Eade, 1981). No statistics have
previously been available for the black coral trade but figures reported to
CITES now give some indication of its level. However, since 1982, when
Antipatharia were listed on CITES, only the USA has regularly reported
C. anguina shipments to CITES (Table 1).

Table 1. Origin countries of imports of C. anguina reported to CITES
1981-1985 by the USA. Numbers of ‘carvings’ - C, and ‘raw corals’ - R.

1980 1981 1982 1983 1984 1985

Philippines Cc - - 1073 162 5830 2657

Taiwan (% - - 157708 190903 158150 167313
+ 52 kg

R - - 1000 - - 2263

Total G - - 158781 191065 163980 169970
+ 52 kg

Eee 2 u = = 2263

eee

303

Cirrhipathes anguina

In addition, consignments of black coral are reported to CITES under the
heading ‘Antipatharia’ and some of these, from countries in the Indo-Pacific,
may include C. anguina (Table 2). The USA is the only country which has
reported significant quantities of trade in Antipatharia, the only exceptions
being Japan, which reported imports of 1000 kg of raw corals from Taiwan in
1982, and Switzerland which imported 1 carving from Italy in 1985 which
reportedly originated in Taiwan.

Table 2. Indo-Pacific countries of export or origin of Antipatharia carvings
and raw corals reported to CITES, 1981-1985 by the USA.
C = carvings, R = raw corals.

1980 1981 1982 1983 1984 1985

Fiji Cc - - 3 - - 12
Hong Kong Cc - - 7346 1350 - -
R - - 492 - - -

Indonesia R - - - 24 - -
Kenya Cc - - - 127 - -
Korea Cc - - 2500 - 13 -
Philippines C - 6869 4726 198 619 7437
R - 117 190 - - -

Taiwan Cc - 5242 169586 134394 24156 147576
R - - 9657 3042 - -

Tonga Cc - - - - 576 93
Total Cc - 12111 184161 136069 25364 155118
R - 117 10339 3066 - -

According to CITES statistics, the USA is a major importer of carved
C. anguina from Taiwan and the Philippines. Unfortunately there is no
information on the operation of the trade in the countries of origin.
Japanese and Taiwanese fishing boats were reported to be taking black coral
from Papau New Guinean waters in the early 1980s and it is known that
C. anguina occurs here. A local fishery provides corals to the main
jewellery firm in the country and coral products are exported mainly to
Australia (Wells, 1982). Black Coral is also harvested commercially in Hawaii
(Grigg, 1976) and small quantities are collected on Guam (Hedlund, 1977) but
these do not include C. anguina. In 1982 the Black Coral harvest in Hawaii
was about 50 per cent lower than in previous years on account of a depressed
market and past stockpiling. Current levels are significantly below the
estimates of a maximum sustainable yield, and in this area at least there is
no danger of over-collection (Grigg, in litt. 5.8.82).

CONSERVATION MEASURES No countries specifically protect C. anguina. The
entire order Antipatharia is listed on Appendix II of CITES and is included in
the IUCN Invertebrate Red Data Book under the category Commercially Threatened
(Taxa not currently threatened with extinction but most or all of whose
populations are threatened as a sustainable commercial resource, or will
become so unless their exploitation in regulated). The order Antipatharia is
listed under the Netherlands legislation for protected non-native species
(Anon, 1980) and black coral is protected in New Zealand (Richardson, 1981).

304

Cirrhipathes anguina

In Hawaii permits are issued for limited commercial harvesting and there is a
minimum size limit of 1.2 m in height and/or 2.5 cm in stem diameter (Grigg,
1976). Caribbean species of black coral receive varying degrees of protection
in Antigua, Barbados, Belize, the British Virgin ‘Islands, Florida (USA),
Jamaica, Mexico, Netherlands Antilles, Trinidad and Tobago, U.S. Virgin
Islands (Wells et al., 1983).

A number of management studies have been or are being carried out on black
corals but these usually concern the branching Antipathes species. For
example, in Hawaii, an optimum annual sustainable yield of 5000 kg has been
estimated for the stock of 84 000 colonies in the Auau Channel (Grigg, 1976).
Management studies have been carried out in Papua New Guinea (Tarr, 1978),
Guam (Hedlund, 1977), the Galapagos (Robinson, 1982) and Mexico (Castorene,
1979) and projects are underway in Tonga (Anon., 1985) and New Zealand
(Grange, 1985).

There is an urgent need for study and monitoring of the black coral trade. As
a result of CITES reporting, the importance of C.-anguina has been revealed
and efforts should now be made to study the ecology, distribution and
exploitation of this species in order to make recommendations for its
sustainable use. A number of studies have been carried out on branching black
corals which provide models for the investigation and management of the
Cirrhipathes fisheries. Since C. anguina appears to be of particular
importance in the trade and may prove to be as vulnerable to intensive
collection as the branching black corals, the need for appropriate information
is considered a high priority. The potential for planting cuttings of
C. anguina has been investigated in Papua New Guinea but no results have
been published (Wells, 1982). This is an area of research which would be of
particular interest to the trade.

REFERENCES

Anon. (1985). Sustainable black coral harvesting potential in Tonga. Paper
presented at 3rd South Pacific National Parks and Reserves Conference and
Ministerial Meeting 24 June - 3 July, Apia, Western Samoa, Ministry of
Lands, Survey and Natural Resources.

Castorena, V. (1979). Coral Negro - una possible estrategia. Tecnica
Pesquera: 20-21.

Castorena, V. and Metaca, M. (1979). El Coral Negro, una riqueza en peligro.
Tecnica Pesquera: 22-27.

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